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► 


AS 


I 


\     ^ 


ttNERAL  LIPPARY, 

UNIV.  Oh  AUCIl. 

M  S4190B 


PROCEEDINGS 

OF  THE 

AMERICAN    PHILOSOPHICAL   SOCIETY 

HELD  AT  PHILADELPHIA 

FOR    PROMOTING   USEFUL   KNOWLEDGE 


Vol.  XLVIL  January-April,  1908.  No.  188. 


CONTENTS. 

Stated  Meeting,  January  j,  igoS i 

Stated  Meeting,  January  I'/ 2 

Tumor  Growth  and  Tissue  Growth.     By  Leo  Loeb 3 

Stated  Meeting,  February  7 , 12 

Stated  Meeting,  February  2 j 12 

Stated  Afeeting,  March  6 13 

Stated  Meeting,  March  20 13 

Stated  Meeting,  April  j 13 

Genera/  Meeting,  April  2j,  24  and  25 13 

The  Law  of  Orestne,  Copernicus  and  Gresham.     By  Thomas  Will- 
ing Balgh iS 

Art  and  Ethnology.     By  Edwin  Swjft  Balch 30 

The  Brain  of  Rhinochimaera.     By  Burt  G.  Wilder » 37 

Observations  Regarding  the    Infliction  of  the  Death    Penalty  by 

Electricity.     By  Edw.  Anthony  Spitzka 39 

Preliminary  Note  on  the  Brains  of  Natives  of  the  Andaman  and 

Nicobar  Islands.     By  Edw.  Anthony  Spitzka 51 

Determination    of    Dominance    in    Mendelian    Inheritance.      By 

Charles  B.  Davenport : 59 

The  Santa  Cruz  Typotheria.     By  William  J.  Sinclair 64 

Notes  on  Some  Chilean  Copper  Minerals.     By  Harry  F.  Keller.,  79 
Progress  of  the  Demarcation  of  the  Alaska  Boundary.     By  O.  H. 

TlTTMANN 86 

The  Most  Primitive  Living  Representative  of  the  Ancestors  of  the 

Plant  Kingdom.     By  George  T.  Moore 91 

The  Comparative  Leaf  Structure  of  the  Sand  Dune  Plants  of  Ber- 
muda.     By  John  W.  Harshberger  97 

Solution  of  Algebraic  Equations  in  Infinite  Series.     By  Pkeston  A. 

Lambert 11 1 

Stated  Meeting,  May  i 134 

Stated  Meeting,  May  l^ 134 

The  Origin  of  Bombshell  Ore.     By  H.  M.  Chance 135 


PIIILADELPHIA 

THE   AMERICAN    PHILOSOPHICAL   SOCIETY 

104  South  Fiitu  Strekt 
1908 


Members  who  have  not  as  ^et  sent  their  photographs  to  the  Societj^  will 
confer  a  favor  hy  so  doing;  cabinet  size  preferred. 


It    is  requested  that  all  correspondence  be  addressed 
To  THE  Secretaries  of  the 

AMERICAN   PHILOSOPHICAL   SOCIETY 
104  South  Fifth  Street 

Philadelphia,  U.  S  A. 


PROCEEDINGS 

OF     THE 

American  Philosophical  Society 

HELD    AT    PHILADELPHIA 

FOR 

PROMOTING  USEFUL   KNOWLEDGE 

VOLUME   XLVII 
1908 


PHILADELPHIA 
THE  AMERICAN   PHILOSOPHICAL  SOCIETY 


3^    V 


PROCEEDINGS 

OP  THX 

AMERICAN  PHILOSOPHICAL  SOCIETY 

HELD  AT  PHILADELPHIA 

FOR  PROMOTING  USEFUL  KNOWLEDGE 
Vol.  XLVII  January-April,  1908.  No.  188- 

Stated  Meeting  January  3,  IQ08, 
Treasurer  Jayne  in  the  Chair. 

The  decease  of  the  following  members  was  announced : 

Dr.  Coleman  Sellers,  at  Philadelphia,  on  December  28,  1907, 
set.  80. 

Prof.  Thomas  Day  Seymour,  at  New  Haven,  on  December  31, 
1907,  set.  59. 

The  judges  of  the  annual  election  of  ofiicers  and  councillors  held 
on  this  day,  between  the  hours  of  two  and  five  in  the  afternoon, 
reported  that  the  following  named  persons  were  elected,  according 
to  the  laws,  regulations  and  ordinances  of  the  Society,  to  be  the 
ofiicers  for  the  ensuing  year. 

President: 

William  W.  Keen. 

Vice-Presidents: 
George  F.  Barker,        William  B.  Scott,        Simon  Newc(Mnb. 

Secretaries: 

I.  Minis  Hays,  ^  James  W.  Holland, 

Arthur  W.  Goodspeed,         Amos  P.  Brown. 

19293'7 


2  MINUTES. 

Curators: 
Charles  L.  DooHttle,  WilUam  P.  Wilson,         Leslie  W.  MUler. 

Treasurer: 
Henry  La  Barre  Jayne. 

Councillors: 

(To  serve  for  three  years.) 

Hampton  L.  Carson,  Harry  F.  Keller, 

Talcott  Williams,  Francis  B.  Gummere. 


Stated  Meeting  January  ly,  iqo8. 
Councillor  Rosengarten  in  the  Chair. 

A  letter  was  received  from  the  American  Institute  of  Electrical 
Engineers,  inviting  the  Society  to  be  represented  at  the  Memorial 
Exercises  in  honor  of  Lord  Kelvin,  to  be  held  on  January  12,  at 
3  P.  M. ;  and  from  Vice-President  Scott  announcing  the  appoint- 
ment of  Mr.  Andrew  Carnegie  and  Professor  Michael  L  Pupin  to 
represent  the  Society  on  the  occasion. 

A  letter  was  received  from  the  Committee  of  Organization  of  the 
First  Congress  of  Chemistry  and  Physics  in  memory  of  the  cele- 
brated Russian  Chemist,  Mendeleeflf,  announcing  that  the  Congress 
will  be  held  at  the  University  of  St.  Petersburg  on  the  second  to 
the  twelfth  of  January,  1908. 

The  decease  was  announced  of  Professor  Charles  Augustus 
Young,  at  Hanover,  N.  H.,  on  January  3,  1908,  aet.  78. 

Professor  Leo  Loeb  read  a  paper  on  *'  Tumor  Growth  and  Tissue 
Growth."    (See  page  3.) 


TUMOR   GROWTH   AND  TISSUE   GROWTH. 

By  LEO  LOEB. 
(Read  January  17,  igo8,) 

In  the  course  of  the  last  five  years,  partly  through  the  aid  of 
their  respective  governments  and  partly  through  private  initiative, 
institutions  have  been  founded  in  the  majority  of  civilized  countries 
for  the  investigation  of  the  causes  and  the  conditions  of  growth  of 
malignant  tumors;  or,  as  briefly  named,  for  the  investigation  of 
cancer.  This  fact  proves  more  clearly  than  anything  else  could  do 
the  widespread  interest  that  has  recently  been  aroused  in  this  part 
of  pathological  research.  Pathological  investigations  share  with 
those  of  other  sciences  a  double  nature.  On  the  one  hand,  their 
problems  are  of  a  practical  character.  Pathology  wants  to  find  the 
causes  of  diseases  and  the  conditions  that  favor  and  inhibit  their 
progress,  in  order  to  lay  a  firm  and  scientific  basis  for  their  cure. 
In  this  respect,  pathology  is  an  ai^lied,  a  technical  science.  On  the 
other  hand,  pathology  desires  to  analyze  the  conditions  that  ulti- 
mately lead  to  death,  in  order  to  recognize  some  of  the  phenomena 
of  life.  In  that  sense,  pathology  is  a  pure  science;  its  aim  is 
philosophical. 

Tempting  as  it  might  be  to  relate  something  of  the  first  attempts 
of  pathology  to  find  the  cause  and  the  cure  of  cancer,  I  shall  here, 
rather,  turn  to  the  purely  theoretical  aspects  of  these  investigations 
and  indicate  some  of  the  results  of  tumor  investigations  that  have 
some  bearing  upon  one  of  the  fundamental  characteristics  of  living 
matter — the  ability  to  grow.  Before  entering,  however,  upon  a  nec- 
essarily very  limited  discussion  of  some  of  the  relations  between 
tissue  and  tumor-growth,  it  might  be  well  to  indicate  what  a  tumor 
is ;  and,  especially,  what  a  cancer  is. 

Perhaps  I  can  best  approach  this  delicate  task  by  stating  some 
varieties  of  growth  that  are  not  included  under  the  term  tumors. 

3 


4  LOEB— TUMOR  GROWTH  AND  TISSUE  GROWTH.     [January  17, 

Our  bodies  consist  of  cells  (that  is  small  parts  of  protoplasm  with 
nuclear  material),  of  products  of  cells  of  different  kinds,  of  decom- 
position products  of  cells  and  of  material  used  for  the  building  up 
of  cells.  Here  we  are  concerned  with  the  two  former  only,  namely, 
with  the  cells  and  their  direct  products.  Now  growth  is  based  upon 
an  increase  in  the  number  or  the  size  of  cells  in  the  locality,  where 
growth  takes  place.  The  increase  in  the  number  of  cells  can  be 
brought  about  in  two  ways :  either  through  the  mult^lication  of  pre- 
existing cells,  or  through  ^  wandering  in  of  new  cells.  Cell-growth 
can  take  place  under  various  conditions.  If  toxic  substances — ^the 
products  of  bacteria  for  instance — or  even  if  inert  substances  foreign 
to  the  body  are  introduced  into  the  organism,  a  certain  proliferation 
of  the  neighboring  cells  and  immigration  of  cells  from  the  blood- 
and  lymph-vessels  take  place.  After  a  certain  period,  such  reactions 
come  to  a  standstill,  and  scar  tissue  develops.  Such  a  cell-prolifera- 
tion we  do  not  call  a  true  tumor ;  but  we  class  it  among  the  inflam- 
matory reactions. 

There  are  other  conditions  in  which  an  unusual  cell-proliferation 
takes  place  in  the  adult  organism ;  in  cases  of  wound  healing.  If, 
for  instance,  a  wound  is  made  in  the  skin,  the  cells  of  the  epidermis 
proliferate  until  the  wound  is  closed;  then  the  additional  prolifera- 
tion ceases.  We  call  this  regenerative  growth.  It  lasts  only  as 
long  as  the  continuity  of  the  epidermis  is  interrupted.  This  is  not 
tumor-growth. 

We  now  come  to  a  third  variety  of  cell-proliferation,  distinct 
from  the  two  former  varieties.  If  a  follicle  of  the  ovary  ruptures 
at  the  time  of  menstruation,  the  follicle  cells  enlarge,  and  proliferate 
much  more  extensively  than  would  be  necessary  in  order  to  insure 
wound-healing.  There  is  formed  a  new  growth,  which  exists  for 
a  limited  period  and  then  disappears.  A  still  more  striking  example 
of  this  new  formation  was  found  in  our  laboratory  in  the  course 
of  the  past  year.  If,  at  a  certain  period  after  cc^ulation  has  taken 
place,  or  at  the  period  of  heat,  the  inner  surface  of  the  uterus  is 
sufficiently  exposed  and  cuts  are  made  in  the  wall  of  the  uterus,  we 
find  that,  instead  of  the  ordinary  wound-healing,  another  process 
takes  place,  namely:  the  development  of  nodules  of  new  tissue, 
which  resembles  closely  the  maternal  part  of  the  placenta — without. 


i9o8.]  LOEB— TUMOR  GROWTH  AND  TISSUE  GROWTH.  5 

however,  an  ovum  being  in  this  case  responsible  for  the  new  forma- 
tion; but  also  in  this  case  the  experimentally  new- formed  decidua, 
as  we  call  this  tissue,  dies. 

The  latter  variety  of  growth  resembles  much  more  closely  the 
real  tumor-growth  than  do  the  former ;  but  in  this  case  also  the  cell- 
proliferation,  and  even  the  life  of  the  newly  formed  cells,  cease, 
when  the  cause  for  the  proliferation  has  disappeared.  The  cause 
for  the  development  of  an  artificial  decidua  is  probably  two- fold: 
in  the  first  place,  a  general  chemical  condition  exists  in  the  body  at 
that  period ;  and,  tmder  these  predisposing  conditions,  a  local  stim- 
ulus suffices  to  produce  the  tumor-like  growth.  These  new  forma- 
tions might  be  called  transitory  tumors,  because  they  have  a  definite 
life-cycle ;  they  grow  for  scmie  time,  and  then  they  disappear. 

In  real  tumors  we  find  a  similar  but  still  more  marked  cell- 
proliferation  ;  and  they  do  not  have  such  a  definite  life-cycle.  Real 
tumors  do  not  retrograde  usually,  and  may  even  g^ow,  more  or  less, 
during  the  lifetime  of  the  bearer.  Furthermore,  we  do  not  know 
the  cause  of  their  origin,  as  we  do  in  the  case  of  the  transitory 
tumor.  They  grow,  and  we  do  not  know  why.  If  such  tumors 
grow  more  rapidly,  and*especially  if  they  grow  deep  into  the  sur- 
rounding tissue,  digesting  it,  if  parts  penetrate  into  the  blood-  or 
lymph-vessels  and  are  carried  away  to  distant  parts  of  the  body,  and 
here  start  a  new  growth,  a  so-called  metastasis,  then  we  call  the 
tumor  malignant,  or  a  cancer. 

We  distinguish  different  varieties  of  cancer,  according  to  the 
tissue  or  variety  of  cells  from  which  these  cancers  originate.  The 
malignant  tumors  derived  from  epithelial  surfaces  or  gland  -cells, 
we  call  carcinomata  and  the  malignant  tumors  derived  from  the 
connective-tissue  cells,  which  unite  the  functionally  more  highly 
develq)ed  cells,  we  call  sarcomata.  But  from  whatever  tissue  these 
malignant  tumors  are  derived,  their  main  characteristics  are  identical. 

During  the  second  half  of  the  last  century,  pathologists  studied 
very  carefully  the  microscopical  character  of  the  different  tumors; 
and  they  determined  quite  accurately  the  genesis  of  these  tumors 
from  normal  tissues.  They  observed  how  cells  began  to  grow  down 
into  the  adjoining  tissues  in  cancer;  they  described  the  general 
spreading  out  of  the  new  formation,  and  the  character  of  the  sec- 


6  LOEB— TUMOR  GROWTH  AND  TISSUE  GROWTH.     [January  17, 

ondary  growth;  they  also  determined  that  a  certain  number  of 
tumors  apparently  originate  in  tissue  that  has  been  misplaced  during 
embryonic  development.  In  other  cases,  long  irritation,  and  occa- 
sionally a  traumatism,  may  be  held  responsible  for  the  origin  of 
cancer.  Apparently,  however,  no  further  progress  could  be  made 
by  these  means  of  observation.  The  investigations  seemed  to  have 
arrived  at  a  dead  point. 

After  a  few  isolated  previous  attempts,  mainly  since  the  year 
1899,  the  attention  of  the  investigators  was  directed  to  the  occur- 
rence of  tumors  in  animals ;  to  the  fact  that  cancer  in  animals  fre- 
quently occurs  endemically.  This  means  that  a  number  of  animals 
are  aflfected  with  cancer  simultaneously  in  a  certain  locality.  Fhr- 
thermore,  they  observed  that  certain  kinds  of  tumors  are  charac- 
teristic for  certain  species  of  animals ;  and  that  the  tumors  occurring 
endemically  in  a  species  of  animals  are  all  of  the  same  type. 

The  most  important  fact,  however,  which  was  fully  developed  only 
within  the  last  eight  years,  is  that  it  is  possible  to  transplant  a  certain 
number  of  cancers  into  other  animals  of  the  same  species.  Many  at- 
tempts have  been  made  to  transplant  cancers  into  animals  of  other  spe- 
cies and  make  them  growl  in  these  animals,  but  without  any  success.  A 
certain  kind  of  cancer  found  in  the  dog  can  be  made  to  grow  in  some 
related  species,  as,  for  instance,  in  the  fox.  Other  tumors  found  in 
white  rats  may  be  transplanted  into  hybrids  between  white  and  gray 
rats,  and  the  cancer  of  white  mice  can  occasionally  be  made  to  grow 
in  gray  mice.  The  cancer  of  a  Japanese  mouse  could  not  be  success- 
fully transplanted  into  white  mice,  however,  but  only  into  the  Japa- 
nese mice.  No  such  tumors  can  be  transplanted  into  more  distantly 
related  animals,  nor  can  the  cancer  of  man  be  transplanted  into  lower 
animals.  A  very  malignant  tumor  from  a  mouse  can  occasionally 
be  made  to  grow  for  a  few  days  in  a  rat,  but  the  growth  soon  stops. 
In  a  similar  way,  normal  tissues  of  the  body,  for  instance  the  epithe- 
lium, may  be  transplanted  into  other  animals  of  the  same  species, 
and  kept  there  alive  after  an  initial  growth ;  but  if  transplanted  into 
an  animal  of  another  species,  it  grows  for  a  short  period  and  then 
it  dies. 

Some  tumors,  and  probably  the  majority  of  them,  can  be  trans- 
planted only  into  the  same  animal  in  which  they  have  originated. 


,9o8il  LOEB— lUMOR  GROWTH  AND  TISSUE  GROWTH.  7 

Here  they  live,  and  even  grow ;  while  in  other  animals  of  the  same 
species,  they  die  very  soon  after  transplantation.  This  probably 
applies  to  most  of  himian  tumors.  The  same  holds  good  of  certain 
animal  tissues  and  organs;  as,  for  instance,  the  ovary.  They  can 
much  more  easily  be  transplanted  into  the  animal  of  which  they 
have  formed  an  integral  part,  than  into  other  animals  of  the  same 
species. 

There  exists  another  point  of  similarity  between  the  transplanta- 
tion of  normal  tissues  and  organs,  on  the  one  hand,  and  of  tumors, 
on  the  other :  in  both  cases,  after  transplantation,  only  the  peripheral 
parts  of  the  transplanted  piece  usually  remain  alive ;  the  central  part, 
which  is  not  well  supplied  with  lymph  or  blood  from  the  host,  soon 
dying.  This  similarity  between  the  behavior  of  normal  tissues  and 
of  tumors  after  transplantation  can  be  easily  explained,  if  we  con- 
sider that  in  both  cases  we  have  equally  to  deal  with  the  inoculation 
of  cells  or  tissues  ivom  an  animal  organism;  and  that  the  trans- 
planted tumor,  as  can  be  readily  shown  by  microscopic  examination, 
grows  merely  from  the  transferred  tumor-cells  themselves,  and  not 
from  the  tissues  of  the  receiving  host-animal. 

On  the  other  hand,  however,  there  exist  also  some  very  interest- 
ing differences  between  the  growth  of  normal  tissues  and  of  tumor- 
tissues  after  transplantation,  the  former  always  growing  only  very 
slowly  for  a  time,  and  then  ceasing  to  grow,  or  merely  remaining 
alive  after  transplantation;  and  the  latter  continuing  to  grow  rap- 
idly, and  sometimes  continuing  to  infiltrate  the  surrounding  host- 
tissue  and  to  make  metastases.  Their  character  is  not  markedly 
modified  through  transplantation.  Eight  years  ago  I  transplanted 
a  sarcoma  of  a  white  rat  into  more  than  forty  generations,  without 
an  appreciable  decrease  in  the  energy  of  growth  of  the  tumor  cells.  The 
fact  that  it  is  possible  to  propagate  tissues  of  the  animal  body  through 
years  and  years  in  other  animals  of  the  same  species,  without  any  loss 
of  vitality  and  power  of  propagation  of  the  tumor-cells,  while  they 
would  long  since  have  died  if  they  had  remained  in  the  animal  to  which 
they  originally  belonged — suggests,  it  seems  to  me,  a  consideration 
of  great  biological  significance,  namely,  the  question  whether  our 
own  body-cells  are  all  equally  mortal,  or  whether  their  death  does 
depend  upon  their  accidental  connection  with  other  cells  and  with 


8  LOEB— TUMOR  GROWTH   AND  TISSUE   GROWTH.     [January  17, 

an  organism  that  dies,  and  because  a  certain  number  of  cells,  espe- 
cially of  nervous  character,  cannot  survive. 

The  inevitable  fate  of  all  metazoan  organisms  is  death;  and 
this  conception  deeply  influenced  all  our  valuations  and  directions  of 
thought,  as  Metchnikoff  only  recently  pointed  out  in  his  book  on  the 
"Nature  of  Man."  Weismann  added  one  consoling  idea:  not  all 
of  our  cells  must  necessarily  die,  but  only  the  so-called  somatic  cells ; 
the  germ-cells,  ova  and  the  sperm-cells,  of  each  individual  may 
propagate  forever,  may  be  immortal.  The  results  of  the  tumor 
investigations  just  mentioned  may,  perhaps,  enlarge  the  number  of 
cells  that  may  remain  alive  for  so  long  a  period  that  we  cannot  see 
the  end  at  present;  ordinary  somatic  cells  may  propagate  through 
many  generations,  long  after  their  brother  cells  that  remained  in 
the  9riginal  organism  have  been  transformed  into  simple  chemical 
substances,  and  who  can  at  present  deny  the  possibility  that  they 
may  have  the  potentiality  of  immortality,  as  well  as  the  germ  cells  ? 
Thus  the  work  on  tumors  leads  us  into  different  realms  of  general 
biology,  and  opens  up  new  fields  that  are  not  without  interest. 

The  experimental  work  on  tumors  has  given  some  other  results 
of  an  unexpected  nature.  One  of  the  great  achievements  of  the  last 
centiiry  was  the  development  of  bacteriological  technique  by  which 
it  is  possible  for  us  not  only  to  cultivate  bacteria  on  artificial  culture- 
media,  but  also  to  influence  markedly  their  behavior,  functions,  vital- 
ity, and  virulence.  It  has  been  found  to  be  possible  to  raise  the  viru- 
lence of  certain  bacteria  by  inoculating  them  into  animals  through 
several  generations ;  on  the  other  hand,  it  is  possible  to  decrease  their 
virulence  by  subjecting  them  to  certain  injurious  chemical  or  physical 
agencies.  Such  a  bacterial  culture  with  artificially  decreased  viru- 
lence has  been  used  as  a  vaccine;  that  means,  as  a  substance  that, 
when  inoculated  into  human  beings  or  animals,  without  causing  the 
disease,  confers  immunity  against  the  virulent  bacilli. 

In  experimenting  with  tumor  cells,  the  surprising  result  was 
obtained  that,  through  successive  transplantations,  by  cutting  out 
pieces  of  tumor,  an  artificial  stimulus  is  given  to  the  tumor  cells, 
so  that  they  begin  to  grow  more  rapidly  and  more  extensively.  In 
other  words,  their  virulence  has  been  increased.  This  is  due  to  a 
direct  stimulating  action  upon  the  tumor  cells,  and  not  to  secondary 


,908]  LOEB— TUMOR  GROWTH   AND  TISSUE  GROWTH.  9 

conditions.  This  explains  a  fact  very  familiar  to  surgeons ;  namely, 
that  after  an  operation  a  recurrent  tumor  is  frequently  more  malig- 
nant than  the  original  tumor. 

But  it  is  also  possible  to  decrease  the  power  of  prq>agation  of 
tumor  cells  without  killing  them  by  exposing  the  cells  to  chemical 
and  physical  injurious  influences,  in  a  way  similar  to  that  pursued 
in  the  case  of  bacteria.  Here,  also,  we  may,  not  without  some  hope, 
look  forward  to  the  preparation  of  some  vaccine  that  may,  some  dSty 
in  the  future,  help  us  to  combat  the  dreaded  disease.  Even  in  this 
case,  however,  tumor  tissue  probably  differs  only  in  degree,  and  not 
in  principle,  from  normal  tissue.  At  least,  this  conclusion  is  indi- 
cated by  the  fact  that  such  an  organ  as  the  normal  thyroid  gland 
may,  without  being  entirely  destroyed,  be  markedly  weakened  in  its 
power  of  growth  through  a  short  exposure  to  the  air  before  trans- 
plantation. 

There  exist,  however,  some  interesting  differences  of  another 
kind  between  timior  tissue  and  normal  tissues  or  organs.  Normal 
organs  have  a  specific  metabolism  and,  in  connection  with  or  as  a 
part  of  this  metabolism,  they  exert  distinct  specific  functions.  We 
understand  by  functions  those  physical  and  chemical  processes  which 
attract  our  attention  by  their  real  or  apparent  significance  for  the 
organism  as  a  whole.  The  normal  female  mammary  gland,  for 
instance,  secretes  milk  under  the  influence  of  certain  chemical 
stimuli  which  are  present  in  the  circulation  at  the  end  of  pregnancy ; 
and  it  also  grows  during  pregnancy,  under  the  influence  of  similar 
stimuli.  If  we  now  transplant  the  mammary  gland  of  a  nonpreg- 
nant animal  into  a  pr^^ant  animal,  the  foreign  transplanted  gland 
may  secrete  milk  at  the  end  of  pregnancy  in  a  similar  way  to  that 
of  the  animal's  own  gland.  The  circulating  chemical  substance 
exerts  the  same  stimulus  upon  the  transplanted  as  upon  the  autoch- 
thonous gland,  and  the  transplanted  gland  responds  to  the  stimulus 
in  the  normal  way. 

There  exist  certain  conditions  in  which  a  tumor-like  hypertrophy 
of  the  mammary  gland  is  found  in  the  white  rat.  The  structure  of 
the  gland  is  slightly  modified,  but  the  tumor  is  not  infiltrating.  We 
do  not  call  it  a  cancer,  but  a  benign  tumor — an  adenoma.  If  we 
transplant  such  a  tumor  to  another  place  in  the  original  animal,  it 


10  LOEB— TUMOR   GROWTH   AND   TISSUE   GROWTH.     [January  17, 

heals,  and  if  the  animal  becomes  pregnant,  it  begins  to  grow  in  the 
same  way  as  the  normal  gland,  but  is  no  longer  able  to  produce  milk. 
It  responds,  therefore,  only  to  certain  stimuli,  but  not  to  others. 

If  we  persist  still  further  and  transplant  a  malignant  tumor,  a 
cancer,  of  the  mammary  gland,  we  find  that  it  no  longer  responds 
to  the  stimuli  of  pregnancy.  Such  tumors  do  not  seem  to  assume 
a  more  rapid  growth,  nor  do  they  ever  secrete  milk.  The  metabo- 
lism of  tumors  diflfers,  however,  only  in  a  greater  or  less  degree 
from  that  of  the  corresponding  normal  tissues ;  and  the  tumor  tissue 
can  even  still  continue  to  secrete  certain  substances  in  a  similar  way 
to  the  normal  tissues.  This  has  been  observed,  for  instance,  in  the 
case  of  the  tumors  of  the  liver  and  of  the  thyroid  gland,  which  latter 
provides  a  so-called  internal  secretion,  without  which  widespread 
changes  would  take  place  in  our  body.  It  seems,  therefore,  in  the 
case  of  the  tumor  tissues  that  there  exists  a  parallelism  between  its 
loss  of  function  and  its  capability  to  respond  to  chemical  stimuli  in 
the  body  that  normally  excite  and  regulate  function  and  growth. 

These  observations  bring  us  also  nearer  to  an  understanding  of 
tissue  growth  and  tumor  growth  in  general.  Just  now  we  men- 
tioned substances  of  various  kinds  circulating  in  the  body  that  rege- 
late the  growth  of  normal  tissues  and  of  tumors ;  but  there  probably 
exist  a  number  of  such  substances.  How  else  could  we  explain  the 
fact  that  the  majority  of  tumors  may  be  successfully  transplanted 
into  the  organism  in  which  the  tumor  had  originated,  but  not  into 
other  individuals  of  the  same  species?  Evidently  there  must  exist 
some  difference  between  the  chemical  composition  of  the  blood  and 
lymph  of  each  individual  of  one  species ;  and  each  tissue  of  one  indi- 
vidual is  more  or  less  adapted  to  its  own  body  fluid.  Furthermore, 
we  have  seen  that  tissues  do  not  grow  in  animals  belonging  to  dif- 
ferent species;  there  must,  therefore,  exist  substances  regulating 
growth,  which  are  the  same  in  the  same  species,  but  differ  in  dif- 
ferent species.  Sometimes,  however,  certain  families  of  white  mice 
differ  among  one  another  to  a  higher  degree  than  the  white  mice 
differ  from  gray  mice. 

Such  substances,  however,  can  merely  regulate  the  growth  of 
normal  tissue  and  of  tumor  tissue;  they  are  not  able  to  transform 
normal  tissue  into  tumor  tissue.    How  the  latter  transformation  is 


1908.]  LOEB— TUMOR  GROWTH  AND  TISSUE  GROWTH.  11 

brought  about,  we  do  not  yet  know ;  and  this  is  one  of  the  problems 
that  remain  before  us.  Of  one  fact  we  may  be  reasonably  certain ; 
namely,  that  the  growth-regulating  substances  to  which  we  referred 
just  now  are,  in  all  likelihood,  not  the  primary  factors  in  the  produc- 
tion of  tumors.  We  draw  this  conclusion  because  the  action  of  such 
substances  has  so  far  not  been  shown  to  be  hereditary.  They  in- 
fluence the  growth  as  long  as  they  are  present.  If  we  liberate  tissues 
or  timiors  from  their  influence  these  substances  lose  their  effect  at 
once  or  relatively  soon.  If,  however,  we  are  able  to  transplant  cer- 
tain tumors  through  forty  generations  of  animals  and  if  the  tumors 
preserve  their  character  as  tumors,  notwithstanding  the  individual 
differences  of  the  different  animals  into  which  they  are  transplanted, 
then  there  must  be  present  some  factor  in  or  near  the  tumor  cells 
themselves  that  constantly  stimulates  their  growth  and  stirs  them 
restlessly  to  new  activity,  until  through  their  activity  they  destroy 
their  host,  and  thus  prepare  their  own  end.  What  the  character  of 
this  local  stimulus  is,  we  do  not  yet  know.  All  the  discoveries  of 
organisms  that  have  been  announced  from  time  to  time  were  found 
to  be  based  upon  erroneous  observations ;  but  that  does  not  exclude 
the  possibility  that,  after  all,  a  microorganism  in  intimate  relation 
with  the  tumor  cell  is  the  local  stimulus  acting  on  the  tumor  cell. 

There  are  two  discoveries  that,  in  themselves  of  interest,  promise 
to  give  us  a  foothold  from  which  to  attack  successfully  this  problem : 
In  the  first  place  the  endemic  occurrence  of  tumors  among  animals, 
to  which  we  alluded  above.  Here  we  can  determine  whether  it  is 
caused  by  hereditary  conditions,  or  whether  it  is  due  to  microorgan- 
isms or  environmental  factors.  Secondly,  the  surprising  fact  we 
learned  three  years  ago,  that  if  we  inoculate  one  kind  of  tumor,  an 
epithelial  tumor,  a  carcinoma,  into  animals,  the  carcinoma,  in  a 
certain  number  of  cases,  causes  the  surroundine  connective  tissue 
to  assume,  likewise,  a  cancerous  growth.  We  have  here,  therefore, 
actually  succeeded  in  producing  a  new  tumor,  a  sarcoma.  Such  a 
fact  was  entirely  unforeseen.  It  could  be  discovered  only  through 
the  experimental  method  of  investigation.  The  more  unexpected  a 
new  fact,  the  more  welcome  it  is ;  the  more  it  promises  to  change 
existing  conceptions  and  to  open  up  new  roads,  where  before  no  way 
out  could  be  seen. 


12  LOEB— TUMOR  GROWTH  AND  TISSUE  GROWTH,     yanuaryir. 

Lastly,  the  first  steps  have  already  been  taken  to  find  a  rational 
way  of  curing  cancer  by  procuring  immunity  in  a  similar  way  to 
that  by  which  we  are  able  to  cure  a  certain  number  of  infectious 
diseases.  Protective  sera  can  not  only  be  prepared  against  bacteria, 
against  toxins,  but  also  against  cells;  and  probably  also  against 
tumor  cells.  The  beginning  has  been  made.  Certain  tumors  in 
animals  have  been  made  to  disappear  in  such  a  way.  Let  us  hope 
that  the  future  holds  still  better  results,  and  that  we  shall  be  able  to 
alleviate  suffering  and  to  gain  a  deeper  insight  into  conditions  that 
determine  the  fate  of  living  matter. 


Stated  Meeting  February  7,  ipo8. 
Councillor  Rosengarten  in  the  Chair. 

A  letter  was  read  from  the  Fourth  International  Congress  of 
Mathematics,  announcing  that  the  Congress  will  be  held  at  Rome, 
April  6-1 1,  1908. 

Professor  Edgar  Odell  Lovett  presented  a  report  on  the 
"Le^on  sur  Tintegration  des  Equations  differentielles  aux  derivees 
partielles  professees,  a  Stockholm  (Fevrier-Mars  1906)  Sur  I'invita- 
tion  de  S.  M.  le  Roi  de  Suede  par  M.  V.  Volterra,  Senateur  du 
RoyaUme  d'ltalie,  Professeur  de  Physique  Mathematique  a  TUni- 
versite  de  Rome."  He  also  presented  a  paper  on  "Integrable 
Oases  of  the  Problem  of  those  Bodies  in  which  the  Force  Function 
is  a  Function  only  of  the  Mutual  Distances." 

Professor  Horace  C.  Richards  and  Professor  Arthur  W. 
Goodspeed  read  a  paper  on  "Recent  Advances  in  Color  Photog- 
raphy." 

Photographs-  by  the  Lumiere  process  were  exhibited  by  Dr. 
Hartzell  and  Dr.  W.  P.  Wilson. 


Stated  Meeting  February  21,  jpo8. 
Treasurer  Jayne  in  the  Chair. 
Dr.  J.  H.  Hart  read  a  paper  on  "  Artificial  Refrigeration. 


i9o8.] 


MINUTES.  18 


Stated  Meeting  March  6,  ipo8. 

Secretary  Holland  in  the  Chair. 

Letters  were  read  from  the  Secretary  of  the  Committee  of 
Organization  of  the  Fourth  International  Congress  of  Mathematics, 
to  be  held  at  Rome,  Italy,  April  6-11,  1908,  inviting  the  Society  to 
be  represented  at  the  congress,  and  Vice-President  Simon  Newcomb 
was  appointed  as  the  Society's  delegate. 


Stated  Meeting  March  20,  ipo8. 

Curator  Miller  in  the  Chair. 

The  death  was  announced  of  Sir  Samuel  Davenport,  of  Adelaide, 
Australia. 

Dr.  George  Byron  Gordon  read  a  paper  on  "Some  of  the 
Results  of  the  University  of  Pennsylvania  Expedition  to  Alaska, 

1907."  

Stated  Meeting  April  3,  1908. 

Councillor  Rosengarten  in  the  Chair. 

Letters  were  received  from  the  Secretary  of  the  Smithsonian 
Institution  informing  the  Society  that  the  Institution  has  learned 
through  the  Department  of  State  that  the  Second  International 
Archaeological  Congress  will  be  held  at  Cairo,  Egypt,  on  the  date  of 
the  Latin  Easter,  1909,  and  requesting  that  the  Institution  be 
apprised  of  the  names  of  scholars  likely  to  attend  the  Congress. 

Dr.  Leonard  Pearson  read  a  paper  on  "  Some  Aspects  of  the 
Production  and  Distribution  of  Milk." 


General  Meeting,  April  23,  24  and  25,  ipo8. 

Vice-President  Scott  in  the  Chair. 

April  23,  Afternoon  Session. 

A  letter  was  received  from  the  College  of  Physicians,  of  Phila- 
delphia, inviting  the  President  to  be  present  on  April  29,  1908,  at 


14  MINUTES. 


[A|iin.4, 


the  laying  of  the  corner-stone  of  the  new  building  of  the  College. 
Owing  to  the  absence  of  the  President  in  Europe,  Secretary  James 
W.  Holland,  M.D,,  was  appointed  to  represent  the  Society  at  the 
ceremony. 

The  following  papers  were  read : 

"  The  Law  of  Orcsme,  Copernicus  and  Gresham,"  by  TaouAS 
Willing  Balch,  of  Philadelphia. 

"The  Dramatic  Function  of  Cassandre  in  the  Oresteia  of 
-Eschylus,"  by  Professor  William  A.  Lambehton,  of  Philadelphia. 

"Goethe's  Private  Library  as  an  Index  of  his  Literary  Inter- 
ests," by  Professor  Waterman  T,  Hewett,  of  Ithaca,  N.  Y. 

"  Art  and  Ethnolc^,"  by  Edwin  Swift  Balch,  of  Philadelphia. 

"  Cytomorphosis,  A  Study  of  the  Law  of  Cellular  Change,"  by 
Professor  Charles  Sedgwick  Minot,  of  Cantbridge. 

"  Preliminary  Report  on  the  Brains  of  the  Natives  of  the  Anda- 
man and  Nicobar  Islands,"  by  Professor  E.  A.  Spitzka,  of  Phila- 
delphia (introduced  by  Professor  J.  W.  Holland). 

"  Observations  regarding  the  Infliction  of  the  Death  Penalty  by 
Electricity,"  by  Professor  E,  A,  Spitzka,  of  Philadelphia  (intro- 
duced by  Professor  J.  W.  Holland). 

"  The  Brain  of  Rhinochimaera,"  by  Professor  Burt  G.  Wildeb, 
of  Ithaca,  N.  Y. 

April  24,  Morning  Session. 

The  following  papers  were  read :  * 

"  A  Comparison  of  the  Albino  Rat  with  Man  in  Respect  to  the 
Growth  of  the  Brain  and  of  the  Spinal  Cord,"  by  Professor  Henry 
H.  Donaldson,  of  Philadelphia.  (See  Journal  of  Comparative 
Neurology  and  Psychology.  Vol.  XVIII,  No.  4,  1908.) 

"  Preliminary   Report  upon  a   Crystallographic   Study  of  the 
Hemoglobins :  A  Contribution  to  the  Specificity  of  Vital  Substances 
■nt  Vertebrates,"  by  Professors  Edward  T.  Reichert  and 
Brown,  of  Philadelphia. 

;ent  Discoveries  in  the  Patholt^y  of  Rabies,"  by  Mazyck 
[el,  M.D,,  of  Madison,  Wis. 

s  Explosion  of  the  Saratoga  Septic  Tank,"  by  Professor 
Pitt  Mason,  of  Troy,  N.  Y. 


,9o8.]  MINUTES.  15 


"Determination  of  Dominance  in  Mendelian  Inheritance,"  by 
Charles  B.  Davenport,  Ph.D.,  of  Cold  Spring  Harbor,  N.  Y. 

"Inheritance  in  Protozoa,"  by  Professor  Herbert  Spencer 
Jennings,  of  Baltimore. 

"  The  Excretory  Organs  of  the  Metazoa :  A  Critical  Review," 
by  Professor  Thomas  H.  Montgomery,  Jr.,  of  Austin,  Texas. 

"The  Classification  of  the  Cetacea,"  by  Dr.  F.  W.  True,  of 
Washington. 

"Additional  Notes  on  the  Santa  Cruz  Typotheria,"  by  W.  J. 
Sinclair,  Ph.D.,  of  Princeton,  N.  J.  (introduced  by  Professor  W. 
B.  Scott). 

Afternoon  Session, 

The  following  papers  were  read : 

"Further  Researches  on  the  Physics  of  the  Earth,  and  espe- 
cially on  the  Folding  of  Mountain  Ranges  and  the  uplift  of  Plateau3 
and  Continents  produced  by  movements  of  Lava  beneath  the  Crust 
arising  from  Secular  Leakage  of  the  Ocean  Bottom,"  by  Dr.  T.  J.  J. 
See,  of  U.  S.  Naval  Observatory,  Mare  Island,  Cal. 

"  Stratigraphic  Observations  in  the  Vicinity  of  Susquehanna 
Gap,  North  of  Harrisburg,  Pa.,  by  Gilbert  van  Ingen,  of  Prince- 
ton, N.  J.  (introduced  by  Professor  W.  B.  Scott). 

"Some  Chilean  Copper  Minerals,"  by  Professor  Harry  F. 
Keller,  of  Philadelphia. 

"Progress  of  Demarcation  of  the  Boundary  between  Alaska 
and  Canada,"  by  Professor  O.  H.  Tittmann,  of  Washington. 

"  The  Leaf  Structures  of  the  Bermuda  Sand  Strand  Plants,"  by 
Professor  John  W.  Harshberger,  of  Philadelphia. 

"The  Influence  of  Heat  and  Chemicals  on  the  Starch  Grain," 
by  Professor  Henry  Kraemer,  of  Philadelphia. 

"  A  Contribution  to  a  Knowledge  of  the  Fungi  of  Pennsylvania ; 
Gasteromycetes,"  by  D.  R.  Sumstine,  of  Wilkinsburg,  Pa.  (intro- 
duced by  Dr.  A.  E.  Ortmann). 

April  25,  Executive  Session. 

The  pending  nominations  for  membership  were  read  and  the 
Society  proceeded  to  an  election,  and  the  teller  of  election  reported 

PROC.  AMER.  PHIU  SOC.,  XLVII.  188  B,  PRINTED  JULY  II,  I908. 


16  MINUTES.  [April  as. 

that  the  following  candidates  had  been  elected  to  membership: 
Residents  of  the  United  States: 

Martin  Grove  Burmbaugh,  Ph.D.,  Philadelphia. 

Walter  Bradford  Cannon,  A.M.,  M.D.,  Boston,  Mass. 

James  Christie,  Philadelphia. 
^  William  Hallock,  Ph.D.,  New  York  City. 

Edward  Washburn  Hopkins,  Ph.D.,  LL.D.,  New  Haven,  Conn. 

Leonard  Pearson,  B.S.,  V.M.D.,  M.D.,  Philadelphia. 

Josiah  Royce,  Ph.D.,  LL.D.,  Cambridge,  Mass. 

Jacob  G.  Schurman,  Ph.D.,  Ithaca,  N.  Y. 

Charles  Henry  Smyth,  Ph.D.,  Princeton,  N.  J. 

Herbert  Weir  Smyth,  Ph.D.  (Gottingen),  Cambridge,  Mass. 

Henry  Wilson  Spangler,  M.S.,  ScD.,  Philadelphia. 

Edward  Anthony  Spitzka,  M.D.,  Philadelphia. 

John  Robert  Sitlington  Sterrett,  Ph.D.  (Munich),  Ithaca,  New 
York. 

Richard  Hawley  Tucker,  Mt.  Hamilton,  California. 

Robert  Williams  Wood,  Ph.D.,  Baltimore. 
Foreign  Residents: 

Ernest  Nys,  Brussels. 

Albrecht  F.  K.  Penck,  Ph.D.,  Berlin. 

Morning  Session,  10,30  o'clock. 
The  following  papers  were  read : 

"The  Solution  of  Algebraic  Equations  in  Infinite  Series,"  by 
Professor  Preston  A.  Lambert,  of  Bethlehem,  Pa. 

"The  Investigation  of  the  Personal  Error  in  Double  Star 
Measures  which  depend  on  the  Position  Angle,"  by  Mr.  Eric  Doo^ 
LITTLE,  of  Philadelphia. 

"  Some  Results  of  the  Ocean  Magnetic  Work  of  the  Carnegie 
Institution  of  Washington,"  by  Dr.  L.  A.  Bauer,  Director  of  the 
Department  of  Terrestrial  Magnetism,  Washington  (introduced  by 
President  Robert  S.  Woodward). 

"Photographs  of  Daniel's  Comet,"  by  Professor  E.  E.  Bar- 
nard, of  Yerkes  Observatory,  Williams  Bay,  Wis. 

"Astronomical  Photography,"  by  Dr.  John  A.  Brashear,  of 
Allegheny,  Pa. 


,9o8.]  MINUTES.  17 

"The  Completion  of  the  Lunar  Theory  and  the  Tables  of  the 
Moon's  Motion  to  be  made  therefrom,"  by  Professor  Ernest  W. 
Brown,  of  New  Haven. 

"  The  Relative  Advantages  of  Various  Forms  of  Telescopes  for 
Solar  Research,"  by  Professor  George  E.  Hale,  of  Solar  Observa- 
tory, Pasadena,  Cal. 

"  Problems  of  Three  Bodies  on  Surfaces,"  by  Professor  Edgar 
Odell  Lovett,  of  Princeton,  N.  J. 

"  A  Living  Representative  of  the  Most  Primitive  Ancestors  of 
the  Plant  Kingdom,"  by  George  T.  Moore,  Ph.D.,  head  of  the  De- 
partment of  Botany,  Marine  Biological  Laboratory,  Wood's  Hole, 
Mass. 

Afternoon  Session, 

The  following  papers  were  read :  . 

"  The  Effect  of  an  Angle  in  a  Wire  Conductor  on  SjlSirk  Dis- 
charge," by  Professor  Francis  E.  Nipher,  of  St.  Louis. 

"  Absorption  Spectra  of  Solutions,"  by  Professor  H.  C.  Jones, 
of  Baltimore  (introduced  by  Professor  Ira  Remsen). 

"The  EflFect  of  Certain  Preservatives  upon  Metabolism,"  by 
Harvey  W.  Wiley,  M.D.,  of  Washington. 

"  A  Vedic  Concordance,"  by  Professor  Maurice  Bloomfield, 
of  Baltimore. 

"  On  the  Lost  Tribes  of  Israel  and  the  Aryan  Ancestry  of  Jesus 
and  His  First  Disciples,"  by  Professor  Paul  Haupt,  of  Baltimore. 

"  The  Sign  and  Name  for  Planet  in  Babylonia,"  by  Professor 
Morris  Jastrow,  Jr.,  of  Philadelphia. 

"Mediaeval  German  Sculpture  in  the  Germanic  Museum  of 
Harvard  University,"  by  Professor  Kuno  Francke,  of  Cambridge. 

"  Notes  on  Greek  Vases  in  the  Museum  of  Science  and  Art  of 
the  University  of  Pennsylvania,"  by  Professor  William  N.  Bates, 
of  Philadelphia  (introduced  by  Professor  Wm.  A.  Lamberton). 


THE  LAW  OF  ORESME,  COPERNICUS  AND  GRESHAM. 

By  THOMAS  WILLING  BALCH. 
{Read  April  23,  1908.) 

Among  the  most  certain  laws  known  to  economic  science  is  the 
one  that,  when  two  moneys  of  unequal  value  are  placed  in  circu- 
lation at  the  same  time  side  by  side  as  currency  of  the  realm,  the 
poorer  or  cheaper  will  drive  the  better  or  dearer  from  circulation. 
This  law,  though  fought  over  most  strenuously  in  this  country  within 
recent  years,  as  if  its  immutable  operation  had  not  been  thoroughly 
demonstrated  in  past  ages  of  humanity,  was  known  in  part  at  least 
to  the  Ancients.  Of  this  there  is  ample  proof  in  the  "  Frogs  "  of 
Aristophanes.  In  that  play,  the  foremost  comic  poet  dramatist  of 
Greece  places  in  the  mouth  of  the  chorus  these  lines : 

"Oftentimes  have  we  reflected  on  a  similar  abuse 
In  the  choice  of  men  for  office,  and  of  coins  for  common  use ; 
For  your  old  and  standard  pieces,  valued  and  approved  and  tried 
Here  among  the  Grecian  nations,  and  in  all  the  world  beside, 
Recognized  in  every  realm  for  trusty  stamp  and  pure  assay, 
Are  rejected  and  abandoned  for  the  trash  of  yesterday; 
For  vile,  adulterate  issue,  drossy,  counterfeit  and  base, 
Which  the  traffic  of  the  city  passes  current  in  their  place."* 

In  Bohn's  Classical  Library  this  passage  is  thus  rendered :  "  The  freedom 
of  the  city  has  often  appeared  to  us  to  be  similarly  circumstanced  witn  regard 
to  the  good  and  honorable  citizens  as  to  the  old  coin  and  the  new  gold.  For 
neither  do  we  employ  these  at  all,  which  are  not  adulterated,  but  the  most 
excellent,  as  it  appears,  of  all  coins,  and  alone  correctly  struck  and  proved 
by  ringing  ever)rwhere,  both  among  the  Greeks  and  the  barbarians,  but  this 
vile  copper  coin,  struck  but  yesterday  and  latterly  with  the  vilest  stamps." 

In  the  above  quotation  it  is  distinctly  shown  that  the  better  coins 
that  had  been  current  were  driven  out  and  replaced  by  pieces  of 
inferior  value.  And  as  a  poetic  mind  like  that  of  Aristophanes 
could  hardly  have  understood,  much  less  have  discovered  such  a 
subtle  unwritten  law  of  money,  had  not  some  knowledge  of  it  been 

*Frere's  translation. 

18 


,9o8.1  COPERNICUS  AND  GRESHAM.  19 

the  common  possession  of  the  intellectuals  of  Greece  in  the  epoch  in 
which  he  lived,  we  can  infer  from  Aristophanes's  statement  of  it,  that 
the  Grecian  states  passed  through  the  ups  and  downs  of  a  change  in 
the  standard  of  value  caused  by  a  debasement  of  the  currency. 

The  same  state  of  affairs  existed  among  the  Romans,  and  the 
amount  of  benefits  and  evils  that  obtained  in  the  reign  of  each  R(Mnan 
emperor  can  in  a  measure  be  judged  by  the  greater  or  less  purity  of 
the  coinage  issued  in  their  respective  reigns. 

The  experiences  of  the  ancient  world  with  money  as  the  mech- 
anism of  exchange  were  largely  known  to  the  peoples  of  the  Middle 
Ages,  and  they  had  to  discover  for  themselves  at  a  great  and  bitter 
cost  that  any  attempt  to  debase  the  currency  only  results  in  the 
good  money  disappearing  from  circulation  to  the  ruin  of  the  com- 
monwealth and  of  its  inhabitants,  especially  of  the  poorer  members. 

Three  men,  exercising  three  different  callings,  but  all  three  pro- 
found students,  and  two  of  them  ranking  among  the  scholars  of  the 
world,  in  three  different  countries,  in  three  distinct  periods  of  time, 
discovered  independently  of  one  another  and  explained  to  their 
respective  sovereigns  that  when  into  the  currency  of  a  country  a 
poorer  or  cheaper  money  is  injected  by  the  side  of  a  better  which  is 
the  standard  of  value,  the  certain  and  immutable  result  will  be  that 
the  currency  of  the  realm  will  be  debased  to  the  standard  of  the 
poorer  money.  For  as  it  will  then  be  possible  to  pay  debts  in  either 
money,  people  will  naturally  pay  them  in  the  cheaper  currency, 
selling  the  better  money  by  weight  at  the  premium  that  it  will  com- 
mand in  the  standard  of  the  poorer  currency. 

These  three  men  were  Nicole  Oresme,  Bishop  of  Lisieux  in 
Normandy,  who  stated  this  subtle  unwritten  law  of  money  for 
Charles  the  Fifth  of  France,  sumamed  the  Wise ;  Nicolaus  Coper- 
nicus of  Thorn  in  Prussia,  the  discoverer  of  the  Copemican  theory 
of  astronomy,  who  expounded  this  same  law  of  the  currency  for 
Sigismund  the  First  of  Poland;  and  Sir  Thomas  Gresham,  a  noted 
English  merchant,  who  explained  it  to  Elizabeth  of  England.  It  is 
proper,  then,  that  in  honor  of  these  three  discoverers  of  an  economic 
truth  that  is  a  precious  thing  for  humanity  to  know,  that  this  law 
should  be  called  the  Law  of  Oresme,  Copernicus  and  Gresham. 

Oresme  and  Copernicus  each  prepared  a  learned  and  comprehen- 


20  BALCH— THE  LAW   OF  ORESME,  [April  .3, 

sive  treatise  for  their  respective  sovereigns  on  the  practical  func- 
tions and  workings  of  money,  and  Gresham  wrote  a  letter  to  his 
Queen  in  which  he  pointed  out  to  her  that  good  and  bad  coin  could 
not  circulate  together.  No  branch  of  science  arises  all  developed  at 
one  bound  from  the  brain  of  a  single  man  as  Minerva  sprang  all 
armed  from  the  head  of  Jove.  It  advances  by  successive  degrees, 
as  one  scholar  after  another,  armed  with  the  knowledge  acquired 
by  his  predecessors,  develops  further  what  the  human  race  knows 
of  the  laws  of  the  universe.  And  as  Hugo  Grotius,  who  assembled 
from  all  points  of  the  compass  the  rules  and  usages  that  princes  and 
cities  observed  in  his  day  in  their  relations  one  with  another  in  his 
monumental  work,  "De  Jure  Belli  ac  Pads,"  and  gave  them  a 
further  advance  in  the  trend  of  a  humane  and  civilized  development, 
has  justly  been  called  ever  since  the  father  of  the  science  of  Inter- 
national Law,  so  Nicole  Oresme  and,  a  greater  man  than  he,  Nicolaus 
Copernicus,  for  their  pioneer  work  in  the  exposition  of  the  true  rules 
that  govern  money  as  the  medium  of  commercial  exchange,  have  just 
as  truly  been  described  by  MacLeod  as  the  Castor  and  Pollux  of 
monetary  science.  They  both  delved  into  the  past  experiences  in 
the  matter  of  money  of  their  respective  countries,  and  probably  made 
use  of  much  of  what  the  Greek  and  the  Roman  publicists  had  said 
on  the  subject.  The  work  of  Grotius  first  redounded  to  the  advan- 
tage of  humanity  by  the  application  of  many  of  the  humane  prin- 
ciples that  he  advocated  by  their  practical  adoption  by  Gustavus 
Adolphus  of  Sweden  in  the  terrible  Thirty  Years  War.  The  light 
shed  by  Oresme  and  Copernicus  on  the  functions  of  currency  first 
helped  to  lighten  the  burdens  of  humanity  through  their  application 
by  Charles  the  Fifth  of  France  and  Sigismund  the  First  of  Poland. 
And  a  generation  after  the  true  expounder  of  our  solar  and  planetary 
system  had  prepared  his  treatise  on  money.  Sir  Thomas  Gresham 
likewise,  through  Elizabeth  of  England,  aided  the  human  race  to 
derive  the  advantages  that  are  conferred  upon  society  by  an  honestly 
maintained  measure  of  value. 

The  importance  of  the  economic  work  of  Nicole  Oresme  was  first 
revealed  to  the  world  at  large  in  1862  by  William  Roscher,  professor 
of  political  economy  in  the  University  of  Leipzig.  Oresme's  master 
work,  "  Tractatus  De  Origine,  Natura,  Jure  et  Mutationibus  Mone- 


xjoB.]  COPERNICUS  AND  GRESHAM.  21 

tarum,"  was  often  referred  to  before  that  time.  But  in  every  case 
before  Roscher  saw  Oresme's  work  in  manuscript,  the  examiners  of 
Oresme's  learned  and  lucid  treatise  failed  to  grasp  its  real  impor- 
tance. When,  however,  it  came  under  the  eye  of  Roscher,  a  trained 
eccMicmnist,  he  saw  at  once  the  profound  significance  of  the  work. 
Under  the  title  of  "A  Great  French  EconcMnist  of  the  Fourteenth 
Century/'  Roscher  called  the  attention  of  the  world  to  Oresme's 
treatise  on  money.  Two  years  later  the  French  naturalized  Pole, 
Louis  Wolowski,  also  signalized  to  his  adc^ted  country  the  work 
of  the  fourteenth  century  economist* 

Nicole  Oresme,  who  may  be  looked  upon  as  the  first  scholar,  so 
far  as  we  now  know,  t6  expound  comprehensively  money  as  the 
mechanism  of  exchange,  was  by  birth  a  Norman.  He  studied  at 
the  University  of  Paris,  where  he  was  classed  in  the  Norman  nation. 
At  the  university,  Oresme  was  reputed  to  be  the  most  able  and 
learned  in  his  knowledge  of  the  sciences  and  the  fine  arts.  He  trans- 
lated at  the  request  of  Charles  the  Fifth  the  "  Ethics,"  "  Politics," 
and  other  works  of  Aristotle.  He  delivered  at  Avignon  on  December 
24,  1363,  before  Pope  Urban  the  Fifth  and  the  members  of  the 
sacred  college  a  sermon  in  which  he  censured  the  high  clergy  of 
France.  Charles  also  conmiissioned  him  to  translate  the  Bible,  in 
order  that  this  vernacular  version  might  be  opposed  to  that  of  the 
Waldensians. 

When  Charles  the  Fifth  succeeded  to  the  throne  of  his  ancestors, 
the  French,  crushed  by  what  was  for  those  times  an  enormous  debt, 
were  groaning  under  the  weight  of  the  accumulated  mismanagement 
of  previous  rulers,  and  the  "royaume  des  lys"  had  shrunk  to  small 
proportions  before  the  English  invasion,  and  was  fast  disappearing 
in  misery  and  anarchy.  Owing  to  the  capture  of  Charles's  father. 
King  John,  by  the  English,  Charles  was  called  upon  to  act  as  regent. 
During  those  years  he  learnt  much  which  later  as  king  he  put  to 
valuable  practical  use.  Reigning  from  1364  to  1380  under  the  title 
of  Charles  the  Fifth,  he  was,  for  his  able  management  of  the  affairs 

'"Traictic  dc  la  premiere  invention  des  Monnoies  dc  Nicole  Oresme" 
textes  frangais  et  latin  d'apres  les  manuscrits  de  la  Biblioth^que  Imp^riale 
et  ''Traits  de  la  Monnoie  de  Copemic,"  texte  latin  et  traduction  franQaise 
publics  et  annot^s  par  M.  L.  Wolowski,  membre  de  Tlnstitut  Paris,  Guil- 
laumin  et  Cie.,  1864. 


22  BALCH— THE  LAW  OF  ORESME, 


[April  as. 


of  his  kingdom,  justly  surnamed  the  Wise.  This  honorary  title, 
Charles  the  Fifth,  who  was  a  capable  and  sagacious  man,  was  enti- 
tled for  in  great  measure  to  the  fact  that  he  surrounded  himself 
and  relied  upon  the  services  of  men  of  first  rate  ability  who  had 
strengthened  their  natural  capacities  by  hard  work,  such  generals 
as  the  Breton,  Bertrand  du  Guesclin,  such  scholars  as  the  Norman, 
Nicole  Oresme.  It  was  Charles  the  Wise,  too,  who,  in  beginning 
the  first  collection  of  manuscripts  in  the  Louvre,  that  afterwards 
became  the  Bibliotheque  Royale,  then  the  Bibliotheque  Imperiale, 
and  to-day  is  known  as  the  Bibliotheque  Nationale,  was  the  founder 
of  what  is  to-day  the  largest  depository  of  learning  in  the  world. 

The  chief  cause  of  the  unhappy  state  in  which  the  French  people 
found  themselves  when  Due  Charles  became  king  in  1364  was  in 
large  measure  due  to  the  tampering  by  their  rulers  with  the  weight 
of  the  value  of  the  coins  of  the  realm.  Many  of  the  French  kings 
had  thought  to  raise  revenue  by  forcing  their  people  to  accept  a 
debased  coinage.  Of  these  royal  false  coiners,  Dante  flays  Philip 
the  Fair  (1285-1314)  in  the  Paradiso  in  these  words: 

''La  si  vedra  il  duol  che  sopra  Senna 
Induce,  f alseggiando  la  moneta." ' 

In  addition  to  debasing  the  coinage,  the  French  sovereigns  ag^in 
and  again  changed  the  mint  price  of  gold  and  silver.  In  the  reign 
of -King  John  the  Second,  the  value  of  the  liz/re  tournois  was  changed 
between  1351  and  1360  no  less  than  seventy-one  times.*  And  what 
made  the  resulting  confusion  from  this  unjustified  and  foolish  med- 
dling with  the  measure  of  commerce  still  worse  was  that  sometimes 
the  value  of  the  ItT^e  tournois  was  raised  and  sc«netimes  it  was  low- 
ered. As  a  result,  far  from  filling  the  coffers  of  the  king,  this 
policy  prostrated  commerce,  and  the  wealth  in  the  realm  of  France 
shrank.  When  Charles  the  Fifth,  upon  his  father's  death,  ascended 
the  throne,  he  called  upon  Nicole  Oresme,  in  order  that  he  might 
reform  the  coinage  of  France,  to  shed  light  upon  the  confused  cur- 
rency of  the  kingdom.  And  thus  it  was  that  Oresme  prepared  his 
most  important  work,  already  referred  to,  the  first  comprehensive 

•"There  shall  be  seen  the  woe  that  he  shall  pour 

Along  the  Seine  by  debasing  the  coinage." 
*  Wolowski. 


S908.]  COPERNICUS  AND  GRESHAM.  23 

treatise  upon  money,  entitled  "  Tractatus  De  Origine,  Natura,  Jure 
et  Mutationibus  Monetarum." 

Of  this  work  many  manuscript  copies  of  the  Latin  original  were 
made,  and  also  of  a  French  translation  by  the  author  himself  under 
the  title  "Traictie  de  la  premiere  invention  des  monnoies."  This 
translation  was  placed  as  early  as  1373  at  least  in  the  library  col- 
lected by  the  direction  of  King  Charles  in  the  Louvre. 

Oresme,  in  stating  the  various  workings  of  money  as  the  mech- 
anism of  exchange,  explained  in  precious  words  to  his  sovereign 
that,  whenever  the  public  currency  was  altered  or  tampered  with  in 
such  a  way  as  to  bring  into  circulation  two  moneys,  bearing  the 
same  designation  but  in  reality  having  two  different  values,  the 
money  of  lower  value  inevitably  drove  the  money  of  higher  value 
out  of  circulation.  For  the  merchants  found  it  to  their  advantage 
either  to  melt  down  the  pieces  of  mc«iey  that  contained  the  higher 
amount  of  metal  and  to  sell  the  bullion  by  weight  or  else  to  export 
the  high  weight  coins  to  other  lands.  Thus  Oresme  says:  "The 
rate  of  exchange  and  the  price  of  the  moneys  must  be  for  the  king- 
dom as  a  law  and  a  firm  ordinance  which  in  no  way  must  alter  or 
change."  And  further  in  speaking  of  the  ratio  of  exchange  be- 
tween gold  and  silver,  Oresme  points  out  that  the  value  or  propor- 
tion in  which  those  metals  are  exchanged  in  their  natural  state,  is 
the  rate  of  exchange  that  must  be  maintained  between  gold  and 
silver  currency.  For  if  a  given  amount  of  gold  is  worth  twenty 
times  as  much  silver,  then  a  livre  of  gold  would  be  worth  twenty 
liT/res  of  silver,  a  nuirk  of  gold  twenty  marks  of  silver.  "But 
always  this  proportion,"  he  says,  "must  follow  the  natural  habit 
or  rate  of  gold  to  silver,  in  value."  The  mutations  of  the  currency 
are  of  great  peril  to  the  national  welfare  "  for  the  injury  which 
comes  by  it,"  he  says,  "  is  not  so  soon  felt  nor  seen  by  the  people, 
as  it  would  be  by  another  tax,  and  nevertheless  no  such  nor  similar 
can  be  more  grievous  or  greater ;  and,  in  addition,  gold  and  silver, 
by  such  mutations  and  changes,  shrink  and  diminish  in  a  kingdom, 
and  in  spite  of  all  vigilance  and  prohibition  that  may  be  taken,  they 
go  abroad  where  they  are  accorded  a  higher  value  for,  by  adventure, 
men  carry  more  voluntarily  their  moneys  to  the  places  where  they 
know  these  have  a  greater  value." 


24  BALCH— THE  LAW  OF  ORESME,  [April  .3, 

The  luminous  treatise  of  Oresme  on  money  opened  the  eyes  of 
King  Charles  to  the  disastrous  results  to  a  country  whose  govern- 
ment attempted  to  alter  the  basic  value  of  its  currency.  As  r^ent 
of  France  during  the  captivity  by  the  English  of  his  father.  King 
John  the  Second,  who  was  captured  at  Poitiers  in  1356,  Charles 
had  not  escaped  the  prevailing  custom  among  rulers  of  that  epoch 
to  fill  the  royal  purse  by  debasing  the  coins  of  the  realm.  In  the 
previous  century  the  great  ordinance  of  1255,  which  the  States  Gen- 
erals of  France,  assembled  at  Paris,  obtained  from  the  king,  Louis 
the  Ninth,  promised  sound  and  stable  money  for  the  whole  kingdcxn 
of  France,  so  that  the  mark  of  silver  should  never  produce  more 
than  six  livre  tournois.  This  royal  promise  was  broken  again  and 
again  by  the  French  sovereigns,  and  Due  Charles,  as  regent  for  his 
captive  father,  said  the  value  of  the  mark  should  be  worth  twelve 
liz/re  tournois.  This  cutting  in  half  of  the  measure  of  value  was 
the  signal  for  the  great  rising  at  Paris  in  1357  under  Etienne  Marcel, 
the  Prevost  of  the  Paris  merchants,  and  it  was  with  difficulty  that 
the  regent  reasserted  the  royal  authofity  in  the  city.*  The  dis- 
tracted and  poverty-stricken  state  of  the  people  and  the  low  ebb  of 
the  kingly  power,  reenforced  by  his  practical  experiences  as  regent, 
caused  Charles  the  Wise,  though  of  a  physique  so  frail  that  he  could 
not  march  at  the  head  of  his  army  in  those  years  of  strife  and  peril, 
yet  endowed  with  a  superior  mind  and  seeking  the  advice  of  sage 
advisers,  to  set  himself  to  reorganize  the  finances  of  France.  The 
luminous  thoughts  expressed  in  the  treatise  of  Oresme  he  made  his 
own,  and  during  his  reign  the  weight  of  the  gold  currency  remained 
a  fixed  and  unchanged  quantity,  and  that  of  silver  was  but  very 
triflingly  altered..  The  resulting  stability  in  the  value  of  money, 
the  measure  of  commercial  exchange,  reestablished  the  regularity 
of  commercial  transactions,  and  furnished  an  important  element  to 
public  prosperity.  The  resources  of  the  realm  augmented  and  with 
them  the  power  of  King  Charles  grew.  To  honor  the  scholar  who 
had  made  plain  the  confusion  that  resulted  from  tampering  with  the 
standard  of  value,  the  money  of  the  realm.  King  Charles  had  Oresme 
elected  in  1377  Count  Bishop  of  Lisieux  in  Normandy.  And  it 
was  there,  two  years  after  the  king's  death  in  1380,  that  the  great 

•  Wolowski. 


X908.J 


COPERNICUS   AND  GRESHAM.  26 


economist  died  on  July  ii,  1382,  regretted  especially  by  the  scholars 
of  his  day. 

The  economic  truths  that  Oresme  so  well  stated  in  his  treatise 
on  money  did  not  become  widely  known,  for  his  work  was  written 
for  his  king's  information,  and  Gutenberg  had  not  yet  made  it  pos- 
sible through  printing  to  give  them  a  wide  circulation.  The  truths 
that  Oresme  taught  and  upon  which  Charles  the  Wise  acted,  to  the 
profit  of  his  kingdom  and  therefore  of  himself,  became  in  great 
measure  forgotten.  A  century  and  a  half  after  Oresme's  death 
they  were  rediscovered  and  restated  a  second  time.  In  the  year 
1526,  in  a  Latin  treatise  entitled  "  Monete  Cudende  Ratio,"  written 
at  the  request  of  Sigismund  the  First,  King  of  Poland,  and  his  Chan- 
cellor, Szydlowiecki,  Nicolas  Copernicus  of  Thorn  in  Prussia,  who 
had  elucidated  for  mankind  some  of  the  celestial  harmonies,  gave 
to  the  world  an  exposition  of  some  of  the  economic  truths.  Inde- 
pendently of  the  work  of  Oresme,  of  which  the  Prusso-Polish 
scholar  knew  nothing,  Copernicus  made  clear  for  his  sovereign  that 
two  moneys  of  unequal  value  could  not  be  kept  in  circulation  at  the 
same  time.  "  Gold  or  silver,"  he  writes,  "  marked  with  an  imprint, 
constitutes  the  money  which  serves  to  determine  the  price  of  things 
that  are  bought  and  sold,  according  to  the  laws  established  by  the 
State  or  the  Prince.  Money  is  therefore  in  some  sort  a  common 
measure  of  estimating  values ;  but  this  measure  must  always  be  fixed 
and  must  conform  to  the  established  rule.  Otherwise,  there  would 
be,  necessarily,  disorder  in  the  State:  buyers  and  sellers  would  at 
all  times  be  misled,  as  if  the  ell,  the  bushel  or  the  weights  did  not 
maintain  constant  quantity. 

"  The  establishment  of  money  has  necessity  for  cause.  Though  in  weigh- 
ing only  gold  and  silver  it  would  have  been  possible  to  practice  exchanges, 
those  metals,  from  the  unanimous  consent  of  men,  being  considered  every- 
where as  precious  things,  nevertheless  there  would  be  numerous  inconveni- 
ences to  have  to  carry  always  weights  along,  and,  all  the  world  not  being  apt 
to  recognize  at.  the  first  glance  the  purity  of  gold  and  silver,  it  is  agreed 
everjrwhere  to  have  money  marked  by  government  ^ith  a  stamp  designed  to 
show  how  much  each  coin  contains  of  gold  and  silver  and  to  serve  as  a 
guaranty  to  public  faith." 

Then  he  explains  how  the  value  of  metal  pieces  is  changed  and 
depreciated. 


26  BALCH— THE  LAW  OF  ORESME,  [April  23, 

"The  value  of  money  is  depreciated  by  various  causes,  either  by  the 
change  of  the  name,  while  the  same  weight  of  metal  contains  a  mixture  of 
copper  which  exceeds  the  measure  desired;  or  because  the  weight  is  wanting, 
although  the  mixture  has  been  accomplished  in  the  right  proportion ;  or,  what 
is  the  worst,  because  the  two  vices  meet  together  at  the  same  time.  The 
value  of  money  diminishes  of  itself  by  reason  of  a  long  service  that  uses  the 
metal  and  diminishes  its  quantity  and  this  reason  suffices  to  cause  to  be 
placed  in  circulation  a  new  money.  This  necessity  is  recognized  by  an  in- 
fallible sign,  when  the  money  weighs  notably  less  than  the  money  intended 
to  be  acquired.  It  is  understood  that  there  results  a  deterioration  of  the 
money." 

At  the  time  Copernicus  prepared  his  treatise  on  the  money  of 
the  realm  for  his  sovereign  liege,  King  Sigismund,  the  Polish  King- 
dom included  Thorn,  Danzig,  and  a  large  part  of  Prussia.  But  a 
portion  of  Prussia,  including  Konigsberg,  had  been  erected  by  the 
treaty  of  Cracow,  concluded  in  1525  between  Sigismund,  King  of 
Poland,  and  Albert,  Margraf  of  Brandenburg,  into  a  hereditary  fief 
for  the  benefit  of  the  latter  and  his  male  descendants,  which  the 
margraf  was  to  hold  of  King  Sigismund.  As  by  this  feudal  tenure 
by  Margraf  Albert  of  part  of  Prussia,  subject  to  the  overlordship 
of  the  Polish  king,  the  two  countries  were  in  a  sense  one,  Coper- 
nicus, in  his  treatise  on  the  money  of  the  realm,  expounded  to  his 
king  what  measures  were  necessary  in  order  to  restore  stability  to 
the  much  depreciated  Prussian  money  and  then  maintain  the  value 
of  the  new  money  on  a  parity  so  that  it  could  circulate  both  in  Poland 
and  Prussia.  After  pointing  out  how  useless  it  was  to  attempt  to 
introduce  into  circulation  by  the  side  of  a  depreciated  currency  one 
of  greater  value,  he  then  explained  how  the  introduction  of  a  cheaper 
measure  of  value  by  the  side  of  a  higher  one  would  drive  the  former 
from  circulation. 

"  If  it  does  not  do  to  introduce  a  new  and  good  money,  while  the  old 
is  bad  and  continues  to  circulate,  a  much  greater  error  is  committed  by  intro- 
ducing alongside  of  an  old  currency,  a  new  currency  of  less  value ;  this  latter 
does  not  merely  depreciate  the  old,  it  drives  it  away,  so  to  speak,  by  main 
force." 

Then  in  answer  to  the  argument  that  a  depreciated  currency  helps 
the  poor,  he  says : 

"We  see  flourish  the  countries  that  possess  a  good  currency,  while 
those  that  only  have  a  depreciated  one,  fall  into  decadence  and  decline.   .   .   . 


,9o8.]  COPERNICUS  AND  GRESHAM.  27 

"  It  is  incontestable  that  the  countries  that  make  use  of  good  currency 
shine  in  all  the  arts,  have  better  workmen,  and  have  of  everything  in  abund- 
ance. On  the  contrary,  in  the  States  which  make  use  of  a  degraded  money, 
reigns  cowardice,  laziness  and  indolence/' 

In  order  to  remedy  the  distress  to  which  Prussia  had  been  brought 
by  the  falsification  and  debasement  of  the  currency,  and  to  draw 
Prussia  and  Poland  closer  together  by  developing  their  commercial 
relations,  it  was  necessary  to  coin  two  moneys  of  equal  intrinsic 
value,  so  that  they  would  circulate  concurrently  in  the  two  lands. 
One  should  bear  one  one  side  the  royal  arms  of  Poland  and  on  the 
other  those  of  the  Prussian  land.  The  other  money  should  likewise 
have  on  one  side  the  royal  arms  of  Poland,  but  on  the  other  the 
imprint  of  the  prince,  that  is,  the  effigy  of  the  king. 

"For  the  first  condition  to  maintain,  is  that  one  and  the  other  currency 
remain  under  the  royal  authority,  and  that  they  be  current  and  accepted  in 
the  whole  kingdom  by  virtue  of  the  prescription  of  His  Majesty;  which  would 
be  not  of  a  mediocre  importance  for  the  conciliation  of  public  opinion  and  for 
reciprocal  transactions. 

"  It  would  be  necessary  that  these  two  currencies  should  be  of  the  same 
degree  of  fineness,  having  a  similar  real  value  and  a  similar  nominal  value, 
so  that,  by  vigilant  care,  the  State  succeeds  to  maintan  perpetually  the  regu- 
lation which  it  is  now  question  to  establish;  it  does  not  belong  to  princes 
to  obtain  any  profit  from  the  money  that  they  shall  coin ;  they  shall  add  only 
so  much  alloy  as  may  be  necessary  for  the  difference  between  the  real  value 
and  the  nominal  value  to  cover  the  cost  of  minting,  which  will  avoid  the 
principal  attraction  to  remelt  it. 

**  It  would  be  necessary,  at  the  time  of  the  emission  of  the  new  money, 
to  demonetise  the  old  and  forbid  entirely  its  use,  allowing  it  to  be  exchanged 
at  the  mints,  in  the  just  proposition  of  the  intrinsic  value.  Otherwise  it 
would  be  labor  lost  to  wish  to  reestablish  good  money;  the  confusion  that 
would  ensue  would  be  perhaps  even  worse  than  the  actual  state  of  affairs. 
The  old  money  would  crush  all  the  advantages  of  the  new." 

Then  Copernicus  explained  that  gold  and  silver  were  the  base 
upon  which  rested  the  value  of  money;  and  went  on  to  show  that 
in  order  to  maintain  them  both  in  circulation  the  ratio  between  them 
must  agree  with  the  commercial  ratio  that  existed  between  them. 

"  It  remains,"  he  went  on,  "  for  us  to  expound  the  manner  of  the  mutual 
exchange  of  gold  and  silver.  In  order  to  pass  from  the  class  to  the  kind 
and  from  the  simple  to  the  composite,  it  is  necessary  first  to  know  the  price 
of  pure  gold  to  pure  silver.     It  is  known  that  the  same  exists  between  pure 


28  BALCH— THE  LAW  OF  ORESME,  [April  .3, 

gold  and  silver,  as  between  gold  and  silver  minted  under  the  same  stamp; 
as  also  that  the  same  ratio  applies  to  gold  coins  and  gold  bars  as  to  silver 
coins  and  silver  bars,  provided  that  they  have  the  same  proportion  of  alloy 
and  that  they  represent  the  same  weight" 

As  Oresme  and  Copernicus  explained  to  their  royal  masters  that 
by  either  debasing  or  raising  the  coins  of  the  ^ealm  disaster  and 
confusion  would  follow,  so  also,  at  the  banning  of  Queen  Eliza- 
beth's reign,  one  of  her  merchants.  Sir  Thomas  Gresham,  pointed 
out  to  his  royal  mistress  this  inflexible  unwritten  law  of  money. 

Of  a  Norfolk  family,  the  son  of  Sir  Richard  Gresham,  who  was 
Lord  Mayor  of  London,  Sir  Thomas  Gresham  was  born  in  that  city 
probably  in  1519,  and  died  there  on  November  21,  1579.  He  was 
educated  at  Cambridge  University,  was  a  Protestant,  and  all  his  life 
took  an  active  part  in  commercial  affairs,  often  representing  in  the 
Low  Countries  the  commercial  interests  of  England.  In  1566  and 
1567  he  built  the  Royal  Exchange  in  London.  He  founded  also 
Gresham  College,  and  provided  that  the  science  of  astronomy  should 
be  taught  there. 

In  a  letter  to  Queen  Elizabeth,  which  is  headed  "  information  of 
Sir  Thomas  Gresham,  Mercer,  touching  the  fall  of  the  exchange, 
MDLVIII,"  and  which  begins,  "To  the  Quenes  most  excellent 
maiestye,"  Gresham  says : 

"  Ytt  may  pleasse  your  majesty  to  understande,  thatt  the  firste  occasion 
off  the  fall  of  the  exhainge  did  growe  by  the  Kinges  majesty,  your  latte 
ffather,  in  abasinge  his  quoyne  ffrome  vi  ounces  fine  to  iii  ounces  fine. 
Whereuppon  the  exchainge  fell  ffrome  xxvis.  viiid  to  xiiis.  ivd.  which  was 
the  occasion  thatt  all  your  fline  goold  was  convayd  ought  of  this  your  realme." 

The  works  on  money  of  these  three  men,  who,  independently  of 
one  another,  expounded  to  their  respective  sovereigns  the  evils 
resulting  to  the  State  from  any  attempt  to  debase  the  coinage,  did 
not  become  generally  known  to  their  contemporaries.  However, 
their  discoveries  through  the  influence  of  their  royal  rulers  gradually 
made  some  impress  upon  mankind,  and  by  the  end  of  the  seventeenth 
century  it  had  become  common  knowledge  among  the  intellectuals  of 
that  day.  In  a  pamphlet  published  in  London  in  1696,  the  Law  of 
Oresme,  Copernicus  and  Gresham,  though  doubtless  the  writer  did 
not  know  directly  of  their  works,  is  thus  stated : 


I908.]  COPERNICUS  AND  GRESHAM.  29 

"When  two  sorts  of  Coin  are  current  in  the  same  nation  of  like  value 
by  denomifiation  but  not  intrinsically  [that  is  in  commercial  value],  that 
which  has  the  least  value  will  be  current,  and  the  other  as  much  as  possible 
will  be  hoarded." 

In  1858  the  British  economist,  Henry  Dunning  MacLeod,  called 
attention  to  Gresham's  statement  of  this  unwritten  law  of  coinage, 
and  suggested  that  it  should  be  known  as  Gresham's  Law.  At  the 
time  he  did  not  know  of  the  more  elaborate  treatises  of  Oresme  and 
Copernicus  on  coinage.  But  when  the  works  on  money  of  those 
two  master  econoniists  were  revealed  through  the  efforts  of  Roscher 
and  Wolowski  in  1862  and  1864  to  the  world  at  large,  MacLeod, 
like  a  true  scholar  who  wishes  to  give  credit  to  whom  honor  is  due, 
sug^gested  that  this  economic  law,  a  law  more  powerful  than  the 
statutes  enacted  by  the  strongest  Parliamentary  bodies,  should  be 
known  after  all  three  of  its  discoverers  as  the  Law  of  Oresme, 
Qjpemicus  and  Gresham. 

During  the  centuries,  many  nations  in  various  parts  of  the  world 
have  had  abundant  experience  to  learn  the  futility  of  attempting  to 
maintain  in  circulation  as  currency  two  moneys  at  a  ratio  different 
from  the  market  or  commercial  ratio  existing  at  that  time  between 
those  two  kinds  of  money.  In  every  case  where  such  an  effort  has 
been  made,  the  money  that  is  underrated  gradually  drives  that 
which  is  overrated  from  the  country.  And  this  nation  has  had  sev- 
eral experiences  with  this  law.  Without  touching  here  upon  the 
works  of  other  economic  scholars,  such  as  Petty,  Locke,  Wolowski, 
Jevons,  Leon  Say,  Horton,  Bamberger,  Laughlili,  White  and  others, 
who  have  added  to  our  knowledge  of  the  unwritten  laws  that  govern 
money  as  the  medium  of  exchange,  it  can  be  safely  said  that  the 
more  the  economic  experience  of  the  human  race  is  studied,  the 
more  does  it  become  clear  that  any  attempt  to  tamper  with  the  cur- 
rency of  a  nation  by  injecting  a  debased  money  into  its  measure  of 
value  is  certain  to  end  in  disaster  through  the  working  of  that  nat- 
ural law  of  finance,  the  Law  of  Oresme,  Copernicus  and  Gresham. 


ART  AND  ETHNOLOGY. 

By  EDWIN  SWIFT  BALCH. 

(Read  April  23,  igo8.) 

Man  has  been  studied  in  many  ways  and  from  many  directions : 
history,  language,  archeology,  anatomy,  natural  history,  geography 
and  other  sciences  have  been  called  upon  in  the  elucidation  of  the 
problems  of  his  history,  descent,  evolution  and  origin.  The  evidence 
which  has  been  gathered  from  these  many  different  sources  about 
man  and  his  history  may  be  divided  into  two  classes:  that  which 
can  be  obtained  from  his  own  personality  or  his  own  remains,  a  class 
I  do  not  need  to  mention  again  in  this  paper ;  and  that  which  can  be 
obtained  from  what  man  has  produced,  and  this  class  of  evidence 
may  be  subdivided  into  three  sub-classes,  namely,  written  records, 
implements  and  art. 

The  most  primarily  available  evidence  in  tracing  the  story  of  the 
human  race  is,  of  course,  written  records,  and  whenever  we  find 
written  records  which  we  can  interpret  we  speak  of  history;  but 
when,  as  in  the  case  of  savages,  there  are  no  written  records,  or 
when,  as  in  the  case  of  Old  Mexico,  we  cannot  read  the  records,  the 
subject  changes  from  history  into  ethnology  and  pre-history. 

When  there  are  no  written  records,  another  class  of  evidence, 
that  obtained  from  implements,  is  largely  resorted  to  by  ethndogists. 
The  term  "implements,"  as  used  in  this  paper,  should  perhaps  be 
defined  as  an  abbreviated  name  for  the  products  of  the  mechanical 
arts,  without  some  of  which  at  least  no  man  can  live.  All  modem 
implements  have  evolved  from  primitive  beginnings,  as,  for  instance, 
the  twelve-inch  shell,  which  is  really  the  most  modem  form  of  the 
flint  arrowhead.  Much  light  has  been  shed  already  and  more  will 
be  shed  on  the  story  of  man  by  a  comparison  of  the  various  imple- 
ments used  in  different  places  and  at  different  times. 

The  other  great  class  of  evidence  is  art,  under  which  term  must 
be  understood  the  fine  arts  of  sculpture,  drawing  and  painting. 

30 


,9o8.J  BALCH— ART  AND  ETHNOLOGY.  81 

Some  use  has  been  made  of  this  class  of  evidence ;  nevertheless,  it  is 
far  below  what  it  should  be  and  usually  it  is  only  local  in  its  deduc- 
tions. There  are  plenty  of  treatises  relating  to  the  art  of  the  white 
races,  of  the  modem  Europeans,  of  the  Romans,  of  the  Greeks ;  scrnie 
on  Egyptian  art ;  others  on  Kaldean  art  and  Assyrian  art ;  some  on 
Old  Mexican  art  and  Peruvian  art,  and  so  forth.  But  so  little  is 
the  subject  worked  out  even  locally,  that  there  is  practically  no 
special  publication  about  African  art  or  Brazilian  art,  and  it  is  only 
within  the  twentieth  century  that  we  find  the  first  serious  attempt 
to  trace  back  the  wonderful  art  of  China.  As  a  subject  of  study, 
either  from  an  artistic  or  an  ethnological  standpoint,  the  art  of  the 
world  as  a  whole  is  so  far  almost  untouched.  Even  in  such  an  excel- 
lent recent  art  history  as  Mr.  S.  Reinach's  "Art  Throughout  the 
Ages,"  one  finds  that  by  art  he  means  European  art  alone  and  that 
Hindu  art  or  Chinese  art  or  Mexican  art  are  left  out  in  the  cold. 
Whether  art  comes  from  only  one  center  or  whether  there  are  sev- 
eral foci  of  dispersion ;  what  relations,  what  resemblances,  and  what 
differences  there  are  in  the  art  of  the  world  as  a  whole,  is  as  yet  an 
almost  virgin  field.  If  I  am  not  mistaken,  only  one  attempt  has 
been  made  (by  the  writer  himself)  to  study  and  classify  art  in  every 
district  of  the  globe. 

Probably  the  main  reason  why  art  in  totality  is  still  so  largely 
unstudied  is  that  it  is  only  recently  that  art  specimens  from  every- 
where have  been  collected,  placed  in  museums,  and  made  accessible. 
But,  connected  with  this  placing  of  art  specimens  in  musetuns,  there 
is  a  curious  fact  which  shows  that  the  art  of  the  world,  at  present, 
appears  to  hang  in  a  sort  of  borderland  between  art  and  science. 
The  specimens  are  divided.  Some  are  placed  in  art  museums,  others 
in  ethnological  museimis.  For  instance,  in  Philadelphia,  art  speci- 
mens are  divided  between  the  Pennsylvania  Academy  of  the  Fine 
Arts  and  the  University  Archeological  Museum;  in  Washington, 
between  the  Corcoran  Gallery  and  the  United  States  National  Mu- 
seum; in  New  York,  between  the  Metropolitan  Museimi  and  the 
American  Museum  of  Natural  History;  in  Boston,  between  the 
Museum  of  Fine  Arts  and  the  Cambridge  Peabody  Museum.  There 
is  no  place  where  anyone  can  go  and  get  a  comprehensive  view  of 
art  from  all  over  the  world. 

PROC.  AMER.  PHIL.  SOC.  XLVH.  l88  C,  PRINTED  JULY  lO,  I908. 


32  BALCH-ART  AND  ETHNOLOGY.  [April  83, 

The  art  of  at  least  half  the  races  of  the  world  has  thus  found  its 
way  into  ethnological  museums.  There  it  is  not  yet  culled  out  as 
art,  but  the  specimens  are  looked  on  rather  as  belonging  to  the  class 
which  can  be  most  briefly  called  implements.  This  is  not  to  be  won- 
dered at.  Ethnologists,  as  a  rule,  have  not  had  any  special  art  train- 
ing. Among  artists  it  is  a  pretty  thoroughly  understood  thing — and 
this  can  be  stated  only  as  a  dictum  and  not  discussed  in  this  paper — 
that  only  a  trained  artist  can  criticize  art  seriously;  in  fact,  the 
present  most  prevalent  opinions  about  art  are  largely  the  consensus 
of  opinion  of  the  many  artist  art  critics  of  modem  times.  Whilst 
possibly  unconscious  of  this  fact,  ethnologists  are  usually  aware  of 
their  inability  to  discuss  the  esthetic  qualities  of  art  specimens,  and 
hence,  while  they  frequently  study  the  decorative  art  of  savages,  its 
patterns  and  its  origins  they  are  apt  to  leave  the  esthetic  qualities 
of  art  alone. 

Whilst  scientists,  therefore,  generally  do  not  give  much  thought 
to  the  esthetic  points  of  the  art  specimens  in  ethnological  museums, 
on  the  other  hand,  artists  and  art  critics  so  far  have  paid  no  atten- 
tion to  such  arts  as  African  or  Australian  art.  In  the  overwhelming 
majority  of  cases,  they  are  doubtless  unaware  of  the  existence  of 
such  arts,  and  if  they  did  know  of  them  they  would  in  many  cases 
despise  them,  because  these  arts  do  not  have  the  qualities  of  Greek 
art  or  Japanese  art  or  French  art  Art  critics  also  usually  know 
nothing  of  ethnology,  and  certainly  care  less.  It  takes  a  good  deal 
of  time  and  thought  and  study  to  learn  something  of  ethnology,  and 
any  scientist  knows  that  only  a  specialist  can  really  give  an  opinion 
about  it.  The  result  of  these  scnnewhat  complex  conditions  is  that 
both  ethnologists  and  art  critics  have  neglected  the  esthetic  arts  of 
perhaps  half  the  races  of  the  world. 

It  seems  as  if  it  should  be  recognized  that  the  present  state  of 
things  leaves  a  gap  in  knowledge.  It  is  time  that  this  gap  should 
be  filled  in  and  that  the  art  of  the  entire  world  should  be  worked 
out  as  a  whole  into  its  proper  divisions  and  its  relations.  Prac- 
tically this  will  amount  to  forming  a  new  branch  of  science,  a  science 
which  might  well  be  termed  comparative  art,  and  it  seems  just  as 
necessary  that  there  should  be  a  science  of  comparative  art  as  a 
science  of  comparative  philology  or  a  science  of  comparative  anat- 


1908.] 


BALCH— ART  AND  ETHNOLOGY.  33 


omy.  It  will  be  a  science  in  which  art  critics  and  ethnologists  will 
have  to  work  hand  in  hand ;  it  will  either  have  to  be  worked  out  by 
trained  artists  and  also  by  ethnologists,  or  better  still,  comparative 
art  must  be  handled  by  men  who  are  something  of  specialists  in 
both  fields. 

Comparative  art  should  not  be  confounded  with  comparative 
archeology.  Although  there  are  points  of  contact,  the  fields  are 
different.  Comparative  archeology  is  mainly  based  on  the  results 
of  digging  with  the  pick  and  the  spade.  It  includes  studies  of  cer- 
tain phases  of  art  and  architecture,  of  inscriptions,  of  implements, 
and  some  other  things.  It  does  not  deal  with  the  work  of  the 
Eskimo,  or  the  Australian,  or  the  Ashantee  of  to-day.  It  is  a  study 
of  past  things. 

Comparative  art,  on  the  contrary,  must  deal,  not  only  with  the 
past,  but  also  with  the  present.  It  will  not  be  a  study  of  written 
inscriptions,  nor  of  implements,  but  it  will  be  a  study  of  art,  and 
architecture  so  far  as  this  is  a  form  of  the  fine  arts,  and  it  must  be 
applied  to  every  district  of  the  globe,  not  only  to  the  remotest  past 
in  which  there  was  art,  but  to  the  actual  present  of  to-day  and  to 
the  future.  It  will  deal  not  only  with  the  art  of  the  Pleistokenes 
and  the  Assyrians,  of  the  Chinese  and  the  Aztecs,  but  also  with  the 
art  of  the  tribes  now  living  in  the  Amazon  and  Kongo  forests,  in 
the  islands  of  the  Pacific,  and  on  the  shores  of  the  Arctic. 

Now  I  do  not  wish  to  claim  that  the  study  of  art  specimens  is 
going  to  clear  up  all  the  problems  of  ethnology,  or  do  away  with 
other  methods  of  studying  man  and  his  history,  or  an3rthing  else  of 
that  kind.  I  only  want  to  say  that  here  is  a  field  still  mainly  untilled, 
in  which  there  is  much  work  to  be  done,  and  from  which,  when  it 
is  properly  plowed  up,  a  valuable  crop  of  scientific  data  may  be 
expected. 

That  comparative  art  will  bring  up  new  problems  and  perhaps 
alter  some  theories  of  the  present  seems  probable.  For  instance,  it 
was  formerly  generally  accepted  that  there  are  five  races  of  men :  a 
white,  a  yellow,  a  brown,  a  red  and  a  black.  Then  other  theories 
were  started :  one  that  there  are  only  three  races,  a  white,  a  yellow 
and  a  black ;  and  another  that  there  are  four  races,  a  white,  a  yellow, 
a  red  and  a  black.    A  study  of  the  art  of  the  world,  however,  tends 


34  BALCH— ART  AND  ETHNOLOGY.  [Apnl  .s, 

to  make  one  revert  to  the  older  theory  of  five  main  races,  if  indeed 
it  does  not  point  to  more  than  five.  For  it  seems  as  if  there  were 
sufficiently  numerous  distinct  arts,  with  sufficiently  individual  racial 
characteristics,  as  to  necessitate  the  classifying  them  provisionally 
into  at  least  five  and  possibly  more  main  classes,  corresponding  to 
the  five  or  more  races  of  man  from  which  these  arts  spring. 

Let  me  now  give  you  some  concrete  examples  of  how  art  can 
help  clear  up  ethnology.  Take  the  Pleistokene  men  of  western  cen- 
tral Europe,  usually  mistakenly  called  the  Cave  men.  We  have  no 
written  records  from  the  Pleistokenes,  but  we  have  implements  and 
art.  Their  implements  show  that  they  must  have  lived  near  the 
edge  of  a  great  ice  sheet  and  that  their  habits  must  have  been  not 
unlike  those  of  the  Eskimo  of  to-day.  But  their  art  tells  us  a  great 
deal  of  which  the  implements  give  no  hint.  In  the  first  place  Pleis- 
tokent  arts  tells  us  the  fauna  amongst  which  these  men  lived.  It 
takes  us  back  to  a  past  geological  epoch,  when  the  mammoth  and 
the  woolly  rhinoceros  tichorinus  roamed  over  western  Europe.  It 
proves  and  is  the  only  proof  that  they  had  domesticated  the  horse 
and  possibly  the  dog  and  that  they  lived  sometimes. in  habitations  not 
unlike  the  teepees  of  the  Red  Ainerinds.  In  the  next  place  Pleisto- 
kene art  reveals  the  fact  that  these  earliest  positively  known  men 
were  unquestionably  advanced  in  some  mental  characteristics.  They 
had  certainly  stopped  hanging  on  by  their  tails.  No  one  who  was 
not  distinctly  intelligent  could  possibly  have  made  their  sculptures, 
their  drawings  and  their  paintings.  Another  fact  their  art  shows  is 
that  in  all  probability  they  were  not  a  Negroid  race.  Ordinary 
Bantu  art,  and  also  the  art  of  Great  Benin,  is  too  unlike  Pleistokene 
art  to  warrant  the  belief  that  its  makers  could  have  been  blood  rela- 
tions of  the  Pleistokenes.  Certain  qualities  of  Pleistokene  art  sug- 
gest early  Greek  art,  but  there  are  more  resemblances  which  suggest 
Chinese  or  Eskimo  work,  so  that  the  evidence  of  art,  and  it  is  the 
strongest  evidence  on  the  subject,  is  that  the  earliest  known  race  was 
a  yellow  race. 

Take  the  case  of  the  eastern  United  States.  Mr.  Henry  C. 
Mercer,  I  believe,  and  many  other  ethnologists  claim  that  there  is 
no  civilization  preceding  that  of  the  Amerinds  or  American  Indians 
on  this  continent.    Dr.  Charles  C.  Abbott  per  contra  claims  that 


,008]  BALCH— ART  AND  ETHNOLOGY.  86 

there  is  an  earlier  geological  horizon  whose  argillite  imfdements  show 
there  was  an  earlier  race.  Unfortunately,  there  are  apparently  no 
art  specimens  known  irom  the  same  horizon  as  these  argillite  imple- 
ments. But  the  lucky  finding  of  a  few,  only  a  few,  works  of  art,  in 
undisturbed  strata^  would  tell  us  positively  whether  those  argillite 
implements  belonged  to  the  Amerinds  or  whether  there  really  was  a 
previous  race.  In  other  words,  art  would  tell  us  what  the  imple- 
ments do  not 

Take  now  the  case  of  the  people  who  inhabit  the  oceanic  fringe 
of  Alaska  and  British  Columbia.  I  believe  ethnologists  consider  that 
they  are  members  of  the  red  race  of  America.  But  their  art  r^ses 
doubts.  Whilst  it  has  certainly  some  resemblances  to  the  art  of  Old 
Mexico  and  of  the  United  States,  it  has  many  more  to  the  art  of 
the  brown  races  of  the  Pacific.  It  is  more  nearly  in  touch  with  New 
Zealand  art,  with  New  Guinea  art,  and  so  forth,  than  it  is  with  the 
art  of  the  rest  of  America.  It  shows  pretty  definitely  that,  even  if 
these  northwestern  tribes  are  not  primarily  a  Pacific  island  race,  yet 
there  must  have  been  some  intercourse  and  some  immigration,  else 
they  could  not  produce  works  of  art  so  similar  to  those  of  some  of 
the  tribes  in  the  southern  Pacific. 

Let  me  give  you  one  more  instance.  The  present  art  of  Japan  is 
an  intrusive  art  which  came  over  from  China  some  fifteen  hundred 
years  ago,  as  is  shown  by  written  records.  Art  critics  are  only  just 
beginning  to  find  out  that  it  has  never  risen  to  the  heights  reached 
by  its  parent  art  of  China.  But  digging  has  revealed  the  fact  that 
there  was  some  more  elementary  art  in  Japan  which  was  prob- 
ably earlier  than  the  Chinese  influence.  This  and  some  of  their 
own  more  recent  work,  their  discarded  suits  of  lacquered  armor 
are  notable  examples,  have  art  qualities  which  are  not  Chinese. 
They  are  much  more  in  touch  with  some  South  Sea  art,  with  that  of 
New  Ireland,  for  instance.  The  evidence  of  their  art  would  tend  to 
show  that  the  Japanese  were  a  brown  race,  who  adopted  much  of 
Chinese  civilization. 

To  sum  up  now  briefly  the  gist  of  this  paper,  I  would  submit  the 
following  main  points : 

I.  Art  is  found  in  every  part  of  the  world. 


36  BALCH— ART  AND   ETHNOLOGY.  [April  03. 

2.  Art  as  a  whole  has  not  been  studied  and  examined  enough 
as  yet. 

3.  The  art  of  the  whole  world  should  be  studied  from  an  esthetic 
point  of  view  not  only  locally  and  individually,  but  in  its  broadest 
relations,  in  its  resemblances  and  its  differences.  This  branch  of 
science  might  well  be  called  comparative  art. 

4.  Comparative  art,  that  is  the  study  of  the  relations  in  the  art 
of  the  world,  must  be  done  from  the  esthetic  standpoint  by  persons 
who  are  trained  art  critics. 

5.  Comparative  art,  properly  worked  out,  may  be  expected  to 
throw  much  light  on  the  story  of  man. 


THE  BRAIN  OF  RHINOCHIMAERA. 

By  BURT  G.  WILDER. 
(Read  April  23,  1908.) 

Four  years  ago,  to  the  small  but  very  peculiar  and  ancient  group 
of  shark-like  fishes  known  as  Chimeroids,  Holocephala  and  Chis- 
mopnea,  Garman  added  a  Japanese  species,  Rhinochimaera  pacifica. 
His  description  of  the  brain  was  brief  and  the  figures  represented 
only  the  general  form  from  the  dorsum,  venter  and  side.  A  well- 
preserved  example  recently  obtained  from  Alan  Owston  of  Yoko- 
hama enables  me  to  confirm  Carman's  account  as  to  the  general 
Chimeroid  character  of  the  brain,  especially  the  cerebellum  and 
adjoining  segments,  and  as  to  the  extraordinary — ^probably  unique — 
slendemess  of  the  other  regions,  due  not  merely,  as  in  Chimaera, 
to  the  elongation  of  the  cerebral  crura,  but  also  to  the  pedunculate 
condition  of  the  olfactory  bulbs,  whose  tracts  or  crura  equal  the 
cerebral  in  length.  The  partial  dissection  of  this  brain  discloses 
additional  features,  as  shown  upon  the  colored  crayon  diagram,  viz., 
(i)  The  cerebral  and  olfactory  cavities.  (2)  The  complete  circum- 
scription of  these  cavities  by  walls  of  moderate  thickness  at  the  olfac- 
tory bulbs  and  parts  of  the  cerebral  hemispheres,  but  mostly  thin 
and  largely  membranous.  (3)  The  olfactory  crura  have  thinner 
walls  than  in  any  brain  known  to  me,  and  the  proper  nervous  sub- 
stance seems  to  be  confined  to  their  outer  or  lateral  sides.  (4)  The 
roof  of  the  undivided  cerebral  cavity  is  wholly  membranous;  like- 
wise a  narrow  mesal  zone  of  the  floor,  but  the  floors  of  the  hemi- 
spheres are  connected  by  a  terma  ("  lamina  terminalis  ")  as  described 
by  me  in  Chimaera  in  1877.  (5)  Each  substantial  wall  of  the  cere- 
bral cavity  begins  as  a  single  broad  band  which  divides  into  a  ventral 
and  a  lateral  portion  as  it  approaches  the  hemisphere ;  this  condition 
has  not  been  observed  by  me  in  any  other  brain.  (6)  There  was 
found  no  trace  of  the  Nervus  terminalis  of  Locy;  nor  has  it  been 
recognized  in  any  other  member  of  the  group. 

37 


38  WILDER— THE  BRAIN  OF  RHINOCHIMAERA.  [AtMn. 

Even  were  our  knowledge  of  structure,  devel<^ment  and  geologic 
records  more  complete,  and  even  were  there  more  substantial  a^ee- 
ment  as  to  the  bearing  of  the  facts  upon  the  affinities,  rank  and  suc- 
cession of  the  forms  concerned,  a  detailed  description  of  this  brain 
and  a  full  discussion  of  the  significance  of  its  resemblances  and  pecu- 
liarities would  be  profitable  before  a  comprehensive  society  like  this 
only  when,  as  ui^ed  by  me  in  this  hall  three  years  ago,  the  concrete 
foundations  of  neurology  are  laid  in  the  primary  schools,  and  when 
no  child  reaches  the  age  of  ten  without  exposing  for  himself,  draw- 
ing, dissecting  and  describing  the  brain  of  a  sharic. 

CoiNELL  Uhivessitv,  April  ao,  19081 


OBSERVATIONS  REGARDING  THE  INFLICTION  OF  THE 
DEATH  PENALTY  BY  ELECTRICITY. 

By  EDW.  ANTHONY  SPITZKA,  M.D. 
(Read  April  23,  1908.) 

A  great  variety  of  methods  of  inflicting  the  death  penalty  has 
been  devised  by  the  inventive  mind  of  man.  The  earlier  forms  are 
chiefly  characterized  by  cruelty,  by  an  intense  and  passionate  desire 
to  wreak  vengeance  and  inflict  pain  upon  the  condemned  and  to 
instill  terror  into  the  minds  of  onlookers.  I  will  not  review  the 
ancient  methods  in  detail.  There  is  the  burning  at  the  stake  by  the 
Romans,  Jews,  ancient  Britons,  Chinese  and  by  the  Spanish  Inqui- 
sition; beating  with  clubs  in  Greece  and  many  African  countries; 
beheading  by  axe  and  block,  the  sword  and  the  guillotine ;  blowing 
from  a  cannon,  either  by  lashing  the  condemned  to  the  muzzle  or 
by  thrusting  him  into  it  as  a  part  of  the  charge ;  boiling  in  water, 
oil,  melted  sulphur,  melted  lead ;  breaking  on  the  wheel;  burial  alive; 
crucifixion,  a  lingering  method  in  which  death  was  sometimes  has- 
tened by  the  thrust  of  a  spear  or  a  blow  with  a  club ;  crucifrangium, 
inflicted  on  Roman  slaves  and  Christian  martyrs  by  laying  the  legs 
of  the  condemned  upon  an  anvil  and  fracturing  the  bones  with  a 
heavy  hammer ;  decimation,  used  upon  mutinous  regiments  by  shoot- 
ing every  tenth  man ;  dichotomy  or  bisecting  the  body  with  a  saw ; 
dismemberment;  drawing  and  quartering;  drowning;  exposure  to 
wild  beasts;  flaying  dive;  flogging;  knouting;  garroting;  impale- 
ment;  the  "Iron  Maiden" ;  "  peine  forte  et  dure" ;  poisoning;  pound- 
ing in  a  mortar;  precipitation  from  a  great  height ;  the  rack;  running 
the  gauntlet;  shooting;  stabbing;  stoning;  strangling;  suffocating — 
in  short,  men  have  exercised  their  utmost  ingenuity  in  devising  means 
for  inflicting  cruel  torture  and  horrible  mutilation  upon  their  victims. 

As  is  well  known,  more  than  two  hundred  offenses  were  punish- 
able with  death  in  England  not  so  very  long  ago.  In  modern  times 
the  penalty  is  now  universally  limited  to  murder,  treason,  piracy  and 

39 


40  SPITZKA— OBSERVATIONS  REGARDING  INFLICTION      [April  .3 

military  offenses.  To  the  credit  of  William  Pemi  and  his  compan- 
ions it  may  be  recorded  that  in  1675,  when  they  founded  Pennsyl- 
vania, the  statutes  prescribing  death  for  all  sorts  of  offenses,  grave 
and  trivial,  were  left  behind  in  darkest  England  with  its  Newgate 
and  London  Tower,  and  the  only  one  retained  was  that  of  death  for 
aggravated  cases  of  murder. 

Electrocution. 

In  the  childhood  of  the  human  race  lightning  and  thunder  played 
an  important  part  in  the  religion  and  the  mental  life  of  the  various 
peoples.  Jupiter  ruled  the  world  by  his  thimderbolts.  The  Norse 
god  Thor  with  mighty  arm  wielded  the  hammer  of  lightning  in 
combat  with  the  enemies  of  the  gods.  Every  ancient  race  and  tribe 
has  been  awed  into  humble  submission  before  the  powerful  divinities 
imagined  to  preside  among  the  clouds  by  this  fascinating  phenome- 
non of  nature.  It  is  even  yet  feared  by  man,  for  is  not  its  dead- 
iiness  and  its  destructiveness  demonstrated  on  every  hand? 

It  is  now  more  than  a  century  and  a  half  ago  that  Benjamin 
Franklin,  accompanied  by  his  son,  went  to  a  field  in  the  neighbor- 
hood of  Philadelphia  as  a  thunder-storm  was  approaching  and  by 
his  famous  kite  experiment  discovered  that  lightning  was,  as  he 
shrewdly  had  surmised,  in  all  respects  similar  to  the  frictional  elec- 
tricity which  man  had  produced  artificially.  In  1760  Franklin 
erected  the  first  lightning  rod  upon  the  house  of  a  merchant  named 
West.  Although  more  than  five  hundred  persons  are  killed  and 
over  eight  hundred  are  injured  annually  in  the  United  States, 
Franklin's  invention,  wherever  used,  has  saved  countless  lives  and 
vast  amounts  of  property.  That  the  sage  Franklin  ever  foresaw 
the  likelihood  of  employing  this  death-dealing  and  mysterious  force 
in  the  infliction  of  capital  punishment  is  apparently  not  on  record. 

Electrocution  (more  properly  electrothanasia),  compounded  from 
"electro-execution,"  is  the  popular  name  for  the  infliction  of  the 
death  penalty  by  passing  through  the  body  of  the  condemned  a  cur- 
rent of  electricity  of  sufficient  intensity  to  cause  death.  The  method 
was  first  adopted  by  New  York  State  in  x888  by  a  law  which  became 
effective  on  January  x,  1889,  and  which  provides  how  many  persons 
may  witness  the  execution,  that  a  post-mortem  examination  of  the 


,9o8.J       OF  THE  DEATH  PENALTY  BY  ELECTRICITY.         41 

body  of  the  convict  be  perfcmned  and  that  the  body,  unless  claimed 
by  relatives,  be  interred  in  the  prison  cemetery  with  a  sufficient 
quantity  of  quicklime  to  consume  it. 

The  first  criminal  to  be  executed  by  electricity  was  William 
Kemmler,  on  August  6,  1890,  at  Auburn  Prison.  Since  that  time 
over  one  hundred  murderers  have  been  executed  in  New  York  State 
and  the  method  has  been  adopted  by  Ohio  (1896),  Massachusetts 
(1898),  New  Jersey  (1907),  and  Virginia  (1907-8). 

Reports  on  the  earlier  cases  have  been  published  by  Drs.  Carlos 
F.  MacDonald,  E.  C.  Spitzka,  E.  W.  Holmes,  and  with  reference 
to  nerve-cell  changes,  by  P.  A.  Fish.  ^ 

My  own  observations  are  based  upon  thirty-one  electrocutions 
(in  the  last  six  and  cme  half  years)  at  Sing  Sing  Prison,  Auburn 
Prison,  Dannemora  Prison  and  Trenton  (State  Penitentiary).  Of 
these  twenty-five  came  to  autopsy  at  my  hands. 

The  apparatus  consists  of  a  stationary  engine,  an  alternating 
dynamo  capable  of  generating  2,000  volts,  a  "  death-chair "  with 
adjustable  head-rest,  binding  straps  and  adjustable  electrodes.  [At 
Trenton  a  2400-volt  current  is  taken  from  the  public  service  wire 
and  lowered  to  the  desired  tension  by  a  rheostat.] 

The  voltmeter,  ammeter  and  switchboard  controlling  the  current 
is  located  in  the  execution-room ;  the  dynamo-room  is  communicated 
with  by  electric  signals.  Before  each  execution  the  apparatus  is 
thoroughly  tested.  When  everything  is  in  readiness  the  criminal  is 
brought  in  unfettered  and  usually  tmassisted,  and  seats  himself  in 
the  chair.  His  head,  chest,  arms  and  legs  are  secured  by  broad 
straps,  an  electrode  thoroughly  moistened  with  saturated  salt-solution 
is  affixed  to  the  head,  another  to  the  calf  of  one  leg,  both  electrodes 
being  molded  so  as  to  assure  good  contact.  The  head  is  not  shaved 
as  is  popularly  thought 

The  application  of  the  current  is  usually  as  follows :  The  contact 
is  made  with  a  high  potential  (1,800  volts)  for  5-7  seconds,  reduced 
to  200-250  volts  until  a  half  minute  has  elapsed;  raised  to  high 
voltage  for  3-5  seconds,  again  reduced  to  low  voltage  until  one 
minute  has  elapsed,  when  it  is  again  raised  to  the  high  voltage  for 
a  few  seconds  and  the  contact  is  broken.    The  ammeter  usually 


42  SPITZKA-OBSERVATIONS  REGARDING  INFLICTION      [April  «3, 

shows  that  from  seven  to  ten  amperes  have  passed  through  the 
criminal's  body. 

A  second  or  even  a  third  brief  contact  is  sometimes  made,  partly 
as  a  precautionary  measure,  but  more  to  completely  abolish  reflexes 
in  the  dead  body. 

The  time  consumed  by  the  strapping-in  process  is  usually  about 
forty-five  seconds  and  the  first  contact  is  made  a  few  secotids  later. 
In  all  about  60-70  seconds  elapse  from  the  moment  the  ccmvict 
leaves  his  cell  until  he  is  shocked  to  death. 

The  convicts  that  I  have  seen  thus  dealt  with  have  usually  slept 
soundly  the  pight  before,  they  have  entered  the  room  calmly  and 
stolidly,  often  with  a  half  smile  on  their  lips,  some  without  uttering 
a  word,  others  repeating  a  brief  prayer,  still  others  with  a  cheerful 
good-bye  to  those  present.  They  usually  seated  themselves  without 
betraying  any  signs  of  fear  or  trembling,  curiously  watching  the 
strapping-in  process  for  a  while,  then  sitting  erect,  looking  straight 
ahead  at  nothing  in  particular. 

The  physician  in  charge  observes  the  respiratory  movements  of 
the  prisoner  and  signals  to  the  electrician  at  a  moment  when  the 
lungs  contain  the  minimum  quantity  of  air.  At  the  moment  that 
the  contact  is  made  the  criminal's  body  stiffens  in  a  state  of  tonic 
muscular  spasm,  restrained  by  the  straps.  This  spasm  abates  some- 
what as  the  voltage  is  reduced,  to  again  attain  its  maximum  with 
each  raise  of  voltage.  When  the  current  is  interrupted  the  body 
collapses  completely.  An  examination  by  the  physicians  usually 
fails  to  elicit  any  signs  of  life.  Occasionally,  there  is  heard  a  turbu- 
lent, incoordinate,  accelerated  heart-beat,  but  apparently  limited  to 
the  auricular  chambers  of  the  heart.  In  only  two  cases  was  there 
any  respiratory  effort  and  this  was  limited  to  a  single  contraction  of 
the  thoracic  respiratory  muscles.  An  additional  brief  contact  or 
two  regularly  abolished  these  reflex  phenomena. 

The  reason  for  making  the  contact  at  the  moment  that  the  convict 
has  expired  air  from  his  lungs  in  the  natural  course  of  his  breathing 
is  this — and  it  will  explain  why  certain  witnesses  of  the  first  electro- 
cution thought  that  life  still  existed  in  Kemmler's  body.  It  must 
be  recalled  that  there  is  created  a  terrifically  powerful  spasmodic  con- 
traction of  all  muscles,  including  the  sphincters  and  the  glottis. 


,9oS.l  OF  THE  DEATH  PENALTY  BY  ELECTRiaTY.  48 

The  closure  of  the  glottis  confines  whatever  air  may  be  in  the  lungs ; 
upon  interrupting  the  current  the  body  becomes  entirely  limp,  the 
glottis  partly  relaxes,  the  thorax  collapses  and  the  contained  air 
rushes  through  the  partly  closed  glottis.  A  sound  resembling  a  sigh 
or  half  g^oan  may  be  thus  produced  upon  the  body  of  any  dead 
animal;  a  little  mucus  present  augments  the  sound  into  a  gurgle. 
It  is  no  wonder  that  inexperienced  persons  then  believe  life  to  be 
still  present. 

The  death  is  undoubtedly  painless  and  instantaneous.  The  vital 
mechanisms  of  life,  circulation  and  respiration,  cease  with  the  first 
contact.  Consciousness  is  blotted  out  instantly  and  the  prolonged 
application  of  the  current  as  it  is  usually  practised  by  Mr.  E.  F. 
Davis,  the  state  electrician  of  New  York,  ensures  the  permanent 
derangement  of  the  vital  functions  so  that  there  could  be  no  recovery 
of  these.  Occasionally,  the  drying  of  the  sponges  through  undue 
generation  of  heat  causes  desquamation  or  superficial  blistering  of 
the  skin  at  the  site  of  the  electrodes,  but  not  often.  Post-mortem 
discoloration,  or  lividity,  often  appears  during  the  first  contact. 
The  pupils  of  the  eyes  dilate  instantly  and  remain  dilated  in  death. 

The  post-mortem  examination  of  "electrocuted"  criminals  re- 
veals a  number  of  interesting  phenomena. 

The  temperature  of  the  body  rises  promptly  and  reaches  as  high 
as  I20®  F.  to  129J''  F.  within  twenty  minutes  in  many  cases.  After 
the  removal  of  the  brain  the  temperature  recorded  in  the  vertebral 
canal  was  often  over  120**  F.  The  development  of  this  high  tem- 
perature is  to  be  regarded  as  resulting  from  the  active  metabolism 
of  tissues  not  (somatically)  dead  within  a  body  where  all  vital 
mechanisms  have  been  abolished,  there  being  no  circulation  to  carry 
off  the  generated  heat.  The  maximum  heat  is  generated  at  the  site 
of  the  leg-electrodes,  where  muscle  (myosin)  coagulation  is  most 
extensive.  Furthermore,  the  release  of  from  ten  to  twenty  horse- 
power of  energy  within  the  body  must  contribute  materially  to  the 
caloric  increase. 

The  heart,  at  first  flaccid  when  exposed  after  death,  soon  con- 
tracts and  assumes  a  tetanized  condition.  This  is  particularly 
marked  in  the  left  ventricle;  on  the  whole  the  organ  assumes  the 
form  of  a  heart  in  systole.     In  one  case  (Koenig)  the  right  ven- 


44  SPITZKA— OBSERVATIONS  REGARDING  INFLICTION      [April  as, 

tricular  wall  of  the  heart  had  ruptured  in  several  places.  In  one 
case  I  was  able  to  elicit  slight  fibrillar  contractions,  limited  to  the 
small  area  stimulated,  by  touching  the  wall  of  the  heart  with  a  cold 
instrument.  In  several  cases  mechanical  irritation  of  the  atrio- 
ventricular  bundle  elicited  slight  contractions  limited  to  the  cdumnse 
cameae  and  the  papillary  muscles  of  the  left  ventricle.  In  experi- 
ments conducted  with  Professor  Coplin  upon  one  of  these  bodies, 
this  mode  of  contraction  could  be  called  forth  by  faradaic  stimula- 
tion, although  no  response  was  elicited  by  direct  stimulation.  In 
the  same  individual  it  was  impossible  to  elicit  any  response  7/ia  the 
nerve  system,  either  through  stimulation  of  the  cortex  (exposed 
within  about  ten  minutes),  the  spinal  cord  or  peripheral  nerves, 
although  muscular  reflexes  could  always  be  called  forth  by  directly 
stimulating  the  muscle. 

The  lungs  are  usually  devoid  of  blood  and  weigh  only  seven  or 
eight  ounces  avoirdupois  each. 

The  blood  is  ,prof oundly  altered  bio-chemically.  It  is  of  a  very 
dark,  brownish  hue,  and  it  rarely  coagulates.  Either  the  fibrinogen, 
or  the  fibrin-ferment,  or  both,  are  destroyed. 

The  maximum  damage  is  undoubtedly  wrought  in  the  nerve 
system  though  this  is  not  always  manifest.  Regarding  the  histo- 
logic changes,  reports  from  various  sources  vary.  There  is  a  gen- 
eral agreement  as  to  the  frequent  occurrence  of  capillary  hemor- 
rhages, disruptive  and  destructive  for  adjacent  tissues.  In  the 
nerve-cells  themselves  there  appears  to  be  no  apparent  change, 
although  there  must  have  resulted  terrific  molecular  change.  P.  A. 
Fish  found  vacuoles  in  one  case,  but  no  visible  changes  in  another. 
Aside  from  the  capillary  hemorrhages  and  the  arterial  anemia  with 
venous  congestion,  the  brain  shows  no  gross  changes  of  appear- 
ance. In  a  case  of  accidental  death  from  contact  with  an  alternating 
current  of  i,ooo  volts  for  about  one  half  minute,  Jellinek  found 
extensive  streaks  of  capillary  hemorrhages  in  the  gray  substance  of 
brain  and  spinal  cord  together  with  more  or  less  destruction  of  the 
nerve  cells,  extrusion  of  the  cell  nucleus,  etc. 

In  the  case  of  StroUo,  I  have  had  sections  made  of  the  pons, 
oblongata  and  spinal  cord  by  my  colleague.  Dr.  Radasch,  and  these 
have  revealed  curious  circular  areas  with  a  peripheral  zone  of  con- 


i9o8.]  OF  THE  DEATH  PENALTY  BY  ELECTRICITY.  45 

densation  which  fades  off  into  the  surrounding  unaffected  areas. 
The  bulk  of  the  central  rarified  portion  shows  a  delicate  network  of 
loose  fibrillae  which  in  all  probability  are  glia  fibers.  The  cellular 
elements  in  the  rarefied  area  are  few  in  number  though  apparently 
free  nuclei  are  scattered  in  this  portion.  These  areas  follow  more 
or  less  closely  the  course  of  the  finer  vessels.  Many  of  them  contain 
an  unruptured  vessel  centrally  located,  while  others  contain  longi- 
tudinal sections  with  the  areas  arranged  in  a  bead-like  manner  along 
such  vessel.  These  areas  are  larger  and  more  nimierous  in  the  pons 
than  in  the  oblongata  and  spinal  cord  and  apparently  distributed  in 
the  longitudinal  directions  more  frequently  than  in  other  directions. 
They  seem  to  resemble  gaseous  emphysema  and  are  possibly  due  to 
the  electrolytic  liberation  of  gas  in  the  peri-vascular  spaces.  One 
is  reminded  of  the  punctures  in  a  piece  of  paper  interposed  in  the 
path  of  the  sparks  of  a  static  machine. 

Through  the  courtesy  of  Mr.  Wilson  H.  Brown,  Sheriff  of 
Philadelphia,  I  was  permitted  to  witness  a  number  of  hangings  and 
thus  was  enabled  to  compare  the  new  method  with  the  old. 

The  preparations  for  the  execution  were  always  swiftly  con- 
ducted. Upon  this  point  comparison  favors  neither  method.  But 
after  the  drop  through  the  trap-door  the  ensuing  seconds  and  even 
minutes  bear  a  different  tale.  In  nearly  all  cases  the  heart  beats  for 
about  thirteen  minutes.  In  no  case  could  fracture  of  a  cervical 
vertebra  or  rupture  of  ligaments  be  determined  in  the  ordinary 
examination. 

In  one  case  only  was  there  no  movement  of  the  body  after  the 
drop,  although  the  heart  beat  the  usual  length  of  time.  This  prisoner, 
a  Chinaman,  apparently  died  in  syncope  or  of  apoplexy.  In  others 
the  unconsciousness  produced  by  the  first  shock  of  the  drop  appeared 
to  abate  and  in  several  instances  there  were  conscious— or  at  least 
semi-conscious — efforts  at  respiration,  efforts  to  reach  the  neck 
where  the  choking  sensation  was  unbearable,  efforts  at  reaching  for 
a  support  for  the  feet  manifested  by  such  vigorous  efforts  that  sev- 
eral witnesses  fainted  at  the  sight. 

They  veritably  "  danced  upon  the  air  "  until  the  asphyxia  (apnea) 
became  so  profound  as  to  blot  out  consciousness,  apparently  after 
one  or  one  and  one  half  minutes  in  some  cases. 


46  SPITZKA— OBSERVATIONS  REGARDING  INFLICTION      [April  .3, 

The  anatomy  of  hanging  has  been  frequently  discussed.  A 
recent  publication^  by  Dr.  Frederic  Wood  Jones  gives  the  results  of 
the  examination  of  the  bodies  of  one  hundred  men  executed  in  Nubia 
in  Roman  and  Byzantine  times.  Sixty-two  were  in  one  trench,  forty 
in  another.  They  were  all  adult  males,  with  cords  binding  the  legs 
and  arms  trussed  to  the  sides.  The  hanging  rc^e  was  still  in  situ 
on  one. 

Not  a  single  case  of  damage  to  the  cervical  vertebrae  was  found. 
The  most  commonly  found  lesion  was  an  oblique  opening  of  the  sutures 
of  the  skull,  so  that  one  portion  of  the  skull,  represented  by  the  occip- 
ital and  temporal  bones  becomes  pulled  aside  from  the  other  portion, 
represented  by  the  facial  part  of  the  skull  and  the  other  temporal 
bone.  The  basilar  suture  in  most  cases  was  also  disunited.  The 
skulls  all  gave  evidence  of  blood  staining. 

This  remarkable  finding  of  evidence  dating  about  2,000  years 
back,  prompted  me  to  examine  the  head  and  neck  bones  of  five  indi- 
viduals executed  by  hanging  and  sent  to  the  JeflFerson  Medical  Col- 
lege for  dissection.  In  not  a  single  instance  could  I  find  a  frac- 
tured cervical  vertebra  or  a  separation  of  any  cranial  suture.  Death 
had  ensued  through  strangulation. 

The  Newgate  Calendar  and  other  criminal  records  are  full  of 
instances  in  which  the  rope  broke  and  the  condemned  had  to  be 
rehanged  and  even  cases  where  the  head  was  severed  from  the  body. 
Furthermore,  there  are  not  a  few  authentic  cases  of  resuscitation  and 
total  recovery  after  hanging. 

Compared  with  hanging  as  well  as  other  methods,  electrocution 
is  the  most  humane,  decent  and  scientific  method  of  inflicting  the 
death  penalty  because  of  its  efficiency,  quickness  and  painlessness, 
and  it  should  be  adopted  by  Pennsylvania  as  well  as  every  state  in 
the  Union.  The  executions  should  take  place  in  a  building  remote 
from  the  penitentiaries  where  other  convicts,  more  or  less  susceptible 
to  reformation,  are  confined.  The  erection  of  scaffolds  in  prison 
corridors  or  the  knowledge  on  the  part  of  other  convicts  that  an 
electrocution  is  in  progress  has  a  bad,  even  brutalizing,  effect  upon 
them. 

*  British  Medical  Journal,  March  28,  1908. 


i9o8.]  OF  THE  DEATH  PENALTY  BY  ELECTRICITY.  47 

At  the  time  when  objections  to  the  hangman's  bungling  were 
most  strongly  urged  in  favor  of  some  better  method,  poisoning  by 
prussic  acid  as  well  as  chloroform  were  suggested.  With  regard 
to  the  injection  of  prussic  acid  by  means  of  the  hypodermic  syringe, 
the  Gerry  Commission  report  says : 

"This  is  open  to  the  very  serious  objection  that  the  use  of  that  in- 
strument is  so  associated  with  the  practice  of  medicine,  and  as  a  legitimate 
means  of  alleviating  human  suffering,  that  it  is  hardly  advisable  to  urge  its 
application  for  the  purposes  of  legal  executions  against  the  almost  unanimous 
protest  of  the  medical  profession." 

It  seems  to  me  that  the  use  of  chloroform,  first  suggested  by 

Wilder  in  1870,  cheap  and  efficient  as  it  would  be,  is  open  to  the 
same  objection.  There  should  be  a  lively  sense  of  violence,  of  mys- 
teriously overwhelming  power,  of  potent  force  and  destructive 
energy  attached  to  the  means  employed  in  putting  the  murderous 
ruffian  out  of  existence.  If  any  sentimentality  is  to  obtain  in  rela- 
tion with  capital  punishment  methods  it  should  not  be  in  favor  of 
the  "plug  ugly"  wielders  of  the  stiletto,  black-jack  and  the  ever- 
ready  revolver. 

Capital  punishment  has  been  abolished  in  Rhode  Island,  Maine, 
Michigan  and  Wisconsin.  Kansas  had  abolished  it  but  restored  it 
after  a  negro  was  burned  at  the  stake  for  an  outrage  upon  a  woman. 
The  states  of  New  York,  Colorado  and  Iowa  deemed  it  wise  to 
reenact  the  death  penalty  after  it  had  once  been  abolished,  owing  to 
the  increase  of  crime.  (The  same  experience  was  met  with  in 
Switzerland  where  the  penalty  was  abolished  in  1874  and  again 
established  in  several  cantons  in  1879.) 

In  Louisiana  the  death  penalty  is  inflicted  for  assault  with  intent 
to  kill,  arson,  burglary  and  administering  poison. 

In  Delaware  and  North  Carolina  arson  and  burglary  are  capital 
crimes. 

In  Missouri  seven  crimes  are  punishable  by  death ;  among  them 
are  murder,  train  robbery,  arson,  perjury  in  a  capital  case  and 
mayhem. 

In  Connecticut  the  law  prescribes  the  death  penalty  for  placing 
obstructions  on  a  railroad  track. 

In  Utah  the  law  provides  that  the  ccMidemned  may  choose  between 
hanging  and  shooting. 

PROC.  AMER.  PHIL.  SOC.  XLVII.  l88  D,  PRINTED  JULY  lO,  I908. 


48  SPITZKA— OBSERVATIONS  REGARDING  INFLICTION      [April  .3. 

The  question  "  Is  capital  punishment  justifiable  "  ?  has  agitated 
the  minds  df  men  ever  since  the  dawn  of  civilization.  Public  opin- 
ion is  never  so  fickle  with  regard  to  any  problem  of  life  as  this  one. 
My  own  opinion  is  a  firm  conviction  in  favor  of  it  for  those  who 
comm.it  premeditated  murder,  arson,  train-wrecking  and  bomb- 
throwing.  Society  needs  this  penalty  for  its  own  protection  and  it 
is  authorized  to  use  it.  The  Mosaic  law  "Thou  shalt  not  kill" 
refers  to  murder  and  not  to  legal  execution.  The  fear  of  death  is 
in  most  men  and  it  is  therefore  the  most  powerful  means  of  intimi- 
dation. Optimists  may  hope  to  see  society  organized  upon  such  an 
enlightened  plane  that  the  penalty  need  not  be  resorted  to — ^but  that 
time  is  not  yet  at  hand.  In  nearly  every  county  or  state  which  abol- 
ished the  penalty,  the  subsequent  increase  in  crime  aroused  a  clamor 
for  its  reestablishment. 

The  opinion  is  held  by  some  that  the  penalty  fails  to  act  as  a  deter- 
rent for  others.  The  argument  is  puerile,  for  this  country  at  least, 
inasmuch  as  only  1.3  per  cent,  of  homicides  are  convicted.  In  Ger- 
many 95  per  cent,  are  convicted,  or,  proportionately,  thirteen  times 
as  many.  Were  the  penalty  as  rigorously  enforced  in  the  case  of 
murder  as  the  whipping-post  is  used  in  Delaware  for  various  crimes, 
its  deterrent  effects  would  soon  become  manifest.  It  is  idle  to  talk 
of  anything  but  prompt  punishment  as  a  deterrent  of  crime. 

Thus,  in  New  York  City,  in  1904,  there  were  147  first  degree 
murders;  but  there  were  only  27  convictions  and  only  two  were 
executed.  In  the  same  year,  in  Philadelphia,  48  murder  trials  re- 
sulted in  only  7  verdicts  of  murder  in  the  first  degree  and  several 
of  these,  on  re-trial,  received  minor  sentences.  London,  with  6,000,- 
000  inhabitants,  had  24  murders ;  9  were  hanged  therefor.  Chicago, 
with  2,000,000  inhabitants,  had  128  murders ;  only  x  was  hanged. 

The  tardy  justice  meted  out  to  murderers  is  the  most  deplorable 
feature  of  our  legal  machinery  to-day.  There  are  too  many  loop- 
holes for  escape — ^long  delays,  endless  appeals,  lots  of  slush  about 
the  "unwritten  law,"  numerous  legal  technicalities  and  sentimental 
juries.  By  the  pettifogging  of  criminal  law  the  great  majority  of 
cases  are  granted  new  trials  in  the  United  States ;  in  Great  Britain 
only  3.5  per  cent.  Nearly  always  the  appeal  is  based  upon  points  of 
pleading  and  practice  and  many  years  elapse  before  the  final  settle- 


N 


X908.I  OF  THE  DEATH  PENALTY  BY  ELECTRICITY.  49 

ment  of  the  case.  Our  administration  of  justice  has  degenerated 
into  a  sort  of  "  rose-water  penology."  Its  demoralizing*  effect  upon 
the  community  is  manifested  by  the  rapid  increase  of  crimes  of 
violence  among  juveniles,  so  ready  to  imitate  and  emulate  their 
seniors  in  crime.  We  have  become  too  much  accustomed  to  failure 
of  justice  in  murder  cases.  This  blot  upon  our  civilization  is  largely 
the  outcome  of  our  indifference  to  the  way  many  criminal  courts 
are  conducted.  Certain  magistrates  make  a  farce  out  of  serious 
business,  lawyers  wrangle  with  each  other  unchecked,  witnesses  are 
brow-beaten  and  bribery  and  corruption  of  political  complexion 
degrade  the  proceedings  to  the  level  of  a  saloon  or  gambling-den 
or  a  policy-shop  rather  than  a  court  of  law. 

The  explanation  is  sometimes  given  that  "  hard  times  "  influence 
this  appalling  increase  of  crime.  That  this  is  not  so  can  be  readily 
shown  by  reference  to  statistics.  I  would  rather  point  to  the  moral 
deterioration  indicated  by  the  manner  in  which  large  sums  of  money 
are  stolen  or  used  for  bribery  and  corruption  and  the  luxury  and 
reckless  extravagance  with  which  some  wealthy  persons  (who  ought 
to  be  in  the  penitentiary)  offend  the  decent  class  of  our  population. 
Add  to  this  the  manner  in  which  the  newspapers  set  forth  the 
details  of  brutal  crimes  and  breed  familiarity  with  thoughts  of  crime. 

Society  has  relaxed  too  much.  The  death  penalty  is  a  necessity 
and  must  not  be  abolished,  else  all  discipline  of  society  will  be  relin- 
quished. Though  society  "  revolts  at  the  old  religious  dogma  of  the 
retribution  of  hell,  the  church  still  retains  it  as  essential  in  its  terrible 
dissuading  appeal  to  the  imagination  of  men"  {New  York  Sun). 
Let  us,  therefore,  in  our  penology,  adhere  to  what  the  test  of  time 
has  proven  to  be  an  efficient  check  if  only  it  be  carried  out  as  has 

been  done  in  Germany  and  Great  Britain. 

Jefferson   Medical  College, 
Philadelphia. 


50  SPITZKA— OBSERVATIONS  REGARDING  INFUCTION      [April  .j. 

BIBLIOGRAPHY. 
KacDonald,  C.  F. 

Report  on  the  execution  by  electricity  of  William  Kemmler,  alias  John 
Hart,    Albany,  N.  Y.,  1890. 

KacDonald,  C.  F. 

The  infliction  of  the  death  penalty  by  means  of  electricity,  being  a  report 
of  seven  cases.    Albany,  N.  Y.,  1893. 

Spitzka,  E.  C. 

Vorlaufige  Mittheilung  betreffs  des  Leichenbefundes  bei  dem  ersten  durch 
Elektricitat  Hingerichteten.  Medicinische  Monatschrift  (New  York), 
August,  1890. 

Fish,  P.  A. 

The  action  of  strong  currents  of  electricity  upon  nerve  cells.  Proc, 
American  Microscopical  Soc,  XVII.,  1895. 

Bell,  CUrk. 

Electricity  and  the  death  penalty.    Alienist  and  Neurologist. 

HolmeB,  E.  W. 

Anatomy  of  Hanging.    Pennsylvania  Med.  Jour.,  July,  1901. 

Gerry,  Southwick  and  Hale. 

Report  of  the  G)mmission  to  Investigate  and  Report  the  Most  Humane 
and  Practical  Method  of  Carrying  into  Effect  the  Sentence  of  Death  in 
Capital  Cases.    Albany,  N.  Y.,  January  17,  1888. 

Spitzka,  E.  A. 

Report  of  the  Postmortem  Examination  of  Leon  Czolgosz.  American 
Journal  of  Insanity,  1902. 

Spitzka,  E.  A. 

Execution  and  Postmortem  Examination  of  the  three  Van  Wormcr 
Brothers.    N.  Y.  Daily  Medical  Journal,  February  i,  1904. 

Spitzka,  E.  A. 

Notes  on  autopsy  of  Toni  Turkofski,  electrocuted  murderer.     Medical 
Critic,  August,  1903. 

Jellinek,  S. 

Elektropathologie.    Stuttgart  (F.  Enke),  1903. 


PRELIMINARY  NOTE  ON  THE  BRAINS  OF  NATIVES  OF 
THE  ANDAMAN  AND  NICOBAR  ISLANDS. 

By  EDW.  ANTHONY  SPITZKA,  M.D. 
(Read  April  23,  igo8.) 

Physical  anthropology,  or  comparative  htunan  morphology,  has 
been  largely  based  upon  cranial  configuration.  Since  the  days  of 
Camper  and  Blumenbach,  the  classification  of  the  human  races  is 
based  on  more  conprehensive  morphologic  foundations,  for  with 
cranial  morphology  as  the  first  criterion,  there  have  been  added 
criteria  derived  from  the  entire  skeleton,  the  soft  tissues  and  the 
brain.  The  last-mentioned  organ  has  been  the  least  studied  because 
it  is  usually  most  difficult  to  obtain,  preserve  and  study.  Never- 
theless, interest  in  this  subject  is  manifestly  increasing  among  anato- 
mists and  anthropologists,  for  they  appreciate  the  fact  that  there  is 
a  pressing  need  for  fruitful  research  in  anthropologic  encephalometry 
among  the  exotic  races,  so  rapidly  becoming  impure  or  even  ex- 
tinct Many  American  Indian  tribes  have  disappeared ;  the  volcanic 
outbreak  in  Martinique  has  wiped  out  nearly  all  Caribs.  The  Aus- 
tralian natives  driven  to  the  desiccated  wastes  of  the  interior,  many 
African  tribes  succumbing  in  the  arid  deserts,  the  Eskimos  deci- 
mated by  epidemics  of  small-pox,  measles  and  pneumonia — all  these 
and  many  others  are  dying  out  and  warn  us  to  make  haste  in  record- 
ing observations  upon  them  while  they  still  exist 

It  has  been  my  good  fortune  to  pursue  comparative  studies  in 
cerebral  morphology  based  upon  the  brains  of  the  white  race,  of 
Eskimos,  Japanese,  Chinese,  Negroes  and  Papuans.  I  am  now  able 
to  add  the  brain  of  a  native  of  the  Andaman  Islands  and  one  of  a 
native  of  the  Nicobar  Islands.  For  this  exceptional  privilege  I  am 
indebted  to  the  eflforts  of  Dr.  W.  W.  Keen,  whose  correspondence 
with  Lord  Curzon,  then  Viceroy  of  India,  opened  the  way  to  com- 
munication with  Mr.  H.  H.  Risley,  Director  of  Ethnc^aphy  for 
India ;  Mr.  W.  R.  H.  Merk,  Superintendent  of  Port  Blair,  and  Major 

51 


52  SPITZKA— PRELIMINARY   NOTE  ON  THE  BRAINS  OF   [April  .3, 

A.  R.  S.  Anderson  (M.  B.  Cantab.)  I.M.S.,  senior  medical  officer  at 
Port  Blair,  Andamans.  To  all  these  I  desire  to  acknowledge  here- 
with my  thanks. 

The  Andaman  Archipelago  is  a  g^oup  of  densely  wooded  islands 
about  1,760  square  miles  in  area,  situated  in  the  Bay  of  Bengal  about 
180  miles  southwest  of  Cape  Negrais,  Burma,  and  about  60  miles 
distant  from  the  more  southerly  Nicobar  Islands.  The  inhabitants 
have  been  considered  a  most  primitive  and  savage  race.  Accoimts 
of  their  cannibalism  are  found  in  the  ancient  Chinese  writings  and 
the  Andamanese  are  probably  referred  to  by  Ptolemy  as  the  "  anthro^ 
pophag^."  Port  Blair  is  a  convict  settlement  and  the  convicts  are 
deterred  from  making  efforts  to  escape  by  their  fear  of  the  natives. 
From  the  observations  of  E.  H.  Man,  who,  more  than  any  other, 
has  made  the  race  a  study,  it  appears  that  the  Andamanese  are 
Negritos  and  not  Papuans.  They  are  wejl  made  and  well  propor- 
tioned. Their  skulls  are  brachycephalic.  Their  lips  are  not  thick, 
their  profiles  are  good  and  they  have  no  peculiar  odor  like  that  which 
is  found  in  the  African  race.  Their  extremities  are  small,  but  the 
heel  projects  slightly  to  the  rear.  The  average  height  of  the  men 
is  149  cm.,  of  the  women  140  cm.  The  average  weight  of  the  men 
is  98  and  93  pounds  respectively.  The  color  of  the  Andamanese  is 
generally  dark  bronze  or  copper  color ;  often  the  color  of  soot  and 
even  quite  black.  The  hair  is  woolly,  but  its  cross-secticm  is  not 
always  elliptical.  In  a  letter  to  the  Smithsonian  Institution,  Dr. 
Abbott  says  of  them :  "  They  are  a  happy,  merry,  little  people,  infan- 
tile both  in  looks  and  behavior.  Unfortunately  they  are  dying  out. 
Contact  with  civilization  is  making  the  women  barren  and  there  are 
comparatively  few  children." 

Mr.  Man  thinks  that  it  has  been  pretty  well  demonstrated  that 
these  Negritos  in  the  Andaman  Archipelago,  so  unlike  any  of  their 
immediate  neighbors,  are  aborig^es  and  have  inhabited  the  group 
from  prehistoric  times.  The  population  in  1901,  Dr.  Anderson 
writes  me,  was  2,200,  including  women  and  children. 

The  Andamanese  wear  no  clothing ;  its  place  is  taken  in  a  measure 
by  necklaces,  circlets  for  the  head,  garters,  bracelets  and  belts. 
They  live  in  thatched  huts  and  sleep  on  mats.  Stones  are  used  as 
anvil  and  hammer,  clam  shells  as  knives.    They  fashion  old  barrel 


,908.]  THE  ANDAMAN  AND  NICOBAR  ISLANDERS.  53 

hoops  from  wrecked  ships  into  jagged  knives.  The  only  thing 
resembling  a  musical  instrument  is  a  wooden  shield-like  drum  upon 
which  the  performer  keeps  time  by  striking  it  with  his  foot.  They 
make  some  pottery ;  the  base  of  the  pot  is  in  the  form  of  a  cup.  To 
this  roll  after  roll  is  added  and  the  sides  built  up,  the  inner  and 
outer  surfaces  are  smoothed  off  with  an  area  shell  and  ornamented 
with  wavy,  checkered  or  striped  designs  by  means  of  a  pointed  stick 
and  baked  by  placing  pieces  of  burning  wood  both  inside  and  arotmd 
the  vessel.  They  make  cane  baskets,  wooden  trays  and  buckets. 
String  is  made  from  vegetable  fiber  (orchid  and  Anadendron)  and 
used  in  making  harpoon  lines,  turtle  nets,  fishing  nets,  bowstrings 
coated  with  wax,  lashings,  reticules  and  necklaces.  Bows  and  ar- 
rows, harpoons  and  fish  spears  are  used  in  hunting.  They  build 
outrigger  canoes  and  simple  dugouts  which  are  propelled  by  pad- 
dles, or,  in  shallow  water,  by  poles  or  the  shaft  of  a  turtle  harpocm. 
Morphologically,  the  Andamanese  form  a  definite  g^oup.  The 
following  criteria  are  given  by  Duckworth  :^ 

Cranial   index    i 82.1 

Alveolar  index   102.0 

Nasal  index  50.9 

Height   index    77.9 

Cranial   capacity    i,^  c.c. 

,  The  skull  is  small  and  round,  with  prominent  jaws.  It  is  crypto- 
zygous,  muscular  ridges  are  not  very  prominent,  the  mastoid  processes 
are  small,  and  the  external  auditory  meatus  is  shallow.  Brow- 
ridges  are  not  developed,  and  sexual  differences  are  often  obscure 
in  these  skulls.  The  face  is  prognathous,  the  prognathism  affect- 
ing chiefly  the  alveolar  maxillary  margin  and  being  therefore  sub- 
nasal.  The  chief  distinction  is  the  association  of  a  highly  brachy- 
cephalic  skull  of  small  capacity  with  dwarf  stature  and  progna- 
thism. The  lumbo-vertebral  index  denotes  simian  affinities.  The 
sacral  curve  is  very  slight,  indicating  a  low  position  among  hominidse. 
The  scapula  is  the  most  pithecoid  amongst  hominidse  with  the  pos- 
sible exception  of  the  Bambut6  dwarfs  of  Africa.  The  proportions 
of  the  limb-bones  are  simian  as  regards  the  radio-humeral  and  the 
tibio- femoral,  but  not  as  regards  the  in termembral  of  humero- femoral 
indices. 

^Duckworth:   "Morphology   and  Anthropology/'   Cambridge   Biological 
Series,  1904. 


54 


SPITZK A— PRELIMINARY  NOTE  ON  THE  BRAINS  OF    lAprilta. 

The  Brain. 


[Preliminary  Report.] 

The  brain  is  that  of  an  Andamanese  named  Juran  of  the  tribe 
called  Aka-yere  or  Aka-Jaro-da.  Juran  was  a  male,  aged  about  45 
years,  who  died  at  Port  Blair  of  pulmonary  tuberculosis  on  June 
30,  1905.  The  brain  was  removed  by  Dr.  Anderson  about  one  hour 
after  death  and  immersed  in  a  mixture  of  formalin  and  water.     Its 


Fig.  I.    Brain  of  Andamanese  (dorsal  view). 


weight  while  fresh  was  not  noted  before  being  sealed  and  trans- 
ported. When  received  about  eight  months  later  it  weighed  1,193 
grams,  as  follows: 

Grams. 

Left   hemicerebrum    *532 

Right   hemicerebrum    525 

Cerebellum,   pons   and    oblongata •_J36 

Total    1,193 

The  specimen  continued  to  lose  weight  slightly  and  in  April,  1908, 
after  removal  of  the  cerebral  pia-arachnoid,  weighed  as  follows : 


,908.]  THE  ANDAMAN  AND  NICOBAR  ISLANDERS.  55 

Grams. 

Left   hemicerebrum    493 

Right   hemicerebrum    490 

Cerebellum,  pons  and  oblongata 130 

Total  7^13 

Various  calculations  indicate  that  the  fresh  weight  of  this  brain 
was  probably  between  1,200  and  1,250  g^ams. 

The  brain  is  broad  and  short.*  The  frontal  lobes  are  less  mas- 
sive than  in  whites.  The  fissuration  is  well  marked  but  not  very 
complex.  The  precallosal  length  is  less  than  in  whites.  The  cal- 
losum  is  of  good  size,  comparing  well  with  those  of  whites.  The 
calcarine  fissure  is  interrupted  on  the  left  side.  The  fissural  pecu- 
liarities must  be  considered  more  fully  in  the  final  publication,  and, 
if  possible,  should  be  based  upon  comparison  with  more  specimens 
from  natives  of  this  race.  The  following  dimensions  may  be  re- 
corded here : 

Centbsimals. 

Brain-length,    left    half 16.1 

Brain-length,   right   half i5-8 

Brain-width    i3-9 

Cerebral  index   86.9 

Horizontal  circumference   47-0 

Width,   left  hemicerebrum 6.9 

Width,  right  hemicerebrum 7-0 

Left  occipito-temporal  length    12.6 

Right  occipito-temporal  length   124 

Length  of  callosum * 7-3 

Percentage  of  callosal  length 45-3% 

Left  centro-temporal  height  10.6 

Right  centro-temporal  height    10.6 

Left  centro-ol factory  height   8.7 

Right   centro-ol  factory   height    8.7 


Arc  Measures. 

Frontal    14.5 

Left      \     Parietal    50 

Occipital    5.5 

Frontal    14.5 

Right     \      Parietal    4-5 

Occipital    6.0 


{ 
{ 

{ 


Cerebral  Indices. 

Frontal    58.0 

Left     {      Parietal    20.0 

Occipital    22.0 

Frontal    58.0 

Right     \      Parietal    18.0 

Occipital    24.0 

'It  was  somewhat  flattened  upon  its  dorsum  during  transportation. 


56 


SPITZKA— PRELIMINARY   NOTE  ON  THE  BRAINS  OF    [April  as, 


Left 
Lateral 
Aspect 


Left 
Mesal 
Aspect 


Right 

Lateral 

Aspect 


Horizontal  Distances   (in  Centesimals). 
From  Frontal  Point  to: 

1.  Tip  of  temporal  lobe 23.0 

2.  Sylvian-presylvian  junction   31.0 

3.  Ventral  end  of  central  fissure 44.1 

4.  Sylvian-episylvian  junction    62.1 

''   6.  Frontal  edge  of  callosum 22.3 

7.  Porta  (Foramen  of  Monro) 42.2 

8.  Dorsal  end  of  central  fissure 64.6 

9.  Dorsal  intersection  of  paracentral  fissure 71.4 

10.  Caudal  edge  of  callosum 67.9 

1 1.  Occipito-calcarine  junction 78.2 

^  12.  Dorsal  intersection  of  occipital  fissure 88.2 

1.  Tip  of  temporal  lobe 21.8 

2.  Sylvian-presylvian  junction   314 

3.  Ventral  end  of  central  fissure 46.8 

4.  Sylvian-episylvian  junction    634 


'   6.  Frontal .  edge  of  callosum 20.5 

7.  Porta  (Foramen  and  Monro) 41.6 

Right         8.  Dorsal  end  of  central  fissure 70.5 

Mesal    <    9.  Dorsal  intersection  of  paracentral  fissure 77.0 

Aspect       10.  Caudal   edge   of   callosum ^.3 

1 1.  Occipito-calcarine  junction    76.2 

L  12.  Dorsal  intersection  of  occipital  fissure 91.6 

Cross-section  area  of  callosum  =  5.85  sq.  ctm. 

NicoBARESE  Brain. 

With  regard  to  the  ethnic  position  of  the  Nicobarese  there  exists 
considerable  doubt.  They  are  very  different  from  the  Andamanese. 
Their  color  is  a  light  brown,  the  hair  is  straight  and  black,  and 
apparently  they  are  of  ancient  Mongolian  origin  with  probably  no 
admixture  of  Papuan  or  Negrito  elements.  Their  stature  is  medium 
(158-163  cm.),  not  small  as  are  the  Andamanese. 

The  brain  is  that  of  an  individual  from  Kar  Nicobar,  a  male, 
aged  25,  who  died  of  hypertrophic  cirrhosis  of  the  liver  and  fatty 
d^eneration  of  the  heart  in  the  hospital  at  Port  Blair.  The  brain 
weighed  49  ounces  avoirdupois  or  1,389  grams.  The  body-weight 
was  136.5  pounds,  while  the  stature  was  170  cm. 

Major  Anderson  injected  about  10  c.c.  of  5  per  cent,  formalde- 
hyde into  the  ventricles  through  the  tuber  and  immersed  the  brain 
in  the  same  mixture.  The  specimen  reached  me  in  March,  1906, 
a  little  over  three  months  after  its  removal  from  the  head. 

Its  present  weight,  divested  of  the  cerebral  pia-arachnoid,  is 
1,257  grams.  The  brain  is  somewhat  flattened  and  elongated.  The 
fissural  pattern  is  fairly  good  but  not  as  complex  as  in  the  average 


«9o8.] 


THE  ANDAMAN  AND  NICOBAR  ISLANDERS. 


57 


Fig.  2.    Brain  of  Nicobarese  (dorsal  view). 

vrhite  brain.  The  calcarine  fissure  is  interrupted  on  both  sides,  the 
interruption  being  somewhat  concealed  on  the  left  side.  The  cal- 
losum  is  small,  a  fraction  over  5  sq.  cm.  The  indusium,  however, 
is  quite  massive,  and  further  study  of  rhinencephalic  parts  may 
prove  interesting.  The  insula  is  slightly  visible  on  both  sides. 
The  measurements  of  this  specimen  are  as  follows : 

Ckntbsimals. 

Brain-length,   left   19.1 

Brain-length,   right    18.5 

Brain-width    '. 13.3 

Cerebral   index    70.O 

Horizontal  circumference 52.0 

Width,  left  hemicerebrum    6.9 

Width,  ri^ht  hemicerebrum   6.4 

Left  occipito-temporal  lobe  14.3 

Right  occipito-temporal  lobe 14.2 

Length  of  callosum   8.5 


58 


SPITZKA— PRELIMINARY  NOTE 


[April  n. 


Percentage  of  callosal  length 44-7% 

Left  centro-temporal   height    9.2 

Right  centro-temporal  height    8.7 

Left  centro-ol factory  height    8.2 

Right  centro-ol  factory  height   8.0 


Left 


Right 


Arc  Measures. 

{Frontal    16.0 

Parietal    4.5 

Occipital    5.5 

Frontal   16.5 

Parietal    5.0 

Occipital    5.0 


( 


Left 


Right 


{ 
{ 


Cerebral  Indices. 

Frontal    61.5 

Parietal    17.3 

Occipital    21.2 

Frontal   62.3 

Parietal    188 

Occipital   .t 18.8 


Horizontal  Distances   (in  Centesimals). 
From  Frontal  Point  to: 

1.  Tip  of  temporal  lobe 25.1 

2.  Sylvian-presylvian  junction    30.3 

3.  Ventral  end  of  central  fissure 41.8 

4.  Sylvian-episylvian  junction    ? 

6.  Frontal  edge  of  callosum 20.0 

7.  Porta  (Foramen  of  Monro) 38.2 

8.  Dorsal  end  of  central  fissure — 

9.  Dorsal  intersection  of  paracentral  fissure 65.4 

ID.  Caudal  edge  of  callosum 64.4 

11.  Occipito-calcarine  junction    81. i 

12.  Dorsal  intersection  of  occipital  fissure 83.2 

I.  Tip  of  temporal  lobe 24.6 

Lateral  -I     ^'  Sylvian-presylvian  j  unction 28.2 

3.  Ventral  end  of  central  fissure 41.3 

4.  Sylvian  episylvian  junction   55.0 

6.  Frontal  edge  of  callosum 18.3 

7.  Porta  (Foramen  of  Monro) 37.7 

8.  Dorsal  end  of  central  fissure 61.2 

9.  Dorsal  intersection  of  paracentral  fissure 63.8 

10.  Caudal  edge  of  callosum 63.8 

1 1.  Occipito-calcarine  j  unction  . , 75.4 

12.  Dorsal  intersection  of  occipital  fissure 874 

Cross-section  area  of  callosum  =  5.02  sq.  ctm. 

Jefferson  Medical  College, 
PhiladelpHla. 


Left 
Lateral 
Aspect 


Left 
Mesal 
Aspect 


Right     r 
Lateral  "K 
Aspect   L 


Right 
Mesal 
Aspect 


DETERMINATION  OF  DOMINANCE  IN  MENDELIAN 

INHERITANCE. 

By  CHARLES  B.  DAVENPORT,  Ph.D. 
(Read  April  25,  igo8.) 

The  longer  one  investigates  the  phenomena  of  heredity  the  more 
one  is  impressed  with  the  grandeur  of  the  discovery  made  over  forty 
years  ago  by  Gregor  Mendel.  His  method  is  not  less  important  than 
its  results.  Following  him,  in  studying  heredity  one  considers  a 
single  character  at  a  time.  One  notes  the  result  in  the  offspring 
when  this  character  assumes  contrasted  forms  in  the  two  parents  or 
when  one  parent  has  the  character  and  the  other  lacks  it  Under 
these  cir<nimstances  one  frequently,  nay,  usually,  finds  that  the  con- 
dition in  one  parent  dominates  over  that  in  the  other  parent,  so  that 
the  offspring  are  all  alike,  and  like  one  parent,  in  respect  to  that 
character.  The  opposite,  or  recessive,  quality  is  not  lost,  however. 
It  persists  in  the  germ  plasm  and  one  half  of  the  germ  cells  of  the 
individuals  belonging  to  the  first  generation  of  hybrids  contain  the 
dominant  and  one  half  the  recessive  quality. 

Dominance,  it  will  be  observed,  it  a  matter  of  the  soma.  The 
hybrid  fertilized  egg  contains  both  contrasting  qualities  and  so, 
probably,  do  all  of  the  cells  of  the  body.  But  only  one  of  the  quali- 
ties ordinarily  makes  its  appearance.  It  has  been  suggested  that  a 
struggle  occurs  between  the  contrasted  qualities  and  the  stronger — 
called  the  dominant — wins.  The  question  is  what  determines  this 
assumed  greater  strength  of  the  dominant  quality  ?  What  determines 
dominance? 

Various  replies  have  been  given  to  this  question.  It  has  been 
suggested  that  the  dominant  quality  is  the  older  and  although  this 
is  sometimes  true  it  so  often  fails  to  be  so  that  age  cannot  be 
regarded  as  the  primary  cause  of  dominance.  Frizzling  and  silki- 
ness  of  fowl's  feathers  are  each  novelties  but  one  dominates  over  the 
ordinary  flat  feather  and  the  other  is  dominated  by  it.     Much  evi- 

59 


60  DAVENPORT— DETERMINATION  OF  DOMINANCE        (April.,, 

dence  of  this  sort  could  be  adduced  proving  the  insufficiency  of  the 
theory  of  the  recessive  nature  of  novelties.  A  different  theory  has 
been  suggested  by  deVries,  namely,  when  an  individual  having  the 
characteristic  patent  Is  crossed  with  one  in  which  it  is  latent  the 
patent  characteristic  is  dominant,  the  latent  recessive.  A  similar 
expression  has  been  proposed  by  Hurst  who  concludes  that  the  pres- 
ence of  a  quality  usually  dominates  over  its  absence.  This  expres- 
sion of  the  facts  is,  in  the  main,  true  but  it  is  too  narrow,  inas- 
much as  it  assures  that  the  mendelian  result  occurs  only  when  a 
character  is  crossed  with  its  absence ;  but  this  I  shall  show  directly 
is  by  no  means  true. 

Two  years  ago  I  suggested  that  a  progressive  variation,  one 
which  means  a  further  stage  in  ontogeny,  will  dominate  over  a  con- 
dition due  to  an  abbreviation  of  the  ontogenetic  process — or  a  condi- 
tion less  highly  developed  than  the  first.    Recent  studies  have  thrown 


Fro.  I. 

additional  light  on  this  matter  and  I  wish  to  treat  it  now  generally. 
First  let  me  present  some  illustrations.  Many  poultry  have  feathers 
on  the  feet ;  these  constitute  the  so-called  boot.  If  a  "  booted  "  bird 
be  mated  with  a  non-booted  all  offspring  are  booted — booting  is 
dominant  over  its  absence.  Booting  occurs,  however,  in  an  iniinity 
of  grades.  For  convenience  I  recognize  ten,  usually  determined  by 
inspection.  If  a  bird  with  a  boot  of  grade  8  or  9  be  crossed  with  a 
bird  with  boot  of  grade  2  or  3,  both  being  pure  dominants,  then  the 
stronger  condition  is  dominant  in  the  offspring,  so  that  their  average 
grade  is  about  8. 

A  second  illustration  may  be  drawn  from  certain  studies  made 
on  the  asparagus  beetle  by  Dr.  F.  E.  Lutz,  of  the  Carnegie  Insti- 
tution of  Washington.  In  the  embryonic  condition  the  outer  wing 
covers  of  this  beetle  are  nearly  pigmentless  or  yellow.     Before 


1908.]  IN  MENDELIAN  INHERITANCE.  61 

emerging  from  the  pupal  condition  black  pigment  is  laid  down.  The 
pigmented  area  is  variable  in  amount.  The  more  extensively  pig- 
mented condition  is  dominant  over  the  less  extensively  pigmented 
(a  over  c,  d  or  e — see  Fig.  i).  In  this  case,  also,  it  is  clear  that  the 
facts  are  better  expressed  by  the  statement  that  the  more  developed 
condition  dominates  over  the  less  developed. 

Still  another  case  is  that  of  human  eye  color.  The  pigmentation 
of  the  iris  is  variable  in  amount.  The  blue  iris  is  without  pigment. 
A  small  amount  of  black  pigment  (with  or  without  yellow)  produces 
the  grays;  still  more  pigment  yields  browns  and  blacks.  Now  it 
appears  that  the  offspring  of  parents  one  of  whom  has  g^ay  eyes 
and  the  other  blue  eyes  will  have  gray  eyes  or  blue  eyes,  but  not 
brown  eyes;  and  gray  will  show  itself  dominant  over  blue.  Simi- 
larly brown  iris  color  is  dominant  over  gray;  the  more  advanced 
condition  of  pigmentation  over  the  less  advanced.  We  have  not 
here  to  do  with  a  qualitative  difference  of  the  presence  of  a  character 
opposed  to  its  absence,  but  of  a  qualitative  difference  only. 

The  heredity  of  human  hair  color  follows  a  similar  law.  In  one 
series  red  pigment  is  absent  in  the  hair  and  such  colors  as  flaxen 
or  tow,  light  brown,  brown,  dark  brown  and  black  may  be  distin- 
guished. The  records  collected  by  Mrs.  Davenport  and  myself 
show  that  two  flaxen-haired  parents  have  flaxen-haired  children  and 
probably  only  such.  Two  parents  with  light  brown  hair  have  chil- 
apparently  only  such.  Two  parents  with  light  brown  hair  have  chil- 
dren of  two  parents  each  with  dark  brown  or  black  hair  produce 
children  with  all  of  the  varieties  of  hair  color.  This  result  means 
that  any  lighter  color  is  recessive  to  any  darker  color. 

The  facts  recited  above  and  many  others  thus  support  the  view 
that,  where  various  stages,  a,  &,  r,  in  the  progressive  development  of 
a  quality  are  found  in  individuals  of  the  same  race  or  species,  the 
more  progressive  condition  will  often  behave  as  a  dominant  toward 
the  less  progressive  condition.  The  extreme  case  is,  of  course,  that 
in  which  the  organ  or  quality  is  absent  in  one  parent  and  present  in 
the  other;  but  this  seems  to  be  only  a  special  case  of  a  more 
general  law. 

As  to  the  universality  of  this  law  it  is  still  early  to  speak  with 
confidence.    We  know  too  little  of  the  developmental  factors  of  an 


62  DAVENPORT -DETERMINATION  OF  DOMINANCE         [April  as, 

organ  to  decide,  in  many  cases,  whether  a  difference  is  due  to  a 
progressive  or  a  retrogressive  change.  For  instance,  the  long  angora 
coat  of  rabbits  is  recessive  to  short  coat ;  and  this  has  been  cited  as 
a  clear  case  of  recessiveness  of  the  advanced  condition.  But  it 
seems  doubtful  if  such  is  the  case.  For  the  angora  coat  retains  an 
embryonic  quality  (viz.,  of  continued  growth)  which  is  present  in 
the  infancy  of  the  short-haired  rabbit  and  is  then  inhibited.  The 
inhibiting  factor  is  present  in  short-haired  rabbits  and  absent  in 
angora  rabbits  and  the  presence  of  the  inhibiting  factor  dominates 
over  its  absence.  At  one  time  I  thought  that  the  dominant  white 
plumage  of  some  poultry  was  a  case  of  dominance  of  absence  of 
color.  But  it  now  appears  that  we  have  among  poultry  recessive 
whites  which  are  true  albinos,  and  the  dominant  whites  which  must 
be  regarded  as  "grays,"  in  which  pigmentaticMi  is  obscured  by  an 
additional  factor  like  that  which  turns  black  hair  gray.  This  gray- 
ing factor  is  dominant  over  its  absence. 

It  is  possible  that  the  future  may  show  that,  in  accordance  with 
the  ideas  of  deVries,  an  advanced  grade  of  a  character  may  be 
regarded  as  a  sum  of  minute  equivalent  elementary  units;  by  the 
dropping  out  of  these  units  one  at  a  time  a  character  passes  through 
a  series  of  degradational  stages.  Then  a  light  brown  hair  may 
have  one  unit  of  melanic  pigment,  brown  hair  two  units,  dark  brown 
three  units,  and  black  hair  four  units.  If  this  should  prove  to  be 
true  then  the  four  unit  condition  would  dominate  over  the  three 
unit  condition,  or  the  fourth  unit  would  dominate  over  its  absence. 
But  such  evidence  as  I  have  at  present  does  not  favor  this  view.  I 
am  inclined  rather  to  the  hypothesis  that  when  the  germinal  deter- 
miner of  greater  intensity  meets  that  of  less  intensity  it  dominates 
over  the  latter.  This  hypothesis  receives  support  from  another  set 
of  facts  which  go  to  prove  that  the  idea  of  varying  intensity  of  a 
determiner  is  a  true  one.  This  set  of  facts  is  derived  from  the 
combs  of  poultry.  In  one  race  of  poultry — Polish  fowl — ^the  comb 
consists  of  a  pair  of  horns  or  broad  flaps  which  lie  far  back  near  the 
base  of  the  beak ;  and  there  is  no  median  comb.  In  the  Minorca  and 
most  other  fowl  there  is  a  single  median  comb.  Now  when  these 
two  races  are  crossed  we  find  that  the  median  comb  dominates  over 
the  absence  of  median  comb ;  sometimes  completely,  running  in  the 


,9ol]  IN  MENDEUAN  INHERITANCE.  63 

hybrid  from  the  base  of  the  beak  to  in  front  of  the  nostrils ;  some- 
times incompletely,  occupying  only  the  anterior  half  or  fourth  of  the 
beak.  It  seems  to  me  clear  that  in  the  varying  proportions  of  this 
median  comb  in  the  hybrids  we  have  at  once  evidence  for,  and  a 
measure  of,  varying  intensity  of  dominance.  Now  it  may  reason- 
ably be  asked  whether,  when  the  long-combed  and  short-combed 
hybrids  are  mated  together,  the  long  comb  dominates  over  the  short 
The  answer  is  complicated  by  the  fact  that  the  Polish  "horns" 
reappear  in  this  second  generation ;  but,  leaving  this  aside,  we  find 
that  there  is  a  greater  preponderance  of  long  median  combs  than 
simple  mendelian  expectation  calls  for  and  this  indicates  that  the 
longer  median  comb  tends,  but  not  always  perfectly,  to  dominate  the 
shorter  median  comb;  or,  in  other  words,  the  more  intense  deter- 
miner dominates  the  less  intense. 

To  sum  up,  I  think  it  is  clear  that  dominance  in  heredity  appears 
when  a  stronger  determiner  meets  a  weaker  determiner  in  the  germ. 
The  extreme  case  is  that  in  which  the  strong  determiner  meets  a 
determiner  so  weak  as  to  be  practically  absent  as  when  a  red  flower 
is  crossed  with  a  white.  In  such  cases  we  have  the  clearest  exam- 
ples of  mendelian  inheritance.  But  there  is  an  entire  gamut  of  cases 
where  the  opposed  determiners  are  of  varying  relative  potency.  The 
phenomenon  of  determinance  is  seen  in  these  cases  also;  but  the 
mendelian  law  in  them  is  sometimes  obscured  and  sometimes  merely 

not  applicable. 

Cold  Spring  Harbor,  Long  Island,  N.  Y.,  April,  1908. 


PROC.  AMER.  PHIL.  SOC  XLVII.  l88  E,  PRINTED  JULY  ID,  I908. 


THE  SANTA  CRUZ  TYPOTHERIA. 

(Figures  i-io.) 

By  WILLIAM  J.  SINCLAIR. 

(Read  April  24,  1908.) 

The  Typotheria  are  a  groupof  semi-ungulate  mammals  of  strictly 
South  American  origin  appearing  first  in  the  Notostylops  beds  of 
Patagonia.*  During  the  Santa  Cruz  epoch  four  genera  are  repre- 
sented but  what  is  lacking  in  generic  and  specific  diversity  is  more 
than  ccmipensated  for  1^  an  abundance  of  individuals.  The  total 
number  of  common  species  apparently  does  not  exceed  e^ht,  but  this 


Fig.  I.  Skull  of  Protypolherium  auslrale  Ameghino,  side  view,  three 
fourths  the  natural  size.  (No.  9565  American  Museum  of  Natural  History 
collection.) 

has  been  increased  to  no  less  than  fifty-one  by  failing  to  estimate 
at  their  true  value  characters  due  to  age  and  others  which  seem  to 
be  of  the  nature  of  individual  variations  in  size,  the  result  no  doubt 
of  the  extremely  fragmentary  character  of  the  material  hitherto 
available.     Even  with  the  large  suites  of  specimens  in  the  collections 

^  Jtotyfotherium,  Efitypotherium. 

64 


X908.]  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  65 

at  Princeton  University  and  the  American  Museum  of  Natural  His- 
tory it  has  been  found  impossible  to  separate  in  a  satisfactory  manner 
the  species  of  the  genus  Protypotherium,  An  almost  exact  inter- 
gradation  in  size  without  appreciable  difference  in  structure  is 
observable  between  the  largest  sptcies  Protypotherium  australe  (Fig. 
I )  and  the  smallest  P.  attenuatum.  As  none  of  the  collections  have 
been  made  with  strict  regard  to  stratigraphic  sequence,  we  are  not 
in  a  position  to  say  whether  these  differences  represent  individual 
variations  or  true  mutations.  The  former  alternative  has  been 
adopted  in  monographing  the  group  (see  the  forthcoming  Volume 
VI.,  Part  I.  of  the  "  Reports  of  the  Princeton  University  Expeditions 
to  Patagonia"). 

Classification  of  the  Santa  Cruz  Typotheria. 

The  Typotheria  are  grouped  by  Scott*  as  a  suborder  of  the 
Toxodontia  and  may  be  defined  as  follows : 

Plantigrade  or  digitig^ade  mammals  with  pentadactyl'  or  tetra- 
dactyl  feet,  strongly  interlocking  carpus  with  os  centrale  and  serial 
or  slightly  interlocking  tarsus  with  hemispherical  astragalar  head. 
Dentition  usually  complete  but  tending  toward  reduction  of  the 
lateral  incisors,  canine  and  anterior  premolars  in  specialized  forms. 
Median  incisors  more  or  less  enlarged  and  functional  as  cropping 
teeth.  Molars  hypsodont,  lophoselenodont  in  crown  pattern,  curving 
inward  above  and  outward  below.  A  clavicle  is  present  in  some 
forms.  Femur  with  third  trochanter.  Fibula  articulating  with 
calcaneum. 

Two  well-marked  families  are  recognizable  among  the  Santa 
Cruz  representatives  of  the  suborder  for  which  the  names  Intera- 
theridae  and  Hegetotheridae  have  priority.  Each  contains  a  large 
and  a  small  genus  of  which,  in  either  case,  the  former  is  the  less 
specialized.  The  following  key  to  the  families  and  genera  may 
facilitate  the  determination  of  new  material : 

*  Scott,  W.  B.,  "The  Miocene  Ungulata  of  Patagonia,"  Rept.  British 
Asso.  Adv,  Sci.,  1904,  pp.  589-590. 

•  Ameghino  figures  a  pentadactyl  manus  in  Pachyrukhos  typicus,  "  Contrib. 
al  conocimiento  de  los  mamiferos  fosiles  de  la  Republica  Argentina,"  Ac  fas 
de  la  Academia  Nacional  de  Ciencias  en  Cdrdoba,  T.  V.,  PI.  13,  fig.  14,  1889, 
and  in  Typotherium,  ibid.,  PI.  18,  fig.  5. 


66  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  [April  m. 

Order  TOXODONTIA  Owen. 
Suborder  TYPOTHERIA  Zittel. 

A.  Family  Interatheridje.    Median  incisors  rooted;  third  and  fourth  pre- 

molars not  completely  molariform,  squamoso-mastoid  region  dilated  and 
cancellous ;  malar  long  and  narrow,  inclosed  between  temporal  process  of 
maxillary  and  squamosal ;  maxillary  orbital ;  carotid  canal  and  foramen 
lacerum  posterius  fused;  tibia  and  fibula  unfused  distally;  pes  par- 
axonic,  digits  II.  and  V.  equally  reduced  and  small,  digits  III.  and  IV. 
large  and  of  equal  length;  astragalar  trochlea  bilaterally  symmetrical; 
no  naviculo-calcaneal  facet;  calcaneum  with  large  fibular  facet 

1.  Protypotherium,     Dental   formula   },  hhi   ^  close  series.      Lateral 

incisors  unreduced;  canine  incisiform;  upper  molars  with  deep  in- 
ternal inflection  and  slight  antero-extemal  ridges;  Ms  externally 
bilobate;  temporal  bar  of  maxillary  with  slight  descending  process; 
humerus  with  internal  epicondylar  foramen;  terminal  phalanges 
laterally  compressed  hoofs  with  slight  clefts  in  manus. 

2.  Interatherium.     Dental  formula  {*  |>  }*  f ,  with  diastemata  between  the 

lateral  incisor,  canine  and  first  premolar,  varying  with  the  species. 
I'  reduced,  often  wanting;  upper  molars  with  deep  internal  inflection 
and  prominent  antero-extemal  ridges;  Ms  externally  trilobate;  tem- 
poral bar  of  maxillary  with  strong  descending  process;  humerus 
without  internal  epicondylar  foramen;  terminal  phalanges  laterally 
compressed  hoofs  with  or  without  clefts. 

B.  Family  HECETOTHERiDiE.     Median  incisors  rootless;  third  and  fourth  pre- 

molars molariform;  mastoid  dilated  inclosing  a  large  hollow  cavity; 
malar  large  excluding  maxillary  from  orbit;  carotid  canal  and  foramen 
lacerum  posterius  widely  separated;  tibia  and  fibula  firmly  fused  both 
proximally  and  distally;  pes  approaching  mesaxonic  with  digit  IIL  the 
longest,  digit  V.  greatly  reduced  and  digits  II.  and  IV.  shorter  than 
III.  but  robust;  astragalar  trochlea  bilaterally  asymmetrical;  navicular 
and  calcaneum  in  articulation;  small  fibulo-calcaneal  facet. 

1.  Hegetotherium.     Dental  formula    f,  |,  J,  f.     Second  and  third  upper 

and  third  lower  incisor  vestigial ;  canine  vestigial ;  upper  molars  inter- 
nally convex,  without  inflection  except  in  M*;  ectoloph  smooth; 
terminal  phalanges  greatly  flattened  transversely  with  prominent 
clefts. 

2,  Pachyrukhos.     Dental  formula  J,  J,  },  j.     All  the  upper  molars  inter- 

nally convex;  ectoloph  smooth;  terminal  phalanges  hoof-like  without 
clefts  in  Santa  Cruz  species. 

The  Santa  Cruz  typotheres  are  animals  of  somewhat  rodent-like 
appearance,  varying  in  size  from  a  cotton-tail  rabbit  to  a  cavy.  A 
review  of  the  more  important  skeletal  characters  of  the  g^oup  may 
be  of  value,  even  though  it  involve  some  repetitioa 


SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA. 


Fre.  2.  Skull  of  Intcratherium  robuitum  Ameghino,  side  view,  three 
fourths  the  natural  size.  <No.  9363  American  Museum  of  Natural  History 
collectioa) 

I.  The  Skull. — ^The  facial  portion  of  the  skull  is  slender  and 
more  or  less  excavated  longitudinally  while  the  brain  case  is  broad 
and  well  expanded.  The  orbits  are  central,  circular  in  outline,  quite 
prominent  in  Hegetoiherium,  Packyrukkos  and  Interatherium  and 
unenclosed  posteriorly.     The  jugal  arches  are  robust  in  all  except 


Fig,  3.    Skull  of  Hegetoiherium  mtrabile  Ameghino,  side  view,  three  fourth! 
the  natural  size.      (No.    15542  Princeton  University  collection.) 

Packyrukhos  and  moderately  expanded.  The  premaxillx  are  short 
and  heavy  with  scarcely  any  ascending  process ;  the  nasals  are  broad 
posteriorly,  tapering  forward  to  blunt  points ;  the  interorbital  tract 
plane  and  the  sagittal  and  lambdoidal  crests  low.     The  most  promi- 


i 


68  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  [Aprils. 

nent  feature  of  the  back  of  the  skull  is  the  greatly  distended  mastoid 
tract  which  may  either  be  filled  with  cancellae  or  lodge  a  large  cavity. 
In  either  case  there  is  direct  ccMnmunication  with  the  tympanic  bulla 
and  the  dilation  appears  to  have  functioned  as  a  secondary  resonator, 
perhaps  associated  with  nocturnal  habits.  The  palate  is  concave 
throughout,  terminating  posteriorly  in  a  pair  of  stout  processes. 
The  mandible  is  heavy  and  deep,  without  trace  of  suture  in  the 
firmly  fused  symphysis. 

2.  Dentition, — Beginning  with  the  normal  incisor  formula  in 
Protypotherium  (Fig.  i)  the  Santa  Cruz  typotheres  show  a  well- 
marked  tendency  toward  an  increase  in  size  of  the  median  incisors 
at  the  expense  of  the  lateral  incisors,  canine  and  anterior  premolar 
until  the  extreme  stage  of  reduction  in  Pachyrukhos  (Fig.  4)  is 
attained.  The  teeth  undergoing  elimination  are  reduced  to  simple 
cylinders.  It  is  not  to  be  understood  that  Protypotherium,  Intera^ 
therium,  Hegetotherium  and  Pachyrukhos  constitute  a  phyletic 
series  because  they  represent  successive  stages  in  the  process  of 
dental  reduction  associated  with  the  hypertrophy  of  the  median  in- 
cisors. As  already  indicated  in  the  key  to  the  genera,  two  divergent 
lines  are  represented  and  not  a  single  progressive  series.  A  rather 
curious  feature  of  the  lower  incisors  in  Protypotherium  is  the  pres- 
ence in  the  first  and  second  of  a  deep  median  cleft  producing  a 
fork-like  structure  recalling  a  somewhat  similar  division  of  the  lower 
incisor  crowns  in  the  Hyracoidea.  In  all  the  Santa  Cruz  typotheres 
the  enamel  layer  on  the  enlarged  incisors  tends  to  be  confined  to  the 
anterior  surface  of  the  crown.  The  molars  in  all  the  genera  are 
constructed  on  much  the  same  plan  but  only  in  Protypotherium  are 
absolutely  unworn  teeth  known,  consisting  essentially  of  a  broadly 
concave  ectoloph  (e,  Fig.  S,  A)  and  a  pair  of  crescents  with  the  con- 
vexity directed  inward  (ac,  pc,  Fig.  8,  A),  of  which  the  anterior 
horns  are  fused  with  the  ectolc^h  inclosing  a  reentrant.  A  crista- 
like  ridge  from  the  ectoloph  (c,  Fig.  8,  ^)  is  separated  frcwn  the 
anterior  crescent  by  a  deep  notch.  A  slight  ridge  (pp,  Fig.  S,  A) 
blocks  the  shallow  valley  inclosed  by  the  posterior  crescent  As  the 
tooth  wears  the  antero-extemal  angle  of  the  crown  elongates  and  is 
channeled  by  a  shallow  groove  producing  the  ridges  noted  in  the 
key  to  the  genera. 


,,oi,j  SINCLAIR— THE   SANTA   CRUZ  TYPOTHERIA.  69 

In  the  lower  molars  the  convexity  of  the  crescents  is  reversed  so 
that  the  reentrant  fold  is  external  (Figs,  i,  3,  9,  A).  A  prominent 
lobe  spanning  the  arc  of  the  posterior  crescent  {pp,  Fig.  g,A)  is  not 
peculiar  to  the  teeth  of  the  Typotheria  alone,  but  is  present  also  in 
Nesodon  (Fig.  9,  S),  Astrapotherium,  Tkeosodon  and  other  extinct 
ungulates  from  South  America.  In  the  last  lower  molar  the  devel- 
opment of  the  third  lobe  present  in  Interatherium  is  accomplished 
by  the  deepening  of  the  shallow  groove  indicated  in  Protypolherium 
at  the  point  marked  pc  in  Fig.  9,  A. 

As  mentioned  in  the  generic  key  the  premolars  are  sometimes 
molariform  and  sometimes  not,  differing  from  the  molars  in  the 
latter  case  in  having  the  anterior  crescentic  lobe  smaller  than  the 
posterior. 

Roots  are  developed  only  in  the  deciduous  molars  but  as  these 
have  been  observed  only  in  Protypotherium  and  Interatherium  it  is 


Fig.  4.    Skull  of  Pachynikhos  moyant  Ameghino,  side  view,  three  fourths  the 
natura.1  size.      (Reconstructed  from  several  specimens.) 

not  altogether  certain  whether  this  character  is  of  family  or  sub- 
ordinal  value.  So  far  as  can  be  ascertained  the  crown  pattern  seems 
to  have  been  the  same  in  the  deciduous  and  pertnanent  series,  the 
milk  teeth  resembling  their  successors.  The  order  of  replacement 
seems  to  have  been  the  normal  one. 

A  thin  layer  of  cement  is  usually  observable  on  the  molars  and 
premolars  of  all  the  genera. 

3,  Axial  Skeleton. — ^The  dorso-lumbar  vertebral  formula  in 
Interatherium  is  twenty-two,  of  which  fifteen  are  dorsals.  It  was 
probably  the  same  in  Protypotherium  but  in  Pachyrukhos  eight  lum- 
bars  are  present  Five  vertebrae  are  coossified,  in  the  sacral  com- 
plex of  which  three  are  true  sacrals  in  contact  with  the  ilium  and 


70 


SINCLAIR— THE  SANTA  CRUZ  TVPOTHERIA. 


[April  *4. 


two  belong  to  the  caudal  series.    The  length  of  the  tail  seems  to 
have  varied.     In  Protypotherium  and  Interatherium  it  is  both  Icmg 
and  heavy  while  in  Pachyrukhos  there  is 
reason  to  believe  that  it  was  quite  short. 
4,  Foot  Structure. — Almost  nothii^ 
has  hitherto  been  known  of  the  struc- 
ture of  the  feet  in  the  Santa  Cruz  typo- 
theres,  but  definite  information  is  now 
available  for  all  the  genera  except  He- 
getotheriutn,  in  which  the  manus  is  still 
unknown,  but  from  the  close  structural 
resemblance  of  Hegetotheriutn  and  Pa- 
chryukhos  it  is  probable  that  it  was  not 
unlike  that  of  the  latter,  which  in  turn 
,'  does  not  differ  materially  from  the  manus 

'f..  of    Interatherium    and    Protypotherium 

^  (Fig-  61  '4)  ■     In  the  Santa  Cruz  forms 

jv  both  manus  and  pes  are  tetradactyl  with- 

"  out  the  slightest  trace  of  an  opposable 

thumb  or  great  toe.*  The  carpus  is 
strongly  interlocking  and  shows  no  trace 
of  the  centrale.  Two  types  of  hind  foot 
^  are  developed  (Figs.  5  and  7,  A)  simu- 

lating the  paraxonic  and  mesaxonic  sym- 
metry of  the  feet  of  the  Artiodactyla 
and  Perissodactyla.  These  are  prolably 
to  be  correlated  in  the  Typotheria  with 
cursorial  and  saltatorial  modes  of  pro- 
gression. Pachyrukhos  was  certainly  a 
jumping  animal  as  shown  by  the  greater 
length  and  strength  of  the  hind  limbs  and  inner  digits  of  the  pes. 
In  fact,  the  structure  of  both  the  fore  and  hind  limbs  in  this  animal 

*  A  pentadactyl  manus  with  separate  centrale  in  the  carpus  and  opposable 
thumb  and  a  pentadactyl  pes  with  large  opposable  hallux  figured  by  Ameghino, 
Revista  Argfnlina  de  Hist.  Nat.,  I.,  pp.  393,  394.  ^S^-  95.  96  and  referred  to 
Interaiherium  (leochilus)  robHslnm  do  not  pertain  to  this  genus.  The  same 
figures  with  the  erroneous  determination  appear  also  in  Zittel's  "Handbuch 
der  Falaeontologie,"  IV.,  p.  493,  iig.  407. 


Fig.  s  Left  huid  foot 
of  Protypotherium  austraU 
Ameghino,  three  fourths 
the  natural  size.  (No. 
9149  American  Museum 
of  Natural  History  col- 
lection.) 


,,,(.]  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  71 

closely  resembles  that  of  the  rabbit.  From  the  numerous  structural 
similarities  between  Pachyrukhos  and  Hegetherium  it  may  be  in- 
ferred that  the  latter  was  also  saltatorial.  Its  broad,  shallow  as- 
tragalar  trochlea  is  in  contrast  with  the  narrow,  more  deeply  incised 
trochlea  of  the  cursorial  Protypotherium  and  Interatherium.  Both 
of  these  genera  have  the  fore  and  hind  limbs  of  approximately  equal 
length.  The  terminal  phalanges  in  the  Santa  Cruz  typotheres  are 
hoof-like  and  in  Hegetotherium  have  prominent  median  clefts. 

Relationships  of  the  Santa  Cruz  Typotheria. 
I.  With  the  Toxodonta. — In  the  evolution  of  the  teeth  and  feet, 
the  Santa  Cruz  Typotheria  are  less  advanced  than'  thetr  contempor- 
aries, the  Nesodons.     The  feet  of  Nesodon  (Figs.  6,  S,  7,  B)  are 


Fig.  6.  A.  Left  fore  foot  of  Protypotherium  austraU  Ameghino,  three 
fourths  the  tiatural  size.  (No.  9149  American  Museum  of  Natural  History 
collection.)  B.  Left  fore  foot  of  Nesodon  imbricatus  Owen,  about  one  fifth 
the  natural  size.      (No.  15460  Princeton  University  collection.) 

tridactyle  with  the  axis  passing  through  the  third  digit.  The  manus 
has  originally  been  tetradactyl  like  that  of  Protypotherium  (Fig. 
6,  A)  but  has  lost  almost  all  trace  of  the  fifth  digit,  a  mere  vestige, 
not  shown  in  the  figure,  remaining.  The  other  bones  of  the  wrist 
and  foot  have  not  suffered  any  displacement  as  a  result  of  this  loss 
but  interlock  in  the  same  way  as  in  Protypotherium.    The  hind  foot 


72 


SINCLAIR— THE   SANTA  CRUZ  TVPOTHERIA. 


(April** 


of  Nesodon  (Fig.  7,  B)  is  the  realization  of  a  structure  already  fore- 
shadowed in  the  pes  of  Hegelotkerium  (Fig.  7,  A).  The  fifth  digit, 
which  is  greatly  reduced  in  Hegetotherium,  has  here  disappeared 
and  the  ento-  and  tneso-cuneiforms  have  united  to  a  single  bone. 
The  shortening  of  the  neck  of  the  astragalus  and  the  increase  in 
size  of  the  fibular  facet  on  the  catcaneum  are,  perhaps,  adaptattcms 
to  the  support  of  weight.    Although  the  molars  of  Nesodon  i^pear 


m 

Fig.  7.  A.  Hegelotkerium  mirabile  Ameghino,  right  hind  foot,  three 
fourths  the  natural  size.  (No.  15542  Princeton  University  collection.)  B. 
Netodon  imbricatHS  Owen,  right  hind  foot,  about  one  third  the  natural  size. 

(No.  15460  Princeton  University  collection.) 


exceedingly  complex,  owing  to  the  development  of  secondary  enamel 
folds,  the  primary  elements  can  be  homologised  with  those  displayed 
in  the  simpler  crown  pattern  of  Frotypotherium-,  as  indicated  by  the 
similar  lettering  in  Figs.  8  and  9.     This  comparison  can  not  yet  be 


.90I.]  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA,  73 

extended  to  the  other  Santa  Cruz  genera,  Hegetotherium,  Packy- 
rukhos  and  Interatherium,  as  unworn  molars  of  these  are  not  avail- 
able. Nesodon  differs  from  the  Typotheria  in  the  enlargement  and 
caniniform  character  of  the  second  incisor  above  and  the  third  below, 
while  in  the  Typotheria  the  median  incisor  in  both  jaws  is  the  only 
one  tending  toward  great  increase  in  size.  In  none  of  the  Santa 
Cruz  Typotheria  is  there  a  trace  of  the  double  deciduous  dentition 
characteristic  of  Nesodon. 


Fig.  8.  A.  Unworn  third  upper  molar  of  a  young  Prolypotherium,  four 
and  one  half  times  the  natural  size.  (No,  9482  American  Museum  of  Natural 
History  collection.)  B.  Nesodon  imbricatus  Owen,  second  and  third  upper 
molars  slightly  worn,  three  fourths  the  natural  size.  (No.  15135  Princeton 
University  collection.)  ae,  antero-intemal  crescent;  pc.  postero -internal 
crescent;  e,  ectoloph;  c,  crista;  pp,  poUerior  pillar. 

From  these  resemblances  in  dentition  and  foot  structure  it  seems 
permissible  to  infer  that  the  Toxodonta  and  Typotheria  had  a  com- 
mon origin,  but  the  facts  at  present  available  do  not  justify  us  in 
saying  more. 

2.  With  Typotherium. — Difficult  as  it  is  to  ascertain  the  relation- 
ship existing  between  the  Santa  Cruz  Typotheria  and  the  Nesodons, 


74 


SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA. 


[April  «4, 


it  is  even  more  so  to  determine  their  degree  of  kinship  with  Typo- 
therium.  From  their  small  size  it  seems  quite  probable  that  none  of 
the  Santa  Cruz  Typotheria  are  in  the  direct  line  of  descent  culmi- 
nating in  this  genus.  This  is  confirmed  by  the  degree  of  specializa- 
tion in  dentition  and  foot  structure  which  Typotherium  displays. 
The  teeth  of  the  latter  show  a  greater  complexity  of  folding  than  is 
attained  by  any  of  the  Santa  Cruz  typotheres,  while  the  feet  are  less 
specialized  with  a  poUex  in  the  manus  which  has  been  lost  in  Pro- 
typotherium,  the  most  generalized  of  the  Santa  Cruz  typotheres 


B 


ac 


Fig.  9.  A.  Unworn  third  lower  molar  of  a  young  Pro  typo  therium,  four 
and  one  half  times  the  natural  size.  (No.  9482  American  Museum  of  Natural 
History  collection.)  B.  Nesodon  imbricatus  Owen,  two  lower  molars,  three 
fourths  the  natural  size.  (No.  15135  Princeton  University  collection.)  ac, 
anterior  crescent;  pc,  posterior  crescent;  pp,  posterior  pillar. 


(Fig.  6,  A),  and  with  digit  V.  of  the  pes  less  reduced  than  in  the 
most  specialized  of  the  latter  (Pachyrukhos) ,  A  poUex  has  been 
figured  by  Ameghino°  in  the  manus  of  Pachyrukhos  typicus,  but 
none  has  been  found  in  any  Santa  Cruz  specimen.  The  manus  in 
Hegetotherium  is  unknown,  so  the  above  statement  regarding  the 

■  Ameghino,  Florentino,  "  Contrib.  al  conoc,  etc.,"  PI.  13,  fig.  14. 


,9o8.]  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  75 

degree  of  specialization  in  foot  structure  displayed  by  Typotherium 
may  require  some  modification  in  the  light  of  fuller  knowledge. 

3.  With  the  Rodents, — In  many  features  of  skull  and  skeleton 
the  Typotheria  resemble  the  rodents.  This  is  most  apparent  in 
Pachyrukhos,  which  seems  to  have  been  a  saltatorial  animal,  but  in 
none  of  the  Typotheria  are  the  following  characters  peculiar  to 
rodents  developed: 

A.  Persistently  growing,  chisel-shaped  incisors  (I.  %  of  the  per- 
manent series,  Weber).*  I.  Yi  of  the  permanent  series  is  enlarged 
in  some  of  the  Typotheria  and  may  g^ow  persistently  but  is  modified 
for  cropping  and  not  for  gnawing. 

B.  More  or  less  antero-posterior  elongation  of  the  mandibular 
condyle  and  corresponding  modification  of  the  glenoid  fossa  to 
permit  backward  and  forward  movement  of  the  lower  jaw.  In  the 
Typotheria  the  condyle  is  approximately  circular  in  outline  with  the 
glenoid  surface  flattened  and  the  movement  of  the  mandible  is  from 
sidp  to  side. 

C.  Frequent  outward  curvature  of  the  crowns  of  the  upper 
molars  and  inward  curvature  of  those  of  the  inferior  series  in  hypso- 
dont  forms.    The  reverse  is  true  in  the  Typotheria. 

D.  Contact  of  ascending  process  of  premaxillary  with  frontal. 
This  process  is  short  and  robust  in  the  Typotheria  and  is  widely 
separated  from  the  frontal  by  the  maxillary. 

E.  Elongation  of  the  mandibular  angle.  The  angle  is  evenly 
convex  in  the  Typotheria. 

F.  The  astragalus  in  rodents  is  characterized  by  a  broad,  short, 
rather  shallow  trochlea  with  the  crests  sharp  and  equally  developed, 
distinct  neck  and  flattened  head,  convex  distally;  trochlea  s)rmmet- 

rical  to  the  vertical  plane ;  fibular  and  internal  malleolar  facets  ver- 

* 

tical ;  body  limited  posteriorly ;  no  astragalar  foramen.  In  the  Santa 
Cruz  Typotheria  the  body  is  deeper  than  in  rodents,  the  crests  may 
or  may  not  be  equally  developed  and  the  head  is  globular  without 
antero-posterior  flattening.  The  symmetry  of  the  trochlea  with 
respect  to  the  vertical  plane  varies  in  the  different  families.  In  the 
other  characters  they  resemble  rodents. 

• "  Die  Saugetiere,"  p.  480,  1904. 


76  SINCLAIR— THE   SANTA  CRUZ  TYPOTHERIA.  [April  .4. 

G.  The  presence  of  a  free  centrale  in  the  carpus  in  all  rodents 
except  the  Hystricidie  and  Caelogenys  and  the  general  fusion  of  the 
scaphoid  and  lunar  in  all  except  the  Bathyergidje,  Ctenodoctilidae  and 
Lagomorpha.^  The  centrale  is  wanting  in  the  carpus  of  the  Typo- 
theria  and  the  lunar  is  always  free. 

H.  The  presence  of  a  tibial  sesamoid  in  all  the  simplicidentate 
rodents.     This  is  not  found  in  the  tarsus  of  the  Typotheria. 

The  Typotheria  resemble  rodents  in  the  elongation  of  the  anterior 
portion  of  the  skull  with  the  reduction  of  the  incisor-canine-premolar 
series  {cf.  Figs.  3  and  4),  in  the  enlargement  and  often  permanent 
growth  of  the  median  incisors  (not  homolc^ous  with  the  enlarged 
incisors  in  rodents,  see  under  A,  above),  in  the  development  of  a 
mastoid  dilation  which  may  be  filled  with  cancellae  ( Interatheridse) 
as  in  many  rodents  and  connected  with  the 
auditory  bulla,  in  the  shape  of  the  proximal 
articular  surfaces  between  the  radius  and  ulna, 
in   the   broad   anteriorly   directed   transverse 
■  processes    of    the    lumbar   vertebras   and    in 

several  other  characters  of  m^inor  importance. 
In  view  of  the  striking  differences  in  struc- 
ture indicated  in  the  preceding  paragraphs, 
it  seems  probable  that  these  resemblances  are 
to  be  explained  as  instances  of  convergence. 
'  4.  With  the  Hyracoidea. — A  more  or  less 

^  intimate  relationship  between  the  Typotheria 

Fic  10.  Left  hind  jjjjj  Hyracoidea  is  commonly  assumed  but 
(Dendrohwax)  arbo-  ^'*'^  ^^  complete  material  now  available  it  is 
rea,  ^  natural  size,  difficult  to  see  on  what  grounds  this  hypothesis 
(No.  365  Princeton  (.3^  be  maintained.  The  hyracoid  carpus  is 
raHoEion."™''*'  arranged  on  the  linear  plan  with  separate 
centrale  while  in  the  tarsus  the  astralagus  is 
unlike  that  of  any  other  mammal  in  possessing  a  large  step-like 
articulation  for  the  internal  tibial  malleolus  (iMg.  10). 

In  striking  contrast  with  hyrax,  the  carpus  in  the  Typotheria  is 
strongly  interlocking  without  centrale,  and  the  internal  tibial  mal- 
leolus is  applied  to  the  lateral  surface  of  the  astragalus  without  trace 
'  Weber,  loc.  cit.,  p.  476. 


1908.J  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  77 

of  the  supporting  shelf  (Figs.  5  and  7,  A),  The  flat  astragalar 
head  in  the  Hyracoidea  and  the  articulation  of  the  fibula  with  the 
astragalus  instead  of  with  the  calcaneum  are  additional  points  of 
difference,  all  of  which  are  more  than  sufiicient  to  offset  similarities 
in  skull  structure  which  are  confined  to  a  few  points,  such  as  the 
cancellous  dilation  of  the  mastoid,  the  shape  of  the  posterior  border 
of  the  palate,  and  the  increase  in  depth  posteriorly  of  the  mandible. 
In  the  Hyracoidea  the  molar  takes  part  in  forming  the  outer  por- 
tion of  the  glenoid  cavity,  the  parietal  enters  into  the  postorbital 
process  and  the  base  of  the  coronoid  just  back  of  the  last  lower 
molar  is  perforated  by  a  large  foramen,  a  superior  branch  of  the 
alveolar  canal.  None  of  these  characters  are  exhibited  by  the  Typo- 
theria.  In  the  hyracoid  dentition,  the  first  upper  incisor  is  a  per- 
sistently growing  downwardly  curved  tusk  of  triangular  cross- 
section.  In  some /of  the  Typotheria  this  tooth  may  grow  persis- 
tently but  it  is  always  antero-posteriorly  compressed,  transversely 
expanded  and  modified  for  cropping,  never  appearing  as  a  tusk. 
The  molars  of  the  Hyracoidea  are  lophoselenodont  and  either 
brachyodont  or  short  hypsodont  while  in  the  Typotheria  they  are 
extremely  hypsodont,  develq)ing  roots  only  in  the  deciduous  series. 
The  crown  pattern  of  the  hyracoidean  molar  bears  more  resemblance 
to  that  of  some  of  the  early  horses  and  rhinoceroses  than  to  the 
molar  pattern  in  the  least  specialized  of  the  typotheres  (Figs.  8 
and  9). 

The  so-called  hyracoids  from  the  Fayum  Province  of  Egypt 
(Saghatherium,  Megalohyrax)  are  as  yet  known  only  from  frag- 
ments of  the  skull  and  dentition  but,  so  far  as  the  available  material 
permits  comparison,  resemble  the  modem  Hyracoidea  and  not  the 
Typotheria  which  would  probably  not  be  the  case  if  the  two  orders 
were  related  as  it  would  naturally  be  expected  that  a  closer  simi- 
larity should  exist  between  the  Eocene  and  Miocene  representatives 
of  an  order  than  between  the  latter  and  the  recent  forms.  All  the 
Egyptian  hyracoids  have  the  base  of  the  coronoid  perforated  by  a 
branch  of  the  dental  canal  as  in  the  recent  forms*  and  unlike  the 
Typotheria. 

•Communicated  by  Mr.  Walter  Granger,  of  the  American  Museum  of 
Natural  History. 


78  SINCLAIR— THE  SANTA  CRUZ  TYPOTHERIA.  [April  m. 

Various  pre-Santa  Cruz  genera  {ArchcBohyrax,  Argyrohyrax) 
have  been  referred  to  the  Hyracoidea.  Their  foot  structure  is  still 
unknown  but  the  skull  and  dentition,  to  judge  from  the  photographs, 
figures  and  descriptions  examined  by  the  writer,  are  not  hyracoidean 
in  character.  Too  little  is  known  of  these  forms  to  warrant  a  dis- 
cussion of  their  relationship  with  the  Santa  Cruz  Typotheria,  but 
there  can  be  little  doubt  that  they  should  be  referred  to  the  same 
suborder. 

Princeton  University,  April,  1908. 


NOTES  ON  SOME  CHILEAN  COPPER  MINERALS. 

By  harry  F.  KELLER. 

(Read  April  24,  1908.) 

Some  time  ago  my  brother,  Mr.  Hermann  A.  Keller,  presented 
me  with  a  fine  suite  of  mineral  specimens  collected  by  him  on  a  pro- 
fessional trip  to  Chilean  mining  localities.  The  minerals,  which  in- 
clude native  sulphur  and  copper,  various  oxides,  chlorides,  sulphates* 
borates  and  silicates,  were  for  the  most  part  readily  identified  by 
their  characteristic  appearance  or  by  simple  tests,  but  some  of  them 
aroused  my  curiosity,  partly  because  of  their  rare  occurrence,  and 
partly  on  account  of  their  beauty  or  exceptional  purity.  I  was  thus 
led  to  make  a  number  of  qualitative  and  quantitative  analyses,  the 
results  of  which  appear  to  me  sufficiently  interesting  to  be  placed  on 
record.  In  the  present  paper  I  shall  confine  myself  to  the  descrip- 
tion of  some  minerals  containing  copper  as  either  a  principal  or 
a  minor  constituent. 

Cupreous  Manganese. 

It  is  well  known  that  in  many  varieties  of  psilomelane  or  wad  the 
manganous  oxide  is  partially  replaced  by  oxide  of  copper,  and  that 
special  names  have  been  given  to  some  of  those  varieties  in  which 
the  proportion  of  the  latter  oxide  is  considerable.  Among  them  is 
the  peloconite  from  Remolinos,  Chile,  which  was  first  described  by 
Richter,^  and  chemically  characterized  by  Kersten.*  Its  quantitative 
composition,  however,  does  not  appear  to  have  been  fully  deter- 
mined. The  material  supplied  by  my  brother  included  several  very 
fine  specimens  of  a  cupreous  manganese  from  Huiquintipa,  Province 
of  Tarapaca,  and  these  are  unquestionably  identical  with  Richter's 
peloconite.  With  the  one  exception  of  the  specific  gravity,  the 
physical  and  chemical  characters  of  the  new  material  are  precisely 

^  Poggendorffs  Annalen,  21,  590. 
'  Sckweigger's  Journal,  66,  7. 

PROC.  AMER.  PHIL.  SOC.  XLVII.   1 88  F,  PRINTED  JULY  lO,  I908. 


80 


KELLER— NOTES  ON  CHILEAN  COPPER  MINERALS.     [April  •4. 


similar  to  those  of  the  Remolinos  occurrence.  The  mineral  is  mas- 
sive and  amorphous,  has  a  conchoidal  fracture,  a  bluish-black  color 
and  a  liver-brown  streak.  Its  hardness  is  between  3  and  4,  and 
the  specific  gravity  3.683  (instead  of  2.5-2.6).  When  brcJcen  into 
small  pieces  and  carefully  picked  with  the  aid  of  a  lens,  the  material 
appeared  quite  homogeneous  except  for  a  few  particles  of  quartz 
and  scMne  green  or  bluish  specks  of  a  copper  compound  on  the  out- 
side and  along  the  crevices.  A  qualitative  analysis  showed  that  it 
contains  the  oxides  of  manganese,  co|^er  and  iron,  together  with 
water  and  varying  amounts  of  admixed  silica.  To  ascertain  whether 
the  mineral  has  a  definite  chemical  composition  analyses  were  made 
of  carefully  selected  samples  from  different  specimens.  It  was 
found  that  the  silica,  which  separates  on  dissolving  the  substance 
in  hydrochloric  acid,  is  not  uniformly  distributed  through  the  mass. 
Its  percentage  varied  from  12  per  cent,  to  32  per  cent,  and  its 
microscopic  examination  showed  that  it  consists  entirely  of  quartz. 
There  could  be  no  douot,  then,  that  it  is  simply  an  admixture,  and 
that  in  calculating  the  composition,  the  silica  (of  which  only  a  trace 
dissolves  with  the  mineral)  should  first  be  deducted  from  the 
amount  of  the  substance  taken.  The  results  of  the  analyses  were 
as  follows: 


I. 

n. 

in. 

IV. 

Ozyireo 

14.37% 
69.61 

5.86 

.48 

.36 
2.05 
1.9a 

5.14 

14.18% 

68.95 
6.05 

.56 

.47 
1.94 
1.91 
5.29 

99.35% 

13.89% 
69.44 
5.69 

not  det 

not  det 
1.89     \ 
2.10     j 

not  det. 

Jo        ........................... 

Maoganous  oxide 

70.61  % 

Cupric  oxide 

6.48 

CocMklt  oxide , 

not  det 

Bariam  oxide 

not  det 

Ferric  oxide 

Alumina , 

4.22 

Water 

99.79% 

It  is  seen  from  these  figures  that  the  proportions  of  the  several 
constituents  of  the  mineral  are  fairly  constant.  The  composition  is 
that  of  psilomelane,  in  which  part  of  the  manganese  is  replaced  by 
cc^per.  It  is  difficult  to  account  for  the  very  constant  prq)ortions 
of  oxide  of  iron  and  alumina. 

Regarding  the  determinations  of  water  and  of  available  oxygen, 
I  may  mention  that  the  former  was  made  by  heating  the  substance 


,9oS.)  KELLER— NOTES  ON  CHILEAN  COPPER  MINERALS.  81 

in  a  current  of  dry  air  and  collecting  the  moisture  in  calcium  chloride, 
while  the  latter  was  estimated  iodometrically  in  I.  and  II.,  and  indi- 
rectly in  III.,  by  heating  a  weighed  portion,  first  in  air  and  then  in 
hydrogen,  and  allowing  for  the  water  and  the  reduction  of  the 
oxides  of  copper  and  iron.  As  a  matter  of  course  this  method  is 
less  reliable,  but  the  result  nevertheless  agrees  quite  well  with  the 
iodometric  determinations. 

Chalcanthite  and  a  Double  Sulphate  of  Copper  and 

Magnesium. 

Among  the  specimens  that  claimed  my  special  attention  there 
was  one'  consisting  of  irregular  and  rounded  masses,  and  which  was 
labeled  "sulphate  of  copper  and  aluminium."  While  the  shape  of 
the  little  lumps  was  about  the*  same,  three  distinct  kinds  of  material 
could  readily  be  picked  from  the  specimen,  even  without  the  help 
of  a  magnifying  glass. 

One  of  these  substances  had  a  deep  blue  color  and  was  recog- 
nized without  difficulty  as  chalcanthite.  The  blue  masses  were  evi- 
dently crystals  which  were  strongly  corroded  and  slightly  effloresced 
on  the  surface.  A  quantitative  analysis  confirmed  the  composition 
CuSO^  +  sHjO,  with  very  small  amounts  of  iron  and  magnesium 
sulphates,  and  a  slight  admixture  of  silicious  matter.     It  gave : 

Fonnd.  Calculated. 

Per  Cent.  Per  Cent. 

Sulphur  trioxide 32.21  32.1 

Cupric  oxide  31.52  31.8 

Ferrous  oxide 30 

Magnesium  oxide 35 

Water    35.79  36.1 

100.19  loo.o 

More  interesting  were  the  bluish-white  masses  which  formed  the 
larger  portion  of  the  specimen.  They  were  earthy  and  friable,  but 
presented  shapes  and  surfaces  exactly  similar  to  those  of  the  chal- 
canthite, suggesting  a  pseudomorph  after  the  latter.  In  composi- 
tion, however,  the  material  was  found  to  differ  from  chalcanthite  in 

•From  Copaquire,  Province  of  Tarapacd. 


82 


KELLER— NOTES  ON  CHILEAN  COPPER   MINERALS.     [April  04, 


that  it  contained  a  large  proportion  of  magnesium  sulphate.    Anal- 
yses of  two  different  samples  yielded: 


Found. 

CalcuUted  for 

I. 

11. 

(Cu,  Mg)SO«+5H.O : 

Sulphur  trioxkle 

35-74% 
12.41 

11.42 

.97 
.23 

trace 
38.45 

35.67% 
12.46 
11.36 
1.05 

.41 
.06 

38.31 

35.84% 

Cupric  oxide 

11.89 

Magnesium  oxide 

11.95 

Ferrous  oxide , 

^J 

Manganous  oxide.. 

Nickel  oxide. 

Water.... 

40.32 

99.22% 

9932% 

100.00% 

The  conclusions  to  be  drawn  from  these  results  are,  first,  that 
the  mineral  is  an  isomorphous  mixture  of  the  sulphates  of  copper 
and  magnesium;  secondly,  that  this  double  salt  contains  five  mole- 
cules of  water  of  crystallization ;  and  thirdly,  that  for  each  molecule 
of  copper  sulphate  there  are  present  (very  nearly)  two  molecules  of 
magnesium  sulphate.  The  shortage  in  the  water  content  is  doubt- 
less owing  to  efflorescence,  and  there  should  be  credited  to  the  mag- 
nesia content  an  amount  equivalent  to  the  percentages  of  the  oxides 
of  iron  and  manganese. 

Under  the  name  of  cupromagnesite  a  double  sulphate  of  copper 
and  magnesium  has  been  described  by  Scacchi.  It  occurs  in  the 
form  of  green  crusts  on  lava  from  the  Vesuvius,  and  is  believed  to 
be  isomorphous  with  melanterite,  containing,  like  the  latter,  seven 
molecules  of  water  of  crystallization.  I  have  seen  no  reference  to 
a  mineral  of  the  same  composition  as  that  above  described. 

Associated  with  the  chalcanthite  and  the  double  sulphate  of  cop- 
per and  magnesium  were  other  little  masses,  dirty-white  in  color  and 
more  or  less  stained  with  ferric  oxide.  They  were  very  hard  and 
consisted  almost  entirely  of  silica,  containing  only  trifling  amounts 
of  oxide  of  iron  and  magnesia.  It  is  puzzling  to  explain  why  these 
masses  should  simulate  the  form  of  the  accompanying  soluble 
sulphates. 

Brochantite(  ?)  Containing  Arsenic  Acid. 

Very  small  quantities  only  were  available  of  an  emerald  green 
mineral  which  was  observed  partly  in  fine  acicular  crystals  dissemi- 


,908. 1         KELLER— NOTES  ON  CHILEAN   COPPER  MINERALS  83 

nated  through  a  silicious  rock,  and  partly  as  an  incrustation  upon 
quartz.  On  account  of  its  physical  characters,  as  well  as  the  strong 
reaction  its  solution  gave  with  barium  chloride,  I  was  first  inclined 
to  regard  it  as  a  typical  brochantite.  This  impression  was  confirmed 
by  rough  estimations  of  the  copper  and  sulphur  trioxide,  but  as  these 
tests  had  been  made  on  impure  material,  I  decided  to  attempt  the 
analysis  of  a  carefully  prepared  sample.  To  obtain  about  .5  grm. 
of  the  substance,  I  found  it  necessary  to  sacrifice  the  best  specimens 
in  my  possession,  and  my  patience  was  put  to  a  severe  test  in  picking 
the  minute  crystals  under  the  lens.  They  were  sorted  over  and  over 
until  the  microscope  showed  only  a  few  remaining  specks  of  quartz 
adhering  to  the  larger  crystals  of  the  coiner  mineral. 

■ 

The  quartzy  material  from  which  this  sample  had  been  picked 
still  contained  considerable  quantities  of  the  copper  mineral,  and  it 
occurred  to  me  that  it  might  serve  for  a  qualitative,  and,  perhaps, 
a  preliminary  quantitative  analysis.  Accordingly  the  material  was 
extracted  with  hydrochloric  acid,  and  the  resulting  green  solutio^i 
divided  into  equal  parts.  When  the  copper  had  been  precipitated 
with  hydrogen  sulphide,  it  was  noticed  that  yellow  flakes  began  to 
form,  and  after  the  liquid  saturated  with  the  gas  had  been  allowed 
to  stand  in  a  warm  place  over  night,  a  considerable  amount  of  the 
yellow  precipitate  had  settled  on  the  black  copper  sulphide.  There 
could  be  no  doubt,  then,  that  arsenic  was  present  in  the  form  of 
arsenic  acid.  The  sulphides  were  separated  and  worked  up  in  the 
usual  manner,  and  the  filtrate  was  searched  for  other  metallic  ions. 
It  yielded  only  traces  of  iron  oxide  and  altmiina.  The  other  half 
of  the  original  solution  was  used  for  the  determination  of  the 
sulphur  trioxide  and  the  arsenic  acid.  The  results  of  the  determi- 
nations, calculated  for  the  entire  amount  of  copper  mineral  dissolved, 

gave: 

Copper   f 5068  grms. 

Sulphur  trioxide    0783  grms. 

Arsenic  anhydride  1309  grms. 

The  question  now  arose  as  to  whether  the  very  large  proportion 
of  arsenic  acid  found  really  constituted  an  integral  part  of  the  sup- 
posed brochantite,  or  whether  it  did  not  belong  to  another  mineral 
contained  in  the  rock.  The  test  previously  made  seemed  to  preclude 
such  an  arsenic  content  of  the  mineral  under  examination. 


84  KELLER— NOTES  ON  CHILEAN  COPPER  MINERALS.     (April  34, 

In  view  of  the  very  limited  amount  of  material  available  for 

« 

analysis,  it  seemed  best  to  dispense  with  the  water  determinaticm 
land  confine  the  characterization  of  the  mineral  to  ascertaining  the 
specific  gravity  and  an  estimation  of  the  base  and  the  acids.  The 
following  results  were  obtained : 

Specific  gravity 3.160 

I.  II. 

Per  Cent.  Per  Cent. 

Sulphur  trioxide   16.32  16.63 

Arsenic  anhydride  2.31  2.40 

Copper  oxide 68.90  68.68 

Silica   1.63  1.18 

If  we  deduct  the  quartz  from  the  substance  takeft  for  analysis, 
the  percentages  of  sulphur  trioxide  and  oxide  of  copper  will  be 
found  to  approach  very  nearly  to  those  in  brochantite  which  contains : 

Per  Cent. 

,  Sulphur  trioxide  17.7 

Cupric  oxide 70.3 

Water   12.0 

and  it  is  difficult  to  explain  the  role  of  the  arsenic  acid  which  is 
equivalent  to  about  4.4  per  cent,  of  sulphur  trioxide.  To  establish 
a  definite  formula  for  the  compound  it  would  be  necessary  to  ascer- 
tain by  further  analyses  whether  or  not  the  proportion  of  arsenic 
anhydride  is  constant,  and  to  complete  the  analysis  by  an  exact  deter- 
mination of  the  water.  The  specimens  in  my  possession,  unfor- 
tunately, are  not  sufficient  for  this  purpose. 

The  specimens  were  collected  at  Copaquire,  Province  of  Tarapaca. 

Atacamite. 

In  conclusion  I  desire  to  call  atl^ntion  to  some  magnificent  speci- 
mens of  atacamite  from  Paposo  in  the  Province  of  Antofagasta. 
They  do  not  show  the  usual  slender  prisms,  but  consist  of  aggre- 
gates of  fairly  large  crystals,  closely  resembling  the  octahedron  of 
the  isometric  system  modified  by  the  cube  and  the  rhombic  dodeca- 
hedron. On  closer  examination,  however,  it  would  seem  that  they 
are  really  combinations  such  as  have  been  observed  on  the  atacamite 


igot.]         KELLER— NOTES  ON   CHILEAN   COPPER   MINERALS. 


Fig.  I.  Atacamjte  from  Paposo,  Chile. 

from  certain  localities  in  South  Australia.*  As  was  to  be  expected, 
the  analysis  showed  the  specimens  to  be  an  atacamite  of  unusual 
purity  and  of  normal  composition.    It  yielded : 

Sp.gr 3738 


FoBBd.       CDa,.]Ca(OH),. 

Ps  Cau.  P«  Col. 

Gilorine    16.51  16.6 

Copper  14-79  I4'9 

Cupric  oxide  SS-aS  sS-8 

Water    12.43  12.7 

Insol 79 

99,79  "00,0 
CEtTntAL  High  School,  Philadelphia. 

*  I  hope  to  verify  this  by  actual  measurnnent. 


PROGRESS  OF  THE  DEMARCATION  OF  THE  ALASKA 

BOUNDARY. 

By  O.  H.  TITTMANN, 

U.  S.  Commissioner. 

{Read  April  24,  1908.) 

The  boundary  between  the  British  and  Russian  possessions  in 
North  America  was  defined  by  the  Treaty  of  St.  Petersburg  of 
1825.  When  the  United  States  purchased  the  Russian  possessions, 
or  Alaska,  in  1867,  it  was  believed  that  the  territorial  jurisdiction 
of  the  United  States  and  Great  Britain  could  not  become  a  matter 
of  controversy.  This  view  is  evidenced  by  the  remark  made  by 
Charles  Sumner  in  his  speech  advocating  the  purchase  of  Alaska. 
"  I  am  glad,"  said  he,  "  to  begin  with  what  is  clear  and  beyond  ques- 
tion.   I  refer  to  the  boundaries  fixed  by  the  treaty." 

The  total  length  of  the  boundary  referred  to  by  Mr.  Sumner  is 
twelve  hundred  miles.  It  divides  itself  naturally  into  two  sections 
of  about  six  hundred  miles  each.  One  is  the  section  bounded  by  the 
141st  meridian,  and  the  other  the  irregular  boundary  delimiting  the 
narrow  coast  strip  of  southeastern  Alaska.  No  dispute  has  ever 
arisen  as  to  that  part  of  the  boundary  defined  as  being  the  141st 
meridian  of  longitude  west  of  Greenwich.  As  is  well  known,  how- 
ever, a  contention  arose  as  to  that  part  of  the  boundary  which 
delimits  the  stretch  of  coast  extending  from  the  neighborhood  of 
Mt.  St.  Elias  southeasterly  to  and  through  the  Portland  Canal.  A 
modus  vivendi  in  1878,  affecting  the  Stikine  River,  and  another  in 
1899,  relating  to  the  country  at  the  head  of  Lynn  Canal,  made  tem- 
porary provision  for  customs  and  police  purposes.  The  dispute 
relating  to  that  part  of  the  boundary  was  happily  settled  by  the  Tri- 
bunal of  London  which  was  constituted  under  a  convention  signed 
at  Washington  January  24,  1903. 

86 


,9o8.]  OF  THE  ALASKA  BOUNDARY.  87 

I 

Hon.  John  W.  Foster,  the  agent  of  the  United  States  in  this 
important  case,  remarks  in  his  report  to  Secretary  Hay : 

**  It  is  a  noteworthy  fact  that  this  important  adjudication  was  brought  to 
a  close  within  less  than  eight  months  from  the  time  when  the  treaty  creating 
the  tribunal  went  into  effect  Such  a  prompt  result  is  almost  without 
parallel  in  the  intercourse  of  nations/' 

Equally  prompt  was  the  action  of  the  governments  in  appointing 
commissioners  in  accordance  with  a  requirement  of  the  convention 
constituting  the  Tribunal.  Within  a  few  months,  that  is,  in  the 
spring  of  1904,  the  commissioners,  Mr.  W.  F.  King,  on  behalf  of 
the  British  Government,  and  your  speaker,  representing  the  United 
States,  began  the  delimitation  of  that  part  of  the  boundary  which 
had  been  in  dispute.  The  commissioners  were  guided  in  their  plans 
by  maps,  accompanying  the  decision,  on  which  the  Tribunal  had 
marked  certain  mountain  p^aks  as  being  the  mountains  contemplated 
by  the  Treaty  of  1825. 

It  is  the  business  of  the  commissioners  to  identify  the  peaks,  to 
establish  their  geographical  position,  to  mark  by  visible  monuments, 
wherever  possible,  the  turning  points  in  the  line  and  such  other 
points  as  may  be  necessary,  and  to  describe  and  define  the  line 
between  the  points  selected  by  the  Tribunal.  There  was  a  stretch 
of  about  one  hundred  and  twenty  miles  where  the  topographic  infor- 
mation was  insufficient,  and  there  the  commissioners  were  directed 
to  make  additional  surveys  and  to  select  mountain  peaks  within  cer- 
tain prescribed  limits  to  define  the  boundary.  The  commissioners 
decided  to  mark  at  once  certain  river  crossings  and  the  mountain 
passes  and  to  connect  all  the  boundary  peaks  by  a  continuous  triangu- 
lation  based  on  the  trigonometric  datum  adopted  by  the  Coast  and 
Geodetic  Survey  for  southeastern  Alaska. 

The  boundary  line,  starting  from  the  rieighborhood  of  Mt.  St 
Elias,  crosses  that  summit  and  other  high  peaks  of  the  St.  Elias 
Alps  and  the  Fairweather  Range.  In  general,  it  lies  amid  perpetual 
snow  and  ice  except  when  it  drops  abruptly  into  the  river  valleys 
only  to  rise  again  into  regions  of  perpetual  snow.  Finally,  it  reaches 
the  head  of  Portland  Canal  and  becomes  a  water  boundary. 

In  the  four  years  since  work  was  begun  on  this  portion  of  the 
boundary  the  commissioners  have  fixed  trigonometrically  all  the  peaks 


88  TITTMANN— PROGRESS  OF  THE  DEMARCATION  [April  .4. 

except  two  near  Mt.  St.  Elias  and  those  in  the  region  between  the 
Whiting  River  and  Devil's  Thumb,  and  some  of  the  peaks  south  of 
the  Unuk  River.  The  passes,  valleys  and  river  crossings  have  been 
monimiented  with  the  exception  of  the  crossing  of  the  Alsek  in  the 
north  and  the  valleys  of  the  affluents  of  the  Iskut  and  the  crossing 
of  the  Le  Due  and  Chicamin  rivers  in  the  south.  The  turning  points 
of  the  water  boundary  in  Portland  Canal  also  remain  to  be  fixed  by 
reference  to  points  on  shore. 

The  141ST  Meridian. 

According  to  the  Treaty  of  1825  the  141st  meridian  west  of 
Greenwich  forms  the  eastern  boundary  of  Alaska  from  the  Arctic 
Ocean  to  near  Mt.  St.  Elias.  It  was  not  until  1889— twenty-two 
years  after  the  acquisition  of  Alaska — ^that  any  steps  were  taken  by 
our  government  towards  establishing  the  location  of  the  141st 
meridian  on  the  ground.  In  that  year  the  Coast  and  Geodetic  Sur- 
vey despatched  one  party  to  the  Yukon  and  another  to  the  Porcupine 
River  to  determine  the  boundary  crossing  of  those  rivers.  The 
Canadian  government  had  previously  sent  an  engineer  to  the  Yukon 
who  made  an  astronomical  determination  of  the  boundary  in  the 
autumn  and  winter  of  1887.  The  country  at  that  time  was  very 
inaccessible  and  the  surveyors  were  compelled  to  determine  the 
longitude  by  moon  culminations  and  occultations,  and  the  American 
parties  spent  a  whole  winter  in  observing  them.  But  the  operations 
of  the  three  parties  were  not  carried  on  under  an  international  agree- 
ment and  the  results  therefore  were  not  reciprocally  binding  on  the 
governments  concerned. 

The  discovery  of  gold  and  the  general  development  of  the  coun- 
try, however,  caused  the  construction  of  a  Canadian  telegraph  line 
overland  to  Dawson  and  beyond,  and  later  the  United  States  govern- 
ment laid  a  cable  from  Seattle  to  Sitka  and  thence  to  Valdez  on 
Prince  William  Sound,  whence  an  overland  line  was  built  by  the 
United  States  War  Department  as  far  as  Fort  Egbert  on  the  Yukon 
near  the  boundary.  Egbert  and  Dawson  were  also  connected  by 
telegraph.  This  important  auxiliary  to  longitude  determination 
made  it  possible  for  the  two  governments  to  determine  the  position 
of  the  141st  meridian  with  all  the  necessary  accuracy  as  soon  as  an 


,9o8.]  OF  THE  ALASKA  BOUNDARY.  89 

agreement  was  reached  and  embodied  in  the  treaty  signed  at  Wash- 
ington in  August,  1906.  This  treaty  provided  for  the  survey  and 
demarcation  of  the  line  and  before  the  end  of  that  summer  the  tele- 
graphic determination  of  the  Yukon  River  crossing  of  the  141st 
meridian  had  been  completed.  The  telegraphic  determination  made 
by  the  Americans  rests  on  the  known  longitude  of  Seattle.  Signals 
were  exchanged  between  Sitka  and  Seattle,  Seattle  and  Valdez, 
Valdez  and  Fort  Egbert,  Fort  Egbert  and  the  boundary.  That  made 
by  the  Canadians  rests  on  the  longitude  of  Vancouver  between  which 
place  and  the  boundary  time  signals  were  exchanged.  As  the  differ- 
ence between  Seattle  and  Vancouver  was  also  determined  by  the 
commissioners,  the  circuit  was  closed  and  a  very  satisfactory  agree- 
ment was  obtained. 

It  is  worthy  of  remark  in  passing  that  the  tracing  of  a  meridian 
or  parallel  on  the  groimd  involves  considerations  which  do  not 
become  apparent  by  an  inspection  of  an  artificial  globe  on  which 
these  lines  are  traced  as  smooth  and  regular  curves.  A  parallel  of 
latitude  must  be  determined  by  astronomical  observations,  but  in 
general  the  circumference  of  a  small  circle  of  the  earth  parallel  to 
the  equator  will  not  lie  in  the  same  astronomical  latitude,  owing  to 
the  so-called  deflection  of  the  vertical.  A  series  of  points  deter- 
mined astronomically  as  being  in  the  same  latitude  or,  as  in  the  case 
of  a  meridian,  in  the  same  longitude,  will  therefore  in  general  pro- 
duce on  the  surface  of  the  earth  a  zig-zag  line  when  they  have  been 
joined  together. 

In  order  to  avoid  all  questions  that  might  arise  from  local  deflec- 
tions of  the  zenith,  it  was  provided  by  the  Treaty  of  1906  that  the 
commissioners  should  determine  by  the  telegraphic  method  a  con- 
venient point  on  the  141st  meridian  and  then  trace  a  porth  and  south 
line  passing  through  the  point  thus  ascertained.  This  provision 
fixed  the  telegraph  crossing  of  the  boundary  as  the  initial  point  for 
the  longitude  determination.  The  commissioners  desired  to  make 
the  determination  as  nearly  on  the  141st  meridian  as* possible,  in 
order  to  avoid  a  deflection  error  which  might  have  been  involved  if 
the  longitude  had  been  obtained  by  linear  measurement  from  a  lon- 
gitude observed  at  some  distance  from  the  boundary.  The  transit 
pier  erected  for  the  purpose  of  exchanging  time  signals  was  found 


90  TITTMANN— PROGRESS  OF  THE  DEMARCATION  [April  04, 

to  be  in  longitude  141  **  00'  oo".4,  a  very  close  hit.  It  is  interesting 
to  note  also  that  the  final  longitude  differed  only  9".43  of  an  arc,  or 
410  feet,  from  that  derived  by  moon  culminations  about  twenty  years 
before. 

The  work  of  tracing  the  boundary  southward  from  the  Yukon 
was  beg^n  in  the  spring  of  1907  and  was  carried  southward  a  dis- 
tance of  about  one  hundred  and  twenty  miles.  Aluminum-bronze 
monuments  were  erected  on  the  north  and  south  banks  of  the  river, 
a  trigonometric  and  topographic  survey  was  made  extending  two 
miles  on  each  side  of  the  boundary  for  a  distance  of  about  forty-five 
miles,  and  a  broad  vista  was  cut  through  the  woods  for  the  same 
distance.  The  work  planned  for  the  coming  season  will  carry  the 
tracing  of  the  line  as  far  as  the  great  mountains  south  of  the  White 
River,  and  the  topographic  survey  and  the  monumenting  will  be 
pushed  until  the  severity  of  the  weather  compels  the  surveyors  to 
abandon  the  work  and  turn  their  faces  homeward. 


THE  MOST  PRIMITIVE  LIVING  REPRESENTATIVE  OF 
THE  ANCESTORS  OF  THE  PLANT  KINGDOM. 

By  GEORGE  T.  MOORE,  Ph.D. 
(Read  April  25,  1908.) 

There  is  but  little  doubt  among  botanists  that  the  land  flora  as 
it  now  exists  has  originated  from  aquatic  ancestors.  Both  from  the 
morphologic  and  palaeontologic  standpoints  the  evidence  corroborates 
this  view.  Indeed,  the  dependence  of  land  plants  upon  an  adequate 
water  supply,  together  with  the  fact  that  in  such  groups  as  the 
Mosses  and  Ferns,  fertilization  itself  can  only  be  accomplished  in 
the  presence  of  water  supplied  from  some  external  source,  gave  rise 
to  the  conclusion  that  the  origin  of  the  vegetable  kingdom  was  from 
primitive  plants  living  in  the  water,  long  before  there  was  the  more 
conclusive  evidence  now  existing. 

It  would  be  interesting  to  inquire  into  the  life  histories  of  certain 
transitional  groups  with  a  view  to  tracing  this  migration  from  water 
to  land.  For  modem  morphological  and  physiological  investigations 
has  enabled  us  to  do  this  with  a  considerable  degree  of  certainty. 
Not  only  would  we  be  able  to  show  that  the  establishment  of  the 
higher  representatives  of  our  land  flora  had  been  brought  about  by 
certain  methods  of  specialization  in  lower  aquatic  or  semi-aquatic 
forms,  but  it  would  be  possible  to  indicate  to  a  certain  extent  at 
least  how  this  process  had  been  carried  on.  However  such  an 
inquiry  would  lead  us  entirely  too  far  afield  at  this  time  and  it  will 
be  necessary  to  grant  without  further  discussion  that  the  facts  are 
sufficient  to  sustain  the  aquatic  origin  of  the  higher  plants. 

Naturally,  in  seeking  for  the  primitive  ancestors  of  the  vegetable 
kingdom,  attention  is  at  once  directed  to  the  algae,  the  g^oup  of 
plants  which  to  a  very  considerable  extent  is  more  dependant 
upon  the  presence  of  external  water  for  the  carrying  on  of  its  vital 
processes  than  any  other.  Furthermore,  in  the  present  state  of  our 
knowledge,  such  an  investigation  would  not  be  devoted  to  the  more 

91 


92  MOORE— MOST  PRIMITIVE  LIVING   REPRESENTATIVE   [Aprils, 

highly  differentiated  brown  or  red  algae,  but  rather  to  the  green  slgst, 
in  which  group  there  exists  the  closest  resemblances  to  the  structure 
of  the  lower  land  plant^.  The  problem  thus  becomes  one  of  discov- 
ering as  nearly  as  may  be  possible  the  most  primitive  member  of  the 
green  algse.  And  by  ''primitive/'  of  course,  is  not  necessarily  meant 
the  simplest  form,  but  that  plant  which  seems  to  be  nearest  to  the 
starting  point  of  the  phylogenetic  tree  and  from  which  certain  defi- 
nite lines  of  ascent  can  be  traced. 

In  considering  the  origin  of  the  g^een  algae,  numerous  theories 
have  been  held  and  it  would  be  impossible  to  give  even  a  mere 
outline  of  the  various  improbable  suggestions  which  have  been 
advanced  regarding  the  evolution  of  this  group.  During  the  past 
ten  years,  however,  a  g^eat  4eal  of  light  has  been  thrown  upon  the 
phylogenetic  relationship  of  the  algae.  Not  only  has  the  increase  in 
our  knowledge  of  the  life  histories  of  the  algae  been  considerable, 
but  the  discovery  of  many  new  genera  and  species  has  made  clear 
the  affinities  of  various  families  as  never  before.  Of  the  275  good 
genera  now  recognized  among  the  green  algae,  one  fourth  have  been 
discovered  and  described  since  the  appearance  of  Engler  and  Prantl's 
"  Pflanzenfamilien  " — ^the  last  complete  work  on  the  subject  and  still 
the  recognized  authority.  The  addition  of  so  many  new  and  in 
many  cases  important  links  to  the  chain  of  development  of  these 
plants,  has  reduced  the  former  chaotic  condition  to  something  like 
order  and  it  is  no  longer  quite  such  a  matter  of  speculation  regard- 
ing the  origin  of  the  main  group  of  the  green  algae. 

Ten  years  ago  Chodat  derived  the  green  algae  from  the  simplest, 
unicellular,  non-motile  forms  then  known,  namely,  the  Palmellaceae. 
Within  this  family  he  included  four  genera  whose  simple  life  his- 
tory showed  three  principal  stages.  From  these  so-called  "condi- 
tions," as  Chodat  pointed  out,  developed  the  three  important  and 
ruling  tendencies  which  have  dominated  the  lower  green  algae. 

These  are :  ( i )  The  zoospore  condition,  or  the  unicellular  motile 
stage,  with  the  other  two  conditions  transient  or  subordinate.  (2) 
The  sporangium  condition,  that  is,  the  unicellular  non-motile  stage, 
with  the  other  two  conditions  accidental  or  transient.  (3)  The 
tetraspora  condition,  where  the  non-motile  cells  are  connected  at 
right  angles  by  the  increasing  consistency  of  the  walls,  giving  rise 


,908.1  OF  THE  ANCESTORS  OF  THE  PLANT  KINGDOM.  93 

to  the  formation  of  a  tissue  or  filament.  The  .other  two  conditions 
are  reduced  or  transient 

Having  established  these  three  principal  "conditions,"  Chodat 
proceeded  to  establish  the  phylogeny  of  the  green  algae  along  these 
lines  and  succeeded  in  clearing  up  considerable  obscurity  which  had 
previously  existed.  However,  the  starting  point  selected  by  Chodat 
has  been  open  to  some  criticism  and  it  remained  for  Blackman  to 
suggest  the  most  satisfactory  explanation  of  the  origin  of  this 
group.  He,  while  following  in  a  general  way  the  theory  of  Chodat, 
took  the  position  that  the  three  "  tendencies  "  had  their  origin  not  in 
the  non-moile  Paltnella  form,  but  in  the  motile  Chlamydomonas  type. 

I  have  had  the  genus  Chlamydomotuxs  under  investigation  for 
several  years,  observing  its  various  species  for  the  most  part  in  pure 
cultures  grown  upon  both  solid  and  liquid  media.  The  vegetative 
cells  of  Chlamydoffwnas  2Lre  variable  in  both  size  and  shape;  in  gen- 
eral, however,  they  are  from  20-35 /n  ^^  length  and  10-20 /n  in 
breadth,  being  elliptic  or  pyriform  in  outline.  One  end  of  the  cell 
is  usually  produced  into  a  colorless  beak,  from  which  two  cilia 
always  protrude.  The  chloroplast  is  quite  variable  in  form  and 
with  one  exception  is  provided  with  a  single  pyrenoid.  Non-sexual 
reproduction  is  by  means  of  zoospores,  which  are  formed  by  the 
division  of  the  contents  of  the  mother  cell,  after  it  has  come  to  rest. 
Sexual  reproduction  is  usually  by  the  conjugation  of  naked  motile 
gametes  of  similar  size  and  in  no  way  distinguishable  from  each 
other.  It  is  interesting  to  note,  however,  that  in  addition  to  this 
method  there  may  also  be  the  conjugation  of  unequal  motile  gametes 
and  in  one  species — ^to  be  referred  to  later — ^there  takes  place  the 
conjugation  of  dissimilar  gametes,  one  of  which,  the  larger,  comes 
to  rest  before  conjugation.  We  thus  have  within  the  limits  of  this 
well  defined  and  natural  genus,  not  only  the  most  primitive  form 
of  gamogenesis,  but  through  anisogamous  conjugation  a  gradual 
approach  to  true  oogamy — the  highest  type  of  sexual  reproduction 
developed  among  the  algae. 

In  abandoning  the  starting  point  of  Chodat's  theory  of  the  devel- 
opment of  the  green  algae,  it  is  not  necessary  to  replace  his  idea 
relative  to  the  three  predominating  tendencies  manifest  in  the  lower 
members  of  this  group.    While  different  names  are  attached  to  these 


1 


94  MOORE— MOST  PRIMITIVE  LIVING  REPRESENTATIVE   [April  .4. 

conditions  as  recognized  at  the  present  time,  they  are  essentially  those 
pointed  out  by  Chodat,  namely: 

1.  A  tendency  towards  the  aggregation  of  motile  vegetative  cells, 
with  a  gradually  larger  and  more  specialized  motile  colony.  This  is 
the  Volvax  type  and  in  no  place  in  the  plant  kingdom  do  we  have 
a  more  perfect  series  of  development  than  from  the  simple  Chlamy- 
domonas  form  to  the  complex  and  highly  differentiated  Volvox  type. 

2.  A  tendency  towards  the  formation  of  an  aggregation  of  non- 
motile  cells  into  a  filament  or  tissue  by  the  repeated  vegetative 
division  of  an  original,  mother  cell.    This  is  the  Tetraspora  type. 

3.  The  Endasphara  type,  where  the  tendency  towards  the  forma- 
tion of  vegetative  divisions  and  septate  cell  formation  is  reduced  to 
a  minimum.  This  is,  of  course,  Chodat's  sporangium  tendency, 
although  not  so  much  importance  is  attached  to  it. 

Without  going  into  details  it  may  be  said  that  various  species  of 
Chlamydomonas  (of  which  there  are  about  thirty,  all  remarkably 
constant  as  regards  their  cytological  characters),  taken  collectively, 
exhibit  all  these  three  tendencies  and  that  the  simpler  forms  of  algae 
which  possess  but  a  single  tendency,  seem  clearly  to  have  diverged 
from  some  one  species  of  this  genus. 

The  endosphaerine  tendency  in  Chlamydomonas  has  given  rise 
to  a  single  family,  EndosptuBra,  This  is  naturally  strictly  unicel- 
lular and  with  no  vegetative  divisions;  the  reproduction  of  the 
species  can  take  place  only  by  the  formation  of  zoospores  or  gametes. 
A  family  so  restricted  as  to  its  vegetative  habit  could  hardly  be 
expected  to  develop  very  far  and  it  is  interesting  to  note  that  prac- 
tically all  the  genera  are  epiphytic  upon  other  algae  or  aquatic  plants, 
and  that  this  habit  of  life  has  undoubtedly  given  rise  to  a  distinct 
group  of  fungi.  The  suggestion  has  been  made  that  the  peculiar 
Siphonales  may  have  developed  from  this  EndospluBva  type,  and 
while  such  a  view  is  reasonable,  it  must  necessarily,  at  the  present 
time,  be  a  mere  matter  of  speculation. 

But  one  family,  the  Volvocaceae,  has  resulted  from  the  develop- 
ment of  the  volvocine  tendency.  While  the  evolution  of  sex  in  this 
group  has  been  carried  to  the  highest  possible  degree,  the  restric- 
tions of  an  enforced  motile  vegetative  condition  did  not  permit  this 
family  to  give  rise  to  anything  further. 


I908.]  OF  THE  ANCESTORS  OF  THE  PLANT  KINGDOM.  95 

It  is  the  tetrasporine  tendency  which  has  been  the  permanent  one 
and  has  resulted  in  producing  the  higher  green  plants.  This  condi- 
tion in  Chlcanydomoncts  resulted  in  the  production  of  a  series  of 
plants  which  gradually  replaced  the  formation  of  zoospores  by  that 
of  vegetative  cell  division.  The  resulting  family  was  the  Palmel- 
laceae,  the  one  which  formed  the  starting  point  in  the  development 
of  the  algae,  according  to  Chodat  While  there  seems  to  be  but  little 
question  that  the  Palmellaceae  have  given  rise  to  most  of  the  other 
families  of  the  green  algse,  there  is  every  evidence  that  it  was  itself 
derived  from  Chlamydomonas,  rather  than  the  reverse,  as  contended 
by  Chodat. 

It  is  impossible  at  this  time  to  even  indicate  the  development  of 
the  higher  algae  from  the  Palmellaceae.  With  the  exception  of  the 
Confervales,  which  seems  to  have  developed  independently  of  the 
typical  green  algae,  and  the  Conjugales,  which  apparently  have  arisen 
directly  from  the  Chlamydomonas  type,  all  the  higher  green  algae 
can  be  traced  back  through  the  Palmellaceae  with  considerable  cer- 
tainty to  their  Chcdmydomonas  ancestor.  The  0>njugales  have 
always  been  a  stumbling  block  in  constructing  any  develq>mental 
line  of  the  algae  from  primitive  forms.  But  granting  that  the  fila- 
mentous Conjugales,  as  well  as  the  desmids,  are  unicellular  (the 
reasons  for  which  can  not  now  be  given)  it  is  comparatively  easy 
to  find  the  origin  of  the  conjugation  habit  so  emphasized  in  this 
group,,  in  Chlamydom^mas  BrautUi,  In  this  species  the  female  or 
receptive  cell  comes  absolutely  to  rest  before  fertilization  and  the 
smaller  or  male  cell  becomes  attached  to  it.  Then  the  entire  con- 
tents of  the  male  cell  passes  into  the  female  cell,  leaving  behind  the 
empty  cell  wall,  just  as  in  some  of  the  desmids  and  in  most  of  the 
filamentous-like  Conjugales. 

The  more  the  genus  is  studied  the  more  reasonable  becomes  the 
conclusion  that  Chlamydomonas  has  not  only  given  rise  to  such  an 
aberrent  group  as  the  Conjugales,  but  that  it  may  safely  be  regarded 
as  the  phylogenetic  starting  point  of  the  various  lines  of  ascent  in 
the  true  green  algae.  At  present  in  the  higher  algae  the  Chlamy' 
domonas  stage  is,  of  course,  retained  in  the  zoospore  and  the  gamete, 
a  vegetative  non-motile  generation  being  interpolated  between  either 

PROa  AMER.  PHIL.  SOC.  XLVII.  l88  G,  PRINTED  JULY  II,  I908. 


96  MOORE— MOST  PRIMITIVE  LIVING   REPRESENTATIVE    [April  a4, 

a  sexual  or  non-sexual  motile  stage.  In  the  very  highest  type  of 
reproduction  in  the  green  algae  the  male  gamete  alone  represents 
the  Chlamydomonas  stage.  Indeed,  we  may  well  asstmie  that  the 
motile  male  gamete  of  the  mosses  and  ferns  constitutes  the  last 
remaining  type  of  the  original  Chlamydomonas  condition,  which 
with  other  more  positive  evidence  points  to  the  origin  of  such  land 
forms  from  a  Chlamydomonas-Vikt  ancestor. 

Although  practically  all  the  evidence  for  the  position  taken  has 
necessarily  been  omitted,  it  is  hoped  that  enough  has  been  said  to  at 
least  indicate  the  unique  and  important  position  occupied  in  the  plant 
kingdom  by  the  alga  Chlamydomonas. 


THE  COMPARATIVE  LEAF  STRUCTURE  OF  THE  SAND 

DUNE  PLANTS  OF  BERMUDA. 

(With  3  plates.) 

By  JOHN  W.  HARSHBERGER,  Ph.D. 

(Read  April  24,  1908,) 

The  writer  has  discussed  the  flora  of  Bermuda  in  two  papers  pub- 
lished in  the  Proceedings  of  the  Academy  of  Natural  Sciences  of 
Philadelphia  and  entitled  "The  Plant  Formations  of  the  Bermuda 
Islands"  (1905:  695-700)  and  "The  Hour-glass  Stems  of  the  Ber- 
muda Palmetto"  (1905:  701-704).  The  study  of  the  flora  pre- 
sented in  these  papers  and  the  study  of  the  microscopic  anatomy  of 
the  leaves  of  the  sand  dune  plants  herewith  given  is  the  result  of  a 
visit  to  the  islands  during  the  month  of  June,  1905. 

The  sand  beaches  and  sand  dunes  are  found  typically  developed 
along  the  south  shore  of  the  main  island  and  in  a  few  isolated  places 
on  the  north  shore,  as  at  Shelly  Bay.  The  largest  sand  beaches  and 
sand  dunes  on  the  south  shore  are  found  in  the  vicinity  of  Tucker- 
town  Bay,  on  the  narrow  strip  of  south  shore  between  Harrington 
Sound  and  the  ocean.  The  sand  dunes  along  the  south  shore  in 
the  parish  of  Paget  are  also  characteristic.  The  sand  dimes,  how- 
ever, in  the  neighborhood  of  Tuckertown  Bay  are  remarkable  in  that 
they  have  encroached  on  the  rocky  shore  line  and  have  invaded  the 
natural  arch  which  is  one  of  the  scenic  wonders  of  the  islands.  The 
sand  has  drifted  beneath  the  arch  and  has  advanced  so  that  it  covers 
part  of  the  top  of  the  arch  itself.     ' 

The  vegetation  of  the  beaches  and  dunes  here  and  in  the  vicinage 
of  the  Devil's  Hole  is  characteristically  Bermudian,  while  the  sand 
dunes  in  Paget  have  been  colonized  in  part  by  plants  introduced 
by  man  into  the  islands,  such  as  the  oleander,  Nerium  oleander,  and 
a  tall  fennel,  Fceniculum  zmlgare.  These  beaches  and  dunes  are 
formed  of  coral  sand  which  represents  the  finely  ground  masses  of 

97 


98  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE     [April  t4, 

coral  and  coralline  (calcareous)  sea- weeds  which  have  grown  on  the 
fringing  coral  reefs.  Bermuda,  geologically  speaking,  is  an  atoll, 
a  ring  of  coral  reefs  surrounding  a  central  lagoon.  The  elevated 
land  was  formed  by  the  raising  of  the  weather  edge  of  the  reef 
above  the  level  of  the  sea.  The  tops  of  the  projecting  corals  were 
broken  off  and  along  with  calcareous  sea-weeds  and  moUusk  shells 
were  ground  by  surf  action  into  a  fine  sand,  which  was  formed  into 
a  beach.  As  the  top  of  the  beach  dried  in  the  sun,  the  sand  was  blown 
off  and  was  deposited  in  the  crevices  of  the  coral  breakwater,  which 
gradually  widened.  Ulti|nately,  by  wind  action,  sand  hills  were 
formed.  The  limestone  rock  found  throughout  the  islands  was  origi- 
nally derived  from  broken-down  coral  and  shells.  These  rocks  vary 
in  texture  from  loose  sand  to  compact  limestone.  The  process  by 
which  the  coral  sand  was  converted  into  limestone  was  very  simple* 
and  it  involved  no  great  lapse  of  time.  As  the  sand  consists  almost 
entirely  of  calcium  carbonate,  it  was  easily  soluble  in  water  contain- 
ing carbon  dioxide.  The  rain  water  took  up  a  little  of  the  calcium 
carbonate  in  the  form  of  bicarbcmate,  and  as  it  percolated  through 
the  sand,  it  lost  its  carbonic  acid  gas  and  evaporating  left  the  dis- 
solved calcium  carbonate  as  a  thin  layer  of  cement  uniting  together 
the  grains  of  sand.  The  rocks  remain  permeable  to  water  and 
soluble,  so  that  this  process  of  solution  and  deposition  goes  on  con- 
stantly until  even  a  marble-like  limestone  may  result.  The  usual 
building  material  consists  of  blocks  of  limestone  sawed  out  of  the 
hillside.  When  built  as  a  wall  sufficient  solution  takes  place  so 
that  the  stones  become  united  together  into  an  almost  solid  piece. 
The  red  soils  of  the  islands  represent  the  one  per  cent,  residue  of 
solid  material  after  the  rain  has  leached  out  all  of  the  other  con- 
stituents. When  the  soluticm,.  owing  to  wave  action  or  constant 
rain  action,  is  excessive,  caverns  with  stalactites  and  sinks  are  formed. 
The  honey-combed  eolian  rock  of  the  shore  line  on  which  charac- 
teristic Bermuda  plants  occur  owed  its  origin  to  similar  water  erosion. 
The  sand  dunes  thus  represent  stages  intermediate  in  the  geologic 
changes  which  have  combined  to  give  the  present  form  to  the  islands. 
They  represent  shifting  masses  of  coral  sand,  forming  flat  surfaces 
in  some  places,  in  other  places  heaped  into  conical  dunes  or  raised 
into  long  ridges.     Frequently  dune  hollows  exist  as  a  result  of  wind 


I908.)  OF  THE  SAND  DUNE  PLANTS  OF  BERMUDA.  99 

action  in  scooping  out  the  sand.    These  dunes  form  the  setting  upon 
which  the  typical  sand  strand  plants  are  distributed. 

Plant  Distribution. — The  upper  beach  at  the  foot  of  the  dunes 
is  characterized  by  the  presence  of  Cakife  (Bqualis,  which  shows  a 
more  decided  branching  habit  than  the  closely  related  species  on  the 
coasts  of  the  American  continent,  Cakile  ff^aritima.  Besides  this 
plant,  the  botanist  sees  clumps  of  Tournefortia  gnaphaJades,  Sccevola 
Plumieri  and  Craton  tnaritimus.  The  shrubs,  however,  grow  most 
luxuriantly  on  the  slopes  and  summits  of  the  dunes.  Iponuea  pes- 
caprcB,  as  elsewhere  in  the  tropics  (Mexico,  the  West  Indies),  is  a 
typical  plant  of  the  upper  beach ;  in  fact,  the  upper  beach  is  char- 
acterized by  its  presence,  with  its  long  runners  growing  down  from 
the  slopes  of  the  dunes  out  upon  the  flat,  sandy  beaches.  On  the 
dune  slopes  in  Bermuda  it  is  associated  with  Sccevola  Plumieri  and 
the  crab  grass,  Stenotaphrum  americanum. 

Back  of  the  dune  crests  are  found  Tournefortia  gnaphdodes, 
Iponuea  pes-capra,  Sccevola  Plumieri,  Juniperus  bermudiana  (wind- 
swept forms),  Sisyrinchium  bermudicmum,  Lepidium  virginicum, 
Euphorbia  buxifolia  (a  prostrate  plant  growing  in  rosettes),  Canch 
vaJia  obtusifolia  (a  l^^minous  vine)  and  the  prickly  pear  cactus, 
Opuntia  vulgaris.  On  the  dunes  at  Tuckertown,  where  the  sand 
covers  the  entrance  to  the  natural  arch,  Sccevola  Plumieri  forms 
extensive  clumps  in  pure  association.  Solidago  sempervirens,  as  in 
the  eastern  United  States,  is  also  a  dune  plant,  together  with  the 
smooth  and  hairy  forms  of  Borrichia  cvrborescens,  Dodoncea  viscosa, 
a  small  tree  with  its  varnished  leaves,  is  also  a  tenant  of  the  dunes. 
The  most  interesting  dune  plant  is  Conocarpus  erectus,  which  is  a 
typical  mangrove  tree  growing  with  its  roots  affected  by  salt  water. 
In  Bermuda,  however,  it  occurs  perhaps  more  frequently  on  the  dry 
upper  slopes  of  the  dunes.  In  one  place  on  the  south  shore,  it  covers 
nearly  a  quarter  of  an  acre.  The  crab  grass,  Stenotaphrum  ameri- 
canum  forms  close  mats  on  the  lee  side  of  the  dunes. 

The  high  dunes  on  the  south  shore  of  the  parish  of  Paget  have 
been  invaded  by  a  number  of  exotic  plants,  introduced  by  man  into 
the  islands,  such  as  Nerium  oleander,  Lantana  camara,  L,  crocea, 
while  Croton  maritimus.  Canon/alia  obtusifolia,  Dodoncea  viscosa, 
Borrichia  arborescens  and  Passiflora  suberosa  are  among  the  most 


•    •  •      ^     i V' 


100  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE      [April  .4, 

abundant  native  plants.  Yticca  aloifolia  forms  clumps  on  low  sand 
dunes  at  Shelly  Bay,  on  the  north  shore,  associated  with  IponuBa  pes- 
capr<B,  Tournefortia  gnaphdodes  and  Opuntia  sp. 

EcoLOGic  Factors. — The  ecologic  factors,  which  have  influenced 
the  distribution  of  the  typical  sand  strand  plants  of  Bermuda,  must 
be  referred  to  briefly.  As  the  plants  of  the  Bermuda  sand  beaches 
and  sand  dunes  in  general  show  xercq)hytic  adaptations,  we  must 
look  upon  these  adaptive  arrangements  as  a  response  to  the  environ- 
ment. The  following  environmental  factors  must  be  considered  as 
influential  in  producing  the  xerophytic  structures  which  the  leaves 
of  the  Bermuda  beach  and  dune  plants  especially  show : 

1.  The  intense  illumination  from  above  is  an  important  ecologic 
factor. 

2.  The  reflection  of  light  from  the  white  coral  sand  and  the 
foam-crested  breakers  beyond  is  important. 

3.  The  action  of  the  strong  winds  that  blow  across  the  islands 
must  be  considered  as  modifying  plant  structure. 

4.  The  action  of  the  salt  spray  blown  inland  by  the  wind  is 
marked  in  the  case  of  some  plants. 

5.  The  permeability  of  the  sand  to  water,  so  that  after  a  rain  the 
surface  layers  quickly  dry  out,  has  its  influence. 

The  most  potent  factor  in  the  modification  of  leaf  structure  has 
been  undoubtedly  the  bright  illumination  from  above  and  below  (by 
reflection)  and  the  physiologically  dry  condition  of  the  soil. 

Structural  Adaptations. — ^The  leaf  adaptations  to  light  are 
found  in  the  increased  number  of  palisade  layers,  their  presence  on 
the  upper  and  under  sides  of  the  leaves,  and  their  arrangement,  so 
that  the  central  part  of  the  leaf  becomes  palisade  tissue  throughout,  a 
typical  staurophyll.  The  depression  of  the  stomata  below  the  sur- 
face, as  in  Sisyrinchium  bermudiantim,  the  distribution  of  the  stomata 
in  pits,  as  in  Nerium  oleander  and  Lantana  involucrata,  the  develop- 
ment of  hairs  as  in  Tournefortia  gnaphdodes,  the  varnished  leaves 
of  Dodoncea  viscosa  and  thick  epidermal  layers  and  cuticle  are  all 
arrangements  to  reduce  transpiration.  The  succulency  of  the  leaves 
of  some  of  the  dune  plants  is  developed  perhaps  for  water  storage 
and  the  presence  of  latex  should  be  mentioned  as  a  means  by  which 


i9o8.]  OF  THE  SAND   DUNE  PLANTS  OF  BERMUDA.  101 

a  dune  plant  is  protected  against  the  untoward  influences  of  its 
environment. 

Light  has  been  most  marked  in  influencing  the  development  of 
leaf  structure  displayed  by  the  typical  sand  dune  plants  of  Bermuda. 
The  stimuli  of  light  have  called  forth  functional  responses  which 
have  produced  changes  in  form  or  structure  of  the  leaves,  or  in  both. 
The  chlorenchyma,  composed  of  chloroplast-bearing  cells,  is  con- 
verted into  two  kinds  of  tissues,  palisade  and  spongy  parenchyma,  as 
a  direct  result  of  the  unequal  illumination  of  the  leaf  surfaces. 
Palisade  tissue  is  formed  as  a  response  to  light,  or  to  low  water 
content,  or  to  both.  When  both  leaf  surfaces  are  equally  illumi- 
nated, the  leaf  may  be  termed  isc^hotic,  when  unequally  illuminated 
diphotic.  Diphotic  leaves  which  are  unequally  illuminated  show  a 
division  into  palisade  and  spongy  parenchyma,  and  such  leaves  are 
called  by  Clements*  diphotophylls.  Isophotic  leaves,  equally  illumi- 
nated on  both  surfaces  have  a  more  or  less  uniform  chlorenchyma. 
Clements  divides  such  leaves  into, three  types:  (i)  The  palisade  leaf, 
or  staurophyll  in  which  the  palisade  tissue  extends  from  the  lower 
to  the  upper  epidermis.  (2)  The  diplophyll,  or  double  leaf,  where 
the  intense  light  does  not  penetrate  to  the  middle  of  the  leaf.  In 
consequence,  the  upper  and  lower  palisade  layers  are  separated  by  a 
central  loose  parenchyma,  which  is  for  water  storage.  (3)  The 
spongophyll,  in  which  the  rounded,  loose  parenchyma  cells  fill  the 
leaf  without  palisade  tissue.  The  influence  of  the  light  and  other 
environmental  conditions  on  leaf  structure  is  perhaps  best  shown  in 
the  thin  and  thick  leaves  of  Conocarpus  erectus  produced  on  diflFerent 
parts  of  the  same  tree  differently  related  to  the  incident  rays  of 
light.  A  detailed  description  of  these  structures  for  each  plant  will 
be  given  at  the  end  of  the  paper.  The  following  is  a  classification 
of  different  leaf  structures  and  the  plants  which  illustrate  such 
adaptive  arrangements : 

Thick  Cuticle. — Nerium  oleander,  Conocarpus  erectus  (thin 
leaf) ,  Sc(Bvola  PlunUeri, 

Thick  Epidermis. — Canavatia  obtusifolia,  Dodon(Ba  viscosa,  Sisy- 
rinchium  bermudianum,  Stenotaphrum  americanum,  Ipomcea  pes- 

^Clements,  F.  E.,  "Research  Methods  in  Ecology,"  138-145;  "Plant 
Physiology  and  Ecology,"  171-184. 


102  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE     rApril«4, 

caprcB,  Cakile  cequalis,  Borrichia  arborescens  (smooth  leaf),  Croton 
maritimus. 

Two  or  Three  Epidermal  Layers. — Euphorbia  buxifolia,  Nerium 
oleander,  Conocarpus  erectus  (thick  leaf),  Croton  ntaritimus,  Tour- 
nefortia  gnaphalodes. 

Two  or  More  Rows  of  Palisade  Cells. — Passiflora  suberosa, 
Dodoncea  viscosa,  Nerium  oleander,  Sesuvium  portulacastrum,  Cakile 
(Bqualis,  Conocarpus  erectus  (thin  leaf  and  thick  leaf),  Sccevola 
Plumieri,  Borrichia  arborescens  (smooth  and  hairy  leaves). 

Stomata  Depressed. — Sisyrinchium  bermudianum  Heliotropium 
curassavicum,  Sesuvium  portulacastrum,  Iponusa  pes-caprce,  Cakile 
(Bqualis,  Conocarpus  erectus  (thick  leaf),  Sccevola  Plumieri,  Borri- 
chia arborescens  (smooth  leaf). 

Stomata  in  Pits. — Lantana  involucrata,  Nerium  oleander. 

Succulent  Leaf. — Sesuvium  portulacastrum,  Cakile  cequalis,  Con- 
ocarpus erectus  (thick  leaf),  Sccevola  Plumieri,  Borrichia  arbor- 
escens (smooth  leaf). 

Hairy  Leaf. — Lantana  involucrata,  Nerium  oleander,  Borrichia 
arborescens  (hairy  leaf),  Croton  maritimus,  Toumefortia  gnaphor 
lodes. 

Varnished  Leaf. — Dodoncea  viscosa. 

Leaf  Becoming  Erect  in  Sun  Position. — Canavalia  obtusifolia, 
Sisyrinchium  bermudianum,  Stenotaphrum  americanum,  Iponuea  pes- 
capra. 

Overlapping  Leaves. — Euphorbia  buxifolia,  Sisyrinchium  bermu- 
dianum, Stenotaphrum  americanum. 

Latex  Tissue. — Euphorbia  buxifolia. 

Gimi-Resin. — Conocarpus  erectus. 

Crystals. — Passiflora  suberosa,  Croton  maritimus. 

Diphotophyll. — Passiflora  suberosa,  Canavalia  obtusifolia.  Eu- 
phorbia buxifolia,  Lantana  involucrata,  Nerium  oleander  =  $. 

Diplophyll. — Dodofuea  viscosa,  Sesuvium  portulacastrum,  Ipomcea 
pes-caprcp,  Cakile  cequalis,  Conocarpus  erectus  (thin  leaf),  Sccevola 
Plumieri,*  Borrichia  arborescens  (smooth  and  hairy  leaves),  Croton 
maritimus,  Tournefortia  gnaphalodes =g. 

*  Scavola  Plumieri  and  Toumefortia  gnaphalodes  are  given  twice,  because 
it  is  difficult  to  decide  whether  their  leaves  arc  diplophyll,  or  staurophyll. 


* 


•  •:   •••  .%   • 


* 


,9o«.]  OF  THE  SAND   DUNE  PLANTS  OF  BERMUDA.  103 

Staurophyll. — Heliotrapium  curassavicum,  Conocarpus  erectus 
(thick  leaf),  Scavola  Plumieri,  Tournefortia  gnaphalodes*  =  4. 

Spongophyll. — Sisyrinchium  bermudianum,  Stenataphrum  ameri" 
c<muin=2.  With  reference  to  the  last  two  plants,  it  should  be 
mentioned  that  the  leaves  of  these  plants  stand  erect,  thus  receiving 
the  incident  rays  of  light  on  the  edge  of  the  leaf,  hence  the  absence 
of  palisade  tissue  and  the  presence  of  spongophyll  structure. 

Detailed  Structure  of  Leaves. — The  sections  of  the  leaves 
which  were  studied  were  made  free-hand  with  a  razor.  After  stain- 
ing, the  sections  were  mounted  for  permanency  in  Canada  balsam. 
The  drawings  of  these  sections  were  made  by  the  use  of  the  micro- 
projection,  electric  lantern,  so  that  in  every  case  with  the  exception 
of  Craton  maritimus,  the  drawings  were  made  on  the  same  scale. 
The  sketches  of  stomata  are  none  of  them  drawn  to  the  same  scale. 
The  description  of  the  histologic  structure  of  the  leaves  of  each 
species  follows. 

Passiflora  suberosa  is  a  small,  slender  species  of  the  genus  found 
growing  over  the  sand  surface  of  the  dunes  in  the  parish  of  Paget. 
Its  flowers  are  small  and  the  branch  tendrils  are  characteristically 
developed.  Histologically  the  leaf  presents  an  upper  epidermis  of 
large 'thin-walled  cells,  and  as  the  whole  plant  is  brilliantly  illumi- 
nated, it  has  two  well-marked  layers  of  palisade  cells.  The  loose 
parenchyma  is  narrow  and  some  of  the  cells  of  it  are  filled  with  con- 
glomerate crystals.  The  stomata  are  slightly  raised  above  the  gen- 
eral epidermal  surface,  and  are  confined  to  the  lower  side  of  the 
leaf.    A  diphotophyll  (Fig.  i,  Plate  II.). 

Canavalia  obtusifolia,  a  trailing  legtiminous  plant,  has  paripinnate 
compound  leaves  with  a  long  petiole  and  broadly  elliptical  leaflets 
with  retuse  apices  and  petiolules,  a  quarter  of  an  inch  long.  The 
upper  epidermis  consists  of  slightly  thickened  cells.  There  are  two 
rows  of  palisade  cells,  a  considerable  amount  of  loose  parench)mia, 
while  the  slightly  raised  stomata  are  found  on  the  upper  and  under 
sides.  The  adaptation  to  thfe  environment  of  the  sand  dunes  seems 
to  be  the  folding  together  of  the  two  sides  of  the  leaves  along  the 
midrib,  so  that  the  edges  of  the  leaves  are  presented  to  the  incident 
rays  of  light.    A  diphotophyll  (Fig.  2,  Plate  II.). 

Euphorbia  buxi folia  is  a  prostrate,  tufted  plant  of  a  rosette  habit. 


104  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE     [April  .4. 

The  taproot  is  large  and  strong  and  from  it  numerous  branches,  six 
to  eight  inches  long,  are  formed.  The  leaves  are  opposite,  small, 
ovate,  with  an  acute  apex  and  barely  petiolate.  The  upper  epidermis 
consists  of  two  rows  of  cells,  the  palisade  is  a  single  layer  and  the 
loose  parenchyma  is  compact.  The  lower  epidermal  cells  are  papil- 
late and  latex  is  present.  The  adaptation  to  the  environment  is 
shown  in  the  latex,  the  two-layered  upper  epidermis  aod  the  over- 
lapping arrangement  of  the  leaves.  A  diphotophyll  (Fig.  3, 
Plate  II.). 

Dodofusa  viscosa, — This  small  sapindaceous  tree  occurs  on  the 
inner  edges  of  the  sand  dunes.  Its  leaves  are  alternate,  spatulate 
with  the  base  narrowed  to  the  point  of  attachment.  The  leaves  are 
varnished.  The  upper  epidermal  cells  are  thick  and  provided  with 
peltate  hairs.  The  palisade  cells  are  disposed  in  two  layers.  The 
loose  parenchyma  is  open,  while  next  to  the  lower  epidermis  there  is 
a  row  of  small  cells  which  may  be  considered  as  a  lower  palisade 
layer.  Hence  the  leaf  is  a  potential  diplophyll.  The  stoniata  of  the 
upper  side  are  slightly  raised  above  the  surface,  while  those  on  the 
under  side  have  developed  a  small  projecting  beak  (Fig.  4,  Plate  II.). 

Lantana  involucrata  is  one  of  the  plants  thait  enters  the  formation 
of  the  Bermuda  scrub.  It  also  invades  the  dunes.  The  leaves  are 
hairy  on  both  surfaces.  A  section  of  a  leaf  shows  that  the  upper 
epidermis  is  without  stomata,  but  is  provided  with  straight,  multi- 
cellular and  capitate,  unicellular  hairs.  The  lower  surface  shows 
depressions  provided  with  the  capitate  hairs,  while  the  raised  por- 
tions of  the  leaf  surface  between  the  depressions  is  covered  with 
both  straight,  muticellular  and  capitate,  unicellular  hairs.  The  pali- 
sade is  a  single  layer.  The  stomata  project  outward  beyond  the 
general  surface  of  the  lower  epidermis,  but  they  always  occur  in 
the  depressions.  The  depressions  provided  with  hairs  and  stomata 
and  thick,  hairy  upper  epidermal  surface  are  structures  which  fit  the 
plant  to  exist  on  the  hot,  sun-exposed  sand  dunes  of  the  islands.  A 
diphotophyll  (Fig.  5,  Plate  II.). 

Nerium  oleander, — ^The  leaf  structure  of  the  oleander,  a  native 
of  the  Mediterranean  flora,  is  well  known.  The  upper  epidermis  is 
in  three  layers  with  thick  cuticle,  the  paliside  tissue  in  two  layers, 
.While  the  under  surface  of  the  leaf  is  pitted,  the  pits  being  filled 


i9o8.]  OF  THE  SAND   DUNE  PLANTS  OF  BERMUDA.  106 

with  straight  hairs  that  form  an  air-still  chamber  into  which  the 
projecting  stomata  open.  The  lower  epidermis  is  two-  to  three- 
layered,  and  the  whole  leaf  is  decidedly  tough  and  leathery,  and  thus 
well  adapted  to  growing  on  the  sand  dunes  of  Bermuda.  A  dipho- 
tophyll  (Fig.  6,  Plate  II.). 

Sisyrinchium  berfnudianum. — ^The  Bermuda  blue-eyed  grass  is 
provided  with  leaves  that  stand  more  or  less  upright,  so  that  the  inci- 
dent rays  of  light  strike  the  edges  of  the  leaves.  The  epidermal 
cells  on  both  the  upper  and  lower  morphologic  sides  of  the  leaf  are 
thick-walled  and  the  stomata  present  on  both  surfaces  are  depressed 
the  entire  width  of  the  epidermal  cells.  There  is  no  palisade  tissue, 
the  loose  parenchyma  filling  the  center  of  the  leaf  between  epidermal 
surfaces.  The  vertical  leaves  are,  therefore,  isophotic  and  the  leaf  is 
Icnown  as  a  spongophyll.  The  vertical  leaves,  the  thick  epidermal 
cells  and  the  depressed  stomata  fit  the  plant  to  its  environment  A 
spongophyll  (Fig.  7,  Plate  II.). 

Stenotaphrum  americanum.  The  Bermuda  crab  grass  is  a  tough, 
wiry  one,  well  fitted  to  survive  in  the  driest  places  on  sand  dunes  and 
rock  faces.  The  leaf  blades  arise  from  sheaths  that,  together  with 
other  overlapping  leaf  sheaths,  form  a  tuft  that  arises  from  the  nodal 
regions  of  the  wiry,  prostrate,  creeping  stem.  The  blades  are  more 
or  less  erect  and  folded  partially  lengthwise,  with  the  upper  side 
innermost.  The  spike  of  closely  set  flowers  is  slightly  bent,  sug- 
gesting a  crab's  claw.  The  upper  epidermis  consists  of  large,  open 
papillate  cells.  The  loose  parenchyma  fills  the  leaf  section  and  the 
under  surface  of  the  leaf  has  a  thick  epidermis  with  numerous 
stomata,  provided  with  small  g^ard  cells  reen  forced  by  two  secon- 
dary cells.  The  bundles  are  toward  the  upper  side.  The  vertical 
isophotic  leaf  consequently  becomes  a  spongophyll.  The  adapta- 
tions to  the  environment  are  upright,  rolled  leaves,  thick  lower  epi- 
dermis and  overlapping,  tufted  leaf  sheaths  (Fig.  8,  Plate  II.). 

Heliotrapium  curassavicum  resembles  in  its  unilateral  cymose 
inflorescence  the  common  heliotrope.  It  is  a  slightly  woody  plant 
that  grows  about  a  foot  or  two  tall,  with  alternate,  narrow,  oblanceo- 
late  leaves.  The  cells  of  both  the  lower  and  upper  epidermis  are 
thin-walled,  with  slightly  sunken  stomata  on  both  sides.  The  chlor- 
ophyll bearing  cells  of  the  leaf  (the  chlorenchyma)  are  arranged  so 


106  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE      [April  .4, 

that  their  long  axes  are  placed  in  a  line  with  the  incident  rays  of 
light  that  strike  the  upper  surface  from  above  and  the  lower  surface 
by  reflection  from  the  sand  below.  A  staurophyll  (Fig.  9, 
Plate  II.). 

Sesuvium  portulacastrum, — ^The  leaf  structure  of  this  member  of 
the  family  Aizoaceae  is  that  of  a  typical  diplophyll,  but  with  a  slight 
indication  of  the  staurophyll  arrangement  of  the  cells.  The  stomata 
present  on  both  sides  of  the  leaf  are  slightly  sunken  and  the  guard 
cells  incline  inward  and  downwards.  The  uf^er  and  lower  palisade 
tissues  show  four  to  five  layers  of  cells.  The  leaves  are  thick  and 
succulent.    A  diplophyll  (Fig.  10,  Plate  II.). 

IponuBa  peS'CaprcB. — ^This  tropical,  seaside  morning  glory  is  a 
typical  plant  of  the  sandy  beaches  in  Mexico,  the  West  Indies  and 
Bermuda.  It  grows  down  off  the  dune  sieves  onto  the  beach  sand 
as  a  creeping  plant,  a  distance  of  twenty  to  thirty  feet  (Fig.  i,  Plate 
I.).  The  leaves  are  alternate,  elliptical,  retuse  at  the  apex  and 
frequently  when  the  sun  is  hot  and  the  reflection  from  the  sand 
intense,  the  leaves  fold  together  along  the  midrib  and  stand  vertically 
so  as  to  receive  the  incident  rays  of  light  on  the  upturned  edges  of 
the  leaves.  The  walls  of  the  epidermal  cells  on  both  sides  of  the 
leaf  are  thick.  The  stomata  on  both  sides  are  sunken  about  half 
the  thickness  of  the  epidermal  cells  and  the  palisade  tissue  is  promi- 
nent on  both  sides,  constricting  the  loose  parenchyma  to  a  narrow 
layer.    The  leaf  is,  therefore,  a  true  diplophyll  (Fig.  11,  Plate  III.). 

Cakile  (Fqualis, — This  cruciferous  plant  grows  on  open,  sandy 
beaches  in  a  more  or  less  scattered  manner.  It  branches  in  a  much 
more  open  way  than  C.  maritima,  found  in  similar  habitats  on  the 
sandy  beaches  of  the  eastern  United  States.  The  leaves  are  fleshy^ 
and  the  walls  of  the  upper  and  lower  epidermal  cells  are  thickened. 
The  stomata,  which  are  partly  sunken,  are  found  on  both  the  upper 
and  the  lower  leaf  surfaces.  The  palisade  tissue  on  both  sides  is 
five  layers  of  cells  thick  and  the  loose  parenchyma  is  restricted  to  a 
narrow  layer  four  cells  thick  in  the  central  part  of  the  leaf.  This 
plant  is  fitted  to  its  environment  by  the  possession  of  succulent 
leaves,  epidermal  cells  with  thick  walls,  and  many-layered  palisade 
tissue.  A  diplophyll  (Fig.  12,  Plate  III.).  Contrast  the  leaf  section 
of  Cakile  maritima  (Fig.  12  A,  Plate  III.). 


i9o8.]  OF  THE  SAND  DUNE  PLANTS  OF  BERMUDA.  107 

Conocarpus  erectus. — ^The  leaves  of  this  small  tree,  which  is  a 
true  mangrove  plant,  but  which  has  adapted  itself  to  growth  on  the 
sand  dunes  in  Bermuda,  are  thin  and  thick.  The  thin  leaves  are 
found  on  the  branches  that  are  placed  above  the  surface  of  the  sand, 
or  in  more  or  less  protected  positions,  while  the  thick,  succulent 
leaves  occur  near  the  surface  of  the  sand,  or  in  exposed,  unshaded 
positions.  There  iff  a  considerable  difference  in  the  anatomical  struc- 
ture. The  cuticle  in  the  thin  leaf  is  thickened  and  the  stomata  on 
both  sides  are  hardly  if  any  sunken  below  the  surface.  The  upper 
leaf  surface  shows  long  palisade  cells,  while  the  palisade  cells  of  the 
lower  side  are  shorter.  The  loose  parenchyma  cells  form  a  broad 
band  in  the  center  of  the  section.  A  diplophyll  (Fig.  13,  Plate  III.). 
The  thick,  succulent  leaf  has  three  rows  of  epidermal  cells  and 
three  rows  of  palisade  cells,  the  cavities  of  which  are  filled  with  a 
gummy,  resinous  material  (not  tested)  of  a  brown  color.  This 
gummy  material  is  found  in  the  lower  palisade  as  well  as  in  the 
upper  palisade  in  both  the  thin  and  thick  leaves  and  also  in  some  of 
the  loose  parenchyma  cells  of  the  thick  leaf.  The  stomata  in  the 
thick  leaves,  by  the  increase  in  the  thickness  of  the  cuticle,  are 
sunken  below  the  surface  with  an  hour-glass  atrium  or  passage  out- 
side of  the  thick-walled  g^ard  cells.  The  parenchyma  cells  of  the 
leaf  center  are  arranged  in  the  direction  of  the  palisade  cells.  A 
typical  staurophyll  (Fig.  13  A,  Plate  III.). 

SccBvola  Plumieri. — This  plant  belongs  to  the  family  Goodeniaceae 
and  forms  dense  clumps  on  the  dune  sieves  (Fig.  2,  Plate  I.). 
Its  leaves  are  alternate,  elliptical,  short  petiolate  and  obtuse.  They 
are  noted  for  their  succulency.  The  epidermal  cells  on  the  upper 
surface  have  a  thick  cuticle  with  numerous  thick  walled,  sunken 
stomata.  The  epidermal  cells  on  the  lower  surface  are  of  the  same 
thickness  as  on  the  upper  surface,  the  stomata  being  likewise  sunken. 
The  palisade  cells  on  the  upper  and  lower  sides  consist  in  each  of 
three  or  four  rows  of  cells,  while  the  loose  parenchyma  is  arranged 
parallel  to  the  palisade  tissue.  Only  a  single  row  of  central  cells 
are  not  so  disposed.  The  leaf  shows,  therefore,  partly  a  staurophyll 
and  partly  a  diplophyll  arrangement  of  cell  (Fig.  14,  Plate  III.). 

Borrichia  arborescens. — This  species  of  the  family  Composite 
exists  in  two  distinct  forms,  if  they  are  not  good  species.    One  ir 


108  HARSHBERGER— COMPARATIVE   LEAF  STRUCTURE     [April  24, 

has  smooth,  thick,  succulent  leaves,  the  other  has  thinner,  densely 
tomentose  leaves,  the  Borrichia  frutescens  of  the  Southern  States. 
The  succulent,  smooth-leaved  form  has  both  thick  upper  and  lower 
epidermal  cells,  with  the  stomata  on  both  sides,  but  more  plentiful 
on  the  lower  side.  The  stomata  are  partly  sunken.  The  palisade 
layers  on  both  sides  are  wide,  but  are  broken  into  mor^  or  less  ex- 
tended patches  by  round  parenchyma  cells,  which  reach  to  the  epi- 
dermis. The  loose  parenchyma  cells  form  a  wide  central  area.  A 
diplophyll  (Fig.  15,  Plate  III.).  What  the  thin  leaf  lacks  in  suc- 
culency,  it  gains  in  hairiness.  Both  sides  are  densely  covered  with 
straight  unicellular  hairs.  The  palisade  layers  are  only  two  in 
number  on  both  sides  of  the  leaf,  and  the  loose  parenchyma  is  also 
much  reduced  in  amount.  The  succulency  of  the  thick  leaf  fits  it 
as  perfectly  as  the  hairiness  of  the  thin  leaf  to  the  trying  seaside 
environment,  where  the  plants  producing  them  grow  side  by  side. 
A  diplophyll  (Fig.  15  A,  Plate  III.). 

Croton  moHHmus. — The  leaves  of  this  plant  studied  by  Kearney* 
are  bifacial,  both  surfaces  densely  covered  with  gray  scale-like  pubes- 
cence, owing  to  presence  of  multicellular,  stalked,  stellate  hairs  that 
cover  them.  The  upper  and  lower  epidermal  cells  have  thick  walls 
and  the  stomata  are  not  sunken.  The  palisade  tissue  in  both  the 
upper  and  the  lower  sides  are  two  cell  layers  in  width  with  a  few 
sclerotic  idioblasts.  The  leaf  in  the  plant  grown  in  the  United  States, 
as  depicted  by  Kearney,  has  only  one  row  of  palisade  cells.  Large 
conglomerate  crystals  of  calcium  oxalate  are  found  in  the  cells  of 
the  loose  parenchyma.  Glandular  capitate  hairs  are  found  on  both 
leaf  surfaces.    A  diplophyll  (Fig.  16,  Plate  III.). 

Tournefortia  gnaphdodes, — ^The  leaves  and  stems  of  this  plant, 
as  well  as  the  calices  of  the  flowers,  are  covered  with  a  dense,  closely 
appressed,  grayish  tomentum,  resembling  that  on  our  common  An- 
tennaria  plant aginifolia  and  edelweiss,  Leontopodium  alpinunk  In 
section  the  hairs  are  unicellular,  straight  and  of  epidermal  origin. 
The  palisade  is  formed  on  the  upper  and  lower  leaf  surfaces  and  is 
two  cells  thick.  The  loose  parenchyma,  occupying  the  center  of  the 
leaf,  suggests  an  arrangement  in  direction  parallel  to  the  long  axis 

*  Kearney,  Thomas  H.  "  Plants  of  Ocracoke  Island,"  Contributions  from 
the  United  States  National  Herbarium  V:  296. 


,908.]  OF  THE  SAND  DUNE  PLANTS  OF  BERMUDA.  109 

of   the    palisade    cells.    Therefore    it   is   a   diplophyll    (Fig.    17, 
Plate  III.). 

Bibliography. — Little  has  been  published  on  the  structure  of 
the  dune  plants  of  tropical  America.  The  following  papers  are 
in  part  a  contributi(»i  to  our  knowledge  of  the  microscopic  struc- 
ture of  the  strand  plants  of  the  American  tropics.  A  few  of  the 
sand  dune  plants  are  of  cosmopolitan  distribution  and  they  are? 
therefore,  described  as  to  their  morphology  in  the  classic  work 
of  A.  F.  W.  Schimper,  "Die  indo-malayische  Strandflora,"  pub- 
lished as  the  third  volume  of  "Botanische  Mittheilungen  aus  den 
Tropen"  in  1891.  Thomas  Kearney  in  1900  published  in  the  Con- 
tributions from  the  U.  S.  National  Herbarium  (V.,  No.  5)  an 
important  paper  on  "The  Plant  Covering  of  Ocracoke  Island;  A 
Study  in  the  Ecology  of  the  North  Carolina  Strand  Vegetation."  A 
chapter  is  devoted  to  the  histological  structure  of  the  plants.  The 
only  plants  which  concern  us  are  Yucca  aloifolia,  Croton  maritimus, 
Borrichia  frutescens,  which  are  common  also  to  the  Bermuda  strand. 
F.  Boergesen  and  Ov^  Paulsen  make  a  contribution  to  "  La  Vegeta- 
tion des  Antilles  Danoises  "  in  Revue  Generale  de  Botanique  (Tome 
XII.,  1900),  in  which  they  discuss  with  figures  the  microscopic 
structure  of  a  few  of  the  typical  strand  plants.  As  throwing  con- 
siderable light  on  the  problems  concerned  in  this  paper  on  the  Ber- 
muda strand  flora  reference  should  be  made  to  these  works  of 
general  import  to  the  botanical  questions  involved. 

Diels,  L. 

Stoffwechsel  ^  und    Structur   der   Halophsrten,   Jahrbucher    fur   wissen- 
schaftlichc  Botanik  XXIIL :  309-322,  1^)8. 

Schimper,  A.  F.  W. 

Pflanzengeographie  auf  physiologischer  Grundlage,  1898. 

Solereder,  H. 

Systematische  Anatomic  der  Dicotyledoncn,  1898-99. 

Wamdng,  E. 

Halofyt  Studier,  Memoires  dc  1'  Acad6mie  Royale  de  Danemark,  ser.  6, 
VIIL,  No.  4.  1897. 


110  HARSHBERGER— COMPARATIVE  LEAF  STRUCTURE      |ApriiM. 


Illustrations. — ^The  reproduced  photographs  (Figs,  i  and  2, 
Plate  I.)  represent  the  dune  vegetation  on  the  south  shore  of 
Bermuda.  The  upper  illustration  shows  the  thicket  of  composite 
vegetation  on  the  crest  of  the  dune  and  the  long,  trailing  stems  of 
IpoffKBG  peS'CarpcB  on  the  upper  beach  with  a  small  clump  of 
Cakile  (Bqualis  to  the  left  in  the  foreground.  The  second  illustra- 
tion depicts  a  clump  of  Sccevola  Plumieri,  with  the  Bermuda  cedar, 
Juniperus  bermudiana,  and  in  the  background  the  grayish-green 
bushes  of  Tournefortia  gnaphaJodes.  Reference  is  made  to  the 
drawings  of  microscopic  structure  in  the  classified  description  of 
dune  plants  throughout  the  paper. 


PROCEEOINOS  Am.  Philos.  8oc.  Vol.  XLVll,  No   1 


PROCEEOINQS  Am.  Philos.  Soc.  VOL.  XLVJt.  No.  181 


PfloctEDiNQs  Am.  Philos.  Soc.  Vol.  XLVII.  No.  188 


^a--.. 


i.'!'^ 


SOLUTION  OF  ALGEBRAIC  EQUATIONS  IN  INFINITE 

SERIES. 

By  PRESTON  A.  LAMBERT. 
(Read  April  ^5,  igo8,) 

I.  Introduction. 

1.  The  object  of  this  investigation  is  to  develop  a  method  for 
determining  all  the  roots,  real  and  imaginary,  of  an  algebraic  equa- 
tion by  means  of  infinite  series. 

2.  Suppose  the  given  equation  to  be  represented  by  f(y)=o. 
The  method  consists  in  introducing  a  factor  x  into  all  the  terms  but 
two  of  the  given  equation ;  expanding  y,  which  now  is  an  algebraic 
function  of  x,  into  a  power  series  in  x ;  placing  x  equal  to  unity  in 
this  power  series.  The  resulting  value  of  y,  if  convergent,  is  a  root 
of  the  given  equation  expressed  in  terms  of  the  coefficients  and  expo- 
nents of  the  equation. 

3.  The  method  presupposes  the  solution  of  the  two-term  equation 

In  fact  the  roots  of  this  equation  when  written  in  the  form 

y  == =  r(cos  0  +  t  sin  0) 

are  found  to  any  required  degree  of  approximation  from  the  formula 

1/        2S'ir  +  0       .  .    2S'ir  +  0\ 
^  =  r »  ^  cos  — ~ -h  ^  sm  — j , 

where 

^=0,1,2,3,4,  ...,n— I. 

4.  The  method  proceeds  step  by  step  from  the  two-term  equation 
to  the  three-term  equation,  from  the  three-term  equation  to  the  four- 
term  equation,  and  so  on. 

PROG.  AMER.  PHIL.  SOC.  XLVII.  l88  H,  PRINTED  JULY  1 8,  I908. 


112 


LAMBERT— SOLUTION  OF   ALGEBRAIC  EQUATIONS      [April  as. 


11.   The  Three-term  Equation. 

5.  In  the  three-term  equation 

the  two  terms  from  which  the  ^  is  to  be  omitted  can  be  selected  in 
three  different  ways.    This  gives  rise  to  the  three  equations 

(i)  ay* -j-fcy*;r-j-c  =  o 

(2)  ay^ -^by^ ']-cx  =  o 

(3)  ay«^-f  fcy*-f  c=o 

each  one  of  which  defines  y  as  an  algebraic  function  of  x. 

6.  Values  of  y  expressed  as  power  series  in  x  may  be  found  from 
each  one  of  these  three  equations  by  any  one  of  the  following  three 
methods,  which,  however,  are  essentially  the  same. 

7.  The  Multinomial  Theorem. — Assume  that  the  power  series 
for  y  is 

(4)  y  =  Po  +  Pi^  +  p2^^  +  P^^  +  Pa^  +'"' 

The  multinomial  theorem  asserts  that  the  coefficient  of  x^  in  the 
expansion  of  y"  is 

provided 

(6)  gi  +  2^2  -f  3^3  -f 

(7)  ^0  +  ^1  +  ^2+-- 


,  p  ff«;^9l+2ffl+S«8+  ...»«• 


q»  =  n. 

The  expansion  of  y*  is  obtained  in  like  manner. 

Assuming  that  the  power  series  (4)  represents  the  algebraic 
function  defined  by  equation  ( i ) ,  the  substitution  of  the  expansions 
of  y*  and  y*  in  equation  ( i )  must  give  an  identity.    This  identity  is 


+^«/rVi 


+*/s 


x+ 


an{n-i)^^__ 


I  -2 

in-l. 


A"-V>? 


(8) 


,    ««(«— iY«— 2)      ,  .x*+ 
^  +        I  •  23       ^^' 


«9o8.] 


IN  INFINITE  SERIES.  113 


In  this  identity  the  coefficient  of  each  power  of  x  equals  zero. 
Hence  />o  is  the  root  of  the  two-term  equation 

The  coefficient  of  the  first  power  of  ;r  equated  to  zero  determines 
P^  uniquely  in  terms  of  p^ ;  the  coefficient  of  x^  equated  to  zero  deter- 
mines p^  uniquely  in  terms  of  p^  and  p^ ;  in  general,  the  coefficient  of 
X*  equated  to  zero  determines  pt  uniquely  in  terms  of  p^^px^Pz*  *••> 
^«_i.  All  the  successive  coefficients  of  the  power  series  (4)  are 
therefore  determined  uniquely  in  terms  of  p^^  any  one  of  the  roots 
of  the  two-term  equation  ap^  +  r=o. 

The  power  series  representing  the  algebraic  functions  defined 
by  equations  (2)  and  (3)  are  determined  in  precisely  the  same  man- 
ner. Unfortunately  if  the  coefficients  of  the  power  series  are  deter- 
mined in  this  way  it  is  difficult  to  recognize  the  law  which  will 
enable  one  to  write  the  general  term  of  the  power  series,  which  is 
necessary  for  the  application  of  a  convergency  test. 

When  X  is  made  unity,  the  equations  (i),  (2)  and  (3)  become 
the  three-term  equation 

and  the  power  series,  if  convergent  when  ^=1,  becomes  the  solution 
of  this  equation. 

If  it  is  known  in  advance  that  some  one  of  equations  (i),  (2), 
(3)  furnishes  a  power  series  which  is  convergent  when  x  =  i,  the 
multinomial  theorem  determines  in  an  elementary  and  direct  manner 
the  coefficients  of  the  power  series. 

8.  Maclaurin's  Series, — ^The  algebraic  function  y  defined  by  the 
equation 

can  be  expanded  into  a  power  series  in  x  by  means  of  Maclaurin's 
series 

^^^  -^     -^^     dx^        dx^  1-2      dx*  I  •  2  •  3 

The  expansion  is  identical  in  form  with  the  expansion  obtained 
by  means  of  the  multinomial  theorem  and  consequently  has  the  same 
disadvantage. 


114  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS       [April  as, 

9.  Lagrang^s  Theorem. — The  equation 
may  be  written 

(10)  y« — ^-y. 

Placing  y**  ^  xr,  whence  y  =  ^*/**,  this  equation  becomes 

c         6  ^ 
(11)  -s== x-z"^. 

Lagrange's  theorem  asserts  that  if 

(12)  ^      J 

If  now 

f{z)^z^,     <K^)^--z\ 

Cm 

and  after  the  derivatives  in  series  (12)  have  been  formed  z/ is  replaced 
by  —  c/a,  there  results,  making  x  unity, 


n 


(13)    +  yy^(i  +  3>& -«)(!  + 3^ -2«)(--)-^ 

In  series  (13)  the  law  of  formation  of  the  successive  terms  is 
evident  and  this  law  is  readily  proved  by  induction  by  using  La- 
grange's theorem. 

Series  (13)  may  be  more  concisely  written  by  placing 


(-i)-- 


J'o. 
so  that  yo  is  a  root  of  the  two-term  equation 


i9o8.] 


IN  INFINITE  SERIES.  115 


and  denoting  the  continued  product 

(i  +  sk  —  n){i+sk  —  2n)(i  +  sk  —  3n)---[i  +sk — (s — i)n] 
by 

(H)  [i+sk^sn\ 

With  these  conventions  series  (13)  beccxnes 

y  =  J'.  +  iyo'*'  +  ^  [I  +  2*  -  «>,>+» 

If  series  (15)  is  convergent,  it  will  furnish  a  root  of  the  three- 
term  equation 

ay*  +  6y*  +  c^o 

for  each  one  of  the  n  values  of  y©. 

ID.  To  test  series  (15)  for  convergency  write  the  first  n  terms  in 
regular  order  in  a  row,  underneath  this  jow  the  succeeding  n  terms 
and  so  on  indefinitely.  The  terms  of  series  (15)  will  now  be  arranged 
in  n  columns  as  follows : 


I 


I 


I 


It 
+ 
.  I 

mm 


k 


+ 


+ 


A 


+ 


+ 


I 

I 


I 


T 


+ 


i 


I 


rm 

+ 


+ 
+ 


I 


i 


I 


1*    4 


I 
CO 


I 

CO 

T 


I 

V* 

i 


I 


+     T 


4- 


4-  <e 

+ 


i 


! 


I 

5 


+ 


I 

+ 

t 


+ 
4- 

f 


^ 


+ 


+ 


1908.] 


IN  INFINITE  SERIES.  117 


This  rearrangement  of  the  terms  of  series  (15)  into  the  n  col- 
umns of  the  table  is  permissible,  inasmuch  as  throughout  this  inves* 
tigation  only  absolute  convergence  is  considered. 

Cauchy's  ratio  test  shows  that  each  one  of  the  n  partial  series 
composed  of  the  terms  in  each  of  the  n  columns  of  the  table  is  con- 
vergent when 

II.  In  like  manner,  if  the  algebraic  functioAs  defined  by  the 
equations 

(2)  ay*»  +  &y*  +  ^'*'  =  o 

(3)  ay*».r  +  6y*  +  c  =  o 

are  expanded  into  power  series  in  x  by  Lagrange's  theorem,  and  if 
X  is  made  unity  in  this  power  series,  it  is  found  that  the  resulting 
infinite  series  are  convergent,  provided 

(18)  -jr^^> 


12.  If  condition  (18)  is  satisfied,  equation  (2)  determines  n — k 
and  equation  (3)  determines  k  roots  of  the  three-term  equation 

Either  condition  (17)  or  condition  (18)  must  be  satisfied,  unless 

If  condition  (19)  is  satisfied,  Raabe's  test  shows  that  the  series 
obtained  from  equations  (i),  (2),  (3)  are  all  convergent. 

13.  The  convergency  conditions  for  equations  (i),  (2),  (3)  may 
be  written  by  following  these  directions : 

(a)  To  the  left  of  the  sign  of  inequality  stands  a  fraction  whose 
numerator  contains  the  coefficient  of  the  middle  term  of  the  three- 
term  equation 

a;/y»_|_ftyf  _|_c=o 

and  whose  denominator  contains, the  product  of  the  coefiicients  of 
the  end  terms,  the  exponent  of  each  coefficient  being  the  difference 
of  the  exponents  in  the  other  two  terms  taken  in  order  from  left 
to  right. 


118  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS      [April  as, 

(6)  The  fraction  to  the  right  of  the  sign  of  inequality  is  obtained 
from  the  fraction  to  the  left  by  replacing  each  coefficient  by  its 
exponent 

(c)  The  sign  of  inequality  is  <  when  the  term  containing  x  is 
between  the  other  two  terms ;  if  the  term  containing  jt  is  an  end  term 
the  sign  of  inequality  is  >. 

14.  The  following  table  exhibits  the  convergency  conditions  for 
the  series  obtained  from  equations  (i),  (2),  (3)  and  the  nimiber  of 
roots  of  the  three-term  equation 

furnished  by  each  one  of  these  series. 

An  |«ii 

(20)  {2)  a)r  +  b^  -^-cx^o      n^k       (    i^  ^ 

The  roots  of  the  three-term  equation  can  always  be  expressed 
in  infinite  series. 

III.  The  Four-term  Equation. 

15.  In  the  four-term  equation 

the  two  terms  from  which  the  factor  x  is  to  be  omitted  can  be 
selected  in  six  diflFerent  ways.    This  gives  rise  to  the  six  equations : 

(21)  a/f^ '\'h'^x-\s,cy^X'\'d'=o 

(22)  fly*-t-6y*  + cy'jT +  djr  =  o 

(23)  ay**jr  +  6y*  +  cy^x  -f  d= o 

(24)  ayi^x  -|-  6y*  •\-cy^  -^dx-^o 

(25)  ay/^x  +  hy^x  +  ry'  +  d  =  o 

(26)  ay*  +  6y*jr +  0^' +  ^'*'  =  o 

Each  one  of  these  six  equations  defines  y  as  an  algebraic  func- 
tion of  X,  The  y  of  equation  (21)  may  be  expanded  into  a  power 
series  in  x  by  any  one  of  the  three  methods  of  articles  7,  8,  9. 
Using  the  symbol  (14)  and  denoting  ( — rf/a)^/"  by  yo,  this  power 
series,  when  x  is  made  unity,  becomes 


x9o8.} 


IN  INFINITE  SERIES. 


119 


+ 

I 

+ 


M 


+ 


M 


tr\ 


+ 


+ 

I 

+ 

+ 


^ 

N 


>5 


M 


tr\ 


+ 


5S 

+ 


II 


IS 

+ 

I 

+ 


^3 


N 


+ 


+    + 


I 


I 


vn 


+ 


120  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS       lAprilts, 

i6.  The  infinite  series  composed  of  the  terms  of  the  left-hand 
column  of  the  value  of  y  is  convergent  when 

and  if  condition  (28)  is  satisfied  this  infinite  series  furnishes  the 
solution  of  the  three-term  equation 

(29)  ay"  +  63;*  +  rf  =  0- 

It  is  found  that  each  one  of  the  infinite  series  composed  of  the 
terms  of  the  respective  coltmins  of  (27)  is  convergent  when  (28)  is 
satisfied.     It  follows  that  (27)  may  be  written  ^ 

(29)      y^X,  +  -^y^X,  +  ^j,y,«X,  +  :^,y-X,+  .-., 

where  X©,  Xi,  X„  Xj,  •••,  stand  for  the  siuns  of  convergent  series. 
If  now  X  is  the  largest  of  the  numbers  Xo,  Xj,  X,,  X,,  •••, 

(30)        ^^  jr(  I +^^.'  +  ^,^,«+^,:v.« +...). 

and  this  last  value  of  y  is  convergent  when 

(31)  ^^•'<^- 

Affecting  both  sides  of  this  inequality  by  the  exponent  n,  this  con- 
vergency  condition  may  be  written 

(32)  •   ^.<«". 

17.  Conditions  (28)  and  (32)  are  sufficient  for  the  absolute  con- 
vergence of  (27).  Condition  (28)  shows  that  the  series  which 
determines  the  roots  of  the  three-term  equation 

(29)  a3f**  +  6y*-j-rf  =  o 

is  found  from 

(33)  ay'''\-by^x-\-d=o. 

The  columns  of  (27)  after  the  first  are  the  qorrections  which 
must  be  applied  to  the  roots  of  the  three-term  equation  (29)  to 
obtain  the  roots  of  the  four-term  equation 


X908.] 


IN  INFINITE   SERIES.  121 


i8.  If  the  two  terms  in  the  second  row  of  (27)  are  interchanged 
and  the  consequent  changes  are  made  throughout  (27),  the  left- 
hand  column  in  the  resulting  value  of  y  is  convergent  if 


«*• 


and  the  entire  expression  for  y  is  convergent  if  in  addition 

(35)  ?^^*<'^"- 

Conditions  (34)  and  (35)  are  sufficient  for  the  absolute  conver- 
gence of  the  new  series  for  y. 

Condition  (34)  shows  that  the  series  which  determines  the  solu- 
tions of  the  three-term  equation  ^ 

(36)  ay»  +  ry  +  d=o 
is  found  from 

(37)  ay»  +  ry^  +  d  =  o. 

This  series  is  the  left-hand  column  of  the  value  of  y. 

Condition  (35)  shows  that  the  series  of  corrections  which  must 
be  applied  to  the  roots  of  the  three-term  equation  (36)  to  obtain 
the  solution  of  the  four-term  equation 

ay*^  _|_  ftyfc  _|_  cy^  -|-  d  =  o 

is  convergent. 

19.  From  equation  (21)  by  omitting  in  succession  each  of  the 
terms  containing  x  are  obtained  the  equations 

(33)  ay»-t-6/jr  +  d  =  o 

(37)  a3f'»  +  fy'jr-f-rf  =  o 

The  convergency  conditions  (28)  and  (34)  may  be  written  from 
equations  (33)  and  (37)  respectively  by  following  the  directions 
(fl),  (fc),  (c)  of  article  13.  The  left-hand  members  of  the  condi- 
tions (32)  and  (35),  together  with  the  character  of  the  signs  of 
inequality,  may  be  written  from  equations  (37)  and  (33)  respec- 
tively by  following  the  same  directions.  The  right-hand  member 
of  conditions  (32)  and  (35)  is  formed  by  writing  the  difference  of 
the  exponents  of  the  two  terms  of  (21)  which  do  not  contain  x  and 


122 


LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS       lAprilas, 


giving  this  difference  an  exponent  equal  to  itself.  It  will  be  found 
that  when  the  sign  of  inequality  is  >  in  convergency  conditions 
corresponding  to  conditions  (32)  and  (35)  the  right-hand  member 
is  the  reciprocal  of  what  it  is  when  the  sign  of  inequality  is  <. 

20.  In  like  manner  two  sets  of  conditions  sufficient  for  the  abso- 
lute  convergence  of  the  infinite  series  giving  the  roots  of  the  four- 
term  equation  obtained  from  each  one  of  the  equations  (21),  (22), 
(23),  (24),  (25),  (26)  may  be  written. 

The  convergency  conditions  for  all  these  infinite  series  may  be 
taken  from  the  following  table,  in  which  the  signs  of  equality  of 
the  limiting  conditions  of  convergence  have  been  omitted. 


(38) 

b^ 

(^ 

ff^  • 

i* 

a^jn-k 

M*-* 

(H-i^*-* 

*«</»-« 

(21)  ay*-}-dj^x+cyjf+d=o 

< 

< 

n* 

(22)  ay^+d}f^+cyx+dx^=o 

> 

> 

{n-k)'-' 

(23)  ay»x-\'fy^+cyx-]-d=ii 

> 

< 

>t* 

(24)  ayx-\-fy^-\'cy+dx—o 

> 

> 

(i-l)'^ 

(25)  ay'x-\'dyx-\-cy-{-d=o 

> 

> 

/« 

(26)  ay*-\'dyx-{-cy+dx=o 

< 

> 

(»-/)--• 

If* 

«• 

(»—/)»-» 

*» 

j^(n—k)^^ 

/'(If-/)"-' 

{i    /)»-*(»    -t)"-» 

fi{i—/)'^ 

In  this  table  the  signs  of  the  two  inequalities  which  constitute 
the  convergency  conditions  of  the  series  obtained  from  the  equa- 
tions (21)  to  (26)  are  placed  to  the  right  of  the  respective  equations. 
The  left-hand  member  of  each  inequality  is  at  the  top  of  the  column 
in  which  the  sign  of  inequality  stands.  The  right-hand  member  of 
one  inequality  must  be  taken  at  the  bottom  of  the  column  in  which 
the  sign  of  inequality  stands ;  the  right-hand  member  of  the  second 
inequality  is  the  expression  at  the  right  of  the  row  in  which  the 
sign  of  inequality  stands  when  the  sign  of  inequality  is  <,  when  the 
sign  of  inequality  is  >  the  right-hand  member  of  the  inequality  is 
the  reciprocal  of  this  expression. 

21.  The.  following  table  exhibits  one  set  of  convergency  condi- 
tions of  the  infinite  series  whfch  give  the  roots  of  the  three-term 
equation 

fly*  +  &y*  +  d  =  o 

together  with  the  equations  from  which  these  series  are  derived  and 


1908.] 


IN  INFINITE  SERIES. 


123 


the  number  of  the  roots  given  by  each  series,  and  also  the  conditions 
sufficient  for  the  absolute  convergence  of  the  series  of  corrections 
which  must  be  applied  to  the  roots  of  this  three-term  equation  to 
obtain  the  roots  of  the  four-term  equation 


(39) 

n 

n-k 

k 

I 
H 

< 
> 
> 

< 

> 

< 

tf 

«" 

I 

>fe* 

kf^n  -  kf-" 

(«  -  kf-" 

22.  The  substitution 

(40)  y=xf*, 

where  ^  is  a  positive  integer,  transforms  the  four-term  equation 

oy*  +  ^y*  -f-  cy '  -f-  d  =  o 
into  the  four-term  equation 

(41)  a^*  +  b^  -f-  cz^*  +  d  =  o. 

The  table  of  convergency  conditions  for  equation  (41)  corre- 
sponding to  table  (39)  is 


(42) 


I  asT+b^x+d^o 
{a:^+bj^+dx=^0 
[aj^x+bz^+d^O 


ns 

ns—ks 

ks 


i— A' 


< 
> 

> 


n      V  « 


\a'd-^') 


< 


(sn)" 


> 


W-') 


\l^{n-k) 
The  three-term  equations 

for  all  values  of  s  have  the  same  convergency  conditions. 


(jw-jyt)*"-*    ('^'^)** 


124  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS      [Apdi.j, 

If  the  inequality 

of  table  (39)  is  not  satisfied,  it  is  always  possible  to  take  s  suffi- 
ciently large  so  that  the  corresponding  inequality 


of  table  (42)  will  be  satisfied. 

In  like  manner,  if  the  inequalities 

_^      «■  _ir-^^        I _£^_ 

a'-d'--  ^  ^(«  -  -t)"-* '     ^-'^-*  ^  («  -ifr)-* '     i'ti'-  <  ^ 

of  table  (39)  are  not  satisfied  simultaneously,  it  is  always  possible 

to  take  s  sufficiently  large  so  that  the  corresponding  inequalities  of 

table  (42) 

a*./-*  ^  ^(«  -  ^)-* '     a'-'c'-"  ^  j"-*(«  -  ^)"-* '     6'd'-'  '^  ^^ 
will  be  satisfied  simultaneously. 

To  the  convei^ncy  conditions  of  table  (42)  must  be  added  the 
limiting  convergency  conditions  obtained  by  replacing  in  the  first 
column  of  inequality  signs  of  table  (42)  each  inequality  sign  by  the 
equality  sign. 

It  follows  that  it  is  always  possible  to  determine  s  so  that  all  the 
roots  of  the  four-term  equation 

(41)  a3^'  +  b^'  +  ce**  +  d  =  o 

may  be  derived  from  the  roots  of  the  three-term  equation 

(42)  as"  +  b^»  -\-d  =  o. 
The  roots  of  the  four-term  equation 

ay"  -|-  ty*  -|-  cy'  -f-  d  ^  o 
from  the  roots  of  equation  (41)  by  substituting  in 

y  =  ^. 
(lile  table  (42)  shows  the  possibility  of  expressing  all  the 


X908.I 


IN  INFINITE  SERIES. 


125 


roots  of  equation  (41)  in  infinite  series,  the  method  of  article  (22) 
requires  the  determination  of  the  ns  roots  of  equation  (41)  to  find 
the  n  roots  of  the  four-term  equation 

ayn  _|.  j/yfc  ^  ^y  ^  d=o. 

This  method  is  therefore  to  be  avoided  in  practice  when  possible. 

The  following  table  exhibits  the  conditions  sufficient  for  the 
absolute  convergence  of  the  infinite  series  which  give  the  roots  of 
the  four-term  equation  obtained  from  the  four  groups  of  equations. 
The  series  obtained  from  each  group  of  equations  determine  all  the 
roots  of  the  four-term  equation.  The  convergency  conditions  must 
be  taken  from  this  table  as  in  article  20,  and  the  limiting  convergency 
conditions  must  be  taken  into  account. 

A  less  inclusive  set  of  conditions  sufficient  for  the  absolute  con- 
vergence of  the  series  which  give  the  roots  of  the  four-term  equa- 
tion derived  from  the  groups  of  equations  of  table  (43)  is  obtained 
by  taking  the  second  member  of  each  inequality  from  the  bottom 
of  the  column  in  which  the  sign  of  inequality  stands. 


(43) 

n 

b* 

b'd'^i 

I      ay*-\-dy^x-\-cyji-\-d—o 

< 

< 

«•» 

(  ay^x-\-by^-\'C)^x-\-d    o 

«— i 

> 

> 

(«-i)-» 

k 

> 

• 

< 

>t* 

III  i  oy''^^y^x~\^c/-\-dx^=Q 
\  ay^x-\-b^X'^c/'\-d—o 

n-'l 

> 

< 

(«-/)-' 

I 

> 

> 

/« 

'  ay^-\-b^-\-c/x'\-dx^=o 

n—k 

> 

> 

(„_^)»-» 

IV     ay^x-\-b^-\-c/-\-dx=o 

k—i 

> 

> 

(,t_/)*-i 

ay^X'\-b^x-\-c^-i^d=o 

I 

> 

> 

/' 

«» 

«» 

(n-n^ 

^ 
/i(j^^/)i^i 

k^(n    >fe)"-* 

/i(«_/)"-i 

(i_/)*-«(«_i)»-» 

It  is  only  when  the  convergency  conditions  of  the  groups  I,  II, 
III,  IV,  together  with  the  corresponding  limiting  convergency  con- 
ditions fail  simultaneously  that  the  use  of  equation  (41)  becomes 
necessary. 

IV.  The  Five-term  Equation. 

24.  In  the  five-term  equation 

ay*  -f  6y*  -j-  ry'  +  dy^  +  /=o 


126  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS      [April  ts, 

the  two  terms  from  which  the  factor  ^  is  to  be  omitted  can  be  selected 
in  ten  different  ways.    This  gives  rise  to  the  ten  equations : 

(44)  'ay*  +  ^y***"  +  cy^^  +  dy^x  +  /=o 

(45)  ay'»  +  6y*4-^y'-*'  +  ^y**^  +  ^-*"=o 

(46)  ay*  +  &y*^  +  ^y'  +  dy^x  4-/^=0 

(47)  ay^  +  ^y*-*"  +  ^y'-*"  +  dy^  4-  /jr=o 

(48)  ay^x  +  fry*  +  cy^  +  (/y**jr  +  lx=^o 

(49)  ay*;r  +  &y*  +  cy^^  +  ^y**  +  ^^=0 

(50)  ay^x  +  &y*^  +  cy^  +  dy^  +  ^^=0 

(51 )  ay";r  +  6y*  +  ^y'-*"  +  rfy**-*"  +  ^= o 

(52)  ay^x  +  6y*^  +  ^y'  +  dy^x  -|-  /= o 

(53)  ay^x  +  &y*^  +  ^y'^  +  dy^  -|-  /= o 

Each  one  of  these  ten  equations  defines  y  as  an  algebraic  func- 
tion of  X  which  may  be  expanded  into  a  power  series  by  any  one  of 
the  methods  of  articles  7,  8,  9. 

25.  The  terms  of  the  power  series  expressing  the  value  of  the 
algebraic  function  defined  by  equation  (44),  using  the  symbol  (14) 
and  placing  yo=  ( — V^)^''">  when  x  is  made  unity,  may  be  arranged 
as  follows: 


I90S.1 


IN  INFINITE  SERIES. 


127 


+ 


•V. 


p» 


+ 


^15 


+ 


•II 
+ 


+ 


II 


+ 
+ 


+ 
+ 


^ 

W 


:i 


W 


+ 


+ 

I 

+ 


e« 


+ 


+ 


m 


+ 


+ 


+ 


+ 


+ 
I 

+ 
+ 


"S 

w 


w 


+ 


+ 

•« 

I 

+ 
+ 


s 

•^ 
w 


nj 


w 


+ 


m 


PROC.  AMKR.  PHIL.  SOC.  XLVII.   1 88  I,  PRINTED  JULY  20,  I908. 


128  LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS       [ApiUas, 

26.  The  first  group  of  terms  of  (54)  is  the  infinite  series  which 
gives  the  solution  of  the  four-term  equation 

obtained  from  the  equation 

ay*  -j-  by^x  +  cy^x  -f-  /  =  o 
provided  the  conditions 


^  «"  ^ 


are  satisfied. 

The  second  group  of  terms  has  the  common  factor 

d 
and  the  successive  groups  of  terms  respectively  the  common  factors 

The  convergency  conditions  of  the  successive  groups  of  terms 
are  identical  with  the  convergency  conditions  of  the  first  group.  It 
follows  that  (54)  may  be  written 

(55)  ^= n  + 1^,^^"+  np^^o*"+  n^^A'"  +  •••. 

where  Fq,  F^,  Fj,  Fg,  F^,  •  ••,  represent  the  sums  of  convergent  infinite 
series. 

If  F  denotes  the  largest  of  the  numbers  F©,  F^,  Fg,  F,,  •••, 

(56)        y^Y[x^^y-^^y^^^^^y^+...). 
The  series  (56)  is  convergent  provided 

(57)  .^•^•''<^- 

If  both  members  of  the  inequality  (57)  are  affected  by  the  exponent 
n,  condition  (57)  becomes 

(58)  ^^^  <  n\ 

The  conditions  sufficient  for  the  absolute  convergence  of  (54)  are 
therefore 


,908.]  IN  INFINITE  SERIES.  129 

ir  «*  r*  ^  d"" 

(59)      -^v^* < -^(n  -  >&)'»-* '     :^V^^^"'     ■^ivi=s^<«"- 

27.  When  the  conditions   (59)  are  satisfied  the  first  group  of 
terms  of  (54)  gives  the  roots  of  the  four-term  equation 

ay "  +  6y*  -f-  ry '  4"  ^ = o 

expressed  in  the  series  obtained  from  the  equation 

ay"  +  h'^x  +  cy^x  +  /= o 

and  the  successive  groups  of  (54)  are  the  series  of  corrections  which 
must  be  applied  to  the  roots  of  this  four-term  equation  to  obtain  the 
roots  of  the  five-term  equation 

ay"  4"  ^y^  +  ^y^  +  ^y^  +  /=o. 

28.  If  in  the  first  row  of  (54)  either  of  the  terms 

c  d 

en  -^^     '     en  -^^ 

is  placed  first  and  the  consequent  changes  in  (54)  are  made,  the 
convergency  conditions  of  the  two  new  series  are  found  to  be 

^  ^  «"  d"" 

,^  ,           ^                      ^                      d""                  nr 
(61)      -jr^  <  n\     -rj^zi  <  «"      ^       


In  the  limiting  convergency  conditions  the  signs  of  inequality  in 
the  first  inequality  of  (59),  in  the  second  inequality  of  (60)  and  in 
the  third  inequality  of  (61)  must  be  replaced  by  the  equality  sign. 

The  conditions  sufficient  for  the  absolute  convergence  of  (54) 
may  be  written  from  equation  (44)  by  the  method  stated  in  article  19. 

In  like  manner  the  conditions  sufficient  for  the  absolute  con- 
vergence of  the  series  obtained  from  equations  (45)  to  (53)  may 
be  written. 

The  convergency  conditions  for  all  these  series  may  be  taken 
from  the  following  table.  The  convergency  conditions  are  taken 
from  the  table  by  the  method  stated  in  article  20,  except  that  the 
right-hand  members  of  two  inequalities  must  be  determined  from 


130 


LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS      [April  .5. 


the  expressions  at  the  right  of  the  row 
stands. 


m 


which  the  sig^  of  inequality 


(62) 


(44) 

(45) 
(46) 

(47) 
(48) 

(49) 
(50) 

(51) 
(5S) 
(53) 


4 

^ 

V 

^ 

> 
^ 

\ 

7 

> 
> 

1 

7 

It 

1 

< 
> 

> 

> 

> 

I 

< 

> 

> 

I 

> 
> 

I 

1 

V 

< 

I 

1 

1 

1 

1 

41 
> 

> 

1 

r 

1 

I 

< 

> 

i 

> 
> 

1: 
T 

It 

r 

1 

< 

4t 

3 

> 

> 

«54 

41 


1 

•^ 


> 
< 
> 


I 


Its 


E 
I 


> 
< 


> 


1 

I 


I 

It 

I 


> 

< 

> 

«5. 

I 

> 

> 
t 

1 

•** 

t. 

s 

> 
< 
> 

I 

;r 

4, 

29.  The  following  table  exhibits  one  set  of  conditions  sufficient 
for  the  absolute  convergence  of  the  infinite  series  which  give  the 
roots  of  the  four-term  equation 

together  with  the  equations  from  which  these  series  are  obtained 
and  the  number  of  roots  given  by  each  series,  and  also  the  conditions 
sufficient  for  the  absolute  convergence  of  the  series  of  corrections 
which  must  be  applied  to  the  roots  of  this  four-term  equation  to 
obtain  the  roots  of  the  five-term  equation 


(63) 


n 


^ 

c^ 

^~/ 

ck 

^» 

^••"^"^W 

akgn—k 

a^gn^l 

ak-lgn-k 

< 

^•%^W^~fW 

ak—m^^k 

< 

> 
> 

< 

> 

< 

> 

nv 

«" 

I 

>fe* 

I 

kk(^n—k)n-k 

(n—k)n-k 

(„_i)*-* 

^* 


g/H^gm-^'i^ 


< 


i^ 


X908.]  IN  INFINITE  SERIES.  131 

30.  The  substitution 

(64)  y  =  z'y 

where  ^  is  a  positive  integer,  transforms  the  five-term  equation 

« 

into  another  five-term  equation 

(65)  ay"*  +  b'f  4-  ry'«  +  dyf^  +  /  =  o. 

An  examination  of  the  table  of  convergency  conditions  for  equa- 
tion (65)  corresponding  to  table  (63),  shows  that  it  is  always  pos- 
sible so  to  determine  s  that  the  convergency  conditions  for  the  series 
obtained  from  the  equation 

(66)  ay""^  +  b'f^x  +  cy^^x  -f-  dy'^'x  +  /  =  o 

or  from  the  pair  of  equations 

(67)  ay*«  +  6y*«  +  cy^*x  +  dy^^x  4-  /;r  =  o 

(68)  ay'^'x  +  by^  +  cy^'^x  +  dy'^^x  +  /  =  o 

are  satisfied.  Hence  it  is  always  possible  to  determine  all  the  roots 
of  a  five-term  equation  by  means  of  series. 

31.  The  method  of  article  30  requires  the  determination  of  the  ns 
roots  of  equation  (65)  in  order  to  find  the  n  roots  of  the  five-term 
equation 

ay"  +  ^y*  +  ^y'  +  ^y^ + ^=o. 

The  use  of  this  method  becomes  necessary  only  when  the  conver- 
gency conditions  of  the  seven  groups  of  equations  of  the  following 
table,  together  with  the  corresponding  limiting  convergency  condi- 
tions fail  simultaneously. 

The  convergency  conditions  must  be  taken  from  this  table  as  in 
article  20. 

A  less  inclusive  set  of  congruency  conditions  may  be  taken  from 
this  table  as  in  article  23. 


132 


LAMBERT— SOLUTION  OF  ALGEBRAIC  EQUATIONS       [April  as. 


LI    I  li    II    II 


tP 


«« — u^ 


M v^tK^ 


A  A 


AA 


V      A 


A 


A 


il 


A       V 


VAAA        AA 


^p 

V           A 

\-^^ 
^ 

V     A           A           AAA 

t-uq 

AV           AAAV        AA 

ut-uq 

A           V     A                  A 

,««(^— «)«»_,( w—y) 

V 


A  A 


(w — u)^m 


uU 


V 


A  A 


AA     A 


A 


!-«(/— ««W 


MM 


VAA 


A 


A 


} 


J      I      S 


o  o  o 

II  II  II 


o  o   o 

II  II  II 


o  o  o 

II  II  II 


o  o  o  o 


(S  IS  IS  X 

4-  +  + 7" 

It     H    I     <^ 

-I-  4-   '    >« 

,   +-r  + 

'      >«     H     H 

^  «*  «^  Q^ 


^?  -  US  S 


t9o8.J  IN  INFINITE  SERIES.  133 

V.   Conclusion. 
32.  In  the  algebraic  equation  of  t  terms 

/(3')=o 

the  two  terms  from  which  the  factor  ;r  is  to  be  omitted  can  be 
selected  in 


ways.  Each  one  of  the  resulting  equations  defines  y  as  an  algebraic 
fimction  of  x,  and  each  algebraic  function  of  x  can  be  expanded 
into  a  power  series  in  x  by  the  methods  used  to  obtain  the  corre- 
sponding expansions  for  the  three-,  four-  and  five-tenfl  equations. 
When  X  is  made  unity  in  these  power  series  the  resulting  series 
become  the  roots  of  the  /-term  equation  and  a  table  of  convergency 
conditions  for  these  series  analogous  to  tables  (20),  (38),  (62)  can 
be  set  up.  In  fact,  this  table  may  be  written  mechanically  by  fol- 
lowing the  directions  of  article  19. 

33.  If  in  the  /-term  equation  the  substitution 

is  made,  a  table  of  convergency  conditions  analogous  to  tables  (39), 
(63)  can  be  set  up,  and  the  value  of  s  can  be  determined  so  that 
this  table  of  conditions  shows  that  it  is  possible  to  obtain  all  the 
roots  of  the  transformed  equation  from  the  series  derived  either 
from  the  equation  in  which  x  is  omitted  from  the  'first  and  last 
terms,  or  from  the  two  equations  in  which  x  is  omitted  from  the 
first  and  second,  and  from  the  second  and  last  terms  respectively. 
The  roots  of  the  given  equation  are  then  found  from  the  roots  of 
the  transformed  equation  by  substituting  in 

34.  Finally,  tables  of  convergency  conditions  analogous  to  tables 
(43),  (64)  can  be  set  up  for  the  /-term  equation,  and  it  is  necessary 
to  use  the  transformed  equation  only  when  the  convergency  condi- 


134  MINUTES. 


[May  IS, 


tions  of  all  the  groups  of  this  table,  together  with  the  corresponding 
limiting  convergency  conditions,  fail  simultaneously. 

35.  It  follows  that  all  the  roots  of  an  algebraic  equation  of  any 
number  of  terms,  that  is,  of  any  algebraic  equation,  can  be  expressed 
in  infinite  series  by  the  method  of  this  investigation. 

Lehigh  IJNiVERsrry,  Bethlehem,  Pa., 
April  2,  1908. 


Stated  Meeting  May  i,  1908, 

Treasurer  Jayne  in  the  Chair. 

Dr.  Martin  G.  Brumbaugh,  a  newly  elected  member,  was  pre- 
sented to  the  chair,  and  took  his  seat  in  the  Society. 

Letters  Arere  read,  accepting  election  to  membership  from 

Martin  Grove  Brumbaugh,  Ph.D.,  Philadelphia. 

Walter  Bradford  Cannon,  A.M.,  M.D.,  Boston,  Mass. 

James  Christie,  Philadelphia. 

Edward  Washburn  Hopkins,  Ph.D.,  LL.D.,  New  Haven,  Conn. 

Josiah  Royce,  Ph.D.,  LL.D.,  Cambridge,  Mass. 

Jacob  G.  Schurman,  Ph.D.,  Ithaca,  N.  Y. 

Edward  Anthony  Spitzka,  M.D.,  Philadelphia. 

Robert  Williams  Wood,  Ph.D.,  Baltimore. 

Mr.  R.  H.  Mathews  presented  some  "Notes  on  Australian 
Laws  of  Descent." 

Professor  Albert  A.  Michelson,  of  Chicago,  was  unanimously 
elected  a  Vice-President  to  fill  the  unexpired  term  of  Professor 
George  F.  Barker,  resigned. 

Stated  Meeting  May  15,  ipo8. 
Curator  Doolittle  in  the  Chair. 

Letters  were  read  accepting  membership  from 
William  HaUock,  Ph.D.,  New  York  City. 
Leonard  Pearson,  M.D.,  Philadelphia. 
Charles  Henry  Smyth,  Ph.D.,  Princeton,  N.  J. 
John  Robert  Sitlington  Sterrett,  Ph.D.,  Ithaca,  N.  Y. 
Ernest  Nys,  Brussels. 


X908.3  IN  INFINITE  SERIES.  135 

From  Professor  Albert  A.  Michelson  accepting  election  to  the 
Vice-Presidency  to  fill  an  unexpired  term. 

From  the  Committee  of  Organization  of  the  Third  Congres 
International  de  Botanique,  announcing  that  the  Congress  will  be 
held  at  Brussels  from  May  14-22,  1910,  and  inviting  the  Society  to 
be  represented  by  delegates. 

Dr.  H.  M.  Chance  read  a  paper  on  "  The  Origin  of  Bombshell 
Ore"  (see  page  135),  which  was  discussed  by  Mr.  Sanders,  Mr. 
Jayne  and  Professor  Doolittle. 


THE  ORIGIN   OF  BOMBSHELL  ORE. 

By  H.  M.  chance. 
(Read  May  15,  1908.) 

The  term  "  bombshell "  ore  is  applied  by  miners  and  iron-masters 
to  hollow  masses  of  limonite — brown  hematite — ^which  sometimes  are 
round  or  oval  but  more  commonly  are  of  any  irregular  shape.  The 
"  bombs  "  may  contain  water,  clay,  sand,  quartz,  flint,  pyrite,  siderite, 
sandstone  or  decomposed  slate,  or  may  be  entirely  empty.  Geolo- 
gists usually  speak  of  such  ore  as  nodular  or  concretionary.  A 
careful  examination  of  the  literature  of  ore  deposits  and  especially 
of  that  relating  to  the  genesis  of  limonite  ores  fails  to  disclose  a 
satisfactory  explanation  of  its  origin  of  mode  of  formation.  By 
many  it  is  assumed  to  be  similar  in  origin  to  silicious  geftdes,  which 
are  supposed  to  be  formed  by  the  deposition  of  silica  or  silicates 
upon  the  walls  of  cavities,  while  others  describe  it  as  of  "concre- 
tionary "  origin  without  attempting  to  explain  the  process  of  forma- 
tion or  the  manner  in  which  it  has  occluded  the  variety  of  materials 
which  are  found  in  the  interior  of  different  specimens  from  the  same 
locality.  That  it  is  not  of  concretionary  origin  is  evident  uppn  even 
cursory  consideration,  for  concretions  are  masses  of  material  ar- 
ranged in  concentric  layers  around  a  central  nucleus.  The  latter 
may  be  a  grain  of  sand,  a  pebble,  fossil,  or  any  substance  around 
which  (as  a  core)  the  concretion  forms,  growing  from  the  center 
by  the  successive  addition  of  concentric  rings.  Concretions-  are 
perhaps  merely  symmetrical  segregations. 

The  peculiarities  of  this  ore  are  well  described  by  Professor  T. 
C.  Hopkins,  Bull,  Geol  Soc.  Am.,  1890,  Vol.  11,  p.  477,  etc.,  as 
follows : 

"  Nodular  ore  consists  of  irregularly  rounded  masses,  varying  in  size 
from  a  fraction  of  a  pound  to  several  hundred  pounds  in  weight.  The 
masses  are  frequently  hollow,  but  some  enclose  a  rounded  or  sub-angular 
rock  fragment,  which  is  sometimes  sandstone,  .  .  .  sometimes  chert,  some- 
times slate,  and  sometimes  clay.    Some  shells  are  filled  with  clay  and  sand, 

136 


1908.]  CHANCE— THE  ORIGIN  OF  BOMBSHELL  ORE.  137 

and  workmen  report  finding  many  of  them  filled  with  water.  Some  are 
filled  with  clay,  which  still  retains  the  laminated  structure  and  appearance 
of  the  original  slate  from  which  the  clay  was  derived,  furthermore,  the 
slaty  structure  was  found  to  extend  through  the  ore  shell,  which  showed, 
besides  the  plain  lamination  of  slate,  a  faint  concentric  structure  as  well. 
.  .  .  While  only  one  shell  was  found  still  retaining  the  laminations  of  the 
clay,  there  were  many  others  containing  clay  and  sand.  Some  of  the  shells 
were  but  thin  crusts,  while  others  were  quite  thick,  almost  solid;  some  have 
a  smooth,  velvety  or  bright  mammillated  inner  surface,  frequently  coated 
with  manganese  oxide.  In  some  instances  the  lining  of  the  shell  is  covered 
with  many  small  stalactites  of  ore.  .  .  .  Many  of  the  shells  arc  lined  with 
a  dense  fibrous  layer,  often  an  inch  or  more  in  thickness.  .  .  .  The  thinner 
shells  have  all  been  broken,  and  we  see  only  the  fragments  of  them  in  the 
clay-ore  masses.  This  shell  form  of  ore  .  .  .  forms  an  appreciable  part 
of  the  ore  body  in  many  cases.  The  small,  irregular,  nodular-like  pieces  of 
ore,  commonly  knows  as  shot  ore,  are  presumably  closely  related  in  origin 
to  the  shells.  .  .  ." 

The  inner  wall  of  many  bombs  consists  of  a  hard,  bright,  brown 
or  jet-black,  glazed  surface,  curved,  rounded  or  botryoidal.  This  is 
frequently  described  as  a  manganese  coating,  but  is  doubtless  a  film 
of  iron  or  manganese  silicate.  Occasionally  the  interior  or  a  part 
of  the  interior  is  lined  with  a  layer  of  extremely  hard,  flinty,  liver- 
colored  iron  silicate,  or  with  quartz  crystals  or  chalcedony,  and  the 
same  silicate  frequently  forms  a  considerable  portion  of  the  body  of 
the  shell  or  of  its  outer  layers,  but  generally  the  shell  is  composed 
of  high-grade  limonite,  of  a  fibrous  structure,  especially  in  those 
layers  forming  the  inner  lining  of  the  shell. 

These  peculiarities  are  satisfactorily  accounted  for  neither  by  the 
theory  that  these  ore  masses  owe  their  origin  to  concretionary  action, 
nor  by  that  which  assumes  the  direct  deposition  of  ferric  hydrate 
upon  the  interior  of  rock  (limestone?)  cavities.  They  may,  how- 
ever, be  explained  by  assuming  that  the  bombs  are  the  residual 
masses,  remaining  after  oxidation,  of  iron  sulphides  or  carbonates 
containing  sand  or  clay  or  both  in  varying  proportions. 

If  the  material  from  which  this  ore  is  formed  consists  of  sand- 
stone, or  of  sandy  slate,  or  of  clay  slate,  impregnated  more  or  less 
completely  with  pyrite  or  siderite,  the  formation  of  bombshells,  con- 
taining just  such  materials  as  are  found  in  these  shells,  may  be 
readily  explained,  especially  if  the  iron  impregnation  be  in  the  form 
of  pyrite  or  marcasite,  that  is,  FeSj. 


138  CHANCE— THE  ORIGIN  OF  BOMBSHELL  ORE.  [May  15, 

If  such  sandstone  or  slate,  is  broken  and  fissured  by  faulting 
and  crushing,  and  by  the  development  of  cleavage  planes,  oxidation 
by  percolating  waters  will  proceed  along  the  joints  or  planes  which 
form  the  channels  through  which  these  waters  circulate,  and  in  each 
fragment  of  the  mineralized  rock  oxidation  will  commence  upon  the 
outside  and  progress  towards  the  center. 

In  this  way  on  outer  skin  or  shell  of  limonite  first  forms  on  the 
outside  of  the  fragment,  for  if  the  iron  be  present  as  pyrite  or  mar- 
casite  while  some  of  it  may  be  removed  as  ferrous  sulphate,  this  salt, 
if  formed,  may  immediately  be  oxidized  and  precipitated  in  situ  as 
ferric  hydrate.  The  sulphuric  acid  formed  by  the  oxidation  of  the 
remaining  molecule  of  sdlphur  will  attack  and  decompose  the  clay 
of  the  gangue,  removing  the  bases  as  sulphates  in  solution;  the 
silicic  acid  also  escaping  in  solution,  or  combining  with  iron  oxides 
to  form  iron  silicates,  remains  as  an  integr^  part  of  the  ore. 

If  clay  be  present  in  large  quantity  a  portion  will  remain  unde- 
composed  in  the  center  of  the  bomb,  together  with  all  of  the  sand 
originally  present  in  the  gangue. 

Hence,  if  the  original  pyritic  material  has  a  clayey  (slate)  gangue, 
bombs  may  form  containing  no  residual  matter,  or  containing  more 
or  less  clay;  if  the  gangue  be  sand  and  clay  (arenaceous  slate),  the 
sand  only,  or  sand  and  clay  may  remain ;  if  the  gangue  be  sand  only, 
some  of  this  will  remain  as  an  impurity  in  the  limonite  forming  the 
body  of  the  shell,  and  some  as  a  partial  filling  of  the  interior  of  the 
bomb. 

It  is  now  well  known  that  pyrite  (of  marcasite)  oxidizing  under- 
ground, whether  by  waters  carrying  free  oxygen,  or  by  waters  con- 
taining no  uncombined  oxygen,  or  by  reactions  involving  hydrolysis, 
does  not  behave  in  the  same  way  as  when  oxidized  by  exposure  to  the 
air  above-ground.  One  of  the  most  common  reactions  above-ground 
is  that  in  which  sulphur  is  set  free,  often  written : 

FeSa  +  40  =  FeSO^  +  S, 

but  this  rarely  occurs  beneath  the  surface,  for  the  gossans  of  pyritic 
veins  seldom  carry  free  sulphur,  although  there  are  a  few  noted 
examples  in  which  large  deposits  of  sulphur  are  found  between  the 
surface  and  the  unoxidized  portions  of  such  veins. 


x9o8].  CHANCE— THE  ORIGIN  OF  BOMBSHELL  ORE.  139 

In  the  absence  of  oxygen,  carbonates  in  solution  may,  as  shown 
by  Dr.  N.  H.  Stokes,  completely  oxidize  the  iron  of  pyrite  or  mar- 
casite  thus : 

8FeSa  +  isNa^COa  =  4Fe,03  +  ^Na^S  +  Na^SaOa  +  15CO,, 

and  tmder  proper  conditions  of  temperature  and  pressure  the  ferric 
oxide  thus  formed  may  be  deposited  as  hydrate ;  but  these  reactions 
do  not  satisfy  the  observed  conditions  and  it  seems  more  probable 
that  oxidation  near  the  surface  has  proceeded  as  indicated  by  some 
of  the  following  reactions : 

FeSa  +  HaO  +  7O  =  FeSO^  +  H^SO^ ; 

3FeS,  +  8H,0  +  22O  =  Fe^Oa,   3HaO  +  FeSO,  +  sH^SO, ; 

2FeS04  +  5H,0  +  O  =  Fe^O,,   sH^O  +  2H,S04 ; 

2FeS,  +  7H,0  +  150  =  Fe^Oa,   SH^O  +  4H,SO, ; 

2FeS2  +  4HjO  +  15O  =  Fe^Oa  +  4HaSO, ; 

2FeS04  +  2H,0  +  O  =  Fe^Oa  +  2HaS04 ; 

6FeSO,  +  30  =  Fe,Oa  +  2Fe,(SO,)a ; 

6FeSO,  +  3H,0  +  30=  Fe,Oa,   3H2O  +  2Fe3  ( SO,)  a ; 

2Fe2(SOj8  +  2FeS2  +  4HjO  +  i20  =  6FeS04  +  4H2SO4. 

The  sulphuric  acid  having  been  formed  in  direct  contact  with  the 
gangue,  it  is  reasonable  to  suppose  that  it  must  at  once  attack  any 
clay  or  other  deoxnposable  material,  and  the  removal  of  th^  soluble 
silicates  and  silicic  acid  by  transfusion  through  the  walls  of  the  bomb 
is  readily  pictured.  It  is,  however,  possible  that  the  colloidal  silicic 
acid  may  be  retained,  and  further  that  it  may  perhaps  often 
be  set  free  in  a  gelatinous  condition.  This  latter  hypothesis  may 
account  for  the  frequent  presence  in  such  ores  of  a  skeleton  of 
amorphous  silica  which  appears  to  completely  ramify  some  parts  of 
the  limonite. 

If  the  oxidation  proceed  according  to  these  equations,  the  succes- 
sive additions  of  layers  of  limonite  to  the  interior  of  the  shells  is 
doubtless  due  to  the  further  oxidation  of  the  ferrous  sulphate  as 
above  shown,  the  oxidation  of  the  solution  occurring  at  or  in  the  wall 
of  the  shell  where  the  solutions,  in  escaping  by  transfusion  through 
the  walls  of  the  shell,  are  met  by  oxidizing  waters  transfusing 
towards  the  center  of  the  shell.    Under  such  conditions  the  ferric 


140  CHANCE— THE  ORIGIN  OF  BOMBSHELL  ORE.  [May  15, 

hydrate  would  be  deposited  in  the  pores  of  the  shell  or  upon  its  inner 
surface. 

In  attempting  to  picture  these  reactions  and  their  results,  it  is 
important  to  remember  the  extremely  slow  rate  at  which  oxidation 
proceeds  under  such  conditions.  Even  at  the  surface  where  decom- 
position is  comparatively  rapid,  the  oxidation  of  pyrite  appears  to 
progress  at  a  very  slow  rate,  perhaps  not  exceeding  an  inch  or  a 
few  inches  in  depth  in  several  hundred  years. 

If  the  iron  be  present  as  carbonate,  a  precisely  similar  series  of 
reactions  may  be  conceived,  in  which  carbonic  acid  transposes  the 
silicates,  freeing  silicic  acid  and  removing  the  bases  as  soluble 
carbonates. 

Other  observers  have  noted  the  occasional  presence  of  a  central 
core  of  siderite  or  pyrite  in  bombshell  ore,  but  have  generally  attrib- 
uted the  presence  of  such  cores  to  concretionary  action  and  replace- 
ment by  sulphates  (accompanied  by  reduction  to  sulphide)  or 
carbonates  in  solution. 

The  foregoing  theory,  advanced  to  account  for  the  origin  of 
bombshell  ore,  is  based  upon  a  study  of  these  deposits  dating  back 
to  1885 — When  the  writer  was  personally  engaged  in  mining  brown- 
hematite  ore — ^and  upon  examinations  of  many  specimens  which 
show  more  or  less  clearly  the  character  of  the  original  material  from 
which  such  ore  is  formed.  It  will  form  an  integral  part  of  a  broader 
statement,  extending  the  application  of  this  theory  to  the  genesis 
of  limonite  ores,  and  including  a  discussion  o{  the  original  sources 
of  the  iron,  methods  of  mineralization,  and  subsequent  decomposition 
and  precipitation. 


I 


MAGELLANIC    PREMIUM 

FotmDED  IN  1786  BY  |OHN  HYACINTH  DE  MAGBLLAN,  OF  LoMDON 

1909 

THE  AMERICAN  PHILOSOPHICAL  SOCIETY 

Held  at  PhiladeiphiAi  for  Promoting  Useful  Knowledge 

ANNOUNCES  THAT  IN 

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Vol.  XLVII.  May-August,  1908.  No.  189. 


CONTENTS. 


The  Sign    and    Name    for    Planet    in    Babylonian,     By  Morris 

Jastrow,  Jr 141 

Further  Researches  on  the  Physics  of  the  Earth,  and  Especially  on 
the  Folding  of  Mountain  Ranges  and  the  Uplift  of  Plateaus  and 
Continents  Produced  by  Movements  of  Lava  Beneath  the 
Crust  Arising  from  the  Secular  Leakage  of  the  Ocean  Bottoms. 
By  T.  J.  J.  See  157 

The  Absorption  Spectra  of  Neodymium  Chloride  and  Praseodymium 
Chloride  in  Water,  Methyl  Alcohol,  Ethyl  Alcohol  and  Mix- 
tures of  these  Solvents.  By  Harry  C.  Jones  and  John  A. 
Anderson 276 

Preliminary  Report  upon  a  Crystallographic  Study  of  the  Hemo- 
globins: a  Contribution  to  the  Specificity  of  Corresponding 
Vital  Substances  in  Different  Vertebrates.  By  Edward  T. 
Reichert  and  Amos  P.  Brown 298 

Influence  of  Preservatives  and  other  Substances  added  to  Foods  upon 

Health  and  Metabolism.     By  Harvey  W.  Wiley 302 

The  Humming  Telephone.     By  A.  E.  Kennelly  and  Walter  L. 

Upson 329 

On  the  After-images  of  Subliminally  Colored  Stimuli.     By  Edward 

Bradford  TrrcHENERand  William  Henry  Pvle 366 

On  the  Classification  of  the  Cetacea.     By  Frederick  W.  True 385 


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PROCEEDINGS 

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AMERICAN  PHILOSOPHICAL  SOCIETY 

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FOR  PROMOTING  USEFUL  KNOWLEDGE 

Vol.  XLVII  May-August,  1908.  No.  189. 

• 

THE  SIGN  AND  NAME  FOR  PLANET  IN  BABYLONIAN. 

y 

By  morris  JASTROW,  Jr. 

(Read  April  25,  igoS.) 

Kpgler  begins  his  valuable  work  on  Babylonian  astronomy^  with 
a  discussion  of  the  ordinary  name  for  planet  in  Babylonian,  namely, 
bibbu,  and  for  which  the  ideographic  designation  is  Lu-Bat.*    He 

*"Stemkunde  und  Stemdicnst  in  Babel"   (Miinstcr,  1907),  I.,  pp.  7-9. 

'That  this  combination  is  used  for  planet  in  general  follows  from  such 
passages  as  (i)  Thompson,  "Reports  of  the  Magicians  and  Astrologers," 
No.  112  Rev.  7;  236  B  Rev.  4,  where  Lu-Bat  occurs  with  the  plural  sign  to 
designate  the  planets  in  general ;  See  also  nos.  88  Obv.  4  and  Rev.  i ;  8p  Rev.  6; 
loi  Obv.  s;  103  Obv.  6,  Rev.  7;  163  Obv.  4;  167  Rev.  i;  172  Rev.  i  and  3; 
17s  Obv.  4;  200  Rev.  s;  216  Rev.  j;  218  Obv.  i;  218  A  Obv.  5;  219  Obv.  i; 
220  Obv.  I ;  222  Obv.  i ;  223  Obv.  i ;  224  Obv.  3 ;  225  Obv.  4 ;  229  A  Obv.  i,  2, 
4;  232  Rev.  I ;  234  Obv.  3;  234  A  Rev.  i ;  235  Obv.  11 ;  244  C  Obv.  6,  where 
Lu-Bat  is  used  for  planet  in  general.  It  is  to  be  noted,  however,  that  the 
only  planets  which  are  regularly  designated  by  means  of  Lu-Bat  are 
Mercury  (Lu-Bat  Gu-Ud)  and  Saturn  (Lu-Bat  Sag-Us).  So  in  the 
famous  list  of  planets  IIR  48 ;  50-54  a-b  and  IIIR  57,  No.  6, 65-67,  and  Thomp- 
son, /.  c,  passim,  though  occasionally  even  in  the  case  of  these  two  planets  the 
element  Lu-Bat  is  omitted,  e.  g.,  Thompson,  /.  c,  Nos.  105  Obv.  8;  215  Obv.  i ; 
217  Obv.  i;  223  Obv.  4;  228  Obv.  i;  (Gu-Ud)  and  167  Rev.  4  (Sag-Us). 
Further  references  in  Kugler,  /.  c,  p.  12.  Occasionally  also  Mars  is  desig- 
nated as  Lu-Bat  Dm  instead  of  (il)  ZAL-BAT-(a-nu),  so  e,  g.,  Thompson  No. 
146  Rev.  4-6,  and  195  Rev.  1-2,  where  in  both  cases  a  gloss  Lu-B at  Dnt=  (il) 
Zal-Bat  (-a-nu)  furnishes  the  proof  for  the  identification.  In  the  later 
period  (after  c.  400  *B.  C.)  Saturn  is  designated  as  GI  and  Mars  as  AN. 
See  Kugler  /.  c,  p.  12,  including  the  note  on  that  page. 

PROC.  AMER.  PHIL.  SOC.  XLVII.   1 89  J,  PRINTED  SEPTEMBER  1 9,   I908. 


142  JASTROW— THE  SIGN  AND  NAME  lAprilas, 

accepts  the  interpretation  proposed  as  long  ago  as  1890  by  Jensen* 
for  the  ideographic  compound  as  "  frei  weidendes,  abseits  weidendes 
Schaf."  This  view  rests  on  the  identification  of  the  first  sign  Lu 
as  "  sheep,"  while  the  second  is  taken  in  the  sense  of  "  to  remove,"* 
the  combination  thus  expressing  the  movement  of  the  planets,  like 
sheep  that  wander  away  from  the  flock.  That  Jensen  was  right  in 
his  explanation  of  the  first  element  as  "  sheep  "  follows  from  various 
considerations,  among  which  the  testimony  of  the  lexicographical 
list  IIR.  6,  4  c-d  by  itself,  Lu-Bat  =  bi-ib-bu,  is  decisive,  since  in 
the  same  list  Lu-Igi  is  explained  as  lu'li-mu  (1.  8)  "  ram  "  or  "  bell- 
wether "  and  ^u  is  the  common  ideogram  for  immcru  the  ordinary 
term  for  *'  sheep.""  In  addition  we  have  the  equation  IIR.  39,  No. 
5,  62  a-b  (t7)  bi-ib'bu  =  (♦/)  Lu-Bat. 

Jensen's  explanation,  however,  of  the  second  element  is  not  satis- 
factory. In  the  first  place  the  equation  Bat  =  nisu  ("to  remove ") 
does  not  represent  the  most  common  value  of  the  ideograph  in  ques- 
tion, for  the  various  meanings  of  which  it  seems  more  reasonable  to 
start  from  the  fundamental  notion  of  "coming  to  completion,"* 
whence  we  have  the  further  development  in  two  directions:  (i) 
"coming  to  an  end"  (gamaru,  katu,  Br.  Nos.  1499,  1512,  etc.). 
"  closing  up  "  (sakku,  sikeru  Meissner,  Assyr.  Ideogramme,  Nos. 
869-872);  "removing"  (nisii,  Br.  No.  1525);  "growing  old" 
(labaru,  Br.  No.  1515) ;  "  die  "  (matu,  etc.,  Br.  Nos.  1517-19,  1527, 
1533)  J  "  set  at  rest "  (pasahu,  Br.  No.  1528) :  (2)  "  Completion 
in  the  sense  of  "  fullness  "  and  "  vitality,"  consequently,  "  life 
(balatu,  Br.  No.  1494)  ;  "being"  (basu,  Br.  No.  1495)  J  "blood 


If 
If 


'Kosmologie  der  Babylonier,  p.  pg.  Hommel  (Aufsatzc  und  Abhand- 
lungen  p.  379)  thinks  that  the  designation  bibbu  which  he  takes  as  "  ram  "  is 
an  allusion  to  the  "  solar  "  character  of  the  planets,  but  this  is  even  less  plau- 
sible than  Jensen's  explanation. 

*Cf.  Briinnow,  No.  1525  (BAT  =  nisu). 

•  See  Muss-Amolt,  Assyr.  Diet.,  p.  61  b.  Note  also  that  in  the  list 
IIR.  6,  5-8  we  have  the  group  bi-ib-bu,  a-tu-du  ("he-goat"),  sap-pa-ru 
("mountain-goat")  and  lu-li-mu. 

•  We  must  bear  in  mind  as  Thureau-Dangin,  "  Recherches  sur  Y  Originc 
de  TEcriture  cuneiforme,"  No.  11,  has  pointed  out,  that  two  originally  dis- 
tinct signs  have  been  confounded  in  Bat,  so  that  all  meanings  associated,  e.  g., 
with  pM  (Br.  No.  1529)  must  be  referred  to  No.  278  (p.  45)  and  explained 
accordingly. 


,9o8.]  FOR   PLANET  IN  BABYLONIAN.  143 

(damu,  Br.  No.  1503),  and  "  rule  "  (belu,  etc.,  Br.  No.  1496;  Meiss- 
ner,  No.  856),  as  manifestations  of  vitality  and  power  as  well  as 
"strong"  (ikdu,  Meissner,  No.  851),  "protect"  (emu,  Meissner, 
No.  853),  etc.  The  idea  of  "  removing  "  falls,  therefore,  in  the  cate- 
gory of  a  secondary  or  tertiary  derivative  from  the  fundamental  value 
of  the  sign  Bat.  In  the  second  place,  it  is  rather  a  violent  transition 
from  the  sense  of  "  removing  "  to  that  of  "  pasturing  by  itself  "  and 
the  like.  Nor  does  the  metaphor  introduced  in  the  Babylonian  crea- 
tion epic^  (Tablet  VII.,  iii,  ed.  King)  where  the  stars,  or  rather 
the  gods,  are  compared  to  sheep  under  the  guidance  of  Marduk 
strengthen  the  conclusion  that  the  planets  are  sheep  that  "  pasture 
aside  "  from  the  stars  in  general,  since  the  passage  does  not  refer 
specifically  to  the  planets.  This  passage,  as  well  as  the  others  ad- 
duced by  Kugler  (/.  c,  p.  7),  merely  justifies  the  interpretation  of 
the  first  element  in  Lu-Bat  as  "  sheep."  For  the  second  element  we 
must  start  from  the  much  more  common  meaning  attaching  to  the 
sign  in  question,  namely,  "  dead  "  (mttu).  The  Babylonians  them- 
selves had  this  equation  in  mind  when  they  explained  Lu-Bat  as  mus- 
mit  bu'lim,  "causing  cattle  to  die"  (VR,  46,  No.  i,  rev.  41)  even 
though  this  explanation  is  to  be  regarded  as  a  fanciful  one.* 

Taking  the  two  signs  as  they  stand,  the  simplest  explanation  is 
to  interpret  them  as  "  dead  sheep  "  in  the  sense  of  a  sacrificial  animal. 
To  the  question  which  now  arises,  what  connection  is  there  between 
the  planets  and  "  dead  sheep,"  the  divination  texts,  I  venture  to  think, 
furnish  a  satisfactory  answer. 

II. 
On  the  basis  of  recent  researches,'  we  must  distinguish  in  Baby- 

*  See  Kugler,  /.  c,  p.  7. 

•Recognized  as  fanciful  by  Jensen,  Kosmologie,  p.  96.  Kugier*s  attempt 
(/.  c.)  to  reconcile  this  e;cplanation  with  the  interpretation  offered  in  astro- 
logical texts  whereby  certain  phenomena  connected  with  the  planets  prognos- 
ticate death  is  very  artificial  and  encounters  a  fatal  objection  from  the  con- 
sideration that  the  prognostication  of  death  in  one  form  or  other,  is  a  com- 
mon interpretation  of  omens,  indeed  one  of  the  commonest  See  examples  in 
Jastrow,  "  Religion  Babyloniens  und  Assyriens,"  II,.  pp.  261,  298,  328,  329,  331, 
333,  343,  etc. 

•  See  Jastrow,  /.  c,  pp.  212  f .,  and  various  papers  by  the  writer  as,  e.  g., 
"  Signs  and  Names  of  the  Liver  in  Babylonian  "  (Zeitschrift  fiir  Assyr.  XX., 
p.  Ill  f.),  "The  Liver  in  Divination  and  the  Beginnings  of  Anatomy"  (Uni- 
versity of  Pennsylvania  Medical  Bulletin,  January,  1908). 


144  JASTROW— THE  SIGN   AND  NAME  [April  25, 

Ionian- Assyrian  methods  of  divining  the  future  two  classes:  (i) 
what  we  may  call  voluntary  divination,  and  (2)  involuntary  divina- 
tion. The  characteristic  feature  of  voluntary  divination  lies  in  delib^ 
erately  seeking  out  some  object  by  means  of  which  an  answer  to  a 
specific  question  regarding  the  future  or  the  outcome  of  an  under- 
taking, a  sickness  or  what-not  is  furnished.  The  signs  furnished  by 
the  liver  of  an  animal  selected  as  a  sacrifice  belong  to  this  category ; 
likewise  the  observation  of  the  flight  of  birds  sent  out  for  the  pur- 
pose of  securing  omens,  the  throwing  of  arrows  before  the  image 
of  a  deity  and  the  like.  Involuntary  divination,  on  the  other  hand, 
is  concerned  with  the  attempt  to  interpret  signs  that  force  themselves 
on  our  notice,  such  as  phenomena  connected  with  the  sun,  moon, 
planets  and  stars,  the  movements  of  clouds,  earthquakes  and  storms ; 
the  actions  of  animals — dogs,  snakes,  locusts,  birds,  etc.,  that  one 
happens  to  encoimter  and  all  the  unusual  or  significant  happenings 
and  accidents  in  human  life,  while  dreams  form  a  special  subdivision 
in  this  class  of  involuntary  divination.  We  might  for  the  sake  of 
convenience  distinguish  the  signs  furnished  by  voluntary  divination 
as  "  omens  "  and  those  of  involuntary  divination  as  "  portents,"  but 
however  we  may  distinguish  them,  the  recognition  of  these  two  dis- 
tinct classes  is  fundamental  to  an  understanding  of  the  general  sub- 
ject of  divination. 

Confining  ourselves  to  Babylonia  and  Assyria,  the  chief  method 
of  voluntary  divination  was  the  inspection  of  the  liver  of  the  sac- 
rificial animal  and  the  chief  method  of  involuntary  divination,  the 
observation  of  phenomena  of  the  heavens.  The  correctness  of  this 
thesis  is  shown  by  the  wide  scope  of  these  methods  as  revealed  in 
the  texts  themselves.^*^  Both  methods  rest  on  a  well-defined  theory, 
the  inspection  of  the  liver  on  the  basis  of  the  primitive  view 
that  the  liver  was  the  seat  of  vitality,  of  the  intellect,  of  both  the 
higher  and  lower  emotions — in  short,  the  seat  of  the  soul,  as  that 
term  was  popularly  understood."  The  deity  in  accepting  the  sacri- 
ficial animal  identifies  himself,  as  it  were,  with  the  animal,  becomes 


1* 


See  Jastrow,  /.  c,  II.,  p.  209  f. — especially  note  i  on  p.  210.    See  parts 
I1-12  of  this  work  for  "  liver  "  omens  and  the  forthcoming  parts  13  and  14  for 
astrological "  omens. 

Jastrow,  /.  c,  pp.  213  f. 


U 


x9o8.] 


FOR   PLANET  IN   BABYLONIAN.  146 


one  with  it  and,  accordingly,  the  liver  of  the  animal  reflects  the  mind 
and  will  of  the  god.  If  one  can  therefore  read  the  liver  correctly,  one 
enters,  as  it  were,  into  the  workshop  of  the  deity.  The  mind  of  the 
animal  and  the  mind  of  the  deity  become  for  this  specific  occasion 
like  two  watches  regulated  to  be  in  perfect  accord. 

The  divining  of  the  future  through  the  observation  of  the  phe- 
nomena in  the  heavens  rests  on  the  identification  (or  personification) 
of  the  gods  with  the  sun,  moon,  planets  and  stars.  The  movements 
of  these  bodies,  the  changes  in  their  aspects  and  the  variations  in 

■ 

their  relationship  to  one  another  represent,  as  it  were,  the  activity 
of  the  gods  and  since,  according  to  the  current  theory,  all  happen- 
ings on  earth  are  due  to  the  gods  or  to  one  god  or  the  other,  a  knowl- 
edge of  what  the  activity  in  the  heavens  portends  furnishes  the 
means  of  foretelling  what  is  to  happen  on  earth.  In  time  no  doubt 
the  theory  was  perfected,  at  least  in  the  theological  circles  of  Baby- 
lonia and  Assyria,  into  a  complete  correspondence  between  occur- 
rences on  earth  and  the  decision  to  bring  about  these  occurrences  by 
the  manifested  activity  of  the  gods  in  the  heavens ;  but  even  without 
the  perfected  theory,  the  repeated  observation  of  the  kind  of  happen- 
ings on  earth  coincident  with  conditions  and  phenomena  in  the 
heavens  would  have  led  to  attaching  importance  to  these  conditions — 
both  those  of  a  usual  order  and  those  of  a  more  or  less  unusual  nature. 
Of  these  two  chief  divisions  of  divination,  it  is  evident  that 
the  inspection  of  the  liver,  connected  as  it  is  with  a  primitive  view 
of  that  organ,  can  be  accounted  for  as  the  distinct  outgrowth  of 
popular  beliefs,  whereas  the  divination  through  the  phenomena  of 
the  heavens  not  only  makes  greater  demands  on  scientific  or  pseudo- 
scientific  knowledge  but  presupposes  also  a  conception  of  world- 
philosophy  which  can  hardly  be  fermed  popular.  The  personifica- 
tion of  the  sun  and  moon  is,  of  course,  an  element  in  all  primitive 
phases  of  belief,  but  the  extension  of  such  personification  to  the 
planets  and  stars  belongs  to  a  higher  order  of  thought,  since  the 
bearings  of  those  bodies  on  the  life,  happiness  and  fate  of  mankind 
are  of  a  more  remote  and  a  more  indirect  character  than  in  the  case 
of  the  two  luminaries ;  and  when  we  come  to  the  projection  of  prac- 
tically all  the  activity  of  the  gods  on  to  the  heavens,  we  have  defi- 
nitely passed  beyond  the  intellectual  range  of  popular  fancy  and 


146  JASTROW— THE  SIGN  AND   NAME  [Aprils, 

• 

have  entered  the  domain  of  distinctly  theological  speculation.  If 
the  views  of  the  school  associated  with  the  names  of  Winckler  and 
Jeremias,  that  the  entire  Babylonian  religion  is  under  the  sway  of 
"astral"  conceptions,  turn  out  to  be  correct,  it  will  also  have  to  be 
recognized  that  the  underlying  "  Weltanschauung  "  is  a  product  of  the 
schools  rather  than  an  expression  of  popular  notions.^^*  I  venture 
to  think  that  one  of  the  weaknesses  of  the  "  astral "  theory,  which 
has  from  other  points  of  view  so  much  in  its  favor,  is  this  failure 
on  the  part  of  its  promoters  to  recognize  the  essentially  "  learned  " 
character  of  what  according  to  them  became  the  prevailing  world- 
philosophy  in  the  ancient  Orient  and  which  must  for  a  long  time  at 
least  have  separated  it  sharply  from  the  much  lower  plane  of 
popular  beliefs  and  fancies. 

Be  this  as  it  may,  the  development  of  a  method  of  divina- 
tion, through  elaborate  observations  of  the  movements  and  positions 
of  sun,  moon,  planets  and  stars,  it  will  be  admitted,  belongs  to  a 
later  stage  in  the  unfolding  of  religious  rites  than  so  primitive  a 
method  as  the  inspection  of  the  liver  of  a  sacrificial  animal.  The 
persistence  of  astrology  among  advanced  cultures  as  in  India  and 
Persia  and  in  western  Europe*^  down  to  the  threshold  of  modem 
times,  whereas  "  liver  "  divination  disappeared  with  advancing  cul- 
ture everywhere  except  among  the  Babylonians  and  Assyrians  and 
the  Greeks,  Romans  and  Etruscans,^^*  clinches  the  argument  in  favor 
of  divination  through  the  liver  as  the  earlier  and  more  primitive 
method.     If  this  be  admitted,  it  would  be  reasonable  to  find  in  the 

"•  See  also  Comont,  "  Les  Religions  Orientales  dans  le  Paganisme 
Romain"  (Paris,  1907),  p.  197. 

"  See  the  summary  by  Jeremias,  "  Das  Alte  Testament  im  Lichte  des 
Orients"  (i  ed.),  p.  7,  note  i. 

"•  Roman  divination  is  dependent  upon  Etruscan,  while  in  the  case  of 
Greek  divination  it  is  still  a  question  whether  we  are  to  assume  direct  in- 
fluence from  Babylonia  or  likewise  through  the  mediation  of  the  Etruscans. 
In  either  case  we  have  only  two  systems  of  "liver"  divination  surviving 
among  cultured  nations — the  Babylonian  and  the  Etruscan;  and  further 
investigations  may  definitely  confirm  the  view  which  on  the  surface  seems 
plausible  that  "  liver "  divination  among  the  Etruscans  stands  in  some  direct 
connection  with  Babylonian  divination.  If  this  be  so,  then  the  single  cause 
to  which  the  persistence  of  "liver"  divination  in  certain  quarters  is  to  be 
ascribed  is  the  elaboration  of  the  complicated  and  ingenious  system  of  inter- 
pretation which  we  owe  to  Babylonian  priests.  See  Jastrow,  II.,  pp.  215 
and  320,  note  3. 


«9o8.] 


FOR  PLANET  IN   BABYLONIAN.  147 


later  method  of  divination  through  the  heavens,  traces  of  the  earlier 
one,  if  not  indeed  some  link  directly  connecting  the  two.  Among 
the  Etruscans  we  actually  encounter  such  a  link  in  the  interesting 
circimistance  that  the  famous  "bronze  liver"  of  Piacenza,^'  pre- 
pared like  the  Babylonian  clay  model  of  a  liver^'*  as  an  object  lesson 
for  instruction  in  the  temple  schools,  is  divided  off  along  the  margin 
into  sixteen  regions,  corresponding  with  the  ordinary  divisions  of  the 
heavens  and  that  the  forty  Etruscan  words  with  which  the  surface 
of  the  liver  is  covered  are  names  of  deities.  Whether  we  accept 
Thulin's  view,^*  who  sees  a  direct  relationship  between  the  enumera- 
tion of  the  gods  and  the  list  and  arrangement  given  by  Martianus 
Capella,  or  follow  Korte,"  in  either  case  the  "  liver  "  reproduces  the 
recognized  divisions  of  the  heavens  and  through  this  combina- 
tion the  liver  becomes,  as  it  were,  a  microcosm  reflecting  the 
macrocosm.  The  much-discussed  problem^'*  whether  this  re- 
markable object  dating  from  about  the  third  century  B.  C.  is  a 
"  liver  "  or,  as  was  first  supposed,  a  "  templum,"  thus  resolves  itself 
into  the  thesis  that  it  is  both.  To  use  the  words  of  Korte  in  his 
paper  in  summarizing  the  results  of  twenty-five  years  of  study  of 
this  object  :^* 

"  The  liver  as  the  seat  of  life  according  to  the  view  of  antiquity  appears 
as  a  minature  reproduction  of  the  universe.  As  the  latter,  so  the  liver  is 
divided  into  a  right  and  left  half,'*  a  day  division  and  a  night  division,  the 
line  of  division  corresponding  to  the  line  dividing  the  universe  into  east  and 
west  As  the  vault  of  heaven,  so  the  edge  of  the  liver  is  divided  into  i6 
regions  in  which  the  gods  who  furnish  portents  dwell." 

"See  Korte,  "Die  Bronzeleber  von  Piacenza"  (MitteiL  d  Kcdserl, 
Deutsch.  Archceolog.'Instituts.,  XX.,  pp.  348-379),  the  latest  and  probably 
final  word  on  the  subject 

"•Cuneiform  Texts,  VI.,  PI.  1-2  and  photograph. 

"*  "  Die  Gotter  des  Martianus  Capella  und  der  Bronzeleber  von  Piacenza  " 
(Giessen,  1906),  pp.  31-59- 

"  Korte,  /.  c,  p.  367  f . 

**•  See  the  references  in  Korte,  p.  349  f .,  to  which  Nicola  Terzaghi,  "  Li 
piu  recente  Interpretazione  dei  Mundus-Templum  di  Piacenza"  {Bolletino 
Storico  Piacentino,  1906,  Maggio-Giugno)  is  to  be  added. 

"  Korte,  /.  c,  p.  362. 

"Referring  to  the  band  on  the  reverse  of  the  object  See  the  illustra- 
tion in  Korte's  article,  p.  356. 


148  JASTROW— THE  SIGN  AND  NAME  I  April  as. 

III. 

If,  therefore,  among  the  Etruscans  we  find  the  unmistakable 
proof  of  a  direct  link  between  the  two  classes  of  divination,  we 
should  be  prepared  to  find  a  similar  association  in  Babylonia  and 
Assyria.  I  believe  that  the  ordinary  name  and  sign  for  planet  in 
Babylonia  points  in  this  direction.  While  already  in  early  days  we 
find  various  animals  and  all  kinds  of  products  dedicated  as  offer- 
ings to  the  gods,^®  for  purposes  of  divination  the  only  animal  set 
aside  was  the  sheep.  This  follows  not  only  from  the  fact  that  the  fa- 
mous clay  model  of  a  liver  found  near  Bagdad  is  that  of  a  sheep,^*  but 
from  the  specific  references  to  sheep  in  "  liver  "  divination  texts  and 
to  no  other  animal.^^  The  sheep  thus  becomes  the  animal  of  divina- 
tion par  excellence,  and  we  can  well  suppose  that  the  word  itself 
should  come  to  be  used  as  synonymous  with  divination.  Such  a 
usage  would  be  paralleled  by  the  extension  of  the  term  auspicium  in 
Latin,  which  from  being  an  omen  derived  through  "  bird  observation  " 
was  applied  to  any  kind  of  an  omen  or  portent,  so  that  an  inspection 

"  See  Thureau-Dangin,  "  Die  Sumerischen  und  Akkadischen  Konigsin- 
schriften"  (Leipzig,  1907)1  PP.  16,  80,  84,  86,  88,  124,  etc.  I  cannot  here 
enter  into  a  full  discussion  of  the  nature  of  sacrifices  among  the  Baby- 
lonians and  Assyrians  but  it  may  be  proper  to  point  out  that  in  an  elaborate 
ritual  controlled  by  an  extended  priestly  organization  we  must  sharply  dif- 
ferentiate between  (i)  offerings  that  constitute  part  of  the  income  of  the 
temples,  (2)  voluntary  gifts,  (3)  sacrifices  offered  in  connection  with  purifica- 
tion or  expiatory  rites  and  (4)  sacrifices  offered  directly  to  and  for  the  god. 
So  far  as  I  can  see  sacrifices  of  the  latter  kind  were  brought  only  when  an 
answer  to  a  specific  question  was  desired,  so  that  it  would  appear  that  divina- 
tion forms  the  starting  point  for  the  development  of  the  whole  idea  of  sacri- 
fice in  the  proper  sense  in  Babylonia. 

"  CT,  VI.,  PI.  I.  See  Jastrow,  /.  c,  II.,  p.  218  note  i,  where  a  reference 
should  have  been  given  to  Stieda,  "Ueber  die  aeltesten  bildlichen  Darstel- 
lungen  der  Leber"  (Bonnet-Merkel,  Anatomische  Hefte  XV.,  p.  697),  who 
shows  that  it  is  (as  also  in  the  case  of  the  bronze  liver  of  Piacenza)  the  liver 
of  a  sheep  and  not  of  a  goat — as  had  been  supposed  by  some  scholars. 

*E.  g.,  CT,  XX.,  I,  I — in  the  opening  line  of  the  first  tablet  of  a  series 
dealing  with  "  liver  "  divination ;  also  Boissier,  "  Documents  Assyriens  relatif s 
aux  Presages,"  p.  97,  n;  212,  27;  also  in  the  "omen "  text  CT.,  IV.,  PI.  34,  9; 
in  the  omen  report  of  the  Cassite  period  (Clay,  Cassite  Archives,  XIV., 
PI.  4,  Obv.  10,  and  lastly  the  constant  mention  of  the  "sheep"  in  the  omens 
attached  to  Knudtzon,  Assyr,  Gebete  an  den  Sonnengott.  Note  also  the 
expression  bel  immeri  "owner  of  the  sheep"'  (CT,  XX.,  33,  93  and  Boissier, 
Documents,  p.  96,  13) .  The  addition  of  NITA  to  Lu  shows  that  a  "  male " 
sheep  was  selected  for  the  purpose. 


,9o8.]  FOR   PLANET  IN   BABYLONIAN.  149 

of  the  liver  of  an  animal  for  the  purpose  of  securing  an  "omen" 
was  also  designated  as  an  auspiciumr^     Similiarly,  in  Greek  the 

word  opvt^,  "  bird,"  is  used  for  any  kind  of  an  omen  and  my  colleague, 
Professor  Lamberton,  has  kindly  called  my  attention  to  the  inter- 
esting passage  in  the  Birds  of  Aristophanes  in  which  this  usage  finds 
a  striking  illustration.  In  the  "  Parabasis,"  after  indicating  all  the 
blessings  that  accrue  to  men  from  the  birds,  the  chorus  turns  to 
divination  and  continues  as  follows  :*^ 

"You  consider  all  things  a  bird,  whatever  gives  a  decision 
through  divination.  With  you  a  word  is  a  'bird,*  and  you  call  a 
sneeze  a  *  bird,'  a  sound  a  '  bird,'  a  sudden  meeting  a  '  bird,'  and  an 
ass  a  '  bird.'    Are  we  not  clearly  a  prophetic  Apollo  to  you  ?  " 

The  sheep,  being  the  animal  of  divination  par  excellence  in 
Babylonia,  would  in  the  same  way  become  the  Babylonian  term  for 
an  "  auspicium  "  in  general.  If  we  assume  that  this  use  of  the  term 
lurks  in  the  application  of  "  sheep  "  as  the  designation  of  a  planet, 
a  satisfactory  explanation  can  be  found  for  the  addition  of  the  sign 
Bat  to  the  sign  for  "  sheep  "  which  has  more  specifically  the  same 
force  in  the  combination  Lu-Bat  as  in  the  combination  IIR.  27, 
No.  2,  Obv.  46,  c-d,  Ur-Bat,  i.  e.,  "  dead  liver  "  in  the  sense  of  the 
liver  of  a  sacrificial  lamb  and  hence  as  the  equivalent  of  ter-tu  sa 
hchse-e,  "  omen  through  the  liver."^' 

The  combination  Lu-Bat  thus  expresses  more  precisely  than  Lu 
alone  the  association  of  an  "  omen  "  with  a  "  sheep,"  and  we  would 
be  justified  in  rendering  the  combination  as  "  sheep  omen,"  and  then 
through  the  association  of  ideas  above  pointed  out,  as  a  general  term 
for  "  omen."" 

"See  Pauly-Wissowa,  "  Real-Encyclopaedie/*  (new  ed.),  H.,  p.  2580  f. 

"LI.  71^22  (cd.  Van  Lceuwcn,  Leiden,  1893).  Dr.  R.  G.  Kent,  of  the 
University  of  Penna.,  also  calls  my  attention  to  the  interesting  passage  in 
Xenophon's  Anabasis  (iii,  2,  9)  where  a  "  sneeze"  as  a  good  sign  is  spoken 
of  as  hiuv6g  or  "  bird  "  in  the  general  sense  of  an  omen. 

*■  On  the  word  hdsii  for  liver  which  may  have  been  used  in  earlier  days 
in  place  of  kabittu  see  Jastrow,  /.  c,  IL,  p.  213,  note  i,  and  p.  276,  note  7. 

••It  is  to  be  noted  that  at  least  in  one  passage  in  a  "liver  divination*' 
the  sign  Bat  is  added  to  Lu-NrrA  "  male  sheep,"  namely  Boissier,  Doc,  Assyr., 
p.  212,  27,  ultu  libbi  Lu-NrrA  Bat  (u)  tertu  (written  Ur-Bat  as  in  the  pas- 
sage IIR  27)  tu-sc-la-a,  i.  e.,  "  Out  of  a  dead  sheep  thou  shalt  bring  forth  an 
omen,"  where  the  phonetic  complement  u  added  to  Bat  suggests  the  reading 
mitu  and  where  "  dead  sheep  "  is  clearly  the  equivalent  of  "  sacrificial  sheep  " 
or  "omen  sheep." 


150  JASTROW— THE  SIGN   AND   NAME  [April  as. 

Now  what  was  the  purpose  for  which  the  movements  of  the 
planets  were  observed  by  the  Babylonians?  What  other  than  to 
secure  through  such  observation,  signs  by  means  of  which  the  future 
could  be  divined?  The  planets  were,  primarily,  regarded  as 
"  omens  "  and  since,  as  has  been  above  set  forth,  divination  through 
the  heavens  follows  in  point  of  time  divination  through  the  liver  of 
the  sheep,  we  would  expect  conceptions  and  terms  used  in  "  liver  " 
divination  to  be  transferred  to  astrological  divination.  The  use  of 
the  term  "  sheep  "  as  the  designation  of  the  planets  observed  to 
secure  omens,  precisely  as  omens  were  furnished  by  means  of  sacri- 
ficial sheep,  I,  accordingly,  take  as  an  illustration  of  this  dependence  . 
of  astrology  upon  hepatoscopy,  forming,  as  it  were,  the  connecting 
link  between  the  two.  It  may  be  noted  in  this  connection  that  the 
interpretations  given  in  astrological  texts  to  signs  observed  are  paral- 
leled in  the  "  liver  "  divination  texts,**  and  there  can  be  little  doubt 
that  they  are  transferred  bodily  from  the  latter  and  earlier  class  of 
texts  to  the  former. 

The  explanation  here  proposed,  according  to  which  Lu-Bat 
as  applied  to  the  planets  conveys  the  notion  that  they  were  regarded 
as  "  omens  "  or  means  of  securing  omens,  throws  a  new  light  upon 
the  statement  in  Diodorus**  that  the  Babylonians  commonly  called 
the  five  planets  ipfxr^vej^,  i.  e»,  "  interpreters,"  adding  as  a  reason 
for  the  designation  that  the  planets  were  regarded  as  "  inter- 
preting "  for  mankind  the  intention  of  the  gods.  Bouche-Leclercq 
("  L'AstroIogie  Grecque,"  p.  40,  note  3),  recognizes  that  the  term 
"  interpreters  "  does  not  embody  a  Greek  tradition,  but  the  notice 
in  Diodorus,  so  far  from  being,  as  he  supposes,  of  "  doubtful  value," 
reflects  the  perfectly  correct  view  that  the  planets  were  used  as 
"  omens  ""  and  the  term  "  interpreters  "  is  evidently  an  attempt  to 

"The  interpretations  in  the  "astrological"  texts  are  in  fact  practically 
identical  with  those  in  "liver"  divination,  furnishing  the  same  references  to 
public  events  and  differing  merely  in  containing  more  references  to  crops,  to 
prices  of  food  and  to  famine.  Cf.,  e.  g.,  Craig,  "  Astrolog.-Astronom.  texts," 
PI.  2,  3;  20,  22  with  CT,  XX.,  26,  Obv.  3;  Boissier,  Doc.  Assyr.,  7,  21; 
Craig  20,  31  with  CT,  XX,,  32,  54;  99,  100  (where  ilu  =  Nergal).  Cf.  Jastrow, 
/.  c,  II.,  p.  342,  note  11). 

"Bibl.  Histor.,  Book  II.  (ed.  Dindorf),  30,  4. 

''To  be  sure,  what  Diodorus  says  in  addition  why  the  planets  and  not 
also  the  other  stars  were  regarded  as  "interpreters"  is  rather  beside  the 


1908.]  FOR   PLANET   IN   BABYLONIAN.  151 

convey  this  idea.  The  term  may,  therefore,  be  regarded  as  a  render- 
ing of  the  Babylonian  designation  "  sheep  omen "  in  the  general 
sense  above  pointed  out. 

The  objection  may  be  raised  at  this  point,  why  should  not  the 
moon  and  sun,  as  playing  an  equally  if  not  more  important  role  in 
divination  lore,  likewise  have  been  designated  as  Lu-Bat  in  the 
generic  sense  of  an  "  omen "  or  "  auspicitmi "  ?  The  answer  is 
obvious.  Sun  and  moon  cults  are  such  ingredient  parts  of  early 
forms  of  religion  everywhere  and  the  dependence  of  human  for- 
tune, life,  health  and  welfare  upon  these  two  luminaries  is  so  direct 
that  other  factors  were  at  work  in  the  development  of  conceptions 
regarding  these  two  deities  than  merely  the  observation  of  their 
movements  and  changing  relationship  to  one  another  as  a  basis 
for  determining  what  these  deities  were  preparing  for  mankind. 
Their  cult  precedes  their  introduction  into  divination  texts,  whereas 
the  planets  were  observed  solely  for  purposes  of  divination.  Since 
the  influence  of  the  latter  on  human  life  was  a  matter  of  speculation 
rather  than  of  direct  experience,  the  basic  and  primary  motive  for 
noting  their  movements  was  in  connection  with  the  view  that,  as  rep- 
resenting gods,  their  movements  indicated  the  activity  of  these  gods 
in  preparing  the  events  that  were  to  happen  on  earth.  The  old 
and  long  established  names  and  designations  for  sun  and  moon  were 
accordingly  retained,  whereas  the  new  term  chosen  for  the  planets 
was  ordinarily  restricted  to  them.  Occasionally,  however,  so,  e,  g., 
III.  R,  57,  No.  6,  65-67,  sun,  moon  and  the  five  planets  are  sum- 
marized as  seven  Lu-Bat  (pi.). 

Thjtt  the  association  of  ideas  did  not,  on  the  other  hand,  lead  to 
the  extension  of  Lu-Bat  to  the  stars  in  general  constitutes  no  valid 
objection  to  the  thesis  here  propounded.  In  the  divination  texts  the 
number  of  stars  introduced,  outside  of  the  planets,  is  not  large  and 
their  role  is  quite  secondary,^*  and  it  is  not  until  we  reach  the  period 
when  astronomy  becomes  more  definitely  diflferentiated  as  a  science 
from  astrology,  when  calculations  are  made  and  "  planet "  tables  are 

mark;  and  shows  that  he  no  longer  fully  understood  the  force  of  the  Baby- 
lonian designation  which  he  here  faithfully  reproduces. 

"In  astrological  texts  proper  as  distinguished  from  astronomical  tablets, 
the  stars  mentioned  are  chiefly  certain  ones  belonging  to  the  constellations  of 
the  ecliptic  and  which  are  frequently  introduced  as  guides  and  indications  for 
fixing  the  position  of  the  planets,  rather  than  as  omens. 


152  JASTROW— THE  SIGN   AND   NAME  [April  .5. 

prepared  independently  Qf  divination,  that  star-lore  assumes  larger 
dimensions.  Besides,  in  securing  omens  the  positions  of  the  stars 
constitute  a  minor  factor  and  are  of  value  chiefly,  if  not  exclusively, 
in  relationship  to  phenomena  connected  with  the  planets — a  condition 
which  is  specially  applicable  to  the  relationship  between  the  planets 
and  the  constellations  of  the  zodiac. 

Attention  has  already  been  called  to  the  fact  that  although  Lu- 
Bat  is  commonly  applied  to  any  planet,  there  are  only  two  planets — 
Mercury  and  Saturn — ^that  regularly  appear  written  with  this  com- 
pound ideograph,^*  the  former  being  designated  as  Lu-Bat  Gu- 
Ud/®  the  latter  as  Lu-Bat  Sag-Us,  while  Mars  occasionally  appears 
as  Lu-Bat  Dir.'^  The  other  planets  appear  in  the  lists  IIR,  48, 
48-54  ab  and  IIIR,  57,  No.  6,  65-67,  as  (il)  Sul-pa-Ud-Du-a 
(Jupiter)  (il)  Dil-Bat  (Venus)  and  ZAL-BAT-a-nu  (Mars),  with 
MuL  =  kakkabu  interchanging  with  an  =  ilu.  Moreover,  the 
phonetic  reading  bi-ib-bu  in  the  latter  list  for  Lu-Bat  Gu-ud  points 
to  Mercury  as  being  the  planet  par  excellence.  Why  should  Mer- 
cury have  been  assigned  to  this  preeminent  position  among  the 
planets?  It  has  been  suggested  to  me'^  that  the  position  of  Mer- 
cury nearest  to  the  sun  may  have  led  to  its  being  looked  upon  as 
the  chief  planet  for  purposes  of  divination  and  it  is  perhaps  not 
without  significance  that  in  Greek  astrology  Mercury,  frequently 
designated  as  ffTtX^wv,  "  shining,"'^  is  closely  associated  with  the 
sun,  and  indeed  at  times  identified  with  Apollo.'*  Certainly,  the 
peculiar  conceptions  connected  with  Mercury  in  the  astrology  of 
the  Greeks  and  of  other  nations — whose  dependence  upon  Baby- 
lonian beliefs  and  speculations  is  generally  admitted — sharply  sepa- 
rate that  planet  from  his  fellows.  While  the  others,  e.  g.,  are  con- 
ceived as  masculine  or  feminine.  Mercury,  and  Mercury  alone,  is 
double  sexed.^°    Qualities  are  heaped  upon  Mercury  in  profusion, 

*•  See  above,  p.  141,  note  2. 

*•  Generally  read  Gud-Ud  but  the  reading  Gu-Ud  seems  preferable. 

**  See  above,  p.  141,  note  2. 

"By  my  friend,  Mr.  H.  H.  Fumess,  Jr.,  whose  suggestion  commended 
itself  to  my  colleague.  Professor  C.  L.  Doolittle,  Director  of  the  Flower 
Observatory   (University  of  Pennsylvania). 

"  Bouche-Leclcrcq,  "  L'Astrologie  Grecque,"  pp.  66  and  100. 

**  Ibid.,  p.  100,  note  5. 

'^  Ibid.,  p.  102.  So  also  in  modem  astrology.  See  Ellen  H.  Bennett, 
"Astrology"  (New  York,  1897),  p.  98. 


,9o8.)  FOR  PLANET  IN   BABYLONIAN.  153 

in  contradistinction  to  the  other  planets  to  whom  generally  a  single 
dominant  trait  is  given.  Intelligence,  thought,  feeling,  eloquence, 
artistic  spirit  are  all  associated  with  Mercury,'*  which  thus  be- 
comes, as  it  were,  the  "  soul "  among  the  planets  and  it  will  not 
seem  far-fetched  to  see  in  the  fancy  which  makes  Mercury  the 
planet  of  revelation  and  of  language'^  a  trace  of  primitive  views 
regarding  the  seat  of  vitality.  In  accord  with  this,  we  actually 
find  Mercury  assigned  to  the  liver'*  as  the  organ  of  revelation, 
though  in  deference  to  later  views  of  the  liver  as  the  seat  of  the 
affections  specifically — and  not  of  all  intellectual  life  and  of  all 
emotions — ^Venus  is  sometimes  identified  with  this  organ.'*  To  be 
sure,  such  associations  of  ideas  have  not  as  yet  been  encountered 
in  Babylonian  texts  and  therefore  a  certain  reserve  is  called  for. 
On  the  other  hand,  the  dependence  of  Greek  astrology  on  Babylonian 
conceptions,  fancies  and  prototypes  is  so  evident  at  every  turn*® 
that  we  are  justified  in  assuming  a  large  measure  of  identity  between 
the  two  systems  of  divination,  just  as,  on  the  other  hand,  modem 
astrology  is  full  of  conceits  and  notions  that  can  be  paralleled  in 
ancient  Greece,  India  and  Persia. 

Another  factor  that  may  have  led  to  assigning  to  Mercury  a 
specially  prominent  place  among  the  planets  for  purposes  of  divi- 
nation is  the  circumstance  that  by  virtue  of  its  close  position  to  the 
sun  and  its  small  size,  it  makes  its  circuit  in  the  short  space  of  twelve 
weeks  and  four  days,  or  87.97  solar  days.  Hence,  since  the  basis  of 
divination  in  the  case  of  the  planets  is  largely  bound  up  with  their 
relative  position  to  the  sun — upper  conjunction,  ascent,  culmination, 
standstill,  descent,  lower  conjunction*^ — Mercury  would  present  a 
far  larger  proportion  of  changes  in  any  given  time  than  any  other 
planet.  In  the  case  of  frequent  observations.  Mercury  would  thus 
play  a  more  prominent  part  than  the  other  planets  whose  movements 
except  for  periods  of  some  duration  would  furnish  less  of  moment 

"  Bouche-Leclercq,  p.  loi ;  Bennett,  p.  99. 
"  Bouche-Leclercq,  pp.  312,  321,  323. 
"  Ibid.,  p.  312  and  323. 
.'^Ibid.,  p.  321. 

**See  Bouche-Leclercq's  summary,  pp.  70-71. 

^See  the  valuable  discussion  in  Kugler,  /.  c,  p.  20  1,  of  the  Babylonian 
equivalents  for  those  terms. 


154  JASTROW— THE  SIGN  AND   NAME  [April  as, 

to  the  observer,  dependent  upon  the  naked  eye.  But  whatever  the 
reasons,  we  can  only  conclude  from  the  fact  that  Mercury  is  the 
"  sheep  "  par  excellence  that  it  was  at  one  time  singled  out  as  the 
planet  of  revelation  and  that,  therefore,  it  was  in  all  probability 
the  first  planet  whose  movements  were  observed  for  the  purpose  of 
securing  through  them  a  means  of  determining  what  events  the  gods 
were  preparing  to  take  place  on  earth. 

The  designation  of  Saturn  as  lulintu,  "  ram,"  I  am  inclined  to 
regard  of  secondary  origin,  that  is  to  say,  dependent  upon  the  appli- 
cation of  bi'ib'bu  to  Mercury — ^the  latter  term  taken  no  longer  in 
the  sense  of  an  "  omen  "  but  already  as  a  specific  and  distinguishing 
designation.  As  companion  piece,  therefore,  to  Mercury  as  a 
"sheep,"  Saturn  was  called  a  "ram"  just  as  the  designation  of 
the  seven  Masi-stars  by  the  determinative  Lu  ("sheep")"  is  a 
secondary  extension  from  Lu-Bat,  limited  originally  to  the  planets. 
Saturn  presents  in  almost  every  respect  a  contrast  to  Mercury.  It 
is  infinitely  larger*'  in  bulk,  at  a  great  distance  from  the  sun,  the 
most  regular  of  the  planets  and  the  slowest  in  its  motion,  taking  10,759 
days  or  29.46  years  to  pass  around  the  sun.  In  Greek  astrology  a 
preeminent  position  is  accorded  to  Saturn,**  which  is  expressed,  for 
example,  by  making  the  planet  the  head  and  "  brain  "  of  the  plan- 
etary world — reflecting  the  later  view  which  placed  the  seat  of  the 
soul  in  the  head,*'  while  the  association  of  Saturn  with  Mercury 


*•  Kugler,  /.  c,  p.  7. 
"Jupiter  alone  is  larger. 


**  Bouch^-Leclercq,  /.  c,  p.  94  f.  It  is  to  be  noted  that  Saturn  is  in 
Babylonian  astrology  called  "  the  star  of  the  sun  "  (as  Diodorus,  II.,  30,  also 
says) — ^which  reminds  one  that  Mercury  (see  above,  p.  152)  was  identified 
with  the  sun  in  Greek  astrology;  the  same  appears  to  have  been  the  case 
with  Saturn.    See  Kugler,  /.  c,  p.  8. 

**Ibid.,  p.  95.  The  soul  was  placed  successively  (a)  in  the  liver,  (6)  in 
the  heart  and  (c)  in  the  brain.  "Liver"  divination  is  the  outward  expression 
corresponding  to  the  first  stage.  The  addition  of  the  "heart"  (and  then 
of  other  organs)  to  the  "  liver  "  in  the  examination  of  the  sacrificial  animal — 
as  among  the  Romans — is  a  concession  to  the  second  stage,  while  phrenology 
is  an  expression — outside  of  the  official  cult — of  the  third  stage.  See  Jastrow, 
"Divination  through  the  Liver  and  the  Beginnings  of  Anatomy"  (University 
of  Pennsylvania  Medical  Bulletin,  January,  1907)-  In  a  special  paper  on  "  The 
Liver  as  the  Seat  of  the  Soul"  I  propose  to  treat  in  detail  of  these  suc- 
cessive views. 


,9o8.]  FOR  PLANET  IN  BABYLONIAN.  155 

crops  out  in  the  belief  which  makes  the  history  of  the  world  begin 
with  the  reign  of  Saturn  and  end  with  that  of  Mercury.**  The 
prominence  of  Saturn  in  Babylonian-Assyrian  astrological  texts 
is  in  accord  with  this  association  with  Mercury  as  a  second  Lu-Bat 
par  excellence,^^^ 

In  modem  astrology  Saturn  continues  to  play  a  particularly  con- 
spicuous role*^ — all  of  which  points  to  its  having  been  the  first  planet 
to  become,  by  the  law  of  contrasts,  associated  with  the  original 
"source"  of  divination  among  the  planets — Mercury. 

Lastly,  a  word  regarding  the  ideographic  designations  of  these 
two  "sheep" — Mercury  and  Saturn.  Kugler,*'  following  in  part 
Jensen,*'  proposes  to  take  the  element  Gu-Ud  in  Lu-Bat  Gu-Ud 
as  karradu  sa  urri,^^  "  warrior  of  the  light,"  because  shortly  after 
his  appearance  in  the  East  day  triumphs  over  night.  The  explana- 
tion seems  forced  and  it  is  hardly  likely  that  a  circumstance  like  this 
should  have  suggested  a  name  for  a  planet.  In  view  of  the  fact 
that  Mercury  and  Saturn  are  the  two  planets  more  particularly  des- 
ignated as  Lu-Bat,  it  is  more  reasonable  to  see  in  Lu-Bat  Gu-Ud 
and  Lu-Bat  Sag-Us  descriptions  of  characteristic  features.  For 
Sag-Us,  fortunately,  the  equivalent,  kora-mor-nu,^^  has  been  defi- 
nitely ascertained  and  the  meaning  "  regular  "  is  also  beyond  doubt. 
The  name  was  clearly  given  to  the  planet  because  of  the  slow  and 
regular  motion  which  is  its  distinguishing  feature.    Mercury,  on  the 

*•  Bouch^-Leclcrcq,  pp.  187,  498  f. 

*^The  statement  of  Diodorus  (/.  c.)  that  Saturn  was  regarded  by  the 
Babylonians  as  the  most  important  for  purposes  of  divination  may  correctly 
reflect  a  later  stage  when  Saturn  assumed  the  preeminent  place  once  occupied 
by  Mercury. 

^  Bennet,  /.  c,  p.  93. 

*•  Kugler,  /.  c,  p.  10.  On  p.  218  he  prefers  the  rendering  "  full  of  light " 
(as  Hommel,  Aufsatze,  p.  381,  does)  but  the  two  ideas  ("  full "  in  the  sense 
of  "  strong  "  and  "  warrior  ")  are  correlated. 

*• "  Kosmologie,"  p.  131,  who  takes  Gud-Up  as  a  single  term  =  ifearrarfu 
"warrior"  (Br.  5742).  It  is  always  to  be  bom  in  mind  that  we  are  to  substi- 
tute Mercury  for  Mars  throughout  Jensen's  volume — ^now  that  it  has  been 
definitely  ascertained  that  Gud-Ud  =  Mercury  and  not  Mars  and  Zal-Bat 
(a-nu)  =    mustabarru  mutanu  =  Mars  not  Mercury. 

■•  GuD  =  karradu    and    UD  =  urru     (Br.    7798)— though    umu  =  "day 
would  suggest  itself  as  more  probable. 

"Sec  Jensen,  /.  c,  p.  114.     Cf.  p^3  in  Amos  5,  26. 


<( 


« 


156  JASTROW— THE  SIGN  AND   NAME  [April  .5, 

Other  hand,  is  marked  by  its  rapid  and  irregular  course  and  I  accord- 
ingly propose  the  equation  GvAJd  =  sahdtu — ^a  common  value  of 
the  compound  ideogram  in  "  liver  "  divination  texts  in  the  sense  of 

hinder,  check,  restrain."'* 
Assuming  the  adjective   formation  sahtu  from  this  stem,  the 

checked  "  one  as  the  designation  of  this  planet  would  form  a  com- 
panion piece  to  kaimanu,  the  "  regular  "  one.  In  contrast  to  Jtoi- 
manu  "  regular,"  the  designation  sahtu  would,  naturally,  convey  the 
notion  of  a  body  checked  and  restrained  and  therefore  "  irregular  " 
in  its  motion.'* 

**  Cf.  Jastrow,  II.,  p.  366,  note  9. 

"The  gloss  in  Hesychius  according  .to  which  pe^^Paroc  is  in  Babylonian 
the  "  fire  "  star  cannot  be  explained  as  Jensen  "  Kosmologie,"  p.  97,  proposes, 
since  he  starts  from  the  false  assumption — since  abandoned  by  him — ^that 
bibbu — the  Lu-Bat  par  excellence  is  Mars,  whereas  it  is  Mercury.  That 
PeXiparo^  designates  Mars  is  however  no  doubt  correct  and  since  the  com- 
mon ideographic  designation  for  Mars  is  Zal-Bat — ^the  addition  of  a-nu 
being  a  phonetic  complement  to  suggest  the  phonetic  reading  mustabarru 
mutdnu,  "the  one  satiated  with  death  " — the  correction  oiPe^iparoc to i^e^iparoc 
suggests  itself  as  a  simple  solution  of  the  problem. 


FURTHER  RESEARCHES   ON   THE   PHYSICS   OF  THE 

EARTH,  AND  ESPECIALLY  ON  THE  FOLDING  OF 

MOUNTAIN    RANGES    AND    THE    UPLIFT    OF 

PLATEAUS   AND   CONTINENTS    PRODUCED 

BY    MOVEMENTS    OF    LAVA    BENEATH 

THE    CRUST    ARISING    FROM    THE 

SECULAR     LEAKAGE     OF     THE 

OCEAN   BOTTOMS. 

By  T.  J.  J.  SEE,  A.M.,  Lt.M.,  ScM.   (Missou.),  A.M.,  Ph.D.   (Berol.), 

Professor  of  Mathematics,  U.  S.  Navy,  in  Charge  of  the 

Naval  Observatory,  Mare  Island,  California. 

(Read  April  24,  IQ08.) 

I.  General  Considerations  on  the  Physics  of  the  Earth,  with 

Especial  Reference  to  the  Secular  Leakage  of  the 

Oceans  and  the  Resulting  Development  of 

Mountains,   Plateaus   and   Islands. 

§  I.  Introductory  Remarks, — In  three  papers  recently  communi- 
cated to  the  American  Philosophical  Society  held  at  Philadelphia 
and  since  published  in  the  proceedings  of  that  Society,^  the  writer 
has  treated  at  some  length  of  the  cause  of  earthquakes,  moun- 

*  I.  "  The  Cause  of  Earthquakes,  Mountain  Formation  and  kindred  phen- 
omena connected  with  the  Physics  of  the  Earth,"  Proc.  Am.  Philos.  Soc., 
1906. 

2.  "  On  the  Temperature,  Secular  G>oling  and  Contraction  of  the  Earth, 
and  on  the  Theory  of  Earthquakes  held  by  the  Anbients,"  Proc.  Am.  Philos. 
Soc,  1907. 

3.  "The  New  Theory  of  Earthquakes  and  Mountain  Formation  as  illus- 
trated by  Processes  now  at  work  in  the  Depths  of  the  Sea,"  Proc.  Am. 
Philos.  Soc.,  1907;  issued  in  March,  1908. 

The  following  shorter  articles  have  also  appeared: 

4.  "  Outline  of  the  New  Theory  of  Earthquakes,"  Popular  Astronomy, 
April,  1908. 

5.  **How  the  Mountains  were  Made  in  the  Depths  of  the  Sea,"  Pacific 
Monthly,  Sept.,  1908. 

PROC.  AMER.  PHIL.  soc.  XLVH.  1 89  K,  PRINTED  SEPTEMBER  21,  I908. 


158  SEE— FURTHER   RESEARCHES  •ON  [April  24. 

tain  formation  and  kindred  phenomena  connected  with  the 
physics  of  the  earth.  In  the  course  of  these  three  memoirs  many 
important  questions  are  considered,  and  it  seems  to  be  rendered 
highly  probable  that  six  great  classes  of  phenomena,  not  heretofore 
closely  associated,  depend  on  a  single  physical  cause,  namely,  the 
secular  leakage  of  the  ocean  bottoms,  and  the  resulting  movement  of 
molten  rock  beneath  the  earth's  crust.  The  six  classes  of  phenomena 
traced  to  a  single  physical  cause  are:  (i)  world-shaking  earth- 
quakes; (2)  the  activity  of  volcanoes;  (3)  mountain  formation; 
(4)  the  formation  of  islands  and  plateaus;  (5)  seismic  sea  waves; 
(6)  the  feeble  attraction  of  mountains  and  plateaus  long  noticed 
in  geodesy. 

The  first  of  the  memoirs  printed  by  the  American  Philosophical 
Society  deals  with  the  problem  of  earthquakes  in  its  general  aspects, 
and  sets  forth  groimds  for  the  theory  that  these  six  classes  of  phe- 
•nomena  are  directly  connected  and  dependent  on  a  single  physical 
cause ;  the  second  examines  the  question  of  the  earth's  temperature, 
secular  cooling  and  contraction,  and  endeavors  to  show  that  the 
traditional  theory  of  the  changes  noticed  on  the  earth's  surface  is 
not  well  founded;  while  the  third  seeks  to  demonstrate  the  more 
important  conclusions  reached  in  the  first  memoir,  by  an  appeal  to 
processes  now  at  work  in  the  depths  of  the  sea,  the  meaning  of  which 
apparently  is  so  plain  as  to  admit  of  no  possible  doubt. 

The  change  in  the  point  of  view  necessitated  by  the  considera- 
tions brought  forth  in  these  papers  is  so  remarkable  as  to  be  worthy 
of  the  attention  of  all  who  are  interested  in  the  grand  science  of 
natural  philosophy.  And  we  therefore  propose  to  consider  in  this 
paper  the  physical  basis  of  the  theory  of  ocean  leakage,  the  folding 
of  mountain  ranges  and  the  uplift  of  plateaus  and  continents  pro- 
duced by  movements  of  lava  beneath  the  crust,  together  with  the 
historical  aspects  of  the  problems  of  the  physics  of  the  earth. 
Heretofore  the  nature  of  the  forces  which  have  folded  mountain 
ranges  and  their  relationship  to  those  slow  movements  which  have 
raised  whole  continents  have  been  equally  mysterious  and  bewilder- 
ing to  the  investigator.  Accordingly  any  light  which  may  be  shed 
on  this  difficult  subject  will  no  doubt  be  exceedingly  welcome  to 
those  who  are  interested  in  the  progress  of  the  physical  sciences. 


«9o8]  THE  PHYSICS  OF  THE   EARTH.  159 

As  the  leakage  of  the  oceans  seems  to  be  clearly  proved  by  the 
movements  noticed  in  earthquakes,  especially  where  mountain  for- 
mation is  now  going  on  in  the  depths  of  the  sea,  and  the  seismic 
disturbances  are  therefore  accompanied  by  the  sinking  of  the  sea 
bottom,  as  shown  by  the  seismic  sea  waves  which  follow  the  earth- 
quakes, it  seems  legitimate  to  appeal  to  these  movements  of  molten 
matter  beneath  the  earth's  crust  as  the  only  available  means  of 
demonstrating  the  porosity  and  other  physical  properties  of  layers 
of  granite  twenty  miles  thick.  Owing  to  the  restricted  conditions  of 
human  life,  no  experiments  on  such  a  grand  scale  can  ever  be  at- 
tempted in  our  laboratories,  however  great  the  facilities  at  our 
command ;  and  our  only  means  of  ascertaining  the  truth  with  regard 
to  the  theory  of  ocean  leakage  is  by  careful  ooservation  in  the  great 
laboratory  of  nature.  The  leakage  of  the  oceans  involves  three 
important  questions:  (i)  The  porosity  of  thick  layers  of  matter 
such  as  those  composing  the  earth's  crust;  (2)  the  penetrability  of 
the  crust  under  steady  fluid  pressure,  by  which  the  capillary  forces 
are  made  to  aid  the  molecular  forces  producing  penetration  of  the 
fluid;  and  (3)  the  accumulation  of  stresses  depending  on  the  forma- 
tion of  steam  in  the  layers  just  beneath  the  earth's  crust. 

The  conditions  existing  in  nature  can  scarcely  be  approximated 
in  our  laboratories,  on  account  of  the  limitations  of  the  forces -at  our 
command,  but  so  far  as  experiments  throw  light  on  these  great 
questions,  the  evidence  tends  to  confirm  the  theory  of  ocean  leakage. 
The  well-known  experiments  of  Daubree,  showing  that  under  the 
action  of  capillary  forces  hot  water  will  penetrate  a  layer  of  sand- 
stone against  a  strong  counter  pressure  of  steam,  and  by  entering 
a  cavity  actually  increase  the  steam  pressure  on  the  further  side,  has 
been  justly  held  to  afford  evidence  of  the  leakage  of  the  earth's 
crust,  and  of  the  probable  mode  of  volcanic  activity.  If  such  action 
is  possible  in  a  minute  way,  it  may  easily  operate  on  a  vastly  greater 
scale  to  produce  the  shaking  of  the  crust  in  earthquakes,  together 
with  the  uplift  of  mountains  and  the  occasional  outbreak  of 
volcanoes. 

Now  if  the  rock  of  the  earth's  crust  is  at  all  as  porous  as  we 
generally  think,  the  constant  pressure  of  the  vertical  column  of 


160  SEE— FURTHER  RESEARCHES  ON  [April  34, 

water,  often  miles  deep,  resting  on  the  ocean  bed  must  tend  to 
force  the  fluid  deeper  and  deeper  into  the  bowels  of  the  earth.  A 
study  of  what  takes  place  on  our  earth  under  the  observed  con- 
ditions constitutes  therefore  one  of  the  grandest  problems  in  nat- 
ural philosophy. 

Indeed  it  may  be  said  that  the  great  laboratory  of  nature  has 
magnificent  experiments  constantly  going  on.  All  that  we  need  to  do 
is  to  interpret  these  experiments  correctly.  The  best  way  to  do 
this  is  to  select  phenomena  in  which  the  processes  are  so  clear  as  to  be 
free  from  doubt ;  after  we  have  found  the  law  of  the  phenomena  in 
cases  which  are  beyond  question,  we  may  then  generalize  and 
interpret  other  phenomena,  in  which  the  relations  are  not  so  obvious. 
By  gathering  principles  and  laws  from  cases  which  are  entirely 
clear,  and  working  by  degrees  to  understand  those  which  are  more 
obscure,  we  may  finally  arrive  at  the  true  processes  even  when  the 
operations  of  nature  are  quite  hidden  from  our  view. 

Laws  thus  established  by  observation  in  the  great  laboratory  of 
nature  will  obviously  hold  true  of  like  experiments  in  the  minute 
physical  laboratories  designed  by  man ;  and  by  noting  the  phenomena 
of  the  globe  we  may  extend  our  knowledge  of  the  universal  proper- 
ties of  matter  under  various  physical  conditions  often  more  extreme 
than  those  ordinarily  witnessed  at  the  surface  of  the  earth. 

§  2.  Heretofore  the  ocean  bottoms  have  been  assumed  to  be  water- 
tight,— The  earth's  crust  is  made  up  chiefly  of  sedimentary,  igneous 
and  granitic  rocks,  and  soil  produced  by  the  decomposition  of  the 
various  kinds  of  rock  under  the  action  of  water  and  the  atmosphere. 
Nearly  all  of  the  sedimentary  rocks  are  quite  leaky,  and  moreover 
they  absorb  a  great  deal  of  moisture  from  the  air;  the  formation  of 
artesian  wells  and  of  natural  springs  depends  primarily  upon  the 
percolation  of  water  through  rocks  and  layers  of  soil  of  various 
kinds.  The  leaky  character  of  the  sedimentary  rocks  is  well  known 
and  has  been  generally  recognized.  But  these  rocks  exist  only 
near  the  surface,  and  do  not  extend  more  than  a  very  few  miles 
deep;  consequently  they  could  admit  the  water  to  but  a  slight  depth 
into  the  earth's  interior.  Below  the  sedimentary  rocks  lies  the 
mass  of  granite  which  makes  up  by  far  the  greater  part  of  the 
earth's  crust.     The  granitic  rocks,  such  as  granite,  andesite,  dia- 


,9o8.]  THE  PHYSICS  OF  THE   EARTH.  161 

base,  etc.,  are  by  no  means  so  penetrable  as  the  sedimentary  rocks, 
and  hence  water  has  more  difficulty  in  passing  through  them.  And 
as  the  layers  of  this  material  composing  the  earth's  crust  aie 
about  fifteen  miles  deep,  it  has  been  generally  held  that  water 
would  have  difficulty  in  making  its  way  down  into  the  heated 
layers  just  beneath  the  crust. ^  Indeed  it  has  been  practically  as- 
sumed that  the  ocean  bottoms  are  water-tight,  in  spite  of  the 
great  fluid  pressure  constantly  exerted  by  the  mere  depths,  of 
the  water  over  a  large  part  of  the  bottom  of  the  sea.  This  fluid 
pressure  in  many  places  is  great  enough  to  throw  a  column  of  water 
to  the  free  surface,  over  five  miles  high;  and  it  operates  not  only 
from  day  to  day,  year  to  year,  but  also  from  century  to  century, 
age  to  age.  If  granite  is  at  all  penetrable  by  water,  is  it  therefore 
any  wonder  that  a  gradual  secular  leakage  should  go  on,  and  at 
length,  by  a  kind  of  slow  perspiration  of  the  stone,  give  rise  to 
sufficient  accumulation  of  steam  beneath  the  crust  to  produce  a 
swelling  of  the  saturated  mass,  and  require  a  readjustment  of  the 
overlying  rocks? 

Now  it  happens  that  by  nature  all  the  granitic  rocks  are  crystal- 
line, and  thus  somewhat  coarse-grained  in  structure ;  so  that  they 
absorb  water  from  the  ground  and  moisture  from  the  air.  The 
crystalline  structure  permits  penetrability  to  a  greater  degree  than 
would  fine-grained  and  very  hard  rocks  such  as  agate;  but  no  rock 
has  such  fine  pores  as  the  metals,  and  especially  vitreous  bodies  like 
glass,  to  which  agate  is  an  approximation.  And  as  ^11  the  metals 
are  proved  by  experiment  to  be  leaky  under  great  fluid  pressure,  and 
glass  is  shown  to  obey  the  same  law,  it  obviously  follows  that  all 
rocks  are  leaky  under  great  fluid  pressure.  Consequently  under  the 
incessant  pressure  of  the  oceans  water  must  make  its  way  into  the 
heated  layer  just  beneath  the  earth's  crust- 
Heretofore  the  possibility  of  earthquakes  cracking  the  ocean 
bottom  has  been  generally  recognized,  but  it  has  been  held  that 

*This  statement  is  perhaps  too  positive,  for  Sir  William  Ramsay,  the 
celebrated  British  Chemist  and  Physicist,  writes  me  that  he  has  long  be- 
lieved that  the  ocean  bottom  leaks  and  that  the  formation  of  minerals 
takes  place  chiefly  in  the  bed  of  the  sea.  Undoubtedly  this  view  will  come 
to  be  generally  accepted.  Similar  views  seem  to  be  held  by  Lord  Rayleigh, 
Sir  Wm.  Huggins,  Arrhenius  and  many  other  eminent  physicists. 


162  SEE— FURTHER   RESEARCHES  ON  [April  .4, 

crevices  thus  formed  would  not  extend  over  five  or  six  miles  deep 
before  they  would  be  closed  by  the  effects  of  pressure,  which  nat- 
urally increases  rapidly  as  we  descend  into  the  earth.  The  belief 
has  therefore  prevailed  that  although  the  bed  of  the  sea  might  be 
rent  by  an  earthquake,  it  would  immediately  close  up  again,  and 
water  would  thus  be  prevented  from  entering  the  bowels  of  the  globe. 

It  scarcely  seems  to  have  occurred  to  investigators  to  consider 
the^  effects  of  the  constant  hydrostatic  pressure  resulting  from  the 
depth  of  the  sea,  in  forcing  the  water  slowly  through  the  fifteen 
miles  of  granite  composing  the  earth's  crust.  A  crevice  is  small, 
and  would  let  in  but  little  water  when  closed  up  quickly;  but  the 
whole  sea  bottom  is  large,  and  unless  it  is  really  water-tight,  even 
a  slow  leakage  over  a  large  area  would  at  length  develop  stresses 
beneath  which  would  necessitate  a  readjustment  of  the  overlying 
blocks  of  the  crust.  This  readjustment  is  ordinarily  called  an 
earthquake.^ 

Moreover  the  great  abundance  of  submarine  earthquakes  has  been 
largely  overlooked  by  previous  investigators.  It  is  the  secular 
effect  of  the  constant  pressure  of  the  oceans  and  of  capillary  forces 
in  promoting  the  downward  movement  of  the  water  which  has  been 
generally  lost  sight  of. 

But  if  we  admit  on  the  basis  of  experimental  evidence  that 
water  can  penetrate  thin  layers  of  granite,  the  question  naturally 
arises:  Can  it  also  penetrate  a  layer  of  granite  fifteen  or  twenty 
miles  thick?  •  It  seems  obvious  that  it  can,  because  for  small  or 
moderate  pressures  water  is  nearly  incompressible  and  would  not 
sensibly  increase  in  density  as  it  went  down  into  the  globe.  The 
fluid  which  passed  through  the  upper  layer  of  granite  would  there- 
fore keep  on  descending,  under  the  increasing  fluid  pressure  from 
above,  and  at  length  the  whole  layer  would  be  saturated,  and  per- 
spiring below  with  a  steady  leakage  which  would  give  rise  to  tre- 
mendous steam  power  in  the  imderlying  molten  rock.  Thus  great 
stresses  due  to  slow  accumulation  of  steam  would  develop  in  the 
layer  just  beneath  the  crust,  and  this  would  give  rise  to  earthquakes 
and  mountain  formation. 

Among  the  practical  men  of  science  to  whom  the  problem  of 
ocean  leakage  was  submitted,  we  might  name  some  of  the  most 


x9o8.]  THE  PHYSICS  OF  THE   EARTH.  163 

eminent  of  living  physicists.  While  disclaiming  especial  authority 
to  pass  upon  such  a  question,  they  expressed  the  opinion  that  it  was 
very  improbable  that  the  ocean  bottom  could  be  water-tight,  unless 
the  nature  of  the  rock  was  greatly  modified  by  pressure,  which  could 
hardly  be  the  case  in  the  first  twenty  miles  of  the  earth's  crust,  where 
the  pressure  does  not  exceed  8,600  atmospheres. 

Whatever  doubt  might  attach  to  this  solution  of  the  problem, 
from  an  experimental  standpoint,  where  positive  knowledge  is  greatly 
lacking,  seems  to  be  dispelled  by  the  phenomena  noticed  in  the 

sea  bottom  in  various  places,  which  show  that  lava  is  expelled  from 

•  

beneath  the  sea  and  pushed  under  the  adjacent  land.  The  phe- 
nomena noticed  in  the  laboratory  of  nature  thus  prove  the  leakage 
of  the  ocean  from  an  observational  standpoint,  because  they  admit 
of  no  other  interpretation. 

§  3.  The  Theory  of  Water-tightness  of  the  Ocean  Bottoms  Dis- 
proved by  the  Expulsion  of  Lava  from  under  the  Sea, — ^Just  south 
of  the  Aleutian  Islands,  a  long,  narrow  and  deep  trench  just  parallel 
to  this  chain  has  been  dug  out  by  the  expulsion  of  lava  from  beneath 
the  sea.  The  nature  of  this  trough  is  illustrated  by  the  accompanying 
Map. 

It  will  be  seen  that  the  island  chain  adjacent  to  the  trough  dug 
out  in  the  sea  bottom  is  really  a  mountain  range  under  water,  with 
only  occasional  peaks  projecting  above  the  water  as  islands.  In 
fact  the  Aleutian  Islands  are  a  continuation  of  the  Alaskan  Moun- 
tains which  are  part  of  the  Rocky  Mountain  System,  and  the  range 
here  continues  into  the  sea.  If  therefore  the  Aleutian  Islands  are 
mountains  now  in  process  of  formation  in  the  sea,  it  would  seem  to 
follow  logically  that  the  Rocky  Mountains  and  Andes,  from  Alaska 
to  the  straits  of  Magellan,  were  formed  in  the  same  way.  What 
then  is  the  process  at  work  forming  the  Aletitian  Islands  ? 

It  is  evident  that  the  deep  trench  south  of  the  islands  has  been 
dug  out  by  the  expulsion  of  lava  from  under  the  sea  and  its  injec- 
tion under  the  Aleutian  ridge;  this  is  accomplished  by  earthquakes, 
and  the  process  is  still  in  full  operation  at  the  present  time.  This 
region  is  a  well-known  breeding-ground  for  world-shaking  earth- 
quakes and  seismic  sea  waves.  Several  islands  have  been  uplifted 
since  1783,  and  one  or  more,  new  volcanoes  have  broken  out  within 


164  SEE— FURTHER   RESEARCHES  ON  [April  24, 

the  historical  period.  The  seismic  sea  waves  following  the  earth- 
quakes which  affect  this  region  indicate  that  the  sea  bottom  often 
sinks  after  these  disturbances.  In  other  words,  when  lava  is  ex- 
pelled from  under  the  trench  and  pushed  under  the  adjacent  ridge, 
the  bottom  gives  down  to  secure  stability.  The  processes  now 
going  on  have  been  at  work  through  immense  ages,  and  have  thus 
dug  out  the  trough  parallel  to  the  Aleutian  Islands,  and  at  the  same 
time  elevated  this  ridge,  till  it  is  now  partly  above  the  water,  thus 
constituting  the  chain  of  islands. 

In  like  manner  the  great  earthquake  at  Yakutat  Bay,  farther 
east,  September  3-20,  1899,  which  was  so  carefully  investigated  by 
Tarr  and  Martin  (Bulletin  of  the  Geological  Society  of  America, 
May,  1906)  raised  the  coast  for  about  100  miles;  the  maximum 
elevation  being  47 J^  feet.  Subsidence  also  occurred  in  a  few  places. 
Such  a  vast  movement  of  the  coast  indicates  an  enormous  expulsion 
of  molten  rock  from  beneath  the  sea  under  the  land.  It  is  these  sub- 
terranean movements  beneath  the  earth's  crust  which  shake  down 
cities  and  devastate  whole  countries.  During  the  earthquake  at 
Yakutat  Bay  the  shaking  was  so  terrible  that  persons  could  not 
stand  on  their  feet;  avalanches  slid  down  the  mountains,  and  gla- 
ciers were  carried  into  the  sea.  This  is  the  true  nature  of  earth- 
quakes, and  one  need  not  therefore  be  surprised  at  the  devastation 
produced.  The  force  which  pushes  lava  under  the  land,  overcoming 
the  weight  of  the  crust,  naturally  destroys  cities  and  all  the  frail 
works  of  man  built  upon  the  surface. 

§  4.  Physical  Experiments  on  the  Porosity  of  Matter, — Modem 
science  presents  many  illustrations  of  the  porosity  of  matter.  In 
fact  so  many  experiments  illustrate  porosity  that  it  is  difficult  to 
find  proof  of  the  general  property  of  impenetrability  cited  by  New- 
ton in  the  "  Principia,"  except  under  the  narrow  limitations  that 
the  matter  in  question  remains  cold  and  the  forces  to  which  it  is 
subjected  are  small.  With  increasing  fluid  pressure  and  rising 
temperature  all  matter  is  leaky ;  and  in  general  a  rise  of  temperature 
expands  and  thus  augments  the  penetrability  and  porosity  of  all  sub- 
stances. We  may  therefore  say  that  all  matter  is  porous  and  leaky 
under  great  fluid  pressure,  and  impenetrability  does  not  exist  except 


,9o8.]  THE  PHYSICS  OF  THE   EARTH.  185 

under  very  restricted  conditions,  so  thai  it  is  not  a  general  property 
of  matter  as  was  once  supposed. 

In  the  early  days  of  physical  science  the  demonstration  of  the 
porosity  of  such  dense  bodies  as  gold,  silver  and  lead  was  considered 
a  great  achievement.  In  1661  some  academicians  at  Florence,  re- 
peating an  earlier  experiment  of  Bacon  with  a  spherical  shell  of 
lead,  filled  a  hollow  sphere  of  solid  gold  with  water,  and,  after 
sealing  it  hermetically,  flattened  the  figure  of  the  spherical  shell  in  a 
hydrostatic  press  so  as  to  diminish  the  volume.  Under  this  deforma- 
tion of  the  sphere  the  water  was  forced  through  the  walls  of  solid 
gold  and  formed  in  drops  on  the  outside.  Corresponding  experi- 
ments were  made  with  spheres  of  silver,  lead,  and  other  metals, 
with  analogous  results.  Modem  engineering  presents  innumerable 
illustrations  of  the  porosity  and  leaky  character  of  structures  made  of 
the  hardest  bodies.  Under  great  pressure  all  pipes  and  pistons  leak, 
and  put  a  limit  to  the  applications  of  hydrostatic  pressure. 

In  1883  Amag^t  forced  mercury  through  plates  of  solid  steel 
three  inches  thick,  under  a  pressure  of  about  4,000  atmospheres. 
This  is  the  highest  pressure  hitherto  applied  in  physical  experiments, 
and  yet  all  rocks  are  subjected  to  such  pressure  at  a  depth  of  only 
ten  miles  below  the  earth's  surface.  In  the  measurement  of  ocean 
depths  it  has  been  found  that  empty  hollow  glass  balls  with*  walls 
half  an  inch  thick  sent  down  with  the  deep  sea  apparatus  come 
up  more  and  more  completely  filled  with  water,  according  to  the 
depth  of  the  sea  and  the  duration  of  the  experiment.  As  glass  is 
the  most  impervious  of  solid  bodies,  this  leakage,  which  it  shows 
under  the  external  application  of  fluid  pressure  from  the  deep  sea, 
is  a  good  illustration  of  what  happens  to  the  bed  of  the  ocean,  which 
is  constantly  subjected  to  this  pressure.  No  rock  is  anything  like 
so  impervious  as  glass,  and  consequently  a  general  leakage  of  the 
ocean  bottom  inevitably  takes  place.  The  water  which  first  enters 
the  bed  of  the  sea  will  keep  on  descending  till  it  comes  into  contact 
with  rock  at  high  temperature,  which  produces  and  readily  absorbs 
steam.  When  the  rock  becomes  saturated  with  steam  it  swells  and 
requires  more  space,  and  this  finally  brings  on  an  earthquake. 
Hence  also  the  preponderance  of  great  earthquakes  under  the  sea 
and  the  almost  total  absence  of  these  disturbances  far  inland. 


166  SEE— FURTHER   RESEARCHES  ON  [April  24, 

§  5.  Important  Criterion  for  the  Nature  of  the  Movement  Be- 
neath the  Earth's  Crust  furnished  by  Seismic  Sea  Waves. — In  the 
paper  on  the  "  Cause  of  Earthquakes "  we  divided  seismic  sea 
waves  into  two  general  classes :  the  first,  due  to  the  sinking  of  the 
sea  bottom,  and  characterized  by  a  withdrawal  of  the  water  after 
the  earthquake,  to  be  followed  later  by  the  return  of  a  g^eat  wave; 
the  Second,  due  to  the  uplift  of  the  bottom,  and  characterized  by 
the  sudden  rise  of  the  sea  without  any  previous  withdrawal  from 
the  shore.  Both  classes  of  these  waves  exist  in  our  seas,  but  those 
of  the  first  class  are  the  most  dangerous  and  the  most  important. 
Most  of  the  great  historical  inundations  by  the  sea  have  been  due 
to  waves  of  the  first  class.  The  phenomena  usually  noted  are: 
first,  a  terrible  earthquake;  second,  after  a  short  interval,  the  sea 
is  noticed  to  be  slowly  draining  away,  laying  bare  the  bottom, 
where  it  ordinarily  is  deep  enough  to  anchor  ships;  third,  after  an 
interval  of  an  hour  or  so,  the  sea  is  seen  to  be  returning  as  a 
mighty  wave,  carrying  everything  before  it,  and  thus  washing  the 
ships  inland  and  stranding  them  high  and  dry ;  fourth,  having  once 
swept  the  shore,  the  sea  again  withdraws  and  lays  bare  the  harbor 
as  before,  and  after  about  the  same  interval  again  returns  as  a 
second  great  wave.  This  periodic  movement  of  the  sea  may  be  kept 
up  for  quite  a  while,  and  sometimes  quiet  is  not  restored  for  a 
day  or  two. 

Among  the  many  well-known  historical  sea  waves  of  the  first 
class  which  might  be  mentioned,  we  shall  cite  only  a  few  typical 
cases:  As  that  which  overwhelmed  Helike  in  373  B.  C.  (see  the 
paper  on  the  "  Temperature  of  the  Earth,"  §  23,  pp.  269-272,  and 
Addendum,  pp.  291-298) ;  the  wave  at  Callao  in  1746;  the  wave 
following  the  Lisbon  earthquake  in  1755 ;  the  waves  of  Arica,  1868, 
and  Iquique,  1877;  the  wave  on  the  Japanese  coast  in  1896.  In  all 
these  cases  the  water  first  withdrew  from  the  shore;  not  suddenly, 
but  slowly,  as  in  the  draining  away  of  a  tide,  though  somewhat 
more  rapidly;  this  of  course  indicated  that  the  sea  bottom  had 
sunk,  and  the  water  was  draining  away  to  fill  up  the  depression  in 
the  level  caused  by  the  drop  of  the  bottom.  When  the  currents  meet 
at  the  center  an  elevation  is  produced  by  their  mutual  impact,  and 


I9081  THE   PHYSICS  OF  THE   EARTH.  167 

when  this  collapses  under  gravity  the  first  great  wave  comes 
ashore.  The  elevation  then  subsides  into  a  depression  as  at  first, 
and  the  currents  again  flow  in  and  force  up  the  level  a  second 
time;  and  with  the  second  collapse  another  wave  is  sent  ashore; 
and  so  the  oscillation  of  the  sea  continues,  sometimes  for  a  day  or 
two  before  it  finally  quiets  down. 

Now  these  sea  waves  of  the  first  class  furnish  an  exceedingly 
important  criterion  as  to  the  nature  of  what  is  going  on  beneath  the 
earth's  crust.  The  sinking  of  the  sea  bottom  often  happens  in 
the  deep  trench  south  of  the  Aleutian  Islands,  and  repeated  drops 
of  this  kind  have  obviously  produced  the  deep  trough  parallel  to 
these  islands.  For  it  is  observed  that  the  earthquake  usually  raises 
one  or  more  of  the  islands  to  the  north,  when  the  sea  bottom  sinks 
to  the  south.  Now  the  islands  could  not  be  upraised  unless  some- 
thing was  pushed  under  them,  and  the  bed  of  the  trough  could  not 
sink  down  unless  it  was  in  some  way  undermined.  Accordingly  it 
follows  that  molten  rock  is  expelled  from  beneath  the  bed  of  the 
trough  to  the  south  and  pushed  under  the  adjacent  islands  to  the 
north,  which  are  thus  uplifted.  The  bed  of  the  sea  often  sinks  during 
the  earthquake  arising  from  this  subterranean  movement,  and  then 
the  water  withdraws  from  the  shore  and  afterwards  returns  as  a 
g^eat  seismic  sea  wave.  It  will  be  observed  that  the  subcrustal  move- 
ment is  from  the  sea  towards  the  land,  because  steam  accumulates 
under  the  ocean,  but  scarcely  at  all  under  the  land. 

Thus  these  seismic  sea  waves  become  very  important  criteria 
for  determining  whether  the  sea  bottom  has  sunk;  and  if  it  has 
sunk  we  know  that  lava  was  expelled  from  under  the  sea  and  pushed 
under  the  land.  Seismic  sea  waves  therefore  may  be  regarded  as 
very  delicate  levels,  for  determining  the  movement  of  the  sea  bot- 
tom; and  from  the  nature  of  this  movement  we  can  often  decide 
what  the  effect  of  the  earthquake  has  been.  Moreover  these  waves 
enable  us  to  tell  with  certainty  that  the  chief  function  of  earth- 
quakes is  the  elevation  of  the  land  along  the  coast  by  the  expulsion 
of  lava  from  beneath  the  bed  of  the  sea.  It  is  not  too  much  to  say 
that  the  true  nature  of  earthquakes  and  their  function  in  the  uplift 
of  mountains  and  plateaus  could  not  be  certainly  made  out  except  for 
the  exceedingly  important  criterion  furnished  by  ^ismic  sea  waves. 


168  SEE— FURTHER   RESEARCHES  ON  [April  24, 

§  6.  Additional  Phenomena  Noticed  near  the  Aleutian,  Kurile 
and  Japanese  Islands,  and  the  Antandes, — South  of  the  Aleutian 
chain,  as  just  remarked,  a  well-known  earthquake  belt  parallels  these 
islands,  and  the  seismic  disturbances  occurring  there  are  frequently 
followed  by  seismic  sea  waves  of  the  first  class.  Soon  after  a  great 
earthquake  the  water  is  seen  to  be  withdrawing  from  the  shore,  and 
after  a  short  interval  of  time  it  again  returns  as  a  mighty  wave 
sweeping  everything  before  it.  Many  volcanoes  have  broken  out  in 
these  islands  and  several  new  islands  have  been  uplifted  within  the 
historical  period.  The  Russians  long  ago  connected  the  earthquakes 
with  the  volcanoes  in  the  Aleutian  Islands.  In  later  years  the 
exact  survey  of  the  sea  bottom  has  shown  that  it  is  sunk  down  into 
a  narrow  trough  right  under  the  earthquake  belt.  Just  parallel  to 
the  trough  the  islands  form  a  real  mountain  ridge  under  water,  with 
only  a  few  of  the  highest  points  projecting  above  the  surface  as 
islands.  The  uplift  of  these  islands  therefore  denotes  the  uplift 
of  mountain  peaks,  some  of  which  have  become  volcanoes. 

Now  if  the  earthquakes  are  accompanied  by  the  uplift  of  islands 
and  the  sinking  of  the  sea  bottom,  as  shown  by  the  seismic  sea 
waves,  it  follows  that  the  uplift  of  the  ridge  is  connected  with  the 
sinking  of  the  adjacent  sea  bottom.  As  the  ridge  is  just  contiguous 
to  the  trench,  and  the  earth  is  terribly  shaken  every  time  these  dis- 
turbances occur,  it  seems  to  indicate  that  matter  is  expelled  from 
under  the  trench  and  pushed  under  the  ridge;  so  that  the  ridge  is 
elevated  and  the  trench  sinks  down  correspondingly.  This  could 
not  occur  without  the  bodily  transfer  of  matter  beneath  the  earth's 
crust,  and  the  shaking  of  the  earth  is.  due  to  this  expulsion  of  lava 
from  under  the  trench,  and  its  injection  under  the  ridge.  This  is 
the  only  possible  explanation  of  the  observed  elevation  of  the  ridge 
and  the  sinking  of  the  trench.  In  this  way  the  trough  near  the 
Aleutian  Islands  has  been  gradually  dug  out.  Similar  troughs  have 
been  formed  by  earthquakes  near  the  Kurile  and  Japanese  Islands, 
as  we  know  by  the  observed  depth  of  the  sea,  the  lay  of  the  earth- 
quake belt  parallel  to  these  islands,  and  the  occurrence  of  the  seismic 
sea  waves,  showing  that  the  sea  bottom  sinks  after  the  earthquakes  j 

by  which  the  region  is  afflicted.    If  the  islands  of  Japan  were  dug  | 

off  and  thrown  into  the  Tuscarora  Deep,  they  would  about  fill  it  up.  ! 


Y    . 


•' 


,9o8.]  THE  PHYSICS  OF  THE  EARTH.  169 

Therefore  all  these  islands  were  formed  by  the  expulsion  of  lava 
from  under  the  sea,  and  the  subsequent  sinking  of  the  sea  bottom 
has  given  rise  to  the  deep  troughs  now  found  in  that  part  of  the 
ocean. 

In  the  same  way  there  is  an  earthquake  belt  between  Samoa 
and  New  Zealand,  and  the  sea  bottom  is  sunk  down  into  a  deep 
trough,  parallel  to  a  ridge  on  the  west,  on  the  opposite  side  of  the 
trench  from  the  ocean.  This  ridge  is  a  new  mountain  range  1,200 
to  1,500  miles  long,  now  forming  on  the  west  of  the  Pacific,  just 
as  tlhe  Andes  were  once  formed  on  the  east.  Lava  is  being  expelled 
from  under  the  trench  and  pushed  from  the  ocean  towards  the  ridge 
on  the  west.  This  is  developing  into  a  new  mountain  range,  which 
we  shall  call  the  Antandes,  because  it  is  being  formed  opposite  to 
the  Andes,  on  the  other  side  of  the  Pacific,  and  in  the  same  manner 
as  the  mighty  mountains  in  South  America  were  in  earlier  geological 
time.  In  the  course  of  immense  ages  the  Antandes  will  rise  above 
the  water  as  a  mighty  chain  on  the  west  of  the  Pacific  just  like  the 
Andes  on  the  east. 

These  phenomena  in  the  sea  bottom  show  the  real  procfss  of 
mountain  formation  at  various  stages  of  its  progress,  and  prove  to 
us  that  most  of  the  folding  observed  in  our  mountain  ranges  now 
on  land  really  took  place  in  the  bed  of  the  sea,  long  before  the  whole 
range  was  raised  above  the  water.  For  this  sinking  and  upheaval 
of  adjacent  portions  of  the  sea  bottom  would  crumple  the  rocks 
exactly  as  they  are  observed  to  be  in  all  mountain  ranges;  and 
moreover  the  several  parallel  ranges  so  often  observed  would  result 
from  the  development  of  several  parallel  troughs,  all  of  which  are 
eventually  uplifted.  It  will  be  observed  that  the  expulsion  of  lava 
is  always  from  the  sea  towards  the  land,  and  this  shows  that  the 
sole  cause  of  the  movement  is  the  leakage  of  the  ocean.  It  thus 
follows  that  mountains,  plateaus,  and  islands  are  uplifted  by  earth- 
quakes depending  on  the  leakage  of  the  oceans,  and  by  nothing  else. 

§  7.  The  Andes  with  their  high  Plateaus  Merely  a  Vast  Wall 
Erected  by  the  Pacific. — It  may  sound  strange  to  say  that  the  Cordil- 
lera of  the  Andes  is  a  vast  wall  erected  by  the  Pacific  Ocean  along 
its  border ;  but  to  the  navigator  who  traverses  the  shore  from  Pan- 
ama to  Cape  Horn  such  a  description  will  seem  most  appropriate. 


SEE— FURTHER   RESEARCHES  ON  [April 


Fig.  I.  Relief  Map  of  South  America.  (From  Frye's  Complete  Geog- 
raphy, by  permission  of  Ginn  &  Co.,  Publishers.)  Notice  that  the  Andes  are 
a  mighly  wall  erected  by  the  Pacific  Ocean  along  its  border.  Professor 
Charles  Burkhalter,  Director  of  the  Chabot  Observatory,  Oakland,  kindly 
suggested  the  use  of  these  relief  maps,  which  are  well  suited  for  bringing 
out  the  leading  characteristics  of  the  different  continents. 


,9o8]  THE  PHYSICS  OF  THE  EARTH.  171 

Throughout  the  length  of  the  continent  the  mountains  are  every- 
where parallel  to  the  coast,  and  run  at  nearly  a  constant  distance 
from  the  shore.  The  Andes  are  not  always  a  simple  chain,  but  they 
are  narrow  relatively  to  their  height,  as  compared  to  the  other 
moimtains.  In  many  places  there  are  two  or  more  ranges  with 
narrow  plateaus  between.  These  plateaus  are  so  interwoven  with 
the  mountains  themselves  that  we  may  feel  sure  they  were  formed 
together  and  represent  a  part  of  one  general  movement.  Unless 
this  were  so  it  is  impossible  to  believe  that  so  many  narrow  and  high 
plateaus  would  be  enclosed  between  mountain  walls  on  either  side. 
The  eastern  cordillera  is  less  volcanic  than  the  western,  and  the 
eastern  slope  is  believed  by  Professor  Solon  I.  Bailey  of  Harvard 
College  Observatory,  who  has  exceptional  opportunities  for  judging 
of  these  mountains,  to  be  two  or  three  times  steeper  than  the  western 
slope. 

If  we  suppose  a  sea  trough  was  first  dug  out  in  the  elevation  of 
the  easterh  range,  and  eventually  when  deep  sediments  had  ac- 
cumulated in  the  trough,  the  western  edge  of  it  was  folded  up  to 
form  the  western  range,  and  the  trough  itself  became  the  plateaus, 
we  shall  have  very  nearly  a  true  picture  of  how  the  Andes  were 
formed.  The  full  details  of  this  process  cannot  be  given  now,  but 
there  is  no  doubt  that  the  Andes  are  a  vast  wall  erected  by  the 
Pacific  along  the  edge  of  the  continent.  This  origin  of  these 
mountains  is  also  indicated  by  the  earthquakes  observed  within 
historical  time;  for  the  coast  has  been  again  and  again  upraised 
by  these  disturbances,  while  the  sinking  of  the  sea  bottom,  indicated 
by  the  accompanying  seismic  sea  waves,  shows  that  the  bed  of  the 
ocean  is  being  undermined  by  the  expulsion  of  lava  under  the  land. 
The  shells,  fossils,  and  other  evidences  of  marine  life  now  found 
at  altitudes  as  high  as  15,000  feet  show  that  the  uplifting  at  present 
going  on  is  but  a  part  of  the  greater  uplift  of  past  geological  ages ; 
so  that  the  great  movement  which  formed  these  mountains  and  pla- 
teaus is  identical  with  the  earthquake  disturbances  noticed  within 
historical  time. 

§  8.  The  uplift  of  mountains  and  plateaus  around  the  margins 
of  the  Pacific,  and  of  islands  in  the  interior,  with  innumerable  sub- 
marine eruptions  everywhere,  is  nature's  way  of  indicating  leakage 


172  SEE— FURTHER   RESEARCHES  ON  [April  .4. 

through  twenty  miles  of  crust, — ^The  peculiar  position  of  the  sea 
bottom  between  a  molten  globe  and  the  overlying  ocean  is  such  that 
any  disturbance  of  the  bottom,  as  in  a  volcanic  eruption,  would 
naturally  excite  our  suspicions  that  the  crust  had  leaked  and  brought 
the  water  into  contact  with  the  underlying  ball  of  fire.  The  situa- 
tion of  the  overlying  ocean,  with  the  fire  so  close  beneath,  is  much 
the  same  as  that  of  the  water  above  the  furnace  of  a  boiler,  in  which 
steam  is  developed;  and  if  one  had  the  molten  globe  for  a  furnace 
and  the  ocean  for  a  reservoir  of  water,  leakage  would  develop  steam 
on  a  grand  scale,  and  give  rise  to  mighty  experiments  exactly  resem- 
bling earthquakes  and  volcanic  eruptions.  Some  of  these  disturb- 
ances might  take  the  form  of  uplifts  of  the  crust  into  islands, 
mountains  and  submarine  volcanoes,  others  near  the  edges  of  the 
sea  would  cause  lava  to  push  out  under  the  land  and  raise  the  coasts. 

Now  the  Pacifip  Ocean  is  everywhere  surrounded  by  high  moun- 
tains, as  if  the  lava  had  been  pushing  out  at  the  margins  of  the  sea. 
,  And  throughout  the  interior  a  vast  number  of  islands  are  raised  up 
in  deep  water,  and  every  part  of  the  ocean  is  from  time  to  time  dis- 
turbed by  terrible  earthquakes.  One  must  therefore  admit  that  the 
ocean  has  the  aspect  which  might  be  expected  to  result  from  a  leak- 
age of  the  ocean  bed.  Moreover  the  Pacific  is  surrounded  nearly 
everywhere  by  volcanoes,  which  emit  chiefly  vapor  of  steam.  If  it 
is  shown  that  mountains  are  formed  by  earthquakes,  chiefly  in  the 
expulsion  of  lava  under  the  land,  and  some  of  the  mountains  break 
out  into  volcanoes,  then  there  will  obviously  be  a  connection  not 
only  between  earthquakes  and  volcanoes,  but  also  between  the  vapor 
of  steam  emitted  from  these  smoking  mountains  and  that  formed 
under  the  ocean  by  the  leakage  of  the  bottom. 

It  is  this  intimate  connection  between  all  the  related  phenomena 
which  tells  so  powerfully  in  favor  of  the  view  that  the  leakage  of 
the  ocean  takes  place  through  a  layer  of  rock  twenty  miles  thick. 
The  height  of  the  mountains  and  plateaus  is  but  a  small  fraction 
of  the  thickness  of  the  crust,  and  movement  in  the  underlying  layers 
therefore  usually  gets  relief  without  breaking  through.  The  crust 
of  the  globe  is  thick  enough  to  oflFer  great  resistance  to  uplift,  so 
that  the  steam  saturated  lava  usually  adjusts  itself  beneath  without 
a  surface  eruption.    Yet  where  the  crust  is  sharply  upheaved  as  in 


i9o8.]  THE  PHYSICS  OF  THE  EARTH.  173 

mountains,  volcanoes  sometimes  break  out.  But  it  is  obvious  that 
earthquakes  are  the  more  general,  volcanoes  the  more  special  phe- 
nomena ;  and  that  both  are  connected  with  mountain  formation,  and 
depend  on  the  sea  for  their  continued  activity. 

§  9.  On  the  Structure  of  Granite  as  a  Typical  Crystalline  Rock 
of  the  Earth's  Crust. — Granite  has  a  thoroughly  crystalline  structure, 
and  is  an  admixture  of  feldspar,  mica,  and  quartz.  The  mica  is  in 
the  form  of  minute  shingles,  or  snowflakes,  embedded  in  the  non- 
crystalline matrix  of  quartz,  which  encloses  the  other  elements.  The 
feldspar  is  chiefly  orthoclase.  The  two  chief  ingredients,  quartz  and 
feldspar,  form  a  granular  aggregate  made  up  of  grains  of  fairly 
equal  size,  varying  all  the  way  from  several  inches  in  diameter  to  a 
structure  so  fine  as  to  be  inseparable  to  the  naked  eye. 

"  Many  granites  contain  irregularly  shaped  cavities  (miarolitic  structure), 
in  which  the  component  minerals  have  had  room  to  crystallize  in  their 
proper  forms,  and  where  beautifully  terminated  crystals  of  quartz  and 
felspar  may  be  observed.  It  is  in  these  places  also  that  the  accessory 
minerals  (beryl,  topaz,  tourmaline,  garnet,  orthite,  zircon  and  many  others) 
are  found  in  their  best  forms.  Not  improbably  these  cavities  were  some- 
what analogous  to  the  steam  holes  of  amygdaloids,  but  were  filled  with  water 
or  vapour  of  water  at  high  temperature  and  under  great  pressure,  so  that 
the  constituents  could  crystallise  under  the  most  favorable  conditions. 
Among  the  component  minerals  of  granite,  the  quartz  presents  a  special 
interest  under  the  microscope.  It  is  often  found  to  be  full  of  cavities  con- 
taining liquid,  sometimes  in  such  numbers  as  to  amount  to  a  thousand 
millions  in  a  cubic  inch  and  to  give  a  milky  turbid  aspect  to  the  mineral. 
The  liquid  in  these  cavities  appears  usually  to  be  water,  either  pure  or  con- 
taining saline  solutions,  sometimes  liquid  carbon-dioxide"  (p.  143).  (Sir  A. 
Geikie,  "  (Geology,"  p.  204.) 

The  cavities  in  crystalline  rock  such  as  granite  may  contain 
either  gas  or  liquid  matter,  and  sometimes  both.  Professor  Tilden's 
researches  have  shown  that  the  included  gases  (hydrogen,  carbon 
dioxide,  carbon  monoxide,  marsh  gas,  nitrogen,  and  water  vapor) 
may  exceed  many  times  the  volimie  of  the  rock  itself.  The  cavities 
have  all  manner  of  forms,  branching,  oblong,  curved,  oval,  spherical 
and  negative  crystalline  shapes,  and  are  often  so  numerous  as  to  give 
a  turbid  aspect  to  the  mineral.  The  intersecting  planes  of  the  crys- 
talline granite  frequently  present  real  fissures  more  or  less  filled  with 
liquid.  Obviously  capillary  forces  may  here  attain  great  importance, 
and  fluid  entering  the  rock  would  be  absorbed  into  these  spaces 

PROC.  AMKR.  PHIL.  SOC.  XLVII.  189  L,  PRINTED  SEPTEMBER  21,   I908. 


174  SEE-FURTHER   RESEARCHES  ON  [April  24, 

with  irresistible  power.  Geikie  remarks  that  the  cavities  in  quartz 
have  all  sizes  from  the  coarse  pores  visible  to  the  naked  eye  to 
minute  spaces  less  than  1/10,000  of  an  inch  in  diameter,  which  can  be 
seen  only  under  high  magnifying  power. 

Now  it  is  worth  while  to  remember  that  small  as  are  the  least 
cavities  and  fissures  which  we  can  see  with  the  microscope,  they  are 
very  large  and  coarse  compared  to  the  molecular  structure  of  a  fluid 
such  as  water  or  of  a  solid  like  glass.  It  is  useful  to  remember  that 
the  limit  of  naked  eye  vision  is  about  1/250  of  an  inch,  and  of  the 
most  powerful  microscope  about  1/100,000  of  an  inch.  The  micro- 
scope therefore  increases  our  power  of  vision  about  400  times. 
(Cf.  Prof.  A.  A.  Michelson's  "  Light  Waves  and  their  Uses,"  p.  30.) 

§  10.  On  Lord  Kelvin's  Determination  of  the  Size  of  Atoms, — 
Jfi  order  to  form  a  clear  conception  of  the  physical  constitution  of 
the  matter  composing  the  crust  of  the  globe,  we  must  recall  the  lines 
of  research  by  which  Lord  Kelvin  has  determined  the  size  of  atoms. 

1.  By  determining  the  work  done  or  heat  produced  in  bringing 
thin  plates  of  zinc  and  copper  together.  The  observed  amount  of 
heat  evolved  when  the  plates  are  made  of  given  thickness  and  after- 
wards imagined  to  be  thinner  and  thinner,  limited  only  by  the  con- 
dition that  the  mass  shall  not  be  melted,  under  the  heat  of  combina- 
tion, which  is  not  indefinitely  great  even  when  brass  is  produced  by 
fusing  zinc  and  copper,  but  corresponds  to  the  mutual  attraction  of 
a  number  of  plates  not  more  ntunerous  than  100,000,000  to  the  milli- 
meter;  hence  it  follows  that  the  molecules  are  at  least  1/1,000,000,000 
cm.  and  probably  more  than  1/400,000,000  cm.  in  diameter.  Lord 
Kelvin  concluded  that  "  Plates  of  zinc  and  copper  1/300,000,000  of 
a  centimeter  thick,  placed  close  together  alternately,  form  a  near 
approximation  to  a  chemical  combination  if  indeed  such  thin  plates 
could  be  made  without  splitting  atoms."  He  fixed  1/1,000,000,000 
of  a  centimeter  as  the  minimum  diameter  of  the  atoms  found  in  this 
way.    It  is  to  be  remembered  here  that  2.54  centimeters  ^  i  inch. 

2.  By  the  study  of  Newton's  rings  on  soap  bubbles  as  they 
become  thinner  and  thinner,  the  thickness  of  the  film  being  reckoned 
from  the  known  wave-length  of  the  reflected  light.  Unless  the 
film  materially  weakened  when  a  certain  limit  is  attained,  it  could 
not  be  stretched  beyond  a  certain  thickness  without  volatilizing,  if 


,9o8.]  THE  PHYSICS  OF  THE  EARTH.  176 

maintained  at  the  same  temperature ;  for  as  it  expands  it  cools,  and 
the  heat  that  would  have  to  be  supplied  to  it  would  be  more  than 
sufficient  to  vaporize  it.  Now  it  is  found  by  observation  that  the 
intensity  of  the  surface  tension  of  the  film  of  water  falls  off  before 
the  thickness  is  reduced  to  1/200,000,000  cm.,  and  hence  there  prob- 
ably are  but  few  molecules  in  that  thickness. 

3.  By  the  phenomenon  of  dispersion  in  the  wave  theory  of 
light.  Cauchy  showed  that  dispersion  of  colors  implied  a  granular 
structure  in  refracting  media,  and  that  the  grains  could  not  be 
indefinitely  small,  but  must  exceed  1/10,000  of  the  shortest  wave 
length;  and  to  produce  the  observed  effect  Lord  Kelvin  concluded 
that  the  number  of  molecules  in  a  wave  length  would  have  to  be 
from  200  to  600.  Nobert  ruled  lines  on  glass  at  the  rate  of  40,000 
to  the  centimeter,*  or  about  two  to  the  wave  length  of  blue  light 

(about  4/100,000  centimeter)  ;  and  as  this  left  the  ruled  surface 
capable  of  reflection,  the  number  of  molecules  in  the  ridges  between 
the  grooves  must  have  been  sufficient  to  give  solid  body  to  the 
sculptured  mass,  and  thus  not  less  than  several  hundred  to  the  wave 
length.  If  the  mean  free  path  in  a  solid  like  glass  be  25  times  the 
diameter  of  the  atom  itself,  this  will  make  the  diameter  of  the 
atoms  of  the  order  of  1/400,000,000  of  a  centimeter. 

4.  By  calculating  the  length  of  the  average  free  path  of  a  mole- 
cule in  a  gas,  according  to  the  kinetic  theory.  Loschmidt  in  1865, 
Stoney  in  1866,  and  Lord  Kelvin  in  1870,  independently  reached 
similar  results,  namely,  for  the  average  free  path  about  1/100,000 
of  a  centimeter,  and  for  the  diameter  of  the  gaseous  molecule  about 
1/500,000,000  of  a  centimeter. 

These  four  methods  of  estimating  the  diameter  of  atoms  thus 
agree  very  closely  among  themselves ;  and  moreover  a  similar  result 
on  the  average  distance  of  molecules  deduced  by  entirely  different 

'Referring  to  Nobert's  lines  Maxwell  says:  "A  cube,  whose  side  is  the 
400th  of  a  millimetre,  may  be  taken  as  the  minimum  visible  for  (microscopic) 
observers  of  the  present  day.  Such  a  cube  would  contain  from  60  to  100 
million  molecules  of  oxygen  or  nitrogen"  (cf.  The  article  "Atom,"  Ency- 
clopedia Britannica,  ninth  edition,  p.  42).  If  there  be  400  molecules  in  a 
line  the  length  of  the  edge  of  the  cube  just  considered,  the  cube  would  con- 
tain 64,000,000,  which  agrees  with  Maxwell's  estimate.  A  line  equal  to  the 
wave  length  of  blue  light  would  thus  contain  250  molecules. 


176  SEE— FURTHER   RESEARCHES  ON  [April  24, 

considerations  was  obtained  by  M.  Lippmann,  in  a  paper  read  to  the 
Paris  Academy  of  Sciences,  October  16,  1882. 

In  his  "  Popular  Lectures  and  Addresses  "  (vol.  i,  p.  224)  Lord 
Kelvin  condenses  his  conclusions  as  follows : 

"The  four  lines  of  argument  which  I  have  now  indicated  lead  all  to 
substantially  the  same  estimate  of  the  dimensions  of  molecular  structure. 
Jointly  they  established,  with  what  we  cannot  but  regard  as  a  very  high 
degree  of  probability,  the  conclusion  that,  in  any  ordinary  liquid,  transparent 
solid,  or  seemingly  opaque  solid,  the  mean  distance  between  the  centres  of 
contiguous  molecules  is  less  than  the  1/5,000,000  and  greater  than  the 
i/i>ooo,ooo,ooo  of  a  centimeter. 

"To  form  some  conception  of  the  degree  of  coarse-grainedness  indi- 
cated by  this  conclusion,  imagine  a  globe  of  water  or  glass,  as  large  as  a 
football,*  to  be  magnified  up  to  the  size  of  the  earth,  each  constituent  mole- 
cule being  magnified  in  the  same  proportion.  The  magnified  structure  would 
be  more  coarse  grained  than  a  heap  of  small  shot,  but  probably  less  coarse- 
grained than  a  heap  of  footballs?* 

§  II.  On  the  Molecular  Constitution  of  Matter  and  on  the  Pene- 
trability of  Solids  by  Fluids, — In  his  address  on  "  Mathematical 
Physics  "  at  the  St.  Louis  Congress  of  Arts  and  Sciences  in  1904, 
Poincare  speaks  of  the  porosity  of  matter  as  follows: 

"The  astronomical  universe  consists  of  masses,  undoubtedly  of  great 
magnitude,  but  separated  by  such  immense  distances  that  they  appear  to  us 
as  material  points;  these  points  attract  each  other  in  the  inverse  ratio  of  the 
squares  of  their  distances,  and  this  attraction  is  the  only  force  which  affects 
their  motion.  But  if  our  senses  were  keen  enough  to  show  us  all  the  details 
of  the  bodies  which  the  physicist  studies,  the  spectacle  thus  disclosed  would 
hardly  differ  from  the  one  which  the  astronomer  contemplates.  There  too 
we  should  see  material  points  separated  by  intervals  which  are  enormous 
in  comparison  with  their  dimensions,  and  describing  orbits  according  to 
regular  laws.  Like  the  stars  proper,  they  attract  each  other  or  repel,  and 
this  attraction  or  repulsiop,  which  is  along  the  line  joining  them,  depends 
only  on  distance."  (Cf.  Bulletin  of  the  American  Mathematical  Society, 
February,  1906,  p.  241;  authorized  translation  by  Professor  J.  W.  Young.) 

Professor  Sir  G.  H.  Darwin's  recent  presidential  address  to  the 
British  Association  for  the  Advancement  of  Science  at  Cape  Town, 
1905,  was  devoted  largely  to  the  discovery  of  electrons.  After 
treating  of  these  subatomic  corpuscles  he  adds : 

"  I  have  not  as  yet  made  any  attempt  to  represent  the  excessive  minute- 
ness of  the  corpuscles,  of  whose  existence  we  are  now  so  confident;  but,  as 
an  introduction  to  what  I  have  to  speak  of  next,  it  is  necessary  to  do  so. 

*  Or  say  a  globe  of  16  centimeters  diameter. 


i9o8.]  THE  PHYSICS  OF  THE   EARTH.  177 

To  obtain  any  adequate  conception  of  their  size  we  must  betake  ourselves 
to  a  scheme  of  threefold  magnification.  Lord  Kelvin  has  shown  that  if  a 
drop  of  water  were  magnified  to  the  size  of  the  earth  the  molecules  of  water 
would  be  of  a  size  intermediate  between  that  of  a  cricket  ball  and  of  a  marble. 
Now  each  molecule  contains  three  atoms,  two  being  of  hydrogen  and  one  of 
oxygen.  The  molecular  system  probably  presents  some  sort  of  analogy  with 
that  of  a  triple  star;  the  three  atoms  replacing  the  stars,  revolving  about  one 
another  in  some  sort  of  a  dance  which  cannot  be  exactly  described.  I  doubt 
whether  it  is  possible  to  say  how  large  a  part  of  the  space  occupied  ^  the 
whole  molecule  is  occupied  by  the  atoms;  but  perhaps  the  atoms  bear  to  the 
molecule  some  such  relationship  as  the  molecule  to  the  drop  of  water  re- 
ferred to.  Finally,  the  corpuscles  may  stand  to  the  atom  in  a  similar  scale 
of  magnitude.  Accordingly,  a  threefold  magnification  would  be  needed  to 
bring  these  ultimate  parts  of  the  atom  within  range  of  our  ordinary  scales 
of  measurement.    .    .    . 

"The  community  of  atoms  in  water  has  been  compared  with  a  triple 
star,  but  there  are  others  known  to  the  chemists  in  which  the  atoms  are  to 
be  counted  by  fifties  and  hundreds,  so  that  they  resemble  constellations." 

Such  general  discussions  by  these  illustrious  physicists,  Kelvin,* 
Poincare  and  Darwin,  are  not  to  be  construed  too  literally,  and  yet 
they  clearly  indicate  the  general  belief  among  the  foremost  men 
of  science  that  the  spaces  between  the  particles  of  matter  are  im- 
mense in  comparison  with  the  dimensions  of  the  particles  themselves. 

From  Lord  Kelvin's  discussion  of  the  size  of  atoms  treated 
in  the  above  section,  we  have  seen  that  the  diameters  of  these  bodies 
is  of  the  order  of  1/500,000,000  of  a  centimeter,  or  1/1,270,000,000 
of  an  inch.  The  average  space  between  the  molecules  being 
1/100,000  of  a  centimeter,  or  about  5,000  times  the  diameter,  is  of 
the  order  of  1/254,000  of  an  inch.     This  is  decidedly  below  the 

'In  a  well-known  paper  on  gravitating  matter,  Lord  Kelvin  compares 
the  stars  of  the  Milky  Way  to  the  atoms  of  a  bubble  of  gas.  For  a  giant  for 
whom  our  suns  would  be  what  atoms  are  to  us,  the  stars  would  be  beyond 
the  reach  of  the  keenest  vision  and  the  Milky  Way  appear  to  behave  as  a 
gaseous  medium.  M.  Poincare  has  discussed  the  problems  of  the  universe 
from  this  point  of  view  in  an  address  to  the  Astronomical  Society  of  France 
(Bulletin  Astronomique  de  la  Societi  Astronomique  de  France,  April,  1906; 
an  excellent  translation  in  Popular  Astronomy  for  October,  1906).  It  is 
remarkable  that  Democritus,  founder  of  the  atomic  theory  among  the 
Greeks  (460-360  B.  C),  should  also  have  recognized  that  the  Milky  Way  is 
composed  of  a  mass  of  stars  too  dense  to  be  seen  separately  by  the  unaided 
vision  (cf.  "Aristotle's  Meteorology,"  Lib.  I.,  Ch.  VIII.,  Sec.  4).  Thus  Lord 
Kelvin's  conceptions  do  not  differ  greatly  from  those  of  Democritus  of 
Abdera,  though  the  modern  theories  are  much  better  established  than  the 
atomic  theories  were  among  the  Greeks. 


178  SEE— FURTHER   RESEARCHES  ON  f  April  24, 

limit  of  resolution  of  the  microscope  which  has  been  estimated  by 
Michelson  at  1/100,000  of  an  inch. 

Now  in  our  discussion  of  the  constitution  of  granites  we  found 
that  the  visible  pores  in  the  .quartz  matrix  have  511  diameters  down 
to  less  than  1/10,000  of  an  inch,  and  thus  practically  to  the  lowest 
limit  visible  in  the  microscope.  These  visible  pores  thus  evidently 
conijpct  directly  with  the  smaller  invisible  spaces  which  separate  the 
molecules.  As  the  diameters  of  the  molecules  in  water  vapor  are 
only  about  1/5,000  of  the  spaces  between  them,  the  triple  atom  of 
hydrogen  and  oxygen  constituting  water  or  water  vapor  would  have 
ample  facilities  for  penetrating  a  spongy  and  cavernous  mass  like 
granite  with  innumerable  holes  frequently  of  large  size  but  always 
at  least  equal  to  the  average  free  path.  If  the  water  or  vapor 
were  under  pressure,  so  as  to  condense  the  fluid  and  thus  increase  the 
number  of  vibrations  of  a  molecule  per  second,  the  rate  of  penetra- 
tion of  the  fluid  obviously  would  be  much  augmented. 

And  since  granite  not  only  is  filled  with  pores  of  these  various 
sizes,  but  also  everywhere  more  or  less  cleft  by  planes  of  crystalline 
structures  which  are  not  really  tight,  but  full  of  fissures  and  thus 
inviting  the  penetration  of  the  fluid  by  the  full  power  of  capillary 
forces,  we  see  that  water  would  necessarily  penetrate  it  at  a  fairly 
rapid  rate.  At  the  same  time  the  influence  of  capillarity  in  such  a 
structure  is  so  great  that  although  water  might  enter  and  slowly  pass 
through  it,  even  the  development  of  steam  pressure  beneath  the 
layer  would  not  force  the  fluid  back,  because  the  steam  pressure  is 
nullified  an  infinitely  small  distance  from  where  it  is  exerted,  on 
account  of  capillary  resistance ;  yet  the  fluid  may  keep  on  descending 
under  the  suction  of  the  capillary  forces  so  long  as  the  supply  from 
above  is  not  cut  off. 

Upon  these  physical  grounds  it  seems  clear  that  there  must  be  a 
secular  leakage  of  the  ocean  bottoms,  and  a  corresponding  develop- 
ment of  steam  beneath  the  earth's  crust.  The  steam  expands  the 
rock  in  which  it  is  absorbed  and  in  seeking  release  thus  brings  on 
earthquakes  and  mountain  formation. 

Even  if  the  pressure  due  to  depth  should  tighten  up  the  struc- 
ture of  the  rock  in  the  lower  layers  of  the  crust,  it  would  not  be 
able  to  obliterate  the  leakage  depending  on  the  pores  and  crystalline 


I908.]  THE   PHYSICS   OF  THE  EARTH.  179 

Structure.  It  is  evident  that  at  depths  such  as  twenty  miles  the 
downward  movement  of  the  fluid  would  continue,  though  very 
slowly.  Hence  the  leakage  of  the  oceans  is  extremely  gradual,  and 
the  recurrence  of  earthquakes  visibly  delayed  after  relief  has  once 
been  obtained.  Thus  while  the  tightness  of  the  earth's  crust  due 
to  the  grain  of  the  rock  and  the  pressure  to  which  it  is  subjected  in 
the  lower  parts  does  not  prevent  ocean  leakage,  it  makes  the  process 
so  slow  and  gradual  as  to  afford  considerable  protection  to  life  upon 
our  planet. 

II.  On  the  Physical  State  of  the  Earth's  Interior,  on  the 
Average  Rigidity  of  the  Globe  as  a  Whole,  and  on  the 
Substratum  of  Plastic  Matter  Beneath  the 
Crust   which    in    Earthquakes    Be- 
haves AS  Fluid. 

§  12.  On  the  Theory  of  a  Fluid  Globe  Held  by  the  Older  Geolo- 
gists, and  on  Hopkins^  Argument  for  Solidity  Based  on  the  Phe- 
nomena of  Precession  and  Nutation, — In  the  early  part  of  the  nine- 
teenth century  it  was  generally  believed  by  geologists  that  the  earth 
was  a  liquid  globe  covered  by  a  rocky  crust  much  thinner  in  pro- 
portion to  the  diameter  than  the  shell  is  to  that  of  an  egg.  This 
supposed  liquid  interior  had  been  suggested  by  the  streams  of  molten 
lava  often  observed  to  issue  from  volcanoes,  and  by  the  igneous 
rocks  so  abundantly  poured  forth  in  many  places.  The  theory  of  a 
fluid  globe  seemed  to  be  confirmed  by  the  observed  increase  of 
temperature  downward,  which  would  give  rise  to  molten  rock  at  a 
depth  of  some  twenty  miles.  The  mountains  and  other  phenomena 
traceable  to  dislocations  of  the  crust  could  all  be  explained  by  a  solid 
layer  of  this  thickness,  and  the  natural  inference  was  that  the  great 
central  nucleus  remained  liquid.  The  consolidation  of  the  globe 
was  ascribed  to  the  progress  of  secular  cooling,  from  the  primitive 
state  of  high  temperature  assumed  by  Laplace  in  the  nebular  hy- 
pothesis postulated  for  explaining  the  origin  of  the  solar  system. 

The  older  geologists  had  not  adequately  considered  the  effects 
of  pressure  in  augmenting  the  solidity  of  the  globe  as  we  go  down- 
ward; for  since  pressure  raises  the  melting  point  of  solids,  the 
matter  of  the  nucleus,  though  highly  heated,  might  be  solid  if  *  the 


180  SEE— FURTHER   RESEARCHES  ON  pipril«4, 

pressure  be  great  enough  to  prevent  fusion  under  the  prevailing 
temperature.  In  order  to  throw  light  upon  this  question,  Hopkins 
of  Cambridge,  England,  took  up  the  problem  in  1839  (Phil.  Trans., 
1839;  "  Researches  in  Physical  Geology,"  1839-1842),  and  sought  to 
prove  from  the  observed  phenomena  of  precession  and  nutation  that 
the  earth  could  not  be  composed  of  a  thin  shell  some  twenty  miles 
thick,  filled  with  liquid.  He  concluded  that  the  crust  could  not  be 
less  than  800  to  1,000  miles  thick,  and  that  the  globe  might  even  be 
solid  to  the  center,  except  some  small  vesicular  spaces  here  and 
there  filled  with  molten  rock. 

In  1868  this  subject  was  examined  by  the  eminent  French  as- 
tronomer, Delaunay,  who  published  a  paper  on  "  The  Hypothesis  of 
the  Interior  Fluidity  of  the  Globe  "  (C.  R.  Acad,  des  Sci.,  Paris, 
July  I3>  1868),  in  which  he  threw  doubt  on  the  views  of  Hopkins, 
and  suggested  that  if  the  earth's  nucleus  were  a  mass  of  sufficient 
viscosity  it  might  behave  as  if  it  were  solid,  and  hence  concluded 
that  the  observed  phenomenon  of  precession  and  nutation  did  not 
necessarily  exclude  a  fluid  nucleus. 

§  13.  Lord  Kelvin's  Earliest  Studies  on  the  Precession  of  a 
Spheroid  Containing  Liquid. — ^Lord  Kelvin  had  already  taken  up  the 
problem  of  the  internal  state  of  the  earth  in  1862,  and  considered  the 
effects  of  a  fluid  nucleus  enclosed  in  a  thih  shell  when  the  whole 
mass  was  subjected  to  tidal  strains.  As  the  shell  must  yield  under 
these  strains  the  land  would  be  carried  up  and  down  with  the  super- 
jacent sea,  and  if  such  yielding  occurred  it  ought  to  be  sensible  to 
observation.  But  since  the  sensible  obliteration  of  the  tides  had  not 
been  observed,  he  naturally  inclined  to  the  view  of  Hopkins  that  the 
earth  is  effectively  rigid  and  behaves  as  a  solid  globe. 

In  reply  to  Delaunay's  criticism  Lord  Kelvin  pointed  out  that  if 
the  French  astronomer  had  worked  out  the  problem  mathematically 
he  could  not  fail  to  see  that  the  hypothesis  of  a  viscous  and  quasi- 
rigid  interior  "  breaks  down  when  tested  by  a  simple  calculation  of 
the  amount  of  tangential  force  required  to  give  to  any  globular 
portion  of  the  interior  mass  the  precessional  and  nutational  motions 
which,  with  other  physical  astronomers,  he  attributes  to  the  earth 
as  a  whole."  {Nature,  February  i,  1872.)  On  making  this  calcula- 
tion Lord  Kelvin  found  that  the  earth's  crust  down  to  depths  of 


,9o8.]  THE   PHYSICS  OF  THE   EARTH.  181 

hundreds  of  kilometers  must  be  capable  of  resisting  a  tangential 
stress  of  nearly  o.i  of  a  gramme  weight  per  square  centimeter;  this 
would  rapidly  draw  out  of  shape  any  plastic  substance  which  could 
be  properly  called  a  viscous  fluid.  "An  angular  distortion  of  8" 
is  produced  in  a  cube  of  glass  by  a  distorting  stress  of  about  ten 
grammes  weight  per  square  centimeter.  We  may  therefore  safely 
conclude  that  the  rigidity  of  the  earth's  interior  or  substance  could 
not  be  less  than  a  millionth  of  the  rigidity  of  glass  without  very 
sensibly  augmenting  the  lunar  nineteen  yearly  nutation."  (Nature, 
February  i,  1872,  p.  258.) 

Notwithstanding  these  early  criticisms  of  Delaunay's  paper,  Lord 
Kelvin  subsequently  concluded  that  the  phenomena  of  precession 
and  nutation  do  not  decisively  settle  the  question  of  the  earth's  in- 
ternal fluidity.  Yet  the  semiannual  and  lunar  fortnightly  nutations 
may  be  considered  to  disprove  absolutely  the  existence  of  a  thin  rigid 
shell  full  of  liquid.  If  the  fluid  were  arranged  in  successive  layers 
of  equal  density,  the  only  nutational  or  precessional  influence  exerted 
upon  it  would  depend  on  the  non-sphericity  of  the  shells.  "  A  very 
slight  deviation  of  the  inner  surface  of  the  shell  from  perfect  spher- 
icity would  suffice,"  according  to  Lord  Kelvin,  "  in  virtue  of  the 
quasi-rigidity  due  to  vortex  motion,  to  hold  back  the  shell  from 
taking  sensibly  more  precession  than  it  would  give  to  the  liquid,  and 
to  cause  the  liquid  (homogeneous  or  heterogeneous)  and  the  shell 
to  have  sensibly  the  same  precessional  motion  as  if  the  whole 
constituted  one  rigid  body."  (Sir  W.  Thomson,  British  Assoc. 
Report,  1876,  Sections,  p.  5.) 

It  will  be  seen  from  this  discussion  that  the  argument  from 
precession  and  nutation  is  only  in  part  conclusive.  If  the  fluid  had 
a  viscosity  approaching  high  rigidity  for  rapidly  acting  forces,  or  it 
were  subjected  to  such  pressure  that  the  particles  in  confinement 
acquired  the  properties  of  a  solid,  there  would  evidently  be  no 
sensible  deviation  from  the  precession  and  nutation  appropriate  to  a 
cold  solid  globe. 

§  14.  On  Lord  Kelvin's  Researches  on  the  Earth's  Rigidity 
Based  on  the  Analysis  of  the  Tides. — ^The  state  of  the  earth's  in- 
terior had  early  engaged  the  attention  of  Lord  Kelvin,  for  the 
propagation  of  heat  through  the  crust  was  before  him  as  early  as 


182  SEE— FURTHER   RESEARCHES  ON  [April  a4. 

1846.  ("De  Motu  Caloris  per  Terrae  G)rpus,"  read  before  the 
faculty  of  the  University  of  Glasgow  in  1846;  also  a  "  Note  on  Cer- 
tain Points  in  the  Theory  of  Heat,"  February,  1844,  published  in 
the  Cambridge  Mathernqtical  Journal,  and  reprinted  in  the  "  Mathe- 
matical and  Physical  Papers  of  Sir  W.  Thomson,"  1882,  Vol.  I, 
Art.  X.) 

In  a  paper  "  On  the  Rigidity  of  the  Earth  "  published  in  the 
Philosophical  Transactions  of  the  Royal  Society  for  May,  1862, 
Lord  Kelvin  pointed  out  that  if  the  matter  of  the  earth's  interior 
yielded  readily  to  the  tidal  forces  arising  from  the  attraction  of  the 
sun  and  moon,  the  crust  itself  would  respond  to  these  forces  in 
much  the  same  way  as  the  waters  of  the  sea ;  and  the  corresponding 
movements  of  the  crust  would  mask  or  largely  reduce  the  height 
of  the  oceanic  tides  calculated  for  a  rigid  earth.  By  actual  analysis 
of  long  series  of  tidal  observations  Kelvin  and  Darwin  subsequently 
found  the  observed  fortnightly  tide  to  have  very  nearly  its  full 
theoretical  height,  and  hence  concluded  that  our  globe  as  a  whole 
possesses  a  very  high  effective  rigidity.  (Cf.  Thomson  and  Tait's 
Natural  Philosophy,"  Vol.  I,  part  II,  §  832-847 ;  also  the  article 
Tides,"  Encyclopedia  Britannica,  ninth  edition,  §  44.) 
Owing  to  the  great  importance  of  this  work  on  the  rigidity  of 
the  earth,  we  must  trace  the  successive  steps  in  the  advancement  of 
our  knowledge.  The  assumption  that  the  earth  is  made  up  of  a 
liquid  nucleus  covered  with  a  thin  crust  stiff  enough  to  maintain  its 
figure  against  the  tide-raising  forces  of  the  sun  and  moon  would 
imply  that  the  crust  has  a  degree  of  strength  and  rigidity  not  pos- 
sessed by  any  known  substance.  It  was  therefore  inferred  by  Lord 
Kelvin  as  early  as  1862  that  the  crust  might  be  2,000  to  2,500  miles 
thick,  in  order  to  resist  distortion  under  the  tide-producing  forces 
arising  from  the  sun  and  moon. 

"If  the  crust  yielded  perfectly,  there  would  be  no  tides  of  the  sea,  no 
rising  and  falling  relatively  to  the  land,  at  all.  The  water  would  go  up  and 
down  with  the  land,  and  there  would  be  no  relative  movement;  and  in  pro- 
portion as  the  crust  is  less  or  more  rigid  the  tides  would  be  more  or  less 
diminished  in  magnitude.  Now  we  cannot  consider  the  earth  to  be  absolutely 
rigid  and  unyielding.  No  material  that  we  know  of  is  so.  But  I  find  from 
calculation  that  were  the  earth  as  a  whole  not  more  rigid  than  a  similar  globe 
of  steel  the  relative  rise  and  fall  of  the  water  in  the  tides  would  be  only 


it 
it 


.908]  THE  PHYSICS  OF  THE  EARTH.  183 

two-thirds  of  that  which  it  would  be  were  the  rigidity  perfect;  while,  if  the 
rigidity  were  no  greater  than  that  of  a  globe  of  glass,  the  relative  rise  and 
fall  would  be  only  two-fifth  of  that  on  a  perfectly  rigid  globe. 

"  Imperfect  as  the  comparison  between  theory  and  observation  as  to  the 
actual  height  of  the  tides  has  been  hitherto,  it  is  scarcely  possible  to  be- 
lieve that  the  height  is  only  two  fifths  of  what  it  would  be  if,  as  has  been 
universally  assumed  in  tidal  theories,  the  earth  was  perfectly  rigid.  It  seems, 
therefore,  nearly  certain,  with  no  other  evidence  than  this  afforded  by  the 
tides,  that  the  tidal  effective  rigidity  of  the  earth  must  be  greater  than  that 
of  glass.  This  is  the  result  taking  the  earth  as  a  globe  uniformly  rigid 
throughout.  That  a  crust  fifty  or  a  hundred  miles  thick  could  possess  such 
preternatural  rigidity,  as  to  give  to  the  mass,  part  solid  and  part  liquid,  a 
rigidity  as  a  whole,  equal  to  that  of  glass  or  steel  is  incredible;  and  we 
are  forced  to  the  conclusion  that  the  earth  is  not  a  mere  thin  shell  filled 
with  fluid,  but  is  on  the  whole  or  in  great  part  solid."  (Paper  read  to 
Geological  Society  of  Glasgow,  February  14,  1878;  Kelvin's  "Popular  Lec- 
tures and  Addresses,"  Vol.  II,  pp.  317-318.) 

In  his  presidential  address  to  the  Mathematical  and  Physical  section  of 
the  British  Association  at  Glasgow,  September  7,  1876,  Lord  Kelvin  remarked 
of  the  earth's  crust  that  "  were  it  of  continuous  steel  and  500  kilometers 
thick,  it  would  yield  very  nearly  as  much  as  if  it  were  india  rubber  to  the 
deforming  influences  of  centrifugal  force  and  of  the  sun's  and  moon's  at- 
tractions." "  The  solid  crust  would  yield  so  freely  to  the  deforming  influence 
of  sun  and  moon  that  it  would  simply  carry  the  waters  of  the  ocean  lip  and 
down  with  it,  and  there  would  be  no  sensible  rise  and  fall  of  water  relatively 
to  the  land."    ("  Popular  Lectures,"  Vol.  II.,  pp.  251-2.) 

Lord  Kelvin's  final  conclusion  was  that  "  the  earth  as  a  whole  is 
certainly  more  rigid  than  glass,  but  perhaps  not  quite  so  rigid  as 

steel." 

§  15.  Darwin's  Researches  on  the  Tidal  Method  of  Evaluating 
the  Earth's  Rigidity. — As  the  natural  successor  of  Lord  Kelvin  in 
the  researches  on  the  physics  of  the  earth,  Professor  Sir  G.  H. 
Darwin  took  up  the  problem  of  the  earth's  internal  physical  con- 
dition and  confirmed  and  extended  these  conclusions  by  several 
important  lines  of  inquiry.  Darwin's  researches  on  the  bodily  tides 
of  viscous  and  semi-elastic  spheroids  and  on  the  oceanic  tides  upon 
a  yielding  nucleus  tended  to  strengthen  the  argument  for  a  high 
eflFective  rigidity  so  decidedly  that  he  concluded  that  "no  very 
considerable  portion  of  the  interior  of  the  earth  can  even  distantly 
approach  the  fluid  condition." 

But  whilst  Darwin's  researches  confirmed  Kelvin's  conclusions 
as  to  the  great  effective  rigidity  of  the  earth,  yet  a  more  critical 


184  SEE— FURTHER   RESEARCHES  ON  [April  24, 

examination  of  the  method  for  calculating  the  fortnightly  tide  led 
to  the  conviction  that  Laplace's  argument  is  regard  to  the  effects  of 
friction  was  unsatisfactory.  That  friction  would  greatly  effect  the 
motion  of  the  water  in  slow  ocean  currents  within  a  few  days  was 
seen  to  be  untenable.  In  consequence  of  this  defect  it  turned  out 
that  long  period  tides  as  short  as  a  fortnight  would  not  enable  the 
physicist  to  evaluate  the  rigidity  of  the  earth,  though  the  18.6  yearly 
tide,  depending  on  the  revolution  of  the  Moon's  nodes,  if  it  can  be 
determined  by  observation,  will  eventually  give  the  desired  result. 
The  height  of  this  18.6  yearly  tide,  however,  is  only  one  third  of  an 
inch  at  the  equator,  and  great  accuracy  will  be  required  for  its 
detection. 

Acting  on  the  old  belief  Darwin  compared  the  lunar  fortnightly 
and  monthly  tides  observed  for  33  years  at  various  Indian  and 
European  ports,  with  the  equilibrium  theory,  and  found  that  the 
tide-heights  were  about  two  thirds  of  the  theoretical  height.  Ac- 
cordingly he  remarks :  "  On  the  whole  we  may  fairly  conclude  that, 
whilst  there  is  some  evidence  of  a  tidal  yielding  of  the  earth's  mass, 
that  yielding  is  certainly  small,  and  the  effective  rigidity  is  at  least 
as  great  as  that  of  steel."  (Thomson  and  Tait's  "  Nat.  Phil.,"  Vol. 
I,  Part  II,  §848.) 

This  was  written  prior  to  the  discovery  of  the  theoretical  defect 
in  the  method  of  calculating  the  height  of  tides  with  periods  not 
exceeding  a  fortnight  in  duration;  yet  even  after  the  discovery  of 
this  defect  it  was  still  possible  to  infer  that  tides  of  long  period  in 
oceans  such  as  ours  must  conform  much  more  nearly  to  the  equi- 
librium laws  than  do  the  tides  of  short  period.  "  Whilst,  then,  this 
precise  comparison  with  the  rigidity  of  steel  falls  to  the  ground, 
the  investigation  remains  as  an  important  confirmation  of  Thomson's 
conclusion  as  to  the  g^eat  effective  rigidity  of  the  earth.  ...  It  ap- 
pears by  numerical  calculation  on  viscous  and  elastico-viscous  tides 
that  in  order  that  the  oceanic  semi-diurnal  tide  may  have  a  value 
equal  to  two  thirds  of  the  full  amount  on  a  rigid  globe,  the  stiffness 
of  the  globe  must  be  about  twenty  thousand  times  as  great  as  that 
of  pitch  at  freezing  temperature,  when  it  is  hard  and  brittle."  (Sir 
G.  H.  Darwin,  article  "  Tides,"  Ency.  Brit.,  §§  44-450 

§  16.  On  the  Rigidity  of  the  Earth  as  found  by  Comparing  the 


f 

,908.]  THE   PHYSICS  OF  THE  EARTH.  185 

Observed  Period  of  the  Polar  Motion  Arising  in  the  Variation  of 
Latitude  with  the  Theoretical  Eulerian  Period  Calculated  for  a  Rigid 
Earth, — The  detection  of  the  variation  of  latitude  by  Kiistner  at 
Berlin  in  1890-91  and  the  subsequent  discussion  by  Chandler  of  long 
series  of  observations  showing  that  the  movement  of  the  pole  in  the 
body  of  the  earth  has  a  period  of  some  427  days,  instead  of  the  305 
days  long  ago  inferred  from  Euler's  theory  of  the  rotation  of  a 
rigid  spheroid,  led  Professor  Newcomb  to  point  out  that  this  ob- 
served prolongation  of  the  theoretical  Eulerian  period  indicates  some 
yielding  of  the  matter  of  the  globe  imder  the  stresses  to  which  it 
is  subjected  by  the  movement  of  the  pole,  and  would  afford  a  new 
method  of  evaluating  the  earth's  rigidity.  In  his  well-known  paper 
on  the  "  Dynamics  of  the  Earth's  Rotation  "  (Monthly  Notices,  R, 
A.  S.,  March,  1892)  Newcomb  showed  that  the  results  already  ob- 
tained decidedly  confirmed  Darwin's  conclusion  that  the  rigidity  of 
the  globe  as  a  whole  is  comparable  to  that  of  steel. 

The  essential  point  in  Newcomb's  explanation  is  that  when  the 
pole  changes  its  position  in  the  body  of  the  globe,  the  distribution 
of  centrifugal  force  shifts  with  respect  to  the  solid  earth,  which  is 
thus  put  into  a  state  of  stress  and  must  yield  to  the  forces  acting 
upon  it,  like  any  other  elastic  solid  body;  the  periodic  deformation 
of  the  earth's  figure  operating  to  lengthen  the  period  of  the  free 
nutation,  by  an  amount  depending  on  the  average  rigidity  of  the 
whole  earth. 

The  continued  investigation  of  the  variation  of  latitude  carried 
out  at  the  various  international  latitude  observatories  by  Albrecht 
and  others  confirms  this  observational  result,  and  the  subject  has 
also  been  examined  theoretically  by  Darwin,  Hough,  Larmor  and 
others;  so  that  the  validity  of  the  method  suggested  by  Newcomb 
is  generally  recognized. 

In  1896  Mr.  S.  S.  Hough  treated  of  the  problem  in  a  very  thor- 
ough manner  in  his  well-known  paper,  "  On  the  Rotation  of  an 
Elastic  Spheroid"  (Phil.  Trans.,  A,  1896).  He  considered  chiefly 
the  case  of  an  incompressible  homogeneous  spheroid,  and  was  en- 
abled to  show  by  rigorous  methods  that  the  rigidity  of  the  earth  in 
all  probability  slightly  exceeds  that  of  steel. 

In  a  remarkable  paper  "On  the  Period  of  the  Earth's  Free 


t 
186  SEE— FURTHER   RESEARCHES  ON  [April  04, 

Eulerian  Precession,"  read  to  the  Cambridge  Philosophical  Society, 
May  25,  1896,  Professor  Larmor  showed  how  to  estimate  the  effect  of 
the  elastic  yielding  of  a  rotating  solid  on  the  period  and  character 
of  the  free  precession  of  its  axis  of  rotation,  and  again  confirmed 
the  high  effective  rigidity  of  the  earth  from  another  point  of  view. 

The  observed  prolongation  of  the  Eulerian  period  is  thus  fully 
explained  by  the  imperfect  rigidity  of  the  earth's  mass,  and  the 
high  rigidity  thus  deduced  has  naturally  strengthened  the  earlier 
conclusions  of  Kelvin  and  Darwin  drawn  from  the  study  of  the 
long  period  tides  of  the  sea. 

This  investigation,  like  those  already  cited,  gives  us  only  an 
average  effect  for  the  earth  as  a  whole,  but  does  not  tell  us  the  law 
of  the  distribution  of  rigidity  within  the  globe.  If  this  law  of  dis- 
tribution of  rigidity  could  be  found,  even  approximately,  it  would  be 
of  great  interest,  because  we  could  then  see  in  what  part  of  the  globe 
the  principal  part  of  the  yielding  takes  place;  and  this  would  give 
us  a  much  better  understanding  of  the  internal  constitution  of  our 
planet  than  heretofore  has  been  considered  possible. 

§  17.  Rigidity  of  the  Earth  Calculated  from  the  Theory  of 
Gravity,  on  the  Hypothesis  that  the  Distribution  of  Rigidity  in  the 
Globe  is  Everywhere  Proportional  to  the  Pressure. — It  has  not  been 
supposed  by  previous  investigators  that  a  method  could  be  devised 
for  deducing  the  rigidity  of  a  body  like  the  earth  from  the  theory 
of  gravity;  but  in  1905  it  occurred  to  the  present  writer  that  such  a 
method  could  be  found  if  we  could  adopt  a  suitable  hypothesis  for 
the  variation  of  the  rigidity  with  the  pressure.  Previous  investi- 
gations of  the  internal  state  of  the  heavenly  bodies  had  justified  the 
law  of  Laplace  as  giving  an  excellent  approximation  to  the  law  of 
density  for  the  earth  and  the  rest  of  the  encrusted  planets;  and 
the  monatomic  law  had  been  found  most  satisfactory  for  the  sun 
and  fixed  stars  (cf.  A.  N.,  4053).  These  laws  enable  one  to  ob- 
tain the  pressure  at  every  point  of  the  radius  of  the  heavenly  bodies. 
For  in  several  ways  Laplace's  law  of  density  is  fairly  well  estab- 
lished for  the  earth,  and  on  equally  good  grounds  the  density  of  the 
sun  is  believed  to  conform  essentially  to  the  monatomic  law. 

From  a  study  of  the  laws  of  density,  pressure  and  temperature 
within  the  heavenly  bodies  it  appeared  to  me  (as  it  had  indepen- 


x9o8.|  THE  PHYSICS  OF  THE  EARTH.  187 

dently  appeared  to  Arrhenius  five  years  before)  that  matter  under 
these  extreme  conditions  must  be  essentially  gaseous;  and  as  it  is 
above  the  critical  temperature,  it  is  made  to  behave  in  confinement 
as  an  elastic  solid.  Now  in  all  gaseous  masses  the  density  is  pro- 
portional to  the  pressure  so  long  as  the  gas  remains  perfect;  and 
the  gas  does  not  cease  to  be  perfect  when  the  temperature  is  above 
the  critical  value,  though  it  may  acquire  in  confinement  the  property 
of  an  elastic  solid  if  the  pressure  be  great  enough  to  bring  the 
molecules  within  a  distance  at  which  the  molecular  forces  become 
effective  in  spite  of  the  high  temperature.  Thus  while  the  property 
of  rigidity  in  cold  solids  depends  wholly  on  molecular  forces 
which  prevent  deformation,  this  property  for  gaseous  matter  in 
confinement  under  such  pressure  that  it  acquires  the  property  of  an 
elastic  solid,  is  due  wholly  to  the  pressure.  The  molecular  forces 
giving  effective  rigidity  must  increase  in  proportion  to  the  pressure, 
or  in  a  higher  ratio. 

If  according  to  hypothesis  the  matter  is  made  solid  by  pressure, 
then  the  molecular  forces  resisting  deformation  in  the  imprisoned 
matter  thus  solidified  cannot  resist  deformation  in  a  less  degree  than 
the  direct  proportion  to  the  pressure  on  which  the  solidification  de- 
pends. And  any  ratio  higher  than  the  direct  proportionality  to  the 
pressure  would  most  likely  depend  on  the  temperature.  Now  the 
temperature  in  the  earth  is  supposed  to  be  everywhere  such  as  to 
make  the  density  conform  essentially  to  Laplace's  law ;  and  the  pres- 
sure resulting  from  this  law  of  density  gives  the  matter  everywhere 
the  property  of  an  elastic  solid,  and  therefore  its  molecular  proper- 
ties must  correspond  to  the  physical  state  determined  by  the  laws  of 
density  and  pressure. 

It  is  of  course  conceivable  that  some  parts  of  the  globe  might 
be  relatively  more  rigid  than  is  required  to  give  solidity,  but  the 
effect  of  this  would  only  increase  the  average  rigidity  of  the  earth 
as  a  whole.  And  since  seismological  and  other  observations  seem 
to  show  that  the  globe  is  solid  throughout,  except  a  thin  layer  just 
beneath  the  crust,  the  h)rpothesis  of  a  rigidity  proportional  to  the 
pressure  will  give  a  true  minimum  value  of  the  earth's  rigidity. 

Now  on  the  hypothesis  that  the  density  follows  Laplace's  law, 


188  SEE— FURTHER   RESEARCHES  ON  lAprila4, 

the  pressure  throughout  the  earth's  mass  is  given  by  the  formula 
(cf.  A.  N.,  4104) 

where  r  is  the  radius  of  the  earth,  g  mean  gravity,  q  the  constant 
for  Laplace's  law,  2.52896  radians  =  144®  53'  55''.2,  cr  the  density 
at  any  point,  8  the  density  at  the  surface,  and  o-i  the  mean  density. 
To  render  this  expression  available  for  integration  throughout 
the  sphere  occupied  by  the  earth's  mass,  we  must  put  for  a*  its  value 

sin*(y;r) 
^  '^  ^«  -g'x^  • 

and  for  8^  its  value 

corresponding  to  the  surface  where  x=i.    Thus  we  obtain 

.  _  3K^)'^  rsin*  {qx)      anVl 

For  the  total  pressure  throughout  a  sphere  of  radius  p=^rx,r  being 
the  external  radius,  and  jr=  {p/r)  =  fraction  of  the  radius,  we  have 


I   p'\in^^ 


rdx 

(3) 


^2^^-(r^-^'^---'r'-). 


which  by  integration  becomes 


/>=MY)^4^(?^:zil!LMf£i(^)_sinV-l    (4) 

As  our  integration  is  to  include  the  whole  sphere  of  the  earth,  we 
put  x=i,  and  then  we  have 

p _  3K^r  ^'T!;*  /  y  -  sin  g  cos  q  _  sin V  \  ,  «. 

The  total  volume  of  the  earth  is  (4/3)»rr*,  and  hence  the  average 
pressure  per  unit  of  area  on  all  concentric  spherical  surfaces  is 


i9o«.]  THE   PHYSICS  OF  THE   EARTH.  189 


(6) 


P  3      C 

_  9i^o^y  •  r  /  ^  —  sin  ^  cos  ^      sin*  g  \ 

If  r  is  expressed  in  meters,  the  mean  pressure  or  mean  rigidity  R 
comes  out  in  kilograms  per  square  meter.  To  reduce  the  result  to 
atmospheres  we  divide  by  10,333.  The  result  for  the  earth  is 
/?  =  748343  atmospheres,  about  the  rigidity  of  wrought  iron. 

This  method  takes  no  account  of  the  earth's  solid  crust,  and  is 
therefore  too  small;  moreover  viscosity  increases  within  the  earth, 
owing  to  the  rise  of  temperature  downward.  We  give  hereafter  an 
approximation  to  the  increase  of  rigidity  by  determining  the  mean 
rigidity  of  the  earth's  matter,  as  distinguished  from  that  of  the 
various  layers  composing  the  globe,  just  found  by  the  above  analysis. 

To  find  the  mean  rigidity  of  the  earth's  matter  we  must  consider 
not  only  the  pressure  but  also  the  density  or  mass  per  unit  volume 
of  the  imprisoned  matter  in  each  layer.  The  result  represents  a 
mean  rigidity  in  which  every  elementary  spherical  shell  composing 
the  globe  is  allowed  a  weight  proportional  to  its  mass,  which  is 
multiplied  by  the  pressure  to  which  it  is  subjected. 

The  theory  of  the  determination  of  the  mean  rigidity  of  the 
earth's  matter  is  as  follows : 

r.       r*        .  ,  sin  (ax) 
p '  4'irr^x^  •  rdx '  <r  =  47rr^(rQ   I    p-x^dx—--^.       (7) 
Jo  ^^ 

Substituting  for  p  its  value  from  (2),  we  get 
The  integral  of  this  last  term  is  —  sin^  q  —— 


q' 


The  value  of  the  first  integral  is  most  conveniently  found  by  quad- 
rature, table  for  which  is  given  in  A.  N.,  4104,  p.  379.  Dividing  out 
the  mass,  or  volume  of  the  sphere  by  the  density,  we  have 

PROC.  AMER.  PHIL.  SOC.  XLVII.   189  M,   PRINTED  SEPTEMBER    22,   I908. 


190  SEE— FURTHER   RESEARCHES  ON  [April  a4. 


F 


3  I    /  •  ^'trr^j^  •  rdx  •  <r 


fTTcr^r*  \irc^r^  (lo) 

On  putting  qx=i44°  53'  55".2,  the  value  of  the  integral  is  found 
by  quadrature  to  be  0.9592502,  and  when  the  rest  of  the  formula  is 
reduced  to  numbers  we  have  (A.  N,,  4104)  : 

R'  =  1028702  atmospheres. 

The  rigidity  of  nickel  steel  is  taken  to  be  1,000,000  atmospheres.  It 
thus  appears  from  this  calculation  that  the  average  rigidity  of  all 
the  earth's  matter  somewhat  exceeds  that  of  nickel  steel.  The 
actual  rigidity  of  the  earth  almost  certainly  lies  between  the  limits 
thus  established,  namely  i?  =  748,843,  based  on  the  rigidity  of  the 
layers  deduced  from  the  pressure  to  which  they  are  subjected,  and 
/?'  =  1,028,  702,  derived  from  the  product  of  the  mass  of  each  layer 
by  the  pressure  acting  upon  it. 

In  the  paper,  "  Researches  on  the  Rigidity  of  the  Heavenly 
Bodies,"  A.  N.,  4104,  the  rigidity  of  the  earth  is  discussed  as 
follows : 

"  When  one  considers  the  effects  of  the  enclosing  crust  and  the  viscosity 
of  the  whole  earth,  which  must  be  assumed  to  increase  towards  the  centre, 
owing  to  the  increasing  density  and  rising  temperature  of  the  imprisoned 
matter,  it  seems  not  improbable  that  the  actual  effective  rigidity  of  our 
globe  may  be  nearer  the  upper  limit  than  the  lower,  and  probably  we  shall 
not  be  far  wrong  in  concluding  that  it  is  approximately  equal  to  that  of 
nickel  steel. 

"  Leaving  aside  the  consideration  of  the  effects  of  the  solidified  crust, 
it  is  evident  from  the  nature  of  the  forces  at  work  that  most  of  the  yielding 
of  our  globe,  due  to  the  periodic  action  of  small  forces,  is  in  the  outer  layers ; 
and  in  general  the  yielding  in  any  concentric  layer  may  be  taken  to  be  in- 
versely as  the  pressure  to  which  the  imprisoned  matter  is  subjected.  It  is 
remarkable  that  the  curve  of  pressure  as  we  descend  in  the  earth  becomes 
therefore  also  the  curve  of  effective  rigidity  for  the  matter  of  which  the 
earth  is  composed.  Thus  the  rigidity  of  the  matter  at  the  earth's  center 
probably  is  at  least  three  times  that  of  nickel  steel  used  in  armor  plate;  as 
we  approach  the  surface  the  effective  rigidity  constantly  exceeds  that  of 
nickel  steel  until  we  come  within  less  than  04  of  the  radius  from  the  sur- 
face, where  the  pressure  is  less  than  1,000,000  atmospheres. 


i9o«.]  THE  PHYSICS  OF  THE  EARTH.  191 

"  To  imagine  a  mechanical  substitute  for  the  earth's  constitution,  without 
the  introduction  of  pressure,  suppose  an  alloy  of  adamant  to  give  the 
material  at  the  centre  of  such  a  globe,  of  the  same  size  but  devoid  of  gravi- 
tation, a  hardness  three  times  that  of  armor  plate.  The  outer  layers  as  we 
approach  the  surface  must  then  be  supposed  softer  and  softer,  until  it  is  like 
armor  plate  at  a  little  over  0.6  from  the  center,  and  finally  a  very  stiff  fluid 
near  the  surface.  In  addition  to  this  arrangement  of  its  effective  internal 
rigidity  the  actual  earth  is  enclosed  in  a  spheroidal  shell  of  solid  rock 
analogous  to  granite.  One  can  easily  see  that  tidal  forces  applied  to  all  the 
particles  of  such  an  artificial  armored  sphere  would  produce  but  very  slight 
deformation,  because  of  the  enormous  effective  rigidity  of  the  nucleus. 

"The  principal  uncertainty  in  this  result  arises  from  the  admissible 
variations  in  the  assumed  Laplacean  distribution  of  density  within  the  earth. 
Both  Radau  and  Darwin  (cf.  Monthly  Notices,  Roy.  Astron.  Soc,  December, 
1899)  have  pointed  out  that  considerable  variations  in  the  internal  distribu- 
tion of  density  are  possible  without  invalidating  the  well-known  argument 
drawn  from  the  phenomenon  of  the  precession  of  the  equinoxes;  yet  on 
physical  grounds  it  seems  clear  that  pressure  is  the  principal  cause  of  the 
increase  of  density  towards  the  earth's  centre.  And  since  this  does  not  vary 
greatly  for  moderate  changes  in  the  law  of  density,  the  principle  of  con- 
tinuity shows  that  the  actual  law  of  density  within  the  earth  cannot  depart 
very  widely  from  that  of  Laplace.  The  above  value  of  tne  theoretical  rigidity 
of  the  earth  may  therefore  be  taken  as  essentially  accurate,  and  I  think  no 
doubt  can  remain  that  the  rigidity  of  our  earth  as  a  whole  considerably 
exceeds  that  of  steel.  The  original  conclusions  of  Kelvin  and  Darwin  are 
therefore  confirmed  by  the  present  dynamical  considerations  based  upon  the 
theory  of  universal  gravitation." 

In  this  connection  we  should  remember  that  the  experimental 
rigidity  of  steel  is  808,000  and  of  glass  235,000  atmospheres.  The 
calculated  rigidity  of  all  the  matter  within  the  globe,  foimd  by  con- 
sidering not  only  all  the  layers,  but  also  the  density  in  each  layer, 
is  found  to  be  1,028,702  atmospheres.  Now  the  average  rigidity 
must  be  greater  than  750,000,  because  the  stiffness  of  the  crust  and 
increase  of  viscosity  downward  is  neglected  in  the  gravitational 
method.  In  fact  this  method  is  not  applicable  to  the  outermost 
layers,  because  the  pressure  there  is  much  less  than  the  rigidity,  and 
only  becomes  equal  to  the  rigidity  at  a  depth  of  something  like  one 
tenth  of  the  radius,  where  the  pressure  is  320,295  atmospheres. 

According  to  the  experiments  of  Milne  and  Gray  the  rigidity  of 
granite  is  about  one  sixth  that  of  steel;  and  as  steel  has  a  rigidity 
of  808,000  atmospheres,  that  of  granite  is  about  135,000  atmospheres, 
or  a  little  more  than  one  half  that  of  glass.    We  may  therefore  take 


192  SEE— FURTHER   RESEARCHES  ON  [April  24. 

the  outer  layers  of  our  globe  to  have  a  rigidity  about  half  that  of 
glass,  and  assume  that  at  a  depth  of  o.i  of  the  radius  it  becomes 
nearly  2.5  times  as  great  as  it  is  at  the  surface. 

Whether  it  becomes  at  a  depth  of  twenty  miles  less  than  it  is  at 
the  surface  we  cannot  tell,  but  such  a  decrease  is  not  impossible, 
perhaps  not  improbable;  because  at  this  depth  the  molten  rock 
moves  in  earthquakes,  and  yet  in  confinement  it  must  have  a  very 
sensible  rigidity,  though  probably  not  more  than  half  that  of  granite. 

Accordingly,  it  looks  as  if  the  rigidity  at  the  surface  is  about 
half  that  of  glass,  at  a  depth  of  20  miles  about  one  half  that  at  the 
surface,  and  at  the  depth  of  40  miles  nearly  the  same,  but  increas- 
ing below  that  depth  and  at  160  miles  again  equal  to  that  at  the 
surface,  and  at  a  depth  of  400  miles  considerably  larger  yet,  or  about 
1.4  times  that  of  glass.  Increasing  below  this  depth  according  to 
the  pressure,  it  becomes  at  the  center  over  3  times  that  of  nickel 
steel  used  in  armor  plate.  The  rigidity  of  steel  is  attained  at  a  little 
over  0.3  of  the  depth  to  the  center  of  the  earth.  If  this  be  the  dis- 
tribution of  rigidity  in  the  earth,  the  curve  of  rigidity  is  as  follows : 

This  postulated  fall  in  the  rigidity  just  beneath  the  crust  is 
probable  for  several  reasons: 

1.  The  temperature  increases  quite  rapidly  as  we  go  downward, 
while  the  pressure  increases  proportionately  more  slowly,  so  that 
a  depth  would  be  reached  at  which  the  matter  would  become  a 
plastic  if  not  a  viscous  fluid. 

2.  The  eruption  of  volcanoes  and  lava  flows  on  a  vaster  scale 
show  that  a  molten  layer  underlies  the  crust,  and  occasionally  is 
forced  to  the  surface. 

3.  This  imderlying  molten  rock  moves  in  world-shaking  earth- 
quakes, and  frequently  is  expelled  from  beneath  the  sea  under  the 
land  to  form  mountain  ranges  along  the  coast. 

4.  We  may  prove  this  expulsion  of  lava  by  the  observed  seismic 
sea  waves  which  indicate  a  sinking  of  the  sea  bottom,  and  by  the 
simultaneous  uplift  of  mountains  and  coasts. 

From  these  considerations  it  follows  that  the  earth  is  most  nearly 
liquid  just  beneath  the  crust,  and  has  the  greatest  rigidity  at  the 
center.  As  the  plastic  or  quasi-viscous  layer  beneath  the  crust  is 
thin,  and  possessed  of  considerable  rigidity,  it  too  remains  quiescent 


i9o8.] 


THE  PHYSICS  OF  THE  EARTH. 


193 


except  when  set  in  motion  by  the  dreadful  paroxysms  of  an  earth- 
quake. 

In  tidal  and  other  observations  the  earth  therefore  behaves  as  a 
solid,  and  the  rigidity  of  the  earth  inferred  by  Kelvin  and  Darwin 
is  confirmed.  Yet  a  layer  of  plastic  matter  or  quasi-viscous  fluid 
exists  just  beneath  the  crust,  and  when  disturbed  by  earthquakes 
gives  rise  to  the  development  of  ridges  in  the  crust  called  moun- 
tains, chiefly  by  the  expulsion  of  lava  from  under  the  sea. 


■ 

• 

"^ 

\ 

'^ 

\ 

^ 

\ 

\ 

k 

\ 

^ 

\ 

\ 

» 

\ 

• 

1 

\ 

\ 

\ 

> 

\ 

\ 

\ 

\ 

\ 

•> 

\ 

V 

C 

1 fl 

1—Q 

3.     .0 

Rad 
f      0 

iuS 
S1-.J? 

2 Q 

rJ 

Fig.  2.    Curve  of  Rigidity  for  the  Earth,  showing  the  plastic  layer 

just  beneath  the  crust. 

§  i8.  Wiecherfs  Researches  on  the  Interior  Constitution  of  the 
Earth  and  on  the  Plastic  or  Viscous  Layer  which  he  Infers  to  Exist 
Just  Beneath  the  Crust  from  Oscillations  of  Long  Period  Noticed 
in  Seismic  Vibrations, — Professor  E.  Wiechert,  of  Gottingen,  has 
devoted  much  attention  to  the  problem  of  the  constitution  of  the 
earth's  interior.  He  long  ago  reached  the  conclusion  that  the  great 
interior  nucleus  probably  is  a  mass  of  iron  covered  with  a  thick 


194  SEE— FURTHER   RESEARCHES  ON  [April  24, 

shell  of  stony  material.  In  the  paper  which  he  recently  presented 
to  the  International  Seismological  Association  in  session  at  the 
Hague,  September  21-26,  1907,  he  estimates  the  depth  of  the  stony 
layer  as  1,500  kilometers,  which  is  nearly  one  fourth  of  the  earth's 
radius. 

This  view  that  interior  of  the  earth  is  metallic*  has  been  en- 
tertained by  many  eminent  physicists,  including  Lord  Rayleigh; 
but  it  is  beset  with  many  difficulties.  We  shall  here  mention  three 
of  the  principal  objections: 

1.  If  this  constitution  of  the  earth  be  admitted,  the  curve  of 
density  will  have  a  sudden  break  at  a  depth  of  about  one  fourth  of 
the  radius;  and,  as  the  pressure  increases  rapidly  as  we  go  down- 
ward, it  seems  improbable  that  the  density  of  the  outer  layer  could 
remain  uniform  and  then  change  suddenly  at  a  depth  of  one  fourth 
of  the  distance  to  the  center.  Such  discontinuity  in  nature  seems 
highly  improbable  for  the  density,  since  there  probably  is  no  sensible 
discontinuity  in  the  laws  of  pressure  and  temperature. 

2.  If  the  central  nucleus  is  metallic,  it  follows  that  the  denser 
elements  have  separated  from  the  rest  of  the  mass.  As  the  matter 
has  been  essentially  solid  and  highly  rigid,  owing  to  the  pressure, 
ever  since  the  globe  attained  anything  like  its  present  dimensions, 
this  sinking  would  not  be  possible,  because  the  resistance  to  the 
motion  would  be  much  too  gjeat.  Thus  owing  to  resistance  to 
motion  arising  from  rigidity  we  can  not  admit  a  separation  of  the 
denser  from  the  lighter  elements  of  such  a  globe.  If  the  metals 
were  all  so  deep  down,  it  would  be  hard  to  account  for  the  veins 
found  in  the  crust  by  any  kind  of  eruptive  process,  since  the  globe 
is  never  fissured  to  a  depth  of  anything  like  one  fourth  of  the 
radius. 

3.  If  in  addition  to  these  mechanical  objections  we  recall  that 
deep  down  the  pressure  is  so  great  as  to  cause  an  interpenetration 
of  all  the  elements,  whatever  be  the  temperature,  but  especially 
under  the  high  temperature  known  to  prevail  in  the  interior  of  the 
globe,  so  that  no  aggregation  or  crystallization  of  substances  would 
be  possible,  and  the  nucleus  would  therefore  be  a  magma  of  all  the 
elements,  it  becomes  inconceivable  that  the  metals  could  separate 
from  the  stony  elements  by  sinking,  while  the  latter  floated  to  the 


,9o8.j  THE  PHYSICS  OF  THE  EARTH.  195 

surface.  Even  if  the  globe  were  a  liquid  mass  of  very  small  viscosity, 
it  is  clear  that  such  a  separation  of  the  elements  could  not  take  place. 

Finally  it  is  to  be  recalled  that  recent  experiments  with  radium 
have  shown  the  probable  transmutation  of  some  of  the  metals,  as 
when  Sir  Wm.  Ramsay  caused  sulphate  of  copper  to  be  partially 
degraded  into  lithium.  If  this  can  occur  for  one  or  two  metallic 
elements,  it  may  eventually  be  possible  for  many  and  perhaps  all  of 
the  metals.  Our  knowledge  of  these  transformations  is  still  in  its 
infancy,  and  we  can  not  yet  ascertain  how  minerals  and  metallic 
veins  have  arisen;  but  it  is  impossible  to  believe  that  the  material 
has  come  up  from  a  pure  supply  at  a  depth  of  1,500  kilometers. 
It  is  much  more  probable  that  the  metallic  elements  have  been  de- 
veloped by  differentiation  and  transformation  from  an  original 
magma,  and  that  the  whole  interior  of  our  planet  is  still  a  magma. 
Differentiation  of  the  elements  appears  to  develop  under  conditions 
met  with  in  the  crust,  but  nowhere  else. 

Accordingly  we  are  obliged  to  dissent  from  the  constitution  of 
the  globe  outlined  by  Professor  Wiechert ;  but  in  the  matter  of  the 
existence  of  a  layer  of  plastic  or  fluid  material  just  beneath  the  crust, 
which  he  infers  from  the  long  seismic  vibrations  with  periods  of 
about  eighteen  seconds,  we  are  in  hearty  accord  with  him.  This  is 
definitely  proved  by  the  phenomena  noticed  in  earthquakes,  as  more 
fully  set  forth  hereafter.  It  is  the  expulsion  of  lava  from  imder  the 
margins  of  the  sea  which  produces  world-shaking  earthquakes  and 
the  upheaval  of  mountains  along  the  sea  coasts. 

§  19.  On  Sir  G,  H.  Darwin's  Researches  on  the  Stresses  in  the 
Interior  of  the  Earth  Due  to  the  Weight  of  Continents  and  Moun- 
tains, — ^We  have  seen  that  the  earth  behaves  as  a  solid  at  all  depths, 
unless  it  is  in  the  thin  layer  just  beneath  the  crust,  in  which  move- 
ments take  place  during  earthquakes.  The  theory  of  an  elastic 
solid  shows  that  when  such  a  body  is  stressed  the  state  of  stress  is 
completely  determined  when  the  amount  and  direction  of  the  three 
principal  stresses  are  known.  No  limit  is  imposed  on  these  stresses 
by  theory,  but  in  practice  nature  fixes  a  limit,  beyond  which  the 
elasticity  breaks  down,  and  t^^e  solid  either  flows  or  ruptures  by 
breaking. 


196  SEE— FURTHER    RESEARCHES  ON  [April  m. 

In  the  "  Nat.  Phil./'  Vol.  I,  part  II,  §  832,  Lord  Kelvin  and 
Professor  Tait  remark  that 

"The  precise  circumstances  under  which  elastic  bodies  break  have  not 
hitherto  been  adequately  investigated  by  experiment.  It  seems  certain  that 
rupture  cannot  take  place  without  difference  of  stress  in  different  directions. 
One  essential  element  therefore  is  the  difference  between  the  greatest  and 
least  of  the  three  principal  stresses.  How  much  the  tendency  to  break  is 
influenced  by  the  amount  of  the  intermediate  principal  stress  is  quite  un- 
known. The  difference  between  the  greatest  and  least  stresses  may  however 
be  taken  as  the  most  important  datum  for  estimating  the  tendency  to  break. 
This  difference  has  been  called  by  Mr.  G.  H.  Darwin  (to  whom  the  investi- 
gation of  which  we  speak  is  due)  the  *  stress-difference.' " 

Stress-difference  is  a  term  which  when  applied  to  matter  within 
the  earth  denotes  the  tendency  to  flow.  For  rupture  is  not  possible 
when  the  matter  is  in  confinement  under  such  pressure  and  at  high 
temperature.  Now  if  the  earth  were  homogeneous,  as  assumed  in 
Darwin's  inquiry,  the  inequalities  of  surface  due  to  the  mountains, 
plateaus,  and  continents  would  give  rise  to  a  stress-difference  in 
the  underlying  layers ;  and  Darwin  showed  that  the  stress-difference 
would  increase  with  the  depth,  being  at  the  center,  for  inequalities 
of  the  type  represented  by  harmonics  of  the  second  order,  eight 
times  what  it  is  at  the  surface. 

If  the  earth  were  not  effectively  solid  throughout,  a  flow  ought 
to  take  place  either  near  the  surface  or  at  greater  depth ;  and  thus 
the  inequalities  of  surface  would  disappear.  But  the  plateaus  and 
mountains  do  not  sink  in,  and  this  fact  proves  that  the  globe  is  not 
fluid,  and  even  that  the  plastic  or  viscous  layer  just  beneath  the  crust 
is  quite  stiff.  As  we  have  seen  that  the  rigidity  increases  very 
rapidly  towards  the  center,  we  easily  see  why  movement  should 
not  occur  at  great  depth,  since  the  rigidity  there  exceeds  that  of 
any  known  substance,  and  at  the  centre  is  about  three  times  that 
nickel  steel  used  in  armor  plate. 

In  the  paper  on  the  "  Temperature  of  the  Earth  "  we  have  shown 
from  the  evidence  of  stability  afforded  by  geological  pinnacles 
millions  of  years  old,  that  no  movements  of  deep  seated  character 
occur  within  the  earth.  This  evidence  supports  the  view  that  the 
earth  is  effectively  solid,  and  has  behaved  as  such  since  the  con- 
solidation of  the  crust. 


,9o8.]  THE  PHYSICS  OF  THE  EARTH.  197 

As  the  rigidity  increases  so  rapidly  towards  the  center  of  the 
earth,  flow  ought  not  to  take  place  at  those  depths ;  and  the  absence 
of  any  evidence  of  deep  seated  movements  among  the  ruins  wrought 
by  geological  time  in  turn  supports  the  theory  of  rigidity  depending 
on  the  pressure. 

Darwin's  hypothesis  of  homogeneity  is  only  a  rough  approxima- 
tion to  the  truth,  and  Laplace's  law  would  no  doubt  give  a  much 
more  exact  representation  of  the  density  and  the  resulting  stress- 
diflference  in  the  earth.  But  this  suggested  change  of  data  would  not 
greatly  modify  the  general  conclusions  already  stated. 

§  20.  The  Theory  of  Isostacy, — ^A  more  important  diflference 
might  arise  from  the  theory  of  isostacy,  the  applicability  of  which 
to  the  earth  seems  to  be  becoming  better  established  by  recent  re- 
searches. In  this  view  the  crustal  inequalities  seen  at  the  surface 
are  compensated  for  by  lighter  or  greater  densities  beneath,  accord- 
ing as  the  crust  is  elevated  or  depressed,  so  that  for  a  certain  thick- 
ness of  crust  equal  blocks  have  equal  mass,  however  unequal  the 
level  of  the  blocks  at  the  surface. 

The  recent  investigations  by  the  U.  S.  Coast  Survey  indicate 
that  the  depth  of  complete  compensation  for  the  United  States  and 
outlying  stations  is  about  71  miles.  No  doubt  a  depth  of  something 
like  this  extent  would  hold  true  for  the  entire  globe.  If  this  view 
be  admissible,  it  will  follow  that  all  inequalities  of  the  crust  cease 
to  be  effective  at  depths  greater  than  71  miles,  and  no  stress-differ- 
ences depending  on  plateaus  and  mountains  would  exist  in  the  globe 
except  in  the  layers  just  beneath  the  crust.  There  would  thus  be  no 
stresses  in  the  deep  interior  depending  on  the  weight  of  continents 
and  mountains. 

This  theory  of  isostacy  is  confirmed  by  the  theory  of  mountain 
formation  developed  in  the  paper  on  the  "  Cause  of  Earthquakes," 
which  shows  clearly  that  these  elevated  ridges  are  underlaid  by 
itiaterial  lighter  than  the  average  rock  of  the  crust.  On  the  one 
hand,  therefore,  if  stress-differences  exist  deep  down,  no  move- 
ment can  take  place,  owing  to  rigidity ;  on  the  other,  if  the  theory  of 
isostacy  be  admissible,  no  stress-differences  can  exist  except  in  the 
outer  layers  of  the  globe,  within  71  miles  of  the  surface. 

We  conclude  therefore  that  in  no  case  could  movements  occur 


198  SEE— FURTHER   RESEARCHES  ON  [April  .4, 

except  in  the  layer  just  beneath  the  crust.  These  superficial  move- 
ments are  called  earthquakes,  and  are  caused  chiefly  by  the  leakage 
of  the  oceans.  Observations  show  that  the  depth  of  such  disturb- 
ances in  all  cases  is  less  than  40  miles.  This  accords  with  the 
theory  of  isostacy,  and  confirms  the  conclusions  drawn  from  that 
theory  that  all  surface  inequalities  are  compensated  for  at  but  a 
slight  depth. 

§  21.  Uplifts  along  the  Andes  show  that  the  mountains  are  not 
sinking  under  their  own  weight. — In  Professor  Sir  G.  H.  Darwin's 
paper  on  the  stresses  in  the  earth,  above  cited,  he  has  also  con- 
sidered harmonics  of  high  order,  corresponding  to  the  case  of  a 
series  of  parallel  mountains  and  valleys,  which  thus  corrugate  a 
mean  level  surface  with  an  infinite  series  of  parallel  ridges  and 
furrows.  Here  the  stress-difference  depends  only  on  the  depth  be- 
low the  surface,  and  is  independent  of  the  position  of  the  point 
considered  with  respect  to  ridge  and  furrow.  Taking  a  series  of 
mountains  13,000  feet  (about  4,000  meters)  above  the  valley  bottoms, 
formed  of  granite  of  density  2.8,  he  shows  that  the  maximum 
stress-difference  is  4  X  10*^  grammes  weight  per  square  centimeter 
(about  the  tenacity  of  cast  tin).  And  when  the  mountain  chains  are 
314  kilometers  apart,  making  the  ridges  about  78  times  wider  than 
they  are  deep,  the  maximum  stress-difference  is  reached  at  a  depth 
of  50  kilometers  below  the  surface,  or  at  a  depth  of  I2j4  times  the 
height  of  the  mountains  above  the  valleys.  Thus  for  mountains  of 
the  height  of  our  average  ocean  depth,  the  maximum  tendency  to 
flow  would  be  at  a  depth  of  about  31  miles.  (Cf.  "  Nat.  Phil," 
Vol.  I,  Part  II,  §832.) 

If  earthquake  shocks  were  due  to  such  flowage  the  mountains 
would  be  gradually  reduced  in  height.  Instead  of  this  settling  oc- 
curring, mountains  like  the  Andes  are  still  rising,  as  we  may  infer 
from  the  fact  that  after  an  earthquake  the  adjacent  sea  coast  often 
is  elevated  and  higher  than  before;  while  the  sinking  of  the  adjacent 
sea  bottom,  indicated  by  the  accompanying  seismic  sea  wave,  shows 
that  the  bed  of  the  sea  was  undermined  by  #ie  expulsion  of  the 
material  pushed  under  the  land  and  mountains.  This  state  of  fact 
emphatically  contradicts  the  view  that  these  great  seismic  disturb- 
ances are  due  to  the  flowage  beneath  the  crust  arising  from  the 


i9o8.) 


THE  PHYSICS  OF  THE   EARTH.  199 


weight  of  continents  and  mountains.  Neither  the  uplift  of  moun- 
tains about  the  sea  coasts,  nor  the  earthquakes  occurring  in  these 
regions  can  be  explained  by  flowage  beneath  the  crust,  because  the 
movement  is  positive  rather  than  negative,  as  required  by  this  theory. 
Whilst  the  investigation  of  Professor  Sir  George  Darwin  there- 
fore does  not  give  us  a  clue  to  the  observed  movement^,  it  is  never- 
theless Asery  valuable  as  furnishing  an  indirect  confirmation  of  the 
present  theory  that  mountain  formation  depends  on  the  sea.  Ob- 
servation shows  that  the  movements  are  positive,  and  as  the  theory 
of  flowage  indicates  that  they  should  be  negative,  we  may  infer 
that  whatever  be  the  stress-differences  existing  beneath  the  earth's 
crust,  the  movements  thus  produced  are  insensible  compared  to  those 
depending  on  the  expulsion  of  lava  from  under  the  sea  by  world- 
shaking  earthquakes. 

III.  The  New  Physical  Theory  of  Earthquakes  and  Moun- 
tain Formation  Based  on  the  Secular  Leakage 

OF  the  Ocean  Bottoms. 

§  22.  On  the  Plastic  and  Perhaps  Viscous  Layers  Just  Beneath 
the  Earth's  Crust. — ^We  have  now  examined  at  length  the  arguments 
in  regard  to  the  constitution  of  the  earth's  interior,  and  have  shown 
that  although  as  a  whole  the  earth  is  solid,  owing  to  the^pressure  to 
which  the  matter  is  subjected,  there  is  a  plastic  layer  just  beneath 
the  crust  which  in  earthquakes  is  made  to  flow  and  behave  almost 
as  a  viscous  fluid.  In  this  layer  just  beneath  the  crust  either  the 
pressure  is  not  great  enough  to  produce  entire  solidity,  with  the 
existing  temperature,  or  else  the  solid  is  made  to  flow  by  the  break- 
ing down  of  the  elasticity  under  the  action  of  the  earthquake  forces, 
which  are  powerful  enough  to  disturb  the  whole  world. 

Although  the  matter  in  this  substratum  appears  to  have  some 
rigidity,  it  seems  probable  that  it  has  not  the  requisite  elasticity  to 
behave  as  a  perfect  solid.  We  know  that  the  layer  must  be  nearly 
solid,  because,  if  it  were  not  so,  there  would  be  a  greater  tendency 
of  the  mountains  to  subside  than  actually  is  observed.  The  stress- 
difference  in  the  layers  just  beneath  the  crust  must  be  very  con- 
siderable ;  and  yet  this  plastic  matter  is  so  stiff  that  it  does  not  flow 
and  allow  the  mountains  and  plateaus  to  sink  in. 


200  SEE— FURTHER   RESEARCHES  ON  [April  24. 

Now  earthquake  disturbances  are  often  complex,  and  consist  in 
horizontal  and  vertical  movements  combined.  We  have  seen  that  in 
the  long  run  the  uplifting  tendency  predominates,  because  it  is  in 
this  way  that  the  mountains  and  plateaus  have  arisen.  Nevertheless 
there  are  numerous  cases  in  which  subsidences  take  place,  and  these 
settlements  often  seem  to  be  somewhat  gradual,  as  if  .the  substratum 
was  slowly  yielding  and  flowing  under  the  stresses  to  which  it  is 
subjected.  These  gradual  subsidences,  of  the  class  that  was  observed 
by  Darwin  and  Fitzroy  at  Conception  in  1835,  seem  to  aflford  con- 
vincing evidence  that  the  layer  beneath  the  crust  is  certainly  plastic, 
perhaps  viscous.*  The  yielding  of  the  layer  beneath  the  crust  is 
shown  not  only  in  movements  noticed  in  earthquakes,  when  lava  is 
expelled  from  under  the  sea  and  pushed  under  the  land ;  but  also  in 
the  subsidences  which  the  sea  trenches  experience  after  earthquakes. 
These  subsidences  have  folded  the  rocks  seen  in  mountain  ranges 
now  on  land ;  and  although  most  of  such  subsidence  is  due  to  the 
imdermining  of  the  troughs  by  the  expulsion  of  lava,  it  s^ems  likely 
that  some  very  gradual  yielding  also  takes  place.  The  layer  under 
the  crust  is  therefore  certainly  plastic,  when  partially  imdermined, 
and  probably  so,  independent  of  the  imdermining,  if  it  is  subjected 
to  great  forces,  as  in  world-shaking  earthquakes,  where  mountains 
are  in  process  of  upheaval.  If  the  matter  is  also  viscous,  the  viscosity 
must  be  very  high.  With  the  matter  imprisoned  beneath  the  earth's 
crust  it  is  difficult  if  not  impossible  to  distinguish  between  plasticity 
and  true  viscosity,  because,  if  the  fluid  is  very  stiff,  it  would  behave 
almost  as  a  solid.  And  the  tests  heretofore  afforded  by  earthquakes 
are  not  decisive.  This  view  of  the  substratum  just  beneath  the  crust 
is  not  essentially  different  from  the  theory  held  by  Arrhenius  with  re- 
gard to  the  interior  of  the  earth  as  a  whole.  But  this  layer  is  the 
only  part  of  the  interior  in  which  movements  may  be  observed,  and 
even  here  movements  would  not  take  place  but  for  the  steam  de- 
veloped beneath  the  crust  by  the  secular  leakage  of  the  oceans.  It 
may  be  that  the  future  study  of  these  movements  will  some  day 

*We  follow  Sir  George  Darwin  in  "distinguishing  viscosity,  in  which 
flow  is  caused  by  infinitesimal  forces,  from  plasticity  in  which  permanent 
distortion  or  flow  sets  in  when  the  stresses  exceed  a  certain  limit."  (Letter 
to  Sir  A.  Geikie,  January  9,  1884.) 


«9o8.] 


THE  PHYSICS  OF  THE   EARTH.  201 


enable  us  to  decide  whether  the  substratum  is  plastic  only,  or 
truly  viscous. 

§  23.  Substratum  Everywhere  Quiescent  Except  when  Disturbed 
by  Earthquakes, — ^The  fact  the  large  areas  of  the  earth's  surface  in 
such  dry  countries  as  Sahara,  our  Western  Plateaus,  and  the  interior 
of  Australia,  are  quite  free  from  earthquake  disturbances,  shows 
what  would  happen  everywhere  but  for  the  presence  of  surface 
water,  and  especially  the  leakage  of  the  crust  depending  on  the  sea. 
The  quiescence  of  the  substratum  in  interior  regions  remote  from 
the  sea  shows  that  imder  normal  conditions  this  layer  is  quite  inert. 
It  is  only  set  in  motion  by  the  vapor  of  steam  which  slowly  develops 
stresses  in  the  rocks  of  the  crust  and  finally  brings  on  earthquakes. 
It  might  be  plastic  enough  to  yield  slightly  under  sufficiently  great 
forces,  but  the  loading  and  unloading  due  to  meteorological  and 
geological  causes  going  on  in  nature  are  not  great  enough  to  have 
any  appreciable  effect,  as  we  may  infer  from  the  universal  quiescence 
of  inland  areas,  especially  in  desert  countries. 

It  seems  to  be  true,  however,  that  when  the  crust  is  broken  and 
upheaved,  in  the  formation  of  mountains  near  the  sea  coast,  some 
slow  yielding  takes  place  beneath.  Yet  at  present  any  changes  of  a 
creeping  nature  can  not  be  entirely  separated  from  those  depending 
on  the  expansion  and  expulsion  of  lava  from  under  the  sea;  and 
we  can  only  feel  sure  of  the  inert  character  of  the  substratum,  except 
where  disturbed  by  water  vapor  entering  from  without.  Along  the 
sea  coasts  the  stresses  in  the  crust  are  constantly  changing,  and  the 
crust  blocks  yielding  more  or  less  to  the  stresses  acting  upon  them ; 
it  is  only  when  sudden  yielding  occurs  that  we  experience  a  shock, 
and  the  greatest  earthquakes  are  characterized  by  molten  rock  ad- 
justing itself  beneath  the  crust.  It  is  probable  that  much  yielding 
takes  place  which  is  exceedingly  gradual  and  produces  no  disturb- 
ances sensible  to  ordinary  observation.  In  dry  regions  remote  from 
the  sea  there  are  no  shocks,  and  therefore  also  no  gradual  yielding 
of  the  crust;  hence  the  substratum  is  inherently  and  naturally  qui- 
escent except  when  disturbed  by  external  forces. 

§  24.  Mountain  Formation  in  the  Sea  and  on  the  Land. — In  the 
paper  on  the  *'  New  Theory  of  Earthquakes  and  Mountain  Forma- 
tion," we  have  cited  certain  cases  of  mountain  formation  now  going 


SEE— FURTHER    RESEARCHES  ON 


Fig.  3.  Relief  Map  of  Norlh  America.  (From  Frye's  Complete  Geog- 
raphy, by  pennission  of  Ginn  &  Co.,  Publishers.)  This  map  illustrates  beauti- 
fully the  recession  of  the  sea  since  the  formation  of  the  Rocky  Mountains, 
which  were  at  one  time  the  eastern  border  of  the  Pacific  Ocean. 


,9o8]  THE  PHYSICS  OF  THE   EARTH.  203 

on  in  the  depths  of  the  sea,  and  directly  connected  with  mountain 
systems  spread  out  on  the  land.  Thus  we  have  shown  that  the 
Aleutian  Islands  are  a  branch  or  part  of  the  Rocky  Mountains  still 
remaining  in  the  depths  of  the  sea.  As  this  part  of  the  chain  is  now 
being  uplifted  by  the  ocean,  we  get  a  very  clear  conception  of  how 
the  whole  Rocky  Mountain  system  was  formed.  We  are  fortunate 
therefore  to  find  a  part  of  a  great  mountain  chain  still  unfinished, 
with  one  end  under  water  and  the  main  body  of  the  system  high 
and  dry  along  the  edge  of  the  continent. 

Now  no  one  believes^  that  mountain  formation  takes  place  far 
inland,  because  the  mountains  generally  follow  the  coast,  and  more- 
over at  present  the  process  is  found  to  be  most  active  in  the  sea, 
as  in  the  region  of  the  Aleutian  Islands  and  the  Antandes.  This 
geographical  distribution  of  mountain-making  is  therefore  a  most 
powerful  argument  for  the  new  theory.  Moreover  it  is  generally 
recognized  that  the  Rocky  Mountains  in  the  United  States  are  a 
good  deal  older  than  the  Andes  in  South  America;  and  as  the 
relative  ages  bear  some  relation  to  the  distances  from  the  sea,  the 
mountains  on  land  give  the  same  indication  as  those  still  in  the 
depths  of  the  sea.  The  recession  of  the  sea  ^bes  on  at  very  unequal 
rates  in  diflFerent  parts  of  the  world,  yet  the  present  positions  of  the 
mountains  show  that  the  older  mountains  are  generally  remote  from 
the  ocean.  The  present  theory  is  therefore  confirmed  by  the  lay  of 
the  older  as  well  as  of  the  younger  mountain  systems;  and  by  the 
situation  of  the  mountains  on  land  as  well  as  of  those  now  being 
formed  in  the  depths  of  the  sea.  All  the  mountain  phenomena  of 
the  globe  are  thus  shown  to  be  consistent.  But  as  direct  observation 
of  mountain  formation  witnessed  with  our  own  eyes  is  the  most 
convincing  of  all  evidence,  it  is  fortunate  that  we  are  able  to  cite 
numerous  cases  of  mountain  ranges  now  developing  in  the  sea.  By  • 
the  study  of  the  sinking  going  on  where  trenches  are  developing, 
we  see  how  the  wrinkles  and  valleys  were  produced  in  mountain 
systems  now  at  a  considerable  distance  from  the  ocean.  Since  the 
sea  recedes  from  the  mountains  in  the  course  of  geological  ages,  it 
follows  that  more  and  more  land  is  constantly  rising  above  the  water, 

*  Compare  §  42  of  this  paper,  where  Leconte's  views  are  quoted  at  length. 
He  held  that  mountain  ranges  are  formed  on  lines  of  thick  sediment  along 
the  shores  of  continents. 


204  SEE-FURTHER  RESEARCHES  ON  [Aprils. 

and  the  continents  growing  larger.  The  mountains  are  formed  by 
earthquakes,  and  earthquakes  are  due  to  the  sea,  which  thus  makes 
more  and  more  land  for  the  development  of  the  higher  forms  of 
life  upon  the  globe. 


Fic  4.    Relief  Map  of  the  United  States.     (From  Fiye's  Complete  Geog- 
especially  how  the  great  plateau   west  of  the  Rocky  Mountains  has  been 


t^AJ  THE  PHYSICS  OF  THE   EARTH,  206 

§  25.  The  Origin  of  Faults  in  the  Earth's  Crust. — It  has  long 
been  recognized  that  faults  in  the  earth's  crust  are  often  displaced 
by  earthquakes.  Now  earthquakes  are  mainly  submarine  or  follow 
the  borders  of  the  continents.    Here  the  mountain  ranges  have  de- 


raphy,  by  permission  of  Ginn  &  Co.,  Publishers.)     Tbe  reader  should  notice 
crumpled  in  the  uplift  from  tbe  sea,  which  has  receded  westward  1,000  miles. 


206  SEE— FURTHER   RESEARCHES  ON  [April  24, 

veloped  or  are  now  developing,  and  in  general  the  faults  run  along 
the  sea  coasts  and  into  the  sea,  where  mountain  formation  is  in 
progress.  Thus  it  is  clear  that  faults  arise  from  the  stresses  and 
movements  of  the  crust  produced  by  earthquakes  and  mountain 
formation,  and  therefore  from  the  secular  leakage  of  the  ocean 
bottoms. 

Sometimes  the  faults  move  but  little,  at  other  times  they  give 
rise  to  conspicuous  changes  of  level ;  and  where  vast  down-throws  or 
uplifts  have  occurred  certain  types  of  mountains  arise  from  normal 
faulting.  The  more  horizontal  movements  of  faults  arise  mainly 
in  the  trenches  along  the  sea  coasts,  which  produce  the  folding  seen 
in  mountain  chains.  The  vertical  movements  are  more  general,  and 
are  especially  conspicuous  in  elevated  plateaus,  like  those  of  our 
western  states. 

In  his  "  Report  on  the  Geology  of  the  High  Plateaus  of  Utah," 
Washington,  1880,  Major  Button  gives  a  description  of  some  of  the 
most  magnificent  faults  in  the  world.  On  page  45  he  indicates  the 
dependence  of  these  faults  on  the  ancient  shore  line  of  the  Eocene 
lake,  thus: 

"It  yet  remains  to  speak  of  another  interesting  relation  of  the  later 
system  of  faults.  They  have  throughout  preserved  a  remarkable  and  per- 
sistent parallelism  to  the  old  shore  line  of  the  Eocene  lake,  following  the 
broader  features  of  its  trend  in  a  striking  manner.  The  cause  of  this  rela- 
tion is  to  me  quite  inexplicable,  so  much  so,  that  I  am  utterly  at  a  loss  to 
think  of  any  subsidiary  facts  which  may  be  mentioned  in  connection  with  it 
and  which  can  throw  light  upon  it." 

What  puzzled  Major  Button  most  was  the  raising  of  the  area  of 
the  lake;  but  as  the  whole  region  was  uplifted  by  the  sea  in  later 
times  this  phenomenon  was  in  no  way  remarkable.  The  rocks  in  such 
disturbed  regions  have  been  broken  and  folded  into  a  series  of 
troughs  and  arches  or  thrown  into  domes  and  basins,  and  probably 
no  two  adjacent  areas  retained  their  relative  levels  throughout.  His 
observation,  however,  confirms  the  present  theory  that  faulting  is 
generally  parallel  to  the  ancient  sea  shore,  and  therefore  produced 
originally  by  the  oceans. 

The  conspicuous  character  of  the  vertical  movement  of  the  crust 
blocks  in  the  region  of  the  Great  Basin  led  several  American  geolo- 
gists to  suggest  that  vertical  forces  had  operated  in  the  uplift  of 


i9o8.]  THE  PHYSICS  OF  THE  EARTH.  207 

these  plateaus.  As  the  whole  region  has  been  raised  from  the  sea  by 
the  injection  of  the  land  with  lava  pushed  under  the  crust  from  be- 
neath the  sea,  it  is  evident  that  the  crust  blocks  ought  to  be  displaced 
unequally  in  different  places,  and  hence  the  various  types  of  faulting 
observed. 

It  should  be  remarked,  however,  that  in  the  elevation  of  a  plateau 
a  mile  high,  only  a  layer  of  lava  a  mile  deep  needs  to  be  injected.  If 
three  miles  high,  the  layer  would  have  to  be  three  miles  thick; 
but  even  this  maximum  height  is  only  about  one  seventh  of  the 
thickness  of  the  crust ;  and  hence  eruptions  would  not  usually  occur 
in  these  uplifts.  The  plateaus  are  all  of  small  height  compared  to 
the  thickness  of  the  earth's  crust,  beneath  which  the  movement 
of  molten  rock  takes  place. 

If  some  faults  should  thus  be  widely  opened,  lava  flows  of  vast 
extent,  like  those  in  Utah  and  Oregon,  might  be  expected  to  occur. 
We  cannot  give  the  details  of  the  cracks  which  produced  these  gi- 
gantic outflows,  but  it  is  evident  that  they  depended  on  the  opening 
of  immense  faults.  Now  the  faults  are  produced  and  moved  by 
earthquakes,  and  earthquakes  are  due  to  the  leakage  of  the  oceans. 
It  follows  therefore  that  the  most  immense  lava  flows  ought  to  take 
place  near  the  sea ;  and  this  seems  to  be  true  both  in  North  America 
and  in  Asia,  where  the  outflow  in  the  plateau  of  Deccan  has  always 
excited  the  wonder  of  the  naturalist. 

That  all  the  faults  of  the  earth's  crust  depend  on  the  sea  and  are 
produced  by  world-shaking  earthquakes,  is  clearly  indicated  by  the 
geographical  distribution  of  these  cracks  in  the  crust.  If  any  other 
cause,  such  as  the  secular  cooling  of  the  globe,  were  at  work,  we 
should  find  a  relatively  greater  predominance  of  faults  far  inland, 
which  is  contrary  to  observation,  especially  in  dry  countries. 

It  is  remarkable  that  geologists  have  referred  so  many  phenomena 
to  faulting,  but  have  made  little  or  no  attempt  to  explain  faulting 
itself.  In  the  present  theory  referring  the  origin  of  faults  to  the 
expulsion  of  lava  from  under  the  sea  we  have  for  the  first  time  a 
satisfactory  and  consistent  view  of  these  phenomena.  Faults  evi- 
dently arise  mainly  from  the  motion  of  lava  in  earthquakes,  by  which 
the  overlying  rocks  of  the  crust  are  broken,  and  often  displaced  along 
the  line  of  fracture. 


208  SEE— FURTHER   RESEARCHES  ON  [April  04, 

When  the  crust  is  thus  rent  into  blocks,  some  of  them  are  re- 
duced to  small  size,  and  eventually  raised  up,  as  in  the  vertical  walls 
of  granite  now  seen  in  Smyth's  channel,  southern  Chile,  the  Straits 
of  Magellan,  Yosemite  Valley,  California,  and  the  fiords  of  Nor- 
way. These  precipitous  walls  of  g^nite  could  be  pushed  up  only 
by  vertical  forces,  in  earthquakes.  It  is  noticeable  that  no  such 
isolated  masses  are  found  towering  up  in  the  plains  of  Kansas,  the 
desert  of  Sahara,  and  other  inland  regions  far  from  the  oceans.  The 
origin  of  faults  and  fault  movements  must  therefore  be  sought  in 
the  leakage  of  the  oceans  and  in  the  resulting  relief,  which  takes 
place  in  the  sea  bottoms  and  along  the  borders  of  the  continents. 

§  26.  On  the  Uplift  of  the  Great  Plateaus  of  the  World  and  on 
the  Gradual  Elevation  of  the  Continents, — ^For  reasons  already  amply 
set  forth  in  §  7,  the  process  involved  in  the  formation  of  the  Andes 
is  clear  and  beyond  dispute.  Now  it  happens  that  the  Andean  pla- 
teaus, such  as  those  of  Quito,  Caxamarca,  Cuzco  and  Titicaca,  arc 
generally  included  between  the  eastern  and  western  ranges  of  the 
Andes,  and  were  evidently  uplifted  by  the  same  forces  which  formed 
the  mountains  themselves.  Accordingly  it  is  tlear  that  a  plateau  such 
as  that  of  Titicaca  was  therefore  uplifted  by  the  expulsion  of  lava 
from  under  the  sea. 

If  now  we  pass  from  the  Andes  to  the  Himalayas,  we  shall  find 
that  in  like  manner  those  great  mountains  of  Asia  were  uplifted 
principally  by  the  Indian  Ocean.  The  plateau  of  Thibet  in  the 
Himalayas  of  Asia  corresponds  exactly  with  that  of  Titicaca  in  the 
Andes  of  South  America;  and  as  the  latter  was  formed  with  the 
Andes,  so  also  the  plateau  of  Tibet  was  formed  with  the  Himalayas. 
This  seems  absolutely  clear  and  incontrovertible.  And  a  similar 
mode  of  development  must  be  ascribed  to  the  table  lands  to  the 
east  and  west  of  Tibet,  so  that  the  principal  plateaus  of  Asia,  Tibet 
and  Iran,  are  clearly  the  work  of  the  sea. 

The  highest  part  of  these  plateaus  is  Tibet,  with  an  average 
elevation  of  about  15,000  feet,  and  a  width  of  about  500  miles  at  the 
highest  part.  At  the  middle  it  is  somewhat  wider,  and  to  the  west 
it  narrows  into  Little  Tioet,  less  than  half  the  width  of  Tibet  proper. 
It  is  evident  that  great  Tibet  was  uplifted  chiefly  by  movements 
from  the  direction  of  the  bay  of  Bengal ;  this  is  shown  by  the  lay  of 


,^]  THE   PHYSICS   OF  THE   EARTH.  209 

the  mountain  chains  south  of  Tibet,  and  by  the  great  earthquake 
belt  still  persisting  in  the  valleys  of  the  Ganges  and  Brahmaputra. 
In  the  case  of  North  America  the  plateaus  are  broader  and  cor- 
respondingly lower  than  those  of  South  America  and  Asia.  But 
if  the  sea  gave  rise  to  the  uplifts  connected  with  the  Andes  and 
Himalayas,  can  anyone  doubt  that  the  plateaus  of  North  America 
are  due  to  the  same  cause?    The  total  volume  of  the  North  American 


Fig.  S-  Relief  Map  of  Asia.  (From  Frye's  Complete  Geography,  by 
permission  of  Ginn  &  Co.,  Publishers.)  The  mountains  along  the  east 
coast  illustrate  the  successive  stages  in  the  recession  of  the  Pacific  Ocean. 
At  sonic  future  time  the  border  of  the  continent  will  extend  to  the  string 
of  islands  running  from  Kamchatka  (o  the  Philippines,  the  shallow  seas  of 
Japan  and  China  becoming  inland  valleys. 


SEE— h;rther  researches  on 


Fig.  6.  Relief  Map  of  Africa.  (From  Frye's  Complete  Geography,  by 
permission  of  Ginn  8f  Co.,  Publishers.)  The  reader  should  notice  how  the 
highest  mountains  along  the  east  coast  face  the  Indian  Ocean,  which  is  a 
of  the  Pacific. 


1908.J  THE   PHYSICS  OF  THE   EARTH.  211 

plateau  is  comparable  with  that  in  Asia,  and  it  is  easy  to  see  how 
the  relief  of  the  Pacific  on  our  side  may  have  taken  the  form  of  a 
table-land  of  greater  width  but  smaller  height.  The  numerous 
parallel  mountain  chains  west  of  the  Rocky  Mountains  show  the 
nature  of  the  mighty  forces  at  work,  and  prove  that  this  uplift  was 
the  work  of  the  Pacific  Ocean. 

§  27.  The  forces  which  have  raised  the  mountains  and  plateaus 
of  the  globe  are  identical  with  those  which  have  raised  the  conti- 
nents above  the  sea,  and  all  these  forces  depend  on  the  leakage  of  the 
oceans, — The  geological  evidence  of  the  slow  operation  of  the  forces 
which  have  uplifted  the  plateaus  and  mountains  shows  the  immeas- 
urable ages  during  which  they  have  been  at  work.  Sometimes  large 
portions  of  a  continent  have  risen  for  a  time,  and  again  slowly 
subsided,  and  thus  have  arisen  the  phenomena  noted  in  the  sedi- 
mentary rocks  studied  in  geology.  These  gentle  movements  often 
are  without  violent  earthquake  shocks,  because  the  yielding  is  very 
gradual,  and  the  crust  is  slowly  raised  up  and  down  without  breaking. 
It  is  only  where  the  expulsion  of  lava  from  under  the  sea  is  rapid 
and  violent  that  breaking  develops  at  such  rate  as  to  form  mountain 
chains  and  plateaus.  The  uplift  of  a  plateau  also  requires  a  large 
amount  of  material.  Where  the  process  is  gentle  and  gradual  a 
whole  continent  may  be  slowly  uplifted,  and  this  process  evidently 
has  raised  the  low  broad  plains  above  the  water.  The  ccmse  of 
epeirogenic  and  of  orogenic  movements  is  everywhere  one  and  the 
same.  The  movements  take  different  forms  according  to  the  sud- 
denness with  which  the  forces  act;  but  both  depend  on  the  leakage 
of  the  oceans,  and  not  at  all  on  the  secular  cooling  of  the  globe,  the 
effect  of  which  is  insensible.^ 

*  Since  this  was  finished  the  writer  has  carefully  recalculated  the  shrink- 
age of  the  earth's  radius  in  2,000  years,  and  finds  that  it  can  not  exceed  1.5 
inches.  This  takes  no  account  of  the  increase  of  the  interior  heat  of  our 
globe  due  to  radio-activity.  If  this  latter  effect  were  taken  into  account 
probably  there  would  be  no  shrinkage  whatever.  Quite  independently  of 
these  effects,  however,  there  is  an  actual  expansion  of  the  globe  due  to  the 
leakage  of  the  oceans. 

In  the  same  way  it  is  found,  by  the  application  of  Fourier's  theory  of 
heat  to  the  cooling  at  the  surface,  that  the  total  shrinkage  in  the  length  of 
a  continent  such  as  North  or  South  America,  assumed  to  be  equal  to  the 
terrestrial  radius  in  length,  is  less  than  1.5  inches.     This  again  takes  no 


212  SEE— FURTHER   RESEARCHES   ON  [April  .4, 

Such  an  inference  seems  justified  by  the  study  of  the  mountains 
and  plateaus  of  the  world,  and  also  by  the  movement  of  the  strand 
line  which  Professor  Suess  has  so  carefully  traced  in  every  country. 
Almost  everywhere  the  level  of  the  sea  has  been  lowered  in  recent 
geological  time. 

During  his  travels  in  South  America,  Darwin  recorded  many 
observations  to  show  that  Patagonia  and  the  whole  end  of  the 
continent  south  of  the  La  Plata  had  been  recently  elevated  above 
the  sea;  and  he  mentions  a  channel  in  the  Andes  quite  a  distance 
north  of  the  Straits  of  Magellan  which  gave  evidence  of  the  former 
passage  of  the  sea  through  it.  In  view  of  these  well-established 
facts,  can  any  one  doubt  that  the  Straits  of  Magellan  will  eventually 
become  dry  and  Tierra  del  Fuego  be  added  to  Patagonia?  This 
whole  region  shows  vast  walls  of  rock  towering  vertically  thousands 
of  feet  above  the  sea;  evidently  they  were  uplifted  by  earthquake 
forces  from  beneath,  sometimes  working  quietly,  and  again  spas- 
modically. 

As  surely  as  Calabria  in  Italy  has  been  uplifted  from  the 
Mediterranean,  by  that  sea,  just  so  surely  has  the  southern  end  of 
South  America  been  raised  up  by  the  southern  ocean.  And  if  an 
end  of  a  continent  can  be  upraised,  obviously  whole  continents  can 
be  uplifted.  Accordingly  in  the  leakage  of  the  oceans  and  the  relief 
taking  place  under  the  land  which  bounds  them  we  have  the  true 
cause  of  continent-making. 

Some  original  inequalities  of  surface  may  have  existed  after  the 
detachment  of  the  moon  from  the  consolidating  globe,  but  these 
have  since  been  enormously  increased  by  the  effects  resulting  from 
the  leakage  of  the  oceans.  As  the  earth  gets  older,  the  lithosphere 
becomes  more  diversified,  and  the  face  of  the  earth  more  and  more 
wrinkled. 

The  situation  of  the  great  plateaus  of  the  world  facing  the  largest 
oceans  gives  a  clear  indication  of  the  nature  of  the  forces  at  work 

account  of  radium,  the  effect  of  which  would  be  to  diminish  this  calculated 

shrinkage,   or  do  away  with   it  entirely.      By  such   comparisons  as   these,  ' 

placed  along  side  of  the  large  horizontal  and  vertical  movements  noticed  in 

earthquakes  near  the  sea,  which  sometimes  amount  to  from  30  to  50  feet  at  a 

single  disturbance,  we  see  the  utter  untenability  of  the  old  theories  heretofore 

current  in  works  on  geology  and  the  related  sciences.     Note  added  July  a8, 

1908. 


THE   PHYSICS  OF  THE  EARTH. 


Fig.  7.  Relief  Map  of  Australia.  (From  Frye's  Complete  Geography, 
by  permission  of  Ginn  &  Co.,  Publishers.)  The  reader  should  notice  how 
the  largest  mountains  along  the  east  coast  face  the  Pacific  Ocean. 


SEE— FURTHER   RESEARCHES   ON 


Fig.  8.  Relief  Map  of  Europe.  (From  Frye's  Complete  Geography,  by 
permission  of  Ginn  and  Co.,  Publishers.)  The  reader  should  notice  how  the 
principal  mountain  chains  face  the  Mediterranean  and  the  Atlantic.  There  is 
a  trough  in  the  sea  bottom  off  the  Scandinavian  coast  to  which  Professor 
Schiaparelli  has  called  a 


1908.] 


THE  PHYSICS  OF  THE  EARTH. 


215 


by  which  these  mighty  uplifts  have  been  produced.  The  complex 
folding  of  the  mountains  to  the  east  of  Tibet  shows  that  the  Pacific 
aided  the  Indian  Ocean  in  producing  this  great  uplift,  but  we  cannot 
yet  determine  the  relative  importance  of  the  parts  played  by  the  two 
oceans.  ^ 

§  28.  On  the  Origin  of  the  Alps  and  on  the  Extreme  Crumpling 
and  Folding  which  They  Exhibit, — ^The  remarkable  crumpling  and 
folding  noticed  in  the  Alps  has  long  been  a  matter  of  surprise  and 
wonder  to  the  naturalist.  This  phenomenon  has  always  presented 
great  difficulty  to  those  who  have  attempted  to  explain  the  origin 
of  the  Alps.  In  the  paper  on  the  "  Cause  of  Earthquakes  "  (§§  14, 
16,  18,  23)  we  have  outlined  the  theory  of  how  the  Alps  were 
formed  by  the  sea,  and  criticised  the  old  theories  as  totally  inade- 
quate to  account  for  the  observed  crumpling.  We  propose  here  to 
develop  the  new  theory  a  little  further,  and  to  show  how  it  accounts 
for  all  the  facts  observed  in  a  range  such  as  the  Swiss  Alps,  which 
are  generally  recognized  as  about  the  most  complex  system  of  moun- 
tains known  upon  the  globe.  If  the  new  theory  will  explain  the 
Swiss  Alps,  it  will  obviously  explain  any  other  mountain  system  in 
the  world.  The  test  of  the  theory  as  applied  to  the  Alps  may  there- 
fore be  regarded  as  an  experimentum  crucis. 


Fig.  9.    Complex  Folding.    Section  Across  the  Alps  from  the  Neighborhood 
of  Zurich  toward  Como;  about  no  miles.     (Heim  and  Prestwich.) 


ITiW^dOt 


Chmpo-LoB^ 


Fig.  10.  Section  through  the  Alps,  Showing  the  Effects  of  Complex 
Folding.  (From  Heim's  Gebirgsbildung.)  The  line  of  the  St.  Gotthard 
Tunnel  and  the  plane  of  equal  temperature,  AB,  beneath  it,  are  compiled 
from  F.  Giordano,  in  Bollctino  del  R.  Cotnitato  Geologico  d* Italia,  Vol.  XL, 
1880,  pp.  408-50. 


216  SEE— FURTHER   RESEARCHES  ON  [April  x,. 

The  accompanying  figures  exhibit :  ( i )  A  general  section  of  the 
Alps  from  Zurich  to  Lake  Como  (Heim  and  Prestwich),  and  (2)  a 
section  on  a  larger  scale  of  a  portion  of  the  central  Alps  (from 
Heim's  Gebirgsbildung)  with  fan-shaped  folds  and  inversion  of 
strata  on  the  two  sides.  It  can  hardly  be  assumed  that  these  illus- 
trations are  extremely  accurate,  but  no  doubt  they  are  free  from 
large  errors  in  exhibiting  the  general  character  of  the  folding,  which 
gives  here  and  there  fan-shaped  structures  with  overturned  dips  at 
the  sides. 

Now  the  explanation  of  such  structures  is  the  most  difficult  prob- 
lem heretofore  presented  to  the  geologist.  They  exhibit  conspicuous 
lateral  and  vertical  movements  which  cannot  well  be  accounted  for  by 
the  contraction  theory.  A  shortening  of  about  74  miles  (Heim) 
in  the  folding,  which  has  amounted  to  50  per  cent,  of  the  whole  span 
of  crust  (Leconte),  can  not  be  accounted  for,  on  the  old  theory, 
without  assuming  that  the  crust  is  loose  from  the  globe,  so  that  a 
vast  amount  of  $lack  could  be  brought  forward  and  concentrated  in 
the  folds  at  one  point,  in  the  Swiss  Alps.  This  is  clearly  unthinkable. 
On  the  other  hand,  the  cone  of  matter  underlying  the  Alps  with 
vertex  at  the  center  of  the  earth  could  not  be  sufficiently  condensed 
to  give  the  required  slack  in  the  overlying  crust  without  increas- 
ing the  density  of  the  cone  by  50  per  cent.,  which  could  easily  be 
detected  by  geodetic  observations,  owing  to  the  resulting  deviations 
of  the  plumb  line.  Accordingly  we  may  feel  sure  that  the  matter 
under  the  Alps  not  only  is  not  denser  than  the  average,  but  actually 
lighter,  by  an  appreciable  amount.  The  crumpling  of  the  Alps 
cannot  therefore  be  due  to  condensation  beneath  these  mountains. 

How  then  did  the  folding  arise  ? 

If  we  cut  a  section  across  the  Aleutian  Islands  perpendicular  to 
the  chain  and  the  parallel  trench  lying  to  the  south,  we  shall  have  a 
figure  something  like  that  shown  in  figure  i  of  the  following  plate. 
Now  in  the  paper  on  the  "  Cause  of  Earthquakes  "  (§  16)  we  have 
shown  how  the  undermining  of  the  sea  bottom  sinks  the  trough  down 
deeper  and  deeper,  and  as  the  expulsion  of  lava  continues  it  eventu- 
ally becomes  easier  to  fold  up  the  side  of  the  trough  towards  the 
ocean  (at  &)  and  make  another  range  of  mountains  parallel  to  the 
first.    And  there  is  nothing  to  prevent  the  process  from  being  re- 


i9o8.] 


THE  PHYSICS  OF  THE   EARTH. 


217 


peated  several  times.  When  several  successive  ranges  of  mountains 
are  thus  developed  in  the  process  of  expulsion  under  the  margin  of 
the  sea,  it  is  easy  to  see  that  the  central  range  may  finally  be  driven 
upward  and  flared  out  at  the  top  exactly  as  in  the  Alps.  Thus  all 
this  movement  occurs  in  the  sea,  and  eventually  the  range  becomes 
like  that  now  seen  in  Switzerland,  as  depicted  by  Heim,  of  Ziirich. 


^gtf  level 


\,  Mountain  /^ange  r/s/ng //f tAe Sea 


2.  Doubfe  Pange  fornNhg, 


3.  Trifile  /fdnge  beginning. 


4.  Movement  from  both  s/cfes. 


5.MoYement/rom  both  sides  ccniinueet' 


^  >Ac- 


6.  iVMe  AfouHdin  Range  ri&ing  from  t/re  ^ea. 


tRismgfromi/fe^ed  continuea,gir/ng/an-stf^pea  structures  and  overturned  dips. 

Fig.  II.  IllustrAtion  of  Formation  of  Complex  Range,  such  as  the  Swiss 
Alps.  The  bending  of  the  crust  has  caused  it  to  pull  apart  at  the  top  and 
bottom  of  the  folds,  where  it  is  largely  covered  by  sedimentary  deposits  and 
filled  by  molten  rock  from  beneath,  so  that  the  breaks  do  not  show  at  the 
surface,  unless  erosion  has  laid  bare  parts  of  the  underljring  structure.  In 
these  figures  the  thickness  of  the  crust  is  less  than  half  the  width  of  the 
folds;  and  for  clearness  the  depth  of  the  sea  is  exaggerated. 


218  SEE— FURTHER   RESEARCHES  ON  [April  34. 

As  the  movement  continues  the  central  range  rises  upwards, 
while  its  flanks  sink  down  on  either  side,  and  thus  the  fan-shaped 
structure  develops,  so  as  to  give  overturned  dip  and  inversion  of 
strata  once  deposited  horizontally  in  the  bed  of  the  sea. 

This  is  a  perfectly  simple  and  direct  explanation  of  one  of  the 
most  mysterious  phenomena  heretofore  encountered  by  naturalists. 

The  new  theory  of  mountain  fomiation  is  proved  to  represent  a 
real  law  of  nature  by  phenomena  now  witnessed  in  the  Aleutian 
Islands,  Japan  and  elsewhere.  The  fact  that  it  perfectly  accounts 
for  the  perplexing  phenomena  seen  in  the  Swiss  Alps,  shows  that 
they  too  were  formerly  under  the  sea,  and  were  uplifted  by  the 
same  force  now  at  work  in  the  Aleutian  Islands  and  the  Antandes. 

Accordingly  it  is  not  remarkable  that  Professor  Suess  should, 
without  knowledge  of  the  true  cause,  describe  the  uplift  of  the  Alps 
from  the  sea  in  words  which  are  almost  prophetic  ("Face  of  the 
Earth,"  Vol.  II,  p.  552)  : 

"As  a  result  of  tangential  thrusts,  the  sediments  of  this  Sea  (Mediter- 
ranean) were  folded  together  and  driven  upward  as  a  great  mountain  range, 
and  the  Alps  have  therefore  been  described  as  a  compressed  sea." 

Without  overestimating  the  significance  of  this  result,  it  seems 
clear  that  neither  parallel  ranges  nor  fan-shaped  structures  with 
inverted  dips  will  hereafter  present  any  further  difficulty  to  the 
geologist.  Now  that  the  true  laws  of  such  phenomena  are  known, 
it  will  be  exceedingly  interesting  to  work  out  the  details  of  all  the 
great  mountain  systems  with  which  the  earth  is  adorned. 

§  29.  All  Complex  Folding  now  seen  in  Mountain  kanges 
Originated  in  the  Sea. — It  is  scarcely  necessary  to  add  that  all  the 
complex  folds  now  seen  in  moimtain  ranges  were  produced  in  the 
sea  by  the  repetition  of  trenches  dug  out  by  earthquakes.  The 
folds  were  frequently  broken  apart  at  both  top  and  bottom,  by  the 
earthquake  movements,  and  thus  the  folded  crust  is  not  shortened  by 
anything  like  so  much  as  has  been  supposed.  Moreover  where  the 
fan-shaped  structures  and  overturned  dips  appear,  the  two  sides 
were  never  joined  together  by  an  arch  above,  as  represented  in  the 
above  figures  by  Heim,  but  were  quite  separated  before  the  range 
arose  to  any  considerable  height.  Accordingly  it  follows  that  erosion 
has  not  worn  off  anything  like  so  much  of  the  top  of  the  range  as 


1908.J 


THE  PHYSICS  OF  THE   EARTH. 


219 


the  theory  of  a  rounded  arch  would  require.  Thus  we  may  not 
only  explain  the  folds  of  the  Alps,  but  also  recognize  that  the  folds 
both  above  and  below  were  less  extensive  than  was  formerly  sup- 
posed; and  this  greatly  simplifies  the  labor  of  the  geologist  in  re- 
storing the  former  structure  of  mountain  chains  as  they  appeared 
before  they  were  greatly  eroded. 


Sect'm  ofi North  AmericA,Eau5t  to  We^t  (Dam). 

A 


5ec/M7ff  of^  douthAT7ierici^Ed5t  to  )Ve^t(/>ana). 


Section  c^/fsia,  Aorth  to  South  (Dana), 


QecthnofAftica./^orth  to  douth  (Danay 


Section aP Africa  Sast  to  mfast  (Danm)* 


HmMlAra^ 


T/bPt^tn  UbielMwi 


/^ips 


Fig.  12.    Sections  of  the  Continents,  and  of  the  Alps  and  Himalaya  on  the 

same  Scale.     (Gen.  Strachey.) 

The  process  of  undermining  the  sea  bottom  in  the  expulsion  of 
lava  arising  from  the  leakage  of  the  ocean,  has  given  rise  to  all  the 
important  folds  of  the  earth's  crust.  Thus  arose  all  the  complicated 
folds  in  the  Alps,  Andes,  AUeghenies  and  other  mountain  ranges. 
And  wherever  we  see  these  folds  sea  trenches  once  existed,  and  the 


220  SEE— FURTHER   RESEARCHES  ON  [Apnl  34. 

crust  was  pushed  hither  and  thither  by  earthquakes,  raising  ridges 
and  undermining  the  troughs,  till  the  rocks  were  crumpled  and 
folded  as  we  find  them  to-day.  The  simplicity  of  this  cause,  and  the 
easy  way  in  which  we  pc^s  from  the  living  troughs  now  being  dug 
out  in  the  sea  to  fossil  troughs  long  since  dead  and  now  far  inland 
give  a  genuine  paleontological  interest  to  the  science  of  mountain 
formation.  What  has  long  been  mysterious  and  nearly  inexplicable 
is  now  as  clear  as  any  theorem  in  geometry. 

§  30.  Application  of  the  New  Theory  to  the  Allegheny  Moun- 

« 

tains. — ^The  Allegheny  Mountains  in  Pennsylvania  and  Virginia  are 
very  remarkable  for  the  great  extent  of  the  folds,  and  it  seems 
worth  while  to  dwell  a  moment  on  the  mode  by  which  these  folds 
were  produced.  We  have  seen  that  they  all  arose  in  the  sea,  and  by 
a  repetition  of  the  earthquake  process  of  digging  out  trenches  along 
the  ancient  shore  line.  As  we  shall  see  in  Part  V,  §  41  of  this  paper, 
Professor  James  Hall  so  long  ago  as  1857  annoimced  to  the  Ameri- 
can Association  in  session  in  Montreal  that  the  enormous  thickness 
of  the  formations  along  the  Appalachian  Chain  in  the  United  States 
was  due  to  the  prolonged  accimiulation  of  sediments  over  a  sinking 
sea  bottom,  at  the  margin  of  the  continent,  where  the  marine  cur- 
rents allowed  the  material  to  deposit. 

Obviously  if  sea  trenches  were  dug  out  by  earthquakes  they 
would  become  the  basins  for  the  accumulation  of  a  vast  amount  of 
detritus.  And  when  several  trenches  were  successively  dug  out  in 
the  sea  bottom  by  earthquakes  depending  on  the  Atlantic,  would  not 
the  resulting  folds  give  us  the  Allegheny,  Tuscarora  and  Blue  Ridge 
Mountains  of  Pennsylvania  and  Virginia?  The  famous  Shenandoah 
Valley  in  Virginia  is  nothing  but  an  ancient  sea  trough ;  and  Penn- 
sylvania has  many  such  valleys  originally  formed  in  the  depths  of 
the  sea.  This  is  clearly  indicated  by  the  beautiful  parallelism  of  the 
mountain  ranges. 

It  is  noticeable  that  the  sea  trench  south  of  the  Aleutian  Islands 
is  remarkably  straight,  and  one  may  easily  predict  that  the  ranges 
hereafter  to  be  formed  in  the  North  Pacific  Ocean  will  be  remark- 
ably parallel  like  those  now  seen  in  Virginia  and  Pennsylvania. 
Under  the  circumstances  can  any  one  doubt  that  the  sea  was  once 
very  deep  near  where  the  Blue  Ridge  stands  to-day  ? 


I908.J  THE  PHYSICS  OF  THE  EARTH.  221 

Excluding  from  consideration  the  crystalline  belt  on  the  east, 
Claypole  estimated  the  shortening  of  the  Appalachians  in  Pennsyl- 
vania at  46  miles.  In  the  same  way  McConnell  estimated  that  of 
the  Laramide  range  in  British  America  at  25  miles,  and  Leconte 
that  of  the  Coast  Range  in  California  at  from  9  to  12  miles.  Cor- 
responding estimates  have  been  made  for  many  other  mountain 
ranges;  but,  for  reasons  already  given  in  dealing  with  the  origin 
of  the  Swiss  Alps,  §  28,  these  estimates  are  too  large.  The  crust 
was  broken  apart  at  both  top  and  bottom  when  the  ranges  were  in 
the  sea,  and  the  folds  heretofore  assumed  to  be  complete  were 
never  really  so.  Consequently  no  slack  in  the  earth's  crust  is  re- 
quired to  explain  these  folds ;  it  was  never  loose  from  the  globe  and 
never  moved  horizontally,  except  when  forced  by  earthquake  move- 
ments proceeding  from  the  underlying  trenches  in  the  sea  bottom. 

The  undermining  and  folding  of  the  crust  has  given  the  Ap- 
palachian Mountains  in  many  places  the  aspect  of  a  series  of  immense 
billows,  running  parallel,  as  if  swept  in  by  a  vast  disturbance  of 
the  sea.  But 'not  even  seismic  sea  waves  of  the  most  imposing 
magnitude  could  approach  the  size  of  these  gigantic  folds,  the  origin 
of  which  heretofore  has  been  so  mysterious.  The  finding  of  a  simple 
and  natural  explanation  of  these  great  billows  of  the  land  will  be 
scarcely  less  interesting  than  the  discovery  of  the  cause  of  seismic 
sea  waves.  Both  depend  on  earthquakes,  though  in  very  different 
ways.  The  land  billows  are  cumulative  products  of  an  infinite  series 
of  seismic  disturbances  along  the  margin  of  the  sea;  the  seismic 
waves  are  small  in  comparison,  and  result  from  a  single  disturbance 
of  the  sea  bottom,  made  in  process  of  shaping  the  vast  billows  of  the 
land,  which  in  all  generations  have  appealed  to  the  imagination  of 
the  painter,  poet,  and  student  of  nature. 

§  31.  Analogy  Between  the  Uplift  of  the  Islands  of  Japan  by  the 
Movement  from  the  Tuscarora  Deep  and  of  the  Plateau  of  Tibet 
from  the  Indian  Ocean. — ^The  uplift  of  the  Islands  of  Japan  now 
going  on  by  the  expulsions  of  lava  from  beneath  the  Tuscarora  Deep 
is  proved  by  the  terrible  earthquakes  and  seismic  sea  waves  afflict- 
ing that  region,  as  well  as  by  the  historical  fact  that  the  east  coast 
of  Japan  is  known  to  be  rising  from  the  sea.  Perhaps  in  general 
the  movement  is  slow  and  insensible,  but  occasionally  earthquakes 

PROC  AMER.  PHIL.  SOC.  XLVII.   189  O,  PRINTED  SEPTEMBER  23,  I908 


222  SEE— FURTHER   RESEARCHES  ON  [April  m, 

have  produced  large  disturbances  of  the  level.  The  nature  of  earth- 
quake movement  in  expelling  lava  from  under  the  sea  is  too  well 
known  to  leave  any  doubt  as  to  what  is  going  on  in  Japan.  And 
the  theory  is  confirmed  by  the  fact  that  if  Nipon  and  Yezo  were 
dug  off  and  thrown  into  the  Tuscarora  Deep  they  would  about  fill 
up  that  profound  abyss  and  leave  the  sea  of  average  depth. 

Now  there  is  a  certain  analogy  between  the  uplift  of  these 
Japanese  Islands,  which  are  considerable  areas,  by  the  Pacific, 
and  of  the  Plateau  of  Tibet  by  the  Indian  and  Pacific  oceans  com- 
bined. Undoubtedly  the  valleys  of  the  Indus,  Ganges  and  Brahma- 
putra are  the  relics  of  ancient  sea  troughs  which  largely  produced 
the  Himalayas  and  the  great  plateau  of  Tibet.  How  much  these 
troughs  have  been  modified  in  later  geological  times  we  cannot 
estimate;  but  even  now  enough  remains  to  tell  the  true  story  ol 
Himalayan  development.  This  is  also  indicated  by  the  preservation 
of  the  earthquake  belt  south  of  the  Himalayas.  The  meaning  of 
these  valleys  and  earthquake  belts  admits  of  no  possible  doubt. 
Just  as  the  whole  island  of  Nipon  is  being  raised  by  movements 
from  the  Tuscarora  Deep,  so  the  whole  of  the  Plateau  of  Tibet  was 
once  raised  by  an  Indian  Deep,  of  which  these  valleys  are  the 
remains. 

In  the  same  way  the  Valley  of  the  Po  is  the  remains  of  the  sea 
Valley  which  was  most  influential  in  uplifting  the  Swiss  Alps.  But 
in  the  case  of  the  Alps,  Geikie  has  shown  that  there  was  also  a  sea 
on  the  north,  which  has  now  quite  disappeared,  though  traces  of  its 
former  existence  still  remain. 

§  32.  The  Origin  of  Volcanoes  and  the  Conditions  of  their  Maxi- 
mum Development, — It  appears  from  the  line  of  proof  developed  in 
this  theory  that  volcanoes  may  break  forth  in  any  region  near  the 
sea  where  there  are  severe  earthquake  disturbances,  by  which  the 
crust  of  the  globe  is  sufficiently  cracked  to  afford  a  vent  for  the 
steam  imprisoned  beneath.  Now  such  vents  are  greatly  facilitated  in 
a  chain  such  as  the  Aleutian  Islands,  in  which  the  crumpling  is 
extreme,  and  the  expulsion  of  lava  from  beneath  the  sea  rapid  and 
violent.  The  crumpling  breaks  the  crust  along  many  lines,  and 
as  the  earthquakes  due  to  the  expulsion  of  lava  are  both  frequent 
and  terrible,  the  chance  of  steam  breaking  through  to  the  surface 


i9o8.]  THE  PHYSICS  OF  THE  EARTH.  223 

is  much  greater  than  in  regions  less  wrinkled  and  less  afflicted 
by  earthquakes.  The  crust  in  the  Andes  was  once  folded  by  the 
sea  in  the  same  way  as  that  in  the  Aleutian  islands,  and  from  this 
circumstance  arises  the  violence  of  the  volcanic  outbreaks  noticed 
all  along  the  west  coast  of  South  America.  From  the  great  simi- 
larity of  the  volcanic  phenomena  in  the  Andes  and  in  the  Aleutian 
Islands,  and  its  enormous  prominence  in  both  ranges,  it  seems 
obvious  that  we  have  here  the  conditions  for  its  maximimi  devel- 
opment. 

Charles  Darwin  believed  that  volcanoes  usually  break  out  in 
regions  of  elevation.  No  doubt  this  is  true,  for  mountain  ranges 
are  the  most  conspicuous  of  rising  areas.  And  according  to  this 
theory  the  tendency  to  rupture  the  crust  is  a  maximum,  when  the 
ranges  are  being  both  folded  and  raised  from  the  sea.  Thus  while 
some  volcanoes  may  break  out  in  less  fractured  regions  of  the  earth's 
crust,  the  greatest  volcanic  activity  develops  where  mountains  are 
being  formed  in  the  sea,  as  in  the  Aleutian  Islands.  This  view  also 
enables  us  to  understand  why  many  volcanoes  in  the  Andes  are  now 
extinct,  though  they  were  formerly  active  for  immense  periods  of 
time,  as  we  know  from  the  thick  deposits  of  volcanic  debris  and  the 
immense  height  of  the  cones  built  up  of  lava,  ashes  and  cinders. 

IV.  Comparison  of  the  New  Physical  Theory  of  Mountain 
Formation  Depending  on  the  Leakage  of  the  Oceans 
with  the  Theory  of  Secular  Cooling  and  Con- 
traction Heretofore  Held  by  Men  of  Science. 

§  33.  General  Remarks  on  the  Method  of  Comparison  Adopted. — 
The  new  physical  theory  of  mountain  formation  depending  on 
the  leakage  of  the  oceans  outlined  in  the  three  memoirs  recently 
published  by  the  American  Philosophical  Society  and  somewhat 
more  fully  developed  in  the  present  paper  might  seem  incomplete  if 
we  failed  to  compare  the  new  theory  with  the  theory  of  secular 
cooling  and  contraction  of  the  globe  heretofore  held  by  men  of 
science  generally.  On  several  grounds  an  examination  of  the  older 
theory  can  hardly  fail  to  be  instructive.  And  if  this  comparison  of 
the  older  theory  with  that  now  adopted  shall  be  the  means  of  har- 


224  SEE— FURTHER   RESEARCHES  ON  [April  t4, 

monizing  in  any  considerable  degree  the  divergent  views  heretofore 
prevailing,  and  of  showing  that  there  is  no  important  geological 
phenomenon  which  the  new  theory  does  not  explain  in  a  more  simpie 
and  direct  manner  than  the  old  theory,  such  a  comparison  will  no 
doubt  seem  quite  justifiable.  For  it  is  highly  desirable  to  establish 
the  adequacy  of  the  new  theory  to  explain  the  geological  as  well  as 
the  physical  phenomena  noticed  at  the  surface  of  the  earth. 

In  making  this  comparison  it  is  necessary  to  bear  in  mind  that 
the  geological  data  on  many  points  are  still  very  incomplete,  and 
therefore  we  should  expect  agreement  with  the  body  of  phenomena 
rather  than  with  the  details,  about  which  much  uncertainty  still 
exists.  Owing  to  the  incompleteness  of  our  knowledge  of  the  mode 
of  origin  of  the  great  mountain  chains  of  the  globe,  the  best  plan 
of  procedure  seems  to  be :  First,  to  give  an  exposition  of  the  views 
of  previous  writers  in  regard  to  the  individual  great  mountain  sys- 
tems; second,  to  add  a  resume  of  the  views  of  certain  great  geolo- 
gists on  mountain  formation  in  general.  Obviously  such  conden- 
sation of  the  views  of  others  should  wherever  possible  be  given  in 
their  own  words. 

As  this  subject  is  extensive  and  widely  scattered  in  a  variety 
of  publications,  we  must  content  ourselves  with  selecting  those 
citations  which  seem  of  most  interest,  without  in  any  way  claiming 
to  exhaust  the  subject.  Indeed  it  may  well  be  that  some  discussions 
of  value  will  be  entirely  overlooked,  but,  as  the  theories  have  been 
but  very  little  changed  for  many  years,  it  is  hoped  that  the  following 
citations  will  be  found  adequate  to  give  an  intelligent  grasp  of  the 
views  heretofore  accepted  by  the  leading  authorities.  If  there  be 
those  who  doubt  the  propriety  of  including  lengthy  quotations 
from  well-known  authors,  I  must  plead  in  extension  of  the  course 
here  adopted,  that  this  memoir  is  intended  for  others  besides  geolo- 
gists, and  that  all  who  are  interested  in  the  physics  of  the  earth, 
whether  they  be  mathematicians,  astronomers,  physicists,  seismolo- 
gists, geologists,  or  even  chemists  and  biologists,  are  entitled  to  have 
a  clear  summary  of  the  principal  theories  heretofore  accepted  in  re- 
gard to  the  development  of  our  globe.  In  dealing  with  a  subject  of 
such  universal  interest  to  all  men  of  science,  any  reasonable  conden- 
sation of  the  previous  theories  may  be  considered  admissible,  and  one 


x9o8]  THE  PHYSICS  OF  THE  EARTH.  225 

may  have  no  hesitation  in  invoking  the  aid  of  many  authors.  If  the 
establishment  of  a  great  law  of  nature  may  be  thus  facilitated,  surely 
no  one  will  doubt  that  the  space  utilized  was  devoted  to  a  most  useful 
purpose.  The  extreme  specialization  characteristic  of  the  science 
of  our  day  makes  such  summaries  both  useful  and  necessary  for 
the  intelligent  study  of  great  problems;  and  if  more  effort  were  made 
in  this  direction  it  might  contribute  materially  to  the  progress  of 
scientific  research. 

(A)  Accounts  of  Particular  Mountain  Systems,  and  Their 

Supposed   Mode  of   Development. 

§  34.  The  Andes, — ^We  shall  begin  with  the  Andes  of  South 
America,  because  this  is  one  of  the  largest,  simplest  and  most 
typical  of  mountain  systems;  and  if  a  theory  will  not  explain  the 
Cordilleras,  we  may  despair  of  its  explaining  the  more  compli- 
cated mountains  of  the  globe.  The  reader  should  carefully  bear 
in  mind  not  only  wfiat  the  author  in  question  says  from  his  own 
point  of  view,  but  also  how  the  facts  he  mentions  accord  with 
the  new  theory  developed  in  this  paper. 

In  the  Encyclopedia  Britannica,  ninth  edition,  under  the  article 
"  Andes,"  we  find  the  following  lucid  exposition  of  Andean  de- 
velopment. It  is  not  signed,  but  is  supposed  to  have  passed  under 
the  review  of  Sir  Archibald  Geikie. 

"The  formation  of  the  Andes  is  due  to  several  causes  operating  at  dis- 
tinct intervals  of  time.  They  consist  mainly  of  stratified  material  which  has 
been  more  or  less  altered.  This  material  was  deposited  at  the  bottom  of  a 
sea,  so  that  at  some  former  time  the  highest  portions  were  submerged, 
probably  in  consequence  to  a  certain  extent,  of  subsidence  of  the  sea  bottom. 
Since  the  latest  deposits  there  has  been  upheaval  and  denudation.  The  range, 
then,  has  resulted  from  the  accumulation  of  sediment  on  a  subsiding  area; 
from  the  subsequent  upheaval  of  such  deposits,  which  have  been  increased 
in  height  by  the  ejection  of  volcanic  products;  and  from  the  operation  of 
denuding  agents. 

"  As  far  as  our  present  knowledge  goes,  it  appears  to  be  probable  that 
the  Andes  mark  an  area  on  which  sedimentary  deposits  have  been  accumulated 
to  a  greater  thickness  than  on  any  other  portion  of  South  America.  It  is 
further  demonstrable  that  these  deposits  belong  to  several  geological  periods, 
the  elevation  having  occurred  at  different  periods,  while  their  axes  extend 
in  different  directions.  Hence  it  is  a  complex  range  of  mountains  formed 
by  the  combination  of  several  distinct  systems  of  ridges.    The  width  of  the 


226  SEE— FURTHER   RESEARCHES  ON  (April  24. 

range  varies  from  about  60  to  300  or  more  miles,  but,  as  compared  with 
other  mountains,  the  Andes  are  for  the  most  part  narrow  relatively  to  their 
height.  Where  their  special  features  are  most  characteristically  developed, 
they  consist  of  a  massive  embankment-like  foundation,  rising  with  a  rapid 
slope  from  the  low  country  on  either  side,  and  having  its  margins  sur- 
mounted by  lofty  ridges  of  ragged  or  dome-like  summits.  These  Cor- 
dilleras, as  they  are  usually  termed,  flank  longitudinal  valleys,  or  plain- 
like depressions  which  form  the  highest  levels  of  the  central  portion  of 
the  gigantic  embankment,  and  which  vary  in  width  from  twenty  to  sixty 
miles.  At  intervals  the  longitudinal  depression  is  broken  up,  either  by  ridges 
connecting  the  Cordilleras,  or  by  lofty  plateau-like  uplands.  In  several 
cases  these  transverse  ridges  and  belts  of  high  ground  form  the  main 
watershed  of  the  country.  They  are  rarely  cut  across  by  the  river  systems, 
whereas  both  the  marginal  Cordilleras  are  intersected  at  numerous  points, 
and  more  especially  by  the  rivers  draining  the  eastern  slope  of  the  country. 
In  no  case  do  these  eastern  rivers  originate  to  the  west  of  the  western 
Cordilleras.  A  few  of  the  central  valleys,  or  plain-like  depressions,  have 
no  connection  either  with  the  western  or  eastern  river  system.  Roughly 
speaking  the  height  of  the  central  plains  or  valleys  is  from  6000  to  11,000 
feet  above  the  sea;  of  the  passes  and  knots,  from  10,000  to  15,000  feet; 
and  of  the  highest  peaks,  from  18,000  to  23,290  feet — the  last  being  the 
altitude  of  Aconcagua  in  Chili,  which  is  generally*  considered  to  be  the 
highest  peak  in  America.  Judging  from  these  estimates,  we  may  regard 
the  bulk  of  the  Andes  as  somewhere  about  that  of  a  mass  4400  miles  long, 
100  miles  wide,  and  13,000  feet  high,  which  is  equivalent  to  5,349,801,600,- 
000,000  cubic  feet.  On  this  basis  we  find  that  the  Mississippi  would  carry 
down  an  equivalent  mass  of  matter  in  785,000  years.  The  rate  of  denuda- 
tion in  certain  river  basins  varies  from  one  foot  in  700  years  to  one  foot 
in  12,000  years.  Assuming  that  similar  rates  would  apply  to  the  Andes, 
they  would  be  denuded  away  in  from  9  to  156  million  years.  In  all  proba- 
bility, much  less  than  9  million  would  suffice.  On  the  other  hand  the 
Andes  would  be  swept  away  in  135,000  years,  supposing  the  denuding  powers 
of  the  globe  were  concentrated  on  them  alone.  From  the  above  data,  and 
assuming  the  average  specific  gravity  of  the  matter  forming  the  Andes  to 
be  2.5,  the  weight  of  the  portion  above  the  sea  may  be  estimated  at  368,- 
951,834,482,750  tons,  giving  an  average  of  about  1,000  tons  on  each  square 
foot  at  the  level  of  the  sea.  Under  Aconcagua  the  pressure  would  be  about 
1,780  tons  per  foot  at  the  same  level,  provided,  of  course,  it  were  not,  as 
it  no  doubt  is,  more  or  less  modified  by  lateral  pressure.  These  figures 
afford  some,  though  at  best  a  vague,  conception  of  the  mighty  grandeur  of 
this  range  of  mountains,  and  of  the  scope  there  is  for  the  exertion  of 
enormous  pressure.  How  vast  then,  must  be  those  forces  which  have 
counteracted  such  pressures,  and  upheaved  the  ocean-spread  sediments  of 
the  continents,  until  the  Andes,  that 

'giant  of  the  Western  Star, 
Looks    from   his   throne   of   clouds 
O'er  half  the  world!' 


x9o8]  THE  PHYSICS  OF  THE  EARTH.  227 

But,  however  vast  the  Andes  may  seem  to  us,  it  should  be  remembered  that 
they  form  but  an  insignificant  portion  of  the  globe  itself.  Aconcagua  is 
about  1/2,000  of  the  earth's  diameter,  which  is  relatively  not  more  than  a 
pimple  1/30  of  an  inch  high  on  the  skin  of  a  tall  man."  (Ency.  Brit.,  Vol. 
II,  pp.  15-16.) 

The  account  here  given  of  how  the  Andes  were  formed  seems 
exceedingly  instructive.  In  the  sea  troughs  formerly  existing  be- 
tween the  ocean  and  the  eastern  range,  which  was  the  first  thrown 
up,  we  have  a  complete  explanation  of  the  extraordinary  depth  of 
sedimentation;  for  in  such  trenches  adjacent  to  a  new  range  the 
rate  of  sedimentation  would  be  a  maximum.  The  subsequent  up- 
lifting of  the  western  side  of  the  sea  troughs,  with  the  vast  lateral 
folding  and  compression  necessarily  accompanying  this  movement, 
accounts  for  the  plateaus,  valleys  and  general  structure  of  the 
Andes,  as  well  as  for  the  violent  volcanic  outbreaks,  which  are  said 
to  greatly  predominate  in  the  range  nearest  the  sea,  from  which 
the  expulsion  of  lava  giving  rise  to  this  mighty  Cordillera  pro- 
ceeded. The  vastness  and  height  of  the  Andes  and  the  terrific 
forces  operating  to  erect  this  gigantic  wall  along  the  shore  of  the 
continent  is  a  true  measure  of  the  secular  leakage  of  the  Pacific 
Ocean,  and  of  the  automatic  relief  it  finds  by  folding  the  earth's 
crust  along  the  border,  in  the  countless  successive  expulsions  of 
lava  from  beneath  the  bed  of  the  sea.  It  is  needless  to  point 
out  how  perfectly  the  new  theory  explains  the  persistence  of  the 
earthquake  belt  along  the  western  shore  of  South  America,  and 
of  the  seismic  sea  waves  by  which  that  region  is  so  often  afflicted. 
It  is  obvious  that  the  forces  which  uplifted  the  mountain  also  car- 
ried up  the  plateaus  enclosed  between  the  various  ranges. 

§  35.  The  Himalayas. — ^The  following  luminous  account  of  the 
Himalayas  by  the  late  Lieutenant  General  Sir  Richard  Strachey, 
Encyclopedia  Britannica,  article  "  Himalayas,"  is  of  extreme  in- 
terest. General  Strachey  resided  in  India  for  many  years,  and  made 
a  life  long  study  of  the  Geology  and  Geography  of  Central  Asia. 
He  was  the  principal  authority  of  his  time  on  this  little  explored 
continent  and  died  February  12,  1908,  at  the  age  of  91  years. 

"Scientific  investigation  has  clearly  shown  that,  so  far  as  the  main 
characteristics  of  the  mountains  are  concerned,  the  natural  boundaries  of 
the  Himalayan  system  must  be  carried  much  farther  than  had  at  first  been 


228  SEE— FURTHER   RESEARCHES  ON  (April  04, 

recognized.  Considerable  obscurity  still  involves  the  eastern  portion  of 
these  mountains,  and  there  is  great  want  of  precise  knowledge  as  to  their 
connection  with  the  ranges  of  western  China,  from  which  are  thrown  off 
the  great  rivers  of  China,  Siam,  and  Burmah.  On  the  west,  however,  it 
has  been  completely  established  that  a  continuous  chain  extends  beyond  the 
Indus  along  the  north  of  the  Oxus,  and  ends  in  that  quarter  about  68"  E.  long. 
In  like  manner  it  is  found  that  no  separation  can  be  established,  except  a 
purely  arbitrary  one,  between  the  Himalaya  as  commonly  defined  and  the 
greatly  elevated  and  rugged  table-land  of  Tibet;  nor  between  this  last  and 
the  mountain  ranges  which  form  its  northern  border  along  the  low-lying 
desert  regions  of  central  Asia. 

"It  thus  appears  that  the  Himalaya,  with  its  prolongation  west  of  the 
Indus,  constitutes  in  reality  the  broad  mountainous  slope  which  descends 
from  the  southern  border  of  the  great  Tibetan  table-land  to  the  lower  levels 
of  Hindustan  and  the  plains  of  the  Caspian;  and  that  a  somewhat  shnilar 
mountain  face,  descending  from  the  northern  edge  of  the  tableland,  leads 
to  another  great  plain  on  the  north,  extending  far  to  the  eastward,  to  the 
northern  borders  of  China.  Towards  its  northwest  extremity  this  great 
system  is  connected  with  other  mountains — on  the  south,  with  those  of 
Afghanistan,  of  which  the  Hindu-Kush  is  the  crest,  occupying  a  breadth 
of  about  250  miles  between  Peshawur  and  Kunduz;  and  on  the  north,  with 
the  mountains  that  flank  the  Jaxartes  or  Sir  on  the  north,  and  the  Thian- 
shan  or  Celestial  Mountains.  The  eastern  margin  of  Tibet  descends  to 
western  China,  and  the  south-eastern  termination  of  the  Himalaya  is  fused 
into  the  ranges  which  run  north  and  south  between  the  95th  and  looth 
meridians,  and  separate  the  rivers  of  Burmah,  Siam,  and  western  China. 

"  Nor  can  any  of  the  numerous  mountain  ranges  which  constitute  this 
great  elevated  region  be  properly  regarded  as  having  special,  definite,  or 
separate  existence  apart  from  the  general  mass  of  which  they  are  the  com- 
ponent parts;  and  Tibet  cannot  be  rightly  described,  as  it  has  been,  as 
lying  in  the  interval  between  the  two  so-called  chains  of  the  Himalaya  and 
the  Kouenlun  or  Kara  Koram.  It  is  in  truth  the  summit  of  a  great  pro- 
tuberance above  the  general  level  of  the  earth's  surface,  of  which  these 
alleged  chains  are  nothing  more  than  the  south  and  north  borders,  while 
the  other  ranges  which  traverse  it  are  but  corrugations  of  the  mass  more 
or  less  strongly  marked  and  locally  developed. 

"The  average  level  of  the  Tibetan  tableland  may  be  taken  at  about 
15,000  feet  above  the  sea.  The  loftiest  points  known  on  the  earth's  sur- 
face are  to  be  found  along  its  southern  or  Himalayan  boundary ;  one  of  them 
falls  very  little  short  of  30,000  feet  in  elevation,  and  peaks  of  20,000  feet 
bound  the  entire  chain.  The  plains  of  India  which  skirt  the  Himalayan 
face  of  the  tableland,  for  a  length  of  rather  more  than  1,500  miles,  along 
the  northern  border  of  British  India,  nowhere  rise  so  much  as  1,000  feet 
above  the  sea,  the  average  being  much  less.  The  low  lands  on  the  north, 
about  Kashgar  and  Yarkend,  have  an  elevation  of  from  3,000  to  4,000  feet, 
and  no  part  of  the  Central  Asiatic  desert  seems  to  fall  below  2,000  feet, 
the  lake  of  Lob-nor  being  somewhat  above  the  level.    The  greatest  dimen- 


x9o8.] 


THE  PHYSICS  OF  THE  EARTH.  229 


sion  of  the  Tibetan  mountain  area  from  east  to  west  may  be  about  2,000 
miles,  while  its  average  breadth  somewhat  exceeds  500  miles;  about  100  miles 
on  either  side  constitute  the  sloping  faces,  the  central  tableland  having  a 
width  of  about  200  miles  on  the  west  and  probably  500  miles  at  its  eastern 
border." 

General  Strachey  thus  shows  that  the  Himalayan  mountains  and 

Tibetan  Plateaus  are  directly  and  intimately  connected  as  merely 

different  parts  of  one  great  continuous  movement  of  the  earth's 

crust. 

After    describing    many    features    of    the    Himalayas,    General 

Strachey  continues: 

"  The  general  conclusion  that  may  be  drawn  from  the  facts  of  structure 
thus  briefly  indicated  is  that  the  elevation  of  the  Himalaya  to  its  present 
great  height  is  of  comparatively  recent  occurrence.  An  area  of  land  must 
have  existed  where  the  main  line  of  snowy  peaks  now  stands,  which  has 
not  been  submerged  since  the  Palaeozoic  period,  and  which  then  had  its 
northern  boundary  somewhere  along  what  has  been  termed  the  Indian  water- 
shed. Evidence  of  a  similar  ancient  sea  on  the  south  also  exists,  but  in 
less  definite  shape;  and  whether  it  was  united  with  the  northern  sea  or  not 
is  still  a  matter  of  conjecture,  though  the  distinctive  character  of  the  fossils 
rather  indicates  that  there  was  no  direct  union.  The  possible  connection  of 
this  ancient  Himalayan  land  area  with  the  pre-Tertiary  land  of  the  peninsula 
of  India  is  also  only  a  matter  for  speculation. 

"  There  is  further  reason  to  infer  that  the  existence  of  the  great  line 
of  peaks  is  rather  due  to  some  previous  line  of  elevation  on  the  ancient 
land,  which  has  continued  to  retain  its  relative  superiority  while  the  whole 
areas  have  been  raised,  rather  than  to  any  special  line  of  energy  of  upheaval 
of  recent  date;  and  that  the  fundamental  features  of  its  former  configura- 
tion of  surface  in  mountain  and  valley  have  been  preserved  throughout. 
There  is  evidence  for  the  conclusion  that  the  chief  rivers  of  the  pre- 
Tertiary  land  issued  from  the  mountains  where  the  present  main  streams  are 
found,  and  this  embryo  Himalaya  may  have  been  of  such  moderate  height 
as  to  have  permitted  the  passage  across  it  of  the  Siwalik  mammals,  the  re- 
mains of  which  appear  both  on  the  border  of  the  Indian  plain  and  in  Tibet. 
It  is  after  the  middle  Tertiary  epoch  that  the  principal  elevation  of  these 
mountains  must  have  taken  place,  and  about  the  same  time  also  took  place 
the  movements  which  raised  the  tablelands  of  Afghanistan  and  Persia,  and 
gave  southern  Asia  its  existing  outlines. 

"The  best  answer  that  can  be  given  to  an  inquiry  as  to  how  changes 
of  level  could  have  arisen,  such  as  those  which  are  observed  in  the  Hima- 
laya, is  that  they  should  be  regarded  as  due  rather  to  secondary  actions 
consequent  on  the  general  contraction  of  the  cooling  terrestrial  sphere  than 
to  direct  elevating  forces,  for  which  no  known  origin  can  be  assigned.  The 
contraction  of  the  cooling  but  now  solid  crust  of  the  earth  must  have  set 
up   great   horizontal    strains,   partly   of    tension   and   partly   of   compression 


230  SEE— FURTHER   RESEARCHES  ON  [April  94. 

which  would  necessarily  have  been  followed  by  rupture  or  crushing  along 
lines  of  least  resistance,  and  the  movements  on  such  lines  are  marked  by 
the  great  mountain  ranges  that  traverse  the  surface.  A  dislocation  of  the 
solid  crust  of  the  earth  once  having  taken  place,  it  would  probably  continue 
to  be  a  line  of  least  resistance  ever  after,  and  a  succession  of  movements 
during  past  geological  periods  may  thus  be  reasonably  expected  along  such 
lines.  Somewhat  in  proportion  as  the  disturbing  forces  are  intense,  and  the 
thickness  of  the  crust  on  which  they  act  is  great,  will  be  the  tendency 
of  the  lines  of  rupture  to  be  continuous  for  a  considerable  distance; 
and  as  the  disturbed  area  is  extended  in  its  dimensions,  the  probability  will 
increase  of  a  repetition  of  a  series  of  similar  dislocations  on  lines  approxi- 
mately parallel  to,  or  at  right  angles  to,  one  another  and  to  the  line  on 
which  the  greatest  compression  and  consequent  tension  take  place.  In  a 
disturbed  area,  one  transverse  dimension  of  which  is  sensibly  greater  than 
the  rest,  the  longitudinal  ruptures  will  predominate  in  the  interior  and  the 
transverse  towards  the  borders.  Almost  all  mountains  give  indications  of 
having  been  shaped  by  forces  thus  related,  and  to  the  action  of  such  forces 
may  the  main  characteristics  of  the  structure  of  the  Himalaya,  and  the 
arrangement  of  its  ridges  and  valleys  be  attributed.  Whatever  may  be  the 
power  of  rivers  in  general  as  instruments  of  erosion,  and  whatever  effect 
the  Himalayan  rivers  have  had  in  removing  the  fragments  of  the  rocks  over 
and  among  which  they  took  their  courses,  it  is  hardly  possible  to  doubt 
that  their  main  directions  were  determined  by  the  anterior  lines  of  dis- 
location which  opened  up  hollows  down  which  they  could  flow,  and  which 
must  invariably  have  been  accompanied  by  a  destructive  and  crushing  action 
on  the  rocks  along  them,  which  has  enabled  the  waters  the  more  readily  to 
sweep  away  the  obstacles  in  their  path.  The  parallelism  of  many  of  the 
great  Tibetan  and  Himalayan  rivers  for  hundreds  of  miles  together,  and 
such  mountains,  seems  wholly  inexplicable  in  any  other  manner.*'  (Ency. 
Brit.,  p.  828.) 

This  account  is  quite  clear  and  satisfactory,  except  that  part  of 
it  which  deals  with  the  cooling  and  contraction  of  the  globe.  Here 
General  Strachey  has  made  the  best  of  a  very  inadequate 
hypothesis. 

Just  as  the  Andes  were  formed  by  expulsions  of  lava  from  under 
the  Pacific,  so  also  here  the  Himalayas  were  formed  by  a  corre- 
sponding movement  due  mainly  to  the  Indian  Ocean,  which  has  also 
raised  high  mountains  along  the  eastern  border  of  Africa.  We  can- 
not yet  give  all  the  details  of  the  Himalayan  development,  but  in 
general  it  is  evident  that  it  was  similar  to  that  of  the  Andes.  The 
uplift  of  the  great  plateau  of  Tibet  corresponds  to  that  of  Titicaca. 
And  the  parallel  ranges  of  the  Himalayas  originated  by  the  usual 
process   of  the   folding  up  of   successive   sea  trenches.     On   the 


x9o8.]  THE  PHYSICS  OF  THE   EARTH.  231 

outside  of  these  mountains  there  still  remain  trough-like  depressions 
where  the  Indus,  Ganges  and  Brahmaputra  now  flow.  The  under- 
mining produced  in  raising  the  Himalayan  embankment  still  shows 
in  the  valleys  to  the  south,  though  the  sea  has  receded;  and  the 
great  earthquake  belt  south  of  the  Himalayas  still  discloses  to  us  the 
nature  of  the  forces  which  produced  this  mighty  uplift. 

The  following  critical  passages  by  General  Strachey  are  also  of 
decided  interest : 

"The  great  peaks  are,  with  few  exceptions,  composed  of  schistose  rock, 
though  granite  veins  may  be  seen  in  the  mountain  faces  to  very  great  ele- 
vations; one  of  these  exceptions  is  the  great  peak  of  Kamet  in  Kumaon, 
which  rises  to  about  25,000  feet  in  what  appears  to  be  a  mass  of  grey  granite. 

"  Passing  to  the  north  of  the  line  of  great  peaks  the  metamorphosed 
schists  are  suddenly  replaced  by  slates  and  limestones,  which  are  in  many 
places  highly  fossiliferous,  exhibiting  what  appears  to  constitute  in  the 
aggregate  a  fairly  continuous  series  from  the  Lower  Silurian  to  the  Cre- 
taceous formations,  though  the  complete  sequence  has  not  been  observed  in 
any  one  locality.  The  western  region  of  the  Himalaya  alone  has  been  suf- 
ficiently explored  to  admit  of  any  positive  statements,  but  the  indications 
gathered  from  such  imperfect  accounts  and  other  data  as  exist  relative  to 
the  eastern  parts  of  the  mountains  leave  little  doubt  that  the  change  ob- 
served in  the  west  on  approaching  and  entering  Tibet  holds  good  on  the 
east  also,  and  that  the  general  physical  features  of  the  whole  tract  are  much 
alike,  though  doubtless  with  many  differences  in  detail. 

"The  fossiliferous  strata  of  western  Tibet  are  continued,  though  per- 
haps with  some  breaks,  to  the  Tertiary  period.  In  certain  localities  num- 
mulitic  rocks,  probably  Eocene,  have  been  observed,  and  from  the  great 
alluvial  deposit  which  forms  the  plain  of  Guge,  already  noticed,  the  remains 
of  mammals,  apparently  of  Siwalik  age,  have  also  been  obtained.  Among  these 
were  bones  of  the  elephant  and  rhinoceros,  the  existence  of  which,  in  the 
present  condition  of  these  regions,  would  be  wholly  impossible;  so  that 
there  is  no  room  to  doubt  that  these  deposits  have  been  raised  from  a  com- 
paratively low  level  to  their  existing  great  elevation  of  upwards  of  15,000 
feet,  since  they  were  laid  out.  As  in  the  case  of  the  plain  of  India,  we 
here,  too,  have  no  complete  proof  of  the  origin  of  these  great  nearly  hori- 
zontal deposits,  but  it  seems  clear,  from  the  materials  of  which  they  are 
formed,  that  they  must  have  been  laid  out  by  the  water,  either  by  the  sea 
or  some  great  inland  lake.  They  are  largely  composed  of  boulder  deposits, 
and  large  boulders  are  strewed  over  the  surface  imbedded  in  the  ground  in 
a  manner  that  seems  only  explicable  as  the  result  of  the  action  of  a  con- 
siderable body  of  water. 

"  Several  lines  of  granitic  and  eruptive  rock  occur  in  western  Tibet,  of 
which  all  that  need  here  be  said  is  that  they  appear  all  to  be  older  than 
the  Tertiary  alluvium,  but  some  of  them  are  possibly  contemporaneous  with 
the  nummulitic  and  older  formations."     (Ency.  Brit.  p.  828.) 


232  SEE— FURTHER  RESEARCHES  ON  (April  24. 

In  an  earlier  passage,  after  comparing  some  of  the  smaller 
Himalayan  ranges  to  the  Swiss  Alps,  General  Strachey  adds : 

"To  obliterate  these  two  ranges  from  the  Himalaya  would  make  no 
very  sensible  inroad  on  it,  though  they  surpass  in  bulk  the  whole  of  the 
Swiss  Alps;  and  it  is  no  exaggeration  to  say  that,  along  the  entire  range 
of  the  Himalaya,  valleys  are  to  be  found  among  the  higher  mountains  into 
which  the  whole  Alps  might  be  cast  without  producing  any  result  that  would 
be  discernible  at  a  distance  of  ten  or  fifteen  miles.  And  it  is  important  to 
bear  in  mind  these  relations  of  magnitude,  for  the  terms  at  our  disposal  in 
the  description  of  the  mountains  are  so  limited  that  it  is  necessary  to  employ 
the  words  chain,  range,  ridge,  spur,  etc.,  rather  with  reference  to  relative 
than  to  absolute  importance,  so  that  the  scale  of  our  nomenclature  changes 
with  the  extent  and  altitude  of  the  mountains  of  which  we  speak."  (Ency. 
Brit.,  p.  827.) 

§  36.  The  Alps, — In  the  Enclycopedia  Britannica,  article  "  Alps," 
by  John  Ball,  we  find  the  following  brief  outline  of  the  salient 
features : 

"  Accurate  knowledge  of  the  Alps  is  so  recent  that  few  attempts  have 
been  made  to  establish  a  general  division  of  the  entire  region,  and  it  can- 
not be  said  that  any  one  arrangement  has  obtained  such  general  recognition 
as  not  to  be  open  to  future  modification;  but  there  is  a  pretty  general 
agreement  as  to  the  main  features  of  that  here  proposed,  to  which  a  few 
general  remarks  must  be  premised. 

"  Whatever  may  have  been  the  original  cause  of  the  disturbances  of  the 
earth's  crust  to  which  great  mountain  chains  owe  their  existence,  it  is  gen- 
erally, though  not  universally,  true  that  the  higher  masses  (formed  of  crys- 
talline rock  and  geologically  more  ancient)  are  found  towards  the  central 
part,  and  that  these  are  flanked  by  lower  ranges,  composed  of  more  recent 
rocks,  which  surround  the  central  groups  very  much  as  an  outer  line  of 
entrenchment  may  be  seen  to  surround  a  fort.  In  most  cases  it  is  not 
possible  to  descend  continuously  in  a  nearly  direct  line  from  the  crest  of  a 
great  mountain  chain  to  the  plains  on  either  side,  for  there  are  usually 
intermediate  valleys,  running  more  or  less  parallel  to  the  central  range,  which 
separate  this  from  outer  secondary  ranges.  These  in  turn,  are  often  ac- 
companied by  external  ranges,  intermediate  between  them  and  the  plains, 
and  related  to  them  as  they  are  to  the  central  ranges.  The  type  of  arrange- 
ment here  described  is  more  or  less  traceable  throughout  the  greater  part 
of  the  Alps,  but  is  most  distinctly  exhibited  in  the  eastern  portion  lying 
between  the  Adige  and  the  frontier  of  Hungary.  We  have  a  central  range, 
composed  mainly  of  crystaline  rock;  a  northern  range,  formed  of  secondary 
rocks,  separated  from  the  first  by  the  great  valleys  of  the  Inn,  the  Salza. 
and  the  Enns;  a  southern  range,  somewhat  similar  to  the  last  in  geological 
structure,  divided  from  the  central  one  by  the  Rienx,  or  east  branch  of  the 
Adige,  and  the  Drave.  Flanking  the  whole,  as  an  external  entrenchment  on 
the  north  side,  are  the  outer  ranges  of  the  Bavarian  Alps,  of  the  Salzkam- 


,9o8.]  THE  PHYSICS  OF  THE  EARTH.  233 

mergut,  and  of  Upper  Austria,  to  which  corresponds  on  the  south  side  the 
Monti  Lessini,  near  Verona,  the  mountains  of  Recoaro,  those  of  the  Sette 
Comuni,  and  the  considerable  masses  crowned  by  the  summits  of  the  Grappa, 
the  Col.  Vicentino,  the  Monte  Cavallo,  the  Monte  Matajur,  and  Monte 
Nanos.  Where,  as  in  the  case  above  mentioned,  the  secondary  ranges  of 
the  Alps  rise  to  a  greater  altitude,  and  are  completely  separated  from  the 
neighbouring  portions  of  the  central  chain,  it  is  impossible  not  to  distinguish 
them  as  distinct  groups;  but  the  outermost  ranges,  which  rarely  rise  above 
the  forest  zone,  are  in  all  cases  regarded  as  appendages  of  the  adjoining 
groups.  These  outer  ranges  are  called  in  German  Voralpen,  and  in  Italian 
Prealpi."     (Ency.  Brit.,  p.  623.) 

Again  on  page  620,  this  author  remarks : 

VIn  every  mountain  system  geographers  are  disposed  to  regard  the 
watershed,  or  boundary  dividing  the  waters  flowing  towards  the  opposite 
sides  of  the  range,  as  marking  the  main  chain;  and  this  usage  is  often 
justified  by  the  fact  that  the  highest  peaks  lie  on,  or  very  near,  the  boundary 
so  defined.  In  applying  this  term  in  the  case  of  the  Alps,  there  are,  however, 
difficulties  arising  from  their  great  extent  and  the  number  of  their  branches 
and  ramifications.  Many  of  the  loftiest  groups  lie  altogether  on  one  side  of 
that  which  we  call  the  main  chain,  and  at  the  eastern  extremity,  where  all 
drainage  is  ultimately  borne  to  the  Black  Sea,  we  must  be  partly  guided  by 
geological  considerations  in  deciding  which  of  several  ranges  deserves  to 
be  considered  pre-eminent."     (Vol.  I.,  p.  620.) 

Sir  Archibald  Geikie's  discussion  of  the  origin  of  the  Alps,  in 
the  article  "  Geology,"  Encyclopedia  Britannica  (pp.  373-374),  bears 
on  the  problem  now  before  us : 

"The  Alps,  on  the  contrary,  present  an  instructive  example  of  the  kind 
of  scenery  that  arises  where  a  mass  of  high  ground  has  resulted  from  the 
intense  corrugation  and  upheaval  of  a  complicated  series  of  stratified  and 
crystalline  rocks,  subsequently  for  a  vast  period  carved  by  rain,  frost,  springs 
and  glaciers.  We  see  how,  on  the  outer  flanks  of  those  mountains  among 
the  ridges  of  the  Jura,  the  strata  begin  to  undulate  in  long  wave-like  ridges, 
and  how,  as  we  enter  the  main  chain,  the- undulations  assume  a  more  gigantic 
tumultuous  character,  until,  along  the  central  heights,  the  mountains  lift 
themselves  towards  the  sky  like  the  storm-swept  crests  of  vast  earth  billows. 
The  whole  aspect  of  the  ground  suggests  intense  commotion.  Where  the 
strata  appear  along  the  cliffs  or  slopes  they  may  often  be  seen  twisted  and 
crumpled  on  the  most  gigantic  scale.  Out  of  this  complicated  mass  of 
material  the  sub-aerial  forces  have  been  ceaselessly  at  work  since  its  first 
elevation.  They  have  cut  valleys,  sometimes  along  the  original  depressions, 
sometimes  down  the  slopes.  They  have  eroded  lake-basins,  dug  out  corries 
or  cirques,  notched  and  furrowed  the  ridges,  splintered  the  crests,  and  have 
left  no  part  of  the  original  surface  unmodified.  But  they  have  not  effaced 
all  traces  of  the  convulsions  by  which  the  Alps  were  upheaved." 


234  SEE— FURTHER   RESEARCHES  ON  lAprila4, 

In  his  account  of  the  Miocene  ("  Text-book  of  Geology,"  p. 
1 261,  edition  of  1903),  Geikie  says: 

"The  Gulf  of  Gascony  then  swept  inland  over  the  wide  plains  of  the 
Garonne,  perhaps  even  connecting  the  Atlantic  with  the  Mediterranean  by  a 
strait  running  along  the  northern  flank  of  the  Pyrenees.  The  sea  washed 
the  northern  base  of  the  now  uplifted  Alps,  sending,  as  in  Oligocene  time, 
a  long  arm  into  the  valley  of  the  Rhine  as  far  as  the  site  of  Mainz,  which 
then  properly  stood  at  the  upper  end,  the  valley  draining  southward  instead 
of  northward.  The  gradual  conversion  of  salt  into  brackish  and  fresh  water 
at  the  head  of  this  inlet  took  place  in  Miocene  time.  From  the  Miocene 
firth  to  the  Rhine,  a  sea-strait  ran  eastwards,  between  the  base  of  the  Alps 
and  the  line  of  the  Danube,  filling  up  the  broad  basin  of  Vienna,  sending 
thence  an  arm  northwards  through  Moravia,  and  spreading  far  and  wide 
among  the  islands  of  southeastern  Europe,  over  the  regions  where  now  the 
Black  Sea  and  Caspian  basins  remain  as  the  last  relics  of  this  Tertiary 
extension  of  the  ocean  across  southern  Europe.  The  Mediterranean  also 
still  presented  a  far  larger  area  than  it  now  possesses,  for  it  covered  much 
of  the  present  lowlands  and  foot-hills  along  its  northern  border,  and  some 
of  its  important  islands  had  not  yet  appeared  or  had  not  acquired  their 
present  dimensions." 

On  pages  137 1-2  of  Geikie's  **  Geology,"  we  find  the  following 
interesting  passages : 

"Alpine  Type  of  Mountain  Structure. — It  is  along  a  great  mountain 
chain  like  the  Alps  that  the  most  colossal  crumplings  of  the  terrestrial  crust 
are  to  be  seen.  In  approaching  such  a  chain,  one  or  more  minor  ridges  may 
be  observed  running  on  the  whole  parallel  with  it,  as  the  heights  of  the 
Jura  flank  the  north  side  of  the  Alps,  and  the  sub-Himalayan  hills  follow 
the  southern  base  of  the  Himalayas.  On  the  outer  side  of  these  ridges,  the 
strata  may  be  flat  or  gently  inclined.  At  first  they  undulate  in  broad  gentle 
folds;  but  traced  towards  the  mountains  these  folds  become  sharper  and 
closer,  their  shorter  sides  fronting  the  plains,  their  longer  slopes  dipping  in 
the  opposite  direction.  This  inward  dip  is  often  traceable  along  the  flanks 
of  the  main  chain  of  mountains,  younger  rocks  seeming  to  underlie  others 
of  much  older  date.  Along  the  north  front  of  the  Alps,  for  instance,  the 
red  molasse  is  overlain  by  Eocene  and  older  formations.  The  inversions 
and  disruptions  increase  in  magnitude  till  they  reach  such  colossal  dimen- 
sions as  those  of  the  Glarnisch,  where  pre-Cambrian  schists,  and  Triassic, 
Jurassic,  and  Cretaceous  rocks  have  been  driven  for  miles  over  the  Eocene 
and  Oligocene  flysch  (pp.  677,  693).  In  such  vast  crumplings  and  thrusts 
it  may  happen  that  portions  of  older  strata  are  caught  in  the  folds  of  later 
formations,  and  some  care  may  be  required  to  discriminate  the  enclosure 
from  the  rocks  of  which  it  appears  to  form  an  integral  and  original  part. 
Some  of  the  recorded  examples  of  fossils  of  an  older  zone  occurring  by 
themselves  in  a  much  younger  group  of  plicated  rocks  may  be  thus  ac- 
counted for. 


)9oe.]  THE   PHYSICS  OF  THE   EARTH.  235 

"  The  inward  dip  and  consequent  inversion  traceable  lowards  the  center 
of  a  mountain  chain  lead  up  to  the  fan-shaped  structure  (p.  678)  where 
the  oldest  rocks  of  a  series  occupy  the  center  and  overlie  younger  masses, 
which  plunge  steeply  under  ihem.  Classical  examples  of  this  structure  occur 
in  the  Alps  (Mont  Blanc,  Fig.  358,  St.  Gothard),  where  crystalline  rocks 
such  as  granite,  gneiss,  and  schists,  the  oldest  masses  of  the  chain,  have 
been  ridged  up  into  the  central  and  highest  peaks.  Along  these  tracts, 
denudation  has  been  of  course  enormous,  for  the  appearance  of  the  granitic 
rocks  at  the  surface  has  been  brought  out,  not  necessarily  by  actual  extru- 
sion into  the  air.  but  more  probably  by  prolonged  erosion,  which  in  these 
higher  regions,  where  many  forms  of  sub-aerial  waste  reach  their  most 
vigorous  phase,  has  removed  the  vast  overreaching  cover  of  younger  rocks 
under  which  the  crystalline  nucleus  doubtless  lay  buried." 


Fig.  13,    Fan-shaped  Structure,  Central  Alps. 

Again  on  page  1372,  we  read: 

"  A  mountain  chain  may  be  the  result  of  one  movement,  but  probably 
in  most  cases  is  due  to  a  long  succession  of  such  movements.  Formed  on 
a  line  of  weakness  in  the  crust,  it  has  again  and  again  given  relief  from  the 
strain  of  compression  by  undergoing  fresh  crumpling  and  upheaval.  Suc- 
cessive stages  of  uplift  are  usually  not  difficult  to  trace.  The  chief  guide 
is  supplied  by  unconformability.  .  .  . 

"  In  most  great  mountain  chains,  however,  the  rocks  have  been  so 
intensely  crumpled,  dislocated,  and  inverted,  that  much  labor  may  be  re- 
quired before  their  true  relations  can  be  determined. 

"  The  Alps  offer  an  instructive  example  of  a  great  mountain  system 
formed  by  repeated  movements  during  a  long  succession  of  geological 
periods.  The  central  portions  of  the  chain  consist  of  gneiss,  schists,  granite, 
and  other  crystalline  rocks,  partly  referable  to  the  pre-Cambrian  series,  but 
some  of  which  (Schistes  lustres,  Biindnerschiefer)  include  metamorphosed 
Palxozoic,  Secondary,  and  in  some  places,  perhaps,  even  older  Tertiary  de- 
posits (pp.  802,  1099).  It  would  appear  that  the  first  outlines  of  the  Alps 
were  traced  out  even  in  pre-Cambrian  times,  and  that  after  submergence, 
and  the  deposit  of  Palatozoic  formations  along  their  flanks,  if  not  over 
most  of  their  site,  they  were  reelevated  into  land.    From  the  relations  of  the 


236  SEE— FURTHER  RESEARCHES  ON  fAprilM. 

Mesozoic  rocks  to  each  other,  we  may  infer  that  several  renewed  uplifts, 
after  successive  denudations,  took  place  before  the  beginning  of.  Tertiary 
times,  but  without  any  general  and  extensive  plication.  A  large  part  of  the 
range  was  certainly  submerged  during  the  Eocene  period  under  the  waters 
of  the  wide  sea  which  spread  across  the  center  of  the  Old  World,  and  in 
which  the  nummulitic  limestone  and  flysch  were  deposited.  But  after  that 
period  the  grand  upheaval  took  place  to  which  the  present  magnitude  of  the 
mountains  is  chiefly  due.  The  older  Tertiary  rocks,  previously  horizontal 
under  the  sea,  were  raised  up  into  mountain-ridges  more  than  11,000  feet 
above  the  sea-level,  and  together  with  the  older  formations  of  the  chain, 
underwent  colossal  plication  and  displacement  Enormous  slices  of  the 
oldest  rocks  were  torn  away  from  the  foundations  of  the  chain  and  driven 
horizontally  for  miles  until  they  came  to  rest  upon  some  of  the  newest 
formations.  The  thick  Mesozoic  groups  were  folded  over  each  other  like 
piles  of  carpets,  and  involved  in  the  lateral  thrusts  so  as  now  to  be  seen 
resting  upon  the  Tertiary  flysch.  So  intense  was  the  compression  and  shear- 
ing to  which  the  rocks  were  subjected  that  lenticles  of  the  Carboniferous 
series  have  been  folded  in  among  Jurassic  strata,  and  the  whole  have  been 
so  welded  together  that  they  can  hardly  be  distinguished  where  they  meet, 
and  what  were  originally  clays  and  sands  have  been  converted  into  hard 
crystalline  rocks.  It  is  strange  to  reflect  that  the  enduring  materials  out 
of  which  so  many  mountains,  cliffs,  and  pinnacles  of  the  Alps  have  been 
formed  are  of  no  higher  geological  antiquity  than  the  London  Qay  and 
other  soft  Eocene  deposits  of  the  south  of  England  and  the  north  of  France 
and  Belgiupi.  At  a  later  stage  of  Tertiary  time,  renewed  disturbance  led 
to  the  destruction  of  the  lakes  in  which  the  molasse  had  accumulated,  and 
their  thick  sediments  were  thrust  up  into  large  broken  mountain  masses, 
such  as  the  Rigi,  Rossberg,  and  other  prominent  heights  along  the  northern 
flanks  of  the  Alps.  Since  that  last  post-Eocene  movement,  no  great  orogenic 
paroxysm  seems  to  have  affected  the  Alpine  region.  But  the  chain  has  been 
left  in  a  state  of  unstable  equilibrium.  From  time  to  time  normal  faults 
have  taken  place  whereby  portions  of  the  uplifted  rocks  have  sunk  down  for 
hundreds  of  feet,  and  some  of  these  dislocations  have  cut  across  the  much 
older  and  more  gigantic  displacements  of  the  thrust-planes  (Fig.  282).  At 
the  same  time  continuous  denudation  has  greatly  transformed  the  surfaces 
of  the  ground,  so  that  now  cakes  of  gneiss  are  left  as  mountainous  outliers 
upon  a  crushed  and  convoluted  platform  of  Tertiary  strata.  Nor,  in  spite 
of  the  settling  down  of  these  broken  masses,  has  final  stability  been  attained. 
The  frequent  earthquakes  of  the  Alpine  region  bear  witness  to  the  strain 
of  the  rocks  underneath,  and  the  relief  from  it  obtained  by  occasional  rents 
propagated  through  the  crust  along  the  length  of  the  chain." 

In  view  of  the  explanation  of  the  folding  of  the  Alps  given 
in  §  28,  we  need  not  comment  on  these  views.  They  confirm  the 
theory  outlined  in  this  paper,  that  the  plications  of  all  such  chains 
must  be  sought  in  the  actions  of  the  sea,  and  mainly  while  the 


1908.J 


THE  PHYSICS  OF  THE  EARTH.  237 


range  is  under  water,  and  not  at  all  in  the  secular  cooling  of  the 
globe. 

V.  Comparison  of  the  Old  and  New  Theory  of  Mountain 

Formation  Continued. 

(B)  Views  of  Eminent  Geologists  on  Mountain  Formation 

IN  General. 

§  37.  Elie  de  Beaumont's  Theory  of  the  Secular  Cooling  and 
Collapse  of  the  Globe, — ^This  venerable  theory  is  thus  condensed 
by  Lyell: 

"The  origin  of  these  chains  depends  not  on  partial  volcanic  action  or  a 
reiteration  of  ordinary  earthquakes,  but  on  the  secular  refrigeration  of  the 
entire  planet.  For  the  whole  globe,  with  the  exception  of  a  thin  envelope, 
much  thinner  in  proportion  than  the  shell  to  an  egg,  is  a  fused  mass,  kept 
fluid  by  heat,  but  constantly  cooling  and  contracting  in  dimensions.  The 
external  crust  does  not  gradually  collapse  and  accommodate  itself  century 
after  century  to  the  shrunken  nucleus,  subsiding  as  often  as  there  is  a 
slight  failure  of  support,  but  it  is  sustained  throughout  whole  geological 
periods,  so  as  to  become  partially  separated  from  the  nucleus  until  at  last 
it  gives  way  suddenly,  cracking  and  falling  in  along  determinate  lines  of 
fracture.  During  such  a  crisis  the  rocks  are  subjected  to  great  lateral  pres- 
sure, the  unyielding  ones  are  crushed,  and  the  pliant  strata  bent,  and  are 
forced  to  pack  themselves  more  closely  into  a  smaller  space,  having  no 
longer  the  same  room  to  spread  themselves  out  horizontally.  At  the  same 
time,  a  large  portion  of  the  mass  is  squeezed  upwards,  because  it  is  in  the 
upward  direction  only  that  the  excess  in  size  of  the  envelope,  as  compared 
to  the  nucleus  can  find  relief.  This  excess  produces  one  or  more  of  those 
folds  or  wrinkles  in  the  earth's  crust  which  we  call  mountain-chains." 

De  Beaumont's  theory  is  given  more  from  its  antiquity  than 
from  its  present  day  importance,  and  yet  in  some  form  it  still  holds 
its  place  in  all  our  treaties  on  geology.  Indeed  the  latest  works 
include  discussions  of  the  strength  of  domes,  as  if  the  nucleus  of 
the  globe  were  shrinking  away  from  the  crust,  and  the  latter  thus 
subjected  to  crushing  from  its  own  weight. 

§  38.  Views  of  Lyell, — This  great  geologist  always  rejected  Elie 
de  Beaumont's  theories  of  mountain  formation,  and  gave  the  most 
cogent  reasons  for  his  course.  He  adopted  the  theory  that  the 
land  is  occasionally  depressed  and  elevated,  by  internal  forces,  but 
did  not  definitely  decide  what  forces  produced  these  progressive 

PROG.  AMER.  PHIL.  SOC.  XLVII.   189  P,  PRINTED  SEPTEMBER  23,   I908. 


238  SEE— FURTHER  RESEARCHES  ON  [A^iI^a, 

or  oscillatory  movements  of  the  earth's  crust.  One  of  Lyell's 
greatest  disciples  was  Charles  Darwin,  whose  views  we  shall  now 
very  briefly  recall. 

§  39-  Views  of  Charles  Darwin, — The  views  of  Darwin  are 
very  briefly  and  lucidly  set  forth  by  Professor  Suess  ("  Face  of 
the  Earth,"  Vol.  I,  p.  104),  as  follows: 

"The  earthquake  of  February  20,  1835  (at  Conception,  Chili),  gave  rise 
to  one  of  the  most  important  works  on  the  elevation  of  mountains,  indeed  I 
may  say  to  the  the  only  attempt,  based  on  direct  observation  of  nature,  to 
establish  more  exactly  the  older  theories  concerning  the  force  which  is  sup- 
posed to  have  raised  up  mountain  chains.  The  author  of  this  work  is  Charles 
Darwin.  Since  that  time  no  second  attempt,  or  at  least  no  attempt  of  equal 
importance,  has  been  made  in  this  direction.  To  day,  more  than  half  a 
century  later,  it  is  possible  to  hold  other  opinions  on  these  questions  and  yet 
to  recognize  the  boldness  of  the  generalization  which  even  then  revealed 
the  master. 

"Darwin  saw  the  awakening  activity  of  the  volcanoes  during  and  after 
the  earthquake;  he  believed  he  saw  elevation,  although  not  uniform  eleva- 
tion of  the  solid  ground;  in  addition  he  saw  the  terraces  along  the  coast. 
But  he  also  knew  that  similar  terraces  occur  on  the  east  coast  of  South 
America,  where  there  are  no  volcanoes  and  no  earthquakes.  The  earthquakes 
must  therefore  have  appeared  to  his  eyes  as  the  local  expression  of  a  uni- 
versal force.  The  secular  contraction  of  the  earth,  a  theory  already  eagerly 
advocated  by  several  investigators,  Darwin  justly  held  to  be  entirely  un- 
suited  to  explain  those  intermittent  elevations  which  the  terraces  betrayed, 
and  thus  he  reached  the  conclusion: 

" '  That  the  form  of  the  fluid  surface  of  the  nucleus  of  the  earth  is  sub- 
ject to  some  change,  the  cause  of  which  is  entirely  unknown  and  the  effect 
of  which  is  slow,  intermittent  but  irresistible/" 

§  40.  Viezvs  of  Professor  James  D,  Dana, — ^The  views  of  this 
eminent  geologist  have  been  carefully  discussed  in  the  paper  on 
"The  New  Theory  of  Earthquakes  and  Mountain  Formation  as 
Illustrated  by  Processes  Now  at  Work  in  the  Depths  of  the  Sea," 
§  13.  The  reader  is  referred  to  that  discussion.  Here  it  must 
suffice  to  say  that,  although  Dana  recognized  that  there  was  a  funda- 
mental relationship  between  the  depth  and  extent  of  an  ocean  and 
the  height  of  the  mountains  which  surround  it,  he  was  unable  to 
define  this  relationship  except  in  very  general  terms,  and  could 
not  assign  any  definite  cause  for  the  law  which  he  pointed  out. 
He  considered  the  oceanic  basins  as  subsiding,  while  the  continents 
were  being  elevated. 


,9o8.]  THE  PHYSICS  OF  THE  EARTH.  239 

Though  Dana's  views  were  somewhat  modified  by  later  study 
and  investigation,  he  always  maintained  that  "  the  principal  moun- 
tain chains  are  portions  of  the  earth's  crust  which  have  been  pushed 
up  and  often  crumpled  or  plicated  by  lateral  pressure  resulting  from 
the  earth's  contraction/'  In  order  to  explain  this  supposed  mode 
of  action  he  held  that  the  oceanic  areas  have  been  "  the  regions  of 
greatest  contraction  and  subsidence,  and  that  their  sides  have  been 
pushed  like  the  ends  of  an  arch,  against  the  borders  of  the  con- 
tinents." 

Even  with  these  arbitrary  assumptions  it  is  not  at  all  clear 
how  the  settlement  of  the  Pacific  Ocean  could  elevate  our  great 
plateau  west  of  the  Rocky  Mountains,  which  is  nearly  a  thousand 
miles  wide.  If  the  subsidence  of  the  ocean  bed  had  pushed  up  the 
margin  of  North  America,  the  crumpling  and  elevation  of  the  land 
could  not  well  extend  one  third  of  the  way  across  the  continent. 
We  need  not,  however,  be  greatly  surprised  at  this  difficulty,  for 
at  best  Dana's  theory  is  vague,  and  he  evidently  could  not  under- 
stand just  how  the  elevation  had  come  about.  Yet  so  fully  was 
Dana  convinced  of  the  dependence  of  the  mountains  on  the  oceans 
adjacent  to  them  that  he  reduced  it  to  calculation  by  the  rule-of- 
three.    He  says: 

"The  relation  of  the  oceans  to  the  mountain  borders  is  so  exact  that 
the  rule-of-three  form  of  statement  cannot  be  far  from  the  truth.  As  the 
size  of  the  Appalachians  to  the  size  of  the  Atlantic,  so  is  the  size  of  the 
Rocky  chain  to  the  size  of  the  Pacific.  Also,  as  the  height  of  the  Rocky 
chain  to  the  extent  of  the  North  Pacific,  so  are  the  height  and  boldness  of 
the  Andes  to  the  extent  of  the  South  Pacific."  ("  Manual  of  Geology,"  1863, 
p.  25.) 

This  was  indeed  a  remarkably  near  approach  to  the  great  law 
of  nature,  that  the  mountains  along  the  coasts  are  formed  by  the 
expulsion  of  lava  from  under  the  sea,  and  are,  therefore,  every- 
where proportional  to  extent  and  depth  of  the  adjacent  oceans. 

§41.  Views  of  James  Hall. — In  1857  this  distinguished  Ameri- 
can geologist  announced  in  a  presidential  address  to  the  Ameri- 
can Association  at  Montreal,  that  the  enormous  depth  of  the  sedi- 
mentation  along  the  Appalachian  chain  was  due  to  the  prolonged 
accumulation  of  sediments  along  a  sinking,  off-shore  line  of  sea 
bottom.     He   reached   this   view    from   the   careful   study   of   the 


240  SEE— FURTHER  RESEARCHES  ON  [April  «4. 

Appalachian  and  other  American  mountain  regions.  To  explain 
such  deposits  he  supposed  that  marine  currents  had  formerly 
traversed  these  regions  and  by  gradually  depositing  sediments  of 
great  weight  had  also  sunk  the  crust  till  at  length  a  great  thickness 
was  attained.  When  the  rocks  thus  formed  had  become  solidified 
and  crystallized  the  borders  of  the  continent  were  afterwards  up- 
raised somehow.  He  did  not  indicate  how  the  uplift  had  come 
about,  nor  did  he  think  that  the  mountain  regions  had  been  raised 
separately.  Denudation  had  then  commenced,  and  finally  given  the 
mountains  the  forms  they  have  today. 

Keferstein,  Sir  John  Herschel,  Dr.  T.  Sterry  Hunt  and  others, 
along  with  Hall,  or  even  before  him,  in  some  cases,  had  de- 
veloped the  theory  of  aqueo-igneous  fusion,  which  was  supposed 
to  produce  a  plastic  zone  between  the  consolidated  crust  and  the 
solid  nucleus.  This  theory  supposed  that  the  isogeotherms  rise  in 
regions  of  heavy  sedimentation.     Hall  held  that  this  would 

"cause  the  bottom  strata  to  establish  lines  of  weakness  or  of  least  resis- 
tance in  the  earth's  crust,  and  thus  determine  the  contraction  which  results 
from  the  cooling  of  the  globe  to  exhibit  itself  in  those  regions,  and  along 
those  lines  where  the  ocean's  bed  is  subsiding  beneath  the  accumulated 
sediments." 

Many  of  the  views  afterwards  more  fully  developed  by  Leconte 
are  here  faintly  traced  by  Hall,  and  for  that  reason  these  early 
views  of  mountain  formation  are  worthy  of  attention. 

§  42.  Views  of  Leconte, — This  veteran  geologist  gave  great 
attention  to  mountain  formation  throughout  a  long  career,  and  his 
residence  on  the  Pacific  Coast  gave  him  exceptional  facilities  for 
studying  the  ranges  of  our  western  states,  and  especially  of  Cali- 
fornia, which  includes  the  most  remarkable  developments  in  North 
America.  The  views  at  which  Leconte  arrived,  as  set  forth  in  his 
"Elements  of  Geology,"  edition  of  1896,  are  as  follows: 

"Mountain  Origin. 
"Leaving  aside  for  the  present  all  disputed  points,  it  is  now  universally 
admitted  that  mountains  are  not  usually  pushed  up  by  a  vertical  force  from 
beneath,  as  once  supposed,  but  are  formed  wholly  by  lateral  pressure.  The 
earth's  crust  along  certain  lines  is  crushed  together  by  lateral  or  horizontal 
pressure  and  rises  into  a  mountain-range  along  the  line  of  yielding,  and  to 
a  height  proportionate  to  the  amount  of  mashing.  But  the  yielding  is  not 
by  rising  into  a  hollow  arch,  nor  into  such  an  arch  filled  beneath  with  liquid 


i9o«.J 


THE  PHYSICS  OF  THE  EARTH. 


241 


(for  in  neither  case  would  the  arch  support  itself),  but  by  mashing  together 
and  in  thickening  and  crumpling  of  the  strata  and  an  upswelling  of  the  whole 
mass  along  the  line  of  greatest  yielding.  That  this  is  the  immediate  or 
proximate  cause  of  the  origin  or  elevation  of  mountains  is  plainly  shown  by 
their  structure.  As  to  the  ultimate  cause — i.  e.,  the  cause  of  the  enormous 
lateral  pressure — this  lies  still  in  the  Held  of  discussion.  We  shall  discuss 
it  briefly  in  its  proper  place"  (pp.  261-2). 

Again,  on  page  264,  we  find  this  account : 

"Proof  of  Elevation  by  Lateral  Pressure  alone:  i.  Folding. — It  is  evi- 
dent that  foldings  such  as  those  represented  in  all  the  above  figures,  and 
which  occur  in  nearly  all  mountains,  cannot  be  produced  except  by  lateral 
pressure,  and  are  therefore  proof  of  such  pressure.  But,  moreover,  it  can 
be  shown  that,  when  we  take  into  consideration  the  immense  thickness  of 
mountain  strata  and  the  degree  of  folding,  lateral  pressure  is  sufficient  to 
account  for  the  whole  elevation,  without  calling  in  the  aid  of  any  upward 
pushing  from  beneath.  For  example,  the  Coast  range  of  California  (Fig. 
228)  is  composed  of  at  least  Ave  anticlines  and  corresponding  synclines.    If 


Fig.  14.     Section  of  a  Portion  of  the  Alps. 

its  folded  strata  were  spread  out  horizontally  in  the  position  of  the  original 
sediments,  they  would  undoubtedly  cover  double  the  space.  Now,  supposing 
the  strata  here  are  only  10,000  feet  thick — a  very  moderate  estimate — in 
mashing  to  one  half  the  extent,  they  would  be  thickened  to  20,000  feet, 
which  would  be  a  clear  elevation  of  10,000  feet  if  they  were  not  subsequently 
eroded  According  to  Renevier,  a  section  of  the  Alps  reveals  seven  anticlines 
and  corresponding  synclines,  and  some  of  them  are  complete  overfolds  (Fig. 
230).  We  are  safe  in  saying  that  Alpine  strata  have  been  mashed  horizontally 
into  one  half  their  original  extent.  Supposing  these  were  originally  30,000 
feet  thick  (they  were  really  much  thicker),  this  would  make  a  clear  eleva- 
tion of  30,000  feet.  Of  course,  most  of  this  has  been  cut  away  by  erosion. 
In  the  Appalachian  range,  according  to  Claypole,  the  foldings  are  so  extreme 
that  in  one  place  95  miles  of  original  extent  have  been  mashed  into  16  miles, 
or  six  into  one,  and  yet  the  Appalachian  strata  are  estimated  as  40,000  feet 
thick.    Cases  of  still  greater  doubling  of  strata  upon  themselves  occur.    In 


242  SEE— FURTHER   RESEARCHES  ON  [Apraa4, 

the  Highlands  of  Scotland  the  strata  by  lateral  thrust  were  broken  and 
slidden  one  over  another  for  ten  miles.  In  the  Canadian  Rocky  Mountains 
there  is  an  overthrust  of  seven  miles,  by  which  the  Cambrian  is  made  to 
override  the  Cretaceous,  and  50  miles  of  strata  are  mashed  into  25  miles 
(McConnell).  In  the  Appalachians  of  Georgia  the  Rome  fault  is  an  over- 
thrust  which  brings  the  Cambrian  in  contact  with  the  Carboniferous  and  the 
fault  under  different  names  may  be  traced  northward  for  275  miles;  and  in 
the  Cartersville  thrust-fault  there  is  an  overriding  of  11  miles  (Hayes).  The 
manner  in  which  this  is  done  is  illustrated  on  a  previous  page  (Fig.  209). 
Evidently,  then,  the  whole  height  of  the  mountains  mentioned  above  is  due 
to  lateral  crushing  alone." 

If  Professor  Leconte  had  been  familiar  with  the  folding  pro- 
duced in  the  sea  trenches  he  could  have  completed  the  theory  of 
mountain  formation  developed  in  this  paper.  As  geologists  have 
for  centuries  recognized  the  fossils  found  in  mountains  as  having 
been  deposited  in  the  sea,  it  is  remarkable  that  the  suggestion  seems 
never  to  have  occurred  to  them  that  the  folding  was  done  in  the  sea 
before  the  land  was  lifted  above  the  water,  and  by  earthquake  proc- 
esses due  to  the  sea  itself.  Leconte,  however,  came  very  near  this 
view,  as  the  following  will  show  (p.  267  et  seq.)  : 

"Mountains  are  made  out  of  lines  of  thick  sediments. — But  the  question 
occurs,  What  determines  the  place  of  a  mountain-range?  The  answer  is, 
A  mountain-range  while  in  preparation — ^before  it  became  a  range — was  a 
line  of  very  thick  sediments.  This  is  a  very  important  point  in  the  theory 
of  mountain  origin,  and  therefore  must  be  proved.  The  strata  of  all  moun- 
tains, where  it  is  possible  to  measure  them,  are  found  to  be  of  enormous 
thickness.  The  strata  involved  in  the  folded  structure  of  the  Appalachian, 
according  to  Hall,  are  40,000  feet  thick,  the  strata  exposed  in  the  structure 
of  the  Wahsatch,  according  to  King,  are  more  than  50,000  feet  thick;  the 
Cretaceous  strata  of  the  Coast  Range,  near  the  Bay  of  San  Francisco, 
according  to  Whitney,  are  20,000  feet  thick;  and  if  we  add  to  this  10,000 
feet  for  the  Eocene  and  Miocene  strata,  the  whole  thickness  is  probably  not 
less  than  30,000  feet,  while  the  Cretaceous  alone  in  Northern  California, 
according  to  Diller,  is  30,000  feet.  The  Alpine  geologists  estimate  the 
thickness  of  the  strata  involved  in  the  intricate  structure  of  the  Alps  as 
50,000  feet.  The  strata  of  Uintah,  according  to  Powell,  are  32,000  feet 
thick. 

"  Now,  it  must  not  be  imagined  that  these  numbers  merely  represent 
the  general  thickness  of  the  stratified  crust;  only  that  in  these  places  the 
strata  are  turned  up  and  their  edges  exposed  by  erosion,  and  thus  their 
thickness  revealed.  On  the  contrary,  it  may  be  shown  that  the  same  strata 
are  much  thinner  elsewhere.  The  same  strata  which  along  the  Appalachian 
range  are  40,000  feet  thick,  when  traced  westward  thin  out  to  4,000  feet  at 
the  Mississippi  River.    The  same  strata  which  along  the  line  of  the  Wah- 


X908.J  THE  PHYSICS  OF  THE  EARTH.  243 

satch  are  30,000  feet  thick,  when  traced  eastward  thin  out  to  2,000  feet  in 
the  region  of  the  plains.  It  is  evident,  therefore,  that  mountain- ranges  are 
lines  of  exceptionally  thick  strata. 

"Mountain-ranges  were  once  Marginal  Sea-Bottoms. — Where,  then,  do 
sediments  now  accumulate  in  greatest  thickness?  Evidently  on  marginal  sea- 
bottoms,  off  the  coasts  of  continents.  The  greater  part  of  the  washings  of 
continents  are  deposited  within  30  miles  of  shore,  and  the  whole  usually 
within  100  miles.  From  this  line  of  thickest  and  coarsest  deposit  the  sedi- 
ments grow  thinner  and  finer  as  we  go  seaward.  But  evidently  such  enor- 
mous thicknesses  as  40,000  feet  cannot  accumulate  in  the  same  place  with- 
out pari  passu  subsidence  such  as  we  know  takes  place  now  whenever 
exceptionally  abundant  sedimentation  is  going  on  (p.  145).  Therefore, 
mountain-ranges  before  they  were  yet  born — while  still  in  preparation  as 
embryos  in  the  womb  of  the  ocean — were  lines  of  thick  off-shore  deposits 
gradually  subsiding,  and  thus  ever  renewing  the  conditions  of  continuous 
deposits. 

"As  this  is  a  very  important  point,  it  is  necessary  to  stop  here  awhile 
in  order  to  show  that  such  was  actually  the  fact  in  the  case  of  all  the 
principal  ranges  of  the  American  Continent — i.  e.,  that  for  a  long  time  before 
they  were  actually  formed,  the  places  which  they  now  occupy  were  marginal 
sea-bottoms  receiving  abundant  sediments  from  an  adjacent  continent  We 
shall  be  compelled  to  anticipate  some  things  that  belong  to  Part  III,  but  we 
hope  to  make  statements  so  general  that  there  will  be  no  difficulty  in  under- 
standing them. 

"  I.  Appalachian. — The  history  of  this  range  is  briefly  as  follows :  At 
the  beginning  of  the  Palaeozoic  era  there  was  a  great  V-shaped  land-mass, 
occupying  the  region  now  covered  by  Labrador  and  Canada,  then  turning 
northwestward  from  Lake  Superior  and  extending  perhaps  to  polar  regions 
about  the  mouth  of  the  Mackenzie  River.  This  is  shown  on  map,  Fig.  269, 
on  page  303.  There  is  another  great  land-mass  occupying  the  present  place 
of  the  eastern  slope  of  the  Blue  Ridge  and  extending  eastward  probably 
far  beyond  the  present  limits  of  the  continent — ^as  shown  in  the  same  figure 
by  dotted  line  in  the  Atlantic  Ocean.  The  western  coast-line  of  this  land- 
mass  was  the  present  place  of  the  Blue  Ridge.  .Westward  of  this  line 
extended  a  great  ocean — *the  interior  Palaeozoic  Sea.'  The  Appalachian 
range  west  of  the  Blue  Ridge  was  then  the  marginal  bottom  of  that  sea. 
During  the  whole  of  the  Cambrian,  Silurian,  and  Devonian,  this  shoreline 
remained  nearly  in  the  same  place,  although  there  was  probably  a  slow 
transference  westward.  Meanwhile,  throughout  this  immense  period  of 
time,  the  washings  from  the  land-mass  eastward  accumulated  along  the 
shore-line,  until  30,000  feet  of  thickness  was  attained.  At  the  end  of  the 
Devonian  some  considerable  changes  of  physical  geography  of  this  region 
took  place,  which  we  will  explain  when  we  come  to  treat  of  the  history 
of  this  period.  Suffice  it  to  say  now  that  during  the  Carboniferous  the 
region  of  the  Appalachian  was  sometimes  above  the  sea  as  a  coal-swamp, 
and  sometimes  below,  but  all  the  time  receiving  sediment  until  pxxx)  or 
10,000  feet  more  of  thickness  was  added,  and  the  aggregate  thickness  became 


244  SEE— FURTHER  RESEARCHES  ON  [April  m. 

40,000  feet.  Of  course,  it  is  impossible  that  such  thickness  could  accumulate 
on  the  same  spot  without  pari  passu  subsidence  of  the  sea-floor.  In  fact, 
we  have  abundant  evidences  of  comparatively  shallow  water  at  every  step 
of  the  process — evidence  sometimes  in  the  character  of  the  fossils,  some- 
times in  the  form  of  shore-marks  of  all  kinds,  sometimes  in  the  form  of 
seams  of  coal,  showing  even  swamp-land  conditions.  Again,  of  course,  the 
sediments  were  thickest  and  coarsest  near  the  shore-line,  and  thinned  out 
and  became  finer  towards  the  open  sea,  i,  e.,  westward.  Finally,  after  40,000 
feet  of  sediments  had  accumulated  along  this  line  the  earth-crust  in  this 
region  gave  way  to  lateral  pressure,  and  the  sediments  were  mashed  together 
and  folded  and  swollen  up  into  the  Appalachian  range.  Subsequent  erosion 
has  sculptured  it  into  the  forms  of  scenic  beauty  which  we  find  there  to-day. 

"  2.  Sierra. — This  was  apparently  the  first-horn  of  the  Cordilleran  family. 
Its  history  is  as  follows:  During  the  whole  Palaeozoic  and  earlier  part  of 
the  Mesozoic,  there  was  in  the  Basin  region  a  land-mass,  whose  form  and 
dimensions  we  yet  imperfectly  know,  but  whose  Pacific  shore-line  was  east 
of  the  Sierra,  The  Sierra  region  was  therefore  at  that  time  the  marginal 
bottom  of  the  Pacific  Ocean.  Probably  the  position  of  this  shore  line  changed 
considerably  at  the  end  of  the  Palaeozoic.  The  extent  of  this  change  we 
will  discuss  hereafter.  Suffice  it  to  say  now  that,  during  the  whole  of  this 
time,  the  Sierra  region  received  sediments  from  this  land-mass  until  an 
enormous  thickness  (how  much  we  do  not  know,  because  the  foldings  are 
too  complex  to  allow  of  estimate)  was  accumulated.  At  last  at  the  end 
of  the  Jurassic,  the  sea  floor  gave  way  to  the  increasing  lateral  pressure 
along  the  line  of  thickest  sediments,  and  th^se  latter  were  crushed  together 
with  complex  foldings  and  swollen  up  into  the  Sierra.  An  almost  incon- 
ceivable subsequent  erosion  has  sculptured  it  into  the  forms  of  beauty  and 
grandeur  which  characterize  its  magnificent  scenery. 

"  3.  Coast  Range. — The  birth  of  the  Sierra  transferred  the  Pacific  shore- 
line westward,  and  the  waves  now  washed  against  the  western  foot  of  that 
range,  or  possibly  even  father  westward  in  the  region  of  the  Sacramento  and 
San  Joaquin  plains.  At  this  time,  therefore,  the  region  of  the  Coast  Range 
was  the  marginal  bottom  of  the  Pacific  Ocean.  During  the  whole  Cretaceous, 
Eocene,  and  Miocene,  this. region  received  abundant  sediments  from  the  now 
greatly  enlarged  continental  mass  to  the  eastward;  until  finally,  at  the  end 
of  the  Miocene,  when  30,000  feet  of  sediments  had  accumulated  along  this 
line,  the  sea-floor  yielded  to  the  lateral  pressure,  and  the  Coast  Range  was 
bom ;  and  the  coast-line  transferred  to  near  its  present  position. 

"4.  Wahsatch. — The  physical  geography  of  the  region  to  the  east  of 
the  Wahsatch  (Plateau  region)  during  Jura-Trias  time  is  little  known.  But 
during  the  Cretaceous  the  region  of  the  Wahsatch  was  the  western  marginal 
bottom  of  the  great  interior  Cretaceous  Sea  (see  map,  Fig.  760,  p.  486), 
receiving  abundant  sediments  from  the  great  land-mass  of  the  Basin  and 
Sierra  region.  This  greatly  increased  the  enormous  thickness  of  sediments 
already  accumulated  along  this  line  in  earlier  times.  At  the  end  of  the 
Cretaceous  the  sediments  yielded,  and  the  Wahsatch  was  bom.  It  is  neces- 
sary, however,  to  say  that  both  the  Sierra  and  Wahsatch  underwent  very 


,9o8.J  THE  PHYSICS  OF  THE  EARTH.  245 

great  changes  of  form  produced  by  a  different  process  and  at  a  much  earlier 
period.    We  shall  speak  of  this  later. 

"5.  Alps, — Mr.  Judd  has  recently  shown  that  the  region  of  the  Alps, 
during  the  whole  Mesozoic  and  Early  Tertiary,  was  a  marginal  sea  bottom, 
receiving  sediments  until  a  thickness  was  attained  not  less  than  that  of  the 
Appalachian  strata.  At  the  end  of  the  Eocene  these  enormously  thick  sedi- 
ments were  crushed  together  with  complicated  foldings  and  swollen  upward 
to  form  these  mountains  and  afterward  sculptured  to  their  present  forms. 

"The  same  may  be  said  of  the  Himalayas  and  nearly  all  other  moun- 
tains. We  may,  therefore,  confidently  generalize,  and  say  that  the  place 
now  occupied  by  mountain-ranges  have  been  previous  to  their  formation, 
places  of  great  sedimentation,  and  therefore  usually  marginal  ocean  bottoms. 
In  some  cases,  however,  the  deposits  in  interior  seas  or  mediterraneans  have 
yielded  in  a  similar  way,  giving  rise  to  more  irregular  ranges  or  groups  of 
mountains."  .  .  . 

"Why  thick  Sediments  should  be  Lines  of  Yielding. — Admitting,  then, 
that  mountains  are  formed  by  the  squeezing  together  of  lines  of  very  thick 
sediments,  the  question  still  occurs.  Why  does  the  yielding  take  place  along 
these  lines  in  preference  to  any  others?  This  is  a  capital  point  in  the 
theory  of  mountain  formation.  The  answer  is  as  follows:  We  have  already 
seen  (p.  231)  that  accumulation  of  sediments  causes  the  isogeotherm  to  rise 
and  the  interior  heat  of  the  earth  to  invade  the  lower  portion  of  the  sedi- 
ments with  their  included  waters.  Now  this  invasion  of  heat  in  its  turn 
causes  hydrothermal  softening  or  even  fusion,  not  only  of  the  sediments, 
but  also  of  the  sea-floor  on  which  they  rest  Thus  a  line  of  thick  sediments 
becomes  a  line  of  softening  and  therefore  a  line  of  weakness,  and  a  line 
of  yielding  to  the  lateral  pressure,  and  therefore  a  line  of  mashing  together 
and  folding  and  upswelling — in  other  words  a  mountain-range.  As  soon  as 
the  yielding  commences  we  have  an  additional  source  of  heat  in  the  crush- 
ing itself.  In  addition  to  this,  upheaval  by  lateral  crush  by  the  tendency 
to  arch  the  strata  would  produce  relief  of  gravitative  pressure,  and  there- 
fore fusion  (p.  103).  It  follows  from  this  that  there  is  or  was  beneath 
every  mountain  a  line  of  fused  or  semi-fused  matter.  This  we  will  call 
the  sub-mountain  liquid.  This  by  cooling  and  solidification  becomes  a  meta- 
morphic  or  granitic  core,  which  by  erosion  forms  the  metamorphic  or  granitic 
axis  and  crest  of  many  great  mountains"  .  .  .  (pp.  271-2). 

"Cause  of  Lateral  Pressure. — ^We  have  thus  proved  that  the  immediate 
cause  of  the  origin  and  the  growth  of  mountains  is  lateral  pressure  acting 
on  thick  sediments,  crushing  them  together  and  swelling  them  up  along  the 
line  of  great  thickness.  But  still  the  question  remains.  What  is  the  ultimate 
cause,  f.  e.,  the  cause  of  the  lateral  pressure?  This,  as  we  have  already  said, 
lies  still  in  the  domain  of  doubt  and  discussion,  but  the  view  which  seems 
most  probable  may  be  briefly  stated  as  follows: 

"In  the  secular  cooling  of  the  earth  there  would  be  not  only  unequal 
radial  contraction,  giving  rise,  as  shown  on  page  175,  to  continents  and 
ocean-basins,  but  also  to  unequal  contraction  of  the  exterior  as  compared 
with  the  interior.    At  first,  and  for  a  long  time,  the  exterior  would  cool 


246  SEE--FURTHER   RESEARCHES  ON  [April  t4. 

fastest;  but  there  would  inevitably,  sooner  or  later,  come  a  time  when  the 
exterior,  feceiving  heat  from  abroad  (sun  and  space),  as  well  as  from 
within,  would  assume  an  almost  constant  temperature,  while  the  interior 
would  still  continue  to  cool,  and  contract.  Thus,  therefore,  after  a  while 
the  interior  nucleus  would  contract  faster  than  the  exterior  shell.  It  would 
do  so,  partly  because  it  would  cool  faster,  and  partly  because  the  coefficient 
of  contraction  of  a  hot  body  is  greater  than  that  of  a  cooler  body.  Now,  as 
soon  as  this  condition  was  reached,  the  exterior  shell,  following  down  the 
shrinking  nucleus,  would  be  thrust  upon  itself  by  a  lateral  or  horizontal 
pressure  which  would  be  simply  irresistible.  If  the  earth's  crust  were  a 
hundred  times  more  rigid  than  it  is  (thirty  times  as  rigid  as  steel,  500  to 
1,000  times  as  rigid  as  granite — Woodward,  Science^  Vol.  XIV,  p.  167,  1889), 
it  must  yield.  Mountain-ranges  are  the  lines  along  which  the  yielding  takes 
place,  and  this  yielding  takes  place  along  the  lines  of  thick  sediments  be- 
cause these  are  lines  of  weakness. 

"There  are  several  serious  objections  which  may  be  brought  against 
this  view:  i.  (Calculations  seem  to  show  that  the  amount  of  crumpling  and 
folding  actually  found  in  the  mountains  is  many  times  greater  than  could 
be  produced  by  the  contraction  of  the  earth  by  cooling.  But  it  may  be 
answered  (i)  that  the  calculations  take  no  account  of  the  greater  coefficient 
of  contraction  at  high  temperatures,  and  therefore  at  great  depths,  (2)  and 
that  there  may  be  other  causes  of  contraction  besides  cooling.  For  example, 
loss  of  constituent  gases  and  vapors  from  the  interior  of  the  earth,  through 
volcanic  vents  and  fissures,  has  been  suggested  by  O.  Fisher  (p.  102). 

"2.  Again,  it  has  been  shown  by  Dutton  that  it  is  impossible  that  the 
effects  of  differential  contraction  should  be  concentrated  along  certain  lines, 
so  as  to  give,  rise  to  mountain-ranges  without  a  shearing  of  the  crust  upon 
the  interior  portions,  which  is  inadmissible  if  the  earth  be  solid.  Instead, 
therefore,  of  conspicuous  mountain-ranges,  the  effects  of  differential  con- 
traction would  be  distributed  all  over  the  surface,  and  be  wholly  impercep- 
tible. But  in  answer  to  this  it  may  be  said  that  there  is  no  difficulty  in  the 
way  of  shearing,  and  therefore  of  such  concentration  of  effects  along  certain 
lines,  if  there  be  a  sub-crust  liquid  or  semi-liquid  layer,  either  universal  or 
else  underlying  large  areas  of  surface. 

"  Still  other  objections  have  been  raised,  but  these  are  so  recent  that  they 
have  not  yet  been  sufficiently  sifted  by  discussion  to  deserve  mention  here.* 
The  origin  of  mountains  by  lateral  pressure  is  a  fact  beyond  dispute.  This  is 
the  most  important  fact  for  the  geologist.  How  the  lateral  pressure  is  pro- 
duced is  a  pure  physical  question  which  must  be  left  to  the  physicists  to 
settle  among  themselves"  (pp.  274-5). 

Leconte  treats  also  of  Monoclinal  mountains,  as  found  in  the 
Great  Basin,  which  he  explains  by  normal  faulting,  or  vertical  move- 
ment of  crust  blocks,  and  finally  adds : 

*  For  a  completer  discussion  of  this  subject,  see  **  Theories  of  Mountain 
Origin,"  Jour.  GeoL,  Vol.  I.,  p.  542,  1893. 


1908. 1 


THE  PHYSICS  OF  THE  EARTH.  247 


"  Thus,  then,  there  are  two  types  of  mountains  strongly  contrasted, 
mountains  of  the  one  type  are  formed  by  lateral  pressure  and  crushing,  of 
the  other  type  by  lateral  tension  and  stretching.  The  one  gives  rise  mainly 
to  reverse  faults,  the  other  always  to  normal  faults.  Mountains  of  the  one 
type  are  formed  by  upswelling  of  thick  sediments,  those  of  the  other  type 
by  irregular  readjustment  of  crust-blocks.  Mountains  of  the  one  type  are 
horn  of  the  sea,  those  of  the  other  type  are  bom  on  the  land.  We  find 
examples  of  the  one  type  in  nearly  all  the  greatest  mountains  everywhere,  but 
especially  in  the  Appalachian,  the  Alps  and  the  Coast  Range.  The  best 
examples,  perhaps  the  only  examples,  of  the  other  type  are  the  Basin  ranges. 
Some  mountains,  as  the  Sierra,  the  Wahsatch,  and  certainly  some  of  the 
Basin  ranges,  belong  to  both  types.  In  their  origin,  they  have  formed  in 
the  first  way,  but  afterward  have  been  modified  by  the  second  way.  Thus 
the  first  is  the  fundamental  method,  and  the  second  only  a  modifying  proc- 
ess" (p.  277). 

These  views  of  Leconte  call  for  no  special  comment,  beyond  the 
remark  that  normal  faulting  itself  is  wholly  unexplained.  If  secu- 
lar cooling  were  the  cause,  such  faults  ought  to  occur  east  of  the 
Rocky  Mountains  as  well  as  west  of  them.  The  important  diflfer- 
ence  is  that  the  Pacific  Ocean  was  on  the  west  pushing  up  the  land, 
and  a  continental  basin  on  the  east,  either  dry  or  covered  by  shallow 
water  and  therefore  doing  little  or  no  pushing  at  all.  In  any  case 
the  great  plateaus  of  the  west  were  certainly  uplifted  by  the  Pacific, 
through  the  expulsion  of  lava  under  the  land.  In  the  Andes  of 
South  America  the  plateaus  are  higher  indeed,  but  also  narrower 
than  those  in  North  America,  because  in  our  continent  the  relief 
resulting  from  the  leakage  of  the  ocean  took  a  broader  and  less  ele- 
vated form.  It  is  impossible  for  any  one  to  doubt  the  identity  of 
the  forces  which  raised  the  Andes  and  their  plateaus,  the  Himalayas 
and  their  plateaus,  and  the  Rocky  Mountains  and  the  mountains  and 
plateaus  of  the  Great  Basin.  The  principle  of  continuity  shows 
clearly  that  the  cause  was  everywhere  one  and  the  same.  Several 
American  geologists  have  suggested  vertical  uplifts  in  the  Great 
Basin,  from  the  way  in  which  the  crust  blocks  are  displaced;  but 
heretofore  no  known  cause  for  such  movements  could  be  assigned, 
because  it  was  held  that  secular  cooling  is  the  chief  if  not  the  only 
cause  operating  in  the  development  of  the  globe. 

§43.  Views  of  Rev.  O.  Fisher, — The  Rev.  O.  Fisher  was  the 
first  to  show  by  long  and  patient  research  the  total  inadequacy  of 
secular  cooling  to  account  for  the  observed  height  of  mountains. 


248  SEE— FURTHER   RESEARCHES  ON  [April  24. 

He  showed  that  the  mountains  are  hundreds  of  times  higher  than 
the  cooling  of  the  earth  will  explain.  On  this  point  his  labors  mark 
a  distinct  advance  in  geological  science;  for  next  in  importance  to 
establishing  true  theories  is  the  overthrow  of  erroneous  ones,  which 
clears  the  ground  for  a  fresh  start.  But  notwithstanding  the  un- 
answerable character  of  Fisher's  argument,  the  old  theories  have 
been  retained  by  geologists  as  the  best  they  could  devise.  Fisher's 
criticisms  of  geological  theories  are  carefully  thought  out,  and 
worthy  of  attention.  He  has  always  denied  the  entire  solidity  of 
the  earth,  holding  that  the  movements  noticed  in  mountains  proved 
the  existence  of  a  mobile  substratum  beneath  a  crust  some  twenty 
miles  thick.  Here  again  he  was  certainly  right,  and  it  is  difficult 
to  see  how  such  an  obvious  proposition  could  be  denied. 

We  need  not  dwell  on  Fisher's  views  of  mountain  formation, 
because  they  imply  convection  currents  within  the  earth,  and  these 
latter  are  certainly  inadmissible,  except  just  beneath  the  crust  in 
earthquake  movements,  as  developed  in  the  theory  set  forth  in  this 
paper. 

§  44.  Views  of  Major  C,  £.  Duttotk — ^Like  the  Rev.  O.  Fisher, 
Major  Dutton  was  one  of  the  earliest  authorities  to  question  the 
adequacy  of  secular  cooling  to  account  for  the  wrinklings  noticed 
in  the  earth's  crust.  Using  the  results  of  Fourier's  solution  for  the 
variation  of  temperature,  as  developed  in  the  work  of  Lord  Kelvin, 
Dutton  found  that 

"  the  greatest  possible  contraction  due  to  secular  cooling  is  insufficient  in 
amount  to  account  for  the  phenomena  attributed  to  it  by  the  contraction 
hypothesis.  By  far  the  larger  portion  of  this  contraction  must  have  taken 
place  before  the  commencement  of  the  Palaeozoic  age.  By  far  the  larger 
portion  of  the  residue  must  have  occurred  before  the  beginning  of  the  Terti- 
ary, and  yet  the  whole  of  this  contraction  would  not  be  sufficient  to  account 
for  the  disturbances  which  have  occurred  since  the  close  of  the  Cretaceous." 

Major  Dutton  concludes  that  "the  determination  of  plications  to 
particular  localities  presents  difficulties  in  the  way  of  the  contrac- 
tional  hypothesis  which  have  been  underrated."  He  held  that  the 
localization  of  the  plications  could  result  only  from  a  large  amount 
of  horizontal  slipping  of  the  crust  over  the  nucleus,  and  the  friction 
involved  in  this  movement  even  over  a  liquid  nucleus  would  be  so 
great  as  to  render  the  assumption  a  physical  absurdity. 


x9o8.] 


THE  PHYSICS  OF  THE  EARTH.  249 


If  wrinkling  resulted  from  uniform  cooling  and  consequently 
uniform  shrinkage,  the  effect  would  be  analogous  to  that  of  a  with- 
ered apple,  with  small  wrinkles  all  over  it,  instead  of  a  surface 
presenting  in  one  region  a  continuous  system  of  folds  extending 
from  Cape  Horn  to  Alaska,  and  in  another,  a  zone  a  thousand  miles 
wide,  from  the  Appalachian  to  the  Rocky  Mountains,  with  scarcely 
any  evidence  of  disturbance  whatever. 

In  these  considerations  Major  Dutton  has  forcibly  expressed 
the  difficulty  of  supposing  that  a  mountain  range  is  formed  by  the 
cooling  of  the  earth  contracting  equally  along  all  its  radii.  Such 
a  supposed  mode  of  formation  of  our  ranges,  folded  and  crumpled 
as  they  are,  is  clearly  impossible;  and  Major  Dutton  shares  with 
the  Rev.  O.  Fisher  the  credit  of  having  been  the  first  to  recognize 
the  total  inadequacy  of  the  contraction  theory. 

It  is  remarkable  that  after  this  antiquated  theory  had  been  thus 
clearly  disproved,  it  should  have  continued  in  use.  No  one  seems 
to  have  been  able  to  frame  a  theory  based  on  any  cause  except  secu- 
lar cooling,  till  the  present  writer  developed  the  theory  based  on  the 
leakage  of  the  oceans  and  the  formation  of  mountains  by  the  expul- 
sion of  lava  under  the  land,  which  perfectly  explains  all  the 
phenomena. 

§  45.  Views  of  Geikie. — In  the  article  "  Geology,"  Encyclopedia 
Britannica^  p.  375,  we  find  the  following  statement  of  the  contrac- 
tion theory: 

"There  still  remains  the  problem  to  account  for  the  original  wrinkling 
of  the  surface  of  the  globe,  whereby  the  present  great  ridges  and  hollows 
were  produced. 

"  It  is  now  generally  agreed  that  these  inequalities  have  been  produced 
by  unequal  contraction  of  the  earth's  mass,  the  interior  contracting  more 
than  the  outer  crust,  which  must  therefore  have  accommodated  itself  to  this 
diminution  of  diameter  by  undergoing  corrugation.  But  there  seems  to  have 
been  some  original  distribution  of  materials  in  the  globe  that  initiated  the 
depressions  on  the  areas  which  they  have  retained.  It  has  been  already 
pointed  out  (ante,  p.  223)  that  the  matter  underlying  the  oceans  is  more 
dense  than  that  beneath  the  continents,  and  that,  partly  at  least,  to  this  cause 
must  the  present  position  of  the  oceans  be  attributed.  The  early  and  per- 
sistent subsidences  of  these  areas,  with  the  consequent  increase  of  density, 
seems  to  have  determined  the  main  contours  of  the  earth's  surface.  .  .  . 

"The  effects  of  this  lateral  pressure  may  show  themselves  either  in 
broad  dome-like  elevations,  or  in  narrower  and  loftier  ridges  of  mountains. 


250  SEE— FURTHER   RESEARCHES  ON  [April  24, 

The  structure  of  the  crust  is  so  complex,  and  the  resistance  offered  by  it 
to  the  pressure  is  consequently  so  varied,  that  abundant  cause  is  furnished 
for  almost  any  diversity  in  the  forms  and  distribution  of  the  wrinkles  into 
which  it  is  thrown.  It  is  evident,  however,  that  the  folds  have  tended  to 
follow  a  linear  direction.  In  North  America,  from  early  geological  times, 
they  have  kept  on  the  whole  on  the  lines  of  meridians.  In  the  Old  World, 
on  the  contrary,  they  have  chosen  diverse  trends,  but  the  last  great  crumplings 
— those  of  the  Alps,  Caucasus,  and  the  great  mountain  ranges  of  central 
Asia — have  risen  along  parallels  of  latitude. 

"  Mountain  chains  must  therefore  be  regarded  as  evidence  of  the  shrink- 
age of  the  earth's  mass.  They  may  be  the  result  of  one  movement,  or  of  a 
long  succession  of  such  movements.  Formed  on  lines  of  weakness  in  the 
crust,  they  have  again  and  again  given  relief  from  the  strain  of  compression 
by  undergoing  fresh  crumpling  and  upheaval." 

Geikie's  views  may  be  considered  the  accepted  views  of  geolo- 
gists generally,  and  it  will  be  seen  that  they  rest  on  the  theory  of 
contraction  due  to  secular  cooling. 

On  the  constitution  of  the  globe  Geikie  quotes  ("  Geology," 
p.  73)  from  the  paper  of  Arrhenius,  "  Zur  Physik  des  Vulcanismus  " 
(1900),  the  following  theory  of  the  illustrious  Swedish  physicist: 

"  If  the  rocks  at  the  earth's  surface  have  a  density  half  that  of  the 
globe  as  a  whole,  and  if  the  density  continues  to  hold  good  for  the  magma 
that  arises  from  the  melting  of  these  rocks,  we  must  conceive  the  existence 
of  a  much  denser  substance  in  the  earth's  interior.  On  various  grounds, 
such  as  the  preponderance  of  iron  in  nature,  both  in  meteorites  and  in  the 
sun,  and  the  phenomena  of  terrestrial  magnetism,  it  may  be  inferred  that 
this  substance  is  metallic  iron.  In  consequence  of  its  greater  density  this 
iron  will  naturally  be  deeper  than  the\  rock  magma,  and  on  account  of  the 
high  temperature  must  exist  in  a  gaseous  condition.  Somewhere  about  a 
half  of  the  planet  therefore  should  consequently  consist  of  iron,  and  of 
other  metals  mingled  with  it  in  smaller  proportions.  The  semi-diameter  of 
this  gaseous  iron-sphere  will  thus  include  about  80  per  cent,  of  the  earth's 
semi-diameter.  Then  will  come  about  15  per  cent,  of  the  gaseous  rock  magma» 
next  to  it  the  liquid  rock-magma  for  a  thickness  of  about  4  per  cent,  of 
the  terrestrial  semi-diameter,  and  lastly  the  solid  crust,  for  which  not  more 
than  I  per  cent,  may  be  claimed"  (pp.  404-5). 

Referring  to  the  light  thrown  on  the  constitution  of  the  interior 
by  the  observation  of  waves  propagated  by  earthquakes,  Geikie  also 
adopts  the  theory  of  Arrhenius,  which  is  as  follows : 

"The  density  of  much  the  largest  part  (reckoned  linearly)  of  this 
interior,  amounting,  as  above  stated,  to  about  80  per  cent,  of  the  radius, 
must  be  nearly  three  times  higher  than  that  of  quartz.  Since  now  the 
mean  velocity  of  transmission  of  earthquake  waves  in  the  interior  of  the 


1908.J 


THE  PHYSICS  OF  THE  EARTH.  261 


earth  has  been  ascertained  to  amount  to  11.3  kilometers  per  second,  the 
compressibility  of  that  region  must  be  31  times  less  than  that  of  quartz, 
that  is,  eight  times  less  than  that  of  solid  steel,  according  to  Voigt.  This 
is  a  figure  of  precisely  that  order  of  magnitude  which  was  to  be  expected. 
We  may  well  believe  that  at  depths  of  more  than  1,000  kilometers  the  com- 
pressibility of  gaseous  iron  sinks  down  to  some  ten  times  less  than  that  of 
steel. 

"The  interior  of  the  earth,  therefore,  with  the  exception  of  a  solid 
crust  about  40  kilometers  thick,  consists  of  a  molten  magma  100  or  200 
kilometers  in  depth  which  shades  continuously  inward  into  a  gaseous  center. 
The  liquids  and  gases  in  the  interior  possess  a  viscosity  and  incompressi- 
bility  such  as  permit  them  to  be  regarded  as  solid  bodies.  From  these, 
however,  they  are  distinguished  in  the  first  place  by  the  fact  that  differentia- 
tions are  possible  to  a  considerable  degree,  the  effects  of  which  may  long 
endure.  In  the  second  place,  long  continued  pressures,  when  acting  on  a 
large  enough  scale,  may  produce  great  deformations.  Further,  the  liquids 
must  possess  the  property  of  great  expansion  on  a  diminution  of  the  high 
pressure,  thereby  readily  becoming  fluid.  The  process  must  thus  differ  but 
little  from  a  normal  melting  with  increase  of  volume,  and  especially  of 
fluidity,  as  well  as  with  absorption  of  heat.  And  yet  the  condition  of  aggre- 
gation is  not  thereby  altered." 

Geikie  remarks  that  the  theory  of  Arrhenius  accords  well  with 

geological  requirements : 

"With  reference  to  the  crust  of  the  earth,  it  meets  the  constantly  re- 
peated objections  of  the  geologists  to  whom  the  existence  of  a  comparatively 
thin  crust  has  always  seemed  an  essential  condition  for  the  production  of 
that  crumpled  and  fractured  structure  which  the  rocks  of  the  land  so  uni- 
versally present.  If  the  solid  crust  of  the  earth  is  allowed  to  be  about  25 
miles  thick,  we  must  conceive  that  in  the  lower  four  fifths  of  its  mass 
the  rocks  are  in  a  condition  of  latent  plasticity.  They  lie  much  beyond  the 
crushing  strength  which  they  exhibit  at  the  surafec.  They  are  not  crushed 
into  powder  as  they  would  be  under  a  similar  strain  above  ground,  but  they 
are  ready  to  yield  to  the  deformations  that  may  arise  consequent  upon  ad- 
justments of  the  gigantic  pressure  to  which  they  are  subjected.  Hence  the 
solid  crust  down  as  far  as  its  structure  has  been  disclosed  abounds  in  proofs 
that  it  has  undergone  colossal  plication  and  fracture,  and  that  higher  por- 
tions of  it  many  square  miles  in  extent  have  been  thrust  bodily  over  each 
other  for  many  miles." 

The  last  view  here  expressed  by  Geikie  as  to  how  the  crust 

becomes  thrust  over  itself  for  many  miles  is  not,  we  think,  well 

founded,  because  it  is  shown  in  this  paper  that  all  this  folding  and 

overlapping  of  the  crust  arises  in  the  trenches  dug  out  in  the  sea 

bottom  by  earthquakes.     This  crumpling  and  overthrusting  of  the 

crust  certainly  would  not  arise  except  for  earthquakes  produced  by 


252  SEE— FURTHER  RESEARCHES  ON  [April  24, 

the  leakage  of  the  oceans,  to  which  mountain  formation  is  due.  Of 
course  the  plasticity  of  this  layer  beneath  the  crust  contributes  to 
the  final  result,  but  the  leakage  of  the  oceans,  with  the  resulting 
earthquakes,  supplies  the  deforming  force. 

§  46.  Views  of  Professor  Suess. — In  the  "  Face  of  the  Earth  " 
(Vol.  I,  p.  107)  we  find  the  following  brief  exposition  of  Professor 
Suess'  views: 

"  The  dislocations  visible  in  the  rocky  crust  of  the  earth  are  the  result 
of  movements  which  are  produced  by  a  decrease  in  the  volume  of  our  planet. 
The  tensions  resulting  from  this  process  show  a  tendency  to  resolve  them- 
selves into  tangential  and  radial  components,  and  thus  into  horizontal  (».  e., 
thrusting  and  folding),  and  into  vertical  (1.  e.,  sinking)  movements.  Dis- 
locations may  therefore  be  divided  into  two  main  groups,  of  which  one  is 
produced  by  the  more  or  less  horizontal,  the  other  by  the  more  or  less 
vertical  relative  displacement  of  larger  or  smaller  portions  of  the  earth's 
crust. 

"There  are  large  areas  in  which  the  first,  and  others  in  which  the 
second  group  predominates,  and  there  are  also  regions  in  which  both  groups 
appear  together,  and  in  which  an  intimate  connection  may  be  recognized 
between  them,  the  resolution  of  the  movements  in  space  having  in  these 
cases  been  less  complete.  This  essential  difference  in  the  movements  of  the 
lithosphere  may  be  clearly  perceived  from  a  comparative  study  of  the  struc- 
ture of  the  Old  World;  nor  has  it  escaped  the  notice  of  American 
geologists. 

"'The  geological  provinces  of  the  Great  Basin,'  remarks  Clarence  King, 
has  suffered  two  different  types  of  dynamic  action:  one  in  which  the  chief 
factor  was  evidently  tangential  compression,  which  resulted  in  contraction 
and  plication,  presumably  in  post -Jurassic  time;  the  other  of  strictly  verti- 
cal action,  presumably  within  the  Tertiary,  in  which  there  are  few  evidences 
or  traces  of  tangential  compression.' 

"Our  colleagues  on  the  other  side  of  the  ocean  have  even  gone  a  great 
deal  further.  After  comparison  of  the  folded  Appalachian  mountains  with 
the  depressed  Basin  Ranges,  Gilbert  had  in  1875  already  suggested  the 
possibility  that  in  the  Appalachians  the  causes  of  movement  were  superficial, 
in  the  ^asin  Ranges  deep-seated.  We  shall  have  an  opportunity,  when  dis- 
cussing the  relation  of  the  Alps  to  their  northern  foreland,  of  determining 
to  what  extent  this  supposition  finds  confirmation  in  Europe.  We  may  how- 
ever state  at  once  that  as  a  rule  it  is  only  the  dislocations  of  the  second 
group  which  are  accompanied  by  volcanic  eruptions." 

§47.  Views  of  Arrhenius. — It  is  well  known  that  this  distin- 
guished Swedish  physicist  holds  that  the  earth's  interior  is  essen- 
tially gaseous  (cf.  §45,  above),  but  under  the  great  pressure  oper- 
ating in  the  globe  made  to  behave  very  nearly  as  a  solid.^    In  his 

*  See  Postscript,  page  274. 


«9o8.] 


THE  PHYSICS  OF  THE  EARTH.  253 


paper  "  Zur  Physik  des  Vulcanismus,"  published  in  1900,  Arrhenius 
points  out  that  in  fluids  at  high  temperature,  where  no  increase  in 
volume  takes  place,  the  internal  friction  of  the  molecules  rises  with 
the  temperature,  so  that  the  viscosity  increases  and  the  'fluidity 
diminishes;  that  a  similar  effect  is  observable  in  both  gases  and 
liquids ;  that  although  gases  have  the  highest  and  solids  the  lowest 
compressibility,  nevertheless  when  a  gas  near  its  critical  tempera- 
ture passes  into  a  liquid,  through  a  trifling  physical  change,  there 
is  practically  no  change  in  the  compressibility.  The  higher  the 
pressure  the  smaller  is  the  compressibility,  and  a  gas  above  the 
critical  temperature  may  be  made  to  acquire  the  properties  of  a 
solid  by  pressure  alone.  Such  a  mass  has  g^eat  density,  small  com- 
pressibility, and  large  viscosity,  so  that  it  has  the  properties  of  a 
solid,  though  really  an  imprisoned  gas. 

At  a  depth  of  40  kilometers  Arrhenius  says  the  temperature  is 
about  1200**  C,  and  the  pressure  about  10,840  atmospheres;  and 
as  these  conditions  would  render  nearly  all  ordinary  minerals  fluid, 
he  concludes  that  below  that  depth  the  matter  is  molten,  in  the 
form  of  a  magma — that  is,  a  viscous  and  nearly  incompressible 
liquid  made  to  act  nearly  as  a  solid  by  pressure. 

At  greater  depths  the  temperature  is  above  the  critical  tempera- 
ture of  every  known  substance,  as  the  pressure  rapidly  increases 
and  the  liquid  magma  becomes  a  gaseous  magma  with  larger  and 
larger  viscosity,  and  smaller  and  smaller  compressibility — in  other 
words,  an  elastic  solid  with  rigidity  increasing  with  the  depth. 

VI.  Abandonment  of  the  Old  Theories  of  the  Physics  of  the 

Earth. 

§  48.  The  Total  Inadequacy  of  the  Old  Theories  to  Account  for 
the  FcMlt  Movements  near  the  Sea,  which  Raise  Vertical  Blocks  and 
Walls  of  Granite^  Thousands  of  Feet  above  the  Water, — ^The  vast 

*  Andesite  is  the  name  used  to  designate  the  kind  of  granitic  rock  found 
in  the  Andes.  Charles  Darwin  showed  that  all  granitic  rocks  are  closely 
related.  In  his  "Text-book  of  Geology,"  edition  of  1903,  book  II,  Part  II, 
^  7»  PP*  230-260,  Sir  Archibald  Geikie  gives  tables  of  the  chemical  compo- 
sitions of  all  these  rocks,  which  show  very  clearly  their  close  relationship. 
When  we  use  the  term  granite  therefore  we  mean  granitic  rock  in  the  wide 
sense. 

PROC.  AMER.  PHIL.  SOC.  XLVII.  1 89  Q,  PRINTED  SEPTEMBER  24,  I908. 


264  SEE— FURTHER   RESEARCHES  ON  (April  24. 

vertical  walls  and  blocks  of  granite  so  often  lifted  thousands  of 
feet  above  the  sea,  with  deep  water  all  around  their  bases,  frequently 
encountered  in  different  parts  of  the  world,  cannot  be  explained 
except- by  the  present  theory.  Thus  along  the  west  coast  of  Chili 
and  Patagonia,  from  Cape  Horn  to  Valparaiso,  in  the  Straits  of 
Magellan,  as  well  as  in  the  ranges  of  the  Andes  further  from  the 
coast,  in  the  Sierras  of  California,  and  elsewhere  these  vertical 
uplifts  are  common.  It  is  obvious  that  they  cannot  possibly  be 
explained  by  the  old  theories  depending  on  the  shrinkage  of  the 
globe.  But  if  lava  is  expelled  from  beneath  the  sea,  owing  to  the 
secular  leakage  of  the  ocean  bottom,  and  the  crust  is  fractured  and 
rent  into  blocks  by  the  earthquake  forces,  some  of  these  blocks  would 
naturally  be  pushed  upward,  leaving  vertical  walls  of  granite  thou- 
sands of  feet  high.  Occasionally  the  blocks  would  be  forced  apart, 
leaving  the  sea  pass  between,  as  so  often  seen  in  Chili,  Patagonia 
and  Tierra  Del  Fuego.  The  Straits  of  Magellan  no  doubt  arose  in 
this  way.  As  already  remarked  in  §  27,  Darwin  describes  similar 
breaks  in  the  Andes  further  north,  through  which  the  sea  once 
flowed,  but  they  are  now  raised  above  the  water.  No  doubt«the 
time  will  come  when  Tierra  Del  Fuego  will  be  joined  solid  to  Pata- 
gonia, by  uplifts  which  will  cause  the  sea  to  withdraw  from  the 
Straits  of  Magellan  and  it  will  become  dry  land,  like  those  ancient 
passages  further  north  mentioned  by  Darwin. 

There  are  many  other  parts  of  the  world  where  similar  phenom- 
ena may  be  seen.  The  origin  of  the  fiords  in  Norway  has  long  been 
a  matter  of  debate.  It  seems  to  be  conceded  that  these  inlets  are 
made  by  mountains  running  into  the  sea,  and  more  or  less  modified 
above  water  by  ice  and  glaciers.  They  are  supposed  to  be  quite 
old,  and  certainly  date  back  of  the  glacial  epochs. 

It  may  no  doubt  be  safely  assumed  that  these  Norwegian  moun- 
tains originated,  like  other  mountains,  by  the  uplift  of  faults,  owing 
to  the  expulsion  of  lava  from  beneath  the  sea.^  Hence  the  precipiti- 
ous  walls  along  the  sea  coast,  with  deep  water  between.    The  blocks 

*  Having  read  the  earlier  papers  of  this  series  with  great  interest,  Pro- 
fessor Schiaparelli  has  kindly  called  my  attention  to  the  trough  in  the  sea 
along  the  Norwegian  coast.  This  confirmation  of  the  theory  by  the  illustrious 
astronomer  of  Milan  is  exceedingly  interesting. 


,9o8.J  THE  PHYSICS  OF  THE  EARTH.  255 

of  the  earth's  crust  were  lifted  vertically  by  the  pushing  of  lava 
beneath  them.  It  is  in  this  way  that  all  such  walls  of  granite  and 
other  towering  rock  are  to  be  explained,  and  the  fact  that  the  sea 
still  encroaches  on  them  shows  how  the  movements  came  about. 
Probably  there  has  been  little  vertical  movement  for  a  long  time 
along  the  coast  of  Norway,  and  subsidence  as  well  as  elevation  may 
have  taken  place,  both  here  and  elsewhere.  Subsidence  is  common 
along  most  sea  coasts,  but  it  does  not  prevail  in  the  long  run,  as 
is  proved  by  Professor  Suess's  work,  showing  a  universal  lowering 
of  the  strand  line  throughout  the  world. 

§  49.  The  Theory  of  Arches  and  Domes  Inapplicable  to  the  Crust 
of  the  Earth,  because  the  Globe  is  not  Shrinking  but  actually  Ex- 
panding.— In  Chamberlin  and  Salisbury's  "  Geology,"  Vol.  I,  p.  583, 
we  find  the  statement  that 

"The  principle  of  the  dome  is  brought  into  play  whenever  an  interior  shell 
shrinks  away,  or  tends  to  shrink  away,  from  an  outer  one  which  does  not 
shrink.  In  this  case  there  is  a  free  outer  surface  and  a  more  or  less  un- 
supported under  surface  towards  which  motion  is  possible.  The  dome  may, 
therefore,  yield  by  crushing  or  by  contortion." 

Owing  to  the  important  part  the  domed  form  of  the  crust  has  played 

in  theories  of  deformation,   these  authors  give  quantitive  results 

calculated  by  Hoskins,  showing  that  such  a  dome  of  continental 

dimensions,  if  unsupported  from  below,  would  sustain  only  1/ 525th 

of  its  o^cn  weight. 

In  his  consideration  of  the  "  Mathematical  Theories  of  the 
Earth'*  (Proc.  Am.  Assoc,  for  Adv.  Sci.,  1889,  p.  49),  Professor 
R.  S.  Woodward  reached  the  analogous  conclusion  that  "If  the 
crust  of  the  earth  were  self-supporting,  its  crushing  strength  would 
have  to  be  about  thirty  times  that  of  the  best  cast  steel,  or  five  hun- 
dred to  one  thousand  times  that  of  granite." 

In  view  of  these  results  it  is  remarkable  that  any  one  should 
have  viewed  the  earth's  crust  as  a  wholly  or  partially  self-support- 
ing dome ;  for  it  could  not  be  supported  even  over  a  very  small 
area.  And  moreover  secular  cooling  is  wholly  inadequate  to  cause 
a  separation  of  the  interior  layers  from  the  crust.  All  that  has  been 
published  on  this  point,  therefore,  is  inapplicable  to  the  earth,  be- 
cause it  rests  on  a  false  hypothesis.  The  supposed  conditions  have 
no  reality. 


266  SEE— FURTHER   RESEARCHES  ON  rAprila4, 

The  earth  is  not  shrinking  and  the  crust  does  not  tend  to  sepa- 
rate itself  from  the  underlayers,  except  where  the  lava  has  been 
expelled  from  beneath  it  by  earthquakes.  The  collapse  of  the  crust 
when  thus  undermined,  however,  shows  that  it  will  not  support  its 
own  weight  even  for  a  short  distance.  Over  such  small  areas  the 
crust  may  be  taken  as  part  of  a  plane,  or  sometimes  as  concave, 
where  subsidence  is  already  at  work,  and  hence  the  theory  of  the 
arch  or  dome  is  scarcely  applicable;  yet  the  observed  collapse  and 
sinking,  even  where  the  area  is  no  larger  than  in  ocean  troughs, 
confirms  the  above  conclusions  regarding  the  total  inability  of  the 
crust  to  support  itself. 

Could  therefore  anything  be  more  absurd  than  to  discuss  the 
stresses  in  the  crust  due  to  the  progress  of  secular  cooling?  Stresses 
arise  only  where  mountain  making  is  in  progress,  and  therefore 
chiefly  near  the  oceans,  but  never  appear  far  inland ;  and  are  wholly 
due  to  the  pressure  arising  from  steam-saturated  rock  and  the  expul- 
sion of  lava  from-  beneath  the  oceans,  or  to  movements  traceable  to 
surface  water  slowly  sinking  into  the  earth.  The  theory  of  arches 
and  domes  therefore  confirms  the  present  theory,  but  this  result  is 
indirect;  and  such  lines  of  thought  did  not  enable  geologists  and 
physicists  to  reach  correct  conceptions  regarding  the  physics  of  the 
earth's  crust. 

§  so.  On  the  Doctrine  that  Earthquake  Movements  depend  on 
Slight  Inequalities  of  Loading,  and  on  the  Abandoned  Theory  that 
the  Earth  is  a  Failing  Structure. — As  the  crust  of  the  earth  is  made 
up  of  solid  rock  and  soil  arising  from  the  disintegration  of  rock  of 
various  kinds,  and  as  this  material  is  elastic  and  yields  under  pres- 
sure, it  naturally  occurred  to  physicists  that  inequalities  of  surface 
loading  deposited  on  adjacent  areas  would  impose  upon  the  under- 
lying crust  unequal  stresses,  and  perhaps  give  rise  to  relative  move- 
ments. Thus  many  physicists,  in  default  of  a  better  theory,  have 
supposed  that  surface  loads,  depending  on  erosion  and  sedimenta- 
tion, tides  and  varying  barometric  pressure,  would  be  adequate  to 
produce -stresses  that  would  cause  readjustment  of  the  surface  strata 
and  perhaps  movements  of  faults  in  earthquakes. 

It  is  undeniable  that  these  varying  loads  do  produce  some  small 
eflFects,  and  very  slight  changes  of  level  may  often  arise  in  this 


i9o8.]  THE  PHYSICS  OF  THE  EARTH.  257 

way.  We  owe  the  establishment  of  these  effects  of  loading  chiefly 
to  the  researches  of  Professor  Sir  G.  H.  Darwin,  whose  labors 
have  so  greatly  advanced  our  knowledge  of  the  physics  of  the  earth. 
They  have  an  extremely  high  importance  in  the  theory  of  bodies 
approximating  elastic  solids.  The  undisturbed  crust  of  the  globe 
fulfills  these  conditions  quite  perfectly. 

But  to  suppose  that  any  of  these  small  surface  effects  could  give 
rise  to  world-shaking  earthquakes  which  would  shake  down  cities, 
raise  sea  coasts,  and  uplift  mountains  and  islands  in  the  sea,  is  too 
severe  a  test  of  credulity  to  be  entertained.  The  class  of  minute 
movements,  due  to  surface  yielding  under  varying  loads  depending 
on  sediments,  tides  and  meteorological  causes,  and  the  class  of  great 
movements,  due  to  the  expulsion  of  lava  from  imder  the  bed  of  the 
sea,  are  quite  distinct.  One  class  of  these  phenomena  is  micro- 
seismic,  the  other  magaseismic.  Previous  investigators  have  gen- 
erally confounded  the  two  classes  of  phenomena,  and  hence  they 
have  been  unable  to  recognize  the  true  cause  of  earthquakes  and 
mountain  formation.  For  that  reason  it  was  necessary  to  restrict 
our  investigation  to  the  great  disturbances,  in  the  first  search  for  the 
cause  of  the  great  movements  of  the  earth's  crust. 

We  repeat  that  both  classes  of  phenomena  are  important  in  a 
complete  theory  of  the  physics  of  the  earth ;  but  the  small  yieldings 
of  microscopic  dimensions  must  be  kept  distinct  from  the  great 
movements  which  have  shaped  the  surface  of  the  globe.  Many  of 
the  small  effects  depend  on  the  greater  movements  of  the  earth, 
while  few  of  the  great  movements  are  influenced  by  surface  forces — 
indeed  none  at  all,  except  where  accumulation  of  subterranean 
stresses  has  already  rendered  the  conditions  highly  unstable.  In 
this  latter  case  small  surface  forces  may  occasionally  accelerate  the 
outbreak  of  an  earthquake,  just  as  a  spark  discharges  a  loaded  gun, 
or  a  shock  explodes  a  charge  of  dynamite. 

*  On  a  par  with  the  theory  that  slight  inequalities  of  surface  load- 
ing produce  earthquakes  is  another  equally  untenable  view  that 
the  earth  is  a  failing  structure.  Such  a  doctrine  might  have  been 
entertained  a  quarter  of  a  century  ago,  when  the  theory  of  secular 
cooling  was  generally  accepted,  but  to-day  such  a  view  is  anti- 
quated and  utterly  indefensible.     Owing  to  the  demonstrated  de- 


258  SEE-FURTHER    RESEARCHES  ON  [April  *4, 

pendence  of  mountain  making  upon  the  sea  the  earth  emphatically 
is  not  a  failing  structure.  So  far  from  failing  "by  collapse,  our 
planet  seems  to  be  expanding  from  lo  to  lOO  faster  than  it  con- 
tracts from  loss  of  heat.  Thus  have  arisen  all  the  highest  moun- 
tains and  plateaus  of  the  globe.  These  great  uplifts  invariably 
face  the  deepest  oceans,  from  which  the  expulsion  of  lava  has  mainly 
proceeded.  Such  antiquated  doctrines  as  that  the  earth  is  a  fail- 
ing structure  are  now  absolutely  without  excuse,  and  practically 
abandoned,  and  the  sooner  they  disappear  from  scientific  literature 
the  better  for  sound  knowledge  of  the  physics  of  the  earth. 

§  51.  Changes  of  the  Force  of  Gravity  in  Regions  Affected  by 
the  Movement  of  Lava  Beneath  the  Crust, — In  view  of  the  demon- 
strated movement  of  lava  streams  beneath  the  crust  of  the  globe, 
it  follows  that  such  bodily  displacement  of  matter  but  a  short  dis- 
tance below  the  surface  may  modify  sensibly  the  observed  intensity 
of  gravity.  A  region  which  is  being  undermined  will  have  the 
intensity  of  gravity  decreased,  and  a  region  which  is  being  filled 
up  will  have  the  attraction  increased.  And  not  only  will  the  in- 
tensity vary,  but  also  the  direction  of  the  vertical,  according  to  the 
movements  which  occur  beneath  the  crust.  And  these  effects  may 
be  large  enough  to  become  sensible  to  very  refined  observation. 

It  is  in  this  way  that  the  anomalies  of  gravity  in  the  neigh- 
borhood of  mountains  have  arisen  in  the  process  of  mountain  form- 
ation. And  in  regions  where  the  expulsion  of  lava  is  still  in 
progress,  both  the  direction  and  intensity  of  gravity  are  subject  to 
change  by  earthquakes.  Thus  in  the  region  of  the  Aleutian  Islands, 
the  east  coast  of  Japan,  and  many  other  places,  such  as  the  west 
coast  of  South  America,  the  direction  and  intensity  of  gravity  is  cer- 
tainly subject  to  change  by  seismic  disturbances. 

As  the  crust  of  the  globe  often  suffers  horizontal  and  vertical 
movement  during  the  greatest  earthquakes,  the  altitude  and  azimuth 
of  places  are  also  subject  to  change;  and  exact  geodetic  triangulS- 
tion  remains  valid  only  for  the  interval  between  great  earthquakes. 
Even  then  there  may  be  a  very  slow  and  gradual  settlement  owing 
to  plastic  yielding  of  the  crust  and  especially  of  the  substratum 
beneath.  Thus  after  earthquakes  such  as  occur  in  Peru  and  Chili, 
Japan  and  Alaska,  gravity  and  geodetic  determinations  need  repeti- 


x9o8.]  THE  PHYSICS  OF  THE  EARTH.  259 

tion,  as  was  done  in  California  after  the  great  earthquake  of  April 
^8,  1906.  And  as  the  disturbance  may  alter  the  direction  and  in- 
tensity of  local  gravity,  this  possibility  must  be  taken  account  of  in 
the  repetition  of  the  observations.  In  order  to  be  entirely  rigor- 
ous the  equations  connecting  the  triang^Iation  should  include  unde- 
termined multipliers  to  take  account  of  possible  variations  in  the 
local  attraction  at  each  point.  If  with  this  general  condition  im- 
posed, the  triangulation  before  and  after  the  earthquake  comes  out 
rigorously  the  same,  within  the  limits  of  errors  of  observation,  it  may 
be  supposed  that  the  surface  effects  of  the  disturbance  are  insen- 
sible; otherwise  the  difference  must  be  attributed  to  disturbances 
due  to  the  earthquake. 

With  the  refinement  now  possible  in  geodesy,  it  is  not  to  be 
doubted  that  these  effects  will  occasionally  prove  to  be  sensible  to 
observation.  The  ,  great  earthquake  in  Assam-Bengal  gave  rise 
to  horizontal  movements  of  the  order  of  20  or  30  feet,  which  may 
affect  the  latitude  by  0^.2  or  o''.3,  and  are  thus  within  the  limits  of 
astronomical  measurement.  But  apparent  changes  in  latitude  may 
result  from  change  in  the  direction  of  gravity  as  well  as  from  actual 
displacements  of  the  crust,  and  both  possibilities  need  to  be  taken 
into  accoimt. 

§  52.  The  Necessity  of  Further  Study  of  the  Contours  and 
Movements  of  the  Sea  Bottom. — In  view  of  the  results  brought  out 
in  this  paper  and  those  which  have  preceded  it,  but  especially  that 
on  "  The  New  Theory  ot  Earthquakes  and  Mountain  Formation  as 
Illustrated  by  Processes  now  at  Work  in  the  Depths  of  the  Sea," 
it  is  scarcely  necessary  to  point  out  the  extreme  importance  of 
further  study  of  the  contours  and  movements  of  the  sea  bottom. 
Our  present  maps  of  the  ocean  depths  are  very  incomplete,  although 
they  afford  a  good  general  idea  of  the  sea  basins.  But  one  can 
scarcely  doubt  that  more  exact  surveys  would  bring  to  light  addi- 
tional mountain  ranges  and  plateaus  in  regions  heretofore  but  slightly 
explored;  moreover  certain  places  in  the  sea  bottom  would  be 
found  to  be  covered  with  a  great  variety  of  peaks  or  submerged 
islands  which  do  not  reach  the  surface. 

Where  the  water  is  deep  the  exact  survey  of  the  bottom  pre- 
sents considerable  difficulty.     As  movements  arising   from  earth- 


260  SEE— FURTHER   RESEARCHES  ON  [April  24. 

quakes  are  extremely  small  in  comparison  with  the  depth  of  the 
sea,  it  would  perhaps  be  very  difficult  to  detect  resulting  changes, 
of  the  sea  bottom,  except  in  cases  where  sinking  takes  place,  and 
the  drop  is  large.  In  some  cases  of  actual  measurement  in  the 
laying  of  cables  the  sinking  has  been  found  to  be  hundreds  of 
fathoms,  which  would  be  very  easily  recognized  if  the  exact  place 
of  former  soundings  could  be  found.  But  as  the  changes  of  level 
in  the  sea  bottom  are  fully  as  capricious  as  on  land,  we  see  that 
regions  where  mountain  formation  is  in  progress  would  present 
extreme  complexity;  and  unless  the  place  were  very  accurately 
known,  one  could  not  be  sure  that  two  soundings  were  over  the 
same  spot.  This  difficulty  would  be  less  near  known  islands  than 
in  the  open  sea,  but  it  would  be  considerable  in  all  places  where 
the  ship  is  at  the  mercy  of  the  winds  and  currents. 

Under  the  circumstances  it  is  clear  that  great  natural  difficulty 
would  arise  in  the  exact  Hydrographic  survey  of  the  deep  sea,  and 
an  economic  difficulty  would  be  added,  on  the  ground  that  such 
surveys  are  not  required  in  practical  navigation.  Yet  the  laying 
and  repair  of  cables  would  necessitate  fairly  accurate  knowledge  of 
the  depths,  and  we  may  hope,  in  spite  of  the  growth  of  the  wire- 
less telegraph,  that  our  ocean  surveys  are  still  in  the  infancy  of 
what  they  will  be  in  another  half  century. 

Where  trenches  are  being  dug  out  by  earthquakes  there  will 
be  the  double  incentive  to  ascertain  the  stage  of  the  process  and 
the  rapidity  and  location  of  the  changes.  These  considerations  may 
contribute  to  our  knowledge  of  particular  regions;  and,  after  all, 
the  changes  in  the  larger  regions  of  the  ocean  bottom  are  small. 

When  the  regions  in  which  trenches  are  forming  are  once 
clearly  recognized,  attention  will  naturally  be  centered  upon  them, 
to  the  neglect  of  less  disturbed  areas.  The  most  interesting  re- 
gions, from  a  seismological  point  of  view,  are  those  in  which 
islands  are  being  uplifted  and  the  sea  bottom  sinking,  as  near  the 
Aleutian,  Kurile  and  Japanese  islands,  the  Antandes,  and  along  the 
west  coast  of  South  America.  But  it  may  also  be  hoped  that 
the  changes  in  depth  near  individual  islands,  such  as  Guam  and 
Martinique,  will  not  be  overlooked.  Here  the  subsidence  of  the 
bottom  often  takes  the  form  of  a  hole  rather  than  of  a  trench.    Yet 


i9o8.] 


THE  PHYSICS  OF  THE  EARTH.  261 


in  time  the  movements  may  give  rise  to  neighboring  islands.  All 
of  these  considerations  show  the  value  of  accurate  knowledge  of  the 
sea  bottom  at  this  epoch. 

§  53.  Greatness  of  the  Forces  which  Uplift  and  fold  the  Earth's 
Crust. — The  tremendous  power  of  earthquake  and  volcanic  forces 
has  been  proverbial  from  the  earliest  ages  of  history,  and  finds 
expression  also  in  the  universal  terror  thus  excited  among  all  liv- 
ing beings.  This  extreme  terror  is  only  too  well  justified  by  the 
vast  extent  of  the  ruin  too  often  wrought  in  different  parts  of  the 
world.  But  probably  only  those  who  have  witnessed  a  great  earth- 
quake can  adequately  appreciate  the  awful  character  of  the  com- 
motion, and  the  gigantic  forces  which  must  underly  it.  This  is 
shown  also  by  the  many  published  attempts  to  belittle  the  signifi- 
cance of  earthquake  disasters. 

Some  writers  of  eminent  mathematical  learning,  but  apparently 
lacking  in  grasp  of  the  larger  physical  phenomena,  have  ascribed 
earthquakes  to  inequalities  of  loading,  changes  of  barometric  pres- 
sure, etc.,  and  have  with  strange  and  almost  marvelous  credulity 
believed  that  the  settlements  of  the  earth  thus  arising  would  shake 
down  cities  and  devastate  whole  countries.  How  these  learned 
authorities  imagined  that  small  subsidences  under  the  steady  action 
of  these  infinitesimal  forces  could  bring  about  such  long  con- 
tinued shaking  and  proportionately  great  havoc  is  difficult  to  un- 
derstand. If  the  forces  are  so  small,  and  act  so  slowly,  is  it 
conceivable  that  the  yielding  could  be  anything  else  than  gradual 
and  insensible?  Such  minute  settlements  evidently  would  be  like 
those  now  experienced  in  dry  inland  regions  free  from  real  earth- 
quakes. 

The  titanic  nature  of  the  forces  which  have  uplifted  islands, 
mountains,  plateaus  and  continents,  can  scarcely  be  realized;  yet 
even  the  ancients  grasped  it  to  some  extent  when  they  described 
the  whole  region  between  Naples  and  Sicily  as  underlaid  by  a  giant, 
whose  movements  disturbed  the  intervening  sea  bottom.  In  his 
account  of  the  Chilean  earthquake  of  1835,  Charles  Darwin  showed 
that  the  entire  region  from  the  island  of  San  Fernandez  to  the 
Andes,  about  450  miles  across,  had  been  moved  together  by  under- 
lying forces.     "  There  was  undoubtedly  a  connection  between  the 


262  SEE-FURTHER  RESEARCHES  ON  [April  24, 

volcanic  forces  acting  under  this  island,  and  under  the  continent,  as 
was  shown  during  the  earthquake  of  1835,"  says  the  great  naturalist. 

As  such  views  have  been  carefully  set  forth  by  the  greatest  of 
original  investigators,  from  Aristotle  to  Darwin,  it  is  remarkable 
to  witness  the  puny  efforts  which  have  been  made  to  belittle  these 
forces.  A  gentleman  holding  a  university  position,  in  a  public 
address  at  Boston,  recently  likened  the  shock  of  an  earthquake  to 
the  jar  experienced  by  an  insect  attached  to  a  reed  which  was  bent 
till  it  snapped.  According  to  this  authority  the  earthquakes  are 
due  to  the  snapping  of  the  rock  of  the  earth's  crust  in  the  bending 
produced  by  secular  cooling.  Is  it  necessary  to  point  out  the  mis- 
leading character  of  the  comparison  made,  and  this  lecturer's  utter 
inability  to  grasp  the  phenomena  of  nature? 

An  equally  common  fallacy  is  to  ascribe  these  tremendous  dis- 
turbances to  inequalities  of  surface  loading,  due  to  geological  and 
meterological  causes.  Such  views  seem  the  more  surprising,  be- 
cause formerly  they  have  proceeded  from  physicists  of  eminent 
learning.  But  at  least  partial  excuse  may  be  found  in  the  universal 
acceptance  of  the  theory  of  secular  cooling  heretofore,  and  in  the 
proved  rigidity  of  the  globe,  which  naturally  led  to  the  supposition 
that  the  crust  was  adjusting  itself  to  the  shrinking  sphere. 

Before  the  development  of  the  theory  of  ocean  leakage  no  ade- 
quate theory  presented  itself  to  investigators,  who  had  unfortu- 
nately not  discriminated  between  the  great  and  small  earthquakes. 
With  a  false  premise  and  such  an  indiscriminate  mixture  of  phe- 
nomena, real  progress  was  difficult,  if  not  impossible. 

§  54.  Darwin's  Remarks  on  the  Forces  which  Uplift  Continents. 
— In  the  extract  quoted  from  Professor  Suess,  §  39,  allusion  has 
already  been  made  to  Charles  Darwin's  attempt  to  explain  the  origin 
of  mountains  by  the  direct  observation  of  nature.  His  paper  "  On 
the  Connection  of  Certain  Volcanic  Phenomena  in  South  America 
and  the  Formation  of  Mountain  Chains  and  Volcanoes  as  the  Effect 
of  the  Same  Power  by  which  Continents  are  Elevated"  {Transac- 
tions of  the  Geological  Society^  Vol.  V,  1838,  pp.  601-631)  led 
Darwin  to  the  conclusion: 

"  That  the  form  of  the  fluid  surface  of  the  nucleus  of  the  earth  is  sub- 
ject to  some  change,  the  cause  of  which  is  entirely  unknown  and  the  effect 
of  xvhich  is  slow,  intermittent,  but  irresistible/' 


,9o8.]  THE   PHYSICS  OF  THE   EARTH.  263 

Again,  in  the  "  Voyage  of  the  NaturaHst/*  Chapter  XIV,  he 
adds : 

"  The  forces  which  slowly  and  by  little  starts  uplift  continents,  and  those 
which  at  successive  periods  pour  forth  volcanic  matter  from  open  orifices, 
are  identical.'* 

It  is  unnecessary  to  dwell  on  the  irresistible  power  which  the  great 
naturalist  correctly  abscribed  to  volcanic  and  earthquake  forces.  It 
is  of  more  interest  to  notice  that  he  declared  them  to  be  identical 
with  those  which  uplift  continents.  The  same  result  is  reached  in 
the  present  paper,  about  three  quarters  of  a  century  later,  and  the 
proof  of  the  proposition  now  seems  overwhelming. 

If  Darwin  had  known  the  cause  of  seismic  sea  waves,  and  had 
seen  how  trenches  are  dug  out  in  the  sea  bottom  by  the  expulsion 
of  lava  from  beneath  the  sea  under  the  land,  can  anyone  doubt  that 
he  would  have  discovered  and  proved  the  leakage  of  the  oceans, 
and  developed  the  correct  theory  of  mountain  formation? 

§  55.  On  the  Oscillatory  Movements  of  the  Crust  Shown  in  the 
Coal  Measures, — In  view  of  the  results  established  in  this  paper  we 
need  not  dwell  on  the  coal  measures,  and  other  evidences  of  the 
oscillation  of  the  earth's  crust.  It  suffices  to  say  that  these  oscilla- 
tions actually  took  place,  as  geologists  have  long  believed.  The 
coal  fields  in  Pennsylvania  were  formed  by  vegetation  growing 
rapidly  and  with  great  luxuriance  over  areas  near  the  sea  level 
which  were  again  and  again  elevated  and  as  often  depressed  by 
earthquakes.  When  the  land  was  under  the  sea  the  vegetation  died 
out,  and  mud  and  shale  were  deposited;  when  the  area  was  again 
upraised  another  layer  of  vegetation  was  produced,  and  sometimes 
it  was  deposited  by  floods,  currents,  and  drifting  where  it  had  not 
grown.  This  was  during  the  Carboniferous  Age,  and  while  all  the 
land  was  near  the  level  of  the  ocean. 

The  details  of  such  inquiries  must  be  left  to  geologists  and 
paleontologists,  who  study  the  flora  and  fauna  of  past  ages.  Our 
aim  in  these  papers  has  been  to  give  a  firm  basis  for  legitimate  study 
and  speculation,  without  which  the  phenomena  of  nature  remain 
unintelligible.  The  progress  of  the  sciences  of  the  earth  requires 
two  conditions :  first,  true  physical  causes ;  and  second,  the  intelli- 
gent and  consistent  application  of  these  causes  to  the  explanation 


264  SEE— FURTHER   RESEARCHES  ON  [April  a4. 

of  the  phenomena,  both  of  the  animate  and  inanimate  world.  The 
physicist  must  content  himself  with  showing  the  mechanical  causes 
at  work  and  their  mode  of  operation,  while  the  geologist  and  paleon- 
tologist may  deal  with  the  evidences  of  life  under  these  known 
conditions. 

§  56.  The  Equilibrium  of  the  Earth  between  the  Land  and  Water 
Hemispheres  Explained  by  the  Intumescence  of  the  Land  Arising 
from  the  Expulsion  of  Porous  Lava  from  under  the  Bed  of  the  Sea. 
— The  remarkable  equilibrium  preserved  by  the  earth  between  the 
land  and  water  hemispheres  has  long  been  a  matter  of  speculation 
among  philosophers.  Sir  John  Herschel  justly  remarked  that  the 
high  altitude  of  the  continents  in  the  land  hemisphere  would  be 
most  easily  accounted  for  by  an  intumescence  of  the  land.  Pratt 
has  since  treated  the  question  in  a  convincing  manner,  and  shown 
that  the  solid  parts  of  the  earth's  crust  beneath  the  water  hemisphere, 
with  pole  in  New  Zealand,  must  be  denser  than  in  the  correspond- 
ing parts  on  the  opposite  side,  otherwise  the  water  would  flow  away 
towards  the  land  hemisphere  and  tend  to  submerge  it  more  com- 
pletely. (Cf.  "  Figure  of  the  Earth,"  3d  edition,  pp.  159-160.) 
Hence  he  concludes  that 

"There  must  therefore  be  some  excess  of  matter  in  the  solid  parts  of  the 
earth  between  the  Pacific  ocean,  and  the  earth's  center  which  retains  the 
water  in  its  place." 

When  Pratt  wrote  this  forty  years  ago  there  was  no  suspicion  of 
an  intumescent  layer  beneath  the  land  due  to  the  expulsion  of  porous 
lava  from  beneath  the  bed  of  the  sea,  and  accordingly  he  added 
that 

"This  effect  may  be  produced  in  an  infinite  variety  of  ways;  and  therefore, 
without  data,  it  is  useless  to  speculate  regarding  the  arrangement  of  matter 
which  actually  exists  in  the  solid  parts  below." 

Now,  however,  it  is  proved  that  the  plateaus  and  continents  have 
been  uplifted  by  intumescent  matter  expelled  from  under  the  sea; 
and  consequently  we  have  data  for  speculating  on  how  the  observed 
effect  is  produced. 

It  is  clear  that  all  the  great  plateaus  of  the  globe  and  even  the 
continents  themselves  are  underlaid  by  material  lighter  than  the 
average  of  the  earth's  crust.     Naturally  the  effects  are  greatest 


x9o8.]  THE  PHYSICS  OF  THE  EARTH.  265 

where  the  plateaus  are  highest,  as  in  Himalayas  and  Tibet,  where 
the  deficiency  in  the  attraction  of  these  elevated  masses  long  ago 
attracted  attention.  In  his  "  Account  of  the  Operations  of  the 
Great  Trigonometric  Survey  of  India,"  Calcutta,  1879,  General 
J.  T.  Walker  says: 

"  There  appears  to  be  no  escape  form  the  conclusion  that  there  is  a 
more  or  less  marked  negative  variation  of  gravity  over  the  whole  of  the 
Indian  continent,  and  that  the  magnitude  of  this  variation  is  somehow  con- 
nected with  the  height. 

"  Pratt's  calculations  had  reference  only  to  the  visible  mountain  and 
oceanic  masses  and  their  attractive  influences — ^the  former  positive,  the  latter 
negative — in  a  horizontal  direction;  he  had  no  data  for  investigating  the 
density  of  the  crust  of  the  earth  below  either  the  mountains  on  the  one 
hand,  or  the  bed  of  ocean  on  the  other.  The  pendulum  observations  fur- 
nished the  first  direct  measures  of  the  vertical  forces  of  gravity  in  different 
localities  which  were  obtained,  and  these  measures  revealed  two  broad  facts 
regarding  the  disposition  of  the  invisible  matter  below;  first,  that  the  force 
of  gravity  diminishes  as  the  mountains  are  approached,  and  is  very  much 
less  on  the  summit  of  the  highly  elevated  Himalayan  table-lands  than  can 
be  accounted  for  otherwise  than  by  a  deficiency  of  matter  below;  secondly, 
that  it  increases  as  the  ocean  is  approached,  and  is  greater  on  islands  than 
can  be  accounted  for  otherwise  than  by  an  excess  of  matter  below.  As- 
suming gravity  to  be  normal  (in  amount)  on  coast  lines,  the  mean  observed 
increase  at  the  islands  stations  was  such  as  to  cause  a  seconds'  pendulum  to 
gain  three  seconds  daily,  and  the  mean  observed  decrease  in  the  interior  of 
the  continent  would  have  caused  the  pendulum  to  lose  25^  seconds  daily  at 
stations  averaging  i^zoo  feet  above  the  sea  level,  5  seconds  at  3,800  feet, 
and  about  22  seconds  at  15400  feet — the  highest  elevation  reached —  in  excess 
of  the  normal  loss  of  rate  due  to  the  height  above  the  sea." 

The  facts  here  mentioned  by  General  Walker  are  recognized  in 
geodesy  as  applying  in  different  degrees  to  all  the  elevated  table- 
lands and  mountainous  regions  of  the  globe.  The  physical  cause 
of  this  deficiency  in  attraction  is  now  established  beyond  all  doubt, 
and  the  intumescence  of  the  land,  first  suggested  by  Sir  John 
Herschel,  is  shown  to  have  arisen  from  the  expulsion  of  lava  from 
beneath  the  sea.  Thus  arises  the  physical  condition  which 
secures  the  equilibrium  of  the  earth  between  the  land  and  water 
hemispheres.  This  must  be  regarded  as  not  the  least  remarkable 
among  several  interesting  results  on  the  physics  of  the  earth  deduced 
from  the  principle  of  the  secular  leakage  of  the  oceans.  Earth- 
quakes, volcanoes,  mountain  formation,  the  uplift  of  islands,  plat- 


SEE— FURTHER    RESEARCHES  ON  [April  .<, 

'i'B  ' 


U  *    M 


■|-  gi 


I  ill 


I  §  J I 
s=  i  E  i 


I908.I 


THE   PHYSICS  OF  THE  EARTH. 


267 


eaus,  and  continents,  seismic  sea  waves,  trenches  and  holes  in  the 
bottom  of  the  sea,  the  feeble  attraction  of  mountains,  and  plateaus, 
the  equilibrium  of  the  globe  between  the  land  and  water  hemispheres, 
are  all  closely  related  and  dependent  upon  a  single  physical  cause. 


Fig.  i6.    Water  Hemisphere,  which  has  the  World  Ridge  around  it,  drawn 

by  W.  R.  Smith,  of  Mare  Island. 


In  view  of  the  order  and  harmony  thus  established  among  these 
varied  phenomena,  who  will  not  concur  in  the  view  of  the  great 
Newton  that  "  Nature  is  pleased  with  simplicity  and  affects  not  the 
pomp  of  superfluous  causes  "  ? 


268  SEE— FURTHER   RESEARCHES  ON  [April  24, 

Conclusions. 

Some  of  the  chief  conclusions  reached  in  this  and  the  preceding 
papers  on  the  physics  of  the  earth  may  be  briefly  summarized  as 
follows : 

1.  The  theory  of  the  secular  leakage  of  the  oceans  explains  satis- 
factorily six  great  classes  of  phenomena,  not  heretofore  closely  asso- 
ciated, namely:  (i)  Earthquakes,  (2)  volcanoes,  (3)  mountain  for- 
mation, (4)  the  formation  of  islands,  plateaus  and  continents,  (5) 
seismic  sea  waves,  (6)  the  feeble  attraction  of  mountains  and 
plateaus  long  noticed  in  geodesy. 

2.  And  the  theory  not  only  explains  the  leading  facts  of  each 
class  of  phenomena  separately,  but  also  in  relation  to  all  the  other 
classes  of  phenomena;  and  this  harmonious  mutual  relationship  of 
all  the  phenomena  proves  the  theory  to  rest  on  a  true  physical  cause. 

3.  A  vera  causa,  once  established,  should  not  only  explain  all 
the  phenomena,  and  all  the  relations,  but  also  exclude  the  considera- 
tion of  other  possible  causes,  by  necessary  and  sufficient  conditions. 
This  alone  ensures  the  entire  validity  of  the  reasoning,  and  the  pres- 
ent theory  meets  this  severe  test  perfectly. 

4.  We  have  traced  the  details  of  the  processes  involved  in  moun- 
tain fomlation,  and  have  exhibited  illustrations  of  its  working  by 
processes  now  observed  in  the  depths  of  the  sea.  All  stages  of 
mountain  formation  are  thus  brought  out,  and  they  are  all  shown 
to  be  consistent  with  this  simple  theory,  which  explains  the  princi- 
pal phenomena  of  the  earth's  crust. 

5.  This  theory  explains  the  distribution  of  mountains  about  the 
continents,  their  great  height  which  the  contraction  theory  cannot 
account  for;  the  formation  of  parallel  ridges  by  the  uplift  of  the 
side  of  the  trough  nearest  the  sea,  when  the  bottom  has  so  far  sub- 
sided that  the  folding  up  of  the  nearer  side  becomes  the  path  of 
least  resistance  in  the  expulsion  of  molten  rocks  from  under  the 
sea. 

6.  Several  successive  troughs  are  often  thus  dug  out,  with  ridges 
forced  up  between  them ;  and  when  the  whole  is  raised  above  the 
water  we  have  a  series  of  parallel  ranges,  such  as  the  Allegheny, 
Tuscarora  and  Blue  Ridge  Mountains  in  Pennsylvania  and  Vir- 


i9o8.]  THE  PHYSICS  OF  THE  EARTH.  269 

ginia.    Heretofore  these  vast  billows  of  the  earth's  crust  have  been 
utterly  bewildering  to  the  naturalist. 

7.  When  several  such  trenches  have  been  dug  out,  and  the  ex- 
pulsion of  lava  is  from  both  sides,  as  happens  when  the  sea  is  thus 
distributed,  the  ridges  may  finally  be  forced  up  and  so  crowded 
together  from  both  sides  that  overturned  dips  and  inverted  strata 
are  produced,  as  in  the  Swiss  Alps.  No  previous  theory  has  been 
adequate  to  account  for  this  amazing  phenomenon,  the  explanation 
of  which  is  thus  seen  to  be  exceedingly  simple.  This  test  may  be 
justly  considered  the  experitnentum  cruets  of  the  theories  of  moun- 

,    tain  formation. 

8.  The  Andes  in  South  America  are  nothing  but  a  vast  wall  or 
embankment  erected  by  the  Pacific  Ocean,  through  the  expulsion 
of  lava,  along  its  border.  Hence  the  persistence  of  the  earthquake 
belt  and  seismic  sea  waves  along  this  coast. 

9.  This  embankment  includes  not  only  the  peaks  and  chains  of 
mountains,  large  and  small,  in  the  Eastern  and  Western  Cordillera, 
but  also  the  intervening  plateaus,  such  as  those  of  Quito,  Caxa- 
marca,  Cuzco,  and  Titicaca. 

ID.  The  molten  rock  expelled  from  under  the  sea  is  lighter  than 
average  material  of  the  layer  below  the  earth's  crust,  and  when  the 
included  vapor  of  steam  is  allowed  to  expand,  as  in  volcanoes, 
pumice  is  formed,  and  often  blow  out  in  vast  quantities.  Pumice 
of  various  degrees  of  density  underlies  the  mountain  chains,  and 
some  of  it  is  blown  out  of  those  mountains  which  become  volcanoes. 

11.  The  way  in  which  these  plateaus  are  interwoven  with  the 
Andes  mountains  shows  that  the  whole  embankment  is  due  to  the 
continued  action  of  one  common  cause.  And  since  the  mountains 
were  uplifted  by  the  expulsion  of  lava  from  under  the  sea,  as  proved 
by  the  uplifting  of  the  land  in  earthquakes  and  the  sinking  of  the 
sea  bottom,  indicated  by  the  accompanying  seismic  sea  waves,  it 
follows  that  the  plateaus  also  are  underlaid  by  matter  lighter  than 
the  average,  which  has  been  expelled  from  under  the  ocean. 

12.  The  total  quantity  of  matter  thus  expelled  from  beneath  the 
ocean  is  very  large,  but  it  is  the  result  of  an  infinite  number  of 
earthquakes  and  seismic  sea  waves  during  past  geological  ages. 
This  circumstance  affords  us  an  idea  of  the  immense  age  of  the 

PROC.  AMER.  PHIL.  SOC.,  XLVII.  189  R,  PRINTED  SEPTEMBER  25,  I908. 


270  SEE— FURTHER  RESEARCHES  ON  [April  24, 

Andes  Mountains,  which  are  the  youngest  of  the  great  mountain 
systems  of  the  globe. 

13.  THe  terrible  fracturing  of  the  crust  in  the  sharp  folding 
involved  in  the  formation  of  the  Andes  enabled  a  vast  number  of 
volcanoes  to  break  out,  and  about  one  hundred  and  five  have  been 
active  within  historical  times. 

14.  The  formation  and  activity  of  the  volcanoes  in  the  Aleutian 
and  Japanese  Islands  is  similar  to  those  in  the  Andes,  and  represent 
conditions  suitable  to  the  maximum  development  of  volcanic  activ- 
ity. These  are  sharp  folds  of  the  crust  near  a  deep  sea  from  which 
the  expulsion  of  lava  is  rapid  and  violent. 

15.  The  connection  of  earthquakes  with  volcanoes  and  of  both 
phenomena  with  the  sea  is  clearly  established  by  the  geographical 
distribution  and  by  the  vapor  of  steam  emitted  by  volcanoes.  The 
nature  of  the  underlying  material  is  shown  by  the  ashes,  cinders, 
pumice  arid  lava  forced  out  by  the  accumulating  subterranean  steam 
pressure. 

16.  Earthquakes,  however,  are  the  more  general,  volcanoes  the 
'more  special  phenomena.    The  mountains  are  formed  by  the  sea, 

but  only  a  few  of  the  peaks  break  out  into  volcanoes.  No  volcano 
long  remains  active  very  far  from  the  ocean  or  other  large  body  of 
water,  because  as  the  lava  hardens  in  the  throat  of  the  volcano  the 
supply  of  steam  is  inadequate  to  maintain  activity. 

17.  If  we  consider  the  innumerable  islands  in  the  sea,  it  is  evi- 
dent that  they  too  have  been  uplifted  by  earthquakes.  Sometimes 
the  sea  bottom  near  them  has  been  undermined  in  the  process  of 
uplifting,  and  afterwards  sunk  down,  making  an  adjacent  hole  in 
the  bottom,  and  producing  seismic  sea  waves  of  the  first  class,  as 
in  mountain  formation  where  trenches  are  being  dug  out  near  the 
continents. 

18.  Seismic  sea  waves  of  the  second  class  are  produced  by  the 
uplift  of  the  sea  bottom,  into  ridges,  or  submarine  plateaus  and 
islands.  In  such  cases  the  water  rises  suddenly  without  previously 
withdrawing  from  the  shore. 

19.  But  seismic  sea  waves  of  the  first  class  due  ta  the  sinking 
of  the  sea  bottom,  after  it  is  undermined  by  the  expulsion  of  lava, 
are  the  most  important  and  most  celebrated.    The  waves  at  Helike, 


«9o8.] 


THE  PHYSICS  OF  THE  EARTH.  271 


373  B.  C;  Callao,  1746;  Lisbon,  1755;  Arica,  1868;  Iquique,  1877; 
Japan,  1896,  were  all  of  this  class. 

20.  We  may  pass  directly  from  the  Andes  to  the  Himalayas,  and 
from  the  high  plateaus  of  South  America  to  those  of  Asia.  Just 
as  the  plateaus  from  Quito  to  Titicaca  were  formed  by  the  expul- 
sion of  matter  from  under  the  Pacific,  so  also  those  of  Tibet  and 
Iran  are  due  mainly  to  the  expulsion  of  lava  from  beneath  the 
Indian  and  Pacific  Oceans. 

21.  In  the  case  of  the  plateau  of  Tibet  the  resulting  uplift  is 
partly  due  to  the  combined  action  of  the  Pacific,  which  thus  folded 
the  ranges  to  the  East.  With  two  oceans  so  large  and  deep  as  the 
Indian  and  Pacific  cooperating  in  this  uplift,  i^  is  no  wonder  that 
the  maximum  effect  was  produced  and  that  Tibet  became  the  highest 
plateau  in  the  world. 

22.  The  Himalayas  are  higher  and  further  from  the  sea  than  the 
Andes,  but  the  earthquake  belt  at  the  base  still  persists  in  both  cases, 
and  the  configuration  in  regard  to  the  sea  shows  that  the  causes  at 
work  to  produce  these  mighty  uplifts  were  absolutely  similar.  And 
if  the  mountains  are  due  to  the  same  cause,  the  plateaus  are  also. 

23.  The  total  height  of  Tibet  is  only  about  one  sixth  or  seventh 
of  the  thickness  of  the  earth's  crust,  and  hence  the  uplift,  great  as 
it  is,  is  not  such  as  would  necessarily  produce  g^eat  volcanic  out- 
breaks at  the  surface. 

24.  Great  lava  flows,  however,  occurred  in  India,  and  some  vol- 
canic phenomenon  are  known  in  the  Himalayas,  but  our  knowledge 
of  these  mountains  is  not  yet  adequate  to  enable  one  to  estimate 
just  how  much  volcanic  activity  developed  there. 

25.  Great  lava  flows  are  due  to  the  rupture  of  the  crust,  by  the 
opening  of  a  fault  near  the  sea,  not  to  volcanic  outbreaks.  These 
flows  are  seen  in  Utah,  Oregon  and  India,  on  a  scale  commensurate 
with  the  forces  which  have  uplifted  the  mountains  and  plateaus. 

26.  One  may  pass  directly  from  the  mountains  and  plateaus  of 
South  America  to  those  of  Asia,  and  then  to  those  on  the  Pacific 
slope  of  North  America,  by  the  most  gradual  stages. 

27.  In  this  transition  the  processes  are  so  similar  and  the  dif- 
ferences so  small,  that  it  is  impossible  to  deny  that  the  mountains 


272  SEE^FURTHER   RESEARCHES  ON  [April 34. 

and  plateaus  west  of  the  Rocky  Mountains  were  all  formed  through 
the  uplift  of  the  land  by  the  Pacific  Ocean. 

28.  The  North  American  Plateau  is  larger,  but  correspondingly 
lower  than  those  in  Asia,  so  that  the  volume  of  material  involved 
in  the  two  uplifts  is  comparable.  Thus  all  the  g^eat  plateaus  of 
the  globe  are  due  to  the  action  of  the  sea,  in  the  course  of  immeas- 
urable ages.  The  slowness  of  the  process  conveys  the  best  con- 
ception of  the  vast  interval  of  time  since  the  consolidation  of  the 
globe. 

29.  Charles  Darwin  long  ago  held  that  "  the  forces  which  slowly 
and  by  little  starts  uplift  continents  and  those  which  at  successive 
periods  pour  forth  volcanic  matter  from  open  orifices  are  identical." 
He  showed  that  the  southern  end  of  South  America  has  recently 
risen  from  the  sea,  and  Professor  Suess  has  shown  that  the  univer- 
sal lowering  of  the  strand  line  throughout  the  principal  coimtries 
gives  a  similar  indication  for  all  the  lands  of  the  globe. 

30.  If  one  end  of  a  continent  can  be  raised  by  earthquake  forces 
depending  on  the  sea,  then  obviously  a  whole  continent  can  be  raised 
by  these  forces ;  and  similar  uplifts  can  occur  for  all  the  continents 
in  both  hemispheres.  The  vast  vertical  walls  of  granite  so  often 
found  rising  from  the  sea  in  South  America  and  elsewhere  have 
clearly  been  uplifted  by  earthquakes. 

31.  We  therefore  reach  the  conclusion  that  the  forces  which 
have  raised  the  mountains,  islands  and  plateaus,  have  also  raised 
the  continents  and  established  the  equilibrium  of  the  globe  between 
the  land  and  water  hemispheres.  This  force  is  nothing  else  than 
common  steam,  operating  through  the  expansion  of  molten  rock  be- 
neath the  crust  and  arises  principally  from  the  secular  leakage  of 
the  ocean  bottoms. 

32.  The  main  effect  of  earthquakes  is  the  production  of  more 
land.  The  continents  are  being  lifted  out  of  the  sea,  in  spite  of 
erosion,  as  we  see  by  the  withdrawal  of  the  oceans  to  a  greater 
and  greater  distance  from  old  mountain  chains,  such  as  the  Rocky 
Mountains  and  Appalachians  in  America  and  the  Alps  in  Europe. 

33.  But  for  this  uplift  of  the  land  by  the  leakage  of  the  oceans 
none  of  the  higher  forms  of  life  could  have  developed  upon  the 
earth.    The  climate  and  drainage  of  all  continents  have  been  largely 


X908.I  THE  PHYSICS  OF  THE  EARTH.  273 

determined  by  these  forces,  which  have  produced  the  mountains 
and  river  systems  of  the  world. 

34.  We  cannot  prove  by  experiments  on  rock  twenty  miles 
thick  that  it  will  leak  under  the  pressure  of  the  ocean,  but  we  can 
observe  the  surface  movements  in  earthquakes  such  as  occur  in 
Alaska,  where  lava  is  being  expelled  from  under  the  ocean  and 
pushed  under  the  land. 

35.  This  movement  is  everywhere  in  the  same  direction,  whether 
in  Alaska,  Japan,  the  Antandes,  South  America,  or  elsewhere — 
namely  from  the  ocean  towards  the  land.  The  reason  of  this  is  that 
much  steam  is  formed  under  the  oceans,  but  scarcely  any  under 
the  land,  and  hence  it  pushes  up  the  crust  along  the  edge  of  the  con- 
tinents and  finally  almost  walls  them  in  with  mountains,  as  was 
long  ago  pointed  out  by  Dana. 

36.  The  old  theory  of  secular  cooling  and  contraction  of  the 
globe  is  false  and  misleading,  and  all  who  have  carefully  examined 
it  agree  that  it  is  totally  inadequate  to  account  for  terrestrial 
phenomena.  In  fact  so  far  from  contracting  it  seems  certain  that 
the  earth  is  actually  undergoing  a  slow  secular  expansion. 

37.  The  Rev.  O.  Fisher  and  Major  Button  were  among  the  earliest 
to  reject  this  theory  as  incapable  of  explaining  mountain  ranges. 
But  it  is  remarkable  that  after  the  contraction  theory  was  proved 
to  be  imsatisfactory,  it  continued  to  be  used  in  all  works  on  geology 
and  kindred  sciences,  and  indeed  still  is  accepted  by  those  who 
adhere  to  the  antiquated  doctrine  that  the  earth  is  a  failing  structure. 
Such  views  had  some  justification  a  quarter  of  a  century  ago ;  today 
they  are  absolutely  without  excuse. 

38.  There  are  the  best  grounds  for  accepting  the  doctrine  of 
Isostacy,  as  approximately  true  for  the  earth  at  all  times;  conse- 
quently there  are  no  sensible  stress-diflferences,  or  tendencies  to 
flow,  except  in  the  layers  just  beneath  the  crust.  At  greater  depths 
the  matter  of  the  earth  is  made  solid  by  pressure,  being  at  the  centre 
about  three  times  more  rigid  than  nickel  steel.  Hence  deep  down 
the  earth  is  now  and  always  has  been  quiescent.  The  only  layer  of 
the  earth  which  is  plastic  and  perhaps  viscous  is  that  just  beneath 
the  crust;  this  layer  flows  under  the  tremendous  forces  at  work  in 
earthquake  movements.     It  is  the  movement  of  this  molten  rock  be- 


274  SEE— FURTHER  RESEARCHES  ON  [April  94. 

neath  the  crust,  chiefly  when  it  is  expelled  from  under  the  sea,  whicS 
shakes  down  cities  and  devastates  whole  countries. 

39.  This  expulsion  of  lava  under  the  land  can  mean  nothing  else 
than  the  secular  leakage  of  the  oceans,  because  the  mountains  along 
the  coast  which  are  rent  by  the  shaking  of  the  earth  till  they  break 
into  volcanoes,  emit  chiefly  vapor  of  steam.  Moreover  the  unsym- 
metrical  shape  of  the  mountain  folds,  showing  the  gentler  slope 
towards  the  shore,  indicates  that  the  folds  of  the  crust  were  pushed 
from  the  direction  of  the  sea.  This  was  produced  by  the  expulsion 
of  lava  under  the  crust  arising  from  the  secular  leakage  of  the 
ocean  bottom. 

40.  By  the  study  of  seismic  and  other  phenomena  now  observed 
in  the  great  laboratory  of  nature  we  may  penetrate  the  deepest 
secrets  hidden  beneath  the  earth's  crust,  to  which  no  mortal  eye  can 
ever  bear  direct  witness.  And  these  researches  may  greatly  increase 
the  safety  of  whole  communities,  and  especially  of  cities  and  of 
commerce,  throughout  the  world,  by  enabling  us  to  guard  against 
the  dangers  of  earthquakes  and  seismic  sea  waves.  This  appro- 
priate use  of  the  laboratory  of  Nature  is  one  of  the  ultimate  objects 
of  natural  philosophy. 

Blue  Ridge  on  Lx)utre, 

Montgomery  City,  Missouri, 
February  19,  1908. 

POSTSCRIPT. 

In  a  paper  read  before  the  Royal  Society  in   1902,  Professor 

J.  H.  Jeans,  formerly  of  Cambridge,  now  of  Princeton  University, 

has  the  following  theory  of  earthquakes : 

"  It  seems  to  be  almost  certain  that  the  present  elastic  constants  of  the 
earth  are  such  that  a  state  of  symmetrical  symmetry  would  be  one  of  stable 
equilibrium.  On  the  other  hand,  if  we  look  backward  through  the  history 
of  our  planet,  we  probably  come  to  a  time  when  the  rigidity  was  so  much 
that  the  stable  configuration  of  equilibriums  would  be  unsymmetrical.  At 
this  time  the  earth  would  be  pear-shaped  and  the  transition  to  the  present 
approximately  spherical  form  would  take  place  through  a  series  of  ruptures. 
It  is  suggested  that  the  earth,  in  spite  of  this  series  of  ruptures,  still  retains 
traces  of  a  pear-shaped  configuration.  Such  a  configuration  should  possess 
a  single  axis  of  symmetry,  and  this,  it  is  suggested,  is  an  axis  which  meets 
the  earth's  surface  somewhere  in  the  neighbourhood  of  England  (or  pos- 
sibly some  hundreds  of  miles  to  the  southwest  of  England).    Starting  from 


i9o8.] 


THE  PHYSICS  OF  THE  EARTH.  275 


England  we  find  that  England  is  at  the  centre  of  a  hemisphere  which  is 
practically  all  land;  this  would  be  the  blunt  end  of  our  pear.  Bounding 
the  hemisphere  we  have  a  great  circle,  of  which  England  is  the  pole,  and  it 
is  over  this  circle  that  earthquakes  and  volcanoes  are  of  most  frequent 
occurrence.  Now,  if  we  suppose  our  pear  contracting  to  a  spherical  shape, 
we  notice  that  it  would  probably  be  in  the  neighbourhood  of  its  equator 
that  the  changes  in  curvature  and  the  relative  displacements  would  be 
greatest,  and  hence  we  would  expect  to  find  earthquakes  and  volcanoes  in 
greatest  number  near  this  circle.  Passing  still  further  from  England,  we 
come  to  a  great  region  of  deep  seas,  the  Pacific,  South  Atlantic,  and  Indian 
Oceans ;  these  may  mark  the  place  where  the  *  waist  *  of  the  pear  occurred. 
Lastly  we  come  almost  to  the  antipodes  of  England,  to  the  Australian  conti- 
nent. This  may  mark  the  remains  of  the  stalk-end  of  the  pear."  (Nature, 
Vol.  LXVII.,  p.  190.) 

After  what  has  been  shown  in  this  series  of  papers,  it  is  un- 
necessary to  dwell  upon  this  hypothesis  of  Professor  Jeans,  which 
has  the  merit  of  originality;  but  we  may  remark  that  if  it  gave  a 
true  view  of  the  physics  of  the  earth,  there  should  be  a  belt 
around  the  globe  of  at  least  the  width  of  the  terrestrial  radius,  over 
which  the  earthquakes  are  about  equally  distributed,  whereas  in 
fact  they  are  felt  principally  along  the  margins  of  the  Pacific 
Ocean.  The  observed  earthquake  belt  on  land  is  so  narrow  that 
it  is  clearly  impossible  to  ascribe  the  effects  to  this  supposed  adjust- 
ment of  the  earth's  figure.  And  of  course  it  fails  totally  to  ac- 
count for  the  sinking  of  the  sea  bottom  and  the  uplift  of  the  coast, 
which  is  typical  of  mountain  formation. 


THE  ABSORPTION  SPECTRA  OF  NEODYMIUM  CHLO- 
RIDE AND  PRASEODYMIUM  CHLORIDE  IN  WATER, 
METHYL  ALCOHOL,  ETHYL  ALCOHOL  AND 
MIXTURES  OF  THESE  SOLVENTS. 

(With  six  plates.) 

Twenty-first  Communication. 

By  harry  C.  JONES  and  JOHN  A.  ANDERSON. 

(Read  April  2$,  1908.) 

(This  is  a  preliminary  report  on  part  of  an  investigation  carried  out  with  the 
aid  of  a  Grant  from  the  Carnegie  Institution  of  Washington.) 

The  absorption  spectra  of  salts  of  cobalt,  nickel,  copper,  iron, 
chromium,  neodymium,  praseodymiimi  and  erbium  have  been  stud- 
ied in  the  present  investigation.  Of  these  the  salts  of  neodymium 
and  praseodymium  are  perhaps  the  most  interesting  and  important. 
This  is  due  to  the  large  number  of  absorption  bands  shown  by  these 
substances,  and,  further,  to  the  very  unusually  sharp  character  of 
these  bands. 

The  method  employed  in  making  the  spectrograms  consists  in 
allowing  light  from  a  spark,  or  from  a  Nernst  filament,  to  pass 
through  the  solution  in  question,  fall  upon  a  grating  and  then  upon 
the  photographic  plate. 

For  visual  work  a  small  direct  vision  grating  pocket  spectroscope 
was  found  very  convenient  and  useful.  For  photographing  the 
spectra  the  vertical  grating  spectroscope  used  by  Jones  and  Uhler^ 
was  employed. 

In  making  the  photographs  the  Seed  L-ortho  film  was  used  for 
the  region  from  A  2000  to  about  A  6000. 

For  photographing  the  red  end  of  the  spectrum  a  Wratten  and 
Wainwright  panchromatic  glass  plate  was  used. 

*  Carnegie  Publication  No.  60. 

276 


i9o8.]  NEODYMIUM  AND  PRASEODYMIUM.  277 

The  Nemst  filament  was  found  to  be  the  most  satisfactory 
source  of  light  from  the  extreme  red  to  the  beginning  of  the  ultra- 
violet It  is  sufficiently  brilliant  to  require  an  exposure  of  only  a 
minute,  but  practically  ceases  at  about  X3200.  For  wave-lengths 
shorter  than  this  some  spark  spectrum  must  be  used. 

The  cadmium-zinc  spark  used  by  Jones  and  Uhler  was  fairly 
satisfactory,  especially  in  the  extreme  ultra-violet,  but  has  the  draw- 
back that  there  are  present  a  limited  number  of  very  intense  lines, 
on  a  rather  faint  continuous  background.  We  tried  to  obtain  a 
spark  spectrum  having  a  very  large  number  of  lines,  but  with  no 
lines  of  very  great  intensity.  We  found  that  tungsten,  molybdenum 
and  uranium  all  satisfied  these  requirements. 

The  terminals  finally  used  were  prepared  by  dipping  pieces  of 
carbon  in  a  concentrated  solution  of  ammonium  molybdatc,  and 
then  heating  in  a  bunsen  burner.  They  were  then  dipped  into  a 
solution  of  uranium  nitrate  and  similarly  heated. 

The  coil  used  to  produce  the  spark  was  a  large  Rontgen  X-ray 
coil. 

Making  a  Spectrogram. 

In  making  a  spectrogram  consisting  of  seven  photographic  strips, 
the  following  mode  of  procedure  was  adopted :  Seven  separate  solu- 
tions were  made  up  of  the  desired  strengths.  The  cell*  to  be  used 
was  filled  to  the  required  depth  with  the  most  concentrated  solution 
of  the  series,  and  the  quartz  plates  determining  the  depth  of  the 
solution  adjusted  to  parallelism.  The  exposure  to  the  Nemst  lamp 
was  then  made,  being  usually  one  minute  long.  An  opaque  screen 
covering  up  the  visible  spectrum  as  far  down  as  X4000  was  then 
interposed  between  the  grating  and  the  photographic  film,  and  the 
exposure  to  the  light  of  the  spark  in  the  ultra-violet  made.  The 
duration  of  this  exposure  was  usually  about  two  minutes.  The 
photographic  film  was  then  moved  into  the  proper  position  for  the 
next  exposure.  The  above  series  of  operations  was  then  repeated 
for  each  of  the  succeeding  strips. 

After  the  film  had  been  exposed  for  each  solution  and  the  spark 
spectrum  impressed,  it  was  necessary  to  make  a  similar  series  of 

'See  Carnegie  Publication  No.  60. 


278  JONES  AND  ANDERSON— ABSORPTION   SPECTRA  OF     [April  as. 

exposures  on  a  panchromatic  plate  for  the  red  end  of  the  spectrum, 
using  the  same  set  of  solutions. 

The  scale  accompanying  the  spectrograms  was  made  by  photo- 
graphing an  ordinary  paper  scale.  Several  photographs  were  taken, 
the  distance  between  the  paper  scale  and  the  lens  of  the  camera 
being  varied  slightly  from  exposure  to  exposure.  The  resulting 
negative  which  fitted  the  majority  of  spectrograms  best  was  selected 
and  used  throughout 

Neodymium  Chloride  in  Water — Beer's  Law. 

(See  plate  i.) 

The  concentrations  of  the  solutions  of  neodymium  chloride  were 
so  chosen  and  the  depths  of  cell  so  selected  that  the  total  amount  of 
coloring  matter  in  the  path  of  the  beam  of  light  was  kept  constant. 
From  Beer's  Law  the  absorption  shown  by  the  several  solutions, 
under  these  conditions,  should  be  the  same.  The  concentrations  of 
the  solutions  used  in  making  the  negative  for  a,  plate  i,  beginning 
with  the  one  whose  spectrum  is  adjacent  to  the  numbered  scale, 
were  3.40,  3.02,  2.72,  2.38,  2.17,  1.90  and  1.70;  the  corresponding 
depths  of  cell  being  12,  13.5,  15,  17,  19,  21.5  and  24  mm.  For  b, 
plate  I,  the  concentrations  were  3.40,  2.55,  1.70,  1.13,  0.80,  0.57  and 
0.43 ;  the  corresponding  depths  of  absorbing  layer  being  3,  4,  6,  9, 
13,  18  and  24  mm. 

The  most  concentrated  solutions  appeared  brownish  yellow  in 
their  bottles,  from  which  the  color  changed  on  dilution  to  a  yellowish 
pink,  the  color  being  extremely  faint  in  the  most  dilute  solutions. 

The  exposures  to  the  light  of  the  Nernst  lamp  and  spark  were, 
respectively,  i  minute  and  2  minutes ;  the  slit  having  a  width  of  o.oi 
cm.  The  exposures  and  slit  width  were  not  varied  in  the  work 
recorded  in  the  present  chapter,  the  object  being  to  make  the  spectro- 
grams as  nearly  comparable  as  possible. 

Both  a  and  b  of  plate  i  show  the  presence  of  some  general 
absorption  in  the  ultra-violet,  which  decreases  quite  rapidly  with 
dilution.  The  absorption  bands  also  narrow  somewhat  with  de- 
crease in  concentration,  especially  from  3.4  normal  to  about  1.7 
normal.  For  concentrations  less  than  about  1.5  normal  Beer's  Law 
seems  to  hold  very  accurately  indeed,  with  the  exception  of  the 


»90«.] 


NEODYMIUM   AND  PRASEODYMIUM. 


279 


shading  towards  the  red  accompanying  the  band  near  X  5800,  which 
seems  to  decrease  somewhat  with  dilution  for  concentrations  of 
one  normal  or  less. 

In  the  following  table  the  measurements  of  the  positions  of  the 
bands  were  made  on  the  seventh  strips  of  a,  plate  i,  and,  therefore, 
refer  to  a  concentration  of  1.7  normal  with  a  depth  of  layer  of  24 
mm.  The  remarks  referring  to  changes  with  dilution  apply  to  a 
change  in  concentration  from  3.4  to  1.7  normal,  the  depths  of  layer 
being  so  varied  that  the  product  of  concentration  and  depth  remains 
constant. 

Remarks. 


k 

Character. 

2810 

Faint  transmission  begins. 

2890-2910 

Band  with  well  defined  sharp 

edges. 

2970-2995 

A  double  band,  strongest  com- 

ponent to  violet 

3220-3330 

Strong  band  of  complete  ab- 

sorption, sharp  edges. 

3^Jo-3400 

Rather    faint   band,   most   in- 

tense towards  red. 

3435-3595 

Complete     absorption,     edges 

sharp. 

4180 

Hazy,  not  very  intense. 

4275 

Very  intense  and  sharp. 

4290 

Narrow  and  faint. 

4330  Hazy. 

4410-4465      Edges  rather  hazy. 


4580-4650 


4665-4710 


Band  with  hazy  edges  not 
completely  separated  from 
X  4665-X  4710. 

More  sharply  defined  on  red 
than  on  violet  side. 


The  observed  narrowing  with 
dilution  perhaps  due  largely 
to  general  U.  V.  absorption. 

Narrows  slightly  with  dilution. 


Narrows  some  with  dilution. 

Narrows  considerably  at  first. 

Between  this  and  X4275  is 
fairly  strong  absorption  in 
the  most  concentrated  solu- 
tion. This  absorption  has 
disappeared  in  the  spectrum 
measured. 

This  band  is  coincident  with 
band  due  to  praseod}rmium, 
and  is  to  be  ascribed  to  this 
element  which  has  not  been 
completely  separated  from 
the  neodymium.  It  does  not 
change  with  dilution. 

Narrows  slightly  with  dilution. 


Partly  due  to  praseodjrmium. 
Does  not  change  with  dilu- 
tion. 


280 


JONES  AND   ANDERSON— ABSORPTION  SPECTRA  OF    [AprUas, 


K  Character. 

4740-4770      Fairly  sharp  edges. 
4820  Hazy  on  violet  side. 


5000-5330 

Red  limit  sharp,  violet  a  little 

hazy. 

566o:-5930 

Violet  limit  sharp.     Red  edge 

hazy. 

6235 

First    and    strongest   band    in 

orange  group. 

6260 

Narrow  and  rather  faint. 

6270-6310 

Faint  band. 

6360-6390 

Faint  band. 

6730 

Faint,  in  shading  of  principal 

red  band. 

6770-6840, 

Principal    red    band.      Edges 

hazy. 

6890 

Band  with  hazy  edges. 

7250 

End  of  transmission. 

Remarks. 

Not  affected  by  dilution. 
Due  at  least  partly  to  prase- 

odjrmium. 
Violet  shading  a  little  greater 

in  concentrated  solutions. 
Shading  on  red  side  decreases 

with  dilution. 
Not  affected  by  dilution. 

Not  affected  by  dilution. 
Not  affected  by  dilution. 
Not  affected  by  dilution. 
Not  affected  by  dilution. 

Not  affected  by  dilution. 

Not  affected  by  dilution. 
Not  affected  by  dilution. 


The  most  marked  change  produced  by  dilution  from  3.4  to  1.7 
normal,  excepting  that  in  the  red  shading  of  the  X  5660-5930  band, 
is  that  taking  place  on  the  red  side  of  the  narrow  absorption  line 
at  X4275.  In  the  spectrum  of  the  most  concentrated  solution  the 
red  edge  of  this  line  falls  at  X4280,  from  which  place  a  uniform 
absorption  extends  to  A  4295.  In  the  third  spectrum,  counting  from 
the  numbered  scale,  the  shading  has  almost  completely  disappeared, 
leaving  a  very  narrow  line  at  approximately  A  4290.  The  width  of 
this  line  is  only  2  or  3  A.  U.  and  it  persists  with  unchanged  intensity 
throughout  the  remaining  strips  of  the  spectrogram.  Its  intensity 
is,  however,  not  sufficient  to  make  it  show  in  the  reproduction,  and 
not  even  great  enough  to  make  it  visible  on  the  negative  for  b, 
plate  I. 

The  limits  of  transmission  for  the  yellow  band,  as  shown  by  the 
spectrum  of  the  most  concentrated  solution,  are  X  5660  and  X  5950 ; 
hence  the  narrowing  of  its  red  side  amounts  to  20  A.  U. 

b,  plate  I,  starts  at  the  same  concentration  as  a,  but  the  effective 
depth  of  absorbing  layer  is  only  one-fourth  of  that  used  in  a. 
Hence  this  spectrogram  represents  the  spectrum  of  a  solution  of 
neodymium  chloride  24  mm.  deep  and  having  a  concentration  of 
0.43  normal.    The  absorption  bands  are  all  much  narrower,  and 


x9o8,J  NEODYMIUM   AND   PRASEODYMIUM.  281 

several  of  them  are  shown  in  the  process  of  breaking  up  into  sim- 
pler bands.  The  bands  in  the  ultra-violet  have  disappeared  except- 
ing the  one  at  X  3435-X  3595,  which  is  still  intense,  and  a  trace  of 
the  one  at  A  3220-A  3330.  Transmission  in  this  region  now  extends 
faintly  to  A  2460.  No  new  absorption  bands  beyond  A  2800  can  be 
seen. 

The  A3435-A3595  band  now  has  the  limits  A3450-A3580,  and 
shows  a  weak  transmission  at  A  3485,  which  increases  somewhat 
with  dilution,  thus  dividing  the  band  into  two. 

The  band  at  A  4180  is  weak  throughout  b,  plate  i. 

The  band  having  its  middle  at  A  4445,  perhaps  due  entirely  to 
praseodymium,  in  a,  plate  i,  has  about  the  same  intensity  as  it  shows 
in  a  solution  of  praseodymium  chloride  having  a  concentration  of 
0.85  and  a  depth  of  absorbing  layer  equal  to  3  mm.  This  indicates 
that  the  percentage  of  praseodymium  in  the  neodymium  salts  used 
was  about  6  per  cent.  The  band  at  A  4825  partly  due  to  praseody- 
mium may  also  be  seen  throughout  the  entire  series  under  consid- 
eration. The  wave-length  of  the  praseodymium  band  being  A  4815,' 
while  that  of  the  band  showing  in  all  the  neodymium  spectra  has  the 
position  A  4825,  showing  that  neodymium  has  a  band  nearly  coin- 
cident with  that  given  by  praseodymium,  but  lying  a  little  closer  to 
the  red  end  of  the  spectrum.  The  remaining  praseodymium  band 
has  the  position  A  4685,  this  nearly  coinciding  with  the  rather  nar- 
row, strong  neodymium  band  whose  position  is  A  4695. 

The  band  which  under  a,  plate  i,  was  recorded  as  having  the 
limits  A4580-A4650,  shows  in  &  as  a  hazy  band  with  its  center  at 
4615,  together  with  a  narrow  faint  line  at  A  4645. 

The  band  which  in  the  table  is  recorded  as  A4740-A4770  has 
in  b,  plate  i,  become  a  slightly  hazy  band  having  its  middle  at 
A  4760.  Its  intensity  is  intermediate  between  that  of  the  bands  at 
A  4695  and  A  4825. 

The  band  which  in  a,  plate  i,  has  the  limits  A  5000-A  5330,  breaks 
up  into  a  rather  complicated  series  of  bands  on  dilution,  some  idea 
of  which  may  perhaps  be  gained  from  the  following:  b,  plate  i, 
shows  some  absorption  throughout  the  region  given,  but  with  a 
deep,  narrow  band  at  A  5090,  and  faint  transmission  at  A  5100  and 
in  the  region  A5i5(>-A5i8o.    Absorption  is  complete  from  A  5105 


282  JONES  AND  ANDERSON— ABSORPTION   SPECTRA  OF     [April  aj, 

to  X5150,  and  from  A  5180  to  A  5270.  There  is  again  incomplete 
absorption   from  A  5270  to  A  5330,  with  indication  of  a  band  at 

A  5315. 

The  limits  of  the  yellow  band  in  b,  plate  i,  are  A5700-A5880, 

in  the  strip  corresponding  to  the  most  concentrated  solution.  The 
band  narrows  by  30  Angstrom  units  on  this  spectrogram,  the  nar- 
rowing being  due  to  a  decrease  in  the  shading  towards  the  red,  with 
decrease  in  concentration. 

The  most  intense  bands  of  neodymium  chloride,  and  hence  the 
ones  which  would  be  most  conspicuous  in  a  very  dilute  solution  are 
the  following :  A  3465,  A  3540,  A  4275,  A  5205,  A  5225,  A  5745,  X  5765 
and  A  7325. 

The  wave-lengths  of  all  the  bands  are  collected  in  the  following 
table,  together  with  a  brief  description  of  the  appearance  of  each 
band.  It  is  to  be  understood  that  this  table  is  not  meant  to  repre- 
sent what  could  be  seen  or  photographed  in  any  one  solution  of 
neodymium  chloride  in  water.  It  merely  records  the  positions  of 
all  the  bands  that  can  be  seen  in  a  layer  from  3  to  12  mm.  deep, 
when  the  concentration  is  varied  from  o  to  3.4  normal. 

A  Description. 

2900  About  20  A.  U.  wide. 

2985  About  25  A.  U.  wide. 

3225  Narrow  and  sharp. 

3390  Narrow,  faint. 

3465  Very  intense,  narrow. 

3505  Rather  wide. 

3540  Very  intense,  narrow. 

3560  Faint,  narrow. 

4180  Faint,  hazy. 

4275  Very  intense  and  sharp. 

4290  Very  narrow,  faint. 

4330  Hazy  edges. 

4615  Rather  wide  and  hazy. 

4645  Very  narrow,  faint 

4695  Narrow,  intense. 

4760  Hazy  edges,  fairly  narrow. 

4825  Narrow  and  fairly  intense. 

5090  Narrow,  intense. 

5125  Rather  wide  and  hazy. 

5205  Very  intense,  narrow. 

5222  Very  intense,  narrow. 


»-*t'      -     ~  —  .      I  » 


4 


,9o8,|  NEODYMIUM   AND  PRASEODYMIUM.  283 

5255  Narrow,  intense. 

5315  Hazy  edges,  faint. 

5725  Narrow,  intense. 

5745  Very  intense. 

5765  Very  intense. 

5795  Intense,  moderately  narrow. 

5830  Very  faint  and  hazy. 

6235  Fairly  narrow. 

6260  Very  narrow,  faint. 

6270-6310  Faint,  hazy  edges. 

6360-6390  Faint,  hazy  edges. 

6730  Faint  band. 

6800  Moderately  intense,  hazy  edges. 

6890  Hazy  edges. 

7325  Very  intense  and  narrow. 

7350  Narrow. 

7390  Rather  wide  band. 

Neodymium  Chloride  in  Methyl  Alcohol — Beer's  Law. 

(See  plate  2.) 

The  concentrations  of  the  solutions  used  in  making  the  negative 
for  a,  beginning  with  the  one  whose  spectrum  is  adjacent  to  the 
numbered  scale  were  0.50,  0.40,  0.315,  0.25,  0.20,  0.16  and  0.125; 
the  corresponding  depths  of  absorbing  layer  being  6,  7.5,  9.5,  12, 
15,  19  and  24  mm.  The  concentrations  for  b  were  in  the  same 
order  0.20,  0.16,  0.13,  o.io,  0.08,  0.06  and  0.05,  the  depths  of  cell 
being  the  same  as  used  in  a. 

There  is  some  absorption  in  the  extreme  ultra-violet,  which  is 
to  be  ascribed  to  the  solvent,  however,  and  not  to  the  neodymium 
chloride. 

No  trace  of  absorption  due  to  the  dissolved  substance  is  visible 
until  we  reach  the  group  of  bands  near  X  3500.  These  are  three 
bands  having  their  centers  at  X  3475,  X  3505,  and  A  3560.  Of  these 
the  one  at  A  3560  is  the  widest  and  also  the  most  intense ;  the  one  at 
^  3475  being  somewhat  fainter  than  that  at  A  3505.  The  bands  are 
all  much  wider  and  hazier  than  those  occurring  near  the  same  place 
in  the  aqueous  solution.  No  change  with  dilution,  indicating  a 
deviation  from  Beer's  Law,  can  be  detected  in  these  or  any  of  the 
other  bands  in  the  alcoholic  solutions  of  the  chloride. 

Tn  the  violet  and  blue  regions  we  find  the  following  band  at 


284  JONES  AND  ANDERSON— ABSORPTION  SPECTRA  OF    [April  a,, 

X  4290,  about  ID  A.  U.  wide  and  only  moderately  intense.  At  A  4325 
a  band  somewhat  wider  and  fainter.  At  X  4460,  a  rather  wide  hazy 
band  with  a  faint  hazy  companion  towards  the  violet.  This  is  the 
band  which  is  perhaps  due  to  praseodymium.  The  much  greater 
concentration  of  the  alcoholic  solutions  of  praseodymium  chloride 
studied  in  this  work,  makes  it  impossible  to  verify  this  by  seeing 
whether  the  praseodymium  band  in  dilute  solution  really  has  this 
general  character. 

Thefe  are  bands  at  A  4700,  A  4780  and  A  4825,  all  of  about  the 
same  intensity;  the  one  at  A 4770  being,  however,  much  narrower 
than  the  other  two,  of  which  A  4825  is  somewhat  the  wider.  Both 
A  4700  and  A  4780  have  faint  companions  to  the  violet. 

The  group  in  the  green  is  made  up  of  six  bands  as  follows: 
A 5125  hazy  and  rather  wide,  moderately  intense;  A 5180,  also  hazy 
but  much  fainter;  A 5220  moderately  intense  and  narrow;  A 5245 
intense  and  with  faint  companion  towards  the  red;  A 5290  narrow 
and  moderately  intense.  Shading  as  far  as  A  5330  with  indications 
of  faint  band  at  A  5315. 

The  yellow  group  is  made  up  of  seven  bands  having  the  fol- 
lowing characteristics :  A  5725  moderately  intense  with  hazy  edges ; 
A  5765  narrower,  but  not  quite  as  intense  as  A  5725;  A  5800  fairly 
narrow,  strong;  A 5835  very  intense;  A 5860  hazy  and  moderately 
intense ;  not  clearly  separated  from  A  5835  shading  to  A  5970,  with 
two  faint  bands  superposed  on  it,  one  at  A  5895  and  the  other  at 

^  5925- 

No  trace  of  bands  is  to  be  seen  in  the  orange,  but  in  the  red 

there  is  a  fairly  narrow  but  faint  band  at  A  6860.  The  spectnmi 
ends  at  A  7355  in  a  deep,  rather  narrow  band.  It  is  evident  that 
the  spectrum  of  neodymium  chloride  when  dissolved  in  methyl  alco- 
hol is  quite  different  from  its  spectrtmi  in  aqueous  solution. 

Neodymium  Chloride  in  Mixtures  of  Methyl  Alcohol  and 

Water.     (See  plates  3  and  4.) 

Since,  as  we  have  just  seen,  the  absorption  spectrum  of  neody- 
mium chloride  in  aqueous  solution  is  so  diflferent  from  that  of  the 
alcoholic  solutions,  it  was  thought  to  be  of  some  interest  to  see  how 
the  change  from  one  to  the  other  would  take  place  if  one  of  the 


,9o8.]  NEODYMIUM  AND   PRASEODYMIUM.  285 

solvents  was  made  to  displace  the  other  gradually.  A  series  of  solu- 
tions was  accordingly  made  up,  the  concentration  of  the  dissolved 
salt  being  constant  and  equal  to  0.5  normal,  but  the  character  of  the 
solvent  varying  as  follows :  The  percentages  of  water  in  the  seven 
solutions  were  o,  16S,  33^,  50,  66f ,  83J  and  100 ;  the  corresponding 
percentages  of  methyl  alcohol  were  100,  83^,  66f,  50,  33^,  i6| 
and  o.  Two  spectrograms  were  made,  namely  a,  plate  3,  where  the 
depth  of  the  cell  was  1.5  cm.  and  b  where  the  cell  had  a  depth  of 
only  5  mm.  a  was  made  in  order  to  show  clearly  the  change  taking 
place  in  the  narrower  and  fainter  bands,  while  b  was  intended  to 
show  the  change  of  structure  of  the  more  intense  bands  such  as  the 
green  and  yellow  ones.  The  strip  which  is  adjacent  to  the  num- 
bered scale  belongs  to  the  solution  in  pure  water,  while  the  one 
nearest  the  narrow  comparison  spark  spectrum  belongs  to  the  solu- 
tion in  pure  methyl  alcohol. 

Plate  3  shows  that  beginning  with  the  strip  nearest  the  scale,  the 
first  six  spectra  are  very  nearly  identical.  From  the  sixth  to  tiie 
seventh  there  is  an  abrupt  change  which  at  first  sight  consists  in  a 
shift  of  all  the  bands  towards  the  red,  but  which  on  closer  examina- 
tion is  seen  to  consist  in  a  disappearance  of  one  spectrum  and  the 
appearance  of  tiie  other.  Since  the  first  strip  is  the  spectrum  of 
the  solution  in  pure  water,  it  follows,  since  the  sixth  is  nearly 
identical  with  the  first,  that  as  large  a  percentage  of  alcohol  in  the 
solvent  as  83  per  cent,  does  not  change  the  absorption  spectrum 
materially;  the  chief  change  taking  place  when  the  percentage  of 
alcohol,  is  varied  from  83  per  cent,  to  100  per  cent. 

It  is  to  be  noted  that  the  apparent  shift  of  the  bands  towards 
the  red  is  in  reality  not  quite  as  great  as  it  appears  at  first  sight 
from  plate  3,  owing  to  the  fact  that  the  film  accidently  shifted 
slightly  towards  the  red  between  the  sixth  and  seventh  exposures. 
The  amount  of  this  mechanical  shift  is  easily  seen,  however,  by 
comparing  the  spark  lines  in  the  ultra-violet.  A  measurement  of 
the  shift  shows  it  to  be  approximately  3  Angstrom  units,  and  the 
same  for  both  a  and  fc,  while  the  "  apparent "  shift  of  the  absorp- 
tion line  at  A  4275  in  aqueous  solution  is  actually  15  Angstrom 
units,  its  position  in  the  alcoholic  solution  being  A  4290. 

PROG.  AMER.  PHIL.  SOC,  XLVII.  1 89  S,  PRINTED  SEPTEMBER  2$,  I908. 


286  JONES  AND  ANDERSON— ABSORPTION  SPECTRA   OF    [April  .5, 

The  slight  changes  taking  place  with  someCof  the  bands  through- 
out the  spectrograms  of  plate  3  are  perhaps  sufficiently  clear  in  the 
reproductions.  However,  as  a  good  deal  of  the  detail  shown  by 
the  negatives  is  lost  even  in  the  most  perfect  processes  of  repro- 
duction, we  give  here  a  description  of  the  changes  taking  place  in 
two  of  the  bands  as  seen  on  the  original  negative.  We  select  the 
bands  at  X  4275,  and  A  4760  from  the  negative  for  a,  plate  3. 

In  the  aqueous  solution  the  A  4275  band  is  very  intense  and  nar- 
row, its  whole  width  being  less  than  5  Angstrom  units.  The  edges 
are  only  very  slightly  shaded.  In  the  alcoholic  solution  the  posi- 
tion of  the  center  of  the  corresponding  band  is  X4290.  It  has  a 
width  of  from  12  to  13  Angstrom  units,  and  is  not  nearly  as  intense 
as  in  the  aqueous  solution. 

Throughout  the  first  six  strips  the  A  4275  band  maintains  its 
position  and  intensity  almost  unchanged.  Its  position  does  not 
change  in  the  least,  but  its  intensity  in  the  sixth  strip  is  a  trifle  less 
than  in  the  others.  In  the  seventh  stfip  there  is  not  the  faintest 
trace  left  of  it.  In  the  third  strip,  corresponding  to  the  solution 
whose  alcohol  content  was  33J  per  cent.,  there  appears  at  A  4285  an 
extremely  faint  and  narrow  line.  In  the  fourth  strip  it  is  some- 
what wider  and  more  intense,  but  its  center  is  still  at  A  4285.  In  the 
fifth  strip  it  is  beginning  to  be  fairly  conspicuous,  and  in  the  sixth 
it  is  a  band  of  moderate  intensity  having  its  center  at  about  A  4287. 
This  band  is  undoubtedly  the  same  one  which  in  the  pure  alcoholic 
solution  has  its  center  at  A  4290  or  very  near  there ;  the  exact  wave- 
length being  perhaps  nearer  to  A  4292.  We  see  then  that  even  when 
the  mixed  solvent  contains  only  about  one-half  alcohol  this  band 
exists  independent  of  and  distinct  from  the  band  characteristic  of 
the  aqueous  solution ;  that  it  is  at  first  only  a  very  narrow  and  faint 
line  which  widens  towards  the  red  as  the  percentage  of  alcohol  is 
increased. 

The  band  whose  center  is  at  A  4760  has  the  following  appear- 
ance in  the  aqueous  solution :  Faint  absorption  begins  at  A  4748  and 
rises  rapidly  to  a  maximum  between  A  4755  and  A  4760,  then  de- 
creases slowly  to  nothing  at  A  4775.  The  band  is  accordingly  a 
trifle  asymmetrical,  the  slope  towards  the  violet  being  considerably 
steeper  than  that  towards  the  red.     The  corresponding  band  in  the 


I908.]  NEODYMIUM   AND  PRASEODYMIUM.  287 

alcoholic  solution  is  double  and  answers  the  following  description: 
Very  faint  absorption  begins  at  A  4753  and  rises  to  a  faint  maxi- 
mum at  about  A  4757,  becoming  again  zero  at  X4760.  It  begins 
again  at  A  4772,  rises  rapidly  to  a  strong  maximum  at  A  4780  and 
falls  to  "zero  at  A  4790.  The  component  whose  center  is  at  A  4757 
is  very  faint  compared  with  the  main  band. 

In  the  first  and  second  strips  we  have  nothing  but  the  band 
corresponding  to  the  aqueous  solution.  In  the  third  strip  the  red 
side  of  the  band  has  increased  slightly  in  intensity,  making  it  appear 
much  more  nearly  symmetrical.  This  change  increases  in  the  fourth 
and  fifth  strips,  the  band  at  the  same  time  widening  considerably. 
In  the  sixth  strip  its  appearance  is  as  follows :  Absorption  begins  at 
A  4748  and  rises  to  a  maximum  just  to  the  violet  side  of  A  4760, 
then  decreases  slightly  towards  A  4770,  after  which  it  increases 
somewhat  to  A  4778,  then  falls  oS  to  zero  at  A  4787. 

It  is  very  evident  from  a  study  of  the  change  in  this  band  that 
the  two  bands  characteristic  of  the  aqueous  and  alcoholic  solutions 
coexist,  and  that  the  band  appearing  in  our  photographic  strips  is  the 
sum  of  the  two  taken  in  different  proportions.  The  proportion  of 
the  alcohol  band  being,  however,  very  much  smaller  than  the  pro- 
portion of  alcohol  in  the  corresponding  solution.  A  similar  de- 
scription might  be  given  for  any  one  of  the  other  bands,  but  this  is 
not  necessary  as  the  changes  are  of  exactly  the  same  nature  as  those 
we  have  already  indicated.  In  every  case  where  the  alcoholic  solu- 
tion has  a  strong  band,  which  differs  somewhat  in  position  from 
any  band  in  the  aqueous  solution,  we  begin  to  see  traces  of  this 
band  when  the  proportion  of  alcohol  in  the  mixture  reaches  50  per 
cent.,  but  the  band  remains  comparatively  faint  even  when  the  pro- 
portion is  as  high  as  83J  per  cent. 

In  order  to  study  the  change  which  takes  place  between  the  sixth 
and  seventh  strips  of  the  spectrograms  of  plate  3,  more  carefully,  a 
series  of  alcohol  solutions  were  prepared  containing  the  following 
percentages  of  water,  o,  2|,  5^,  8,  io§,  13^  and  16.  The  concen- 
tration of  the  neodymium  chloride  was  constant  and  equal  to  0.5 
normal.  Two  spectrograms  were  made,  one  with  a  depth  of  absorb- 
ing layer  of  1.5  cm.,  in  order  to  show  the  fainter  bands,  and  the 
other  with  the  depth  of  the  cell  only  5  mm.  in  order  to  show  as 


288  JONES  AND  ANDERSON— ABSORPTION   SPECTRA  OF     [April  a$, 

much  as  possible  of  the  structure  of  the  larger  bands.  The  first 
spectrogram  is  reproduced  as  a,  the  second  as  b,  plate  4.  The 
strips  corresponding  to  the  pure  alcohol  solutions  are  adjacent  to 
the  numbered  scale,  the  spectnim  of  the  solution  containing  16  per 
cent,  water  being  next  to  the  comparison  spark  spectnmi. 

Although  we  found  in  considering  plate  3  that  some  slight  change 
in  the  spectrum  takes  place  where  the  percentage  of  alcohol  is 
changed  from  o  to  83  per  cent.,  yet  this  change  is  so  small  and 
the  bands  due  to  the  aqueous  solution  are  so  strong  that  we  may 
regard  the  spectrum  of  a  solution  containing  16  per  cent,  of  water 
as  practically  that  of  the  aqueous  solution.  Accordingly,  the  spec- 
trograms on  plate  4  may  be  taken  to  show  very  nearly  the  whole 
change  which  takes  place  when  the  solvent  of  neodymium  chloride 
is  gradually  changed  from  pure  water  to  pure  methyl  alcohol. 

In  a  the  ultra-violet  band  is  rather  too  intense  to  allow  its 
structure  to  be  seen.  Accordingly,  we  see  the  whole  band  remains 
sensibly  unchanged  as  the  water  is  varied  from  16  per  cent,  to  8 
per  cent.,  and  then  shifts  towards  the  red  with  increasing  rapidity  as 
the  water  is  reduced  to  zero ;  the  whole  apparent  shift  amounting  to 
about  twenty  Angstrom  units.  On  the  negative  the  intense  band 
at  A  3465  may,  however,  be  clearly  seen,  and  its  intensity  decreases 
very  slowly  from  the  first  to  the  third  strips,  counting  from  the 
narrow  comparison  spark  spectrum.  In  the  fourth  strip  its  inten- 
sity is  about  half  of  what  it  was  in  the  first  strip,  and  from  this  it 
decreases  rapidly,  vanishing  entirely  in  the  strip  nearest  the  scale. 
In  b  the  structure  of  this  band  is  seen  very  distinctly,  and  we  find 
that  the  bands  characteristic  of  the  aqueous  solution  gradually  de- 
crease in  intensity,  especially  from  the  third  to  the  sixth  strips,  while 
the  wider  bands  characteristic  of  the  alcoholic  solutions  increase  in 
intensity,  the  two  sets  existing  together.  The  change  in  the  band 
at  A  4275  is  the  one  that  shows  the  best,  because  here  the  two  bands 
belonging  to  the  aqueous  and  alcoholic  solutions,  respectively,  are 
both  intense  and  narrow  and  clearly  separated  from  each  other. 
The  alcoholic  band  is  clearly  visible  in  the  first  strip,  and  it  in- 
creases continuously  in  intensity  as  the  amount  of  water  is  de- 
creased, but  more  rapidly  from  the  fourth  to  the  seventh  strips 
than  from  the  first  to  the  fourth.     Its  position  also  shifts  somewhat 


,9o8.]  NEODYMIUM  AND  PRASEODYMIUM.  289 

towards  the  red  from  the  first  to  the  fourth  strips,  the  wave  lengths 
of  its  center  for  the  two  strips  being,  respectively,  A  4287  and  A  4292. 
Accompanying  this  shift  is  a  change  in  its  character  which  may  be 
gathered  from  the  following  statements:  In  the  first  strip  it  has 
the  appearance  of  an  unsymmetrical  band,  the  maximum  intensity 
being  nearer  the  violet ;  in  the  third  strip  it  extends  from  A  4280  to 
A 4295  and  has  about  the  same  intensity  throughout;  in  the  fourth 
strip  the  intensity  of  its  violet  edge  has  decreased,  while  that  of 
the  red  edge  has  increased  considerably,  giving  it  the  appearance 
of  an  unsymmetrical  band  with  the  maximum  intensity  towards  the 
red.  In  the  fifth  strip  the  violet  shading  from  A  4280  to  about 
A  4284  has  disappeared,  leaving  a  band  very  nearly  symmetrical 
about  A  4290.  It  appears,  therefore,  that  we  are  really  dealing  with 
two  unresolved  bands,  one  having  its  center  at  about  A  4285,  and  the 
other  at  A  4292. 

The  band  at  A  4275,  due  to  the  aqueous  solution,  decreases  in  in- 
tensity throughout,  but  more  rapidly  from  the  third  to  the  sixth 
strips  than  at  first.  Its  position  remains  the  same  throughout.  As 
near  as  the  eye  can  judge,  this  band  has  had  its  intensity  reduced 
to  about  half-value  when  the  fourth  strip  is  reached,  corresponding 
to  8  per  cent,  of  water  in  the  solution.  The  alcohol  band  at 
A  4292  also  has  about  50  per  cent,  of  its  final  intensity  in  the  same 
solution. 

The  band  at  A  4760  shows  the  same  kind  of  a  change  that  we 
described  in  some  detail  above,  only  the  change  is  much  more  gradual 
and  easy  to  follow  here.  It  also  shows  about  equal  intensity  for  the 
two  sets  of  bands  when  the  amount  of  water  is  8  per  cent,  of  the 
whole. 

The  green  and  yellow  bands  are  not  sufficiently  resolved  in  a 
to  allow  the  change  in  the  individual  bands  to  be  followed,  and 
hence  these  apparently  show  only  a  gradual  shift  towards  the  red 
with  decrease  in  the  amount  of  water.  In  b,  however,  they  are 
sufficiently  resolved  to  enable  us  to  follow  the  change  in  each 
individual  band,  which,  although  a  little  difficult,  on  account  of 
their  large  number  and  the  incompleteness  of  their  separation  in 
some  cases,  may  still  be  done.  The  change  is  in  every  respect  the 
same  as  we  have  found  for  the  other  bands;  namely,  those  due 


290  JONES  AND  ANDERSON— ABSORPTION   SPECTRA  OF     [Aprilas, 

to  the  aqueous  solution  diminish  in  intensity  and  reach  about  half 
value  in  the  8  per  cent,  aqueous  solution,  while  those  belonging  to  the 
alcoholic  solution  increase  in  intensity,  as  the  amount  of  water 
is  decreased. 

The  band  in  the  red  near  A  6800  shows  the  change  very  well 
indeed,  the  "  water  "  band  having  the  position  A  6800,  while  that 
pertaining  to  the  alcoholic  solution  is  situated  at  A  6860,  and  hence 
the  two  are  well  separated.  Here  the  point  of  equal  intensity 
appears  to  be  reached  in  the  solution  containing  io|  per  cent,  of 
water,  but  this  is  due  to  the  fact  that  the  alcoholic  band  has  a 
considerably  greater  intensity  than  that  due  to  the  aqueous  solu- 
tion, conditions  as  to  concentration  and  depth  of  layer  being  the 
same.  Taking  this  into  account  it  is  seen  that  this  band  obeys 
substantially  the  same  rule  as  the  others. 

The  change  in  the  band  at  X  7325  is  more  difficult  to  follow,  on 
account  of  the  small  intensity  of  the  photographic  action  on  the 
less  refragible  side  of  this  position.  The  band  belonging  to  the 
aqueous  solution  may  be  seen  very  clearly  even  in  the  strip  corre- 
sponding to  the  2§  per  cent,  water  solution,  but  is  of  course  en- 
tirely absent  in  the  alcoholic  solution.  Its  intensity  in  the  2§  per 
cent,  solution,  however,  seems  a  little  greater  than  we  should  ex- 
pect from  the  behavior  of  the  other  bands,  but  this  is  perhaps  due 
to  the  rather  weak  photographic  action  in  this  part  of  the  spectrum, 
combined  with  the  great  intrinsic  intensity  of  the  band.  The  alco- 
holic solution  transmits  light  as  far  as  A  7355  where  its  spectrum 
ends  abruptly  in  a  band. 

Throughout  this  description  we  have  laid  great  stress  on  the 
fact  that  on  plate  4  the  two  sets  of  bands  coexist ;  the  bands  due  to 
the  aqueous  solution  decreasing,  while  those  belonging  to  the  alco- 
holic solution  increase  in  intensity  with  decrease  in  the  percent- 
age of  water.  We  have  also  called  attention  to  the  fact  that  the 
two  sets  of  bands  have  about  one-half  their  full  intensity  in  a  solu- 
tion containing  about  8  per  cent,  of  water.  This  was  for-  a  0.5 
normal  solution. 


i9o8.]  NEODYMIUM  AND  PRASEODYMIUM.  291 

Praseodymium  Chloride  in  Water — Beer's  Law. 

(See  plate  5.) 

The  concentrations  of  the  solutions  used  in  making  the  nega- 
tive for  a,  beginning  with  the  one  whose  spectrum  is  adjacent  to 
the  numbered  scale  were  2.56,  1.92,  1.25,  0.85,  0.60,  042  and  0.32. 
For  b  the  concentrations  were  0.85,  0.63,  0.42,  0.28,  0.2b,  0.14  and 
0.1 1 ;  the  depths  of  absorbing  layer  being,  respectively,  3,  4,  6,  9,  13, 
18  and  24  mm. 

The  solutions  of  praseodymium  chloride  are  all  green  or  yellow- 
ish green,  only  the  intensity  of  the  color  changing  with  change  in 
the  concentration. 

For  these  solutions  Beer's  Law  holds  very  exactly,  excepting 
for  the  extreme  ultra-violet  absorption  in  a,  and  the  yellow  bands  in 
the  two  or  three  most  concentrated  solutions  of  a. 

The  limits  of  transmission  in  the  ultra-violet  for  the  most  con- 
centrated and  most  dilute  solutions  of  a  are,  respectively,  X2720 
and  A  2650.  The  edge  is  fairly  sharp,  indicating  the  presence  of 
a  rather  intense  band.  This  is  also  indicated  by  b,  where  the  spec- 
trum ends  abruptly  at  A  2630,  the  limit  being  the  same  for  all  the 
solutions. 

The  absorption  bands  shown  in  a  are  as  follows:  A 4380  to 
X4480,  strong  band  with  red  edge  somewhat  shaded;  X4640  to 
X4710,  sharp  on  red  side,  quite  diffuse  towards  the  violet;  A 4800 
to  A 4830,  sharply  defined  on  both  sides;  A 5860  to  A 5950,  both 
edges  diffuse ;  A  5985,  fairly  narrow  band  with  diffuse  edges.  The 
region  between  this  band  and  the  principal  yellow  one  shows  very 
strong  absorption. 

b  shows  the  following':  A 4410  to  A 4465,  both  edges  a  little  dif- 
fuse ;  A  4685,  fairly  narrow  band,  still  more  diffuse  towards  the  vio- 
let, although  somewhat  shaded  also  towards  the  red;  A 4815,  narrow 
band  with  edges  slightly  shaded ;  A  5900,  wide  hazy  band ;  absorp- 
tion not  complete  even  at  its  middle ;  A  5985,  rather  faint,  hazy  band. 

The  greenish  tinge  of  the  solutions  would  suggest  that  there  is 
considerable  general  absorption  in  the  red,  because  the  absorption 
in  the  yellow  is  not  sufficient  to  impart  any  marked  color  to  the  solu- 
tion, and  the  bands  in  the  violet  and  blue  could  only  give  it  a  yellow 


292  JONES  AND  ANDERSON— ABSORPTION   SPECTRA  OF    [April  ts. 

tint.  The  negative  for  a  does,  in  fact,  show  pretty  strong  general 
absorption  from  k  7100  to  the  end  of  the  red,  but  no  doubt  a  spectro- 
photometric  study  of  the  solutions  would  show  general  absorption 
much  farther  down  into  the  red.  The  negative  for  b  shows  no  sign 
of  this  absorption  for  very  obvious  reasons. 

Praseodymium  Chloride  in  Mixtures  of  the  Alcohols  and 

Water. 

(See  plate  6.) 

The  concentrations  of  the  praseodymium  chloride  was  constant 
throughout  and  equal  to  0.5  normal.  The  percentages  of  water  in 
the  solutions,  beginning  with  the  one  whose  spectrum  is  adjacent 
to  the  numbered  scale,  were  o,  2J,  5I,  8,  loj,  13J  and  16.  The 
depth  of  absorbing  layer  was  i.o  cm. 

Methyl  alcohol  was  the  chief  solvent  in  the  solutions  pertaining 
to  a,  while  ethyl  alcohol  was  used  in  the  solutions  used  in  making 
the  negative  for  b.  The  two  spectrograms  are  identical,  except  for 
a  little  greater  general  absorption  in  the  ultra-violet  with  the  ethyl 
alcohol. 

The  most  striking  feature  of  the  spectrograms  is  the  appearance 
of  the  intense  absorption  band  near  X3000  as  the  percentage  of 
water  is  g^dually  decreased.  Only  a  faint  trace  of  this  band  is 
visible  with  16  per  cent  of  water  in  the  solution,  and  the  band  is 
comparatively  weak  even  with  only  8  per  cent,  of  water.  From 
this  point  it  increases  very  rapidly  in  width  and  intensity  with 
decrease  in  the  amount  of  water,  until  in  the  pure  alcohol  solutions 
its  limits  (transmission)  are  X2970  and  X3230,  being  by  far  the 
most  intense  band  in  the  whole  spectrtun. 

The  bands  in  the  violet  and  blue  apparently  shift  somewhat 
towards  the  red,  this  being,  however,  due  to  the  fact  that  the  alco- 
hol bands  are  a  little  nearer  the  red  end  of  the  spctrum,  and  that 
when  the  percentage  of  water  changes  from  16  to  o,  the  two  sets 
of  bands  coexist,  but  are  far  from  being  separated.  The  change  is 
exactly  the  same  in  character  as  the  one  described  in  detail  in  dis- 
cussing the  X4760  band  in  mixtures  of  alcohol  and  water  for 
neodymium  chloride.  The  positions  of  the  bands  in  the  solution, 
containing  16  per  cent,  of  water,  are  as  follows:  X4390  to  X4470, 


,9o8.]  NEODYMIUM  AND  PRASEODYMIUM.  293 

X  4660  to  A  4700,  X  4800  to  X  4825.  In  the  solution  in  pure  alcohol 
they  are  X  4410  to  X  4480,  X  4690  to  X  4715,  X  4810  to  X4840.  Hence 
it  appears  that  the  two  most  refragible  bands  have  a  slightly  greater 
width  in  the  aqueous  solution,  while  the  X4815  band  is  more  intense 
in  the  alcoholic  solutions. 

The  bands  in  the  yellow  show  very  well,  indeed;  the  fact  that 
here  as  in  the  spectrum  of  neodymium  chloride  we  have  the  coex- 
istence of  two  sets  of  bands  when  the  water  content  of  a  one-half 
normal  solution  is  in  the  neighborhood  of  8  per  cent.  The  band 
in  the  yellow  has  already  been  described  under  Beer's  Law,  but  as 
the  concentration  and  depth  of  layer  is  different  here,  the  following 
will  serve  to  indicate  what  the  spectrum  of  the  16  per  cent,  aqueous 
solution  shows. 

Absorption  begins  at  X5850  and  rises  to  a  maximum  at  about 
X  5900,  then  decreases  to  a  minimum  at  X  5950,  from  which  it  again 
rises  to  a  maximum  at  about  X  5980,  falling  off  to  zero  at  X  6000. 
The  solution  in  pure  alcohol  shows  the  following :  Weak  absorption 
begins  at  X5800,  and  continues  without  material  change  up  to 
X5880,  where  it  falls  almost  to  nothing.  At  X5900  il  begins  to 
increase  and  reaches  a  strong  maximum  at  X  5955,  falling  oflf  gradu- 
ally to  zero  at  X6000.  The  intermediate  solutions  show  the  gradual 
disappearance  of  the  bands  characteristic  of  the  aqueous  solution, 
and  the  increase  in  intensity  of  those  belonging  to  the  alcoholic 
solution  as  the  percentage  of  water  is  gradually  decreased.  The 
maximum  change  takes  place  from  the  fifth  to  the  third  strips, 
counting  from  the  numbered  scale,  indicating  here  as  with  neody- 
mium chloride  that  the  two  sets  have  about  half  their  normal  in- 
tensity when  the  water  content  of  the  solution  is  about  8  per  cent., 
or  when  the  solution  contains  about  ten  molecules  of  water  per 
molecule  of  the  dissolved  substance. 

Discussion  of  the  Results. 

The  results  established  by  these  plates  may  be  briefly  summar- 
ized as  follows: 

1.  The  absorption  spectra  of  a  salt  in  different  solvents  are,  in 
general,  different. 

2.  When  a  salt  is  dissolved  in  mixtures  of  two  solvents  the 


294        ■   JONES  AND  ANDERSON— ABSORPTION  SPECTRA  OF    [Aprils, 

relative  percentages  of  which  are  varied,  there  is  not  a  gradual 
change  of  one  spectrum  into  the  other,  but  the  spectrum  given  in 
the  mixture  is  a  superposition  of  the  two  spectra,  the  two  sets  of 
bands  existing  together.  If  the  salt  is  one  whose  spectrum  changes 
considerably  with  its  state  of  dissociation,  we  have  in  addition  to 
the  above  phenomena  the  changes  due  to  the  varying  dissociation 
of  the  dissolved  salt  produced  by  the  varying  composition  of  the 
mixture. 

A  study  of  all  the  plates  (eighty  in  number)  obtained  in  this 
work  shows  that  deviations  from  Beer's  Law  is  the  rule  rather 
than  the  exception;  only  a  limited  number  obey  Beer's  Law  even 
approximately.  Beer's  Law  could  only  hold  in  cases  where  the 
relative  concentrations  of  the  different  kinds  of  absorbers  in  solu- 
tion do  not  change  with  the  dilution,  or  where  the  different  kinds 
of  absorbers  have  the  same  kind  of  absorption.  The  first  condition 
is  perhaps  never  realized,  while  the  second  is  undoubtedly  closely 
approached  with  sucji  salts  as  neodymium  chloride  and  praseody- 
mium chloride. 

The  rule  is  that  the  different  absorbers  have  different  absorbing 
powers,  and  the  problem  of  absorption  spectra  is  to  determine  which 
kind  of  absorbers  in  solution  are  responsible  for  the  different  bands. 

The  theory  of  Ostwald,  which  would  refer  absorption  in  solution 
mainly  to  the  ions  present,  has  been  found  to  be  entirely  insufficient 
to  account  for  the  facts  established  in  this  investigation. 

The  other  theories  which  aim  to  account  for  the  deviations  are 
of  two  kinds,  viz. : 

1.  Those  that  assume  that  the  increased  absorption  in  concen- 
trated solutions  is  due  to  the  formation  of  aggregates  of  the  mole- 
cules of  the  dissolved  substance,  or  of  the  molecules  and  the  ions 
into  which  they  break  down  in  dissociation. 

2.  Those  that  assume  that  the  deviation  is  due  to  the  formation 
of  solvates,  that  is,  combinations  of  the  parts  of  the  dissolved  sub- 
stance with  the  molecules  of  the  solvent. 

Now,  it  has  been  shown  by  Hartley  and  other  workers,  who 
have  studied  the  change  in  the  absorption  with  change  in  tempera- 
ture, that  the  bands  which  widen  with  increase  in  concentration 
(conditions  for  Beer's  Law  assumed  to  obtain)   also  widen  with 


,9o8.]  NEODYMIUM   AND   PRASEODYMIUM.  295 

rise  in  temperature;  that  is,  a  rise  in  temperature  produces  very 
much  the  same  effect  as  increase  in  concentration.  This  seems  to 
us  pretty  conclusive  evidence  against  the  theories  which  are  based 
on  the  formation  of  aggregates,  for  it  is  well  known  that  the  change 
in  the  aggregates  produced  by  rise  in  temperature  is  not  the  same 
as  that  produced  by  increase  in  concentration,  but  exactly  the 
opposite. 

The  theories  which  assume  the  formation  of  solvates  are  not 
open  to  this  objection,  because  it  is  well  known  that  the  change  in 
the  solvates  produced  by  rise  in  temperature  is  in  general  the  same 
as  that  produced  by  increase  in  concentration.  As  a  solution  be- 
comes more  concentrated  the  solvates  become  simpler  and  simpler, 
that  is,  fewer  molecules  of  the  solvent  are  combined  with  each  part 
of  the  dissolved  substance.  Rise  in  temperature  also  breaks  down 
complex  solvates  into  simpler  ones.  Of  course,  it  does  not  follow 
that  the  solvates  of  a  solution  of  concentration  c^  at  temperature  tj^ 
are  exactly  the  same  as  those  in  a  solution  of  concentration  c^  at  a 
temperature  /,;  since  under  the  changed  conditions  it  may  happen 
that  the  particular  solvates,  which  were  most  stable  when  the  con- 
ditions were  c^  and  t^,  may  be  less  stable  than  solvates  of  nearly 
the  same  composition  at  C2,  t^. 

For  this  reason  and  also  because  our  work  on  neodymium  and 
praseodymium  salts  in  mixed  solvents  seems  almost  conclusive  evi- 
dence in  favor  of  the  existence  of  solvates,  we  have  used  the  solvate 
theory  as  a  working  hypothesis  throughout  this  work.  That  it  is 
not  far  from  being  correct  is  shown  by  the  fact  that  all  the  phenom- 
ena observed  in  the  great  number,  about  1,200,  of  solutions  studied, 
are  accounted  for  without  anything  but  the  simplest  assumptions  in 
regard  to  the  behavior  of  the  solvates  in  question. 

The  most  interesting  and  important  results  were  obtained  from 
the  study  of  the  salts  of  neodymium  and  praseodymium,  especially 
those  of  the  former.  These  substances  have  not  only  very  many 
absorption  bands,  but  they  are  remarkably  narrow  and  sharp,  and, 
hence,  peculiarly  suitable  for  spectrographic  study.  The  chief  ex-  • 
perimental  results  were  the  following: 

I.  The  absorption  spectrum  of  aqueous  solutions  of  the  chloride 
and  bromide  of  neodymium  changes  very  little  with  change  in  con- 


296        JONES  AND  ANDERSON— ABSORPTION  SPECTRA  OF         [April  •$ 

centration,  and  the  two  are  nearly  identical,  throughout,  excepting 
for  the  fact  that  the  absorbing  power  of  the  bromide  appears  to  be 
somewhat  greater  than  that  of  the  chloride. 

2.  Solutions  of  the  salts  in  non-aqueous  solvents  give  spectra 
which  are  not  only  different  for  different  salts,  but  the  spectrum 
of  any  one  salt  is  different  in  the  different  solvents.  An  apparent 
exception  is  the  spectrum  of  neodymium  or  praseodymium  chloride 
in  methyl  and  ethyl  alcohols,  which  are  almost  exactly  alike. 

3.  When  a  salt  like  neodymium  chloride  is  dissolved  in  mixtures 
of  water  and  one  of  the  non-aqueous  solvents,  and  the  relative 
amounts  of  the  two  solvents  in  the  mixture  is  varied,  no  marked 
change  in  the  spectnun  is  observed  when  the  amount  of  water  is 
changed  from  100  per  cent,  to  about  15  or  20  per  cent.  As  the 
amount  of  water  is  still  further  reduced  we  find  that  the  solution 
gives  a  spectrum  which  consists  of  a  superposition  of  the  spectra 
belonging  to  the  aqueous  and  the  non-aqueous  solutions ;  the  former 
decreasing  in  intensity  while  the  latter  increases  as  the  amount  of 
water  is  decreased.  The  composition  of  the  mixed  solvents,  which 
will  show  the  two  spectra  with  about  one-half  their  normal  inten- 
sity, depends  upon  the  concentration  of  the  salt  in  solution;  and  a 
constant  ratio  between  the  number  of  molecules  of  water  and  those 
of  the  dissolved  salt  were  indicated  by  the  experiments,  this  ratio 
having  the  value  10. 

Praseodymium  chloride,  dissolved  in  mixtures  of  water  and 
methyl  or  ethyl  alcohol,  shows  in  general  the  same  kind  of  change 
in  the  spectrum  as  neodymium  chloride ;  but  in  addition  there  appears 
in  the  alcoholic  solutions  an  entirely  new  band  having  no  analogue 
in  the  aqueous  solution.  In  the  former  this  new  band  in  the  ultra- 
violet is  by  far  the  most  intense  in  the  entire  spectrunu  It  disap- 
pears entirely  on  addition  of  water,  having  about  half  its  normal 
intensity  for  a  half  normal  solution  when  the  water  content  of  the 
solvent  is  about  8  per  cent. 

These  facts  seem  to  us  inexplicable  on  any  other  hypothesis  than 
the  one  we  have  made,  namely,  that  when  a  salt  of  one  of  these 
elements  is  dissolved  in  any  solvent,  both  the  molecules  of  the  salt 
and  the  ions  formed  from  these  become  solvated,  that  is,  they  com- 
bine with  a  certain  number  of  molecules  of  the  solvent.    While  in 


,9o8.]  NEODYMIUM  AND   PRASEODYMIUM.  297 

the  case  of  some  salts  the  spectra  point  to  the  existence  of  solvates 
of  varying  complexity,  in  the  case  of  salts  of  neodymium  and 
praseod)miium  they  indicate  rather  the  existence  of  one  definite 
hydrate.  A  more  extended  study,  including  the  changes  in  the 
spectra  produced  by  changes  in  temperature,  may,  however,  some- 
what modify  this  conclusion. 

Granting  the  existence  of  solvates  all  of  the  facts  observed  in 
connection  with  the  absorption  spectra  of  neodymium  and  praseody- 
mium salts  can  be  readily  explained. 

Physical  Chemical  Laboratory, 
Johns  Hopkins  University, 
June,  1908. 


PRELIMINARY  REPORT  UPON  A  CRYSTALLOGRAPHIC 

STUDY  OF  THE  HEMOGLOBINS :  A  CONTRIBUTION 

TO  THE  SPECIFICITY  OF  CORRESPONDING 

VITAL  SUBSTANCES  IN  DIFFERENT 

VERTEBRATES. 

By  EDWARD  T.  REICHERT  and  AMOS  P.  BROWN. 

(Read  April  24,  1908.) 

The  primary  object  of  this  research  was  to  determine  whether 
or  not  corresponding  proteins  are  identical  in  different  species. 
Hemoglobin  was  selected  as  a  favorable  substance  to  begin  such  a 
study  upon  because  of  its  being  readily  obtained  in  a  state  of  com- 
parative purity,  and,  in  many  cases,  readily  isolated  in  crystals. 
When  a  sufficient  supply  of  blood  wa3  available,  it  was  nearly  always 
possible,  by  the  use  of  suitable  methods,  to  produce  well  formed 
crystals  that  could  be  satisfactorily  examined  and  studied  by  the 
method  adopted.  The  crystallographic  method  was  chosen  because, 
by  its  means,  differences  in  substances  may  be  observed  that  would 
elude  the  ordinary  methods  of  analysis  employed  by  the  chemist. 
Moreover,  it  is  comparatively  rapid  and  therefore  well  adapted  to 
the  study  of  a  substance  so  liable  to  alteration  as  hemoglobin.  In 
the  method  employed  it  was  not  even  necessary  to  remove  the  crys- 
tals from  the  mother  liquor  for  examination.  In  studying  the  crys- 
tals and  measuring  the  crystallographic  constants  the  petrographic 
microscope  was  used,  but  in  the  case  of  these  crystals  of  hemoglobin 
we  have  this  advantage  over  the  petrographer  in  his  examination 
of  rock  sections,  in  that  these  crystals  are  not  imbedded  in  an  opaque 
or  semi-opaque  matrix,  but  are  in  a  transparent  medium  and  are 
usually  isolated  from  each  other.  Moreover,  hundreds  and  often 
thousands  of  crystals  are  open  to  observation  in  a  single  slide, 
and  these  present  almost  all  possible  orientations,  allowing  the  opti- 
cal characters  to  be  determined  with  much  greater  accuracy  than 

298 


,9o8  ]       A  CRYSTALLOGRAPHIC  STUDY  OF  THE  HEMOGLOBINS.       299 

is  usually  the  case  with  minerals  in  rock  sections.  Measurements 
within  the  limit  of  error  of  the  instrument  could  frequently  be  ob- 
tained, and,  as  various  orientations  were  available,  the  results  of 
the  angular  measurements  often  furnished  complete  data  for  the 
calculation  of  the  axial  ratios.  The  crystals  examined  were  usually 
complete  and  often  geometrically  perfect,  so  that  the  symmetry  and 
'  crystal  habit  could  be  determined  as  readily  as  in  the  case  of  ordi- 

nary mineral  substances  occurring  in  isolated  crystals. 

A  chemical  substance,  possessing  a  rational  composition,  tends 
to  arrange  its  parts  in  an  orderly  manner  so  that  a  definite  struc- 
ture is  assumed,  which  results  in  a  definite  external  form.  This  is 
so  universally  true  that  the  crystalline  condition  is  the  normal  one 
for  matter  of  definite  composition.  Differences  of  crystalline  form 
hence  indicate  differences  of  substance ;  and,  by  the  crystallographic 
method  of  investigation,  obscure  differences,  such  as  those  between 
isomerides,  may  readily  be  detected. 

Photographic  records  of  the  crystals  were  secured  and  upwards 
of  2,500  negatives  have  been  made.  The  hemoglobins  of  more  than 
one  hundred  species  have  been  examined  and  data  secured  in  regard 
to  their  crystals.  From  a  study  of  these  records  certain  facts  stand 
out  very  prominently. 

I.  The  Constancy  of  Generic  Characters  in  the  Crystals, — The 
crystals  of  the  species  of  any  genus  belong  to  a  crystallographic 
group.  When  their  characters  are  tabulated,  they  at  once  .recall 
the  crystallographic  groups  of  minerals.  The  crystals  of  the  genus 
Felis  iorm  an  isomorphous  group;  as  strictly  isomorphous,  in  fact, 
as  the  group  of  the  rhombohedral  carbonates  among  minerals.  The 
genus  Cams  is  even  more  strictly  isomorphous,  but  the  crystals  of 
hemoglobin  from  the  two  genera  are  perfectly  distinct,  the  one  from 
the  other. 

As  an  example  of  the  individuality  of  these  generic  characters 
the  following  may  be  cited :  A  sample  of  blood,  marked  as  that  of 
a  certain  species  of  baboon  was  received  from  one  of  our  Zoological 
Gardens.  Upon  making  preparations  and  examining  the  crystals,  it 
was  at  once  evident  that  they  did  not  correspond  to  any  species  of 


300     REICHERT  AND  BROWN— PRELIMINARY  REPORT  UPON    (April  34, 

baboon  thus  far  examined,  nor  did  they  show  the  characters  of  the 
genus  Papio.  They  were  identified  by  their  crystallogfraphic  char- 
acters as  belonging  to  the  cats  (genus  Felts)  but  not  to  any  species 
that  we  had  examined  up  to  that  time.  Inquiry  at  the  2^1ogical 
Garden  from  which  the  blood  was  received  showed  that  the  animal 
recorded  as  being  subjected  to  a  post-mortem  examination  on  the 
date  when  the  blood  was  collected  was  a  species  of  the  genus  Felis, 
but  not  one  of  which  we  had  previously  examined  the  blood.  Other 
similar  cases  of  incorrect  labelling  of  specimens  were  detected,  in 
which  the  wrongly  labelled  blood  was  one  that  had  been  examined 
and  the  species  known  from  other  specimens. 

2.  Specificity  in  the  Crystals  of  a  Genus, — ^The  crystals  of  the 
different  species  of  a  genus,  when  they  are  favorably  developed  for 
good  measurement,  can  usually  be  distinguished  from  each  other  by 
definite  differences  of  angle,  etc. ;  while  preserving  their  isomorphous 
character  as  belonging  to  a  definite  genus.  In  cases  where,  on 
account  of  difficulty  of  measurement,  the  differences  cannot  be  given 
a  quantitative  value,  variations  in  the  habit  of  the  crystals  and  in 
their  mode  of  growth  will  often  show  specific  differences. 

•3.  The  Occurrence  of  Several  Types  of  Crystals  of  Oxyhemo- 
globin in  Many  Species. — In  some  species  the  oxyhemoglobin  is 
dimorphous  (crystallizing  in  two  systems  or  with  two  axial  ratios), 
in  other  tases  even  trimorphous.  Where  several  types  of  crystal 
occur  in  this  way  in  the  species  of  any  genus,  the  crystals  of  each 
type  may  be  arranged  in  an  isomorphous  series.  In  other  words, 
certain  genera  are  isodimorphous  or  isotrimorphous. 

4.  The  Constant  Recurrence  of  Certain  Angles,  Plane  or  Dihe^ 
drcd,  in  the  Oxyhemoglobin,  Hemoglobin  and  the  "  Methemoglo- 
bins  "  of  Various  Species,  even  when  these  Species  are  Widely  Sep- 
arated Zoologically  and  when  their  Crystals  Belong  to  Various  Crys- 
tal Systems. — This  appears  to  indicate  a  common  substance  in  hemo- 
globins or  a  common  structure  in  the  various  hemoglobin  molecules. 

5.  The  Constant  Recurrence  of  Certain  Types  of  Twinning  in 
the  Hemoglobins,  and  the  Prez^alence  of  Mimosie  in  these  Crystals. 
— ^This  also  indicates  a  common  structure  in  the  various  hemoglobin 
molecules. 


«9o8.l       A  CRYSTALLOGRAPHIC  STUDY  OF  THE  HEMOGLOBINS.       301 

6.  Differences  between  Oxyhemoglobin  and  Reduced  Hemoglobin 
in  Certain  Species, — ^Undoubted  diflferences  between  the  crystals  of 
these  two  substances  in  the  same  species  have  been  observed. 

We  have  gathered  additional  evidence  that  other  corresponding 
proteins,  as  well  as  certain  fats  and  carbohydrates,  will  be  found  to 

exhibit  similar  specificities. 

UNivERsmr  OP  Pennsylvania. 
AprU  23,  1908. 


PROC.  AMKR.  PHIL.  SOC.,  XLVII.  189  T,  PRINTED  SEPTEMBER  26,  I908. 


INFLUENCE  OF  PRESERVATIVES  AND  OTHER 

SUBSTANCES  ADDED  TO   FOODS  UPON 

HEALTH  AND  METABOLISM. 

By  HARVEY  W.  WILEY,  M.D. 
(Read  April  25,  /poS.) 

In  connection  with  studies  of  food  adulteration,  which  have  been 
conducted  during  the  past  twenty-five  years  under  my  direction  in 
the  Bureau  of  Chemistry,  frequent  evidence  was  obtained  of  the 
addition  of  certain  preserving  agents  and  coloring  matters  to  food 
products.  These  bodies  are  not  of  the  character  known  as  condi- 
mental ;  on  the  contrary,  as  a  rule,  they  possess  neither  appreciable 
taste  nor  odor  in  the  quantities  in  which  they  are  employed. 

In  so  far  as  preservatives  are  concerned,  therefore,  the  consumer 
would  have  no  certain  knowledge  of  their  presence,  and  in  respect 
to  coloring  matters,  he  would  likewise  be  ordinarily  deceived,  since 
such  coloring  matters  are  often  used  to  imitate  the  natural  tints 
found  in  food  products.  Thus  there  would  be  practiced  upon  the 
consumer  a  fraud  in  that  in  the  purchase  and  consumption  of  foods 
he  was  buying  and  consuming  articles  which  are  distinctly  not  foods 
and  the  presence  of  which  is  a  just  cause  of  suspicion. 

The  use  of  chemical  preservatives  and  artificial  colors  in  foods 
IS  of  quite  recent  date.  I  think  I  may  say  with  safety  that  if  one 
could  go  back  thirty,  or  at  most,  forty  years,  he  would  find  a  food 
supply  practically  free,  both  from  chemical  preservatives  and  artifi- 
cial colors.  The  rapid  development  of  organic  and  tinctorial  chem- 
istry during  the  past  forty  years  has  made  it  possible  to  offer  to 
manufacturers  chemical  preservatives  of  high  potency,  and  colors 
of  great  beauty  and  persistence,  at  prices  which  make  it  entirely 
possible  to  use  them  freely  in  food  products.  Inasmuch  as  the  use 
of  these  bodies,  whatever  the  claims  may  be  in  regard  thereto,  has 
for  its  chief  purpose  either  to  cheapen  the  product  itself  or  to  sell 

302 


'f 


,908.]  UPON   HEALTH  AND  METABOLISM.  303 

it  at  a  higher  price  than  it  really  should  command,  it  is  evident  that 
unless  the  pecuniary  conditions  attending  the  use  of  these  bodies 
were  favorable  they  would  not  be  employed. 

When  the  claims  which  are  made  by  manufacturers  respecting 
the  use  of  these  substances  are  carefully  considered,  we  find  that 
most  of  them  are  without  foundation.  In  regard  to  the  supposed 
general  preference  for  artificial  color,  I  would  say  that  an  experi- 
ment performed  on  a  large  number  of  totally  unbiased  people  has 
convinced  me  beyond  any  reasonable  doubt  that  the  great  majority 
of  American  consumers  would  prefer  uncolored  foods.  The  ex- 
periment mentioned  was  made  on  about  sixty  different  men  during 
a  period  of  five  years  to  determine  whether  or  not  they  preferred 
an  artificially  colored  food  or  one  in  its  natural  tint.  Butter,  which 
is  perhaps  the  one  food  product  most  universally  colored  in  this 
country,  was  used.  The  subjects  on  whom  the  experiment  was  tried 
had  been  in  the  habit  of  using  nothing  except  colored  butter,  hence, 
if  there  was  any  prejudice  existing  in  their  minds  it  must  have  been 
in  favor  of  the  article  which  they  had  constantly  consumed.  More- 
over, the  test  was  made  in  the  winter  time  when  the  uncolored  but- 
ter has  the  least  tint  of  the  whole  year,  being  almost  white.  No 
attempt  was  made  to  inform  the  men  of  the  nature  of  these  prod- 
ucts. The  natural  butter  and  the  colored  butter  were  moulded  in 
the  same  forms  and  placed  upon  the  same  plate,  and  offered  with- 
out comment  of  any  kind.  At  first  very  few  of  the  men  would  do 
more  than  look  at  the  uncolored  butter.  A  very  common  expres- 
sion was,  "  This  is  oleomargarine."  A  few  made  a  trial  of  its 
properties.  Little  by  little,  without  any  propaganda  of  any  kind, 
the  whole  attitude  of  these  men  changed.  In  the  course  of  four 
or  five  months  nine  tenths  of  them  were  using  the  uncolored  butter 
and  they  expressed  a  most  decided  antipathy  to  the  use  of  the  col- 
ored butter  when  at  certain  times  the  supply  of  the  uncolored  butter 
was  exhausted. 

I  believe  that  this  completely  refutes  the  arguments  of  those  who 
claim  that  they  color  butter  to  meet  the  demand  of  the  consumer. 
In  point  of  fact,  the  color  in  butter  has  been  almost  from  the  first 
a  fraudulent  process.  It  is  a  common  belief  that  the  best  butter  of 
the  year  is  produced  during  the  early  spring  months,  and  especially 


304  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  as, 

in  June,  when  the  cows  have  access  to  the  succulent  pastures.  Dur- 
ing this  time,  owing  to  the  oxidation  of  the  chlorophyll  of  the  grass, 
a  xanthophyll  is  produced,  imparting  to  the  cream  a  rich  golden, 
or  yellow,  tint  which  is,  of  course,  perpetuated  in  the  butter.  Dur- 
ing the  winter  months,  when  the  chlorophyll  is  withdrawn  practi- 
cally from  the  diet  of  the  cow,  this  natural  coloring  matter  is  absent. 
The  use  of  the  artificial  color,  therefore,  is  to  simulate  for  winter 
butter  the  color  of  the  butter  in  June,  and  thus  to  conceal  what  is 
at  least  believed  to  be  inferiority. 

Again,  in  experimental  observations  of  a  less  extended  oharacter, 
I  have  found  that  the  American  consumer  does  not  prefer  his  foods 
preserved  with  chemical  preservatives.  In  a  large  number  of  in- 
stances which  have  come  under  my  own  personal  observation  the 
consumer  has  stopped  eating  an  article  as  soon  as  he  has  found  that 
it  contains  a  chemical  powerful  enough  to  inhibit  fermentative 
action.  The  users  of  chemical  preservatives,  however,  do  not  as 
a  rule  claim  that  they  use  them  at  the  demand  of  the  consumer. 
A  careful  study  of  manufacturing  data  made  by  one  of  the  most 
conscientious  manufacturers  in  the  West  shows  that  it  costs  more 
to  make  a  food  product  without  a  preservative  than  it  does  with  a 
preservative.  In  very  extensive  practical  experiments  on  tomatoes 
this  manufacturer  found  that  it  was  necessary  to  charge  from  fif- 
teen to  twenty  cents  more  for  ketchup  per  case  made  without  a 
preservative  than  with  a  preservative.  Thus  I  think  it  is  well 
established  by  this  experimental  study  that  the  real  reason  which 
the  manufacturers  have  for  using  chemical  preservatives  is  to 
cheapen  the  cost  of  production.  This  of  itself  would  be  a  most 
worthy  object,  because  presumably  the  cheapening  of  the  cost  of 
production  would  lower  the  price  to  the  consumer.  If,  therefore, 
a  food  product  of  equal  nutritive  value  and  equal  wholesomeness 
could  be  produced  with  the  aid  of  chemical  preservatives,  such  a 
process  should  meet  with  the  approbation  of  all.  But  a  very  seri- 
ous problem  of  a  different  kind  is  presented  here.  A  chemical  pre- 
servative is  eflFective  usually  by  reason  of  its  inhibitive  action  on 
fermentation.  Very  extensive  studies  of  this  action  of  chemical 
preservatives  have  led  to  the  general  conclusion  that  while  these 
bodies  inhibit  the  fermentative  action  giving  rise  to  the  ordinary 


,9o8j  UPON   HEALTH  AND   METABOLISM.  305 

evidences  of  decay  and  putrefaction,  and,  as  a  rule,  stop  most  effec- 
tively those  fermentations  which  produce  alcohol  and  carbon  dioxid, 
they  do  not  have  the  same  restrictive  influence  on  those  processes 
resulting  in  the  general  degradation  and  decay  of  organic  matter, 
due  chiefly  to  that  class  of  chemical  reactions  which  is  represented 
by  the  term  hydrolysis.  In  other  words,  the  ferments  which  break 
down,  for  instance,  nitrogeneous  tissues  into  more  soluble  and  finally 
more  dangerous  forms  of  combination,  are  not  so  particularly  in- 
hibited as  is  the  first  class  of  ferments  mentioned. 

This  fact  might  well  be  used,  however,  as  a  justification  of  the 
employment  of  chemical  preservatives,  since  if  they  prevent  the 
ordinary  processes  of  fermentation  which  produce  evident  indica- 
tions of  decay  and  putrefaction,  it  might  be  held  that  they  would 
not  interfere  with  that  other  class  of  fermentations  or  hydrolytic 
processes  peculiarly  exercised  by  the  digestive  ferments.  It  will 
probably  not  be  contested  at  the  present  time  that  there  is  some 
justification  for  this  plea,  since  it  has  been  well  established  that 
an  amount  of  a  preservative  which  will  for  instance  prevent  alco- 
holic fermentation  will  not  interfere  in  anything  like  so  serious 
a  manner  with  the  action  of  such  ferments  as  the  diastatic  fer- 
ments of  the  saliva,  of  the  stomach,  and  of  the  pancreas.  On  the 
other  hand,  it  is  well  established  that  in  any  notable  quantities  these 
preservatives  do  interfere  with  even  the  latter  class  of  ferments. 

But  the  problem  which  is  of  most  importance  in  this  con- 
nection is.  What  is  the  chief  effect  of  these  preservatives  upon  the 
health  of  those  who  constantly  use  them  and  upon  the  metabolism  re- 
sulting from  the  normal  functions  of  the  body?  To  answer  this 
question,  there  was  begun  in  an  experimental  way  in  the  Bureau  of 
Chemistry,  under  my  direction,  a  few  years  ago,  a  series  of  studies 
having  for  their  purpose  the  elucidation  of  this  problem.  The  gen- 
eral plan  of  the  experimental  work  was  extremely  simple.  It  con- 
templated the  selection  of  a  number  of  young  men  between  the  ages 
of  twenty  and  thirty,  in  excellent  health,  who  had  suffered  from 
no  serious  disease  in  the  immediate  past,  who  were  of  steady  habits, 
who  were  not  addicted  to  the  use  of  alcohol,  and  whose  character 
was  such  as  to  warrant  especial  confidence  and  trust  in  their 
veracity  and  general  conduct.     Such  young  men  evidently  are  to  be 


306  WILEY— INFLUENCE  OF  PRESERVATIVES  rAprilas, 

found  among  those  who  pass  the  examinations  for  the  civil  ser- 
vice of  the  United  States.  In  these  examinations  the  very  quali- 
ties which  were  looked  for  in  the  young  men  in  question  must 
be  present  or  they  could  not  receive  the  vouchers  for  character  and 
conduct  which  are  necessary  to  entitle  them  to  compete  in  the  ex- 
aminations. These  young  men  were  subjected  to  a  careful  physical 
examination  similar  to  that  exercised  upon  those  who  apply  for 
policies  in  life  insurance  companies.  This  examination  showed  them 
free  from  organic  diseases  and  not  to  have  suffered  within  a  year, 
usually  not  at  all,  from  any  serious  disturbance  of  health.  The  sub- 
jects were  placed  upon  their  honor,  by  a  formal  pledge,  that  they 
would  obey  all  the  rules  established  for  the  experimental  work 
and  abstain  from  any  form  of  food  and  drink  except  that  offered 
in  the  regular  course  of  the  investigation.  Those  who  used  to- 
bacco, tea  and  coffee  were  permitted  to  continue  to  do  so  in  the 
regular  manner  so  as  not  to  change  the  habits  of  their  previous 
daily  life.  They  were  also  limited  by  their  pledge  to  a  regular 
course  of  exercise  which  they  undertook  to  follow  without  varia- 
tion, and  also  regular  hours  of  work  and  sleep.  As  a  justification 
of  the  faith  and  confidence  reposed  in  these  young  men  I  think  it  is 
sufficient  to  say  that,  although  during  the  five  years  of  the  experi- 
ments we  have  had  about  sixty  young  men  under  observation,  only 
three  have  been  found  to  have  violated  their  pledges. 

The  subjects  so  selected  were  first  placed  on  a  generous  diet 
of  the  kind  and  character  to  keep  them  in  equilibrium;  that  is,  to 
maintain  the  weight  of  their  body  without  notable  changes.  The 
part  of  the  experiment  devoted  to  this  purpose  was  known  as  the 
"  fore  period."  Each  one  was  allowed  to  determine,  within  cer- 
tain limits,  the  character  of  the  diet  from  the  foods  offered ;  that  is, 
a  relative  amount  of  meat,  bread,  potatoes,  butter,  milk,  coffee,  tea, 
etc.,  to  suit  his  taste  and  to  conform  with  his  previous  habit  of 
life.  Only  in  those  cases  where  an  excess  of  some  particular  kind 
of  food  seemed  to  be  preferred  was  any  restriction  placed  upon  this 
matter.  This  fore  period,  therefore,  enabled  us  to  determine  the 
magnitude  of  the  ration  which  would  preserve  the  body  equilibrium 
and  presumably  be  in  entire  conformity  with  the  normal  digestive 
functions. 


,908.1  UPON   HEALTH   AND  METABOUSM.  307 

The  study  of  the  food  ingested  and  of  the  excreta  secured  estab- 
lished a  chemical  control  whereby  it  would  be  easy  to  determine 
any  variation  in  the  quantity  of  food  consumed  should  any  of  the 
young  men  attempt  to  evade  the  conditions  of  their  pledge.  Hav- 
ing thus  established  the  normal  conditions  of  the  body  and  ascer- 
tained the  normal  metabolic  processes,  there  was  introduced  into  the 
same  ration  varying  quantities  of  the  preservative  which  was  to  be 
studied.  It  was  thus  evident  that  any  change  taking  place  in 
health  or  metabolism  could  be  due  only  to  the  one  factor  which 
was  varied  in  the  method  of  life,  namely  the  injection  of  the 
chemical  preservative.  This  period,  during  which  a  drug  was  used, 
was  known  as  the  "  preservative  period,"  and  lasted,  according  to 
circumstances,  from  twenty  to  sixty  days,  depending  upon  the  char- 
'acter  and  magnitude  of  the  effects  produced.  As  soon  as  any  de- 
cided disturbance  of  health  was  produced,  clearly  traceable  to  the 
administration  of  the  preservative,  its  future  use  was  discontinued 
since  it  was  not  the  purpose  to  seriously  or  permanently  affect  the 
health  of  the  subject,  but  only  to  secure  positive  diagnostic  data. 
Then  followed  an  "  after  period,"  during  which  the  chemical,  or 
drug,  was  withdrawn  from  the  food  and  the  normal  ration  con- 
tinued as  in  the  fore  period,  the  object  being  to  correct,  if  pos- 
sible, any  disturbances  of  metabolism  which  had  been  produced 
and  restore  the  subject  again  to  normal  conditions  of  health  and 
digestion  and  also  to  study  the  after  effects  of  the  preservative 
should  such  persist.  This  period  of  observation  was  called  the 
"  after  period."  Thus  each  series  of  experimental  investigations 
were  divided  into  these  three  periods. 

During  the  progress  of  the  experiment  the  following  substances 
were  added  to  the  foods  for  the  purposes  mentioned  above:  Boric 
acid,  borates,  salicylic  acid,  salicylates,  benzoic  acid,  benzoates,  sul- 
phurous acid,  sulphites,  fonnaldehyde,  sulphate  of  copper  and  potas- 
sium nitrate.  There  is  given  in  the  accompanying  table  a  condensed 
statement  of  the  effects  which  were  produced  in  these  various  cases. 
It  is  not  the  purpose  of  this  paper  to  go  into  the  experimental  detail 
of  this  matter.  The  amount  of  chemical  analysis  incident  to  this 
study  was  enormous.  A  great  many  chemists  gave  their  entire  time 
during  the  whole  period  of  observation  to  these  analytical  problems, 


308 


WILEY— INFLUENCE  OF  PRESERVATIVES 


[April  as. 


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UPON   HEALTH  AND  METABOLISM. 


309 


and  in  addition  to  that  a  number  of  calculators  were  employed  to 
tabulate,  classify,  and  average  the  data.  The  experimental  data 
which  were  obtained  are  published  in  Bureau  of  Chemistry  Bulletin 
84,  which  when  completed  will  contain  the  entire  series  of  studies. 
Part  I,  of  Bulletin  84,  is  devoted  to  the  detailed  study  of  the  effect 
of  borates  and  boric  acid  upon  health  and  metabolism.  This  part 
of  the  Bulletin  consists  of  477  pages.  Part  II  is  devoted  to  the 
study  of  salicylic  acid  and  salicylates  and  contains  283  pages.  Part 
III  contains  the  data  relating  to  sulphurous  acid  and  sulphites,  and 
contain  281  pages,  making  a  total  of  published  matter  of  1,041 
pages. 

Changes  in  the  Urinary  Nitrogen  and  Sulphur  Compounds.' 


PresenratiYe. 

•  S  >* 

• 

J 

< 

Kreatiain. 

1 

a 

< 

• 

B 
•S 

S 

X 

• 

1 

s 

S5 

Total 
Sulphates. 

Inorganic 
Sulphates. 

Ethereal 
Sulphates. 

Borax... 

0.5  -3.0 
0.5  -5.0 

0.21-2.0 
0.17-0.4 
0.22-0.76 
1.0  -2.5 
1.0  -2.5 
0.1  -0.2 

O.OC-O.  IC 

Boric  acid 

Salicylic  add  and  sali- 
cylates   

{- 

— 

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0 

+ 

+ 

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1 

Sulphites 

1 
0 

Benzoic  acid 

0 

Benzoates 

0 

— 

1 

0 

CoDDcr  sulphate 

4- 
0 

1 

+ 

Potf^ium  nitrate •  o.  ic-o.6 

** 

0 

~-  -  J 

The  data  relating  to  benzoic  acid  and  benzoates  are  in  press. 
These  data,  together  with  those  relating  to  the  other  parts  of  the 
study  which  have  been  completed  and  submitted  for  publication, 
will  make  a  volume  of  approximately  2,100  pages.  All  that  I  can 
give  in  this  paper  will  be  the  general  conclusions  relating  to  each 
part  of  the  study. 

Conclusions. 

Boric  Acid  and  Borates, — In  the  consideration  of  the  action  of 
preservatives  of  a  mineral  nature,  such  as  borax  and  boric  acid, 
it  must  be  remembered  that  the  animal  as  well  as  the  plant  possesses 

^  Minus  and  plus  signs  indicate  decreased  or  increased  total  excretion  in 
preservative  period  as  compared  with  the  fore  period. 


310  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  asi 

a  certain  mineral  hunger.  In  other  words,  mineral  substances  play 
a  double  role  in  animal  and  plant  nutrition:  First,  they  may  serve 
as  real  foods,  necessary  to  the  formation  and  nutrition  of  the  tissue. 
In  the  animal  economy  this  is  especially  true  of  phosphoric  acid 
and  lime.  In  the  second  place,  they  are  necessary  to  the  func- 
tional activity  of  the  various  organs  of  the  body,  irrespective  of  any 
part  they  may  take  in  direct  nutrition. 

The  necessity  of  saline  solutions  in  the  blood  is  known  to  every 
physician  and  physiologist.  If  the  blood  were  deprived  of  all  of  its 
saline  constituents  the  circulation  would  be  impeded,  restricted,  or 
stopped,  and  death  would  result.  In  cases  of  collapse  in  disease 
saline  injections  in  the  blood  are  often  used  as  a  restorative  measure. 
These  salts  in  solution  stimulate  the  heart's  action  and  undoubtedly 
are  active  in  the  osmotic  operations  of  the  cells.  This  is  one  of  the 
facts  which  show  the  intimate  relation  existing  between  physical 
chemistry  and  physiology. 

Common  salt  is  the  most  frequent  and  most  abundant  of  the 
saline  constituents  of  the  blood,  but  the  alkalinity  of  the  blood  is  not 
due  of  course  to  the  common  salt,  which  is  a  neutral  substance. 
The  existence  of  alkaline  carbonates  or  other  alkaline  salts  is  neces- 
sary to  the  vital  functions.  While  it  is  true  that  the  digestion  in  the 
stomach  takes  place  in  an  acid  solution,  it  is  likewise  true  that  any 
excessive  acid  must  be  neutralized  and  enough  of  alkali  added  in  the 
small  intestine  in  order  that  the  further  digestion  of  the  food  may 
properly  take  place.  That  saline  bodies  other  than  common  salt  or 
the  alkaline  carbonates  may  be  useful,  however,  in  the  perform- 
ance of  the  vital  functions  cannot  be  denied,  though  it  might  be 
difficult  to  demonstrate  their  absolute  necessity.  Hence  the  intro- 
duction of  saline  bodies,  which  may  or  may  not  be  of  an  antiseptic 
character,  may,  within  certain  limits,  have  a  favorable  influence  upon 
health  and  digestion.  At  the  same  time  it  should  not  be  forgotten 
that  all  excess  of  such  bodies  imposes  upon  the  excretory  organs  an 
additional  burden,  which,  while  it  might  not  impair  their  efficiency 
even  for  a  number  of  years,  might  finally  produce  a  condition  of 
exhaustion  which  would  be  followed  by  serious  consequences. 
Especially  is  this  remark  true  of  the  kidneys,  which  appear  to  be  a 


,9o8.]  UPON   HEALTH  AND  METABOLISM.  311 

general  clearing  house  for  all  the  surplus  of  saline  matters,  ingested 
in  the  foods. 

The  most  interesting  of  the  observations  which  were  made  dur- 
ing the  progress  of  the  experiments  was  in  the  study  of  the  direct 
effect  of  boric  acid  and  borax,  when  administered  in  food,  upon 
the  health  and  digestion.  When  boric  acid,  or  its  equivalent  in 
borax,  is  taken  into  the  food  in  small  quantities,  not  exceeding  half 
a  gram  (7^  grains)  a  day,  no  notable  effects  are  immediately  pro- 
duced. The  medical  symptoms  of  the  cases,  in  long-continued  ex- 
hibitions of  small  doses  or  in  large  doses  extending  over  a  shorter 
period,  show  in  many  instances  a  manifest  tendency  to  diminish  the 
appetite  and  to  produce  a  feeling  of  fullness  and  uneasiness  in  the 
stomach,  which  in  some  cases  results  in  nausea,  with  a  very  general 
tendency  to  produce  a  sense  of  fullness  in  the  head,  which  is  often 
manifested  as  a  dull  and  persistent  headache.  In  addition  to  the 
uneasiness  produced  in  the  region  of  the  stomach  there  appear  in 
some  instances  sharp  and  well-located  pains,  which,  however,  are 
not  persistent.  Although  the  depression  in  the  weight  of  the  body 
and  some  of  the  other  symptoms  produced  persist  in  the  after 
periods,  there  is  a  uniform  tendency  manifested  after  the  with- 
drawal of  the  preservative  toward  the  removal  of  the  unpleasant 
sensations  in  the  stomach  and  head  above  mentioned. 

The  administration  of  boric  acid  to  the  amount  of  4  or  5  grams 
per  day,  or  borax  equivalent  thereto  continued  for  some  time,  results 
in  most  cases  in  loss  of  appetite  and  inability  to  perform  work  of 
any  kind.  In  many  cases  the  person  becomes  ill  and  unfit  for  duty. 
Four  grams  per  day  may  be  regarded,  then,  as  the  limit  of  exhibi- 
tion beyond  which  the  normal  man  may  not  go.  The  administration 
of  3  grams  per  day  produced  the  same  symptoms  in  many  cases, 
althr>ugh  it  appeared  that  a  majority  of  the  men  under  observa- 
tion were  able  to  take  3  grams  a  day  for  a  somewhat  protracted 
period  and  still  perform  their  duties.  They  commonly  felt  injurious 
effects  from  the  dose,  however,  and  it  is  certain  that  the  normal  man 
could  not  long  continue  to  receive  3  grams  per  day. 

In  many  cases  the  same  results,  though  less  marked,  follow  the 
administration  of  borax  to  the  extent  of  2  grams  and  even  of  i 
gram  per  day,  although  the  illness  following  the  administration  of 


312  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  n, 

borax  and  boric  acid  in  those  proportions  may  be  explained  in  some 
cases  by  other  causes,  chiefly  grippe. 

The  administration  of  borax  and  boric  acid  to  the  extent  of  one 
half  gram  per  day  yielded  results  markedly  different  from  those 
obtained  with  larger  quantities  of  the  preservatives.  This  experi- 
ment, Series  V,  conducted  as  it  was  for  a  period  of  fifty  days,  was 
a  rather  severe  test,  and  it  appeared  that  in  some  instances  a  some- 
what unfavorable  result  attended  it.  On  the  whole,  the  results 
show  that  one  half  gram  per  day  is  too  much  for  the  normal  man  to 
receive  regularly.  On  the  other  hand,  it  is  evident  that  the  normal 
man  can  receive  one  half  gram  per  day  of  boric  acid,  or  of  borax 
expressed  in  terms  of  boric  acid,  for  a  limited  period  of  time 
without  much  danger  of  impairment  of  health. 

It  is,  of  course,  not  to  be  denied  that  both  borax  and  boric  acid 
are  recognized  as  valuable  remedies  in  medicine.  There  are  certain 
diseases  in  which  these  remedies  are  regularly  prescribed  for  both 
internal  and  external  use.  The  value  which  they  possess  in  these 
cases  does  not  seem  to  have  any  relation  to  their  use  in  the  healthy 
organism  except  when  properly  prescribed  as  prophylactics.  The 
fact  that  any  remedy  is  useful  in  disease  does  not  appear  to  logically 
warrant  its  use  at  any  other  time. 

It  appears,  therefore,  that  both  boric  acid  and  borax,  when  con- 
tinually administered  in  small  doses  for  a  long  period  or  when  g^ven 
in  large  quantities  for  a  short  period,  create  disturbances  of  appe- 
tite, of  digestion,  and  of  health. 

Salicylic  Acid  and  Salicylates, — In  the  conclusions  based  upon 
the  general  observations  the  same  conservatism  must  be  observed 
and  the  same  general  reservations  made  as  are  found  in  Part  I 
concerning  boric  acid  and  borax.  While,  as  described  in  the  borax 
report,  the  attempt  has  been  made  to  control  as  far  as  possible,  all 
the  conditions  of  the  experimental  work,  the  difficulties  attending 
the  task  are  so  enormous  that  it  is  not  possible  that  complete  suc- 
cess should  be  secured.  There  has,  however,  been  no  attempt  made 
to  discriminate  in  the  choice  of  data,  all  the  observations  being 
recorded  and  the  discussion  of  the  individual  data  based  upon  the 
tabular  statements  being  without  prejudice  and  without  bias.  The 
general  assumption  has  been  made,  as  in  the  previous  cases,  that. 


,9o8]  UPON   HEALTH  AND  METABOLISM.  313 

by  reason  of  the  regular  habits  of  life  which  were  imposed  upon  the 
subjects,  the  amount  of  energy  developed  and  the  quantity  of  nour- 
ishment expended  therein  are  reasonably  constant  throughout  the 
experimental  period.  If  these  factors  vary,  as  they  necessarily  must 
to  a  certain  degree,  it  is  evident  that  they  vary  uniformly  above  or 
below  the  average,  and  hence  these  variations  could  not  possibly 
produce  any  notable  effect  upon  the  final  result. 

There  has  been  a  general  consensus  of  opinion  among  scientific 
men,  including  the  medical  profession,  that  salicylic  acid  and  its 
compounds  are  very  harmful  substances,  and  the  prejudice  against 
this  particular  form  of  preservative  is  perhaps  greater  than  against 
any  other  material  used  for  preserving  foods.  This  is  due  not  only 
to  the  belief  in  the  injurious  character  of  salicylic  acid,  but  perhaps 
is  especially  due  to  the  fact  that  it  has  in  the  past  been  so  generally 
used  as  an  antiseptic.  That  salicylic  acid  should  be  singled  out 
especially  for  condemnation  among  preservatives  does  not  seem  to 
be  justified  by  the  data  which  are  presented  and  discussed  in  this 
bulletin.  That  it  is  a  harmful  substance,  however,  seems  to  be  well 
established  by  the  data  taken  as  a  whole,  but  it  appears  to  be  a  harm- 
ful substance  of  less  virulence  than  has  been  generally  supposed. 
There  is  no  doubt  of  the  fact  that  salicylic  acid  is  a  drug  which  is 
often  indicated  in  diseases  well  established  and  also  perhaps  in  cer- 
tain conditions  wjiich,  while  verging  on  disease,  might  still  be  re- 
garded as  a  state  of  health.  But  the  administration  of  salicylic  acid 
as  a  medicine  should  be  controlled  exclusively  by  the  medical  pro- 
fession, and  while  it  is  a  remedy  well  established  in  the  Pharma- 
copoeia and  especially  prized  for  its  eflFect  upon  rheumatism  and  gout, 
it  does  not  seem  that  there  should  be  any  warrant  in  this  fact  for  its 
promiscuous  use  in  foods,  even  if  it  were  harmless. 

The  data  show  very  clearly  that  salicylic  acid  and  salicylates 
appear  to  exert  an  exciting  influence  upon  the  activities  which  take 
place  in  the  alimentary  canal,  stimulating  the  organs  to  greater  effort, 
and  this  stimulation  leads  at  first  to  increased  solubility  and  absorp- 
tion of  the  foods  which  are  introduced  into  the  stomach.  In  the 
light  of  the  data  which  are  exhibited  salicylic  acid  may  be  said  to 
increase  the  solubility  and  absorption  of  the  food  in  the  alimentary 


314  WILEY— INFLUENCE  OF  PRESERVATIVES  [AprUas, 

canal,  so  that  larger  parts  of  the  nutrients  taken  into  the  stomach 
actually  enter  the  circulation. 

The  data  which  show  the  effect  just  noted  also  indicate  that 
the  general  effect  upon  the  system  is  depressing,  in  that  the  tissues 
are  broken  down  more  rapidly  than  they  are  built  up,  and  thus  the 
normal  metabolic  processes  are  interfered  with  in  a  harmful  way. 
The  administration  of  the  salicylic  acid  is  attended  by  a  gradual 
decrease  in  the  weight  of  the  subjects,  although  the  quantity  of  food 
elements  administered  during  the  preservative  and  after  periods  is 
slightly  increased,  which  fact,  together  with  the  greater  degree  of 
absorption  of  the  food  elements,  should  have  resulted  in  a  slight 
increase  in  weight.  This  increase  in  weight,  however,  does  not 
occur,  and  the  disturbing  influence  of  the  salicylic  acid  upon  meta- 
bolism, although  not  very  great,  is  specifically  demonstrated. 

The  final  conclusion  in  this  matter,  therefore,  is  that  the  un- 
enviable position  which  salicylic  acid  has  heretofore  held  among 
preservatives,  in  being  regarded  as  the  most  injurious  of  all,  is  to  a 
certain  extent  undeserved.  Like  other  ordinary  preservatives,  it  is 
not  one  which  can  be  classed  as  a  poison  in  the  usual  sense  of  the 
word.  When  used  as  a  medicine  in  many  cases  of  derangement  of 
health  it  is  like  the  other  chemical  preservatives,  often  highly  bene- 
ficial when  properly  prescribed  by  a  competent  physician.  It  is  when 
used  in  the  food  at  first  an  apparent  stimulant,  increasing  the  absorp- 
tion and  solubility  of  the  common  food  elements  from  the  alimen- 
tary canal.  It  soon,  however,  loses  its  stimulating  properties  and 
becomes  a  depressant,  tending  to  break  down  the  tissues  of  the  body 
more  rapidly  than  they  are  built  up.  It  disturbs  the  metabolic  proc- 
esses, in  most  cases  producing  conditions  which  are  not  normal  and 
which,  apparently,  are  not  beneficial.  It  has  a  tendency  to  diminish 
the  weight  of  the  body  and  to  produce  a  feeling  of  discomfort  and 
malaise,  which,  while  not  marked,  is  distinctly  indicative  of  injury. 
In  some  cases  these  symptoms  of  malaise  approach  illness,  and  while 
not  always  diagnostic  are  sufficiently  common  to  point  unmistakably 
to  the  salicylic  acid  as  their  origin.  It  places  upon  the  excretory 
organs,  especially  the  kidneys,  an  additional  burden  which  they  are 
not  able  to  bear  and  which  cannot  possibly  result  in  any  good,  but 
on  the  contrary  must  necessarily  finally  result  in  injury,  though  per- 


,5o8.]  UPON   HEALTH   AND  METABOLISM.  316 

haps  with  the  use  of  very  small  quantities  of  the  preservative  these 
organs  would  continue  to  perform  their  function  for  many  years 
before  finally  breaking  down. 

This  work  is  offered  as  an  unbiased  study  of  all  the  data  re- 
corded, both  of  those  which  appear  to  be  in  favor  of  the  use  of 
salicylic  acid  and  those  which  appear  to  be  against  its  use,  and  leads 
to  the  inevitable  conclusion  that  salicylic  acid  is  a  substance  which, 
when  added  to  foods  even  in  small  quantities,  exerts  a  depressing  and 
harmful  influence  upon  the  digestion  and  health  and  the  general 
metabolic  activities  of  the  body.  Further,  there  appears  to  be  no 
necessity  for  its  use,  as  food  can  be  preserved  in  unobjectionable 
ways  without  its  aid.  Its  indiscriminate  use  would  tend  to  care- 
lessness in  the  quantities  employed,  thus  increasing  the  dangers  to 
which  the  consumer  is  subjected.  Also  its  use  in  the  preservation 
of  foods  tends  to  induce  carelessness  and  indifference  on  the  part 
of  the  manufacturer,  as  when  a  chemical  antiseptic  is  employed 
many  of  the  processes  necessary  to  the  proper  selection,  cleaning, 
and  preservation  of  foods  may  be  omitted. 

The  addition  of  salicylic  acid  and  salicylates  to  foods  is  there- 
fore a  process  which  is  reprehensible  in  every  respect,  and  leads  to 
injury  to  the  consumer,  which,  though  in  many  cases  not  easily 
measured,  must  finally  be  productive  of  great  harm. 

Sulphurous  Acid  and  Sulphites. — From  a  careful  consideration 
of  the  data  in  the  individual  cases  and  the  summaries  of  the  results, 
it  appears  that  the  administration  of  sulphurous  acid  in  foods,  either 
in  the  form  of  sulphurous  acid  gas  in  solution  or  in  the  form  of 
sulphites,  is  objectionable  and  produces  serious  disturbances  of  the 
metabolic  functions  and  injury  to  health  and  digestion.  This  injury 
manifests  itself  in  a  number  of  different  ways,  both  in  the  produc- 
tion of  clinical  symptoms  which  indicate  serious  disturbances, 
malaise,  or  positive  suflFering,  and  also  by  inducing  certain  changes 
in  the  metabolic  processes  which  are  not  manifested  in  the  way  of 
ordinary  clinical  symptoms,  and  are  only  detected  by  careful  chem- 
ical and  microscopical  study  of  the  excretory  products.  It  can 
safely  be  said  from  the  evidence  adduced  that  the  administration 
of  sodium  sulphite  and  sulphurous  acid  as  above  indicated  produces 
a  marked  influence  of  an  unfavorable  character  on  metabolism.     As 


316  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  .5. 

a  result  of  this  action  an  assimilation  of  food  materials  containing 
organic  phosphorus  is  retarded,  while  there  is  evidence  of  increased 
sulphur  katabolism.  The  sulphur  balance  sheets  show  what  an 
immense  burden  has  been  added  to  the  already  overworked  kidneys, 
which  are  called  upon  in  this  case  to  rjmove  nearly  all,  if  not  quite 
all,  of  the  added  sulphur  frcwn  the  body,  previously  converted,  in 
great  part  to  sulphuric  acid.  It  is  not  possible  that  placing  upon 
the  kidneys  this  increased  work  of  excreting  sulphur  can  result  in 
anything  but  injury.  The  fact  that  the  microscc^ic  crystalline  and 
amorphous  bodies  in  the  urine  are  increased  in  number  under  the 
influence  of  the  added  sulphur,  is  another  indication  of  the  extra- 
ordinary demands  made  upon  the  kidneys  in  such  circumstances. 

This  increase  is  interesting  in  respect  of  the  effect  which  the 
continued  exhibition  of  sulphurous  acid  must  eventually  have  upon 
the  structure  of  the  kidney.  It  is  reasonable  to  suppose  that  the 
continued  use  of  a  body  which  produces  such  results  would  cause 
lesions  of  a  histological  character  which  eventually  would  develop 
conditions  which  would  give  serious  apprehension.  In  the  nature 
of  these  experiments  it  was  not  possible  to  examine  the  organs  of 
the  body  histologically  and  hence  the  above  conclusion  is  only  based 
upon  experience  of  a  similar  character  where  the  organs  in  question 
have  been  subject  to  such  examinations.  While  there  might  be  no 
distinguishable  lesion  of  the  kidneys  produced  during  a  period  of 
twenty  or  thirty  days,  or  even  longer,  it  is  plain  that  sooner  or  later 
lesions  of  a  very  serious  character  producing  organic  diseases,  pos- 
sibly of  an  incurable  type,  would  be  induced.  The  further  observa- 
tion that  there  is  a  marked  tendency  to  the  production  of  albu- 
minuria, although  of  an  incipient  character,  is  an  indication  of  the 
unfavorable  results  of  the  administration  of  the  sulphurous  acid. 
It  is,  therefore,  evident  that  by  increasing  the  burden  upon  the 
excretory  organs,  the  administration  of  sulphur  in  the  form  men- 
tioned is  highly  detrimental  to  health. 

All  of  these  tendencies  cannot  be  interpreted  as  being  other  than 
of  a  decidedly  harmful  nature.  Another  effect  which  the  adminis- 
tration of  the  sulphur  produced,  and  one  of  a  more  serious  character 
still,  is  found  in  the  impoverishment  of  the  blood  in  respect  of  the 
number  of  red  and  white  corpuscles  therein.    The  administration 


,9o8.]  UPON  HEALTH   AND  METABOLISM.  317 

of  a  substance  which  diminishes  by  a  notable  percentage  these  im- 
portant component  particles  of  the  blood  must  be  regarded  in  every 
sense  as  highly  prejudicial  to  health.  Some  of  the  most  important 
functions  of  the  blood,  as  has  been  well  established  by  careful 
physiological  studies,  are  intimately  connected  with  the  number  and 
activity  of  both  the  red  and  white  corpuscles.  The  bleaching  effect 
of  the  sulphurous  acid  upon  the  color  of  the  blood  is  a  matter  of 
less  consequence  and  no  great  effect  is  produced  upon  the  hemo- 
globin, but  the  diminution  of  the  number  of  red  and  white  cor- 
puscles is  a  matter  of  serious  concern. 

The  variations  of  the  metabolic  processes  from  the  normal,  as 
indicated  in  this  series  of  experiments,  were  never  of  a  character 
favorable  to  a  more  healthy  condition  of  the  system,  but,  on  the 
other  hand,  all  these  variations,  in  so  far  as  the  effect  of  the  changes 
could  be  distinguished,  are  of  a  prejudicial  character.  There  is  no 
evidence  whatever  that  the  sulphur  added  to  the  foods  in  the  form 
of  sulphurous  acid,  or  sulphites,  takes  any  part  in  the  nutrition  of 
the  tissues  of  the  body  containing  sulphur,  namely,  the  proteids; 
hence,  no  claim  of  food  value  can  be  established  for  these  bodies. 
The  evidence  all  points  to  the  fact  that  they  are  purely  drugs,  devoid 
of  food  value,  having  no  favorable  effects  upon  the  metabolic  proc- 
esses, but,  on  the  other  hand,  exerting  deleterious  and  harmful 
effects.  The  conclusion,  therefore,  is  inevitable  that,  as  a  whole, 
the  changes  produced  in  metabolic  activity  by  the  administration  of 
sulphur  in  the  forms  noted  above  in  the  comparatively  short  time 
covered  by  the  experiments  are  decidedly  injurious. 

The  verdict  which  must  be  pronounced  in  this  case  is  decidedly 
unfavorable  to  the  use  of  this  preservative  in  any  quantity  or  for 
any  period  of  time,  and  shows  the  desirability  of  avoiding  the  addi- 
tion of  any  form  of  sulphurous  acid  to  products  intended  for  human 
food. 

Benzoic  Acid  and  Benzoates, — From  a  careful  study  of  the  data 
in  the  individual  cases  and  of  the  summaries  of  the  results,  it  is 
evident  that  the  administration  of  benzoic  acid,  either  as  such  or 
in  the  form  of  benzoate  of  soda,  is  highly  objectionable  and  pro- 
duces a  very  serious  disturbance  of  the  metabolic  functions,  attended 
with  injury  to  digestion  and  health. 

PROG.  AMER.  PHIL.  SOC.,  XLVII.   189  U,  PRINTED  SEPTEMBER  26,   I908. 


318  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  .5. 

As  in  the  case  of  boric  acid,  salicylic  acid,  and  sulphurous  acid, 
this  injury  manifests  itself  in  a  number  of  diflFerent  ways,  both  in 
the  production  of  unfavorable  symptoms  and  in  the  disturbance  of 
metabolism.  These  injurious  effects  are  evident  in  the  medical  and 
clinical  data  which  show  grave  disturbances  of  digestion,  attended 
by  phenomena  which  are  clearly  indicative  of  irritation,  nausea, 
headache,  and  in  a  few  cases  vomiting.  These  symptoms  were  not 
only  well  marked,  but  they  were  produced  upon  healthy  individuals 
receiving  good  and  nourishing  food  and  living  under  proper  sani- 
tary conditions.  It  is  only  fair  to  conclude,  therefore,  that  under 
similar  conditions  of  administration  of  benzoic  acid  or  benzoate  of 
soda  in  the  case  of  weaker  systems,  or  less  resistant  conditions  of 
health,  much  more  serious  and  lasting  injury  would  be  produced. 

It  was  also  noticed  that  the  administration  of  benzoic  acid  and 
benzoate  of  soda  was  attended  with  a  distinct  loss  of  weight,  indica- 
tive of  either  a  disturbance  of  assimilation  or  an  increased  activity 
in  those  processes  of  the  body  which  result  in  destruction  of  tissue. 
The  production  of  a  loss  of  weight  in  cases  of  this  kind  must  be 
regarded  as  indicative  of  injurious  effects. 

The  influence  of  the  benzoic  acid  and  benzoate  of  soda  upon 
metabolism  was  never  of  a  character  indicative  of  a  favorable  change 
therein.  While  often  the  metabolic  changes  were  not  strongly 
marked,  such  changes  as  were  established  were  of  an  injurious 
nature.  It  is  evident  that  the  administration  of  these  bodies,  there- 
fore, in  the  food  tends  to  derange  metabolism  in  an  injurious  way. 

An  important  fact  in  connection  with  the  administration  of  these 
bodies  is  found  in  the  efforts  which  nature  makes  to  eliminate  them 
from  the  system.  In  so  far  as  possible  the  benzoic  acid  is  converted 
into  hippuric  acid.  There  is  a  tendency  usually  manifested,  how- 
ever, to  retain  the  benzoic  acid  in  the  body  for  a  notable  length  of 
time,  and  this  is  much  more  marked  in  the  case  of  benzoate  of  soda 
than  in  the  case  of  benzoic  acid. 

While  the  administration  of  both  these  bodies,  therefore,  is 
undoubtedly  harmful,  the  injurious  effects  are  produced  more  rap- 
idly in  the  case  of  benzoic  acid  than  they  are  in  the  case  of  benzoate 
of  soda ;  the  data,  however,  will  show  that  the  total  harmful  effect 
produced  in  the  end  is  practically  the  same  in  both  cases,  hence  there 


i9o8.]  UPON   HEALTH  AND   METABOLISM.  319 

appears  to  be  no  reason  for  supposing  that  the  administration  of  the 
preservative  in  the  form  of  benzoate  of  soda  can  be  justified  by  any 
argument  relating  to  the  less  injurious  eflfect  thereof  upon  health. 

The  occurrence  of  microscopic  bodies  in  the  urine  is  undoubt- 
edly increased  under  the  administration  of  benzoic  acid  in  all  its 
forms,  thus  showing  conclusively  the  tendency  to  stimulate  the 
destructive  activities  of  the  body. 

Coming  to  the  final  consideration  of  all  these  different  phases 
of  the  subject,  there  is  only  one  conclusion  to  be  drawn  from  the 
data  which  have  been  presented  and  that  is  that  in  the  interests  of 
health  both  benzoic  acid  and  benzoate  of  soda  should  be  excluded 
from  food  products.  This  conclusion  is  reached  independently  of 
any  consideration  of  the  conditions  which  it  is  alleged  surround  the 
processes  of  manufacture  and  which  result  in  the  demands  of  manu- 
facturers to  be  allowed  to  continue  the  use  of  this  body.  This  is  a 
subject  which  must  be  discussed  from  an  entirely  different  point  of 
view  and  has  no  bearing  whatever  upon  the  general  conclusions 
which  have  been  reached,  namely,  that  both  benzoic  acid  and  ben- 
zoate of  soda  are  bodies  which,  when  added  to  foods,  are  injurious 
to  health. 

Formaldehyde, — A  general  study  of  all  the  data  leads  to  the 
conclusion  that  the  admixture  of  formaldehyde  with  food  is  injuri- 
ous to  health,  even  in  the  case  of  healthy  young  men.  It  is  fair  to 
conclude,  therefore,  that  in  the  case  of  infants  and  children  the 
deleterious  effects  would  be  more  pronounced.  The  metabolic  func- 
tions are  disturbed  in  a  notable  way,  both  by  the  retardation  of  the 
nitrogen  and  sulphur  metabolism,  and  the  acceleration  of  phos- 
phorus metabolism.  There  seems  to  be  a  tendency  to  an  increased 
absorption  from  the  alimentary  canal,  especially  in  the  cases  when 
the  formaldehyde  had  stood  in  contact  with  the  milk,  and  hence  it 
is  fair  to  presume  that  in  so  far  as  the  enzymic  action  in  the  intes- 
tinal canal  is  concerned,  transforming  solid  food  into  soluble  mate- 
rials which  may  enter  the  circulation,  there  is  evidently  a  stimulat- 
ing effect  produced. 

There  are,  however,  many  varying  conditions  which  must  be 
considered  in  properly  interpreting  the  data.  The  uniformly  in- 
creased absorption  of  the  proteid  elements  of  the  food,  and  also  of 


320  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  as, 

the  sulphur  and  phosphoric  acid,  accompanied  in  the  first  two  in- 
stances by  a  decrease  in  the  metabolized  elements  excreted  and  in 
the  last  instance,  namely,  phosphoric  acid,  by  a  pronounced  increase 
in  metabolism,  makes  the  explanation  of  the  data  rather  difficult. 
Attention  should  be  called  to  the  fact  that  while  the  variations  from 
normal  metabolism  are  not  very  wide,  the  individual  data  are  re- 
markably uniform  and  consistent. 

The  conditions  which  are  noted  in  the  case  of  the  proteins  would 
lead  one  to  expect  a  gain  in  the  body  weight.  This  expectation, 
however,  is  not  realized  for  either  class  of  subjects,  although  the 
losses  in  weight  are  so  slight  as  to  be  practically  negligible.  The 
ratio  of  the  food  weight  to  the  body  weight  was  uniformly  main- 
tained throughout  the  experiment,  and,  hence,  if  no  variations  in 
metabolic  activity  had  occurred  a  fair  presumption  would  have  been 
that  the  body  weight  would  remain  constant.  That  the  change  of 
weight  was  slight  in  the  view  of  the  disturbances  of  the  metabolic 
functions  may  be  accounted  for  by  the  inhibiting  or  retarding  influ- 
ence of  the  preservative  upon  the  nitrogen  and  sulphur  katabolism, 
or  by  the  slight  increase  in  water  in  the  urine  and  feces.  It  cannot 
be  maintained,  however,  that  a  retarded  katabolism  is  beneficial 
to  health.  On  the  contrary  a  more  rapid  renewal  of  the  tissues 
within  the  limits  of  healthy  activity  would  be  more  likely  to  pre- 
serve a  normal  condition.  The  old  tissues  cannot  be  expected  to 
functionate  as  perfectly  as  those  which  are  newer,  and  hence,  within 
reasonable  limits,  a  change  of  the  tissues  of  the  body  must  be  con- 
sidered as  necessary  to  a  healthy  condition,  and  the  maintenance  of 
a  normal  vitality. 

The  medical  data  indicate  plainly  that  formaldehyde,  even  when 
given  in  small  quantities,  is  an  irritating  substance  to  the  mucous 
membrane,  and,  therefore,  the  normal  organs  are  at  first  actively 
stimulated  to  rid  themselves  of  the  irritating  foreign  substance.  It 
is  not  strange,  therefore,  that  this  preservative  had  a  marked  stimu- 
lative action  on  those  organs  and  cells  secreting  the  various  diges- 
tive juices.  It  is  evident  that  when- the  digestive  and  excretory 
organs  of  the  body  are  excited  to  unusual  activity  by  such  an  ex- 
traneous body  having  neither  food  nor  condimental  value,  they  act 
in  self  defence,  and  it  would  be  wholly  illogical  to  conclude  from 


x9o8.]  UPON   HEALTH  AND  METABOLISM.  321 

this  increased  excitation  that  these  bodies  were  helpful  to  digestion 
and  conducive  to  health.  The  nature  of  the  investigation  made  it 
impossible  to  determine  whether  any  organic  change  took  place  in 
the  various  organs  aflFected,  but  it  may  be  assumed  that  any  such 
change  which  these  organs  had  undergone  in  the  limited  time  was 
not  sufficient  to  disturb  in  any  notable  way  their  normal  functions 
which  they  would  perform  until  the  continued  administration  of  the 
drug  produced  disease  due  to  the  excessive  stimulation. 

In  the  case  of  pTiosphoric  acid,  the  increased  katabolic  activity 
is  difficult  of  definite  interpretation,  though  it  is  established  beyond 
doubt  that  such  an  effect  is  produced.  The  formaldehyde  may 
exert  a  selective  action  for  those  proteid  bodies  high  in  phosphorus, 
rendering  them  insoluble,  but  in  this  case  there  would  be  an  excess 
of  phosphorus  in  the  feces,  which  is  not  found.  Or  the  formalde- 
hyde may  induce  a  change  in  the  process  of  digestion  whereby  the 
phosphorus  of  the  food  is  changed  into  a  soluble  and  easily  exerted 
form  without  passing  through  the  tissues  of  the  body.  This  might 
easily  be  the  case  if  in  the  process  of  digestion  the  glycerol-phos- 
phoric  acid  formed  is  transformed  into  soluble  inorganic  salts, 
which  are  readily  excreted.  Whatever  may  be  the  explanation,  the 
changes  indicated  in  normal  metabolism,  accompanied  as  they  are 
by  the  development  of  the  symptoms  described,  can  only  be  consid- 
ered as  prejudicial  to  health. 

The  general  tendency  to  produce  a  slight  decrease  in  the  tem- 
perature of  the  body,  assuming  for  the  moment  that  the  data  war- 
rant the  conclusion  that  such  a  condition  of  affairs  existed,  might 
well  be  due  to  the  inhibition  of  cell  activity  shown  by  the  retarda- 
tion in  the  breaking  down  of  tissues.  The  normal  functions  of  the 
body  would  doubtless  be  disturbed  by  such  a  condition,  aside  from 
the  irritating  and  other  disturbing  influences  exerted  by  the  exhib- 
ited drug. 

The  tendency  of  the  preservative  to  produce  albumin  in  the 
urine,  while  not  well  marked,  is  at  least  worthy  of  attention.  The 
fact  that  only  slight  changes  take  place  in  the  body  weight  is  suffi- 
ciently explained  in  the  data,  and  cannot  be  urged  in  favor  of  the 
exhibited  preservative. 

Apart  from  the  injurious  eflFects  of  formaldehyde  itself,  its  use 


322  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  as, 

as  a  food  preservative  would  be  especially  inadvisable  in  milk  or 
cream,  because  its  addition  in  dilute  solution  prevents  the  growth 
of  acid-forming  bacteria,  but  has  no  effect  in  retarding  the  action 
of  many  harmful  organisms ;  in  other  words,  the  milk  is  prevented 
from  becoming  sour  and  thus  indicating  its  age  and  the  danger  sig- 
nal is  thus  removed,  while  the  other  organisms  which  are  capable 
of  producing  disease  continue  to  multiply  in  the  milk  with  practically 
the  same  degree  of  rapidity  as  if  the  formaldehyde  was  not  present. 

The  final  conclusion,  therefore,  is  that  the  addition  of  formalde- 
hyde to  foods  tends  to  derange  metabolism,  disturb  the  normal  func- 
tions, produce  irritation  and  undue  stimulation  of  the  secretory 
activities,  and,  therefore,  it  is  never  justifiable. 

Sulphate  of  Copper. — The  data  which  have  been  collected  in  the 
course  of  this  experiment  have  led  to  the  conclusion  that  the 
administration  of  sulphate  of  copper  even  in  the  extremely  small 
quantities  in  which  it  has  been  given  has  a  very  distinctly  unfavor- 
able effect  upon  health  and  digestion,  as  indicated  by  the  ordinary 
clinical  and  medical  summaries.  Severe  pains  are  produced  in  the 
stomach  accompanied  often  with  nausea  and  sometimes  with  vomit- 
ing, there  is  a  general  tendency  to  malaise,  often  a  development  of 
headache,  and  other  unfavorable  symptoms  of  a  more  or  less  per- 
sistent and  uniform  character.  Further  than  this,  the  symptoms 
which  are  usually  not  developed  for  about  a  week  continue  in  some 
instances  for  a  number  of  days  into  the  after-period  after  the  sul- 
phate of  copper  has  been  withdrawn.  The  data  indicate  that  cop- 
per, like  many  other  metals,  is  likely  to  produce  a  cumulative  effect, 
and  that  its  administration  in  even  much  smaller  quantities  than 
those  indicated,  or  less  than  those  which  would  be  ingested  in  the 
regular  consumption  of  coppered  vegetables,  is  attended  with  more 
or  less  danger  on  this  account. 

There  was  a  very  small  loss  of  weight  in  nine  of  the  subjects, 
while  the  three  who  showed  the  greatest  tolerance  of  the  copper 
sulphate  gained  in  weight.  No  definite  conclusions  can,  therefore, 
be  formed  respecting  the  general  effect  upon  the  weight  of  the 
body,  except  that  in  the  cases  where  uniform  effects  are  produced 
there  is  a  slight  loss  of  weight 

The  copper  salt  which  was  used  in  this  experiment  diflFers  from 


X908.1  UPON   HEALTH   AND  METABOLISM.  323 

Other  chemicals  which  have  been  used  in  this  series  of  investigations 
in  that  its  excretion  falls  only  partly  upon  the  kidneys.  The  effect 
produced  on  the  urine,  therefore,  cannot  be  ascribed  directly  to  the 
copper  salt  employed,  but  only  to  such  derangements  of  the  metab- 
olism due  thereto  as  would  incidentally  aflfect  the  composition  of 
the  urine. 

The  eflFect  upon  the  general  metabolism  is  of  a  character  which, 
though  not  very  pronounced,  is  indicative  of  a  retardation  of  normal 
metabolic  processes.  Inasmuch  as  a  small  quantity  of  sulphur  was 
introduced  into  the  system  through  the  copper  salt,  the  quantity  of 
this  sulphur  must  be  taken  into  consideration  in  studymg  the  effect 
on  metabolism.  There  is  seen  to  be  quite  a  uniform  tendency  to 
derange  the  ratio  of  the  metabolized  sulphur  and  nitrogen. 

The  apparent  increase  in  the  relative  quantities  of  sulphur  ex- 
creted is  due  rather  to  the  diminution  in  the  nitrogen  than  to  an 
actual  increase  in  the  sulphur  over  that  which  would  be  expected 
from  the  ingestion  of  the  sulphuric  acid  in  the  copper  salt.  The 
most  marked  change  in  the  sulphur  compounds  is  in  the  case  of 
neutral  sulphur,  which  shows  a  decided  and  uniform  increase  dur- 
ing the  administration  of  the  copper  salt  and  in  some  cases  for 
several  days  thereafter. 

The  effect  produced  upon  the  metabolism  of  nitrogen  is  more 
important.  Under  the  administration  of  sulphate  of  copper  there 
is  a  marked  and  constant  decrease  in  the  excretion  of  urea,  which 
is  a  matter  of  great  significance.  Such  a  decrease  can  only  be 
regarded  as  an  indication  of  a  retarding  effect  on  nitrogen  metab- 
olism.  At  the  same  tune  the  quantity  of  uric  acid  and  xanthin 
bases  are  increased  during  the  administration  of  the  copper  salt  and 
'  the  increase  in  xanthin  is  still  very  marked  in  the  after-period. 
These  two  important  observations  indicate  that  the  nitrogen  metab- 
olism is  disturbed  in  a  way  which  must  be  considered  injurious  to 
health. 

There  is  also  a  notable  effect  produced  upon  the  phosphoric  acid 
metabolism.  There  is  a  marked  decrease  in  the  total  metabolized 
phosphorus,  and,  while  the  non-metabolized  phosphorus  is  less  uni- 
formly  aflFected,  there  is  a  decided  tendency  shown  to  decrease  the 


324  WILEY— INFLUENCE  OF  PRESERVATIVES  [April  ts 

total  excretion  of  phosphoric  acid  under  the  influence  of  the  copper 
sulphate. 

The  final  conclusion,  based  on  the  medical  and  clinical  data  and 
on  the  study  of  the  effect  of  the  copper  sulphate  upon  metabolism, 
is  that  the  administration  of  this  salt  is  prejudicial  to  health. 

Potassium  Nitrate. — It  is  evident  that  the  administration  of 
small  quantities  of  potassium  nitrate  induce  only  slight  disturbances 
in  the  metabolic  processes,  and  indicate  only  to  a  slight  degree  harm- 
ful or  deleterious  effects  as  noted  in  the  medical. and  clinical  data. 
It  is  evident  moreover  that  with  the  exception  of  one  instance, 
namely,  the  increase  of  the  number  of  red  corpuscles  in  the  blood, 
that  no  beneficial  effect  can  possibly  be  attributed  to  the  exhibition 
of  this  chemical. 

While  the  data  are  in  this  case  far  less  conclusive  than  those  in 
any  of  the  preceding  cases,  they  are  of  a  character  to  warrant  the 
suggestion  that  so  far  as,  health  and  digestion  are  concerned  it  is 
safer  to  omit  a  body  of  this  kind  from  the  food.  There  are  some 
foods  which  naturally  contain  small  quantities  of  potassium  nitrate. 
Its  very  poisonous  action  when  taken  in  large  doses,  however,  is  a 
warning  which  should  cause  great  care  in  its  use  even  in  small 
quantities  and  deter  any  one  charged  with  the  protection  of  the 
public  health  fit>m  expressing  any  favorable  opinion  in  respect  to 
its  use. 

It  is  evident  that  potassium  nitrate  in  the  quantities  used  has 
neither  a  preserving  effect  nor  has  it  any  condimental  value.  What- 
ever may  be  said  to  the  contrary,  it  is  perfectly  evident  that  the  sole 
purpose  of  its  use  is  the  intensification  of  the  red  color  of  meats 
after  preservation.  Whatever  may  be  the  ethical  principle  under- 
lying this  use  of  potassium  nitrate  is  a  question  which  is  not  the- 
subject  of  discussion  in  a  bulletin  of  this  kind,  but  it  is  only  due 
to  the  consumer  that  the  real  purpose  of  using  potassium  nitrate  in 
the  curing  of  meats  should  be  revealed. 

The  further  question  arises  as  to  whether  or  not  the  coloring  of 
preserved  meats  in  this  way  in  order  that  they  may  have  the 
color  of  fresh  meats  is  a  violation  of  the  Food  and  Drugs  Act, 
which  forbids  the  coloring  of  food  products  for  the  purpose  of  con- 
cealing damage  or  inferiority. 


1908.1  UPON   HEALTH  AND  METABOLISM.  325 

While,  therefore,  the  data  which  have  been  accumulated  are  not 
such  as  to  warrant  a  sweeping  condemnation  of  potassium  nitrate  in 
foods,  they  are  sufficiently  indicative  to  justify  the  conclusion  that 
its  presence  in  foods  is  undesirable  and  open  to  suspicion. 

General  Considerations. 

Having  thus  set  forth  the  general  results  of  this  long  and  labor- 
ious study,  it  is  seen  that  if  the  conclusions  based  upon  the  experi- 
mental data  are  correct  that  there  can  be  no  justification  of  the  proc- 
ess of  adding  chemical  preservatives  to  human  foods.  Successful 
manufacturing  establishments  have  demonstrated  beyond  peradven- 
ture  that  better,  more  wholesome,  and  more  permanent  forms  of  food 
products  can  be  produced  without  the  aid  of  any  preservative  what- 
ever. Sterilization  will  preserve  sweet  cider  better  than  benzoate  of 
soda.  Proper  care  in  handling  fruits  and  in  conducting  the  manu- 
facturing processes  for  preserves,  jams  and  marmalades  will  make 
a  more  palatable  product  and  one  that  keeps  better  than  the  use  of 
salicylic  acid.  Careful  curing  of  meats  and  proper  care  in  trans- 
portation will  preserve  these  meats  better  than  boric  acid.  The 
natural  color  of  the  pea  kept  in  a  sanitary  can  where  its  color  is 
not  lost  by  action  due  to  imperfections  of  the  tin  will  make  a  far 
more  palatable  article  than  will  the  use  of  sulphate  of  copper,  and 
so  on  to  the  end  of  the  list.  There  is  no  single  food  product  which 
is  not  more  palatable  and  of  equal  if  not  better  keeping  qualities 
when  made  carefully  without  the  use  of  preservatives.  There  is, 
therefore,  absolutely  no  commercial  necessity  for  the  use  of  these 
bodies,  but  it  is  urged  by  those  who  employ  them  that  even  though 
considerable  quantities  of  these  bodies  are  injurious  to  the  health, 
which  no  one  denies,  yet  in  the  minute  quantities  in  which  they  are 
used  in  foods  they  can  not  be  regarded  as  in  any  way  deleterious. 
It  is  easy  to  show  that  such  an  opinion  is  without  scientific  basis. 
It  is  quite  impossible  for  any  expert  who  holds  this  opinion  to  indi- 
cate to  any  jury,  much  more  to  the  great  jury  of  the  American  peo- 
ple any  point  in  the  addition  of  the  preservative  to  food  at  which 
it  remains  harmless,  or  the  point  at  which  it  beginsfto  be  harmful. 
Unless  such  a  point  could  be  fixed  and  demonstrated  upon  reliable 


326  WILEY— INFLUENCE  OF  PRESERVATIVES  lAprlTas, 

experimental  data,  it  is  evident  that  no  scientific  reason  can  be  urged 
for  the  use  of  limited  quantities  of  a  preservative,  which  is  acknowl- 
edged to  be  harmful,  on  the  ground  that  in  such  quantities  it  is  not 
injurious. 

Inasmuch  as  a  preservative  is  not  a  food,  and  as  it  does  not  in 
any  way  take  part  in  the  nourishment  of  the  body  nor  in  the  res- 
toration of  waste  or  growth ;  and  further  as  it  is  necessarily  elimi- 
nated, either  unchanged  or  in  other  forms  which  may  be  even  more 
harmful  than  the  original,  by  the  excretory  organs  of  the  body,  thus 
imposing  upon  them  an  unnecessary  and  injurious  burden  and  affect- 
ing more  or  less  the  constitution  of  the  ultimate  cells  thereof  in  an 
imfavorable  way,  it  is  evident  that  the  argument  which  would  per- 
mit their  use  in  small  quantities  is  wholly  illegitimate. 

The  fallacy  of  the  argument  that  small  quantities  of  an  injurious 
substance  are  not  injurious  may  perhaps  be  best  represented  graph- 
ically. The  chart  which  accompanies  this  discussion  shows  theoret- 
ically the  normal  and  lethal  dose  of  a  food  and  a  drug  or,  as  in  this 
case,  a  chemical  preservative.  The  chart  shows  two  curves,  one 
representing  a  chemical  preservative  and  one  representing  a  food. 
The  normal  dose  of  a  food  is  that  quantity  of  food  which  maintains 
a  healthy  adult  body  in  equilibrium.  It  is  represented  on  the  right 
of  the  chart  by  the  number  lOO.  If  the  quantity  of  food  necessary 
to  maintain  the  equilibrium  in  a  healthy  adult  body  is  slightly  dimin- 
ished, no  apparent  change  is  at  first  experienced  and  possibly  even 
no  discomfort.  If,  however,  the  quantity  of  food  be  still  further 
diminished  progressively,  as  indicated  by  following  the  curve  down 
to  the  left,  the  point  is  finally  reached  when  no  food  is  given  at  all 
and  death  ensues,  represented  by  zero  on  the  left  hand  of  the 
diagram  designated  "  lethal  dose."  As  the  curve  begins  to  deviate 
from  the  perpendicular  on  the  right  the  degree  of  injury  is  very 
readily  noticed  and  starvation  or  symptoms  of  starvation  are  set  up. 
Thus,  if  you  follow  the  perpendicular  on  the  right  downward  to  the 
point  80,  the  divergence  of  the  corresponding  point  of  the  curve  is 
already  measurable.  As  you  descend  to  zero  the  magnitude  of  the 
measurement  increases.  It  requires  but  very  little  further  illustra- 
tion to  show  how  easily  the  effect  of  diminishing  the  normal  dose  of 


x9o8.] 


UPON   HEALTH   AND   METABOLISM. 


327 


a  food  can  be  measured  immediately  after  the  curve  begins  to  vary 
appreciably  from  the  perpendicular  on  the  right. 

Let  us  now  consider  the  perpendicular  on  the  left,  which  is 
marked  at  the  top  imder  the  term  "  lethal  dose,"  viz. ;  a  quantity  of 
the  added  preservative  sufficient  to  destroy  life.  The  normal  dose  of 
such  an  added  chemical  preservative  is  o,  and  is  shown  at  the  base 
line  to  the  right  marked  "  normal  dose."  If  you  add  a  very  minute 
quantity  of  a  chemical  preservative,  the  curve  representing  it  varies 


LETHAL  00S£ 
100  > 


lerHAL  D05£ 


NOfiMAL  OOSi 
100 


75 


NORMAL  DOSE 

Fig.    I.    Graphic   chart   representing   the  comparative   influences   of    foods 

and  preservatives. 


so  slightly  from  the  horizontal  base  as  to  be  impossible  of  measure- 
ment by  ordinary  means.  If  we  follow  along  to  the  number  75,  on 
the  horizontal  base,  we  see  the  deviation  of  the  curve  is  sufficiently 
great  to  measure.  At  50  it  is  still  greater,  at  25  still  greater,  while 
at  the  left  of  the  basic  line  it  is  a  maximum,  extending  from  o  to 
100,  or  the  lethal  dose.  It  is  easy  to  show  by  mathematical  data  that 
no  matter  how  small  the  quantity  of  an  injurious  substance  or  pre- 
servative is,  it  will  still  produce  an  injurious  effect,  which  may  be 
infinitely  small  if  the  dose  be  infinitely  small.     It  follows  then,  as  a 


328  WILEY- INFLUENCE  OF  PRESERVATIVES  [April  .5, 

mathematical  demonstration,  that  any  quantity  of  an  injurious  sub- 
stance added  to  a  food  product  must  of  necessity  be  injurious,  pro- 
vided it  is  in  the  nature  of  a  drug  and  the  body  is  in  a  perfectly 
healthy  normal  condition. 

Hence  the  argument  which  has  been  so  persistently  urged  in 
favor  of  a  chemical  preservative  that  if  in  small  quantities  it  is 
harmless  is  shown  to  be  wholly  untenable.  Where  there  is  no  neces- 
sity for  the  addition  of  a  harmful  substance,  where  no  particular 
benefit  is  secured  thereby,  and  where  there  is  no  disturbance  of  the 
normal  state  of  health  there  can  be  no  possible  excuse  of  a  valid 
nature  to  offer  for  the  exhibition  of  even  minute  quantities.  That 
these  minute  quantities  would  not  be  dangerous,  in  so  far  as  pro- 
ducing any  fatal  effect  is  concerned,  is  conceded,  but  that,  in  the 
end,  they  do  not  produce  any  injury,  even  in  these  small  quantities, 
is  certainly  to  be  denied. 

The  course  of  safety,  therefore,  in  all  these  cases  is  to  g^ard  the 
opening  of  the  door.  If  the  use  of  small  quantities  is  permitted, 
then  there  can  never  be  any  agreement  among  experts  or  others 
respecting  the  magnitude  of  the  "  small  quantity,"  and  continued 
litigation  and  disagreement  must  follow.  On  the  other  hand,  when 
the  harm  fulness  of  any  substance  which  it  is  proposed  to  add  to 
food  is  established  and  no  reason  for  its  use  can  be  given  other  than 
the  convenience,  carelessness,  or  indifference  of  the  manufacturer, 
the  exclusion  of  such  bodies  entirely  from  food  products  follows  as 
a  logical  sequence  and  a  hygienic  necessity. 


THE  HUMMING  TELEPHONE, 

A  Contribution  to  the  Theoretical  and  Practical  Analysis 

OF  ITS  Behavior. 

By  a.  E.  KENNELLY  and  WALTER  L  UPSON. 
(Received  July  20,  1908.) 

The  following  paper  describes  the  salient  features  of  an  experi- 
mental research  on  the  humming  telephone,  conducted  in  the  Gradu- 
ate School  of  Applied  Science  of  Harvard^  University  during  the 
year  1907-08,  and  discusses  an  elementary  mathematical  theory 
which  the  observations  appear  to  indicate  and  support. 
'     Definition, — ^A  "  humming  telephone  "  is  a  connection  of : 

1.  A  telephone  receiver,  or  ordinary  hand  'phone. 

2.  A  telephone  transmitter,  or  ordinary  carbon  microphone. 

3.  A  source  of  electric  power,  such  as  a  voltaic  battery  and  tele- 
phone induction  coil,  with  the  receiver  in  such  electric  and  acoustic 
relation  to  the  transmitter,  that  it  is  able  to  emit  a  sustained  note 
or  hum.  This  auto-excited  hum  may  be  so  loud  as  to  be  heard  in 
a  distant  room  through  several  partitions. 

Historical  Outline. — The  fact  that  a  telephone  receiver  held, 
either  in  contact  with,  or  close  to,  the  face  of  its  transmitter  may 
cause  the  production  of  a  hum  or  singing  tone,  appears  to  have  been 
first  observed  by  Mr.  A.  S.  Hibbard.^  This  experimental  fact  is 
now  well  known  to  telephonists.  In  many  cases,  it  is  only  necessary 
to  lift  a  subscriber's  telephone  from  its  hook,  and  hold  it  face  to 
face  with  its  transmitter,  in  order  to  produce  a  loud  hum. 

The  only  published  investigation  of  the  humming  telephone  that 
the  authors  have  succeeded  in  finding  is  an  important  paper  by  Mr. 

* "  Investigation  of  the  Phenomena  of  *  The  Humming  Telephone,' "  by 
Walter  L.  Upson,  a  thesis  towards  the  degree  of  master  of  science  in  elec- 
trical engineering,  Harvard  University,  1908. 

*  September,  1890.     See  Gill's  paper  hereafter  referred  to. 

329 


330  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         [Julyao, 

F.  Gill,'  read  before  a  meeting  of  the  Dublin  Local  Section  of  the 
Institution  of  Electrical  Engineers  in  April,  1901.  Very  briefly,  the 
salient  experimental  facts  reported  in  this  valuable  paper  are: 

1.  The  reversal  of  the  telephone  receiver  connections  in  the  cir- 
cuit alters  the  pitch  of  the  auto-excited  tone,  the  pitch  being  higher 
for  one  direction,  and  lower  for  the  other  direction,  of  connection. 

2.  The  pitch  of  the  tone  may  also  be  altered  by  changing:  (a) 
the  inductance,  capacity  or  resistance  of  the  circuit,  or  circuits;  (b) 
the  strength  of  current  in  the  microphone  transmitter;  (c)  the  dis- 
tance between  the  receiver  and  transmitter  diaphragms;  (d)  pres- 
sure on  either  of  the  diaphragms. 

The  Gill  paper  does  not  discuss  the  theory  of  the  subject  beyond 
suggesting  that  the  phase  retardation  of  the  acoustic  impulses  reach- 
ing the  transmitter  from  the  receiver  has  a  controlling  influence  on 
the  pitch  of  the  tone. 

The  research  reported  in  this  paper  may  be  regarded  as  extend- 
ing the  investigation  from  the  stage  reached  in  Gill's  paper  to  a 
stage  which  admits  of  a  first  approximation  theory.  A  large  amount 
of  research  remains,  however,  to  be  carried  on  in  the  future,  before 
the  experimental  and  theoretical  analysis  of  this  fascinating  but 
.complex  phenomenon  can  be  regarded  as  satisfactorily  nearh 
complete. 

Method  of  Observation  Employed, — As  pointed  out  in  Gill's 
paper,  the  pitch  of  the  note  emitted  by  the  humming  telephone, 
although  substantially  constant  under  fixed  conditions,  is  affected  by 
almost  any  change  in  the  apparatus,  in  a  seemingly  most  intricate 
manner.  In  order,  therefore,  to  study  the  effect  of  varying  one 
particular  variable  at  a  time,  the  device  was  hit  upon  of  acoustically 
connecting  the  receiver  and  transmitter  diaphragms  in  a  definitely 
controllable  way  by  means  of  telescoping  tubes  fitting  on  to  the 
receiver  and  transmitter  faces.  These  tubes,  and  also  the  standard 
electric  connections  employed,  are  indicated  in  Fig.  i. 

The  transmitter  was  kept  stationary,  with  one  end  of  the  tube 
covering  and  secured  to  its  cone.  The  receiver  was  fastened,  on 
a  sliding  wooden  carriage,  to  the  other  end  of  the  telescoping  tube. 

* "  Note  on  a  Humming  Telephone,"  by  F.  Gill,  Journal  of  the  Institution 
of  Electrical  Engineers,  1901-02,  Vol.  XXXI.,  No.  153,  pp.  388-399. 


1908.] 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


331 


The  distance  between  the  faces  of  the  two  instruments  could  be 
varied  at  will  by  pulling  out,  or  pushing  in,  the  telescoping  tube- 
sections.  The  average  current  in  the  primary  circuit  was  measured 
with  a  Weston  d.c.  milliammeter.  The  pitch  of  the  humming  note 
was  measured  approximately  by  the  ear,  with  the  aid  of  a  number 
of  short  organ  pipes,  and,  in  some  instances,  with  the  aid  of  a  violin. 
The  voltaic  battery  used  consisted  of  a  selected  number  (from  two 
to  nine,  but  usually  four)  of  2S-ampere-hour  lead  storage  cells.  The 
reversing  switch  in  the  secondary  circuit  enabled  the  receiver  ter- 
minals to  be  reversed  at  will. 


Transmitter, 


Reeeiirer, 


rrft 


Tulie  Lengtti 


ftil'/lmmeter 


fl'it 


^.!^ 


Battery.  induehon  Coil,  ftwtning  Smteli, 

Fig.  I.    Diagram  of  Humming  Telephone  Connections. 

The  Telescoping  Tubes, — The  tubes  were  made  of  heavy  wrap- 
ping paper.  Their  internal  diameters  varied  from  5  cm.  (2  in.)  to 
6  cm.  (2J  in.).  They  were  used  in  lengths  of  65  cm.  (25^  in.), 
with  a  few  shorter  and  longer  sections  for  special  measurements. 
The  substance  of  which  the  tubes  was  composed  did  not  appreciably 
affect  the  observations.  It  Was  found,  however,  that  if  the  tele- 
scoping sections  did  not  fit  fairly  tightly,  erratic  results  were  ob- 
tained.   Closely  fitting  sections  were  used. 

The  Transmitters. — The  transmitters  used  were  of  the  standard 
Western  Electric  Co.  type  and  manufacture.  The  diaphragm  in 
these  instruments  was  of  aluminum,  6.32  cm.  (2.49  in.)  in  total 
diameter,  and  0.55  mm.  (0.022  in.)  thick,  over  a  coating  of  Japan 
varnish  on  one  face.  The  diaphragm  was  loaded  at  its  center  with 
one  of  the  disk  electrodes  of  the  carbon  microphone.  The  dia- 
phragm was  damped  by  being  clamped  between  rubber  rings  to  an 


332 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         [Ja'y«>. 


internal  diameter  of  4.8  cm.  (1.9  in.),  and  also  by  the  application 
of  a  pair  of  rubber-tipped  flat  metal  springs  to  areas  between  the 
center  and  edge.  The  resistance  of  the  microphone  varied  between 
the  approximate  limits  of  20  ohms  when  quiescent,  and  no  ohms 
when  in  powerful  vibration. 

The  Receivers, — The  receivers  used  in  most  of  the  measurements 
were  of  the  standard  bipolar  Western  Electric  Co.'s  type,  known  as 
No.  122,  having  poles  1.4X0.2  cm.  (0.55  X0.08  in.),  separated 
by  0.82  cm.  (0.325  in.).  They  had  a  resistance  of  210  ohms,  and 
an  inductance  of  0.025  henry,  at  a  frequency  of  1,000  r^.  With 
steady  currents,  their  resistance,  at  15®  C,  was  about  70  ohms.  The 
diaphragm  of  varnished  ferrotype  iron  had  an  external  diameter  of 
5.5  cm.  (2.17  in.),  a  clamping  diameter  of  4.95  cm.  (1.95  in.)  and 
a  thickness,  over  varnish,  of  0.292  mm.  (0.0115  in.).  Its  weight 
was  4.0  grammes. 

The  Induction  Coil, — The  induction  coil  used  was  of  the  standard 
Western  Electric  Co.'s  type,  known  as  No.  13.  Its  resistances  and 
inductances  were  taken  as  follows:* 


Table  I. 


Freauency 

Cycles  per 

Second. 

Resistance  at  i8<>  C. 

'                Self-Inductance. 

Mutual 

Primary 
Ohms. 

Secondary 
Ohms. 

1 

Primary 
Henrys. 

Secondary 
Henrys. 

Inductance, 
Henrys. 

0 
I,CXX) 

1.62 
3-2 

1 
20.3 
48.1 

0.0044 

0.0765 

0.0172 

The  principal  dimensions  of  the  coil  were:  Length  over  all  8.2 
cm.  (3.16  in.).  Interflange  6.3  cm.  (2.5  in.).  Diameter  over  out- 
side cover  2.5  cm.  (i  in.).  Internal  diameter  of  core  tube  0.75  cm. 
(0.296  in.).  Diameter  of  iron  wires  in  core  0.0356  cm.  (0.014  in.). 
Total  number  of  iron  wires  in  core  about  75. 

Observation  Series  No  /.  Effect  of  Shortening  the  Tube. — 
Commencing  with  the  connections  of  Fig.  i,  a  battery  of  8.6  volts, 
and  a  tube  length  of  267  cm.,  as  indicated  in  Fig.  2  on  the  scale  of 
abscissas,  a  loud  steady  note  between  G"*  and  A"  (850  ^)  was  sus- 
tained in  the  telephone.    The  pitch  of  this  note  is  shown  at  P  on  the 

*The  data  for  the  coil  at  1,000 —  were  kindly  supplied  by  the  engineering 
department  of  the  Western  Electric  Co. 


X9o8.] 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


333 


upper  ziz-zag  line  /.  The  current  strength,  on  the  d.c.  milliammeter, 
as  shown  at  p  on  the  lower  ziz-zag  line  I,  was  130  milliamperes. 
When  the  telescopic  tube  was  gradually  shortened,  the  pitch  of  the 
note  steadily  rose,  until  it  reached  Q,  at  A!'t  (92O'-'),  with  240 
cm.  of  tube-length,  and  a  primary  current  strength  q  of  200  mas. 
The  intensity  of  the  note  near  920 /^  was  ordinarily  somewhat 
weaker  than  when  near  825  ^,  On  continuing  to  shorten  the  tube, 
the  pitch  suddenly  broke  from  Q,  at  920  r^,  to  R  at  825  r^.    Pushing 


fA 


mo 


^    mo 
o 


I     I     I     I     I     I     I     I     I '«'''' 


so       so 


''''»' 


409 


mo       mo      MO      im>       too      mmo      zmo     moo 
TUB£  LENGTH  •  CENTIMETERS 

Fig.  2.    Effect  of  Shortening  Tube,  and  of  Reversing  Receiver  Connections. 

in  the  tube  further,  the  pitch  would  again  climb  steadily  to  T,  at 
201  cm.,  with  a  new  maximum  of  current.  Beyond  this  point,  the 
pitch  would  break  suddenly  to  U  at  810  r^.  Again  it  would  climb 
to  W,  at  170  cm.  and  suddenly  collapse  to  X,  Continuing  in  this 
manner,  the  pitch  would  alternately  rise  to  maxima  and  break  sud- 
denly to  minima,  along  the  pitch  zig-zag  /.  At  the  breaks  of  pitch, 
the  current  would  sometimes  break  to  a  lower  value,  as  at  t,  u;  or 
break  to  an  upper  value,  as  at  w,  x;  or  vary  suddenly  in  rate  of 
change,  without  discontinuity  in  magnitude,  as  at  q.  Repeating  the 
experiment,  the  zig-zag  lines  of  pitch  and  of  current  would  be 
repeated,  not  exactly  but  substantially,  the  variations  being  due  not 

PROC.  AMER.  PHIL.  SOC.,  XLVII.  189  V,  PRINTED  OCTOBER  2,  I908. 


334  KENNELLY  AND  UPSON- HUMMING  TELEPHONE.         [Julyao, 

merely  to  observational  error,  but  also  to  variations  in  the  behavior 
of  the  transmitter. 

The  zig-zag  pitch  line  PQRST  is  found  to  be  somewhat  irregu- 
lar. 'The  slants  are  by  no  means  r^^larly  parallel.  The  breaks 
QTW  are  neither  regularly  elevated,  nor  regularly  spaced.  The 
only  substantial  regularity  is  in  the  spacing  along  the  pitch  line 
G"t  of  825  r^.  The  intersections  of  the  ascending  branches  with 
this  line  lie /approximately  40  cm.  apart,  at  no,  150,  190,  230  and 
270  cm.,  or  in  accordance  with  the  series  30  +  4om  cm.,  where  m 
is  any  positive  integer. 

As  regards  the  current  curve  pqrst,  its  points  of  minima  p,  r, 
V,  etc.,  correspond  fairly  well  to  the  ascending  intersections  of  the 
pitch  line  with  the  line  of  G"^  825  r^.  The  points  of  maxima  q,  t,  x, 
etc.,  occur  near  to  the  breaks  in  the  pitch  Q,  T,  W,  etc.  Minimum 
primary  current  was  noticed  to  be  associated  with  maximum  micro- 
phonic activity  of  vibration.  Feeble  action  in  the  microphone,  on 
the  other  hand,  was  found  to  be  associated  ordinarily  with  increase 
of  primary  current. 

Observation  Series  2,  Effect  of  Shortening  the  Tube  with  Re- 
versed Receiver  Terminals, — Curves  II,  in  Fig.  2,  represent  the 
behavior  of  note  pitch  and  primary  current,  as  the  tube  was  short- 
ened from  265  cm.  to  80  cm.,  with  the  terminals  of  the  receiver 
reversed.  Their  general  characters  are  similar  to  those  of  curves  /. 
The  two  sets  of  curves  indicate  the  effect  which  would  be  produced 
by  reversing  the  receiver  terminals  at  any  particular  tube-length 
within  the  above  range.  Thus,  at  5",  or  220  cm.,  a  reversal  would 
lower  the  pitch  from  870  ^  on  curve  /  to  K,  at  810  r^,  on  curve  //. 
On  the  other  hand,  a  reversal  made  on  curve  /,  at  V,  of  825  /^, 
would  raise  the  pitch  to  N  of  900  r^  on  curve  II,  so  that  whether 
the  reversal  produces  a  rise  or  fall  of  pitch  depends,  in  general, 
upon  whether  the  reversal  is  effected  above  or  below  the  mean  pitch 
of  G"«,  825  ^. 

The  only  apparent  regularity  in  the  pitch  line  //  lies  in  the 
spacing  of  the  ascending  intersections  with  the  line  of  mean  pitch 
G't  (825  ^).  These  occur  near  to  90,  130,  170,  210  and  250  cm. 
of  tube-length,  or  according  to  the  series  10  + 40m  cm.  On  the 
mean-pitch  line,  the  ascending  intersections  of  one  curve  lie  ap- 


X908.1 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


335 


proximately  20  cm.  from,  or  midway  between,  those  of  the  other 
curve. 

The  note  frequencies  and  primary  current  strengths  for  tubes 
of  less  than  60  cm.  in  length  are  given  in  Fig.  3,  commencing  at 
60  cm.  and  shortening  down  to  about  i  cm.,  when  the  receiver  face 
came  into  contact  with  the  transmitter  face  (cone  removed),  and 
so  prevented  closer  approach.    Curves  /  and  //  of  Fig.  3  correspond 


Fig.  3. 


to  20  90  to  so 

TUBE  LENGTH  -  CENTIMETERS 

Humming  Note  Frequencies  and  Primary  Current  Strengths  with 

Short  Tubes. 


to  curves  /  and  //  of  Fig.  2,  respectively,  and  indicate  the  effect  of 
reversing  the  receiver  terminals.  It  may  be  observed  that  follow- 
ing the  pitch  line  I,  the  ascending  branch  intersects  the  mean  fre- 
quency line  of  825  r^,  at  a  tube-length  of  30  cm.,  for  the  last  time. 


336  KENNELLY  AND  UPSON-HUMMING  TELEPHONE.        [July  «>, 

Shortening  the  tube  beyond  this  point,  the  pitch  rises  until  it  reaches 
e'"  of  1,300  r^y  at  12.5  cm.,  and  at  a  primary  current  strength  of 
300  mas.  Here  the  note  breaks  without  descending  to  a  new  low 
note.  There  is  silence  with  this  connection  of  the  receiver  between 
12.5  cm.  and  o  cm.  With  the  transmitter  and  receiver  touching 
each  other,  kt  was  possible  to  produce  almost  any  note  between 
620 '-^^  and  1,300 /-',  by  giving  suitable  opening  to  the  air  at  one 
side.  If,  however,  the  outside  air  was  shut  off,  and  the  air  between 
the  transmitter  and  receiver  diaphragms  was  cylindrically  enclosed, 
by  bringing  their  faces  into  full  opposition  and  contact,  no  note 
could  be  obtained. 

If  we  follow  pitch  curve  //,  we  find  that  the  ascending  branches 
intersect  the  mean-frequency  line  at  50  cm.  and  at  10  cm.  The 
pitch  866  r^  was  obtained  steadily  when  the  transmitter  and  receiver 
faces  were  in  full  contact,  corresponding  to  a  "tube-length"  of 
I  cm.  With  this  connection  of  receiver  terminals,  no  other  note, 
or  variety  of  notes,  could  be  obtained  at  contact. 

A  telescoping  tube  of  9  meters  (29.5  ft.)  total  length  was  used 
in  one  series  of  measurements,  and  the  results  appear  in  Fig.  4. 
They  were  all  obtained  with  diminishing  tube-lengths,  or  with  com- 
pression of  the  telescoping  tube.  The  small  crosses  indicate  dis- 
continuities produced  at  the  removal  of  sections  of  tube  when 
finished  with.  In  regard  to  the  pitch  line,  it  will  be  seen  that  it 
corresponds  to  curve  /  of  Figs.  2  and  3.  That  is,  it  crosses  the 
mean- frequency  line  of  825  r^  ascendingly  at  30  +  40m  cm.  with 
a  fair  degree  of  precision.  With  the  shortest  tube,  the  range* in 
pitch- frequency  was  from  740  ^-^^  to  1,060 '--',  or  through  320^--'. 
At  the  full  length  of  9  meters,  this  range  fell  to  75  ^,  The  ultimate 
limit  tended  apparently  to  the  mean-pitch  frequency  of  G"^  825  r^. 
The  average  note  was  above  this  pitch;  but  this  was  probably  be- 
cause the  tube  was  being  compressed.  Reference  to  Fig.  5  will 
show  that,  when  shortening  the  tube,  the  average  pitch  lies  above 
the  mean  of  825  r^ ;  while  in  lengthening  the  tube,  the  average 
pitch  lies  below. 

The  primary  current  strength  in  Fig.  4  tends,  in  general,  to 
minima  at  the  mean- frequency  pitch  of  825  r^,  and  to  maxima  at 
the  breaks.    The  differences  in  current  strength  become,  however. 


.jol]  KENNELLY   AND  UPSON— HUMMING  TELEPHONE.  337 


iiiiiiii* 

-iM3Mm  310N        taiuirrmiii  -  iNJumi 


338 


KENNELLY   AND  UPSON— HUMMING  TELEPHONE. 


[July 


less  marked  as  the  tube  is  longer,  the  minimum  currents  rising, 
as  the  length  increases,  by  about  40  mas.  in  9  meters,  indicating 
steadily  reduced  action  in  the  transmitter  with  increasing  distance. 
Since  the  current  rose  to  260  mas.  when  the  transmitter  diaphragm 
was  entirely  out  of  action,  we  should  expect,  at  this  rate,  to  be  able 
to  sustain  the  humming  note  to  a  total  tube-length  of  40  meters; 
but  no  tests  were  actually  made  beyond  9  meters. 


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aO  MfO  mo  490  900         M90 

TUBE  LENGTH  -  CENTIMETERS. 

Fig.  5.    Effect  of  Lengthening  and  Shortening  the  Tube. 


Observation  Series  5.  Effect  of  Lengthening  the  Tube. — Fig.  5 
indicates  the  relative  effects  produced  by  lengthening,  as  compared 
with  shortening,  the  telescoping  tube  joining  the  transmitter  and 
receiver  in  Fig.  i,  using  the  same  apparatus  and  connections  as  in 
Figs.  I,  2,  3  and  4.  The  heavy  or  continuous  lines  in  Fig.  5  show 
the  effects  of  shortening  the  tube,  or  correspond  to  curves  /  in  Fig. 
2.  The  broken  lines  show  the  effects  of  lengthening  the  tube.  It 
will  be  observed  that  the  points  of  maximum  and  minimum  current 
agree  fairly  well.  The  ascending  intersections  of  the  pitch  lines 
with  the  mean-frequency  line  of  G">  825  ^,  lie  near  together,  and 
approximately  conform  to  the  series  30  +  40m  cm.  of  tube-length. 
The  points  of  break  in  pitch  do  not,  however,  agree,  and  the  dis- 


i9o8.] 


KENNELLY  AND  UPSON— HUMMING   TELEPHONE. 


839 


tances  between  corresponding  pairs  of  breaks  in  pitch  increase  as 
the  tube-length  is  greater,  being  4  cm.  at  A,  9  at  B,  11  at  C,  13 
at  D,  and  15  at  E.  Although  not  shown  in  Fig.  5,  owing  to  limi- 
tations of  space,  it  was  found  that  these  distances  between  corre- 
sponding breaks  continued  to  increase  until  they  reached  about  20 
cm.,  after  which  they  shortened  again  to  commence  a  new  expand- 
ing series. 


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TUBE  LENGTH  -  CENTIMETERS. 


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Fig.  6.    Humming  Cycles  with  Cyclic  Changes  in  Tube-length. 

Observation  Series  4.  Effect  of  Alternately  Reversing,  or  Re- 
ciprocating, the  Motion  of  the  Tube.  Humming  Cycles. — If,  when 
compressing  the  telescopic  tube,  and  when  the  note  broke  from  a 
higher  to  a  lower  pitch,  the  tube  was  immediately  extended  again, 
the  note  would  continue  to  lower  in  pitch  for  a  little  while,  and  then 
break  back  to  a  higher  pitch.    By  moving  the  tube  in  and  out,  like 


340  KENNELLY   AND  UPSON— HUMMING  TELEPHONE.         [july«o, 

a  concertina,  over  this  range,  the  pitch  would  break  to  and  fro  in 
a  very  regular  way.  The  corresponding  reverse  action  would  also 
occur  if  the  motion  commenced  with  extension.  These  conditions 
are  shown  in  Fig.  6.  Commencing  at  the  point  O,  with  no  cm. 
of  tube-length,  on  the  mean  frequency  of  825  ^,  if  we  shorten  or 
compress  the  tube  to  90.5  cm.,  we  reach  P  at  900  ^,  near  A">. 
The  note  then  breaks  to  Q  at  780^,  Increasing  the  tube-length 
back  to  95  cm.,  we  reach  R  at  770  ^,  The  note  then  breaks  up- 
wards to  5*  at  880  ^.  This  humming  cycle  PQRS,  could  be  repeated 
indefinitely  with  a  considerable  degree  of  precision  as  to  pitch  and 
tube-length;  but  with  a  more  moderate  degree  of  precision  as  to 
primary  current  strength.  Similarly,  the  cycle  TUVW,  of  10.5  cm. 
amplitude  in  length,  and  100  ^^  amplitude  in  pitch,  might  be  re- 
peated indefinitely.  The  amplitudes  and  areas  of  these  humming 
cycles  vary  at  different  breaking  points. 

•  Purity  of  Humftting  Tone. — ^With  the  greater  tube-lengths, 
shortly  before  the  break  of  pitch  occurred,  there  was  frequently 
noted  an  appearance  of  the  new  tone  in  advance.  As  the  breaking 
point  was  approached,  the  old  tone  dwindled,  while  the  new  tone 
strengthened.  At  the  break,  the  old  tone,  already  faint,  would  sud- 
denly cease.  Consequently,  before  breaking,  both  the  old  and  new 
tones  might  be  recognized,  forming  a  sort  of  trill,  or  combination 
tone.  This  association  of  simultaneous  tones  had  the  effect  of  main- 
taining the  primary  current  strength  more  nearly  uniform.  With  the 
shorter  tube-lengths,  which  involved  a  greater  jump  of  frequency 
at  the  breaks,  these  combination  tones  were  rarely  heard,  and  the  old 
note  would  break  suddenly  into  the  new  note  without  any  suggestion 
of  a  trill. 

In  some  of  the  observations,  the  notes,  aside  from  the  above- 
mentioned  trilling  near  to  the  breaking  points,  gave  acoustical  evi- 
dence of  multiple  tones.  Occasionally,  the  principal  tone  was  accom- 
panied by  an  octave  overtone.  The  octave  might  be  either  the  first 
octave  below,  or  the  first  octave  above,  the  principal  tone.  Such 
overtones  were  comparatively  faint.  At  other  times,  the  superposed 
tone,  instead  of  being  harmonic  to  the  principal  tone,  appeared  to 
differ  therefrom  by  only  about  one  tone  on  the  musical  scale.  This 
inharmonic  superposed  tone  was  also  relatively  faint  with  respect 


\ 

( 


I908.J  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.  341 

to  the  principal  tone.  Generally,  however,  no  superposed  tones  could 
be  discerned,  and  the  note  was  clear  and  flute-like  in  quality.  Irregu- 
larities in  the  fitting  of  the  telescoping  tube-sections,  or  in  other 
acoustic  connections,  were  found  to  be  productive  of  superposed 
notes. 

Effects  of  Electrical  Changes. 

Observation  Series  5.  Effect  of  Kesistance  in  Primary  or  Sec- 
ondary Circuit, — In  this  test  a  single  tube  of  constant  length  (86.5 
cm.  or  34  in.)  was  used.  It  was  of  pasteboard,  had  an  internal 
diameter  of  5.1  cm.  (2  in.)  and  weighed  IJ3.5  gm.  This  length 
happens  to  be  about  midway  between  the  ascending  intersections  of 
pitch  lines  /  and  //  in  Fig.  2  measured  on  the  mean-frequency  line 
.of  825  ^.  That  is,  the  tube-length  selected  favored  each  of  the  lines 
/  and  //  nearly  equally.  The  battery  e.m.f.  of  8.6  volts  was  the  same 
as  in  all  the  above  descrioed  measurements.  The  same  telephone 
receiver  and  induction  coil  were  also  used.  Substantially  non-induc- 
tive resistance  was  introduced,  by  rheostat,  into  either  the  primary, 
or  the  secondary,  circuit  at  will,  leaving  the  connections  of  Fig.  i 
otherwise  unchanged. 

After  starting  the  loud  humming  note  with  no  extra  resistance 
in  either  circuit,  resistance  was  gradually  inserted  into  the  primary 
circuit  until  the  note,  diminishing  in  amplitude,  finally  disappeared. 
The  extra  resistance  in  the  circuit  at  the  extinction  of  the  tone  was 
recorded,  under  the  name  of  "  extinguishing  resistance."  Resistance 
was  then  withdrawn  from  the  primary  circuit,  and,  after  the  loud 
note  had  been  reestablished,  was  introduced  gradually  into  the  sec- 
ondary circuit,  until  again  the  note  was  extinguished.  The  second- 
ary extinguishing  resistance  was  likewise  recorded.  The  same  tests 
were  repeated  with  the  telephone  receiver  terminals  reversed. 

It  was  found  that  both  the  primary  and  secondary  extinguishing 
resistances  repeated  themselves  very  fairly  (within  about  5  per  cent.) 
in  successive  trials.  In  order  to  obtain  the  best  comparative  results 
in  successive  tests,  it  was  found  desirable  to  tap  the  transmitter 
gently  when  approaching  the  condition  of  extinction. 

The  pitch  of  the  tone  when  enfeebled  almost  to  extinction  by 
extra  resistance,  in  either  the  primary  or  secondary  circuit,  was 
always  close  to  the  mean  frequency  of  825  ^, 


342  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         Uuly«, 

The  amount  of  either  the  primary  or  secondary  extinguishing 
resistance  was  found  to  depend  upon  the  adjustment  and  operative 
condition  of  the  transmitter,  keeping  the  receiver,  tube-length  and 
all  other  conditions  unaltered.  This  led  to  a  trial  of  this  method 
as  a  practical  test  of  microphone  transmitters. 

Observation  Series  50,  Test  /)f  Transmitter  by  Hum^extin- 
guishing  Resistances, — A  nmpber  of  transmitters,  some  good  and 
others  imperfect,  were  tested  under  the  conditions  above  outlined. 
These  transmitters  were  kindly  loaned  for  this  purpose  by  the  West- 
em  Electric  Co.  Twelve  were  regular  standard  instruments  that 
had  already  satisfactorily  passed  the  factory  tests.  These  were 
labelled  T^  to  Tj,  respectively.  Four  more  were  marked  defective 
and  "  down  in  volume."  They  were  labelled  Tj,,  T15,  T^^  and  T2^. 
Four  more  were  marked  defective  and  "  thick  in  quality."  These' 
were  labelled  T^,  T^^,  T22  and  Tjs.  Yet^nother  four  were  marked 
defective  and  "  burning."  These  were  labelled  Tig,  Tj,,  Tjo  and  T^^. 
Defective  transmitters  "  down  in  volume  "  are;  recognized  as  weak. 
Those  which  are  of  "thick  quality"  are  strong  but  defective  in 
articulation.  Those  which  are  "burning"  produce  slight  arcing, 
at  or  near  the  electrodes,  when  subjected  to  normal  conditions  of 
operation. 

The  results  of  the  tests  on  these  24  transmitters  are  given  in  the 
accompanying  table;  where  R  represents  the  primary,  and  r  the 
secondary,  extinguishing  resistance,  when  the  transmitter  was  gently 
tapped.  Care  was  taken  that  the  observer  in  this  test  did  not  know 
the  label  number,  or  reported  condition,  of  the  transmitter  under 
trial.  It  will  be  seen  that  with  the  good  transmitters,  the  mean 
primary  extinguishing  resistances  were  all  included  between  26.5 
and  58.5  ohms,  their  mean  secondary  extinguishing  resistances  be- 
ing between  1,925  and  4,150  ohms.  All  of  the  defective  transmitters 
lay  outside  these  limits,  the  "  down  in  volume  "  being  low,  and  the 
"  thick  quality  "  high,  in  their  extinguishing  resistances ;  except  two 
of  the  "  burning  "  type,  which  fell  within  the  good  secondary  extin- 
guishing resistance  limits.  It  would  seem,  therefore,  that  this  re- 
sistance method  constitutes  a  possible  practical  application  of  the 
humming  telephone  to  transmitter  testing ;  except  that  "  burning  " 
transmitters  may  require  a  separate  test  for  their  detection. 


«9o8.|* 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


343 


The  results  recorded  in  the  last  two  columns  of  Table  11.  are  pre- 
sented graphically  in  the  target  diagram  of  Fig.  7.  The  square  in- 
cludes all  the  good  instruments  and  none  of  the  bad.  The  mean  of 
the  good  transmitters  is  indicated  by  the  solid  black  circle. 

Table  II. 

Table  of  Comparative  Hum-Extinguishing  Resistances  for  12  Good  and  12 

Defective  Transmitters. 


Transmit- 
ter. 

Extinguishing  Resistances. 

Average. 

Quality  of 
Transmitter. 

xst  Position  of  Rec'r. 

ad  Position  of  Rec'r. 

Average, 
r. 

^(Pri.). 

r  (Sec.). 

i?(PrL). 

50 
44 
43 
31 
40 
69 
26 

30 

44 
46 

74 
21 

r(Sec.). 

OK. 

5 

P 

7' 

r" 

''11 

Mean 

43 
23 

55 
25 
45 
31 
27 
30 
31 
47 
43 
ZZ 

36.1 

3,100 
3.050 

4,300 
2,300 

3,900 

3,000 

2,600 

2,600 

2,700 

3,900 

4,100 

2,600 

3,180 

2,300 
3,000 
4,000 
1,900 
2,600 
3,800 
1,600 
1,700 
2,300 
3,900 
3,800 
1,250 

46.5 

33.5 

49 
28 

42.5 

50 

26.5 

30 

37.5 

46.5 

58.5 

27 

2,800 
3,025 
4,150 
2,100 

3,250 
3,400 
2,100 
2,150 
2,500 
3.900 
3,950 
1,925 

43.2 

2,696 
700 

1,000 
1,500 

39.6 

2,937 

Down 

in 

Volume. 

Mean 

10 
6 
6 

17 

750 

390 

900 

1,400 

860 

10 

12 
20 

10 
6 

9 
18.5 

725 

390 

950 

1,450 

9.75 

14 

1,070 

10.9 

879 

Thick 
Quality. 

5* 

Mean 

62 
50 
52 
66 

4,900 
9,000 
7,000 
5,300 

6,550 

66 
70 
75 
78 

2,300 

10,000 

8,000 

6,700 

6.750 

64 
60 

63.5 
72 

3,600 
9,500 
7,500 
6,000 

57.5 

72.5 

64.75 

6,650 

Burning. 

54 
61 
82 
60 

3.800 
4,700 
5,300 
3,900 

68 

93 
102 

61 

3,400 
6,900 
5,000 
2,900 

61 

77 
92 
60.5 

72.6 

3,600 
5,800 

5,150 
3,400 

Mean 

64.25 

4,425 

81 

4,550 

4,487 

Observation  Series  6.  Effect  of  Varying  the  E.M.F.  in  the 
Primary  Circuit, — Among  so  many  variables  and  variations  as  are 
displayed  in  preceding  diagrams,  it  is  comforting  to  find  one  variable 
which  produced  relatively  little  effect  within  certain  practical  limits. 
Fig.  8  shows  the  frequencies  and  primary  currents  for  tube-lengths 


^ 
* 


844 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         Utolyw, 


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4iM^        iM#        #iM0        7JI«r 

RESISTANCE  IN  SECONDARY  -  OHMS. 

Target   Diagram  of  Transmitter  Tests   by  the   Method   of   Hum 
Extinguishing  Extra  Resistance. 


{2 


TUBE  LENGTH  •  CENTIMETERS.  ^ 

Fig.  8.    Frequencies  and  Primary  Currents  for  Different  Primary  E.M.F's. 


1908.1 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


345 


steadily  reduced  from  260  to  70  cm.,  with  batteries  of  3,  4  and  5 
storage  cells,  respectively,  in  the  primary  circuit  (6.5,  8.5  and  10.5 
volts).  The  transmitter,  induction  coil,  receiver  and  transmitter 
were  all  as  in  Figs,  i  to  6.  It  will  be  seen  that  the  primary  cur- 
rents have  their  respective  maxima  and  minima  in  substantial  agree- 
ment, the  range  of  variation  being  naturally  greatest  for  the  largest 
battery,  and  least  for  the  smallest.  The  ascending  intersections  of 
the  frequency  line  with  the  mean-frequency  line  of  825  ^  are  the 
same  throughout,  and  conform  to  the  series  30  +  40t»  cm.,  in  agree- 
ment with  line  /  of  Fig.  2.  The  breaks  in  pitch  do  not  all  coincide ; 
but  the  differences  in  this  respect  are  not  great,  nor  can  it  be  said 


CO 


i 

M  m  liO  tH$  /Si  J99  iO$ 

TUBE  LENGTH  •  CENTIMETERS. 
Fig.  9.    Frequencies  and  Primary  Current  Strengths  for  Different  Condensers 

in  Secondary  Circuit. 

that  the  biggest  battery  always  produced  the  most  retarded  break. 
Moreover,  excepting  perhaps  the  break  at  240  cm.,  the  variations 
in  breaking  points  are  within  the  limits  of  variation  obtained  in  suc- 
cessive series  with  one  and  the  same  battery. 

Observation  Series  7.  Effect  of  a  Condenser  in  the  Secondary 
Circuit. — It  was  found  that  a  certain  magnitude  of  condenser 
capacity  inserted  in  series  in  the  secondary  circuit  had  a  marked 
effect  on  the  behavior  of  the  humming  telephone.     The  results  are 


346  KENNELLY  AND  UPSON—HUMMING  TELEPHONE.        [Juiyao, 

indicated  in  Fig.  9,  for  a  tube-length  commencing  at  270  cm.  and 
steadily  reduced  to  75  cm.,  with  8.6  volts  in  the  primary  circuit  and 
the  same  instruments  as  before.  Three  sets  of  curves  are  given,  for 
OJ2  f*f.  (microfarad),  0.5  f*f.,  and  00  ftf.  (condenser  short-circuited), 
respectively.  Referring  to  the  pitch  lines,  it  will  be  seen  that  there 
is  not  much  difference  between  the  cases  of  00  and  0.5  /if.  The 
ascending  branches  of  the  zig-zags  cut  the  mean  frequency  line  of 
G"*  at  90,  132.5,  175  and  210  cm.  pr  fairly  in  conformity  with  the 
series  10  -|-  40W,  as  in  curve  //  of  Fig.  2.  With  0.2  ftf .,  however, 
the  intersections  with  this  line  are  at  100,  145  and  185  cm.,  or  more 
nearly  in  conformity  with  the  series  22  -|-  40m  cm. ;  that  is,  at  points 
displaced  about  12  cm.  further  along  the  tube.  Moreover,  the  breaks 
occur  at  higher  frequencies  by  about  40  ^. 

As  regards  primary  current  strengths,  the  minima  in  each  series 
occur  at  substantially  the  points  where  the  pitch  line  intersects  ascend- 
ingly  with  the  G"*  line.  That  is,  the  minima  of  00  and  0.5  f*f.  are 
fairly  close  together ;  while  those  for  0.2  ftf .  are  displaced  about  12 
cm.  further  along  the  tube.  Maximum  currents  occur  near  breaking 
points,  as  usual. 

Effects  of  Mechanical  Changes  in  Instruments. 

Observational  Series  8,  Effects  of  Modifying  the  Transmitter, 
— In  order  to  study  the  influence  of  changes  in  the  transmitter  upon 
the  humming  note,  three  similar  Western  Electric  transmitters  were 
selected,  of  standard  type  and  quality,  already  referred  to  as  T^  T^ 
and  Til,  in  connection  with  Fig.  7.  The  receiver,  induction-coil, 
battery  and  connections  were  as  in  previous  tests.  The  comparative 
results  with  these  three  transmitters  are  shown  in  Fig.  10,  for  tube- 
lengths  steadily  reduced  from  260  to  70  cm.  It  will  be  noted  that 
the  ascending  intersections  of  the  pitch  lines  all  intersect  the  mean- 
frequency  line  of  825  ^  in  substantial  conformity  with  the  series 
30  +  40W,  or  in  accordance  with  curve  /  of  Fig.  2.  The  breaking 
points  do  not  agree,  No.  8  always  breaking  last  at  a  higher  pitch. 
No.  5  next  at  a  medium  pitch  and  No.  11  first  at  a  lower  pitch.  It 
may  also  be  noted  that  in  the  hum-extinguishing  resistance-test  of 
these  three  transmitters,  as  given  in  Table  II.,  and  in  Fig.  7,  their 
order  of  succession  was  the  same. 


X908.], 


KENNELLY   AND  UPSON— HUMMING  TELEPHONE. 


347 


3 


Fig.  10. 


ii0  m  i40  490  999 

TUBE  LENGTH  -  KNTIMETERS. 

Comparative  Behavior  of  Three  Regular   Standard  Transmitters 

with  Reduced  Tube-lengths. 


The  test  indicates,  therefore,  that  different  standard  transmitters 
in  normal  adjustment  do  not  alter  the  mean-frequency  tube-lengths ; 
but  that  variations  in  breaking  lengths  may  be  expected  within  cer- 
tain limits. 

A  further  test  was  made  of  the  effect  of  modifying  the  trans- 
mitter, by  selecting  for  experiment  a  particular  Western  Electric 
Co.'s  standard  type  of  transmitter  which  had  been  used  in  the  labora- 
tory for  some  years,  and  was  not  in  the  best  adjustment.  A  test 
was  made  with  this  instrument  (using  the  same  receiver,  coil,  battery 
and  connections  as  in  preceding  tests),  first  without  any  extra  load 
on  its  diaphragm,  second  with  a  load,  and  third  with  the  load  re- 
moved.  The  load  consisted  of  a  small  brass  disk  1.5  cm.  (0.59  in) 
in  diameter,  and  0.2 'cm.  (0.079  "*•)  thick,  clamped  at  its  center 
between  the  two  small  nuts  at  the  center  of  the  external  surface  of 
the  diaphragm.  This  added  a  mass  of  2.7  gm.  to  the  vibrating 
system  of  the  transmitter.  The  results  are  seen  in  Fig.  11.  Curves 
I  and  3  represent  the  behavior  of  the  system  unloaded,  before  and 
after  loading  respectively,  the  tube-length  being  steadily  diminished 


348 


KENNELLY   AND  UPSON— HUMMING   TELEPHONE.        IJ»ly«>, 


from  200  to  80  cm.    Curves  2  represent  the  corresponding  behavior 
when  the  diaphragm  was  loaded.    The  primary  currents  were  all 
unusually  large,  probably  owing  to  the  imperfect  adjustment  of  the. 
transmitter. 


:/ 


t^O  ^ 


mo  _ 


in 


80        m        m        IH4        m        m 

TUBE  LENGTH  •  CENTIMETERS. 

Fig.  II.    Test  of  a  Transmitter  with  its  Diaphragm  Loaded  and  Unloaded. 

It  will  be  observed  that  the  loading  did  not  appreciably  alter  the 
ascending  intersections  of  the  pitch  lines  with  the  G"^  mean- 
frequency  line,  which  occur  in  conformity  with  the  series  20  +4om 
cm.  The  loading  seems  to  have  somewhat  lowered  the  range  of 
pitch  as  a  whole;  or  to  have  modified  the  conditions  at  breaking, 
without  materially  affecting  the  conditions  at  mean-frequency 
(825--). 

A  number  of  trials  with  further  modifications  of  the  transmitter 
diaphragm  substantiated  the  above  stated  results.  In  one  case,  a 
new  experimental  diaphragm  of  tinned  sheet  iron,  0.38  mm.  thick 


I908.] 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE. 


349 


(0.015  in.),  with  parallel  and  opposite  symmetrical  sectors  sliced 
off,  was  substituted  for  the  regular  diaphragm  in  the  test  transmit- 
ter. The  primary  current  strength  during  activity  was  thereby  in- 
creased ;  but  the  G"*  tube-lengths  remained  substantially  unchanged 
at  30  +  40m  cm.  Adding  loads,  altering  the  damping-spring  pres- 
sure, or  varying  the  other  mechanical  adjustments  of  the  transmitter 
produced  either  complete  silence;  or  else  the  usual  G"*,  at  30  + 
40W  cnf . 

The  tests  showed  that  modifying  the  transmitter  alters  the  range 
and  limits  of  pitch  variation,  as  well  as  the  primary  current 
strengths;  but  does  not  sensibly  alter  the  tube-leng^s  for  mean- 
frequency. 


90 


Fig.  12. 


/MS  M/0  Mt  /t0  ifi9 

TUBE  LENGTH  •  CENTIMETERS. 

Comparative  Frequencies  and  Currents  with  Three  Different 

Receiver  Diaphragms. 


Observation  Series  p.  Effect  of  Altering  the  Receiver, — In 
order  to  determine  the  influence  of  the  telephone  receiver  diaphragm 
on  the  hum,  three  special  receiver  diaphragms  were  made  up,  ea^ 
of  soft  transformer  steel,  0.355  ^^'  (0.014  in.)  thick,  and  5.5  cm. 
(2.16  in.)  in  diameter,  labeled  D^^  D^  and  D^  respectively.    D^  was 

PROC.  AMER.  PHIL.  SOC,  XLVII.  1 89  W,  PRINTED  OCTOBER  2,  I908. 


350  KENNELLY   AND  UPSON -HUMMING  TELEPHONE.         [Julyao, 

left  circular,  D^  and  D^  had  symmetrical  sectors  cut  from  opposite 
sides,  reducing  their  width  to  4  cm.  (1.57  in.)  and  3  cm.  (1.18  in.) 
respectively.  In  clamping  these  strip  diaphragms  in  front  of  the 
bipolar  magnet  of  the  standard  receiver,  their  angular  position  did 
not  appear  to  affect  the  system  appreciably. 

The  results  obtained  with  these  three  diaphragms  are  indicated 
in  Fig.  12,  for  tube-lengths  diminished  steadily  from  270  to  80  cm. 
It  will  be  seen  that  the  receiver  diaphragm  influences  the  hum  pro- 
foundly. Thus,  the  circular  diaphragm  D^^  developed  a  mean- 
frequency  of  1,100/^  or  c'"<,  judging  by  the  points  of  minimum 
primary  current,  and  its  pitch  zig-zag  formed  ascending  intersections 
with  this  line  at  95,  125,  155,  185,  215  and  245  cm.,  approximately, 
in  conformity  with  the  series  5  -|-  30m  cm.  The  sectored  diaphragm 
2?2  developed  a  mean- frequency  of  A"if,  at  920  ^,  with  ascending 
intersections  nearly  in  conformity  with  the  series  36W  cm.  The 
narrowest  diaphragm  D^  developed  a  mean- frequency  of  F",  at 
705  /-',  and  ascending  intersections  in  substantial  conformity  with 
the  series  33  -|-  47m  cm. 

It  will  be  observed  that  there  are  double  breaks  in  pitch  on  zig- 
zag jDi.  This  tendency  was  found  to  follow  irregularity  in  the  dia- 
phragm, or  in  its  mounting.  Thus,  the  ordinary  standard  diaphragm 
used  in  all  the  preceding  tests  was  observed  to  develop  similar  double 
breaks  when  the  clamping  screw-cover  was  slackened,  so  as  to  leave 
the  diaphragm  somewhat  loosely  clamped. 

The  pitch  zig-zag  of  D^  shows  gaps.  These  gaps  seemed  to  be 
due  to  the  enfeebled  condition  of  the  electromagnetic  vibrating  sys- 
tem in  the  receiver  when  used  with  the  experimental  diaphragm 
£>8-  A  very  marked  case  of  such  gaps  is  presented  in  Fig.  13,  which 
indicates  the  frequencies  and  currents  obtained  with  a  particular 
single-pole  telephone  receiver,  the  remainder  of  the  apparatus  being 
unchanged,  and  the  tube-length  being  steadily  reduced  from  165  to 
80  cm.  The  line  of  mean-frequency  is  at  1,025  /--',  and  the  ascend- 
ing intersections  with  this  line  are  formed  at  points  conforming 
with  the  series  28  +  32m  cm.  Only  short  pieces  of  the  zig-zag 
were,  however,  obtainable,  and  these  only  with  the  aid  of  a  condenser 
in  the  secondary  circuit.  The  dotted  segments  RS  and  TV  were 
obtained  with  the  receiver  terminals  reversed,  and  correspond  ap- 


i9o8] 


KENNELLY   AND  UPSON— HUMMING  TELEPHONE. 


361 


proximately  to  ascending  intersections  of  the  series  13  +  32m  cm. 

Various  other  modifications  of  receiver  and  receiver  diaphragm 
were  tried.  Loading  the  diaphragm  with  a  small  central  mass  low- 
ered the  mean  humming  frequency.  By  selecting  suitable  dia- 
phragm dimensions,  the  mean-frequency  of  the  hum  could  be  varied 
between  wide  limits. 


"±1/959 


9^ 


u. 


\ 


6 


{2 
8S 


tM 


uj  in 


J \ L L 


80 


90 


;^        m         S9$         m  100         foa 

^      TUBE  LENGTH  •  CENTIMETERS. 
Fig.  13.    Discontinuous  Frequencies,  or  Large  Gaps  in  Curves,  for  Case  of 

Singe-pole  Receiver. 

Conclusions  Directly  Derivable  from  the  Experiments. — ^The  fol- 
lowing more  prominent  conclusions  are  indicated  by  the  experiments 
themselves,  independently  of  any  theory : 

1.  The  mean-frequency  of  the  humming-telephone  note  is  deter- 
mined solely  by  the  receiver  diaphragm,  and  its  natural  free  rate 
of  vibration. 

2.  The  ascending  intersections  of  the  frequency  zig-zag  with  the 
mean-frequency  line  will  be  formed  approximately  at  tube-lengths 
of  (i  +  m)  v/n^  cm.  for  one  connection,  and  of  (J  +  m)  v/n^  cm. 
for  the  other  connection,  of  the  receiver;  where  v  is  the  velocity 
of  sound  in  air  (33,cxx)  cm.  per  sec.  nearly),  n^  is  the  mean  fre- 
quency in  cycles  per  second,  and  m  is  any  positive  integer,  within 


352  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.        [July*,. 

the  working  range  of  the  tube.    The  constants  f  and  i  may  be 
modified  by  the  presence  of  condensers,  and  other  circumstances. 

3.  The  range  of  pitch  variation,  and  the  breaking  positions,  are 
determined  by  the  transmitter,  and  by  the  reinforcing  capability  of 
the  system.  For  systems  that  are  weak,  either  electrically  or  acous- 
tically, the  range  of  pitch,  above  or  below  the  mean,  will  be  small. 

4.  The  primary  current,  as  measured  by  a  d.c.  instrument,  is 
ordinarily  a  minimum  at  the  mean  frequency,  and  a  maximum  at 
a  break. 

5.  Transmitters  may  be  tested  for  effectiveness,  by  measunng 
their  hum-extinguishing  resistances  in  the  primary  or  secondary 
circuit.  The  tube-length  should  be  such  as  to  produce  mean  fre- 
quency if  one  connection  of  receiver  only  is  used,  but  should  favor 
both  connections  equally,  if  both  connections  of  receiver  are  used. 

Outline  of  Theory  of  the  Humming  Telephone. 

Preliminary  Considerations.  Simple  Orbital  Motion  and  Simple 
Unretarded  Vibration. — Let  a  particle  of  mass  m  grammes  de- 
scribe a  simple  plane  circular  orbit  zab.  Fig.  14,  about  the  center 


b 

Fig.  14.    Vector  Diagram  of  Free  Undamped  Vibration. 

O.  Let  the  radius  Oz  =  r  cm.,  and  let  OX  be  the  initial  line  of 
reference.  At  time  ^  =  0  seconds,  let  the  particle  occupy  the  posi- 
tion 2 ;  so  that  its  initial  radius  vector  is  Os.  Let  w  be  the  uniform 
angular  velocity  of  the  particle  about  the  center  O,  in  radians  per 
second.  Then,  after  the  lapse  of  t  seconds,  the  particle  will  occupy 
a  point  in  the  plane  defined  by  the  vector  displacement 


,9o8.]  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.  363 

f  =  r€'w«  cms.   Z       (l) 


where  ;'=  V —  i>  2ind  f  is  the  displacement  of  the  particle  in  cms. 
from  O  at  the  angle  wf,  measured  positively,  or  counter-clockwise, 
from  the  initial  line  OX, 

Let  the  particle  be  acted  upon  by  a  centrally  directed  elastic  force 

F=  — Ai  =  —  ma(  =  —  marf,^^*  dynes   Z      (2) 

proportional  to  and  opposing  the  displacement,  as  represented  by 
the  vector  OF  in  Fig.  14.  Let  there  be  no  other  forces  except 
those  of  inertia,  acting  on  the  particle;  so  that  the  movement  is 
f  rictionless.    Then  the  velocity  of  the  particle  at  any  instant  t  will  be 

V  =  $  =^  jutrt^^*  cms./sec.   Z     (3) 

The  direction  of  the  velocity  will,  therefore,  be  perpendicular  to  the 
radius  vector,  or  parallel  to  the  instantaneous  tangent,  as  indicated 
by  the  dotted  line  Ov,  90°  ahead  of  Os  in  phase  displacement 
The  acceleration  of  the  particle  will  be,  at  any  instant  t, 

c  =  v  =  $  =  —  wVe'w*  cms./sec'  Z     (4) 

That  is,  the  acceleration  will  be  directed  oppositely  to  the  displace- 
ment.. Thus  at  time  /  =  o,  represented  in  Fig.  14,  the  acceleration 
will  be  directed  along  OF.  The  virtual  reactive  force  of  inertia 
will  be 

/  =  —  mc  =  —  m^  =  w<tt*rc'«*  dynes  Z      (5) 

In  Fig.  14,  this  reactive  force  of  inertia  is  represented  by  Of. 

In  order  that  the  circular  orbital  motion  shall  be  stable,  the 
sum  of  the  forces  OF  and  O/,  of  elasticity  and  inertia  must  be  zero ; 
or 

OF  +  Of  =  o  dynes  Z 

.*.    —  mart^*^*  +  mwVc'w*  =  o  dynes  Z 

whence 


ft,  =  yJA/m  =  y/a  radians/sec.     (6) 


354  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         [J".y  «>, 

If,  therefore,  the  angular  velocity  of  the  motion  be  numerically 
equal  to  the  square  root  of  elastic  force  per  unit  of  mass,  the 
orbit  will  be  circular  and  stable,  and  Fig.  14  may  represent  its 
vector  diagram.  The  particle  z  rotates  about  O,  at  constant  radius 
with  uniform  angular  velocity  w,  and  the  pair  of  equilibrating 
forces  OF  and  Of  rotate  in  synchronism  with  it.  The  entire 
system,  Fig  14,  may  be  imagined  as  pivoted  about  an  axis  through 
O  perpendicular  to  the  orbital  plane,  and  spun  about  this  pivot  with 
uniform  angular  velocity  w. 

By  a  well  known  proposition  connecting  simple  harmonic  vibra- 
tion with  circular  orbital  motion,  the  displacements  in  the  former 
are  the  projections  of  the  displacements  in  the  latter,  upon  a  straight 
line  passing  through  the  center  of  the  system.  In  other  words,  to 
every  case  of  simple  circular  orbital  motion  in  two  dimensions 
corresponds  a  case  of  simple  harmonic  vibration,  its  projection  in 
a  single  dimension.  Consequently,  at  time  t,  we  have  for  the  dis- 
placement in  the  case  of  simple  vibration, 

^  =  rc'w«  cms.     (7) 

measured  along  the  initial  line  OX  by  projection.  The  real  part 
only  of  ^  is  retained,  and  the  imaginary  part  ignored.  Similarly, 
the  vibratory  velocity  will  be 


y 


v  =  ^^=  /wrc'w*  cms./sec.     (8) 

taking  only  the  real  part  of  the  equation,  or  the  projected  value 
along  YOX.    Again,  the  vibratory  acceleration  will  be 

c= — «V€^«*  cms./sec*     (9) 

retaining  only  the  real  or  projected  part.  Similar  reasoning  ap- 
plies to  the  forces  of  elasticity  and  inertia.  The  same  equations 
appear  as  in  the  circular  orbit  case ;  but  only  their  real,  or  horizon- 
tally projected  values,  are  retained.  Consequently,  we  deduce  that 
the  vibration  of  a  particle  possessing  elasticity  and  inertia  without 
frictional  retardation  will  be  stable  and  self  sustained  under  the 
condition 


X908.J  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.  366 


0) 


=  2wn  =  yj  A/m  =  y/a  radians/sec.     ( lo) 


where  n  is  the  frequency  of  the  vibration  in  cycles  per  second. 

If,  for  example,  the  diaphragm  of  a  telephone  receiver  had 
simple  elasticity  and  inertia  without  f  rictional  retardation,  such  that 
the  elastic  intensity  a  =  26.87  X  10*  d)mes  per  cm.  of  displace- 
ment and  per  gramme  mass,  then  any  displacement  released  would 
be  followed  by  an  indefinitely  sustained  angular  velocity 


0) 


=  V26.87  X  io«  =  5,184 


radians  per  second,  corresponding  to  n  =  825  cycles  per  second.  If 
the  initial  displacement  were  r  =  o.oi  cm.,  the  corresponding  simple 
circular  orbit.  Fig.  14,  would  have  a  radius  of  o.oi  cm.,  an  angular 
velocity  of  5,184  radians  per  second,  an  orbital  velocity  of  51.84 
cm.  per  second,  and  an  acceleration  of  268,700  cm.  per  second.  If 
the  elastic  force  A  were  1.3435  X  10*  dynes  per  cm.  of  displace- 
ment and  the  effective  mass  were  0.05  gm.,  the  elastic  force  OF 
would  be  13,435  dynes,  and  the  centrifugal  force  Of  13,435  dynes, 
the  two  being  equal  and  in  complete  opposition. 

Case  of  Free  Vibration  Damped  and  Unreinforced.  Spiral 
Orbital  Motion. — In  the  case  of  the  particle  moving  about  a  center, 
let  the  motion  be  retarded  by  a  force  f,  proportional  to  the  velocity, 
defined  by  the  relation 

f  =  —  Tv  =  —  2myv  dynes  Z     (11) 

Then  the  orbital  displacement  at  any  time  t  becomes 

^  =  r€<-^*^«>*  cms.  I     (12) 

The  orbital  velocity  is 

v  =  i  =  r( — y +  /a))€^-7+i«>*  cms./sec.  Z      (13) 

The  orbital  acceleration  is 

c  =  z/='^  =  r(— y  +  ya))V-7+iw)*        cms./sec.*  Z      (14) 


356  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         [Julyao, 

Each  of  the  above  equations  defines  an  equiangular  spiral,  an  in- 
wardly directed  spiral  in  which  the  curve  mak^  a  constant  direction 
—  y  +  y<«>  with  the  radius  vector. 

The  vector  diagram  for  this  case  is  indicated  in  Fig.  15.    Let 


T 

Fig.  15.    Vector  Diagram  of  Free  Damped  Vibration. 

z  be  the  position  of  the  particle  at  any  instant.    The  velocity  at  this 
instant  will  have  the  vector  OV,  parallel  to  the  tangent  at  z,  where 

tan  <^  =  o)/y  (15) 

The  acceleration  at  the  same  instant  will  be  directed  along  OY, 
the  angles  XOV  and  VOY  being  each  equal  to  the  supplement  of  ^. 
The  virtual  force  of  inertia  will  be  directed  along  Of,  The  retard- 
ing force,  opposing  the  velocity,  will  be  directed  along  Of.  At  any 
instant  the  vector  sum  of  the  three  forces  of  elasticity,  retardation 
and  inertia  must  be  zero.    That  is, 

OF  +  Of +  0/  =  o  dynes   Z 

or 

—  fnar€^-y*^^^*  —  2ywr( —  y  +  ;o>)€^"7+i«>* 
—  mr(— y +  ya))V-7+i«)*  =  o         dynes  Z 
whence 


0) 


=  ^/a  —  y^  =  V<«»o*  —  y'  =  «o  sin  ^      radians/sec.     (16) 


where  wo  is  the  unretarded  angular  velocity.  That  is,  the  angular 
velocity  of  orbital  rotation  has  been  reduced  by  the  retardation  in 
the  ratio  of  sin  ^,  Fig.  15,  and  the  displacement  or  radius  vector  r 
continually  dwindles  with  time  by  €"7*. 

In  the  corresponding  case  of  free  damped  vibration,  the  above 


,9o8.|  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.  867 

equations  apply;  but  their  real  parts  only  are  taken.  In  Fig.  15,  the 
projections  of  the  vectors  on  a  straight  line  through  O,  are  se- 
lected.  The  dwindling  vibrations  of  a  tuning  fork,  or  the  oscillatory 
discharge  of  a«condenser  through  a  circuit  containing  resistance  and 
inductance,  obey  this  law.  In  the  last  named  case,  the  inductance 
corresponds  to  the  mass  m,  the  reciprocal  of  the  capacity  corresponds 
to  the  elastic  coeiBcient  A,  and  the  resistance  corresponds  to  the 
velocity-resisting  coefficient  r.     The  .condenser-charge,  or  electric  / 

quantity,  corresponds  to  the  vibratory  displacement,  the  electric 
current  to  the  vibratory  velocity,  the  discharging  electromotive  force 
to  the  elastic  force  OF,  the  resistance  e.m.f.  to  O/',  the  e.m.f.  of 
self-induction  to  Of,  and  the  impedance  of  the  discharging  circuit 
to  the  vector  nua^  L  ^,  or  yJMA   Z  ^  =  r/2  -|-  /mw. 

Case  of  Retarded  Free  Vibration  Reinforced,  Restored  Circu- 
lar Orbit, — In  order  to  sustain  stable  orbital  motion  in  a  particle 
retarded  with  a  force  proportional  to  the  velocity,  it  is  necessary 


Fig.  16.    Vector  Diagram  of  Reinforced  Vibration. 

to  supply  energy  continuously  to  the  particle  and  to  act  upon  it  with 
a  force  equal  but  opposite  to  the  velocity-resisting  force.  The  orbit 
will  then  be  restored  from  an  inmoving  spiral  to  a  simple  circle. 
The  displacement,  velocity  and  acceleration  of  the  particle.  Fig.  16, 
will  then  be  severally  expressed  by  equations  (i),  (2)  and  (3)  ap- 
plied to  Fig.  14. 

Let  OR,  Fig.  1-6,  be  an  outwardly  directed  force  from  the  center 
O,  the  magnitude  of  OR  being  some  function  mx(r)  of  the  radius 
of  displacement,  and  0  the  phase  retardation  behind  the  displace- 


368  KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         [July«o, 

ment,  reckoned  positively  in  the  direction  indicated,  or  clockwise. 
The  restoring  force  OR  Z.B  may  be  analysed  into  two  components 
OT=zOR  sin  6,  and  OS=OR  cos  0  along  the  directions  OV  and 
OX  respectively.  The  component  OT  may  be  calkd  the  velocity 
component,  or  T  component,  since  it  acts  in  the  direction  of  the 
velocity  Ov,  and  against  the  retarding  force  Of.  The  component 
Os  may  be  called  the  5"'  component,  or  the  new  elastic  component. 
It  coacts  with  the  elastic  force  OF  that  resists  displacement. 

In  order  that  the  circular  orbit  may  be  retained,  it  is  necessary 
and  sufficient  that  the  T  component  of  the  restoring  force  shall 
equilibrate  the  velocity-resisting  force  Of;  or,  if  i?  be  the  restor- 
ing force,  that 

R  sin  $  +  f  =  o   ^   dynes 

If  the  T  component  should  be  less  than  the  velocity-resisting  force, 
the  system  will  lose  energy.  The  orbit  will  spiral  inwards  until  the 
velocity  has  been  sufficiently  diminished  to  equilibrate  the  T  com- 
ponent, and  permit  a  stable  circular  orbit  of  reduced  radius  to  be  re- 
stored. If,  on  the  contrary,  the  T  component  exceeds  the  velocity- 
resisting  force,  the  system  will  accumulate  energy,  and  the  orbit  will 
spiral  outwards  until  the  radius  and  velocity  of  the  motion  are  suffi- 
cient to  restore  equilibrium  and  permit  a  circular  orbit  of  enlarged 
radius  to  be  maintained. 

In  the  condition  of  equilibrium  represented  in  Fig.  i6,  we  have 
four  forces  acting  on  the  particle,  forming  two  separate  equili- 
brating pairs;  namely,  a  pair  along  the  displacement  vector  Os, 
which  we  may  call  the  displacement  pair,  and  a  pair  perpendicular 
thereto,  which  we  may  call  the  velocity  pair.  Both  these  pairs  rotate 
together  at  some  uniform  angular  velocity  w,  which  will  in  general 
diifer  from  that  which  would  hold  for  unretarded  motion  wo,  as  in 
Fig.  14,  or  from  that  which  would  hold  for  retarded  unrein  forced 
motion,  as  in  Fig.  15. 

Considering  the  displacement  pair,  the  first  member  is  the  elastic 
force  OF,  modified  by  the  new  elastic  force  05"^  to  OF',  Fig.  16. 
The  new  virtual  force  of  inertia  is  Of.     Consequently 

OF'  +  0/  =  o    dynes 


«9o8.]  KENNELLY   AND  UPSON— HUMMING   TELEPHONE.  359 

or 

—  mar€^^*  -\-fnx(r)  cosO  €^«*  +  wuoVe^^*  =  o        dynes  Z 

whence 


0)  =  Va — cos6  •  X (^)  A  radians/sec.     (17) 

Considering  the  velocity  pair,  the  first  member  is 

Of  =  —  Tv  =  —  2myt;  =  —  J2myu>r€^^*  dynes  Z 

The  second  member  is  the  T  component : 

0T== — ywx(r)  sin^-c'«*  dynes   Z' 

For  equilibrium 

Of-\-OT  =  o 
or 

—  ;2Wywrc^«*  —  jfnx(r)  sinS  •  c'«*  =  o 
From  which 

x(r)  sin  0  =  —  2rya)        dynes  (18) 

and 


(0 


=  V«o^  +  y^  cot^  ^  +  y  cot  d  radians/sec.     (19) 


It  follows  that  0),  the  new  angular  velocity  under  reinforcement, 
is  independent  of  the  force  function  R  =  mx(r),  and  depends  only 
on  the  natural  angular  velocity  wo,  the  phase  retardation  $  of  the  re- 
storing force  and  the  magnitude  of  the  damping  coefficient  y.  Some 
curves  of  w  as  a  function  of  6  for  four  particular  values  of  T  be- 
tween 50  and  500  dynes  per  cm.  per  sec. ;  i.  e.,  of  y  between  500  and 
5,000  dynes  per  cm.  per  sec.  and  per  g^m.,  are  given  in  Fig.  17.  It 
may  be  seen  that  for  all  values  of  the  damping,  a)  =  c.)o  for  ^  =  270**. 
That  is,  the  angular  velocity  of  reinforced  motion  is  the  same  as 
that  of  unretarded  motion  when  the  restoring  force  is  applied  at  270** 
of  phase  lag,  or  exactly  in  phase  with  the  velocity,  as  seen  in  Fig. 
16.  If  the  phase  retardation  ^  is  between  180®  and  270**,  the  new 
angular  velocity  will  be  greater  than  the  natural  angular  velocity  «o ; 
but  if  0  is  between  270®  and  360®,  must  be  less  than  «o. 

Applying  the  above  principle  to  the  corresponding  case  of  rein- 
forced vibration,  by  taking  the  projections  or  real  parts  of  the  rotat- 


360 


KENNELLY  AND  UPSON— HUMMING  TELEPHONE.         U^\y  ao. 


ing  vectors,  it  follows  that  if  any  automatically  reinforced  vibrating 
system,  such  as  an  electromagnetic  bell,  electromagnetic  tuning  fork, 
or  humming  telephone,  is  propelled  by  an  elastic  force  proportional 
to  the  displacement,  reinforced  by  a  cyclic  force  some  function  of  the 
displacement,  and  damped  by  a  force  proportional  to  the  velocity,  it 
is  subject  to  equations  (17),  (18)  and  (19)  which  appear  to  be  new. 
In  the  series  of  measurements  on  the  humming  telephone  above 


im 


Fig.  17.    Reinforced  Frequency  in  Relation  to  the  Phase  of  the  Reinforcement. 

outlined,  the  force  function  /?  =  wx(r)  was  not  measured.  The 
restoring  electromagnetic  force  on  the  receiver  diaphragm,  due  to  the 
action  of  the  transmitter,  will  manifestly  diminish  when  the  tube- 
length  is  increased.  For  a  fixed  tube-length,  moreover,  it  cannot 
increase  indefinitely  in  simple  proportion  to  the  displacement  of  the 
diaphragm,  or  to  its  amplitude  of  vibration.  If  we  assume  pro- 
visionally that  R  increases  as  the  square  root  of  the  amplitude  of 


,9o«.]  KENNELLY  AND  UPSON—HUMMING  TELEPHONE.  361 

receiver-diaphragm  vibration;  so  that  for  a  fixed  tube-length, 
R  =  bm^/r;  or  ;((r)=rr6yr;  where  b  is  the  numerical  constant 
1.036  X  io*  dynes  per  gm.  and  per  Vcm. ;  then  for  the  example 
already  considered,  if  r=ioo;  or  y  =  1,000,  we  find: 

The  displacement  r  =  o.oi  cm.  when  ^  =  270**. 

The  reinforced  angular  velocity  is  w  =  5,184  radians  per  second ; 
n  =  825  r^. 

The  maximum  cyclic  values  of  the  vibratory — 

velocity  z/=        51.84  cm.  per  sec. 

acceleration  c  =  268,74(j  cm.  per  sec* 

damping  force  Of=     5,184  dynes. 

restoring  force  0T=     5,184  dynes. 

elastic  force  0F'=    13435  dynes. 

inertia  force  Of=   13,435  dynes. 

As  the  phase  $  of  reinforcement  changes  from  180**  to  360®,  the  line 
of  y  =  1,000  in  Fig.  17  shows  the  change  in  frequency;  while  the 
dotted  line  indicates  the  computed  amplitude  of  vibration,  which 
reaches  a  maximum  near  280**. 

According  to  the  theory,  therefore,  if  the  phase  of  the  displace- 
ment is  270®  behind  the  displacement  of  the  receiver  diaphragm,  the 
reinforced  frequency  coincides  with  the  natural  frequency.  This 
condition  is  substantially  borne  out  in  all  of  the  observations.  For 
example,  in  Fig.'  2,  taking  the  pitch  line  No.  i,  with  a  natural  fre- 
quency of  Ho  =  825  r^  and  a  sound-velocity  in  air  of  33,000  cm.  per 
sec,  the  wave-length  A.  =  33,000/825  =  40  cm.  corresponding  to 
360*^  of  phase.  A  lag  of  270**  would  be  represented  by  30  cm. ;  so 
that  we  should  expect  the  reinforced  frequency  to  be  825  ^  at  30 
cm.  of  tube-length,  and  at  every  40  cm.  beyond ; «.  e.,  in  accordance 
with  the  series  30  +  40m,  as  was  substantially  observed.  Moreover, 
by  reversing  the  receiver  terminals,  the  phase  of  the  reinforcement 
is  necessarily  changed  180** ;  so  that  with  this  change  of  connec- 
tion, 270**  of  phase  lag  would  be  altered  to  90**  of  phase  lag,  or  10 
cm.  of  tube-length.  The  natural  frequency  of  825  r^  should  then 
occur  in  conformity  with  the  series  10-I-40W  cm.,  as  was  substan- 
tiallv  observed. 


362  KENNELLY   AND  UPSON— HUMMING  TELEPHONE.         [July  20, 

When  the  phase  retardation  $  of  the  restoring  force  (Fig.  16) 
is  less  than  270®,  we  should  expect,  according  to  the  theory,  that  the 
pitch  should  rise ;  because  the  elastic  resilience  of  the  diaphragm  is 
virtually  increased  by  the  OS  component  of  the  new  forte,  and  when 
$  >  270®,  on  the  contrary,  the  pitch  should  fall.  This  was  always  the 
case  in  the  observations.  We  have  to  bear  in  mind,  however,  that 
with  any  given  tube-length,  an  alteration  of  pitch  involves  a  change 
of  wave-length,  and  therefore  a  change  of  phase  in  transmission 
through  the  air-column,  besides  any  electrical  change  in  phase  due 
to  change  in  current  frequency.  In  Figs.  3,  4  and  6,  the  sloping 
dotted  lines  are  drawn  to  indicate  constant  acoustic  phase  retardation 
of  270®  for  all  of  the  frequencies  within  the  range  considered. 
Taking,  for  instance,  Fig.  6,  the  break  at  P  occurred  103*^  in  phase 
from  the  dotted  line  of  270**,  and  the  return  at  S  occurred  77°  in 
phase  from  the  dotted  line.  According  to  the  theory,  assuming  no 
electric  change  of  phase,  each  of  these  angles  should  be  something 
less  than  90**,  since  the  phase  retardation  must  be  something  more 
than  iSo*'  on  the  side  of  increasing  pitch.  The  discrepancy  here  is 
not  serious;  for  the  mean  of  the  two  angles  is  90®.  At  T  and  IV, 
however,  the  corresponding  angles  are  153**  and  119°,  with  a  mean 
of  136®,  which  should  be  something  less  than  90**,  a  greater  diver- 
gence from  the  theory  than  observation  errors  can  explain.  While, 
therefore,  the  theory  accounts  for  all  of  the  experimental  results  in 
a  general  way,  it  can  only  be  regarded  as  a  first  approximation.  For 
example,  it  is  possible  that  superposed  harmonic  currents  might  have 
to  be  considered ;  or  that  in  estimating  the  damping  forces,  the  in- 
clusion of  higher  powers  of  the  velocity  than  the  first  might  be 
necessary. 

Setting  aside  unexplained  deviations,  as  the  tube-length  is  short- 
ened from  a  point  of  ^  =  270°,  the  phase  retardation  of  the  electro- 
magnetic reinforcement  on  the  receiver  diaphragm  is  diminished. 
This  causes  the  pitch  to  rise,  and  incidentally  readjusts  the  phase 
change  to  a  lower  value  than  if  the  pitch  were  kept  steady.  The 
amplitude  of  vibration  diminishes  until  the  diaphragm  suddenly 
selects  a  lower  pitch  for  the  same  tube-length,  to  which  the  ampli- 
tude will  be  greater.  In  other  words,  the  receiver  diaphragm  auto- 
matically seeks  to  maintain  the  greatest  amplitude  that  the  condi- 


2908.] 


KENNELLY   AND  UPSON— HUMMING  TELEPHONE. 


363 


tions  of  reinforcement  will  permit.  If  a  lower  tone,  with  a  pha^e 
lag  $  more  than  270°,  will  give  more  amplitude  than  the  higher  note 
to  which  it  has  been  driven,  with  $  less  than  270**,  it  will  break 
pitch  downwards.  This  process  will  continue  down  to  the  first 
wave-length  of  tube,  or  40  cm.  in  the  case  examined.  For  connec- 
tion /  of  the  receiver,  it  can  break  to  no  lower  note  after  passing 
270°,  and  the  tone  will  rise  to  such  a  pitch  that  the  amplitude  be- 
comes insufficient  to  excite  the  transmitter,  so  that  silence  should 
ensue  at  or  near  the  length  12  cm.,  as  actually  observed  in  Fig.  3. 
The  curve  /  of  frequency  between  12  and  50  cm.  accords  fairly  well 
with  the  curve  y  =  1,000  in  Fig.  17. 


^O'tf 


Fig.  18. 
Fig.  19. 


E  -= 


Fig.  18.  Fig.  19. 

Diagram  of  Electrical  Connecfions  with  Step-up  Induction  Coil. 
Equivalent  Diagram  of  Connections,  with  Level   Induction  Coil. 


JmUt 


Fig.  20.  Fig.  21. 

Figs.  20  and  21.    Equivalent  Conductive  Connections  with  Level  Induction 

Coil  and  Alternating  E.M.F. 

With  reference  to  the  influence  of  capacity  in  the  secondary  cir- 
cuit, Figs.  18  to  21  show  the  successive  steps  by  which  the  secondary 
circuit  may  be  treated  as  a  conductive  branch  of  the  primary  circuit. 
Using  the  constants  given  in  Table  I.,  ignoring  any  capacity  exist- 
ing between  the  windings  of  the  coils,  and  assimiing  that  the  eifect 
of  the  transmitter  in  the  primary  circuit  is  equivalent  to  an  alter- 
nating e.m.f.  e,  working  through  a  transmitter  resistance  R,  of  50 
ohms,  we  find  a  coupling  coefficient  for  the  coil  of  if  =  0.937,  ^^d 


364 


KENNELLY  AND  UPSON—HUMMING  TELEPHONE.         [Julyao, 


an  inductance  ratio  5"  =  0.0575  between  primary  and  secondary 
windings.  Proceeding  in  this  way,  the  following  table  has  been 
arrived  at,  giving  the  conductances  which  when  multiplied  by  the 
equivalent  transmitter  e.m.f.  yield  the  current  strength  in  the  sec- 
ondary circuit  of  Fig.  21. 

Table  III. 


Frequency  Cycles  per 
Second,    n. 


637 
796 

956 

637 
796 
956 


Angular  Velocity 
Radians  per  Second. 


4,000 
5,000 
6,000 

4,000 

5,000 

•  6,000 


Capacity  in  Secondary 
Circuit,    id. 


oc 

oc 

0.2 
0.2 
0.2 


Conductance  of  Secon- 
dary.   Mhoa. 


0.009  723** 

0.0087  /17** 

0.0093  1^^^ 

0.0054  \I40^ 

0.0095  \i22l 

O.II  \liSl 


Although  the  assumptions  employed  do  not  anticipate  a  high 
degree  of  accuracy  in  the  conclusions  above  tabulated^  yet  we  may 
safely  infer  that  when  no  condenser  is  used  in  the  secondary  circuit 
(/if.  =  oc ) ,  the  secondary  current  will  lead  the  impressed  primary 
e.m.f.  by  a  small  angle,  and  this  current  will  have  substantially  the 
same  strength  and  phase  for  all  frequencies  between  600 /-^  and 
1,000  r^.  When,  however,  a  condenser  of  0.2  fii.  is  inserted  in  the 
secondary  circuit,  the  current  in  the  receiver  will  be  advanced  in 
phase  about  no®  or  nearly  a  third  of  a  cycle;  while  the  strength  of 
this  current  will  be  considerably  greater  at  the  highpr  frequencies 
than  at  the  lower  frequencies. 

Since  the  total  lag  in  phase  of  the  restoring  electromagnetic  force 
behind  the  displacement  of  the  receiver  diaphragm  includes  (i)  the 
electric  current  lag;  (2)  any  hysteretic  electromagnetic  lag  in  the 
receiver  cores;  (3)  any  mechanical  inertia  lag  of  the  transmitter 
diaphragfm;  (4)  the  acoustic  lag  in  the  air  column  of  the  tube;  it 
follows  that  the  total  lag  with  a  condenser  of  0.2  /if.  should  be  about 
1 10®  less  than  with  short-circuited  condenser ;  while  the  higher  fre- 
quency notes  should  be  favored,  and  the  lower  frequency  notes  dis- 
favored. Fig.  9  shows  that  both  these  effects  took  place,  the  acous- 
tic lag  had  to  be  increased  by  about  12  cm.,  or  about  no**,  in  order 
to  produce  mean  frequency,  and  compensate  for  the  current  lead. 


i9o8.]  KENNELLY   AND  UPSON— HUMMING  TELEPHONE. 


365 


Also  the  range  of  frequency  is  moved  bodily  towards  higher  notes. 

All  of  the  experimental  series  of  observations  appear  to  be  ac- 
counted for  and  explained  by  the  above  theory  to  a  first  approxima- 
tion ;  although  in  matters  of  quantitative  detail  there  remains  much 
room  for  further  development. 

In  conclusion,  the  authors  desire  to  express  their  mdebtedness 
to  the  Western  Electric  Co.  for  the  loan  of  apparatus  used  in  the 
tests. 


PROC.  AMER.  PHIL.  SOC,  XLVII.  189  X,  PRINTED  OCTOBER  3,  I908, 


ON  THE  AFTER-IMAGES   OF   SUBLIMINALLY 

COLORED  STIMULI. 

By  EDWARD  BRADFORD  TITCHENER  and  WILUAM  HENRY  PYLE. 

(Received  July  23,  igo8.) 

We  attempt,  in  the  present  paper,  to  answer  the  question  whether 
a  subliminally  colored  stimulus  may  arouse  a  colored,  negative  or 
complementary  after-image.  This  question  has  been  answered  in 
the  affirmative  both  for  direct  and  for  indirect  vision,  and  in  indirect 
vision  for  all  three  of  the  retinal  zones.  Our  own  experiments,  on 
the  other  hand,  have  led  us  to  answer  it  in  the  negative.  Provided 
that  the  subliminally  colored  stimulus  appears  on  a  neutral  (black, 
gray  or  white)  background,  and  provided  that  the  retina  is  achro- 
matically  adapted,  we  find  no  trace  of  the  colored  after-image  in 
either  direct  or  indirect  vision,  with  either  light  or  dark  adaptation. 

Previous  Experiments. 

^* 

I.  Direct  Vision. — In  a  paper  entitled  Das  Anpassungsproblem 
in  der  Physiologic  der  Gcgcnwart  (1904),  A.  Tschermak  compares 
the  course  of  excitation  in  the  retina  with  the  eifects  produced  by 
the  constant  current  in  a  nerve-muscle  preparation.  The  passage  is 
as  follows: 

"  Haben  wir  doch  gerade  in  der  Anwendung  des  constanten  Stromes  auf 
Nerv  und  Muskel  ein  vorziigliches  didaktisches  Mittel,  um  die  Grundbegriffe 
der  allgemeinen  Reiz-  und  Adaptationslehre  zu  veranschaulichen  und 
einzupragen.  Am  besten  demonstrieren  wir  als  Gegenstuck  zugleich  die 
Wirkung  eines  massig  satten  Farbglases  auf  das  Auge:  die  Phase  der 
Reizwirkung,  individuell  verschieden  lang,  und  dadurch  erinnernd  an  die 
verschiedenrasche  Adaptation  des  Praeparates  vom  Warmfrosch  und  Kalt- 
frosch  an  den  constanten  Strom  —  weiterhin  das  Stadium  der  vollendeten 
Adaptation,  endlich  den  gegensinnigen  Oeffnungseffect.  Nicht  minder 
lehrreich  ist  die  Parallele  des  subjectiven  und  des  objectiven  Erscheinungs- 
gebietes  fiir  das  Phaenomen  des  Einschleichens  d.  h.  des  Ausbleibens  einer 
sinnfalligen  Reizwirkung,  wenn  der  Reiz  so  langsam  anwachst,  dass  das 
Adaptationsvermogen  folgen  kann  —  gleichwohl  hat  auch  nunmehr  Wegfall 

366 


,908.]  OF  SUBLIMINALLY  COLORED   STIMULI.  367 

des  'Reizes*  cine  gegensinnige  Oeffnungswirkung.  Analoges  gilt  vom  Aus- 
schleichen,  also  vom  Ausbleiben  eines  sinnfalligen  Oeffnungseffectes.  Zum 
optischen  Versuche  schiebt  man  zweckmassig  eine  schwach  tingierte  Glas- 
platte  vor  die  andere  oder  benutzt  einen  Keil  farbigen  Glases."* 

The  observation  here  briefly  mentioned  was  apparently  made  in 
light-adaptation.  The  observer,  we  may  suppose,  looked  through  a 
vertical  slit  in  a  cardboard  screen  towards  a  window.  The  thin 
end  of  the  colored  glass  wedge,  viewed  through  the  slit,  appeared 
colorless.  The  wedge  itself  was  slowly  pushed  forward  —  so  slowly 
that  progressive  adaptation  prevented  its  color  from  being  per- 
ceived. Presently  the  observer  turned  his  eye  to  the  cardboard 
screen,  and  there  saw  the  negative  colored  after-image,  the  "  gegen- 
sinnige Oeffnungswirkung  "  that  followed  the  "  Einschleichen  des 
Reizes." « 

2.  Indirect  Vision. — In  the  Studies  from  the  Psychological 
Laboratory  of  Mount  Holyoke  College  for  1905,  Miss  G.  M.  Femald 
reports  the  arousal  of  colored  after-images  in  the  peripheral  or 
black-white  zone  01  the  retina.  "  A  further  point  worth  mention- 
ing " — so  the  passage  runs — "  is  the  fact  that,  in  the  case  of  several 
colors,  exposure,  beyond  the  limits  where  any  color  is  seen,  is  fol- 
lowed by  a  very  clear  [colored]  after-image.  This  was  repeatedly 
found  to  be  true  with  red,  orange,  green  and  blue  and  often  with 
yellow  [stimuli].  This  after-image  for  the  first  three  and  for  yel- 
low was  blue,  and  for  bltfe  a  very  clear  yellow.  This  may  explain 
the  '  gegenfarbige '  zone  found  by  Hellpach  in  his  dark-room  work, 
as  under  those  conditions  there  would  have  been  no  way  of  telling 
whether  the  color  came  exactly  at  the  time  of  exposure  or  inynedi- 
ately  afterwards." '     No  further  details  are  given. 

^Archives  des  sciences  biologiques,  XL,  Supplement  (Festschrift  for  Pro- 
fessor J.  P.  Pavloff),  82  f. 

*The  procedure  is  sketched  by  H.  Abels,  Zeits.  f.  Psychol,  XLV.,  IQ07, 
86.  "  Man  kann  .  .  .  einen  schwach  gefarbten  Glaskeil  so  langsam  vor 
das  Auge  schieben,  von  der  Kante  gegen  den  Rucken  fortschreitend,  dass 
uberhaupt  keine  Farbenempfindung  zustande  kommt;  und  dennoch  haben 
wir  bei  plotzlichem  Entfernen  desselben  und  Betrachten  einer  indifferent 
gefarbten  Flache  die  deutliche  Empfindung  der  komplementaren  Farbe." 
Abels  is  here  quoting  a  conversation  with  Tschermak;  there  is  no  evidence 
that  he  himself  performed  the  experiment. 

•"The  Effect  of  the  Brightness  of  Background  on  the  Extent  of  the 
Color  Fields  and  on  the  Color  Tone  in  Peripheral  Vision,"  Psychol.  Review, 
XIL,  November,  1905,  405. 


\ 


368  TITCHENER   AND   PYLE— ON   THE   AFTER-IMAGES        [July  93, 

These  observations  would,  no  doubt,  have  been  repeated,  and 
their  interpretation  discussed  by  other  experimenters,  had  not  Baird 
published,  earlier  in  the  same  year,  his  study  of  the  color  sensitivity 
of  the  peripheral  retina.  "  There  seems  to  be  no  doubt,"  Baird  had 
written,  "  that  Hellpach's  zone  of  complementariness  is  an  artifact, 
and  that  its  discovery  is  wholly  due  to  the  experimenter's  failure 
to  avoid  retinal  fatigue  [chromatic  adaptation]  in  his  explorations."  * 
Nevertheless,  one  of  the  present  writers  (T)  made  in  1906. a  fairly 
long  series  of  campimetrical  observations  (some  200  in  all)  with 
the  view  of  testing  Miss  Femald's  conclusion.  The  colored  stimuli 
were  Hering  papers,  R,  Y,  G  and  B ;  the  backgrounds  were  white, 
neutral  gray  and  black.  In  no  case  was  "  exposure,  beyond  the  lim- 
its where  any  color  is  seen,"  followed  by  a  colored  after-image,  clear 
or  obscure.  All  four  colors,  if  they  gave  an  after-image  at  all, 
gave  a  colorless  image,  indistinguishable  from  the  after-images  of 
gray  stimuli — ^as  these  gray  stimuli  themselves  were  indistinguish- 
able from  the  colored  papers.  It  therefore  seemed  probable — in- 
deed, it  seemed  practically  certain — ^that  the  Mount  Holyoke  results 
were  due  to  a  defect  of  method.  Since  Baird's  disproof  of  the 
"  gegenfarbige  Zone  "  was  deemed  complete  and  final,  the  Cornell 
observations  were  not  published. 

However,  in  the  following  year,  1907,  a  second  paper  from  the 
Mount  Holyoke  laboratory  reported  the.  same  phenomenon.  "  At 
the  extreme  periphery  it  sometimes  happened:  (a)  that  a  stimulus 
which  was  clearly  seen  produced  no  after-image.  ...  (6)  On  the 
other  hand  there  were  118  cases  in  which  a  subliminal  stimulus  pro- 
duced an  after-image  which  was  perfectly  distinct  in  color.  .  .  . 
That  this  somewhat  unusual  result  was  not  the  outcome  of  imag- 
ination or  suggestion  seems  proved  by  the  fact  that  these  invisible 
colors  gave  rise  to  their  appropriate  after-images."  *  The  authors, 
the  Misses  H.  B.  Thompson  and  K.  Gk)rdon,  found  no  indication 
of  Hellpach's  zone  of  complementarism.  They  refer  the  images  to 
the  enhancing  influence  of  a  light  background. 

*  J.  W.  Baird,  "  The  Color  Sensitivity  of  the  Peripheral  Retina,"  Carnegie 
Institution  of  Washington,  Publication  No.  29,  May,  IQ05,  73. 

• "  A  Study  of  After-images  on  the  Peripheral  Retina,"  Psychol  Review, 
XIV.,  March,  1907,  126  f.,  129  f. 


1908.]  OF  SUBLIMINALLY  COLORED   STIMULI.  369 

Again,  in  1908,  in  a  continuation  of  her  former  study,  Miss 
Femald  writes :  "  In  agreement  with  the  observations  already  made 
in  our  first  paper,  and  later  in  the  work  of  Miss  Thompson  and  Miss 
Gordon,  our  results  show  that  in  many  cases  a  characteristic  colored 
after-image  follows  an  unperceived  color  stimulus.  In  general  this 
after-image  is  perfectly  clear  and  distinct.  .  .  .  That  the  phenomena 
here  described  are  genuine  after-images  is  shown  by  the  fact  that 
the  color  is  in  every  case  the  color  complementary  to  the  stimulus 
as  [it  would  be]  perceived  either  in  central  or  in  peripheral  vision, 
although  the  observer  was  kept  in  complete  ignorance  concerning 
the  nature  of  the  stimuli  employed,  and  so  had  no  clew  as  to  what 
after-image  was  to  be  expected  in  cases  in  which  the  [color  of  the] 
stimulus  was  not  seen.  Moreover,  gray  and  white,  though  fre- 
quently used  as  stimuli,  were  never  followed  by  colored  after- 
images."  •  Hellpach  here  drops  out  of  sight  altogether,  while  the 
range  of  the  subliminally  aroused  after-image  is  extended,  from 
"the  extreme  periphery,"  to  include  both  the  B-Y  and  the  R-G 
zones. 

New  Experiments. 

I.  Direct  Vision:  (a)  Light-Adaptaiion. 

Experiment  /.;  The  Glass  Wedge. — ^We  wished  to  begin  our 
own  experiments  by  repeating  Tschermak's  observation  with  the 
faintly  colored  glass  wedge.  However,  the  difficulty  of  finding  a 
suitable  glass  proved  to  be  so  great  that  this  Exp.  I.  was,  as  a  mat- 
ter of  fact,  performed  last  of  all.  After  many  delays  we  were  able, 
through  the  kind  assistance  of  Professor  J.  A.  Brashear,  to  secure 
a  wedge  of  light  blue  glass,  5  by  20.5  cm.,  the  thin  end  of  which 
was  almost  colorless  in  clear  daylight.  Although  the  color  might 
well  have  been  still  fainter,  we  found  it  possible,  with  an  observation- 
slit  of  22  by  5  mm.,  and  with  a  white  muslin  screen  stretched  be- 
tween the  glass  wedge  and  the  white-screened  windows  from  which 
our  illumination  was  derived,  to  take  observations  of  2  to  5  min. 
duration,  in  which  the  wedge  was  moved,  for  the  practised  observ- 
ers, from  1.5  to  4  cm.,  and  for  the  impractised  from  5  to  10  cm. 

•"Studies  from  the  Bryn  Mawr  College  Laboratory:  The  Effect  of  the 
Brightness  of  Background  on  the  Appearance  of  Color  Stimuli  in  Peripheral 
Vision,"  Psychol.  Review,  XV.,  January,  1908,  33  flF. 


370  TITCHENER   AND  PYLE— ON   THE   AFTER-IMAGES        [July  .3, 

We  made  no  long  series  of  tests,  since  the  question  at  issue  had 
already  been  answered,  so  far  as  we  could  answer  it,  by  the  follow- 
ing Exps.  II.-V.  The  experiments  were,  however,  carefully  con- 
ducted. The  work  was  done  in  a  long  gray-tinted  light-optics  room, 
with  achromatic  adaptation;  the  observers  were  the  writers  (T,  P), 
Mr.  L.  R.  Geissler  (G),  assistant  in  psychology,  and  two  unprac- 
tised students,  Mrs.  G.  L.  de  OUogni  and  Mr.  E.  M.  Stevens ;  and 
the  experimenter  had  acquired  great  skill,  from  Exps.  II.  and  IV., 
in  moving  the  wedge  slowly  and  steadily  forward.  In  general,  the 
stimulus-background  was  black,  and  the  field  for  the  projection  of 
the  after-image  was  white,  though  these  relations  were  occasionally 
changed. 

As  we  had  expected,  there  was  no  trace  of  color  in  the  after- 
image ;  this  result  was  uniform.  In  control  experiments,  in  which 
(after  a  period  for  the  recovery  of  the  eye)  the  glass  was  exposed 
for  30  sec.  at  the  point  finally  reached  in  the  adaptation  experiments, 
the  after-image  showed  a  brief  period  of  dirty  orange  or  brownish 
yellow,  followed  by  gray. 

Experiment  II.:  The  Marbe  Color  Mixer, — ^The  observations 
with  Tschermak's  wedge  could  not,  in  any  case,  be  regarded  as  more 
than  preliminary.  For  systematic  work  we  employed,  first,  the 
Marbe  color  mixer,  which  permits  the  change  of  a  colored  sector 
during  rotation  of  its  discs,  and  thus  gives  scope  for  progressive 
adaptation. 

The  observer,  head  in  rest,  was  seated  at  a  distance  of  i  m.  from 
a  black  cardboard  screen.  The  rotating  discs  were  observed 
through  a  circular  opening,  2  cm.  in  diameter,  cut  in  the  screen  at 
the  level  of  the  eyes.  The  observation  was  monocular,  and  was 
continued  for  S  to  7  min.  The  discs  were  made  up  of  white,  with 
a  sector  of  colored  paper  (Zimmermann  R,  Y,  G,  B,  V)  ;  the  color 
at  the  outset  was  subliminal  for  the  achromatically  light-adapted 
eye,  and  was  gradually  increased  in  amount  as  the  observation  pro- 
ceeded. The  after-image  was  projected  upon  a  fixation-point 
marked  on  awhite  cardboard  dropped  in  front  of  the  black  screen.* 

'For  comparative  purposes,  a  few  observations  were  taken  with  a  gray 
screen,  and  with  projection  upon  a  black  or  gray  background.  Nothing  new 
resulted. 


1908.] 


OF  SUBLIMINALLY  COLORED  STIMULI. 


371 


The  regular  observers  were  T,  P,  G,  and  Mr.  T.  Nakashima,  gradu- 
ate scholar  in  psychology  {N).  A  few  observations  were  secured 
from  Professor  I.  M.  Bentley  (S),  and  from  an  unpractised  ob- 
server, Mr.  H.  J.  Bool;  single  observations  were  made  by  several 
visitors  to  the  laboratory. 

In  intention,  the  procedure  was  without  knowledge.  In  practice, 
the  experimenter  found  it  impossible,  in  the  early  stages  of  the 
work,  to  regulate  the  size  of  the  colored  sector  in  precise  accordance 
with  the  course  of  adaptation.  The  observer  was  therefore  in- 
structed to  tap  on  the  table  with  a  pencil  whenever  he  perceived 
a  color  in  the  stimulus.  If  a  tap  was  given,  the  experimenter  ran 
the  colored  sector  back  through  five  or  ten  degrees,  and  continued 
the  experiment  from  that  point.  The  results  of  these  interrupted 
observations  varied,  according  to  the  frequency  of  the  taps  and  the 
insistence  of  the  color  in  the  stimulus.  The  following  are  typical 
records. 

A,    No  Color  Seen  in  Stimulus. 

Duration  of  Obs. 


Observer. 

Color  in  Disc. 

After-image. 

T 

205^  B 

Gray 

i8o^  V 

Gray 

P 

120^   G 

Gray 

140^  V 

Gray 

G 

135^  R 

Gray 

B 

190^  B 

Gray 

7  m 
6  m 
5  m 

5  m 

6  m 
6  m 


n. 

n. 

n. 

n.  IS  sec. 

n. 

n. 


After  a  period  for  recovery,  the  stimuli  were  exposed  at  their 
final  color-strength  for  30  sec,  and  the  after-image  was  projected 
as  before.    The  results,  in  the  above  instances,  were  as  follows : 


> 


Obsenrer. 

Color  Seen. 

After-ima^. 

T 

Blue 

Brownish  yellow 

Bluish  violet 

Dirty  olive  yellow 

P 

Green 

^ 

Pink 

Bluish  violet 

Dingy  yellow 

G 

Red 

Gray 

B 

Blue 

Qear  yellow 

B,    Color  Seen 

in  Stimulus 

• 

• 

Observer. 

Color  in  Disc. 

After-ima^. 

Duration  of  Obs 

T 

140*  G 

Gray 

5  min. 

G 

120^   Y 

Dark  blue 

5  min. 

220*   B 

?  Orangish 

7  min. 

N 

155*R 

Gray 

7  min. 

130*  Y 

Blue 

6  min. 

372  TITCHENER  AND   PYLE— ON  THE  AFTER-IMAGES        [July  23. 

The  control  experiments,  with  30  sec.  exposure,  gave  the  results : 

Obsenrer.  Color  Seen.  After  image. 

T  Green  Purple 

G  Yellow  Dark  blue 

Blue  Yellow 

N  Pink  ?  Violet 

Yellow  Blue 

The  general  results  of  these  experiments  may  be  summed  up  in 
the  following  propositions. 

1.  With  every  one  of  our  observers,  regular  and  casual,  we  have 
been  able  to  raise  a  color-component  in  the  stimulus  from  a  sub- 
liminal to  a  normally  supraliminal  value,  while  the  stimulus  ap- 
peared throughout  as  gray.  In  no  instance  of  this  kind  has  the 
observer  found  the  complementary  color  in  the  after-image.  Our 
results  thus  stand  in  direct  opposition  to  the  observation  of 
Tschermak. 

2.  There  are,  however,  marked  individual  differences  among  the 
observers.  In  the  7  min.  which  represented  the  limit  of  our  obser- 
vations, it  was  difficult,  with  G  and  N,  to  increase  the  color-com- 
ponent, without  detection,  to  a  normally  supraliminal  amount :  with 
T,  P  and  B  there  was  no  such  difficulty.  The  control  images,  on 
the  other  hand,  were  obtained  most  readily  from  T  and  P. 

The  observer  N  is  of  the  subjective  type,  and  is  often  misled 
by  an  "  expected  "  or  "  imagined  "  color.  Thus  a  disc  containing 
17s**  G  was  seen  as  B  with  a  rim  of  Y;  the  after-image,  after  6 
min.,  was  a  Y  of  irregular  form,  larger  than  the  stimulus.  We 
recur  to  these  "  imagined  "  colors  later.  The  remaining  observers 
were  of  a  distinctly  objective  type. 

3.  There  were  also,  as  might  be  expected,  marked  differences  in 
the  "coloring  power"  of  the  Zimmermann  papers.  Experiments 
of  the  form  A  were  easiest  with  B,  less  easy  with  V;  then  follow 
in  order  R,  G,  Y.  The  last-mentioned  color,  indeed,  gave  results 
only  with  entirely  naive  and  unpractised  observers.  The  R  and  G, 
when  seen  as  color,  usually  appeared  first  as  Y. 

4.  As  a  rule,  the  after-images,  whether  colored  or  gray,  devel- 
oped very  slowly.  The  gray  images,  in  particular,  might  appear 
only  after  a  blank  interval  of  15  to  30  sec.    They  usually  showed 


I 

■ 

t 

I 

f 


,5o8]  OF  SUBLIMINALLY   COLORED   STIMULI.  373 

two  Stages,  dark  and  light.  The  colored  images,  both  of  the  regular 
and  of  the  control  experiments,  passed  oflf  as  gray. 

Experiment  III.:  The  Color  Mixer  with  Unchanged  Discs. — So 
far  we  have  followed  and  systematised  Tschermak's  method;  the 
amount  of  color  in  the  stimulus  has  increased,  during  the  single 
observation,  and  has  been  compensated  by  a  progressive  adaptation. 
In  the  present  experiments  the  amount  of  color  in  the  discs  is  in- 
creased from  subliminal  to  normally  supraliminal,  step  by  step,  in 
successive  observations. 

The  rotating  discs  were  observed,  as  before,  through  a  circular 
opening  in  a  black  or  neutral  gray  screen.  The  discs  themselves 
were  made  up  of  neutral  gray  (identical  with  that  of  the  screen), 
with  a  colored  sector  (Hering  R,  Y,  G,  B).  The  stimulus  was 
fixated  for  i  min.,  and  the  after-image  was  projected  upon  a  neutral 
g^ay  or  black  background.  P,  G  and  N  served  as  regular  observers : 
a  few  observations  were  also  taken  from  B  and  T.  The  following 
are  typical  results. 

Color  in 
After-ima^. 

None 

None 

Pinkish 

None 

None 

None 

Yellow 

None 

None    * 

Green 

None 

None 

Dark  blue 

In  the  above  observations,  the  black  screen  and  the  neutral  gray  back- 
ground were  employed.  Other  arrangements  of  screen  and  background  gave 
similar  results. 

In  no  case  was  a  colored  after-image  obtained  from  a  sublimi- 
nally  colored  stimulus.  On  the  contrary,  the  image  appeared  only 
when  the  stimulus-color  was  distinctly  supraliminal. 


Observer. 

Color  in  Disc. 

Color  Seen. 

G 

4'  G 

None 

10^  G 

Green 

14^  G 

Green 

N 

9^  B 

None 

12^  B 

None 

20*  B 

?  Pinkish 

50^  B 

Blue 

P 

6^  R 

?  Ruddy 

10**  R 

Red 

30^  R 

Red 

T 

6**  Y 

None 

12**  Y 

?  Yellowish 

20*  Y 

Yellow 

374  TITCHENER  AND  PYLE-ON  THE  AFTER-IMAGES       [July  as, 

(&)  Dark-Adaptation, 

Experiment  IV.:  The  Glass  Wedge. — Besides  furnishing  the 
light  blue  wedge  of  Exp.  L,  Professor  Brashear  supplied  us  with 
smaller  and  more  highly  colored  wedges  of  claret,  red,  orange, 
green  and  blue  glass.  With  these,  or  with  combinations  of  them, 
we  proceeded  as  follows. 

A  sheet  of  ground  glass  was  inserted  in  the  Hering  window  of 
a  large  dark-room:  the  width  of  the  strip  could  be  regelated  at 
will.  Some  2.50  m.  before  the  window  was  a  table,  on  which  stood 
a  large  screen  of  white  cardboard.  Immediately  behind  a  vertical 
slit  in  this  screen  (3  by  25  mm.)  lay  a  grooved  strip  of  wood,  in 
which  the  wedge  or  wedges  could  be  moved.  Observations  were 
made  in  dark-adaptation.  The  thick  end  of  the  wedge  was  first 
shown ;  it  appeared  as  black  or  as  dark  gray.  The  wedge  was  then 
moved  along,  very  slowly:  if  the  observer  saw  its  color,  he  tapped 
with  a  pencil,  and  the  experimenter  withdrew  it  a  trifle,  to  start 
again  after  a  few  seconds.  At  a  given  signal,  the  observer  looked 
away  from  the  slit  to  the  cardboard  screen,  or  to  a  black  surface 
directly  below  the  screen,  and  watched  the  development  of  the  after- 
image. The  regular  observers  were  T,  P,  G  and  N;  a  few  observa- 
tions were  also  made  by  B. 

Owing  to  the  difficulty  of  procuring  the  large  glass  wedge  of 
Exp.  I.,  these  dark-room  observations  were  the  first  taken.  And, 
in  our  desire  to  do  justice  to  Tschermak's  method,  we  spent  more 
time  and  trouble  upon  them  than  we  like  to  recall.  The  observer's 
head  was  fixed  securely  in  a  head-rest;  the  height  of  the  screen 
was  carefully  adjusted ;  generous  time  was  allowed  for  adaptation ; 
the  admission  of  light  was  rigorously  controlled,  beforehand,  by  the 
experimenter;  the  tmiform  movement  of  the  wedge  was  assiduously 
practised.  We  were  rewarded,  however,  by  the  unequivocal  charac- 
ter of  the  results.  Though  observation  might  be  continued  for  5 
min. ;  though  during  this  period  the  observer  might  tap  his  glimpse 
of  color  no  less  than  seven  times ;  and  though  in  the  control  experi- 
ments, with  immediate  observation  of  the  part  of  the  wedge  finally 
exposed,  a  good  complementary  after-image  might  be  obtained  in 


,9o8]  OF   SUBLIMINALLY  COLORED  STIMULI.  375 

20  sec. :  we  did  not  once,  in  the  course  of  the  principal  experiments; 
obtain  a  record  of  color  in  the  after-image.  Sometimes  the  after- 
image failed  to  appear  at  all ;  more  often  it  appeared,  and  obstmately 
remained,  as  gray. 

The  duration  of  a  single  observation  varied  between  the  limits  of  2  min. 
30  sec.  and  5  min.;  most  of  the  exposures  were  about  3  min.  The  number 
of  taps  varied  from  o  to  7;  the  average  for  all  observers  was  4.  The  color 
was  thus  much  more  insistent  than  in  Exp.  II.  —  partly,  no  doubt,  because  the 
range  of  possible  movement  was  only  about  one-third  of  that  allowed  by  the 
Marbe  mixer.  In  the  control  experiments,  T  and  P  obtained  the  colored 
after-image  fairly  easily ;  G,  N  and  B  often  failed  to  secure  it 

Experiment  V,:  Colored  Papers. — These  observations  were  also 
made  in  the  dark-room  and  with  dark-adaptation.  A  number  of 
Milton-Bradley  colored  papers,  4  by  8  cm.,  were  pasted  upon  white, 
neutral  gray  and  black  grounds.  The  Hering  window  was  so  ad- 
justed that,  for  the  experimenter,  the  color  of  the  particular  paper 
exposed  was  just  subliminal.  The  observers  {T,  P,  G,  N  and  occa- 
sionally B)  fixated  the  colored  strip  at  a  distance  of  i  m.  for  40 
sec,  and  projected  the  after-image  upon  a  white,  neutral  gray  or 
black  surface.  All  possible  combinations  of  stimulus-ground  and 
projection-ground  were  employed. 

The  observer  was  instructed  to  report  the  quality  of  the  stimulus 
as  it  appeared  at  first  fixation,  and  to  mention  any  qualitative  change 
that  it  might  undergo  in  the  course  of  an  observation.  In  most 
cases  the  color  was  subliminal ;  and  the  subliminally  colored  stimulus 
never  gave  a  colored  after-image.  In  the  cases  in  which  the  color 
of  the  strip  was  seen,  the  after-image  was  sometimes  colored,  some- 
times gray. 

The  direct  judgment  of  color  under  these  conditions  is  extremely 
difficult,  and  the  observer  is  sorely  tempted  to  avail  himself  of 
secondary  criteria — brightness,  velvetiness,  depth,  shimmer,  etc.  An 
observer  of  the  objective  type  soon  learns,  however,  to  distinguish 
between  vision  and  imagination :  "  I  can  see  nothing,"  he  will  say, 
"  but  I  should  guess  that  it  is  red  "  or  what  not.  The  guesses  were 
confined — probably  from  the  analogy  of  the  immediately  preced- 
ing Exp.  III. — to  the  four  colors  R,  Y,  G,  B ;  and,  as  we  had  the 
full  set  of  Milton-Bradley  papers  at  our  disposal,  they  were  more 


376  TITCHENER  AND   PYLE— ON  THE  AFTER-IMAGES        LFulyaa, 

often  wrong  than  right.®  Their  influence  upon  the  after-image  ap- 
peared only  in  the  case  of  the  subjective  observer  N.  Thus,  R  seen 
on  W  was  judged  by  N  to  be  "  red  or  blue  " ;  and  the  af ter-unage, 
also  on  W,  was  a  large  irregular  disc  of  yellow.  R  seen  on  Bk  was 
judged  to  be  "  bluish  " ;  and  the  after-image,  on  gray,  was  green- 
blue  with  a  vag^e  yellow  rim.  B  seen  on  W  was  judged  "  blue  or 
red  "  ;  and  the  after-image,  on  gray,  was  red  above  and  blue  below, 
with  a  yellow  patch  between.  It  is  noteworthy  that  here,  as  in  Exp. 
II.,  after-images  of  the  "  supposed  "  or  "  imagined  "  color  Invariably 
diflfered  in  form  and  size  from  those  of  the  true  color.  The  ob- 
server did  not  realise  the  significance  of  this  difference,  though  in 
time  he  would  doubtless  have  learned  to  use  it  as  a  secondary 
criterion.  ' 

II.   Indirect  Vision. 

We  have  already  mentioned  the  experiments  made  by  T  in  1906 
with  the  view  of  testing  the  conclusions  of  Miss  Fernald's  first 
paper.  The  observations  were  rigorotisly  confined  to  the  Bk-W 
zone,  and  their  outcome  was  definitely  negative.  In  the  meantime, 
however,  the  arousal  of  a  colored  after-image  by  a  subliminally 
colored  stimulus  had  been  maintained  for  both  the  B-Y  and  the 
R-G  zones.  Unsystematic  observations  made  in  the  Cornell  Labora- 
tory failed  to  confirm  this  result.  It  seemed  worth  while,  however, 
to  obtain  further  testimony;  and  Professor  J.  W.  Baird,  of  the 
University  of  Illinois,  very  kindly  consented  to  investigate  the 
subject.* 

•One  of  the  observers  remarked  that  the  experiments  showed — ^what  he 
had  never  fully  understood  before — how  it  is  that  a  case  of  partial  color- 
blindness may  remain  undetected  both  by  the  color-blind  person  himself  and 
by  the  normal  persons  in  his  surroundings.  In  principle,  the  remark  was 
correct  enough;  but  in  practice  the  observer  would  have  had  to  revise  and 
extend  his  criteria  very  considerably. 

•All  the  observations  in  indirect  vision  mentioned  in  this  paper  were 
carried  out  with  light-adaptation.  Peripheral  after-images  in  dark-adapta- 
tion are  practically  non-existent.  In  op.  cit.,  56  f.,  Baird  writes:  "After- 
images— in  the  ordinary  sense  of  the  term — were  almost  invariably  absent 
from  our  experiments.  They  were  reported  in  less  than  one  per  cent,  of 
our  exposures;  and  when  they  did  occur,  they  were  aroused  by  the  stimula- 
tion of  paracentral,  never  of  peripheral,  regions  of  the  retina."  And  in  a 
personal  letter  he  adds:  "There  is  an  interesting  difference  of  function  in 


,9o8.]  OF  SUBLIMINALLY   COLORED  STIMULI.  377 

The  experiments  were  carried  out  by  means  of  a  simplified  form 
of  the  Zimmermann  perimeter,  which  permitted  an  accurate  record 
of  the  degree  of  eccentricity  at  which  the  stimulus  was  exposed. 
Exploration  was  confined  to  the  horizontal  nasal  meridian  of  each 
eye.  The  stimulus  was  a  beam  of  light  from  an  electric  (i6  c.  p.)* 
lamp,  transmitted  through  appropriate  combinations  of  gelatines  and 
colored  glasses;  the  colors  employed  were  (non-equated)  B  and  Y, 
R  and  G.  Six  of  the  most  reliable  laboratory  students*®  acted  as 
observers,  and  Professor  Baird  had  personal  charge  of  the  entire 
work.  The  after-images  were  projected  upon  white,  gray  and  black 
grounds.  The  experiments  proper  were  preceded  by  a  careful  de- 
termination of  the  outermost  limits  of  color  vision  for  the  stimuli 
used,  and  all  pains  were  taken  to  avoid  chromatic  adaptation. 

The  following  may  serve  as  a  sample  of  method  and  results. 

Determination  of  Outermost  Limits  of  Blue  Vision:  Observer  Bu. 

Right  Eye, 

Prelftninary.  Series  x. 

90*-75*  Nothing  72*-58*  Black 

70*-S5*  Dark  gray  56*-SO*  Bluish 

50^-45*  Bluish  48**-       Blue 

40**-  Blue 

Series  a.  Series  3. 

75*-63'  Black  73*-6i*  Very  dark  gray 

61^-53**  Bluish  59**-       Bluish 

51*-       Blue 

Outermost  limit  (bluish  or  blue)  :  61*. 

Left  Eyey^ 

90*-^5*  Nothing  62*-5o*  Black 

6o*-40*  Dark  gray  48**-44*  Bluish 

35**-       Bluish  to  blue  ^  42**-       Blue 

6o**-48*  Black  58*-44*  Black 

46**-        Bluish  42*-        Bluish 

Outermost  limit  (bluish  or  blue)  :  48*. 

the  peripheral  retina  in  light-adaptation  and  in  dark-adaptation.  In  the 
latter  case,  after-images — ^both  uncolored  and  colored — are  faint  or  wholly 
lacking.  In  the  former  case  they  are  readily  perceptible.  Yet  even  in 
light-adaptation  they  are  less  perceptible  than  arc  the  primary  images 
aroused  by  the  given  stimuli." 

"  The  Misses  M.  Miller,  A.B.,  and  B.  Scoggin :  and  Messrs.  C.  B.  Busey» 
A.B.,  R.  Garrett,  O.  L.  Hemdon  and  A.  C.  Schertz,  A.B. 

"  The  visual  acuity  of  the  left  eye  was  less  than  that  of  the  right 


378  TITCHENER   AND   PYLE— ON  THE  AFTER-IMAGES        [Julyaa. 

* 

Perimetrical  Experiments. 


Stimulus. 

Duratioo. 

Perception. 

After-image. 

po**  Right 

30  sec. 

Nothing 

None 

90**  Left 

30  sec. 

Nothing 

None 

8o*  Right 

30  sec. 

Nothing 

None 

8o^  Left 

30  sec. 

Nothing 

None 

70**  Right 

40  sec. 

Dark  gray 

None 

70^  Left 

40  sec. 

Nothing 

None 

60**  Right 

40  sec. 

Bluish,  then  black 

None 

60**  Left 

40  sec. 

Gray 

None 

50**  Right 

40  sec. 

Dark  bluish, 

then  gray 

Yellowish,  then  gray 

50**  Left 

40  sec. 

Trace  of  bluish,  then  gray 

Gray 

» 


It  does  not  seem  necessary  to  print  the  full  set  of  results,  though 
the  data  are  at  the  disposal  of  anyone  who  may  wish  to  consult 
them.  The  net  outcome  of  the  enquiry,  in  Professor  Baird's  words, 
is  as  follows :  "  In  not  a  single  instance  did  any  stimulus  give  a 
colored  after-image  at  a  retinal  region  where  it  gave  an  uncolored 
image,"  t.  e,,  where  it  was  seen  as  black  or  gray.  He  proceeds: 
"  I  have  tried  every  variation  of  the  conditions  (with  fxclusion  of 
chromatic  adaptation)  v/hich  my  ingenuity  could  devise;  and  the 
result  is  in  every  instance  n^rgative,  so  far  as  the  contention  of  the 
Misses  Femald,  Thompson  and  Gordon  is  concerned." 

Criticism  and  Interpretation. 

1.  The  positive  outcome  of  Tschermak's  observations  with  the 
glass  wedge  must,  in  our  opinion,  be  explained  by  the  prepossession 
of  the  observer  and  the  roughness  of  the  method  employed.  Had 
Tschermak  been  in  doubt  as  regards  the  after-image,  he  would  have 
had  recourse  to  a  more  refined  instrument,  as  the  Marbe  color-mixer. 
And  had  he  adopted  a  better  method,  we  cannot  doubt,  on  our  side, 
that  the  outcome  of  his  observations  would  have  been  negative.  We 
may,  perhaps,  venture  to  express  the  hope  that  he  will  now  submit 
his  hypothesis  to  a  stricter  test. 

2.  It  is  less  easy  to  account  for  the  peripheral  results.  The 
experimentum  cruets,  in  positive  regard,  would  seem  to  be  the  pro- 
duction of  a  colored  after-image,  in  the  achromatically  adapted  eye, 
at  a  point  lying  well  beyond  the  limits  of  B-Y  vision.  It  must  be 
remembered  that  in  all  liminal  determinations  an  unnoticed  varia- 
tion in  physical  or  physiological  conditions,  or  in  the  conditions  of 


i9o«.]  OF   SUBLIMINALLY   COLORED   STIMULI.  379 

attention,  may  lead  to  a  serious  variation  of  numerical  result.  It 
is,  for  instance,  exceedingly  doubtful  if  any  but  the  most  careful 
and  most  highly  practised  observers  can  maintain  their  fixation  so 
accurately  as  to  ensure  a  precise  localisation  of  the  retinal  area 
affected  by  a  given  stimulus.  Moreover,  we  are  here  dealing  with 
a  retinal  function  which  tails  oflf  gradually  from  center  to  periph- 
ery: so  that  a  very  slight  shift  of  regard,  or  a  momentary  lapse 
of  attention,  or  a  minimal  change  in  adaptation  or  in  illumination 
may  be  enough  to  vitiate  an  observation.  An  illustration  may  be 
taken  from  the  records  of  the  observer  Sm.,  quoted  above.  The 
outermost  limit  of  B-vision,  in  the  left  eye,  was  determined  as  48**. 
Nevertheless,  the  observer  reported,  in  the  experiments  proper,  a 
"  trace  of  bluish,  then  gray  "  with  the  stimulus  at  50°.  There  was 
no  colored  after-image.  But  suppose  a  tinge  of  blue-adaptation: 
then  we  might  have  had  a  perception  of  gray,  and  a  yellow  after- 
image ;  and  we  should  still  have  been,  apparently,  beyond  the  limit 
of  B-Y  vision.  It  was  only  the  care  taken  to  avoid  chromatic 
adaptation  that  prevented  the  positive  result. 

It  is,  of  course,  precisely  this  crucial  experiment  which  is  de- 
scribed affirmatively  by  Miss  Fernald  in  1905,"  and  which  came  out 
negatively  in  Vs  experiments  of  1906.  The  question  then  arises 
as  to  the  accuracy  of  determination  of  the  zonal  limits^^  And  on 
this  point  we  may  quote  specimen  results  from  Miss  Fernald's 
tables. 

I.  R  stimulus  on  light  gray  background." 

io*'-73®  Stimulus  uniformly  seen  as  red. 

74.5**  No  color  seen, 

76**  Red  seen  in  two  observations. 

80*  Red  seen  in  four,  no  color  seen  in  two  observations. 

82.5®  No  color  seen. 

84®  No  color  seen. 

"  We  follow  the  phrasing  of  the  Psychol  Review  of  1905 :  "  Exposure, 
beyond  the  limits  where  any  color  is  seen,  is  followed  by  a  very  clear  after- 
image." In  the  Journ.  Philos.,  Psychol.  &  Sci.  Meth.,  iii.,  1906,  352  (Report 
of  Sec.  of  N.  Y.  Acad,  of  Sciences),  the  report  reads:  "After-images  were 
perceived,  almost  without  exception,  as  far  out  as  any  color  could  be  dis- 
tinguished, and  in  many  cases  were  clearly  seen  though  the  stimulus  color 
was  not  recognised." 

"  Psychol.  Review,  XII.,  408.     Italics  ours. 


'  380  TITCHENER  AND  PYLE— ON  THE  AFTER-IMAGES        [July  aj. 

85.5*  Red  seen  once,  no  color  seen  once. 
87**  Red  seen  twice,  no  color  seen  twice. 
The  conditions  can  hardly  have  remained  constant  from  74.5*  to  87*. 
Again,  R  on  Hering  gray  no.  7  is  seen  colorless  at  37**,  while  it  is  seen  red 
at  39**,  41.5*  (twice),  and  even  at  47*  (twice)."  And  yet  again,  G  on  the 
same  gray  is  seen  colorless  at  82'',  green  at  84**,  and  once  colorless  and  once 
green  at  87*.**     Instances  of  this  irregularity  might  easily  be  multiplied. 

2.  If  we  turn  to  the  special  table  for  the  limits  of  B  and  Y,  we  find  a 
greater  uniformity  of  result,  but  a  certain  arbitrariness  in  the  selection  of 
the  limiting  values.  Thus,  on  various  backgrounds  and  for  different  ob- 
servers, the  limits  for  Y  are  taken  as 

(fl)  97''»     although  at  98.5*  the  color  is  seen  3  times  out  of  14, 
(b)  88.5®,  although  at  92.5*  the  color  is  seen  once  in  3  times, 
(<c)  ^.5*,  although  at  98.5*  the  color  is  seen  once  in  3  times, 
(rf)  92.5®,  although  at  95.5*  the  color  is  seen  3  times  out  of  10, 
and  so  on.     Similarly,  the  limits  for  B  are  taken  as 

(a)  88.5®,  although  at  91.5*  the  color  is  seen  once  in  4  times, 

(fi)  97^>     although  at  99.5^  the  color  is  seen  once,  and  one  observation 

is  doubtful. 
ic)  97*,     although  at  99.5*  the  color  is  seen  3  times  out  of  9,  with  one 
observation  doubtful, 
and  so  on." 

Now  in  her  second  paper,  of  1908,  Miss  Femald  states  that  the 
paradoxical  after-images  "  are  perceived  most  frequently  either  just 
inside  or  just  beyond  the  regular  limits  for  the  color."*^  If  this 
statement  may  be  applied  to  the  limits  of  color  vision  at  large,  ».  e., 
to  the  work  of  1905,  we  must  conclude  that  the  crucial  experiment 
has  not  been  adequately  performed ;  for  the  limits  given  are,  as  we 
have  seen,  irregular  and  arbitrary. 

Each,  however,  if  we  maintain  that  7^s  results  are  conclusive  for 
the  Bk-W  zone,  we  have  still  to  account  for  the  colored  after-images 
of  subliminally  colored  stimuli  in  the  B-Y  and  R-G  zones.**     Miss 

"  Ibid,,  4;22. 

''Ibid,,  416. 

^Ibid.,  402. 

"  Psychol.  Review,  XV.,  33. 

"Miss  Femald  uses  the  term  " unperceived,"  not  subliminal.  The  latter 
word  is,  however,  employed  by  the  Misses  Thompson  and  Gordon,  whose 
results  Miss  Femald  assimilates  to  her  own.  That  "  unperceivcd "  really 
means  "  imperceptible  "  is  shown  also  by  a  passage  in  a  letter  received  from 
Miss  Fernald :  "  I  should  be  very  much  afraid  of  my  observer's  life,  if  it 
depended  on  his  identification  of  the  stimulus  color,  in  all  cases  in  which  a 
clearly  colored  after-image  is  seen.  In  fact,  when  forced  to  say  what 
stimulus  he  thought  was  used,  he  guessed  at  B  for  O  as  often  as  O  for  O, 
insisting  all  the  while  that  he  did  not  see  any  color." 


x9o8.] 


OF  SUBLIMINALLY  COLORED  STIMULI. 


381 


/ 


Femald  has  been  good  enough  to  send  us  an  account  of  the  condi- 
tions under  which  her  observers  found  the  after-image,  and  to  make 
a  special  series  of  observations,  with  Mr.  C.  E.  Ferree  as  observer. 
"  The  head,"  she  says,  "  must  be  held  firm  (my  method  is  the  bit, 
with  the  impression  of  the  teeth).  The  background  must  be  light, 
and  the  illumination  good.  The  observer  must  hold  the  fixation 
steadily  after  the  stimulus  is  removed.  The  after-image  screen 
must  be  white  to  obtain  Y  or  B  after-images  and  black  to  obtain 
R  after-images.  A  very  slight  change  in  conditions  makes  a  great 
difference  in  results,  which  seem  to  me  to  depend  wholly  on  bright- 
ness." Professor  Baird  was  acquainted  with  these  conditions  be- 
fore he  undertook  his  perimetrical  observations. 
The  new  set  of  observations  is  as  follows. 

Observer:  C.  K  Ferree.  Full  illumination  on  bright  day  (May  17,  1908). 
Nasal  meridian,  right  White  ground.  Projection  field  white,  except  in  obs. 
14-17,  when  it  was  black.  Stimulus,  13  sq.  mm.  Distance  from  eye  to 
stimulus,  25  cm. 


ion  Point.      Stimulus. 

Color  Seen. 

After-image. 

8o* 

0 

Dark  gray 

Unsaturated  light  blue 

85* 

B 

Just  dark 

Wash  of  unsaturated 
yellow 

85* 

Y 

Nothing 

Nothing 

8o* 

Y 

Tinge  of  dirty  yellow 

Very  pale  blue 

8o* 

Medium  gray 

Dark 

White 

8o* 

0 

Indefinite  gray 

Nothing 

8o* 

Light  gray 

Dark 

White 

75' 

Y 

Reddish  yellow 

Good  blue 

75* 

B 

Good  blue 

Good  yellow 

75* 

B 

Good  blue 

Good  yellow 

65' 

0 

Yellowish  red 

Unsaturated  blue 

65* 

Y 

Reddish  yellow 

Blue 

6o^ 

G 

Indefinite  greenish  gray 

Uncertain 

65* 

G 

Greenish  yellow 

Dark  red,  more  satu- 
rated than  stimulus 

8o* 

Medium  gray 

Dark 

Nothing 

80'' 

Medium  gray 

Dark 

Nothing 

65* 

G 

No  color 

Flash  of  red 

65* 

R 

No  color 

Blue 

Positive  results  occur  in  the  two  first  and  two  last  observations 
of  the  series.  The  former  may  be  explained  in  terms  of  chromatic 
adaptation.    If,  as  the  illtunination  suggests,  the  observer  began  the 

PROC.  AMER.  PHIL.  SOC.,  XLVII.  I89  V,  PRINTED  OCTOBER  3,  I908. 


X 


382  TITCHENER   AND   PYLE— ON   THE   AFTER-IMAGES        [July  23, 

work  in  Y-adaptation,^*  the  first,  blue  after-image  would  naturally 
follow.  If  the  second  observation  was  taken  at  too  short  an  interval 
of  time,  the  resulting  B-adaptation  would  show  itself  as  a  yellow 
after-image.  The  two  final  observations  suggest  a  shift  of  condi- 
tions. G  is  seen  at  65°  as  greenish  yellow,  and  as  colorless;  at  60 ** 
as  indefinite  greenish  g^ay.  It  is  possible  that,  in  the  case  in  which 
"  no  color  "  is  reported,  the  G  simply  escaped  notice ;  peripheral 
colors  at  the  limit  of  vision  often  appear  as  momentary  flashes. 
Again,  R  is  reported  at  65°  as  "  no  color,"  although  "  reddish  yel- 
low "  had  been  seen  as  far  out  as  75°.  It  is  possible  that  the  flash 
of  red  escaped  notice ;  it  is  also  possible  that  R-adaptation,  from  the 
preceding  after-image,  brought  out  the  blue. 

The  puzzling  thing  is  that  the  positive  outcome  should  be  thus 
definite  in  the  Mount  Holyoke  and  Bryn  Mawr  laboratories,  while 
neither  Professor  Baird  nor  ourselves — ^though  working  with  full 
knowledge  of  conditions,  and  though  trying  various  possibilities 
which  have  not  been  reported  in  detail**' — ^are  able  in  a  single  case 
to  obtain  the  colored  after-image.  We  can  only  guess  at  an  expla- 
nation ;  and  we  oflfer  the  following  guesses  in  what  seems  to  us  to 
be  the  order  of  their  likelihood:  (i)  chromatic  adaptation;*^  (2) 
the  momentary  and  flash-like  appearance  of  colors  at  the  limit  of 
vision;  (3)  the  phenomenon  of  "fluctuation  of  attention";  (4)  de- 
fective method  and  unsystematic  procedure  in  the  determination 

"These  observations  were  taken  "after  the  limits  had  been  roughly  de- 
termined in  previous  experiments."  If  the  determination  of  limits  was  made 
at  the  same  sitting,  and  if  the  last  test-color  employed  was  O,  there  would 
be  additional  reason  for  an  initial  Y-adaptation. 

"Thus,  Mr.  Ferree  wrote  to  us:  "After-images  seem  to  occur  most  in- 
tensively when  the  stimulus  is  removed  while  adaptation  is  still  going  on. 
If  one  carries  the  stimulation  to  a  stationary  point  in  adaptation,  the  after- 
image will  weaken  in  proportion  to  the  length  of  time  during  which  the 
stimulus  is  regarded  before  the  after-image  is  evoked.  This  is  true  whether 
one  uses  intensive  or  slightly  supraliminal  stimuli."  We  thought  that  it 
might  possibly  be  true  of  subliminal  stimuli,  and  accordingly  made  brief 
observations  both  in  light  and  in  dark  adaptation.  But  we  never  saw  the 
after-image. 

"On  chromatic  adaptation,  see  Baird,  op,  cit.,  57  if.,  64  if.,  73  f.;  Joum. 
Philos,,  Psychol.  &  Sci.  Meth,,  II.,  1905,  21. 


,9o8]  OF  SUBLIMINALLY   COLORED   STIMULI.  383 

of  zonal  limits ;  (s)  unnoticed  variations,  physical,  physiological  or 
psychological,  in  the  conditions  of  observation  during  a  series.** 

We  are  well  aware  that  negative  experiments  are  logically  in- 
conclusive.** The  fact  that  we  have  failed  to  find  the  colored  after- 
image does  not  prove  that  this  after-iijfage  is  non-existent.  We 
have,  however,  attempted  a  positive  explanation:  for  Tschermak's 
result,  in  terms  of  prepossession  and  inaccurate  method;  for  Miss 
Femald's  result,  in  terms  (predominantly)  of  chromatic  adaptation. 
Further  experimentation  by  other  observers  must  show  whether  our 
hypotheses  are  correct. 

We  are  aware,  also,  that  the  charge  of  prepossession  is  double- 
edged,  and  that  we  may  ourselves  be  accused  of  an  initial  bias.  We 
freely  confess  that  we,  as  well  as  Professor  Baird,  approached  the 
peripheral  experiments  in  a  sceptical  attitude  of  mind.  On  the 
other  side,  we  may  point  out  that  the  scepticism  was  positively  based 
upon  the  results  of  Baird's  Carnegie  Institution  research,  and  that 
the  student-observers  at  the  University  of  Illinois  knew  nothing  of 
the  question  at  issue. 

In  the  case  of  Tschermak's  observation,  however,  our  initial  bias 
was  positive ;  we  were  surprised  at  the  uniformly  negative  character 
of  our  results  with  the  Marbe  mixer.  Tschermak's  position  seemed 
to  accord  well  with  current  visual  theory.  Moreover,  we  knew 
that  a  contrast-color  may  be  more  saturated,  may  appear  more 
*'real,"  than  the  inducing,  objective  color.  We  knew  that  Hey- 
mans,  in  his  experiments  on  "psychische  Hemmung,"  had  some- 
times seen  the  contrast-color  while  the  inducing  color  was  still  un- 
perceived.**    We  knew  of  Helmholtz'  statement,  "dass  die  gesat- 

"  In  a  letter  to  T  Miss  Femald  remarks :  "  You  will  see  that  colored  after- 
images were  seen  in  less  than  one  third  of  the  total  number  of  cases  in  which 
the  stimulus-color  was  not  seen."  In  a  communication  made  to  Professor 
Baird,  she  estimates,  roughly,  that  the  phenomenon  appeared  in  about  five 
per  cent,  of  her  exposures  upon  the  peripheral  retina.  This  sporadic  and 
fortuitous  character  of  the  after-images  suggests  that  they  are  the  product 
of  some  variable  condition  which  has  not  been  taken  account  of  in  the 
investigations. 

•J.  S.  Mill,  "A  System  of  Logic,"  1884,  SiS;  W.  S.  Jcvons,  "The  Prin- 
ciples of  Science,"  1900,  434. 

**G.  He)rmans,  Untersuchungen  tiber  psychische  Hemmung,  i.  Zeits,  f. 
Psychol,  u.  Physiol,  d.  Sinnesorgane,  XXL,  1899,  328.  "Wo  mit  weissep 
Sectorenscheiben  experimentirt  wurde,  kam  es  ofters  vor,  dass  ehe  noch  der 
Ring  die  Farbe  des  Papierstuckes  erkennen  Hess,  sich  im  Hintergrunde  schon 
die  Contrastfarbe  bemerklich  machte." 


384  TITCHENER   AND   PYLE— ON  THE   AFTER-IMAGES        [July  aj, 

tigsten  objectiven  Farben,  welche  existiren,  die  reinen  Spectral- 
farben,  im  unermiideten  Auge  noch  nicht  die  gesattigste  Farben- 
empfindung  hervornifen,  welche  iiberhaupt  moglich  ist,  sondem  dass 
wir  diese  erst  erreichen,  wenn  wir  das  Auge  gegen  die  Complemen- 
tarfarbe  unempfindlich  machen."  ***  There  was,  then,  no  a  priori 
reason  to  doubt  Tschermak's  result ;  on  the  contrary,  we  thought  it 
probable  that  under  conditions  which  were  unfavorable  to  the  ap- 
pearance of  the  stimulus-color,  but  favorable  to  the  appearance  of 
its  complementary,  the  subliminally  colored  stimulus  would  give  a 
perceptibly  colored  after-image.^*    As  a  matter  of  fact,  it  did  not. 

""Physiol.  Optik/'  1867,  370;  1896,  520.  Cf.  W.  Wundt,  "Physiol. 
Psychol.,"  II.,  1902,  146. 

"This  possibility  was  considered,  also,  in  the  peripheral  work;  so  that 
even  for  that  our  bias  was  not  wholly  negative. 


ON   THE   CLASSIFICATION   OF   THE   CETACEA. 

By  FREDERICK  W.  TRUE. 

(Read  April  24,  1908.) 

In  this  communication  I  wish  to  call  attention  to  the  various 
changes  in  the  generally-accepted  classification  of  the  Cetacea  pro- 
posed by  Professor  Dr.  O.  Abel,  of  the  University  of  Vienna,  in 
connection  with  his  recent  study  of  the  Miocene  toothed  whales — 
chiefly  those  obtained  from  the  vicinity  of  Antwerp,  and  now  in 
the  museurn  of  Brussels.^  Professor  Abel's  classification  (1905) 
is  as  follows : 

OdontocHes 
+  Archeocetes 
+  Squalodontidae 

Physeteridae 

Ziphiidae 


Argyrocetinae 
Acrodelphinae 
Iniinae 
.  Beluginae 


-j-  Eurinodelphidae 

Acrodelphidae  .  . 
+  Saurodelphidae 

Platanistidae 

Delphinidae 

I  would  call  attention  particularly  to  the  following  features  to 
which  my  remarks  will  mainly  relate: 

1.  The  use  of  the  term  "  Odontocetes  "  for  all  toothed  whales 
and  zeuglodonts. 

2.  The  subordination  of  the  "  Arch^ocetes  "  to  the  "  Odonto- 
cetes/' 

3.  The  new  family  Eurinodelphidae. 

4.  The  new  family  Acrodelphidae  (should  be  Iniidae). 

5.  The  inclusion  of  Delphinapterus  and  Monodon  in  this  family 
instead  of  in  Delphinidae,  and  the  inclusion  of  Stenodelphis  and 
Pontistes, 

^Mim.  Mus.  Roy.  Hist.  Nat.  Belgique,  i,  1901  and  3,  1905. 

385 


386         TRUE— ON  THE  CLASSIFICATION  OF  THE  CETACEA.    [April  .4 

6.  The  new  family  Saurodelphidae. 

7.  The  family  Platanistidae,  consisting  of  Platanista  only. 

The  zeuglodonts  are  included  in  the  order  Cetacea  by  the  ma- 
jority of  cetolog^sts,  though  they  were  rejected  from  the  great 
"  Osteography  "  of  Van  Beneden  and  CJervais,  as  these  authors  did 
not  consider  them  to  be  cetaceans.  Brandt  placed  them  with  the 
squalodonts  as  families  in  a  tribe  subordinate  to  the  Odontoceti,*  but 
nearly  all  other  authorities  have  considered  them  as  a  distinct  sub- 
order,— Archaeoceti,  or  Zeuglodontes. 

It  seems  to  be  generally  agreed  that  the  zeuglodonts  have  been 
proven  by  the  researches  of  Dawes,  Fraas,  Stromer,  Andrews  and 
others  to  be  derived  from  the  creodonts.  I  do  not  know  from  what 
particular  creodont  they  are  supposed  to  have  sprung,  and  whether 
the  connection  is  good  in  that  direction  is  for  those  ftost  familiar 
with  the  creodonts  to  decide.  The  chief  argument  appears  to  be 
that  in  some  zeuglodonts  some  of  the  molars  are  three-rooted. 

Whatever^  may  be  the  truth  as  regards  that  connection,  various 
zoologists  have  proposed,  in  more  or  less  definite  terms,  to  unite  the 
zeuglodonts  to  the  ordinary  cetaceans  through  the  squalodonts, 
which  are  clearly  cetaceans,  but  with  two-rooted  or  three-rooted 
teeth  having  serrated  crowns.  Professor  Abel  advances  the  con- 
crete proposition  of  uniting  the  zeuglodonts  and  squalodonts* 
through  the  small  form  from  the  Caucasus,  described  by  Lydekker 
under  the  name  of  Zeuglodon  caucasicus,^  and  afterward  made  the 
basis  of  a  new  genus,  Microzeuglodon,  by  Von  Stromer.  Of  this 
only  a  part  of  the  lower  jaw,  the  humerus  and  a  caudal  vertebra,  are 
known.  The  upward  turn  of  the  superior  margin  of  the  jaw  pos- 
teriorly, and  the  form  of  the  humerus — particularly  the  quite  good 
articular  facets, — ^appear  to  me  to  indicate  that  this  is  a  zeuglodont, 
with  no  very  strong  leaning  toward  Squalodon,  If  this  be  con- 
ceded, there  is  no  way  at  present  in  which  to  connect  the  Cetacea 
with  any  group  of  land  mammals. 

I  would  point  out  in  this  connection  that  while  Microzeuglodon 
is  from  the  Eocene  and  is  of  small  size,  and  Squalodon  is  from  the 

*Mitn.  Acad.  Imp.  Set.  St.  Petersburg,  VII*  Serie,  XX,  1873,  P-  vii. 

•L.  c,  p.  34. 

*  Proc.  Zool.  Soc.  London,  1892,  p.  558,  pi.  36. 


,9o8]         TRUE— ON  THE  CLASSIFICATION   OF  THE   CETACEA.  387 

Miocene  and  Pliocene  and  is  of  comparatively  large  size,  there  is 
an  American  form  of  squalodont  which  is  either  from  the  Oligocene 
or  Lower  Miocene,  and  is  of  small  size. 

This  is  the  genus  Agorophius.  It  is  based  on  a  skull  from  South 
Carolina.  It  has  serrate  teeth  like  Squalodon,  but  what  is  especially 
remarkable,  the  parietals  occupy  a  long  area  on  the  top  of  the  skull, 
while  in  Squalodon  and  existing  cetaceans  the  f  rontals  and  occipital 
come  together  at  the  vettex  so  as  to  entirely,  or  almost  entirely, 
exclude  the  parietals."  The  very  remarkable  conformation  of 
Agorophius  led  Van  Beneden  and  Gervais,  and  also  Cope,  to  suspect 
that  it  might  possibly  be  the  progenitor  of  the  whalebone  whales.  I 
do  not  think  this  is  likely,  but  Agorophius  appears  to  indicate  that 
Squalodon  may  have,  and  probably  did,  originate  from  forms  very 
unlike  Zeuglodon, 

It  might  be  supposed  that  the  whole  argument  concerning  the 
derivation  of  the  Cetacea  from  the  zeuglodonts  was  negatived  by 
the  occurrence  of  various  characteristic  forms  of  Cetacea  in  the 
Eocene  and  even  earlier  formations,  and  hence  contemporaneously 
with,  or  earlier  than,  Zeuglodon,  In  all  such  cases,  however,  so  far 
as  I  have  traced  them,  the  forms  reported  are  really  from  the  Mio- 
cene. A  notable  case  is  that  of  the  various  important  forms  from 
Chubut,  Patagonia,  described  by  Lydekker  in  1893.  These  include 
such  genera  as  Scaldicetus  and  Paracetus,  which  certainly  occur  in 
the  Miocene  of  North  America  and  Europe,  and,  indeed,  I  under- 
stand the  deposits  at  Chubut  to  be  assigned  at  present  without  dis- 
pute to  the  Miocene. 

The  matter  of  the  history  and  development  of  Squalodon  is 
especially  important,  as  Professor  Abel  derives  four  families  of 
cetaceans  from  the  squalodonts,  namely,  Physeteridae,  Ziphiidae, 
Eurinodelphidae  and  Acrodelphidae  (or  Iniidae),  and  one  of  them — 
the  Physeteridae — directly  from  Squalodon  itself.  The  main  argu- 
ment in  the  latter  case  is  that  the  teeth  of  some  species  of  Scaldi- 
cetus  (or  Physodon) — an  intermediate  genus — have  a  ridge  on  the 
crown.  This  seems  an  unimportant  character  relatively,  and  does 
not  balance  the  difficulty  of  deriving  the  extremely  concave  skull  of 
Physeter  from  the  extremely  flat  skull  of  Squalodon, 

•See  True,  "Remarks  on  the  Type  of  the  Fossil  Cetacean  Agorophius 
Pygmaus  (Miiller),"  Smithsonian  Publ.,  No.  1694,  1907,  with  i  plate. 


388  TRUE— ON   THE  CLASSIFICATION   OF  THE  CETACEA.     [April  24 

I  think  that  we  shall  in  the  end  come  to  agree  with  the  c^inion 
expressed  many  years  ago  by  Dr.  Theo.  Gill,*  that  the  origin  of  the 
Cetacea  dates  much  further  back  than  is  generally  believed,  and  that 
the  forms  above  mentioned  are  sideshoots  from  a  stem  reaching  into 
a  much  more  remote  past. 

However  it  may  be  as  to  the  origin  of  the  families  mentioned, 
Professor  Abel  is  correct,  I  believe,  in  following  the  course  of  Gray^ 
and  Gill'  in  separating  the  sperm  whales  and  the  beaked  whales  into 
two  families,  the  Physeteridae  and  the  Ziphiidae.  Abel's  line  of  de- 
velopment for  Physeter  through  Scaldicetus,  Physeterula,  Prophy- 
seter  and  Placoziphius  seems  excellent,  except  that  it  ignores  Hypo- 
cetus  Lydek.  (or  Diaphorocetus  Amegh.)  of  North  and  South 
America,  which  is  certainly  an  ancestor  of  Physeter  or  Kogia,  and 
probably  the  former. 

The  family  Eurinodelphidae  of  Abel  is  quite  certainly  distinct. 
While  obviously  allied  to  the  Ziphiidae,  Eurinodelphis  has  distinc- 
tive characters  of  its  own,  such  as  the  small  pterygoids,  very  long 
toothless  premaxillae,  a  delphinoid  prenarial  region,  etc.  I  suc- 
ceeded in  discovering  a  skull  of  this  genus  in  the  Miocene  of  Mary- 
land last  year  and  thus  introducing  the  family  into  the  American 
fauna. 

Abel's  family  Acrodelphidae,  which,  as  Professor  Eastman  re- 
cently pointed  out,  should  be  called  Iniidae®,  while  not  entirely  new, 
is  a  very  interesting  assemblage.  It  comprises  the  following  sub- 
families and  genera : 

Family  Iniid^  Gill  (Acrodelphidae  Abel). 

Argyrocetus, 


Argyrocetinae 


Acrodelphinae 


Cyrtodelphis, 
Pontivaga, 
Ischyorhynchus, 
Champsodelphis. 

fAcrodelphis. 
Heterodelphis. 


*  Amer.  Nat.,  7,  1873,  P-  2. 

*Cat.  Seals  and  Whales  Brit.  Mus.,  2d  ed.,  1866,  p.  326. 
•Smithsonian  Misc.  Coll.,  11,  1872,  p.  15. 

*  Bull.  Mus.  Comp.  Zool,  51,  1907,  p.  86. 


i9o8]         TRUE— ON   THE  CLASSIFICATION   OF  THE  CETACEA.  389 


Iniinae 


Pontistes. 
Stenodelphis. 


Beluffinae  I  Beluga, 

The  partial  breaking  up  of  the  currently-accepted  families  Plat- 
anistidae  and  Delphinidae  here  shown  is  quite  radical.  Usually  Plaia- 
nista,  Inia  and  Stenodelphis  (the  so-called  "  river-dolphins  ")  are 
united  to  form  the  family  Platanistidae,  but  Professor  Abel  leives 
only  the  genus  Platcmista  in  that  family.  The  limits  of  the  family 
have  always  been  uncertain,  and  Sir  Wm.  Flower,  though  accept- 
ing it  provisionally  in  its  usual  form,  remarked :  "  There  are  three 
distinct  genera,  which  might  almost  be  made  the  types  of  families, 
but  it  is  probably  more  convenient  to  keep  them  together,  only  regard- 
ing them  as  representing  three  subfamilies."*® 

Stenodelphis,  although  having  separate  cervicals  and  broad  lum- 
bar diapophyses  like  Inia,  has  involuted  pterygoids,  ossified  sternal 
ribs,  and  the  articulations  of  the  ordinary  ribs  with  the  vertebrae  as 
in  Delphinidae.  Associated  with  it  is  the  fossil  genus  Pontistes  of 
South  America,  which  resembles  Stenodelphis  very  closely,  but  is 
larger.  The  prenarial  region  in  these  genera,  as  well  as  the  form 
and  position  of  the  nasals  and  the  form  of  the  zygomatic  processes, 
recall  Phoccena  and  also  Inia,  but  I  have  been  unable  to  satisfy  my- 
self of  the  importance  of  these  resemblances. 

The  most  radical  feature  of  Professor  Abel's  classification  is 
the  removal  of  the  white  whale  and  narwhal  (Delphinapterus  and 
Monodon)  from  the  Delphinidae  to  the  Iniidae,  although  it  is  true 
tha^  these  forms  had  previously  been  considered  as  constituting  a 
separate  subfamily  of  the  Delphinidae  by  Gill,  Flower  and  myself. 
They  agree  with  Inia  in  having  no  dorsal  fin,  a  broad  pectoral^  and 
separate  cervical  vertebrae,  and  the  diapophyses  of  the  lumbars  are 
somewhat  expanded.  On  the  other  hand,  the  sternal  ribs  are  ossi- 
fied, the  sternum  is  shaped  as  in  other  Delphinidae,  the  ribs  articu- 
late with  the  vertebrae  in  the  same  manner  as  in  that  family,  and 
the  enamel  of  the  teeth  is  smooth.     This  combination  of  charac- 

*•  Flower  and  Lydekker,  "  Mammals  Living  and  Extinct/*  1891,  p.  258. 


390         TRUE— ON  THE  CLASSIFICATION   OF  THE  CETACEA.    [April  m 

ters    recalls    Stenodelphis  rather  than  Inia,  although    the    former 
has  a  dorsal  fin. 

Professor  Abel's  chief  reason  for  rejecting  Delphinapterus  and 
Monodon  from  the  Delphinidae  appears  to  be  that  the  cervical  ver- 
tebrae are  separate.  He  says  that  on  this  account  they  cannot  be 
derived  from  Delphinidae.*^  This  seems  to  me  illogical,  for  it  must 
be  true  that  the  existing  Delphinidae  with  extremely  thin,  more  or 
less  rudimentary,  and  anchylosed  cervicals  were  derived  from  forms 
with  well-developed,  separate  cervicals.  Hence,  one  might  expect 
to  find  some  forms  still  existing  in  which  the  cervicals  are  distinct 
I  do  not  think  that  on  that  account  alone  they  should  be  rejected 
from  among  die  Delphinidae. 

In  this  connection,  the  genus  Lophocetus  from  the  Miocene  of 
Maryland  is  of  interest.  This  is  represented  by  a  skull  and  cervical 
vertebrae.  The  skull,  which  is  long-beaked,  is  delphinoid  in  general 
appearance,  especially  in  the  prenarial  region,  but  the  temporal 
fossae  are  large  and  the  supraoccipital  narrow,  and  shaped  somewhat 
as  in  Inia,  The  teeth  are  lacking,  but  appear  to  have  had  simple 
cylindrical  roots.  The  cervical  vertebrae  are  separate.  They  are, 
however,  imbedded  in  the  matrix,  so  that  little  can  be  determined 
regarding  their  characters. 

This  genus  has  been  associated  with  Inia  in  the  Platanistidae 
by  Cope;**  and  Dr.  C.  R.  Eastman,  who  has  recently  given  a  new 
description  of  it,*'  also  regards  it  as  allied  to  Inia,  while  Brandt 
and  Abel  have  considered  it  closely  allied  to  Delphinapterus.  I  am\ 
myself  inclined  to  the  latter  view,  although  conceding  that  the  shape 
of  the  supraoccipital  is  inioid.  H  this  be  accepted,  we  have  in 
Lophocetus  a  Miocene  delphinoid  form  with  separate  cervicals. 

On  account  of  the  combination  of  characters  presented  by  Steno- 
delphis,  Delphinapterus,  Monodon  and  Lophocetus,  three  courses 
are  possible  as  regards  their  classification.  They  may  be  included 
in  the  family  Iniidae,  or  made  the  basis  of  a  separate  family  Steno- 
delphidae,  or  included  in  the  family  Delphinidae.  The  latter  course 
seems  to  me  best  at  present, 

^Mim.  Mus.  Roy,  Hist.  Nat,  Belgique,  3,  1905. 
^Amer.  Nat,,  1890,  pp.  606  and  615. 
^Bull.  Mus.  Comp,  Zool,  51,  1907,  p.  79. 


i9o8]  TRUE— ON   THE  CLASSIFICATION  OF  THE  CETACEA.  391 

Professor  Abel  has  described  a  delphinoid  form  from  the  Upper 
Miocene  of  Antwerp — Pithanodelphis — in  which  the  atlas  and 
axis  are  united  as  in  existing  genera.  It  would  appear,  from  this 
and  other  evidence,  that  the  family  Delphinidae  was  differentiated 
^  as  early  as  the  Miocene  and  that  both  forms  with  separate  cervicals 

and  forms  with  united  cervicals  were  then  existing. 

The  family  Saurodelphidae  of  Abel  comprises  the  single  genus 
Saurodelphis  Burmeister,  from  the  banks  of  the  Parana  River, 
Argentina.  The  geological  horizon  is  understood  to  be  Pliocene. 
Professor  Abel  considers  that  it  cannot  be  associated  at  present 
with  any  group  of  toothed  whales,  but  it  appears  probable  from 
Burmeister's  figures  that  the  skull  has  a  maxillary  hood  and  other 
characters  resembling  those  of  Platanista,  and  the  teeth  are  also 
similar  in  some  respects,  especially  as  regards  the  growth  of  irreg- 
ular roots  with  age,  etc.  For  these  reasons,  I  think  it  should 
be  assigned  to  the  Platanistidae,  at  least  provisionally. 

The  modifications  which  I  have  proposed  in  the  classification 
of  the  toothed  whales  are  summed  up  as  follows : 

Cetacea. 
Odontoceti. 
+  Squalodontidae. 

^,  . ,  f  Physeterinae. 

Physetendae \  ^^     .. 

•^  ( Kogiinae. 

Ziphiidae. 

+  Eurinodelphidae. 

'  Iniinae. 

Iniidae -I  Argyrocetinae. 

Acrodelphinae. 

r  Stenodelphinae. 
Delphinidae J  Delphinapterinae. 

[  Delphininae. 
Eurinodelphidae. 


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JAMES  W.  HOLLAND 

AMOS  P.  BROWN 

Secre/aries, 


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Vol.  XLVIl  Sept.-Dbc.,  1908.  No.  190. 

HEREDITY,  VARIATION  AND  EVOLUTION  IN 

PROTOZOA.    II. 

Heredity  and  Variation  of  Size  and  Form  in  Paramecium, 
WITH  Studies  of  Growth,  Environmental  Action 

AND  Selection.* 

By  H.  S.  JENNINGS. 

(Read  April  24,  igo8.) 

TABLE  OF  CONTENTS. 
(See  pages  544-546.) 

I.   INTRODUCTORY. 

The  first  of  this  series  of  studies^  gave  a  general  introduction  to 
the  investigations,  and  dealt  with  the  fate  of  new  or  acquired  char- 
acters in  protozoa,  showing  that  these  are  as  a  rule  not  inherited  and 
that  there  is  no  difference  in  principle  on  this  point  between  protozoa 
and  metazoa.  The  present  paper  takes  up  heredity  and  variation  in 
size  and  form  in  Paramecium. 

Our  present  questions  are  then  mainly  as  follows :  In  what  respects 
do  the  individuals  of  Paramecium  resemble  each  other?    In  what 

*From  the  Laboratory  of  Experimental  Zoology,  Johns  Hopkins  Uni- 
versity, Baltimore,  Md. 

*  Journal  of  Experimental  Zoology,  Vol.  5,  1908,  pp.  577-632. 

PROG.  AMER.  PHIL.  SOC  XLVII.  I90  Z,  PRINTED  JANUARY  8,  I909. 


3i>4  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  34, 

respects  do  they  diifer?  What  are  the  causes  of  the  resemblances 
or  differences,  as  the  case  may  be? 

The  attempt  is  made  to  treat  these  questions  broadly,  determin- 
ing experimentally  the  different  classes  of  causes  concerned,  without 
prejudice  as  to  their  relative  importance.  External  and  internal  fac- 
tors are  therefore  equally  considered,  the  purpose  of  the  investiga- 
tion being  to  give  as  complete  an  analysis  of  the  phenomena  of 
resemblances  and  differences  as  possible.  Our  problem,  then,  requires 
an  analysis  from  this  point  of  view  of  all  things  which  may  result 
in  producing,  increasing  or  decreasing  the  similarities  and  differences 
between  individuals — reproduction,  growth,  conjugation,  the  effects 
of  environment,  of  selection,  and  the  like. 

The  investigation  will  be  best  introduced  by  proposing  at  once 
what  is  really  the  central  problem — that  concerning  heredity.  Is 
size  inherited  in  Paramecium? 

How  would  heredity  of  size  be  shown?  If  certain  individuals 
differ  in  size,  and  the  progeny  of  these  individuals,  under  identical 
conditions,  show  corresponding  differences,  this  is  what  would  com- 
monly be  called  heredity  of  size.  "  Heredity  is  a  certain  degree  of 
correlation  between  the  abmodality  of  parent  and  off  spring  "  (Daven- 
port, 1899,  p.  35 ).  Do  large  individuals  of  Paramecium  produce, 
under  the  same  conditions,  larger  progeny  than  do  small  ones?  Is 
it  possible  to  obtain  by  selection  large  and  small  races  of  Parameciaf 

To  study  this  question,  we  must  first  examine  the  variations  in 
size  commonly  found  in  Paramecium, 

II.    PRELIMINARY  STUDY  OF  VARIATION  IN 

PARAMECIUM. 

We  owe  our  present  knowledge  of  variation  in  Paramecium 
mainly  to  Pearl  and  his  co-workers  (see  Pearl,  1907;  Pearl  and 
Dunbar,  1905).  A  more  extensive  work  by  Pearl  on  variation  in 
Paramecium  has  been  mentioned  as  in  prospect ;  I  learn  from  per- 
sonal communication,  however,  that  this  is  not  to  appear.  I  shall 
therefore  publish  my  own  results  more  fully  than  I  should  otherwise 
have  done.  Certain  points  in  connection  with  variation  in  Para- 
mecium have  been   dealt  with  by  Simpson    (1902)   and   Pearson 


,9o8.]  JENNINGS— HEREDITY   IN    PROTOZOA.  395 

( 1902)  ;  also  by  McClendon  ( 1908) .    But  we  have  at  present  nothing 
like  a  thorough  analysis  of  the  matter,  based  on  extensive  data. 

I.   General  Methods  of  Work;  Statistical  Treatment  and 

Its  Uses. 

Before  we  can  study  experimentally  the  nature  and  causes  of  the 
existing  variations,  we  must,  of  course,  know  their  extent,  character 
and  distribution.  To  this  end  I  have  made  a  statistical  study,  con- 
structed frequency  polygons,  and  determined  the  more  important 
constants  of  variation  and  correlation.  This  has,  of  course,  not  been 
done  because  of  belief  in  any  occult  virtue  in  mathematical  treat- 
ment. Statistical  methods  have  been  used  in  this  preliminary  survey 
merely  because  they  form  the  most  natural  and  direct  way  of  discov- 
ering and  displaying  the  problems  on  which  we  wish  to  work ;  I  doubt 
whether  the  most  determined  critic  of  the  use  of  such  treatment  in 
biology  could  suggest  any  other  way  for  our  material.  But  I  am 
fully  convinced  that  "crucial  evidence  is  always  individual  in  the 
last  analysis  "  (Whitman)  ;  that  the  preHminary  statistical  examina- 
tion of  the  facts  requires  development  as  soon  as  possible  into  precise 
experimental  knowledge.  It  is  valuable  to  know  just  how  many 
men  out  of  a  thousand  will  die  in  a  given  period,  but  it  is  infinitely 
more  valuable  to  know  which  ones  will  die  if  the  conditions  are  not 
changed,  and  why;  and  the  latter  knowledge  includes  the  former. 
I  have  therefore  advanced  at  once  from  the  descriptive  statistical 
work  to  experimental  treatment.  A  curve  or  polygon  of  variation 
(such  as  Diagram  i)  or  a  correlation  table  (such  as  Table  I.)  is  to 
be  looked  upon  as  a  mass  of  problems.  The  place  occupied  in  the 
polygon  or  table  by  any  individual  is  due  to  certain  causes,  and  it  is 
these  causes  that  we  seek. 

In  seeking  these  causes  by  experimental  methods,  statistical 
treatment  is  again  found  to  be  of  the  greatest  value  for  detecting 
and  registering  the  effects  of  single  factors,  under  complex  condi- 
tions. This  method  may  be  compared  to  a  microscope;  it  enables 
us  to  detect  and  deal  with  causes  and  effects  which  we  could  not 
handle  without  it.  I  am  convinced  that  it  is  a  great  mistake  to  hold 
that  the  only  or  the  main  use  of  statistical  treatment  is  for  "  dealing 


396  JENNINGS— HEREDITY  IN   PROTOZOA.  [April  24, 

with  the  sphere  of  indefinitely  numerous  small  causes — amenable 
only  to  the  calculus  of  chance,  and  not  to  any  analysts  of  the  indir 
victual  instance."  Such  treatment  is  a  most  valuable  instrument  for 
precisely  such  analysis  as  will  bring  out  the  effects  of  individual 
factors  when  we  are  unable  to  experimentally  disengage  them  com- 
pletely from  others;  it  aids  us  most  essentially  in  the  "analysis  of 
the  individual  instance."  Of  this  I  hope  the  present  paper  may  fur- 
nish illustrations.  As  Johannsen  (1906,  p.  98)  has  well  expressed 
it,  the  mathematical  treatment  must,  to  give  valuable  results,  be 
"  based  upon  an  accomplished  sorting  of  the  special  facts  and  a 
biological  setting  out  of  the  premises  which  are  to  be  treated." 
Davenport  (1899)  states  that  "the  statistical  laws  of  heredity  deal 
not  with  the  relations  between  one  descendant  and  its  parent  or 
parents,  but  only  with  the  mean  progeny  of  mean  parents."  The 
object  of  the  present  work  is  precisely  to  discover  so  far  as  possible 
the  relation  between  one  descendant  and  its  parent  (or  other  rela- 
tives) ;  for  this,  stat;istical  methods  show  themselves  most  useful. 

2.  A  Typical  Culture. 

We  will  then  first  examine  a  typical  culture  of  Paramecium,  made 
in  the  usual  way  with  pond  water  and  decaying  vegetation,  in  a 
circular  glass  vessel  about  nine  inches  across  and  three  inches  deep. 
This  culture  we  will  call  Culture  i. 

Inspection  showed  that  Paramecia  of  markedly  different  size 
were  found  in  this  culture,  so  that  it  seemed  a  favorable  one  for  a 
study  of  inheritance  in  size.  Cursory  examination  seemed  to  indi- 
cate the  existence  of  two  sets  of  individuals,  those  of  one  set  being 
nearly  double  the  length  of  the  others. 

Of  this  culture  a  large  number  were  killed  on  April  10,  1907,  and 
four  hundred  specimens,  taken  at  random,  were  measured  as  to 
length  and  breadth. 

3.   Methods  of  Measuring  and  Recording. 

The  animals  were  killed  with  Worcester's  fluid,  which  is  known  to  cause 
practically  no  distortion  when  properly  used.  Worcester's  fluid  consists  of 
ten  per  cent  formalin  saturated  with  corrosive  sublimate.  In  using  it,  a 
large  number  of  the  infusoria  must  be  brought  into  one  or  two  drops  of 


,9o8.j  JENNINGS— HEREDITY   IN  PROTOZOA.  397 

water,  then  these  must  be  overwhelmed  with  a  considerable  quantity  of  the 
fluid.  If  the  infusoria  are  in  a  larger  quantity  of  water,  the  killing  takes 
place  more  slowly,  the  animals  have  time  to  contract,  and  distortion  results. 

The  measurements  were  made  on  the  slide,  the  organisms  being  either 
still  in  the  killing  fluid  or  in  ten  per  cent,  formalin.  Transference  to  the 
latter  has  no  effect  on  the  form  of  the  fixed  animals.  Most  of  the  meas- 
urements were  made  directly  with  an  ocular  micrometer.  In  the  case  of 
cultures  of  large  individuals,  however,  the  form  was  projected  on  paper  with 
the  camera,  in  the  way  described  by  Pearl  (1907),  the  extremities  of  length 
and  breadth  marked  with  the  pencil,  then  these  were  measured  with  a  scale 
made  by  projection  of  the  ocular  micrometer. 

Such  combinations  of  lenses  were  used  that  one  division  of  the  microm- 
eter scale  was  equal  to  4  microns  (or  in  a  few  cases,  which  will  be  expressly 
noted,  to  3!  microns).  The  measurements  were  thus  recorded  in  units,  each 
of  which  was  equal  to  4  microns,  so  that  the  recorded  units  are  multiplied 
by  four  to  give  results  in  microns.  When  the  measurements  fell  between  two 
lines  of  the  micrometer,  the  line  nearest  the  actual  measure  was  that  re- 
corded; if  the  measurement  fell  just  half  way  between  two  lines,  the  higher 
line  was  recorded.  Thus,  the  recorded  unit  45  included  all  measurements 
beginning  with  44^,  and  less  than  45^.  In  the  tables,  the  measurements, 
given  in  microns,  are  therefore  grouped  about  such  values  that  each  group 
includes  values  from  two  microns  below  to  two  microns  above  the  one 
recorded.  Thus,  in  Table  i,  the  length  180  includes  all  the  specimens  meas- 
uring from  178  up  to  (but  not  including)  182. 

It  will  be  well  to  summarize  here,  once  for  all,  the  method  of  treating 
the  data  obtained  in  the  measurements.  For  most  of  the  tables  the  con- 
stants computed  (and  recorded  below  the  tables)  were  the  following;  the 
mean,  standard  deviation,  and  coefficient  of  variation,  for  length  and  for 
breadth;  the  mean  index  or  ratio  of  breadth  to  length;  and  the  coefficient 
of  correlation.  The  computation  of  the  constants  was  based  on  the  well- 
known  formulae  that  have  been  brought  together  by  Davenport  (1904)  and 
others.  I  used  as  a  rule  the  actual  methods  set  forth  so  clearly  by  Yule 
(1897).  The  computations  were  made  by  the  aid  of  seven-place  logarithms 
and  of  Crelle's  and  Barlow's  tables.  Two  independent  computations,  at 
considerable  intervals  of  time,  were  made  in  each  case.  While  I  cannot 
hope  that  errors  in  computation  are  excluded,  I  believe  that  such  as  may 
exist  do  not  in  any  way  affect  the  conclusions  to  be  drawn. 

Certain  points  of  detail  should  be  mentioned.  While,  as  will  appear, 
most  of  the  tables  do  not  give  symmetrical  curves,  I  have  used  only  the 
simple  statistical  methods  applicable  in  strictness  to  such  curves;  the  methods 
are  quite  sufficient  as  a  basis  for  the  comparisons  we  wish  to  make. 

In  computing  the  standard  deviation,  Sheppard's  correction  of  the 
second  moment  was  used  throughout.  That  is,  if  we  employ  the  method 
of  Yule  (1897), 

a^  l/2(7^«y-rf'«-. 08333. 
or  using  the  signs  employed  by  Davenport  (1904) 


398 


JENNINGS— HEREDITY   IN  PROTOZOA. 


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,9o8.J  JENNINGS— HEREDITY   IN   PROTOZOA.  399 

The  mean  index  given  below  the  tables  is  the  mean  of  the  quotient 

breadth 

r— :  it  shows  essentially  what  percentage  the  breadth  is  of  the  length. 

length 

This  mean  was  found,  without  computing  the  index  for  each  individual,  by 

the  following  formula: 

,  =  ^(i-hC2.«-rC2rCi). 

Where  t  is  the  mean  index,  As  is  the  mean  breadth,  Al  the  mean  length, 
Cb  the  coefficient  of  variation  for  breadth,  Cl  the  same  for  length,  and  r  is 
the  coefficient  of  correlation  between  length  and  breadth. 

I  am  greatly  indebted  to  Dr.  Ra3rmond  Pearl  for  assistance  in  the  mathe- 
matical treatment  of  the  data. 

The  results  of  the  measurements  of  a  random  sample  of  400  of 
Culture  I  are  given  in  Table  I. 

It  is  evident  on  inspection  of  this  table  that  the  individuals  fall 
into  two  well-marked  groups,  one  set  varying  in  length  from  84  to 
144  microns,  the  other  set  varying  from  164  to  240  microns,  while 
between  these  groups,  in  the  region  from  144  to  164  microns,  only 
two  specimens  are  found.  The  mean  length  for  the  entire  sample 
falls  at  165.840  microns,  almost  precisely  in  the  region  where  no 
specimens  are  found.  The  smaller  set  have  their  mean  length  at 
125.420  microns :  the  larger  set  at  200.972  microns. 

These  results  are  shown  as  frequency  polygons  in  the  lower  por- 
tions of  piagrams  i  and  2. 

4.   Method  of  Constructing  the  Polygons. 

In  making  the  polygons  for  length,  three  units  of  measurement  (12 
microns)  were  grouped  together  to  make  a  single  unit  of  the  abscissa  of  the 
polygon.  This  was  done  in  order  to  destroy  any  irregularities  due  to  un- 
conscious prejudice  on  the  part  of  the  measurer  for  certain  numbers.  Thus, 
in  measuring  a  large  number  of  individuals,  it  may  be  found,  for  example, 
that  few  are  recorded  at  51,  while  at  50  there  are  many;  or  the  reverse  may 
occur.  This  is  due  only  to  the  fact  that  in  doubtful  cases  falling  between 
these  numbers  the  measurer  unconsciously  gives  the  preference  regularly  to 
one  of  them.  The  error  thus  introduced  is  extremely  small  (it  can  ^hardly 
be  more  than  one  micron  in  any  case),  but  if  the  polygon  is  made  without 
grouping  together  adjacent  classes,  there  appear  extreme  irregularities  in 
its  outline,  irregularities  that  are  quite  without  significance.  When  three 
units  are  thrown  together,  any  marked  irregularities  remaining  in  the  poly- 
gons are  almost  certainly  due  to  peculiarities  in  the  material  itself.  It  is 
of  course  possible  that  small  peculiarities  really  existing  may  be  hidden  in 


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Length  in  Microns. 

Diagram  i.  Polygons  of  variation  for  length  in  Culture  i  and  its  de- 
scendants. A  and  a  form  together  the  polygon  for  400  specimens  taken  at 
random  from  the  original  culture  i,  on  April  10,  1907.  B,  polygon  for  100 
descendants  of  ten  of  the  larger  individuals  of  Culture  i.  D,  polygon  for 
100  descendants  of  the  single  large  individual  D,  from  culture  i.  b,  polygon 
for  100  descendants  of  fifty  smaller  individuals  from  culture  i.  c,  polygon 
for  100  descendants  of  the  single  small  individual  c,  from  culture  i. 


]9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA. 


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Breadth  in  Microns. 

Diagram  2.  Polygons  of  variation  for  breadth  in  culture  i,  and  in  its 
descendants  from  selected  specimens.  The  letters  have  the  same  significance 
as  in  Diagram  i. 


402  JENNINGS— HEREDITY   IN   PROTOZOA.  fApnia*. 

this  way,  but  it  was  thought  wiser  to  be  conservative  in  this  matter.  Thus 
the  space  between  two  perpendicular  lines  of  the  polygons  includes  three 
of  the  groups  of  the  correlation  table,  and  is  marked  at  its  base  with  the 
middle  value  of  the  three  groups  which  it  includes. 

In  making  the  polygons  for  breadth,  it  was  found  that  there  was  little 
evidence  of  error  due  to  unconscious  preference  for  certain  numbers  in 
making  this  measurement.  This  is  probably  due  to  the  comparatively  small 
numbers  of  units  in  the  breadth  measurement,  and  to  the  fact  that  it  is 
possible  to  hold  both  limits  of  the  measurement  on  the  scale  sharply  in  the 
eye  at  once,  while  this  is  hardly  possible  in  measuring  length.  In  the  poly- 
gons for  breadth,  therefore,  one  unit  of  the  polygon  was  made  to  correspond 
to  one  unit  of  measurement  (four  microns). 

In  all  the  polygons  the  numbers  to  the  left  indicate  percentages  of  the 
entire  number,  so  that  all  the  polygons  are  of  equal  area,  whatever  the 
number  of  specimens  on  which  they  are  based.  The  only  exception  to  this 
is  in  the  case  of  the  double  polygons  a  and  A,  of  Diagram  i,  resulting  from 
plotting  the  random  sample  of  Table  I.  Since  this  sample  falls  into  two 
groups,  the  entire  (double)  polygon  was  made  of  fwice  the  area  of  the 
other  polygons.  Each  half  polygon  therefore  becomes  approximately  equal 
to  any  one  of  the  single  polygons  of  the  other  diagrams,  thus  permitting 
ready  comparison. 

The  numbers  at  the  foot  of  the  diagrams  are  the  dimensions  in  microns. 
Each  number  corresponds  to  the  value  of  the  center  of  the  column  beneath 
which  it  stands. 

5.   Two  Groups  of  Paramecia. 

Thus  the  Paramecia  in  our  natural  culture  I  fall  into  two  groups 
which  are  almost  completely  separated,  so  far  as  length  is  concerned, 
but  which  overlap  a  certain  amount  in  breadth.  Characteristic  out- 
lines of  varied  members  of  the  two  groups,  drawn  to  the  same  scale, 
are  shown  in  Fig.  i. 

Are  these  two  groups  permanent  differentiations,  such  as  might 
be  called  distinct  species,  or  are  the  differences  possibly  due  merely 
to  temporary  dimorphism  of  some  sort?  To  answer  this  question 
individuals  of  the  two  sizes  were  isolated  and  allowed  to  multiply 
separately,  in  cultures  made  of  boiled  hay.  After  varying  periods 
of  time  100  individuals,  taken  at  random,  were  measured  from  each 
of  these  pure  cultures,  and  the  frequency  polygon  derived  from  these 
was  compared  with  the  two  (nearly  distinct)  polygons  from  the 
original  culture.    The  following  cultures  were  made  and  measured : 

I.  Fifty  of  the  smaller  individuals  were  selected  from  the  orig- 
inal culture,  placed  together,  and  allowed  to  multiply  for  twelve  days 
(from  April  10  to  April  22),    The  measurements  of  100  of  this 


1908.J 


JENNINGS— HEREDITY   IN   PROTOZOA. 


403 


Fig.  I.  Outline  of  characteristic  specimens  from  the  original  wild  culture 
I,  April  10,  1907.  The  upper  row  shows  examples  of  the  larger  "  caudatum 
form";  the  lower  row  examples  of  the  smaller  " aurelia  form."  d,  Young 
of  the  caudatum  form ;  h,  dividing  specimen  of  the  aurelia  form.    All  X  235. 

culture  are  shown  in  curve  b  (broken  line),  Diagrams  i  and  2;  their 
dimensions  are  given  in  the  correlation  Table  II.  It  is  evident  that 
this  group  corresponds  in  a  general  way  with  the  smaller  group  of 
the  original  culture,  though  its  mean  length  and  breadth  are  some- 
what lower  (96.280  X  29.080  microns  instead  of  125.42  X  33-396), 
and  it  shows  a  little  less  variation. 

2.  Ten  of  the  larger  individuals  selected  from  the  original  cul- 
ture were  likewise  allowed  to  multiply  in  the  same  vessel  for  twelve 
days,  then  100  were  measured.  The  results  are  shown  in  curve  B, 
Diagrams  i  and  2,  and  in  the  correlation  Table  III.     It  is  evident 


404 


JENNINGS- HEREDITY   IN   PROTOZOA. 


[April  94, 


Table  II. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  from  De- 

cendants  of  50  of  the  Smaller  Individuals  from  Culture  i,  allowed  to 

Multiply  for  12  Days.     {See  polygons  b,  Diagrams  i  and  2.) 


en 

C 

o 
•—  20 

S  24 

.S  28 

A  32 
^  36 

ki 

PQ 


Length  in  Microns. 
80  84  88  92  96  100  104  108  112  116  120  124  128 


I 

3 
6 


2 

4 
10 

9 


5 
8 

8 


5 

4 
I 


I 
I 

3 
3 


I 
2 

3 


I    o 


10   14    25 
96.280  ±  .S52/i 


21    10     8      6 


3 

16 

I 

40 

I 

33 

I 

8 

I 

2 

100 

Length — Mean, 

St.  Dev.,  8. 160 ±.388^ 

Coef.  Var.,      7678  ±  .368 
Mean  Index,  or  Ratio  of  Breadth  to  Length,  27428  per  cent.;  Coef.  of 
Cor.,  .3768  ±  .0579. 


Breadth — Mean,  29.080  ±  .212M 

St.  Dev.  3.320  ±  .i68m 

Coef.  Var.    12.100  ±  .585 


that  the  progeny  of  these  ten  correspond  to  the  larger  set  {A)  of 
the  original  culture,  though  with  slight  differences  in  the  means  and 
in  the  amount  of  variation. 

3.  A  single  smaller  individual,  c,  was  selected  from  the  original 
culture.     As  near  as  could  be  measured  when  alive,  this  individual 

Table  III. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  from  De- 

scendants  of  10  of  the  Larger  Individuals  from  Culture  i,  allowed 

to  multiply  for  12  Days.    (See  polygons  B,  Diagrams  i  and  2.) 

Length  in  Microns. 
144  148  152  156  160  164  168  172  176  180 184  188  192  196  200  204  208  212 


•S.  48 

I 

I 

I 

I 

4 

2  5» 

I 

3 

I 

I 

I 

I 

3 

I 

I 

I 

I 

I 

16 

C  56 

I 

I 

I 

4 

I 

I 

I 

I 

3 

I 

15 

S  60 

2 

I 

I 

2 

2 

I 

8 

I 

2 

2 

I 

23 

.S  64 

2 

I 

I 

I 

I 

2 

I 

I 

I 

II 

2   68 

I 

I 

2 

3 

3 

I 

I 

2 

I 

15 

•S  7» 

I 

2 

2 

I 

2 

2 

I 

2 

13 

2  76 

I 

I 

2 

£80 

I 

I 

I 

I 

2 

5 

8 

2 

4 

7 

5 

7 

8 

12 

12 

10 

6 

3 

3 

4 

100 

Length — Mean,  182.760  ±  1.096/i       Breadth— Mean,  61.360  ±  496M 

St.  Dev.,  16.264  ±    .776/i  St.  Dev.,        7-376  ±.332M 

Coef.  Var.,      8.899  ±    .428  Coef .  Var.,  11. 912  ±  .576 

Mean  Index  or  Ratio  of  Breadth  to  Length,  33.652  per  cent. ;  Coef.  Cor., 
.5288  ±  .0486. 


x9o8.] 


JENNINGS— HEREDITY   IN   PROTOZOA. 


405 


was  I20  microns  in  length.  It  was  allowed  to  propagate  in  a  culture 
free  from  all  other  Paratnecia,  from  April  9  to  June  11  (thus  a  little 
more  than  two  months).  Now  a  random  sample  gave  the  polygons 
shown  at  c.  Diagrams  i  and  2 ;  the  measurements  are  given  in  Table 
IV.     This  group  corresponds  very  closely  to  the  smaller  group  a  of 

Table  IV. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  Descended 

from  the  single  small  Individual  c,  taken  from  Culture  i  and  allowed 

to  Multiply  63  Days.    (See  Polygons  c,  Diagrams  i  and  2.) 

Length  in  Microns, 
c  104  108  112  116  120  124  128  132  136  140  144  148  152  156 


TOO 


Breadth — Mean,  36.280  ±  .260^1 

St.  Dev.,        3.880  ±  .184M 
Coef.  Var,  10.700 ±.516 

Mean  Index  or  Ratio  of  Breadth  to  Length,  27.913  per  cent. ;  Coef.  Cor., 
.5208  ±  .0492. 


Length — Mean, 

St.   Dev., 
Coef.  Var., 


130.120  ±  .628M 
9.284  ±  443M 
7.134  ±  .342 


the  original  culture,  though  with  slight  differences  in  breadth. 

4.  A  single  very  large  specimen,  D,  approximately  250  microns 
in  length,  was  isolated  from  the  original  culture  on  April  12  and 
allowed  to  propagate  freely  till  June  11  (two  months) :  100  speci- 
mens taken  at  random  then  gave  the  measurements  shown  in  the 
polygon  D,  Diagrams  i  and  2,  and  Table  V. 

Examination  of  the  polygons  and  tables  shows  that  the  two 
forms  retain  their  essential  characteristics  when  isolated  and  propa- 
gated. The  results  shown  in  the  diagrams  are  typical  of  many 
others.  I  have  kept  distinct  strains  of  each  of  these  groups  for 
periods  (at  the  present  time)  of  more  than  eighteen  months,  and 
measurements  made  at  frequent  intervals  during  that  time  show  that 
they  have  always  remained  quite  distinct. 

Thus  it  is  clear  that  these  colorless  Paratnecia  fall  into  two  dis- 
tinct  groups,  which  are  at  least  relatively  permanent.  As  is  well 
known,  two  species  of  colorless  Paratnecia  have  long  been  distin- 


406  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  24, 

Table  V. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  Descended 

from   the  Single  Large  Individual  D,   taken  from   Culture   i, '  and 

allowed  to  Multiply  60  Days.    (See  polygons  D,  Diagrams  i  and  2.) 

Length  in  Microns. 

k^  1.^  k^         k-i         ^^  a.^  k^  k«l  ^U  b^         k^  k^  k^  ^^  ^^  1-^  t—i  t_i  mm         A>  A>         Al  AA  Ml  Al  JU 


WWWCIWC*      «      c« 


28  j  I  I    I 

^-  32 1  I  II 

c  36  :  I        2        I    2  6 

2  40  I  23431  13 

-  44  1  I             3   4   2    I  I              I                         13 

'^  48   1  II                             4    I     3    5     I     2  I     I           2     I                       '23 

.5  52  I     I           3          2    2    3    2  2     I                             I    I    18 

^56'  I                 12                22II2I  I    13 

^  60   I  12     1  I                      is 

g  64   I  I  I     I     I     I                            ,5 

i;  68  I  I  I 

!    o 

I  I     I 


72 
76 


loooooi    I    I    14211614189946262101   100 

Length— Mean,  188.360  ±  .980M  Breadth— Mean,  49.000  ±  .548M 

St.    Dev.,       14.532  ±  .692M  St.  Dev.,        8.144  ±  .388/:* 

Coef.  Var.,      7.715  ±  .370  Coef.  Var.,  16.618  ±  .814 

Mean  Index,  26.029  per  cent. ;  Coef.  Cor.,  .4188  ±  .0556. 

guished  under  the  names  Paramecium  aurelia  Miiller  and  Para- 
mecium caudatum  Ehr.  The  two  groups  we  have  found  correspond 
to  the  descriptions  heretofore  given  of  the  two  species,  the  sijialler 
set  representing  Paramecium  aurelia,  the  larger  Paramecium  cau- 
datum. Besides  the  diflferences  in  size  certain  other  characteristics 
have  been  held  to  distinguish  the  two  species,  and  these  distinguish- 
ing characteristics  are  evident  in  our  two  groups.  Paramecium 
aurelia  is  described  as  having  two  micronuclei  and  P.  caudatum  but 
one;  this  is  true  for  our  larger  and  smaller  groups  respectively. 
Paramecium  aurelia  is  said  to  be  more  rounded  behind,  while  P. 
caudatum  is  pointed.  In  spite  of  many  variations  in  form  within 
each  group,  it  is  clear  that  our  smaller  group  corresponds  in  this 
respect  also  with  P.  aurelia,  the  larger  one  with  P.  caudatum. 

Calkins  (1906)  has  brought  forward  evidence  tending  to  show 
that  the  supposed  distinction  into  permanently  differentiated  forms 
is  not  well  based,  so  that  there  are  not  two  species,  the  different  sizes 
being  merely  variants  of  one.     Calkins  based  his  doubts  as  to  the 


X908.J  JENNINGS— HEREDITY   IN   PROTOZOA.  407 

really  specific  distinctness  of  P,  aurelia  and  P.  caudatum  on  the  fact 
that  in  one  of  his  pedigree  cultures  of  P.  caudatum  the  number  of 
micronuclei  changed  from  one  to  two,  remained  at  two  for  many 
generations,  and  finally  changed  back  again  to  one. 

The  results  here  published  tend  to  indicate  that  the  distinction 
into  two  groups  is  not  without  some  sort  of  foundation.  But  it  will 
be  best  to  reserve  the  discussion  of  species  until  we  have  more  data 
at  hand.  We  may  temporarily  speak  of  the  smaller  set  as  the 
aurelia  group,  the  larger  one  as  the  caudatum  group.  In  a  later 
part  of  the  paper  the  question  of  distinguishing  species  will  be  taken  » 
up  in  detail,  in  the  light  of  full  data. 

6.   Are  Differences  in  Size  Hereditary  within  Each  of  the 

Two  Groups? 

We  ^ave  found  that  among  the  variations  of  Paramecium  in  size 
I  are  two  groups,  limited  by  internal  causes,  so  that  even  under  the 

same  external  conditions  they  differ  in  size;  these  two  groups  have 
heretofore  been  considered  two  species.  But  within  each  of  these 
groups  we  find  likewise  many  variations  in  size,  so  distributed,  how- 
ever, as  to  produce  a  curve  with  a  single  apex  (Diagrams  i  and  2, 
etc.).  These  variations  are  at  times  very  considerable,  as  will  be 
evident  from  an  examination  of  the  polygons  shown  in  Diagrams  3 
and  6  (pages  413,  470),  or  the  tables  numbered  VII.  (page  412)  and 
XX.  (page  466).  The  next  question  to  be  considered  is:  Are  the 
differences  in  size  within  such  a  group  hereditary?  That  is,  do  the 
differences  in  size  depend  upon  internal  conditions,  of  such  a  char- 
acter that  the  differences  will  persist  in  the  progeny,  even  when  the 
external  conditions  remain  the  same? 

The  experimental  answer  to  this  question  is  to  be  obtained  by 
isolating  individuals  of  different  size  belonging  to  one  of  the  two 
groups  (either  "aurelia"  or  "caudatum"),  allowing  these  to  mul- 
tiply and  determining  whether  the  progeny  show  differences  in  size 
corresponding  to  those  in  the  parents.  Can  we  by  selection  and 
propagation  produce  within  the  limits  of  a  single  group  races  of 
I  different  mean  size? 

j  Experiments  designed  to  answer  this  question  were  undertaken 

in  the   following  way.     As  representing  the  caudatum  group   I 


408  JENNINGS— HEREDITY  IN   PROTOZOA.  [April  a^, 

selected  the  cultures  descended  from  the  individual  D;  while  the 
progeny  of  c  represented  the  aurelia  group.  Now,  from  each  of 
these  groups  the  largest  and  smallest  individuals  were  isolated  and 
allowed  to  multiply,  under  uniform  conditions.  Thus,  the  selected 
large  and  small  individuals  of  a  given  group  were  all  progeny  of  a 
single  individual,  forming  thus  a  "  pure  line  " ;  this  fact  is  of  great 
importance,  as  the  sequel  will  show. 

A  large  number  of  experiments  gave  throughout  negative  results. 
The  progeny  of  large  and  of  small  individuals  (within  a  given  pure 
line)  showed  no  characteristic  differences  in  size.  Large  specimens 
of  the  caudatum  form  produced  progeny  on  the  whole  no  larger 
than  those  produced  by  small  specimens  of  the  same  form,  and  the 
same  was  true  in  the  aurelia  group.  In  many  experiments  a  single 
large  and  a  single  small  specimen  were  isolated,  and  their  progeny 
compared;  in  other  cases  a  number  of  large  specimens  were  placed 
together  in  one  vessel,  a  number  of  small  ones  in  another,  and  their 
progeny  compared  after  lapse  of  a  considerable  period.  Since  the 
results  of  these  experiments  were  throughout  negative,  I  will  give 
the  details  of  but  a  single  illustrative  experiment : 

On  July  27  ten  large  and  ten  small  specimens  were  selected  from 
a  lot  of  the  caudatum  group,  all  being  descendants  of  a  single  indi- 
vidual D,  The  ten  large  specimens  measured,  as  nearly  as  could  be 
determined  while  alive,  approximately  250  microns  each,  and  were 
thick  in  proportion  to  the  length.  The  ten  small  specimens  were 
about  150  microns  long,  and  were  thin.  The  two  sets  were  placed 
in  equal  quantities  of  the  same  culture  fluid. 

At  the  end  of  three  days  the  large  set  had  produced  many  indi- 
viduals. Fifty  of  these  taken  at  random  gave  a  mean  length  of 
189.040  microns,  a  mean  breadth  of  60.560  microns. 

The  smaller  individuals  did  not  increase  rapidly  and  five  of  them 
died  before  dividing,  so  that  all  the  progeny  came  from  six  indi- 
viduals. The  six  increased  in  size  before  dividing.  At  the  end  of 
three  days  there  were  twenty-one  individuals.  The  mean  length 
was  205.140  microns,  the  mean  breadth  56.570  microns. 

Thus  the  smaller  specimens  had  produced  progeny  that  were  a 
little  longer,  but  not  quite  so  broad,  as  those  resulting  from  the 
larger  set.     The  existing  differences  are  clearly  without  significance. 


x9o8]  JENNINGS— HEREDITY   IN    PROTOZOA.  409 

In  Other  cases  there  was  more  variation  in  size  among  the  dif- 
ferent sets  of  progeny  of  D,  particularly  if  the  measurements  were 
made  after  but  few  fissions  had  occurred.  But  sometimes  the 
progeny  of  the  large  specimens  were  smaller,  sometimes  larger,  than 
those  of  the  small  specimens.  On  the  whole,  both  large  and  small 
specimens  produced  progeny  of  about  the  mean  size  for  the  group, 
under  the  given  conditions. 

Thus  it  is  apparent  that  the  differences  in  size  shown  within 
such  a  polygon  as  D,  Diagram  i,  are  not  due  mainly  to  hereditary 
internal  factors.  Before  we  can  determine  with  certainty  whether 
any  such  factors  are  involved,  we  must  make  an  analysis  of  the 
variation  polygon,  determining  so  far  as  possible  the  different  fac- 
tors, external  and  internal,  which  go  to  make  it  up. 

7.   Proposed  Analysis  of  the  Polygons  of  Variation. 

Our  present  task  is  then  to  determine,  so  far  as  possible,  what 
factors  produce  such  polygons  of  variation  as  ace  shown  in  Dia- 
gram I ;  to  define  what  the  individuals  of  different  sizes  and  propor- 
tions really  are,  and  to  what  their  particular  characteristics  are  due. 

There  are  several  sets  of  problems  to  be  considered;  these  we 
may  classify  as  follows : 

1.  What  are  the  causes  and  the  significance  of  the  variations 
shown*  in  a  single  variation  polygon,  such  as  D,  Diagram  i  ?  Why, 
in  a  group  of  Paramecia  grown  under  the  same  conditions,  and 
perhaps  all  descended  from  the  same  ancestor,  do  certain  indviduals 
show  the  mean  length,  while  others  are  larger  and  others  smaller? 
Each  size  must  have  its  determining  factors. 

2.  In  different  polygons  from  Paramecia  of  the  same  general 
group  and  even  when  all  are  progeny  of  the  same  individual,  the 
mean  size  differs  much.  Thus,  in  Diagram  6  (page  470)  the  mean 
length  for  polygon  8  is  146.108  microns;  for  polygon  11  it  is  191.360 
microns,  though  both  represent  descendants  of  the  individual  D,  of 
the  caudatum  group.  What  are  the  causes  of  such  variations  in 
mean  size  among  different  sets  of  individuals  ? 

3.  In  different  sets  of  individuals  belonging  to  the  same  general 
group,  or  descended  from  the  same  individual,  the  amount  and  range 

PROC.  AMER.  PHIL.  SOC.  XLVII.  I90  AA,  PRINTED  JANUARY  8,  I909. 


410  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

of  variation  differs  much.  This  is  readily  evident  to  the  eye  on 
comparing  the  polygon  8  of  Diagram  6  and  its  correlation  table, 
XIX.  (page  466),  with  polygon  9  (Diagram  6)  and  its  table,  XX. 
In  the  former  the  length  ranges  only  from  120  to  176  microns,  and 
the  coefficient  of  variation  is  7.003,  while  in  the  latter  the  range  of 
length  is  from  120  to  220,  and  the  coefficient  of  variation  is  12.767. 
What  is  the  cause  of  these  great  differences  in  the  variation  of 
different  groups  ? 

4.  In  different  sets  belonging  to  the  same  general  group  the 
correlation  between  length  and  breadth  differs  greatly.  Thus,  in 
Table  XX.  (page  466)  the  correlation  is  high  and  positive,  a  differ- 
ence in  one  dimension  being  accompanied,  with  much  r^fularity,  by 
a  corresponding  difference  in  the  other.  In  Table  XXXI.  (appen- 
dix), on  the  other  hand,  there  is  almost  no  correlation,  while  in 
Tables  XXIX.  and  XXXII.  the  correlation  is  marked,  but  negative 
— an  increase  in  length  being  associated  with  a  decrease  in  breadth, 
and  vice  versa.  What  are  the  causes  and  significance  of  these  dif- 
ferences in  correlation  found  in  different  sets  ? 

In  dealing  with  these  questions,  there  are  three  main  sets  of 
possible  factors  to  be  examined,  2^3  follows : 

1.  Hereditary  Factors. — Some  of  the  factors  concerned  may  be 
internal  and  largely  independent  of  the  environment — so  that  the 
differences  in  size  are  hereditary.  The  existence  and  nature  of  such 
factors  form  our  main  problem,  but  they  can  be  dealt  with  only 
after  the  other  factors  are  investigated. 

2.  Growths — Some  of  .the  variations  in  size,  and  in  proportions, 
may  be  due  to  different  stages  of  growth,  so  that  this  matter  must 
be  carefully  examined. 

3.  Environmental  Influences, — It  appears  probable  that  the  dif- 
ferences in  the  means,  the  differences  in  the  range  and  amount  of 
variation,  and  in  the  correlation,  may  depend  partly  on  the  nature 
of  the  environment. 

We  shall  take  up  in  detail  these  three  sets  of  factors,  beginning 
with  growth. 


,9o8.J  JENNINGS— HEREDITY   IN   PROTOZOA.  411 

III.    GROWTH  IN  PARAMECIUM. 

One  significant  fact  was  noted  in  the  breeding  experiments 
described  in  a  previous  section.  Whenever  a  large  and  small  speci- 
men (belonging  to  a  given  group)  were  isolated  at  the  same  time, 
the  large  specimen  as  a  rule  divided  first.  Often  at  the  end  of 
forty-eight  hours  the  large  specimen  had  produced  eight  or  sixteen 
progeny,  while  the  small  specimen  had  either  not  divided  at  all,  or 
had  produced  but  a  single  pair. 

This  suggests  that  the  differences  in  size  may  be  largely  matters 
of  growth;  that  the  small  specimens  may  be  young  ones,  and  that 
the  variations  shown  in  the  frequency  polygons  may  be  largely 
growth  differences.  It  is  clear  that  a  study  of  growth  in  Para- 
mecium is  imperative  before  intelligent  work  can  be  done  with 
variation.  The  subject  of  growth  in  the  Protozoa  is  an  interesting 
one  in  itself,  so  that  this  study  will  be  made  as  thorough  as  possible 
for  its  own  sake,  as  well  as  for  the  light  it  throws  on  variation. 

Growth  was  studied  by  three  different  methods :  ( i )  By  obser- 
vation of  abnormal  specimens  bearing  localized  appendages,  noting 
the  changes  in  position  during  growth ;  (2)  by  following  the  changes 
of  form  and  size  in  living  specimens;  (3)  by  a  statistical  examina- 
tion of  the  dimensions  of  individuals  of  known  age. 

The  observations  on  growth  in  abnormal  specimens  have  been 
described  in  my  first  communication  (Jennings,  1908).  By  obser- 
vations on  the  living  specimen  it  is  not  possible  to  obtain  precise 
measurements.  It  will  be  best  therefore  to  begin  our  account  with 
the  statistical  examination,  taking  up  the  observations  on  the  living 
specimens  by  way  of  control. 

Effects  of  Growth  on  a  Variation  Polygon. 

If  our  suspicion  that  growth  differences  make  up  an  important 
part  of  the  observed  variations  in  size  of  Paramecium  is  justified, 
then  cultures  rapidly  multiplying  and  growing  should  be  more  vari- 
able than  those  that  are  stationary.  To  test  whether  this  is  true, 
two  lots  were  removed  from  a  rather  old  culture  of  descendants  of 
D,  in  which  inspection  showed  that  the  individuals  were  not  multi- 
plying rapidly.     One  of  these  lots  was  killed  at  once,  while  the  other 


412 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  84, 


was  placed  in  fresh  culture  fluid.  Twenty-four  hours  later  this 
second  set  was  found  to  be  multiplying  rapidly ;  a  portion  of  it  was 
then  killed.  The  measurements  of  the  two  lots  are  given  ih  Tables 
VI.  and  VII.,  while  the  facts  are  graphically  represented  in  the 

Table  VI. 

Correlation  Table  for  Lengths  and  Breadths  of  a  Random  Sample  from 
a  Culture  of  Descendants  of  D,  in  which  Multiplication  was  not  in 
Progress.  For  comparison  with  Table  VI J.  (See  also  Diagram  3.)  (Row 
3,  Table  XVIII.) 

Length  in  Microns. 

148  152  156 160 164 168  172  176  180  184  188  192  196  200  204  208  212 

c  32 

t3  36 

g  40 

c  ^ 
•^  48 

-  52 

S  60 

u 

I      I     0     4     3    II    17    10  15    13    9    II    10   II    5  8     6    135 

Length — Mean,           185.008  ±  .836^  Breadth— Mean,  43-556  ±  .392M 

St.  Dev.,        14.420 ± .592/i  St  Dev.,  6.748 ±.276^ 

Coef.  Var.,      7.794 ±.324,  Coef.  Var.,  15.490 db .651 

Mean  Index,  23.517  per  cent.;  Coef.- Cor.,  .5955  ±  .0375. 

Table  VII. 
Correlation  Table  for  Lengths  and  Breadths  of  a  Random  Sample  of  De- 
scendants of  D,  at  a  Time  when  Rapid  Multiplication  xvas  in  Progress. 
For  comparison  with   Table   VL      (See   also   Diagram   3.)      (Row  4, 

Table  XVIII.) 

Length  in  Microns. 


I 

2 

I 

I 

3 

I 

9 

2 

6 

5 

I 

2 

2 

2 

20 

1 

I 

I 

3 

8 

4 

4 

4 

3 

I 

3 

3 

35 

I 

I 

2 

2 

2 

3 

3 

4 

I 

3 

22 

I 

I 

3 

2 

I 

3 

4 

3 

I 

4 

2 

,  25 

- 

I 

I 

I 

I 

I 

2 

I 

I 

I 

2 

2 
2 

3 
I 

4 

;  ^5 

1  4 
5 

M       » 

H      »<4       H4       l> 

i4IH^4H4^4»irtlH^4t>4t>4 

1.^ 

^^ 

.Hl-IIHMI-IIHMI^CICICIC«C«C« 

.  32 

2 

2 

s  36 

I    I    I    4    I 

4 

I 

12           I 

17 

2  40 

I         I    I    I    2   2 

2 

3   3   5   2   3    I    I    I 

29 

.y  44 

I 

111111114 

2 

3    I    5   6  2  6    I    2 

41 

s  48 

2     I 

112 

I 

II        24544431 

37 

.s  52 

I     I 

I                  I 

I    2    I    5   4   I    5   5   2 

31 

1^ 

I 

I    2   3    I    3   2  3   2  4   I 

23 

I 

I             12                     2   3    I    I 

12 

8  64 

I        I 

2 

rS    ^ 

0 

W    72 

I 

I 

203    20    I     I    3    I    4469882    711  II  15  22  15  14  14  13  II   5     I     I     i;i9 

Length—Mean,  176.124  ±  1.128^         Breadth— Mean,  47.364  ±  .344M 

St.  Dev.,        23.360  db   .797M  St  Dev.,        7.132  db  .244M 

Coef.   Var.,   13.262  ±   461.  Coef.  Var.,  15.057  ±  .526 

Mean  Index,  27.153  per  cent;  Coef.  Cor.,  .3945  ±  .0408. 


i9o8.] 


JExNNlNGS -HEREDITY   IN   PROTOZOA. 


418 


polygons  of  Diagram  3.  It  is  evident  that  the  variability  has  become 
much  greater  in  the  rapidly  growing  culture.  The  range  of  variation 
of  length  in  the  stationary  culture  is  from  148  to  212  microns;  in 
the  growing  culture  it  is  from  104  to  220  microns,  so  that  in  the 
latter  the  range  has  almost  doubled  in  extent.  The  coefficient  of 
variation  in  length  has  likewise  almost  doubled,  changing  from  7.794 
when  the  culture  was  stationary  to  13.262  when  it  was  growing. 
For  breadth  the  range  of  variability  has  likewise  increased  consid- 
erably, though  the  coefficient  of  variability  shows  little  change.  The 
correlation  between  length  and  breadth  has  become  considerably  less 
in  the  rapidly  multiplying  culture,  decreasing  from  .5955  to  .3945. 
The  mean  length  has  slightly  decreased,  the  mean  breadth  slightly 
increased,  in  the  growing  culture. 

35 


30 

• 
en 

to 

C 

o 

u 

V  20 


>.»5 
o 

c 

s 
8*10 

u 


K 

/ 

/ 

Vv 

1 
f 

/ 

/ 

/^ 

;^... 

1 
1 

/ 

\ 

\ 

A 

ij 

i 

\ 

\ 

• 

/ 

-  / 

1 

\  \ 

^ 

^ 

/ 

y 

104   116   128 


140  152   164   172 
Length  in  Microns. 


188 


200     212 


224 


Diagram  3.  Polygons  of  variation  in  length  for  (a)  a  culture  of  de- 
scendants of  D  that  is  rapidly  multiplying  and  (6)  one  that  is  not.  The 
continuous  line  represents  the  rapidly  multiplying  culture  of  Table  VII.;  the 
broken  line  the  stationary  culture  of  Table  VI. 


From  this  example  it  is  clear  that  growth  and  multiplication  may, 
and  probably  do,  play  a  large  part  in  determining  the  character  and 
distribution  of  the  variations,  as  well  as  in  determining  the  mean 
dimensions  and  their  correlations.     We  shall  now  attempt  to  deter- 


414  JENNINGS— HEREDITY  IN  PROTOZOA.  [April  .4, 

mine  as  accurately  as  possible  what  this  part  is  by  a  systematic  study 
of  growth. 

Material  and  Methods  of  Work. 

In  order  to  exclude  possible  differences  due  to  different  ancestry,  the 
study  of  growth  was  made  with  the  progeny  of  a  single  individual  for  each 
of  the  two  groups.  Of  the  caudafum  group  a  single  individual  D  was  iso- 
lated April  12.  This  individual  was  a  large  one,  measuring  approximately 
250  microns  in  length.  From  it  many  cultures  were  made  under  various 
conditions,  and  all  the  results  on  growth  in  this  group  were  reached  with 
progeny  of  this  individual  D,  save  in  cases  where  the  contrary  is  expressly 
stated.  In  the  same  way  the  results  for  the  aurelia  group  were  reached  with 
the  progeny  of  a  single  individual  c,  unless  otherwise  noted. 

The  method  of  work  in  the  statistical  study  of  growth  was  as  follows: 
Numbers  of  dividing  Paramecia  of  known  descent  were  isolated  and  kept 
for  varying  periods,  so  that  the  age  of  the  individuals  was  known  to  within 
a  few  minutes  or  even  less.  The  individuals  were  then  killed  at  different 
ages  by  the  use  of  Worcester's  fluid,  and  measured.  In  this  way  the  usual 
size  at  various  ages  was  determined,  and  those  variations  in  size  that  are 
due  only  to  varying  age  of  the  individuals  were  excluded.  By  pursuing  this 
method,  an  approximate  curve  of  growth  is  obtained  and  the  part  played  by 
growth  in  the  observed  variations  elucidated;  much  light  is  in  this  way  cast 
on  many  obscure  matters. 

To  persons  who  have  worked  with  Paramecium  it  is  unnecessary  to  point 
out  the  extremely  laborious  and  time-consuming  character  of  the  operations 
required.  Dividing  specimens  must  be  sought  for  with  the  microscope, 
among  hundreds  of  their  rapidly  moving  fellows;  they  must  be  taken  up 
with  the  capillary  tube,  isolated,  placed  in  culture  fluid,  and  the  time  of 
capture  noted.  They  must  then,  after  lapse  of  the  proper  interval,  be  killed 
and  measured;  this  is  the  smallest  part  of  the  work.  To  thus  deal  with 
individuals  of  known  age  by  the  hundred  involves  an  incredible  amount  of 
exhausting  labor,  so  that  if  the  mathematical  student  finds  in  any  stage  the 
numbers  employed  not  always  as  large  as  would  be  ideally  desirable,  he 
will  realize  that  there  is  good  reason  for  this.  But  it  is  hoped  that  the 
numbers  used  are  amply  sufficient,  on  the  whole,  for  the  purposes  designed; 
the  results  are  drawn  from  the  measurement  of  over  1,500  specimens  of 
known   age;   together   with  control   cultures  of  mixed  ages  in  still   larger 

number. 

Especially  in  the  study  of  individuals  that  are  very  young  (up  to  the 
age  of  half  an  hour  or  so),  there  is  very  great  difficulty  in  dealing  with 
large  numbers  owing  to  the  fact  that  the  time  required  for  picking  them 
out  is  very  large  in  proportion  to  the  amount  of  time  they  are  to  be  kept, 
so  that  but  few  can  be  dealt  with  at  once.  Another  great  difficulty  lies  in 
the  fact  that  to  be  strictly  comparable,  the  sets  of  different  ages  must  be 
chosen  on  the  same  day  from  the  same  culture;  otherwise  differences  due 
to  cultural  conditions  show  themselves,  confusing  our  results.  No  culture 
remains  the  same  for  two  successive  days,  and  the  differences  quickly  show 


ijoB.]  JENNINGS— HEREDITY   IN   PROTOZOA.  415 

themselves  in  the  statistical  results.  The  condition  just  mentioned  cannot 
be  absolutely  fulfilled,  but  much  effort  was  directed  toward  filling  it  as 
completely  as  possible,  and  where  it  could  not  be  fulfilled,  strict  account  of 
that  fact  was  taken. 

The  fixing  and  measurement  of  the  specimens  was  done  by  the  methods 
already  described  (p.  396). 

1.  Description  of  Different  Stages  of  Growth. 

First  Stage:  the  Young  Before  Separation  is  Complete. 

In  the  earliest  stage  recognizable,  the  young  Paramecium  forms 
half  of  a  dividing  specimen.  Before  the  constriction  appears  the 
macronucleus  has  become  band-like,  and  the  mother  infusorian  is 
shorter  and  thicker  than  the  specimens  not  preparing  to  divide  (see 
Fig.  2,  a).  The  oral  groove  and  other  differentiated  parts  have 
become  less  marked.  At  the  first  appearance  of  the  constriction  the 
anterior  and  posterior  halves  still  retain  something  of  their  charac- 
teristic form,  and  the  body  of  the  mother  has  extended  a  little  (Fig. 

2,  &).  The  constriction  does  not  pass  squarely  across  the  body,  but 
is  a  little  oblique,  being  farther  back  on  the  oral  side  (Fig.  2,  c,  d,  e). 
As  a  result,  when  the  two  halves  are  measured  separately,  they  will 
seem  to  differ  in  length,  according  to  the  place  where  the  measure- 
ment is  taken.  Thus,  if  d,  Fig.  2,  is  measured  from  the  ends  to  the 
constriction  along  the  oral  side,  the  anterior  half  measures  96 
microns,  the  posterior  half  84  microns,  while  if  the  measurements 
are  taken  along  the  aboral  side  these  proportions  are  exactly  reversed. 
Measurements  taken  from  one  of  the  lateral  sides  give  the  same 
length  for  the  two  halves.  The  Paramecia  may  lie  in  various  posi- 
tions and  this  obliqueness  of  the  constricting  groove  is  not  always 
evident.  Misled  by  this  fact,  I  took  great  pains  to  measure  the 
precise  length  of  each  half  in  a  large  number  of  cases,  finding  con- 
siderable differences,  though  without  any  marked  preponderance  of 
either  half.  But  I  am  now  convinced  that  in  early  stages  of  fission 
the  most  accurate  measurements  of  the  young  are  to  be  obtained  by 
considering  each  to  be  one  half  the  length  of  the  two  together. 

The  breadth  of  the  two  halves  frequently  differs  a  little,  the 
posterior  half  being  at  times  slightly  broader  than  the  anterior  half. 

As  the  constriction  deepens,  the  two  halves  lengthen  (Fig.  2, 
btofigtol,  etc.).     This  lengthening  progresses  with  the  advancing 


416 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  94, 


Fig.  2.  Dividing  specimens  of  the  caudatum  form,  descended  from  the 
individual  D.  Note  the  increase  in  length  and  decrease  in  breadth  as  the 
constriction  deepens.    Anterior  ends  above.    All  X  235. 


,9o8]  JENNINGS— HEREDITY   IN   PROTOZOA.  417 

constriction  until  the  two  halves  separate.  This  lengthening  is  clearly 
evident  in  the  figures  and  in  the  correlation  table  giving  depth  of 
constriction  with  length  of  body  (Table  XL,  page  441).  As  Table 
XI.  shows,  there  is  a  period  at  the  beginning,  before  the  constriction 
reaches  a  depth  of  about  10  microns,  when  there  is  little  relation 
between  the  length  of  the  body  and  depth  of  constriction,  showing 
that  in  this  period  the  halves  have  not  yet  begun  to  lengthen.  We 
may  therefore  take  the  length  of  the  young  at  this  period  as  that 
characteristic  for  the  young  individuals  in  their  earliest  recognizable 
condition,  before  growth  has  begun.  By  dealing  with  these  alone 
we  are  able  to  compare  the  variability  of  the  young  with  that  of 
the  adults,  or  with  random  samples  including  all  ages.  In  the  further 
treatment,  therefore,  the  measurements  of  the  unseparated  young  are 
divided  into  two  classes:  (a)  those  before  lengthening  has  begun; 
(b)  those  after  lengthening  has  begun. 

(a)  The  Unseparated  Halves  before  Lengthening  Has  Begun. — 
Studies  were  made  of  the  young  of  three  lots  of  the  caudatum  group 
(descendants  of  the  individual  D),  and  of  two  lots  of  the  aurelia 
group  (descendants  of  the  individual  c).  Each  "lot"  included 
individuals  taken  on  th^  same  day  from  the  same  small  culture.  In 
most  of  the  lots  there  were  examined:  (i)  The  unseparated  young 
before  growth  had  begun;  (2)  the  unseparated  young  after  growth 
had  begun;  (3)  a  random  sample,  including  all  sorts  of  individuals 
found  in  the  culture.  The  results  of  these  measurements  are  given 
in  Table  VIII.,  page  418. 

(i)  The  caudatum  Form  (Descendants  of  D). — The  most 
thorough  study  was  made  of  lot  i,  of  the  caudatum  group;  the 
results  there  reached  are  typical,  and  perhaps  more  reliable  than  any 
others,  owing  to  the  large  numbers  examined.  We  shall  therefore 
make  the  results  on  this  lot  the  basis  of  our  discussion,  afterward 
bringing  out  points  of  difference  and  resemblance  shown  in  the 
other  lots. 

From  this  lot  i,  I  measured  313  dividing  specimens,  which,  of 
course,  included  626  unseparated  young;  a  random  sample  of  200 
individuals  not  dividing  was  likewise  measured.  A  correlation  table 
for  the  313  dividing  specimens,  giving  the  depth  of  the  constriction 
below  the  general  body  surface  and  the  length  is  given  on  page  441 


418 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  24, 


Table  VIII. 

Mean  Dimensions  and  Constants  of  Variation  for  Youngest  Stages,  in  Corn- 
is  for  convenience  of  reference  in  the  text.  The  column  headed 
which  fuller  data  are  given  on  the  lot  in  question.  A  "  Lot "  consists  of 
Table  X.,  page  428. 


8 


A.  Progtny  of  D{  Caudatum 
Form). 


P 

i- 

55  JS 


Lot  I.  Young    halves,  ! 

where    depth     of    cod-  | 

striction  is  4;/  or  less |  262 

Lot  I.  Halves,     where 

depth  of   constriction  is 

more  than  \^ 


364 
3'  Lot  I.  Random  sample \  200 


Lot  2.  Halves,    where 
depth  of   constriction  is 

less  than  5^  breadth 

5!  Lot  2.  All  halves  of  divid- 

I      ing  specimens 

6  Random  sample 


8 


10 


II 


12 


'3 
14 


Lot  3. ^Halves,  depth  of 
constriction  less  than 
%  breadth 

Lot  3.  Adults  24  hours  old. 

B.   Progeny  ofe{  aurtlia 
form). 

Lot  4.  Halves,  where 
depth  of  constriction  is 
less  than  %  breadth 

Lot  4.  Halves,  lengthening 
begun  (constriction  more 
thanX  breadth) 

Lot  4.  Random  sample 


80 

124 
200 


84 
300 


Lot  5.  Halves,  where  con- 
striction is  less  than  )^ 
breadth 

Lot  5.  Random  sample 


Lots  4  and  5.  All  halves 
where  constriction  is  less 
than  %  breadth  (com- 
bination of  rows  9  and 
12) 


132 

106 
225 


76 
100 


Mean. 


(62) 
14 


(43) 

(42) 
30 


(44) 
41 


(47) 


(63) 
49 


48 
50 


87.848zh.278 


93.o33±.355 
i99.96o±:.740 


82.600zb.468 

85.774rb.593 
184.100i.776 


83.810^1.498 
168.532^1.419 


208  i     — 


51.868zb.325 


60.692zb.527 
114.163^1.784 


56.666db:.425 
1 14.033^.820 


53.6224r.300 


Length. 


Standard 
Deviation. 


Coefficient  of 
I       Variation. 


I 


4.7i6rb.i97     5.368zb.224 


7.i04zb.25i 
J5.528dz.524 


4.394^.332 

6.924zb.42o 
16.264zb.548 


4.782zb.352 
10.768zb.296 


7.636db.27i 
7.765:^.263 


5.320dz.402 

8.o72zb.492 
8.834zb.3oo 


5.7o6rb.42i 
6.389zb.i75 


3.912^2.190 

5.684^.372 
17.443^.555 


3.889zb.302 
12.140zb.580 


4.535±.2i2 


7.541  ±.445 


9.3652^.613 
15.279^.497 


6.862zb.533 
10.646zb.513 


8.459:^.398 


(Table  XL).  In  131  of  these  specimens  the  constriction  had  sunk 
less  than  one  unit  of  the  micrometer  (4  microns)  below  the  surface, 
while  in  the  other  182  the  depth  of  the  constriction  was  greater.  We 
may  take  the  131  specimens  in  which  constriction  had  barely  begun 


«9o8.J 


JENNINGS— HEREDITY   IN   PROTOZOA. 


419 


Table  VIII. — Continued. 

parison  with  Random  Samples  and  Adults,  (The  column  headed  "  Row  " 
"Table"  gives  the  number  of  a  table  found  elsewhere  in  the  paper,  in 
specimens  all  taken  from  the  same  culture  on  the  same  day.)     Compare 


Mean. 


55^8orb.297 


49.540i.215 
50.220zb.308 


50.700^1.364 

50.3881b.307 
46.020^1.251 


65.716dr.706 
40.3201b. 230 


34.850rb.287 


34.590^.383 
34.207rb.241 


45.263^1.597 
47.300rb.437 


38.653dr.437 


Brotdtb. 

Standard 
Deviation. 


Coefficient  of 
Variation. 


5.040db.2IO  9.o82rb.382 


4.296=b.i52 
6468d=.2i8 


3.584rb.2i7 
5.256Hb.i77 


6.784db.499 
5.892  Hb.162 


4.i47±.273 
5.363=b.i7i 


5.463^.423 
6490d=.3io 


6.607  rb.3 10 


8.671  rb.309 
12.877rb.441 


3.532db.26o    I      6.769  ir.5 13 


7.II2dr.433 

II.42Izb.390 


10.322  rb. 768 

14.615dr.411 


3.453±.203    I      9.9"  :t. 587 


1 1. 9891+=. 797 

15.683rb.511 


12.071  rb.947 
13.720rb.667 


17.089  db.822 


Ratio  of 

Breadth  to 

Lenf  th«  or 

Mean  Index 

Per  Cent. 


63.136 


53.592 
25.114 


61.530 

59.166 
25.084 


78.563 
23.899 


67.246 

57.296 
30.177 


79.806 
41.455 


71.835 


Coefficient  of  Corre- 
lation. 


.6546rb.o337 


.0938dr.0496 
.6o64rb.o302 


.I048rb.l055 

.ii36rb.o840 
.4282^.0389 


.22l5rb.0999 
.5496dr.0272 


.6502^1.0479 


.3ioorb.o837 
.6757dt.0244 


.6744^.0597 

.8l52dr.0226 


.7476rb.0292 


as  types  of  the  earliest  stage  of  fission,  and  their  262  halves  as  young 
Paramecia  in  the  earliest  stage.  The  lengths  and  breadths  of  these 
262  halves  are  given  in  Table  IX.  The  constants  derived  from  the 
measurements  of  these,  as  well  as  from  the  measurements  of  the  364 


420 


JENNINGS— HEREDITY   IN   PROTOZOA. 


lApril  24^ 


Table  IX. 

Correlation  Table  for  Length  and  Breadth  of  262  Unseparated  Halves  of 
Dividing  Specimens,  in  which  the  Depth  of  Constriction  was  less  than 
four  microns.  All  descendants  of  the  single  individual  D,  and  taken  from 
the  same  culture  on  the  same  day. 

Length  in  Microns. 

80    82    84    86    88    90    92    94    96    98    100    102 


78 


c 
o 

u 
o 

s 

c 


u 


44 
48 

52 
56 
60 

64 
68 

72 


2 
6 
2 
6 


12 
4 
4 
4 


2 
6 
16 
8 


8 
12 
20 

2 
2 


2 

4 

14 
10 


12 

30 
16 

4 


2 
2 

4 
2 

6 


2 

4 
2 

2 


2 

4 

4 
2 


2 
2 

4 


4 
38 
54 
94 

44 
22 

4 
2 


16 


Length — Mean, 

St.   Dev., 
Coef.   Van, 


24    34    44    30 

87.848  db  278M 
4.716  ±  .197M 
5.368  ±  .224 


62    16    10    12     8       2        2    262 

Breadth— Mean,  55480  ±  .297M 
St.  Dev.,        s.040±.2i0M 

Coef.  Var.,  9.082  ±  .382 


Mean  Index,  63.136  per  cent.;  Coef.  Cor.,  6546 ±.0337. 

halves  in  which  lengthening  had  begun,  and  of  the  random  sample, 
are  given  in  the  first  three  rows  of  Table  VIIL* 

We  will  for  the  present  limit  the  discussion  to  the  relations 
shown  by  comparing  the  youngest  stages  (row  i)  with  the  random 
sample  (row  3)  which  consists  mainly  of  adults.  The  following 
important  facts  are  shown : 

1.  The  mean  length  of  the  youngest  stages  of  the  new  individuals 
is  considerably  less  than  one  half  of  the  mean  length  of  the  indi- 
viduals that  are  not  dividing.  The  mean  length  of  the  young  is 
87.848  microns,  while  that  of  the  individuals  not  dividing  is  199.960 
microns,  or  24.264  microns  more  than  twice  the  mean  length  of  the 
young  individuals.  This  remarkable  relation  will  be  taken  up  later, 
in  discussing  the  measurements  of  dividing  specimens  (page  443). 

2.  The  mean  breadth  of  the  youngest  stages  is  slightly  greater 
than  that  of  adults  not  dividing — 55.480  microns,  in  place  of  50,220 
microns. 

•  In  Tables  VIII.  and  IX.  the  measurements  were  made  and  the  constants 
were  first  computed,  for  the  entire  dividing  specimens.  The  constants  for 
the  halves  were  of  course  readily  obtained  from  these;  they  are  the  same, 
save  that  the  mean  and  standard  deviation  for  length  are  halved,  and  the 
mean  index  is  doubled.  The  computation  of  the  probable  errors  was  based 
on  the  number  of  dividing  specimens,  not  on  the  number  of  halves. 


,9o8.J  JENNINGS— HEREDITY   IN   PROTOZOA.  421 

3.  The  mean  index,  or  ratio  of  breadth  to  length,  is  considerably 
more  than  twice  as  great  in  the  young  as  in  the  adults ;  in  the  former 
it  is  63.136  per  cent ;  in  the  latter  25.114  per  cent. 

4.  The  variability  in  length  is  less  in  the  earliest  stages  of  the 
young  than  in  the  individuals  that  are  not  dividing.  In  the  former 
the  coefficient  that  measures  the  variability  is  but  5.368,  while  in  the 
latter  it  is  7.765. 

5.  The  variability  in  breadth  is  likewise  much  less  in  the  youngest 
stages — ^the  coefficient  being  9.082  in  place  of  12.877. 

6.  The  correlation  between  length  and  breadth  is  nearly  the  same 
in  the  youngest  stage  as  in  the  random  sample,  being  .6546  in  the 
former,  .6064  in  the  latter. 

From  the  other  lots  smaller  numbers  were  examined.  These 
gave  on  the  whole  similar  results,  though  with  certain  significant 
diflferences.    The  facts  are  as  follows : 

From  lot  2  (descendants  oi  D),  124  halves  were  obtained.  On' 
account  of  the  small  number,  I  threw  together  all  in  which  the  depth 
of  the  constriction  was  less  than  one  fourth  the  breadth,  and  consid- 
ered these  the  earliest  stage  (the  depth  of  constriction  and  length  are 
given  for  the  entire  dividing  specimens  in  Table  XLIL,  appendix). 
There  were  thus  obtained  eighty  young  individuals  (dimensions 
for  the  entire  dividing  specimens  in  Table  XLIII.,  appendix). 
It  is  evident  that  this  lot  includes  individuals  varying  more  in 
age  and  growth  than  in  lot  i,  since  in  Idt  2  we  have  included  those 
having  a  much  greater  depth  of  constriction.  The  results  are  shown, 
in  comparison  with  a  random  sample  of  the  same  lot,  in  rows  4  and 
6  of  Table  VIII.  The  facts  are  in  the  main  parallel  with  those  for 
lot  I.  As  compared  with  the  random  sample,  the  mean  length  of 
the  young  is  less  than  one  half,  the  mean  breadth  a  little  greater,  the 
mean  ratio  of  breadth  to  length  more  than  double,  the  coefficients 
of  variation  for  length  and  breadth  much  less.  A  striking  differ- 
ence between  this  set  and  the  young  of  lot  i  is  that  in  the  present 
case  the  correlation  between  length  and  breadth  has  decreased  to 
such  an  extent  that  the  coefficient  computed  (.1048)  is  without  sig- 
nificance, being  less  than  its  probable  error  (.1055).  This  is  due, 
as  we  shall  clearly  see  later,  to  the  fact  that  we  have  included  in  the 


422  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

young  of  row  4  individuals  older  (constriction  deeper)  than  in  those 
of  row  I. 

From  a  third  lot  of  descendants  of  D,  154  halves  were  obtained  ^ 
in  84  of  these  the  constriction  was  less  than  one  fourth  the  breadth. 
Unfortunately  no  random  sample  of  this  culture  was  preserved. 
But  300  individuals  just  twenty-four  hours  old  were  taken  from  it 
for  other  purposes,  and  the  young  halves  may  be  compared  with 
these  (rows  7  and  8,  Table  VIII.).*  It  should  be  noted,  however^ 
that  the  adults  of  row  8  had  been  kept  for  twenty-four  hours  in  a 
rather  small  quantity  of  water,  where  food  was  relatively  scarce,  so- 
that  they  were  smaller  than  would  have  been  the  case  if  they  had 
lived  throughout  under  the  same  conditions  as  the  dividing  specimens. 

In  general,  the  same  relations  are  shown  here  as  in  the  other 
lots.  A  striking  peculiarity  is  the  great  breadth  of  the  young  halves 
(65.716  microns),  as  compared  with  that  of  the  adults  (40.320- 
microns),  so  that  the  ratio  of  breadth  to  length  (the  "  mean  index  *') 
is  more  than  three  times  as  great  in  the  young  as  in  the  adults 
(78.563  per  cent,  in  the  former,  23.899  per  cent,  in  the  latter). 
Owing  to  the  inclusion  of  older  halves,  in  which  lengthening  has 
begun,  the  correlation  between  length  and  breadth  is  again  low 
(.22x5  ±.0999). 

(2)  The  aurelia  Form  (Descendants  of  c), — Two  lots  of  divid- 
ing specimens  of  the  aurelia  form  were  examined,  the  first  including 
132  halves  in  which  lengthening  had  hardly  begun,  the  second  76. 
The  constants  for  these,  in  comparison  with  random  samples  of 
those  not  dividing,  are  given  in  rows  9  to  14  of  Table  VIII.  These 
show  the  same  relations  that  we  have  already  seen  in  the  caudatum 
group,  with  one  exception.  In  the  smaller  collection  (lot  5),  the 
mean  breadth  of  the  halves  was  a  little  less,  instead  of  greater,  than 
that  of  the  random  sample.  In  this  culture  the  animals  were  extra- 
ordinarily broad,  the  mean  ratio  of  breadth  to  length  in  the  random 
sample  being  41.455  per  cent.,  in  place  of  the  usual  ratio  of  about  30 
per  cent.  This  was  due  to  the  fact  that  these  animals  had  been 
placed  twenty- four  hours  before  in  a  rich  nutrient  solution  and  had 

*The  dimensions  of  the  entire  dividing  specimens  of  which  row  7  are 
the  halves  are  given  in  Table  XLIV.  of  the  Appendix;  the  dimensions  of  the 
300  just  twenty- four  hours  old  are  given  in  Table  XLI. 


Z90D.J 


JENNINGS— HEREDITY   IN   PROTOZOA. 


4*Jfc 


become  very  plump.  The  point  of  interest  is  that  the  breadth  of  the 
young  individuals  in  the  earliest  stages  tends  toward  a  constant 
dimension,  becoming  greater  when  the  adults  are  thin,  less  when  the 
adults  are  plump.  Outlines  of  dividing  specimens,  and  of  those  not 
dividing,  from  this  culture,  are  shown  in  Fig.  3,  a  to  /;  the  great 
difference  in  breadth  is  noticeable. 


Fig.  3.  Outlines  of  specimens  of  the  aurelia  form  (descendants  of  c), 
from  Lot  5,  Table  VIII.  c  to  /,  Successive  stages  of  fission.  Note  the 
greater  breadth  of  the  specimens  not  dividing  (a  and  b).  Same  magnifica- 
tion as  Fig.  2.     (235  diameters.) 

In  row  14,  Table  VIII.,  are  given  the  constants  for  all  the  young 
halves  examined  of  the  aurelia  group ;  that  is,  for  the  sum  of  rows 
9  and  12.  The  coefficients  of  variation  are,  as  might  be  expected, 
increased  by  adding  these  two  dissimilar  groups.  The  fact  that  the 
correlation  between  length  and  breadth  is  likewise  increased,  as  com- 
pared with  what  we  find  in  either  group  taken  alone,  might  not, 
perhaps,  be  anticipated.  These  changes  in  variation  and  correlation 
are  environmental  eflfects,  to  be  studied  later. 

(b)  The  Unseparated  Halves  after  Lengthening  Has  Begun, — 
As  we  have  already  seen,  the  length  of  the  halves  increases  as  the 
constriction  deepens  (see  the  correlation  tables  for  length  with  depth 
of  constriction,  Nos.  XI.  (page  441),  XLV.,  XLVI. ;  compare  also 
the  outlines  of  dividing  specimens,  Figs.  2  and  3).  The  coefficient 
of  correlation  between  depth  of  constriction  and  length  is,  for  the 
626  halves  of  Table  XL,  .6882;  with  each  increase  of  10  microns 
in  depth  of  constriction  the  length  increases  4.30  microns.  If  we 
include  only  the  individuals  in  which  lengthening  has  clearly  begun 
(thus  omitting  the  uppermost  row  of  Table  XL),  we  find  that  for 


f24  JENNINGS— HEREDITY  IN   PROTOZOA.  [April  94, 

these  364  halves  the  correlation  between  depth  of  constriction  and 
length  is  greater,  amounting  to  .7818;  while  the  increase  in  length 
with  each  10  microns  of  increase  in  depth  of  the  constriction  is 
5.598  microns. 

While  the  length  thus  increases,  the  breadth  decreases.  This  is 
evident  on  inspection  of  Table  XII.  The  correlation  between  depth 
of  constriction  and  breadth  of  body  is  therefore  negative;  its  coeffi- 
cient, in  the  case  of  Table  XII.,  is  —  .5232.  With  each  increase  of 
10  microns  in  the  depth  of  constriction  the  breadth  of  body  decreases 
2.630  microns.  If  again  we  take  into  consideration  only  the  364 
halves  in  which  lengthening  has  decidedly  begun,  omitting  thus  the 
uppermost  row  of  Table  XII.,  we  find  that  the  correlation  decreases 
to  —  .3316,  and  the  decrease  in  breadth  for  an  increase  of  10  microns 
in  depth  of  constriction  is  but  1.252  microns.  This  appears  to  indi- 
cate that  a  large  part  of  the  decrease  in  breadth  occurs  in  the  first 
stages  of  constriction. 

If  we  compare  with  the  means  of  the  262  halves  in  which  length- 
ening has  not  begun,  the  means  of  the  364  in  which  lengthening  has 
begun  (Table  VIII.,  rows  i  and  2),  we  find  that  the  length  has 
increased  from  87.848  to  93.033  microns,  while  the  breadth  has 
decreased  from  55.480  to  49.540  microns.  If  we  examine  the  means 
at  successively  older  stages,  we  find,  of  course,  greater  diflferences. 
Thus,  when  the  constriction  has  reached  a  depth  of  36  microns,  the 
10  specimens  in  that  stage  show  the  mean  length  increased  to  101.200 
microns,  while  the  mean  breadth  is  but  46.400  microns.  Similar 
relations  are  to  be  observed  if  we  compare  the  means  of  the  younger 
and  older  sets  of  each  lot  shown  in  Table  VIII. 

Since,  while  the  length  is  increasing,  the  breadth  is  decreasing, 
the  growth  tends  to.  decrease  the  correlation  between  length  and 
breadth  or  even  to  make  it  negative.  Thus,  while  in  the  stage  before 
lengthening  has  begun  (row  i.  Table  VIII.)  the  correlation  is  .6546, 
in  the  364  specimens  of  the  same  lot,  after  lengthening  has  begun 
the  correlation  has  decreased  to  — .0938  (row  2,  Table  VIIL).  In 
a  second  lot,  containing  124  halves,  when  we  throw  all  the  halves 
together  the  coefficient  of  correlation  between  length  and  breadth 
becomes  — .1136  (row  5,  Table  VIII.).  In  the  aurelia  form,  106 
halves  after  lengthening  has  begun  give  a  positive  correlation  between 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  425 

length  and  breadth  of  .3100  (row  'lo,  Table  VIII.).  Why  there 
should  sometimes  be  a  slight  positive  correlation,  sometimes  a  nega- 
tive one,  at  this  stage,  will  be  discussed  in  the  section  where  we  deal 
with  the  various  factors  determining  correlation. 

A  variation  polygon  for  the  youngest  stage  of  lot  i  of  Table 
VIII.  is  shown  in  Diagram  4,  p.  440,  at  a. 

The  changes  above  set  forth  from  statistical  data  were  in  a  num- 
ber of  cases  observed  in  living  individuals.  These  observations  give 
a  number  of  additional  points  of  importance,  so  that  they  will  be 
described.  The  facts,  as  illustrated  mainly  by  a  typical  specimen  of 
the  aurelia  form,  are  as  follows : 

Some  time  before  fission  the  body  thickens  and  becomes  shorter, 
taking  the  form  shown  at  a.  Fig.  2,  or  c,  Fig.  3.  The  form  and  dimen- 
sions differ  very  noticeably  from  those  of  the  specimens  not  preparing 
to  divide.  How  long  before  the  appearance  of  the  constriction  these 
preparatory  changes  in  form  begin  it  is  not  possible  to  say,  because 
it  is  not  possible  to  distinguish  with  certainty  whether  a  given  speci- 
men is  to  divide  or  not  until  we  can  see  the  constriction,  and  this  is 
at  a  relatively  advanced  stage  of  the  process.  At  the  time  the  con- 
striction first  appears  the  anterior  and  posterior  halves  still  differ  in 
form,  though  they  are  losing  their  characteristic  features. 

As  the  constriction  deepens  the  two  halves  become  longer  (Fig. 
2,  b  to  /,  Fig.  3,  r  to  d).  A  specimen  of  the  aurelia  form  (descend- 
ant of  c)  was  at  about  the  stage  shown  at  d.  Fig.  3,  at  12.05;  each 
half  measured  very  nearly  80  microns  in  length. 

Ten  minutes  later  (at  12.15)  the  connecting  portion  had  become 
smaller,  while  the  two  halves  had  lengthened,  so  that  each  measured 
about  85  microns  in  length.  The  anterior  half  was  more  pointed 
and  slightly  more  slender  than  the  posterior  half  (/,  Fig.  3)  ;  this  is 
regularly  the  case. 

Six  minutes  later  (at  12.21)  the  posterior  half  measured  about 
90  microns,  the  anterior  half  94.  The  connecting  band  was  now 
extremely  slender. 

Five  minutes  later  (at  12.26)  the  two  halves  separated.  The 
anterior  half  was  still  clearly  distinguishable  from  the  posterior  one 
by  its  pointed,  somewhat  pear-like  form.     It  measured   100X44 

PROC.  AMER.  PHIL.  SOC.  XLVU.  I90  BB,  PRINTED  JANUARY  9,  I9O9. 


426  JENNINGS— HEREDITY   IN   PROTOZOA.  [AprUa4, 

microns,  while  the  posterior  halt  was  shorter,  but  thicker,  measuring 
96  X  52  microns.  The  succeeding  changes  of  form  will  be  described 
in  the  next  section. 

Thus  from  the  condition  shown  at  d,  Fig.  3,  to  the  completion 
of  fission  a  period  of  twenty-one  minutes  elapsed.  From  the  earliest 
appearance  of  the  constriction  the  time  till  separation  is  usually  a 
little  more  than  one  half  hour. 

Second  Stage:  the  Young  Immediately  after  Fission  up  to  the  Age 

of  Ninety  Minutes. 

Observation  of  LiiAng  Specimens. — Immediately  after  separation 
of  the  two  halves,  growth  occurs  rapidly,  and  the  shape  changes,  both 
halves  becoming  more  pointed  at  both  ends.  In  the  specimens  of  the 
aurelia  form  under  description  at  the  close  of  the  last  section,  the 
posterior  half  had  two  minutes  after  fission  increased  in  size  from 
96  X  52  microns  to  104  X  48  microns.  Eight  minutes  after  separa- 
tion both  halves  measured  112  microns  in  length,  so  that  they  had 
during  that  period  increased  respectively  12  and  16  microns  in  length. 
The  difference  between  anterior  and  posterior  individuals  was  still 
marked. 

Now  followed  a  period  of  slower  growth.  At  12.53,  twenty- 
seven  minutes  after  division,  each  half  measured  approximately  120 
microns  in  length.  They  had  taken  nearly  the  characteristic  adult 
form  and  it  was  no  longer  possible  to  distinguish  the  anterior  product 
from  the  posterior  one. 

At  2  P.  M.  (one  hour  and  thirty-four  minutes  after  separation) 
the  length  was  about  135  microns  and  the  progeny  were  similar  to 
the  adult  specimens  of  the  aurelia  form. 

Thus,  at  the  time  of  separation  the  two  individuals  have  some- 
what more  than  half  the  adult  length ;  they  grow  rapidly  at  first,  then 
slowly,  and  in  an  hour  and  a  half  have  reached  nearly  the  adult  size. 
(As  later  statistical  studies  show,  growth  continues  for  a  long  time 
still.) 

Observation  on  the  growth  of  living  specimens  of  the  caudatum 
form  gave  a  parallel  series  of  phenomena  (see  Fig.  4).  Thus,  in  a 
descendant  of  D,  the  length  of  each  half  at  the  time  of  separation 


,9o8.j  JENNINGS— HEREDITY   IN    PROTOZOA.  427 

was  about  120  microns ;  width  48  microns.  Five  minutes  later  the 
length  had  increased  to  132  microns,  while  the  width  was  still  48 
microns.  Nine  minutes  later  the  length  of  the  anterior  product  was 
148  microns ;  that  of  the  posterior  product  144  microns.  The  width 
had  decreased  a  little ;  it  was  now  about  44  microns. 

After  thus  increasing  in  fourteen  minutes  by  nearly  one  fourth 
the  original  length,  growth  became  less  rapid.  JForty  minutes  later 
(fifty-four  minutes  after  separation)  the  length  was  about  156 
microns.  During  two  succeeding  hours  no  increase  in  length  could 
'  be  detected.  The  form  was  that  of  the  normal  adult,  though  the 
adult  size  was  not  yet  reached. 

We  may  summarize  as  follows:  Some  time  before  fission  (per- 
haps a  half  hour)  the  body  shortens  and  thickens,  so  that  each  half 
is  at  first  less  than  half  the  adult  length.  As  the  constriction  deepens 
the  two  halves  grow  longer,  till  at  the  time  of  separation  they  are 
somewhat  more  than  half  the  adult  length.  For  five  to  twenty  min- 
utes after  separation  growJth  in  length  is  very  rapid,  while  the  thick- 
ness remains  stationary  or  decreases.  Then  follows  a  period  of 
several  hours  of  slower  growth,  till  the  adult  size  is  reached. 

This  somewhat  indefinite  account,  based  on  the  observation  of 
living  specimens,  will  now  be  supplemented  by  a  statistical  investi- 
gation of  a  large  number  of  individuals  at  various  ages.  The  main 
results  of  this  statistical  investigation  are  brought  together  in  Table  X. 

(c)  Age  o  to  5  Minutes  (Table  XXIX.). — A  large  number  of 
dividing  specimens,  all  descendants  of  the  individual  D  (caudatum^ 
form),  were  removed  from  a  rapidly  multiplying  culture  and  kept 
for  from  o  to  5  minutes  in  a  watch-glass  of  culture  fluid,  then  killed 
and  measured.  The  method  of  work  was  to  spend  five  minutes  in 
picking  out  dividing  specimens  with  the  capillary  tube  and  placing 
them  in  the  watch-glass ;  at  the  end  of  the  five  minutes  the  lot  was 
killed.  Then  other  lots  were  prepared  in  the  same  way.  In  each 
lot  killed,  therefore,  there  occurred  specimens  that  were  in  the  early 
stages  of  fission ;  others  that  had  separated  at  the  moment  of  removal 
and  were  hence  just  five  minutes  old;  and  all  stages  intermediate 
between  these  two.  All  together,  62  unseparated  pairs  and  59  sepa- 
rated individuals  were  secured  in  this  way.  The  latter  set  consists 
of  individuals  from  o  to  5  minutes  old  (reckoning  from  the  moment 


428 


JENNINGS—HEREDITY  IN   PROTOZOA. 


[April  34. 


Table  X. 

Dimensions  and  Constants  of  Variation  for  Paramecia  of  Various  Ages,  in 
taken  from  the  same  culture  on  the  same  day.  The  lots  where  identical 
column  headed  "Row"  is  for  convenience  of  reference.  The  column 
elsewhere,  in  which  fuller  data  are  given  on  the  lot  in  question.) 


8 

OS 


j 


3 
4 

5 

6 

7 
8 

9 
lo 

II 

12 

13 
15 


i6 

17 
i8 

19 

20 
21 


22 

23 

24 


25 


1*3 -3' 

\u  § 

Xs  Progtny  of  D  {Caudaium      ,JS2 
Form).  ,     \B.t 


» 


(2 


Length. 


Mean  in 
Microns. 


Lot  I.  Youngest  unseparated 
halres,  constriction  begin- 
ning 1262 

Lot    I.    Halves,    lengthening  | 
begun 364 

Lot  I.   Random  sample , 


Lot  2.  From  beginning  of  con- 
striction to  5  minutes  after 
separation 

Lot  2.  o  to  5  minutes  after 
separation 

Lot  2.  Random  sample 


Lot  6.  Age  o  to  19  minutes. ... 
Lot  6.  Age  18  to  28  minutes... 
Lot  6.  Age  35  to  45  minutes... 
Lot  6.  Age  75  to  90  minutes... 
Lot  6.  Age  o  to  90  minutes 

(sum  of  rows  7-10) 

Lot  6.  Random  sample 


Lot  7.  Age  o  to  19  minutes. 


Lots  6  and   7.    All  o  to  19 
(sum  of  rows  7  and  13) 


Lots  6  and  8.  Age  18  to  28 
minutes  (sum  of  row  7,  and 
of  57  of  another  lot) 


Lot  9.  Age  3  to  4  hours 

Lot  9.  Age  4.20  to  5  hours 

Lot  9.  Age  3  to  5  hours  (sum 

of  rows  16  and  17) 

Lot  9.  Random  sample 


Lot  10.  Age  12  hours 

Lot  10.  Age  12  hours  (same 
as  row  20,  but  omitting  2 
smallest) 

Lot  10.  Age  18  hours 

I^t  3.  Age  24  hours 

Lot  3.  Early  fission,  depth  of 
constriction     less    than     ^ 
breadth 


Lot  I.  Early  fission,  constric- 
tion 4^  or  less 


200 


183 

59 
200 

24 
49 
25 
42 

140 
100 

39 


63 


106 

93 
95 

188 
'95 


(62) 


29 
30 

31 
33 
35 
36 


51 
32 


34 

37 
38 


73  39 


71 
105 

300 


42 


131 


39 
40 
41 


44 


'3 


87.8484:  .278 

93.o33±  .355 
i99.96o=b  .740 


92.940±  .718 

107.660dzi.296 
i84.ioo±  .776 

128.ooozhi.908 
«43.348zfc  .624 
149.920dbi.012 
16i.524dbi.004 

i47.544±  .824 
i84.68o±  .848 

134.256dri.663 


I3^872ifci.288 


Standard 

Deviation  in 

Microns. 


Coefficient  of 
Variation. 


4.7i6dz  .197,  5.368dz  .224 


7.i04±  .251 
I5.528dz  .524 


7.636zb  .271 
7.765 ±  .263 


14.400±  .508  15.494:!=  .559 


i4.78odz  .916 
i6.264±  .548 

13.856dbi.348 
6.48odz  .440 
7.5'2db  .716 
9.648 db  .712 

14.464^  .584 
i2.596±  .600 

15.394dbi.176 


I5.i76zb  .912 


5.769^  .268    1 1 2-1 68 


143.82  zb  .544!  8.296±  .384 

I49.636zfc  .688   9.856dz  .488 
i86.736±  .652   9.4i6dz  .460 


i68.384zb  1.028  20.904dz  .727  I2.4i5dz  .438 
i76.i24±i.i28  23.36odz  .797  i3.262dr  .461 


72-132 

76-132 
140-216 

108-152 
132-160 
132-160 
140-180 

128-180 
156-224 

ii.468±  .857    io8-i6o 


13.7291b  .868 
8.8344=  .300 

io.825±i.o66 
4.52 1  db  .309 
5.oiozb  .479 
S.974=b  .441 

9.803  d-  .399 
^6.82 1  rb  .327 


Range  of 

Variation 

in  Microns. 


78-102 

80-112 
148-240 


1 1.507 dz   .701 


108-160 


6.58743   .327 
5.043^=  .247 


132-176 
164-216 

132-216 
104-220 


188.9884::  .996  i2.6i2dz  .704   6.672dz  .3741  136-216 


I90.424db  .752  9.388dz  .53'  4.9304:  .280' 
I99.048dz  .38oii.844zt  .552  5949^  .278! 
i68.532±:  .419  io.768dz  .629   6.389^=  .175 


1 67.620 dz   .996 


I75.696dr  .556 


9.564^  .704 


9432  zh  .393 


5.7o6zb  .421 


5.368dz  .224 


164-216 
168-228 
140-200 


152-192 
156-240 


i9o3.| 


JENNINGS— HEREDITY   IN   PROTOZOA. 


429 


Table  X. — Continued. 

Comparison  with  Random  Samples.  (Each  "Lot"  consists  of  specimens 
with  those  of  Table  VIII.  are  numbered  the  same  as  in  Table  VIII.  The 
headed  "Table"  gives  the  number  of  a  table  found  in  the  appendix  or 


Breadth. 

Mean  Index, 

Ratio  of 

Breadth  to 

Length. 

Per  Cent. 

Mean  in 
Microns. 

Sundaid 

Deviation  in 

Microns. 

Coefficu  et  of 
Variation. 

Range  of 

Variation 

in  Microns. 

44-72 

Coefficient  of 
Correlation. 

55.480rb.297 

5.040rb.2IO 

9.082  4- .382 

63.136 

.6546^.0337 

49.54Oi.2i5 
50.22Orb.308 

4.296^.152 
6.468^.218 

8.67 1  ±.309 
i2.877dr.44i 

40-68 
36-72 

53592 
25.114 

— .0938rb.0496 

.6o64dr.0302 

48.852dr.2iO 

4.216-I-.I49 

8.633^.307 

36-64 

54.080 

— .3625rb.0433 

46.3724-.332 
46.020-h.251 

3.8o4dr.236 
5.256-^-.i77 

8.200±:.524 
1 1.42 1  dr. 390 

36-56 
36-60 

44.037 
25.084 

-.3i38dr.o792 

.4282dr.0389 

60.l68dr.788 

54.284dr.364 
55.84Odr.636 
54.192  dr.6oo 

5.7'2di.556 
3.788 -<-.26o 
4.724-^-.452 

5.752dr.424 

9.495  ±933 

6.976dr478 

8.461  dr. 813 

10.6174-.790 

52-76 
48-64 
48-64 
40-68 

47.573 
37.921 
37.296 
33.558 

— .0337dz.i375 
.1937-+- .0927 
.27994^.1243 

.52324-.0756 

55.544rb.308       5.4l6dr.220 
64.880-l-.580   '   8.624-1- .412 

9.748^-.397 
13.2924- .645 

40-76 
44-88 

38.038 
35.131 

--.o844dr.o566 
.6469d:.0392 

46.768-+- .408     3.792-J-.288 

8.io8rb.623 

36-52 

35.616 

—  .2546^:. 1010 

5 1. 872-+- .680     7.980-^.480 

1 

15.382-^.946 

36-76 

40.028 

—.24764^.0798 

50.832dr.320 

4.9004- .228 

9.640-^- .45 1 

36-64 

35.438 

.1319^.0644 

5 1.568 -+-.322 
60.168rb.360 

4.752=b.236 

5.224dr.256 

9.2I2i:.459 

8.6794-.428 

40-64 
52-76 

34.546 
32.225 

.320I4:.o628 

.5557 -^-.0478 

55.916-^- .324 
47.364-^- .344 

6.588dr.229 
7.I324r.244 

11.7854r.416 
15.057^.526 

40-76 
32-72 

33.372 
27.153 

.7 1 32  4- .0242 
.3945^.0408 

62.796 -4- .464 

5.8724r.328 

9.350db.526 

48-80 

33.275 

.4868dr.o6o2 

63.156dr.443 
56.496  dr.292 

40.320de.230 

5.536^.313 

4.428rb.Io8 

5.8924-.I62 

8.763-^.500 

7.837-^-.367 
14.615^.411 

48-80 
48-68 
28-56 

33.197 
28.427 

23.899 

.3474d=.o704 
.43044-.0536 
.5496db.0272 

65.7i6-<-.7o6 

6.7844r.499 

10.322  4-.768 

48-80 

39.286 

.221543.0999 

* 

55.480^.207 

q.o40-i-.2io 

0.082-^.382 

1        44-72 

31.568 

.6546=^.0337 

430 


JENNINGS—HEREDITY   IN   PROTOZOA, 


[April  34. 


Table  X. — Continued. 


& 


26 


27 
28 


29 


30 
31 
32 


33 


34 
35 

36 
37 

38 


A.  Prcgtnyof  D  (  Caudatum 
Form). 


Lot  I.  Fission,  all  stages  but 
earliest 

Lot  I.  Random  sample 

Lot  I.  Largest  specimens  of 
random  sample,  all  more 
than  196  long 

Lot  I.  Combination  of  early 
6ssion  with  largest  of  random 
sample  (sum  of  rows  25  and 

a8) 

Lot  2.  Early  stages  of  fission .. 

Lot  2.  All  stages  of  fission 

Lot  2.  Random  sample 

B.  Progeny  of  c   {aurelia 
form). 

Lot  4.  Early  fission,  depth  of 
constriction  less  than  j^ 
breadth 

Lot  4.  Later  stages  of  fission... 
Lot  4.  Random  sample 


Lot  5.  Early  fission 

Lot  5.  Random  sample 

Lots  4  and  5.  All  in  early  fis- 
sion ( sum  of  rows  33  and  36 ) . 


Number  of 
Individuals. 

• 

I 

182 

62 

200 

14 

134 

— 

264 

— 

40 
62 

44 
42 

200 

30 

66 

47 

'53 

63 

225 

49 

38 

48 

100 

50 

104 

— 

Length. 


Mean  in 
Microns. 


l86.o66dr  .710 
l99.96ozb  .740 

—  2o8.268db  .566 

—  192.108 

i65.2oo±  .936 
i7L548±:i.i88 
i84.iooih  .776 


io3.737±:  .650 

121.383zbi.053 
ii4.i63rt  .784 

II3.333:±:  .850 
ii4.o33±  .820 


—    1 07.243  dz  .600 


Standard 

Deviation  in 

Microns. 


14.208:^.502 
i5.S28dr.524 

9.720zb400 


18.904 

8.788ih.664 
13.848i.840 
i6.264±:.548 


7.823±.379 

11.367db.743 
i7.443±.55S 

7.778=b.6o3 
12.140dr.580 


9.o7odr.423 


Coefficient  of 
Variation. 


7.636^.271 
7.765^.263 


5.320±402 
8.072Hb.492 


Range  of 

Variatioo 

in  Microns. 


160-224 
148-240 

196-240 


153-192 
144-212 


8.834it.300    140-216 


7.541  ±.445 


83.3-126.7 


9.365^.613    100-156.7 
15.279^.497  '73.3-160 

6.862=b.533  1933-' 26.7 
10.643^.513  86.7-146.7 


8.459^=.398  83.3-126.7 


of  separation  of  the  two  halves).  The  measurements  of  these  59 
young  specimens  are  given  in  Table  XXIX.,  while  the  polygon  of 
variation  for  length  appears  at  b,  Diagram  4.  For  control,  Table 
XXX.  gives  the  measurements  of  a  random  sample  of  the  culture 
from  which  these  young  specimens  were  selected.  The  constants 
deduced  from  the  measurements  of  the  young  and  of  the  random 
sample  are  shown  in  Table  X.,  rows  4  to  6. 

The  following  are  the  important  facts  which  result  from  the 
examination  of  the  young,  in  comparison  with  the  adults  (rows  5 
and  6,  Table  X.). 

I.  The  mean  length  of  the  young  (o  to  5  minutes  old)  is  consid- 
erably more  than  half  that  of  the  culture  as  a  whole,  being  107.660 
microns  as  compared  with  184.100  microns.     Of  course,  the  culture 


x9o8.| 


JENNINGS— HEREDITY  IN   PROTOZOA. 


431 


Table  X. — Continued. 


Breadth. 


Mean  in 
Microns. 


49.540^.215 
50.22Odr.308 

52.360^.348 


53.908 

50.700dr.364 
50.388d1.308 

46.020d=.25i 


34.850rb.287 

34.590dr.383 
34.207rfc.241 

45.263rfc.597 
47.300dr437 


38.653^.437 


Standard 

Deviation  in 

Microns. 


4.296rfc.i52 
6468dr.2l8 

5.964rfc.246 

5.752 

3.432dr.260 

3.584^.216 

5.256dr.i77 


3453±.203 

4.i47±.273 
5.363^.171 

5.463^.423 

6490dr.3IO 


6.607  dr. 3 10 


Coefficient  of 
Variation. 


8.67 1  dr. 309 
i2.877dr.44i 


6.769dr.5i3 

7.iiirfc.433 

Ii.42idr.390 


9.9"dt.587 

1 1.989^.797 
15.683dr.511 

12.071  dr. 947 
13.720dr.667 


17.089dr.029 


Range  of 

Variation 

in  Microns. 


40>68 
36-72 

40-72 


3<2| 


p5"^ 


26.796 
25.114 


48-80 
40-60 
36-60 


26.7-43.3 

26.7-46.7 
20-50 

33.3-56.7 
36.7-66.7 


26.7-56.7 


30.765 

29.583 
25.084 


33.623 

28.648 
30.177 

39.903 
41.455 


Coefficient  of 
Correlation. 


— .0938d:.0496 

.6o64dr.0302 

.468idr.0455 
.0350dr.04i5 

.I048dr.I055 

— .ii36d=.o840 

.4282dr.0389 


.6502dr.0479 

.3ioodr.o837 
.6757zfc.o244 

.6744dr.o597 

.8l52dr.0226 


.7476rfc.0292 


as  a  whole  contains  a  large  number  of  young  specirpens,  so  that  the 
mean  of  the  adults  would  be  greater  than  that  of  the  random  sample. 

2.  The  mean  breadth  of  the  young  is  almost  exactly  the  same  as 
that  of  the  culture  as  a  whole. 

3.  The  relative  variation  in  length  is  much  greater  for  the  young 
than  for  the  culture  as  a  whole,  the  coefficient  being  13.729  for  the 
former  as  compared  with  8.  834  for  the  latter.  Moreover,  the  coeffi- 
cient of  variation  is  almost  three  times  as  great  as  in  the  very  young- 
est stages  before  separation  (Table  X.,  row  i),  or  in  the  first  stages 
of  fission  (Table  X.,  rows  25,  30,  33,  36). 

This  great  variability  of  the  young  at  this  age  indicates  that  they 
are  growing  rapidly  in  length ;  those  five  minutes  old  are  considerably 
longer  than  those  that  have  just  separated,  so  that  when  all  are  taken 


432  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  24, 

together  the  variation  is  great  in  proportion  to  the  mean  length. 
While  the  statistical  data  are  themselves  open  to  other  interpretations, 
observation  of  the  changes  in  living  indnriduals,  as  described  earlier, 
shows  that  this  explanation  is  the  correct  one. 

The  absolute  variation  of  the  young,  as  shown  by  the  standard 
deviation,  is  less,  as  might  be  expected,  than  that  of  the  culture  as  a 
whole,  though  the  difference  is  not  great. 

4.  The  variation  in  breadth,  both  absolute  and  relative,  is  less  in 
the  young  than  in  the  culture  as  a  whole.  The  fact  that  it  is  still 
considerable  perhaps  indicates  that  changes  in  breadth  are  taking 
place  during  growth.     To  this  we  shall  return  immediately. 

5.  The  correlation  between  length  and  breadth  is  negative  in  the 
young,  while  in  the  culture  as  a  whole  it  is  positive.  In  the  former 
the  coefficient  is  — .3138;  in  the  latter  it  is  -f-  .4282. 

The  fact  that  the  correlation  is  negative  in  young  specimens 
(greater  length  associated  with  less  breadth)  indicates  that  while  the 
animals  are  growing  in  length  they  are  becoming  more  slender. 
With  an  increase  of  10  microns  in  length  the  decrease  in  breadth  is 
.757  micron.  If  we  group  together  the  unseparated  halves  (124  in 
niunber)  with  the  separated  ones  (59),  we  find  that  the  negative 
correlation  between  length  and  breadth  is  still  greater,  becoming 
—  .3625  (see  row  4,  Table  X.). 

6.  The  mean  ratio  of  breadth  to  length  ("  mean  index  ")  is  much 
greater  in  the  young  than  in  the  random  sample.  In  the  former  the 
breadth  is  44.037  per  cent,  of  the  length ;  in  the  latter  but  25.084  per 
cent.  If  we  include  the  unseparated  halves  with  those  under  five 
minutes  old,  the  breadth  is  54.080  per  cent,  of  the  length  (row  4, 
Table  X.),  while  in  the  unseparated  halves  alone  it  is  59.166  per 
cent.,  and  in  the  earliest  stages  of  the  unseparated  halves  it  is  61.530 
per  cent,  (see  Table  VIII.,  rows  4  and  5).  There  is  thus  a  steady 
reduction  of  the  ratio  of  breadth  to  length ;  to  this  is  due  the  negative 
correlation  of  the  two,  when  those  of  different  ages  are  thrown 
together. 

(d)  Age  0  to  ip  Minutes  ( Tables  XXXI .  and  XXXII . ) .—From 
another  culture  composed  of  descendants  of  the  individual  D,  speci- 
mens were  taken  on  June  14  and  kept  to  several  different  ages.    The 


,9o80  JENNINGS— HEREDITY   IN   PROTOZOA.  433 

various  ages  and  measurements  are  given,  with  those  of  a  random 
sample  of  the  culture  in  lot  6,  Table  X. 

The  first  set  taken  consisted  of  but  24  specimens,  aged  from  o 
to  19  minutes.  Though  the  number  is  small  it  is  worth  while  to 
work  out  the  constants  for  comparison  with  other  stages  in  this  same 
culture;  it  must  be  remembered  that  it  is  extremely  difficult  to  get 
large  numbers  at  any  one  time  of  individuals  so  young.  The  meas- 
urements are  given  in  Table  XXXI.,  while  the  constants  are  shown 
in  row  7,  Table  X.  For  comparison  with  these  a  second  lot  of  the 
same  age,  but  containing  39  specimens,  was  taken  from  the  same 
culture  two  weeks  later.  The  measurements  are  given  in  Table 
XXXII. ;  the  constants  in  row  13,  Table  X.  The  constants  for  the 
two  sets  taken  together  (63  specimens  aged  o  to  19  minutes)  are 
given  in  row  14,  Table  X. 

Comparing  these  with  the  specimens  but  o  to  5  minutes  old,  we 
find  that  the  mean  length  has  increased  by  36  to  40  microns.  The 
breadth  is  about  the  same  in  one  of  the  lots  (row  13,  Table  X.),  but 
is  much  greater  in  the  other  (row  7).  This  difference  is  due  to 
environmental  effects.  The  coefficient  of  variability  in  length  shows 
a  decided  decrease,  indicating  that  growth  is  relatively  more  rapid 
during  the  first  five  minutes  than  later.  The  correlation  between 
length  and  breadth  is,  as  might  be  expected,  negative  in  the  sets  o  to 
19  minutes  old,  as  it  was  in  the  set  still  younger. 

A  number  of  specimens  were  killed  at  precisely  known  ages,  and 
the  measurements  taken.  Thus,  from  lot  7  (row  13,  Table  X.)  a 
typical  pair  of  young  at  the  moment  of  separation  measured  no  X  52 
microns.  At  the  age  of  one  minute  the  two  members  of  a  pair 
measured  each  124  X  52  microns;  at  two  minutes  another  pair  were 
each  120  X  52  microns.  At  three  minutes  one  member  of  a  pair 
measured  120  X  48  microns,  the  other  124  X  44-  At  five  minutes 
the  lengths  of  the  two  resulting  from  a  certain  fission  were  respect- 
ively 124  X  48  and  112  X  44  microns.  Five  specimens  kept  till  they 
were  precisely  nineteen  minutes  old  measured  respectively  160  X  48 
microns;  160X44;  152X36;  152X40;  156X44.  The  mean  di- 
mensions were  thus  156  X  42.4  microns. 

Outlines  of  individuals  from  o  to  19  minutes  old,  showing  the 


434 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[April  a4. 


relative  sizes,  are  given  in  Fig.  4.    These  may  be  compared  with  the 
adults  of  this  race,  a  to  c,  Fig.  i. 


Fig.  4.  Young  Paramecia,  descendants  of  D  (caudatum  form),  from 
inmiediately  after  separation  to  the  age  of  19  minutes,  a  has  just  separated; 
b,  c  and  d  are  two  to  three  minutes  old;  i  and  /  are  19  minutes  old;  the 
others  are  intermediate.  These  should  be  compared  with  the  adults  a  to  r 
of  Fig.  I  (page  403),  which  are  drawn  to  the  same  scale.    All  X  235. 

(e)  Age  18  to  28  Minutes  {Tables  XXXIII.  and  XXXIV.),— 
The  first  lot  of  this  age  (row  8,  Table  X.)  contained  49  specimens 
(Table  XXXIII.)  and  came  on  the  same  day  from  the  same  lot  as 
the  first  lot  of  24  of  the  preceding  stage,  so  that  the  two  are  strictly 
comparable.  The  mean  length  has  increased  in  the  period  of  about 
thirteen  minutes  by  nearly  16  microns,  while  the  mean  breadth  has 
decreased  7  to  8  microns.  The  ratio  of  breadth  to  length  has  decreased 
almost  10  per  cent.  The  correlation  between  length  and  breadth  is 
in  the  present  lot  positive  though  small  (.1937).  If  we  should  throw 
together  the  two  lots  (rows  7  and  8,  Table  X.),  the  correlation 
would,  of  course,  be  decidedly  negative. 

A  second  lot  of  57  specimens  aged  18  to  28  minutes  was  taken 
from  the  same  culture  about  two  weeks  later.     If  we  throw  the  two 


x9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  435 

lots  together  (Table  XXXIV.)  we  have  io6  specimens  at  this  age 
(row  15,  Table  X.) ;  the  mean  length  is  143.82  microns,  the  mean 
breadth  50.832  microns,  while  the  mean  ratio  of  length  to  breadth  is 
35438  per  cent. 

The  polygon  for  variation  in  length  at  this  age  is  shown  at  c, 
Diagram  4,  p.  440. 

(/)  Age  25  to  45  Minutes  (Table  XXXV.), — From  the  same  lot 
6  (Table  X.)  from  which  came  the  first  sets  aged  o  to  19  and  18  to 
28  minutes,  there  were  taken  on  the  same  day  25  specimens  that  were 
allowed  to  reach  the  age  of  35  to  45  minutes  (row  9,  Table  X.). 
Growth  has  now  become  much  slower.  These  specimens  average 
17  minutes  older  than  the  last  set,  yet  they  have  increased  in  length 
only  about  6.5  microns.  The  breadth  remains  about  the  same;  the 
slight  increase  shown  in  the  figures  is  probably  not  significant,  since 
it  disappears  at  the  next  stage.  The  mean  ratio  of  breadth  to  length 
continues  to  decrease,  reaching  now  37.296  per  cent.  The  correla- 
tion between  length  and  breadth  is  more  strongly  positive  than  before 
(.2799),  indicating  that  these  dimensions  are  not  changing  so  decid- 
edly in  opposite  ways. 

The  polygon  for  variation  in  length  at  this  age  is  shown  at  d, 
Diagram  4. 

(g)  Age  75  to  gto  Minutes  {Table  XXXV L). — Forty-two  speci- 
mens of  this  age  were  measured,  taken  on  the  same  day  from  the 
same  lot  from  which  came  the  sets  last  described  (lot  6,  Table  X.). 
The  specimens  average  about  twice  the  age  of  those  in  the  last  set, 
the  absolute  increase  being  45  minutes,  yet  the  growth  in  length  has 
been  only  about  12  microns,  which  is  about  the  same  as  the  growth 
in  the  first  five  minutes  after  separation.  The  breadth  still  remains 
about  the  same;  it  is  notably  less  than  in  the  very  earliest  stages. 
The  ratio  of  breadth  to  length  continues  to  decrease,  reaching  now 
33.558  per  cent.  Meanwhile  the  correlation  between  length  and 
breadth  has  increased  greatly,  till  now,  at  .5232,  it  is  not  much  below 
that  of  the  culture  as  a  whole  (.6469). 

(A)  Age  0  to  90  Minutes. — From  a  single  culture  of  D,  on  a 
single  day,  we  have  thus  measured  140  young  specimens,  varying  in 
age  from  o  to  90  minutes.  The  constants  for  variability  and  corre- 
lation of  such  a  collection  are  of  interest;  they  are  therefore  given 


436  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a4, 

in  Table  X.,  row  ii.  The  variability,  as  measured  by  its  coefficient^ 
is  less  in  both  length  and  breadth  than  in  the  random  sample,  or  in 
the  collection  of  young  specimens  including  only  those  under  nineteen 
minutes  in  age.  There  is  practically  no  correlation  in  the  collection 
taken  as  a  whole  between  length  and  breadth.  This  is  because 
breadth  at  first  decreases  while  length  increases  (giving  negative 
correlation) ;  later  they  increase  together  (giving  positive  correla- 
tion) ;  the  two  tendencies  about  cancel  each  other  in  the  collection 
as  a  whole. 

Third  Stage:  Three  to  Pipe  Hours  Old  (Tables  XXXVII ,  and 

XXXV II L). 

Three  days  later  than  the  sets  shown  in  lot  6,  Table  X.,  and 
under  as  nearly  the  same  conditions  as  possible,  I  took  from  the  same 
culture  of  progeny  of  D  two  sets  of  young,  keeping  the  first  set  till 
the  age  was  between  3  and  4  hours,  the  second  set  till  the  age  was 
between  4.20  and  5  hours  (see  lot  9,  Table  X.).  The  culture  was,, 
however,  in  a  diflFerent  condition  from  that  of  lot  6;  it  contained  a 
very  large  number  of  young  and  dividing  specimens.  A  random 
sample  of  this  culture,  containing  195  specimens,  is  shown  in  Table 
VII.  (page  412),  while.the  constants  for  this  sample  are  shown  in 
row  19,  Table  X.  The  entire  left  portion  of  Table  VII.,  up  to  the 
length  of  about  160  microns,  or  more,  evidently  consists  of  young 
individuals  in  various  stages  of  growth.  This  decreases  the  main 
length  (176.124  microns)  and  the  correlation  (.3945),  while  it  greatly 
increases  the  variability  in  length  (13.262,  as  against  6.821  for  the 
random  sample  of  the  previous  lot). 

(t)  Age  3  to  4  Hours  {Table  XXXVIL).—The  effects  of  dif- 
ferent environmental  conditions  are  at  once  seen  on  comparing  this 
set  of  93  specimens  (Table  X.,  row  16)  with  the  set  75  to  90  minutes 
old,  from  the  previous  culture  (Table  X.,  row  10).  The  specimens 
of  the  present  lot,  though  li  to  2I  hours  older  than  the  others,  are 
shorter,  the  length  (149.636  microns)  being  less  by  about  16  microns. 
The  breadth  is  about  the  same  as  in  the  previous  set ;  the  correlation 
between  the  two  is  rather  low  (.3201). 

(/)  Age  4,20  to  5  Hours  {Table  XXX F///.).— Ninety-five 
specimens  kept  for  about  an  hour  longer  than  those  in  the  foregoing 


/ 


,9o«.)  JENNINGS— HEREDITY   IN   PROTOZOA.  437 

set  showed  a  rapid  growth  in  length  and  breadth.  The  length  now 
reaches  186.736  microns,  the  breadth  60.168;  both  dimensions  are 
considerably  greater  than  the  mean  of  the  random  sample.  Thus, 
the  animals  at  this  age  had  reached  about  the  average  size  of  the 
infusoria  in  a  collection  of  the  same  descent  taken  at  random.  Table 
VI.  (page  412)  shows  a  sample  of  this  same  culture  taken  twenty- 
four  hours  earlier,  at  a  time  when  little  division  was  occurring ;  the 
mean  length  is  very  nearly  the  same  as  that  of  the  young  of  the 
present  set.  The  correlation  between  length  and  breadth  has  con- 
siderably increased. 

Certain  peculiar  facts  are  brought  out  by  considering  these  two 
sets  together  (Table  X.,  row  18).  Here  we  have  a  collection  of  188 
young  individuals  taken  at  practically  the  same  time  from  a  small 
watch-glass  culture.  The  variability  and  correlation  depend  in  a 
high  degree  on  the  length  of  time  we  keep  these.  If  they  are  all 
kept  three  to  four  hours  (row  16)  or  4.20  to  5  hours  (row  17),  the 
variability  in  length  is  about  5  to  6,  in  breadth  about  9.  But  when 
we  keep  part  of  them  for  the  shorter  period,  part  for  the  longer,  the 
variability  rises  to  about  12.5  for  length  and  12  for  breadth.  Again, 
the  correlation  between  length  and  breadth  is  but  .3201  and  .5557 
in  the  two  lots  taken  separately,  but  when  we  take  them  together  the 
correlation  is  much  greater,  rising  to  .7132.  These  relations  show 
the  important  part  which  may  be  played  by  growth  in  determining 
observed  variability  and  correlation ;  their  significance  will  be  taken 
up  again  in  our  general  sections  on  these  topics. 

Fourth  Stage:  12  to  18  Hours  Old  {Table  X.,  Lot  10). 

From  the  same  culture  of  the  progeny  of  D  from  which  came  the 
lots  last  described,  but  three  days  later  were  taken  two  lots  of  young, 
of  73  and  105  specimens,  respectively,  which  were  kept,  the  former 
to  the  age  of  12  hours,  the  latter  to  the  age  of  18  hours. 

{k)  Age  12  Hours  {Table  XXXIX,,  and  rows  20  and  21,  Table 
X,), — There  is  a  still  further  increase  in  both  length  and  breadth,  as 
■compared  with  the  specimens  4.20  to  5  hours  old  (see  Table  X.,  rows 
20  and  21).  Among  the  73  specimens  of  this  lot  were  two  of  about 
the  same  size  which  were  much  smaller  than  the  others  (see  Table 
XXXIX.).    There  is  little  doubt,  I  believe,  that  these  are  the  prod- 


438  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  24, 

ucts  of  a  second  division;  either  one  of  the  twelve-hour  specimens 
had  divided,  or  there  was  accidentally  taken  with  them  an  older 
specimen  which  divided.  In  either  case  these  two  specimens  do  not 
belong  in  the  twelve-hour  lot,  as  they  are  much  younger.  On  this 
account  I  have  calculated  the  constants  for  this  twelve-hour  lot  twice, 
once  including  these  two  small  specimens  (row  20,  Table  X.),  the 
second  time  excluding  them  (row  21).  ,The  variability  in  length  is 
much  reduced — from  6.672  to  4.930 — ^by  the  omission  of  these  two. 
At  the  same  time  the  correlation  between  length  and  breadth  is  like- 
wise reduced  from  .4868  to  .3474. 

(i)  Age  18  Hours  {Table  XL,,  and  row  22,  Table  X,), — Growth 
in  length  continues,  though  very  slowly;  in  six  hours  the  increase 
has  been  less  than  during  the  first  five  minutes  after  separation.  The 
animals  at  this  age  are  decidedly  longer  than  the  mean  for  the  cul- 
ture as  a  whole,  as  judged  from  the  random  sample  of  Table  VII. 
(page  — ),  taken  three  days  earlier.  The  mean  breadth  of  the 
eighteen-hour  specimens,  while  greater  than  that  of  the  random 
sample,  has  decreased  as  compared  with  that  of  those  only  twelve 
hours  old. 

The  variability  of  these  two  lots  (12  and  18  hours  old)  of  adult 
size  is  less  than  that  of  the  random  samples  (for  examples,  rows  3, 
6,  12,  19,  Table  X.). 

Fifth  Stage:  24  Hours  Old  (Table  XLL,  and  row  23,  Table  X.). 

A  final  lot  of  300  specimens  was  selected  while  dividing  and 
these  were  kept  till  they  were  24  hours  old.  Th^e  were  progeny 
oi  D,  but  were  taken  from  the  culture  somewhat  more  than  a  month 
later  than  those  o  to  18  hours  old.  To  understand  their  measure- 
ments it  is  necessary  to  take  into  consideration  the  cultural  condi- 
tions. These  animals  were  living  in  an  ordinary  hay  culture,  which 
was  getting  old,  so  that  they  were  not  dividing  rap'idly;  they  were 
rather  slender  in  form.  Now  a  large  number  of  these  was  placed 
in  a  fresh  decoction  of  hay  and  left  there  for  24  hours.  They 
increased  in  size  and  began  to  divide  rapidly.  Now  150  dividing 
specimens  (producing,  of  course,  300  young)  were  taken  out  and 
returned  to  the  original  culture  fluid.  This  was  for  the  purpose  of 
preventing  a  second  division  before  the  end  of  the  period  of  twenty- 


,^.]  JENNINGS— HEREDITY   IN   PROTOZOA.  439 

four  hours.  As  a  result  of  this  treatment  they  did  not  grow  so 
rapidly  as  did  the  twelve-  and  eighteen-hour  lots,  and  are  smaller 
than  these.  The  purpose  in  studying  this  group  (as  well  as  other 
groups)  was  mainly  to  determine  the  variability  and. the  correlation 
between  length  and  breadth.  Both  are  less,  as  Table  X.  shows,  than 
is  usually  the  case  in  random  samples. 

The  specimens  12,  18  and  24  hours  old  may  be  taken  as  types  of 
adult  Paramecia  of  this  strain  (progeny  of  D\  caudatum  form) 
before  the  changes  leading  to  fission  have  begun. 

Diagram  4  gives  polygons  of  variation  for  the  different  ages,  in 
descendants  of  D,  as  compared  with  a  random  sample;  it  shows 
clearly  the  part  played  in  the  observed  variations  by  the  presence 
of  different  stages  of  growth. 

Sixth  Stage:  Preparing  for  Fission. 

As  Table  X.  shows,  the  adults  of  the  progeny  of  D  (caudatum 
form)  reach  a  mean  length  of  168.532  to  199.048  microns  (rows  23 
and  22)  under  the  cultural  conditions  employed,  while  the  mean 
breadth  varies  from  40.320  (row  23)  to  62.796  microns  (row  20). 
But  the  maximum  length  is  (under  the  same  conditions),  of  course, 
much  greater  than  the  mean.  In  the  random  samples  we  find  indi- 
viduals up  to  224  microns  in  length  and  88  in  breadth  (see,  for 
example.  Table  LI.) ;  and  among  those  18  hours  old  (Table  XL.) 
we  find  a  length  of  228  microns. 

Now,  when  we  compare  these  large  adults  with  the  specimens 
actually  beginning  fission  (which  are  supposedly  the  oldest  of  all), 
certain  peculiar  facts  appear.  The  specimens  beginning  fission  are 
by  no  means  the  longest  of  the  lot;  a  given  culture  contains  many 
specimens  much  longer  than  those  showing  the  first  signs  of  division. 
Thus,  in  the  "  Lot  i "  of  Table  VIIL,  we  find  131  specimens  in  the 
very  earliest  stages  of  fission  (Table  XIII.,  page  442).  The  mean 
length  of  these  is  175.696  microns  (row  25,  Table  X.),  and  the 
longest  specimen  is  204  microns  long.  But  in  the  random  sample 
of  the  specimens  that  are  not  dividing,  from  this  same  lot  (taken  at 
the  same  time)  the  mean  length  is  199.960  microns  (row  27,  Table 
X.)»  and  certain  individuals  reach  a  length  of  240  microns  (Table 
XIV.,  page  443).     Of  the  two  hundred  specimens  of  the  random 


440 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  84. 


sample,  69,  or  more  than  one  third,  are  longer  than  the  longest  of 
the  specimens  beginning  fission.  Only  nine  of  the  entire  200  falls 
below  the  mean  length  of  the  specimens  beginning  division. 


128     140     152      164 
Length  in  Microns. 


176 


224 


Diagram  4.  Polygons  of  variation  in  length  for  descendants  of  indi- 
vidual D,  at  various  ages.  A  (heavy  line),  Random  sample,  195  specimens 
(row  19,  Table  X.).  a,  youngesf  halves,  constriction  beginning  (row  i. 
Table  X.).  b,  age  o  to  5  minutes  (row  5,  Table  X.).  c,  age  18  to  28  minutes 
(row  8,  Table  X.).  d,  age  35  to  45  minutes  (row  9,  Table  X.).  e,  age  75 
to  90  minutes  (row  10,  Table  X.).  /,  age  4.20  to  5  hours  (row  17,  Table 
X.).    g,  age  12  hours  (row  21,  Table  X.).    h,  age  18  hours  (row  22,  Table 

X.). 


,9o8.j  JENNINGS— HEREDITY   IN   PROTOZOA.  441 

Since  then  the  specimens  beginning  fission  are  not  the  longest  of 
the  culture,  it  is  clear  that  the  length  decreases  before  fission  begins. 
This  is  borne  out  by  the  form  of  the  specimens  beginning  fission; 
though  their  mean  length  is  less  than  that  of  the  random  sample, 
their  mean  breadth  is  greater  (mean  breadth  50.220  microns  in  the 
random  sample,  55.480  in  those  beginning  fission).     While  then  the 

Table  XI. 

Correlation  Table  for  Depth  of  Constriction  and  Total  Length  in  313  Dividing 
Specimens  from  a  Single  Culture  of  Descendants  of  D, 

All  taken  the  same  day. 

Total  Length  of  Body,  in  Microns. 


tf) 


g  156  160 164  168  172  176  180  184  188  192  196  200  204  208  212  216  220  224 


.y  4 

S  8 

c  12 

.0  20 

o  24 

'B  28 

c3  36 

u.  40 

o  44 


8     12 
I     2 


2     I 
2 

I      I 


22 

IS 

31 

8 

5 

6 

4 

I 

I 

131 

8 
10 

5 
9 

7 
3 

5 
3 

2 

I 

I 

37 
30 

3 

5 
3 

4 

I 

4 
3 
3 

4 
I 

4 

2 

I 
5 

3 

I 
2 

2 

I 

29 
12 
16 

I 

I 

4 

2 

5 

I 

2 

2 

2 

I 

I 

22 

I 

3 

I 

2 

4 

3 

I 

I 

17 

2 

2 
2 

2 

I 
I 

I 

I 

I 
2. 

I 
I 

I 
I 

10 
8 
I 

g*  I     10    17   29  44  38  47  30  20  22   16   10   10    6     3     6     2     2  I313 

Q 

Length— Mean,  181.725  ±  .512M      Depth  of  Constriction— Mean,      13.265/* 

St.  Dev.,        13.446  ±  .362M  St  Dev.,  2.721M 

Coef .  Var.,     7.399  ±-  -201 
Coef.  of  Cor.  between  Depth  of  Constriction  and  Length,  .6882  ±  .0201  ; 

Increase  in  Length  for  i  unit  of  depth,  .86om;  Coef.  of  Cor.  if  first  row  is 

omitted,  .7818  ±  .0194. 

length  decreases  preparatory  to  fission,  the  breadth  increases  at  the 
same  time.  How  long  before  fission  this  change  of  dimensions 
begins  I  can  see  no  way  of  determining.  The  period  may  perhaps 
be  one  or  two  hours. 

Thus,  the  longest  individuals  of  the  culture  are  the  adults  that 
have  not  begun  the  changes  preparatory  to  fission.  These  decrease 
in  length  and  increase  in  breadth  before  fission. 

PROC.  AMER.  PHIL.  SOC.  XLVII.   I90  CC,  PRINTED  JANUARY  9,   I909. 


442 


JENNINGS— HEREDITY   IN   PROTOZOA. 


fApri!.,, 


Table  XII. 
Correlation  Table  for  Depth  of  Constriction  and  Breadth  of  Body,  in  313  Di- 
viding Specimens  from  a  Single  Culture  of  Descendants  of  D. 
(Same  lot  shown  in  Tables  XL,  XIII.  and  LXII.) 
Breadth  in  Microns. 
o  40    44    48    52    56    60    64    68    72 


s    * 

2 

IQ 

27 

47 

22      11        2         I 

11 

■( 

15 
.1 

4 

7 

4 

' 

?  .w 

1.  ,tt> 

—  40 

T 

0  44 

I 

5     32    90    ?8    65    28 


Breadth— Mean, 

St.  Dev.. 


52.026  ± 
5-473  i 


Coef.  Var.,  10.544  ±  J87 


Depth  of  Constriction— Mean,      13.265ft 
St.  Dev,  372ifi 


Coef.  of  Cor.  between  Depth  of  Constriction  and  Breadth,  —  .5232  ±  .0277 
Decrease  in  Breadth  with  Increase  of  io*i  in  Depth,  2.63o;>. 

Omitting  uppermost  row:  Coef.  of  Cor,  — .3316 ±  .0445'';  Decrease  i 
Breadth  with  Increase  of  itv  in  Depth,  1.252/1. 

Table  XIII. 

Correlation  Table  for  Length  and  Breadth  of  131  Specimens  of  Lot  I  in  th 

Earliest  Stages  of  Fission.     (Descendants  of  D,  Table  X.,  row  25,) 

Length  in  Microns. 

156  160  164  168  172  176  r8o  184  188  iga  rg6  200  204 


;  48 


36341      I 
1286261   I 

3      a      4     10     7     15     2      2      I      I 

2      I      1      5      8      1      I      2      1 

I  23122 


19 


I      8     12    17    22    IS    31     8      5     6      4      I      I  J131 
VIean,  175-696  ±  .5S6f*  Breadth— Mean,  55.48a  ±  .297/" 

5t.   Dev.,        9432  ~  .393**  St.   Dev.,       5.040  ±  .2ioi« 

:oef.  Var..      5.368  ±  .224.  Coef.  Var.,  gxj8a  ±  .382 

Index,  31.568  per  cent. ;  Coef.  Cor.,  .6546  ±  .0337. 


x9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA. 


443 


Seventh  Stage:  Fission. 

Some  of  the  data  bearing  on  the  dimensions  during  iission  have 
been  incidentally  taken  up  in  the  account  of  the  young  in  the  earliest 
stages,  before  the  two  halves  have  separated. 

(w)  Beginning  Fission,  Descendants  of  D  {caudatum  Form). 
— Four  lots  of  dividing  specimens  descended  from  the  individual  D 
were  studied.  These  lots  were  taken  at  diflFerent  times;  the  first 
included  313  dividing  specimens  (Tables  XI.  and  XII.,  and  rows 
25-29,  Table  X.)  ;  the  second  62  (Tables  XLIL,  XLIII.  (appendix) 
and  rows  30-32,  Table  X.) ;  the  third  77  (Table  XLIV.,  and  rows 
23-24,  Table  X.)  ;  the  fourth  37.  The  dimensions  of  random  sam- 
ples of  the  same  lots  are  given  in  Table  X. 

The  large  lot  containing  313  dividing  specimens  may  be  described 

as  typical;  the  others  show  the  same  relations,  except  as  hereafter 

noted. 

Table  XIV. 

Correlation  Table  for  Random  Sample  of  Specimens  not  Dividing,  of  Lot 
I  (from  which  came  the  diiHding  specimens  of  Table  XI I L).     (See 

Table  X.,  row  27.) 

Length  in  Microns. 

^iOiOnOnO^    r^t^OOOOOO     OnOnO     op     •-•    •-«    «    «    «    KJK)^ 


36 

40 

44 

48 

52 

56 

60 

64 

68 
W   72 


en 

c 
o 

u 
o 


s 


I                               z 

2 

I                12 

131          2           I 

I          2                            I 

354  10  5442        21 

I          I 

11537424332        I 

222566773421 

221112331431 

I             2   I   2  4  3    I 

I                       121            I 

I                          I 

212OI3O3 

6  II  16  22  19  28  17  18  II  15  13  7  3  0  0  2 

Length— Mean,  199.960  ±  .74(W*  Breadth— Mean,  50.220 

St.  Dev.,         15.528  ±  .524A*  St.  Dev.,        6468 

Coef .  Var.,      7765  ^  263  Coef .  Van,  12.877 

Mean  Index,  25.114  per  cent.;  Coef.  Cor.,  .6064  ±  .0302. 


4 
12 

44 
38 
47 
33 
14 
6 

o 

2 

200 

:.3o8m 
.218^ 

Ml 


In  the  dividing  specimens  the  length  of  the  body  increases  as  the 
depth  of  the  constriction  between  the  two  halves  becomes  greater; 
this  is  well  shown  in  Fig.  2,  page  416.  In  order  to  include  only  the 
earliest  stages  of  fission  we  shall,  of  course,  have  to  take  the  speci- 


444  JENNINGS— HEREDITY  IN   PROTOZOA.  [Apnl  a4, 

mens  in  which  constriction  is  beginning.  Among  the  313  dividing 
specimens  of  lot  i  (Table  XL)  there  were  131  in  which  the  depth 
of  the  constriction  below  the  body  surface  was  less  than  one  unit 
of  the  micrometer  scale  (less  than  4  microns).  These  may  be  taken 
'as  representing  the  earliest  stages  of  fission.  The  depth  of  the  con- 
striction is  in  these  specimens  less  than  one  twelfth  the  breadth. 
Their  measurements  are  given  in  Table  XIII.,  while  the  constants 
deduced  from  the  measurements  are  shown  in  row  25,  Table  X. 
These  should  be  compared  with  the  measurements  and  constants  for 
the  random  sample  of  the  specimens  not  dividing  in  this  same  culture 
(Table  XIV.,  and  row  27,  Table  X.). 

Examination  of  these  tables  shows  the  following  remarkable  facts : 

1.  The  mean  length  of  the  specimens  beginning  fission  (175.696 
microns)  is  much  less  than  the  mean  length  of  the  random  sample 
(199.960  microns) — although  the  latter  must  contain  many  specimens 
that  have  not  reached  adult  size. 

2.  The  range  of  variation  in  length  is  much  less  in  the  specimens 
beginning  fission  than  in  the  culture  as  a  whole.  In  those  beginning 
division  the  range  is  from  156  to  204  microns ;  in  the  random  sample 
it  is  from  148  to  240  microns. 

3.  The  longest  specimens  beginning  fission  are  36  microns  shorter 
than  the  longest  of  the  random  sample.  In  the  random  sample,  34.5 
per  cent,  of  all  the  specimens  are  longer  than  the  longest  of  those 
beginning  fission,  while  95.5  per  cent,  are  longer  than  the  mean  length 
of  the  specimens  beginning  fission. 

4.  The  variation  in  length  is  decidedly  less  in  the  specimens 
beginning  fission  than  in  the  random  sample.  In  the  lot  beginning 
fission  the  coefficient  of  variation  is  but  5.368,  while  in  the  random 
sample  it  is  7.636. 

It  may  here  be  noticed  that  coefficient  of  variation  in  the  speci- 
mens beginning  fission  is  less  than  that  for  conjugating  specimens,  as 
studied  by  Pearl  (1907).    To  this  matter  we  shall  return  later. 

5.  In  the  specimens  beginning  fission  the  mean  breadth  (55.480 
microns)  is  greater  than  the  mean  breadth  of  the  random  sample 
(50.220  microns). 

6.  The  variation  in  breadth  is  much  less  in  the  specimens  begin- 


,9o8.J  JENNINGS-HEREDITY  IN   PROTOZOA.  445 

ning  fission  than  in  the  others.     In  the  former  the  coefficient  is  but 
9.082,  while  in  the  latter  it  is  12.877. 

7.  The  mean  index,  or  ratio  of  breadth  to  length,  is  much  greater 
in  the  specimens  beginning  fission;  in  these  it  is  31.568  per  cent.,  as 
contrasted  with  25.114  per  cent,  in  the  random  sample. 

8.  The  correlation  between  length  and  breadth  is  high  in  the 
specimens  beginning  fission;  it  is  somewhat  greater  than  in  the 
random  sample.     In  the  former  it  is  .6546 ;  in  the  latter  .6064. 

Owing  to  the  smaller  numbers  in  the  other  lots  of  dividing  speci- 
mens, I  included  in  the  group  "  beginning  fission  "  all  those  in  which 
the  depth  of  the  constriction  below  the  body  surface  was  less  than 
one  fourth  the  breadth  of  the  animal.  Thus,  all  specimens  with 
constriction  12  microns  deep,  or  less,  were  included.  Of  course, 
these  groups  contained  specimens  in  decidedly  more  advanced  stages 
of  fission  than  in  the  large  group  we  have  been  considering.  The 
numbers  of  specimens  in  early  stages  of  fission  thus  secured  were 
respectively  40  (Table  XLIII.)  and  42  (Table  XLIV.).  The  con- 
stants for  these,  in  comparison  with  random  samples  or  adults,  are 
shown  in  Table  X.  (rows  24  and  30). 

As  the  tables  show,  these  manifest  in  most  particulars  the  same 
relations  which  we  have  brought  out  above  for  the  larger  and  more 
precise  set  containing  131  specimens.  The  differences  between  the 
dividing  specimens  and  the  other  individuals  (as  shown  by  the  random 
samples,  etc.)  are  in  the  main  somewhat  less  in  amount  than  in  our 
first  example.  This  is  because  in  the  smaller  lots  specimens  are 
included  in  which  lengthening  and  narrowing  had  begun,  causing  the 
dimensions  to  approach  those  of  the  specimens  not  dividing. 

The  most  striking  difference  between  our  large  lot  (Table  X., 
row  25)  and  the  smaller  ones  (Table  X.,  rows  24  and  30)  is  in  the 
correlation  between  length  and  breadth.  While  in  the  larger  lot  the 
correlation  was  high,  in  the  smaller  ones  it  is  small  or  quite  lacking. 
This  is  again  due  to  the  inclusion  of  more  advanced  stages  in  the 
smaller  lots ;  as  the  length  increases  the  breadth  decreases,  tending  to 
destroy  the  correlation. 

Descendants  of  c  (aurelia  Form). — Two  lots  of  dividing  speci- 
mens were  examined  from  the  descendants  of  the  small  individual  c. 
The  first  contained  119  specimens   (Table  XLV.)  ;  the  second  63 


446  JENNINGS— HEREDITY   IN  PROTOZOA.  [April  .4. 

specimens  (Table  XLVI.).**  Selecting  from  these,  as  representing 
the  early  stages  of  fission,  all  those  in  which  the  depth  of  constriction 
is  less  than  one  fourth  the  diameter  of  the  body,  we  obtain  from  the 
larger  lot  66  specimens  (Table  XLVIL) ;  from  the  smaller  lot  38 
specimens  (Table  XLVIII.).  The  constants  for  these,  in  compari- 
son with  those  for  random  samples,  are  given  in  Table  X.  (lots  4 
and  5,  rows  33  to  38).  The  measurements  of  the  random  samples 
are  shown  in  Tables  XLIX.  and  L. 

These  specimens  of  the  aurelia  form  show  the  same  relations 
that  are  found  in  the  caudatum  form,  with  one  exception.  In  lot  5 
(Table  X.,  row  36)  the  mean  breadth  of  the  specimens  beginning 
fission  is  less  than  that  of  the  random  sample,  instead  of  greater  as 
in  all  other  cases.  But  this  peculiarity  is  due  to  environmental  con- 
ditions. In  lot  5  the  breadth  was  very  great  in  proportion  to  the 
length,  as  is  shown  by  the  dimensions  of  the  random  sample  (Table 
L.,  and  row  37,  Table  X.).  In  this  lot  the  breadth  was  41.555  per 
cent,  of  the  length,  while  in  most  cases  it  is  near  to  30  per  cent. 
This  was  due  to  the  recent  transference  of  the  animals  to  a  nutritive 
solution;  they  became  very  plump.  Evidently,  when  preparing  to 
divide  the  body  tends  to  return  to  a  constant  form;  in  this  case, 
therefore,  it  becomes  narrower  instead  of  broader. 

In  the  specimens  of  the  aurelia  form,  as  in  the  caudatum  form, 
all  dimensions  are  less  variable  in  the  specimens  beginning  fission. 
This  difference  in  variability,  as  compared  with  the  random  samples, 
is  very  great  in  some  cases.  Thus,  while  the  coefficients  of  variation 
in  length  for  the  random  samples  of  lots  4  and  5  are  15.279  and 
10.643,  fo^  those  of  the  same  lots  beginning  fission  they  are  but  7.541 
and  6.862,  respectively.  Had  we  included  in  the  lots  beginning  fission 
only  specimens  in  which  the  depth  of  constriction  was  still  less,  the 
coefficients  of  variation  would  have  been  still  smaller. 

The  constants  for  all  specimens  of  c  that  are  beginning  fission, 
taken  together,  are  shown  in  row  38,  Table  X.  The  standard  devia- 
tions and  coefficients  of  variation  are,  of  course,  greater  than  for 

■In  making  these  measurements  of  descendants  of  c,  a  higher  power  of 
the  microscope  was  used,  so  that  the  single  unit  of  measurement  was  3J 
microns.  This  caused  the  tables  (in  the  appendix)  to  take  a  somewhat  differ- 
ent appearance  from  those  of  the  descendants  of  D. 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  447 

each  of  the  two  component  lots  taken  separately,  since  the  two  lots 
diflfered  as  a  result  of  different  environmental  conditions. 

(n)  Later  Stages  of  Fission, — As  the  constriction  deepens  the 
animal  as  a  whole  becomes  more  elongated,  while  the  breadth  de- 
creases slightly.  These  relations  are  shown  both  for  the  descendants 
oi  D  (caudatunt  form)  and  the  descendants  of  c  (aurelia  form)  in 
Table  X.  (rows  25  and  26;  30  and  31 ;  33  and  34).  In  the  large  lot 
I  of  dividing  descendants  of  D,  comprising  313  specimens  (Table 
XL)  the  correlation  between  length  of  body  and  depth  of  constric- 
tion below  the  surface  is  .6882.  The  length  increases  8.6  microns 
with  every  increase  of  10  microns  in  the  depth  of  constriction.  The 
correlation  between  breadth  and  depth  of  constriction  (Table  XII.) 
is  —  .5232,  the  breadth  decreasing  2.63  microns  for  each  10  microns 
increase  in  depth  of  constriction.  If  we  include  only  the  specimens 
in  which  lengthening  has  decidedly  begun  (thus  omitting  the  earliest 
stages,  in  the  uppermost  rows  of  Tables  XL  and  XII.),  then  the 
correlation  between  length  and  depth  of  constriction  is  .7818;  between 
breadth  and  depth  of  constriction,  — .3316.  With  an  increase  of  10 
microns  in  depth  of  constriction  the  length  now  increases  11. 195 
microns,  while  the  breadth  decreases  1.252  microns.  In  this  same 
culture  while  the  mean  length  of  the  131  specimens  beginning  fission 
is  175.696  microns,  that  of  the  seven  specimens  having  a  connecting 
portion  but  4  microns  wide  is  212.572  microns.  Thus,  the  increase 
in  length  before  separation  takes  place  is  36.876  microns,  or  about 
21  per  cent,  of  the  length  at  the  time  fission  begins.  The  breadth 
has  decreased  from  55.480  microns  at  the  beginning  of  fission  to 
43.428  microns  in  the  seven  specimens  with  the  narrowest  connec- 
tions— a  decrease  of  about  21  per  cent.  The  ratio  of  breadth  to 
length  decreases  from  31.568  per  cent,  at  the  beginning  of  fission  to 
20.430  per  cent,  just  before  separation. 

Corresponding  relations  are  shown  in  other  lots  of  dividing  speci- 
mens ;  some  of  the  data  are  given  in  Table  X. 

2.  Summary  on  Growth  in  Paramecium  with  a  Growth  Curve. 

We  have  thus  followed  the  growth  from  the  time  when  the  indi- 
vidual is  but  half  a  constricting  specimen  to  the  period  when  it  is 
again  ready  to  separate  into  two  new  individuals.     We  are  ready. 


448  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a^, 

therefore,  to  outline  the  main  features  of  the  growth  of  Paramecium, 
and  to  construct  curves  which  shall  give  an  idea  of  the  processes 
involved.  In  spite  of  an  incredible  amount  of  work  devoted  to  col- 
lecting the  data,  certain  of  the  less  important  features  of  the  growth 
curves  must  remain  obscure,  but  the  main  facts  are  clear. 

The  main  outlines  of  the  changes  due  to  growth  are  as  follows : 
From  the  time  the  constriction  appears  in  the  mother  until  a  few 
minutes  after  separation  takes  place,  the  length  increases  rapidly, 
while  the  breadth  decreases  a  little.  A  few  minutes  after  separation 
the  processes  become  less  rapid.  The  breadth  soon  reaches  its  mini- 
mum, then  begins  to  increase  like  the  length,  though  more  slowly. 
Growth  in  length  continues  for  at  least  eighteen  hours;  the  time 
undoubtedly  varies  with  the  conditions.  The  breadth  continues  to 
increase  for  some  time,  but  it  undergoes  marked  fluctuations,  due  to 
environmental  conditions.  In  lot  lo  (Table  X.)  it  decreased  between 
the  ages  of  12  and  18  hours;  this  is  probably  an  environmental  effect, 
not  one  due  to  the  normal  growth  processes. 

As  the  time  for  fission  approaches  the  animals  are  considerably 
more  than  twice  as  long  as  the  original  halves  from  which  they  devel- 
oped. Now  as  fission  comes  on  they  shorten  and  thicken,  all  tending 
to  approach  a  uniform  length  and  thickness.  There  is  thus  much 
less  variation  in  the  dimensions  at  the  beginning  of  fission  than  in 
specimens  taken  at  random.  Now  the  constriction  appears  and  the 
animal  begins  to  narrow  and  extend  in  the  way  already  described, 
finally  separating  into  two  parts. 

If  from  our  data  we  construct  curves  showing  these  changes,  we 
get  such  results  as  are  shown  in  Diagram  5. 

Method  of  Constructing  the  Curves, — The  horizontal  scale  repre- 
sents the  time  in  hours,  while  the  vertical  scale  represents  the  meas- 
urements of  the  animals  in  microns.  The  upper  curve  shows  the 
length,  the  lower  one  breadth,  as  measured  from  the  base  line. 
Fission  is  assumed  to  take  place  once  in  twenty- four  hours,  which  is 
an  approximation  to  a  rate  commonly  occurring.  The  time  between 
the  appearance  of  the  constriction  and  the  actual  separation  of  the 
two  halves  is  taken  as  one  half  hour. 

The  relative  distances  of  the  two  curves  from  the  base  line  shows 
the  relative  dimensions  of  length  and  breadth.     The  vertical  rise  of 


i9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA 


449 


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450  JENNINGS— HEREDITY   IN    PROTOZOA.  lApril  a4, 

the  curve  of  length  shows  the  actual  proportion  of  growth  to  the 
original  length.  The  distance  from  the  base  to  the  curves  is  357 
times  the  actual  dimension  at  the  given  time. 

In  order  to  show  changes  due  to  growth  alone  all  the  data  for  such  a 
curve  should  be  measurements  from  a  single  uniform  culture  on  a  single  day ; 
otherwise  environmental  differences  complicate  the  matter,  as  we  shall  see 
more  clearly  in  the  next  division  of  this  paper.  Now,  it  is  impracticable  to 
obtain  from  a  single  culture  on  a  single  day  measurements  of  all  the  required 
stages.  We  are  compelled  therefore  to  make  certain  corrections  in  some  of 
the  measurements,  to  compensate  so  far  as  we  can  for  environmental  differ- 
ences. As  Table  X.  shows,  the  mean  dimensions  of  random  samples  differ 
much  in  (for  examples)  lots  i  (row  3)  and  6  (row  12).  It  will  not  do, 
therefore,  to  compare  directly  the  young  of  these  two  lots.  Since  we  have 
from  lot  6  the  greatest  number  of  different  stages,  it  is  best  to  make  the 
measurements  from  this  the  basis  for  the  curve,  correcting  others,  so  far  as 
possible,  to  compare  with  this.  In  lot  2  the  mean  length  (Table  X.,  row  6) 
is  almost  exactly  the  same  as  for  lot  6,  so  that  we  may  use  the  measurements 
of  lot  2  without  correction,  so  far  as  length  is  concerned.  On  this  account 
we  shall  employ  lot  2  for  the  earliest  stages,  in  place  of  lot  i,  though  the 
latter  is  based  on  a  larger  number  of  specimens. 

Since  the  mean  breadth  of  the  sample  of  lot  6  is  64.880  microns,  while 
that  of  lot  2  is  but  46.020  microns,  it  is  necessary  to  correct  the  breadth  for 
lot  2.  At  first  thought  it  would  seem  that  the  proper  method  of  making: 
this  correction  would  be  by  multiplying  the  breadths  of  the  different  sets  of 
lot  2  by  the  ratio  64.880/46.020.  This  would  be  the  proper  method  of  pro- 
cedure if  we  were  dealing  with  the  same  stages  of  growth  in  the  two  lots; 
the  specimens  of  lot  2  would  be  made  plump,  like  those  of  lot  6.  But  the 
stage  with  which  we  are  dealing  is  that  of  the  beginning  of  fission.  Now, 
we  have  already  seen  that  when  the  specimens  not  dividing  are  plump,  the 
breadth  does  not  increase  at  the  approach  of  fission  nearly  so  much  as 
when  the  specimens  not  dividing  are  thin.  Indeed,  if  the  specimens  are  very 
plump,  there  is  an  actual  decrease,  instead  of  an  increase,  at  the  approach 
of  fission.  Our  problem  is:  What  would  be  the  breadth  of  specimens  be- 
ginning fission,  in  which  the  length  is  82.600,  and  the  animals  are  very  plump, 
as  in  lot  6?  This  problem  can  best  be  solved  by  asking  what  is  the  ratio 
of  breadth  to  length  in  specimens  beginning  fission,  in  a  very  plump  culture? 
In  lot  3  (row  7,  Table  VIII.)  we  have  such  a  plump  culture,  and  we  find  that 
the  ratio  of  breadth  to  length  is,  in  the  earliest  stage  of  fission,  78.563  per 
cent.  We  therefore  take  this  as  the  ratio  of  breadth  to  length  for  the 
earliest  stage  of  lot  2,  from  which  the  corrected  breadth  is  found  to  be  64.893. 
If  this  decreases  at  the  same  relative  rate  as  actually  occurred  in  lot  2,  then 
the  breadth  15  minutes  after  the  beginning  of  constriction  would  be  64493 
microns. 

We  are  compelled  to  use,  further,  lots  9  and  10  (Table  X.).  In  lot  9 
both  length  and  breadth  require  correction  to  make  them  comparable  with 
the  measurements  of  lot  6.     The  correction  is  made  by  multiplying  the 


i9o8.] 


JENNINGS— HEREDITY   IN   PROTOZOA. 


451 


dimensions  by  the  ratio  between  the  length  of  the  random  samples  of  the 
two  lots.  In  lot  9  we  use  only  the  average  of  the  two  sets,  as  given  in  row 
18,  Table  X. 

In  lot  10,  since  we  unfortunately  have  no  random  sample,  we  are  unable 
to  make  a  correction. 

Owing  to  the  very  great  difference  in  the  environmental  conditions  of 
lot  3  (rows  23  and  24,  Table  X.)  we  are  unable  to  use  the  24-hour-old 
specimens  of  that  lot,  although  we  need  measurements  at  that  age.  The 
older  portions  of  the  curve  (beyond  18  hours,  at  the  right)  cannot  be 
plotted  from  exact  data,  and  there  are  certain  features  of  much  importance 
for  which  it  appears  that  the  collection  of  such  data  would  be  almost 
impossible.  As  we  have  shown,  before  fission  the  animals  shorten  and 
thicken.  How  long  before  fission  this  begins  it  is  not  possible  to  say;  in 
making  the  curve  the  period  is  arbitrarily  taken  as  two  hours. 

When  we  make  the  corrections  above  described,  we  have  the  following 
mean  dimensions  at  different  ages,  as  data  for  the  construction  of  our  curve. 
The  ages  given  are  the  average  ages  for  the  lots  considered;  thus  the  age 
for  row  8,  Table  X.  (18  to  28  minutes)  is  taken  as  23  minutes. 


Table  XV. 

Dimensions  in  Microns  of  Paramecia  {Descendants  of  D)  at  Different  Ages, 

Corrected  (so  far  as  possible)  to  Correspond  with  Those  of  Lot  6, 

Table  X,    Data  used  in  making  the  Curves  of  Growth. 


Age. 


Beginning  constriction 

Fifteen  minutes  after  beginning 

constriction 

2^  minutes  after  separation 

9^  minutes 

23  minutes 

40  minutes 

82^  minutes 

4  hours 

12  hours 

18  hours 

Beginning  constriction 


Lot. 


Mean  Length 
in  Microns. 


«« 
« 
«< 
(I 
tt 
t« 
<« 
ti 
(« 
<( 


5» 
5» 
7. 
8. 

9, 
10, 

18, 

20, 

22, 

50» 


able 

tVIII. 

82.600 

64.893 

«i 

VIII. 

85-774 

64493 

i< 

X. 

107.660 

59.355 

tt 

X. 

128.000 

60.168 

<< 

X. 

143348 

54.284 

<i 

X. 

149.920 

55.840 

t< 

X. 

161.524 

54. 192 

t< 

X. 

176.560 

58.922 

(« 

X. 

188.988 

62.796 

<< 

X. 

199.048 

56.496 

<< 

X. 

165.200 

64.893 

Mean  Breadth 
in  Microns. 


When  we  lay  off  on  the  vertical  scale  the  distances  corresponding  to  the 
lengths  and  breadths  at  the  different  periods,  as  given  in  the  above  table, 
and  connect  these  points,  we  obtain  the  curves  given  in  Diagram  5. 

Characteristics  of  the  Curves. — As  the  curves  show,  the  length 
increases  with  great  rapidity  for  about  twenty  minutes  after  fission; 
continues  less  rapidly  for  about  an  hour,  and  still  less  rapidly  for 
four  or  five  hours.  Now  the  increase  continues,  though  very  slowly, 
till  a  maximum  is  reached  at  a  length  considerably  greater  than  twice 
the  original  length;  later  the  length  decreases  in  preparation  for 


452  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  24, 

fission;  this  decrease  continues  till  the  length  is  just  twice  the  original 
length.  Now  the  constriction  appears,  so  that  the  animal  may  be 
looked  on  as  two;  the  length,  therefore,  drops  in  a  straight  line  to 
the  original  length  found  at  the  beginning  of  the  curve.  The 
breadth  decreases  from  the  beginning  till  about  an  hour  after  fission ; 
then  slowly  increases;  it  shows  in  the  course  of  the  twenty- four 
hours  many  fluctuations  which  are  doubtless  mainly  due  to  diflFerences 
in  the  environment — especially  to  differences  in  the  amount  of  food 
taken.  In  preparation  for  fission  the  breadth  increases  at  the  same 
time  that  the  length  decreases. 

The  curve  of  length  is  much  the  more  interesting  of  the  two,  since 
it  is  the  one  which  represents  mainly  the  actual  growth.  It  is  of 
great  interest  to  find  that  this  curve  of  growth  in  a  single  cell  is  of 
essentially  the  same  form  and  character  as  those  which  have  been 
obtained  for  the  growth  of  many  higher  organisms,  composed  of 
many  cells.  A  number  of  such  curves  are  brought  together  in  the 
recent  interesting  paper  of  Robertson  (1908).  Inspection  shows  at 
once  that  the  curve  of  growth  in  Parmecium  closely  resembles  that 
for  growth  of  the  rat,  as  worked  out  by  Donaldson  ( 1906)  ;  for 
growth  of  man,  and  for  gro)vth  in  various  other  organisms. 

The  curve  of  growth,  as  is  well  known,  is  a  logarithmic  curve 
in  the  cases  where  it  has  been  worked  out  mathematically.  While 
the  growth  in  Paramecium  has  merely  been  plotted  empirically,  it  is 
evident  that  it  is  essentially  a  similar  logarithmic  curve;  this  could 
doubtless  be  worked  out  from  the  data  given. 

The  fact  that  the  curve  of  growth  is  essentially  the  same  in  the 
unicellular  organism  as  in  the  animal  composed  of  millions  of  cells 
is  in  some  respects  surprising.  In  the  brain  of  the  rat,  or  in  its  body, 
the  curve  of  growth  is  the  resultant  of  the  growth  of  many  different 
groups  of  cells,  some  groups  growing  at  one  period,  some  at  another ; 
yet  the  resultant  curves  are  of  the  same  character  as  when  there  is 
growth  in  but  a  single  cell. 

The  temporal  relations  shown  in  the  curves  are  likewise  of  much 
interest.  As  our  diagram  shows,  that  portion  of  the  curve  showing 
the  greatest  curvature  requires  in  Paramecium  about  four  hours 
from  the  beginning.  In  the  rat  the  corresponding  part  of  the  curve 
takes  several  months,  while  in  man  it  requires  several  years.     It 


,9o8j  JENNINGS— HEREDITY  IN   PROTOZOA.  453 

seems  extraordinary  that  a  process  following  the  same  laws  should 
in  some  cases  be  measured  by  hours,  in  other  cases  by  m6nths,  in 
others  by  years. 

3.  Effects  of  Growth  on  the  Observed  Variation. 

A  random  sample  of  an  ordinary  culture  of  Paramecium  contains 
specimens  falling  in  all  parts  of  the  growth  curves  represented  in 
Diagram  5.  If  we  measure  the  various  members  of  such  a  sample, 
as  was  done  by  Pearl  (1907),  we  shall  then  find  many  variations  in 
size,  which  variations  consist  to  a  considerable  extent  of  diflFerent 
growth  stages.  Not  all  the  observed  variations  are  due  to  this 
factor,  but  its  importance  is  very  considerable.  This  will  best  be 
appreciated  by  running  through  the  columns  headed  "  coefficients  of 
variation"  in  Table  X.  If  we  take  samples  including  specimens 
falling  in  the  early  parts  of  the  growth  curve,  when  the  absolute  size 
is  small  but  the  changes  with  growth  are  very  marked,  then  the 
coefficients  of  variation  in  length  are  high ;  thus  in  rows  4  and  5  they 
are  15.494  and  13.729,  respectively,  while  in  the  random  sample  of 
the  same  culture  the  coefficient  is  but  8.834  (row  6).  On  the  other 
hand,  if  we  take  specimens  restricted  to  a  very  small  portion  of  the 
curve,  the  coefficient  of  variation  becomes  very  low;  thus  in  a  lot 
whose  age  falls  between  18  and  28  minutes  the  variation  is  but  4.521 
(row  8)  ;  at  the  age  of  4.20  to  5  hours  is  5.043  (row  17),  though  the 
variation  for  a  random  sample  of  this  same  culture  is  13.262  (row  19). 
The  effects  of  growth  on  variation  are  shown  to  the  eye  in  Diagram 

4,  p.  440.  • 

Variation  at  Fission, — iThe  effects  of  growth  on  the  observed 
variation  are  likewise  seen  when  we  compare  random  samples  with 
individuals  that  are  at  a  definite  stage  in  the  life  history.  Thus,  if 
we  fake  specimens  at  the  beginning  of  fission,  when  the  constriction 
first  appears,  we  find  the  coefficient  of  variation  very  low,  as  com- 
pared with  those  of  random  samples  of  the  same  cultures.  This  is 
readily  seen  in  the  following  tabulation  of  the  coefficients  of  varia- 
tion for  the  four  cultures  of  Table  X.  in  which  the  specimens  begin- 
ning fission  were  studied  (see  next  page). 

Variation  in  Conjugants. — Again,  the  same  thing  appears  when 
we  compare  conjugating  individuals  with  random  samples  of  the 


454 


JENNINGS- HEREDITY   IN   PROTOZOA. 


[April  34, 


same  cultures.  Conjugation  does  not  occur  till  a  certain  stage  of 
growth  has  been  reached,  and  the  conjugants  do  not  include  speci- 
mens undergoing  the  changes  preparatory  to  fission.  The  conjugants 
would  then  fall  in  those  portions  of  the  growth  curve  that  are  nearly 
straight ;  that  is,  there  would  be  in  these  little  variation  due  to  growth. 

Table  XVI. 
Coefficients  of  Variation. 


Length. 

Breadth. 

Lot. 

Bennning 
Fission. 

Random  Sample. 

Bqpnning 
Fission. 

Random  Sample. 

I 

2 

4 
5 

5.368 
5.320 

7.541 
6.862 

7.765 

8.834 

15.279 
10.643 

9.082 
6.769 

9.91 1 
12.071 

12.877 

II.42I 
15.683 
13.720 

Pearl  (1907)  has  already  shown  that  the  observed  variability  of  con- 
jugants is  less  than  that  of  random  samples  of  the  same  culture.  I 
have  made  extensive  studies  of  conjugants  and  find  the  same  thing. 
Details  regarding  the  relation  of  conjugation  to  variation  and  heredity 
are  to  be  taken  up  in  a  later  communication ;  here  I  give  merely  the 
coefficients  of  variation  for  certain  cases,  as  compared  with  those  of 
random  samples. 

Table  XVII. 

Coefficients  of  Variation  for  Conjugants,  as  compared  with  those  for  random 

samples  of  non-conjugants  of  the  same  culture. 


Lot. 

Length. 

Breadth. 

Conjugants. 

Non-Conjugants. 

8.185 

9-123 
11.578 
11.026 

Conjugants. 

Non-Conjugants. 

Ay  Pearl. 

C,      " 

a,  Jennings. 

6.668 

7-439 

7.39* 
7.678 

9-398 

7.910 

12.409 

15.766 

II. 112 
10.894 
19.176 
18.142 

On  comparing  the  coefficients  of  variation  in  conjugants,  as  given 
in  Table  XVII.,  with  those  for  specimens  beginning  fission  (Table 
XVI.),  and  those  for  specimens  at  definite  ages  (Table  X.),  it  is 
found  that  in  the  conjugants  the  variation  is  not  so  small  as  it  is  in 
specimens  at  definite  growth  stages.  This  shows  clearly  that  nothing 
is  required  to  explain  the  low  variation  of  conjugants,  save  the  fact 
that  a  certain  number  of  growth  stages  (the  earlier  and  later  ones) 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  455 

are  lacking  in  these.  There  is  no  evidence  of  an  unusually  low 
degree  of  congenital  variation  in  the  conjugants,  for  the  non-conju- 
gating specimens  beginning  fission  show  a  still  lower  variability 
(Table  XVI.). 

It  appears  highly  probable  that  if  we  could  examine  a  large 
number  of  individuals,  derived  from  the  same  parent,  cultivated 
under  identically  the  same  conditions,  and  all  in  precisely  the  same 
stage  of  growth,  we  should  find  coefficients  of  variation  considerably 
smaller  than  the  smallest  we  have  found,  which  is  4.521  (row  8, 
Table  X.).  Indeed,  if  we  could  further  exclude  all  inaccuracies  of 
measurement,  it  is  quite  possible  that  the  coefficient  of  variation 
would  approach  closely  to  zero,  if  it  did  not  reach  it  completely. 
This  would,  of  course,  mean  that  the  variations  observed  among  the 
progeny  of  a  single  individual  are  not  congenital,  but  are  all  due  to 
growth  and  environmental  action.  Further  evidence  of  this  will 
come  out  later  in  this  paper. 

4.  Effects  of  Growth  on  the  Observed  Correlation  Between 
Length  and  Breadth. 

As  Diagram  5  shows,  the  curves  of  length  and  breadth  diverge 
at  the  beginning,  then  run  for  a  considerable  distance  nearly  parallel, 
then  finally  approach  each  other.  That  is,  at  first  the  breadth  decreases 
while  the  length  increases;  later  they  increase  together;  and  still 
later  the  breadth  increases  while  the  length  decreases.  If  a  collec- 
tion of  specimens  includes  individuals  in  various  different  stages  of 
growth  (as  is  usually  the  case),  then  these  various  relations  of 
breadth  to  length  will  deeply  affect  the  amount  of  correlation  observed 
between  the  two  dimensions. 

Thus,  if  we  take  a  collection  composed  of  various  ages  under  one 
hour,  when  the  length  is  increasing  while  the  breadth  is  decreasing, 
then  on  the  whole  greater  length  will  be  associated  with  less  breadth, 
so  that  the  correlation  between  them  will  tend  to  be  negative.  This 
is  the  explanation  of  the  negative  correlation  shown  in  Table  X., 
rows  2,  4,  5,  7,  II,  13,  14.  Next  follows  a  period  (from  about  the 
end  of  the  first  hour  to  the  fourth)  in  which  the  inclusion  of  indi- 
viduals of  different  ages  tends  to  cause  a  certain  degree  of  positive 
correlation,  since  the  two  dimensions  are  increasing  together.     Then 


456  JENNINGS—HEREDITY   IN   PROTOZOA.  [April  24, 

comes  a  long  period  in  which  both  dimensions  remain  nearly  the 
same — the  length  increasing  slowly^  while  the  breadth  fluctuates. 
Different  growth  stages  during  this  period  have  little  marked  eflfect 
on  the  coefficient  of  correlation  between  length  and  breadth;  they 
tend  to  prevent  its  reaching  i.ooo,  but  this  it  would  not  reach  for 
other  reasons. 

Now,  for  a  certain  period  before  fission  (taken  as  two  hours,  in 
.  our  curves) ,  the  length  decreases  while  the  breadth  increases.  Greater 
breadth  will  then  be  associated  with  less  length,  tending  to  produce 
again  a  negative  correlation.  If  we  make  a  collection  of  individuals 
representing  various  stages  in  this  process,  we  should,  therefore, 
expect  to  find  the  correlation  much  less  than  in  collections  taken  ( i ) 
either  before  these  processes  have  begun,  or  (2)  after  they  are 
ended.  We  can  realize  this,  in  the  main,  by  taking  from  a  large 
random  sample  all  the  largest  specimens  (which  are,  of  course,  the 
older  ones)  and  combining  these  into  a  single  correlation  table  with 
specimens  from  the  same  culture  that  are  beginning  fission  (the 
oldest  specimens  of  the  culture).  I  performed  this  operation  for 
lot  I  of  Table  X.  This  collection  contains  131  specimens  beginning 
fission  (row  25,  Table  X.),  and  134  specimens  (not  dividing)  that 
are  196  microns,  or  more,  in  length  (row  28,  Table  X.)  ;  throwing 
these  together,  we  have  a  collection  of  264  of  the  oldest  specimens  in 
the  culture  (row  29,  Table  X.).  For  the  131  specimens  beginning 
fission  the  coefficient  of  correlation  is  -|--6S46;  for  the  134  large 
specimens  it  is  -f-  4681.  When  the  two  are  taken  together  the  corre- 
lation disappears.  The  computation  gives  us  a  coefficient  of  -f-  -0350, 
but  this  is  less  than  its  probable  error  (.0415),  so  that  the  figures 
have  no  significance ;  no  correlation  appears. 

The  effects  of  the  inclusion  of  various  growth  stages  on  the 
observed  correlation  shows  itself  in  many  other  ways,  which  will 
become  evident  to  anyone  who  carefully  examines  the  data  of  Table 
X..  in  connection  with  our  curves  of  growth  (Diagram  5),  and  the 
relations  brought  out  in  the  foregoing  paragraphs.  Note,  for  exam- 
ple, the  coefficients  of  correlation  for  lot  9  (rows  16-18,  Table  X.). 
For  the  specimens  3  to  4  hours  old  the  coefficient  is  but  .3201,  and 
for  those  4.20  to  5  hours  old  it  is  .5557.  When  we  throw  these  two 
lots  together,  so  as  to  include  a  much  greater  proportion  of  the 


i9o8.)  JENNINGS— HEREDITY  IN   PROTOZOA.  457 

growth  curve,  the  correlation  rises  to  .7132.  In  this  larger  collection 
the  short  specimens  are  much  the  narrower,  the  large  specimens  much 
broader — ^giving  high  positive  correlation.  Slight  changes  in  one 
dimension  may  not  be  accompanied  by  notable  changes  in  the  other, 
while  great  changes  in  one  are  always  accompanied  by  changes  in  the 
other.    This  is  a  relation  which  we  shall  meet  again. 

While  thus  growth  has  a  very  great  effect  on  the  correlation  to 
be  computed  from  the  measurements  of  a  collection  of  Paramecia, 
it  is  important  to  bear  in  mind  the  fact  that  it  is  by  no  means  the 
only  factor  concerned  in  correlation.  This  becomes  evident  as  soon 
as  we  take  a  collection  in  which  the  specimens  are  all  in  nearly  the 
same  stage  of  growth;  the  coefficient  of  correlation  is  then  high. 
This  is  perhaps  best  realized  by  considering  specimens  in  the  begin- 
ning of  fission.  As  we  have  before  noticed,  in  the  collection  of  131 
specimens  beginning  fission,  from  lot  i,  great  pains  were  taken  to 
include  only  a  single  stage  in  the  process.  This  collection  gives  a 
high  positive  correlation  of  .6546.  This  correlation  can  be  due  only 
to  the  fact  that  in  specimens  at  a  single  growth  stage  the  length  and 
breadth  tend  to  bear  a  certain  proportion  to  each  other.  The  effects 
of  this  are  clearly  seen  in  many  other  collections  of  Table  X.  Thus, 
in  rows  8,  9  and  15  the  specimens  all  fall  in  the  period  when  length 
is  increasing  while  breadth  is  decreasing ;  yet  there  is  in  each  case  a 
small  positive  correlation.  This  is  due  to  the  fact  that  the  period  of 
growth  over  which  each  collection  extended  was  small,  so  that  the 
negative  correlation  due  to  growth  was  more  than  counterbalanced 
by  the  inherent  proportionality  of  length  to  breadth.  A  collection 
including  only  specimens  that  were  all  in  the  same  stage  of  growth 
would  undoubtedly  (other  things  begin  equal)  show  a  high  corre- 
lation between  length  and  breadth,  no  matter  what  point  on  the 
growth  curves  they  represented.  This  signifies,  of  course,  that  in 
any  given  stage  of  growth  the  relation  of  length  to  breadth  tends  to 
be  the  same  in  all  specimens — although  in  different  stages  of  growth 
this  is  often  not  the  case.  Other  factors  which  modify  the  correla- 
tion will  be  considered  in  the  later  sections  of  this  paper ;  a  summary 
of  all  these  factors  will  be  presented  in  a  special  section. 

With  this  we  conclude  our  study  of  growth  in  Paramecium; 

PROC.  AMER.  PHIL.  SOC.  XLVII.   I90  DD,  PRINTED  JANUARY  II,   I909. 


468  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  24. 

being  prepared  to  understand  the  part  played  by  this  in  the  observed 
variations  and  correlations,  we  may  pass  to  other  factors  afifecting 
these. 

IV.  THE  EFFECTS  OF  ENVIRONMENTAL  CONDITIONS 
ON  DIMENSIONS,  VARIATION  AND  CORRELATION. 

The  data  for  the  study  of  growth,  just  concluded,  show  inci- 
dentally that  environmental  conditions  affect  profoundly  the  dimen- 
sions, variation  and  correlation  in  Paramecium.  As  we  have  seen, 
samples  taken  from  the  same  culture  on  two  successive  days  are  not 
strictly  comparable  for  determining  matters  relating  to  growth, 
because  of  the  environmental  changes  from  day  to  day,  inducing 
marked  changes  in  the  organisms.  Thus,  in  a  given  culture  we  found 
that  the  mean  length  at  the  age  of  li  to  li  hours  was  161.524  microns ; 
three  days  later  specimens  more  than  twice  as  old,  from  the  same 
culture,  were  smaller,  measuring  but  149.636  microns.  We  wish 
now  to  investigate  the  causes  of  such  diflferences. 

We  shall  not  attempt  at  present  a  systematic  investigation  of  the 
effects  of  different  chemical  and  physical  agents  on  size,  form  and 
variation,  though  this  is  a  matter  which  much  needs  study.  Our 
present  object  is  rather  to  examine  the  eflfects  of  altered  nutritional 
conditions  and  of  the  commoner  "favorable"  and  "unfavorable" 
conditions.  We  shall  study  the  variations  from  the  standpoint  of 
interest  in  the  organism  rather  than  in  the  agents  inducing  them,  the 
purpose  being  to  form  a  conception  of  the  changes  which  may  be 
looked  for  in  Paramecium  as  a  result  of  common  alterations,  mainly 
nutritional,  in  its  cultural  conditions.  One  of  the  results  of  this  study 
will  be  to  show  that  we  cannot  assign  a  definite  effect  to  each  agent 
taken  in  any  absolute  way.  What  effect  a  given  agent  will  have 
depends  on  the  previous  condition  of  the  organisms  on  which  it  acts. 
The  same  agent  produces  at  one  time  an  increase  in  size,  at  another 
a  decrease;  at  one  time  it  increases  the  variability;  at  another  it 
decreases  it.  A  given  agent  may  either  increase  the  positive  corre- 
lation between  length  and  breadth,  or  it  may  decrease  it  or  convert 
it  into  a  negative  correlation.  In  succeeding  days  the  same  agent 
may  produce  these  diverse  effects  on  the  same  set  of  Paramecia. 


,5^.]  JENNINGS— HEREDITY   IN   PROTOZOA.     .  459 

Yet,  of  course,  these  results  are  not  produced  haphazard ;  what  we 
wish  to  study  are  the  laws  they  follow. 

The  effects  of  the  environment  were  studied  mainly  on  the  same 
animals  that  served  for  the  study  of  growth.  Two  strains  were 
used;  one  consisted  of  descendants  of  the  individual  D,  of  the  cau- 
datum  form,  the  other  of  descendants  of  c  (aurelia  form).  The 
results  show  the  extent  of  the  variations  producible  through  environ- 
mental action  in  the  progeny  of  single  individuals  multiplying  by 
fission.  No  conjugation  occurred  in  the  D  strain  during  the  time  it 
was  under  experimentation.  On  a  given  date,  therefore,  the  age  of 
the  individuals,  as  measured  in  generations  of  the  "  cycle,"  was  about 
the  same. 

Table  XVIII.  gives  a  summary  of  the  statistical  results  in  the 
experiments  on  the  effects  of  the  environment ;  it  will  be  referred  to 
frequently  in  the  following  account  (see  next  page). 

I.   Progeny  of  D  (caudatum  Form). 

The  individual  D  was  isolated  April  12,  1907;  it  measured,  as 
nearly  as  could  be  determined  when  alive,  about  250  microns.  It 
was  placed  in  culture  fluid  made  of  boiled  hay  and  the  progeny  were 
kept  in  such  cultures  for  months.  Characteristic  progeny  of  D  are 
shown  in  Fig.  i,atod. 

The  experiments  with  the  descendants  of  D  may  be  divided  into 
three  series. 

First  Series. 

Old  Large  Culture, — On  June  11  a  sample  of  100  of  the  descend- 
ants oiD  was  killed,  from  a  hay  culture  that  had  stood  several  weeks 
and  was  flourishing,  though  multiplication  was  not  occurring  actively. 
This  culture  was  in  a  vessel  about  nine  inches  across.  The  measure- 
ments of  this  sample  are  given  in  Table  V.  (page  406),  while  the 
constants  are  found  in  row  i,  Table  XVIII. 

Effects  of  Fresh  Hay  Infusion. — Three  days  after  these  measure- 
ments were  taken,  a  number  of  individuals  of  this  culture  were 
removed  and  placed  in  a  fresh  hay  infusion,  in  a  watch-glass;  in 
this  they  were  allowed  to  remain  24  hours.  The  increased  food  in 
the  fresh  infusion  caused  them  to  increase  much  in  breadth  (from 
49.000  microns  to  64.880  microns),  and  at  the  same  time  to  begin  to 


460 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[Apiiia^, 


s 


I 

2 

3 

4 

5 

6 


7 
8 

9 
lo 

It 

12 

13 
14 

15 

i6 
17 


Table  XVIII. 

Effects  of  Environmental  Conditions  on  Dimensions  and  Constants  of  Varia- 
same  culture  at  the  same  time  (except  in  rows  12,  15  and  20).  The 
appendix  or  elsewhere,  in  which  fuller  data  are  given  for  the  lot  in 


A.  Progtny  p/D, 
First  Strits. 


RaDdom  sample  of  Z>,  June  1 1, 

1907 

Same  after  24   hours  in  fresh 

hay  infusion,  June  15 

Two  days  after  last;  culture 
fluid  not  renewed,  June  17... 

Same,  after  24  hours  in  fresh 
hay  infusion.  Rapid  multi- 
plication, June  18 

Same,  one  week  later;  bac- 
teria multiplied  injuriously, 
June  25 

Starvation,  same  as  row  2,  but 
left  X I  days  in  small  quantity 
of  fluid,  June  25 

Second  Series. 

24  hours  in  fresh  hay  infusion ; 

rapid  multiplication,  July  17.. 
Same  as  last,   but  starved   a 

week,  July  24 

Same  as  last,  but  24  hours  in 

fresh  hay  infusion,  July  25... 
Same  as  last,  but  kept  I  week 

without  change  of  fluid,  July 

31 

Same  as  last,  but  kept  48  hours 

in  fresh  hay  infusion,  Aug.  3.. 

Rows  8,  10  and  il  combined... 

Third  Series, 

Slender,  old  culture,  in  large 
jar,  September  15 

Same  as  last,  after  48  hours  in 
fresh  hay  infusion,  Septem- 
ber 15 

Rows  13  and  14,  combined 

B.  Progeny  of  c. 

Random  sample  of  r,  June  11, 

1907 

Random  sample  of  r,  August  9.. 


3-6 

2i^ 


100 


100 


135 


19s 


178 


100 


200 
150 

350 

150 

1150 
450 


100 


100 

200 


ICO 

100 


.0 


5 

51 
6 


52 


53 


30 
19 


21 
22 


54 


55 


Length. 


Mean  in 
Microns. 


Standard 

Deviation 

in  Microns. 


Coefficient  of 
Variation. 


i88.36o±  .980 

i84.68ort  .848  I2.596d=  .600 

i85.oo8±  .836  I4.420±  .592 


I4.532±  .692  ^715^.370 

6.821  zb.327 

7.794^.324 


176.124^=1.12823.360^  .797 


2oi.888d:i.i47 


22.680zfc  .81 X 


i49.36ozb  .736  io.896db  .520 


I  .13  o  .776 
i46.io8zb  .563 


i6.264±:  .548 
1 0.228 dr  .398 


20  163.932^  .754  2o.928d=  .533 


I74.400rh  .819  I4.876dr  .579 


191.360^:  .943  I7.ii6d:  .666 
i8o.624zb  .74823.537^  .529 


202.2801b X. 03 1  X 5.284 db   .729 


i75.320dr  1.060' i5.7o8dr  .749 
1 88.800  d=  .980  2o.540d=  1.092 


4     il30.I20±   .628    9.284  rb   .443 

56  ;x23.666rb  .8x3 ;i2.040rb  .573 


X3.262i46x 

IX.233zb.407 

7.296db.35o 

8.834^.300 

7.oo3db.274 
x2.767rb.331 

8.530zb.335 

8.945^.351 
I3.795:i=.3i6 

7.556rb.362 


8.959±.43i 
10.8791b.371 


7.134^.342 
9.736±.469 


Range  of 

Variation 
in  Bficroos. 


28-228 
56-224 
48-2x2 

04-220 

40-256 

28-188 

40-216 
20-176 
20-220 

32-212 

36-240 
20-240 

60-232 

24-316 
24-232 


04-156 
00-160 


\ 


x9o8.] 


JENNINGS— HEREDITY   IN   PROTOZOA. 


461 


Table  XVIII.'-Continucd. 

tion  in  Paramecium.  Each  row  consists  of  specimens  taken  from  the 
column  headed  "Table"  gives  the  number  of  a  table  found  in  the 
question. 


Mean  in 
Microns. 


49.OOOrb.548 
64.8804r.580 

43.556db.392 
47.364dr.344 
56.Ii2i.395 

38.080db.356 

46.02Orfc.25i 
3i.l8Orfc.2i2 
46.684rfc.488 

44.80Odz.429 

54.880  dr  .431 
43.60Orfc.377 

49.60Orfc.298 


63.l60rfc.472 
56.380  rfc.427 


36.280  rb.260 
33.60Ort.4Oo 


Breadth. 


Standard 

Deviation 

in  Microns. 


8.144^.388 
8.624dr4I2 

6.748dr.276 

7.132^.244 

7.808  rfc.279 
5.288  rfc.252 

5.256rfc.i77 

3.88idb.i5i 

13.484i.344 

7.79^^.304 

7.824dr.3o5 
11.852rfc.266 

4.4I2rfc.2IO 


Coefficient  of 
Variation. 


l6.6l8rfc  .814 
I3.292rfc   .645 

I5.490±  .651 

i5.o57±  .526 

13.9131b  .507 

13.88 1  =fc  .675 

iM2iifc  .390 
I2.473rfc  .493 
28.879d=  .793 

I7.397±  .698 

14.255  db  .566 
27.x84dr  .654 

8.896rfc  .428 


Range  of 

Variation 

in  Microns. 


7.ooorb.334     ii.o83dr  .535 
8.956rfc.302  ;  I5.884rfc  .549 


3.88odb.i84i  io.7oorfc  .516 
5.917^3.283  ;  i7.6o8dr  .865 


28-76 

44-88 
*  32-60 

32-72 
36-80 
28-52 

36-60 
20-40 
20-80 

32-68 

36-84 
20-84 

40-60 


44-80 
40-80 


28-44 
23.3-50 


Mean  Index, 

or  Ratio  of 

Breadth  to 

Length. 

Per  Cent. 

Coefficient  of 
Correlati<m. 

26.029 

.4i88d=.o556 

35.131 

.6469rfc.0392 

23.517 

.5955-^-.o375 

27.153 

.3945  ±0408 

27.850 

.677irfc.o274 

25.515 

.448izfc.o539 

25.084 

.42824-.0389 

21.337 

.39o6rfc.0467 

28.236 

.8463-4- .0102 

25.657 

.5704dfc.0372 

28.639 

.7364^.0252 

24.593 

36.123 
30.350 


27.913 
27.136 


.4o85rfc.o562 


.5376=fc.o48o 

— .26l3dr.04X4 


.5208rfc.0492 
.6258rt.04lO 


462 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  94, 


Table  XVllL-^ontinued. 


i 


18 


19 


20 


21 
22 

23 
24 

25 

26 
27 


B.  Progeny  of  c. 
Continued. 


Same  as  last,  but  24  hours  after 
addition  of  boiled  grass,  Au- 
gust 10 

Same  as  row  17,  but  24  hours 
in  fresh  hay  infusion,  August 
12  f 

Rows  17  and  19,  together ;  same 
animals,  half  in  old  fluid, 
half  in  new 

Conjugating  culture,  large  ves- 
sel, September  25 

Same  culture,  5  days  after,  food 
getting  scarce 

Large,  old  culture,  January  23, 
1908; 

Same,  two  days  later,  January 
25,  1908 

Another  old  culture,  January 
23,  1908 

Same  as  row  23,  but  starved  3 
weeks,  February  14 

Same  as  row  23,  but  cultivated 
in  small  watch  glass,  January 
30-February  15,  1908 


225 


100 


2CX> 


200 


100 


100 


50 


100 


37 


•3 


49 


50 


57 
58 
59 


Length 


Mean  in 
Microns. 


Standard 

Deviation 

in  Microns. 


Coefficient  of 
Variation. 


Range  of 

Variadon 

in  MicroQ^ 


114.163d-  .78417.443^  .555  i5.279±.497  73.3-160 


114.033=^  .820 


1 1 8.850  i  .622 


X2.i4o±  .580  10.6461^.513  '86.7-146.7 


i3.o37±  .440 


10.698zb.374  86.7-160 


l58.8oo=h  .877  i8.384zt:  .620  11.578rb.396 


129.640^1  .867 
144.880rbi.097 
130.640^  1.227 


i2.848dr  .613 

l6.264rb   .776 

i2.863dr  .868 


I37.200zb  .842  I2.488rb  .596 

102.594 rb 1. 1 61  10.4671b   .821 


IOq    60   ,I00.320rb    .528 


7.828rb  .373 


124-200 
100-152 
100-176 


9.9i^cb.477 

XI. 224  dr. 542 

I 

9.846rb.670 ,  104-156 
9.io2db.438    104-162 

1 0.202 rb. 808  I     76-128 

7.8o4rb.3741    7^120 


multiply.  The  measurements  of  a  sample  of  100  of  these  are  given 
in  Table  LI.  (appendix),  while  the  constants  are  found  in  row  2, 
Table  XVIII.  The  increased  breadth,  with  little  change  in  the 
length,  of  course,  results  in  an  increase  of  the  mean  index  or  ratio  of 
breadth  to  length ;  while  in  row  i  this  was  but  26.029  per  cent.,  in  the 
present  lot  it  is  35.131  per  cent.  It  is  worthy  of  notice  that  with  the 
increase  in  ratio  of  breadth  to  length  there  is  an  increase  in  the  cor- 
relation between  length  and  breadth  from  .4188  to  .6469. 

Scarcity  of  Food, — The  watch-glass  culture  just  described  (row 
2,  Table  XVIII.)  was  now  allowed  to  stand  for  three  days  (till  June 
17)  without  renewing  the  culture  fluid.  The  animals  had  multiplied 
greatly,  so  that  food  became  scarce;  as  a  result  they  became  thin. 
The  measurements  are  given  in  Table  VI.  (page  412)  and  the  con- 
stants in  row  3,  Table  XVIII.  While  the  length  remained  about  the 
same,  the  mean  thickness  of  the  body  decreased  from  64.880  to  43.556 
microns.     The  mean  ratio  of  breadth  to  length  fell  from  35.131  per 


1908.] 


JENNINGS— HEREDITY  IN    PROTOZOA. 


•    463 


Table  XVllL-^ontinued, 


Breadth. 

Mean  in 
Microns. 

Standard 

Deviation 

in  Microns. 

Coefficient  of 
Variation. 

Range  of 

Variation 

in  Microns. 

Mean  I 

or  Rati 

Breadt 

Leng 

PerO 

Coefficient  cf 
Correlation. 

34.207 -4-.24I 

5.363-^-.i7i 

X5.683d=  .511 

20-50 

30.177 

.6757rh.0244 

47  300^.437 

6.490-I-.3XO 

I3.720ih  .667 

36.7-66.7 

41.455 

.8l524r.0226 

40.4SOzb.44i 

9.247-1-.312 

22.857-+-  .810 

23.3-66.7 

.i758rh.0462 

38.S601h.353 

7.396^.249 

19.176^1  .670 

16-60 

24.244 

.7135^.0234 

35440-I-.400 

5.928 -^-.283 

l6.730rh   .820 

20-48 

27.262 

.7576-f-.o287 

54.1601h.765 

11.3464-.541 

20.948  ±1.042 

32-84 

37.106 

1 

.8500-+- .01 87 

37.760 -t-.639 

6.697^452 

17.7361hl.233 

28-52 

28.97s 

.4141^.0790 

37.960i.413 

6.I28-h.292 

16.142-f-  .790 

24-56 

27.625 

.6691-+- .0373 

23.892H-.644 

5.8o4±.455 

24.291 -4-2.014 

16-40 

23.067 

.80 1 8-+- .0396 

26:480 -1-.266 

3.944±.i88 

14.895  ±  .753 

X6-36 

26.321 

.767i-t-.o278 

cent,  to  23.517  per  cent.,  and  at  the  same  time  correlation  between 
the  two  fell  from  .6469  to  .5955. 

Thus,  within  a  week  we  find  enormous  fluctuations  in  breadth, 
due  to  changes  in  the  amount  of  food,  while  the  length  remains  about 
the  same.  The  breadth  is  much  more  affected  by  nutritional  changes 
than  is  the  length. 

Rapid  Multiplication, — To  the  watch-glass  culture  just  described 
(row  3)  new  hay  infusion  was  added.  Twenty- four  hours  later 
(June  18)  multiplication  was  occurring  actively;  stages  of  fission 
and  all  the  stages  of  growth  were  numerous.  Measurements  of  195 
specimens,  taken  at  random  at  this  time  (Table  VII.,  page  412,  and 
row  4,  Table  XVIII.)  show  a  very  great  increase  in  the  range  and 
amount  of  the  variability  in  length,  while  there  is  little  change  in  the 
breadth.  This  is,  of  course,  due  to  the  fact  that  the  culture  contains 
many  young ;  these  differ  much  from  the  adults  in  length,  but  little 
in  breadth.     The  mean  length  decreases  from  185.008  to  176.124 


464       •  JENNINGS— HEREDITY  IN   PROTOZOA.  [April  24, 

microns,  and  the  variability  in  length  almost  doubles,  increasing  from 
7.794  to  13.262.  Owing  to  the  inclusion  of  many  young  individuals, 
in  which  the  length  is  increasing  while  the  breadth  is  stationary  or 
decreasing,  the  correlation  between  length  and  breadth  decreases  to 
.3945.  Inspection  of  Tables  VI.  and  VII.  (page  412)  shows  at 
a  glance  the  great  effect  of  nutrition  and  division  on  the  range  and 
distribution  of  variations  in  size  and  form. 

Injurious  Bacteria. — ^A  remarkable  effect  of  what  may  be  called 
"  bad  "  conditions  is  shown  in  this  series  of  experiments.  The  same 
watch-glass  culture  shown  in  row  3,  Table  XVIII.,  was  allowed  to 
stand  for  a  week,  till  June  25.  Bacteria  of  a  certain  character  mul- 
tiplied greatly,  and  seemed  to  get  the  upper  hand  of  the  Paramecia. 
The  latter  became  opaque  and  abnormal  in  appearance,  and  some  of 
them  died,  disintegrating  into  shapeless  masses.  It  was  now  observed 
that  many  of  the  specimens  still  living  were  very  large,  and  that 
variation  in  size  was  extreme.  The  distribution  of  the  variations  is 
shown  in  Table  LII. ;  the  constants  in  row  5,  Table  XVIII.  Though 
no  multiplication  is  occurring,  so  that  no  young  are  present,  the 
range  of  variation  is  from  140  to  256  microns,  while  in  row  3,  from 
which  this  lot  is  derived,  the  range  is  only  from  148  to  212  microns. 
The  mean  length  has  increased  to  201.888  microns,  one  of  the  greatest 
mean  lengths  ever  observed  in  progeny  of  D.  The  maximum  size 
for  descendants  of  D  was  likewise  reached  in  this  culture;  in  no 
other  case  were  specimens  256  microns  long  observed. 

Starvation, — In  striking  contrast  with  the  effects  of  much  nutri- 
tion (row  4,  Table  XVIII.)  and  of  injurious  bacteria  (row  5)  are 
the  results  of  starvation  (Table  LIII.,  and  row  6,  Table  XVIII.). 
The  starving  culture  consisted  of  individuals  from  the  same  culture 
as  row  I,  placed  in  fresh  hay  infusion  June  14.  The  constants 
before  they  were  placed  in  the  hay  infusion  are  given  in  row  i.  Table 
XVIII.,  while  the  immediate  effects  of  the  infusion  are  shown  in 
row  2  of  the  same  table.  The  same  animals  were  left  in  this  fluid 
for  eleven  days,  till  June  25.  They  had  evidently  begun  to  starve; 
they  were  small  and  thin  and  almost  half  of  them  had  died.  The 
dimensions  are  given  in  Table  LIII.,  and  the  constants  in  row  6, 
Table  XVIII.  The  length  had  fallen  from  184.680  to  149.360 
microns ;  the  breadth  from  64.880  to  38.080  microns.     The  breadth 


i9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  466 

decreases  with  lack  of  food  proportionately  more  than  does  the 
length,  so  that  the  ratio  of  length  to  breadth  has  fallen  from  35.131 
per  cent,  to  25.515  per  cent.  It  is  to  be  noticed,  however,  that  this 
greater  proportionate  decrease  of  breadth  takes  place  in  the  first 
days  after  the  withdrawal  of  abundant  food,  since  after  the  animals 
had  been  only  three  days  without  new  food  the  ratio  of  breadth  to 
length  fell  to  23.517  per  cent,  (row  3,  Table  XVIII.) ;  it  did  not 
decrease  farther  after  starvation  b^an. 

A  comparative  inspection  of  Tables  VII.  (page  412)  and  LIIL 
(appendix)  shows  to  the  eye  the  very  great  effects  of  nutrition  on 
size  and  variation. 

Second  Series. 

After  the  series  of  experiments  described  above,  the  progeny  of 
D  were  kept  in  large  culture  jars  of  hay  and  water  for  about  three 
weeks.  Then  followed  an  exceedingly  instructive  series  of  experi- 
ments on  the  effects  of  environmental  conditions,  the  results  of  which 
are  shown  in  Tables  XIX.-XXII.  and  in  the  large  Table  XVIII., 
rows  7  to  12.  Mere  inspection  of  the  correlation  tables  shows  the 
effects  in  such  a  striking  way  that  I  have  placed  the  main  tables 
together  in  the  text,  instead  of  relegating  them  to  the  appendix. 

Fresh  Hay  Infusion. — On  July  16,  1907,  specimens  from  the 
large  cultures  were  placed  in  a  watch-glass  of  hay  infusion  and 
allowed  to  remain  twenty-four  hours.  This  induced  rapid  multipli- 
cation ;  while  this  was  occurring  a  random  sample  of  200  specimens 
was  measured,  with  the  results  shown  in  Table  XXX.  (appendix), 
and  in  row  7,  Table  XVIII. 

Starvation. — Next  these  were  allowed  to  starve  for  a  week ;  then 
150  specimens  were  measured  (Table  XIX.,  and  row  8,  Table 
XVIII.).  The  results  may  be  compared  with  our  other  starving 
culture  of  Table  LIII.,  and  row  6,  Table  XVIII.  It  will  be  noticed 
that  for  both  length  and  breadth  the  amount  of  variation  is  not 
great ;  that  the  absolute  dimensions  are  small ;  that  the  ratio  of  breadth 
to  length  (21.337  per  cent.)  is  the  least  we  have  even  seen,  and  that 
the  correlation  between  length  and  breadth  is  very  low  (.3906). 

Effects  of  Abundant  Food  on  a  Starving  Culture. — Now  this 
starving  culture  (Table  XIX.)  was  placed  for  twenty-four  hours  in 


466 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  a4. 


Table  XIX. 

Correlation  Table  for  Length  and  Breadth  of  a  Starving  Culture  of  De- 
scendants of  D.     (Row  8,  Table  XVIII.) 

Length  in  Microns. 


S-^OOWSOO^OOWSOQ^OQCiSO 


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2    2    8    61126162122236    4    I     I     I 


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54 
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150 


Length — Mean,  146.108  ±  .563/*  Breadth — Mean,  31.180  ±  j2I2a« 

St.  Dev.,        10.228  ±.398^  St.  Dev.,        3.881:!:  .15  im 

Coef.  Var.,     7.003  dt  .274  Coef .  Var.,  12473  —  A93 

Mean  Index  or  Ratio  of  Breadth  to  Length,  21.337  per  cent. ;  Coef.  Cor., 

.3906  ±  .0467. 


Table  XX. 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  D  when 
Starving  Culture  (Table  XIX.)  is  placed  for  24  Hours  in  Fresh 
Hay  Infusion,     (Row  9,  Table  XVIII.) 

Length  in  Microns. 

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Length — Mean,  163.932  ±  .754/*  Breadth— Mean,  46.684  di  488^ 

St.  Dev.,       20.^±.533A*  St.  Dev.,  13484 ±.344M 

Coef.  Var.,    12.767  ±  .331  Coef.  Var.,  28,879^  .793 

Mean  Index,  28.236  per  cent. ;  Coef.  Cor.,  .8463  ±  .0102. 


«9o8.] 


JENNINGS— HEREDITY   IN   PROTOZOA. 


467 


Table  XXI. 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  D,  after  Re- 
maining One   Week  in   Hay  Infusion,   Unchanged,     (Row    lo, 

Table  XVIII.) 
Length  in  Microns. 


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Length — Mean,  174400  ±  .819^  Breadth — Mean,  44.800  ±  .429^ 

St.  Dev.,        14.876  ±  .579^  St  Dev.,        7796  ±'  304^ 

Coef.  Van,      8.530  ±  -335  Coef .  Var.,  17397  ±  698 

Mean  Index  or  Ratio  of  Breadth  to  Length,  25.657  per  cent.;  Coef.  Cor., 

.5704  ±  .0372. 

Table  XXII. 

Correlation  Table  for  Length  and  Breadth,  after  the  Culture  shown  in  Table 
XXL  has  remained  48  hours  in  Fresh  Hay  Infusion. 

(Row  II,  Table  XVIII.) 

Length  in  Microns. 

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Length— Mean,           191.360  ±  .943^*           Breadth— Mean,  54-88o  ±  431A1 

St.  Dev.,        I7.ii6±.666/*                           St.  Dev.,  7.824 ±.305^ 

Coef.  Var.,     8.945  ±  -351                             Coef.  Var.,  14.255  ±  .566 
Mean  Index,  28.639  per  cent. ;  Coef  Cor.,  7364  ±  .0252. 


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468 


JENNINGS -HEREDITY   IN   PROTOZOA. 


[April  94, 


a  fresh  hay  infusion.  At  once  the  culture  "  spread  out "  greatly,  in 
a  way  that  will  appear  on  comparing  Table  XIX.,  for  the  starving 
culture,  with  Table  XX.,  for  those  twenty-four  hours  in  nutritive 
fluid.  Many  of  the  animals  began  to  grow  at  once  after  they  were 
placed  in  the  nutritive  fluid,  so  that  the  maximum  length  increased 
from  176  to  220  microns,  the  maximum  breadth  from  40  to  80 
microns  (see  rows  8  and  9,  Table  XVIII.).  Others  had  not  yet 
begun  to  increase  when  the  sample  of  Table  XX.  was  taken,  so  that 


Fig.  5.  Characteristic  forms  and  sizes  from  a  culture  of  descendants  of 
D  (caudatum  form),  that  had  been  starved  for  a  week  (Table  XIX.),  then 
was  left  twenty- four  hours  in  fresh  hay  infusion  (Table  XX.).  a  and  b. 
Starved  specimens,  c,  d,  e,  f,  transitional  forms,  becoming  large  and  plump 
in  the  abundant  food ;  g,  characteristic  large,  plump  form,  a  to  ^  from  Tabic 
XX.  h,  characteristic  form  a  week  later  (Table  XXI.)  ;  animals  becoming 
thinner  again,  but  retaining  the  increased  length.    All  X  235. 


,9o8.1  JENNINGS— HEREDITY  IN   PROTOZOA.  469 

the  minimum  size  remained  as  before;  and  between  these  extremes 
all  intermediate  gradations  were  found.  Fig.  5  shows  characteristic 
forms  and  sizes  from  this  culture,  a  and  b  showing  the  starving  con- 
dition, while  ctof  show  various  stages  in  the  transition  to  the  largest 
size,  one  of  which  is  shown  at  g. 

As  a  result  of  these  changes,  the  variability  has  increased  enor- 
mously. The  coefficient  of  variation  in  length  has  increased  in 
twenty- four  hours  from  7.003  to  12.767;  that  for  breadth  has  more 
than  doubled,  increasing  from  12.473  to  28.879.  The  mean  size  has 
likewise  increased  greatly,  while  the  ratio  of  breadth  to  length  has 
changed  from  21.337  P^^  cent,  to  28.236  per  cent.  Perhaps  the  most 
striking  change  is  in  the  correlation  between  length  and  breadth.  In 
the  starving  culture  this  is  but  .3906 ;  twenty- four  hours  later  it  has 
become,  in  the  growing  culture,  .8463 — one  of  the  highest  coefficients 
of  correlation  that  I  have  ever  found  in  Paramecium,  It  is  evident 
that  breadth  and  length  are  increasing  proportionately,  on  the  whole, 
so  that  the  inclusion  of  different  degrees  of  increase  in  size  in  Table 
XX.  gives  a  high  coefficient  of  correlation.  Furthermore,  the  fact 
that  fission  had  not  begun  in  this  lot  permits  the  correlation  to  remain 
high ;  if  there  were  many  young  included,  the  correlation  would,  of 
course,  be  lowered.  With  every  increase  of  10  microns  in  length  the 
breadth  increases  5452  microns. 

Fluid  Unchanged  for  a  Week. — Now  the  same  culture  was  kept 
for  a  week  in  the  same  fluid.  The  animals  had  reached  more  nearly 
a  condition  of  equilibrium ;  the  variability,  and  with  it  the  correla- 
tion, had  greatly  decreased,  while  the  mean  length  had  increased 
(Table  XXI.,  and  row  10,  Table  XVIII.).  It  is  noticeable  here,  as 
in  many  other  cases,  that  the  coefficient  of  correlation  decreases  when 
the  ratio  of  breadth  to  length  decreases. 

Forty-eight  Hours  in  New  Culture  Fluid. — ^The  addition  of  new 
hay  infusion  to  the  culture  just  described  caused  in  forty-eight  hours 
a  considerable  increase  in  mean  length  and  breadth,  while  the  varia- 
tion did  not  change  greatly  (Ts^ble  XXII.,  and  row  11,  Table 
XVIII.).  Again,  as  the  ratio  of  breadth  to  length  increases,  the 
correlation  between  the  two  likewise  increases. 

RSsumS. — Polygons  showing  the  changes  in  the  animals  of  this 
series,  from  the  starving  condition  of  Table  XIX.  to  the  well-fed 


470 


JENNINGS -HEREDITY   IN   PROTOZOA. 


[April  a4. 


condition  of  Table  XXII.  are  given  in  Diagram  6;  these,  taken  in 
connection  with  Fig.  5  and  with  Tables  XIX.  to  XXII.  give  a  good 
idea  of  the  changes  in  dimensions  and  variation  that  may  be  pro- 


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Length  in  Microns. 


236 


Diagram  6.  Polygons  of  variation  in  length  for  a  culture  of  descendants 
of  the  individual  D  when  subjected  successively  to  varied  conditions  of  nu- 
trition. The  numbers  above  the  highest  points  of  the  polygons  correspond 
to  rows  of  Table  XVIII.,  in  which  are  given  the  constants  for  the  different 
polygons.  8,  culture  starved  a  week.  9  (heavy  broken  line),  same  as  8,  but 
after  24  hours  in  fresh  hay  infusion.  10,  same  after  one  week  in  the  same 
fluid,  unchanged,  it,  same  after  48  hours  in  fresh  hay  infusion,  x,  polygon 
for  combination  of  8,  10  and  11,  showing  its  resemblance  to  the  polygon  for 
9  alone. 

The  correlation  tables  for  these  polygons  are  numbers  XIX.  to  XXII., 
pages  466,  467. 

duced  in  a  short  time  by  changes  in  the  conditions  of  nutrition. 
Evidently  Table  XX.,  taken  twenty-four  hours  after  the  starving 
specimens  were  placed  in  the  fresh  hay  infusion,  is  a  transitional 


1 


,9o8.j  JENNINGS— HEREDITY   IN   PROTOZOA.  471 

condition,  including  representatives  of  the  small,  starving  condition, 
the  well  grown  condition,  and  intermediate  states;  it  is  a  sort  of  a 
resume  of  the  variations  due  to  nutrition.  If  we  add  together  the 
tables  given  by  the  starving  culture  (earlier  than  Table  XX.)  and 
the  two  well-fed  cultures  (later  than  Table  XX.),  we  get  a  collection 
of  450  individuals,  in  which  the  variation  in  length  and  breadth  is 
about  the  same  as  for  Table  XX.  (see  row  12,  Table  XVIIL).  For 
Table  XX.  the  coefficients  of  variation  for  length  and  breadth  are 
12.767  and  28.879;  ^^^  corresponding  coefficients  for  the  three  lots 
combined  are  13.795  and  27.184. 

Although  the  animals  are  all  descended  from  the  same  parent  and 
have  lived  under  the  same  conditions  save  for  the  ten  days  during 
which  these  experiments  lasted,  we  find  that  in  the  period  just  men- 
tioned the  polygons  of  distribution  of  variations  in  length  have  so 
changed  that  the  one  for  the  end  of  the  ten  day  period  (11,  Diagram 
6)  hardly  more  than  overlaps  at  one  end  that  for  the  beginning  of 
the  period  (8,  Diagram  6). 

Addition  of  fresh  hay  infusion  causes  in  these  cases  an  increase 
in  length,  in  breadth,  in  variation,  and  in  the  correlation  between 
length  and  breadth.  But  whether  these  results  shall  follow  depends 
upon  the  previous  condition  of  the  animals.  This  is  illustrated  by 
the  fact  that  there  is  one  exception  to  the  statement  just  made;  the 
variability  in  breadth  decreased  in  place  of  increasing  in  the  transi- 
tion from  Table  XXI.  to  Table  XXII.  The  effect  of  the  previous 
condition  is  better  seen  in  the  experiments  of  the  third  series,  to  be 
described  next. 

Third  Series, 

A  culture  of  the  descendants  of  D  was  rather  ill-fed,  though  not 
starving;  the  animals  were  long  and  slender  (Fig.  6,  a  and  b).  Half 
of  these  were  allowed  to  remain  in  the  old  fluid,  while  half  were 
placed  in  fresh  hay  infusion.  After  forty-eight  hours,  a  random 
sample  of  each  set  was  measured.  The  measurements  of  the  set  in 
the  old  fluid  are  given  in  Table  LIV.,  the  constants  in  row  13,  Table 
JCVIII.  The  results  of  keeping  the  animals  forty-eight  hours  in  the 
fresh  infusion  are  shown  in  Table  LV.,  and  in  row  14,  Table  XVIII. 
The  animals  grew  plump  and  multiplied ;  the  mean  breadth  increased 


472 


JENNINGS— HEREDITY  IN   PROTOZOA. 


(April  34, 


from  49.6CX)  microns  to  63.160  microns  (characteristic  form  shown 
ate.  Fig.  6).  But  the  mean  length  decreased  from  202.280 to  175.320 
microns.  This  is  probably  due  to  rapid  multiplication ;  the  animals 
now  divide  before  they  reach  the  length  which  they  had  at  first.  As 
a  result  of  the  increase  in  breadth  and  decrease  in  length,  of  course, 


Fig.  6.  a  and  b,  characteristic  slender  specimens  from  row  13,  Table 
XVIII.  c,  characteristic  short  plump  specimen  from  row  14,  Table  XVIII.; 
produced  by  allowing  those  of  row  13  to  remain  24  hours  in  fresh  hay  in- 
fusion.   Descendants  of  D  (caudatutn  form).    All  X235. 

the  mean  ratio  of  breadth  to  length  increased  greatly,  from  24.593 
per  cent,  to  36.123  per  cent.  With  the  increase  of  this  ratio,  the 
correlation  likewise  increased,  as  is  usually  the  case.  The  variation 
increased,  both  in  breadth  and  in  length. 

These  are  the  results  if  we  consider  separately  the  two  samples, 
taken  forty-eight  hours  apart.  But  if  we  throw  them  together,  look- 
ing at  them  merely  as  a  sample  of  the  descendants  of  D,  taken  at 
intervals,  we  get  a  surprising  effect  on  the  correlation  between  length 
and  breath.  The  marked  positive  correlation  in  the  two  samples 
taken  separately  disappears  and  is  replaced  by  a  negative  correkUion. 
In  the  first  sample  the  correlation  is  -f-  .4085 ;  in  the  second  it  is 
+  -5376;  in  the  two  together  it  is  —  .2613.  (The  constants  for  tfic 
two  together  are  given  in  row  15,  Table  XVIII.)  The  negative 
correlation  is,  of  course,  due  to  the  fact  that  the  nutritive  fluid  causes 
the  breadth  to  increase  and  the  length  to  decrease,  so  that,  on  the 


,9o8.]  JENNINGS— HEREDITY  IN   PROTOZOA.  473 

whole,  when  the  two  samples  are  taken  together,  greater  breadth  is 
associated  with  less  length. 

2.   Progeny  of  c  (aurelia  Form). 

With  the  smaller  Paramecia,  progeny  of  the  small  individual  c,  a 
similar  series  of  experiments  was  undertaken.  The  individual  c 
came  from  the  same  wild  culture  as  D;  its  length,  as  nearly  as  could 
be  determined  in  life,  was  120  microns.  It  was  isolated  April  8, 
1907.  Fig.  3  shows  some  examples  of  the  descendants  of  c,  drawn 
to  the  same  scale  as  the  figures  of  the  descendants  of  D. 

Random  Sample. — On  June  1 1  one  hundred  of  the  progeny  of  c 
gave  the  measurements  shown  in  Table  IV.,  page  405,  the  constants 
being  given  in  row  16,  Table  XVIII. 

Effect  of  Adding  Boiled  Hay, — On  August  9  a  fairly  flourishing 
culture  of  the  descendants  of  c  was  examined,  with  the  results  shown 
in  Table  LVL,  and  in  row  17,  Table  XVIII.  To  this  culture  a  quan- 
tity of  boiled  grass  was  added;  this  caused  rapid  multiplication. 
Twent)r-four  hours  later  a  sample  of  225  specimens  was  measured, 
with  the  results  shown  in  Table  XLIX.,  and  row  18,  Table  XVIII. 
The  added  nutrition  has  caused  the  mean  length  to  decrease,  while 
the  mean  breadth  remains  nearly  the  same.  This  is  due  to  the  fact 
that  the  main  effect  of  the  nutrition  was  to  cause  rapid  multiplication 
rather  than  growth  in  size.  The  coefficient  of  variation  in  length 
increased  greatly,  from  9.736  to  15.279,  while  the  variation  in  breadth 
remained  about  the  same,  though  with  a  slight  decrease.  This  pecu- 
liar result  is  mainly  due  to  the  fact  that  the  culture  after  the  addition 
of  the  grass  (row  18)  contains  many  young  specimens,  which  diflfer 
from  the  adults  greatly  in  length,  but  little  in  breadth.  As  usual,  we 
find  that  an  increase  in  the  ratio  of  breadth  to  length  is  accompanied 
by  an  increase  in  the  correlation  between  the  two. 

Effect  of  Fresh  Hay  Infusion. — The  next  day  (August  11) 
another  lot  from  the  culture  shown  in  Table  LVL  (row  17,  Table 
XVIII.)  was  placed  in  a  fresh  hay  infusion  and  left  twenty- four 
hours.  This  nutritive  fluid  caused  the  animals  to  become  very 
plump,  while  at  the  same  time  a  moderate  amount  of  fission  was 
induced.    The  results  are  shown  in  Table  L.,  and  in  row  19,  Table 

PROC.  AMER.  PHIL.  SOC.  XLVII.   I90  EE,  PRINTED  JANUARY  II,   I909. 


474  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a*, 

XVIII.  As  there  appears,  the  mean  breadth  increased  from  33.600 
to  47.300  microns.  The  length,  on  the  other  hand,  decreased  from 
123.666  to  114.033  microns.  The  mean  ratio  of  breadth  to  length 
thus  increased  very  greatly,  from  27.136  per  cent,  to  41.455  per  cent. 
The  latter  is  the  largest  mean  index  I  have  ever  observed  in  Para- 
mecia  not  selected  with  relation  to  the  age  of  the  individuals;  it  is 
exceeded  only  by  the  mean  index  of  the  young  halves  during  fission 
(see  Table  X.).  With  the  increase  in  the  mean  ratio  of  breadth  to"" 
length,  there  is  as  usual  an  increase  in  the  correlation  between  the 
two  dimensions ;  this  reaches  the  unusually  high  value  of  .8152.  The 
nutritive  fluid  left  the  variation  in  length  about  the  same,  but  con- 
siderably decreased  the  variation  in  breadth.  This  is  undoubtedly 
due  to  the  fact  that  before  the  hay  infusion  was  introduced  some  of 
the  specimens  were  well  fed,  some  poorly  fed,  as  the  chances  of  the 
daily  life  determined;  while  after  the  infusion  was  introduced  all 
were  well  fed,  so  that  there  was  less  variation  in  breadth  than  before. 
Characteristic  forms  after  the  infusion  was  introduced  are  shown  in 
Fig.  3,  a  to  c  (page  423). 

The  facts  in  these  cases  are  nearly  parallel  with  those  observed 
in  the  third  series  of  experiments  on  the  progeny  of  D  (Table  XVIII., 
rows  13-15).  If  we  combine  the  two  samples  of  c  (row  20,  Table 
XVIII.),  as  we  did  those  of  D,  the  effect  is,  as  in  the  case  of  D,  to 
decrease  greatly  the  correlation  between  length  and  breadth  But  in 
the  present  case  the  very  high  positive  correlation  of  the  two  samples 
taken  separately  is  not  entirely  overcome  by  combining  them,  though 
the  correlation  falls  to  .1758.  The  actual  numerical  coefficient  just 
given  is  the  resultant  of  a  number  of  conflicting  factors.  In  the  two 
samples  taken  separately  greater  length  is  associated  on  the  whole 
with  greater  breadth,  giving  high  positive  correlation,  which  in  pass- 
ing from  Table  LVI.  to  Table  L.  an  increase  in  breadth  is  associated 
with  a  decrease  in  length,  tending  to  diminish  the  correlation.  The 
facts  show  clearly  that  the  observed  statistical  correlation  does  not 
involve  any  necessary  and  constant  relation  of  the  one  dimension  to 
the  other;  both  dimensions  depend  on  various  factors,  which  some- 
times act  in  the  same  way  on  both,  sometimes  differently. 

Combining  the  two  samples  of  c  (as  in  row  20,  Table  XVIII.), 
gives,  of  course,  increased  variation,  illustrating,  like  most  of  our 


,9o8.l  JENNINGS— HEREDITY  IN   PROTOZOA.  475 

results,  the  fact  that  a  definite  coefficient  of  variation  cannot  be  con- 
sidered characteristic  of  a  given  species  or  race.  The  observed 
variation  depends  oi?  many  factors. 

Conjugating  Culture. — The  progeny  of  c  I  divided  into  two  sets, 
both  of  which  were  kept  in  larger  culture  vessels  and  maintained  by 
adding  boiled  hay  at  intervals.  September  25  one  of  these  cultures 
was  found  to  be  undergoing  an  epidemic  of  conjugation  (though,  of 
course,  all  were  progeny  of  a  single  individual).  The  details  regard- 
ing the  relation  of  conjugation  to  the  phenomena  we  are  studying  are 
to  be  taken  up  in  a  later  communication,  but  I  will  give  here  the 
essential  facts  regarding  dimensions  and  constants  of  variation,  in 
order  that  our  picture  of  the  changes  undergone  by  the  c  line  may  be 
as  complete  as  possible.  A  random  sample  of  the  non-con jugants 
of  this  conjugating  culture  gave  the  results  shown  in  row  21,  Table 
XVIIL,  and  in  Table  LVII.  The  mean  length  (158.800  microns) 
was  considerably  greater  than  has  been  observed  in  any  other  culture 
of  c.  Whether  this  fact  has  any  relation  to  the  occurrence  of  conju- 
gation, or  whether  it  is  merely  a  matter  of  the  environmental  condi- 
tions must  remain  for  the  present  a  question. 

Scarcity  of  Food  After  Conjugation, — This  conjugating  culture 
was  allowed  to  stand  five  days.  All  conjugation  ceased  and  the  food 
began  to  get  scarce.  Now  a  sample  gave  the  results  shown  in  row 
22,  Table  XVIIL,  and  in  Table  LVIII.  The  length  had  decreased 
from  1 58.800  to  129.640  microns.  Breadth  likewise  decreased,  though 
not  in  so  great  a  proportion  as  length,  so  that  the  ratio  of  breadth  to 
length  increased.  As  is  usual  when  this  ratio  increases,  the  coeffi- 
cient of  correlation  likewise  increased. 

Variation  in  Different  Divisions  of  the  Same  Pure  Line  on  the 
Same  Date. — After  the  observations  just  described,  the  two  cultures 
composed  of  the  progeny  of  c  were  maintained  for  several  months. 
On  January  23,  1908,  samples  from  each  were  measured,  giving  the 
results  shown  in  rows  23  and  25,  Table  XVIII.  As  is  evident,  the 
two  differed  considerably.  The  details  do  not  demand  attention, 
save  that  in  one  of  these  old  cultures  (row  23,  and  Table  LIX.)  the 
coefficient  of  correlation  between  length  and  breadth  was  the  highest 
I  have  ever  observed  in  Paramecium,  reaching  .8500.  Both  these 
cultures  were  flourishing  and  well  fed. 


476  JENNINGS— HEREDITY  IN  PROTOZOA.        [April.,, 

Effects  of  Lack  of  Food. — From  the  culture  shown  in  row  23, 
Table  XVIII.,  a  large  number  of  specimens  were  removed  and  placed 
in  a  small  watch-glass,  which  was  allowed  to  stand  for  two  days. 
The  food  decreased  rapidly  and  the  animals  became  smaller,  giving 
the  results  shown  in  row  24,  Table  XVIII.  The  mean  length  had 
decreased  10.174  per  cent ;  the  mean  breadth  33.024  per  cent.  These 
were  now  allowed  to  stand  for  three  weeks  more  in  the  watch-glass, 
without  adding  food.  At  the  end  of  this  time  they  were  in  the 
extremes  of  starvation,  and  only  37  specimens  remained  of  the  many 
hundreds  originally  present.  These  37  gave  the  results  shown  in 
row  26,  Table  XVIII,  As  compared  with  the  original  condition  of 
row  23,  the  mean  length  had  decreased  30.638  per  cent,  the  mean 
breadth  55.8S6  per  cent.  A  peculiar  fact  is  that  this  starving  culture 
shows  a  very  high  coefficient  of  correlation  between  length  and 
breadth  (.8018),  while  in  our  other  starving  cultures  this  has  not 
been  the  case  (see  rows  6  and  8,  Table  XVIII.). 

From  the  culture  of  large  specimens  shown  in  row  23  another 
lot  was  removed  January  30  and  kept  in  a  small  watch-glass,  new 
hay  infusion  being  added  at  intervals.  In  spite  of  this  addition  of 
new  food  material,  and  the  fact  that  they  continued  to  flourish  and 
multiply,  these  decreased  in  length  even  more  than  in  the  starving 
culture,  the  mean  being  100.320.  This  is  the  smallest  mean  length 
observed  in  any  lot  of  the  c  line.  The  data  for  this  lot  are  given  in 
row  27,  Table  XVIII.,  and  in  Table  LX. 

3.     SUMMABV   ON    THE    EFFECTS   OF    ENVIRONMENT. 

The  facts  given  above  show  that  the  nature  of  the  environment 

affects  greatly  the  dimensions,  proportions  and  variations  of  Para- 

mecium,  and  that  these  effects  are  produced  with  great  ease  and 

---■"■■     by  such  changes  as  are  common  in  any  culture  of  these 

1.     Some  of  the  more  important  effects  may  be  summarized 

vs: 

:t  on  Length. — Under  the  influence  of  varied  nutritional  con- 
he  length  varies  extremely.  In  the  line  descended  from  the 
al  D  the  mean  length  varied  under  different  conditions  from 
to  202.280  microns — the  difference  being  38.445  per  cent,  of 
lest  mean  length.     In  the  c  line  the  variation  in  mean  length 


,9o8.]  JENNINGS-HEREDITY   IN   PROTOZOA.  477 

under  the  influence  of  the  environment  was  from  100.320  to  158.800 
microns,  or  58.293  per  cent,  of  the  lowest  mean.  The  extreme 
lengths  in  each  line,  of  course,  differed  still  more ;  in  the  D  line  the 
extreme  variation  in  length  was  from  104  to  256  microns,  or  146.153 
per  cent,  of  the  least  length ;  in  the  c  line  it  was  from  73.3  to  200 
microns,  or  172.851  per  cent,  of  the  minimal  length. 

Effect  on  Breadth. — The  breadth  (the  thickness  of  the  body) 
varies  under  different  environmental  conditions  more  readily  and  in 
a  higher  degree  than  does  the  length.  In  the  D  line  the  mean 
breadth  varied  in  different  cultures  from  31.180  to  64.880  microns, 
or  by  108.08  per  cent,  of  the  lowest  mean ;  the  extreme  variation  in 
breadth,  under  different  conditions,  was  from  20  to  88  microns,  or 
340  per  cent,  of  the  minimal  breadth.  In  the  c  line  the  mean  breadth 
varied  un(|er  different  conditions  from  23,892  to  54.160  microns,  or 
by  126.69  per  cent,  of  the  lowest  mean;  the  extreme  variation  in 
breadth  was  from  16  to  84  microns,  or  425  per  cent,  of  the  minimal 
breadth.  The  greater  variability  of  the  breadth,  as  compared  with 
the  length  is  seen  in  the  coefficients  of  variation  of  the  single  cultures. 
The  largest  coefficient  of  variation  for  length  is  15.279,  while  for 
breadth  it  is  28.879. 

Relation  of  Length  to  Nutrition, — In  general,  increased  nutrition 
increases  the  length.  But  the  result  is  not  always  the  same,  because 
increased  nutrition  has  two  main  effects :  to  increase  directly  the  size 
of  the  adults,  and  to  bring  about  multiplication.  The  latter  effect,  of 
course,  decreases  the  mean  length  of  the  individuals  of  a  culture^ 
since  it  induces  the  presence  of  many  specimens  that  are  young,  and 
therefore  small.  Increase  in  mean  length  due  to  added  nutrition  is 
seen  in  Table  XVIII.,  rows  8  to  9,  10  to  11.  Decrease  in  mean 
length,  due  to  added  nutrition  is  seen  in  the  same  table  on  comparing 
rows  I  and  2;  3  and  4;  13  and  14;  17  and  18.  This  decrease  is  due 
to  the  fact  that  in  the  nutritive  fluid  the  animals  divide  before  they 
reach  the  length  of  those  in  the  poor  fluid. 

Decrease  of  length,  due  to  decrease  of  nutrition,  is  seen  in  Table 
XVIIL,  by  comparing  rows  2  and  6;  7  and  8;  21  and  22;  23  and  24; 
23  and  26. 

Relation  of  Breadth  to  Nutrition, — ^The  relation  of  breadth  to 
nutrition  is  simpler  than  that  of  length ;  in  all  cases  increase  of  nutri- 


478  JENNINGS— HEREDITY   IN   PROTOZOA.  [April .4, 

tion  increases  the  breadth ;  decrease  of  nutrition  decreases  it.  The 
response  of  breadth  to  changes  in  nutrition  is  immediate  and  very 
marked.  Within  twenty-four  hours  increased  nutrition  caused  in 
the  D  line  an  increase  of  49.724  per  cent,  in  breadth  (rows  8  and  9, 
Table  XVIII.)  ;  in  the  c  line  it  caused  in  twenty- four  hours  an 
increase  of  40.778  per  cent,  (rows  17  and  19,  Table  XVIII.). 

But  the  decrease  of  breadth  with  decrease  of  nutrition  does  not 
vary  directly  with  the  time;  when  plump  individuals  are  left  without 
food,  they  decrease  much  more  rapidly  at  first  than  later.  Thus,  in 
the  series  shown  in  Table  XVIIL,  rows  2,  3  and  6,  the  breadth 
decreased  in  the  first  forty-eight  hours  21.324  microns,  or  32.867  per 
cent. ;  in  nine  days  more  of  lack  of  food  the  breadth  decreased  only 
5.476  microns,  or  8.440  per  cent.  more. 

Proportion  of  Breadth  to  Length, — Since  changes  in  nutritional 
and  other  conditions  act  more  readily  and  more  strongly  on  breadth 
than  on  length,  and  since  the  same  agent  may  increase  the  breadth 
while  decreasing  the  length,  the  proportion  of  breadth  to  length  varies 
greatly  under  different  conditions.  The  mean  index,  or  ratio  of 
breadth  to  length,  varies  in  different  cultures  of  the  D  line  from 
21.337  per  cent,  to  36.123  per  cent.;  in  the  c  line  from  23.067  per 
cent,  to  41.45s  per  cent.  Since  the  breadth  is  more  dependent  on 
nutritive  conditions  than  is  the  length,  we  find  the  lowest  ratio  of 
breadth  to  length  in  the  starving  cultures  (rows  8, 26,  Table  XVIII.)  ; 
the  highest  ratio  in  well-fed  cultures  (rows  2, 14, 19,  Table  XVIII.). 
An  increase  of  nutrition  causes  uniformly  an  increase  of  the  ratio 
of  breadth  to  length;  a  decrease  of  nutrition  has  almost  uniformly 
the  reverse  effect.  A  single  exception  to  the  relation  last  mentioned 
is  seen  in  the  change  from  row  21  to  row  22,  Table  XVIII. ;  here 
other  causes,  connected  with  conjugation,  were  probably  at  work. 
Whenever  the  mean  breadth  increases,  the  mean  ratio  of  breadth  to 
length  likewise  increases.  (The  only  exception  is  the  case  just  men- 
tioned, where  conjugation  was  involved.)  It  must  be  understood 
that  this  does  not  mean  that  in  all  cases  the  mean  ratio  of  breadth 
to  length  varies  directly  with  the  mean  breadth ;  if  we  compare  rows 
6  and  7,  Table  XVIIL,  for  example,  we  find  that  this  is  not  the 
case.  But  whenever,  as  a  matter  of  experimental  procedure,  the 
mean  breadth  was  caused  to  increase,  the  mean  ratio  of  breadth  to 


I 

,^.1  JENNINGS- HEREDITY   IN   PROTOZOA.  479 

length  likewise  increased.  This  is  due  to  the  two  facts  mentioned 
in  the  first  sentence  of  this  paragraph. 

Effect  of  Environment  on  Variation. — The  amount  of  observed 
variation,  as  measured  by  the  coefficient  of  variation,  depends  largely 
on  environmental  conditions;  this  is  true  both  for  length  and  for 
breadth.  In  the  D  line  the  coefficient  of  variation  for  length  varies 
in  different  cultures  from  6.821  to  13.262;*  for  breadth  it  varies 
from  8.896  to  28.879.  Ij^  th^  ^  ^^^^  ^^^  coefficient  varies  for  length 
from  7.134  to  15.279;  for  breadth  from  10.700  to  24.291. 

The  effects  on  the  coefficient  of  variation  of  changes  in  nutrition 
vary  much  in  different  cases ;  increased  nutrition  sometimes  increases 
the  coefficient,  sometimes  decreases  it,  sometimes  produces  first  one 
effect,  then  the  other.  There  are  evident  physiological  reasons  for 
the  different  effects.  In  a  starving  culture  the  first  effect  of  rich 
nutrition  is  to  cause  many  of  the  individuals  to  increase  in  size, 
while  those  individuals  in  which  the  effects  of  starvation  had  gone 
far  do  not  at  first  take  food  and  change.  Hence  there  is  a  great 
increase  in  the  coefficients  of  variation ;  in  changing  from  row  8  to 
row  9  (Table  XVIII.)  both  coefficients  approximately  doubled  in 
twenty-four  hours.  Later,  though  the  animals  were  kept  in  the 
same  fluid,  the  coefficients  decreased  again — all  of  the  specimens 
having  reached  more  nearly  a  condition  of  equilibrium.  If  the 
animals  are  fairly  well  fed  before  the  additional  nutrition  is  met,  an 
early  effect  is  to  cause  rapid  multiplication ;  the  consequent  presence 
of  both  young  and  old  individuals  in  the  culture  increases  the  coeffi- 
cients of  variation,  and  particularly  that  for  length.  An  example 
of  this  is  seen  in  the  change  from  row  3  to  row  4,  Table  XVIII.  A 
little  later,  when  the  multiplication  has  ceased,  the  coefficients  of 
variation  become  small  again.  The  coefficients  of  variation  are 
likely  to  be  small  in  starving  cultures,  owing  to  the  fact  that  there  is 
little  multiplication  and  the  adults  have  reached  a  condition  of  rela- 
tive equilibrium.  By  taking  into  consideration  the  immediate  and 
the  remote  effects  of  a  given  agent  on  growth  and  multiplication,  its 
effects  on  the  coefficients  of  variation  usually  become  intelligible. 

•Of  course  the  cultures  contain  specimens  in  all  stages  of  growth;  as 
we  have  previously  seen,  the  coefficient  of  variation  becomes  much  less  when 
the  animals  are  selected  with  reference  to  age. 


480  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a4. 

It  is  not  necessary  to  emphasize  the  fact  that  since  different 
environmental  conditions  produce  diflferent  dimensions,  the  coeffi- 
cients of  observed  variation  will  be  much  increased  by  throwing 
together  specimens  from  diflferent  environments,  or  those  taken  at 
diflferent  times  from  the  same  culture.  Examples  of  this  are  seen 
in  rows  12,  15  and  20,  Table  XVIII. 

The  question  may  be  asked,  How  can  we  account  for  the  large 
coefficients  of  variation  in  given  lots,  taken  all  from  the  same  envir- 
onment (as  in  the  various  "rows"  of  Table  XVIII.)?  Surely,  it 
may  be  said,  the  age  diflferences  among  the  individuals  are  not  suffi- 
cient to  account  for  coefficients  of  12,  13,  20,  etc.,  such  as  we  actually 
find.  This  is  undoubtedly  true,  and  it  becomes  still  more  striking 
when  we  consider  cases  like  Table  XLI.  (appendix),  where  the  indi- 
viduals are  all  of  practically  the  same  age,  and  all  come  at  one  time 
from  the  same  small  watch-glass  of  hay  infusion,  yet  we  find  the 
coefficients  of  variation  to  be  respectively  6.389  and  14.6x5.  The 
'considerable  variation  is  to  be  understood  only  by  realizing  that  even 
a  small  mass  of  fluid  constitutes  a  relatively  large  and  varied  envir- 
onment for  Paramecium.  A  watch-glass  of  hay  infusion  is  a  micro- 
cosm to  this  animal.  Bacteria  gather  on  the  surface,  while  they 
may  not  be  found  on  the  bottom  or  through  the  middle.  The  bac- 
terial zoogloea  may  become  thicker  at  one  edge  than  at  the  other, 
owing  to  the  accidents  of  the  original  distribution  of  the  seed  bacteria 
or  of  the  infusoria.  Some  of  the  Paramecia  thus  get  more  food 
than  the  others,  perhaps  at  a  critical  period  of  growth;  they  thus 
get  a  start,  which  enables  them  perhaps  to  obtain  more  food  than 
the  others,  even  under  uniform  conditions.  Some  of  the  individuals 
get  crowded  away  from  the  bacterial  zoogloea,  and  remain  against  a 
rough  spot  on  the  glass  instead,  where  they  get  no  food.  In  short, 
even  in  a  few  drops  of  water  the  conditions  are  no/  uniform  through- 
out; some  of  the  animals  are  well  nourished,  others  poorly  nour- 
ished, and  the  results  show  in  the  variations  of  their  measurements. 

The  question  whether  some  of  the  variations  in  such  cases  are 

■ 

not  congenital  and  hereditary  will  be  taken  up  later;  we  shall  find 
little  evidence  that  this  is  the  case. 

It  is  clear  that  no  particular  coefficient  of  variation  can  be  con- 
sidered characteristic  of  a  particular  race,  except  as  the  conditions 


x9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  481 

are  very  precisely  defined.  If  all  conditions  of  environment  and 
growth  were  made  absolutely  the  same,  there  is  reason  to  believe  (as 
we  shall  see  farther)  that  for  a  given  line  (descended  from  a  single 
individual)  the  coefficient  of  variation  would  be  very  close  to  zero. 
Its  actually  observed  value  in  a  given  lot  then  depends  almost  entirely 
on  environmental  and  growth  diflferences. 

Effect  of  Environment  on  Correlation, — ^The  observed  correlation 
between  length  and  breadth  varies  greatly  under  diflferent  environ- 
mental conditions.  In  the  D  line  the  coefficient  which  measures 
correlation  varies  in  different  cultures  from  .3906  to  .8463 ;  in  the 
c  line  from  4141  to  .8500  (see  Table  XVIII.).  Such  differences 
are  easily  and  quickly  produced  by  environmental  changes ;  thus  the 
two  extremes  just  mentioned  for  the  D  race  were  found  in  samples 
of  the  same  lot  of  Paramecia  taken  twenty-four  hours  apart — one 
before,  the  other  after,  the  addition  of  a  nutritive  fluid. 

The  correlation  between  length  and  breadth  expresses  the  accu- 
racy with  which  length  and  breadth  vary  proportionately.  The 
actual  proportion  of  one  to  the  other,  in  a  given  lot,  is,  of  course,  of 
no  consequence ;  length  and  breadth  might  be  the  same,  or  one  might 
be  50  per  cent,  or  i  per  cent,  of  the  other;  the  correlation  would 
still  be  complete  (i.ooo)  provided  this  same  proportion  were  main- 
tained throughout  the  particular  lot  examined.  Any  factor  which 
causes  the  proportion  of  breadth  to  length  to  vary  in  a  given  lot,  of 
course  causes  the  correlation  to  fall  below  i.ooo.  If  in  a  given  lot 
many  different  ratios  of  breadth  to  length  are  represented,  the  cor- 
relation is,  of  course,  lowered.  In  such  a  lot,  any  factor  which  tends 
to  make  the  proportion  of  breadth  to  length  more  constant,  of  course, 
increases  the  correlation. 

Examining  the  various  factors  which  have  the  effects  just  men- 
tioned, we  find  that  the  observed  correlation  depends  upon  many 
things. 

(a)  In  considering  the  effects  of  growth  (page  455),  we  saw 
that  the  proportion  of  breadth  to  length  differs  in  different  stages. 
Some  of  the  effects  of  the  environment  on  correlation  are  due  to  its 
effect  on  multiplication  and  growth. 

(6)  Certain  environmental  agents  (as  increased  nutrition)  increase 
the  breadth  while  decreasing  the  length.    Now,  if  this  happens  at 


482  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  ^4, 

the  same  time  and  in  the  same  proportion  in  all  the  individuals,  then 
at  any  given  moment  the  coefficient  of  correlation  will,  of  course, 
not  be  altered  by  it.  But  if  for  any  reason  the  changes  occur  more 
quickly  or  strongly  in  certain  individuals  than  in  others  (as  is  usually 
the  case),  then,  of  course,  the  coefficient  of  correlation  will  be 
decreased.  Or,  if  we  throw  together  individuals  taken  at  different 
stages  of  the  process,  the  correlation  becomes  greatly  decreased ;  it 
may  even  become  negative.  For  examples,  see  rows  15  and  20, 
Table  XVIII. 

(c)  Even  if  a  given  agent  causes  a  change  in  the  same  direction 
(e,  g,,  an  increase)  in  both  length  and  breadth,  the  inclusion  of 
different  stages  in  the  process  may  reduce  the  correlation  (if  it  is 
already  high).  This  will  occur  (i)  if  the  two  dimensions  are  not 
changed  proportionately  to  each  other,  and  (2)  if  the  change  in  a 
given  dimension  varies  at  different  stages  of  the  process.  Both  these 
conditions,  as  we  have  seen,  are  fulfilled  in  the  changes  in  dimensions 
induced  by  the  environment.  Under  almost  any  environmental 
change  breadth  is  altered  more  than  the  length.  Furthermore,  when 
nutrition  is  decreased,  breadth  decreases  more  rapidly  at  first  than 
later.  The  inclusion  of  different  stages  of  the  process  in  a  collection 
therefore  results  in  the  inclusion  of  various  different  proportions  of 
breadth  to  length — lowering  the  correlation. 

(d)  If  the  correlation  is  already  low,  indicating  the  presence  of 
many  different  ratios  of  length  to  breadth,  then  varied  changes  in 
these  ratios  may  compensate  some  of  the  existing  differences,  causing 
an  increase  in  the  correlation.  Whether  this  shall  or  shall  not  occur 
depends  upon  the  condition  of  affairs  before  the  changes  are  made, 
and  on  the  nature  of  the  changes  themselves.  A  special  case  of  this 
comes  up  in  the  next. 

(e)  When  a  culture  containing  thin,  poorly  fed  individuals  is 
given  added  nutriment,  the  correlation  between  length  and  breadth 
increases  (compare,  in  Table  XVIII.,  rows  i  and  2;  8  and  9;  10  and 
II ;  13  and  14;  17  and  18;  17  and  19,  etc.).  This  is  because,  when 
fresh  nutriment  is  added,  the  thinnest,  poorest- fed  individuals  nat- 
urally take  more  food  than  do  the  individuals  that  are  already  plump 
and  well-fed ;  they  therefore  increase  most  in  breadth.  As  a  result, 
existing  differences  in  breadth  are  compensated ;  all  the  animals  take 


,9o8.j  JENNINGS— HEREDITY  IN   PROTOZOA.  483 

on  that  relative  proportion  of  breadth  to  length  that  belongs  to 
well-fed  specimens. 

Thus,  we  find  almost  throughout  that  an  increase  in  the  ratio  of 
breadth  to  length  is  accompanied  by  an-increase  in  the  coefficient  of 
correlation ;  a  decrease  in  the  ratio  of  breadth  to  length  by  a  decrease 
in  the  coefficient  of  correlation.  Examining  these  two  constants,  in 
the  last  two  columns  of  Table  XVIII.,  we  find  this  relation  to  hold 
in  every  case  of  experimental  procedure  save  one.  (In  the  change 
from  row  3  to  row  4  it  does  not  hold ;  this  is  due  to  another  factor, 
to  be  taken  up  later.)  If  without  regard  to  experimental  pro- 
cedure, we  merely  compare  the  mean  index  (or  ratio  of  breadth  to 
length)  with  the  coefficient  of  correlation,  we  find  the  relation  a  little 
less  general,  though  still  marked;  a  large  mean  index  is  usually 
accompanied  by  a  high  coefficient  of  correlation. 

Since,  as  we  have  previously  seen,  greater  breadth  is  usually 
accompanied  by  a  higher  mean  index,  it  follows  that  greater  breadth 
is  likewise  usually  accompanied  by  a  higher  correlation  between 
breadth  and  length.  This  is,  on  the  whole,  evident  on  inspection  of 
Table  XVIII.,  though  since  other  factors  are  involved,  the  relation 
is  not  without  exception.  But  in  general,  broader  specimens  tend 
to  show  a  more  constant  proportion  of  breadth  to  length  than  do 
thin  ones. 

(/)  In  poorly-fed  cultures,  as  we  have  just  seen,  the  breadth  is 
apt  to  be  variable  in  proportion  to  the  length  (giving  low  correla- 
tion) because  some  of  the  individuals  get  more  food  than  others. 
But  if  all  are  reduced  to  an  actually  starving  condition,  then  this 
source  of  variation  is  removed,  and  we  may  again  get  high  corre- 
lation between  breadth  and  length.  This  condition  appears  to  be 
realized  in  row  26  of  Table  XVIII.  Here  a  large  culture  had  been 
reduced  by  starvation  to  a  population  of  but  37,  and  these  give  the 
very  high  correlation  of  .8018  ±  .0396. 

(g)  When  a  given  agent  causes  rapid  multiplication,  so  that  the 
sample  taken  includes  many  diflferent  stages  of  growth,  with  their 
different  proportions  of  breadth  to  length,  the  correlation  becomes 
low.  This  is  the  reason  for  the  marked  decrease  in  correlation  in 
changing  from  row  3  to  row  4  in  Table  XVIII. 

All  together,  it  is  clear  that  no  particular  coefficient  of  correlation 


484  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  m. 

• 

can  be  taken  as  characteristic  of  a  particular  race  of  Paramecia; 
certainly  not  without  very  precise  definition  of  the  conditions.  It 
appears  probable  that  if  all  conditions  of  environment,  growth,  food 
taken,  etc,  could  be  made  absolutely  the  same  for  individuals  derived 
from  the  same  ancestor,  the  coefficient  of  correlation  would  be  close 
to  i.ooo.**  By  varying  these  conditions  any  degree  of  positive  cor- 
relation, down  to  zero,  and  many  degrees  of  negative  correlation 
can  be  attained. 

V.   INHERITANCE  OF  SIZE. 

Having  examined  the  effects  of  growth  and  of  environment  on 
size  and  form,  we  are  now  prepared  to  investigate  how  far  these  are 
determined  by  internal  factors,  handed  on  from  parent  to  progeny. 
Without  such  a  preliminary  study  of  growth  and  environmental 
action  it  would  be  impossible  to  investigate  successfully  the  heredity 
of  size  and  form. 

We  have  already  seen  that  not  all  differences  in  size  are  due  to 
growth  and  environment;  in  the  first  culture  examined  (Table  I., 
page  398)  there  were  at  least  two  sets  of  individuals  of  characteris- 
tic diflferent  sizes,  and  these  diflferences  in  size  are  lasting.  Progeny 
of  the  two  typical  individuals  D  and  c,  from  these  two  sets,  still 
retain  their  characteristic  relative  sizes  after  more  than  a  year  of 
culture  under  all  sorts  of  conditions. 

The  differences  between  these  two  sets  are  about  the  same  as 
those  which  have  been  described  as  distinguishing  two  species,  D 
corresponding  to  the  accounts  of  Paramecium  caudatum,  c  to  Para- 
mecium  aurelia.  The  next  problem  is  to  determine  whether  there 
are  still  other  races  of  Paramecium,  distinguishable  on  the  basis  of 
differences  in  size,  independently  of  the  environment.  Can  we  by 
selecting  individuals  of  differing  sizes  isolate  races  of  corresponding 
sizes?  Can  we  find  races  of  all  sorts  of  sizes  intermediate  between 
the  largest  and  smallest  adult  representatives  of  such  a  heterogeneous 
culture  as  is  shown  in  Table  I.  ? 

The  clear  grouping  of  the  culture  of  Table  I.  into  two  sets  seems 
to  indicate  that  we  have  present  simply  two  races  or  species.    My 

•'Of  course  if  all  variation  disappeared,  as  would  perhaps  be  the  case, 
then  the  concept  of  correlation  would  have  no  further  application. 


x9o8.]  JENNINGS— HEREDITY  IN   PROTOZOA.  486 

first  experiments  consisted  of  attempts  to  break  the  two  lines  derived 
respectively  from  D  and  c  into  other  races  of  different  sizes  by  selec- 
tion and  breeding  of  individuals  of  different  sizes.  This  led  inci- 
dentally, as  we  have  seen,  to  the  study  of  the  effects  of  growth  and 
environment  on  size;  it  was  found  that  the  observable  differences 
between  different  members  of  either  race  were  due  to  these  factors, 
so  that  selection  of  such  members  did  not  lead  to  the  establishment 
of  races  of  different  sizes.  The  results  of  a  large  amount  of  time- 
consuming  work  along  this  line,  done  before  the  investigation  of 
growth  and  environmental  action,  were  throughout  negative.^ 

As  a  result  of  this  work,  I  was  disposed  toward  the  belief  that 
the  characteristic  sizes  of  D  and  c  represent  conditions  of  stability, 
which  have  properly  been  distinguished  as  two  species,  and  that  races 
of  other  sizes  were  not  to  be  found  or  produced. 

But  the  work  thus  far  has,  of  course,  been  based  on  "  pure  lines," 
in  the  sense  in  which  that  expression  is  used  by  Johannsen  ( 1903, 
1906).  The  lines  D  and  c  are  each  derived  from  a  single  individual, 
reproducing  asexually,  so  that  no  admixture  from  outside  has  entered 
them  during  the  experiments.  Now,  while  it  appears  difficult  or 
impossible  to  produce  other  races  within  these  pure  lines,  there 
remains,  of  course,  the  possibility  that  still  other  lines  exist  in  nature. 
Can  we  find  in  a  "wild"  culture,  by  proper  selection  of  differing 
individuals,  still  other  races  of  differing  size  ?  This  was  the  question 
next  investigated. 

I.   Selection  for  Different  Races  in  a  Wild  Culture. 

(a)  Races  Isolated  from  Cultures  Not  Conjugating, 

Attempts  to  separate  out  other  races  than  those  represented  by 
D  {" caudatum  form")  and  c  {'* aurelia  form")  were  first  made 
with  a  wild  culture  which  I  called  OL  This  culture  developed  in 
decaying  vegetation  from  a  marsh.  It  contained  two  well  marked 
sets  of  individuals:  (i)  very  large  individuals,  corresponding  in 
many  respects  to  the  D  line,  but  with  a  mean  length  on  January  3, 
1908,  of  238.280  microns;  these  we  will  designate  £;  (2)  smaller 

*To  the  experiments  on  selection  within  a  pure  line  we  return  in  a  later 
section. 


486  JENNINGS— HEREDITY   IN   PROTOZOA.  rAprila4, 

individuals  corresponding  in  many  respects  to  the  c  line,  with  a 
mean  size  on  November  14,  1907,  of  140.133  microns.  These  two 
sets  occurred  mixed,  but  each  reached  its  maximum  development  at 
the  dates  mentioned.  Isolated  samples  of  the  two  sets  retained  their 
characteristic  differences  in  size,  just  as  happened  in  the  case  of 
D  and  c. 

But. the  interesting  condition  showed  itself  in  the  smaller  set 
An-ong  these  were  individuals  of  such  different  sizes,  that  in  spite 
of  our  knowledge  of  the  great  diflferences  produced  by  growth  and 
environment,  it  seemed  worth  while  to  try  to  isolate  and  breed  them. 
In  a  random  sample  of  60  specimens  the  length  varied  from  96  to 
176  microns — the  smaller  sizes  being  grouped  about  120  microns,  the 
larger  about  160  microns. 

Accordingly,  on  November  9,  1907,  I  separated  two  lots,  one 
containing  ten  of  the  smaller  specimens,  the  other  ten  of  the  larger 
ones.  These  were  placed  in  watch-glasses  with  equal  quantities  of 
the  same  culture  fluid,  and  kept  under  identical  conditions,  where 
they  were  allowed  to  multiply.  One  week  later  (November  16) 
thirty  specimens  measured  from  each  showed  mean  dimensions  of 
125.600  X  36.200  microns  for  the  progeny  of  the  larger  ten,  96.400 
X  30.00  microns  for  the  progeny  of  the  smaller  ten.  On  November 
27,  a  random  sample  of  100  from  each  gave  for  the  progeny  of  the 
larger  ten,  dimensions  of  134.320  X  36.280  microns;  for  the  smaller 
set,  92.240  X  26.920  microns.  Thirty-seven  days  later  (January  2, 
1908)  the  two  lots  still  showed  their  characteristic  differences,  though 
cultivated  under  identical  conditions.  The  mean  dimensions  of  the 
two  sets  (from  random  samples  of  100)  were  now  134.360  X  33-440 
microns  (for  the  larger),  and  104.208X26.583  microns  (for  the 
smaller). 

Thus,  we  have  clearly  two  sets,  with  differences  in  size  persisting 
from  generation  to  generation  (in  spite  of  fluctuations  in  each  due 
to  environmental  changes),  and  both  falling,  in  a  general  way,  in  the 
dimensions  previously  found  for  the  line  c.  It  is  evident,  therefore, 
that  D  and  c  did  not  represent  the  only  existing  different  lines. 

Since  the  two  sets  under  experimentation  had  come  each  from 
ten  individuals  which  may  be  of  heterogeneous  origin,  I  isolated 
from  each,  as  soon  as  it  was  evident  that  they  were  retaining  their 


,9o8]  JENNINGS— HEREDITY   IN   PROTOZOA.  4*87 

differences,  a  single  characteristic  individual.  This  was  done  on 
November  13.  The  specimen  from  the  larget  set  I  called  g;  it 
measured  approximately  130  to  140  microns  in  length.  The  speci- 
men from  the  smaller  set  I  called  «;  its  length  was  about  90  to  95 
microns.  These  two  individuals  were  kept  under  the  same  condi- 
tions and  allowed  to  multiply. 

The  small  specimen  i  multiplied  more  rapidly  than  the  large  one 
g.  On  November  16  there  were  but  seven  progeny  of  g,  while  i  had 
produced  a  large  number.  Two  typical  specimens  of  g  were  killed 
and  gave  measurements  of  160  X  48  microns  and  164  X  56  microns. 
Five  typical  specimens  of  i  ranged  in  size  from  92  X  36  to  128  X  44 
microns,  with  a  mean  of  103.2  X  39-2. 

Evidently,  therefore,  the  progeny  of  g  and  i  tend  to  retain  the 
differences  in  size  characteristic  of  the  parents.  The  two  lines  were 
kept  for  a  long  time,  under  the  same  conditions ;  at  intervals  random 
samples  were  measured.  The  measurements  at  different  dates,  with 
the  number  of  specimens  on  which  they  are  based  are  given  in 
Table  XXIIL,  p.  488.  (The  small  numbers  of  specimens  employed 
on  certain  dates  are  due  to  the  fact  that  only  a  small  number  ex- 
isted at  that  time.) 

The  great  fluctuations  in  the  dimensions  of  each  line  will  of 
course  surprise  no  one  who  has  examined  that  part  of  this  paper 
which  deals  with  the  effects  of  the  environment.  These  fluctua- 
tions are  due  mainly  to  differences  in  nutritional  conditions.  At 
intervals  it  was  necessary  to  add  new  culture  fluid ;  the  dimensions 
in  both  lines  thereupon  rose  at  once;  they  then  gradually  declined 
till  new  fluid  was  added.  Details  on  this  matter  are  not  necessary 
for  our  present  purpose. 

The  important  fact  is,  that  in  spite  of  all  fluctuations,  the  lines  g 
and  i  retained  throughout  the  three  months  in  which  they  were 
under  observation  their  characteristic  relative  sizes.  Multiplication 
was  probably  at  the  rate  of  about  one  fission  a  day,  so  that  the  table 
represents  90  to  100  generations.  We  have  here  two  lasting  races 
comparable  to  the  two  races  from  our  first  culture,  which  we  called 
D  and  c.  It  is  clear  that  neither  g  nor  i  is  identical  with  D,  since 
the  latter  is  much  larger;  whether  either  is  the  same  as  c  we  shall 
inquire  later. 


486 


JENNINGS— HEREDITY   IN   PROTOZOA. 


I  April  a4. 


Table  XXIII. 

Comparative  Sises  in  Microns  of  g  and  i  and  their  Progeny  at  Different 
Dates,  when  Cultivated  under  the  Same  Conditions. 


£'  and  Its  Progeny 

1 

/and  Its  Progeny. 

Date. 

No.  of 

Mean 

Mean 

No.  of 

Mean                 Mean 

Specimens. 

Length. 

Breadth. 

Specimens. 

Length.             Breadth. 

1907 

Nor.   13 

I 

130-140 

35-40? 

I 

90-95             30-40? 

"      16 

2 

162.000 

52.000 

5 

103.200             39.200 

"      18 

7 

140.000 

40.000 

12 

101.666             35.666 
88.268      1       30.268 

,«      23 

30 

129.333 

34.933 

30 

*'      26 

100 

137.120 

38.720 

100 

99.560             28.200 

Dec.     7 

61 

120.590 

4I.IIO 

96 

98.709             34.208 

«*      16 

17 

127.059 

38.588 

23 

98.608      ,       29.739 

"     30 

40 

112.600 

31300 

64 

86.756             22.062 

X908 

1 

Jan.    2 

100 

146.640 

40.600 

100 

106.680             26.400 

Feb.    5 

57 

II6.9I2 

36.070 

43 

93.583             27.500 

It  will  be  recalled  that  in  the  original  culture  from  which  came 
g  and  i,  there  was  a  still  larger  set  which  we  called  £.  Ten  of  these 
were  selected  and  cultivated  under  the  same  conditions  as  g  and  ». 
They  retained  throughout  their  much  larger  size  (numerical  results 
are  given  later),  so  that  from  this  culture  we  have  isolated  three 
lines  or  races  which  retain  their  differences  in  size  under  the  same 
external  conditions. 

At  this  period,  then  (January  i,  1908),  I  had  in  the  laboratory  a 
number  of  lines  or  races  which  had  been  studied  with  care.  These 
formed  two  sets,  so  far  as  our  knowledge  of  them  up  to  this  point  is 
concerned.  The  two  lines,  D  and  c,  from  culture  /,  were  clearly 
distinct  even  under  identical  conditions.  The  three  lines,  g,  i  and  E, 
from  the  second  wild  culture  01,  are  likewise  clearly  distinct  from 
each  other.  But  the  relation  of  g,  i  and  E  to  D  and  c  is  uncertain ; 
we  may  have  on  hand  five  distinct  lines,  or  only  four,  or  three. 

To  determine  whether  any  of  these  five  lines  are  identical,  it  is 
necessary  to  cultivate  all  five  under  the  same  conditions.  A  certain 
number  must  be  selected  from  each ;  these  must  be  brought  into  the 
same  culture  fluid  and  allowed  to  multiply  in  the  same  environment. 

It  is  extraordinary  what  difficulties  are  presented  in  carrying  out  this 
apparently  simple  plan.  The  different  lines  have  become  adapted  to  certain 
diverse  nutritive  conditions;  if  now  they  are  brought  at  once  into  the  same 
culture  fluid,  some  of  them  die.  In  the  present  case,  g  and  i  had  been  living 
in  comparatively  fresh  hay  infusion,  D  and  c  in  different  old  hay  cultures, 
£  in  a  culture  of  decaying  pond  weeds.    When  all  were  brought  into  fresh 


,9o8]  JENNINGS— HEREDITY   IN   PROTOZOA.  489 

hay  infusion,  E  died  at  once,  c  after  a  day  or  two ;  D  multiplied  slowly,  then 
died  in  the  course  of  a  week  or  so,  while  g  and  •  throve  and  multiplied. 

It  was  therefore  necessary  to  bring  the  different  lines  gradually  into  the 
new  fluid,  by  mixing  some  of  it  with  the  fluid  in  which  they  lived,  increasing 
the  proportion  of  new  fluid  at  intervals.  This  was  found  to  be  a  very  deli- 
cate undertaking.  Certain  of  the  lines  would  thrive  for  a  time,  under  this 
procedure,  then  would  begin  to  degenerate;  in  this  way  much  time  was  lost. 
Finally,  however,  the  different  sets  were  induced  to  thrive  in  the  same  hay 
infusion. 

Procedure  Necessary  for  Making  the  Conditions  Identical  for  Different 
Lines. — ^The  procedure  followed,  in  order  to  be  certain  that  the  cultural  con- 
ditions were  the  same  for  all,  was  as  follows:  From  each  race  ten  typical 
individuals  were  selected.  These  were  mixed  with  gradually  increasing 
amounts  of  hay  infusion,  in  the  way  just  set  forth — while  at  the  same  time 
of  course  they  multiplied  in  number.  After  they  had  all  gotten  accustomed 
to  the  infusion,  it  was  necessary  to  take  measures  to  assure  the  identity  of 
the  solutions  in  which  the  different  sets  were  living.  For  this  it  is  not 
sufficient  merely  to  transport  the  individuals  to  definite  quantities  of  the  same 
nutritive  solution.  For  up  to  this  point  each  set  has  been  living  in  a  solution 
which  has  received  an  admixture  of  the  original  culture  for  that  set.  Now, 
these  different  original  cultures  contained  different  kinds  of  bacteria.  On 
transferring  the  infusoria  to  the  hay  infusion,  they  of  course  carried  some  of 
their  own  bacteria.  By  repeated  changes  the  number  of  bacteria  introduced 
could  be  much  reduced.  Nevertheless  different  kinds  were  brought  in  in 
different  cases,  so  that  we  still  have  the  different  lines  in  cultures  of  diverse 
bacteria.  From  this  fact  naturally  diverse  chemical  properties  may  develop 
in  the  different  cultures,  though  the  basic  nutritive  solution  is  the  same. 
These  diverse  chemical  properties  would  of  course  modify  the  organisms, 
making  it  impossible  to  compare  them  with  regard  to  inherited  size.  To 
make  the  conditions  of  existence  the  same,  it  is  not  sufficient  to  attend 
merely  to  the  basic  fluid;  the  bacteria  in  the  fluid  must  also  be  the  same. 
This  is  a  principle  of  wide  practical  importance  in  all  experimental  work  with 
such  infusoria.  It  is  not  a  mere  theoretical  requirement;  death  frequently 
results  from  the  introduction  of  a  certain  kind  of  bacteria  into  a  certain 
culture,  while  another  culture  of  identically  the  same  fluid  flourishes,  be- 
cause the  bacterial  infection  is  different. 

This  requirement  was  met  in  the  following  way:  After  the  different 
sets  had  become  acclimatized  to  the  same  hay  infusion,  ten  of  each  were 
removed  with  a  fine  capillary  pipette,  and  washed  twice  in  fresh  hay  in- 
fusion. The  second  washing  of  the  different  sets  was  done  in  the  same  mass 
of  fluid, — a  small  watch-glass  full.  The  different  sets  might  of  course  each 
carry  with  them  a  few  of  the  bacteria  characteristic  of  their  original  culture. 
After  all  had  been  washed  in  the  same  mass  of  fluid,  this  fluid  would  of 
course  be  infected  with  bacteria  from  all  the  different  sets.  Now,  after  the 
washing  was  finished,  a  definite  quantity  of  this  fluid  in  which  all  had  been 
washed  was  added  to  the  final  culture  fluid  for  each  lot. 

Thus  each  lot  of  ten  is  in  the  same  quantity  of  the  same  nutritive  fluid, 

PROC.  AMER.  PHIL.  SOC.  XLVII.   I90  FF,  PRINTED  JANUARY  12,  I909. 


490  JENNINGS— HEREDITY  IN   PROTOZOA.  [April  m. 

and  infected  with  the  same  bacteria  as  all  the  others.  All  are  kept  in  watch- 
glasses  of  the  same  form  and  size,  close  together  in  the  same  moist  chamber. 
Any  characteristic  differences  in  the  resulting  progeny  must  then  be  due  to 
conditions  within  the  animal,  and  not  to  differences  in  the  environment  If 
we  reach  the  same  result,  not  merely  in  one  experiment,  but  in  a  series  con- 
ducted in  this  manner,  we  can  be  sure  of  our  results. 

Cultures  of  the  five  lines,  D,  c,  g,  %  and  E,  prepared  in  the  way 
just  described,  were  set  in  progress  January  19,  1908.  In  order  to 
determine  with  certainty  how  much  effect  possible  environmental 
differences  might  have  on  the  results  (as  well  as  for  certain  other 
purposes),  two  lots  each  of  D,  g  and  i  were  used.  If  the  two  lots 
of  g,  for  example,  show  differences  as  great  as  those  between  g  and 
c,  then,  of  course,  we  have  no  ground  for  considering  g  and  c  inher- 
ently different ;  the  environmental  differences  account  for  all.  These 
lots  were  allowed  to  multiply  till  February  5.  Then  a  sample  of 
each  was  killed  and  measured.  Now  a  new  lot  of  ten  of  each  set 
was  prepared  by  the  methods  given  above,  and  the  animals  were 
again" allowed  to  multiply  till  February  15,  when  samples  were  again 
measured. 

It  will  be  recalled  that  £  is  a  lot  derived  from  ten  specimens  of  possibly 
diverse  ancestry,  from  the  culture  01,  with  an  original  mean  length  of 
238.280  microns;  that  the  line  D  has  shown  in  repeated  determinations  a 
highest  mean  length  of  202.280  microns  (Table  XVIII.)  ;  that  r,  g  and  i  are 
smaller  lines,  derived  from  single  individuals;  g  is  known  to  be  larger  than 
1,  but  the  relation  of  c  to  these  is  unknown. 

The  results  of  these  breeding  experiments  are  given  in  the  fol- 
lowing Table  XXIV. 

The  experimental  results  given  in  this  table  show  certain  things 
clearly. 

I.  The  method  of  culture  is  adequate  for  bringing  out  the  inher- 
ent differences  in  different  lines  without  confusion  due  to  environ- 
mental effects.  This  is  shown  by  the  fact  that  when  two  cultures 
are  made  from  certain  single  lines,  these  show  themselves  after 
breeding  for  many  generations  to  be  nearly  identical,  while  the 
different  lines  give  diverse  results.  In  only  one  case  (Z>  on  Feb- 
ruary 15)  is  there  a  notable  difference  btween  the  two  samples  of  a 
single  line,  but  this  is  much  less  than  the  difference  between  that  line 
and  any  other. 


i9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA. 


491 


Table  XXIV. 

Mean  Dimensions  in  Microns  of  the  Five  Lines  E,  D,  c.  g  and  i,  when  CulH' 
voted  under  the  same  Conditions,  January  19  to  February  5  and 

February  5  to  February  15, 


Dunensions  of  E, 

Dimensions  of  D.                      Dimensions  of  c. 

Date. 

43 

100 
100 

it 
1 

*a 

57 
19 

100 
100 

100 

it 

89 

*a 

60 
100 

Feb.     5 

Feb.  15 
Feb.  27 

169.395X52.930 

200.320X52.400 
172.040X55-520 

(1)  169.754X46.877 

(2)  169.895X43-579 

(1)  180.240X46.880 

(2)  173.240X49.760 

175-3^X47.160 

99.667x26.333 
100.320x26.480 

Dimensions  of/*. 

Dimensions  of  t. 

Date. 

^1 

50 

57 

jIOO 

si 

so 
48 

100 

Feb.     5 
Feb.  15 

(1)  114.720X33-920 

(2)  116.912X36.070 

125.240X35440 

(1)  92.000X26.960 

(2)  93.583X27.500 

1           95.440X3<^040 

2.  At  least  four  distinct  lines  are  present,  D,  c,  g  and  »;  these 
maintain  their  relative  different  sizes  throughout  the  experiments, 
which  lasted  about  twenty-five  generations. 

3.  The  lines  E  and  D  are  nearly  or  quite  the  same.  On  February 
5  they  show  nearly  the  same  measurements,  but  on  February  15 
there  was  a  marked  difference.  To  test  the  meaning  of  this  these 
two  were  cultivated  twelve  days  more;  then  on  February  27  they 
gave  again  nearly  the  same  measurements.  It  will  doubtless  be 
sa^fest  to  consider  them  the  same. 

We  have  now,  therefore,  four  different  lines  or  races  of  Para- 
mecium, characterized  by  persisting  relative  differences  in  size. 
One  of  these  {D  and  E)  belongs,  from  its  size,  to  the  " caudatum 
group  " ;  the  other  three  are  much  smaller  and  fall  in  the  "  aurelia 
group."  Of  these,  g  is  the  largest,  i  the  smallest,  while  c  is  inter- 
mediate. Under  a  similar  change  in  the  environment  these  all  change 
in  a  corresponding  way,  as  is  shown  by  the  fact  that  on  February  15 
all  were  somewhat  larger  than  on  February  5.     It  may  be  noted  that 


492  JENNINGS— HEREDITY   IN   PROTOZOA.  (April  m, 

the  differences  in  size  among  these  four  lines  were  very  evident  to 
the  eye  on  inspection  with  the  low  power  of  the  microscope,  and 
that  the  difference  was  clearly  present  at  all  ^riods  between  the 
dates  when  the  measurements  were  made.  The  measurements 
merely  make  precise  what  is  evident  to  the  eye  without  them. 

Before  attempting  to  determine  whether  still  other  lines  can  be 
isolated,  and  particularly  whether  it  is  possible  to  fill  the  wide  gap 
between  the  caudatum  group  and  the  aurelia  group,  another  question 
must  be  investigated — a  question  which  strikes  at  the  foundation  of 
our  conclusions  up  to  this  point.  This  is  the  question  of  the  relation 
of  these  lines  of  diverse  size  to  conjugation  and  the  life  cycle. 

(&)  Are  the  Lines  of  Different  Size  Merely  Different  Stages  in  the 

Life  Cycle? 

Calkins  (1906)  and  others  have  set  forth  the  fact  that  Para- 
mecium  and  other  infusoria  show  different  dimensions  in  different 
stages  of  the  life  cycle — ^the  cycle  which  begins  with  conjugation, 
extends  over  many  generations  of  reproduction  by  fission,  and  ends 
with  another  conjugafion.  The  question  arises,  therefore,  whether 
our  lines  of  diverse  dimensions  are  not  merely  different  stages  in  the 
life  cycle ;  whether  they  would  not,  if  brought  to  the  same  stage  of 
the  cycle,  show  the  same  dimensions.  This  possibility  must  be 
investigated  before  we  proceed  farther. 

The  details  of  the  relation  of  conjugation  and  the  life  cycle  to 
variation,  inheritance,  etc.,  are  to  be  dealt  with  in  a  separate  paper 
of  this  series.  But  since  the  question  which  stands  at  the  head  of 
this  section  is  an  absolutely  fundamental  one  for  the  proper  inter- 
pretation of  the  results  of  the  present  paper,  it  must  be  dealt  with  here. 

To  answer  this  question,  it  is  evidently  necessary  to  proceed  as 
follows:  Cultures  showing  epidemics  of  conjugation  must  be  exam- 
ined for  conjugating  pairs  of  diverse  sizes.  If  such  are  found,  the 
individuals  must  be  isolated  and  allowed  to  multiply,  in  order  to 
determine  whether  the  progeny  retain  the  diverse  sizes  characteristic 
of  the  parents.  If  from  a  conjugating  culture  we  can  obtain  diverse 
lines  standing  all  in  the  same  relation  to  conjugation  and  the  life 
cycle,  then  evidently  our  diverse  lines  represent  something  more 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  493 

than  different  stages  in  the  life  cycle.  The  problem  also  can  be 
attacked  in  certain  other  ways,  which  will  be  described. 

The  relation  of  diverse  sizes  to  conjugation  and  the  life  cycle 
was  studied  with  special  thoroughness  in  the  case  of  a  culture  in 
which  there  was  an  epidemic  of  conjugation  January  29, 1908.  This 
culture  was  found  in  decaying  vegetation  from  a  small  pond  near 
Baltimore ;  I  called  it  culture  M.  Table  LXI.  (appendix)  shows  a 
random  sample  of  this  culture,  including  both  conjugants  ahd  non- 
con  jugants;  of  the  238  specimens  in  the  table,  38  were  conjugants, 
200  non-con  jugants. 

From  this  culture  M  a  large  number  of  pairs  were  isolated,  for 
various  purposes,  and  allowed  to  multiply.  Without  going  here  into 
the  details  of  the  experiments,  on  February  21  I  had  from  this  cul- 
ture eight  sets  or  lines,  each  descended  from  a  single  equal  pair  or 
a  single  ex-con jugant;  these  lines  were  designated  in  my  notes  L2, 
Gi,Ai,A2, 1,  C2,  Fi  and  F2,  (The  designations  are  the  same  as  those 
given  to  the  original  pair  or  individual  from  which  the  lines  came.) 
In  addition  to  these  eight  "  pure  lines,"  I  had  two  cultures  derived 
each  from  eight  pairs  of  conjugants  of  approximately  the  same  size; 
these  were  called  Ki  and  K2,  A  final  culture  was  derived  from  ten 
small,  nearly  equal,  non-con  jugants  from  the  same  culture;  it  was 
designated  H. 

It  is,  of  course,  unfortunate  that  it  is  not  possible  to  measure 
accurately  the  original  living  individuals  from  which  the  different 
lines  are  derived,  but  this  will  not  alter  in  any  way  the  results  on 
the  problem  in  which  we  are  at  present  interested.  The  essential 
question  is  whether  the  lines  derived  from  the  different  pairs  or 
individuals  ar^  identical  or  diverse  in  size. 

These  various  cultures  were  kept,  so  far  as  possible,  in  the  same 
nutritive  fluid  and  under  the  same  conditions.  Marked  differences 
in  size  were  apparent  on  examining  the  different  sets  with  low  power 
of  the  microscope.  On  February  21  fifty  individuals  of  each  of 
these  eleven  different  sets  were  brought,  with  all  the  precautions 
mentioned  on  page  489,  into  the  same  culture  fluid,  while  at  the  same 
time  fifty  specimens  each  of  D  and  g  of  our  earlier  pure  lines  (see 
page  491)  were  brought  into  the  same  fluid.  These  were  all  allowed 
to  multiply  till  February  26,  when  a  random  sample  of  100  or  more 


494 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  24, 


of  each  was  killed  and  measured.  Later,  on  March  7,  twenty  indi- 
viduals were  taken  anew  from  each  of  these  thirteen  lots,  brought 
again  with  elaborate  precautions  into  the  same  culture  fluid,  kept 
under  the  same  conditions  and  allowed  to  multiply,  part  till  March 
13,  part  till  March  19,  when  other  samples  were  killed  and  measured. 
From  our  previous  extensive  experience  with  i  and  g  (Table  XXIII., 
page  488)  and  with  five  lines  of  Tables  XXIV.  (page  491),  we  can 
be  assured  that  two  sets  of  measurements  taken  at  such  intervals 
will  give  us  reliable  data  as  to  the  existence  of  any  considerable 
lasting  differences  among  the  different  lines.  The  results  of  the 
measurements  of  the  thirteen  different  sets  are  given  in  classified 
form  in  Table  XXV. 

Table  XXV. 

Mean  Dimensions  in  Microns,  of  the  Thirteen  Sets  Described  in  the  Text, 
after  Cultivation  under  the  Same  Conditions,  February  21  to  February 
26,  and  March  7  to  March  13  {or  March  19),  (The  conditions  before; 
and  in  intervening  periods  were  essentially  the  same,  but  elaborate  pre- 
cautions were  taken  for  the  periods  specified).  All  are  from  the  con- 
jugating culture  M,  of  January  29,  save  the  last  two  sets. 


^1 

^1 

IJne. 

i 

February  96. 

§3 

March  13. 

March  19. 

^1 

^1 

==a 

(I) 

Dcsc 

endants  of  Pairs. 

Z2 

100 

206.360  X  60.840 

100 

220.560  X  59.960 

Gl 

100 

201.400  X  52.400 

100 

210.960  X  52.200 

A  1 

ICO 

193-560  X  51840 

100 

203.640X52.560 

A  2 

100 

184.640x50-760 

100 

187.878  X  44.490 

/ 

100 

132.880X41.960 

100 

138.880  X43.I20 

€2 

100 

128.880X40.400 

100 

119.200X37.280 

(2)  Descendants  of  Single  Ex-conjugants. 

Fl 

100 

193.000X50.840 
182.200  X  51.040 

56 

209.643  X  56.643 

F2 

100 

100 

199.960  X  50.120 

(3)  Descended  each  from  8  Equal  Pairs. 

K\ 

100 

133-680  X  39400 

Kz 

100 

125.920X37.040 

100 '  125.000  X  42.520 

(4)  Descended  from  10  Small  Non  conjugants. 

H 

100 

131.400X43.000  1          1                                   •  100' 128.840X41.360 
(5)  Older  Lines,  not  from  Culture  M, 

D 

III 

176.901  X  50.018 

120 

187.033X49.100 

S 

100 

124.440X35-920 

140.800  X  39.640 

Examination  of  this  table  shows  that  lines  derived  from  different 
conjugating  pairs  or  different  ex-con jugants  do  differ  from  each 
other  at  the  same  periods  in  the  life  cycle,  even  though  living  under 


,9o8|  JENNINGS— HEREDITY  IN   PROTOZOA.  495 

identical  conditions.  The  differences  are  fully  as  marked  as  tliose 
found  among  diverse  lines  derived  from  individuals  not  conjugating 
and  taken  without  reference  to  the  period  in  the  life  cycle  in  which 
they  happen  to  be. 

Besides  this  general  result  on  our  main  problem,  the  following 
important  facts  are  brought  out  by  the  table : 

1.  The  six  lines  derived  from  the  six  different  pairs  (first  six  of 
the  table)  are  clearly  distinct.  They  show  parallel  differences  in 
both  sets  of  tests ;  the  order  of  dimensions  from  largest  to  smallest 
is  the  same  in  both  the  first  and  the  second  measurements,  though 
these  are  separated  by  at  least  fifteen  generations. 

2.  The  two  lines,  Fi  and  F2,  derived  from  single  ex-con jugants, 
are  likewise  distinct  from  each  other.  So  far  as  the  measurements 
go,  Fi  may  possibly  be  the  same  as  Ai,  F2  as  A2, 

3.  Certain  different  sets  are  likewise  found  in  the  other  lots  of 
the  table. 

4.  The  different  sets  fall  into  two  very  distinct  groups,  whose 
dimensions  are  separated  by  a  wide  interval.  To  the  large  group 
belong  L2,  Gi,  Ai,  A 2,  Fi,  F2  and  D.  To  the  small  group  belong 
the  others.  The  greatest  mean  length  of  any  set  of  the  smaller 
group  (140.800  microns)  differs  widely  from  the  least  mean  length 
of  any  set  of  the  larger  group  ( 176.901  microns).  These  two  groups 
correspond  in  general  to  what  we  have  heretofore  called  the  "  aurelia 
form  "  and  the  "  caudatum  form." 

As  there  was  no  danger  of  confusing  any  lot  of  the  larger  group  with 
any  lot  of  the  smaller  one,  the  second  measurements  of  the  two  groups  were 
not  made  for  the  same  day;  the  lots  of  the  larger  group  were  killed  March 
13,  while  those  of  the  smaller  group  were  not  killed  till  March  19,  as  the 
table  shows.  This  was  done  on  acount  of  the  great  labor  involved  in  select- 
ing, with  capillary  pipette,  killing  properly,  and  preserving,  so  many  different 
sets  on  the  same  day.  This  difference  of  treatment  of  course  does  not  alter 
the  comparability  of  the  different  sets  within  a  given  group,  which  is  all  that 
we  require. 

5.  How  shall  we  decide  which  of  the  thirteen  different  sets  form 
distinct  lines?  For  this  it  will  be  best  to  take  into  consideration 
mainly  the  length,  since  we  know  from  our  earlier  studies  that  little 
significance  is  to  be  attached  to  difference  in  breadth,  owing  to  the 
extreme  changes  in  that  dimension  with  slight  differences  in  food. 


496  JENNINGS— HEREDITY   IN   PROTOZOA.  [Aprils. 

If  any  two  sets  differ  in  length  in  the  same  way  at  both  measure- 
ments (taken  many  generations  apart)  and  if  the  differences  between 
them  are  each  time  decidedly  greater  than  the  sum  of  the  probable 
errors  of  the  measurements  of  the  two,  then  we  can  be  assured  that 
we  are  dealing  with  really  differentiated  sets.  Now,  examination  of 
the  extensive  series  of  measurements  in  Tables  X.  and  XVIII.  shows 
that  the  probable  error  of  the  mean  length  never  reaches  two  microns, 
even  when  the  number  of  specimens  is  much  smaller  than  in  our 
present  measurements,  and  when  conditions  are  of  the  most  varied 
character.  It  is  practically  certain  that  the  probable  error  of  the 
mean  length  would  not  amount  to  one  micron  in  any  of  the  compara- 
tively homogeneous  sets  with  which  we  are  here  dealing.  If,  then, 
we  require  a  difference  of  four  microns  between  the  mean  lengths 
of  the  two  sets,  this  difference  to  have  the  same  sign  (-(-or  — )  at 
*both  measurements,  we  shall  be  within  safe  limits.  Applying  this 
test,  we  find  four  lines  clearly  distinct  in  the  larger  or  "  caudatum  " 
group,  while  in  the  smaller  or  "  aurelia  "  group  we  can  be  certain  of 
but  two  distinct  lines  (represented  best  perhaps  by  /  and  C2),  We 
have  previously  found  three  distinct  lines  in  the  aurelia  group  (c,  g 
and  i,  Table  XXIV.),  so  that  all  together  we  now  have  at  least  seven 
different  lines  of  Paramecium,  showing  constant  relative  differences 
in  length.  It  is  probable  that  very  exact  tests  would  show  the  dis- 
tinctness of  some  other  lines  of  Table  XXV. 

The  striking  difference  between  adults  of  different  races,  under 
varied  conditions,  is  shown  in  Fig.  7.  Here  we  have  two  adults,  one 
belonging  to  our  smallest  race  (i)  ;  the  other  to  one  of  the  large  races. 


Fig.  7.  Extreme  adult  sizes  from  different  pure  lines  of  Paramecium, 
a,  large  individual  from  a  large  line,  h,  small  individual  from  the  small  line 
f  of  Table  XXIII.,  page  488.    Both  magnified  235  diameters. 


tgos.]  JENNINGS— HEREDITY  IN  PROTOZOA.  497 

It  is  clear,  then,  that  the  question  placed  at  the  head  of  the 
present  section  is  to  be  answered  in  the  n^ative.  The  diverse  lines 
of  diflferent  size  are  not  merely  diflferent  stages  in  the  life  cycle. 

(c")  Other  Evidences  of  Permanent  Differentiation  in  Size,  Inde- 
pendent of  the  Life  Cycle. 

The  proof  just  given,  that  lines  beginning  with  conjugants  are 
differentiated  in  size  even  in  the  same  portion  of  the  life  cycle  and 
under  the  same  conditions,  is  conclusive.  But  it  may  be  worth  while 
to  give  briefly  certain  other  evidences  of  the  same  thing. 

1.  First  we  have  the  fact  that  in  a  given  culture  the  conjugants 
themselves  differ  in  size;  this  has  already  been  shown  by  Pearl 
(1907).  In  a  certain  Culture  IV.,  I  found  conjugants  varying  in 
dimensions  from  148  X  44  to  260  X  60  microns.  I  have  found  (not 
in  the  same  culture)  conjugants  with  length  as  low  as  100  microns. 
It  is  clear,  therefore,  that  not  all  individuals  are  of  the  same  size 
at  conjugation.  There  is  no  reason  to  expect  them  to  be  so,  there- 
fore, at  other  definite  periods  in  the  life  cycle;  as  we  have  seen,  they 
are  not.  Selection  of  small  pairs  gives  small  progeny ;  of  large  pairs, 
large  progeny. 

2.  In  certain  of  my  pure  lines  whose  history  was  followed  for  a 
long  time  and  whose  dimensions  were  taken  at  intervals,  conjugation 
occurred  at  times,  but  the  dimensions  at  such  times  were  not  very 
different  from  the  dimensions  at  other  periods  in  the  life  history. 
Thus,  in  the  earlier  sections  of  this  paper  we  have  dealt  with  two 
pure  lines,  D  and  c;  the  former  showed  usually  a  mean  length  of 
about  180  microns,  the  latter  a  mean  length  of  about  130  microns 
(see  Table  XVIII.).  At  a  certain  time  an  epidemic  of  conjugation 
arose  in  c.  The  mean  dimensions  were  indeed  higher  than  usual 
at  that  time,  the  mean  length  of  the  conjugants  rising  to  158.496 
microns.  But  this  does  not  by  any  means  bring  it  up  to  the  ordinary 
mean  of  D,  and  immediately  after  conjugation  (in  five  days)  the 
mean  length  of  c  fell  back  to  129.640  microns.  Again,  in  the  small 
race  g,  of  Table  XXIII.,  conjugation  occurred  in  a  number  of  cases ; 
a  typical  pair  measured  but  no  microns  in  length.  In  other  lines  I 
have  found  for  the  conjugants  means  as  high  as  199.024  and  as  low 


498  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

as  116.856,  and  these  were  correlated  with  corresponding  measure- 
ments throughout  the  series. 

These  facts,  of  course,  do  not  show  that  the  size  may  not  change 
at  the  time  of  conjugation  or  before  or  after.  What  they  do  show 
is  that  any  diflferences  thus  produced  do  not  account  for  the  perma- 
nent differentiations  we  have  found  among  different  lines.  We  may 
distinguish  (i)  differences  in  size  due  to  growth;  (2)  those  due  to 
nutrition  and  other  environmental  conditions;  (3)  those  due  to 
different  stages  in  the  life  cycle  (as  a  rule  not  marked  in  comparison 
with  the  others)  ;  (4)  inherent,  hereditary  differences  in  size,  per- 
sisting when  all  other  conditions  are  made  the  same. 

(d)  Lines  Intermediate  Between  the  Two  Main  Groups,     The 

Question  of  Species  in  Paramecium. 

As  we  have  already  noted,  the  seven  differentiated  lines  which 
we  have  thus  far  distinguished  fall  into  two  main  groups,  separated 
by  a  wide  interval.  In  Table  XXV.  we  find  one  group  with  mean 
lengths  varying  from  119.200  to  140.800  microns,  while  in  the  other 
group  the  mean  lengths  vary  from  176.901  to  220.560  microns. 
Between  the  two  there  is  thus  a  gap  of  36.101  microns  in  which  none 
are  found.  Is  this  gap  constant  and  characteristic,  so  that  our  two 
large  groups  are  permanently  differentiated?  If  so,  we  should  have 
some  real  basis  for  the  common  distinction  into  two  species,  Para- 
mecium caudatum  (larger)  ^nA Paramecium  aurelia  (smaller).  The 
fact  that  we  find  in  nature  such  cultures  as  that  shown  in  Table  I. 
(page  398),  in  which  the  individuals  are  distinctly  separated  into  the 
two  groups,  seems  to  raise  a  presumption  that  the  groups  are  natural 
ones,  not  due  to  accidents  of  selection. 

For  a  long  time  I  found  no  pure  lines  that  were  intermediate 
between  these  groups.  It  is  possible  that  this  was  partly  due  to  a 
tendency  to  choose  for  breeding  the  largest  and  smallest  specimens, 
rather  than  intermediate  ones,  since  my  purpose  at  first  was  to  deter- 
mine whether  there  were  any  permanent  differentiations  at  all;  for 
this,  marked  differences  were  desirable. 

In  the  course  of  work  on  certain  problems  connected  with  con- 
jugation, I  came  in  possession  of  a  pure  line,  Nf2,  descended  from  a 
single  ex-conjugant.     This,  when  cultivated  in  the  usual  hay  infu- 


,9o8.j  JENNINGS— HEREDITY   IN   PROTOZOA.  499 

sion,  gave,  under  various  different  conditions,  the  following  mean 
lengths  in  microns  (each  mean  is  based  on  measurements  of  lOO 
individuals)  :  148.197,  151.920,  158.760,  153.320,  160.852,  156.482. 

It  is  evident  that  these  means  fall  in  the  gap  separating  the 
*' caudatum"  group  from  the  " aurelia"  group.  I  therefore  decided 
to  cultivate  these  under  identical  conditions  with  a  typical  repre- 
sentative of  each  of  the  two  main  groups.  For  this  purpose  I  chose 
D  and  c,  the  two  lines  longest  cultivated,  which  I  had  used  for  the 
study  of  growth,  environmental  action,  etc.  (Tables  X.,  XVIII., 
etc.).  Twenty-five  specimens,  each  of  the  three  lines,  D,  c  and  Nf2, 
were  brought  on  May  i,with  the  precautions  described  on  page  489, 
into  the  same  quantity  of  the  same  hay  infusion  and  allowed  to  mul- 
tiply tin  May  5.  On  that  date  a  random  sample  of  each  was  killed. 
Though  the  samples  were  large,  extrinsic  conditions  prevented  my 
measuring  more  than  the  numbers  mentioned  below;  larger  numbers 
would  not  have  altered  the  results  by  more  than  one  or  two  microns 
in  any  case.  The  mean  dimensions  of  these  three  lines,  cultivated 
under  identical  conditions,  were 

D  (31  specimens),  202.710  X  51.871  microns. 

^/^  (33  specimens),  168.970X48.970  microns. 

c  (43  specimens),  126.605  X  44-930  microns. 

Thus,  the  dimensions  of  AT/^  lie  almost  precisely  half  way  between 
those  of  D  and  c  (the  dimensions  exactly  half  way  between  would 
be  164.658X48.401).  We  have,  therefore,  in  Nf2  an  eighth  pure 
line,  intermediate  between  the  "  caudatum"  and  *' aurelia''  groups 
formed  by  the  other  seven.  These  two  groups  are  then  not  sepa- 
rated by  an  unbridged  gap. 

The  other  character  which  had  been  held  to  separate  Paramecium 
caudatum  from  Paramecium  aurelia  was  the  presence  of  but  a  single 
micronucleus  in  the  former,  while  the  latter  had  two.  Calkins 
(1906)  showed  that  in  the  same  pure  line  we  sometimes  have  two 
micronuclei,  sometimes  but  one,  so  that  this  is  not  sufficient  ground 
for  distinguishing  two  species.  Though  the  present  study  has  shown 
that  differences  in  size  among  different  lines  are  more  permanent 
than  the  data  available  to  Calkins  had  seemed  to  indicate,  this  does 
not  give  any  better  basis  for  distinguishing  two  species,  since  we 


500  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

have  been  able  to  isolate,  not  merely  two  permanently  differentiated 
lines,  but  eight.  Of  course,  it  would  require  merely  more  extensive 
and  intensive  work  to  isolate  others;  doubtless  the  number  to  be 
isolated  would  depend  only  on  the  accuracy  of  the  methods  used. 

To  my  great  regret,  I  was  unable  to  take  the  steps  necessary  to 
determine  the  number  of  micronuclei  in  the  various  pure  lines  with 
which  I  worked.  The  animals  multiply  so  rapidly  that  with  several 
lines  in  progress  it  is  quite  impossible  even  to  keep  up  with  the  data 
for  size  alone;  probably  half  my  experiments  were  lost  on  this 
account,  after  much  work  had  been  spent  on  them.  It  was  then  out 
of  the  question  to  carry  on  at  the  same  time  the  staining  processes 
necessary  to  determine  with  certainty  the  number  of  micronuclei. 
For  work  of  the  kind  presented  in  this  paper,  a  syndicate  of 'investi- 
gators is  needed  for  keeping  track  of  the  various  important  aspects 
of  the  matter.  In  the  case  of  two  of  my  lines  the  number  of  micro- 
nuclei  was  determined;  D  (larger)  had  one;  c  (smaller)  had  two. 

I  may  be  permitted  to  add  to  the  precise  data  thus  far  given  a 
personal  impression  or  surmise.  Though,  as  I  have  shown,  inter- 
mediate lines  occur,  I  believe  it  will  be  found  that  most  Paramecia 
can  be  placed  in  one  of  the  two  groups  that  we  have  called  "  cau- 
datum  "  and  "  aurelia"  In  other  words,  if  my  impression  is  correct, 
most  lines  will  have  a  mean  length  either  below  145  microns  or  above 
170  microns;  rarely  will  lines  be  found  whose  mean  falls  between 
these  values.  Such  at  least  has  been  my  experience  in  a  large 
amount  of  work.  Furthermore,  I  am  inclined  to  believe  that  those 
belonging  to  the  smaller  group  (mean  length  below  145  microns) 
will  be  found  to  have  as  a  rule  two  micronuclei ;  those  belonging  to 
the  large  group  but  one  micronucleus.  This  matter  is  worthy  of 
special  examination. 

{e)  Do  the  Diverse  Lines  Differ  in  Other  Respects  Besides 

Dimensions? 

In  the  investigations  above  set  forth  the  dimensions,  and  espe- 
cially length,  were  made  the  basis  of  study,  simply  because  they  were 
the  characters  most  readily  examined.  Most  other  characteristics 
are  not  easily  handled  in  so  minute  and  relatively  undiflferentiated 
an  animal  as  Paramecium,     But  there  is,  of  course,  no  reason  to 


X908.J  JENNINGS— HEREDITY  IN   PROTOZOA.  501 

suppose  that  the  relations  we  have  brought  out  are  limited  to  length 
alone.  Probably  other  differentiated  pure  lines  could  be  distin- 
guished on  the  basis  of  other  characteristics. 

The  only  other  characteristic  on  which  our  data  might  give  results 
is  that  of  form,  as  distinguished  from  size.  Are  some  races  broader, 
some  narrower,  in  proportion  to  the  length  ? 

We  may  first  examine  this  question  with  reference  to  the  two 
main  groups  into  which  most  of  our  lines  fall.  Is  there  any  general 
difference  in  the  proportion  of  breadth  to  length  when  we  compare 
the  larger  races  ("  caudatum  group  ")  with  the  smaller  ones  ("  aurelia 
group  ")  ?  The  experiments  whose  results  are  summarized  in  Table 
XXV.,  page  494,  give  us  data  for  a  number  of  different  lines  of  both 
groups,  cultivated  under  the  same  conditions.  We  may,  therefore, 
determine  the  proportion  of  breadth  to  length  in  these.  The  more 
accurate  way  of  doing  this  would  be  by  means  of  the  formula  given 
on  page  399.  This,  however,  would  involve  much  computation  not 
made  for  other  purposes ;  and  we  may  reach  very  nearly  the  same 
results  by  simply  dividing  the  mean  breadth  by  the  mean  length.  If 
the  differences  between  the  different  races  are  not  sufficient  to  show 
clearly  under  this  treatment,  they  are  doubtful  and  inconsequential. 
The  following  table  gives  the  ratio  of  mean  breadth  to  mean  length 
in  the  different  lines  represented  in  Table  XXV.;  the  lines  are 
arranged  according  to  relative  size,  so  as  to  exhibit  any  differences 
between  the  large  and  small  groups. 

The  table  shows  that  the  ratio  of  breadth  to  length  is  almost 
uniformly  greater  in  the  small  or  aurelia  group  than  in  the  larger. 
The  lowest  ratios  of  the  aurelia  group  are,  indeed,  a  little  below  the 
highest  of  the  caudatum  group,  but  the  difference  between  the  groups 
as  a  whole  is  unmistakable.  The  first  column  of  the  table  is  the  most 
satisfactory  in  this  respect,  since  both  sets  were  killed  at  the  same 
time.  In  the  second  column  the  difference  between  the  ratios  for 
the  two  groups  is  still  more  decided,  but  environmental  differences 
may  play  some  part  in  this  case.  The  average  ratio  for  the  cau- 
datum group  is,  from  the  first  column  27.473  per  cent. ;  from  the 
second  25.679  per  cent.  For  the  aurelia  group  the  averages  are: 
first  column  30.441  per  cent;  second  column  31.319  per  cent.    The 


602 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[April  24, 


Table  XXVI. 

Ratio  of  Mean  Breadth  to  Mean  Length  in  the  Lines  and  Races  of  Table 
XXV.,  page  494,  Cultivated  under  Identical  Conditions. 


X.  Caudatum  Group. 

Febniary  a6. 

March  13. 

Percent. 

Per  Cent. 

L2 

29482 

27.185 

Gi 

26.018 

24.744 

Ai 

26.782 

25.810 

Fi 

26.342 

27.019 

A2 

27491 

23.680 

F2 

27.921 

25.065 

D 

28.275 

26.252 

a.  Aurelia  Group. 

Febniary  a6. 

March  19. 

Percent. 

Percent. 

Ki 

29.473 

I 

31.577 

31.048 

H 

31.967 

32.102 

C2 

31.347 

31.275 

K2 

29.416 

34.016 

g 

28.865 

28.153 

general  average  for  the  caudatum  group  is  26.576  per  cent. ;  for  the 
aurelia  group  30.840  per  cent. 

In  Table  XXIV.,  page  491,  we  have  data  for  certain  other  mem- 
bers of  the  two  groups  when  cultivated  under  similar  conditions. 
If  we  determine  the  ratio  of  mean  breadth  to  mean  length  for  this 
table,  the  results  are  not  so  clear  as  in  the  cases  we  have  just  con- 
sidered.    They  are  given  in  Table  XXVII. 


Table  XXVII. 

Ratio  of  Mean  Breadth  to  Mean  Length  for  the  Races  of  Table  XXIV,, 

page  491. 


I.  Caudatum  Group. 

E 
D 
Average 

a.    Aurelia  Group. 
C 

g 
1 

Average 


February  5. 
Per  Cent. 

31.245 
/  27.615 
1 25.651 

28.170 

26482 
/  29.568 

I  30.853 
/  29.303 
I  29.386 

29.118 


February  15. 
Per  Cent. 

26.158 
/  26.010 

26.964 


26.396 
28.298 

31.852 
28.849 


February  27. 
Per  Cent.  ; 

26458 
26.893 
26.675 


In  this  table  the  averages  for  the  aurelia  group  are  again  higher 
throughout  than  for  the  caudatum  group.  But  the  highest  ratio  is 
given  by  one  of  the  caudatum  group,  and  the  line  c  of  the  aurelia 


,9o8.J  JENNINGS— HEREDITY    IN   PROTOZOA.  503 

group  gives  in  both  cases  a  low  ratio.  But  taking  the  averages,  in 
connection  with  those  of  Table  XXVL,  it  is  clear  that  the  smaller 
races  are  as  a  rule  slightly  broader  in  proportion  to  the  length  than 
are  the  larger  races. 

Turning  now  to  the  question  whether  there  are  differences  in  the 
proportion  of  breadth  to  length  in  different  races  of  the  same  group, 
we  have  full  data  only  for  the  lines  g  and  i^  as  given  in  Table  XXIII., 
page  488.  Beginning  with  the  data  for  November  23  (since  before 
that  date  the  number  of  individuals  is  small),  we  can  make  determi- 
nations for  seven  different  dates  of  the  ratio  of  mean  breadth  to 
mean  length,  the  two  sets  being  on  each  date  as  nearly  as  possible 
under  identical  conditions. 

Table  XXVIII. 
Ratio  of  Mean  Breadth  to  Mean  Length  for  g  and  i  (Table  XXIII.). 

November      November     December      December      December       January        February 

83.     Per         96.     Per         7.     Per         x6.     Per         30.    Per         a.     Per  5.    Per 

Cent.  Cent.  Cent.  Cent.  Cent.  Cent.  Cent. 

g        27.011        28.238        34091        30.370        27.797        27.686        30.853 
i        34.291        28.325        34.655        30.159        25.430        24.747        29.386 

Thus,  in  the  first  three  determinations  the  ratio  was  greatest  in 
the  line  i ;  in  the  last  four  it  was  greatest  in  the  line  g.  Evidently 
there  is  no  constant  difference  in  proportions  between  these  two  lines. 

For  other  lines  our  data  are  not  sufficient  to  test  this  matter. 
Our  only  positive  result  on  this  point  then  is  that  the  smaller  races 
are  as  a  rule  proportionately  broader  than  the  larger  ones. 

2.   Results  of  Selection  within  Pure  Lines. 

We  have  seen  that  an  ordinary  "  wild  **  culture  of  Paramecium 
contains  many  lines  or  races,  which  are  differentiated  in  size.  By 
selection  it  is  possible  to  isolate  these  diverse  lines;  so  that  in  this 
way  we  can  obtain  cultures  in  which  the  mean  size  is  large  or  small, 
or  intermediate,  as  we  prefer.  In  this  case  selection,  of  course,  acts 
by  isolating  lines  that  already  exist,  and  allowing  them  to  propagate 
unmixed. 

How  do  these  diverse  lines  arise  ?  Can  we  obtain  them  by  selec- 
tion within  the  limits  of  a  single  line?  If  from  among  the  progeny 
of  a  single  individual  we  select  the  larger  and  the  smaller  specimens, 


604  JENNINGS— HEREDITY  IN   PROTOZOA.  lAprilM. 

will  we  obtain  two  diverse  lines,  one  showing  a  greater  mean  size 
than  the  other  ? 

As  we  have  already  seen,  our  first  attempts  to  do  this  failed. 
But  these  first  experiments  were  made  before  our  study  of  growth  and 
environmental  effects,  so  that  the  basis  of  selection  was  wrong.  The 
smaller  specimens  selected  were  as  a  rule  the  younger  ones;  they 
grew  to  full  size,  then,  of  course,  produced  progeny  of  the  same  size 
as  other  adults. 

After  the  thorough  study  of  growth,  it  appeared  possible  that  a 
more  adequate  method  of  selection  might  be  found.  The  propor- 
tions of  the  young  differ  from  those  of  the  adult  (as  our  account  has 
shown),  so  that  after  long  practice  one  comes  to  recognize  the  young 
specimens  with  some  accuracy.  It  appeared  worth  while,  therefore, 
to  attempt  to  select  larger  and  smaller  adults  for  further  propagation. 

(a)  Differences  Due  to  Environmental  Action  Not  Inherited. 

It  is,  of  course,  easy  to  obtain  within  a  pure  line  adults  ot  differ- 
ent size,  by  subjecting  them  to  different  environments.  An  analysis 
of  our  section  on  the  effects  of  the  environment  shows  that  as  a  rule 
these  are  not  inherited.  Thus,  if  we  examine  Table  XVIII.  (page 
460),  we  find  that  the  same  set  that  gave  on  July  17  a  mean  length 
of  184.100  microns  (row  7)  gave  one  week  later,  under  different 
conditions,  a  mean  of  146.108  microns ;  one  day  later  163.932  microns ; 
one  week  later  174.400  microns;  two  days  later  191.360  microns. 
The  breadth  changed  even  more,  and  the  extremes  of  size  in  a  given 
culture  showed  corresponding  changes.  There  was  no  difficulty  in 
changing  the  dimensions  back  and  forth  in  the  most  varied  ways. 
The  entire  Table  XVIII.  is  an  illustration  of  the  general  lack  of 
continued  inheritance  of  environmental  effects. 

Many  experiments  directed  precisely  on  this  point  gave  the  same 
results.     When,  for  example,  the  small  specimens  of  row  8  (Table 
XVIII.)  were  cultivated  under  the  same  conditions  as  large  speci-. 
mens  from  row  9,  the  resulting  cultures  were  soon  indistinguishable. 

Thus,  it  is  clear  that  such  environmental  action  as  is  summarized 
in  Table  XVIII.  is  not  as  a  rule  inherited.  But  I  wish  to  point  out 
and  emphasize  certain  facts  regarding  the  experiments  on  the  action 
of  the  environment.     ( i )  In  all  the  experiments  thus  far  tried,  the 


X908.1  JENNINGS— HEREDITY   IN   PROTOZOA.  505 

differential  action  of  the  diverse  environments  lasted  but  a  short 
time.  (2)  The  experiments  were  directed  toward  determining 
whether  the  differences  produced  were  permanently  inherited.  Crit- 
ical investigations  have  not  yet  been  made  to  determine  whether  the 
environmental  effects  may  not  persist  for  one  or  a  few  generations 
after  transference  to  the  new  fluid;  nor  whether  long  continued 
action  of  a  certain  environment  may  not  produce  more  lasting  results 
than  brief  action. 

To  these  points  I  hope  to  devote  special  and  extended  investiga- 
tions. The  purpose  in  the  present  paper  is  to  show  on  this  matter 
the  main  general  result;  this  unquestionably  is  that  environmental 
action  is  not  as  a  rule  inherited  in  any  lasting  way. 

(b)  Selection  from  Among  Differing  Individuals  in  the  Same 

Environment. 

Besides  the  differences  among  individuals  under  different  envir- 
onments, we  likewise  find  differences  among  individuals  of  the  same 
pure  line  in  the  same  culture,  as  a  glance  at  the  tables  of  the  appendix 
will  show.  What  will  be  the  effect  of  selecting  for  breeding  larger 
and  smaller  specimens  from  such  a  culture,  avoiding,  so  far  as  pos- 
sible, different  stages  of  growth  ? 

In  order  to  make  the  selections  properly,  certain  things  must  be 
considered.  ( i )  It  is  well  to  bring  the  culture  into  as  stable  a  con- 
dition as  possible — a  condition  where  there  is  little  or  no  multipli- 
cation— in  order  that  we  may  not  be  confused  by  different  stages  in 
growth.  (2)  It  must  be  remembered  that,  so  long  as  conjugation 
does  not  occur,  the  same  results  that  selection  would  produce  are 
brought  about  in  the  ordinary  course  of  events,  save  that  the  large 
and  small  specimens  remain  mixed.  That  is,  if  there  is  congenital 
variation,  producing  large  and  small  individuals,  this  must  occur  in 
the  same  way  whether  the  different  sizes  are  isolated  or  not.  The 
progeny  of  every  individual  forms  a  "  pure  line,"  quite  unmixed  with 
any  other,  so  long  as  no  conjugation  occurs.  If,  then,  by  variation 
a  large  individual  a  and  a  small  one  b  are  produced,  and  these  differ- 
ences are  inherited,  then  later  we  shall  find  a  mixture  of  two  strains 
instead  of  a  single  strain.    We  should  then  expect  the  progeny  of  a 

PROG.  AMER.  PHIL.  SOC.  XL VII.   I90  GG,  PRINTED  JANUARY  12,  I909. 


506  JENNINGS— HEREDITY   IN   PROTOZOA.  [ApriU4. 

single  individual  to  show  more  and  more  variation  as  the  strain 
became  older;  it  would  break  into  several  or  many  strains,  which 
would,  however,  remain  intermingled. 

Therefore,  the  best  method  of  procedure  will  be  to  take  an  old 
strain,  which,  derived  from  a  single  individual,  has  for  a  long  time 
been  multiplying  freely  without  conjugation.  From  this  the  largest 
and  the  smallest  individuals  should  be  separated  and  allowed  to 
propagate  under  identical  conditions.  If  hereditary  variations  in 
size  have  occurred,  we  should  in  this  way  reach  the  same  result  as 
by  actual  selection  and  isolation  through  many  generations.  Physio- 
logical isolation  has  been  as  complete  as  would  be  experimental 
isolation. 

A  race  fulfilling  these  conditions  we  have  in  the  pure  line  derived 
from  the  individual  D,  on  which  most  of  the  work  described  in  the 
first  parts  of  this  paper  was  done.  On  January  19,  1908,  large 
cultures  of  D  had  been  multiplying  without  conjugation  since  April 
12, 1907,  a  period  of  about  nine  months.  During  this  time  about  250 
generations  must  have  been  produced;  these  had  remained  physio- 
logically isolated.  The  superfluous  individuals  had  been  removed  by 
periodic  "  catastrophic  "  destruction ;  the  greater  part  of  the  culture 
was  thrown  out,  and  a  remnant  saved,  without  selection,  for  a  new 
culture. 

On  January  19,  1908,  I  took  from  the  large  stock  culture  of  D 
(i)  the  ten  largest  individuals  that  I  could  find;  (2)  the  ten  smallest 
individuals  I  could  find.  They  were  separated  in  two  watch-glasses 
and  kept  under  identical  conditions.  The  difference  between  the  two 
sets  was  very  marked ;  the  smaller  lot  were  certainly  not  more  than 
two-thirds  the  length  of  the  larger,  and  they  were  very  slender,  while 
the  large  ones  were  both  long  and  broad.  It  was  clear  that  both  sets 
were  adults. 

It  was  found  that  the  smaller  lot  multiplied  much  less  rapidly 
than  the  large  lot,  and  some  of  the  small  ones  died.  By  January 
30  there  were  but  twenty  of  the  small  lot,  while  a  very  large  number 
had  arisen  from  the  large  lot.  On  this  date  the  culture  fluid  was 
changed  and  but  fifty  of  the  larger  lot  retained.  The  small  lot  con- 
tinued to  multiply  very  slowly.     It  is  clear  that  the  small  specimens 


x9o8.]  JENNINGS— HEREDITY  IN   PROTOZOA.  507 

are  weak,  sickly  ones,  and  the  physiological  difference  persists  at  least 
for  some  generations  (a  matter  for  further  study). 

On  February  5  about  half  of  each  lot  was  killed  and  measured. 
This  gave  57  specimens  from  the  larger  lot,  19  from  the  smaller. 
The  mean  dimensions  were,  for  the  larger  lot,  169.754  X  46.877 
microns ;  for  the  smaller  lot,  169.895  X  43-579  microns. 

Thus  the  two  were  practically  identical ;  one  could  not  expect  a 
closer  approximation  in  two  identical  lots  kept  separate  for  seventeen 
days.  The  slight  difference  in  breadth  is  only  what  we  might  expect 
when  we  consider  the  extreme  sensitiveness  of  that  dimension  to 
faint  environmental  differences.  The  most  striking  differences  that 
we  can  find  as  a  result  of  physiological  isolation  for  250  generations 
have  equalized  themselves  in  a  short  time,  when  we  got  both  sets  to 
multiplying  freely  under  the  same  conditions. 

It  seems  hardly  worth  while  to  continue  this  series,  since  the  two 
sets  have  now  become  equalized.  However,  they  were  continued 
for  some  time,  and  samples  of  100  each  were  measured  on  February 
15  and  February  27.  In  these  two  measurements  we  find  certain 
differences  between  the  two  sets,  but  these  are  in  opposite  directions 
in  the  two  cases.    The  means  are  as  follows : 

February  15.  February  37. 

Large  D  180.240  X  46.880  i75-36o  X  47.100 

Small   D  173.240  X 49760  193.680 X  52.320 

Evidently  slight  environmental  differences  between  the  two  cultures 
had  crept  in.  It  is  clear  that  the  two  sets  show  no  constant  differ- 
ences, such,  for  example,  as  we  find  between  the  two  lines,  g  and  i, 
in  Table  XXIII.,  page  488. 

Another  set  of  experiments  dealt  with  the  two  differentiated 
lines,  g  and  i.  The  line  g  consists  of  individuals  that  are  constantly 
larger  than  those  of  the  line  i,  when  the  two  are  under  the  same 
conditions  (see  Table  XXIII.,  p.  488).  The  experiments  consisted 
in  an  attempt  to  separate  these  races  still  farther  by  propagating 
continually  from  the  largest  specimens  of  g  and  from  the  smallest 
specimens  of  t.  Thus,  if  selection  is  effective,  g  must  become  larger, 
i  smaller.  The  length  was  the  dimension  mainly  attended  to  in  these 
sdections. 

On  November  23, 1907,  the  mean  size  for  g  was  129.333  X  34-933 


508  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a4, 

microns ;  for  i  it  was  88.268  X  30.268.  On  this  date  I  placed  in 
separate  watch-glasses  the  ten  largest  specimens  of  g  and  the  ten 
smallest  specimens  of  i,  keeping  them  under  the  same  conditions. 

On  November  29  I  again  selected  from  the  progeny  of  these  the 
ten  largest  g  and  the  ten  smallest  i,  destroying  the  others. 

On  December  7  the  same  selection  was  repeated ;  the  remainder 
of  each  lot  was  killed  and  measured.    The  mean  measurements  were 

g,  120.590  X  41-115  microns. 
1,   98.709  X  34.208  microns. 

Thus,  in  spite  of  the  fact  that  for  at  least  fourteen  generations 
we  have  selected  for  propagation  the  largest  of  g  and  the  smallest  of 
i,  g  has  become  smaller  and  i  has  become  larger!  The  results  of 
selection,  if  there  are  any,  quite  disappear  in  comparison  with  the 
effects  of  slight  environmental  differences. 

In  spite  of  this  discouraging  result,  the  experiment  was  con- 
tinued. On  December  16  I  selected  the  five  largest  g  and  the  five 
smallest  %  and  again  measured  the  rest  of  each.    The  results  were 

g,  127.059  X  38.588  microns. 
i,  98.608  X  29.739  microns. 

Thus,  f  retains  the  same  length,  while  g  has  increased,  but  has  not 
regained  the  length  it  had  at  the  beginning  of  the  experiment. 

On  December  25  the  five  largest  g  and  the  five  smallest  i  were 
again  selected  for  propagation. 

On  December  30,  thirty-seven  days  after  the  beginning  of  the 
experiment,  I  again  measured  all  but  the  five  largest  of  gr  and  the 
five  smallest  of  i.    The  results  are 

g,  1 12.600  X  30.300  microns. 
h   86.756  X  22.062  microns. 

Thus,  f  has  decreased  as  compared  with  its  original  length,  while 
g,  which  was  selected  for  increase  of  size,  has  decreased  a  great 
deal  more!  The  decrease  in  length  of  i  is  less  than  two  microns; 
the  decrease  in  g  is  more  than  sixteen  microns!  And  this  is  the 
result  of  five  selections,  taking  for  g  the  largest,  for  i  the  smallest, 
specimens  produced  in  the  course  of  at  least  thirty  generations  !* 

•The  number  of  specimens  on  which  the  measurements  are  based  will 
be  found  in  Table  XXIIL,  page  488,  which  includes,  for  another  purpose,  the 
measurements  from  these  experiments. 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  509 

Evidently,  selection  is  having  no  effect  that  can  be  detected. 
The  fluctuations  in  the  two  sets  are  precisely  what  would  be  expected 
from  unavoidable  changes  in  conditions  of  nutrition ;  they  show  no 
relation  to  selection. 

Later  another  experiment  in  selection  was  tried  with  these  same 
races,  g  and  i.  On  January  19  I  selected  from  a  large  culture  that 
had  been  multi^ying  freely  for  a  month  (i)  the  ten  largest  speci- 
mens of  g  that  I  could  find;  (2)  the  ten  smallest  specimens  of  g\ 
(3)  the  ten  largest  specimens  off;  (4)  the  ten  smallest  specimens  of  1. 

These  were  allowed  to  multiply  under  identical  conditions  till 
February  5.  Then  a  sample  of  fifty  of  each  was  measured.  The 
results  are  as  follows : 

Large  gr^  114.720  X  33.920  microns.' 
Small  ^,  116.912  X  36.070  microns. 
Large  i,  92.000  X  26.960  microns. 
Small  i,  93.583  X  27.500  microns. 

The  difference  between  the  two  sets  of  each  is  slight  and  without 
significance,  but  such  as  is  found  is  in  favor  of  the  progeny  of  the 
smaller  specimens  in  each  case. 

Evidently,  we  are  not  making  a  start  with  any  effect  of  selection, 
and  it  is  useless  to  continue  the  experiment. 

Many  other  attempts  were  made  to  break  a  pure  line  by  selection 
into  several  strains ;  on  this  point  an  immense  amount  of  work  was 
directed.  But  in  most  cases  the  difference  between  the  two  sets 
became  equalized  almost  at  once,  so  that  the  experiments  were  not 
carried  farther.  As  soon  as  two  unequal  sets  become  quite  equalized, 
there  is  little  opportunity  for  further  selection.  In  the  experiments 
described  above,  though  their  futility  seemed  evident  from  the  first 
results,  the  work  was  continued  for  many  generations,  in  order  that 
failure  might  not  be  due  to  lack  of  perseverance. 

One  other  set  of  experiments  deserves  to  be  described,  because 
in  these  the  basis  for  selection  was  changed.  Among  the  progeny 
of  a  certain  individual  Nf2  conjugation  occurred.  The  conjugants 
varied  in  size.    This  offered  an  opportunity  to  make  a  selection 

•These  measurements  are  found,  for  another  purpose,  in  Table  XXIV., 
page  491. 


510  JENNINGS— HEREDITY   IN    PROTOZOA.  [April  24, 

based  on  specimens  that  were  evidently  adults;  possible  confusion 
due  to  growth  differences  could  be  avoided. 

On  March  31  I  killed  and  measured  all  but  the  largest  and 
smallest  pairs  of  conjugants;  the  length  was  found  to  vary  from  124 
to  148  microns.  The  smallest  and  largest  pairs  were  reserved  for 
propagating;  the  former,  of  course,  measured  not  more  than  124 
microns,  the  latter  not  less  than  148  microns.  Thes^were  allowed  to 
multiply  separately,  but  under  the  same  conditions,  till  April  10. 

On  April  10  I  measured  a  random  sample  of  100  specimens  of 
the  progeny  of  each  of  these  pairs.    The  results  are  as  follows : 

Larger  pair,    151.920  X  43.840  microns. 
Smaller  pair,  158.760  X  38.120  microns. 

Thus,  the  difference  in  size,  whatever  its  cause,  does  not  corre- 
spond to  the  difference  between  the  ancestors ;  selection  for  size  has 
had  no  evident  effect. 

Another  experimefit  on  the  progeny  of  Nf2  consisted  in  com- 
paring the  descendants  of  a  single  small  conjugant  with  those  of 
several  large  non-con jugants.  Details  of  this  and  similar  experi- 
ments will  be  reserved  for  our  paper  on  the  relation  of  conjugation 
to  variation  and  heredity.  But  since  it  has  a  certain  bearing  on  our 
present  problem,  the  results  may  be  given  here. 

At  the  same  time  with  the  cultures  last  described  (on  March  31), 
I  isolated  ten  of  the  largest  non-conjugant  progeny  of  the  same 
individual  Nf2.  A  sample  of  thirty-four  of  these  had  given  a  mean 
length  of  147.412  microns,  so  that  this  may  be  taken  as  the  mean 
length  of  these  ten  specimens.  With  the  progeny  of  these  was  com- 
pared the  progeny  of  the  smaller  pair  mentioned  in  the  preceding 
experiment.  As  we  have  seen,  this  pair  measured  not  more  than 
124  microns  in  length.  The  greatest  pains  were  taken  to  cultivate 
the  two  sets  under  identical  conditions.  On  April  20  I  killed  a 
sample  of  108  of  each.  The  mean  measurements  were  as  follows : 
Progeny  of  small  pair  (124  microns)  —  160.852X42.036  microns. 
Progeny  of  ten  large     (147  microns)  —  156.482  X  43.815  microns. 

Thus,  again,  there  is  no  correspondence  between  the  differences 
in  size  of  the  parents  and  those  of  the  progeny.  The  determining 
factor  in  the  size  is  the  fact  that  both  sets  belong  to  the  same  pure 


X908.J  JENNINGS— HEREDITY  IN   PROTOZOA.  511 

line ;  the  variation  of  the  parents  from  the  type  of  the  pure  line  has 
no  effect.  The  difference  in  the  figures  above  is  either  purely  statis- 
tical in  character  or  means  a  faint  variation  in  the  culture  fluid. 

(c)  Summary  on  Selection  within  Pure  Lines. 

Thus,  we  come  uniformly  to  the  result  in  all  our  experiments, 
that  selection  has  no  effect  within  a  pure  line ;  the  size  is  determined 
by  the  line  to  which  the  animals  belong,  and  individual  variations 
among  the  parents  have  no  effect  on  the  progeny. 

But  for  our  results  with  different  lines,  it  might  be  maintained 
that  the  reason  why  we  get  no  constant  differences  between  the 
progeny  of  different  individuals  of  the  same  line  is  because  the 
effects  of  environment  are  so  much  greater  than  the  effects  of  selec- 
tion that  the  latter  are  covered  up  and  obscured.  But  as  soon  as 
we  are  dealing  with  lines  that  are  really  different  (though  by  but  a 
small  amount)  we  have  no  such  difficulty;  the  different  lines  retain 
their  relative  sizes  in  spite  of  environmental  action.  This  is  clearly 
shown  in  Tables  XXIII.  and  XXV.,  pages  488  and  494. 

The  significance  of  these  results  will  be  dealt  with  in  the  next 
section. 

VI.   SUMMARY  AND  DISCUSSION. 

I.   Resume  of  the  Investigatioi^s. 

The  present  paper  is  an  experimental  study  of  the  factors  involved 
in  variation  and  inheritance  of  size  in  the  infusorian  Paramecium, 
in  the  period  when  reproduction  is  taking  place  by  fission,  without 
conjugation. 

I.  The  first  question  proposed  is  whether  the  differences  in  size 
among  different  individuals  of  a  culture  are  inherited.  The  pre- 
liminary study  showed  that  in  a  typical  culture  there  were  two 
permanently  differentiated  groups  of  large  and  small  individuals, 
respectively,  corresponding  to  what  had  been  described  as  the  two 
species,  Paramecium  caudatum  and  Paramecium  aurelia.  But  when 
a  culture  was  produced  from  a  single  individual  of  either  of  these 
groups,  forming  thus  a  '^j>ure  line,"  it  was  found  that  though  the 
different  individuals  of  the  single  pure  line  differed  much  in  size. 


612  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  a4, 

these  differences  were  not  inherited.    Large  and  small  specimens  of 
a  single  pure  line  produced  progeny  of  the  same  mean  size. 

2.  The  next  question  then  was:  What  are  the  causes  and  the 
nature  of  the  variations  in  size  among  the  different  individuals 
of  a  culture  of  Paramecium?  Even  in  a  pure  line  the  indi- 
viduals differ  greatly.  The  "  polygon  of  variation  "  of  a  given  cul- 
ture was  looked  upon  as  a  mass  of  problems  for  analysis.  What 
determines  the  position  which  any  given  individual  holds  in  such  a 
polygon,  or  in  a  correlation  table?  And  why  dp  different  lots  of 
Paramecia  differ  in  mean  dimensions ;  in  the  amount  of  variability ; 
in  proportions,  and  in  the  correlation  between  length  and  breadth  ? 

The  analysis  of  the  factors  in  variation  led  to  a  detailed  study 
of  (i)  growth,  (2)  the  effect  of  the  environment;  (3)  inherited 
differences  in  size.  To  these  three  matters  the  three  main  divisions 
of  the  paper  are  devoted.  To  one  or  the  other  of  these  three  cate- 
gories most  of  the  variations  in  size  were  found  to  belong.  A 
fourth  category,  consisting  of  variations  connected  with  conjugation, 
is  reserved  for  consideration  in  a  later  paper. 

3.  A  large  share  of  the  differences  in  size  to  be  observed  in  a 
given  culture  are  differences  in  growth.  In  study  of  variation  in 
protozoa  it  is  as  necessary  to  take  growth  into  consideration  as  it  is 
in  the  study  of  higher  animals ;  the  part  played  by  it  is  fully  as  great 
in  the  protozoa  as  elsewhere.  The  paper  gives  a  detailed  study  of 
growth,  based  on  the  measurements  of  1,500  specimens  of  various 
known  ages,  in  comparison  with  large  numbers  of  "random  sam- 
ples." In  this  way  a  curve  of  growth  was  plotted  (Diagram  5,  page 
449)  ;  this  curve  resembles  essentially  the  curves  of  growth  of  higher 
animals,  as  the  rat,  or  man.  In  different  parts  of  this  curve  of 
growth  individuals  show  different  lengths,  different  breadths,  and,  of 
course,  different  proportions  of  breadth  to  length.  A  flourishing 
culture  contains  individuals  in  all  stages  of  growth;  so  that  this 
affects  largely  the  mean  dimensions,  the  observed  variations,  and 
the  correlations  between  length  and  breadth.  The  precise  effects  of 
growth  on  each  of  these  matters  are  dealt  with  in  detail  in  the 
paper;  they  will  be  summarized  in  later  paragraphs.  A  summarized 
account  of  growth  and  its  effects  is  found  in  the  body  of  the  paper, 
pages  447  to  458 ;  the  constants  for  dimensions  and  variation  in  dif- 


,9o8]  JENNINGS— HEREDITY  IN   PROTOZOA.  513 

ferent  stages  of  growth  are  brought  together  in  Table  X.,  page  428. 

4.  Environmental  conditions  were  found  to  play  a  very  large 
part  in  determining  dimensions,  variations  and  correlation  in  Para- 
mecium, Conditions  of  nutrition  were  found  to  be  particularly 
effective.  By  changes  in  nutrition  the  mean  length  of  a  given  culture 
could  be  changed  in  a  week  from  146  microns  to  191  microns ;  the 
breadth  from  31  to  54  microns;  in  twenty- four  hours  the  coefficient 
of  variability  for  length  was  thus  changed  from  7.003  to  12.767,  for 
breadth  from  12.473  to  28.879;  ^he  coefficient  of  correlation  from 
.3906  to  .8463.  Changes  of  the  most  varied  sort  could  be  produced 
and  reversed  with  the  greatest  ease  in  short  periods ;  many  examples 
of  this  are  summarized  in  Table  XVIII.,  page  460.  Within  a  given 
culture  at  a  given  time  many  of  the  differences  between  individuals 
are  due  to  slight  environmental  differences  in  different  regions. 
The  breadth  is  more  sensitive  to  environmental  changes  than  the 
length;  to  such  an  extent  is  this  true  that  it  is  difficult  to  use  the 
breadth  dimensions  for  accurate  study  of  any  other  factors.  A  sum- 
mary on  the  effects  of  the  environment  on  dimensions,  proportions, 
variation  and  correlation  is  found  on  pages  476  to  484. 

5.  After  the  study  of  growth  and  environmental  action,  an  inves- 
tigation was  made  of  the  internal  factors  in  dimensions  and  variation ; 
of  the  inheritance  of  size.  Are  all  the  observed  differences  between 
the  individuals  of  a  culture  mere  matters  of  growth  and  environ- 
ment? Or  may  we  find  different  races  or  lines  that  retain  their 
relative  sizes  even  in  the  same  stage  of  growth  and  in  the  same 
environment? 

A  thorough  experimental  study  showed  that  a  given  "  wild " 
culture  usually  contains  many  different  lines  or  races,  which  maintain 
their  relative  sizes  throughout  all  sorts  of  changing  conditions. 
Eight  of  these  differing  pure  lines  were  isolated  and  propagated; 
these  varied  in  mean  length  from  a  little  less  than  100  to  a  little 
more  than  200  microns  (see  Tables  XXIII.  and  XXV.).  Other 
lines  could  unquestionably  be  distinguished  by  sufficiently  accurate 
experimentation. 

These  different  lines  fall  usually  into  two  main  groups,  one  group 
having  a  mean  length  greater  than  170  microns,  the  other  having  a 
mean  length  below  140  microns.    These  two  groups  correspond  to 


514  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

the  distinction  that  has  been  made  between  two  species,  the  larger 
ones  representing  the  supposed  species  caudatum,  the  smaller  ones 
aurelia.  But  a  line  or  race  was  found  with  mean  length  lying  mid- 
way between  these  groups,  at  about  150  to  160  microns. 

The  smaller  or  aurelia  lines  were  found  to  be,  under  the  same 
conditions,  as  a  rule  a  little  broader  in  proportion  to  the  length  than 
the  larger  or  caudatum  lines.  But  the  difference  is  slight  and  the 
two  sets  overlap  extensively  in  this  matter;  slight  differences  in 
environment  quite  obscure  the  difference  in  proportions. 

The  differences  among  the  different  lines  were  found  not  to  be 
due  to  different  periods  of  the  life  cycle.  By  beginning  with  con- 
jugating pairs  of  different  sizes,  distinct  pure  lines  were  as  readily 
isolated  as  by  beginning  anywhere  else  in  the  cycle. 

6.  After  becoming  thoroughly  familiar  with  differences  due  to 
growth,  to  environment,  and  to  divergent  ancestry,  a  further  attempt 
was  made  to  change  by  selection  the  characteristics  of  pure  lines,  or 
to  break  such  lines  into  strains  of  differing  size.  In  spite  of  much 
work  directed  on  this  point,  it  was  found  that  selection  within  a  pure 
line  was  quite  without  effect.  Large  individuals  of  the  line  produce 
progeny  of  the  same  mean  size  as  do  the  small  individuals.  To  this 
matter  we  return  in  later  paragraphs. 

2.   Determining  Factors  for  Dimensions,  Variations  and 

Correlations. 

Based  on  the  analysis  of  the  factors  in  variation  above  set  forth, 
a  summary  can  be  given  of  the  various  determining  causes  of  the 
different  dimensions,  the  proportions,  the  amount  of  variation  and 
the  correlations  observed  in  samples  of  different  cultures  of  Para- 
mecium. We  may  take  as  an  example  such  a  sample  as  is  shown  in 
Table  LXI.  (appendix)  from  a  "wild"  culture. 

I.  The  various  different  lengths  depend  upon  the  following  factors : 
(a)  The  collection  embraces  a  number  of  different  races  or  lines, 
having  different  lengths  even  when  all  conditions  are  the  same.  We 
have  seen  that  different  lengths  varying  from  less  than  100  to  more 
than  200  microns  may  be  included  as  a  result  of  this  fact.  The 
mean  length  may  not  represent  any  of  these  races  (this  is  the  case  in 
Table  I.). 


i9o8.]  JENNINGS—HEREDITY   IN   PROTOZOA.  615 

(b)  The  collection  includes  various  growth  stages  of  each  of  the 
lines  represented.  The  youngest  stages  of  each  line  are  little  more 
than  half  the  lengths  of  the  adults;  all  intermediate  stages  may  be 
present,  and  the  adults  themselves  shorten  again  as  they  approach 
fission.  A  very  wide  range  of  variation  in  length  may  be  brought 
about  by  these  growth  stages,  all  within  the  limits  of  a  single  pure 
line  or  race.  Of  course  when  many  diflferent  lines  are  present,  an 
immense  number  of  combinations  are  thus  produced. 

(c)  The  collection  includes  individuals  of  the  various  races  that 
have  lived  under  slight  or  considerable  differences  in  environment, 
particularly  in  the  matter  of  nutrition.  Those  that  have  been  able 
to  get  more  food  will  be  much  larger  and  will  multiply  more  fre- 
quently (thus  giving  more  young)  than  those  that  get  less.  Even 
slight  environmental  differences  make  decided  differences  in  dimen- 
sions. While  the  environment  shows  its  effects  most  strongly  on 
comparison  of  different  cultures,  even  within  the  same  culture,  and 
when  all  the  individuals  are  of  one  race  and  of  approximately  the 
same  age,  there  are  marked  diflferences  due  to  this  cause.  This  is 
shown,  for  example,  in  Table  XLI.  (appendix)  ;  here  variations  in 
length  from  140  to  200  microns  must  be  considered  environmental 
effects.  A  few  drops  of  water  form  a  varied  microcosm  to  the 
infusoria.  When  diverse  pure  lines,  diverse  growth  stages,  and 
diverse  environmental  conditions  are  found  in  a  culture  (as  is  usually 
the  case),  of  course,  the  number  of  different  sizes  and  forms  due  to 
the  varied  combinations  of  all  these  factors  are  very  great  The 
same  sizes  may,  of  course,  be  produced  in  different  ways ;  two  diverse 
lines  in  different  stages  of  growth  or  in  different  environments,  or 
in  some  combination  of  the  two,  may  produce  forms  outwardly  iden- 
tical. The  actual  variety,  as  defined  by  the  physiological  conditions, 
is  therefore  much  greater  than  the  measurements  show,  for  the  latter 
throw  together  heterogeneous  combinations. 

Combinations  of  all  the  three  factors  inducing  diversity  might 
give  us  in  a  single  collection  individuals  varying  in  length  from  50 
microns  to  332  microns.  While  these  are  the  extremes  given  by  our 
data,  presumably  the  actual  extremes  would  be  still  more  divergent. 

(d)  In  different  collections  the  observed  mean  lengths  depend 
upon  the  three  different  sets  of  factors  just  mentioned.    The  inclu- 


513  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  «4. 

sion  of  different  lines  or  races,  even  if  conditions  of  growth  and 
environment  are  essentially  the  same,  may  give  us,  as  we  have  seen, 
mean  lengths  of  somewhat  less  than  lOO,  or  somewhat  more  than 
200  microns,  or  any  intermediate  length.  Different  stages  in  growth 
ma>  give  us,  in  the  same  line  and  in  the  sahie  environment,  means 
differing  to  such  an  extent  that  one  is  nearly  twice  the  other,  or  any 
intermediate  condition.  The  absolute  extreme  values  will,  of  course, 
depend  upon  the  race  employed ;  in  the  line  i  the  variation  of  mean 
length  caused  by  growth  might  be  from  about  50  to  about  100 
microns ;  in  D  it  was  from  about  100  to  about  200  microns ;  in  L  it 
would  be  from  about  117  to  234  microns.  Different  enznronmental 
conditions  give  us,  within  the  same  lines,  mean  lengths  differing  to 
such  an  extent  that  the  greater  is  25  to  30  per  cent,  more  than  the 
less  (lines  c  and  D).  In  different  "wild"  cultures  we  shall  have 
different  combinations  of  all  these  factors,  resulting  in  extreme 
diversities  in  different  cases.  Fig.  7  shows  two  extreme  sizes  drawn 
to  the  same  scale  (page  496). 

2.  The  various  different  breadths  depend  upon  the  same  factors 
as  the  different  lengths.  There  are  certain  differences,  however. 
As  compared  with  length,  the  breadth  is  affected  much  less  by 
growth;  about  the  same  (though  a  trifle  less)  by  diversity  of  race; 
and  much  more  by  environmental  differences.  Environmental  dif- 
ferences produced  within  the  races  D  and  c  such  differences  in  mean 
breadth  that  the  greater  was  about  twice  the  less. 

3.  The  observed  variation,  as  measured  by  the  coefficient  of 
variation,  of  course,  depends  upon  the  three  sets  of  factors  enumer- 
ated above  as  affecting  the  length  and  breadth.  If  a  collection 
consisted  of  several  different  lines  or  races,  all  in  the  same  condition 
as  regards  growth  and  environmental  conditions,  this  would,  of 
course,  give  us  a  considerable  coefficient  of  variation.  For  example, 
if  a  collection  consisted  of  ten  individuals  each  of  all  the  different 
lines  represented  in  Table  XXVI.,  page  502,  and  if  all  of  each  set  of 
ten  had  the  mean  dimensions  for  its  line  (thus  excluding  differences 
due  to  growth  and  environment  within  the  lines),  the  coefficient  of 
variation  when  computed  in  the  same  way  as  for  the  actual  collections 
given  in  the  text  is  found  to  be  for  length  19.689;  for  breadth  15.679. 

If  a  collection  consists  of  individuals  all  belonging  to  the  same 


1908.]  JENNINGS— HEREDITY   IN   PROTOZOA.  517 

line  or  race,  and  in  the  same  environment,  then  the  coefficient  of 
variation  depends  largely  upon  the  stages  of  growth  it  contains.  By 
taking  specimens  nearly  in  the  same  stage  of  growth  we  were  able  to 
reduce  the  coefficient  of  variation  in  length  in  some  cases  to  4.521, 
in  breadth  to  6.976,  while  by  taking  collections  including  various 
ages,  under  similar  conditions,  coefficients  were  found  as  high  as 
13.729  for  length  and  13.292  for  breadth  (Table  X.).  The  most 
carefully,  selected  lots  contain  specimens  differing  a  certain  amount 
in  age,  otherwise  the  coefficient  of  variation  could  be  still  further 
reduded  in  this  way.  Specimens  beginning  fission  or  undergoing 
conjugation  include  few  growth  stages,  hence  they  show  a  low  coeffi- 
cient of  variation.  The  coefficient  for  those  beginning  fission  is  less 
than  for  conjugants  (see  page  453). 

The  coefficient  of  variation  for  a  given  line  is  tremendously 
affected  by  environmental  conditions.  Thus,  we  see  this  coefficient 
changed  in  twenty-four  hours,  by  a  change  in  environment,  from 
7.003  to  12.767  for  length;  from  12.473  ^^  28.879  for  breadth. 
Different  environments  give  us  all  sorts  of  values  between  such 
extremes. 

It  is  evident  that  no  particular  coefficient  of  variation  can  be 
considered  characteristic  of  Paramecium,  or  of  any  line  of  Para- 
mecium; certainly  not  unless  the  conditions  as  t©  growth,  envir- 
onment, etc.,  are  very  precisely  defined.  We  have  seen  that  the 
variations  found  among  different  individuals  of  the  same  pure  line 
do  not  show  themselves  to  be  heritable.  This,  along  with  all  the 
rest  of  the  evidence,  indicates  that  if  all  conditions  of  growth  and 
environment  were  made  identical  throughout  a  sample  of  Paramecia 
belonging  to  a  pure  line,  the  coefficient  of  variation  would  be  very 
near  to  zero.  In  other  words,  all  the  variations  that  we  have  been 
able  to  detect  with  certainty  in  a  pure  line  are  due  to  growth  and 
environment.  Presurtiably  other  variations  (congenital  and  heredi- 
tary) must  occur  at  times,  but  they  appear  to  be  so  rare  that  it  is 
difficult  to  detect  them  and  they  would  have  little  effect  on  the 
coefficient  of  variation.  By  properly  varying  the  conditions,  we  may 
get  in  a  pure  line  all  coefficients  of  variation  in  length,  from  a  limit 
near  zero  up  to  20  or  more. 

4.  The  ratio  of  breadth  to  length  (serving  to  partly  define  the 


518  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  .4, 

form  of  the  body),  of  course,  varies  in  dependence  upon  all  the  three 
sets  of  factors  with  which  we  have  dealt — difference  of  race,  growth 
and  environmental  conditions.  The  smaller  races  are  found  to  show, 
under  the  same  conditions,  a  slightly  greater  ratio  of  breadth  to 
length  (see  Table  XXVI.).  Within  the  same  race  different  stages 
of  growth  show  different  ratios ;  in  general,  the  proportion  of  breadth 
to  length  is  greatest  in  the  young,  and  gradually  decreases  with  age ; 
it  increases  again  very  rapidly  in  preparation  for  fission.  Environ- 
mental agents  affect  in  most  marked  and  varied  ways  the  proportion 
of  breadth  to  length ;  this  is  connected  with  the  fact  that  such  agents 
act  more  upon  the  breadth  than  upon  the  length.  A  detailed  sum- 
mary of  the  different  effects  of  the  environment  on  the  proportion 
of  breadth  to  length  is  found  on  pages  478  and  479.  The  most  im- 
portant general  relation  is,  that  increase  of  nutriment  increases  the 
proportional  breadth;  decrease  of  nutriment  produces  the  opposite 
effect.  Any  agent  which  suddenly  increases  the  breadth  likewise,  as 
a  rule,  increases  the  ratio  of  breadth  to  length. 

5.  The  coefficient  of  correlation  between  length  and  breadth  is 
the  measure  of  the  accuracy  with  which  breadth  and  length  vary 
proportionately.  If  the  proportion  of  breadth  to  length  is  the  same 
in  all  individuals  of  a  collection,  then  the  coefficient  of  correlation 
of  that  collectioa  is  i.ooo.^®  Since,  as  we  have  just  seen,  the  pro- 
portion of  breadth  to  length  is  altered  by  many  factors,  it  follows 
that  all  these  factors  modify  the  correlation,  tending  to  reduce  it 
below  1. 000.  The  correlation  is  affected  by  all  the  three  categories 
of  factors  that  affect  the  dimensions  in  essentially  the  following  ways : 

(a)  The  inclusion  of  different  races  in  a  collection,  particularly 
if  some  of  the  smaller  and  some  of  the  larger  races  occur,  makes  the 
correlation  less  than  i.ooo,  because  the  proportion  of  breadth  to 
length  is  greater  in  the  smaller  races.  The  reduction  in  correlation 
produced  by  this  alone  is  very  slight.     If  we  make  a  collection  by 

"  It  is  perhaps  not  necessary  to  point  out  that  the  "coefficient  of  correla- 
tion "  is  descriptive;  it  shows  the  observed  condition  in  a  given  set  of  meas- 
urements. The  cause  of  this  condition  is  a  matter  to  be  determined.  Corre- 
lation  is  often  conceived  physiologically  as  an  underlying  something  that 
binds  two  things  together,  so  that  they  must  change  correspondingly.  The 
descriptive  correlation  of  the  statistician  may  be  the  resultant  of  many 
factors. 


,9o8.]  JENNINGS— HEREDITY   IN   PROTOZOA.  519 

throwing  together  ten  each  of  the  different  lines  of  Table  XXV. 
(page  494),  giving  the  individuals  of  each  line  the  mean  dimensions 
of  its  line  (thus  nearly  excluding  variations  due  to  growth  and 
environment),  then  calculate  the  coefficient  of  correlation  in  the 
same  way  as  for  our  other  collections,  we  find  it  to  have  the  high 
value  of  .9735. 

(&)  The  inclusion  of  different  stages  of  growth  in  a  collection 
reduces  the  correlation  below  i.ooo,  since  different  growth  stages 
have  different  ratios  of  breadth  to  length.  A  detailed  summary  of 
the  effects  of  growth  on  correlation  is  found  on  pages  455  to  457; 
here  we  can  notice  only  the  main  points.  In  the  earliest  stages  of 
growth  the  length  is  increasing  while  the  breadth  is  decreasing; 
hence  if  we  take  a  collection  including  various  stages  within  this 
period,  the  correlation  between  length  and  breadth  becomes  negative ; 
it  may  fall  to  a  value  of  — .3138  (see  Table  X.).  The  inclusion  of 
various  early  stages  in  a  collection  of  adults  decreases  the  positive 
correlation  shown  by  the  adults.  In  later  growth,  length  and  breadth 
increase  together ;  the  inclusion  of  various  stages  at  this  period  has 
little  effect  on  the  correlation;  it  does,  however,  tend  to  reduce  it 
slightly,  since  length  and  breadth  do  not  increase  at  the  same  ratio. 
In  old  specimens,  beginning  fission,  the  length  decreases  while  the 
breadth  increases;  a  collection  including  different  stages  in  this 
process  tends  again  to  give  negative,  correlation,  or  to  reduce  the 
positive  correlation  due  to  other  causes.  In  a  collection  from  the 
same  pure  line,  in  which  all  specimens  are  in  the  same  stage  of 
growth,  the  correlation  between  length  and  breadth  is  high;  this 
would  be  true  no  matter  what  stage  of  growth  is  the  one  represented. 
Random  samples  from  any  culture  usually  contain  many  stages  of 
growth ;  this  lowers  the  correlation  between  length  and  breadth. 

(c)  Environmental  differences,  like  growth,  affect  length  and 
breadth  differently  or  in  different  proportions;  if  individuals  thus 
diversely  affected  are  included  in  a  sample,  this  tends  to  decrease  the 
correlation  between  length  and  breadth.  A  detailed  analysis  of  the 
many  and  important  effects  of  environmental  action  on  the  corre- 
lation'will  be  found  on  pages  481  to  484;  here,  again,  we  can  but 
summarize  the  important  points. 

I.  Certain  environmental  agents  increase  the  breadth  while  decreas- 


520  JENNINGS— HEREDITY   IN   PROTOZOA.  [ApriU*, 

ing  the  length.     Inclusion  of  different  stages  of  this  process  in  a 
sample  reduces  the  correlation ;  it  may  make  it  zero  or  negative. 

2.  Most  environmental  agents  change  the  breadth  more  than  the 
length,  even  when  both  are  changed  in  the  same  direction  The  inclu- 
sion of  different  stages  then  reduces  correlation. 

3.  Samples  in  which  some  of  the  specimens  are  well-fed  and 
plump,  others  ill-fed  and  thin,  of  course,  show  low  correlation,  since 
the  ratio  of  breadth  to  length  is  not  uniform.  This  is  usually  the 
case  in  cultures  where  food  is  scarce. 

4.  Addition  of  abundant  nutriment  causes  the  thin  specimens  to 
increase  in  breadth,  by  taking  food,  while  the  plump  ones  change 
little.  As  a  result  the  proportion  of  breadth  to  length  becomes  nearly 
uniform  throughout  the  lot;  the  correlation  is  therefore  increased. 
As  a  rule,  any  agent  which  increases  the  mean  breadth  likewise  (  for 
the  reason  just  set  forth)  increases  the  correlation  between  breadth 
and  length. 

Decrease  of  nutriment,  for  the  converse  reason,  decreases  the 
correlation. 

5.  Any  agent  that  causes  rapid  multiplication  decreases  the  cor- 
relation between  length  and  breadth  for  the  period  of  multiplication. 
This  is  owing  to  the  inclusion  in  the  collection  of  many  stages  of 
growth,  showing  different  proportions  of  length  to  breadth. 

6.  Slight  differences  in  one  dimension  may  be  produced  without 
corresponding  differences  in  the  other,  so  that  in  a  collection  varying 
little  in  length  the  correlation  may  be  low.  But  considerable  changes 
in  one  dimension  are  usually  accompanied  by  corresponding  changes 
in  the  other.  Hence,  when  two  groups  of  differing  lengths  are 
thrown  together,  the  correlation  may  become  higher  than  in  either 
one  taken  separately  (for  example,  see  page  437). 

In  any  ordinary  sample  of  Paramecium  all  these  varied  factors 
are  at  work  in  determining  the  observed  correlation.  It  is  clear  that 
no  particular  coefficient  of  correlation  can  be  considered  character- 
istic for  Paramecium  or  for  any  particular  race  of  Paramecium,  for 
by  various  combinations  of  these  factors  we  may  get  any  coefficient 
of  correlation  ranging  from  a  pronounced  negative  value  upward 
through  2ero  to  a  high  positive  value.     In  Tables  X.  and  XVIII.  we 


,,o8.]  JENNINGS— HEREDITY  IN   PROTOZOA.  521 

see  varied  collections  showing  extremes  of  value  for  the  coefficient 
of  correlation,  from  — .3138  to  +  .8500.^^ 

3.   Results  on  Variation,  Inheritance  and  the  Effects  of 

Selection. 

Our  general  results  with  regard  to  variation,  inheritance  and  the 
effects  of  selection  are  then  as  follows : 

In  a  given  "pure  line"  (progeny  of  a  single  individual)  all 
detectible  variations  are  due  to  growth  and  environmental  action, 
and  are  not  inherited.  Large  and  small  representatives  of  the  pure 
line  produce  progeny  of  the  same  mean  size.  The  mean  size  is 
therefore  strictly  hereditary  throughout  the  pure  line,  and  it  depends, 
not  on  the  accidental  individual  dimensions  of  the  particular  pro- 
genitor, but  on  the  fundamental  characteristics  of  the  pure  line  in 
question. 

In  nature  we  find  many  pure  lines  differing  in  their  characteristic 
mean  dimensions. 

Our  results  with  the  infusorian  Paramecium  are,  then,  similar  to 
those  reached  recently  by  certain  other  investigators  working  with 
pure  lines  of  other  organisms.  Johannsen  (1903)  showed  that  in 
beans  and  in  barley  many  pure  lines,  slightly  differentiated  from 
each  other,  exist  in  nature,  but  that  selection  within  a  pure  line  has 
no  effect  upon  its  characteristics.  These  plants  are  self-fertilized, 
so  that  there  is  no  intermingling  of  different  lines.  Hand  (1907) 
has  recently  found  the  same  state  of  affairs  in  Hydra  when  multi- 
plying by  budding.  Certain  lines  tend  to  have  a  higher  mean  number 
of  tentacles,  others  a  lower  mean  number.  But  within  a  given  line 
selection  of  parents  with  more  or  fewer  tentacles  has  no  effect  on 
the  progeny ;  selection  has  no  effect  within  the  pure  line. 

It  is  doubtless  too  early  to  draw  any  very  positive  conclusions 
from  these  facts.  While  the  results  with  Paramecium  seem  clear, 
I  intend  to  test  them  further  in  every  way  possible.  It  is  pos- 
sible that  selection  may  be  made  on  some  other  basis,  with  a  better 

"This  fact  of  course  does  not  render  the  study  of  the  coefficient  of 
correlation  valueless.  Its  examination  under  varied  experimental  conditions 
is  of  the  utmost  importance  for  determining  the  real  effects  of  various  agents, 
and  in  many  other  ways  it  furnishes  a  valuable  datum. 

PROC  AMER.  PHIL.  SOC.  XLVH.  190  HH,  PRINTED  JANUARY  I3,  I909. 


522  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  14. 

chance  of  avoiding  differences  due  to  environment  and  growth.  It 
is  conceivable  that  congenital  hereditary  variations  exist,  but  that 
they  are  few  in  number  compared  with  those  due  to  environment 
and  to  slight  differences  in  ways  of  living,  so  that  in  our  selection 
we  always  get  the  mere  environmental  variations.  There  are  decided 
differences  between  the  specimens  of  the  same  line  beginning  fission, 
as  Table  XIII.  (page  442)  well  shows;  here  the  length  varied  from 
156  to  204  microns.  It  is  possible  that  selection  among  specimens 
beginning  fission  might  have  a  better  chance  for  success.  I  have 
attempted  this,  but  it  is  extremely  difficult ;  I  hope  to  return  to  it. 

We  must  consider,  however,  that  if  the  non-inheritable  differ- 
ences are  so  much  more  mmierous  and  markecl  than  the  inheritable 
ones  as  to  render  conscious  selection  by  human  beings  ineffective,  they 
would  apparently  have  the  same  effect  on  selection  by  the  agencies 
of  nature.  The  same  ground  for  selection  offered  by  heritable  varia- 
tions is  offered  so  much  more  fully  by  those  not  heritable  that  there 
would  be  as  little  effect  in  selection  by  nature  as  in  selection  by  man. 

Certainly,  therefore,  until  someone  can  show  that  selection  is 
effective  within  pure  lines,  it  is  only  a  statement  of  fact  to  say  that 
all  the  experimental  evidence  we  have  is  against  this.  The  results 
set  forth  in  the  present  paper  tend  to  strengthen  that  explanation  of 
the  observed  facts  regarding  selection,  regression,  etc.,  in  mixed 
populations,  which  is  set  forth  by  Johannsen  (1903).  We  need  not 
discuss  these  in  detail  here ;  they  are  essentially  as  follows : 

1.  Selection  in  a  mixed  population  consists  in  isolating  the 
various  different  lines  already  existing. 

2.  If  selection  is  made,  not  of  single  individuals,  but  of  consid- 
erable numbers  having  a  certain  characteristic,  then  by  repeated 
selection  it  will  be  possible  to  approach  nearer  and  nearer  to  a 
certain  end. 

Thus,  if  we  select  from  such  a  heterogeneous  collection  as  is  rep- 
resented in  Table  LXI.  all  the  larger  individuals,  we  shall  have  taken 
representatives  of  many  different  lines.  Our  selection  will  include 
the  larger  individuals  of  lines  of  median  size,  as  well  as  the  average 
individuals  of  lines  of  large  size.  The  progeny  of  this  selected  lot 
will  then  consist  of  various  lines,  some  larger,  some  smaller,  but  with 
the  average  higher  than  in  the  original  collection.    Another  selection 


^ 


,9o8.1  JENNINGS— HEREDITY  IN  PROTOZOA.  523 

will  raise  the  average  still  further  by  getting  rid  of  some  of  the 
smaller  lines,  etc. 

3.  It  has  been  noticed  that  in  many  cases  continued  selection  will 
not  carry  a  character  beyond  a  certain  point.  This  is  due  (on  the 
view  we  are  setting  forth)  to  the  fact  that  we  have  finally  isolated 
that  line  (or  lines)  of  the  original  collection  which  had  this  character 
most  strongly  marked,  and  since  selection  of  the  fluctuations  has  no 
eflFect  within  the  pure  line,  we  can  make  no  farther  progress. 

4.  The  phenomenon  of  so-called  regression  finds  its  explanation 
in  the  same  way.  It  is  found  that  when  extremes  are  selected,  the 
progeny  of  these  extremes  stand  nearer  the  mean  than  did  the  par- 
ents, though  they  diverge  in  the  same  direction  as  the  parents.  The 
reason  for  this  may  again  be  seen  by  considering  such  a  hetero- 
geneous collection  as  that  of  Table  LXL,  with  the  effects  of  selecting 
the  extremes  of  size.  If  we  select  the  largest  and  the  smallest  indi- 
viduals, we  shall  have  taken  ( i )  the  largest  individuals  of  the  largest 
lines,  and  (2)  the  smallest  individuals  of  the  smallest  lines.  But 
these,  when  they  propagate,  produce,  as  we  have  seen,  merely  the 
means  of  the  lines  to  which  they  belong.  The  largest  individuals 
will  produce  then  progeny  that  average  smaller  than  themselves; 
the  smallest  individuals  progeny  that  are  larger  than  themselves; 
both  sets  will  then  approach  the  mean  of  the  original  collection  as 
a  whole. 

In  working  with  populations  reproducing  by  cross  fertilization 
among  the  different  lines,  the  conditions  on  which  these  results 
depend  become  quite  obscured,  owing  to  the  introduction  of  new 
factors,  the  union  of  different  factors,  the  appearance  of  mendelian 
results,  etc.  Work  with  pure  lines  perhaps  shows  the  real  cause 
for  the  observed  phenomena  above  set  forth. 

It  must  be  admitted,  then,  that  the  work  with  pure  lines,  indi- 
cating that  selection  of  fluctuations  within  the  lines  is  powerless, 
leads  to  a  simple  and  consistent  explanation  of  many  of  the  observed 
facts.  But,  of  course,  it  gives  no  explanation  of  the  origin  of  the 
different  pure  lines.  Clear  proof  of  the  effectiveness  of  selection 
even  within  a  pure  line  would  therefore  be  of  the  greatest  interest, 
and  the  present  writer  would  find  great  pleasure  in  being  the  first  to 
present  such  proof.     But  until  such  proof  is  forthcoming,  it  must  be 


524  JENNINGS— HEREDITY   IN   PROTOZOA.  lAprila4. 

admitted  that  the  experimental  results  go  strongly  against  the  effect- 
iveness of  selection  among  slight  fluctuating  variations  in  producing 
new  inherited  characteristics. 

How,  then,  do  the  different  pure  lines  rise?  This  is  after  all  the 
main  problem.  Toward  its  solution  further  investigations  of  this 
series  will  be  directed.  It  is  proposed  to  study  in  detail  (i)  the 
effects  of  conjugation  on  variation,  heredity  and  the  production  of 
new  races;  (2)  the  effects  of  long-continued  differences  in  environ- 
mental action  on  different  divisions  of  the  same  line;  (3)  the  ques- 
tion whether  the  different  lines  arise  from  something  like  mutations. 
Further,  (4)  additional  different  way  of  exercising  selection  within 
a  single  line  will  be  tested.  The  question  may  be  raised  whether  the 
production  "  by  mutation  "  of  such  slight  differences  in  size  as  we 
are  here  dealing  with  would  not  be  essentially  the  same  as  their 
production  by  the  inheritance  of  slight  variations — since  the  extent 
of  the  "  mutations  "  would  not  be  greater  than  what  we  should  call 
slight  variations  in  size.  The  difference  between  the  two  conceptions 
almost  or  quite  vanishes  when  we  come  to  deal  with  such  minute 
changes  in  characteristics  as  those  we  find  in  the  different  lines  of 
Paramecium.  The  "  mutation  "  would  be  merely  a  rare,  heritable, 
variation,  and  it  is  now  clear  that  heritable  variations  in  size  are 
much  rarer  than  had  been  supposed;  their  number  is  so  small  that 
in  Paramecium  they  are  not  statistically  detectible  among  the  many 
non-heritable  fluctuations  due  to  the  environment. 

Raquette  Lake,  New  York, 
August  22,  1908. 


,9o8.]  JENNINGS— HEREDITY  IN   PROTOZOA.  526 


LIST  OF  LITERATURE. 
Calkiiis,  6.  N. 

1906.    The  Protozoan  Life  Cycle.    Biol.  Bui.,  11,  229^244. 
Calkins,  6.  N. 
Z906.    Paramecium  aurelia  and  Paramecium  caudatum.    "  Biological  Studies  " 

by  the  Pupils  of  Wm.  T.  Sedgwick,  Chicago. 
Davenport,  C.  B. 
1899.    Statistical  Methods,  with  Special  Reference  to  Biological  Variation, 

New  York. 
Z904.    Idem.     Second  edition. 
Donaldson,  H.  H. 

1906.  A  Comparison  of  the  White  Rat  with  Man  in  Respect  to  Growth.  Boas 
Memorial  Volume,  New  York,  pp.  5-26. 

Hanel,  Blise. 

1907.  Vererbung  bei  ungeschlechtlicher  Fortpflanzung  von  Hydra  grisea. 
Jenaische  Zeitschr.,  43,  321-372. 

Jennings,  H.  S. 

1908.  Heredity,  Variation  and  Evolution  in  Protozoa.  I.  The  Fate  of  New 
Structural  Characters  in  Paramecium,  with  Special  Reference  to  the 
Question  of  the  inheritance  of  Acquired  Characters  in  Protozoa.  Journ. 
Exp,  ZooL,  5,  577-632. 

Johannsen,  W. 

1903.    Erblichkeit  in  Populationen  und  in  reinen  Linien.    68  pp.    Jena. 

Johannsen,  W. 

1906.  Does  Hybridisation  Increase  Fluctuating  Variability?  Report  of  the 
Third  International  Conference  (1906)  on  Genetics.    London. 

McClendon,  J.  F. 

1908.    Protozoan  Studies,  I.    Journ,  Exp.  Zool.,  6. 
Pearl,  R. 

1907.  A  Biometrical  Study  of  Conjugation  in  Paramecium,  Biometrika,  S, 
213-297. 

Pearl,  R.,  and  Dunbar,  F. 

1905.    Some  Results  of  a  Study  of  Variation  in  Paramecium.    Seventh  Re- 
port Michigan  Acad.  Sci.,  pp.  77-86. 
Pearson,  K. 

190a.    Note   on   Dr.    Simpson's   Memoir   on   Paramecium   caudatum.     Bio- 
metrika, I,  404-407. 
Robertson,  T.  B. 

1908.  On  the  Normal  Rate  of  Growth  of  an  Individual  and  its  Biochemical 
Significance.    Arch,  f,  Entw.-mech.,  25,  582-614. 

Simpson,  J.  Y. 

190a.    The  Relation  of  Binary  Fission  to  Variation.    Biometrika,  i,  400-404. 

Yule,  G.  U. 

1897.    On  the  Theory  of  Correlation.    Journ.  Roy.  Statistical  Society,  60,  1-44. 


526 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[April  a4. 


APPENDIX. 


Tables  of  Measurements. 


The  first  twenty-eight  tables  are  distributed  through  the  text 
Tables  XXIX.  to  LXIII.  follow. 


Table  XXIX. 

Correlation  Table  for  Length  and  Breadth  of  59  Specimens,  Age 
Minutes,     (See  Lot  2,  Table  10.)     Descendants  oi  D. 

Length  in  Microns. 
76  80  84  88  92  96  100  104  108  112  116  120  124  128  132 


o  to   s 


CO 

C 
O 
u 
o 

c 


36 

I 

40 

I      I 

I 

2 

I 

44 

2           4 

I 

2 

5 

I 

3 

2 

I 

48 

I     2    3     I      3 

I 

2 

2 

I 

2 

I 

I 

52 

I     I                3 

I 

I 

I 

I 

I 

56 

I 

2112828      3 

Length — Mean,  107.660  ±.  i,2f^ 
St  Dev.,  14,780  ±  .916A1 
Coef .  Van,  13.729  ±   .868 


I 

6 

21 

20 

10 

I 


53657331  59 
Breadth— Mean,  46.372  ±-  .332M 

St.  Dev.,       3.804^^36^ 
Coef.  Var.,   8.200^1.524 


Mean  Index,  44.037  per  cent. ;  Coef.  Cor.,  —  .3138  ±  .0792. 


Table  XXX. 

Correlation  Table  of  Length  and  Breadth  for  a  Random  Sample  of  Lot  2, 

Table  X. — Same  Lot  from  which  came  Specimens  in  Tables  VIL  and 

XXIX,    Descendants  of  D,    (24  hours  in  fresh  hay  infusion:  July  17.) 

Length  in  Microns. 

«     w     ct     w     « 


I    o    2    2    7    14   9    8   14  13  14  15  18  22  17  17  12   7    5    3  200 

Length — Mean,  184.100  ±  .776At  Breadth — Mean,  46.020  ±.  .251M 

St.   Dev.,       16.264  ±  .548A*  St.  Dev.,        5.256  ±.177/* 

Coef.  Var.,     8.834  ±-  .300  Coef.  Var.,  11421  ±  .390 

Mean  Index,  25.084  per  cent. ;  Coef.  Cor.,  .4282  ±.  .0389. 


X9o8.] 


JWNINGS— HEREDITY  IN   PROTOZOA. 


627 


Table  XXXI. 

Correlation  Table  for  the  Length  and  Breadth  of  the  Young  of  Lot  6,  between 
the  Ages  of  o  and  19  Minutes,    (Sec  Table  X.,  row  7.) 

Length  in  Microns. 


in 

^  60 

.S  64 

J5  68 

^  72 

S  76 

u 


108  112  116  120  124  128  132  136  140  144  148  152 


I 

2 


I 
I 


I 
I 


Length—Mean,  128.000  ±  1.908/* 
St  Dev.,  13.856  ±  1.348A* 
Coef.  Var.,  10.825  ±.  1.066 

Mean  Index,  47.573  per  cent. ;  Coef.  Cor.,  —  .0337  ±  .1375. 


J 
I 

I      3      1 
I       I 

I 

I 

3 
6 

8 

4 
I 
I 
I 

I      2      6      I       I       I    i  24 

Breadth— Mean,           60.168  -+■  .788M 
St  Dev.,        5.712 ±.S56m 
Coef.  Var.,  9-495^.933 

Table  XXXII. 

Correlation  Table  for  Length  and  Breadth  of  Young  of  Lot  7,  between  the 
Ages  of  0  and  19  Minutes,  Descendants  of  Individual  D,    (See  Table 

X.,  row  13.) 

Length  in  Microns. 


Vi 

§ 

108 

112 

116  120  124  128  13 

u 





— — 

i  36 

•^  40 

I 

.S  44 

3 

121 

s  48 

I       I       2      I 

^52 

I 

I 

I      3       I       I 

I 

4 

I      3      7      4      I 

I 
4 


I 
I 
I 
2 
I 


I 
I 


I 

3 
12 

14 
39 


Length— Mean,         134^56  ±  1.663A*  Breadth— Mean,  46.768 

St   Dev.,     15.394  ±  1.176A*  St.  Dev.,        3.792 

Coef.  Var.,  1 1468  ±   .857  Coef.  Var.,    8.109 

Mean  Index,  35.643  per  cent. ;  Coef.  Cor.,  —  .2546  ±  .1010. 


uto8M 
.288M 
.623 


528 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[April  14, 


Table  XXXIIL 

Correlation  Table  for  Length  and  Breadth  of  Young  of  Lot  6,  between  the 
Ages  of  i8  and  28  Minutes,    (Sec  Table  X.,  row  8.) 

Length  in  Microns. 


i 

132 

136 

140 

144 

148 

152 

156    160 

■^  48 

.s  56 
«  64 

2 

4 

4 

I 

6 

3 
2 

5 
7 
2 

2 

I 

I 
I 

I 
I 
I 

2 

I        I 

I 

5 
22 

13 

7 

2 

0 

Length — Mean, 

St.  Dev., 

2       9 

143.348  ± 
6.480  ± 

II 

.624Ai 

440/i 

14 

5        3 
Breadth- 

4        I 

-Mean, 
St.  Dev., 

49 

• 

54.284  ±  .364M 
3.788  ±  .260^ 

Coef.  Var.,     4.521  ±.  .309  Coef.  Var.,    6.976  ±.  478 

Mean  Index,  37.921  per  cent. ;  Coef.  Cor.,  1937  ±  .0927. 


Table  XXXIV. 

Correlation   Table  for  Length  and  Breadth  of  106  Specimens,  Age   18-28 
Minutes.     (See  row  15,  Table  X.)     (Descendants  of  D,  but  taken 

part  one  day,  part  another.) 

Length  in  Microns. 


112  116  120  124  128 132  136  140  144  148  152  156 160  164  168 


in 

C 

o 

u 
u 

c 


PQ 


36 

40 

44 
48 
52 
56 
60 

64 


I 

I 

I 

3 

2 

I 

2 

I 

I 

I 

6 

16 

3 

3 

2 

I 

5 

6 

8 

3 

4 

3 

I 

4 

4 

3 

2 

3 

I 

I 

2 

I 
I 

I 

I 
I 

I 

I     2     7     II    19  29    13    10    9     3 


I 
I 

14 
33 
31 
17 

7 

2 


106 


Length— Mean,  143.812  ±  .544A* 

St.  Dev.,         8.296  ±  .384/* 
Coef.  Var.,     5.769  ±  .268 

Mean  Index,  35438  per  cent.;  Coef.  Cor.,  1319^  .0644. 


Breadth — Mean,  50.832 

St.  Dev.,        4.900 
Coef.  Var.,    9.640 


.3«V* 
.228^ 

451 


igo8.) 


JENNINGS— HEREDITY  IN  PROTOZOA. 


529 


Table  XXXV. 

Correlation  Table  for  Length  and  Breadth  of  Young  of  Lot  6,  between  thf 
Ages  of  35  and  43  Minutes,    (See  Table  X.,  row  9.) 


o 

•S  48 

^  52 

.S  56 

^  64 
u 


Length — Mean, 

St.  Dev., 


132 


Length  in  Microns. 
136    140    144    148    152    156    160 


I 

2 
1 


I 

3 

I 

I 

I 

6 

2 

I 

2 
I 

9 

3 

I 

I 

I 

4 

5 

5 

3 

5 

25 

149.920  ±  I.0I2M 
7.512  ±    .716M 


Breadth — Mean, 

St.   Dev., 


55.840  ±  .636M 
4.724  ±  452M> 


Coef.  Van,    5.010  ±   479 


Coef.  Var.,  8461  ±  .8ia 


Mean  Index,  37.296  per  cent. ;  Coef.  Cor.,  .2799  ±  .1243. 


Table  XXXVI. 

Correlation  Table  for  Length  and  Breadth  of  Young  of  Lot  6,  between  the 
Ages  of  75  and  90  Minutes,     (See  Table  X.,  row  10.) 

Length  in  Microns. 


xn 


140   144   148   152   156   160   164   168   172   176   180   ^ 


o  40 

M  44 

s  48 

c  52 

.S  56 

rS  60 

2i  68 

PQ 


I 

I 

2 

I 

I 

I 

I 

I 

I 

2 

I 

6 

I 

4 
4 

I 

T 

I 

I 

2 

I 

I 

2 

I 

I 

I 

I 

1 

12 


I 
O 

8 

16 

6 
8 
o 

_3 
42 


Length— Mean,         161.524  ±  1.004/i  Breadth — Mean,  54.192  db  .600M 

St.  Dev.,        9.648  ±    .712/i  St.   Dev.,       5.752^424/* 

Coef.  Var.,  5.974  ±   441  Coef.  Var.,  10.617  ±  .790 

Mean  Index,  33.558  per  cent. ;  Coef.  Cor.,  .5232  ±  .0756. 


530 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  M. 


Table  XXXVII. 

Correlation  Table  for  Length  and  Breadth  of  Young  of  Lot  p,  between  the 
Ages  of  3  and  4  Hours,    (Sec  Table  X.,  row  16.) 

Length  in  Microns. 
132    136    140    144    148    152    156    160    164    168    172    176 


CO 

2 

o 

.S 

t 


40 

I 

44 

I 

I 

3 

I 

2 

48 

2 

I 

? 

I 

3 

3 

4 

3 

52 

5 

6 

4 

3 

2 

3 

4 

56 

3 

2 

I 

3 

3 

3 

60 

2 

I 

I 

64 

I 

I 

2 


I 
I 


19 


12 


II 


12 


10 


I 

9 

i    27 
30 

i  17 

I     6 

I     3 

93 


Length— Mean,  149.636  ±  .688^ 

St  Dev.,        9.856  ±488/* 
G)ef.  Var.,    6.587 ±.327 

Mean  Index,  34.546  per  cent;  Coef.  G>r.,  .3201  ±  .0628. 


Breadth— Mean,  51.568 

St.    Dev.,       4,752 
Coef.  Var.,    9.212 


.322M 
.236^ 

459 


Table  XXXVIII. 

Correlation  Table  for  the  Length  and  Breadth  of  Young  of  Lot  9,  between 
the  Ages  of  4,20  and  5  hours,    (See  Table  X.,  row  17.) 

Length  in  Microns. 


o 

u 

o 


V. 

PQ 


164  168  172  176  180  184  188  192  196  200  204  208  212  216 


9     15     13     18    12     8 


95 


Length— Mean,  186.736  ±  .652M 

St   Dev.,        9416  ±460^ 
Coef.  Var.,      5.043  ±  .247. 

Mean  Index,  32.225  per  cent. ;  Coef.  Cor.,  .5557  ±  .0478. 


Breadth— Mean,  6o.i68±  .360^ 

St   Dev.,       5.224  ±.256** 
Coef.  Van,  8.679  ±428 


i9o8.] 


JENNINGS— HEREDITY  IN  PROTOZOA. 


631 


Table  XXXIX. 

Correlation  Table  for  Length  and  Breadth  of  Paramecia  at  the  Age  of  12 
Hours.    (Descendants  of  D;  See  Table  X.,  rows  20  and  21.) 

Length  in  Microns. 


s  48 

2  52 

a  56 

a  60 

.S  64 

?  72 

8  76 
£80 


so      O^OOWVOQ^OQ 

vO 

06  06  00 

M    \0     Q 

1 

1 

VO 

I                                                      I 

2 

I 

I 

2 

I 

I 

I 

3 

2 

1X2 

12 

2 

I 

2     2 

2     I     4 

I 

15 

I 

2     3 

4 

I    5 

5    4    2 
332 

I 

I 

3 
I 

I 

21 

7 
0 

I 

IIOOOOOII 

2 

4 

4 

7   14 

12  10  10 

5 

0 

0 

I 

73 

Length— Mean,  188.988  ±  .996M  Breadth— Mean,  62.796  ±.464^ 

St  Dev.,        12.6x2 ±.704^  St  Dev.,        5.872 ±.328^ 

Coef .  Var.,      6.672  ±  .374  Coef .  Var.,    9.350  :*:  526 

Mean  Index,  33.275  per  cent. ;  Coef.  Cor.,  4868  ±  .0602. 


Table  XL. 

Correlation  Table  for  Length  and  Breadth  of  Paramecia  at  the  Age  of  18 
Hours.    (Descendants  of  D;  See  Table  X.,  row  22.) 


i 

.S  60 
I  68 


Length  in  Microns. 
168  172  176  180  184  188  192  196  200  204  208  212  216  220  224  228 


I 

2 

I 

I 

I 

I 

I 

I 

5 

2 

6 

5 

4 

3 

I 

I 

V 

I 

I 

4 

2 

4 

6 

5 

2 

5 

2 

I 

I 

I 
I 

5 

I 

4 
4 

4 
I 

2 

2 
I 

5 

I 
2 

I 

6      7      8      6     14    16    12    13     9      6      3 


5 
31 
30 
26 
II 

2 

1 105 


Length— Mean,            199.048 ±  .780^1          Breadth— Mean,  56^(96 ±.292^ 

St  Dev.,        xx.844±.552M                         St  Dev.,  4.428 ±.2o&« 

Coef.  Var.,      5.949 ±.278                          Coef.  Var.,  7837 ^3^7 

Mean  Index,  28427  per  cent. ;  Coef.  Cor.,  4304  ±  .0536. 


532 


JENNINGS— HEREDITY   IN  PROTOZOA. 


[April  «4. 


Table  XLI. 

Correlation  Table  for  Length  and  Breadth  of  300  Paramecia  at  the  Age  of 
24  Hours,     (Descendants  of  D;  Sec  Table  X.,  row  23.) 

Length  in  Microns. 


5 

43 
66 

69 
65 
36 
10 
6 

300 

Length— Mean,  168.532  ±  uticy*  Breadth— Mean,  40.320  d::  .23CMB1 

St.  Dev.,        10.768  ±  .296A1  St  Dev.,       5.892  d=  .i62« 

Cocf.  Var.,      6.389  ±  .175  Coef.  Var.,  14.615  ±411 

Mean  Index,  23.899  per  cent. ;  Coef.  Cor.,  .5496  ±  .0272. 


• 

140 144  148  152 

156  160  164  168  172  176  180  184  188 192  196  200 

§  28 

I 

2*     I 

I 

t3  32 

I 

4 

2      5 

8 

8762 

*s  36 

5 

8 

8     9    15    10    7     2     I                  I 

.S  44 

I     2 

5 

3    10   16   12    7     9     3     I 

I 

I 

2 

69    10   12    7664           I 

5  48 

2 

24318636                 I 

"S  52 

233                 2 

2i  56 

cq 

I           2                  III 

2 

r 

5    14 

2<J 

27  40   52   39  32   26   14    12    3     2     I 

Table  XLII. 

Correlation  Table  for  Length  and  Breadth  of  62  Dividing  Specimens  of  Lot 

2.    (Descendants  of  D;  See  Table  X.,  row  31.) 

Length  in  Microns. 


• 

C 

144 

148 152  156  160  164  168  172  176  180  184  188  192  196  200  204  208  212 

t3  40 

2  44 

-  52 

5  56 
•S  60 

I 

I 

I 
22                                                         I 
111213322111                       I 
I     2     2     7     3     3     5     I                                        I 
121                       121 

I 

cq 

I 

I 

247124694421201     I     I 

I 

6 

20 

26 

8 
I 

62 


Length- Mean,           171.548  ±  i.i88fi         Breadth— Mean,  50.388  ±  .308^ 

St.  Dev.,        13.848  ±  .840^                        St.  Dev.,  3.584  ±.2i6i» 

Coef.  Var.,      8.072  ±   ut92                           Coef.  Var.,  7.iii±u|33. 

Mean  Index,  29.583  per  cent.;  Coef.  Cor.,  — .ii36±  .0840. 


I908.J 


JENNINGS— HEREDITY   IN   PROTOZOA. 


533 


Table  XLIII. 

Correlation  Table  for  Length  and  Breadth  of  Specimens  in  Early  Stages  of 
Fission:   Constriction  less  than  one-fourth  Breadth.    Lot  2,     (See 

Table  10,  row  30.) 


c 
o 

a  44 

s  48 

.s  52 

^  56 

■^  60 


Length  in  Microns. 
144    148    152     156    160    164    168    172    176    180 


I 

2 

I 

I 

I 

I 

I 

3 

3 

I 

I 

2 

2 

6 

I 

3 

I 

I 

I 

2 

I 

I 

I 

Length-— Mean,  165.200  ±  .936^1 

St.  Dev.,         8.788  ±  .664M 
Coef .  Var,      5.320  ±  .402 


10 


4 
12 

18 

5 
I 


40 


Breadth — Mean,  50.700  dr  .364^* 

St.  Dev.,        3.432  ±  .260M 
Coef.  Van,    6.769^.513 


Mean  Index,  30.76^  per  cent;  Coef.  Cor.,  .1048 ±.1055. 


Table  XLIV. 

Correlation  Table  for  Length  and  Breadth  of  Early  Stages  of  Fisson,  in  Lot 
3.     {Depth  of  Constriction  less  than  one-fourth  Breadth.)     (See 

Table  X.,  row  24.) 


152 


Length  in  Microns. 
156  160  164  168  172  176  180  184  188  192 


S  48 

§  52 

s  ^^ 

2 

S  60 

.S  64 

I 

n  68 

I 

^  72 

S  76 

^  80 

2 

3 
I 


2 
2 
I 

3 


8   8 


I 
2 

3 
2 


8 


I 

2 


I 
o 
6 

4 
10 

II 

4 

5 
I 

42 


Length—Mean,  167.620  ±  .996^^  Breadth— Mean,  65.716  ±  .706^ 

St.  Dev.,         9-5^  -  -704^  St.  Dev.,        6.784  ±  .499M 

Coti.  Van,      5.706  ±  421  Coef.  Van,  10.322  ±  .76a 

Mean  Index,  39.286  per  cent. ;  Coef.  Cor.,  .2215  ±  .0999. 


534 


JENNINGS— HEREDITY   IN   PROTOZOA. 


I  April  S4* 


Table  XLV. 

Correlation  Table  for  Length  of  Body  and  Depth  of  Constriction  in  119 

Dividing  Specimens  of  the  Aurelia  form,  Descended  from  c, 

(Sec  Lot  4.    Tables  VIII.  and  X.) 


Length  in  Microns. 


c 
o 

u 

i 

s     3.3 

=2   6.7 

c   10.0 

•2  13.3 
.a  16.7 
^  20.0 

g      23.3 

cj  26.7 
^  30.0 

®  33.3 

■*-• 

o 

Q 


«oi*»o   «oi*»o   coi*»o   «oi*»o   «oi*»o   «oi*»o   cot^o   «Ol*» 

O    «0  NO    O    «0  NO    O    CO  so    O    «0  so 


•ovo  6  «oxO  0  covO   o  covo   o'  «ovd  c>  «ovd  d  «osd  o*  «osd 

00  00    ON  On  OS   O    O    O    I-"    •-•**'""     " 


I    I    I    5   3  10  |i  6   9   3 

I      4331 

I 


2     I 


I    3 


I    I 


I 

2 


2 

I 


2 
2 

1 
I 


I 

2 
2 

4 
I 

I    I 


I 

2 


3 
I 

3 


2    I 


i  54 

12 

'     8 

I    ID 

I     7 

13 

'  \ 

I  I      2 


I     I    63   15  16  13  13  87    II   6382    1-2    00    I    O    I   119 


Length— Mean,  111.541  ±  .797^*       Depth  of  Constriction,  Mean,  iaSQ4Ai 

St.  Dev.,        12.898  ±  .564/*  St  Dev.    a43i/i 

Coef.  Van,    11.563  ±  .512 
Coef .  Cor.,  7862  ±  .0236.    Increase  in  length  with  iom  increase  in  depth 

of  Constriction,  12.027/*. 


Table  XLVI. 

Correlation  Table  for  Length  of  Body  and  Depth  of  Constriction  in  63 
Dividing  Specimens  of  the  Aurelia  form,  Descended  from  c,    (See 

Lot  5,  Tables  VIII.  and  X.) 

Length  in  Microns. 


Vi 

C 

2 

•  v4 

«o 

• 

l^ 

^ 

• 

8 

«o 

d 

*o 

• 

»o 

• 
NO 

d 

to 

• 

«o 

no' 

d 

«o 

• 

«o 

no' 

d 

«o 

• 

«o 

• 

so 

• 

0 

«o 

• 

*o 

ON 

0 

^4 

^4 

^4 

1^ 

I 

«o 

^4 

«o 

^4 

*o 

1^ 

•^        3.3 

I 

3 

7 

4 

5 

4 

1    25 

c     6.7 

I 

2 

2 

I 

2 

2 

10 

e    ^^' 

I 

2 

3 

0    13.3 

2 

I 

I 

'     4 

t;  16.7 

I 

I 

n  20. 

2 

2 

S  23.3 

I 

I 

2 

0  26.7 

^  30- 

I 

2 

I 

I 

I 
I 

I 
I 

3 
6 

•s  33.3 

I 

2 

I 

I 

I 

6 

I  36.7 

I 

I 

T3 

I 

0 

3 

0 

8 

I 

8 

7 

8 

3 

6 

5 

2 

I 

2 

3 

I 

2 

2 

63 

Q 

i9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA. 


535 


Table  XLVII. 

Correlation  Table  for  Length  and  Breadth  of  Dividing  Specimens  of  Lot  4, 
in  which  the  Depth  of  Constriction  was  Less  than  one-fourth  the 
Breadth,  (Aurelia  form.  Descendants  of  c.)  (Sec  Table  X.,  row 
33.) 

Length  in  Microns. 


CO 


«o     t^ 


CO  l^ 


CO  t^ 


• 

c 

00 

S 

a 

«. 

8 

? 

^ 

0 

CO 

>-• 

vO 

s 

CO 

VO 

0 

^4 

•H 

"^ 

M 

*^ 

^4 

^4 

►i^ 

H* 

a    26.7 
•^    30. 

I 

I 

I 

I 

2 

I 

3 

2 

I 

.s  33.3 

2 

II 

6 

2 

2 

I 

42    36.7 

2 

2 

5 

4 

4 

2 

I    ! 

•S  40. 

I 

2 

2 

I 

I 

1 

8  43.3 

2 

I 

1 

cq 

I 

I 

I 

6 

3 

14 

14 

9 

10 

3 

I 

0 

2 

I    1 

Length^Mean,  103.737^1.650^  Breadth — Mean,  34-850 

St.  Dcv.,  7.823 ± .379^  St.  Dev.,        3453 

Coef.  Var.,      7.541  ±  445  Coef.  Var.,    9.91 1 

Mean  Index,  33.623  per  cent. ;  Q)ef .  Cor.,  .6502  ±  .0479^1. 


I 
II 

24 
20 

7 
3 

66 

:.297M 
:.203M 

:.587 


Table  XLVIII. 

Correlation  Table  for  Length  and  Breadth  of  Diznding  Specimens  of  Lot 
S,  in  which  the  Depth  of  Constriction  was  Less  tham  one-fourth  the 
Breadth.     (Aurelia  form.  Descendants   of  c)      (See  Table  X.,  row 

36.) 

Length  in  Microns. 


CO 


CO 


cn 

C 

o 


g   ^    8     ?   ^    2 


CO       l^ 

CO     vd 


o 


CO       l^ 

CO         NO 


5 


33.3 
36.7 

I 

I 

40. 

I 

43.3 
46.7 

I 

50. 

53.3 
56.7 

2 

3 

2 

I 

8 


I 

5 

2 


8 


I 
I 

3 
2 


I 
I 

I 

2 
5 


I 
I 


I 
2 


3 
I 

6 

4 

13 

9 
o 

2 
38 


Length— Mean,  113.333  =t  .8s<W*  Breadth— Mean,  45263 

St.  Dev.,  7.778  ±  .603M  St  Dev.,        SA'^S 

Coef.  Var.,     6.8(52  ±  .533  Coef.  Van,  12.071 

Mean  Index,  39.903  per  cent. ;  Coef.  Cor.,  .6744  ±  .0507. 


.597M 
^3M 
.947 


536 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  84, 


C    20. 

§  23.3 
a  26.7 
•^    30- 

.5  33.3 
36.7 
40. 


8  43.3 
^  46.7 


m 


50. 


Table  XLIX. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  of  Lot  4. 
(See  Table  10.    Aurelia  form.  Descendants  of  c.    Many  dividing.) 


Length  in  Microns. 


Ot^  «Ot^  cot^  «Ot^  cOl^  «Ot^  cOl*»  «Ot^  «*5I^ 

OvOO«i)4?      Q.«!>Si.S      SPS?      O      *OvO      O      rfi  \p      O     «OvO      O      cOvQ      O      «OnOQ 
^l*»CO0000      OvO>a^O      O      O      >-*      m      i^      M      nw      fOcOtO^rJ-^iOiOiONi 


CO 


I 

I 

I 

I 

I 

' 

I 

2 

2 

I 

2 

6 

2 

3 

I 

2 

I 

I 

2 

2 

I 

5 

6 

5 

6 

7 

6 

6 

I 

I 

2 

I 

1 

1 

I 

I 

I 

3 

4 

3 

2 

5 

7 

7 

6 

2 

4 

2 

3 

2 

I 

I 

2 

2 

3 

3 

2 

7 

5 

5 

2 

4 

I 

3 

I      ! 

I 

I 
I 

2 

I 

4 

3 

I 

I 

6 
I 

I 

4 
2 

I 

2 

I 

I 
2 

2 
I 

I 

3 

I 

I 

2 

I 
I    I 

Length— Mean,  Ii4.i63±  .784M  Breadth— Mean,  34-207 

St.  Dev.,        17443  ^  'SSSf^  St.  Dev.,        5.363 

Coef.  Var.,    15.279  ±  .497  Coef.  Var.,  15.683 

Mean  Index,  30.177  per  cent. ;  Coef.  Cor.,  .6757  ±  .0244. 


±.24If» 

±.171/* 
±511 


I 
4 
23 
52 
53 
42 

30 
10 

9 
I 


2    2    o    7    6    2    7   12  19  13  18  16  17  19  16  10  12  12  8    5    4    7    3    2    2    3    i  225 


Table  L. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  of  Lot  5 
(Table  X.).    Aurelia  form;  Descendants  of  c.    24  Hours  in  a  Fresh 

Hay  Infusion, 


ro     l*» 


Length  in  Microns. 


«0     l*»  CO     l*»  fO     l*» 


«o    t>. 


«o   l^ 


So     tOx©     o     tovp     O     tOvO     O     «OvO     O     tOvO     O     «OvO 
Q\     O^     0\     O      00      •-'      "^      <-•      «      W      «      eOfOrO^^^ 


§  36.7 

g  40. 

§  43.3 

•^  46.7 

.S  50. 

M  53.3 

^  56.7 

8  60. 

c5  63.3 
66.7 


2 

3 

I 

I 

3 

I 

3 

3 

I 

I 

2 

3 

I 

6 

2 

2 

3 

2 

I 

I 

5 

5 

I 

I 

4 

5 

5 

I 

I 

2 

I 

4 
I 

2 


I    8    7    5    13  4    12  14  10  8    5    3 


I 

I 

I 

2 

2 

I 

2 

I 

I 
I 

I 

I 

I 

5 

3 

2 

2 

I 

2 

I 

7 

14 
22 

13 

19 

13 
6 

4 
I 

I 
100 


Length— Mean,  114.033^.820^  Breadth— Mean,  47300 

St  Dev.,       12.140  ±  .580^  St  Dev.,        6.490 

Coef.  Var.,    10.646 ±.513  Coef.  Var.,  13.720 

Mean  Index,  4x455  per  cent. ;  Coef.  Cor.,  .8152  ±  .0226. 


.437M 
.3io#* 

.667 


1908.1 


JENNINGS -HEREDITY  IN  PROTOZOA. 


537 


Table  LI. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  of  the 

Culture  from  which  came  the  Young  of  Lot  6,  Table  X.,  after  24 

hours  in  fresh  hay  infusion,    (See  row  2,  Table  XVIII.) 

Length  in  Microns. 

156  160 164  168 172  176 180  184 188 192  196  200  204  208  212  216  220  224 


I     4     2     5     9     9     8    18   13 

Length— Mean,  184.680  ±  .848/* 

St.  Dev.,        12.596  ±  .600M 
Coef.  Var.,     6.821  ±  .327 


885630 

Breadth— Mean,  64.880  ±  .580^ 

St.  Dev.,       8.624  ±4i2M 
Coef.  Van,  13.292^.645 


Mean  Index  or  Ratio  of  Breadth  to  Length,  35.131  per  cent. ;  Coef.  Cor., 
.6469  ±  .0392. 

Table  LIL 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  D,  in  Culture 
Fluid  where  Injurious  Bacteria  have  Multiplied.    June  2$.    (See  row 

5,  Table  XVIII.) 

Length  in  Microns. 


•H       H 

4MMMMMIHMMI-4MMM 

•^ 

« 

w 

« 

M     M     C4     C4     C4     (i 

w    «    fi 

« 

w   ti 

.  36 
s  40 

S  44 
.H  48 

S  53 

I 

I 

• 

I                                                  I 

I                       I    2    I     I     I     I 
I    2    I    3    I     I    6 

I          238542 

I 

5 

2 

3 

I 

2 

3 

I 
I 

I       I 
I       I 

I 

2 

12 

24 
36 

•s|^ 

II                                  411 

2 

6 

I 

5 

I    5   4    I    I    3 

I     I 

39 

*  5° 

I                 12     1 

I 

3 

2 

32243 

I 

26 

^  5t 

I 

2 

232           2 

I           I 

I 

15 

S  68 

80 

I 

I 
I 

2 

I     I     I     I     2 

I   I 

3   2 

I    I 
I 

I 

I 

15 

4 

3" 
s 

I   o  I   o  3   I   4  I   6  8  14 15  6  5  II 15 10 II  7  13 10  8  5  7  7  4  2   I    I    I 

Length— Mean,  201.888  ±.  1.147^*  Breadth — Mean,  56.112  ±  .395/* 

St.  Dev.,        22.680  ±   .81 IM  St.  Dev.,        7.808  ±.279^ 

Coef.  Var.,    11.233  ±   407  Coef.  Var.,  13.913  ±  .507 

Mean  Index  or  Ratio  of  Breadth  to  Length,  27.850  per  cent. ;  Coef.  Cor., 

.6771  ±  .0274. 

PROC.  AMER.  PHIL.  SOC.  XLVII.   I90  IF,   PRINTED  JANUARY  I3,    I909. 


178 


538 


JENNINGS— HEREDITY   IN   PROTOZOA. 


[April  •4, 


Table  LIII. 

Correlation  Table  for  Length  and  Breadth  of  a  Starving  Culture  of  De- 
scendants of  D,    Eleven  days  in  small  watch  glass  of  hay  infusion, 
not  renewed.    (Sec  row  6,  Table  XVIII.) 

Length  in  Microns. 

.    128  132  136  140  144  148  152  156  160  164  168  172  176  180  184  188 


S  28 

3 

I 

2 

6 

a  32 

I 

I 

6 

2 

5 

I 

2 

18 

S  36 

I 

2 

6 

7 

2 

4 

3 

I 

I 

27 

.s  40 

I 

I 

I 

I 

2 

4 

b 

I 

2 

3 

I 

I 

24 

J5  44 
?  48 

3 

I 

3 

6 

2 

I 

2 

I 

19 

I 

t 

2 

4 

8  52 

I 

I 

2 

& 

3 

4 

4 

16 

12 

IS 

20 

7 

7 

4 

4 

2   0 

I   0 

I 

TOO 

Length— Mean,  149.360  ±  ,7^ 

St.  Dev.,        10.896 ±.520^ 
Coef.  Van,     7.296 ±.350 


Breadth— Mean,  38.080  ±  .356#» 

St.  Dev.,        5.288  ±.252M 
Coef.  Var.,  13.881  ±  .675 


Mean  Index  or  Ratio  of  Breadth  to  Length,  25.515  per  cent. ;  Coef.  Cor., 
.4481  ±  u}539. 


Table  LIV. 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  D,  in  a  rather 
Ill-fed  Culture.    September   15.     (See  row   13,  Table  XVIII.) 

Length  in  Microns. 


M 

§ 

U 

s 
.s 

ja 
•€ 


Q      T^OO      WVOOjS-OO      Sl^OO      ^CO      MVO      O      -^00      W 

\ov0^t^t>«coaoooo>a^OOO^M^c4M«o 

MMMMMMMMMMC4C4C4C4C4f4C4C4M 


40 

222 

I 

44 

I     113 

12    2 

48 

I 

I  I  2  2 

7  6  I  5  I  2 

52 

I  I  2  3  3 

3    2433 

50 

121 

I  2    I 

60 

I 
I 

3 
I 


4 
I 

I 


3 
2 


looi    I    37   12  8   12  95   II   556653 


7 
13 
30 
35 

13 

2 


100 


Length— Mean,  202.280  ±  1.031M         Breadth— Mean,           49.600  ±  .298;^ 

St.  Dev.,  15.284 ±    .729/^                           St.  Dev.,        4.412 ±.2io/a 

Coef.  Var.,  7556 ±   .362                           Coef.  Van,    a896 ± .428 

Mean  Ratio  of  Breadth  to  Length,  24.593  per  cent.;  Coef.  Con.  4085 
±.0562. 


1908.1 


JENNINGS— HEREDITY  IN   PROTOZOA. 


539 


Table  LV. 

Correlation  Table  for  Length  and  Breadth  of  the  Same  Lot  Shown  in 
Table  LIV,,  but  after  48  hours  in  fresh  hay  infusion,    September  15. 

(See  row  14,  Table  XVIII.) 

Length  in  Microns. 


■^  00 

CO 

VO 

»o 

%5 

% 

loioNOvo^   t^t^oooooo    o^a^p    QP   1^   ^ 

^      ^      ^      ^      k^      ^      ^      ^      k^      k^      krf      ^      M      M      M      ra      fM 

C.44 

I 

0  48 

I 

.y  52 

I  I 

II           12     1           2 

S  56 

223           I           I           I 

c  60 

II33234232           I 

*  ^ 

I 

3          III          I     I    4    I 

M  <>8 

I 

3        3   4   4    I    I    2                      I 

8  7f 

II         I         I    3   4   2    I    I 

A  76 

« 80 

I    I         I 

I 

loioi    103759  10  68  10  9  12  852    I   00    I 

Length— Mean,  175.320  ±  1.060^         Breadth— Mean,  63.160: 

St  Dev.,        15.708  ±   .749^  St.  Dev.,        7.000 : 

Coef.  Var.,      8.959  ±   -431  Coef.  Var.,  11.083: 

Mean    Ratio    of    Breadth    to    Length,    36.123    per    cent.;    Coef. 
.5376  ±  .0480. 


I 

I 

10 
10 

25 

14 
20 

15 

3 
I 

100 

:472M 
:.334/* 
:.535 
Cor., 


Table  LVI. 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  c.    August  9. 

(See  row  17,  Table  XVIII.) 


Length  in  Microns. 


fO   i*» 


ro     t>» 


«0     l*»  ro     l*» 


8«OvO      O      rovO      O      **^sO      O      coso      Q«5*0^0      «OsO      Q 


•  ••••• 

Q      «0   ^P      O      «0   so 


en 

C 

o 

u 
u 


23.3 
26.7  I 

30.    ; 
33.3 
36.7  ! 
40.   I 

43.3 
46.7 

50. 


2 
I 


2 

I 

2 


5 
I 

2 


I 
I 

3 

2 

2 


I 

3 
I 

3 
I 

I 


4 
2 

4 
2 


I 
I 

2 

4 
I 


I 
2 


I 

4 
I 

5 
2 


I 

3 

3 
I 


I   3 


I 
I 


I 
2 
I 


589  10  12  95  13  85344000   I 


9 
14 

13 
21 

22 

12 

7 
I 

I 
100 


Length — Mean,  123.666  ±  .813M  Breadth— Mean,           33-6oo  ±  .400M 

St.  Dev.,  12.040  ±  .573^                           St.  Dev.,        5.9x7  ±  .283M 

Coef.  Var.,  9.736  ±  469^                             Coef.  Var.,  17.608  ±  .865 

Mean  Ratio  of  Breadth  to  Length,  27.136  per  cent;  Coef.  Cor.,  .6528 
±  .0410. 


540 


JENNINGS— HEREDITY* IN   PROTOZOA. 


[April  94 


Table  LVII. 

Correlation  Table  for  Length  and  Breadth  of  a  Sample  of  the  Non-Conju" 
gants  of  a  Conjugating  Culture  of  Descendants  of  the  IndizHdual  c. 
Flourishing  culture  in  a  large  vessel,  September  25,  1907,  (See  row 
21,  Table  XVIII.) 

Length  in  Microns. 


w 

C 

o 


.S 


PQ 


00 

^4 

to 

% 

M 

M 

so 

vo  so  vd 

M   ►-•   M 

M 

do  06  00 

M   M   M 

8s  «s 

M   M 

§ 

16 

I 

I 

20 

I 

I 

24 

2 

I 

I 

2 

6 

28 

2 

4 

4 

I 

5 

I 

I 

I 

19 

3? 

I 

3 

2 

I 

3 

3 

3 

3 

2 

I 

I 

1 

24 

36 

I 

5 

5 

8 

5 

2 

7 

2  5 

I 

I 

42 

40 

2 

3 

2 

9 

5 

7  3 

I 

5 

4 

2 

3  I 

I 

48 

44 

2 

2 

I 

2  4 

2 

5 

I 

3 

2 

I 

25 

48 

I 

I 

2 

I 

4 

2 

2 

3  3 

2 

I 

22 

52 

I 

I 

I 

3 

I 

2 

9 

56 

I 

I 

60 

19 

6 

12  5 

2 

2 

2 

3 

10 

13 

II 

19 

14 

18 

II  15 

8 

7 

14 

7 

3 

3 

200 

Length— Mean,  158.800  ±  .877^1  Breadth— Mean,  385^  —  •353/» 

St.  Dev.,        18.384  ±  .620^  St  Dev.,        7.396  ±  .249^ 

Coef.  Van,    11.578 ±.396  Coef.  Var.,  19. 176 ±.670 

Mean  Ratio  of  Breadth  to  Length,  24.244  per  cent.;  Coef.  Cor.,  .7135 
±.0234. 

Table  LVIII. 

Correlation  Table  for  Length  and  Breadth  of  Descendants  of  c.  Five  Days 
after  Cessation  of  Conjugation.    Food  getting  scarce.    September  so, 

1907,    (See  row  22,  Table  XVIII.) 

Length  in  Microns. 
100  104  108  112  116  120  124  128  132  136  140  144  148  152 


CO 

C 

o 
u 

ii 


PQ 


20 

I 

I 

24 

I 

• 

I 

2 

28 

4 

5 

I 

3 

I 

32 

2 

3 

4 

3 

I 

2 

2 

2 

I 

36 

I 

I 

I 

2 

2 

5 

4 

I 

2 

40 

I 

I 

I 

3 

5 

10 

6 

2 

2 

44 

4 

2 

I 

48 

I 

I 

I 

I 

I 

6 

10 

7 

5 

7 

7 

10 

13 

16 

9 

6 

2 

2 

4 

14 
20 

19 

31 

7 

_3 

100 


Length — Mean,  129.640  ±  .867^1  Breadth — Mean,  35440 ±  u|00^ 

St  Dev.,        12.848  ±  .613M  St  Dev.,        5.928  ±,  .283M 

Coef.  Van,      9.9ii±477  Coef.  Van,  16.730  dr  .820 

Mean  Ratio  of  Breadth  to  Length,  27.262  per  cent;  Coef.  Con,  .7576 
±  .0287. 


«90«.] 


JENNINGS— HEREDITY   IN   PROTOZOA. 


541 


Table  LIX. 

Correlation  Table  for  Length  and  Breadth  in  a  Large,  Old  Culture  of 
Descendants  of  c,  January  23, 1908,    (Sec  row  23,  Table  XVIII.) 

Length  in  Microns. 

8^0QW\OO'*00«vOO^00«vOQ'^0QW\O 


M    M 

M   M   M 

M 

M    M 

»-• 

•H 

M 

M 

M 

H« 

M 

M 

M 

M 

M   M 

32 

2 

2 

36 

I  3 

I 

5 

40 

I 

I 

I 

I 

2 

I 

7 

2 

44 

2  2 

3 

I 

I 

9 

48 

I 

I 

I 

2 

4 

2 

7 

I 

19 

S 

52 

I 

I 

2 

I 

3 

I 

I 

10 

c 

56 

I 

I 

2 

I 

I 

5 

2 

I 

14 

60 

2 

2 

I 

I 

3 

I 

10 

•C 

•0 

CO 

64 

I 

I 

I 

I 

I 

2 

I 

8 

68 

I 

2 

3 

6 

72 

I 

2 

I 

4 

PQ 

76 
80 

84 

I 

I 

I 

2 
I 

4 
0 

2 

10014524898  13  79387461  lioo 

Length— Mean,  144.880  dt  1.097M         Breadth — Mean,         54.160  ±.   .765^ 

St  Dev.,        16.264  ±   .776^  St.  Dev.,    11.346  ±   .541M 

Coef.  Var.,    11.224  ±   .542  Coef.  Van,  20.948  ±  1.042 

Mean  Ratio  of  Breadth  to  Length,  37.106  per  cent.;  Coef.  Cor.,  .8500 
±  .0187. 


Table  LX. 

Correlation   Table  for  Length  and  Breadth  of  Descendants  of  c.     Same 

Culture  shown  in  Table  LIX,  but  cultivated  in  small  watch  glass; 

January  so  to  February  13,  1908,    (See  row  27,  Table  XVIII.) 

Length  in  Microns. 


m 

C 

%    16 

2  20 
•S  28 

I  36 

n 


y6     80  84  88  92  96  100  104  108  112  116  120 


I 

I 

2 

6 

2 
6 

3 
II 

I 
6 

3 

2 

t 

3 

I 

II 

II 

II 

I 

I 

3 

5 

I 

4 
2 

I 

I 

I 

0 

3 

6 

II 

15 

21 

19 

14 

7 

2 

I 

I 

13 
30 

39 
13 

4 

100 


Length — Mean,  100.320  ±  .528^  Breadth— Mean,  26.480  ±  .266M 

St.  Dev.,         7.828 ±.373^  St.  Dev.,        3.944^.188^ 

Coef.  Van,      7.804  ±  .374  Coef.  Van,  14.895  ±  .753 

Mean  Ratio  of  Breadth  to  Length,  26.321 ;  Coef.  Cor.,  .7671  ±  .0278. 


1 


542 


JENNINGS— HEREDITY  IN   PROTOZOA. 


[Apriis4, 


28 

32 
.36 

S  40 

P  44 


t3 

8 

PQ 


48 
52 
56 
60 

64 
68 

72 
76 
80 

84 
88 


Table  LXI. 

Correlation  Table  for  Length  and  Breadth  of  a  Random  Sample  of  the 
"Wild"  Conjugating  Culture  M,  January  29,  1908.    200  Non-con- 

s'ugants,  38  Conjugants, 

Length  in  Microns. 

«%0     O     ^00     MvO     Q     -^OQ     WVOO^OO     fix©     Q     ^OO     «vO      O     "^00      «%OQ 
«0«0'*'^^»0»0\OvO^      l*»t^CO0000      OvO^O      op      ^      HI      «      «      W      •OfO'V 

MMMHIMMHIHII-)MI-IMHIMl-IMMC«C«C4C«(iM(iC«         ««« 


I 

I 

2 

I 

2 

I 

3 

2 

2 

3 
7 

I 
4  4 

I  I 
5  I  2 

I 

I 

I 

2 

I 

5 

6  7 

5  4  I 

2  6 

2 

I 

2 

I 

I 

I 

I 

I 

3 

I 

4  5 

562 

6  7 

I 

5 

2 

I 

I 

I 

I 

I 

5  2 
I 

4  3  I 

I  3 
2 

2 

2  5 
2  4 

2 

3 

I 
2 

6 

I 
2 
2 

2 

I 

I 
I 

I 
I 

I 
I 

4 
I 

I 

3 

2 

I 

.3 
I 

I 

2 

I 

I 

I 

I 

I 

I 

I   2 

O 

I  3 

3 

7 

7 

i8 

2o  19  24  i8  9 

17  26  18 

12 

10 

8 

5 

3 

4 

0 

2001 

I 

3 
II 

31 
47 
52 

40 
18 

4 
10 

II 
2 

4 
2 
0 
2 

238 


Table  LXII. 

Correlation  Table  for  Length  and  Breadth  of  Dividing  Specimens  of  Lot  i 
(Table  X.),  in  which  Lengthening  had  begun.    (Constriction  more 

than  4  microns  deep.) 

Length  in  Microns. 


160 164 168  172  176  180  184  188  192 196  200  204  208  212  216  220  224 


Vi 

o  40 

b  44 

5  48 

'^  56 

6  60 

•S  64 

t  68 

PQ 


Length— Mean,  186.066  ±  .7iOjtt  Breadth— Mean,  49-S40 

St.  Dev.,        14.208  ±  .502A1  St.  Dev.,        4^ 

Coef .  Van,      7.636  ±  .271  Coef .  Var.,    ^^1 

Mean  Ratio  of  Breadth  to  Length,  26.796  per  cent.;  Coef.  Cor., 
±.0496. 


2 

I 

I 

I 

5 

2 

I 

3 

2 

2 

5 

2 

4 

2 

2 

3 

I 

I 

30 

2 

10 

9 

5 

7 

8 

8 

8 

4 

3 

2 

3 

2 

71 

2 

2 

10 

10 

'4 

12 

2 

2 

4 

I 

I 

I 

5; 

3 

4 

2 

I 

I 

3 

3 

I 

18 

I 

I 

3 

I 

I 

6 
0 
I 

2 

5 

12 

22 

23 

16 

22 

15 

16 

12 

9 

9 

6 

3 

6 

2 

2 

182 

.1521^ 

.309 
.0938 


.9o8.] 


JENNINGS— HEREDITY  IN   PROTOZOA; 


543 


Table  LXIII. 

Correlation  Table  for  Length  and  Breadth  of  Dividing  Specimens  of  the 
Aurelia  Form  (Descendants  of  c),  in  which  Lengthening  had  begun. 

(Sec  Lot  4,  Tables  VIII.  and  X.) 
Length  in  Microns. 


«o  t^ 


«o  t^ 


«o  t^ 


«o  t^ 


fO       tNi 


CO 

S  26.7 

5  30. 

z,  33.3 

.S  36.7 

^  40. 

8  46.7 


8«p\rj    o'    fovd    p    «osp    d    «ovd    d«o>pd    «o!vd 


I 

I 

I 

I 

I  ' 

I 

I 

2 

2 

I 

I 

112 

2 

2 

2 

4 

I 

I 

I 

I 

I 

3 

5 

2 

I 

2 

I 

I 

2 

I 

I          I 

I 

124 

3 

5 

6 

II 

4 

I 

8 

3 

I 

2    0    0    I    0    I 

3 
10 

18 

II 

9 

I 

I 
53 


Length— Mean,  121.383  dt  1.053M  Breadth— Mean,  34.590  dr  .3^3^ 

St  Dev.,        11.367^  .743M  St.  Dcv.,        4.147  ±^3Ai 

Cotf.  Var.,     9-365=*:  -613.  Coef.  Var.,  11.9^ ±.797, 

Mean  Ratio  of  Breadth  to  Length,  28.648  per  cent;  Coef.  Cor^  .3100 
±  .0837. 


544  JENNINGS— HEREDITY   IN   PROTOZOA.  [April  04, 

CONTENTS. 

Page 

I.  Introductory   393 

II.  Preliminary  Study  of  Variation  in  Paramecium 394 

1.  General  Methods  of  Work:  Statistical  Treatment  and  its  Uses.  395 

2.  A  Typical  Culture  396 

3.  Methods  of  Measuring  and  Recording 396 

4.  Method   of   Constructing   the    Polygons    399 

5.  Two  Groups  of  Paramecia   402 

6.  Are  Differences  in  Size  Hereditary  within  Each  of  the  Two 

Groups  ?    407 

7.  Proposed  Analysis  of  the  Polygons  of  Variation 409 

III.  Growth  in  Paramecium 4^1 

Effects  of  Growth  on  a  Variation  Polygon,  p.  411;  Material 
and  Methods  of  Work,  p.  414. 

1.  Description  of  the  Different  Stages  of  Growth 415 

First  Stage :  The  Young  before  Separation  is  Complete 415 

a.  The  Unseparated  Halves  before  Lengthening  has  Begun.  417 
(i)  The  caudatum  form  (descendants  of  Z>),  p.  417; 
(2)  the  aurelia  form  (descendants  of  c),  p.  422. 

h.  The  Unseparated  Halves  after  Lengthening  has  Begun.  423 
Second  Stage:  The  Young  Immediately  after  Fission,  up  to 

the  Age  of  Ninety  Minutes 426 

c.  Age  o  to  5  Minutes 427 

d.  Age  o  to  19  Minutes 432 

e.  Age  18  to  28  Minutes 434 

/.  Age  35  to  45  Minutes 435 

g.  Age  75  to  90  Minutes 435 

h.  Age  o  to  90  Minutes 435 

Third  Stage :  Three  to  Five  Hours  Old 43^ 

I.  Age  3  to  4  Hours 43^ 

y.  Age  4.20  to  5  Hours 43^ 

Fourth  Stage:  Twelve  to  Eighteen  Hours  Old 437 

k.  Age  12  Hours  437 

/.  Age  18  Hours  43^ 

Fifth  Stage:  Twenty-four  Hours  Old  438 

Sixth  Stage :  Preparing  for  Fission  439 

Seventh  Stage :  Fission  443 

m.  Beginning  Fission    , 443 

Descendants  of  D  {caudatum  form),  p.  443;  Descend- 
ants of  c  {aurelia  form),  p.  445. 
n.  Later  Stages  of  Fission  447 

2.  Summary  on  Growth  in  Paramecium,  with  Curves  of  Growth..  447 

Method  of   Constructing  the   Curves,  p.   448;   Characteristics 
of  the  Curves,  p.  451. 

3.  Effects  of  Growth  on  the  Observed  Variations  in  Dimensions.  45.^ 

Variation  at  Fission,  p.  453;  Variation  in  Conjugants,  p.  453. 


,9o8.J  JENNINGS— HEREDITY   IN   PROTOZOA.  545 

Pace 

4.  Effects  of  Growth  on  the  Observed  Correlation  between  Length 

and  Breadth 455 

IV.  The    Effects    of    Environmental    Gjnditions    on    Dimensions, 

Variation  and  Correlation  45^ 

1.  Progeny  of  D   (caudatum  form)    459 

First    Series    459 

Old  Large  Culture,  p.  459;  Effects  of  Fresh  Hay  Infusion, 
p.  459;  Scarcity  of  Food,  p.  462;  Rapid  Multiplication,  p. 
463;  Injurious  Bacteria,  p.  464;  Starvation,  p.  464. 

Second  Series  4^5 

Fresh  Hay  Infusion,  p.  465;  Starvation,  p.  465;  Effects  of 
Abundant  Food  on  a  Starving  Culture,  p.  465;  Fluid 
Unchanged  for  a  Week,  p.  469;  Forty-eight  Hours  in 
New  Culture  Fluid,  p.  469;  Resume,  p.  469. 

Third    Series    47' 

2.  Progeny  of  c  (aurelia  form) 473 

Random  Sample,  p.  473 ;  Effect  of  Adding  Boiled  Hay,  p.  473 ; 
Effect  of  Fresh  Hay  Infusion,  p.  473;  Conjugating  Culture, 
p.  475 ;  Scarcity  of  Food,  after  Conjugation,  p.  475 ;  Variation 
in  Different  Divisions  of  the  Same  Pure  Line,  on  the  Same 
Date,  p.  475;  Effects  of  Lack  of  Food,  p.  476. 

3.  Summary  on  the  Effects  of  the  Environment 47^ 

Effect  on  Length,  p.  476;  Effect  on  Breadth,  p.  477;  Relation  of 
Length  to  Nutrition,  p.  477;  Relation  of  Breadth  to  Nu- 
trition, p.  477;  Proportion  of  Breadth  to  Length,  p.  478; 
Effect  of  Environment  on  Variation,  p.  479;  Effect  of  En- 
vironment on  Correlation,  p.  481. 
V.  Inheritance  of  Size 484 

1.  Selection  for  Different  Races  in  a  Wild  Culture 4^5 

a.  Races  Isolated  from  Cultures  Not  Conjugating — Procedure 

Necessary  for  Making  the  Conditions  Identical  for  Dif- 
ferent Lines   485 

b.  Are  the  Lines  of  Different  Size  Merely  Different  Stages  in 

the  Life  Cycle? 492 

c.  Other  Evidences  of  Permanent  Differentiation  in  Size,  Inde- 

pendent of  the  Life  Cycle  497 

d.  Lines  Intermediate  between  the  two  Main  Groups.     The 

Question  of  Species  in  Paramecium 49^ 

e.  Do   the   Diverse   Lines   Differ   in   Other   Respects  besides 

Dimensions  ?    500 

2.  Results  of  Selection  within  Pure  Lines'. 503 

a.  Differences  Due  to  Environmental  Action  Not  Inherited 504 

b.  Selection  from  among  Differing  Individuals  in  the  Same 

Environment   505 

c.  Summary  on  Selection  within  Pure  Lines 511 


i 


546  JENNINGS— HEREDITY  IN  PROTOZOA.  [April  .4. 

Pack 

VI.  Summary  and  Discussion 5'' 

1.  Resum6  of  the  Investigations 5*' 

2.  Determining    Factors    for    Dimensions,    Variations    and    Cor- 

relations       5*4 

3.  Results  on  Variation,  Inheritance,  and  the  Results  of  Selection.  521 

List  op  Literature 524 

Appendix  :  Tables  of  Measurements    (Tables  XXIX.-LXIII.) 5*^ 


ON  THE  MORPHOLOGY  OF  THE  EXCRETORY  ORGANS 
OF  METAZOA:    A  CRITICAL  RE^TIEW. 

By  THOS.  H.  MONTGOMERY,  Jr. 
(Read  April  24,  1908,) 

The  desire  to  acquaint  myself  with  modem  ideas  as  to  the 
homologies  of  the  excretory  organs  has  led  to  the  present  review. 
These  organs  constitute  a  chapter  in  comparative  anatomy  that  is 
one  of  the  most  compendious  and  intricate  of  all.  Their  relations 
are  so  broad  and  manifold  that  no  morphologist  can  go  far  without 
touching  upon  them,  and  one  need  not  wonder  at  this  because  their 
function  is  above  all  others  necessary  to  the  continuance  of  the  life 
of  the  individual. 

Among  those  who  have  contributed  largely  to  this  subject  are 
Balfour,  Bergh,  Burger,  Cuenot,  Eisig,  Goodrich,  Hatschek,  Lang, 
Eduard  Meyer,  Sedgwick  and  Vejdovsky,  but  the  bibliography  shows 
how  many  well-known  investigators  have  added  to  our  knowledge. 
There  is  a  wealth  of  accumulated  facts  that  have  never  been  treated 
critically  in  their  entirety,  and  on  that  account  the  present  bringing 
together  of  them  may  be  of  help  to  future  students. 

This  memoir  is  divided  into  two  portions:  (^)  a  descriptive 
one,  in  which  the  groups  of  Metazoa  and  their  particular  excretory 
organs  are  treated  in  succession;  and  (S)  a  comparative  one,  in 
which  all  the  excretory  organs  are  reduced  to  certain  types,  and 
then  the  homologies  of  these  discussed.  It  is  in  this  second  part 
that  a  standpoint  is  reached  different,  so  far  as  I  know,  from 
previous  ones,  one  that  I  hope  puts  the  facts  in  a  clearer  light. 

A.   DESCRIPTIVE. 

The  following  is  a  brief  summary  of  our  knowledge  of  the  gen- 
eral structure  and  embryogeny  of  special  excretory  organs  con- 
sidered separately  for  each  group.      Histological  details  are  not 

547 


548  MONTGOMERY— MORPHOLOGY   OF  THE  [April  94 

entered  upon.  I  have  endeavored  to  consult  all  the  more  important 
literature  up  to  1907,  but  at  the  same  time  to  refer  in  the  citations 
to  only  the  more  comprehensive  accounts;  the  literature  references 
therefore  do  not  by  any  means  represent  complete  bibliographies, 
but  refer  the  reader  to  the  more  important  memoirs. 

The  Orthonectida,  Dicyemida,  Cnidaria  and  Fori  f  era  lack  special 
excretory  organs ;  and  such  structures  are  still  unknown  for  Cephalo- 
discus,  Rhabdopleura,  the  Qiaetosomatidae,  Desmoscolicidae,  and 
Pentastomida. 

I.  Ctenophora. 

Here  there  are  short,  presumably  entoblastic,  canals  that  connect 
the  aboral  canal  (funnel  canal)  of  the  gastro- vascular  cavity  with 
the  aboral  surface  of  the  body ;  there  may  be  two  or  four  of  these 
openings;  these  discharge  injected  carmine,  while  there  is  no  evi- 
dence that  water  is  taken  in  through  them  (Chun,  1880). 

2.  Plathelminthes. 

These  possess  branching,  tubular  organs  whose  finest  branches 
(capillaries)  have  intracellular  cavities  and  terminate  in  closed 
flame  cells,  the  latter  being  very  small  and  numerous.  Nothing  is 
known  as  to  their  embryonic  origin,  except  the  one  observation  of 
Lang  (1884)  that  in  Polyclades  a  pair  of  solid  ingrowths  of  the 
ectoblast  seems  to  represent  their  beginnings.  The  main  structural 
variations  are  with  regard  to  the  number,  ramification  and  degree 
of  anastomosis  of  the  main  canals,  and  the  number  and  position  of 
nephridiopores  and  excretory  canals. 

(i)  Turbellaria. 

Polycladidea. — Discovered  by  Max  Schultze  (1854)  these  organs 
have  received  subsequent  description  only  by  Lang  (1884),  who 
found  that  the  terminal  flames  are  unicellular  and  who  could  trace 
the  supposed  excretory  canals  of  Thysanozoon  to  the  dorsum,  but 
could  not  find  their  openings  there.  Accordingly,  a  complete 
knowledge  of  their  structure  is  still  a  desideratum.  I  have  not 
been  able  to  find  them  on  sectioned  material. 

Rhabdocoelida, — Here   they  appear  to  be   absent  only   in   the 


,9oS.l  EXCRETORY  ORGANS  OF  METAZOA.  549 

Acoela.  Three  chief  types  have  been  distinguished  (v.  Graff,  1882)  : 
a  single  main  canal  with  a  single  nephridiopore  (Stenostoma) ;  a 
pair  of  main  canals  with  independent  nephridiopores,  and  a  pair 
of  main  canals  with  a  common  nephridiopore.  In  Bothrioplafia 
(Vejdovsky,  1895)  there  are  two  pores  different  in  structure,  one 
at  the  middle  and  the  other  at  the  anterior  end  of  the  body;  into 
the  former  open  two  main  canals,  each  of  the  latter  divides  into  an 
anterior  and  a  posterior  branch,  and  these  anterior  branches  con- 
nect also  with  the  anterior  nephridiopore.  In  the  Eumesostomina 
(Luther,  1904)  the  main  canals  open  independently  either  on  the 
surface  of  the  body,  or  into  the  mouth,  or  into  the  genital  atrium 
(all  these  being  ectoblastic),  and  besides  the  terminal  flame  cells 
there  are  other  flames  (without  nuclei)  interpolated  in  the  course 
of  the  canals.  In  these  forms  there  is  never  more  than  one  pair 
of  main  canals,  or  more  than  two  nephridopores. 

Tricladidea. — In  Planaria  but  more  specially  Gunda  Lang 
(1881)  described  two  main  ducts  on  each  side  of  the  body,  each 
bearing  numerous  capillaries  ending  in  flame  cells ;  there  are  anasto- 
moses between  the  former  but  not  between  the  latter;  from  each 
main  duct  proceeds  a  series  of  excretory  ducts  each  of  which  opens 
dorsally  by  a  small  contractile  vesicle.  And  Bohmig  (1906)  adds 
to  this  account  of  Gunda  by  the  discovery  of  four  pairs  of  main 
canals,  and  of  ventral  as  well  as  dorsal  nephridiopores.  In  Dendro- 
caelum  Ijima  (1885)  found  similar  relations,  though  he  held  there 
to  be  but  one  main  canal  on  each  side;  while  Wilhelmi  (1906) 
found  two  of  them  with  a  segmental  arrangement  of  eight  pairs 
of  excretory  ducts,  and  (in  opposition  to  the  earlier  observations 
of  Chichkoff)  no  openings  into  the  pharynx.  The  Tricladidea 
differ  from  the  Rhabdocoelida  in  the  presence  of  numerous  serial 
excretory  ducts. 

(2)  Tretnatoda, 

The  chief  characteristic  of  the  excretory  organs  is  their  dendritic 
branching  and  their  degree  of  anastomosis.  In  the  Monogenea 
there  are  usually  paired  nephridiopores  (in  Gyrodactylus  a  single 
one)  placed  in  most  cases  at  the  anterior  end  but  sometimes  at  the 
posterior.     The  excretory  vesicle  of  the  Digenea  is  at  the  posterior 


550  MONTGOMERY— MORPHOLOGY  OF  THE  [AprU«4. 

end,  terminal  or  dorsal,  and  into  it  open  usually  two  but  sometimes 
four  or  even  six  main  canals  (Braun,  1893).  I^  ^^^  larva  (mira- 
cidium)  of  DistomUm  there  is  a  single  large  flame  cell  on  each  side 
of  the  body  with  a  capillary  opening  on  the  surface  (Coe,  1896). 
Bugge  (1902)  has  shown  that  each  flame  cell  and  its  capillary  "  de- 
velop out  of  one  cell  and  are  to  be  compared  with  a  unicellular 
gland,"  a  practical  confirmation  of  Lang's  (1884)  suggestion. 

(3)  TemnocephalecB, 

The  excretory  system  of  these  curious  forms  has  been  made 
known  particularly  by  Weber  (1889).  There  is  a  pair  of  sepa- 
rated dorsal  nephridiopores,  each  communicating  with  main  canals 
that  branch  and  anastomose  with  those  of  the  opposite  side,  so 
that  the  general  arrangement  is  like  that  of  the  Trematoda. 

(4)  Cestoda. 

Here  the  main  canals  have  no  dendritic  branching  but  frequent 
anastomoses,  so  that  quite  generally  each  proglottid  has  one  or  two 
pairs  of  transverse  canals  connecting  the  main  lateral  ones ;  the  main 
lateral  canals  open  by  a  common  contractile  vesicle  at  the  posterior 
end  of  the  ripest  proglottid  (Pintner,  1896).  In  the  most  detailed 
contribution  on  the  subject  (Bugge,  1902)  muscle  fibrils  of  the  main 
canals  are  described  and  also  valves  within  them  (the  latter  dis- 
covered by  Kohler  in  1894) ;  in  the  cysticercus  stage  foramina 
secondaria  were  found  connecting  the  main  canals  with  the  surface 
of  the  body.  Bugge  uses  the  term  "  Wimperflamme  "  to  include 
the  "  Terminalzelle "  ("  Geisselzelle,  Deckzelle"),  with  the 
"Wimper"  and  "Trichter"  and  "Capillare."  He  traced  such 
Wimperflammen  as  outgrowths  from  the  walls  of  the  main  canals : 
a  cell  of  the  latter  projects  outwards  then  divides  into  a  group  of 
four;  of  these  four  one  forms  three  Trichter  and  the  capillary 
(the  cavities  of  these  parts  being  intracellular),  while  each  of  the 
three  others  becomes  a  flame  cell  with  a  ciliary  flame. 

Anatomically  considered  there  are  two  main  kinds  of  excretory 
organs  in  the  Platyhelminthes :  ( i )  with  numerous  serial  excretory 
canals,  found  only  in  the  Tricladidea;  and  (2)  with  only  one  or 


X908.J  EXCRETORY  ORGANS  OF  METAZOA.  551 

two  excretory  canals,  found  in  all  the  others  (though  the  relations 
are  not  yet  known  for  the  Polyclades). 

3.  Nemertini. 

From  the  comprehensive  treatment  given  by  Biirger  (1895), 
based  largely  upon  his  own  researches,  it  follows  that  the  excretory 
organs  are  as  a  rule  in  the  form  of  two  main  canals  parallel  with 
the  lateral  blood  vessels  and  not  communicating  together;  each 
opens  to  the  exterior  of  the  body  by  one,  or  more  rarely  by  a  series 
of  several  (up  to  about  forty),  excretory  ducts;  the  main  canals  are 
usually  restricted  to  the  region  of  the  stomach,  but  in  some  genera 
they  extend  the  length  of  the  body.  From  them  proceed  delicate 
capillaries  that  terminate  blindly  in  multicellular  "  Endkolbchen  " ; 
the  latter  may  project  into  the  walls  of  blood  vessels,  but  (contrary 
to  the  earlier  opinion  of  Oudemans)  there  is  no  open  communica- 
tion of  any  portion  of  the  nephridia  with  these  vessels  or  other  in- 
ternal cavities.  In  the  ^eshwater  Stichostemma  I  showed  (1897) 
that  an  unusual  condition  obtains,  in  that  in  the  adult  instead  of  a 
single  canal  on  each  side  there  is  a  series  of  them,  some  with  and 
some  without  excretory  canals;  and  then  Bohmig  (1898)  demon- 
strated that  the  latter  are  produced  by  a  secondary  segmentation 
of  originally  continuous  ducts.^  Punnett  (1900)  and  Coe  (1906) 
found  in  Tcmiosoma  besides  excretory  pores  opening  on  the  surface 
of  the  body  others  that  connect  with  the  oesophagus ;  the  latter  are 
clearly  embryonic  ducts  persisting  in  the  adult. 

The  larvae  do  not  possess  special  excretory  organs.  The  defini- 
tive ones  arise,  according  to  Biirger,  as  a  pair  of  hollow  evaginations 
of  the  ectoblastic  stomodaeum  of  the  larva,  soon  abstrict  from  the 
oesophagus  and  then  open  into  the  amniotic  cavity  at  a  ventral  point 
near  the  mouth,  a  position  quite  different  from  that  of  the  adult 
excretory  pores.      The  origin   of  the  latter  is  not  known,  and 

*I  had  described  the  terminal  bulbs  of  this  genus  as  closed  from  the 
capillaries,  with  an  internal  cuticular  lining  but  no  flame,  while  Bohmig 
found  them  essentially  as  described  by  Burger  except  that  each  consists  of 
usually  not  more  than  two  cells.  I  have  recently  had  opportunity  to  ex- 
amine living -material  and  to  compare  it  with  my  former  sections,  and  find 
I  had  overlooked  the  true  flame  cells  and  that  Bohmig  had  described  them 
correctly.    Each  terminal  bulb  consists  of  from  one  to  five  cells. 


652  MONTGOMERY— MORPHOLOGY  OF  THE  lAprUa4. 

Burger  suggests  they  may  either  be  secondary  invaginations  of  the 
epidermis,  "  or  the  nephridium  itself  must  break  a  new  way  through 
the  body  wall.  Judging  from  the  adult  animal  the  first  alternative 
must  be  the  case." 

4.  Gastrotricha. 

There  is  a  single  pair  of  much  convoluted  tubules,  lateral  from 
the  intestine,  opening  near  each  other  on  the  ventral  surface  with- 
out excretory  vesicles.  Each  ends  internally  with  a  single  closed 
bulb,  but  it  is  not  determined  whether  it  contains  a  single  flagellum 
or  a  row  of  cilia  (Zelinka,  1889). 

5.  Rotatoria. 

The  excretory  organs  have  been  most  carefully  described  for 
the  Philodinidae  (Zelinka,  1886,  1888,  1891,  Plate,  1889),  Floscu- 
lariidae  (Montgomery,  1903,  Gast,  1900),  Melicertidae  (Hlava,  1904, 
1905),  Atrochidae  (Wierzejski,  1893)  and  Asplanchnidae  (Hudson 
and  Gosse,  1886,  Rousselet,  1891,  Weber,  1898).  There  is  always 
a  right  and  left  main  canal ;  the  flame  cells  may  be  directly  attached 
to  these  (most  Philodinidae),  but  more  usually  are  placed  at  the 
ends  of  capillaries,  branches  of  a  main  capillary  that  open  into  the 
main  canal  at  one  or  two  points.  The  number  of  flame  cells  on 
eash  side  of  the  body  is  small,  usually  from  three  to  six,  and  in 
that  case  they  are  relatively  large;  but  in  the  Asplanchnidae  there 
are  some  fifty  of  them  on  each  side  sessile  on  a  main  capillary. 
Their  great  number  here  may  be  due  to  the  large  size  of  these 
species.  The  main  canals  unite  posteriorly  into  a  short  unpaired 
duct  that  opens  into  the  cloaca ;  and  anteriorly  they  are  usually  con- 
nected by  a  transverse  commissure  (absent  in  some  Philodinidae). 
The  main  canals  have  an  intracellular  cavity,  are  composed  of  a 
few  cells  and  are  usually  without  cilia ;  terminal  flame  and  capillary 
is  a  single  cell,  the  termination  of  which  is  entirely  closed  from  the 
body  cavity  and  contains  an  internal  flame  of  cilia  and  (in  As- 
planchna)  has  a  couple  of  long  flagella  on  the  outer  surface. 

The  early  development  of  these  structures  has  not  been  de- 
termined (Zelinka,  1891). 


,9o8.|  excretory  organs  of  metazoa.  553 

6.  Endoprocta, 

Joliet  (1880)  described  for  Pedicellina  and  Loxosoma  a  pair  of 
short  ciliated  canals  with  a  common  nephridiopore,  and  with  their 
inner  ends  open  to  the  body  cavity.  Prouho  (1890)  leaves  the 
question  unsettled  whether  these  ends  are  open  or  closed.  All 
other  observers  describe  the  inner  termination  of  each  canal  as 
closed  by  a  flame  cell:  so  Foettinger  (1887)  and  Ehlers  (1890)  for 
Pedicellina,  Harmer  (1885,  Loxosoma),  and  Davenport  (1893, 
Urnatella).  The  cavity  of  these  canals  is  intracellular,  and  only  in 
Loxosoma  are  there  paired  nephridiopores. 

Besides  this  "  Chief  excretory  apparatus  "  Davenport  found  in 
the  stalk  of  Urnatella  "  elongated  spaces  terminating  blindly  at  one 
end  in  structures  which  must  be  regarded  as  flame  cells  ...  I  have 
not,  however,  been  able  in  any  instance  to  trace  an  individual  tubule 
to  any  considerable  distance,  or  until  it  opens  into  any  other  organ." 

Accordingly,  all  Endoprocta  seem  to  have  a  pair  of  nephridia 
internally  closed  that  do  not  serve  as  genital  ducts,  and  in  one 
genus  flame  cells  seem  to  occur  in  the  stalk. 

In  regard  to  their  development,  Hatschek  (1877)  found  in  the 
full-formed  larva  of  Pedicellina  a  pair  of  ciliated  canals  like  those 
of  the  adult,  but  did  not  determine  either  their  structure  or  origin. 
It  would  seem  probable  that  this  excretory  organ  persists  in  the 
adult 

7.  Rhodope. 

For  this  curious  form  that  has  been  variously  related  to  the 
Turbellaria  and  the  opisthobranch  mollusks,  Bohmig  (1893)  de- 
scribed a  nephridiopore  on  the  right  side  just  anterior  to  the  anus, 
into  which  opens  a  "  Urinkammer  " ;  into  the  latter  discharge  rami- 
fied ducts,  and  to  each  of  these  are  attached  about  forty  flame- 
bearing  terminal  organs,  each  completely  closed  from  the  body 
cavity  and  consisting  of  from  four  to  eight  cells.  Nothing  is  known 
of  the  development. 

8.   ACANTHOCEPHALA. 

The  excretory  organs  of  this  group  are  known  only  from  the 
observations  of  Kaiser  (1892,  1893).  They  occur  only  in  the 
large  Echinorhynchus  gigas  and  seem  to  be  absent  in  the  smaller 

PROC  AMER.  PHIL.  SOC.,  XLVII.  I90  JJ,  PRINTED  JANUARY  I4,  I909. 


664  MONTGOMERY— MORPHOLOGY  OF  THE  [April  94. 

species  (I  also  have  looked  for  them  in  vain  in  a  number  of  Amer- 
ican species).  In  the  female  there  is  a  pair  of  them  discharging 
into  the  oviduct;  each  is  a  broad  spade-shaped  organ  composed  of 
three  cells,  the  free  end  branched  dendritically,  each  finest  sub- 
division of  which  terminates  in  a  perforated  membrane  bearing 
on  the  luminal  side  a  tuft  of  long  cilia;  there  are  about  five  to  six 
hundred  of  these  terminal  flames  to  each  nephridium,  though  the 
whole  organ  it  will  be  recollected  is  composed  of  only  three  cells. 
The  Acanthocephala  are  specially  characterized  by  the  small  number 
but  great  degree  of  specialization  of  their  cells.  In  the  male  the 
nephridia  open  into  the  ductus  ejaculatorius,  and  are  similar  to 
those  of  the  female  save  that  the  terminal  flames  are  less  numerous. 
Kaiser  supposes  that  in  the  smaller  species  lacking  these  organs  the 
oviduct  is  excretory,  since  in  them  the  uterus  bell  is  open  to  the 
body  cavity. 

They  arise  conjointly  with  the  genital  ducts  from  the  ectoblast. 

9.  Chjetognatha. 

No  excretory  organs  were  found  by  Hertwig  (1880),  while 
Grassi  (1883)  suggests  that  a  pair  of  small  glands  opening  at  the 
junction  of  the  head  and  prepuce  may  be  urinary. 

The  genital  ducts  are  not  comparable  with  nephridia  because 
they  do  not  develop  until  maturity,  and  because  the  vasa  deferentia 
are  ectoblastic  and  the  oviducts  are  outgrowths  of  the  ovaries 
(Doncaster,  1902). 

10.    KiNORHYNCHA. 

The  genus  Echinoderes  exhibits  one  pair  of  short,  pyriform 
canals,  ciliated  throughout,  with  enlarged  closed  inner  ends,  that 
open  separately  and  dorso-laterally  (Reinhard,  1887). 

II.  Nematoda. 

As  first  made  known  by  Anton  Schneider  (1866)  and  confirmed 
by  most  subsequent  writers  there  is  usually  an  excretory  duct  in 
each  lateral  line  (though  one  may  be  wanting)  that  extend  from 
the  posterior  region  of  the  body  to  the  oesophagus,  where  they  con- 
verge and  open  by  a  single  median  nephridiopore.     The  inner  ends 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  555 

of  these  canals  are  closed,  and  each  is  lined  by  a  cuticula. '  Four 
types  of  these  have  been  distinguished  according  to  their  form  and 
position  of  the  nephridiopore  (Jagerskiold,  1898).  A  more  careful 
description  has  been  furnished  by  Goldschmidt  (1906)  for  Ascaris 
lumbricoides,  who  found  that  the  whole  apparatus  is  composed  of 
but  two  cells,  with  a  single  nucleus  for  both  main  canals  and  one 
for  the  anterior  unpaired  duct.  Goldschmidt  further  considers 
these  canals  to  be  simply  for  discharge,  and  that  a  peculiar  solid 
tissue  of  the  lateral  lines  is  the  true  secretory  portion ;  these  gland- 
ular masses  are  multinuclear  and  do  not  touch  the  walls  of  the 
canals,  but  fine  pores  appear  to  extend  towards  them  from  the 
lumina  of  the  canals.  In  his  own  words :  "  The  excretory  system 
of  Ascaris  .  .  .  consists  of  the  excretory  gland  proper  (analogous 
to  a  kidney)  that  lies  within  the  lateral  lines,  and  of  the  discharge 
duct  (analogous  to  a  ureter)  that  consists  of  two  horseshoe-shaped 
limbs  composed  of  a  single  cell  and  of  an  unpaired  terminal  portion 
represented  by  one  cell." 

Little  is  known  of  the  development  of  these  canals.  They  lie 
within  the  lateral  lines,  and  Zur  Strassen  (1892)  has  shown  that 
the  latter  are  mesoblastic.  Conte  (1902)  found  the  excretory 
apparatus  to  arise  from  a  single  mesoblast  cell  that  becomes  sec- 
ondarily placed  in  the  lateral  line. 

12.    GORDIACEA. 

For  this  group  specific  excretory  organs  are  still  unknown, 
though  it  has  received  much  study.  Vejdovsky  (1886,  1894)  has 
interpreted  the  peri-intestinal  cavity  as  excretory,  but  this  has  no 
opening  to  the  exterior ;  and  he  has  suggested  that  the  oviducts  and 
vasa  deferentia  are  modified  nephridia, — a  conclusion  drawn  from 
his  idea  that  the  Gordiacea  are  degenerate  annelids,  a  standpoint 
that  has  been  combated  by  me  (1903a).  In  late  embryonic  stages 
he  found  a  "  braune  Driise  *'  opening  into  the  intestine  near  the 
mouth ;  this  is  not  found  in  the  adult,  unless  the  problematical  supra- 
intestinal  orgon  described  by  me  for  Paragordius  may  be  an  ex- 
cretory organ  conveying  fluids  from  the  peri-intestinal  space  to  the 
intestine.      It  is  probable  that  excretion  must  take  place  through 


556  MONTGOMERY— MORPHOLOGY  OF  THE  [April  .4. 

either  the  genital  ducts  or  the  intestine,  sincie  the  thick  cuticula  on 
the  surface  of  the  body  is  hardly  permeable. 

The  gland  of  the  larva  construed  by  Villot  (1874)  as  an 
excretory  organ  has  been  considered  by  me  (1904)  to  be  rather 
a  poison  gland ;  I  have  shown  that  its  body  develops  as  an  abstriction 
of  the  entoblast,  and  that  its  duct  opens  at  the  base  of  the  pro- 
boscideal  stilets;  it  is  completely  closed  from  the  body  cavity  and 
does  not  possess  cilia.* 

13.    ECTOPROCTA. 

For  the  Phylactotemata  the  fullest  description  is  that  of  Cori 
(1893,  Cristatella) ,  according  to  whom  there  is  a  nephridium  just 
above  the  anus,  between  the  body  wall  and  the  peritoneum,  con- 
sisting of  two  ciliated  nephrostomes  opening  into  the  coelom,  con- 
necting with  an  enlarged  sac  that  has  a  single  nephridiopore  near 
the  cerebral  ganglion.  He  proved  experimentally  that  lymphocytes 
ingest  waste  particles,  and  then  are  discharged  by  this  organ. 

In  the  Gymnolaemata  there  is  in  some  species  an  organ  discov- 
ered by  Hincks  (1880),  and  more  fully  described  by  Prouho  (1892) 
who  names  it  the  "  organe  intertentaculaire  " ;  this  occurs  only  in 
sexual  individuals,  is  primarily  a  genital  duct,  and  is  a  ciliated 
canal  with  an  inner  nephrostome.  In  most  Gymnokemata  special 
excretory  organs  are  absent  (Ostroumoff,  1886,  Harmer,  1891). 
Harmer  concludes  from  injection  experiments  that  excretion  is  per- 
formed "  partly  by  the  cells  which  I  have  described  as  leucocytes, 
partly  by  the  walls  of  the  alimentary  canal,  and  partly  by  the 
funicular  tissue,"  while  he  and  Ostroumoff  have  proved  that  the 
formation  of  the  "  brown  body  "  and  the  death  of  the  polypid  is 
due  to  an  accumulation  of  waste  substances  especially  in  the 
intestine. 

The  larvae  lack  excretory  organs,  and  the  development  of  those 
of  the  adult  has  not  been  described. 

14.    SiPUNCULIDA. 

There  are  as  a  rule  two  "  excretory  tubes,"  but  within  the  same 
genus  either  two  or  one  may  occur.    In  most  cases  each  of  these  has 

•In  the  marine  Nectonema,  that  shows  some  similarity  to  the  diplobiotic 
Gordiacea,  excretory  organs  are  unknown. 


X908.]  EXCRETORY  ORGANS  OF   METAZOA.  557 

a  nephridiopore  on  the  ventral  surface  of  the  body,  and  a  ciliated 
nephrostome  placed  at  the  inner  end  of  the  tube  or  else  near  the 
external  opening.  But  in  Sternaspis  Goodrich  (1897)  found  no 
nephridiopores,  and  in  an  immature  individual  of  Phascolosoma 
proki  Sluiter  (18826)  found  no  nephrostomes.  In  all  cases  these 
serve  as  genital  ducts.  Metalnikoff  (1900),  in  the  most  detailed 
memoir,  concludes  that  the  nephrostome  cannot  serve  excretion  but 
acts  merely  to  swallow  the  germ  cells,  while  excretion  must  be 
accomplished  by  osmosis  through  the  wall  of  the  organ  that  is 
lined  by  cells  resembling  the  chloragogue  of  annelids.  Goodrich 
holds  these  are  not  true  nephridia,  but  "  peritoneal  funnels  peculiarly 
modified." 

The  embryological  data  are  conflicting.  In  Phascolosoma 
Gerould  (1906)  found  no  excretory  organs  in  the  trochophore,  and 
in  the  "  larva "  (that  succeeds  the  trochophore)  the  definitive 
nephridia  arise  as  solid  ectoblastic  ingrowths  ("  a  pair  of  ingrowths, 
probably  of  ectoderm"),  to  which  are  added  funnels  of  mesoblastic 
origin.  In  Sipunculus  Hatschek  (1883)  described  a  pair  of 
"  Nierenzellen "  in  the  mesoblast  of  the  embryo ;  each  of  these 
divides  into  four  cells  which  acquire  an  intracellular  cavity,  then 
one  end  of  each  cell  cord  becomes  attached  to  the  ectoblast  while 
the  other  opens  into  the  coelom.  Gerould's  account  is  the  much 
more  detailed  and  thorough,  and  renders  it  probable  that  both 
ectoblast  and  mesoblast  enter  into  these  nephridia.  The  trocho- 
phore lacks  nephridia. 

15.  Priapulida. 

For  these  animals  we  have  only  the  brief  description  of  Schauins- 
land  (1886),  unaccompanied  by  figures.  From  each  side  of  the 
posterior  end  there  is  said  to  invaginate  a  pair  of  ectoblastic  tubes. 
Then  a  series  of  short  excretory  tubules  grow  out  from  the  walls 
of  these ;  the  "  Endorgane  "  are  multicellular,  closed  from  the  body 
cavity,  each  cell  with  a  long  flagellum.  Still  later  other  folds 
evaginate  from  the  walls  of  the  main  ducts,  and  their  cells  become 
the  reproductive  elements.  According  to  this  description  this 
would  be  a  unique  ectoblastic  organ,  not  unlike  that  of  the  Plathel- 
minthes,  that  proliferates  germ  cells. 


658  MONTGOMERY— MORPHOLOGY  OF  THE  [April  .4, 

16.  Phoronidea. 

In  the  adult  just  behind  the  transverse  septum  Cori  (1890) 
found  a  pair  of  ciliated  canals  with  open  nephrostomes,  and  deter- 
mined that  their  function  is  both  genital  and  urinary.  In  Phoronis 
australis  Benham  (1889)  found  that  each  tube  has  two  nephro- 
stomes, and  a  similar  relation  was  discovered  by  Cowles  (1905). 

There  is  quite  general  agreement  that  the  larval  nephridia  are 
ectoblastic  (Ikeda,  1901,  Longchamps,  1902,  Shearer,  1906,  Cowles, 
1905)  ;  from  a  nephridial  pit  at  the  posterior  end  grow  out  the  two 
canals  whose  cavity  is  intercellular.  The  observers  already  cited 
together  with  Caldwell  (1882)  and  Goodrich  (1903),  in  contradic- 
tion to  Masterman  (1897),  agree  further  that  the  nephridia  of  the 
actinotrocha  are  closed  at  their  inner  ends  from  the  blastocoel  in 
which  they  lie ;  and  Shearer,  who  gives  the  most  complete  account 
of  the  development  of  these  structures,  shows  that  their  inner  ends 
are  closed  by  a  group  of  solenocytes  that  represent  outgrowths  from 
the  tubes.  Longchamps  states  that  these  larval  organs  persist  into 
the  adult ;  this  is  assumed  by  Shearer  who  decides  that  these  canals 
"  acquire  openings  into  the  coelom  by  means  of  ciliated  funnels  of 
unknown  origin  " ;  while  Ikeda  concludes :  "  We  may  assume  that 
the  formation  of  the  infraseptal  nephridial  funnels  of  the  adult  is 
due  to  secondary  outgrowths  of  the  infraseptal  portion  of  the 
atrophied,  larval  nephridial  canals."  The  only  point  not  fully 
decided  is  that  of  the  origin  of  the  funnels. 

17.  Brachiopoda. 

According  to  the  monographs  of  Van  Bemmelen  (1883),  Bloch- 
mann  (1900)  and  Morse  (1902)  there  is  usually  one  pair  of  sup- 
posed excretory  organs,  with  nephrostomes  and  nephridiopores, 
that  serve  as  genital  ducts;  in  Hemithyris  and  Rhynchonella  there 
are  two  pairs. 

Nothing  is  known  of  their  development,  and  there  appear  to  be 
no  larval  nephridia. 

18.  Echinodermata. 

Crinoidea. — Special  excretory  structures  are  unknown. 
Echinoidea, — The  axial  organ  (ovoid  gland)  has  been  consid- 


,9o8.|  EXCRETORY  ORGANS  OF  METAZOA.  569 

ered  an  excretory  organ  (Hamann,  1887,  Sarasin,  1888,  Ludwig, 
1889)  and  proved  to  be  so  by  carmine  injection  (Kowalevsky, 
1889),  while  to  it  has  also  been  ascribed  the  function  of  producing 
coelomic  cells  (Leipoldt,  1893).  '^  ^s  a  slender  axial  sac,  the  oral 
end  of  which  ends  blindly,  opening  by  a  delicate  canal  under  the 
madreporite  close  to  the  stone  canal ;  it  is  composed  of  a  meshwork 
of  trabeculae  of  connective  tissue,  covered  internally  by  an  epi- 
thelium, in  the  meshes  of  which  lie  amoeboid  cells  (Ludwig). 
Hamann  described  its  cavity  as  communicating  with  blood  lacunae 
and  the  Sarasins  as  connecting  with  the  body  cavity  by  nephro- 
stomes,  but  these  results  have  not  been  confirmed  and  the  bulk  of 
evidence  points  to  its  being  closed  from  other  body  cavities. 

Ophiuroidea, — Here  both  respiration  and  excretion  take  place 
osmotically  through  the  walls  of  the  genital  bursae  (Cuenot,  1888). 

Asieroidea, — By  injection  Kowalevsky  (1889)  found  that  the 
bodies  of  Tiedemann  are  the  excretory  organs  of  the  ambulacral 
system.  Cuenot  (1901)  distinguished  (i)  amoebocytes,  floating 
cells  in  the  coelom,  blood  vessels  and  ambulacral  system,  that  are 
first  phagocytic,  and  when  they  become  laden  with  excretory  prod- 
ucts leave  the  organism  by  passing  through  the  walls  of  the  gill 
sacs;  and  (2)  nephrocytes.  Of  the  latter  he  distinguished:  those 
that  take  up  indigo  (epithelium  of  the  intestinal  caeca),  and  those 
that  ingest  carmine  (peritoneum,  epithelia  of  perihaemal  spaces  and 
ambulacral  vessels,  inner  cells  of  septal  organs). 

Holothurioidea. — In  the  Synaptids  the  "  ciliated  funnels  "  have 
been  proved  to  collect  waste  products,  by  their  ciliary  action  and 
agglutinating  secretion ;  such  products  and  amoebocytes  loaded  with 
them  become  caught  in  these  organs,  and  ultimately  make  their  way 
through  the  solid  tissues  to  become  deposited  beneath  the  skin 
(Schultz,  1895,  Cuenot,  1902).  These  funnels  are  generally  ar- 
ranged in  rows  on  either  side  of  the  mesenteric  radix,  and  project 
into  the  coelom  either  separately  or  in  groups.  Each  is  a  some- 
what spoon-shaped,  flattened  prominence,  with  a  concave  ciliated 
surface,  attached  to  the  wall  of  the  coelom  by  a  slender  stalk,  both 
plate  and  stalk  being  composed  of  solid  connective  tissue  covered 
by  peritoneum.  Thus  they  are  really  not  funnels  at  all,  but  solid 
projections  into  the  body  cavity,  and  cannot  in  any  way  be  compared 


560  MONTGOMERY— MORPHOLOGY   OF  THE  [April  24, 

with  the  peritoneal  funnels  (peritoneal  evaginations)  of  other 
forms.  In  the  Pedata  the  respiratory  trees  have  been  considered 
as  in  part  excretory  (Schultz,  1895) ;  and  the  organs  of  Cuvier, 
tubes  that  also  open  into  the  cloaca  behind  the  preceding,  have  been 
regarded  as  excretory  by  Herouard  (1893),  ^^^  ^^  ^s  proven  that 
these  are  rather  eversible  defensive  structures  (Minchin,  1892, 
Russo,  1889). 

The  ambulacral  system  of  the  echinoderms  seems  to  mainly 
subserve  locomotion,  respiration  and  nutrition;  but  the  bodies  of 
Tiedemann,  as  mentioned  above,  that  occur  in  it  are  excretory,  and 
the  Polian  vesicle  in  holothurians  may  contain  an  "  irregular  non- 
living mass  of  brown  spherules"  which  may  be  waste  substances 
derived  from  the  brown  wandering  cells  occurring  elsewhere  in 
this  system  (Gerould,  1896). 

The  larvae  lack  nephridia,  and  there  appear  to  be  no  organs  in 
this  group  comparable  with  excretory  organs  in  others.  The  only 
representatives  of  peritoneal  funnels  are  ciliated  evaginations  from 
the  embryonic  hydrocoel  that  join  secondarily  with  ectoblastic  in- 
vaginations ;  there  is  usually  only  one  of  these  and  it  persists  as  the 
stone  canal,  but  there  may  be  two;  Field  (1892)  compared  the 
enterocoels  with  nephridia  that  have  secondarily  come  into  the 
service  of  locomotion. 

There  is  little  known  of  the  development  of  the  genital  organs 
of  Holothurioids.  In  Asteroids  they  have  been  described  as  coming 
from  a  solid  mesenchyme  mass  that  invaginates  the  peritoneum; 
only  in  Echinoids  is  the  gonad  stated  to  be  peritoneal,  a  proliferation 
of  cells  of  the  left  posterior  enterocoel.  Accordingly,  there  is  no 
evidence  that  the  gonads  or  their  ducts  stand  in  relation  to  nephridia. 

15.    TUNICATA. 

Special  organs  of  excretion  fail  in  the  Appendiculariae  (Seeliger, 
1893),  ^"^  I  have  not  found  them  described  for  the  Doliolidae.  For 
other  forms  Dahlgriin  (1901)  has  distinguished  the  following 
kinds:  (i)  Scattered  excretory  cells,  in  the  visceral  region  (in 
Botryllus,  Botrylloides,  Polycyclus,  Ciona,  Salpa) ;  (2)  vesicles, 
rather  numerous  in  the  connective  tissue,  each  with  a  wall  formed 
of  prismatic   cells   and   with   fluid   or   solid   contents    (Ascidiella, 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  561 

Ascidia) ;  (3)  sacs,  less  numerous,  on  both  sides  of  the  body  below 
the  mantle,  with  walls  of  cubical  cells  {Cynthia,  Microcosmus) ; 
and  (4)  renal  organs,  a  single  voluminous  sac  on  the  right  side  of 
the  body  with  epithelial  wall  (Molgula).  Todaro  (1902a,  b)  de- 
scribed them  for  the  Salpidse  as  hollow  vesicles  in  the  number  of 
three  pairs,  to  which  waste  products  are  carried  by  the  blood  cor- 
puscles. 

Thus  in  the  majority  of  Tunicates  they  are  vesicles  without 
ducts  placed  in  the  mesenchyme.  Van  Beneden  and  Julin  (1886) 
found  them  to  be  derived  from  mesench)mie,  and  concluded  that 
this  embryonic  tissue  is  a  modification  of  what  was  ancestrally 
enterocoelic  mesoblast;  Conklin  (1905),  however,  has  shown  that 
all  the  mesoblast  is  peristomial,  consequently  the  tissue  from  which 
these  organs  develop  may  be  mesectoblast. 

The  genital  ducts  are  outgrowth  of  the  gonads,  therefore  prob- 
ably have  no  relation  to  nephridia. 

16.    DiNOPHILEA. 

Korschelt  (1882)  described  for  Dtnophilus  apatris,  and  Weldon 
(1887)  for  D'  9^9^^,  a  nephridial  system  of  the  platyhelminthan 
type,  though  both  of  them  saw  clearly  only  the  flame  cells.  Subse- 
quent observations  have  demonstrated  that  there  are  metamerically 
arranged,  separated  nephridia.  Thus  Schimkewitsch  (1895)  found 
in  D,  vorticoides  four  pairs  of  these  in  the  male  and  five  pairs  in  the 
female;  Harmer  (1889)  and  Shearer  (1906)  for  D.  tceniatus, 
Nelson  (1907)  for  D,  conklini,  and  E.  Meyer  (1887)  for  D,  gyro- 
ciliatus  discovered  five  pairs.  These  are  ciliated  tubes  each  with  its 
own  nephridiopore,  closed  internally,  and  (according  to  Shearer) 
beset  with  solenocytes.  In  D.  conklini  the  first  pair  is  much  more 
complex  than  the  others  and  consists  of  a  considerable  number  of 
cells ;  each  of  those  of  D,  gyrociliatus  is  described  by  Meyer  as  con- 
sisting of  only  two  cells. 

Schimkewitsch  considered  the  genital  ducts  of  the  male  to  be  a 
fifth  pair,  and  the  corresponding  ducts  of  the  female  to  be  a  sixth 
pair  of  nephridia,  and  Harmer  regarded  the  seminal  vesicles  as 
segmental  organs.  This  is,  however,  little  more  than  a  supposition, 
since  the  genital  ducts  are  quite  different  in   structure  from  the 


562  MONTGOMERY— MORPHOLOGY  OF  THE  [April  24. 

nephridia  and  are  in  connection  with  the  coelom  (genital  chamber), 
and  since  the  development  of  the  nephridia  is  imknown. 

The  mid-gut  has  also  been  demonstrated  to  be  excretory 
(Schimkewitsch,  1884). 

17.    HiRUDINEA. 

Adult  Meganephridia, — There  is  a  series  of  separated  pairs,  less 
numerous  than  the  somites.  Nephrostomes  may  be  lacking  as  in 
the  case  of  five  out  of  the  seventeen  pairs  of  Hirudo  (McKim, 
1895)  the  three  most  anterior  pairs  of  Nephelis  (Graf,  1893),  ^^'^ 
all  of  Branchellion  (Bourne,  1884).  Leuckart  (1894)  discovered 
the  anatomical  connection  of  the  nephridia  with  the  nephrostomes, 
and  this  has  been  corroborated  by  Voinov  (1896),  McKim,  Graf 
and  Schultze  (1883),  in  opposition  to  the  results  of  Bolsius  (1892) 
that  the  "  organes  cilies  "  have  no  connection  with  the  loop.  But 
even  when  they  are  connected  there  need  not  be  an  open  communi- 
cation between  the  two  (Graf,  1899).  When  present  the  funnel 
lies  in  the  segment  preceding  that  of  the  loop.  The  cavity  of  the 
nephridia  is  much  branched  and  intracellular.  An  excretory  bladder 
may  be  present  as  a  part  of  the  excretory  duct,  but  this  is  lacking 
in  Clepsine, 

The  nephridia  arise  from  segmentally  arranged  mesoblastic 
nephroblasts,  that  lie  deep  below  the  embryonic  epidermis.  Each 
of  these  divides  into  two  cells,  the  anterior  of  which  gives  rise  to 
the  funnel  and  the  posterior  to  a  cord  of  cells  that  forms  the  secre- 
tory portion  of  the  loop;  the  cavity  into  which  the  nephrostomes 
open  is  a  true  coelom;  the  excretory  ducts  and  vesicles  are  ecto- 
blastic  ingrowths  (Biirger,  1891,  1894,  1902,  Bergh,  1891,  McKim, 
189s).  Biirger  is  very  positive  with  regard  to  the  mesoblastic 
origin  of  the  nephridia,  in  opposition  to  the  earlier  view  of  Whit- 
man (1887). 

Adult  Plectonephridia, — Bourne  (1884)  first  found  net-like 
nephridia  in  Branchellion,  Pontobdella  and  Piscicola;  in  Pontobdella 
they  consist  of  a  network  of  canals  extending  from  the  ninth  to  the 
nineteenth  s^ment,  with  ten  pairs  of  nephridiopores,  while  in 
Branchellion  they  have  only  one  pair  of  such  openings.  They  have 
been  redescribed  by  Johansson  (1898),  and  I  am  acquainted  with 


«9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  563 

his  account  only  from  the  citation  given  by  Lang  (1903,  p.  103). 
^'  In  Pontobdella  the  nephridia  consist  of  very  richly  branched  and 
reticularly  anastomosing  tubes,  among  which  one  cannot  distinguish 
main  trunks.  The  two  nephridia  of  the  same  segment  are  many 
times  joined  together,  and  the  nephridia  of  the  several  segments 
equally  so.  In  Cystobranchus  each  nephridium  has  attained  a  com- 
plete independence  and  connects  neither  with  the  other  nephridia 
of  the  same  segment,  nor  with  those  of  neighboring  segments.  It 
consists  then  also  only  of  a  single,  coarse,  unbranched  tube.  The 
remaining  genera  correspond  in  this  relation  more  or  less  with 
Pontobdella;  one  can,  however,  always  distinguish  particular 
trunks.  In  Piscicola  one  part  of  the  nephridium,  that  is  much  more 
strongly  developed  than  the  remaining  part,  corresponds  exactly 
in  position  with  the  nephridium  of  Cystobranchus,  Pontobdella 
departs,  finally,  from  all  the  other  genera  in  this,  that  the  nephridia 
have  inner  openings."  Nothing  is  yet  known  of  the  development 
of  these  reticular  organs. 

Genital  Ducts. — These  were  considered  by  Nusbaum  (1885)  ^o 
be  modified  nephridia.  Burger  first  (1894)  opposed  this  com- 
parison, but  later  (1902)  he  maintained  that  the  female  genital 
apparatus  and  the  terminal  portions  at  least  of  the  vasa  deferentia 
are  possibly  homologous  with  nephridia  in  developing  from  gono- 
blasts  that  are  homodynamous  with  nephroblasts. 

Larval  Nephridia. — In  the  Hirudinea  three  of  the  blastomeres 
of  the  4-cell  stage  give  rise  to  a  larval  body  that  later  perishes, 
while  the  fourth  blastomere  alone  produces  the  adult  body 
(Brandes,  1901).  This  larval  body  produces  no  nephridia.  The 
"  Umieren  "  arise  from  the  germ  band  that  develops  within  this 
larval  body,  and  they  last  only  as  long  as  the  latter  does.  Bergh 
(1884,  1901)  has  shown  that  there  are  three  pairs  of  these  in 
Aulastoma  and  Hirudo  and  two  pairs  in  Nephelis,  all  developing 
from  the  germ  band;  and  he  and  Sukatchoff  (1900)  demonstrated 
that  the  inner  ends  are  closed  and  the  cavity  intracellular.  These 
larval  nephridia  arise  from  cell  rows  of  the  germ  band  that  are 
generally  considered  mesoblastic,  though  this  point  is  hardly  finally 
settled. 

Excretophores. — Excretory   cells   within   the   connective   tissue 


564  MONTGOMERY— MORPHOLOGY   OF  THE  [April  34. 

(Graf,  1899),  that  develop  from  the  splanchnic  layer  of  the  meso- 
blast  (Burger,  1902). 

Chloragogue  (Botryotdal  Tissue). — Excretory  cells  placed  upon 
the  blood  vessels  (Graf,  1893). 

18.   OLIGOCHiETA. 

Adult  Nephridia, — There  are  two  main  kinds  of  these  which 
it  will  be  convenient  to  consider  separately:  meganephridia,  larger 
and  in  separated  pairs;  and  plectonephridia,  networks  of  smaller 
nephridia. 

Meganephridia, — Of  these  there  is  usually  one  pair  to  each  trunk 
segment,  though  exceptions  are  very  numerous ;  each  has  a  preseptal 
open  funnel  and  a  postseptal  loop  with  intracellular  cavity;  their 
nephridiopores  are  usually  separated  and  placed  latero-ventral.  The 
smallest  number  known  is  two  pairs  (Bdellodrilus,  Moore,  1897). 
In  Brachydrilus  there  are  two  pairs  to  each  somite  (Benham,  1888). 
The  anterior  five  pairs  open  into  the  pharynx  in  Dichogaster  (Bed- 
dard,  1888&),  and  probably  also  in  Eminea  (according  to  Benham, 
1890&,  who  terms  this  a  "peptonephridium  ").  In  Limnodrilus 
the  two  anterior  pairs  perforate  septa  while  the  others  do  not 
(Rybka,  1899).  Libyodrilus  is  characterized  by  the  nephridia 
opening  into  a  tubular  system  situated  in  the  musculature,  consisting 
of  four  main  longitudinal  vessels  extending  from  segment  to  seg- 
ment and  of  segmental  ring  vessels,  there  being  numerous  excretory 
ducts  from  the  latter;  this  integumental  network  is  secondary  and 
develops  after  hatching  (Beddard,  1891).  Numerous  other  devia- 
tions from  the  general  type  are  known  that  it  is  not  necessary  to 
mention  here,  beyond  the  fact  that  nephrostomes  are  lacking  in  the 
Chaetogastrids  (Vejdovsky,  1885). 

Plectonephridia, — A  plectonephridium  is  a  complex  that  in  each 
segment  is  composed  of  numerous  micronephridia,  without  nephro- 
stomes, that  are  joined  by  a  network  of  canals.  In  Acanthodrilus 
there  is  one  such  micronephridium  to  each  of  the  eight  setae  of  each 
posterior  segment,  and  in  each  anterior  segment  there  are  about  one 
hundred  nephridiopores;  somewhat  similar  relations  obtain  in 
Typhceus  (Beddard,  1888a).  In  Megascolides  there  are  a  great 
number  of  bundles  of  micronephridia  which  clothe  the  body  wall 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  566 

except  medially,  these  opening  into  a  network  of  intracellular  ducts 
placed  outside  of  the  peritoneum,  and  the  latter  discharge  at  the 
surface  by  irregularly  arranged  canals  (Spencer,  1889).  In 
Mahbenus  each  micronephridium  has  its  own  excretory  duct 
(Bourne,  1894).  The  network  of  fine  canals  may  be  continuous 
from  segment  to  segment,  as  in  Perichcsta,  or  only  the  micro- 
nephridia  of  one  and  the  same  segment  may  be  so  connected  as 
exemplified  by  Deinodrilus,  Acanthodrilus,  and  Dichogaster  (Bed- 
dard,  18886). 

Both  of  these  kinds  of  nephridia  may  occur  in  the  same  animal 
and  even  in  the  same  segment,  as  in  Megascolides;  and  in  this  genus 
there  is  a  pair  of  ventral  longitudinal  canals  continuous  from  seg- 
ment to  segment  into  which  both  open  (Spencer).  In  Dichogaster 
the  posterior  segments  contain  both  kinds  (Beddard,  18886). 

Development  of  the  Meganephridia. — ^With  great  hesitation  I 
attempt  to  give  a  brief  review  of  this  subject,  that  has  proved  the 
Austerlitz  of  many  a  theory.  Kowalevsky  (1871)  was  the  first 
to  demonstrate  the  mesoblastic  origin  of  these  organs  in  Euaxes 
(Rhynchelmis)  and  Lumbricus.  Vejdovsky  and  Bergh  have  fur- 
nished more  observations  on  the  subject  than  any  other  writers. 
Vejdovsky's  results  (1885,  1892a,  1900)  on  Rhynchelmis,  Stylaria 
and  Tubifex  are  as  follows:  Each  nephridium  arises  from  three 
separated  anlages:  (i)  A  large  preseptal  funnel  cell,  giving  rise  to 
the  nephrostome;  (2)  a  cord  of  small  cells  budded  off  behind  the 
former,  producing  the  secretory  loop;  and  (3)  an  ectoblastic  in- 
vagination that  joins  with  the  latter  and  forms  the  distal  canal 
and  the  excretory  vesicle.  Bergh's  studies  (1888,  1890,  1899)  on 
Lumbricus,  Criodrilus  and  Rhynchelmis  differ  from  those  of 
Vejdovsky  mainly  in  deriving  each  nephridium  from  a  single  meso- 
blastic anlage  instead  of  from  three  parts ;  in  his  mind  the  organ  is 
essentially  an  embryonic  unit.  Wilson  (1889)  concluded  for 
Allolobophora  that  the  funnel  arises  from  a  large  mesoblstst  cell, 
and  the  loop  from  a  postseptal  mass  of  cells  that  is  continuous  with 
the  ectoblastic  nephridial  cell  cords,  though  he  admits  the  loop  may 
nevertheless  be  mesoblastic.  And  Lehman  (1887,  Allolobophora) 
derived  the  nephridium  from  a  large  preseptal  cell. 

These  researches  agree  in  finding  that  the  nephridia  arise  seg- 


566  MONTGOMERY— MORPHOLOGY  OF  THE  [April  .4, 

mentally,  to  which  the  conclusions  of  Roule  (1889)  alone  are 
opposed,  and  that  their  first  beginning  is  the  preseptal  funnel  cell. 
But  there  is  considerable  conflict  of  opinion  as  to  what  germ  layer 
produces  these  cells  and  the  cords  that  arise  behind  them.  They 
arise  in  that  cell  row  of  the  germ  band  formed  by  proliferation  of 
the  posterior  nephroblasts.  The  germ  band  is  covered  by  a  thin 
ectoblast,  and  the  funnel  cells  lie  at  points  where  the  mesoblastic 
dissepiments  meet  the  ectoblast;  they  are  blastocoelic  in  position. 
Bergh  is  positive  that  funnel  cells  and  nephridial  cords  are  meso- 
blastic, derived  from  what  he  terms  the  "innere  Muskelplatten," 
and  Lehmann  and  Roule  express  the  same  opinion.  Wilson  hesi- 
tates to  decide  whether  the  nephridial  cords  are  ectoblastic,  though 
he  ascribes  this  origin  to  the  funnel  cells.  Vejdovsky  considers 
that  at  this  early  stage  of  the  embryo,  when  these  parts  are  first 
definable,  there  is  no  mesoblast  but  only  the  two  primary  germ 
layers  and  that  the  funnel  cells  may  have  emigrated  from  the 
ectoblast.  It  is  to  be  noted  in  this  connection  that  the  funnel  cells 
when  they  are  first  distinguishable  have  never  been  seen  actually 
in  the  ectoblast,  but  always  beneath  it.  And  the  nephridial  develop- 
ment is  so  correspondent  with  that  of  the  Hirudinea,  for  which 
Burger  shows  so  convincingly  that  the  nephridia  are  mesoblastic, 
that  the  view  of  Bergh  would  seem  to  be  correct.  Consequently 
Goodrich  (1895)  in  his  summary  of  the  literature  on  this  subject 
would  seem  to  have  misunderstood  the  facts  of  the  case.  We  may 
at  least  conclude,  that  in  light  of  the  evidence  at  hand  all  the  inner 
portion  of  the  nephridium  is  mesoblastic,  and  only  its  distal  outer 
termination  comes  from  the  ectoblast. 

Remarkable  postembryonic  changes  have  been  described  by  Rosa 
( 1903a)  for  Lumbricus.  In  a  newly  hatched  individual  two  canals 
extend  through  the  whole  trunk  and  join  posteriorly  into  an  ampulla 
that  opens  dorsally  into  the  intestine  (for  which  reason  the  describer 
compares  it  with  the  nephridia  of  Rotatoria).  From  each  of  these 
canals  tubes  branch  off  segmentally  and  connect  with  the  nephridia 
of  the  corresponding  segments,  while  the  nephridia  still  lack 
nephridiopores ;  later  in  each  segment  a  diverticulum  grows  out 
from  each  canal  and  opens  on  the  surface  in  the  position  wherein 
the  adult  the  nephridiopore  lies,  while  in  each  segment  the  main 


,9o8.]  EXCRETORY   ORGANS  OF  METAZOA.  567 

canals  swell  into  a  pair  of  vesicles;  in  the  adult  these  longitudinal 
canals  have  disappeared,  probably  by  segmenting  into  segmental 
excretory  vesicles  and  nephridiopores. 

Development  of  the  Plectonephridia. — In  Megascolides  each  seg- 
ment has  one  pair  of  nephridial  anlages,  each  consisting  of  a 
preseptal  cell  and  a  postseptal  cord;  so  far  the  development  is  like 
that  of  the  meganephridia ;  then  the  postseptal  cord  originates  many 
loops  and  by  a  rupture  of  their  connecting  bridges  the  micro- 
nephridia  result;  the  longitudinal  canals  connecting  the  latter  arise 
later  and  are  therefore  secondary  (Vejdovsky,  18926).  In  Mah- 
benus  Bourne  (1894)  described  an  essentially  similar  process:  that 
the  funnels  degenerate,  that  the  loops  form  secondary  and  the 
latter  tertiary  branches,  until  each  segment  comes  to  contain  about 
fifty  micronephridia.  These  observations  indicate  clearly  that  the 
plectonephric  condition  is  a  modification  of  the  primary  macro- 
nephric  by  a  subdivision  of  originally  single  organs.  This  is  the 
position  taken  by  Vejdovsky,  Bourne  and  Beddard  (1892)  which 
is  contrary  to  the  hypothesis  of  Benham  (1890,  1891a),  Spencer 
(1889)  and  Beddard  (1891)  that  the  plectonephric  condition  is 
primitive  and  comparable  with  that  of  the  Plathelminths.  Micro- 
nephridia lack  nephrostomes  because  they  are  division  products  of 
the  loops  only,  and  not  of  the  funnels.  Therefore  Vejdovsky  is 
probably  correct  in  his  conclusion  that  the  micronephridia  are 
homologous  with  the  meganephridia,  because  both  arise  from  a 
common  anlage,  comparable  with  the  embryonic  pronephridium 
of  Rhynchelmis, 

In  Acanthodrilus  deverticula  grow  out  from  the  intestine,  at  a 
region  probably  anterior  to  the  proctodaeum,  and  join  with  the 
plectonephridia  of  that  region  of  the  body;  this  connection  is  sec- 
ondary (Beddard,  1889,  1890,  1892). 

Embryonic  Nephridia, — For  Rhynchelmis  three  sets  of  em- 
bryonic excretory  structures  have  been  found  by  Vejdovsky 
(1892a).  These  are  (i)  "  Schluckzellen,"  cleavage  cells  containing 
canals,  which  had  been  previously  considered  to  digest  the  albumen 
of  the  tgg;  (2)  larval  pronephridia,  "  Kopfnieren  "  placed  between 
the  germ  band  and  the  ectoblast;  and  (3)  embryonic  nephridia, 
which  later  change  into  the  definitive  nephridia.      Bergh   (1888) 


568  MONTGOMERY— MORPHOLOGY  OF  THE  [April  a4. 

found  in  Criodrilus  a  pair  of  tubes  closed  internally  that  he  called 
Urnieren,  though  on  account  of  the  lateness  of  their  origin  Vejdovsky 
considered  they  are  rather  embryonic  nephridia.  Wilson  (1889)  de- 
scribed for  Allolobophora  a  pair  of  head  kidneys,  and  Hoffmann 
(1899)  found  these  opened  into  the  head  cavity.  In  the  opinion  of 
Vejdovsky  the  larval  nephridia  develop  either  from  the  Schluck- 
zellen,  or  else  come  from  mesenchyme  of  ectoblastic  origin.  But 
it  is  yet  by  no  means  decided  from  what  germ  layer  these  kidneys 
originate. 

Genital  Ducts, — It  was  Williams  (1858)  who  first  indicated  the 
homology  of  the  genital  ducts  with  nephridia,  and  he  held  the 
excretory  function  to  be  secondary.  Claparede  pointed  to  the 
typical  absence  of  nephridia  in  the  genital  segments  as  evidence  that 
the  genital  ducts  are  modified  nephridia.  Then  Lankester  (1865), 
reasoning  from  the  condition  in  the  Lumbricids,  suggested  that 
genital  ducts  represent  the  sole  traces  of  a  ventral  set  of  nephridia 
that  must  originally  have  existed  together  with  the  dorsal  set  in  all 
the  segments ;  according  to  this  view  the  primitive  relation  would  be 
two  pairs  of  these  organs  to  each  segment.  This  idea  was  adopted 
by  Benham  (i886a^  b)  who  maintained  that  in  Lumbricus,  Titanus 
and  Pontodrilus  the  ventral  series  of  nephridia  disappears  except 
those  that  change  into  genital  organs,  and  that  in  Rhinodrilus, 
Eudrilus,  Anteus,  Urochceta  and  Moniligaster  just  the  opposite  con- 
dition obtains.  But  Balfour  (1885),  as  most  students  after  him 
concluded  that  one  pair  of  nephridia  to  a  segment  is  primitive,  and 
that  "  in  the  generative  segments  of  the  Oligochaeta  the  excretory 
organs  had  at  first  both  an  excretory  and  a  generative  function,  and 
that,  as  a  secondary  result  of  this  double  function,  each  of  them 
has  become  split  into  two  parts,  a  generative  and  an  excretory." 
Here  it  is  to  be  recalled  that  two  pairs  of  nephridia  to  a  segment  is 
unusual,  and  that  only  in  the  Lumbricidae  do  both  genital  ducts  and 
nephridia  occur  in  the  same  segment ;  anatomical  relations  therefore 
do  not  bear  out  Lankester*s  theory.  With  regard  to  the  embryogeny 
of  the  genital  ducts,  Vejdovsky  (1885)  found  them  to  arise  inde- 
pendently of  the  nephridia,  though  he  considered  they  might  be 
wholly  or  in  part  homodynamous  with  the  latter ;  at  least  the  funnels 


X908.]  EXCRETORY  ORGANS  OF  METAZOA.  569 

of  the  two  he  considered  to  have  this  relation.  Similar  results  were 
reached  by  Bergh  (1886),  Roule  (1889)  and  Beddard  (1892). 
Lehmann  (1887)  opposed  the  idea  of  homodynamy  on  the  grounds: 
(i)  That  two  pairs  of  nephridia  to  a  s^ment  is  not  typical;  (2) 
that  in  the  embryo  nephridia  develop  in  the  genital  segments; 
and  (3)  that  the  genital  ducts  arise  later  than  the  nephridia.  Finally 
there  may  be  mentioned  the  view  of  Benham  (1904)  according  to 
whom  the  phylogenetic  series  is  as  follows:  (i)  The  nephridia 
acted  as  genital  ducts;  then  (2)  a  special  coelomostome  became 
added  to  the  nephridia,  forming  a  nephromixium ;  finally  (3)  the 
coelomostome  formed  "  its  own  coelomo-duct,  which  may  either  co- 
exist in  the  genital  segment  with  the  nephridium  (as  in  most '  terri- 
coline'  Oligochaetes),  or  the  nephridium  .  .  .  disappears  from  the 
segment  during  or  before  the  development  of  the  genital  duct  (as 
in  '  limicoline '  Oligochaetes  and  Protodrilus).  We  have,  then,  to 
some  extent  a  parallel  series  of  phenomena  analogous  to  those  de- 
scribed with  so  much  care  by  Goodrich  in  the  Polychaeta." 

There  is  much  in  these  relations  that  is  still  puzzling.  But  at 
least  the  funnels  of  both  organs  seem  to  be  homodynamous  since 
they  have  an  approximately  similar  mode  of  growth.  In  the 
Lumbricids  the  two  organs  of  a  genital  segment  might  well  have 
arisen,  as  Balfour  intimated,  as  division  products  of  a  common 
embryonic  anlage.  And  in  those  species  where  nephridia  are  want- 
ing in  the  genital  segments,  the  genital  ducts,  as  Vejdovsky  argued, 
are  to  be  considered  as  in  part  at  least  modifications  of  the  nephridia 
of  such  segments. 

Chloragogue  {Pericardial  Gland), — This  is  peritoneal  in  origin 
and  particularly  excretory  (Grobben,  1888,  Rice,  1902,  Rosa,  1903a). 

Peritoneum  and  Coelomic  Fluid, — These  have  been  considered 
excretory  by  Grobben  (1888),  who  holds  that  the  coelomic  fluid  is 
in  great  part  an  excretory  product  though  at  the  same  time  it  has 
the  functions  of  blood  and  lymph. 

Other  Excretory  Organs, — Here  are  to  be  reckoned  the  bacter- 
oidic  cells  of  the  connective  tissues,  the  yellow  cells  of  the  intestine, 
and  the  amoebocytes  of  the  blood  (Cuenot,  1897). 

PROC.  AMER.  PHIL.  SOC.,  XLVII.   I90  KK,  PRINTED  JANUARY  I4,  I909. 


570  MONTGOMERY— MORPHOLOGY  OF  THE  [April  24, 

19.    POLYCHJETA. 

Adult  Nephridia. — There  is  usually  one  pair  to  each  trunk  seg- 
ment. In  the  Phyllodocidae,  Glyceridae  and  Nephthyidae  their  inner 
ends  are  closed  and  the  loops  are  beset  with  solenocytes,  each  of 
which  is  a  cell  projecting  into  the  body  cavity  "  containing  a  deeply- 
staining  rounded  or  oval  nucleus,  attached  by  a  sort  of  neck  to  the 
extremity  of  a  thin  tube  which  opens  at  its  opposite  end  into  the 
lumen  of  the  nephridial  canal  .  .  .  Working  inside  the  tube  and 
attached  at  its  distal  end  is  a  single  long  flagellum,  which  passes  far 
down  the  nephridial  canal  "  (Goodrich,  1900).  In  the  other  families 
the  inner  end  is  open  to  the  coelom,  with  the  exception  of  Poly- 
gordius  (Hempelmann,  1906)  where  the  first  pair  is  closed.  The 
nephridiopores  usually  open  separately.  Each  pair  of  nephridia 
stands  in  relation  to  two  segments  in  Archiannelids,  Alciopidae, 
Typhloscolecidae,  certain  Nereids  (Eisig,  1887),  Terebelloids  and 
Girratulidae  (Meyer,  1887),  Aphroditidae  (Darboux,  1900)  and 
Disomidae  (Allen,  1904)  ;  in  the  other  families,  therefore  in  the 
majority  of  species,  to  only  a  single  segment.  Some  of  the  main 
deviations  from  this  tjrpe  are  the  following : 

(a)  In  Capitellids  each  nephridium  may  have  several  nephro- 
stomes,  there  may  be  several  pairs  to  a  s^ment  and  they  may  dis- 
charge into  the  skin  and  not  on  the  surface  of  the  latter  (Eisig, 
1887).  In  Lattice  and  Ploimia  the  fourth  segment  possesses  two 
pairs  (Meyer,  1887). 

(&)  In  the  Terebelloid  Lanice  conchilega  the  three  anterior  pairs 
of  nephridia  connect  with  a  pair  of  longitudinal  canals  from  each 
of  which  a  single  nephridiopore  discharges  on  the  surface;  while 
the  four  following  pairs  of  nephridia  open  into  a  longer  pair  of 
posterior  canals  which  end  blindly  at  about  the  sixteenth  thoracal 
s^ment,  and  each  of 'which  discharges  by  four  nephridiopores. 
Ploimia  presents  quite  similar  relations.  Meyer  (1887)  who  de- 
scribed these  conditions  holds  it  probable  that  the  longitudinal  canals 
are  formed  secondarily  by  a  meeting  and  fusion  of  separate 
nephridial  loops,  incipient  stages  of  which  are  to  be  noted  in  other 
genera.  Also  in  Owenia  (Gilson,  1894)  do  the  nephridia  open  into 
longitudinal  canals,  that  are  here  described  as  formed  by  an  infold- 
ing of  the  epidermis. 


i9o8.]  EXCRETORY   ORGANS  OF  METAZOA.  571 

(c)  In  the  Terebelloids  an  impervious  dissepiment  separates  the 
anterior  from  the  posterior  thoracal  cavity ;  in  the  former  there  are 
no  germ  cells,  and  the  three  pairs  of  nephridia  have  small  funnels ; 
in  the  posterior  space,  which  communicates  with  the  abdominal 
coelom,  occur  germ  cells,  and  there  the  nephridia  have  large  nephro- 
stomes  (peritoneal  funnels)  for  the  discharge  of  these  cells.  In 
the  Cirratulids,  Serpulacea  and  Hermellids  only  the  first  pair  of 
nephridia  are  strictly  excretory,  and  the  others  serve  as  genital 
ducts  (Meyer,  1887). 

(d)  In  Hermellids  and  Serpulacea  the  pair  of  thoracal  nephridia 
unite  dorsally  into  an  unpaired  duct  that  opens  near  the  anterior 
end  of  the  trunk  (Meyer,  1887).  And  in  Dybozvscella  the  pair  of 
the  "  head  "  has  a  single  medio-dorsal  pore  (Nusbaum,  1901). 

Development  of  the  Definitive  Nephridia. — The  nephrostome  of 
Polymnia  (Meyer,  1887)  arises  as  a  fold  of  the  peritoneum  that 
grows  backward  to  join  the  loop;  the  latter  developes  independently, 
simultaneously  or  a  little  later,  from  retroperitoneal  tissue  (whether 
mesectoblastic  or  mesentoblastic  was  not  determined)  that  is  at  first 
solid  and  later  acquires  a  cavity ;  the  distal  excretory  duct  is  prob- 
ably ectoblastic.  In  Psygmobranchus  (Meyer,  1888)  there  first 
appears  in  the  unsegmented  larva  a  pair  of  large  cells  in  the 
blastocoel,  apposed  to  the  ectoblast  and  separated  from  the  meso- 
blast,  these  two  cells  become  placed  between  the  two  layers  of  the 
first  dissepiment  and  give  rise  to  the  tubes,  while  there  evaginates 
to  meet  each  of  them  a  peritoneal  funnel.  Meyer  holds  that  all  the 
funnels  of  Terebelloids  must  have  originally  been  parts  of  dissepi- 
ments, and  with  the  degeneration  of  the  latter  have  either  become 
independent  organs  or  else  have  become  grafted  upon  nephridia.  In 
what  is  the  most  detailed  account  of  any  polychaetous  nephridium, 
Lillie  (1905)  finds  for  Arenicola  that  the  nephridia  arise  seg- 
mentally  and  independently,  entirely  from  the  somatic  layer  of  the 
mesoblast;  at  first  they  are  small  tubes  with  intracellular  cavities 
and  a  minute  opening  into  the  coelom ;  "  the  anterior  region  of  these 
organs  .  .  .  together  with  a  portion  of  the  adjoining  septum,  con- 
stitutes the  primitive  nephrostome,  from  which  the  adult  nephro- 
stome is  directly  derived."  The  terminal  vesicle  is  also  not  ecto- 
blastic, but  "  is  formed  as  a  differentiation  of  the  most  posterior 


572  MONTGOMERY— MORPHOLOGY   OF  THE  [April  i^, 

portion  of  the  primitive  nephridium.  There  is  no  ectodermal  in- 
vagination," but  the  terminal  portion  comes  from  a  region  where 
mesoblast  and  ectoblast  join,  probably  from  a  region  that  was  orig- 
inally ectoblastic. 

The  work  of  Meyer,  Fraipont  and  Woltereck  shows  that  Hat- 
schek  (1878)  was  entirely  wrong  in  deriving  the  nephridia  from  a 
continuous  anlage,  and  in  stating  the  adult  nephridia  of  Polygordius 
arise  as  branches  of  longitudinal  ducts  of  larval  nephridia. 

Larval  Nephridia, — There  is  one  pair  of  these  in  Polymnia 
(Meyer,  1887),  each  with  a  long  flagellum  placed  upon  the  outer 
surface  of  the  closed  inner  end,  on  which  region  follows  a  loop  com- 
posed of  two  cells  and  then  an  excretory  canal  with  intercellular 
cavity ;  these  persist  until  the  first  definitive  nephridia  function.  In 
Psygmobranchus  (Meyer,  1888)  there  is  also  one  pair,  each  com- 
posed of  two  cells  and  probably  without  internal  opening,  that  open 
on  the  ectoblast  and  do  not  touch  the  mesoblast ;  they  belong  to  the 
first  somite  (that  just  behind  the  hietastomium).  Meyer  (1887) 
has  figured  the  larval  nephridia  of  Nereis  as  internally  closed 
canals;  Hatschek  (1885)  finds  this  structural  relation  in  Eupomatus, 
and  holds  the  nephridia  to  be  mesoblastic.  In  Hydroides  the  head 
kidney  opens  into  the  proctodaeum  (Wilson,  1890).  Drasche 
(1884,  Pomatoceros)  held  the  head  kidneys  to  have  funnels,  and  to 
be  mesoblastic. 

The  larva  about  which  there  has  been  the  most  discussion  is  that 
of  Polygordius.  For  F.  neapolitanus  Hatschek  (1878)  described 
the  branched  head  kidney  as  having  open  nephrostomes  and  being 
joined  by  longitudinal  canals  with  the  trunk  nephridia,  a  condition 
that  has  led  to  manifold  comparisons  with  platodan  relations.  But 
Fraipont  (1888)  and  Meyer  (1901)  found  that  such  longitudinal 
canals  do  not  exist,  and  that  tlje  inner  ends  of  these  tubes  do  not 
possess  funnels  but  are  beset  with  slender  cells  (solenocytes)  that 
project  into  the  blastocoel.  Meyer  described  also  a  second  pair 
of  larval  nephridia  behind  these,  which  differ  from  trunk  nephridia 
only  in  the  lack  of  funnels.  Then  Woltereck  (1905)  in  disagree- 
ment with  these  writers  states  that  the  two-branched  first  pair  of 
larval  nephridia  belong  to  the  second  somite,  are  mesenchymatous 
and  degenerate  entirely;  while  the  second  larval  nephridium  belongs 


,9oi.J  EXCRETORY  ORGANS  OF  METAZOA.  573 

to  the  third  somite  and  consists  of  two  parts :  ( i )  A  mesenchymatous 
portion,  composed  of  two  "  Kopfchenzellen "  beset  with  ciliated 
tubes,  that  later  degenerates,  and  (2)  a  segmental  portion,  at  least 
in  part  ectoblastic  in  origin,  that  joins  with  the  mesenchymatous 
part.  Woltereck  finds  this  second  pair  to  become  the  first  pair  of 
definitive  nephridia  that  differs  from  the  others  in  the  absence  of 
funnels. 

In  Polygordius  lacteus  Woltereck  (1902)  found  also  two  pairs 
of  larval  nephridia:  (i)  Hauptnephridia,  close  to  the  epidermis  of 
the  ventral  hyposphere,  beset  proximally  only  with  tube-cells;  and 
(2)  Seitennephridia,  lined  with  such  cells  along  most  of  their 
lengths.  In  the  adult  of  this  species  also  one  of  these  pairs  must 
persist,  since  the  foremost  definitive  nephridia  lack  nephrostomes 
(Hempelmann,  1906). 

The  present  evidence  is  that  the  head  kidneys  are  closed  inter- 
nally, and  Meyer  accounts  for  this  by  the  lack  of  a  dissepiment  in 
front  of  them  from  which  a  nephrostome  could  form.  But  while 
Meyer  and  Woltereck  incline  to  an  ectoblastic  and  mesenchymatous 
origin,  Lillie  concludes  a  mesoblastic.  There  is  no  evidence  that  the 
adult  nephridia  are  division  products  of  larval  ones,  but  when  there 
is  a  second  pair  of  larval  nephridia  it  may  persist  in  the  adult. 

Provisory  Nephridia. — Following  on  the  larval  nephridia  and 
before  the  adult  one  are  formed  there  are  in  the  Capitellids  (and 
so  far  as  is  known  only  here  among  the  Polychaeta)  provisory 
nephridia,  each  of  which  participates  in  two  segments  (Eisig,  1887). 

Relation  of  Genital  Ducts  and  Nephridia. — This  question  has 
been  so  ably  reviewed  by  Goodrich  (1895,  1900),  and  his  investi- 
gations have  contributed  so  much  to  its  solution,  that  I  need  ta 
discuss  it  only  briefly.  Williams  (1858)  held  that  these  organs 
are  homologous,  and  derived  from  a  common  "  viscus."  Then 
Cosmovici  (1880)  concluded  that  the  segmental  organs  of  Annelids 
are  of  two  kinds :  excretory  organs  (organs  of  Bojanus),  and  genital 
ducts,  and  that  the  two  may  be  separated  or  may  be  united.  It  is 
the  particular  service  of  Eisig  (1887)  and  Meyer  (1887  ^^^  later 
'papers)  to  have  demonstrated  by  their  anatomical  and  embryological 
studies  that  the  peritoneal  funnels,,  the  original  genital  ducts,  are 
evaginations  caudad  of  dissepiments,  and  that  they  may  or  may  not 


674 


MONTGOMERY— MORPHOLOGY   OF  THE 


[April  S4» 


join  secondarily  with  the  nephridium  proper  that  develops  inde- 
pendently from  retroperitoneal  tissue.'  But  it  is  Groodrich  who  has 
made  the  most  comprehensive  comparative  investigation  of  these 
relations  (1895,  1897,  1898,  1900).  He  calls  the  peritoneal  tunnel 
(Genitalschlauch)  a  coelomoduct,  and  its  opening  a  coelomostome ; 
when  the  latter  preserves  its  original  strictly  genital  function  it  is 
a  gonostome.  According  to  his  terminology,  further,  a  nephridium 
is  an  excretory  organ  with  its  own  inner  opening,  and  the  latter 
is  a  nephridiostome.  The  coelomoducts  may  open  on  the  surface 
of  the  body  entirely  separate  from  the  nephridia,  the  primitive  con- 
dition, and  in  this  case  the  nephridia  are  purely  excretory  and 
possess  small  nephridiostomes ;  or  the  coelomostomes  may  become 
secondarily  grafted  upon  the  nephridia,  forming  compound  nephro- 
mixia  which  are  geni to-urinary  and  possess  large  funnels  (coelomo- 
stomes). These  relations  in  the  Polychaetes  he  tabulates  as  fol- 
lows ( 1900) : 


Nephridium   f  Genital    funnel   distinct,   but   open- 
closed       <       ing  into  nephridia!  canal  may  be  ■* 
internally.     I       acquired  at  maturity. 


Phyllodocidae. 

Glyceridae. 

Nephthyidae. 


Nephridium 

open 
internally. 


Genital  funnel  with  independent  ex-  (  Capitellidee. 
ternal  opening.  (  ?  Nereidae  (Lycoridea). 


Genital  funnel  becomes  connected 
with  the  nephrostome,  and  loses 
its  primitive  opening  to  the  ex- 
terior. 


Hesionidae  (all?). 

Syllidae. 

Aphroditidae. 

Eunicidae. 

Spionidae. 

Terebellidae. 

Sabellidse. 

Etc.,  etc. 

Goodrich  adduces  the  various  evidence  for  this  conclusion  and 
adds :  "  Moreover,  it  must  be  remembered  that  the  two  organs  are 
mutually  exclusive;  never  do  we  find  a  separate  genital  funnel  in 
those  forms  which  possess  wide-mouthed  excretory  organs;  and 
conversely,  with  the  one  possible  exception  of  Polygordius,  never 
do  we  find  Polychaetes  having  nephridia  with  only  small  true  nephro-  * 

•Meyer  (1890)  has  shown  that  Kleinenberg  (1886)  was  mistaken  in 
deriving  the  genital  ducts  from  the  ectoblast. 


i9o«]  EXCRETORY  ORGANS  OF  METAZOA.  575 

stomes  without  genital  funnels."*  Allen  (1904)  has  demonstrated 
that  in  Pcecilochcetus  both  kinds  of  organs  occur,  nephridia  with 
small  nephridiostomes  in  the  anterior  somites,  and  nephromixia  with 
large  funnels  in  the  posterior. 

Thus  the  evidence  is  convincing  that  coelomoduct  and  nephridium 
are  two  distinct  organs,  with  originally  separate  origins  and  func- 
tions, but  that  the  two  frequently  unite  to  produce  a  compound 
nephromixium. 

Mid-gut. — This  is  excretory  in  the  Polynoidae  (Schimkewitsch, 
1884),  and  so  are  the  intestinal  caeca  in  the  Aphroditidae  (Darboux, 
1900). 

Chloragogue, — Schaeppi  (1894)  found  the  chloragogue  of  only 
the  peritoneum,  nephridia  and  intrasinous  connective  tissue  is  ex- 
cretory (contains  guanin).  In  Arenicola  some  of  the  vessels  have 
caeca  whose  walls  possess  chloragogue  cells  (Willem,  1899).  I^^r 
the  chloragogue  of  peritoneal  origin  (peritoneal  glands)  Meyer 
(1901)  uses  the  term  "  phagocytic  organs." 

Eisig  (1887)  has  made  the  most  thorough  study  of  excretion  in 
the  Polychaetes ;  he  determined  that  carmine  is  taken  up  by  the  mid- 
gut, then  by  the  peritoneum,  and  that  the  haemolymph  is  the  vehicle 
of  its  transport  to  the  nephridia,  blood  vessels  being  absent  in  the 
Capitellids ;  it  ultimately  reaches  also  the  setal  glands  and  the  skin ; 
the  skin  is  not  excretory  though  it  becomes  the  seat  of  excretory 
substances,  and  it  is  by  the  accumulation  of  such  material  that  the 
skin  in  necessitated  to  undergo  moults. 

20.    ECHIURIDA. 

Segmental  Organs, — These  serve  mainly  if  not  wholly  as  genital 
ducts  and  in  Bonellia  the  male  lives  within  those  of  the  female. 
Bonellia  has  but  a  single  one,  while  in  Echiurus  and  Thalassema 
there  are  from  one  to  four  pairs.  Structurally  (Greef,  1879, 
Spengel,  1880)  these  are  long  tubes  each  with  a  nephrostome  close 
to  a  nephridiopore.  Nothing  seems  to  be  known  of  their  de- 
velopment. 

*Hempelniann  (1906)  has  since  shown  that  in  Polygordius  the  nephridio- 
stomes are  too  small  for  the  discharge*  of  the  germ  cells,  and  that  the  latter 
escape  by  rupture  of  the  posterior  end  of  the  body. 


576  MONTGOMERY— MORPHOLOGY  OF  THE  [April  m. 

Anal  Tubes. — ^There  is  one  pair  of  these  opening  into  the  most 
posterior  portion  of  the  intestine.  On  their  surfaces  there  are 
numerous  "  Wimpertrichter,"  and  Greef  supposed  these  not  to  open 
directly  into  the  coelom,  but  Spengel  demonstrated  that  they  do  make 
such  a  direct  connection  and  that  their  ciliated  lining  is  continuous 
with  the  peritoneum.  Their  fimction  is  not  ascertained.  From 
their  position  Spengel  concluded  them  to  be  ectoblastic,  but  not  to 
be  homodynamous  with  the  segmental  organs.  But  Hatschek 
(1880)  describes  them  as  arising  not  from  the  rectum  but  from  the 
somatic  mesoblast  of  the  telson ;  and  according  to  this  account  they 
form  first  the  Wimpertrichter,  then  later  the  external  pores  that  lie 
lateral  from  the  anus. 

Larval  Nephridia, — These  are  known  only  from  Hatschek's  ac- 
count ( 1880)  of  Echiurus;  the  first  origin  of  these  "  Kopfnieren  " 
was  not  determined;  each  becomes  a  much  branched  organ  with 
intracellular  cavity,  from  the  surface  of  which  delicate  blind  capil- 
laries grow  out.  Torrey  (1903)  was  unable  to  find  larval  nephridia 
in  Thalassema,jaLnA  determined  that  in  this  form  excretion  is  accom- 
plished by  certain  mesenchyme  cells. 

21.  Myzostomida. 

The  single  pair  of  nephridia  were  first  recognized  as  such  by 
Beard  (1894),  and  their  structure  particularly  described  by  Wheeler 
(1896)  and  Stummer-Traunfels  (1903).  Their  relations  differ 
somewhat  in  different  species:  they  may  be  separated  from  each 
other,  or  their  open  and  large  nephrostomes  may  be  united,  \heir 
nephridiopores  may  be  separated  or  united;  in  one  species  nephro- 
stomes appear  to  be  absent.  In  some  species  they  are  purely  excre- 
tory, in  others  also  spermiducal.  From  their  development  Wheeler 
concluded  that  they  originally  opened  on  the  surface  of  the  body  and 
not  into  the  cloaca  (their  usual  termination  in  the  adult),  because 
in  one  species  the  unpaired  excretory  duct  opens  "  on  the  surface 
of  the  body  through  a  papilla  lying  just  ventral  to  the  cloacal 
orifice."* 

•The  segmental  sacs  (suckers)  supposed  by  Nansen  (1885)  to  be  nc- 
phric,  have  been  shown  by  Wheeler  to  be  probably  sensory. 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  677 

22.   EnTEROPNEUSTA. 

Nephridia. — ^There  is  a  left  canal  (or  a  right  and  left)  con- 
necting the  coelom  of  the  proboscis  with  the  exterior,  a  pair  of 
similar  canals  in  the  collar  region,  and  in  Spengelia  (Willey,  1899) 
rudimentary  pores  along  the  whole  trunk.  Spengel  (1893)  con- 
sidered them  to  take  in  water  from  without  and  to  subserve  loco- 
motion ;  Willey  regarded  them  as  having  lost  their  former  excretory 
function,  while  Bateson  (1884)  showed  by  carmine  injection  that 
the  collar  pores  are  excretory. 

An  ectoblastic  origin  of  these  structures  was  the  result  of  the 
study  of  Spengel  and  Morgan  (1894).  But  DawydofF  (1907), 
examining  those  of  the  proboscis  in  the  process  of  regeneration, 
found  that  they  develop  from  a  peritoneal  evagination  that  connects 
with  an  ectoblastic  ingrowth,  and  from  this  concluded  that  they 
are  true  nephridia — a  view  previously  reached  by  Schimkewitsch 
(1888). 

The  genital  ducts  seem  to  bear  no  relation  to  nephridia,  and  the 
larva  (tornaria)  lacks  special  excretory  organs. 

Glomerulus. — ^A  vascular  structure  connected  with  the  peri- 
cardium, considered  the  only  excretory  organ  in  the  adult  (Willey, 
1899)  ;  I  have  not  seen  the  original  description  and  consequently  am 
unable  to  add  further  details. 

23.   MOLLUSCA. 

Adult  Nephridia. — I  have  not  attempted  to  labor  through  the 
compendious  literature  on  the  anatomy  of  these  organs,  but  shall 
simply  give  a  brief  summary  drawn  mainly  from  the  excellent  treat- 
ment by  Hescheler  (1900).  These  are  essentially  similar  and 
homologous  throughout  the  group,  and  consist  typix:ally  of  a  pair 
of  sacs  which  communicate  internally  by  open  nephrostomes 
(renopericardial  apertures)  with  the  ccelom  (pericardial  cavity), 
and  externally  by  nephridiopores  with  the  mantle  cavity.  They  are 
paired  in  all  the  groups  except  the  Gasteropods,  and  among  the  latter 
in  most  of  the  diotocardial  prosobranchs ;  among  living  forms  there 
is  more  than  one  pair  only  in  Nautilus.  They  may  be  simple  tubes, 
or  may  be  twisted  or  excessively  ramose.      Functionally  they  may 


578  MONTGOMERY— MORPHOLOGY  OF  THE  [April  a*, 

be  exclusively  excretory,  the  usual  condition,  or  mainly  genital 
(Solenogastra),  or  genito-urinary. 

Development  of  the  Adult  Nephridia. — ^According  to  one  view 
the  glandular  portion  of  the  nephridium  arises  as  a  peritoneal  funnel, 
an  evagination  of  the  pericardium,  this  joining  later  with  an  ecto- 
blastic  ingrowth,  the  duct  or  ureter;  in  support  of  this  view  is  the 
work  of  Rabl  (1879,  Planorbis),  Erlanger  (1891a,  Paludina), 
Biitschli  (1877,  Paludina),  Salensky  (1885,  Vertnetus),  Schimke- 
witsch  (1888,  Limax),  Drummond  (1902,  Paludina),  Ahting  (1901, 
Pelecypods),  Pelseneer  (1901,  Helix),  and  Stauffacher  (1898, 
Cyclas).  That  these  organs  are  wholly  mesoblastic  is  the  opinion 
of  Salensky  (1872,  Calyptrcea),  Erlanger  (18926,  Bythinia), 
Georgevitch  (1900,  Aplysia),  and  Faussek  (1900,  Loligo).  The 
third  view  is  that  they  are  altogether  ectoblastic:  Fol  (1875,  Ptero- 
pods),  Bobretzky  (1877,  Nassa),  Joyeux-Laflfuie  (1882,  Onchid- 
ium),  Sarasin  (1882,  Bythinia),  and  Meisenheimer  (1898,  Limax, 
1901a,  Dreissensia,  igoib,  Cyclas). 

The  first  of  these  views  has  the  greatest  support,  pointing  to  the 
pericardial  origin  of  the  funnel  and  glandular  portion,  and  to  ecto- 
blastic origin  of  some  portion  of  the  ureter  only.  For  the  third 
view,  wholly  ectoblastic  origin,  it  will  be  noted  that  the  only  recent 
work  is  that  of  Meisenheimer.  Now  almost  all  the  writers  con- 
clude a  common  origin  of  the  glandular  portion  of  the  nephridium 
and  the  pericardium  and  Meisenheimer  does  so  likewise,  but  in 
opposition  to  almost  all  preceding  study  he  regards  the  pericardium 
and  heart  as  ectoblastic  abstrictions.  Meisenheimer  must  surely  be 
incorrect  in  interpreting  the  peritoneum  and  with  it  the  nephridiimi 
as  ectoblastic,  i,  e.,  he  must  have  defined  the  germ  layers  quite  diflfer- 
ently  from  other  embryologists,  since  the  pericardial  cavity  is  justly 
considered  coelomic  yet  in  no  other  animal  group  is  the  ccelom  re- 
garded as  lined  by  ectoblast. 

These  definitive  nephridia  seem  to  arise  independently  of  the 
larval  ones,  save  that  Rho  (1888)  and  Mazzarelli  (1892,  1898)  state 
that  the  mesoblastic  anal  kidneys  of  opisthobranch  larvae  become 
transformed  into  the  adult  ones.  I 

Genital  Ducts, — "  Relations  between  the  nephridial  and  genital 
system,  similar  to  those  in  the  Worms,  exist  in  the  Solenogastrids 


,9o8]  EXCRETORY   ORGANS  OF  METAZOA.  679 

where  the  nephridia  function  as  discharge  ducts  for  the  genital 
products.  .  .  .  And  again  in  some  Lamellibranchs,  Diotocardians 
and  the  Scaphopods  there  exist  relations  between  sex  glands  and 
nephridia  in  that  the  sex  glands  open  into  the  nephridia,  so  that  a 
shorter  or  longer  portion  of  the  latter  functions  not  only  as  kidney 
or  ureter  but  also  as  discharge  duct  for  the  genital  products  ' 
(Hescheler,  1900).  In  those  prosobranchs  with  only  one  adult 
nephridium,  Drummond  (1902)  has  shown  for  Paludina,  and  after 
a  full  discussion  of  the  literature,  that  the  right  nephridium  of  the 
embryo  persists  as  the  left  one  of  the  adult,  in  agreement  with 
Erlanger,  but  contrary  to  his  results  she  finds  the  left  nephridium 
of  the  embryo  does  not  disappear  but  becomes  the  genital  duct. 

Larval  Nephridia. — These  are  known  only  in  Gasteropods  and 
Pelecypods  (Lamellibranchs),  and  it  will  be  most  convenient  to 
treat  separately  the  groups  in  which  they  occur. 

(a)  Prosobranch  Gasteropods, — ^Two  kinds  of  these  have  been 
described,  (i)  External  nephridia  (Aussennieren,  excretory  cells). 
These  are  ectoblastic,  unicellular  or  multicellular  organs,  usually 
projecting  from  the  surface  of  the  body  just  behind  the  velum; 
there  is  one  pair  of  them,  and  their  cavity  communicates  with  the 
blastoccel;  sometimes  they  have  an  opening  to  the  exterior.  They 
have  been  described  most  carefully  for  Crepidula  (Conklin,  1897) 
and  Fasciolaria  (Glaser,  1905),  also  for  Nassa,  Natica,  Fusus 
(Bobretzky,  1877),  Paludina  and  Bythinia  (Sarasin,  1882,  who 
calls  them  "ansae"),  Fasciolaria  and  Fulgur  (McMurrich,  1886), 
Fissurella  (Boutan,  1885),  and  Capulus  (Erlanger,  1892a).  Glaser 
has  demonstrated  that  they  are  first  digestive,  later  serve  as  reser- 
voirs for  waste  products,  and  subsequently  fall  off  from  the  surface 
of  the  larva ;  Sarasin  and  McMurrich  supposed  they  were  originally 
parts  of  the  preoral  velum,  and  that  with  excretory  specialization 
they  separated  off  from  it;  but  Conklin  and  Glaser  show  that  they 
arise  independently  of  and  before  the  velum.  As  "  secondary 
outer  kidneys  "  Glaser  has  described  certain  excretory  cells  placed 
in  the  velum  and  the  head  vesicle. 

(2)  The  second  kind  of  larval  excretory  organs  of  the  proso- 
branchs are  mesoblastic.  These  arise  from  a  mesoblastic  anlage 
that  is  at  first  solid,  while  more  or  less  of  the  duct  is  ectoblastic; 


580  MONTGOMERY— MORPHOLOGY   OF  THE  [April  a4, 

they  are  ciliated  with  exterior  apertures.     These  have  been  found 
in  Bythinia  and  Paludina  (Butschli,  1877,  Erlanger,  1891a,  1892&). 

(&)  Opisthobranch  Gasteropods, — Here  there  are  distinguished 
nephrocysts  and  anal  kidneys.  The  nephrocysts  were  discovered 
and  named  by  Trinchese  (1881)  for  Ercolania,  Amphorina,  Bergia 
and  Doto;  and  were  described  also  by  Mazzarelli  (1892)  for  Aplysia 
and  by  Casteel  ( 1904)  for  Fiona.  These  are  rounded  bodies  lying 
anterior  to  the  anus  in  the  blastocoel,  without  external  ducts ;  nothing 
positive  is  known  of  their  origin,  and  Trinchese  supposes  them 
mesoblastic  simply  from  their  position..  They  may  occur  in  the 
same  embryo  together  with  the  following  organs.  The  anal  kidneys 
were  first  interpreted  as  excretory  by  Langerhans  (1873,  Doris  and 
Accra),  They  are  a  pair  of  single  cells,  or  groups  of  cells,  that 
originate  near  the  anus  but  may  migrate  further  forward.  Trinchese 
(1881)  and  Guiart  (1901)  derived  them  from  the  mesoblast,  and 
so  also  did  Mazzarelli  (Aplysia,  1892,  1898)  who  ascribed  the  occa- 
sional unpaired  condition  to  the  fusion  of  a  pair.  But  Lacaze- 
Duthiers  and  Pruvot  (1887)  described  them  as  ectoblastic,  and  this 
conclusion  was  reached  also  in  the  careful  studies  of  Heymons 
(1893,  Umbrella)  and  Casteel  (1904,  Fiona).  Casteel's  work  is  the 
most  thorough  on  any  opisthobranch,  and  he  states :  "  There  is  no 
point  regarding  the  cytogeny  of  Fiona  of  which  I  am  more  certain 
than  that  the  group  of  cells  constituting  the  anal  kidney  is  of  ecto- 
dermal origin." 

(c)  Pulmonate  Gastcropods. — Here  again  there  are  two  kinds 
of  larval  kidneys.  The  external  kidneys  (aussere  Nieren)  occur 
one  on  either  side  of  the  body,  each  a  projecting  group  of  vacuo- 
lated cells  forming  part  of  the  ectoblastic  velum.  These  were  dis- 
covered by  Butschli  (1877),  and  have  been  described  by  Fol  (1880) 
and  Rabl  (1879)  for  Planorbis.  Much  more  attention  has  been 
given  to  the  head  kidneys  (Umieren).  The  most  detailed  descrip- 
tion of  these  in  their  perfected  condition  is  that  of  Meisenheimer 
(1898,  1899)  :  in  the  Basommatophora  {Ancylus,  Physa,  Planorbis, 
Limncea)  these  are  much  alike,  each  consisting  of  but  four  cells 
with  intracellular  cavity,  the  innermost  of  which  closes  the  canal 
against  the  blastocoel  and  bears  a  ciliary  flame.  In  the  Stylom^ 
matophora   (Limax,  Succinca,  Helix,  Arion)   the  cells  are  much 


x9o8]  EXCRETORY   ORGANS   OF   METAZOA.  581 

more  numerous  and  the  inner  end  is  composed  of  a  number  of  large 
amoeboid  cells  all  of  which  have  long  cilia;  for  a  while  the  inner  end 
may  be  open  (as  described  by  Rabl,  1879,  and  Erlanger,  1894)  since 
the  cells  there  may  become  loosened  from  their  epithelial  connection, 
but  later  this  end  becomes  completely  closed  even  though  at  places 
by  a  very  thin  membrane.  These  are  the  most  complicated  larval 
nephridia  found  in  Gasteropods ;  they  subsequently  degenerate  com- 
pletely. As  to  the  development  of  these  head  kidneys:  Rabl  (1879) 
and  Holmes  (1900)  considered  them  mesoblastic;  Erlanger  (1893) 
interpreted  them  as  mainly  mesoblastic  with  a  portion  of  the  duct 
ectoblastic,,and  Pelseneer  (1901)  stated  that  the  large  distal  portion 
is  ectoblastic.  But  Fol  (1880),  Wolf  son  (1880)  and  Meisenheimer 
(1898)  concluded  that  they  are  entirely  ectoblastic;  the  last  named 
investigator  speaks  of  them  as  arising  as  paired  tubular  invaginations 
at  the  level  of  the  proctodaeum. 

(d)  Pelecypods  {Lamellibranchs) , — In  Teredo  there  is  a  pair 
of  ciliated  Urnieren  in  the  young  larva  (Hatschek,  1880).  Only 
the  left  one  is  developed  in  Cyclas,  and  opens  externally  in  the 
region  of  the  head  vesicle;  it  consists  of  three  highly  complex  cells 
with  intracellular  cavity,  the  innermost  branched  cell  closing  it 
from  the  blastocoel  (Stauffacher,  1898).  In  Dreissensia  each  of  the 
larval  kidneys  consists  of  three  cells,  the  innermost  provided  with 
a  ciliary  flame  and  closing  the  canal,  the  next  forming  the  tube,  and 
the  third  constituting  a  duct  connecting  with  the  surface  (Meisen- 
heimer, 1901a).  With  regard  to  the  embryogeny,  Hatschek  de- 
scribed these  organs  as  appearing  first  at  the  anterior  ends  of  the 
mesoblastic  bands,  at  first  with  no  connection  with  the  ectoblast, 
and  concluded  that  the  nephridium  of  each  side  "  is  probably  derived 
from  only  one  or  a  few  mesoderm  cells  " ;  Stauffacher  held  that  in 
Cyclas  only  the  innermost  cells  is  mesoblastic  and  the  others  ecto- 
blastic; while  Meisenheimer  (Dreissensia,  1901a,  Cyclas,  19016) 
described  thehi  as  arising  conjointly  with  the  heart  and  pericardium 
from  the  ectoblast. 

Homologies  of  the  Larval  Nephridia. — Salensky  (1872)  and 
Bobretzky  (1877)  homologized  the  outer  kidneys  of  prosobranchs 
with  the  Urnieren  of  Pulmonates.  Biitschli  ( 1877)  suggested  that 
the  Urnieren  of  Paludina  are  possibly  homologous  with  those  of 


582  MONTGOMERY—MORPHOLOGY  OF  THE  [April  a4, 

the  pulmonates,  but  that  there  is  no  homology  between  the  outer 
kidneys  of  these  groups.  Rabl  (1879)  concluded  that  the  outer 
kidneys  of  Planorbis  are  probably  comparable  with  the  outer  kid- 
neys of  freshwater  prosobranchs,  but  not  with  the  Umieren.  Fol 
( 1880)  maintained  that  the  outer  kidneys  of  Pulmonates  are  homo- 
logous with  the  Urnieren  of  prosobranchs.  Erlanger  (1893)  ^^ 
garded  all  the  larval  nephridia  as  homologous  with  each  other  and 
probably  also  with  the  head  kidneys  of  Annelids,  and  distinguished 
the  following  kinds:  (i)  Outer  ectoblastic  kidneys  (marine  proso- 
branchs) ;  (2)  inner  mesoblastic,  and  these  either  (a)  purely  meso- 
blactic  (opisthobranchs),  or  (6)  mesoblastic  with  the  canal  at  least 
in  part  ectoblastic  (pulmonates,  pelecypods,  freshwater  proso- 
branchs). Mazzarelli  (1904)  considered  the  Urnieren  of  pele- 
c3rpods,  pulmonates  and  freshwater  prosobranchs  to  be  homologous, 
but  the  external  nephridia  of  marine  prosobranchs  to  be  different 
structures ;  and  the  nephrocysts  of  opisthobranchs  to  be  organs  that 
have  secondarily  lost  their  ducts  and  that  correspond  with  the  ex- 
cretory cells  of  the  Umieren  of  other  MoUusks.  Finally  Glaser 
(1905)  has  given  a  good  review  of  the  question,  and  maintains  there 
are  at  least  three  distinct  and  dyshomologous  larval  excretory 
organs  ( i )  Urnieren,  mesectoblastic  structures  of  prosobranchs  and 
pulmonates;  (2)  Aussennieren,  modified  ectoblastic  cells  of  proso- 
branchs and  pulmonates;  and  (3)  excretion  cells,  those  of  Umbrella 
placed  near  the  anus ;  the  Urnieren  are  further  of  two  kinds  because 
some  of  them  appear  to  be  wholly  ectoblastic. 

There  is  so  much  confusion  of  opinion  with  regard  to  the  de- 
velopment of  even  the  same  kind  of  excretory  organ  in  the  same 
species,  that  I  fully  agree  with  Casteel  (1904)  "that  much  more 
work  must  be  done  upon  these  organs  of  molluscan  larvae  before  we 
are  ready  to  come  to  definite  conclusions  regarding  their  mutual 
relations  and  homologies,  if  such  exist."  There  are  certainly  two 
distinct  kinds  that  may  occur  at  the  same  stage  in  the 'same  species, 
and  that  on  account  of  their  differences  in  position,  structure  and 
origin  are  not  homodynamous,  and  these  are:  (i)  Projecting  vesi- 
cles, wholly  ectoblastic,  forming  part  of  or  placed  near  to  the 
velum;  and  (2)  vesicular  or  tubular  organs  placed  below  the  ecto- 
blast  and  behind  the  preceding,  which  in  most  cases  appear  to  be 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  683 

in  part  mesoblastic.     All  those  of  the  first  kind  may  well  be  homo-    / 
logous,  but  those  of  the  second  kind  are  more  probably  hetero- 
geneous structures. 

Other  Excretory  Organs, — According  to  Cuenot  (1899)  the  fol- 
lowing structures  are  excretory :  in  the  Amphineura  and  Scaphopoda 
connective  tissue  cells;  in  prosobranch  and  opisthobranch  Gas- 
teropoda similar  cells  as  well  as  cells  of  the  liver ;  in  the  Pelecypoda 
pericardial  glands;  and  in  the  Cephalopoda  phagocytes  and  the 
gill-hearts. 

24.  Tardigrada  (Arctiscoidea.) 

A  pair  of  glands  opening  into  the  rectum  were  supposed  by 
Plate  (1888)  to  be  excretory,  and  he  compared  them  with  the 
Malpighian  vessels  of  the  Acarina.  But  neither  he  nor  Basse 
(1905),  who  has  furnished  a  fuller  description,  were  able  to  find 
excretory  products  in  these  organs.  Nothing  is  known  of  their 
development. 

25.  Pycnogonida  (Pantopoda). 

Dohm  (1881)  has  described  problematical  "  Excretionsorgane  " 
within  the  cavity  (blastocoel)  of  the  fourth  or  fifth  joint  of  the 
second  extremity,  or  the  third  or  fourth  joint  of  the  third;  each 
has  an  external  opening  placed  upon  a  small  tubercle;  in  genera 
where  the  named  extremities  are  absent,  these  organs  are  found 
in  the  wall  of  the  body  at  points  opposite  the  missing  extremities. 
These  organs  lie  in  extremities  that  lack  reproductive  organs,  and 
for  that  reason  Dohm  suggested  they  may  have  some  homodynamic 
relation  to  the  latter. 

Kowalevsky  (1892)  found  by  injections  of  acid  fuchsine  that 
the  stain  is  taken  up  by  small  hypodermal  glands  placed  in  Phoxi- 
chilus  on  the  borders  of  the  three  anterior  segments  and  on  the  bases 
of  extremities  fourth  to  seventh,  and  in  Pallene  and  Ammothea  in 
the  lateral  processes  of  trunk  segments  and  in  the  first  joints  of  the 
extremities. 

26,  Crustacea. 

Shell  Glands  {Maxillary  Glands). — These  have  been  described 
for  the  Phyllopoda  (Leydig,  i860,  Weismann,  1874,  Claus,  1875, 
Dohrn,  1870,  Nowikoff,  1905),  Copepoda  (Claus,  1877,  Nettovich, 


584  MONTGOMERY— MORPHOLOGY  OF  THE  [April  34, 

1900),  Isopoda  (Vejdovsky,  1901 ;  and  Nemec,  1896,  who  states 
that  in  Ligidium  they  are  modified  into  salivary  glands),  Cirripedia, 
(Bruntz,  1903,  Bemdt,  1903;  in  Balanus  they  communicate  with  the 
ccelom  only  in  the  cypris-stage  according  to  Gruvel,  1894),  Stoma- 
topoda  (Bruntz,  1903),  and  freshwater  Ostracoda  (Claus,  1895, 
Daday,  1895).  These  open  at  or  near  the  base  of  the  second 
maxillae,  each  has  a  closed  enlarged  end  sac  lined  by  an  excretory 
epithelium,  and  they  are  placed  in  the  shell  duplicature  except  in 
Leptodora  where  the  greater  portion  of  the  organ  lies  in  the  thorax. 
According  to  Richard  (1892)  their  ducts  are  longest  in  freshwater 
and  shortest  in  brackish  water  species.  In  freshwater  Cladocera 
(Simocephalus)  I  have  found  that  the  end  sac  takes  up  injected 
carmine  at  the  end  of  a  few  hours. 

Antennal  Glands, — These  have  been  described  for  the  larvae  (but 
not  adults)  of  Copepoda  and  Phyllopoda  (Grobben,  1881),  for 
Amphipods  (Grobben,  1881,  Bonnier,  1891,  Bruntz,  1903,  Vejdov- 
sky, 1901,  Delia  Valle,  1893),  Schizopoda  (Grobben,  1881,  Bruntz, 
1903) >  Ostracoda  (Claus,  1890,  1895),  Cirripedia  where  they  are 
modified  into  cement  glands  but  may  still  continue  excretory 
(Koehler,  1890),  Isopoda  (in  Asellus  where  they  are  degenerate, 
Nemec,  1896),  and  Decapoda  (Marchal,  1892,  Waite,  1889).  The 
antennal  glands  are  essentially  similar  to  the  maxillary.  Both  have 
closed  end  sacs,  are  without  cilia,  and  both  (Vejdovsky,  1901) 
possess  at  the  junction  of  the  gland  and  duct  a  narrow  "  Trichter  " 
composed  of  a  few  large  cells  with  a  peripheral  muscular  sphincter. 

Development  of  the  Preceding  Organs. — According  to  the  earlier 
observers  (Reichenbach,  1886,  Ischikawa,  1885)  the  shell  and 
antennal  glands  are  ectoblastic,  but  other  studies  (Kingsley,  1889, 
Waite,  1899,  Grobben,  1879,  Lebedinsky,  1891)  show  that  each 
arises  as  a  reduced  coelomic  sac  (or  portion  of  one)  connecting  with 
an  ectoblastic  duct.  The  end  sac  of  the  adult  thus  corresponds  to 
the  coelomic  sac  of  the  embryo. 

Maxillipedal  Glands. — In  Diaptomus  there  is  a  pair  of  these 
opening  at  the  basis  of  the  first  maxilHpeds;  their  structure  is  like 
that  of  the  preceding  glands  (Richard,  1892).  It  is  probable  that 
some  of  the  glands  described  as  maxillary  are  really  maxillipedal. 

Coxal  Glands. — In   Gammarus   (Delia  Valle,   1893)   there  are 


.9o8]  EXCRETORY  ORGANS  OF  METAZOA.  586 

small  groups  of  gland  cells,  that  take  up  carmine,  placed  at  the  bases 
of  the  maxillipeds,  thoracic  and  abdominal  extremities.  Similar  ap- 
pear to  be  the  "  Segmentalorgane  "  of  the  Ostracoda  (G.  W.  MuUer, 
1894),  which  in  Paradoxostoma  lie  above  each  leg  pair,  and  in 
Bairdia  above  the  first  pair ;  and  the  glands  opening  on  the  maxil- 
lipeds of  Cyprids  (Claus,  1890). 

Genital  Ducts. — The  first  origin  of  these  seems  to  have  been 
little  investigated,  but  Pedaschenko  (1899)  finds  them  to  arise  from 
a  proximal  mesoblastic  and  a  distal  ectoblastic  portion. 

Homologies  of  the  Preceding  Organs. — ^The  maxillary,  antennal 
and  maxillipedal  glands  are  probably  homodynamous,  and  seem  to 
differ  only  in  antero-posterior  position.  Sometimes  they  occur  at 
the  same  time  in  the  same  individual,  or  (as  in  Phyllopods  and 
Copepods)  the  antennal  gland  is  the  larval  and  the  shell  gland  the 
adult  excretory  organ.  Sometimes  both  antennal  and  shell  glands 
are  absent  in  the  adult,  as  in  some  Copepoda  (Nemec,  1896).  Waite 
(1899)  has  discussed  these  homologies  at  some  length,  and  resumes : 
"  The  nephridium  of  Annelids  is  probably  represented  in  Crustacea 
in  the  second  (antennal)  segment  by  the  antennal  gland  of  Mala- 
costraca;  in  the  fifth  (second  maxillary)  segment  by  the  shell  gland 
of  Entomostraca  and  some  Malacostraca ;  in  the  sixth  (first  maxil- 
lipedal) segment  of  some  Malacostraca  by  the  '  Segmentalorgan '  of 
Lebendinski ;  it  is  possibly  represented  in  the  fourth  (first  maxillary) 
segment  by  the  excretory  organ  described  by  Boutchinsky,  and  in 
the  sixth  to  thirteenth  (maxillipedal  and  pareiopodal)  segments  in 
part  by  the  branchial  glands,  and  in  part  (in  the  eleventh  and 
thirteenth  segments)  by  the  genital  ducts." 

Nephrocytes, — ^Bruntz  ,(1903)  has  found  these  excretory  cells 
to  be  distributed  as  follows :  they  are  absent  in  the  Qadocera ;  there 
is  one  cephalic  pair  in  the  Isopoda,  Amphipoda  and  Cirripedia;  up 
to  eight  pairs  placed  in  the  thorax  in  the  Schizopoda,  Decapoda  (in 
the  gills),  and  Copepoda  parasitica  (diffuse);  from  one  to  eight 
pairs  in  the  abdomen  in  the  Isopoda  and  Stomatopoda  (in  the  legs)  ; 
and  eleven  pairs  in  the  thorax  and  abdomen  in  the  Amphipoda. 

Other  Excretory  Organs. — As  such  have  been  described  the  fer- 
ment cells  of  the  liver  of  Decapoda,  Amphipoda  and  Isopoda,  and 

PROC.  AMER.  PHIL.  SOC.,  XLVII.  I9O  LL,  PRINTED  JANUARY  I4,  I909. 


686  MONTGOMERY— MORPHOLOGY  OF  THE  [April  a4, 

the  mid-gut  caeca  of  Amphipoda  (Bruntz,  1903)  ;  the  mantle  in  the 
Cirripedia  (Gruvel,  1894) ;  and  connective  tissue  cells  of  Copepoda 
when  the  antennal  and  maxillary  glands  are  lacking  (Nemec,  1896). 

27.  Onychophora  (Protracheata). 

Nephridia, — According  to  the  observations  of  Balfour  (1883) 
and  subsequent  investigators,  one  pair  of  nephridia  occurs  in  each 
trunk  somite,  «.  e.,  one  pair  to  each  pair  of  legs,  except  in  the 
penultimate  or  antepenultimate  segment.  Each  opens  ventrally  at 
the  basis  of  a  leg,  and  consists  of  an  outermost  excretory  bladder, 
a  loop  and  a  nephrostome  that  opens  into  the  ccelom ;  but  the  portion 
of  the  ccelom  that  has  such  a  connection  is,  as  in  the  case  of  the 
antennal  and  maxillary  glands  of  the  Crustacea,  completely  ab- 
stricted  from  the  remainder  of  the  ccelom  and  with  excretory  func- 
tion, therefore  each  such  coelomic  sac  may  rightly  be  considered  a 
closed  inner  end  sac  of  the  nephridium.  This  is  in  agreement  with 
the  facts  of  the  embryogeny,  as  detailed  by  Sedg>yick  (1885-8)  and 
Evans  (1901),  according  to  whom  each  right  and  left  coelomic  sac 
pinches  into  a  dorsal  and  a  ventral  portion,  and  the  latter  portion 
sends  an  outgrowth  reaching  to  and  opening  at  the  leg. 

The  salivary  glands  and  genital  ducts  develop  like  the  nephridia 
and  represent  them  in  segments  where  they  are  lacking,  are  accord- 
ingly homodynamous  with  them  (Sedgwick)  ;  and  the  receptaculum 
ovorum  is  homodynamous  with  an  end  sac  of  a  nephridium  (Evans). 

Anal  Glands, — These  also  have  been  considered  homologous  with 
nephridia  by  v.  Kennel  (1885).  But  Purcell  (1900)  has  indi- 
cated that  the  so-called  "  accessory  glands  "  of  the  postgenital  seg- 
ments may  rather  be  dyshomologous ;  that  while  those  (anal  glands) 
of  the  American  Peripatus  are  nephridia,  those  of  other  genera  are 
probably  ectoblastic  crural  glands. 

Nephrocytes, — ^There  are  medio-dorsal  bands  of  these,  also 
masses  of  them  near  the  bases  of  the  legs  (Bruntz,  1903). 

28.  Insecta. 

Malpighian  Vessels, — These  are  absent  in  Japyx  (Grassi,  1888) 
and  also  in  the  Collembola  where  Folsom  and  Welles  ( 1906)  found 
that  the  whole  ventriculus  is  excretory  and  periodically  moults  its  epi- 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  587 

thelium ;  they  are  not,  as  generally  supposed,  absent  in  the  Aphidse, 
for  Witlaczil  (1882)  has  shown  that  the  so-called  pseudovitellus 
represents  them.  In  all  other  Insects  these  vessels  are  present,  and 
are  usually  delicate,  cylindrical  tubes,  rarely  varicose  or  ramose, 
with  their  inner  ends  closed  and  the  distal  ends  joining  with  the 
intestine  usually  at  the  junction  of  the  mid-gut  and  proctodaeum, 
and  they  may  insert  there  singly  or  by  one  or  several  common  ducts. 
In  some  cases  there  are  two  different  kinds  in  the  same  species. 
Their  number  is  often  constant  for  a  group  as  may  be  seen  from 
the  following  summaries  taken  from  the  observations  of  Dufour 
(1833,  1841,  1851):  in  the  Diptera  there  are  usually  four,  rarely 
five  (Culex)y  and  never  more  than  four  in  the  Hemiptera;  there  are 
generally  less  than  eight  in  the  Coleoptera;  six  in  Phryganids, 
Termes,  Megaloptera  (Corydalis,  Sialis),  Panorpa,  eight  in 
Hemerobia  and  Myrmeleo;  they  are  much  more  numerous  in  the 
Orthoptera,  Hymenoptera,  Libellulidae  and  Ephemeridae.* 

While  Dufour  called  them  "  organes  hepatiques  ou  biliaires," 
subsequent  work  has  proved  conclusively  that  they  are  the  main 
excretory  organs. 

According  to  the  majority  of  investigators  they  arise  as  evagina- 
tions  of  the  ectoblastic  proctodaeum,  and  only  in  some  Hymenoptera 
do  they  first  appear  as  ectoblastic  evaginations  at  the  posterior  end 
before  the  proctodaeum  forms.  The  largest  number  known  in  any 
embryo  is  ten  (Melanoplus,  Packard),  which  seems  to  be  the  single 
case  not  in  agreement  with  Wheeler's  conclusion  (1893a)  that  no 
more  than  six  occur  in  embryos.  Wheeler  concludes  that  six  is 
the  primitive  number,  while  others  have  reasoned  this  to  be  foun 
Only  in  the  Termites  are  they  more  numerous  in  the  larvae  than 
in  the  adults. 

Homologies  of  the  Malpighian  Vessels, — These  have  been  com- 
pared specially  with  the  sericteries  and  tracheae  and  more  generally 
with  nephridia  of  the  annelidan  type;  and  it  is  most  convenient  to 
treat  these  relations  at  this  place.  Biitschli  (1870)  showed  that 
the  sericteries  and  Malpighian  vessels  develop  like  the  tracheae,  re- 

•A  good  review  of  their  numerical  and  other  relations  is  given  by 
Packard  (1898).  In  theThysanura  (except  Japyx)  their  number  was  found 
by  Grassi  (1888)  to  vary  from  eight  to  sixteen. 


688  MONTGOMERY— MORPHOLOGY  OF  THE  [April  m. 

garded  the  sericteries  and  tracheae  as  homologous,  but  questioned 
whether  the  Malpighian  vessels  are  related  to  them.  Then,  follow- 
ing Semper's  (1874)  suggestion  that  the  tracheae  are  metamor- 
phosed segmental  organs,  Mayer  (1875)  went  further  in  concluding 
that  the  tracheae,  sericteries  and  Malpighian  vessels  are  homo- 
dynamous  and  all  homologous  with  nephridia  of  Annelids.  Grassi 
(1885)  has  in  the  main  supported  Mayer,  in  reasoning  that  the 
Malpighian  vessels,  sericteries,  the  two  transitory  invaginations  on 
the  head  and  the  homodynamous  tracheae  are  all  probably  excretory 
in  the  larva;  and  (1888)  supports  the  idea  of  the  homology  of 
Malpighian  vessels  with  tracheae  on  the  ground  that  the  former 
occur  in  segments  where  the  latter  are  lacking  and  are  most  abundant 
when  the  latter  are  least  numerous.  But  several  strong  objections 
have  been  made  to  these  comparisons,  and  especially  by  those  who 
have  studied  the  embryogeny  more  in  detail.  Thus  Hatschek 
(1877&)  has  argued  against  the  homology  of  the  sericteries  and 
salivary  glands  with  the  tracheae,  that  in  the  segments  where  the 
former  occur  tracheal  invaginations  are  formed  independently  of 
them.  Then  Palmen  (1877)  concluded  that  the  Malpighian  vessels, 
developing  from  the  proctodaeum,  were  originally  hypodermal 
glands  that  have  come  to  group  themselves  around  the  inner  end 
of  the  proctodaeum  and  that  their  number  is  "  in  no  way  dependent 
upon  the  number  of  particular  body  segments";  while  against  the 
homology  of  the  tracheae  with  nephridia,  he  adducted  the  case  of 
their  coincident  segmental  occurrence  in  Peripatus,  Wheeler  also 
(1893a)  judged  that  if  the  Malpighian  vessels  are  homologous  with 
nephridia  they  can  be  only  with  the  ectoblastic  portion  of  the  latter ; 
and  that  they  are  not  homodynamous  with  tracheae,  but  rather  with 
the  mass  of  oenocytes  that  represent  the  ectoblastic  remains  of 
nephridia.  Heymons  (1896)  also  concluded  that  the  Malpighian 
vessels  are  not  to  be  compared  with  nephridia,  that  they  are  only 
local  evaginations  of  the  hind-gut. 

The  evidence  is  that  the  Malpighian  vessels  are  certainly  not 
homologous^  with  annelidan  nephridia,  because  they  are  strictly 
ectoblastic  and  are  not  segmental.  Their  resemblance  to  the 
sericteries  and  tracheae  is  only  a  very  general  one  in  that  all  of 
these  are  ectoblastic  invaginations,  so  that  at  the  most  we  mtist 


i9o8]  EXCRETORY   ORGANS  OF  METAZOA.  689 

conclude,  with  Palmenj  that  while  these  may  all  have  had  an  essen- 
tially similar  beginning^no  one  of  them  has  been  derived  from  the 
others.  The  Malpighian  vessels  may  well  have  been  hypodermal 
glands  that  have  invaginated  with  the  proctodaeum,  and  for  this 
speaks  their  independent  origin  in  the  embryos  of  some  Hymen- 
optera.  In  this  connection  it  is  interesting  to  note  the  conditions 
in  the  larvae  of  Phryganids,  as  described  by  Henseval  ( 1896)  :  here 
there  are  three  pairs  of  ventro-median  glands  (glands  of  Gilson)  ; 
and  Henseval  regards  the  Malpighian  vessels  as  homologous  glands 
of  the  last  segment,  and  the  proctodaeum  as  their  unpaired  portion 
that  has  secondarily  joined  with  the  mid-gut.  If  we  omit  this  ex- 
planation of  the  proctodaeum  as  being  problematical,  the  comparison 
of  Malpighian  vessels  with  segmental  glands  placed  anteriorly  on 
the  hypodermis  might  well  hold.^ 

Homologues  of  Nephridia, — Here  there  are  in  the  first  instance 
the  genital  ducts,  that  develop  as  coelomic  evaginations  (Wheeler, 
18931  Nassonow,  1886)  ;  Wheeler  has  shown  that  all  the  abdominal 
coelomic  sacs  develop  such  peritoneal  funnels,  but  that  only  those 
of  one  particular  somite  reach  the  exterior  and  become  functional 
genital  ducts.  He  also  (1893a)  holds  that  the  oenoc)rtes  represent 
ectoblastic  remains  of  nephridia.  The  prothoracic  gland  of 
Dicranura  has  been  considered  homologous  (Latter,  1897).  Nasso- 
now (1886)  has  concluded  a  like  relation  for  the  head  glands  of 
Campodea,  all  salivary  glands,  the  maxillary  glands  of  Lepisma,  and 
the  extensible  vesicles  of  the  Thysanura;  but  Oudemans  (1887)  and 
Haase  (1889)  combat  this  view  and  regard  the  extensible  glands 
at  least  as  not  nephridial  but  as  respiratory  skin  glands.  Wheeler 
(1893a)  considers  the  fat-body  to  represent  mesoblastic  remains  of 
nephridia;  some  of  its  cells  are  proved  to  be  excretory  (Wheeler, 
Cuenot,  1895,  Bruntz,  1903),  and  Anglas  (1901)  suggests  that 
such  cells  compose  an  "  accumulating  kidney  "  that  functions  during 
the  substitution  of  Malpighian  vessels  in  the  metamorphosis. 

Nephrocytes. — According  to  Bruntz  (1903)  these  cells  are 
labial  in  MachiJis,  and  in  it  as  ih  Lepisma  are  found  also  on  the 
fat-body;  in  larval  Neuroptera  on  the  wing  muscles;  in  Ephemera 

^  Other  ectoblastic  glands  regarded  as  excretory  are  the  segmental  globi- 
form  glands  of  Ocypus  (Georgevitch,  1898). 


690  MONTGOMERY— MORPHOLOGY  OF  THE  [April  14, 

on  the  fat-body;  in  the  Hymenoptera,  Hemiptera  and  Coleoptera 
on  the  pericardium;  in  the  Lepidoptera  usually  dorsal  in  the 
abdomen ;  in  the  Diptera  along  the  heart.  The  pericardial  cells  of 
Cuenot  (1895)  are  perhaps  to  be  reckoned  with  these. 

29.    DiPLOPODA. 

Malpighian  Vessels. — One  pair  proved  to  be  excretory  by 
Kowalevsky  (1896)  and  Bruntz  (1903). 

Homologues  of  Nephridia, — Here  are  to  be  placed  the  genital 
ducts,  that  develop  like  those  of  Peripatus  (Heathcote,  1888) ;  and 
probably  the  salivary  glands  that  are  mesoblastic  in  origin  (Heath- 
cote), and  which  on  account  of  their  closed  end  sacs  are  named 
"  rein  labial  "  by  Bruntz. 

Fat-body  and  nephrocytes  have  been  shown  to  be  excretory 
(Bruntz).* 

30.  Chilopoda. 

Malpighian  Vessels, — There  is  one  pair  of  these  in  all  genera 
(Verhoeff,  1902),  and  they  develop  as  outgrowths  from  the 
proctodaeum  (Sograf,  1883,  Heymons,  1901). 

Homologues  of  Nephridia, — ^The  genital  ducts  are  mesoblastic 
and  to  be  compared  with  nephridia  (Heymons)  ;  and  Herbst  (1891) 
has  described  for  Lithobius  a  pair  of  glands  with  thin-walled  end 
sacs  opening  behind  the  second  maxillae,  and  has  suggested  that  these 
may  be  modified  nephridia.  The  salivary  glands  are  ectoblastic 
and  not  to  be  compared  with  nephridia  (Heymons,  1898). 

31,  Symphyla  (Scolopendrella). 

There  is  one  pair  of  Malpighian  tubules;  the  ventral  sacs  are 
simply  respiratory  skin  glands  (Haase,  1889). 

32.  Pauropoda. 

Malpighian  Vessels, — There  is  one  pair  of  these  in  Eurypauropus 
but  apparently  only  in  the  female  (Kenyon,  1895).  In  Pauropus 
they  are  absent  (Sclimidt,  1895),  ^^^  ^"  this  genus  there  are  groups 
of  cells  in  the  fat-body  that  may  be  excretory  (Kenyon). 

■Haas€  (1889)  has  demonstrated  that  the  ventral  sacs  are  neither  ex- 
cretory in  function  nor  nephridia!  in  origin. 


:9o3.1  EXCRETORY  ORGANS  OF  METAZOA.  591 

33.  XiPHOSURA  (LiMULUS). 

Coxal  Glands. — A  very  thorough  account  has  been  given  by 
Patten  and  Hazen  (1900).  The  adult  gland  consists  of  four 
nephric  lobes  at  the  bases  of  the  second,  third,  fourth  and  fifth 
legs,  respectively,  and  these  are  connected  medially  by  a  stolon  of 
collective  tubules ;  the  duct  lies  dorso-lateral  from  the  latter,  is  much 
convoluted  and  opens  at  the  basis  of  the  fifth  leg.  The  duct  arises 
from  a  plate  of  cells  of  the  somatic  mesoblast  of  the  fifth  somite, 
this  plate  invaginating  to  produce  a  funnel  opening  into  a  thin- 
walled  end  sac  that  represents  the  fifth  ccelomic  sac;  the  distal  end 
of  the  duct  is  formed  by  an  ectoblastic  invagination.  Outgrowths 
of  the  end  sac  finally  unite  with  cell  chains  of  adjacent  nephric  lobes. 
In  each  of  the  six  thoracic  somites  a  mass  of  nephric  cells  arises 
independently  of  the  duct  from  the  somatic  mesoblast,  and  these 
masses,  of  which  the  first  and  sixth  ultimately  disappear,  form  the 
nephric  lobes;  offshoots  from  the  four  persisting  masses  produce 
the  canals  of  the  stolon.  Thus  there  are  in  the  embryo  six  pairs 
of  coxal  glands,  but  only  four  of  them  persist  in  the  adult. 

The  genital  ducts  arise  as  deverticula  of  the  opercular  meso- 
blastic  sacs,  and  are  to  be  compared  with  nephridia  (Patten  and 
Hazen). 

34.  Arachnida. 

(i)  Araneida, 

Malpighian  Vessels, — These  are  excessively  dendritic  and  their 
delicate  end  branches  form  a  fine  felt-work  around  the  liver  lobes ; 
by  a  pair  of  main  ducts  these  open  into  the  intestine  just  anterior 
to  the  rectal  vesicle.  They  have  been  proved  to  be  excretory 
(Marchal,  1889,  Bruntz,  1903).  Balfour  (1880)  and  Morin 
(1888)  described  them  as  arising  from  the  ectoblastic  proctodeum; 
but  with  the  exception  of  Kishinouye  (1890,  1894)  who  derived 
them  from  the  mesoblast,  the  other  embryologists  (Lx)man,  1887, 
Schimkewitsch,  1897)  find  that  they  develop  from  the  entoblastic 
mid-gut.  Locy  (1886)  described  them  as  coming  from  the  prester- 
coral  tube,  but  though  the  latter  is  probably  entoblastic  its  origin 
was  not  definitely  settled.  Renewed  investigation  is  needed  on  this 
question,  but  the  entoblastic  origin  seems  to  be  best  authenticated. 


592  MONTGOMERY— MORPHOLOGY   OF  THE     ,  [Apr.l  m. 

Coxal  Glands. — Evidently  these  are  not  functional  but  are  de- 
generate in  the  adult;  Bruntz  (1903)  has  proved  they  are  excretory. 
In  the  young  of  Atypus  there  is  a  pair  of  these  opening  on  the  third 
coxae  (Sturany,  1891),  but  the  duct  is  lacking  in  the  adult  (Sturany, 
Bertkau,  1885).  In  the  young  of  Mygale  Lx)man  (1888)  states  it  is 
degenerate,  while  Pelseneer  (1885)  finds  no  ducts  but  on  each  side 
of  the  body  a  four-lobed  gland  corresponding  to  the  four  extremities 
of  the  thorax.  Sturany  and  Hansen  and  Sorensen  (1904)  state 
that  in  the  Tetrapneumones  it  opens  behind  the  fifth  extremity 
(third  leg)  and  in  the  Dipneumones  behind  the  third  (first  leg). 
Kishinouye  (1890)  maintained  that  these  organs  arise  from  the 
ectoblast,  though  he  showed  that  the  anlage  opens  by  a  funnel  into 
the  coelom. 

Genital  Ducts. — Purcell  (1895)  ^^s  shown  that  these  arise  as 
evaginations  of  the  coelomic  sacs ;  "  the  similarity  of  their  develop- 
ment with  that  of  the  coxal  glands  in  Arachnids  generally  indicates 
their  nephridial  origin." 

Hind-gut. — This  is  said  to  serve  as  an  excretory  organ  until  the 
Malpighian  vessels  are  developed  ( Bertkau ).• 

(2)  Scorpionidea. 

Malpighian  Vessels. — These  are  branched,  four  in  number 
(Dufour,  1854)  ;  though  generally  supposed  to  have  the  same  func- 
tion as  those  of  other  arachnids  they  are  stated  by  Bruntz  (1889) 
to  be  not  urinary.    They  arise  from  the  entoblastic  mid-gut  (Brauer, 

.1895). 

Homologues  of  Nephridia. — The  genital  ducts  develop  like  and 

are  homodynamous  with  the  coxal  glands   (Brauer,  1895).      The 

latter  are  in  one  pair  and  open  behind  the  fifth  extremity  (third 

leg)  ;  Bruntz  has  shown  that  they  have  an  excretory  function.    These 

have  each  a  narrow  duct  and  an  enlarged  inner  end  sac.     Bernard 

(1893)  held  these  glands  to  be  ectoblastic,  independent  of  the  coelom, 

homologues   of   acicular  glands.      But  the   researches   of   Laurie 

(1890),  Sturany  (1891)  and  Brauer  (1895)  have  demonstrated  that 

they  arise  each  as  an  outpushing  of  the  somatic  mesoblast  that 

•The  spinning  glands  are  ectoblastic,  and  may  be  equivalent  to  crural 
glands,  but  are  neither  excretory  nor  nephridial. 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  693 

reaches  to  and  opens  upon  the  skin,  then  later  loses  this  opening ; 
Brauer  found  that  a  series  of  them  arise,  in  segments  third  to  sixth, 
inclusive,  but  that  all  but  those  of  the  fifth  segment  soon  disappear. 

(3)  Cyphophthalmidea, 

Malpighian  Vessels, — There  is  one  pair  of  these  in  Gibocellum, 
opening  at  the  junction  of  the  mid-gut  and  hind-gut;  they  are  of 
great  size  and  each  is  remarkable  in  having  a  net-like  branching  at 
its  middle  only  (Stecker,  1876). 

Coxal  Glands, — Sturany  (1891)  holds  what  Stecker  called 
"  Speicheldriisen  "  to  be  probably  coxal  glands ;  there  is  one  pair 
of  them  on  the  sides  of  the  stomach. 

(4)  Phalangida, 

There  are  here  no  Malpighian  vessels,  and  their  absence  is  due, 
according  to  Loman  (1888),  to  the  functional  persistence  of  the 
coxal  glands.  The  latter  are  organs  with  an  inner  closed  end  sac 
(Faussek,  1892),  that  open  in  the  Opiliones  laniatores  behind  the 
third,  and  in  the  Opiliones  palpatores  and  Chelonethi  behind  the 
fifth  extremity.  They  develop  as  mesoblastic  outgrowths  of  the 
particular  extremities  in  which  they  are  placed  (Sturany,  1891, 
Faussek,  1892). 

(5)  Pseudoscorpionidea  {Chernetidce). 

Here  also  there  are  no  Malpighian  vessels.  The  coxal  glands 
are  stated  to  have  no  exterior  openings,  to  lie  at  the  base  of  the  fifth 
extremity,  and  to  be  of  mesoblastic  (nephridial)  origin  (Sturany, 
1891).  The  spinning  glands  that  have  two  pairs  of  opening  on  the 
chelicera  are  considered  by  Bertkau  (1888)  to  be  homologous  with 
them. 

(6)  Solifugce  (Galeodida). 

Malpighian  Vessels, — These  are  one  pair  of  branched  tubes. 

Coxal  Glands, — ^There  is  one  pair  placed  between  the  third  and 
fourth  coxae;  Bernard  (1893)  considered  the  end  sacs  to  be  pro- 
longations of  the  ducts,  but  his  account  is  not  convincing.  Loman 
(1888)  has  suggested  that  the  poison  glands  are  homologous  with 
them. 


594  MONTGOMERY— MORPHOLOGY  OF  THE  [April  a4. 

(7)  Microthelyphonida  (Palpigradi). 

There  are  no  Malpighian  vessels  but  the  adult  excretory  organs 
are  the  coxal  glands,  and  have  been  described  by  Rucker  (1901) 
and  Bomer  (1904).  There  is  one  pair  of  these  extending  forward 
from  the  third  abdominal  segment  to  their  opening  between  the 
second  and  third  legs;  the  great  size  of  these  Bomer  gives  as  the 
explanation  for  the  loss  of  Malpighian  vessels. 

(8)  Pedipalpi  (Thelyphonida), 

Malpighian  Vessels. — According  to  Bomer  (1904)  there  is  one 
very  ramose  pair  of  these;  they  develop  from  the  entoblastic  ster- 
coral pocket  near  its  posterior  end  (Laurie,  1894). 

Coxal  Glands. — These  are  strongly  developed,  function  in 
postembryonic  life,  and  their  ducts  open  on  the  third  pair  of  coxae 
(Borner). 

(9)  Acartna. 

My  account  of  this  group  is  necessarily  very  defective  because 
for  the  most  part  I  have  seen  only  reviews  of  the  literature. 

Malpighian  Vessels. — These  seem  to  be  absent  in  many  species, 
but  a  pair  of  them  has  been  described  for  Ixodes  (Wagner,  1894), 
Gamasidae  (Michael,  1892,  Winkler,  1888),  Halarachne  (Kraemer, 
1885),  and  Tyroglyphfdae  (Nalepa,  1884,  1885,  Haller,  1880).  In 
the  nymphs  of  Gamasids  these  penetrate  deep  into  each  leg.  For 
Bdella  Karpelles  (1893)  ^^^  described  an  unpaired  excretory  organ 
of  entoblastic  origin  opening  into  the  rectum. 

Caudal  (Proctodceal)  Excretory  Organs. — ^These  are  urinary 
structures  opening  at  the  posterior  end  of  the  trunk  without  con- 
nection with  the  mid-gut,  and  are  tubular  or  saccular,  closed  in- 
ternally. These  may  be  present  ( i )  when  the  intestine  is  provided 
with  an  anus,  as  in  Hydrodroma  (Schaub,  1888)  ;  or  (2)  when  the 
mid-gut  ends  blind  and  has  no  anus,  as  in  Prostigmata  (Thor, 
1904),  Gamasidae  (Michael,  1892,  1895),  and  Trombidium  (Crone- 
berg,  1879,  Henking,  1882).  The  suggestion  was  made  by  Thor 
that  the  second  type  probably  represents  a  rectal  bladder  with 
Malpighian  vessels  that  have  become  separated  from  the  mid-gut. 
But  the  first  type,  that  has  an  opening  separate  from  the  anus,  can- 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  595 

not  have  been  so  formed,  but  would  rather  seem  to  be  ectoblastic 
like  the  Malpighian  vessels  in  Insects. 

Unicellular  Glands  of  the  Intestine. — Nalepa  (1888)  has  de- 
scribed for  Phytopids  three  large  unicellular  glands  in  connection 
with  the  rectum,  and  supposed  they  may  be  excretory. 

Coxal  Glands, — In  Limnocharis  Thon  (1905)  found  a  pair  of 
glands  in  the  region  of  the  second  coxae;  in  Eulais  they  are  most 
active  in  the  nymphal  stage  while  they  degenerate  in  the  adult  (by 
substitution  of  the  proctodaeal  organ),  but  in  Limnochares  they 
function  even  in  late  life.  Supposed  coxal  glands  have  also  been 
described  by  With  (1904)  for  the  Notostigmata,  by  Sturany  (1891) 
for  Trombidium,  by  Winkler  (1888)  for  Gamasidae,  and  by  Michael 
(1883)  for  Oribatids.  The  lateral  abdominal  glands  of  Gamasids, 
Tyroglyphids  and  Oribatids  may  be  homodynamous.  The  develop- 
ment of  these  various  glands  seems  to  be  quite  unknown,  so  that 
nothing  can  be  said  of  their  homologies. 

35.  Leptocardii.*® 

The  nephridia  in  Amphioxus  were  discovered  by  Weiss  (1890) 
and  particularly  described  by  Boveri  (1892).  The  latter  found 
them  to  be  segmentally  arranged,  in  about  ninety  pairs  in  the 
branchial  region,  there  being  one  pair  to  every  two  branchial  arches. 
Each  nephridium  was  described  by  Boveri  as  a  canal  with  one  open- 
ing into  the  ectoblastic  atrium,  and  several  into  the  coelom  (sub- 
chordal  cavity)  ;  inserting  into  the  orifice  of  each  of  these  nephro- 
stomes,  but  not  into  that  of  the  nephridiopore,  is  a  tuft  of  long 
Fadenzellen.  Goodrich  (1902)  has  reinvestigated  these  organs, 
and  while  he  confirmed  the  preceding  account  in  most  particulars, 
he  found  that  the  Fadenzellen  are  solenocytes,  each  hollow  with  a 
long  cilium  and  each  closed  from  the  body  cavity,  and  that  there 
are  no  open  communications  of  nephridia  with  the  coelom :  **  These 
tubules  are  situated  '  morphologically*  outside  the  coelom,  being 
covered  with  coelomic  epithelium;  the  solenocytes  alone  push 
through  into  the  coelomic  cavity."     And  he  concluded  "  that  in  their 

"The  Leptocardii  exhibit  so  many  morphological  peculiarities  that  they 
are  to  be  removed  from  the  group  of  the  Vertebrata;  the  Craniota  by  them- 
selves compose  a  homogeneous  assemblage. 


596  MONTGOMERY— MORPHOLOGY   OF  THE  [April  .4. 

segmental  arrangement,  in  their  function,  and  in  their  histological 
structure,  the  excretory  organs  of  Amphioxus  and  the  nephridia  of 
Phyllodoce  are  in  all  essentials  identical."  In  a  second  communica- 
tion Boveri  (1904)  maintained  the  occurrence  of  true  nephrostomes, 
and  held  the  solenoc)rtes  to  be  modified  peritoneal  cells  and  not  to  be 
covered  by  a  peritoneal  investment. 

Unfortunately  nothing  is  known  of  the  development  of  these 
structures. 

36.  Vertebrata  (Craniota). 

With  regard  to  the  excretory  organs  of  this  group  I  shall  deal 
reather  summarily,  because  they  have  been  much  more  studied  than 
the  excretory  organs  of  other  animals,  and  because  most  of  the  larger 
contributions  on  the  subject  deal  extensively  with  the  literature. 

Nephridia. — Qood  reviews  of  the  embryogeny  of  these  structures 
have  been  presented  particularly  by  Riickert  (1892),  Boveri  (1892), 
Wheeler  (1899)  and  Brauer  (1902).  There  are  three  kidney  sys- 
tems which  occur  in  the  ontogeny  in  the  order  of  their  naming;  the 
pronephros,  mesonephros  and  metanephros.  The  first  two  occur 
in  all  vertebrates,  the  third  in  amniotes  only.  The  pronephros  is 
purely  an  embryonic  structure  except  in  Bdellostoma,  Lepidosteus 
and  some  Teleosts  {e,  g.,  Fierasfer)  in  which  it  functions  also  in 
the  adult.  The  mesonephros  is  the  adult  kidney  of  all  other  anam- 
niotes,  and  the  metanephros  of  the  amniotes.  All  these  organs  are 
paired  and  segmented. 

Pronephros. — This  develops  in  the  anterior  trunk  segments  as 
serial  solid  thickenings  of  the  somatic  mesoblast,  each  of  which 
secondarily  becomes  tubular  and  pushes  towards  and  opens  into  the 
coelom.  Their  lateral  ends  unite  to  form  the  collecting  tubule.  The 
arterial  connection  is  in  most  cases  by  a  paired  glomus,  an  unseg- 
mented  vascular  inpushing  of  the  dorsal  peritoneum  medial  from  and 
opposite  the  nephrostomes.  The  duct,  generally  known  as  the  seg- 
mental duct,  also  as  the  pronephric  or  Wolffian  duct,  arises  just 
lateral  from  the  tubules  and  grows  back  from  them  to  open  into  the 
cloaca;  in  the  Selachii  and  Mammals,  possibly  also  in  Lepidosteus, 
it  is  ectoblastic  and  joints  secondarily  with  the  tubules ;  in  all  other 
forms  it  arises  from  the  somatic  mesoblast  in  conjunction  with  the 


i9o8.j  EXCRETORY  ORGANS  OF  METAZOA.  597 

tubules  and  like  them  is  at  first  solid.  Some  of  the  more  impor- 
tant papers  on  the  development  of  these  structures  are  the  follow- 
ing: for  the  Amphibia,  Fiirbringer  (1878),  MoUier  (1890),  Field 
(1891),  Semon  (1891),  and  Brauer  (1902);  for  the  Cyclostomes, 
Wheeler  (1899),  Price  (1897);  for  the  Selachii,  Balfour  (1881), 
Van  Wyhe  (1889),  Riickert  (1888),  Rabl  (1896)  ;  for  the  Teleostei, 
HoflFmann  (1886),  Henneguy  (1888),  H.  V.  Wilson  (1891),  Swaen 
and  Brachet  (1901) ;  for  the  Ganoidei,  Parker  and  Balfour  (1882), 
Beard  (1889)  ;  for  the  Reptiles,  Hoffmann  (1889),  Gregory  (1900)  ; 
for  Aves,  Sedgwick  (1881),  Balfour  (1881),  Renson  (1883),  Felix 
(1891);  and  for  the  Mammals,  Spee  (1884),  Flemming  (1886), 
KoUman  (1891),  Martin  (1888). 

Mesonephros, — These  tubules  develop  usually  in  the  segments 
behind  the  pronephroi,  but  there  are  certain  segments  that  may  con- 
tain both  of  them,  and  they  are  more  numerous  and  more  diflfer- 
entiated  that  the  pronephroi.  To  understand  their  origin  it  is  neces- 
sary to  recall  that  the  coelom  becomes  divided  into  the  dorsal 
myoccels  (cavities  of  the  myotomes  or  somites),  the  middle  neph- 
rocoels,  both  of  these  being  segmented  and  paired,  and  the  large 
unsegmented  hypocoel  that  is  imperfectly  paired;  these  relations 
were  established  particularly  by  Van  Wyhe.  Very  early  the 
myoccels  pinch  oflF  from  the  nephrocoels,  whereby  the  latter  are  left 
as  short  tubes,  the  dorso-lateral  end  of  each  ending  blindly  while 
the  ventral  opens  into  the  hypocoel.  These  peritoneal  nephrocoels 
become  the  mesonephroi  and  grow  laterad  to  join  with  and  open 
into  the  segmental  duct,  for  they  develop  no  duct  of  their  own. 
The  arterial  connection  is  segmental :  From  the  aorta  a  vessel  grows 
towards  each  tubule  and  ends  in  a  capillary  glomerulus  against  the 
wall  of  the  latter  above  the  nephrostome;  the  wall  of  the  tubule 
forms  a  partial  sheath  (capsule  of  Bowman)  around  the  glomerulus, 
In  Petromyzon  there  is  a  larval  as  well  as  a  definitive  set  of  these 
tubules,  and  there  may  be  several  in  each  segment  (Wheeler). 

The  principal  studies  on  the  mesonephros  are  these :  For  Selachii, 
Ruckert  (1888),  Van  Wyhe  (1889),  Rabl  (1896);  for  Teleostei, 
Felix  (1897);  for  Cyclostomata,  Wheeler  (1899),  Price  (1897), 
Maas  (1897);  for  Amphibia,  Semon  (1891),  Brauer  (1902),  Hall 
(1904) ;  for  Reptiles,  Gregory  (1900),  Mihalkovics  (1885),  Wieder- 


598  MONTGOMERY— MORPHOLOGY   OF  THE  [Aprils. 

sheim  (1890);  for  Aves,  Sedgwick  (1880),  Felix  (1891)  ;  and  for 
Mammals,  Janosik  (1887),  Martin  (1888),  H.  Meyer  (1890). 

Metanephros  {Kidney  of  Amniotes), — This  consists  of  the  duct 
or  ureter,  and  the  kidney  proper,  both  developing  behind  the  meso- 
nephros.  The  ureter  is  a  dorsal  outgrowth  from  the  segmental  duct. 
There  are  two  views  concerning  the  origin  of  the  glandular  kidney. 
According  to  the  first  and  older  of  these  the  kidney  tubules  arise 
as  evaginations  from  the  anterior  end  of  the  ureter  (KoUiker,  1861, 
Waldeyer,  1870).  There  is  much  more  evidence  for  the  second 
view,  origin  independent  of  the  ureter  from  mesoblastic  tissue 
(Emery,  1883,  Hoffmann,  1889,  Wiedersheim,  1890).  The  ureter 
grows  forward  into  an  embryonic  cell  mass  known  as  the  kidney 
blastema,  of  somewhat  uncertain  origin,  but  possibly  homodynamous 
with  the  anterior  mesonephric  anlage  (Wiedersheim).  According 
to  the  description  of  Emery  (1883)  the  so-called  collective  tubules 
of  the  kidney  arise  as  bhnd  outgrowths  of  the  ureter,  and  these  join 
with  the  secretory  tubules  that  arise  independently  from  the  kidney 
blastema.  There  is  still  much  to  be  decided  concerning  the  exact 
method  of  formation  of  the  kidney,  but  certainly  a  considerable 
portion  of  it  arises  independent  from  the  ureter  from  somatic  meso- 
blast.  Each  tubule  of  the  metanephros  commences  proximally  with 
a  Malpighian  corpuscle,  that  is,  a  vascular  glomerulus  enclosed  in 
a  capsule  of  Bowman,  a  vascular  relation  like  that  of  the  meso- 
nephroi;  metanephric  tubules  lack  nephrostomes  or  other  connec- 
tions with  the  coelom.^* 

Relations  of  these  Nephridial  Systems. — That  the  pronephros 
and  mesonephros  are  homodynamic  is  the  view  of  Balfour  (1881), 
Sedgwick  (1881),  Price  (1897)  and  Brauer  (1902).  '  Field  (1891) 
argued  that  the  two  are  differentiated  parts  of  one  ancestral  organ, 
that  differ  structurally  because  they  develop  at  different  periods. 
But  the  majority  of  investigators  hold  them  to  be  not  homodynam- 
ous, and  here  may  be  mentioned  W.  Miiller  (1875),  Fiirbringer 
(1878),  Van  Wyhe  (1889),  Riickert  (1892),  Semon  (1891),  Rabl 
(1896),  Wheeler  (1899),  and  Maas  (1897).  If  we  omit  the  con- 
ditions in  the  Gymnophiones  in  which  the  relations  of  the  pronephros 

"Adult  mesonephric  tubules  may  still  maintain  their  nephrostomes,  or 
may  lose  them ;  cf .  Spcngel,  1876. 


i^]  EXCRETORY  ORGANS  OF  METAZOA.  599 

appear  strongly  modified  or  at  least  quite  different  from  those  in 
other  groups,  then  it  is  highly  probable  that  these  two  organ  systems 
are  not  strictly  homodynamous.  For  the  pronephroi  arise  as  solid 
thickenings  of  the  somatic  mesoblast,  that  later  become  tubular  and 
only  secondarily  join  with  the  ccelom;  and  their  vascular  supply  is 
an  uns^mented  glomus  opposite  their  nephrostomes.  On  the  other 
hand  the  mesonephroi  are  abstricted  portions  of  the  ccelom  (nephro- 
coels),  they  are  from  the  start  peritoneal  and  in  open  communication 
with  the  ccelom;  and  the  vascular  connection  of  each  is  a  Malpighian 
corpuscle.  The  pronephroi  are  retroperitoneal,  the  mesonephroi, 
peritoneal  funnels  in  the  main;  the  former  develop  in  close  con- 
nection with  the  segmental  duct,  while  the  latter  arise  much  later 
than  it  and  join  it  secondarily.  In  view  of  these  differences  pro- 
nephros and  mesonephros  are  probably  only  incompletely  homo- 
dynamous. 

As  to  the  metanephros,  its  ureter  being  an  outgrowth  of  the 
segmental  duct  is  a  new  structure ;  while  the  glandular  kidney  arises 
from  mesoblast  that  may  represent  a  late  generation  of  mesonephric 
tubules.  Accordingly,  the  metanephros  can  be  only  in  part  homo- 
dynamous with  the  mesonephros. 

Homologues  of  Nephridia, — Here  are  to  be  placed  the  genital 
organs  that  I  will  treat  very  briefly.  Particular  genital  ducts  are 
absent  in  the  Cyclostomes,  Lcemargus  and  certain  Teleostei;  here 
the  genital  cells  fall  into  the  ccelom  and  are  discharged  through 
peritoneal  canals,  supposed  peritoneal  funnels  (Weber,  1886),  the 
development  of  which  has  not  been  studied. 

In  the  males  of  Teleosts  and  certain  other  fishes  the  genital 
ducts  are  simply  outgrowths  of  the  gonads,  while  in  all  other  forms 
the  segmental  ducts  (or  portions  of  them)  are  urogenital.  The  vasa 
efferentia  of  the  testis,  the  paradidymis  and  the  hydatid  of  Morgagni 
are  modified  mesonephric  tubules. 

In  the  females  of  all  forms  except  most  Teleosts  and  Lepidosteus, 
where  the  ducts  are  outgrowths  of  the  gonads,  the  oviduct  (with 
uterus  when  present)  is  distinct  from  the  urinary  canal  (segmental 
duct  or  ureter)  and  is  known  as  the  Miillerian  duct.  This  is  paired 
and  arises  in  the  Selachii  as  a  longitudinal  abstriction  of  the- seg- 
mental duct,  but  in  other  forms  as  a  structure  independent  of  the 


600  MONTGOMERY— MORPHOLOGY  OF  THE  [Apiila4, 

latter,  i.  e.,  as  a  longitudinal  peritoneal  groove,  showing  sometimes 
(Reptiles)  traces  of  segmental  origin,  that  becomes  a  tube  closed 
from  the  coelom  except  at  its  anterior  end  (ostium).  These  two 
kinds  of  Miillerian  ducts  cannot  be  homologized,  for  the  first  is  an 
abstriction  from  the  segmental  duct,  while  the  second  arises  as  a 
peritoneal  infolding  and  may  be  compared  with  an  elongated  peri- 
toneal funnel  or  with  a  series  of  them.  The  ovaries  diflfer  from 
testes  in  lacking  vasa  elferentia  connecting  them  with  the  ducts,  but 
other  remnants  of  mesonephric  tubules  are  found  in  amniotes  In 
form  of  the  epoophoron  and  paroophoron. 

Other  Excretory  Organs. — The  liver  forms  urea,  while  the 
sudoriparous  glands,  respiratory  organs  and  skin  aid  in  the  dis- 
charge of  waste  substances. 

B.    GENERAL  COMPARISONS. 

I.  Main  Types  of  Excretory  Organs. 

We  use  the  idea  homology  to  denote  that  relation  between  a 
certain  organ  of  one  animal  and  a  certain  organ  of  another,  which 
is  dependent  upon  derivation  from  a  common  ancestral  organ.  In 
other  words,  homology  denotes  community  of  descent  of  parts. 
To  elucidate  such  relations,  to  demonstrate  change  of  both  form 
and  use  of  parts,  is  the  first  object  of  comparative  anatomy;  later  all 
such  knowledge  may  be  so  compounded  as  to  give  the  general  his- 
tory of  phylogeny.  When  one  considers  such  manifold  and  diverse 
organs  as  those  that  subserve  excretion,  difficulties  of  interpretation 
that  are  almost  insuperable  arise  to  perplex  and  bewilder,  yet  at 
the  same  time  compel,  the  attention.  Any  conclusions  with  regard 
to  the  homologies  of  these  organs  must  be  tentative  because  our 
knowledge  of  them  is  so  very  imperfect;  in  fact  for  most  of  the 
animal  groups  only  the  outlines  have  been  made  known.  Therefore 
the  following  attempt  to  arrange  the  excretory  organs  according 
to  their  genetic  relations  should  be  regarded  as  only  an  essay. 

The  criteria  of  homology  are  still  a  matter  of  dispute.  I  have 
discussed  this  matter  in  another  place  (1906),  and  will  simply  state 
here  that  similarity  of  relative  position  to  other  parts  seems  to  be 
the  surest  criterion,  together  with  general  similarity  in  mode  of 


I908.I  EXCRETORY   ORGANS  OF  METAZOA.  601 

ontogenetic  formation.  We  shall  place  first  relative  position  with 
regard  to  the  outer  skin,  the  blastocoel  and  coelom,  the  intestine  and 
the  genital  organs.  These  relations  involve  genetic  connections 
with  the  particular  germ  layers,  and  a  word  of  discussion  may  be 
in  place  with  regard  to  these.  The  concept  of  the  essential  homol- 
ogy of  the  primary  germ  layers  has  been  many  times  attacked  since 
its  formulation  by  Huxley  and  Kowalevsky.  Yet  these  objections 
have  been  weakened  by  much  of  the  more  recent  work.  Ectoblast 
always  furnishes  nervous  elements,  entoblast  originates  digestive 
and  assimilative  parts,  from  the  mesoblast  come  the  reproductive 
cells;  these  are  cardinal  distinctions  that  seem  to  hold  throughout 
the  Metazoa.  Therefore  it  is  no  valid  objection  to  the  idea  of  the 
homology  of  these  layers  to  cite  the  observations  of  Chun  on 
Ctenophores,  that  in  the  process  of  gemmation  an  ectoblastic  out- 
pushing  gives  rise  to  both  ectoblast  and  entoblast.  This  observa- 
tion can  rather  prove  only  that  such  an  ectoblastic  bud  is  not  purely 
ectoblastic  but  mixed  in  its  nature.  And  when  He)mion's  studies 
on  Insects,  resulting  in  the  completely  ectoblastic  formation  of  the 
whole  intestine,  are  brought  up  as  an  objection,  it  may  be  answered 
that  the  observational  distinction  of  the  germ  layers  in  insects  is 
very  difficult,  and  also  that  these  conclusions  have  not  been  corro- 
borated by  all  subsequent  examiners.  The  oft-cited  case  of  the 
Trematodes,  to  the  effect  that  the  embryo  throws  off  its  whole  ecto- 
blast, must  now  be  allowed  to  drop  since  Goldschmidt  has  demon- 
strated that  it  is  not  the  true  ectoblast  but  only  a  follicle  cell  layer 
that  becomes  so  moulted.  For  these  and  other  reasons  those  critics 
are  becoming  fewer  who  maintain  that  ectoblast  is  not  always 
homologous  with  ectoblast,  and  entoblast  with  entoblast  throughout 
the  Metazoa;  and  the  most  painstaking  of  all  embryological  work, 
that  on  cell-lineage,  bears  out  most  strongly  the  well-founded  general 
homologies  of  these  primary  layers.  The  discussion  has  shifted 
rather  to  the  significance  of  the  mesoblast,  the  existence  of  which 
was  so  stoutly  denied  by  Kleinenberg.  This  long  and  wearying  dis- 
cussion has  brought  out  the  result,  first  clearly  stated  by  Meyer, 
that  two  kinds  of  mesoblast  are  to  be  sharply  distinguished,  the 
primary  or  mesectoblast,  and  the  secondary  or  mesentoblast.     The 

PROC.  AMER.  PHIL.  SOC.,  XLVIl     I90  MM,  PRINTED  JANUARY  I4,  I909. 


602  MONTGOMERY— MORPHOLOGY  OF  THE  [April  m. 

probable  correctness  of  this  distinction  is  amply  substantiated  by 
the  cell-lineagists,  and  the  arguments  for  it  have  been  well  presented 
by  Torrey.  The  mesectoblast  is  of  ectoblastic  origin,  it  is  in  part 
equivalent  to  the  mesenchyme  of  the  Hertwigs ;  it  forms  larval  and 
to  less  extent  adult  structures,  but  never  gives  rise  to  germ  cells. 
The  mesentoblast  form  adult  structures  and  contains  the  germ  cells. 
These  again  are  fundamental  differences,  so  that  it  is  no  longer 
sufficient  to  state  a  part  is  mesoblastic,  it  is  necessary  to  know 
whether  it  is  mesectoblastic  or  mesentoblastic.  The  mesectoblast  is 
in  reality  an  emigrant  or  delaminant  of  the  ectoblast,  it  is  genetically 
related  with  that  layer  and  not  with  the  mesoblast. 

Relation  of  position  to,  and  origin  from,  these  four  embryonic 
layers  gives  then  a  primary  criterion  for  deciding  the  homologies  of 
the  excretory  organs.  And  these  relations  of  position  involve  also 
place-relations  with  regard  to  the  primary  cavities  of  the  body :  The 
blastocoel,  the  space  between  ectoblast  and  mesoblast ;  the  ccelom,  the 
space  lined  by  mesentoblast;  and  the  gastroccel,  the  space  lined  by 
entoblast. 

Using  the  relations  of  position  and  origin  as  of  primary  im- 
portance, and  anatomical  and  histological  relations  as  of  secondary, 
we  will  proceed  to  arrange  the  excretory  organs  in  genetic  groups. 
Many  of  the  organs  described  in  the  preceding  part  of  this  paper 
could  not  be  entered  here  on  account  of  the  insufficiency  of  our 
knowledge  concerning  them;  and  some  others  have  to  be  marked 
doubtful  for  the  same  reason.  It  is  at  the  best  a  hazardous  under- 
taking to  classify  other  men's  results,  and  the  danger  is  multiplied 
when  descriptions  are  imperfect." 

(a)  Wholly  Ectoblastic  Excretory  Organs,  not  Opening  into  the 
Cwlom  and  not  Serving  as  Genital  Ducts, 

I.  Hypodermal  skin  glands.  These  are  perhaps  the  most 
primitive  excretory  organs,  and  are  of  wide  distribution.  Excre- 
tory function  of  them  has  been  proved  for  Pycnogonids,  Insects, 
Arachnids,  Vertebrates  and  certain  others ;  but  probably  most  hypo- 
dermal  glands  are  rather  secretory  than  excretory. 

"Here  may  be  mentioned  a  generalized  embryonic  excretory  organ,  the 
blastocoel,  which  Kofoid  has  shown  to  have  the  value  of  a  discharging  vesicle 
and  to  continue  that  function  up  to  the  gastrula  stage ;  Meisenheimer  has  ac- 
cepted Kofoid*s  conclusions. 


i9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  603 

2.  Evaginated  vesicles,  open  to  the  blastocoel.  Here  are  to  be 
reckoned  the  outer  nephridia  of  prosobranch  and  pulmonate  mol- 
luscan  embryos,  and  probably  the  anal  kidneys  of  opisthobranchs. 
The  latter  have  a  method  of  formation  similar  to  that  of  the  others, 
but  th^  differ  in  position. 

3.  Tubular  invaginations  terminating  blindly  in  flame  cells,  with 
the  cavities  of  at  least  the  capillaries  intracellular.  Their  origin 
from  the  ectoblast  has  been  proved  only  in  the  case  of  the  Nemertini 
and  Acanthocephala  and  with  some  doubt  in  the  Polycladidea.  Here 
are  to  be  placed  the  definitive  nephridia  of  the  Platodes,  Nemertini, 
Gastrotricha,  Rotatoria,  Rhodope,  Acanthocephala,  and  the  larval 
nephridia  of  Phoronis;  probably  those  of  the  Endoprocta  should  be 
placed  here  (if  they  are  not  mesectoblastic),  and  perhaps  those  of 
the  Priapulida  and  the  head  kidneys  of  some  Molluscan  larvae. 
This  type  of  excretory  organ  has  been  named  by  Hatschek  (1888) 
protonephridium,  though  he  extended  this  term  to  cover  also  organs 
of  mesectoblastic  and  even  mesentoblastic  origin.  This  is  a  very 
natural  group  of  excretory  ofgans,  showing  great  similarity  in  both 
structure  and  development.  The  only  case  of  a  larval  or  head 
kidney  among  them  is  that  of  Phoronis,  yet  here  this  kidney  persists 
into  the  adult  though  it  later  joins  with  a  coelomostome.  Kaiser 
(1892)  is  inclined  to  compare  the  organs  of  the  Acanthocephala 
with  those  of  Annelids  or  even  with  the  anal  kidneys  of  Bonellia, 
but  their  strictly  ectoblastic  origin  renders  this  view  unlikely ;  while 
those  of  the  Acanthocephala  open  into  the  genital  ducts,  so  also  do 
those  of  certain  Turbellaria,  consequently  this  relation  does  not 
speak  against  their  community. 

4.  Tubular  invaginations  with  wholly  intercellular  cavity,  with- 
out flame  cells  of  cilia.  These  are  the  Malpighian  vessels  of 
Insects  and  Chilopods  (?  and  of  other  Myriopods),  the  proctodaeal 
organs  of  the  Acarina,  and  possibly  the  rectal  tubes  of  the  Tardi- 
grada.  All  of  these  either  open  into  the  proctodaeum  or  upon  the 
surface  of  the  body  near  the  anus;  it  is  probable  they  secondarily 
acquired  the  proctodseal  position  when  the  ectoblast  invaginated  to 
produce  the  end-gut.  These  tubes  are  usually  unbranched,  but  in 
some  Insects  they  are  dendritic.  They  differ  from  type  3  mainly  in 
lacking  cilia  and  in  possessing  a  wholly  intercellular  cavity ;  but  the 


604  MONTGOMERY—MORPHOLOGY   OF  THE  [April  .4. 

lack  of  ciliated  epithelia  is  a  histological  characteristic  of  the  groups 
that  possess  them. 

(b)  Mesectoblastic  Organs, 

Here  are  to  placed  the  following  structures : 

5.  Scattered  excretory  cells,  such  as  connective  tissue  elements 
of  the  Mollusca,  and  possibly  the  bacterioidic  cells  of  the  Oligochaeta. 

6.  Closed  vesicles,  the  kidney  sacs  of  Tunicata,  and  possibly 
the  nephrocysts  of  nudibranch  Mollusca.  These  seem  to  act  as 
centers  of  accumulation  of  waste  substances. 

7.  Tubes  communicating  with  the  exterior,  the  inner  ends  blind 
and  terminating  with  a  flame  cell  or  solenocj'tes.  In  all  probability 
the  larval  nephridia  (head  kidneys)  of  Oligochaeta  and  Polychaeta 
belong  here  (in  the  latter  sometimes  a  portion  of  the  duct  is  strictly 
ectoblastic) ;  possibly  the  nephridia  of  the  Dinophilea  fall  also  into 
this  category,  but  nothing  is  known  as  yet  of  their  development.  In 
their  structure  these  are  very  similar  to  the  organs  of  type  3,  the 
protonephridia  in  the  restricted  sense,  the  only  difference  being  that 
the  one  come  directly  from  the  ect6blast,  the  others  from  the 
mesectoblast. 

(c)  Organs  Wholly  or  Partially  Mesentoblastic. 

These  represent  the  more  specialized  kinds  of  excretory  organs, 
correspond  in  part  to  the  metanephridia  of  Hatschek,  and  may  be 
subdivided  into  the  following  main  t)rpes: 

8.  An  ectoblastic  invagination  joining  directly  (without  partici- 
pation of  retroperitoneal  mesentoblast)  with  a  ccelomostome  (peri- 
toneal funnel),  the  involved  portion  of  the  ccelom  not  exclusively 
excretory.  Examples  are  the  adult  nephridia  of  Phoronis,  and 
the  head  and  collar  pores  of  the  Enteropneusta ;  homologous  with 
these  is  the  stone  canal  of  the  Echinodermata.  The  present  evi- 
dence does  not  allow  us  to  decide  whether  the  segmental  organs  of 
the  Sipunculida,  Ectoprocta,  Brachiopoda,  Echiurids  and  Myzo- 
stomes  belong  with  this  type  or  with  type  11. 

9.  An  ectoblastic  invagination  joining  directly  (without  partici- 
pation of  retroperitoneal  mesentoblast)  with  a  reduced  ccelomic 
sac,  the  latter  being  an  exclusively  excretory  end  sac.     There  are 


' 


,9o8)  EXCRETORY  ORGANS  OF  METAZOA.  605 

two  main  kinds  of  these:  (i)  The  ectoblastic  portion  very  small, 
and  the  end  sac  representing  only  a  portion  of  the  coelom  of  a  seg- 
ment, as  in  the  case  of  the  salivary  glands,  nephridia,  and  genital 
ducts  of  the  Protracheata.  And  (2)  the  ectoblastic  portion  rela- 
tively larger,  the  end  sac  being  a  whole  ccelomic  sac,  as  in  the  case 
of  the  coxal  glands  of  Arachnids,  Xiphosura,  Crustacea,  the  salivary 
glands  of  Diplopods,  and  the  antennal,  maxillary  and  maxillipedal 
glands  of  Crustacea. 

10.  An  ectoblastic  tube  joining  with  retroperitoneal  mesento- 
blast,  the  latter  neither  joined  with  a  ccelomostome  nor  serving  as 
a  genital  duct ;  the  inner  end  is  either  quite  closed  or  else  has  a  small 
opening  (nephridiostome)  into  the  ccelom;  the  cavity  is  usually 
intracellular.  Here  belong  the  larval  nephridia  of  the  Hirudinea, 
and  the  definitive  nephridia  of  the  Hirudinea,  Oligochaeta  and  some 
Polychseta  (Phyllodocidae,  Glyceridae,  Nephthyidae,  Capitellidae, 
and  perhaps  the  Nereidae).  Probably  the  anal  kidneys  of  Echiurids 
belong  here,  and  perhaps  also  the  nephridia  of  the  Nematoda.  In 
essential  agreement  with  this  type  is  the  pronephros  of  the  Verte- 
brata,  which  also  consists  of  a  retroperitoneal  mesentoblastic  tube 
whose  inner  end  opens  secondarily  into  the  coelom  (not  by  a  peri- 
toneal funnel)  and  whose  outer  end  joins  with  the  segmental  duct 
that  is  of  either  mesentoblastic  or  ectoblastic  origin.  Possibly  the 
nephridia  of  the  Leptocardii  are  also  homologous,  as  Boveri  has 
suggested,  but  nothing  is  known  of  their  development;  it  will  be 
recalled  that  Boveri  homologized  the  atrial  chamber  of  the  Lepto- 
cardii  with  the  segmental  duct  of  the  Vertebrata.^'  There  is  no- 
homology  between  the  s^mental  duct  of  Vertebrates  and  the  longi- 
tudinal  canals  of  the  Polychaetes  Lattice  and  Ploitnia,  for  the  latter 
seem  to  be  formed  by  a  late  fusion  of  the  secretory  portions  of  the 

"As  to  the  phylogeny  of  this  segmental  duct,  Balfour  considered  it  to 
be  the  foremost  modified  pronephric  tubule,  and  Field  has  accepted  this 
view.  Haddon  (1886)  and  Beard  (1887)  suggested  that  the  pronephroi  first 
opened  separately  into  an  open  ectoblastic  groove,  that  later  closed  to  become 
the  segmental  duct.  Ruckert  (1888)  also  concluded  that  originally  the  pro- 
nephric tubules  opened  independently  to  the  exterior,  and  that  they  ex- 
tended through  the  whole  trunk;  he  maintained  that  the  segmental  duct 
arose  by  the  meeting  and  fusion  of  their  lateral  ends,  that  is,  by  a  back- 
ward growth  of  collective  tubules. 


606  MONTGOMERY— MORPHOLOGY   OF  THE  [April  24. 

nephridia.     Indeed,  the  segmental  duct  of  Vertebrates  appears  to 
have  originated  in  this  class. 

11.  An  ectoblastic  tube  (though  this  portion  may  be  very  small) 
joining  with  retroperitoneal  entomesoblast,  and  the  latter  con- 
necting with  a  coelomostome ;  these  are  generally  either  urogenital 
or  homodynamous  with  genital  ducts,  and  the  cavity  is  usually 
intercellular.  The  inner  end  is  widely  open  at  least  in  the  embryo. 
These  correspond  to  type  10,  with  the  addition  of  a  coelomostome. 
In  this  type  fall  the  nephridia  of  the  Mollusca,  and  those  of  most 
Polychaeta.  As  mentioned  above,  the  segmental  organs  of  the 
Sipunculida,  Ectoprocta,  Brachiopoda,  Echiurida  and  Myzostomida 
probably  belong  either  here  or  with  type  8.  Essentially  homologous 
are  the  mesonephroi,  therefore  probably  also  the  metanephroi,  of 
the  Vertebrates,  which  consist  to  great  extent  of  peritoneal  funnels. 
And  Boveri  has  argued  that  the  gonads  of  the  Leptocardii  may  be 
homologous  with  these  mesonephroi. 

12.  Non-tubular  peritoneal  differentiations  of  excretory  nature. 
Here  are  the  so-called  ciliated  funnels  of  the  Holothurians,  that 
are  not  funnels  (ccelomostomes)  at  all,  and  the  widely  represented 
peritoneal  glands  (phagocytic  organs,  chloragogue  in  parte). 

13.  Non-tubular  retroperitoneal  mesentoblastic  cell  masses. 
With  these  belong  a  variety  of  structures  the  development  of  most 
of  which  has  been  little  examined,  such  as  the  excretophores  of  the 
Hirudinea  and  the  fat-body  of  Insects  (the  latter  perhaps  repre- 
senting, as  Wheeler  has  suggested,  the  remains  of  nephridia). 

(d)  Entoblastic  Excretory  Organs, 

14.  These  are  relatively  few  in  number  and  seldom  have  an 
exclusively  excretory  function.  In  the  first  place  there  are  tubular 
evaginations  of  the  mid-gut,  as  the  Malpighian  vessels  of  Arachnida, 
then  the  mid-gut  coeca  of  the  Polycladidea  and  Amphipoda  and 
probably  of  the  Arachnida;  these  are  all  essentially  homologous. 
The  whole  mid-gut  has  been  shown  to  be  excretory  in  the  CoUem- 
bola,  Dinophilus  and  the  Ectoprocta;  it  seems  to  be  specially  so 
only  when  other  excretory  organs  are  wanting,  and  in  that  case 
there  is  either  periodical  moulting  of  the  lining  of  the  mid-gut 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  607 

(Collembola),  or  when  this  fails  there  is  rapid  death  of  the  indi- 
vidual from  poisoning  of  the  intestinal  tract  (Ectoprocta). 

2.  Homologies  of  the  Preceding  Types. 

The  entoblastic  type  (14)  is  sui  generis  and  not  related  to  the 
others.  T)rpes  12  (peritoneal  glands)  and  13  (retroperitoneal  dif- 
ferentiations) are  so  generalized  in  both  structure  and  function, 
that  it  is  hardly  advisable  to  attempt  to  draw  homologies  between 
them;  and  the  same  holds  for  types  i  (ectoblastic  skin  glands),  2 
(ectoblastic  vesicles),  5  (scattered  mesectoblastic  cells)  and  6 
(mesectoblastic  vesicles).  There  remain  then  for  consideration  all 
those  distinctly  tubular  organs,  nephridia  proper,  into  the  composi- 
tion of  which  entoblast  does  not  enter.**  The  earliest  and  most 
uniform  of  these  are  those  of  type  3,  ectoblastic  invaginations  ter- 
minating in  flame  cells,  which  are  referable,  as  argued  by  Lang,  to 
still  simpler  skin  glands.  Type  4,  ectoblastic  invaginations  like  3 
but  without  cilia,  are  essentially  similar;  for  no  one  would  hesitate 
to  homologize  the  mid-gut  of  the  Turbellaria  and  the  Insects,  though 
the  former  is  ciliated  and  the  latter  is  not ;  therefore  one  should  not 
object  to  drawing  homology  between  the  water  vascular  system  of 
the  former  and  the  Malpighian  vessels  of  the  latter.  The  lack  of 
cilia  is  not  a  characteristic  merely  of  these  vessels,  it  marks  all  the 
tissues  of  the  Insects.  The  only  differences  between  types  3  and  4 
is  the  lack  of  cilia  in  the  latter,  and  this  is  a  difference  that  is  of 
little  homological  importance,  a  merely  histological  character.  And 
essentially  similar  to  both  of  these  is  type  7,  tubes  of  mesectoblastic 
origin ;  they  do  not  come  immediately  from  the  ectoblast,  but  from 
tissue  of  ectoblastic  derivation  which  is  but  a  step  removed.  These 
three  types,  accordingly,  3,  4  and  7  are  anatomically  and  embryolog- 
ically  essentially  alike,  they  are  to  be  considered  homologous;  they 
stand  in  no  relation  to  the  coelom,  never  conduct  the  genital  prod- 

"The  term  nephridium  has  been  used  very  variously  since  its  coinage 
by  Lankester  (1877).  It  might  be  well  to  limit  it  in  the  future  to  tubular 
excretory  organs  not  containing  entoblast.  In  the  descriptive  part  of  the 
paper  I  have  discussed  special  homologies  of  excretory  organs  within  the 
same  group,  such  as  relations  of  embryonic  to  adult  nephridia,  of  mega- 
nephridia  and  plectonephridia,  homologies  of  tracheae,  etc.;  these  need  not 
be  repeated  here. 


608  MONTGOMERY— MORPHOLOGY  OF  THE  [April  24. 

ucts,  and  contain  no  mesentoblast.  I  would  propose  that  Hatschek's 
(1888)  term  protonephridium  be  limited  to  them. 

From  such  protonephridia  the  other  types  of  nephridia  have 
probably  been  derived  by  the  persistence  of  only  the  discharge  ducts, 
or  portions  of  them,  of  the  former  and  by  the  substitution  of  mesen- 
toblastic  elements  for  their  other  portions.  The  only  elements  of  the 
protonephridia  that  have  been  retained,  it  should  be  repeated,  are 
their  distal  nephridiopores  with  more  or  less  of  the  connectant  dis- 
charge ducts,  while  the  remainder  of  the  protonephridia,  all  the 
excretory  portion  proper,  has  been  replaced  by  mesentoblastic  ele- 
ments. Accordingly,  the  two  other  main  kinds  of  nephridia  of 
which  we  shall  have  to  speak  can  be  at  the  most  compared  only  in 
part  with  these  protonephridia,  only  their  distal  nephridioporal  ends 
can  be  so  compared.  The  more  specialized  kinds  of  nephridia  have 
probably  originated  from  the  protonephridia,  not  as  further  special- 
izations of  them  but  rather  by  addition  of  extraneous  elements; 
on  the  whole  they  are  not  homologous. 

These  more  specialized  nephridia  with  mesentoblastic  consti- 
tuents fall  into  two  main  groups. 

The  first  of  them  consists  of  types  8  and  9,  both  of  which  have 
in  common  the  union  of  an  ectoblastic  duct  with  the  peritoneum  but 
have  no  retroperitoneal  mesentoblast.  They  are  either  urogenital, 
or  are  homodynamous  with  genital  ducts  ( ?also  in  the  Entero- 
pneusta).  Their  main  difference  is  that  in  type  8  the  peritoneal 
invagination  is  more  pronounced  as  a  rule,  and  that  in  type  9  the 
connectant  coelom  has  become  exclusively  excretory.  These  differ- 
ences are  not  important,  and  these  two  types  are  in  general  homol- 
ogous. Until  retroperitoneal  elements  are  discovered  for  them 
they  must  be  considered  distinct  from  the  following;  and  to  them 
the  name  coelonephridium  might  be  given. 

The  second  kind  of  the  more  specialized  nephridia  comprises 
types  10  and  11,  both  characterized  by  the  union  of  ectoblast  with 
retroperitoneal  mesentoblast.  Type  11  differs  from  10  by  the 
addition  of  a  coelomostome  (peritoneal  funnel),  in  the  manner  made 
known  particularly  by  the  studies  of  E.  Meyer  and  Goodrich. 
Their  essential  peculiarity  is  the  retroperitoneal  mesentoblast,  not 
the  peritoneal  funnel.     Hatschek  (1888)  classed  these  together  with 


,9o8]  EXCRETORY  ORGANS  OF  METAZOA.  609 

the  preceding  as  metanephridia,  and  diagnosed  them  by  the  presence 
of  a  coelomostome ;  but  the  difference  with  regard  to  the  retroperi- 
toneal element  seems  to  me  so  important  that  these  should  be  held 
distinct  from  the  preceding,  and  in  that  case  it  would  be  well  to 
limit  the  term  metanephridium  to  types  lo  and  ii. 

The  three  main  kinds  of  nephridia  that  these  considerations  lead 
us  to  distinguish  may  be  briefly  compared  as  follows :  Protonephrid- 
turn  (types  3,  4,  7),  wholly  ectoblastic  or  mesectoblastic  (possibly 
in  some  cases  both  ectoblastic  and  mesectoblastic) ;  ccelonephridium 
(types  8, 9),  distal  ectoblastic  portion  joining  directly  with  a  ccElomo- 
stome;  metanephridium  (types  10,  11),  distal  ectoblastic  portion 
joining  with  retroperitoneal  mesentoblast,  and  the  latter  connecting 
or  not  connecting  with  a  coelomostome.  Only  the  second  and  third 
of  these  ever  serve  as  genital  ducts  or  are  homodynamous  with  them. 
The  metanephridium  is  the  most  complex  because  it  may  consist  of 
as  many  as  three  elements,  and  it  contains  the  smallest  amount  of 
the  ectoblastic  constituent. 

The  protonephridium  in  the  course  of  transmutation  and  division 
of  labor  has  not  become  entirely  replaced,  but  it  has  rather  become 
reduced  in  amount  by  the  substitution  of  other  elements  for  certain 
of  its  parts.  And  there  have  been  two  paths  in  this  process.  By 
the  one,  a  relatively  larger  portion  of  the  protonephridium  has  per- 
sisted and  a  coelomostome  has  become  directly  connected  with  it, 
exemplified  by  the  coelonephridium.  By  the  other  a  relatively 
smaller  portion  of  it  has  maintained  itself,  to  this  has  been  added 
a  secretory  tube  of  retroperitoneal  mesentoblastic  tissue,  and  to  the 
latter  in  some  cases  a  coelomostome,  as  illustrated  by  the  meta- 
nephridium. The  coelomostome  is  homologically  a  genital  funnel, 
as  demonstrated  by  Meyer  and  Goodrich,  comparable  with  a  genital 
duct  of,  e.  g.,  a  Nemertean.  But  what  the  retroperitoneal  mesento- 
blastic element  was  originally,  before  it  attached  itself  to  a  proto- 
nephridium, we  are  unable  to  decide;  it  may  have  originated  from 
the  outer  layer,  that  outside  of  the  peritoneum,  of  a  primitive 
gonadal  f)OUch. 

We  have  now  to  see  how  these  conclusions  relate  themselves 
to  the  views  of  other  students.  It  will  not  be  necessary  to  attempt 
a   full  historical  review  of  the  various  opinions  because  a  good 


610  MONTGOMERY— MORPHOLOGY  OF  THE  [ April  a*, 

discussion  of  them  has  been  recently  furnished  by  Lang  (1903). 
There  are  two  main  views:  (i)  That  the  nephridia  of  all  the 
Metazoa  are  essentially  homologous,  and  (2)  that  those  of  the 
higher  Metazoa  are  dyshomologous  with  the  protonephridia. 

The  first  of  these  has  been  maintained  particularly  by  Lang 
(1881,  1884,  1903).  To  him  the  starting  point  is  the  condition  in 
the  Turbellarian  Gunda,  where  there  are  continuous  longitudinal 
main  trunks,  and  more  or  less  regularly  arranged  excretory  ducts. 
He  holds  that  such  a  condition  has  maintained  itself  in  the  case  of 
the  plectonephridia  of  the  Hirudinea  and  Oligochaeta,  but  that  it 
has  become  modified  in  other  Annelids  by  the  segmentation  of  the 
longitudinal  trunks.  This  idea  is  in  a  sense  a  necessary  corollary 
of  his  view  of  the  close  relationship  of  the  Turbellaria  and 
Hirudinea.  Besides  the  similarity  in  the  Turbellaria  and  the 
Hirudinea  above  mentioned,  he  adduces  the  following  main  anatom- 
ical resemblances,  (i)  Hatschek*s  contention  that  in  Polygordius 
the  adult  nephridia  develop  as  outgrowths  from  a  continuous  longi- 
tudinal canal;  the  error  of  this  observation  has  since  been  pointed 
out  by  Fraipont,  Meyer,  and  Woltereck.  (2)  The  presence  of  net- 
like nephridia  (plectonephridia)  in  the  Annelids;  I  have  entered 
into  the  question  of  the  homologies  of  these  in  the  descriptive 
section  upon  the  Oligochaeta,  and  here  need  only  recall  that  Vejdov- 
sky's  embryological  studies  have  shown  that  the  plectonephric  con- 
dition is  secondary,  derived  from  the  meganephric.  (3)  The 
similarity  in  histological  structure  of  the  two  kinds  of  nephridia. 
(4)  Occurrence  of  serial  provisory  larval  nephridia  in  Polychaetes, 
that  closely  resemble  protonephridia;  that  these  are  homologous 
with  larval  protonephridia  as  well  as  with  the  definitive  ones,  accord- 
ingly, that  the  protonephridia  are  homologous  with  segmental  or- 
gans. Thus  Lang  derived  (1903)  "all  the  segmental  nephridia  of 
the  Annelids  from  the  segmental  portions  of  the  water  vascular 
system  that  open  externally,  on  the  premise  that  in  the  Annelids 
those  canals  have  not  persisted  which  joined  the  successive  seg- 
ments of  the  water  vascular  system.  Such  a  nephridial  segment 
would  have  consisted  in  the  ancestors  of  the  Annelids  of  a  pair  of 
water  vascular  trees  with  excretory  ciliated  cells  on  the  terminal 
ends  of  the  capillary  branches,  and  of  a  trunk  opening  outward. 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  Gil 

.  .  .  Since  in  the  development  of  the  Annelids  the  head  end  of  the 
body  precedes  and  the  trunk  with  its  successive  segments  first  later 
comes  to  formation,  so  develops  first  the  first  nephridial  tree  pair, 
the  head  kidney  adapted  to  the  larval  body,  whose  homology  with 
the  water  vascular  system  is  not  contended  even  by  the  opponents 
of  the  unit  theory,  later  perhaps  a  second  and  possibly  still  a  third 
similar  pair  with  reduced  branching.  This  most  anterior  pair  of 
nephridial  trees  that  functions  during  the  earliest  larval  life,  at  a 
time  when  there  is  still  no  secondary  body  cavity  developed  in  the 
regions  concerned,  became  in  the  phylogeny  a  transitory  provisory 
structure,  as  can  be  demonstrated  on  so  many  larval  organs,  while 
the  succeeding  nephridial  pairs  of  the  trunk  segments  changed  to 
s^^ental  organs." 

The  other  main  view  is  that  represented  by  Bergh  (1885).  Ac- 
cording to  him  the  larval  nephridia  of  the  Coelomata  are  homologous 
with  the  protonephridia,  while  the  adult  nephridia  of  the  Annelids 
are  homologous  with  the  gonadal  ducts  of  the  Platodes  but  not 
homologous  with  the  protonephridia.  Thus  he  concluded  (as 
Williams  did  long  before)  that  the  segmental  organs  of  Annelids 
were  originally  genital  ducts  and  later  changed  into  excretory 
organs;  while  the  protonephridia  do  not  communicate  with  the 
coelom  and  never  serve  as  genital  ducts. 

Goodrich  has  recently  represented  a  view  that  in  the  main  sup- 
ports Lang's.  To  him  there  are  "  nephridia  "  proper  that  never 
serve  as  genital  ducts ;  he  considers  all  of  these  ectoblastic  invagina- 
tions and  essentially  homologous.  Then,  adding  materially  to  the 
discoveries  of  Eisig  and  E.  Meyer,  amplifying  them,  he  find  that 
upon  such  a  nephridium  a  ccelomostome  (peritoneal  funnel,  genital 
funnel)  may  become  grafted,  giving  rise  then  to  a  complex  "  nephro- 
mixium."  To  Goodrich  all  nephridia  are  essentially  homologous, 
they  differ  only  in  being  combined  or  not  combined  with*  a  ccelomo- 
stome.^' His  argument  like  Lang's  is  rather  anatomical  than  em- 
bryological.  Both  of  these  investigators  also  lay  great  stress  upon 
the  presence  in  Annelid  nephridia  of  the  solenocytes,  cells  similar  to 
the  flame  cells  of  protonephridia;  Goodrich  argues  that  such  com- 

"In  the  descriptive  part  under  the  caption  of  Polychaeta,  Goodrich's 
ideas  are  given  more  in  extenso. 


612  MONTGOMERY- MORPHOLOGY  OF  THE  (April  84, 

plex  cells  could  not  have  arisen  independently  in  the  two  groups, 
rather  that  their  presence  in  them  means  homology  of  the  organs 
concerned. 

It  will  be  seen  that  my  views  do  not  coincide  exactly  with  any 
of  the  preceding.  I  agree  entirely  with  Meyer  and  Goodrich  that 
the  coelomostome  is  an  organ  of  origin  independent  from  the 
nephridium,  one  that  in  some  cases  may  connect  with  the  latter. 
This  coelomostome  is  equivalent  to  the  genital  duct  of  a  lower  meta- 
zoan,  as  shown  by  Bergh.  I  agree  also  with  Lang  that  the  excretory 
ducts  of  the  protonephridia  have  maintained  themselves  in  part  in 
the  higher  Metazoa,  and  that  the  longitudinal  canals  have  dis- 
appeared. But  I  have  tried  to  show  that  while  sometimes  such  an 
excretory  duct  joins  directly  with  a  coelomostome,  forming  what  I 
call  a  coelonephridium,  in  other  cases  it  joins  with  retroperitoneal 
mesentoblastic  tissue  and  the  latter  may  secondarily  join  with  a 
coelomostome  (metanephridium).  In  other  words,  we  have  to 
reckon  with  a  retroperitoneal  element  that  frequently  forms  the 
greater  portion  of  the  nephridium,  and  this  is  what  Lang  and  Good- 
rich have  failed  to  take  into  account.  And  I  differ  from  Bergh 
in  concluding  that  the  metanephridium  is  not  in  its  entirety  equiv- 
alent to  a  genital  duct,  but  that  only  a  portion  of  it  (the  coelomo- 
stome) is.  Groodrich's  mistake,  if  my  interpretation  is  correct,  is  in 
assuming  that  there  are  only  two  elements,  ectoblastic  tube  and 
peritoneal  coelomostome;  he  entirely  neglects  the  retroperitoneal 
tissue,  and  yet  this  is  just  what  shows  the  dyshomology  of  proto- 
nephridium  and  metanephridium.  It  is  a  mistake  that  has  resulted 
from  too  exclusive  reliance  upon  phenomena  of  adult  structure 
with  neglect  of  comparative  embryology.  And  the  arguments  from 
histological  similarity,  intracellular  cavity,  similarity  of  solenocytes 
to  flame  cells,  etc.,  can  have  little  weight  now  that  we  are  acquainted 
with  still  more  striking  cases  of  histological  convergence  as  notably 
the  case  of  the  Malpighian  vessels  of  Insects  and  those  of  Arachnids. 
Goodrich  has  excellently  analyzed  the  history  of  the  coelomostome 
and  has  thereby  greatly  clarified  our  knowledge  of  nephridia.  But 
he  has  omitted  entirely  from  his  general  conclusions  the  retroperi- 
toneal element  which  has  come  to  supplant  the  protonephridium 


igo8.]  EXCRETORY   ORGANS  OF  METAZOA.  613 

almost  entirely  thus  excluding  the  homology  of  the  protonephridium 
and  metanephridium. 

It  will  be  noted  that  in  my  considerations  I  have  entirely  ex- 
cluded the  argument  from  the  side  of  the  recapitulation  theory,  for 
I  have  maintained  (1906)  that  this  theory  is  fundamentally  errone- 
ous. I  have  compared  corresponding  stages,  adult  or  embryonic, 
of  the  different  groups,  have  stressed  embryological  resemblances, 
but  have  not  compared  an  adult  stage  of  one  organ  with  an  em- 
bryonic one  of  another. 

It  might  be  expected  that  I  should  now  enter  upon  the  question 
of  the  phylogenetic  significance  of  the  ccelom,  because  this  space 
has  so  often  a  close  anatomical  connection  with  nephridia.  But  I 
have  nothing  new  to  add  to  the  discussion,  and  for  a  good  repre- 
sentation of  it  would  refer  to  the  treatments  by  E.  Meyer  (1901) 
and  Lang  (1903).  I  need  only  state  that  there  are  three  main 
theories  in  explanation  of  the  origin  of  the  ccelom.  The  oldest  was 
founded  by  Sedgwick,  and  is  to  the  effect  that  the  coelom  is  an 
enterocoelic  diverticulum,  referable  to  a  gastral  pocket  of  an 
anthozoan.  This  has  deservedly  received  little  support.  Next  came 
the  gonocoel  theory,  foreshadowed  by  Hatschek,  elaborated  partic- 
ularly by  Bergh  and  E.  Meyer,  and  more  recently  supported  by 
Lang  and  Goodrich;  it  concludes  that  the  ccelomic  sac  of  a  higher 
metazoan  is  the  amplified  derivative  of  the  genital  pouch  (gonad) 
of  such  a  form  as  a  Platode,  therefore  that  the  mesentoblast  is 
referable  to  germ  cells  The  third  view  is  the  nephroccEl  theory, 
founded  by  Faussek  (1901)  and  Ziegler  (1898),  that  the  ccelom 
was  originally  an  excretory  organ  and  that  the  germ  cells  have 
associated  themselves  secondarily  with  it.  Of  these  three  theories 
the  gonocoel  theory  seems  to  me  to  receive  the  fullest  support  from 
the  facts  of  anatomy  and  embryology. 

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X904.    The  Anatomy  of  Poecilochaetus,  Claparede.    Quart.  Journ.  Micr,  Set. 
(2),  48. 


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Anglas,  J. 

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Beard,  J. 

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Beddard,  F.  E. 

1888a.    On  the  occurrence  of  numerous  nephridia  in  the  same  segment  in 

certain  Earthworms,  etc.    Quart.  Joum.  Micr.  Sci.   (2),  28. 
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gaster  n.  g.,  etc.    Ibid.,  29. 

1889.  On  the  possible  origin  of  the  Malpighian  tubules  in  the  Arthropoda. 
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etc.    Quart.  Journ,  Micr.  Sci.  (2),  32. 
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Ibid.,  33. 

1902.  A  note  upon  the  Gonad  Ducts  and  Nephridia  of  Earthworms  of  the 
Genus  Eudrilus.    Proc.  R.  Soc.  London. 

Bemmelen,  J.  F.  Van. 

1883.  Untersuchungen  iiber  den  anatomischen  und  histologischen  Bau  der 
Brachiopoda  Testicardinia.    Jena.  Zeit.,  16. 


L 


,^.]  EXCRETORY  ORGANS  OF  METAZOA.  615 

Beneden,  E.  Van  et  Julin,  C. 

1886.    Recherches   sur  la   morphologic  des   Tuniciers.    Arch,  de  Biol,  6. 

Benham,  W.  B. 

1886.    Studies  on  Earthworms.     Quart.  Journ.  Micr.  Set.   (2),  26,  27. 

1888.  Note  on  a  new  Earthworm.    Zooi  Ans.,  11. 

1889.  The  Anatomy  of  Phoronis  Australis.    Quart.  Journ.  Micr.  Sci,   (2), 

30. 

1890a.    An  attempt  to  classify  Earthworms.    Ibid.,  31. 

1890b.  Report  on  an  earthworm  collected  for  the  Natural  History  Depart- 
ment of  the  British  Museum,  etc.    Journ.  R.  Micr.  Soc.  London. 

1891.  The  nephridium  of  Lumbricus  and  its  blood-supply;  with  remarks  on 
the  nephridia  in  other  Chaetopoda.    Quart.  Journ,  Micr.  Sci.  (2),  32. 

1904.  On  a  new  species  of  the  genus  Haplotaxis;  with  some  remarks  on  the 
genital  ducts  in  the  Oligochaeta.    Ibid.,  48. 

Bdxgli,  R.  S. 

1884.  Ueber  die  Metamorphose  von  Nephelis.    Zeit.  w.  Zooi,  41. 

1885.  Die  Excretionsorgane  der  Wiirmer.    Kosmos,  17. 

1886.  Untersuchungen  iiber  den  Bau  und  die  Entwickelung  der  Geschlechts- 
organe  der  Regenwiirmer.    Zeit.  w.  Zooi,  44, 

1888.  Zur  Bildungsgeschichte  der  Excretionsorgane  bei  Criodrilus.  Arb. 
Zooi  Inst.  Wiirzburg,  8. 

1890.  Neue  Beitrage  zur  Embryologie  der  Anneliden.  1.  Zur  Entwickelung 
und  Differenzirung  des  Keimstreifen  von  Lumbricus.    Zeit.  w.  Zooi,  50. 

1891.  Die   Schichtenbildung  im   Keimstreifen   der  Hirudincen.    Ibid.,  52. 

1899.  Nochmals  iiber  die  Entwickelung  der  Segmentalorgane.    Ibid.,  66. 
1901.    Kleinere  histologische   Mittheilungen.    Ibid.,  69. 

Bernard,  H.  M. 

1893.    The  Coxal  Glands  of  Scorpio.    Ann.  Mag.  Nat.  Hist.  (6),  11. 
1896.    The  Comparative  Morphology  of  the  Galeodidse.     Trans.  Linn.  Soc. 
London  (2),  6. 

Bemdt,  W. 

1903.  Zur  Biologic  und  Anatomic  von  Alcippe  lampas  Hancock.  Zeit.  w 
Zooi,  74-  ' 

Bertkau,  P. 

1885.    Ueber  die  Coxaldriiscn.     Verh.  Nat.  Ver.  Bonn,  41. 

1888.    Ueber  die  Chemctiden  oder  Pseudoskorpione.    Ibid.,  44, 

Blochmann,  F. 

1900.  Untersuchungen  iiber  den  Bau  der  Brachiopoda.    II  ter  Theil.  Jena. 
Bobretzky,  N. 

1877.  Studien  iiber  die  embryonale  Entwicklung  der  Gastcropoden.  Arch, 
mikr.  Anat,  13. 

Bdhmig,  L. 

1893.    Zur  feineren  Anatomic  von  Rhodope  Veranii  Kolliker.    Arb.  zooi  Inst. 

Graz,  5. 
1898.    Beitrage  zur  Anatomic  und  Histologic  der  Nemcrtincn.    Ibid.,  6. 
1906.    Tricladenstudien,  I.    Ibid.,  7. 


616  MONTGOMERY— MORPHOLOGY  OF  THE  [April  .4. 

BolsiuSy  H. 

189a.    Les  organes  cili^s  des  Hirudinees.    La  Cellule,  7. 

Bonnier,  J. 

1891.  La  glande  antennale  chez  les  Amphipodes  de  la  famille  des  Orches- 
tidae.    C,  R.  Acad.  Sci.  Paris,  113. 

Bdrner,  C. 

Z904.  Beitrage  zur  Morphologie  der  Arthropoden,  i.  Ein  Beitrag  zur  Kennt- 
niss  der  Pedipalpen.    Zoologica,  42. 

Bomne,  A.  6. 

1884.  Contributions  to  the  Anatomy  of  the  Hirudinea.  Quart.  Joum.  Mier. 
Sci.  (2),  24. 

1894.  On  certain  Points  in  the  Development  and  Anatomy  of  some  Earth- 
worms.   Ibid.,  36. 

Boutan,  L. 

1885.  Recherches  sur  Tanatomie  et  la  d^veloppement  de  la  Fissurella.  Arch. 
Zool.  ex  ph.  gin.  j,  supplement. 

BoTeri,  T. 

189a.    Die  Nierencanalchen  des  Amphioxus.    Zool.  Jahrh.,  5. 
1904.    Bemerkungen   uber   den    Bau    der   Nierencanalchen    des   Amphioxus. 
Anat.  Am.,  25. 

Brandes,  6. 

1901.  Die  Parasiten  des  Menschen    (Leuckart).    2te  Aufl.  Leipzig. 
Brauer,  A. 

1895.  Beitrage  zur  Kenntnis  der  Entwickelungsgeschichte  des  Skorpions,  2. 
Zeit.  wiss.  Zool.,  59. 

1902.  Beitrage  zur  Kenntnis  der  Entwickelung  und  Anatomic  der  Gymno- 
phionen,  3.    Die  Entwickelung  der  Excretionsorgane.    Zool.  Jahrh.,  16. 

Braun,  M. 

1893.  Trematodes.  Bronn's  Klassen  und  Ordnungen  des  Thierreichs,  4, 
Leipzig. 

BrontZy  L. 

Z903a.    Contribution  4  Tetude  de  Texcretion  chez  les  Arthropodes.    Arch,  de 

Biol,  20. 
X903b.    Excretion  et  phagocytose  chez  les  Onychophores.    C.  R.  Acad.  Sci. 

Paris,  136. 

Bugge,  6. 

1902.  Zur  Kenntnis  der  Exkretionsgefasssystems  der  Trematoden  und 
Cestoden.    Zool.  Jahrh.,  16. 

Biirger,  0. 

1891.  Beitrage  zur  Entwickelungsgeschichte  der  Hirudineen.  Zur  Embry- 
ologie  von  Nephelis.    Ihid.,  4. 

1894.  Neue  Beitrage  zur  Entwickelungsgeschichte  der  Hirudineen.  Zur 
Embryologie  von  Hirudo  medicinalis  und  Aulastomum  gulo.  Zeit.  wiss. 
Zool,  58. 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  617 

1895.    Nemertinen.  Fauna  u.  Flora  d.  Golfes  v.  Neapel,  22. 
190a.    Weitere  Beitrage  zur  Entwickelungsgeschichte  der  Hirudineen.     Zur 
Embryologie  von  Clepsine.    Zeit.  wiss.  ZooL,  72. 

Bfitschli,  0. 

1870.    Zur  Entwickelungsgeschichte  der  Biene.    Ibid, 

1877.    Entwicklungsgeschichtliche  Beitrage.    Ueber  Paludina  vivipara.    Ibid., 
29. 

Caldwell,  E.  H. 

1882.    Preliminary   Note  on  the   Structure,   Development  and   Afl5nities  of 
Phoronis.    Proc,  R.  Soc.  London,  34, 

Casteel,  D.  B. 

1904.  The  Cell-lineage  and  early  larval  Development  of  Fiona  marina,  a 
Nudibranchiate  Mollusk.    Proc,  Acad.  Nat,  Set.  Philadelphia. 

Chun,  C. 

1880.    Die  Ctenophoren  des  Golfes  von  Neapel.    Fauna  u.  Flora  d.  Golfes 
V.  Neapel,  i. 

Glaus,  C. 

1875.    Die  Schalendriise  der  Daphnien.    Zeit.  wiss.  ZooL,  25. 

1877.    Die  Schalendriise  der  Copepoden.    Sitzber.  Akad,  Wiss.  Wien,  74. 

1890.    Ueber  die  Organisation  der  C)rpriden.    Anz.  Akad.  Wiss,  Wien,  27. 

1895.  Beitrage  zur  Kenntnis  der   Siisswasser-Ostracoden.    Arb,  zool,  Inst, 
Wien,  II. 

\/0e,  Mr  ■  iCa 

1896.  Notizen  iiber  den  Bau  des  Embryos  von  Distomum  hepaticum.    Zool, 
Jahrb.,  9. 

1906.    A  peculiar  Type  of  Nephridia  in  Nemerteans.    Biol.  Bull.,  11. 

Conklin,  E.  6. 

1897.  The  Embryology  of  Crepidula.    Journ.  Morph.,  13. 

1905.  The  Organization  and  Cell-Lineage  of  the  Ascidian  Egg.    Journ.  Acad. 
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Conte,  A. 

1902.    Contributions  i  Tembryologie  des  Nematodes.    Ann.  Univ.  Lyon  (2),  i. 

Cori,  C.  J. 

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Phoronis.    Zeit.  miss.  ZooL,  51. 
1893.    I^ie  Nephriden  der  Cristatella.    Ibid.,  55. 

Cosmovici,  L.  C. 

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Arch.  Zool.  exph.  gin.,  8. 

Cowles,  R.  P. 

1905.    Phoronis  architecta.    Mem.  National  Acad.  Sci.  Washington,  10. 

Cronebexg,  A. 

1879.    Ueber  den  Bau  von  Trombidium.    Bull.  Soc.  Imp.  Nat.  Moscou,  2. 

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Culnot,  L. 

1888.    £tudes  anatomiques  et  morphologiques  sur  les  Ophiures.    Arch.  ZooL 

expSr,  gin.  (2),  5  bis. 
1895.    £tudes  physiologiques  sur  les  Orthopteres.    Arch,  de  Biol.,  14. 
Z897.    £tudes  physiologiques  sur  les  Oligochetes.    Ihid.,  15. 

1899.  L'excr6tion  chez  les  Mollusques.    Ibid.,  16. 

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190a.    Organes  agglutinants  et  organes  cilio-phagocytaires.    Ibid.,  la 

Daday,  E.  y. 

1895.  Die  anatomischen  Verhaltnisse  der  Cyprois  dispar  (Chyz.).  Termisx. 
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X901.  Untersuchungen  iiber  den  Bau  der  Excretionsorgane  der  Tunicaten. 
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Danielssen,  D.  C.  and  Koren,  J. 

1881.    Gephyrea.    Norske  Nordhavs-Expedition.    1876-1878,  4,  Christiania. 

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1900.  Recherches  sur  les  Aphrodities,  Lille. 

Dayenport,  C.  B. 

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Delia  Valle,  A. 

1893.    Gammarini.    Fauna  u.  Flora  d.  Golfes  v.  Neapel,  20. 

Dohm,  A. 

1870.    Untersuchungen  uber  Bau  und  Entwicklung  der  Arthropoden,  Leipzig. 

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Doncaster,  L. 

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Z884.    Beitrage  zur  Entwicklung  der  Polychaeten,  i,  Wien.  i 

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Hymenopteres  et  les  Neuropteres.    Ibid.,  7. 
1851.    Recherches  anatomiques  et  physiologiques  sur  les  Dipteres.    Ibid.,  11. 
1854.    Histoire  anatomique  et  physiologique  des  Scorpions.    Ibfd.,  14. 


.go8.]  EXCRETORY  ORGANS  OF  METAZOA.  619 

EhlerSy  E. 

Z890.    Zur  Kenntniss  der  Pedicellineen.    Abh.  Ces.  IViss.  Gottingen,  j6. 

Eisigy  H. 

1887.    Monographie  der  Capitelliden.    Fauna  u.  Flora  d.  Golfes  v.  NcapeL 

Emeryy  C. 

1883.    Ricerche  embriologiche  sul  rene  dei  Mammiferi.    Mem.  Accad,  Lincei. 

15. 

ErUnger,  R.  y. 

i89xa.    Zur  Entwicklung  der  Paludina  vivipara.    Morph,  Jahrb.,  17. 

1891b.    Zur  Entwicklung  von  Paludina  vivipara.    II  Theil.      Ibid, 

x892a.    Mittheilungen   tiber  Bau   und   Entwicklung  einiger  marinen   Proso- 

branchier,  I.  Ueber  Capulus  hungaricus.    ZooL  Anz.,  15. 
i89ab.    Beitrage  zur  Entwicklungsgeschichte  der  Gastrc^oden.    Erster  Theil. 

Zur     Entwicklung     von     Bytjiinia     tentaculata.    Mittheil,     sooL     Stat 

Neapel,  10. 

1893.  Bemerkungen    zur    Embryologie    der    Gasteropoden,    i.    Ueber    die 
sogenannten  Umieren  der  Gasteropoden.    Biol.  Centralbi,  13. 

1894.  Bemerkungen  zur  Embryologie  der  Gasteropoden,  II.  Ibid.,  14. 

Evans,  R. 

1901.    On    the   Malayan   Species   of    Onychophora,    Part   2.    Quart.   Journ. 
Micr.  Sei.  (2),  45. 

Faussek,  V. 

1892.    Zur  Anatomic  und  Embryologie  der  Phalangiden.    Biol.  Centralbi,  12. 
1900.    Untersuchungen  uber   die   Entwicklung  der   Cephalopoden.    MittheiL 
sool.  Stat.  Neapel,  14. 

Felix,  W. 

1891.    Die  erste  Anlage  des  Excretionssystems  des  Hiihnchens.    Festschr.  f. 

Nageli  u.  Kolliker,  Zurich. 
1897.    Beitrage  zur  Entwickelungsgeschichte  der  Salmoniden.    Anat.  Hefte,S. 

Field,  G.  W. 

189a.    The  Larva  of  Asterias  vulgaris.     Quart.  Journ.  Micr.   Sci.    (2),  34. 

Field,  H.  H. 

1891.    The  development  of  the  pronephros  and  segmental  duct  in  Amphibia. 
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Flemming,  W. 

1886.  Die   ektoblastische   Anlage   des   Urogenital-systems   beim   Kaninchen. 
Arch.  Anat.  Physiol. 

Foettinger,  A. 

1887.  Sur  Tanatomie  des  Pedicellines  de  la  cote  d'Ostende.    Arch,  de  Biol.,  7. 

Fol,  H. 

1875.    £tudes   sur   le   d^veloppement    des    Pt^ropodes.    Arch.    Zool.    expkr, 

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i88a    fitudes  sur  le  developpement  des  Gasteropodes  pulmones.    Ibid.,  8. 


620  MONTGOMERY— MORPHOLOGY  OF  THE  [Aprils, 

Folsom,  J.  W.  and  Welles,  M.  IT. 

1906.    Epithelial    Degeneration,   Regeneration,   and   Secretion   in   the    Mid- 
Intestine  of  Collembola.     Univ,  Illinois  Bull.  4. 

Fraipont,  J.  * 

1888.    Le  genre  Polygordius.    Une  monographic.    Fauna  u.  Flora  d.  Golfes 
V.  Neapel,  14. 

Fiirbringer,  M. 

1878.    Zur  vergleichenden   Anatomie   und   Entwicklungsgeschichte   der    Ex- 
cretionsorgane  der  Vertebraten.    Morph.  Jahrb.,  4. 

Cast,  R. 

1900.    Beitrage    zur    Kenntniss    von    Apsilus    vorax    (Leidy).    Zeit.    wiss. 
Zooi,  67, 

Geoxgevitchy  J. 

1898.    Die  Segmentaldriisen  von  Ocypus.  *Zool.  Ans.,  21. 

Georgevitch,  P.  M. 

1900.    Zur  Entwickelungsgeschichte  von  Aplysia  depilans  L.    Anat  Ans.,  18. 

Gerould,  J.  H. 

X896.    The  Anatomy  and  Histology  of  Caudina  arenata.     Gould.  Bull.  Mus. 

Zool.  Harvard,  29. 
1906.    The  Development  of  Phascolosoma.    Zool.  Jahrb.,  23. 

Gilson,  6. 

1894.  The  Nephridial  Duct  of  Owenia.    Anat.  Am.,  10. 

Glaser,  0.  C. 

1905.  Ueber  den   Kannibalismus   bei   Fasciolaria  tulipa    (var.   distans)    und 
deren  larvale  Exkretionsorgane.    Zeit.  wiss,  Zool.,  80. 

'Goldsclimidt,  R. 

1906.  Mittheilungen  zur  Histologic  von  Ascaris.    Zool.  Anz.,  29. 

Ooodricli,  E.  S. 

1895.  On  the  Coelom,  Genital  Ducts,  and  Nephridia.    Quart.  Journ.  Micr.  Sci 

(2),  37. 
1897a.    Notes  on  the  Anatomy  of  Stemaspis.    Ibid.,  40. 
1897b.    On  the  Nephridia  of  the  Polychaeta,  Part  I.  On  Hesione,  Tyrrhena, 

and  Nephthys.    Ibid.,  40. 
1898.    On  the  Nephridia  of  the  Polychaeta,  Part  II.  Glycera  and  Goniada. 

Ibid.,  41. 
1900.    On   the   Nephridia   of   the   Polychaeta,    Part   III.  The    Phyllodocidae, 

Syllidae,  Amphinomidae,  etc.,  with  Summary  and  Conclusions.    Ibid.,  43. 

1902.  On  the  Structure  of  the  Excretory  Organs  of  Amphioxus.    Ibid.,  45. 

1903.  On   the   Body   Cavities   and    Nephridia   of   the   Actinotrocha   larva. 
Ibid.,  47. 

Graf,  A. 

1893.    Beitrage  zur  Kenntniss  der  Exkretionsorgane  von  Nephelis  vulgaris. 

Jena.  Zeit.,  28. 
Z899.    Hirudineen-studien.    Nova  Acta  Acad.  Leop.  Carol.  72. 


' 


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Graff,  L.  t. 

1 88a,    Monographie  der  Turbellarien,  i.  Rhabdocoelidea.    Leipzig. 

Grassi,  G.  B. 

1883.    I  Chetognati.    Fauna  u.  Flora  d.  Golfes  v.  Neapel,  $. 
1B85.    Intomo  alio  sviluppo  delle  api  nell'  uovo.    Atti.  Ac  cad,  Gioenia,  18. 
1886.    Intomo  ad  un  nuovo  aracnide  artrogastro.    Boll.  Soc.  entom.  ital.,  18. 
1888.    Anatomia  comparata  dei  Tisanuri.    Reale  Accad.  de  Lincei  (4),  4. 

Greef  ,  R. 

1879.  Die  Echiurcn  (Gephyrea  armata).  Nova  Acta  Leop.  carol.  Akad. 
Halle,  41. 

Gregory,  B.  R. 

1900.  Observations  on  the  Development  of  the  Excretory  System  in  Turtles. 
Zool.  Jahrh.,  13. 

Grobben,  C. 

1879.  Die  Entwickelungsgeschichte  der  Moina  rectirostris.  Arh,  sooL  Inst, 
JVien,  2. 

1881.    Die  Antennendruse  der  Crustaceen.    Ibid.,  3. 

1888.  Die  Pericardialdruse  der  chsetopoden  Anneliden,  nebst  Bemerkungen 
iiber  die  perienterische  Flussikkeit  derselben.  Sitzher.  Akad.  Wiss. 
Wien,  97. 

Gmveli  A.  < 

1894.    Contribution  i  I'^tude  des  Cirrhipedes.    Arch.  Zool.  expkr.  gkn.  (3),!. 

Guiart,  J. 

1901.  Contribution  a  I'etude  des  Gasteropodes  opisthobranches  et  en  par- 
ticulier  des  C^phalaspides.    Mkm.  Soc,  Zool.  France,  14. 

Haase,  B. 

1889.  Die  Abdominalanhange  der  Insekten  mit  Beriicksichtigung  der  Myrio- 
poden.    Morph.  Jahrh.,  15. 

Haddon,  A.  C. 

1886.  Suggestion  respecting  the  epiblastic  origin  of  the  Segmental  Duct. 
Proc.  R.  Soc.  Dublin  (2),  5. 

Hall,  R.  W. 

1904.  The  Development  of  the  Mesonephros  and  the  Miillerian  Duct  in 
Amphibia.    Bull.  Mus.  Zool.  Harvard,  45. 

Haller,  G. 

1880.  Zur  Kenntnis  der  Tyroglyphen  und  Verwandten.    Zeit.  wiss.  Zool.,  34. 

Hamann,  0. 

1887.  Beitrage  zur  Histologie  der  Echinodermen.    Heft  3.    Jena.  Zeit.,  21. 

Hansen  and  Sorensen. 

1904.    On  two  Orders  of  Arachnids,  etc.    Cambridge. 

Harmer,  S.  F. 

1885.  On  the  structure  and  development  of  Loxosoma.  Quart.  Joum,  Micr, 
Set.  (2),  25. 


622  MONTGOMERY— MORPHOLOGY  OF  THE  [Aprils, 

1889.    Notes  on  the  Anatomy  of  Dinophilus.    Joum.  Marine  Biol.  Assoc. 

London  (2),  i. 
1891.    On  the  Nature  of  the  Excretory  Processes  in  Marine  Polyzoa.    Quart. 

Joum.  Micr.  Sci.  (2),  33. 

Hatschek,  B. 

zSyya.    Embryonalentwicklung  und  Knospung  der  Pedicellina  echinata.    Zeit. 

wiss.  ZooL,  29. 
z877b.    Beitrage     zur     Entwickelungsgeschichte     der    Lepidopteren.     Jena. 

Zeit,  II. 
1878.    Studien  iiber  Entwicklungsgeschichte  der  Anneliden.    Arb.  sool.  Inst. 

IVien,  I. 
z88oa.    Ueber  Entwicklungsgeschichte  von  Teredo.    Ibid.,  3. 
z88ob.    Ueber  Entwicklungsgeschichte  von  Echiurus  und   die  systematische 

Stellung  der  Echiuridse  (Gephyrei  Chaetiferi).  Ihid.,  3. 
1883.  Ueber  Entwicklung  von  Sipunculus  nudus.  Ihid.,  5. 
1885.    Entwicklung    der    Trochophora    von    Eupomatus    uncinatus    Philippi 

(Serpula  uncinata).    Ibid.,  6. 
1888.    Lehrbuch  der  Zoologie,  Jena. 

Heathcote,  F.  6. 

1888.    The  Post-embryonic  Development  of  Julus  terrestris.    Proc.  R.  Soc. 
London,  43. 

Hempelnunn,  F. 

1906.    Zur  Morphologie  von  Polygordius  lacteus  Schn.  und  Polygordius  tri- 
estinus  Woltereck,  nov.  spec.    Zeit.  zviss.  ZooL,  84. 

Henking,  H. 

z883.    Beitrage    zur    Anatomic,    Entwicklungsgeschichte    und    Biologic    von 
Trombidium  fuliginosum  Herm.    Ibid.,  37. 

Hennegiiy,  F. 

x888.    Recherches  sur  le  developpement  des  Poissons  osseux.    Embryogenie 
de  la  Truite.    Joum.  Anat.  Physiol.,  24. 

Hensevaly  M. 

1896.    £tude  compar^e  des  glandes  de  Gilson.    Organes  m^tameriques  des 
larves  d'Insectes.    La  Cellule,  12. 

Herbst,  C. 

X891.    Beitrage  zur  Kenntnis  der  Chilopoden.    Bibl.  ZooL,  9. 

H6roiiard,  B. 

X893.    Recherches  sur  les  Holothuries  de  la  mer  Rouge.    Arch.  ZooL  expir. 
gin.  (3),  I. 

Hertwig,  0. 

x88o.    Die  Chaetognathen.    Eine  Monographic.    Jena.  Zeit.,  14. 

Hertwig,  0.  und  R. 

1881.    Die  Colomtheorie,  Jena. 

Hescheler,  E. 

1900.    Mollusca.    Lang's  Lehrbuch  der  vergleichenden  Anatomic  der  wirbel- 
losen  Thiere.    2te  Aufl.  Jena. 


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H^monSy  R. 

X893.    Zur  Entwicklungsgeschichte  von  Umbrella  meditcrranea  Lam.    Zeit. 

wiss,  Zool. 
1896.    Grundzuge  der  Entwickelung  und  des  Korperbaues  von  Odonaten  und 

Ephemeriden.    Anh,  Abh.  Akad.  Wiss.  Berlin, 
X898.    Entwickelungsgeschichte    der    Chilopoden.    Sits.    Ber.    Akad.    Wiss. 

Berlin. 
X901.    Die  Entwickelungsgeschichte  der  Scolopender.    Zoologica,  33. 

Hincks,  T. 

1880.    A  History  of  the  British  Marine  Polyzoa,  London. 

Hlava,  S. 

1904.  Einige  Bemerkungen  uber  die  Exkretionsorgane  der  Radertierfamilie 
Melicertidae  und  die  Aufstellung  eines  neuen  Genus  Conchiloides.  Zool. 
Anz.,  27. 

1905.  Ueber  die  Anatomic  von  Conchiloides  natans  (Seligo).  Zeit.  wiss. 
Zool.,  So. 

Hoflhnann,  C.  K. 

1886.    Zur  Entwickelungsgeschichte  der  Urogenitalorgane  bei  den  Anamniem. 

Ibid.,  44. 
X889.    Zur  Entwicklungsgeschichte  der  Urogenitalorgane  ber  den  Reptilien. 

Ibid.,  48. 

HoffmAnn,  R.  W. 

1899.    Beitrage  zur  Entwickelungsgeschichte  der  Oligochaten.    Ibid.,  66. 

Holmes,  S.  J. 

X900.    The  early  Development  of  Planorbis.    Joum.  Morph.,  16. 

Hudson,  C.  T.  and  Gosse,  P. 

X88&-9.    The  Rotifera ;  or  Wheel-Animalcules,  London. 

Ijima,  J. 

1885.  Ueber  Bau  und  Entwickelung  der  Siisswasserplanarien  (Tricladen). 
Zeit.  wiss.  Zool. 

Ikeda,  I. 

X901.  Observations  on  the  Development,  Structure  and  Metamorphosis  of 
Actinotrocha.    Joum.  Coll.  Sci.  Univ.  Tokyo,  13. 

Ischikawa,  C. 

X885.  On  the  Development  of  a  freshwater  macrourous  Crustacean,  Atyephyra 
compressa  de  Haan.    Quart.  Joum.  Micr.  Sci.  (2),  25. 

JigerskiSld,  L.  A. 

1898.  Ueber  die  biischelformigen  Organe  bei  Ascarisarten.  Centralbl.  Bakt. 
Parasitenk.,  24. 

Janosik,  J. 

X887.    Zwei  junge  menschliche  Embryonen.    Arch.  mikr.  Anat.,  30. 

Jobansson,  L. 

1898.  Einige  systematisch  wichtige  Theile  der  inneren  Organisation  der 
Ichthyobdelliden.    Zool.  Am.,  21. 


624  MONTGOMERY— MORPHOLOGY  OF  THE  [April  24, 

Joliet,  L. 

1880.    Organe  segmentaire  des  Bryozoaires  endoproctes.    Arch.  Zool.  expSr, 
ght.,  8. 

Joyenx  Laffuie. 

188a.    Organisation  et  devcloppement  de  rOncidie.    Ibid.,  10. 

Kaiser,  J. 

189a.    Die  Nephridien  der  Acanthocephalen.    Centralbl.  Bakt.  Parasitenk.,  11. 

1893.    Die  Acanthocephalen  und  ihre  Entwickelung.    Bibl.  Zool,  7. 

Kaipelles,  L. 

1893.    Zur  Anatomic  von  Bdella  arenaria   (Kramer).     Verh.  sool.-hot.  Ges. 
Wien,  43. 

Eennely  J.  v. 

1885.  Entwicklungsgeschichte   von    Peripatus   edwardsii    und    P.   torquatus. 
Art.  sool.  Inst.  Wurzburg,  7,  8. 

Eenyon,  F.  C. 

1895.    The  Morphology  and  Classification  of  the  Pauropoda,  with  Notes  on 
the  Morphology  of  the  Diplopoda.    Tufts  College  Studies,  4. 

King8ley,  J.  S. 

1889.  The  Development  of  Crangon  vulgaris.    Bull.  Essex.  Inst.,  21. 

Kishinouye,  K. 

1890.  On  the  Development  of  Arancina.    Journ.  Coll.  Sci.  Univ.  lapan,  4. 
x894«    Note  on  the  Ccelomic  Cavity  of  the  Spider.    Ibid.,  6. 

Kleinenberg,  N. 

1886.  Die   Entstehung  des   Annelids  aus   der  Larve  von  Lopadorhjmchus. 
Zeit.  wiss.  Zool.,  44. 

Eoehler,  R. 

1890.  Recherches  sur  Torganisation  des  Cirrhip^des  (Lepadides  et  Balanes). 
Arch,  de  BioL^  10. 

Eof  oid,  C.  A. 

1895.    On  the  early  development  of  Limax.    Bull.  Mus.  Zool.  Harvard,  27. 

Eolliker,  A.  v. 

1861.    Entwickelungsgeschichte    des    Menschen    und    der    hoheren    Thierc, 
Leipzig. 

Eollman,  J. 

1891.  Die  Rumpfsegmente  menschlicher  Embryonen  von  13  bis  35  Urwirbeln. 
Arch.  Anat.  Physiol. 

Eorschelt,  E. 

z88a.    Ueber    Bau    und    Entwicklung    des    Dinophilus    apatris.    Zeit.    wiss, 
Zool.,  37. 

Eowalevsky,  A. 

1871.    Embryologische  Studien  an  Wiirmem  und  Arthropoden.    MSm.  Acad. 
Sci.  St.  PStersbourg. 


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1889.    Ein  Beitrag  zur  Kenntnis  der  Excretionsorgane.    Biol.  Centralbl.,  9. 
X893.    Beitrag  zur  Kenntniss  der  Excretionsorgane  der  Pantopoden.    Mhn. 
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1896.  £tude  des  glandes  lymphatiques  de  quelques  Myriapodes.    Arch.  Zool. 
expSr.  gin.  (2),  3. 

Kramer,  P. 

1885.  Ueber  Halarachne  Halichoeri  AHm.    Zeit.  Naturw.  Halle,  58. 

Lacaze-Duthiers,  H.  et  Pmvot,  6. 

1887.  Sur  un  ceil  anale  larvaire  des  Gasteropodes  opisthobranches.    C.  R. 
Acad.  Set.  Paris,  105. 

Lang,  A. 

1881.    Der  Bau  von  Gunda  segmentata.    Mittheil.  zool.  Stat.  Neap  el,  3. 
1884.    Die  Polycladen  (Seeplanarien)  des  Golfes  von  Neapel.    Fauna  u.  Flora 
d.  Golfes  V.  Neapel,  11. 

1888.  Lehrbuch  der  vergleichenden  Anatomie,  Jena. 
1903.    Beit  rage  zu  einer  Trophocoltheorie,  Jena. 

Langerhans,  P. 

1873.    Zur  Entwicklung  der  Gastropoda  Opisthobranchia.  Zeit.  zviss.  Zool,  23. 

Lankester,  E.  R. 

1865.    The  Anatomy  of  the  Earthworm.    Quart.  Joum.  Micr.  Sci. 
1877.    Notes  on  the  Embryology  and  Classification  of  the  Animal  Kingdom, 
etc.    Ibid.,  17. 

Latter,  0.  H. 

1897.  The  Prothoracic  Gland  of  Dicranura  vinula,  and  other  Notes.    Trans. 
Ent.  Soc.  London. 

Laurie,  M. 

1894.    On  the  Morphology  of  the  Pedipalpi.    Joum.  Linn.  Soc.  London,  25. 

Lebedinsky,  J. 

1891.    Die  Entwicklung  der  Daphnia  aus  dem  Sommereie.    Zool.  Ans.,  14. 

189a.    Die  Entwicklung  der  Coxaldriise  bei  Phalangium.    Ibid.,  15. 

Lehmann,  0. 

1887.    Beitrage   zur   Frage   von   der   Homologie   der   Segmentalorgane   und 

Ausfiihrgange    der    Geschlechtsproducte    bei    den    Oligochaeten.    Jena. 

Zeit.,  21. 

Leipoldt,  F. 

1893.    Das  angebliche  Excretion sorgan  der  Seeigel,  untersucht  an  Sphaere- 
chinus  granularis  und  Dorocidaris  papillata.    Zeit.  wiss.  Zool,  55. 

Leuckart,  R. 

Z894.    Die  Parasiten  des  Menschen.    2te  Aufl.,  Leipzig. 

Leydig,  F. 

i860.    Naturgeschichte  der  Daphniden,  Tubingen. 

Locy,  W.  A. 

1886.  Observations  on  the  Development  of  Agelena  naevia.    Bull.  Mus.  Zool. 
Harvard,  12. 


626  MONTGOMERY— MORPHOLOGY   OF  THE  [April  24, 

Loman,  J.  C.  C. 

1887.  Ueber  die  morphologische  Bedeutung  der  sogenannten  Malpighischen 
Gefasse  der  echten  Spinnen.    Tijdschr,  Nederl.  Dierk,  Ver,  (2),  i. 

x888.    Altes  und  Neues  uber  das  Nephridium  (die  Coxaldruse)  der  Arach- 
niden.    Bijd.  Dierk.  Amsterdam,  14. 

Longchamps,  de  Selys. 

1902.    Recherches  sur  le  developpement  des  Phoronis.    Arch,  de  Biol.,  18. 

L0088,  A. 

1894.    Die  Distomen  unserer  Fische  und  Frosche.    Bibl.  Zool.,  16. 

Lttdwig,  H. 

1889-94.    Echinodermen   (Stachelhauter),  Bronn's  Klassen  u.  Ordnungen  d. 
Thierreichs.    2,  Leipzig. 

Luther,  A. 

1904.    Die  Eumesostominen.    Zeit.  wiss.  Zool.,  77. 

Maas,  0. 

1897.    Ueber   Entwicklungsstadien   der   Vorniere   und  Urniere   bei    Myicine. 
Zool.  Jahrb.,  10. 

McKim,  W.  D. 

X895.    Ueber    den    nephridialen    Trichterapparat    von    Hirudo.    Zeit.    wiss. 
Zool.,  59. 

McMurrich,  J.  P. 

1886.    A  Contribution  to  the  Embryology  of  the  Prosobranch  Gastropods. 
Stud.  Biol.  Lab.  Johns  Hopkins  Univ.,  3. 

ISarchal,  P. 

1889.    L'acide  urique  et  la  fonction  renale  chez  les  Invertebres.    Mim.  Soc. 

zool.  France,  3^ 
i89aa.    La  glande  coxale  du  Scorpion  et  ses  rapports  morphologiques  avec 

les  organes  excr^teurs  des  Crustac^s.    C.  R,  Acad.  Sci.  Paris,  115. 
i89ab.    Recherches   anatomiques   et   physiologiques    sur   Tappareil    excr^teur 

des  Crustaces  d^capodes.    Arch.  Zool.  exph.  gSn.  (2),  10. 

Martin,  £• 

1888.  Ueber    die    Anlage    der    Urniere    beim    Kaninchen.    Arch.    Anat. 
Physiol. 

Masterman,  A.  T. 

1897.  On  the  Diplochorda.    Quart.  Journ.  Micr.  Sci.  (2),  40. 

Mayer,  P. 

1875.    Ueber  Ontogenie  und  Phylogenie  der  Insekten.    Jena.  Zeit.,  10. 

Mazzarelli,  6. 

189a.    Intorno  al  preteso  occhio  anale  delle  larve  degli  Opisthobranchi.    Rend. 
Ac  cad,  Lincei,  i. 

1898.  Bemerkungen  iiber  die  Analniere  der  freilebenden  Larven  der  Opis- 
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1904.    I  reni  primitivi  dei  Molluschi.    Monit.  Zool.  Ital.,  14. 


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Meisenheiiner,  J. 

1898.  Entwicklungsgeschichte  von  Limax  maxitnus.    Zeit  zinss.  ZooL,  63. 

1899.  Zur  Morphologic  der  Umiere  der  Pulmonaten.    Ibid.,  65. 

1901a.    Entwickelungsgeschichte  von  Dreissensia  polymorpha  Pall.    Ibid,,6g. 
1901b.    Die  Entwickelung  von  Herz.    Pericard,  Niere  und  Gcnitalzellen  bci 
Cyclas  im  Verhaltniss  zu  den  tibrigen  Mollusken.    Ibid.,  69. 

Metalnikoff,  S. 

1900.  Sipunculus  nudus.    Ibid.,  68. 

Meyer,  B. 

1887.  Studien  iiber  den  Korperbau  der  Anneliden.  Mittheil.  zool.  Stat. 
Neapel,  7. 

1888.  Studien  uber  den  Korperbau  der  Anneliden,  IV,  Ibid.,  8. 

1890.    Ueber  die  Nephridien   und  Geschlechtsorgane   von   Lopadorh)mchus. 

Biol.  Centralbl,  10. 
190X.    Studien   uber   den   Korperbau   der   Anneliden.    Mittheil.   zool.   Stat. 

Neapel,  14. 

Meyel^i  H. 

X890.    Die  Entwickelung  der  Urniere  beim  Menschen.    Arch.  mikr.  Anat.,$6. 

Michael,  A.  D. 

1883.  Observations  on  the  Anatomy  of  the  Oribatidx.  Journ.  R.  Micr. 
Soc,  (2),  3. 

1892.  On  the  Variations  in  the  Internal  Anatomy  of  the  Gamasinse,  etc 
Trans.  Linn,  Soc,  London,  5. 

X895.  A  Study  of  the  Internal  Anatomy  of  Thyas  petrophilus,  an  unre- 
corded Hydrachnid  found  in  Cornwall.    Proc.  R.  Soc.  London. 

Mihalkovics,  6.  v. 

1885.  Untersuchungen  iiber  die  Entwickelung  der  Ham-  und  Geschlechts- 
apparates  der  Amnioten.  Intemat.    Monatschr.  Anat.  Physiol,  2. 

Minchin,  B.  A. 

1892.  Notes  on  the  Cuvierian  Organs  of  Holothuria  nigra.  Ann.  Mag.  Nat 
Hist. 

Mollier,  S. 

1890.  Ueber  die  Entstehung  des  Vomierensystems  bei  Amphibien.  Arch. 
Anat.  Physiol. 

Montgomery,  T.  H.  Jr. 

X897.    On  the  Structure  of  the  Nephridia  of  Stichostemma.    Zool.  Jahrb.,  10. 
X903a.    The  adult  Organisation  of  Paragordius  varius  (Leidy).    Ibid.,  i8w 
X903b.    On  the  Morphology  of  the  Rotatorian  Family  Flosculariidse.    Proc. 

Acad,  Nat.  Sci.  Philadelphia, 
X904.    The  Development  and  Structure  of  the  Larva  of  Paragordius.    Ibid. 
X906.    The  Analysis  of  Racial  Descent  in  Animals,   New  York. 

Moore,  J.  P. 

X897.    On  the  Structure  of  the  Discodrilid  Nephridium.    Journ.  Morph.,  13. 

Morgan,  T.  H. 

X894.    The  Development  of  Balanoglossus.    Ibid.,  9. 


628  MONTGOMERY— MORPHOLOGY  OF  THE-  [April 24, 

Moiiiiy  J. 

x888.    Studien    uber   die    Entwicklung   der    Spinnen.     (In    Russian.)     Abh. 
neuruss.  Naturf.  Odessa,  13. 

Monei  £•  S. 

1902.    Observations  on  Living  Brachiopoda.    Mem.  Boston  Soc.  Nat.  Hist.,  5. 

Wmet,  6.  W. 

X894.    Die  Ostracoden  des  Golfes  von  Neapel.    Fauna  u.  Flora  d.  Golfes  v. 
Neapel,  21. 

Mmier,  W. 

1875.    Has  Urogenitalsystem   des   Amphioxus  und   der  Cyclostomen.    Jena. 
Zeit.,  9. 

Nalepa,  A. 

1884.  Die  Anatomie  der  Tyroglyphen,  2.    Sitz.  Ber.  Akad.  Wiss.  Witn,  90. 
X885.    Die  Anatomie  der  Tyroglyphen,  2.    Ibid.,  92. 

1888.    Die  Anatomie  der  Phytopoden.    Ibid.,  96. 

Nansen,  F. 

1885.  Bidrag  til  Myzostomemes  Anatomie  og  Histologic  Bergen. 

Nassonow,  N. 

x886.    Welche  Insectenorgane  durften  homolog  den   Segmentalorganen  der 
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1900.  Zur  Kenntniss  der  phagocytaren  Organe  bei  den  parasitischen  Nema- 
toden.    Arch.  mikr.  Anat.,  55. 

Nelson,  J.  A. 

X904.    The  early  Development  of  Dinophilus.    Proc.  Acad.  Nat.  Sci.  Phila- 
delphia. 
X907.    The  Morphology  of  Dinophilus  Conklini.    Ibid.,  59. 

Nemec,  B. 

1896.    Ueber    Excretionsorgane    und    Geschlechtsverhaltnisse    einiger    Iso- 
poden.    Zool.  Ans.,  19. 

Nettovich,  L.  v. 

X900.    Neue  Beitrage  zur  Kenntnis  der  Arguliden.    Arb.  zool.  Inst.  Wien,  13. 

Nowikoff,  H. 

X905.    Untersuchungen  iiber  den  Bau  der  Limnadia  lenticularis.    Zeit.  wiss. 
Zool.,  78. 

Nttsbamn,  J. 

X885.    Zur   Entwickelungsgeschichte   der   Geschlechtsorgane   der   Hirudinecn 
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1901.  Dybowscella  baicalensis  nov.  gen.  spec.    Biol.  Centralbl.,  21. 

Ostroumoff,  A.  A. 

x886.    Contribution  i  T^tude  zoologique  et  morphologique  des  Bryozoaires 
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Oudemans,  J.  T. 

X887.    Bijdrage  tot-  de  kennis  der  Thysanura  en  Collembola,  Amsterdam. 


,9o8.]  EXCRETORY  ORGANS  OF  METAZOA.  629 

Packard,  A.  S. 

X898.    A  Text-Book  of  Entomology,  New  York. 

Palm6ii,  J.  A. 

1877.    Zur  Morphologie  des  Tracheensystems,  Helsingfors. 

Patten,  W.  and  Hazen,  A.  P. 

1900.  The  Development  of  the  Coxal  Gland,  Branchial  Cartilages,  and  Geni- 
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Pedaschenko,  D.  D. 

1899.  Embryonalentwickelung   und   Metamorphose  von   Lemaea  branchialis 
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Pelseneer,  P. 

1885.    On  the  Coxal  Glands  of  Mygale.    Proc.  Zool.  Soc.  London,  2. 

1901.  fitudes  sur  des  Gastropodes  pulmones.    Mhn.  Acad.  Set.  Belg.,  54. 

Pintner,  T. 

1896.  Studien  iiber  Tetrarh)mchen,  II.  Sitsber.  Akad.  Wiss.  Wien,  105. 

Plate,  L.  H. 

1888.  Beitrage  zur  Naturgeschichte  der  Tardigraden.    Zool.  Jahrb.,  3. 

1889.  Ueber  die  Rotatorien fauna  des  bottanischen   Meerbusens,   etc.     Zeit. 
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Price,  G.  C. 

1897.  Development  of  the  E'xcretory  Organs  of  a  Myxinoid   (Bdellostoma 
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Prooho,  H. 

1890.  Contributions   i   Thistoire   des   Loxosomes.    Arch.   Zool.   exph.  gin. 

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189a.    Contribution  a  Thistoire  des  Bryozoaires.    Ibid.,  10. 

Punnett,  R.  C. 

X900.    On  a  Collection  of  Nemerteans  from  Singapore.    Quart.  Journ.  Micr. 

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The  Enteropneusta.    Fauna  and  Geography  of  the  Maldive  and  Lac- 

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Purcell,  W.  P. 

1895.  Note  on  the  Development  of  the  Lungs,  Entapophyses,  Tracheae  and 
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1900.  On  the  Anatomy  of  Opisthopatus  cinctipes,  Pure,  with  Notes  on  other, 
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Rabl,  C. 

1879.    Ueber  die  Entwicklung  der  Tellerschnecke.    Morph.,  Jahrb.,  5. 

1896.  Ueber  die  Entwicklung  des  Urogenitalsy stems  dei^  Selachier.    Ibid.,  24. 

Reichenbach,  H. 

x886.    Studien  zur  Entwicklungsgeschichte  des  Flusskrebses.    Abh.  Senckenb. 
Nat.  Ges.,  14. 


630  MONTGOMERY— MORPHOLOGY  OF  THE  [Aprils, 

Reinhard,  W. 

1887.  Kinorhyncha  (Echinoderes)  ihr  anatotnischer  Bau  und  ihre  Stdlung 
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Renson,  6. 

1883.    Contributions  a  Tembryologie  des  organes  d'excr^tion  des  oiseaux  et 
des  Mammif^res,  Bruxelles. 

Rho,  F. 

1888.  Studii    sullo    sviluppo   della    Chromodoris   elegans.    AtH  Acad.   Set. 
Napoli,  I. 

Richard,  J.  1 

1892.    Recherches  sur  le  systeme  glandulaire  et  sur  le  systeme  nervcux  des 
Copepodes  libres  d'eau  douce,  etc.    Ann,  Set.  Nat  Paris  (7),  12. 

Rosa,  D. 

1903a.    II    cloragogo    tipico    degli    Oligocheti.    Mem,    Accad,    Sci.    Torino 

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1903b.    Nefridii  di  Rotifero  in  giovani  Lombrichi.    Boll,  Mus.  Zool.  Anat. 

Com  p.  Torino,  18.  n  . 

Roule,  L. 

1889.  fitudes  sur  developpement  des  Ann^lides  et  en  particulier  d'un  oligo- 
chsetc  limicole  marin   (Enchytraeoidcs  Marioni).    Ann.  Sci,  Nat.  (7),  7, 

Rottsselet,  C. 

1891.    On  the  vibratile  tags  of  Asplanchna  amphora.    Joum.  Quekett  Micr. 
Club,  4. 

Rucker,  A. 

1901.    The  Texan  Kcenenia.    Amer.  Nat,,  35. 

Riickert,  J. 

1888.  Ueber   die   Entstehung   der   Excretionsorgane   bei    Selachicm.    Arch. 
Anat.  Physiol, 

Z892.    Entwicklung  der  Excretionsorgane.    Ergebn.  Anat.  Entw.,  i. 

Russo,  A. 

1889.  Sul   valore   morfologice    e    funzionale   degli   organi    di    Cuvier   delle 
Oloturie.    Monit.  Zool.  Ital.,  10. 

Salensky,  W. 

1873.    Beitrage  zur  Entwicklung  der  Prosobranchiaten   und  Referat.    Zeit, 

wiss.  Zool.,  12. 
1885.    fitudes  sur  le  developpement  du  Vermet.    Arch,  de  Biol,  6. 

Sarasin,  C.  F.  und  P.  B. 

1888.    Ueber  die  Niere  der  Seeigel.    Zool.  Anz.,  11. 

Sarasin,  P. 

z883.    Die  Entwicklungsgeschichte  der  Bythynia  tentaculata,  Wiesbaden. 

Schaeppi,  T. 

1894.    Das  Chloragogen  von  Ophelia  radiata.    Jena,  Zeit,,  28. 


x9o8.]  EXCRETORY   ORGANS  OF   METAZOA.  631 

Schaub,  R.  v. 

1888.    Ueber  die  Anatomic  von  Hydrodroma  (C.  L.  Koch).    Sitz,  Ber.  Akad, 
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Schauinslandy  H. 

1886.    Die  Excretions-  und  Geschlechtsorgane  der  Priapuliden.    Zool,  Ans.,g 

Schimk^witsch,  W. 

1884.    Ueber   die    exkretorische    Thatigkeit    des    Mittelarmes   der    Wurmer. 

Biol.  CentralbL,  14. 
1888.    Ueber  Balanoglossus  Mereschkovskii.    ZooL  Am.,  11. 
1895.    Zur  Kenntnis  des  ^aues  und  der  Entwicklung  des  ^Dinophilus  vom 

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1897.    Ueber    die    Entwickelung    des    Darmcanals    bei    einigen    Arachniden. 

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Schmidt,  P. 

1895.    Beitrage  zur  Kenntnis  der  niederen  Myriopoden.    Zeit.  wiss.  Zool.,  $9- 

Schneider,  A. 

1866.    Monographic  der  Nematoden,  Berlin. 

SchttltZy  B. 

1895.    Ueber  den  Process  der  Excretion  bei  den  Holothurien.    Biol.  Cen- 

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19038.    Aus  dem  Gebiete  der  Regeneration,  III.  Zeit.  wiss.  ZooL,  75. 
1903b.    Aus  dem  Gebiete  der  Regeneration,  IV.  Ibid. 

Schultze,  M. 

1854.    Bericht  iiber  einige  in   Herbst   1853  an  der  Kuste  des  Mittelmeeres 

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1883.    Beitrage  zur  Anatomic  des  Excretionsapparates  (Schleifencanale)  der 
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Sedgwick,  A. 

1880.    Development  of  the  Kidney  in  relation  to  the  Wolffian  Body  in  the 

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X885-8.    Development  of  the  Cape   Species  of   Peripatus.    Ibid.,  25-28. 

Seeliger,  0. 

1893.    Tunicata    (Mantelthiere).    Bronn's  Klassen   u.   Ordnungen  d.   Thicr- 
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Seinoiiy  R. 

1891.    Studien  uber  den   Bauplan   des   Urogenitalsystems   der  Wirbelthiere. 

Dargelegt  an  der  Entwicklung  dieses  Organsystems  bei  Ichthyophis  glu- 

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Semper,  C. 

1874.    Die  Stammesverwandtschaft  der  Wirbelthiere  und  Wirbellosten.    Arb. 
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632  MONTGOMERY— MORPHOLOGY  OF  THE  [April  a*. 

Shearer,  C. 

1906a.    On  the  Structure  of  the   Nephridia   of  Dinophilus.    Quart,  lourn. 

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1906b.    Studies  on  the  Development  of  Larval  Nephridia,  Part  i.  Phoronis. 

Mittheil.  zool.  Stat.  NeapeL  17. 

Shipley,  A.  B. 

1890.  On  Phymosoma  varians.    Quart.  Journ.  Micr.  Set.   (2),  31. 

1892.  On  Onchesoma  Steenstrupii.    Ibid.,  33. 

Sluiter,  C.  P. 

1882a.    Beitrage    zu    der    Kenntnis    der    Gephyreen    aus    dem    Malayschen 

Archipel,  i.    Nat.  Tijdsehr,  Nederl.  Indie,  14. 
i88ab.    Idem.,  2.    Ibid. 
1884.    Idem.,  3.    Ibid.,  43. 

Sograf,  N. 

1883. ,  Materiah'en  zur  Kenntnis  der  Embryonalentwicklung  von   Geophilus 

ferrugineus  L.  K.  und  Geophilus  proximus  L.   K.  Nachschr.   Ges.   Fr. 

Naturw.  Moskau,  43. 

Spee,  F. 

1884.  Ueber  direkte  Betheiligung  des  Ektoderms  an  der  Bildung  der  Umi- 
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Spencer,  W.  B. 

X889.    The  giant  Earthworm  of  Gippsland.    Nature,  39. 

Spengel,  J.  W. 

1876.    Das  JLJrogenitalsystem  der  Amphibien,  L  Theil.    Arb.  sool.-soot.  Inst. 

WUrzburg,  3. 
1880.    Beitrage  zur  Kenntnis  der  Gephyreen.    Mittheil.  200I.  Stat.  Neapel,  i. 

1893.  Enteropneusta.    Fauna  u.  Flora  d.  Golfes  v.  Neapel,  18. 

Stauffacher,  H. 

1898.    Die  Umiere  bei  Cyclas  cornea  (Lam.).    Zeii.  wiss.  Zool.,  63. 

Stecker,  A. 

1876.  Anatomisches  und  Histologisches  uber  Gibocellum,  eine  neue  Arach- 
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Stummer-Traunfels,  R.  t. 

1903.  Beitrage  zur  Anatomic  und  Histologic  der  Myzostomen,  i.  Zeit.  wiss 
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Sturany,  R. 

1891.  Die  Coxaldriisen  der  Arachnoiden.    Arb.  sool.  Inst.  Wien,  9. 

Sukatschoff,  B. 

1900.  Beitrage  zur  Entwickelungsgeschichte  der  Hirudineen,  i.  Zur  Kennt- 
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Swaen,  A.  et  Brachet,  A. 

1901.  fitudes  sur  les  premieres  phases  du  developpcment  des  organcs  de- 
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Thon,  K. 

X905.    Neue  Exkretionsorgane  bei  der  Hydrachniden-familie  Limnocharidae 

Kramer.    Zeit,  wiss,  Zool,  79. 
Thor,  S. 
1904.    Recherches   sur   Tanatotnie   compar^e   des   Acariens   prostigmatlques. 

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Todaro,  F. 

xgoaa.    Sopra  gli  organi  escretori  delle  SalpidL    Rend,  Accad  Lincei  (5),  11. 
1909b.    Sur  les  organes  excr^teurs  des  Salpid6s   (Salpidae  Forbes).    Arch, 
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Torrey,  J.  C. 

1903.    The  early  Embryology  of  Thalassema  mellita   (Comi).    Ann,  New 
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Trinchese,  S. 

x88z.    Par  la  fauna  marittima  italiana.    Aeolididae  e  Familie  affini.    Atti  R. 
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VejdOYsk;^,  P. 

1885.  System  und  Morphologie  der  Oligochaeten  Prag. 

1886.  Zur  Morphologie  der  Gordiiden.    Zeit  wiss.  Zool.,  43, 

X890.    Entwickelungsgeschichtliche  Untersuchungen.    Heft  2,    Die  Entwicke- 

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lides  australis.    Arch,  mikr,  Anat.,  40. 
1894.    Organogenie  der  Gordiiden.    Zeit,  zviss.  Zool.,  57. 
Z895.    Zur  vergleichenden  Anatomie  der  Turbellarien,  2.    Ibid.,  60, 
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Verhoeff,  C. 

1902.    Myriopoda.    Bronn's  Klassen  u.  Ordnungen  d.  Thierreichs,  5,  Leipzig. 

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X874.    Monographie  des  Dragonneaux.    Arch.  Zool,  exph.  gin,,  3. 

Voinov,  D.  N. 

X896.    Les  n^phridies   de  Branchiobdella  varians    (V.  astaci).    Mim,  Soc. 
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Wagner,  J. 

i894«    Die   Embryonalentwickelung  von   Ixodes   calcaratus   Bir.   Arb.   soot. 
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Waite,  P.  C. 

Z899.    The  Structure  and  Development  of  the  Antennal  Glands  in  Homarus 
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Waldeyer,  W. 

Z870.    Eierstock  und  El  Leipzig. 

PROC.  AMER.  PHIL.  SOC.,  XLVH.   I90  CO,  PRINTEDBR  FEUARY  6,  I909. 


634  MONTGOMERY— MORPHOLOGY  OF  THE  [April  .4, 

Weber,  E.  P. 

1898.  Faune  rotatorienne  du  bassin  du  Leman.    Rev,  suisse  ZooL,  5. 

Weber,  M. 

x886.    Die  Abdominalporen   der   Salmoniden  nebst   Bemerktmgen   uber  die 

Geschlechtsorgane  der  Fische.    Morph.  Jahrb.,  12. 
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Z874.    Ueber  Bau  und  Lebenserscheinungen  von  Leptodora  hyalina,  Lillje- 
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Weiss,  B. 

Z890.    Excretory  Tubules  in  Amphioxus.    Quart,  Joum,  Micr.  Set.  (2),  31. 

Weldon,  W.  F.  R. 

Z887.    On  Dinophilus  gigas.    Ibid,,  27. 

Wheeler,  W.  M. 

18938.    The  primitive  number  of  Malpighian  vessels  in  Insects.    Psyche,  6. 
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Z899.    The  Development  of  the  Urinogenital  Organs  of  the  Lamprey.    ZooL 
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Whitman,  C.  0. 

Z887.    A  Contribution  to  the  History  of  the  Germ-layers  in  Qepsine.    Joum. 
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Wiedersheim,  R. 

Z890.    Ueber  die  Entwicklung  des  Urogenitalapparates  bei  Crocodilen  und 
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Wienejski,  A. 

1893.    Atrochus  tentaculatus  nov.  gen.  et  spec    Zeit.  vnss.  ZooL,  55. 

Wilhelmi,  J. 

1906.    Untersuchungen  tiber  die  Excretionsorgane  der  Susswassertricladen. 
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Willem,  V. 

1899.  Observations    sur    Texcr^tion    chez    I'Ar^nicole.    Trav,    Stat,    sooL 
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Z899.    Enteropneusta  from  the  South  Pacific,  with  Notes  on  the  West  Indian 
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Williams,  T. 

1858.    Researches   on   the    Structure   and   Homology   of    the   Reproductive 
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Wilson,  H.  V. 

X89Z.    The  Embryology  of  the  Sea  Bass  (Serranus  atrarius).    Bull.  U.  S. 
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Winkler,  W. 

1888.  Anatomie  der  Gamasiden.    Arh.  sooL  Inst,  Wien.,  7. 

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1904.    The  Notostigmata,  a  new  suborder  of  Acari.    Vid.  Medel,  Nat  For. 
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Witlacxil,  B. 

i88a.    Zur  Anatomie  der  Aphiden.    Arh,  sool,  Inst,  Wien,  4. 

Wolf  son,  W. 

x88o.    Die   embryonale    Entwicklung   des   Limnaeus    stagnalis.    Bull.   Acad, 
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Woltereck,  R. 

1909.    Trochophora-Studien.  i.  Ueber  die  Histologie  der  Larve  und  die  Ent- 
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Z889.    Ueber  die  Mesodermsegmente  des  Rumpfes  und  die  Entwicklung  des 
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Zelinka,  C. 

1886.    Studien  uber  Raderthiere,  i.    Arb.  xool,  Inst,  Gras,  i. 

z888.    Idem.,  2,    Ibid,,  2, 

1889.  ^ic  Gastrotrichen.    Ibid,,  3. 

X891.    Studien  uber  Raderthiere,  3.    Ibid.,  3. 

Ziegler,  H.  B. 

1898.    Ueber  den  derzeitigen  Stand  der  Colomfrage.    Verh,  deutsch.  Zool 
Ges, 

Zur  Strassen,  0. 

1892.    Bradynema  rigidum  v.    Sieb.  Zeit,  wiss.  Zool.  54. 


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Bv  KUNO  FRANCKR 
(Read  April  2$,  1908.) 

There  is  a  curious  anomaly  in  the  equipment  of  German  uni- 
versities, an  anomaly  accounted  for  partly  by  the  traditional  cosmo- 
politanism of  German  scholarship,  partly  by  the  somewhat  belated 
development  of  Germany  into  a  united  and  powerful  nation. 

Whereas  for  sudents  of  classical  archaeology  there  is  provided  in 
nearly  every  university  of  the  fatherland  a  well-planned  and  sys- 
tematically arranged  museum  of  casts  of  Greek  sculptures,  the 
student  of  German  history  would  not  find  at  a  single  one  of  these 
universities  any  collection  which  would  offer  to  him  a  fairly  accurate 
representation  of  the  artistic  development  of  his  own  country. 
Even  in  the  German  capital  with  its  wealth  of  ethnological  and 
archaeological  exhibits  from  Troas  and  Pergamon,  from  Egypt  and 
Assyria,  from  India  and  South  America,  no  attempt  has  as  yet  been 
made  to  bring  together,  in  reproductions,  the  great  artistic  landmarks 
of  Germany  herself.  It  has  been  reserved  to  an  American  university 
to  make  at  least  a  beginning  of  such  an  undertaking,  but  it  is  inter- 
esting to  note  that  the  Germanic  Museum  of  Harvard  University 
could  not  have  achieved  whatever  success  it  has  had  thus  far,  had  it 
not  been  for  the  generous  interest  bestowed  upon  it  by  His  Majesty 
the  German  Emperor.  So  that  this  museum,  although  established 
on  non-German  soil,  is  after  all  in  its  way  another  symptom  of  the 
long  strides  which  modem  Germany  has  made  toward  national  great- 
ness and  international  influence. 

The  bulk  of  the  collections  of  the  Germanic  Museum  at  Cam- 
bridge is  devoted  to  German  sculpture  of  the  Middle  Ages  and  the 
Renaissance,  and  particular  stress  is  laid  upon  a  good  representation 
of  the  thirteenth  century. 

It  is  not  as  generally  acknowledged  as  it  should  be  that  the  thir- 

636 


X908.J    IN  THE  GERMANIC  MUSEUM  OF   HARVARD  UNIVERSITY.    637 

teenth  century  marks  a  truly  classic  epoch  in  the  development  of 
German  plastic  act.  German  sculpture  between  1220  and  1250  is 
fully  on  a  level  with  the  great  creations  of  the  lyric  and  epic  poetry 
of  chivalry;  and  no  one  who  is  susceptible  to  the  peculiar  beauty 
of  Walther  von  der  Vogelweide's  minne-song  or  is  impressed  with 
the  heroic  figures  of  the  Nibelungenlied,  of  Kudrun,  of  Parzival,  or 
Tristan,  can  fail  to  observe  their  affinity  of  spirit  with  the  plastic 
monuments  of  Wechselburg  and  Freiberg,  of  Naumburg  and  Hal- 
berstadt,  of  Bamberg  and  Strassburg.  Here  as  well  as  there  we 
find  a  high  degree  of  refinement  and  measure ;  a  strenuous  insistence 
on  courteous  decorum ;  intense  moral  earnestness  linked  to  a  strange 
fancifulness  of  imagination;  a  curious  combination  of  scrupulous 
attention  to  certain  conventional  forms  of  dress,  gesture,  and  ex- 
pression, on  the  one  hand,  and  a  free  sweep  in  the  delineation  of 
character,  on  the  other.  Here  as  well  as  there  we  find  a  happy 
union  of  the  universally  human  with  the  distinctively  mediaeval ;  a 
wonderful  blending  of  the  ideal  human  type  with  the  characteristic 
features  of  the  portrait.  As  the  art  of  Phidias  and  Praxiteles  is  an 
indispensable  supplement  to  the  art  of  ^Eschylus  and  Sophocles,  for 
oyr  understanding  of  Attic  culture  in  its  prime,  so  these  works 
of  German  sculpture  of  the  thirteenth  century  stand  to  us  (or 
should  stand  to  us)  by  the  side  of  the  great  productions  of  the 
chivalric  poets,  as  incontrovertible  proofs  of  the  free  and  noble 
conception  of  humanity  reached  by  mediaeval  culture  at  its  height. 

A  brief  review  of  a  few  at  least  of  these  sculptures  may  serve 
to  elucidate  this  statement  somewhat  more  fully. 

Among  the  earliest  plastic  monuments  of  the  thirteenth  century 
are  the  pulpit  and  the  Crucifixion  group  of  the  Church  of  Wechsel- 
burg in  Saxony,  executed  probably  between  12 10  and  1220.  In  both 
monuments  it  seems  as  though  the  artist  was  still  grappling  with  the 
problem  of  form.  In  the  relief  from  the  front  of  the  pulpit — 
Christ  seated  on  the  throne  as  Judge  of  the  world,  surrounded  by  the 
symbols  of  the  Evangelists — mastery  of  form,  classic  solemnity, 
exalted  repose  have  indeed  been  attained.  In  the  more  animated 
scenes  of  the  side  reliefs — the  sacrifice  of  Isaac  and  the  healing  of 
the  Jews  by  the  brazen  serpent — there  is  a  curious  contrast  between 
grandeur  and  awkwardness,  sweetness  of  feeling  and  naive  natural- 


638  FRANCKE— MEDIAEVAL  GERMAN   SCULPTURE  (April  as. 

ism.  And  a  similar  contrast  is  found  in  the  Crucifixion  group.  The 
figures  of  Mary  and  John  standing  under  the  Cross,  as  well  as  that 
of  Joseph  of  Arimathea  holding  out  the  cup  to  receive  the  blood 
of  the  Saviour,  are  remarkable  for  nobility  of  outline,  depth  of  feel- 
ing, and  measured  beauty  of  expression.  There  is  a  fine  sweep  of 
movement  in  the  two  angels  on  the  cross-beam,  gentle  sadness  in  the 
figure  of  Christ,  and  a  mild  tenderness  in  the  attitude  of  God  the 
Father  appearing  above.  The  symbolical  figures,  however — ^prob- 
ably Jewdom  and  Pagandom — on  which  John  and  Mary  are  stand- 
ing, are  tortuous  and  forced.  Apparently,  here  is  an  artist  who 
looks  at  life  about  him  with  a  keen,  penetrating,  and  receptive  eye, 
but  who  at  the  same  time  is  impelled  to  subject  reality  to  certain 
canons  of  measure  and  proportion  which  he  has  not  yet  made  fully 
his  own. 

A  decided  step  in  advance  is  made  in  the  sculptures  of  the  Golden 
Gate  of  the  Cathedral  of  Freiberg,  likewise  in  Saxony.  In  the 
arrangement  of  plastic  figures,  both  on  the  sides  of  the  portal  and 
on  the  archivolts,  French  influence  is  clearly  seen.  But  these  plastic 
figures  seem  here  much  more  independent  of  the  architectural  frame- 
work than  is  common  in  the  French  sculptures,  e.  g,,  those  of  Char- 
tres  Cathedral,  which  served  as  models  to  the  German  artist ;  and  the 
human  type  and  bodily  proportions  are  unnrtstakably  original. 

A  thoroughly  satisfactory  interpretation  of  all  the  figures,  human, 
animal  and  fantastic,  which  cover  the  sides  of  the  portal,  the  tympa- 
num and  the  archivolts,  and  of  the  fundamental  conception  under- 
lying them,  has  not  yet  been  g^ven,  although  Anton  Springer  has 
done  a  great  deal  for  the  identification  of  individual  personages. 
Springer  thinks  that  the  fundamental  conception  of  the  whole  is 
the  mystic  marriage  between  Christ  and  the  Church,  and  that  all  the 
scenes  and  figures  of  the  portal  may  be  interpreted  as  symbolic  of 
this  mystic  idea.  Simpler  and  more  plausible  it  seems  to  me  to 
find  in  this  portal  a  plastic  counterpart  to  dramatic  scenes  from  the 
cycle  of  the  Christmas  plays,  the  popularity  of  which  in  the  thir- 
teenth century  is  proved,  for  Germany,  by  a  particularly  complete 
example,  the  Benediktbeuren  Christmas  Play.  Clearly  a  scene  from 
the  Christmas  cycle  is  the  one  represented  in  the  tympanum  of  the 
portal :  the  Adoration  of  the  Magi,  the  three  kings  approaching  from 


i9o8.]   IN  THE  GERMANIC  MUSEUM  OF  HARVARD  UNIVERSITY.    639 

the  left,  Mary  with  the  child  enthroned  in  the  middle,  the  archangel 
Gabriel  and  Joseph  at  the  right.  And  no  less  plausibly  than  this 
scene  may  the  eight  somewhat  under  life-size  figures  which  Hank 
both  sides  of  the  portal  be  connected  with  the  subject  of  the  Christ- 
mas plays.  Prophet  and  Sibyl  scenes  were  very  frequently  used  as 
introducing  the  Nativity  play  proper,  one  prophet  or  Sibyl  after 
another  entering  to  testify  to  the  coming  of  the  Saviour.  While 
retaining  most  of  the  names  suggested  by  Springer  for  these  eight 
figures, — ^John  the  Baptist  and  John  the  Evangelist,  David  and  Solo- 
mon, the  Queen  of  Sheba  and  Bathseba,  David  and  Aaron, — we  may 
call  them  collectively  witnesses  to  Christ's  Nativity. 

As  to  the  plastic  representations  on  the  four  archivolts  encircling 
the  tympanum,  they  are,  to  be  sure,  not  taken  f  rc«n  any  actual  scene 
of  a  Christmas  play ;  but  they  are  entirely  in  keeping  with  the  joy- 
ous, idyllic  character  of  these  plays.  On  the  innermost  archivolt, 
nearest  to  the  Adoration  of  the  Magi,  there  are  at  the  sides  the  four 
archangels,  in  worshipful  attitude ;  in  the  middle,  the  Coronation  of 
Mary  by  Christ.  The  next  archivolt  contains  six  apostles,  three  at 
each  side,  and  in  the  center  Abraham  with  a  soul  of  the  blessed  in 
his  lap,  while  an  angel  reaches  out  another  soul  toward  him.  The 
third  archivolt  shows  eight  figures  of  .apostles  and  in  the  center 
the  dove  of  the  Holy  Ghost  surrounded  by  angels.  On  the  outer- 
most archivolt,  finally,  the  resurrection  of  the  flesh  is  represented  by 
ten  figures  rising  from  their  graves  with  manifoldly  varying  expres- 
sions of  faith,  hope  and  exultation ;  while  the  central  group,  an  angel 
receiving  by  either  hand  a  saved  soul,  fittingly  symbolizes  the  last 
and  highest  stage  of  human  redemption.  All  these  sculptures,  as 
well  as  those  of  the  tympanum  and  the  sides  of  the  portal,  are  dis- 
tinguished by  a  remarkable  symmetry  and  adjustment  to  architec- 
tural demands,  and  by  a  wonderful  mellowness  and  purity  of  form 
and  an  exquisite  sweetness  and  serenity  of  expression,  making  an 
artistic  whole  of  extraordinary  beauty  and  perfection. 

The  climax,  however,  of  North  German  art  of  the  thirteenth 
century  is  reached  in  the  Portrait  Statues  of  Founders  and  Patrons 
of  Naumburg  Cathedral  from  the  west  choir  of  that  church,  a  series 
of  works  which  may  be  definitely  assigned  to  the  middle  of  the  thir- 
teenth century.    These  statues,    together    with  that    of  a    young 


640  FRANCKE— MEDIi€VAL  GERMAN  SCULPTURE  [Ap«il«s, 

ecclesiastic  from  the  same  church,  are  a  striking  refutaticm  of  what 
since  Jacob  Burckhardt's  "  Kultur  der  Renaissance  in  Italien  "  has 
come  to  be  a  popular  axiom,  the  assumption,  namely,  that  modern 
individualism  had  its  origin  in  the  era  of  the  rinascimento ;  they 
show  conclusively  that  Burckhardt*s  phrase  of  "  the  discovery  of  the 
individual "  by  the  great  Italians  of  the  quatro-cento  is  misleading, 
that,  in  other  words,  the  Middle  Ages  themselves  contain  the  germs 
of  modern  individualism.  There  is  nothing  in  the  art  of  the  Renais- 
sance which  surpasses  these  Naumburg  statues  in  fulness,  distinct- 
ness, and  vigor  of  individual  life.  Every  one  of  these  figures  is  a 
type  by  itself,  a  fully  rounded  personality.  The  two  pairs  of  princely 
husband  and  wife,  one  of  the  men  full  of  power  and  determination, 
the  other  of  youthfully  sanguine  appearance,  one  of  the  wcwnen 
broadly  smiling,  the  other,  with  a  gesture  full  of  reserved  dignity, 
drawing  her  garment  to  her  face ;  the  canoness  standing  erect,  but 
with  slightly  inclined  head,  thoughtfully  gazing  down  upon  a  book 
which  she  supports  with  one  hand  while  the  other  turns  over  its 
leaves ;  the  princess  drawing  her  mantle  about  her ;  the  young  eccle- 
siastic with  his  carefully  arranged  hair  flowing  from  his  tonsure, 
holding  the  missal  in  front  of  him ;  the  various  knights,  one  looking 
out  from  behind  his  shield,  another  supporting  his  left  on  the  shield 
and  shouldering  the  sword  with  his  right  hand,  a  third  resting  both 
shield  and  sword  in  front  of  him  on  the  ground,  while  with  his  right 
hand  he  gathers  his  mantle  about  his  neck,  others  in  still  different 
postures  and  moods, — there  is  not  a  figure  among  them  which  did 
not  represent  a  particular  individual  at  a  particular  moment,  and 
which  did  not,  without  losing  itself  in  capricious  imitation  of  acci- 
dental trifles,  reproduce  life  as  it  is.  It  is  impossible  in  the  face 
of  such  works  of  sculpture  as  these  not  to  feel  that  they  proceeded 
from  artists  deeply  versed  in  the  study  of  human  character,  fully 
alive  to  the  problems  of  human  conduct,  keenly  sensitive  to  im- 
pressions of  any  sort — in  other  words,  fully  developed,  highly  or- 
ganized, complicated  individuals.  One  feels  that  here  are  seen  the 
mature  artistic  fruits  of  the  great  Hohenstaufen  epoch — an  epoch 
rent  by  tremendous  conflicts  in  church  and  state,  and  convulsed  by 
the  throes  of  a  new  intellectual  and  spiritual  birth. 

Almost  contemporary  with  these  statues,  though  probably  some- 


,9o8.1    IN  THE  GERMANIC  MUSEUM    OF  HARVARD  UNIVERSITY.    641 

what  younger,  is  the  Naumburg  Rood  Screen  separating  the  west 
choir  of  the  Cathedral  from  the  nave.  The  sculptures  of  this  rood 
screen  form  an  interesting  contrast  to  the  sculptures  of  the  Freiberg 
Golden  Gate,  analyzed  before.  While  the  Freiberg  sculptures  pre- 
sent a  plastic  counterpart  to  the  mediaeval  Christmas  plays,  we  have 
in  the  Naumburg  rood  screen  a  plastic  counterpart  to  the  Passion 
plays.  On  the  middle  beam  of  the  door  leading  through  the  screen, 
which  has  the  shape  of  a  cross,  the  figure  of  the  dying  Saviour  is 
suspended,  while  on  each  side  of  the  door  there  stand  in  niches  the 
over  life-size  figures  of  Mary  and  John.  The  other  scenes  of  the 
Passion,  from  the  Last  Supper  to  the  Bearing  of  the  Cross,  are 
brought  to  view  in  high  reliefs  which  as  a  continuous  frieze,  crowned 
by  a  Gothic  canopy,  give  to  the  whole  structure  a  most  impressive 
attic-like  top.  These  sculptures  seem  to  mark  a  stage  of  develop- 
ment somewhat  beyond  that  reached  by  the  Naumburg  portrait 
statues.  They  are  signalized  by  intense  dramatic  power.  Some  of 
the  scenes  of  the  frieze  in  particular  impress  one  as  direct  transpo- 
sitions into  stone  of  scenes  from  the  Passion  Play  stage.  They 
excel  even  the  portrait  statues  in  freedom  and  sweep  of  movement 
and  in  keenness  of  realistic  characterization.  On  the  other  hand, 
they  show  a  tendency  toward  exaggeration,  which  occasionally  (as 
in  John  and  Mary)  leads  to  a  strained  and  distorted  expression  of 
feeling;  and,  in  the  portrayal  of  the  vulgar  and  the  commonplace, 
they  occasionally  (as  in  the  representatives  of  the  Jewish  rabble) 
diverge  into  caricature.  They  are,  then,  clear  anticipations  of  the 
ultra-naturalistic,  and  therefore  unnatural  tendency  of  later  Gothic 
sculpture. 

We  may  properly  close  our  review  by  selecting  at  least  one 
group  of  South  German  sculptures  affording  a  striking  example  of 
the  strong  influence  exerted  by  French  Gothic  art  upon  this  part  of 
Germany :  I  mean  the  "  Death  of  Mary  "  and  the  "  Ecclesia  and 
Synagoga  "  from  the  Romanesque  portal  of  Strassburg  Cathedral. 
The  Death  of  Mary  is  one  of  the  noblest  creations  in  the  whole 
history  of  art.  The  Virgin  is  represented  reclining  on  a  couch, 
wrapped  in  a  garment  which  reveals  with  rare  delicacy  the  lines  of 
her  body.  Her  face  is  majestic,  Juno-like.  Although  the  moment 
represented  is  after  her  death,  her  eyes  are  still  open  and  have  a 


642  FRANCKE— MEDIAEVAL  GERMAN   SCULPTURE  [April  25. 

look  of  heavenly  exaltation.  Behind  her  couch,  in  the  middle  of 
the  tympznvm,  stands  Christ,  holding  Mary's  soul  (in  the  form  of 
an  infant)  in  his  left  hand,  his  right  hand  raised  in  blessing.  Mary 
Magdalen  cowers  in  front  of  the  couch,  wringing  her  hands,  her 
face  expressing  deepest  sorrow.  The  space  at  the  sides  and  back 
of  the  death-bed  is  filled  with  the  figures  of  the  Disciples,  seme  of 
them  giving  way  to  grief,  others  contemplative,  others  transfigured, 
all  of  them  filled  with  holy  awe  and  deep  religfious  feeling.  The 
graceful  vine  which  runs  along  the  edge  of  the  Romanesque  arch 
of  the  tympanum  gives  to  the  whole  composition  a  fitting  enclosure. 
In  this  monument  the  French  sense  of  form  and  German  feeling 
seem  most  happily  blended. 

Of  no  less  refinement  are  the  statues  of  Ecclesia  and  Synagoga. 
To  contrast  the  Church  triumphant  and  the  Synagogue  defeated  was 
a  very  common  conception  both  in  the  religious  sculpture  and  in 
the  religious  drama  of  the  Middle  Ages.  Noteworthy  instances  of 
their  occurrence  in  sculpture  are  the  statues  of  Rheims  Cathedral, 
the  north  portal  of  Bamberg  Cathedral,  and  the  vestibule  of  the 
Cathedral  of  Freiburg  im  Breisgau;  of  their  introduction  into  the 
drama,  the  part  played  by  them  in  the  Ludus  de  Antichristo  and 
the  Alsfeld  Play.  Of  all  plastic  representations,  these  Strassburg 
statues  are  the  most  exquisite.  The  Church,  with  wide-flowing 
mantle,  the  crown  on  her  head,  her  right  hand  holding  the  standard 
of  the  cross,  her  left  bearing  the  communion  chalice,  stands  erect 
and  dignified  at  the  left  side  of  the  portal,  looking  with  pride  and 
disdain  at  her  adversary  on  the  opposite  side.  The  Synagogue 
wears  neither  crown  nor  mantle ;  in  her  left  hand  she  holds  the  table 
of  the  Mosaic  law  turned  downward,  in  the  right  a  standard,  the 
shaft  of  which  is  broken  in  many  places;  her  eyes  are  bandag^ 
(to  indicate  that  she  does  not  see  the  true  light),  and  her  face  is 
turned  away  from  the  Church  and  is  bent  slightly  down.  In  spite 
of  her  humiliation,  she  appears  more  human  and  lovable  than  her 
victorious  rival.  Both  figures  together  are  perhaps  unsurpassed  in 
mediaeval  sculpture  for  gjace  and  delicacy  of  outline;  only  in  the 
somewhat  coquettish  twist  of  the  hips  there  is  observable  a  slight 
indication  that  the  highest  point  in  the  classic  epoch  of  plastic  art 


f9o8.]  MINUTES.  643 

has  already  been  passed  and  that  the  age  of  extravagant  emotion 
and  artificiality  is  setting  in. 

When,  in  November,  1903,  these  and  other  precious  gifts  of  the 
German  Emperor  were  temporarily  installed  in  the  insig^nificant 
little  building  which  Harvard  University  could  spare  for  them  as  a 
scanty  shelter,  it  was  hoped  that  only  a  short  time  would  elapse 
before  a  new  and  worthy  museum  building  would  have  been  erected 
through  the  liberality  of  American  friends  of  German  culture. 
These  hopes  have  not  yet  been  fulfilled.  Here  is  the  opportunity 
for  our  fellow  citizens  of  German  origin  to  prove  to  the  world  that 
they  do  not  leave  their  ideals  at  home  when  they  leave  the  father- 
land; and  here  is  a  chance  for  all  Americans  to  show  their  appre- 
ciation of  what  German  culture  has  given  to  this  country. 
Cambridge,  Mass. 


644  MINUTES.  [Oaober  a. 

Stated  Meeting  October  2,  ipo8. 

Secretary  Holland  in  the  Chair. 

Dr.  E.  A.  Spitzka,  a  newly  elected  member,  was  presented  to  the 
chair  and  took  his  seat  in  the  Society. 

Letters  accepting  membership  were  read  from : 
Prof.  Richard  Hawley  Tucker. 
Prof.  Albrecht  F.  K.  Penck. 
Prof.  Herbert  Weir  Smyth. 
Letters  were  received 
From  the  City  of  Faenza  inviting  the  Society  to  be  represented 

at  the  Torricelli  tercentenary. 
From  the  University  of  Cambridge  inviting  the  Society  to  par- 
ticipate in  the  commemoration  of  the  centenary  of  Charles 
Darwin's  birth  in  June,  1909,  and  Prof.  Henry  F.  Osbom 
was  appointed  to  represent  the  Society  on  the  occasion. 
From  the  Physico-Medical  Society  at  Erlangen,  thanking  the 
Society  for  its  congratulatory  address  on  the  occasion  of  its 
centenary  celebration. 
The  decease  was  announced  of : 
Hon.  Grover  Cleveland,  at  Princeton,  N.  J.,  on  June  25,  1908, 

aet.  71. 
Prof.  F.  L.  Otto  Rohrig,  at  Pasadena,  Cal.,  on  July  14,  1908, 

aet.  89. 
Dr.  Ainsworth  Rand  Spofford,  at  Holderness,  N.  H.,  on  August 

12,  1908,  set.  83. 
Prof.   Antoine  Henri   Becquerel,  at   Croisic,  in   Brittany,   on 

August  25,  1908,  aet.  56. 
Prof.  E.  Mascart,  at  Paris,  on  August  26,  1908,  aet.  71. 
Prof.  Dr.  Hugo  von  Meltzel,  of  Koloszvar,  Hungary. 
The  following  papers  were  read: 

"  The   Humming  Telephone,"   by   Prof.   A.   E.   Kennelly   and 
Walter  L.  Upson.     (See  page  329.) 

"  On  the  After-images  of   Subliminally  Colored   Stimuli,"   by 
Edward  Bradford  Titchener  and  William  Henry  Pyle.     (See  page 

366.) 


I908]  MINUTES.  646 

Stated  Meeting  October  i6,  ipo8. 
President  Keen  in  the  Chair. 

The  decease  was  announced  of  President  Daniel  Coit  Oilman,  at 
Norwich,  Conn.,  on  October  15,  1908,  aet.  77. 

Dr.  Edward  O.  Hovey  read  a  paper  entitled  "  A  contribution  to 
the  History  of  Mont  Pelee,  Martinique." 


Stated  Meeting  November  6,  ipo8. 
President  Keen  in  the  Chair. 

A  letter  was  received  from  the  Board  of  Curators  and  Faculty 
of  the  University  of  Missouri,  inviting  the  Society  to  be  represented 
at  the  inauguration  of  Albert  Ross  Hill,  LL.D.,  as  president  of  the 
University,  at  Columbia,  Mo.,  on  December  10  and  11,  1908,  and 
Dr.  William  Trelease  was  appointed  to  represent  the  Society  on  the 
occasion. 

The  decease  was  announced  of  Prof.  Otis  T.  Mason,  at  Washing- 
ton, D.  C,  on  November  5,  1908,  aet.  70. 

Dr.  Alexis  Carrel  read  a  paper  entitled  "  Recent  Studies  in 
Transplantation  of  Organs  in  Animals  "  (see  page  677),  which  was 
discussed  by  President  Keen,  Dr.  CopHn,  Dr.  Eshner  and  Dr.  Carrel. 


HEPATOSCOPY  AND  ASTROLOGY  IN  BABYLONIA  AND 

ASSYRIA. 

By  morris  JASTROW,  Jr. 
(Read  December  4,  1908.) 

In  any  general  study  of  the  subject  of  divinatiwi  we  must  dis- 
tinguish between  two  forms  which  for  want  of  a  better  designation 
we  may  distinguish  as  voluntary  and  involuntary.  Under  voluntary 
divination  is  meant  the  act  of  deliberately  seeking  out  some  object 
or  means  through  which  one  may  hope  to  pierce  the  unknown  future, 
hidden  from  the  ordinary  gaze.  The  placing  of  marked  arrows 
before  the  image  of  a  deity,  and  according  to  the  ones  drawn  by  lot, 
to  determine  what  the  god  may  have  in  mind  or  what  his  pleasure 
may  be  is  an  illustration  of  voluntary  divination  as  practiced  among 
the  ancient  Arabs.^  Sending  out  birds  selected  for  the  purpose  and 
noting  the  direction  and  manner  of  their  flight*  may  be  instanced  as 
another  procedure  of  direct  divination.  Among  the  Babylonians 
and  Assyrians,  the  common  method  of  voluntary  divination  was 
the  examination  of  the  liver  of  the  sacrificial  animal — invariably 
for  this  purpose  a  sheep — ^and,  according  to  signs  noted  in  the 
various  parts  of  that  organ,  to  diagnose  the  intentions  of  the  gods 
as  the  arbiters  of  human  fate  and  as  the  powers  presiding  over  all 
occurrences  on  earth. 

Involuntary  divination,  on  the  other  hand,  rests  on  the  interpre- 
tation of  all  manner  of  signs  and  phenomena  that  without  being 
sought  out  force  themselves  on  our  notice.  Preeminent  among  such 
signs  is  the  observation  of  the  phenomena  of  the  heavens,  primarily 
the  movements  and  aspects  of  the  sun,  moon  and  planets  with  the 
gradual  extension  to  the  observation  of  clouds,  of  constellations  and 
of  single  particularly  prominent  stars — as  practiced  by  the  cultural 

*Wellhausen,  "Reste  Arabischen  Heidenthums,"  p.  126. 
*Wissowa,  "Religion  der  Romer/'  p.  457,  note  3. 

646 


.908.] 


IN  BABYLONIA  AND  ASSYRIA.  647 


nations  of  antiquity.^  In  addition  to  this  branch  of  involuntary 
divination,  we  have  the  significance  attached  to  diverse  occurrences 
that  by  their  more  or  less  unusual  or  striking  character  attract  at- 
tention or  that  for  any  other  reason  were  regarded  as  fraught  with 
some  special  importance.  The  interpretation  of  dreams  falls  within 
this  category.  Monstrosities  among  human  beings  and  animals 
form  another  subdivision,  while  peculiar  actions  among  animals — 
snakes,  dogs,  ravens,  locusts  and  the  like — further  extend  the  scope 
of  involuntary  divination  until  it  becomes  practically  boundless. 
All  the  little  mishaps  and  accidents  of  daily  life  were  looked  upon 
as  sig^s,  indicative  of  the  disposition  of  the  gods  towards  men,  and 
in  a  still  larger  sense,  as  aflfecting  the  general  welfare,  were  storms, 
floods,  swollen  streams,  climatic  disturbances  and  more  the  like. 

In  order  to  differentiate  between  these  two  methods  of  divination 
we  may  designate  the  signs  derived  from  voluntary  divination  as 
omens,  and  those  obtained  from  involuntary  divination  as  portents, 
while  within  the  field  of  involuntary  divination  two  broad  divisions 
may  be  recognized,  the  one  represented  by  portents  connected  with 
the  phenomena  of  the  heavens,  including  clouds,  storms  and  rains, 
and  such  as  are  connected  with  terrestrial  phenomena.  In  grouping 
the  portents  derived  from  the  observation  of  the  phenomena  of 
nature  under  the  general  heading  of  astrology,  it  must  therefore  be 
borne  in  mind  that  the  term  includes  more  than  the  mere  study  of 
the  stars,  but  so  far  at  least  as  Babylonia  and  Assyria  are  concerned, 
there  is  no  distinction  between  the  character  of  the  interpretations 
offered  for  the  phenomena  of  the  heavens  in  the  narrower  sense,  and 
such  phenomena  as  are  merely  associated  with  the  heavens.  For\ 
the  Babylonians  and  Assyrians,  as  for  the  nations  of  antiquity  in 
general,  heaven  is  not  very  far  removed  from  the  earth.*  It  was/ 
supposed  to  begin  where  the  solid  earth  came  to  an  end  and  indeed 
the  tops  of  mountains  so  frequently  enveloped  in  clouds  appear  to 
have  been  regarded  as  bordering  on  the  domain  of  heaven  if  not 

•Dr.  J.  G.  Frazer  calls  my  attention  to  the  fact  that  astrology  in  any 
proper  sense  is  not  found  among  peoples  of  primitive  culture. 

*This  view  underlies  the  Biblical  story  of  the  building  of  the  Tower 
of  Babel  (Genesis,  chapter  XL),  as  is  shown  by  the  circumstance  that  the 
task  of  building  a  tower  which  should  reach  to  heaven  is  not  looked  upon 
as  an  impossible  task  but  as  a  wicked  one. 


648  JASTROW— HEPATOSCOPY   AND   ASTROLOGY         [December  4. 

actually  a  part  of  it.  Hence  to  place  the  seat  of  the  gods  on  the 
tops  of  mountains,  as  was  so  frequently  done  by  nations  of  antiquity, 
was  equivalent  to  assigning  them  to  the  heavens. 

Confining  ourselves  to  Babylonia  and  Assyria,  we  find  that  al- 
though divination  through  the  interpretation  of  terrestrial  phenom- 
ena— dreams,  monstrosities,  actions  of  animals,  mishaps,  swollen 
streams,  etc.,  etc. — also  play  a  prominent  role  and  that  within  the 
field  of  voluntary  divination  we  have  by  the  side  of  hepatoscppy  (or 
divination  through  the  liver),  other  procedures  such  as  the  interpre- 
tation of  the  action  of  oil  bubbles  in  a  basin  of  water,"  the  two  chief 
niethods  of  divination,  forming  part  of  the  official  cult,  are  Hepa^ 
toscopy  and  Astrology."  Both  forms  were  developed  into  elaborate 
/  systems  marked  by  definite  rules  of  interpretation,  consistently  and 
logically  applied.  Extensive  collections  of  omens  and  portents  were 
compiled  by  Babylonian  and  Assyrian  priests  attached  to  the  temples, 
in  which  all  signs  noted  on  the  liver  of  sheep  and  all  manner  of 
phenomena  observed  in  connection  with  sun,  moon,  planets,  con- 
t  stellations  and  stars  on  the  one  hand,  and  with  clouds,  storms,  rains 
.  and  floods  on  the  other,  were  entered  together  with  the  interpretation 
^of  the  signs.  The  evident  endeavor  of  the  compilers  was  to  make 
the  collections  as  comprehei^sive  as  possible  so  as  to  provide  for  all 
/  contingencies,  since  the  purpose  of  the  collections  was  to  serve  as 
guides  and  handbooks  for  the  priests  in  their  practical  labors  as  well 
as  text-books  in  instructing  the  pupils  of  the  temple  schools.  As  a 
consequence,  considerable  skill  and  ingenuity  were  displayed  in  ar- 
ranging the  omens  and  portents  systematically  so  as  to  facilitate 
their  use.  On  the  other  hand,  while  the  signs  noted  were  primarily 
based  on  actual  cases,  the  theoretical  factor  enters  largely  into  play. 
This  led  to  many  signs  being  entered  in  both  classes  of  divination 

•See  Hunger,  " Becherwahrsagung  bei  den  Babyloniern *'  (Leipzig,  1903). 

•For  details  with  copious  translations  of  texts  see  the  writer's  "Religion 
Babyloniens  und  Assyriens,"  parts  10  to  14,  as  well  as  various  articles  on 
special  points  such  as  "  The  Signs  and  Names  for  the  Liver  in  Babylonian  *' 
(Zeitschrift  fiir  Assyriologie,  XX.,  pp.  105-129)  ;  "  The  Liver  in  Antiquity 
and  the  Beginnings  of  Anatomy "  ( University  of  Pennsylvania  Medical 
Bulletin,  January,  1908,  and  Trans,  of  the  Phila.  College  of  Physicians,  3d 
Series,  XXIX.,  pp.  1 17-138)  ;  "  Sign  and  Name  for  Planet  in  Babylonian " 
(Proceedings  of  the  Amer.  Philos.  SoaETV,  XLVIL,  pp.  141-156). 


x9o8.]  IN   BABYLONIA   AND   ASSYRIA.  649 

wJWch  r^resent  such  as  in  the  opinion  of  the  priests  might  occur. 
Certain  rules  of  interpretation  having  been  devised,  based  on  actual 
occurrences  following  upon  the  signs  noted,  these  rules  were  ap- 
plied to  contingent  cases  which  might  occur;  and  often  in  astro- 
logical texts,  signs  are  even  entered  which  have  no  practical  sig- 
nificance at  all  but  purely  a  theoretical  interest  as  illustrations  of 
the  extremes  to  which  the  system  of  interpretation  was  pushed. 

In  the  case  of  both  methods  the  interpretations  have  reference 
almost  exclusively  to  the  general  welfare  and  not  to  the  individual, 
to  crops,  war,  pestilence,  victory,  defeat,  famine,  plenty,  favorable 
or  unfavorable  climatic  conditions  and  the  like.  The  individual 
plays  a  very  minor  role,  and  when  he  is  introduced,  in  most  cases  it 
is  the  king  who  is  directly  mentioned  or  indirectly  referred  to. 
Even  the  welfare  of  the  king  is  bound  up  with  the  welfare  of  the  ^ 
country  under  the  view  of  kingship  which  continues  to  hold  good  till 
the  end  of  the  Babylonian-Assyrian  control  and  according  to  which 
the  king's  welfare,  because  of  his  peculiar  relationship  to  the  gods, 
conditions  the  general  prosperity  and  happiness;^  and  this  applies/ 
also  to  signs  connected  with  a  member  of  the  royal  household.  Jt  is 
because  of  this  bearing  of  both  forms  of  divination  on  the  general 
welfare  that  they  form  integral  parts  of  the  official  cult.  Especially 
is  this  the  case  with  the  rites  of  hepatoscopy  which,  as  texts  from 
the  days  of  the  Assyrian  empire  show,  formed  part  of  a  regular 
ritual.* 

More  important,  however,  than  this  aspect  of  hepatoscopy  and 
astrology  in  Babylonia  and  Assyria  is  the  circumstance  that  both 
methods  rest  upon  a  well-defined  theory  and  are  therefore  not  to  be 
viewed  as  merely  arbitrarily  chosen  devices.  In  the  case  of  hepa- 
toscopy the  underlying  theory  may  be  summed  up  as  follows.  The 
sacrificial  animal  on  being  accepted  by  the  deity  to  whom  it  is 
oflFered  is  assimilated  to  the  deity.  The  deity  becomes  one  with  it, 
much  in  the  same  way  as  the  one  who  partakes  of  an  animal  becomes 
part  of  that  animal,  or  the  animal  part  of  him.  The  soul  of  the 
animal  is  thus  put  in  harmonious  accord  with  the  soul  of  the  god. 

*  See  J.  G.  Frazer,  "  Lectures  on  the  Early  History  of  Kingship." 
•See  Jastrow,  "Religion  Babyloniens  und  Assyriens,"  II.,  pp.   174  scq. 
and  300  seq. 

PROG.  AMER.   PHIL.  SOC.  XI.VII.   I90  PP,  PRINTED  FEBRUARY  6,  I9O9. 


660  JASTROW— HEPATOSCOPY  AND  ASTROLOGY        [Dec^ber4. 

• 

The  two  agree  as  two  watches  regulated  to  be  in  perfect  unison. 
If,  therefore,  one  can  read  the  soul  of  the  animal,  one  enters  at  the 
same  time  into  the  inner  being  of  the  god.  Now  according  to  a 
view  widespread  still  among  people  living  in  a  state  of  primitive 
culture,  the  seat  of  life  is  in  the  liver,  which  is  not  only  the  organ 
of  emotional  activity  but  of  intellectual  fimctions  as  well,  the  source 
of  all  emotions  high  and  low,  of  thought,  will  and  all  manifestations 
of  what  we  ordinarily  call  soul  life.*  From  this  point  of  view  the 
liver  is  the  seat  of  life  and  of  the  soul,  as  the  ancients  conceived 
\  vitality  and  its  inward  and  outward  phenomena. 

The  combination  of  these  two  conceptions  (i)  of  the  liver  as  the 
seat  of  the  soul  and  (2)  of  the  assimilation  of  the  soul  of  the  sacri- 
ficial animal  to  the  soul  of  the  deity  to  whom  it  is  oflfered  and  who 
accepts  it,  leads  to  the  conclusion  that  if  one  is  able  to  read  the  soul 
of  the  animal  as  revealed  in  the  condition  of  the  liver  and  of  the 
sig^s  thereon,  the  soul  including,  therefore,  the  will  and  intention  of 
.  the  deity  is  revealed.  Through  the  liver  of  the  sacrificial  animal  cme 
enters  as  it  were  into  the  workshop  of  the  gods.  The  mind  of  the 
god  is  reflected  in  the  liver  of  the  sacrificial  animal  like  an  image  in 
a  mirror — ^to  use  the  figure  introduced  by  Plato  in  an  interesting 
passage  of  the  Timaeus^®  bearing  on  divination  through  the  liver. 

As  for  the  system  of  interpretation  of  the  signs  noted  it  revolves 
largely  around  a  more  or  less  natural  association  of  ideas. /The 
chief  parts  of  the  liver  to  which  attention  was  directed  being  the 
right  and  left  lower  lobes,  the  upper  lobe  with  its  two  appendices, 
the  larger  one  known  as  the  processus  pyramidalis  and  the  smaller 

•For  further  details  regarding  this  view  of  the  liver  which  also  under- 
lies hepatoscopy  among  the  Etruscans,  Greeks  and  Romans  see  Jastrow,  "  Re- 
ligion Babyloniens  und  Assyriens/*  II.,  pp.  213  seq.  In  a  special  article 
(shortly  to  be  published)  on  "  The  Liver  as  the  Seat  of  the  Soul "  I  have  set 
forth  the  historical  development  of  the  location  of  the  soul  in  the  liver,  in 
the  heart  and  in  the  head  successively.  The  second  stage,  though  reached  by 
the  Babylonians  and  Assyrians,  never  found  expression  in  Hepatoscopy, 
whereas  among  the  Romans  from  a  certain  period  on,  the  heart  and  occa- 
sionally the  lungs  and  even  the  milt  were  also  examined.  The  third  stage 
was  reached  too  late  for  incorporation  into  the  divination  rites,  but  in  phre- 
nology as  an  extra-official  pseudo-scientific  form  of  divination  we  have  the 
outward  expression  of  the  belief  which  placed  the  soul  in  the  brain. 

**§7iC. 


,5o8.]  IN   BABYLONIA  AND   ASSYRIA.  661 

one  as  the  processus  papillaris,  the  gall-bladder,  the  cystic  duct, 
the  hepatic  duct,  the  common  bile-duct,  the  hepatic  vein  and  the 
"liver  gate"  {porta  hepatis).  A  swollen  gall-bladder  was  inter- 
preted as  pointing  to  an  enlargement  or  increase  of  power,  a  long 
cystic  duct  to  a  long  reign,  a  depression  in  the  liver  gate  to  a  de- 
crease in  power  and  so  forth.  Through  the  further  distinction  be- 
tween right  and  left,  the  former  representing  the  favorable  side,  the 
latter  the  unfavorable  side,  the  signs  in  question  referred  to  the 
king's  side  or  to  the  enemy's  side,  as  the  case  might  be.  Besides  the \ 
parts  of  the  liver,  markings  on  the  liver — ^holes,  lines,  and  depres- 1 
sions — due  largely  to  the  traces  on  the  liver  surface  of  the  subsidiary  I 
ducts  and  veins,  were  accorded  a  special  significance.  According  tc/ 
the  shape  of  these  markings,  frequently  fantastically  pictured  as 
weapons  of  the  gods,  an  interpretation,  likewise  based  on  association 
of  ideas,  was  oflfered  and  in  this  way  the  field  of  hepatoscopy  was. 
further  extended.  No  two  livers  were  ever  exactly  alike,  and  it  will 
readily  be  seen  how  in  the  course  of  time  the  collections  of  signs 
with  their  interpretation  would  grow  to  huge  proportions,  and  the 
opportunity  thus  given  for  the  imagination  and  fancy  of  the  divining 
priest — the  bdru  or  "  inspector  "^^  as  he  was  called,  to  roam  over  a 
boundless  territory.  To  the  credit  of  the  Babylonian  and  Assyrian 
priests  be  it  said  that  so  far  as  the  evidence  goes,  they  applied  the 
elaborate  and  complicated  system  devised  by  them  logically  and 
consistently.  They  did  not  hesitate  to  announce  to  the  kings  an 
unfavorable  result  of  the  examination  of  the  signs.  Grouping  all 
the  signs  noted  together,  if  the  unfavorable  signs  predominated,  a 
second  sheep  was  oflfered  and  the  liver  examined,  and  if  the  result  of 
this  diagnosis  was  also  unfavorable,  the  omens  were  taken  for  a 
third  time.  The  frequency  with  which  in  oflScial  reports  to  the 
kings  unfavorable  prognostications  are  set  forth*^  warrants  the  con- 

"  The  underlying  stem  is  the  common  term  for  "  to  see."  The  bdru 
as  "the  seer"  was  the  one  who  by  means  of  an  "inspection"  foretold  the 
future.  The  term  was  extended  also  to  the  "inspector"  of  the  heavens  or 
the  astrologer.  In  Hebrew  we  have  as  an  equivalent  ro'ih  and  in  an  article 
"Ro'ih  and  Hosih"  (Journal  of  Biblical  Literature,  Vol.  XXVIIL,  part  I) 
I  have  tried  to  show  that  the  ro'ih  like  the  bdril  was  originally  an  "  inspector  " 
of  some  object  through  which  the  future  was  divined. 

"Jastrow,  0.  c,  II.,  p.  287  seq.  for  examples. 


652  JASTROW— HEPATOSCOPY   AND   ASTROLOGY         [December 4 

.elusion  that  the  diviners  were  far  removed  from  resorting  to  decep- 
tion and  to  tricky  devices  such  as  are  reported  of  augurs  among 

I  Greeks  and  Romans.^*    Indeed  the  mere  circumstance  that  hepatos- 

I  copy  prevailed  uninterruptedly  from  the  earliest  to  the  latest  periods, 
and  that  on  all  important  occasions  it  was  resorted  to  as  the  official 
means  of  ascertaining  the  will  and  intentions  of  the  gods,  is  a  testi- 
mony to  the  conscientious  manner  in  which  the  priests  must  have 

\carried  out  their  tasks. 

In  passing  from  hepatoscopy  to  astrology — the  term  always  used 
in  the  larger  sense  above  pointed  out** — we  pass  also  from  the  do- 
main of  popular  and  to  a  large  extent  primitive  beliefs  to  a  domain 
of  speculation  that  in  comparison  justly  merits  the  designation 
scientific.  Astrology  in  Babylonia  and  Assyria  rests  on  the  identi- 
fication of  the  heavenly  bodies  with  the  gods  of  the  pantheon.  While 
in  the  case  of  the  personification  of  the  sun  and  moon  as  deities  we 
are  still  within  the  province  of  popular  and  primitive  conceptions, 
we  pass  beyond  this  province  in  the  extension  of  such  personifica- 
tion to  the  planets  and  stars.  It  lies  in  the  nature  of  animism,  which 
is  certainly  to  be  regarded  as  a  stage  in  the  development  of  religious 
beliefs,  even  if  it  is  not  admitted  to  be  the  starting-point  of  such 
development,  not  to  distinguish  sharply  between  the  manifestation 
of  a  personified  power  and  the  seat  of  that  power.  The  sun  is  at 
once  the  sun-god  and  the  seat  of  that  god;  and  the  same  applies  to 
the  moon.  Both,  accordingly,  have  their  places  in  the  heavens. 
Storms,  rains,  thunder  and  lightning  likewise  come  from  the  heavens 
and  hence  the  gods  representing  the  personification  of  these  powers 

.also  have  their  seats  in  the  heavens.  Such  conceptions  are  a 
direct  outcome  of  popular  and  primitive  methods  of  thought,  and  we 
may  perhaps  go  a  step  farther  and  assume  that  by  analogy  other 
powers  whose  manifestations  proceeded  from  a  hidden  source  were 
assigned  to  the  heavens,  but  this  step  is  far  removed  from  the  identi- 
fication of  all  the  stars  with  deities  and  still  farther  from  projecting 

"See,  e.  g.,  the  anecdotes  related  by  Polyaenus,  "  Strategicon,"  IV.,  20, 
and  Frontinus,  "  Strategematicon,"  I.,  XL,  15.  Compare  also  Hippolytus, 
Refutatio,  IV.,  40. 

"The  earliest  reference  occurs  in  the  inscriptions  of  Gudea  (c.  2500 
B.  C),  the  latest  in  the  inscriptions  of  Nabonidus,  the  last  king  of  Baby- 
lonia.    See  Jastrow,  0.  c,  II.,  p.  273  and  247  seq. 

"  See  above,  p.  647. 


,9o8.]  IN   BABYLONIA   AND  ASSYRIA.  653 

the  seats  of  all  gods  and  goddesses  on  to  the  heavens.    Again,  the  \ 
influence  of  moon  and  sun,  as  well  as  storms  with  their  accompany-  j 
ing  phenomena,  on  the. fate,  welfare  and  happiness  of  mankind  wasi 
so  apparent  as  to  force  itself  upon  the  notice  even  of  people  living! 
in  a  state  of  primitive  culture ;  and  when  we  pass  to  the  higher  stages  / 
of  nomadic,  semi-nomadic  and  agricultural  life,  the  dependence  of{ 
the  country's  prosperity  and  of  the  individual's  welfare  upon  sun,  I 
moon  and  climatic  conditions  would  be  correspondingly  increased.] 
The  observation  of  the  movements  and  aspects  of  sun  and  moon 
would  follow  as  a  natural  consequence,  and  we  may  suppose  that  at 
a   comparatively   early   stage   in   cultural   development   crude   and  V 
sporadic  attempts  might  be  made  on  the  basis  of  empirical  observa- 
tions to  select  the  favorable  moment  for  such  actions  as  the  under- 
taking of  a  journey,  for  hunting  or  war,  for  the  planting  of  seeds, 
for  the  gathering  of  the  harvest  or  even  for  the  pairing  of  domesti- 
cated animals.      The  influence  of  the  planets  and  stars,  however,  ^ 
would  be  less  obvious  and  indeed  until  a  comparatively  advanced 
stage  of  intellectual  development  would  not  be  recognized  at  all. 
Astrology  in  the  proper  sense,  therefore,  is  not  found  among  peoples 
of  primitive  culture**   who  at   the  most  are   guided  by   certain 
empirical  considerations  in  their  enterprises. 

The  projection  of  the  seats  of  all  the  gods  on  the  heavens  can  ^ 
only  have  arisen  in  people's  minds  as  the  outcome  of  theoretical 
speculation.  This,  to  be  sure,  represents  merely  the  extension  by 
analogy  of  the  primitive  conception  of  sun,  moon  and  storms,  but  an 
extension  which  for  the  very  reason  that  it  is  neither  obvious  nor 
the  result  of  actual  experience,  lies  outside  of  the  range  of  early^ 
thought.  The  views  of  Cumont*^  and  Boll'®  may,  therefore,  be  un- 
hesitatingly accepted  that  astrology  everywhere  represents  a  scientific 
view  of  the  universe — scientific  of  course  in  a  relative  sense,  and  in 
comparison  with  the  conceptions  that  underlie  hepatoscopy  or  with 
the  significance  attached  to  universal  occurrences  on  earth  or  to  the 

"See  above,  p.  647,  note  3. 

"Les  Religions  Orientales  dans  le  Paganisme  Romain"  (Paris,  1907), 
pp.  197  seq. 

""Die  Erforschung  der  antiken  Astrologie"  (Neue  Jahrhucher  fur  das 
Klassische  Alter  turn,  I.,  Abt.,  Bd.  XXI.),  p.  108  seq. 


654  JASTROW— HEPATOSCOPY   AND   ASTROLOGY  [December  4. 

mishaps  and  accidents  of  daily  life.  Indeed,  one  may  safely  go  a 
step  further  and  set  up  a  contrast  between  hepatoscopy  and  astrology 
corresponding  to  the  difference  nowadays  between  the  popular  views 

I  of  the  universe  which  are  still  so  largely  controlled  by  superstitious 
beliefs  and  crude  speculations — instance  the  hold  that  astrolog>% 
k  phrenology,  chiromancy,  clairovoyance,  dreams  and  belief  in  the 
power  of  ghosts  still  have  upon  the  masses, — and  those  held  by 
scientific  thinkers.  The  astrological  system  of  Babylonia  and  As- 
syria, which  is  the  earliest  known  to  us,  might  be  described  as 
taking  the  place  in  antiquity  that  in  modern  times  is  taken  by  the 
"  Darwinian  "  theory  of  evolution  in  so  far  as  it  is  the  product  of 
the  schools  and  not  of  popular  conceptions. 

It  may   reasonably   be   supposed   that   the   recognition   of   the 
;  regular  movements  of  the  planets  and  that  within  certain  periods 
\  they  pass  through  a  well-defined  course  as  do  the  sun  and  moon, 
'  was  the  decisive  step  which  led  to  the  departure  from  along  the 
^  lines  of  popular  conceptions.     With  the  planets  thus  placed  on  a 
par  with  sun  and  moon,  it  was  a  natural  sequence  to  regard  them 
also  as  gods,  or,  what  amounted  to  the  same  thing,  as  the  seats  of 
gods,  and  to  endow  them  with  the  power  to  control  occurrences 
/on  earth.    In  the  oldest  astrological  texts,  as  a  matter  of  fact,  we 
find  the  five  planets  already  identified  with  the  chief  gods  of  the 
/  Babylonian-Assyrian  pantheon,  Jupiter  being  known  as  Marduk, 
I  Venus  as  Ishtar,  Saturn  as  Ninib,  Mercury  as  Nebo  and  Mars 
(  as  Nergal.^*    This  identification  in  itself  is  sufficient  to  establish  the 
advanced  character  of  the  entire  astrological  lore,  for  the  gods  in 
question,  according  to  the  popular  conceptions  and  even  in  the 
ofllicial  cult,  stand  in  no  connection  with  the  stars.    Marduk,  Ninib 
and  Nergal  are  originally  solar  deities.    Nebo  appears  originally  to 
have  been  a  water  deity ,*^  while  Ishtar  is  the  earth  goddess,  the  sym- 
bol and  personification  of  fertility  in  general.  In  thus  being  identified 

4- 

with  the  planets,  the  original  character  of  the  deities  in  question  is  , 
entirely  lost  sight  of.     The  i'dentification,  therefore,  represents  a 
break  with  popular  conceptions  and  with  the  traditions  that  had 

*•  Kugler,  "  Stemkunde  und  Sterndienst  in  Babel/'  I.,  p.  8  with  the  cor- 
rections on  pp.  221  and  286. 
•Jastrow,  o.  c,  I.,  p.  118. 


i9o8.] 


IN   BABYLONIA   AND  ASSYRIA.  655 


gathered  around  these  deities.    In  view  of  this,  it  is  clear  that  in 
dealing  with  Babylonian-Assyrian  astrology  we  have  to  do  with  the  \ 
theories  of  the  theologians  or  priests  as  the  representatives  of  ad-  - 
vanced  and  abstract  thought,  and  not  with  popular  notions.    More-  \ 
over,  the  choice  of  the  deities  in  question  and  the  order  in  which  they 
are  enumerated  when  introduced  as  equivalents  of  the  planets  are 
further  indications  of  the  speculative  spirit  which  led  to  their  iden- 
tification with  the  planets,  and  also  of  the  time  when  this  identifica-  , 
tion  took  its  rise.    Jupiter-Marduk  is  always  mentioned  first  and  this 
precedence  is  evidently  a  reflection  of  the  period  when  Marduk  was 
regarded  as  the  head  of  the  pantheon,  *.  e,,  the  period  after  Ham- 
murabi with  whom  as  the  unifier  of  the  Euphratean  states,  the  city 
of  Babylon  as  the  capital  of  the  empire  assumes  the  definite  position 
it  continued  to  hold  till  the  destruction  of  the  neo-Babylonian  king- 
dom by  Cyrus  in  539  B.  C.    The  pantheon  as  constituted  during  of 
after  the  days  of  Hammurabi  assigns  to  Marduk  as  the  patron  deity 
of  Babylon  the  first  position.    Marduk  takes  the  place  held  by  Enlil . 
of  Nippur  and  subsequently,  as  would  appear,  by  Ninib.*^   The  other; 
great  gods  of  the  pantheon,  as  found  in  the  Hammurabi  period,  are 
precisely  the  ones  identified  with  the  remaining  four  planets,  Ishtar, 
Ninib,  Nebo  and  Nergal  together  with  Sin  the  moon-god,  Shamash 
the  stm-god  and  Adad-Ramman  the  storm-god.     The  basis  upon 
which  Babylonian-Assyrian  astrology  rests  thus  assumes  the  defi- 
nite formation  of  a  pantheon  and  moreover  the  particular  form  of 
the  pantheon  that  marks  the  Hammurabi  period,  i,  e.,  after  2000  B.  C. 
This  does  not  necessarily  mean  that  astrology  dates  in  Babylonia 
from  this  period,  for  it  is  possible  that  there  was  an  earlier  series  of 
identification  of  gods  with  planets,  but  that  the  astrological  texts 
known  to  us  do  not  revert  to  originals  older  than  the  days  of  Ham- 
murabi.   There  are  indeed  references  in  the  inscriptions  of  Gudea> 
which  would  point  to  the  practice  of  interpreting  the  signs  of  the  ' 
heavens  at  this  earlier  period**  and  it  may  well  be  therefore  that  the 
priests  long  before  Hammurabi  had  started  on  the  course  of  specu- 
lation which  culminated  in  placing  the  seats  of  all  the  gods  in  the 
starry  firmament.     But  whatever  the  age  of  Babylonian-Assyrian 

**  See  Jastrow,  o.  c,  I.,  p.  452  seq. 
"See  Jastrow,  o.  c,  II.,  p.  423. 


656  JASTROW— HEPATOSCOPY  AND  ASTROLOGY        [December  4, 

/  astrology  may  be,  it  must  have  involved  the  dissociation  of  the 
gods  identified  with  planets  and  stars  from  their  original  character 
as  solar,  agricultural,  water  or  chthonic  deities,  and  it  is  also  reason- 
I  able  to  assume  that  it  is  subsequent  to  the  period  when,  by  a  process 
of  selection,  certain  deities,  though  originally  local  in  character,  were 
differentiated  from  the  many  other  local  gods  and  became  members 
of  a  definitely  constituted  pantheon  consisting  of  a  limited  number 
of  great  gods  and  of  a  larger  number  of  minor  deities. 

Before  passing  on  to  another  phase  of  the  subject,  it  may  be 
proper  to  point  out  the  more  specific  factors  involved  in  the  identi- 
fication of  the  planets  with  certain  gods — ^all  confirmatory  of  the 
general  thesis  that  astrology  represents  a  system  devised  in  the 
schools,  and  that  its  very  artificial  character  is  indicative  of  its 
being  a  "  scientific  "  and  not  a  "  popular  "  product.     Marduk  was_ 
/identified  with  Jupiter  by  the  natural  association  which  led  to  assign- 
/  ing  the  head  of  the  pantheon  to  the  most  striking  of  the  planets 
\known  to  the  ancients.^    In  the  case  of_Venus  it  was  probably  her 
/double  character  as  morning  and  evening  star  that  suggested  the 
/  identification  with  Ishtar,  who  as  the  goddess  of  fertility  likewise 
\  presents  two  aspects  in  the  two  divisions  of  the  year — the  producer 
of  life  and  vegetation  in  the  spring  and  summer,  and  the  one  who 
withdraws  her  favors  in  the  fall  and  winter.^*    The  dark-red  color 
of  Mars  appears  to  have  been  the  factor  which  prompted  the  identi- 
fication with  Nergal,  the  god  of  the  burning  summer  solstice,  of 
pestilence  and  death.     Nebo  becoming  in  the  pantheon  of  Ham- 
•  murabi  the  son  of  Marduk,?**  a  natural  association  of  ideas  would 
(lead  to  assigning  him  to  the  smallest  of  the  planets.    There  would 

"See  Kugler,  o.  c,  p.  14,  note  i. 

"This  double  character  of  Ishtar  underlies  the  famous  myth  commonly 
known  as  Ishtar's  descent  into  the  lower  regions.  See  Jensen,  "Keilin- 
schriftliche  Bibliothek,"  VI.,  i,  pp.  80-91.  The  destructive  character  of  Ishtar 
appears  also  in  the  myth  of  the  slaying  of  Tammuz  and  in  the  other  capacity 
of  Ishtar  as  a  goddess  of  war.    See  Jastrow,  o.  c,  I.,  pp.  82  seq. 

"See  Jastrow,  0.  c,  I.,  p.  120.  As  a  concession  to  the  predominance  of 
the  Nebo  cult  in  the  days  of  the  neo-Babylonian  dynasty,  we  find  in  the 
«stronomical  texts  of  the  latest  period  (after  400  B.  C.)  a  change  in  the 
order  of  the  planets,  Nebo-Mercury  assuming  the  third  place,  ».  e.,  after 
Marduk  and  Ishtar,  instead  of  Ninib-Satum  who  is  assigned  to  the  fourth 
place.    See  Kugler,  0.  c,  p.  13. 


I908.) 


IN   BABYLONIA   AND  ASSYRIA.  657 


thus  remain  for  Ninib  the  planet  Saturn  whose  large  size  would 
have  been  regarded  as  appropriate  for  a  solar  deity  once  occupying 
the  position  that  afterwards  was  assumed  by  Marduk. 

The  planets  thus  representing  the  great  gods  of  the  pantheonX 
the  prominent  fixed  stars  were  associated  with  the  minor  deities  and  / 
while  in  the  case  of  many  of  the  stars  occurring  in  the  purely  as- 
tronomical texts  which  belong  to  the  later  and  latest  periods  of 
Babylonian  culture,**  no  definite  association  with  'specific  deities  was 
worked  out,  yet  it  is  to  be  borne  in  mind  that  all  the  stars  were 
regarded  as  gods  in  a  logical  and  consistent  extension  of  the  prin- 
ciple which  gave  rise  to  astrology  as  a  system  of  divination.  It  is 
one  of  the  many  merits  of  Hugo  Winckler*'  to  have  demonstrated 
as  one  of  the  tenets  of  the  Babylonian-Assyrian  conception  of  the 
universe  a  perfect  correspondence  between  occurrences  on  earth  and 
phenomena  in  heaven.*^*  Earth  and  heaven  stand  related  to  each 
other  as  a  reflection  in  a  mirror  to  the  original  which  is  reflected. 
Since  all  that  happens  is  due  to  the  gods,  it  follows  from  the  specu- 
lative view  which  places  the  gods  in  the  heavens  that  occurrences  on 
earth  are  prepared  in  the  heavens.  What  one  sees  in  the  heavens  isj 
therefore  the  activity  of  the  gods  preparing  the  events  on  earth.] 
The  constantly  changing  aspect  of  the  starry  universe  thus  finds  a 
natural  explanation.  The  movements  of  sun,  moon  and  planets  as 
well  as  the  ever-varying  aspects  of  clouds  and  all  other  phenomena 
of  a  striking  character  were  the  external  symptoms  of  the  never- 

"  See  Kugler,  "  Sternkunde,"  p.  2  and  elsewhere  whose  views  have  been 
accepted  by  Boll,  Eduard  Meyer,  Schmidt  and  many  others.  See  Jastrow, 
11.,  p.  432,  note  I,  where  I  have  set  forth  my  own  position  on  the  important 
question  as  to  the  age  of  astronomy  in  Babylonia  and  Assyria  with  an  en- 
deavor to  do  justice  to  both  sides  of  the  burning  problem. 

""'Himmels  und  Weltenbild  der  Babylonier"  (Leipzig,  1893,  2'*  Auflage) 
and  numerous  other  monographs  of  this  scholar.  See  Jastrow,  o.  c,  II.,  p. 
418,  note  2. 

"'  The  same  view  prevails  among  the  Indians  of  Mexico  according  to 
Preuss  "Die  Astral  religion  in  Mexico  in  vorspanischer  Zeit  und  in  der 
Gegenwart"  (Transactions  of  the  3d  International  Congress  for  the  History 
of  Religions  I.,  p.  36  seq.).  It  is  to  be  noted  that  also  among  the  Mexican 
Indians  the  astral  cult  included  the  worship  of  storm  and  rain  deities  (/.  c. 
p.  38  seq.).  Preuss  is  mistaken,  however,  in  regarding  this  astral  religion  as 
"primitive."  On  the  contrary,  it  betrays  all  the  earmarks  of  a  cult  devised 
by  priests  on  the  basis  of  elaborate  cosmical  speculations. 


\ 


658  JASTROW— HEPATOSCOPY   AND  ASTROLOGY         [December  4. 

ceasing  divine  activity.  The  theory  of  the  correspondence  between 
<2/  heaven  and  earth  was  carried  by  the  theologians  of  the  Euphrates 
Valley  to  its  logical  consequences.  Myths  and  legends  were  so 
shaped  under  the  influence  of  the  theory  as  to  admit  of  a  double 
interpretation,  the  one  having  reference  to  the  movements  and  as- 
pects of  the  heavenly  bodies,  the  other  to  occurrences  whose  scene 
is  placed  on  earth.  A  series  of  acts  of  creation  on  earth  is  counter- 
balanced by  a  corresponding  series  in  the  heavens.**  The  heavens 
were  divided  off  into  districts  with  mountains,  rivers  and  cities 
corresponding  to  those  on  earth.  The  famous  Gilgamesh  Epic — a 
composite  tale  with  almost  equal  proportions  of  nature  myth,  legen- 
dary lore  and  dimned  historical  traditions — admits  likewise  of  a 
double  interpretation,  the  scenes  applying  equally  to  the  movements 
of  heavenly  bodies  and  to  events  on  this  globe  ;***  and  the  same  holds 
good  for  such  tales  as  the  story  of  Etana  and  the  Adapa  myth  which, 
besides  betraying  the  work  of  theological  schools  in  making  the 
tales  the  medium  of  conveying  doctrinal  teaching,*"  are  so  con- 
structed as  to  conform  with  the  fundamental  principle  of  a  corre- 
spondence  between  heaven  and  earth. 

Corresponding,  therefore,  to  the  theory  underlying  Babylonian- 
Assyrian  hepatoscopy  as  above  set  forth,  we  have  in  the  case  of 
astrology  likewise  a  theory  which  lifts  the  endeavor  to  divine  the 
future  through  the  observation  of  the  planets  and  stars  beyond  mere 
/caprice  and  arbitrary  guesswork.    Granted  the  underlying  assump- 
*  tion  that  there  is  a  perfect  correspondence  between  heaven  and  earth, 
it  follows  that  if  one  can  grasp  the  meaning  of  the  aspects  and 
movements  of  the  heavenly  bodies  one  can  recognize  clearly  what 
the  gods  are  doing,  and  hence  what  the  future  is  to  be,  which, 
since  it  is  in*the  hands  of  the  gods,  is  merely  the  outcome  of  their 
1     activity  as  revealed  in  the  heavens.     Astrology  is,  therefore,  like 

1       V 

\   hepatoscopy  a  means  of  entering  into  the  workshop  of  the  divine 

"  See  Zimmem,  "  Biblische  und  Babylonische  Urgeschichte  "  (3'*  Auflage, 
Leipzig,  1903). 

"•See  Kugler,  die  Stemenfahrt  des  Gilgamesch  (Stimmen  aus  Maria- 
Laach,  1904.    Heft.  4). 

•See  Jastrow,  "Religion  of  Babylonia  and  Assyria"  (Boston,  1898), 
PP-  5i9-555»  and  in  greater  detail  in  the  writer's  next  volume.  "Temples, 
Myths  and  Cults  of  Babylonia  and  Assyria." 


,5o8J  IN   BABYLONIA   AND  ASSYRIA.  659 

will  and  intention.  Through  the  planets  and  stars  or  rather  in  the 
planets  and  stars  one  sees  the  gods  at  work  and  if  one  knows  what 
they  are  contriving,  one  knows  what  occurrences  will  take  place  on 
earth.  Again,  as  in  the  case  of  hepatoscopy,  past  experience  and 
association  of  ideas  are  the  two  main  factors  involved  in  the  system 
of  interpretation  gradually  devised  by  the  Babylonian-Assyrian 
baru  priests  or  "  inspectors "  in  their  capacity  as  astrologers  or 
**  inspectors  "  of  the  heavens.  A  favorable  event  or  a  favorable  out- 
come of  a  crisis  following  upon  certain  aspects  of  the  heavenly 
bodies  would  be  made  the  basis  of  a  favorable  prognostication  on 
another  occasion  when  the  same  conditions  presented  themselves; 
and  the  prognostication  would  be  made  without  reference  to  the  par- 
ticular event  following  upon  the  original  observation.  It  was  not^ 
the  event  that  was  of  importance  but  merely  the  circumstance 
whether  it  was  favorable  or  unfavorable.  On  the  basis  of  this  ex- 
perience phenomena  were  entered  as  pointing  to  favorable  or  un- 
favorable occurrences,  and  these  entries  served  as  a  guide  to  the 
priests  in  the  task  imposed  upon  them  of  divining  the  future.  But 
while  the  principle  of  post  hoc  propter  hoc  entered  largely  into  the 
formation  of  collections  of  astrological  omens — ^as  it  did  in  the  col- 
lections of  hepatoscopical  omens'® — ^the  natural  or  artificial  associa- 
tion of  ideas  was  even  a  more  prominent  factor.  Normal  conditions' 
as  a  rule  were  interpreted  as  favorable.  Thus,  if  the  moon  and  sun 
appeared  in  conjunction  at  the  proper  time,  a  favorable  prognosti- 
cation was  indicated.  If  the  conjunction  took  place  at  a  time  earlier 
or  later  than  the  expected  moment  it  forboded  disaster  of  some 
kind.  Again,  by  a  perfectly  logical  association,  in  case  the  new 
moon  was  seen  on  the  first  day  of  the  month,  i,  e,,  was  not  obscured 
by  clouds,  the  omen  was  of  a  favorable  character ;  if,  however, 
clouds  obscured  it  so  that  the  new  moon  was  not  visible,  diflSculties 
of  some  kind  might  be  expected.  Days  were  entered  as  favorable 
or  unfavorable  according  to  these  and  numerous  other  indications  ] 
and  though  in  the  case  of  a  specific  inquiry  of  the  gods  recourse 
was  had  to  hepatoscopy  in  order  to  ascertain  what  a  deity  had  in 
mind  with  regard  to  the  particular  situation  in  question,  the  signs 

"  See  Jastrow,  "  Religion  Babyloniens  und  Assyriens,"  II.,  p.  251  seq.,  for 
examples. 


660  JASTROW— HEPATOSCOPY    AND   ASTROLOGY         [December  4, 

involuntarily  forced  on  one's  notice  by  conditions  prevailing  in  the 
heavens  were  not  and  could  not  be  neglected.  A  few  examples  from 
astrological  texts  will  suffice  for  our  purposes.  Thus  we  read  in  an 
official  report  of  the  Assyrian  period  :'^ 

"If  the  moon  is  seen  on  the  first  day,  good  faith  and  stable  conditions 
in  the  land.  If  the  first  day  is  abnormally  long,  the  king  will  have  a  long 
reign." 

The  prognostication  is  clearly  based  on  a  natural  association  of 
ideas.  From  the  fact  that  the  new  moon  is  visible  on  the  day  set 
for  it,  the  conclusion  is  drawn  that  as  the  moon  kept  good  faith, 
as  it  were,  so  the  king  may  expect  those  entrusted  with  any  mission 
to  be  faithful  and  that  his  subjects  in  general  will  be  loyal.  By 
a  still  clearer  association  long  days  point  to  a  long  reign. 

Another  report  states  :** 

"If  the  moon  is  seen  out  of  the  expected  time,  prices  will  be  low."  The 
moon  was  seen  with  the  sun  on  the  twelfth  day.  If  moon  and  sun  are  seen 
together  at  an  abnormal  time,  a  strong  enemy  will  oppress  the  land,  but  the 
king  of  Babylonia  will  accomplish  the  overthrow  of  his  enemy." 

The  normal  period  when  moon  and  sun  should  be  seen  at  the 
same  time  in  the  heavens  is  on  the  fifteenth  day — ^the  moment  of 
opposition.  The  fcoru-priest  reports,  however,  that  the  appearance 
of  moon  and  sun  took  place  already  on  the  twelfth  day— earlier, 
therefore,  than  was  expected.  The  abnormal  condition  points  to 
some  misfortune  and  two  omens  that  are  to  be  regarded  as  extracts 
from  actual  collections  are  introduced,  the  one  referring  to  economic 
conditions,  the  other  to  political  affairs,  and  though  both  are  un- 
favorable, yet  in  the  second  instance  it  is  added  that  ultimately  the 
enemy  will  be  overthrown.  In  the  case  of  such  specific  prognosti- 
cations we  are  perhaps  justified  in  concluding  that  they  rest  on  past 
experience.  In  other  words,  on  some  occasion  when  sun  and  moon 
were  seen  together  in  the  heavens  earlier  than  the  fourteenth  or 
fifteenth  day  of  the  month,  prices  went  down  or  an  enemy  entered 
the  land  but  was  eventually  vanquished.     The  occurrences  were 

"Thompson,  "Reports  of  the  Magicians  and  Astrologers  of  Nineveh 
and  Babylon"  (London,  1900),  Vol.  I.,  No.  i. 

"0.  c,  No.  119. 

"Low  prices  were  regarded  as  an  unfavorable  condition  in  Babylonia 
and  Assyria. 


,9o8.]  IN   BABYLONIA   AND   ASSYRIA.  661 

accordingly  entered  as  unfavorable  in  the  collections,  and  when  the 
same  conditions  again  took  place,  the  fact  was  reported  to  the  king 
who  would  thus  be  warned  either  against  undertaking  an  expedition 
or  at  least  would  be  prepared  for  some  disaster  or  discomfiture. 

To  even  partially  enumerate  the  phenomena  noted  in  the  astro- 
logical collections  would  carry  us  too  far,  and  it  will  easily  be  seen 
how  in  the  course  of  time  the  collections  would  grow  to  huge  pro- 
portions.'* Halos  around  the  moon  or  sun,  moon  and  sun  eclipses, 
thunder  in  certain  months  or  on  certain  days,  one  planet  or  the 
other  standing  within  the  halo  around  the  moon,  the  appearance  of 
Venus  or  some  other  planet  at  the  heliacal  rising  or  at  some  other 
point  in  its  course,  the  appearance  of  the  moon's  horns  or  crescent, 
the  position  or  appearance  of  a  certain  planet  or  of  a  certain  star 
are  among  the  phenomena  entered  and  here  the  prognostications  vary 
according  to  the  season  of  the  year,  according  to  the  month  or  day 
of  the  month.'** 

Without  losing  sight  of  the  purely  artificial  character  of  the^ 
system  of  interpretation  devised  by  the  Babylonian  theologians,  one 
should  not  withhold  one's  meed  of  praise  for  the  consistency  with  | 
which  the  elaborate  system  was  carried  out  for  a  long  stretch  of  \ 
centuries,  as  well  as  for  the  patience  displayed  in  the  compilation  of  j 
the  extensive  collections  of  omens  of  which  only  portions  have  come  i 
down  to  us.     Moreover,  the  Babylonian- Assyrian  astrology  shows ) 
that  even  a  superstition  can  harbor  an  exalted  idea,  for  the  result  of/ 
the  continuous  observation  of  the  movements  and  aspects  of  planets 
and  stars  must  have  been  to  impress  at  all  events  the  priests  with  the 
realization  of  the  reign  of  law  in  the  universe;  and  it  is,  assuredly, 
a  decided  gain  to  realize  that  even  the  activity  of  the  gods  is  under 
the  sway  of  a  fixed  order.     In  striking  contrast  to  hepatoscopy\ 
which  rests  upon  the  arbitrary  nature  of  the  gods  and  merely  aims  ' 
to  fathom  their  caprice,  astrology  starts  with  the  recognition  of  the  | 

"The  best  known  of  such  astrological  collections  in  Ashurbanapal's 
famous  library  is  ^  series  known  from  the  opening  words  as  "  When  Anu  and 
Enlil"  and  comprising  more  than  seventy  tablets.  See  Jastrow,  o.  c,  II., 
p.  424,  notes  3  and  4,  and  copious  examples  beginning  p.  458. 

"In  their  ambition  to  make  the  collections  as  complete  as  possible,  the 
baru'pritsts  even  enter  phenomena  that  never  occurred,  and  some  that  never 
could  have  occurred. 


662  JASTROW— HEPATOSCOPY  AND  ASTROLOGY         [December  4, 

fact  that  the  gods  as  represented  by  the  planets  and  stars  act  in 
concert.  The  phenomena  of  the  heavens  suggest  united  action  in 
place  of  individual  caprice,  and  the  general  regularity  of  the  move- 
)  ments  of  heavenly  bodies  must  soon  have  suggested  to  the  priests 
j  the  view  that  divine  government  of  the  universe  rests  at  least  to  a 
1  large  extent  upon  law  and  order.  We  may  properly  assume  that  this 
\^  aspect  of  astrology  by  which,  through  constant  observation,  the 
permanent  impression  of  awe  and  reverence  for  the  grandeur  of 
heavenly  phenomena  was  deepened,  was  an  important  factor  in 
maintaining  the  faith  in  the  stars  as  manifestations  of  the  divine 
will  and  of  the  intentions  of  the  gods  towards  mankind.  The 
Babylonian  fcdriJ-priest  could  reecho  the  sentiment  of  the  Psalmist 
(19,  1-2)  who,  carried  away  by  the  sight  that  greeted  him  in  the 
heavens,  exclaimed,  **  The  heavens  declare  the  glory  of  God  and 
the  firmanent  sheweth  his  handywork.  Day  unto  day  uttereth 
speech  and  night  unto  night  sheweth  knowledge."  To  the  baru- 
priest  the  heavens  spoke  by  day  and  night,  and  it  was  his  privilege 
to  interpret  to  others  the  knowledge  revealed  to  him. 
'^  Attention  has  already  been  directed**  to  the  fact  that  in  the  case 
of  both  hepatoscopy  and  astrology  the  interpretations  of  the  omens 
have  reference  exclusively  to  the  public  welfare,  to  the  condition 
of  the  crops,  to  pestilence,  to  war  or  victory  and  that  the  introduction 
of  the  king  likewise  falls  within  this  category.  More  than  this,  the 
interpretations  in  both  systems  are  substantially  the  same,  so  that  a 
dependence  of  one  system  upon  the  other  becomes  at  least  a  probable 
hypothesis-  A  detailed  study  of  the  two  systems  leads  indeed  to  a 
confirmation  of  this  thesis  and  since  hepatoscopy,  as  has  been  shown, 
is  an  outcome  of  popular  conceptions  and  exists  in  full  force  in  the 
arliest  period  of  Babylonian  history,  it  is  reasonable  to  suppose  that 
t  was  the  first  to  be  developed  and  that  the  astrological  system  repre- 
ents  an  adaptation  of  the  principles  underlying  the  interpretation  of 
ig^s  on  the  liver  to  signs  noted  in  the  heavenly  bodies.  The  "  scien- 
tific "  view  of  the  universe  that  is  closely  bound  up  in*  the  astrological 
system  represents,  as  is  obvious,  a  later  stage  in  cultural  development 
^than  the  "popular"  conception  upon  which  hepatoscopy  rests.     In 

"See  above,  p.  649. 


,908]  IN  BABYLONIA  AND   ASSYRIA.  663 

the  name  g^ven  to  the  planets  m  Babylonia  we  have,  I  venture  to 
think,  a  direct  proof  of  this  dependence  of  astrology  upon  hepa- 
toscopy.  It  has  always  been  a  puzzle  to  scholars  that  the  common 
designation  for  planet  should  have  been  a  compound  ideograph,*^ 
the  two  elements  of  which  signify  "  sheep  "  and  "  dead."  Attempts 
to  furnish  a  satisfactory  explanation  have  failed  and  the  interpreta- 
tion offered  by  Babylonian  scribes  as  "  causing  the  death  of  cattle,''** 
while  confirming  the  division  of  the  sign  into  the  two  elements  in 
question,  is  purely  fanciful  and  is  of  value  chiefly  as  showing  that  the 
real  origin  of  the  designation  had  already  in  ancient  times  become 
obscured.  Through  a  syllabary  (II.  Rawlinson,  PI.  6,  4  c-d)  we 
learn  that  the  compound  sign  (Lu-Bat)  is  to  be  read  bi-tb-bu  and 
the  context  in  which  the  word  occurs'"  is  sufficient  to  show  that  it 
is  one  of  the  names  for  "  sheep."  This,  moreover,  is  confirmed 
by  the  fact  that  the  first  element,  Lu,  with  or  without  the  addition 
of  the  sign  for  "  male  "  designates  the  **  sheep."  Now,  the  second 
element  (Bat)  has  also  the  force  of  tertu,  "omen,"*®  the  explanation 

''Lu-Bat.  For  a  full  discussion  see  a  special  article  by  the  writer 
"  The  Sign  and  Name  for  Planet  in  Babylonian  "  in  the  Proceedings  of  the 
American  Philosophical  Society,  Vol.  XLVII.,  pp.  141-155.  It  is  also  to  be 
noted  that  while  all  the  planets  are  designated  as  Lu-Bat  or  bibbu,  there  are 
two,  Mercury  and  Saturn,  to  whom  the  designation  is  specially  applied.  On 
the  reason  for  this  as  well  as  for  the  explanation  of  the  Babylonian  names 
for  Mercury  (Lu-Bat  Gu-Ud)  and  Saturn  (Lu-Bat  Sag-Us)  see  the  article 
just  referred  to,  in  which  on  p.  142  a  reference  should  have  been  added  to 
Zimmem,  "  Keilinschriften  u.  das  alte  Testament,"  p.  622,  seq. 

**mus-mit  bu-lim  (V.  Rawlinson,  PI.  46,  Nr.  i  (rev.),  41),  in  which 
equation  Lu  is  entered  as  the  equivalent  of  bulu  "cattle"  and  Bat  as  IIL, 
I  of  mdtu  "  cause  the  death  "  or  "  kill."  The  artifical  character  of  the  expla- 
nation is  revealed  by  the  unwarranted  extension  of  Lu  in  the  general  sense 
of  "cattle,"  nor  can  Bat  without  some  further  qualifying  prefix  mean  "cause 
to  die"  but  merely  "to  die"  or  "to  be  dead."  Lu-Bat  could  have  the 
force  of  "sheep  that  is  dead"  or  "sheep  that  is  killed,"  but  never  "sheep 
(or  'cattle')  that  kill." 

"It  is  followed  by  a-tu-du  "goat"  and  sap-pa-ru  "mountain  goat." 
Note  also  that  1.  i,  a-b  Lu  =  kir-ru—Si  common  term  for  "  lamb."  Dr.  Ru-  . 
dolf  Eisler  finds  in  the  double  sense  of  the  Semitic  stem  Omar  "  word "  and 
"sheep"  a  further  support  for  the  thesis  here  set  forth  ("Origin  of  the 
Eucharist,"  p.  10— an  address  before  the  Third  International  Congress  for 
the  History  of  Religions  at  Oxford,  Sept.  18,  1908.) 

*•  See  IL,  Rawlinson,  PI.  27,  No.  2,  46  obv.  c-d.     Ur-Bat  =  ter-tum  sa 
ha-se-e,  i.  e.,  "omen  of  the  liver,"  the  first  element  (Ur)  being  the  common 


r 


664  JASTROW— HEPATOSCOPY   AND  ASTROLOGY         [December  4. 

for  which  is  to  be  sought  in  the  circumstance  that  through  the  sacri- 
ficial animal,  killed  for  the  purpose,  an  omen  was  secured.  The 
combination  Lu-Bat,  "  dead  sheep,"  is  therefore  intended  to  convey 
the  notion  of  a  "  sacrificial  sheep,"  offered  to  the  deity  as  a  means  of 
securing  an  "  omen."  So  prominent  is  the  part  played  by  hepa- 
toscopy  in  the  Babylonian-Assyrian  religion  as  shown  not  merely 
by  the  extensive  omen  texts,  dealing  specifically  with  divination 
through  the  liver,*^  but  by  the  frequent  allusions  to  the  rite  in  his- 
torical inscriptions  that  one  is  tempted  to  set  up  the  thesis  that  the 
original  purpose  of  sacrifice  among  the  inhabitants  of  the  Euphrates 
Valley  was  to  ascertain  through  the  sacrificial  animal  what  the  future 
had  in  store  or  what  the  gods  had  in  mind, — this  purpose  taking 
precedence  of  other  views  of  sacrifice  such  as  tribute  or  alliance  with 
the  deity .^^  However  this  may  be,  the  animal,  so  far  as  the  evidence 
goes,  invariably  chosen  for  purposes  of  divination  was  the  "  sheep,"** 
and  there  is  one  instance**  in  which  the  combination  Lu  and  Bat 
occurs  in  a  "  liver "  divination  text  to  designate  the  "  sacrificial 
sheep  "  the  liver  of  which  is  to  be  examined  as  a  means  of  divination. 
It  is  with  this  use  of  the  term  that  I  propose  to  connect  the  designa- 
tion Lu-Bat  for  "planet."  The  sheep  being  the  common  animal 
of  divination,  the  term  acquired  the  general  force  of  an  "  omen  " 
precisely  as  in  Latin  we  have  auspicium,  originally  an  augury 
through  "  bird  observation,"  i.  e,,  the  noting  of  the  flight  of  birds, 
becoming  the  generic  term  for  any  kind  of  an  augury,  because  of  the 
prominence  of  "  bird  observation  "  as  a  means  of  divination.    Still 

ideograph  for  "liver"  (see  Jastrow,  "Signs  and  Names  for  the  Liver  in  Baby- 
lonian," in  Zeits.  fur  Assyr.,  XX.,  p.  105,  seq.  and  p.  127)  and  the  combina- 
tion thus  having  the  force  of  "liver  omen."  The  association  leading  from 
"dead"  to  "omen"  thus  becomes  intelligible,  since  the  "dead"  or  "sacri- 
ficed "  animal  is  the  medium  for  procuring  an  omen. 

**Over  1,000  of  the  circa  30,000  fragments  of  the  royal  Library  of 
Ninevah  are  "liver"  divination  texts.  See  Jastrow,  "Religion  Babyloniens 
und  Assyriens,"  II.,  p.  211,  note  i,  and  p.  222,  note  2. 

**  See  Jastrow,  0.  c,  II.,  p.  217. 

**So,  e.  g.,  in  the  case  of  the  official  reports  to  Assyrian  Kings,  in  the 
prayers  connected  with  the  divination  rite  as  well  as  in  the  omen  collections. 
See  Jastrow,  o.  c,  II.,  pp.  281,  289,  301,  307,  308,  etc ;  "  Cun.  Ttexts,"  XX.,  PI. 
I,  i;  Boissier,  "Documents  assyriens  relatifs  aux  Presages,"  p.  97,  11. 

**Boissier,  /.  c,  p.  212,  27.    Lu(Nita)  Bat  (u)  :=immeru  mitu. 


x9o8.] 


IN  BABYLONIA  AND  ASSYRIA.  666 


more  striking  is  the  analogy  offered  by  the  usage  in  Greek  where 
the    word    for    bird,    6pvi^  or  ^io>vo$,    has    acquired    the    force    of 
"  omen."*'    The  planets,  accordingly,  wete  called  "  sheep  "  because 
the  purpose  for  which  they  were  observed  was  to  serve  as  "  omens," 
and  this  view  is  confirmed  by  a  statement  of  Diodorus  (Bibl.  Hist. 
II.,  30)  that  the  Babylonians  (or  "  Chaldeans  "  as  he  calls  them) 
called  the  planets    ijpftcveis,  "  interpreters,"  because  "  they  reveal  (or 
"  interpret)  the  intention  of  the  gods  to  men."    The  term  used  by 
Diodorus  accurately  reproduces  the  force  of  Lu-Bat  in  the  sense  of 
an  "  omen  "  or  "  interpretation  "  of  the  will  and  purpose  of  the  gods. 
If  this  explanation  be  admitted,  we  would  thus  have  a  direct  evidence 
of  the  dependence  of  astrology  upon  hepatoscopy,  in  accord  with 
the  reasonable  assumption  on  a  priori  grounds  of  the  rise  of  astrology 
subsequent  to  hepatoscopy.    The  justification  for  thus  assuming  a\ 
bond  uniting  astrology  and  hepatoscopy  is  furnished  by  the  evidence 
for  an  analogous  condition  among  the  Etruscans  whose  method  of 
hepatoscopy  has  many  points  in   common  with   the   Babylonian-j 
Assyrian  rite.**     On  the  famous  bronze  model  of  a  liver  found  near] 
Piacenza*^  and  which,  dating  from  about  the  third  century  B.  C, 
was  used  as  an  object  lesson  for  instruction  in  hepatoscopy,  precisely 
as  the  clay  model  of  a  liver  dating  from  the  Hammurabi  period  was 
used  in  a  Babylonian  temple  school,*®  we  find  the  edge  of  the  liver 
divided  into  sixteen  regions  with  the  names  of  the  deities  inhabiting 
them,  corresponding  to  divisions  of  the  heavens  in  which  the  gods- 
have  their  seats,  while  on  the  reverse  side  there  is  a  line  dividing 

•See  the  passage  in  the  Birds  of  Aristophanes  11.  719-22  to  which  my 
colleague  Prof.  Lamberton  directed  my  attention  and  Xenophon,  Anabasis, 
III.,  2,  9,  which  Dr.  R.  G.  Kent,  of  the  University  of  Pennsylvania,  kindly 
pointed  out  to  me. 

**See  Thulin,  "  Die  Etruskische  Disciplin,"  I.  (Goteborg,  1905),  p.  xii,  seq. 

*  It  is  sufficient  for  our  purposes  to  refer  to  two  recent  treatises  on 
this  remarkable  object  (a)  Thulin,  "die  Gotter  des  Martianus  Capella  und  der 
Bronzeleber  von  Piacenza"  (Giessen,  1906),  and  Korte,  "die  Bronzeleber  von 
Piacenza,"  in  Mitt,  d,  Kais.  Deutsch.  Arch.  Instituts  (Romische  Abteilung), 

XX.,  pp.  349-379. 

*•  Published  in  "Cun.  Texts,"  VI.,  Pis.  i  and  2  (with  photograph). 
See  Boissier's  first  attempt  at  an  interpretation,  "  Note  sur  un  Monument 
babylonien  se  rapportant  d  TExtispicine  *'  (Geneve,  1899).  I  hope  ere  long 
to  publish  the  results  of  my  study  of  the  inscription  on  this  object. 

*•  See  Korte,  /.  c,  p.  356. 

PROC.  AMER.  PHIL.  SOC.  XLVII.    I90  QQ,  PRINTED  FEBRUARY  8,  I909. 


666  JASTROW— HEPATOSCOPY  AND  ASTROLOGY        [December  4. 

the  liver  into  "  day  "  and  "  night.""  Professor  Korte,  in  a  study 
of  this  remarkable  object,  summing  up  the  results  of  many  years  of 
research,  explains  this  by  showing  that  the  liver  was  regarded  as  a 
microcosm  reflecting  the  macrocosm,*^**  or,  in  other  words,  the  liver 
of  the  sacrificial  animal  from  being  originally  a  reflection  of  the 
soul  or  mind  of  the  god  to  whom  the  animal  was  offered,  was 
brought  into  connection  with  the  observation  of  the  heavenly  bodies 
revealing  the  intention  of  the  gods  acting  in  concert.  This  combina- 
tion of  hepatoscopy  with  astrology  likewise  points  to  the  latter  sys- 
tem of  divination  as  the  later  one,  dependent  in  some  measure  upon 
the  earlier  method  of  divining  through  the  liver. 

This  leads  us  to  the  last  two  points  to  be  considered  here,  the 
relationship  of  Babylonian- Assyrian  astrology  to  astronomy  and  the 
spread  of  astrology  from  the  Euphrates  Valley  to  other  peoples. 
While  astrology  even  in  its  most  primitive  phases  assumes  some 
knowledge  of  astronomy,  it  stands  to  reason  that  since  the  sole  pur- 
pose for  which  the  planets  and  stars  were  observed  was  as  a  means 
of  securing  omens,  there  could  be  no  genuine  interest  in  astronomical 
lore,  pure  and  simple.  As  the  scope  of  astrology  increased,  more 
stars  were  added  to  the  field  of  observation,  with  each  succeeding 
ages  further  details  of  the  movements  of  the  planets  were  noted,  and 
groups  of  stars  were  combined  into  constellations  of  a  more  or  less 
fanciful  character.  It  became  necessary  for  purposes  of  instruction 
in  astrology  to  systematize  and  synthesize  the  knowledge  thus  ac- 
quired from  empirical  observation.  In  the  course  of  time  a  con- 
siderable body  of  "  school  "  literature  thus  took  shape  in  the  form  of 
lists  of  stars,  with  attempts  to  locate  them  and  to  set  forth  some  of 
the  phenomena  connected  with  them.*^  For  the  practical  purpose  of 
regulating  the  calendar  further  pedagogical  aids  were  devised,  and 

■•Korte  (p.  362)  expresses  himself  as  follows  "Die  Leber,  der  Sitz  des 
Lebens  nach  antiker  Auflassung,  erscheint  als  ein  Abbild  des  Weltganzen  im 
kleinen,  Wie  dieses  ist  sie  in  eine  rechte  und  in  eine  linke  Halfte,  eine 
Tages — und  Nachtseite  geteilt.  Die  Trennungslinie  entspricht  der  Ost- 
Westlinie  des  Weltalls.  Wie  das  Himmelsgewolbe  ist  ihr  Rand  in  16 
Regionen  geteilt,  in  denen  Gotter  walten  und  Zeichen  geben  konnen." 

"As  examples  of  such  lists  see  II.  Rawlinson,  PI.  49,  Nos.  i,  3,  4;  III. 
Rawlinson,  PI.  57,  No.  6,  and  the  texts  entered  in  the  Index  to  Bezold's 
"Catalogue  of  the  Cuneiform  Tablets  of  the  Kouyunjik  Collection,"  p.  2096. 
These  lists  in  the  royal  library  of  Nineveh  revert  to  older  Babylonian  originals. 


,9o8J  IN   BABYLONIA  AND  ASSYRIA.  667 

thus  at  a  comparatively  early  age  the  seeds  for  a  genuine  science  of 
astronomy  were  planted.  The  fact,  however,  is  significant  that, 
with  perhaps  some  exceptions,  we  have  in  the  library  of  Ashur- 
banapal,  representing  to  a  large  extent  copies  from  older  originals, 
no  texts  that  can  properly  be  called  astronomical.'^*  For  this  reason 
a  reaction  has  set  in  among  Assyriologists  against  the  view 
formerly  held  that  astronomy  was  cultivated  at  an  early  period  in 
Babylonia  and  Assyria.*^  It  is  certainly  significant  that  the  astronom- 
ical tablets  so  far  found  belong  to  the  latest  period  and  in  fact  to  the 
age  following  upon  the  fall  of  the  Babylonian  empire.'^*  While  we 
must  be  warned  against  pressing  the  argument  ex  silentio  too  far, 
still  there  is  sufficient  evidence  to  warrant  the  conclusion  that  the 
most  glorious  period  of  Babylonian  astronomy  falls  in  the  fourth  to 
the  second  centuries  before  this  era,  that  is  to  say,  within  the  period 
of  the  Greek  occupation  of  the  Euphrates  Valley.  According  to 
Kugler,*"  the  oldest  dated  genuinely  astronomical  tablet  belongs  to 
the  seventh  year  of  Cambyses,  i.  e.,  522  B.  C,  although  it  shows  evi- 
dence of  having  been  revised  on  the  basis  of  an  older  original.  We 
also  find  evidence  of  changes  both  in  the  astronomical  terminology 
and  in  the  order  of  the  planets  after  c.  400  B.  C.,^*  so  that  while  we\ 
are  justified  in  going  back  to  the  neo-Baby Ionian  dynasty  as  the  point 
of  departure  for  the  beginnings  of  a  genuine  astronomical  science, 
it  would  be  rash  to  go  much  farther  back  than  this.    At  all  events 

"K.  9794  appears  to  be  purely  astronomical.  See  Bezold,  0.  c.  Vol.  V., 
p.  XXV.  and  iii.,  p.  1039 ;  also  Jeremias,  "  das  Alter  der  babylonischen  Astro- 
nomie"  (Leipzig,  1908),  p.  21. 

"For  a  fuller  discussion  of  the  recent  literature  on  the  subject  sec 
Jastrow,  0,  c,  XL,  pp.  232-434.  Kugler,  in  "  Kulturhistorische  Bedeutung  der 
Babylonischen  Astronomic"  {Vereinsschriften  der  Gorres-Gesellschaft,  1907, 
III.,  pp.  38-50),  maintains  the  late  origin  of  Babylonian  astronomy.  His 
views  have  been  accepted  by  Boll,  "  die  Erforschung  der  Antiken  Astrologie  " 
in  Neue  Jahrhiicher  fur  das  Klassische  Alter  turn,  i.  Abteilung,  Bd.  XXI., 
pp.  103-126)  and  others,  while  Jeremias  ("das  Alter  der  babylonischen 
Astronomic  ")  and  the  adherents  of  the  Winckler  school  cling  to  the  view  that 
astronomy  took  its  rise  in  the  early  period  of  Babylonian  history.  For  a 
general  summary  of  our  present  knowledge  of  Babylonian  astronomy,  on  the 
basis  chiefly  of  Kugler's  researches,  see  the  two  articles  by  Schiaparelli  in  the 
Rivista  di  Scienza,  III.,  pp.  213-259,  and  IV.,  pp.  24-54, 

•*  See  Kugler,  "  Stemkunde  und  Stemdienst  in  Babel,"  I.,  p.  2. 

••  Sternkunde,  p.  61. 

••0.  c,  pp.  12,  13,  22,  62,  etc. 


■/ 


668  JASTROW— HEPATOSCOPY  AND  ASTROLOGY         [December  4, 

it  is  not  until  we  reach  the  days  of  the  Seleucidian  and  Arsacidian 
dynasties  that  we  find  astroncwnical  calculations  of  the  movements 
\  and  of  the  position  of  the  moon  and  planets  in  full  swing. 

It  can  hardly  be  regarded  as  accidental  that  the  flourishing  period 
of  Babylonian  astronomy  should  thus  be  coincident  with  the  time 
when,  according  to  definite  evidence,  Babylonian  astrology  passed 
over  into  Greece.  "  The  conquest  of  Alexander,"  as  Bouche-Leclercq 
tersely  puts  it,  "  threw  down  the  barriers  hitherto  separating  races 

• 

and  civilizations."*^  To  Berosus,  the  "  Chaldaean  "  priest  who  wrote 
in  Greek  a  history  of  Babylonia  and  Assyria,  the  Greeks  themselves 
ascribe  the  introduction  of  astrology  into  their  midst.  Settling  in  the 
island  of  Cos,  the  home  of  Hippocrates,  Berosus  himself  taught  the 
Babylonian  system  to  the  students  whom  the  fame  of  the  great  phys- 
ician had  attracted  to  that  place.'^®  The  fragments  preserved  of  the 
writings  of  Berosus,*®  few  as  they  are,  suffice  to  show  that  he 
gathered  his  material  direct  from  the  sources,  and  there  is  therefore 
no  reason  to  question  that  he  followed  conscientiously  the  methods 
laid  down  in  the  Babylonian  collections  of  astrological  omens. 
While  it  is  of  course  possible  and  indeed  probable  that  through  the 
contact  with  the  Persians  the  Greeks  may  have  heard  of  the  Baby- 

y  Ionian  system  of  divining  the  future  through  the  stars,  it  is  certain 
/  that  astrology  did  not  take  a  definite  hold  on  the  Greeks  and  become 
I    part  of  their  intellectual  outfit  until  the  days  of  Berosus,  i.  e,,  till 

\  about  the  beginning  of  the  third  century  B.  C.  A  few  centuries 
sufficed  to  transform  Babylonian  astrology  under  the  influence  of 
the  Greek  spirit  from  the  character  of  an  "oriental  religion " 
which  as  Bouche-Leclercq*®  recognised  it  had  at  the  time  of  its 
adoption,  into  the  appearance  of  a  science.  Already  advanced  stu- 
dents of  astronomy,  the  Greek  physicists  combined  astrology  with 
the  principles  and  speculations  of  mathematics  and  brought  it  into 
accord  with  the  current  systems  of  philosophy  until  it  became  a 
genuine  expression  of  the  Greek  spirit  and  an  integral  part  of 
Greek  culture.    A  feature  which  the  Greeks  introduced  and  which 

■^  UAstrologie  Grccque/'  p.  35. 

"Vitruvius,  de  Architectura,  IX.,  6.      See  also  Bouche-Leclercq,  o,  c.» 
pp.  2  and  37. 

"Cory,  "Ancient  Fragments,"  pp.  51-69. 
^  0.  c,  p.  I. 


,9o8.|  IN   BABYLONIA  AND  ASSYRIA.  669 

of  itself  served  to  change  the  aspect  of  the  Babylonian  system  was 
tKe  perfection  of  a  method  whereby  the  fate  of  the  individual  was 
brought  into  connection  with  the  stars.  The  science  of  genethli- 
alogy®^  or  the  casting  of  the  individual  horoscope  from  the  position 
of  the  stars  at  the  time  of  an  individual's  birth  is  a  distinctly  Greek 
contribution.  The  insignificant  role  that  the  individual  plays  in  all 
phases  of  divination,  except  in  the  case  of  the  accidents  and  unusual 
incidents  that  happen  to  him  and  which  were  therefore  looked  upon 
as  signs  sent  by  the  gods  to  the  individual  as  such,  prevented  the 
rise  of  the  thought  that  the  activity  of  the  gods  as  shown  in  the 
heavens  had  any  bearing  on  the  fate  of  the  individual.  As  we  have\ 
seen,  astrology,  just  as  hepatoscopy,  concerned  itself  in  Babylonia  /  k 
and  Assyria  with  the  general  welfare  and  the  public  state.  There/ 
was  no  place  in  either  of  the  two  great  systems  of  divination  for  the 
individual  and  we  may  go  a  step  farther  and  assert  that  it  was  con- 
trary to  the  entire  spirit  of  the  Babylonian-Assyrian  religion  to  sup- 
pose that  the  gods  concerned  themselves  with  the  individual  suffi- 
ciently to  give  him  as  such,  through  the  stars  or  through  the  liver  of 
a  sacrificial  animal,  an  indication  of  what  they  purposed  doing.**  It 
was  diflFerent  in  Greece  where  long  before  the  time  that  Babylonian 
astrology  was  assimilated  to  Greek  culture,  the  individual  had  as- 
serted himself  to  an  extent  undreamed  of  in  the  Euphrates  Valley. 
Instead  of  an  intellectual  oligarchy  with  all  learning  confined  to 
priestly  circles,  corresponding  to  the  concentration  of  all  political 
power  in  the  hands  of  a  few  privileged  families,  we  have  in  Greece 
a  republic  of  letters  with  an  independence  of  thought  only  surpassed 
by  the  strength  of  individualism  in  the  political  sphere.  Religion  had 
long  ceased  to  be  the  controlling  factor  or  at  least  the  predominant 

~Bouche  Leclercq,  /.  r.,  p.  49,  while  noting  that  there  is  no  trace  of  the 
application  of  the  astrology  to  the  individual  horoscope  in  cuneiform  texts,  is 
disposed  to  attribute  this  to  the  dearth  of  material.  Since  he  wrote  his  great 
work  that  material  has  largely  increased,  and  it  is  perfectly  safe  to  conclude 
that  this  phase  of  astrology  never  existed  in  the  Euphrates  Valley. 

"If  in  a  few  very  late  texts  (cf.  Bouch^-Leclercq,  /.  c.,  p.  50)  we  find 
entries  of  the  birth  of  a  child  with  the  mention  of  the  aspect  of  the  moon, 
planets  and  constellations,  this  is  to  be  ascribed  to  Greek  influence  as  Bouch6- 
Leclercq  himself  suggests.  Some  Greek  astrologers  even  went  so  far,  accord- 
ing to  Vitruvius  (/.  r.),  as  to  cast  the  horoscope  of  an  individual  from  the 
time  of  conception. 


670  JASTROW—HEPATOSCOPY   AND  ASTROLOGY         [December  4, 

factor  in  Greek  civilization.  One  science  after  the  other  had  freed 
itself  from  the  thraldom  of  religious  tradition  and,  accordingly, 
astrology,  when  introduced  into  Greece,  did  not  become  a  part  of 
the  Greek  religion  but  an  element  of  Greek  science.  Passing  on  to 
the  Romans**'  as  an  integral  part  of  Greek  culture,  and  becoming 
with  the  spread  of  Roman  authority  the  general  possession  of  the 
ancient  world,  astrology,  because  of  its  indissoluble  association  with 
astronomy,  mathematics,  and  the  philosophical  systems  of  Greece,  be- 
came part  of  the  heritage  of  Greece  to  the  world  and  took  on  in 
time  the  aspects  of  a  religious  cult.*^  With  the  revival  of 
Greek  influence  through  the  intellectual  movement  following  upon 
the  rise  and  spread  of  Islamism,  astrology  took  a  firm  hold  on  the 
choice  minds  of  mediaeval  Europe  by  the  side  of  such  a  force  as 
Aristotelianism,®*  and  continued  to  sway  men's  minds  till  the  thresh- 
old of  modern  scientific  thought,  when  it  was  swept  away  with  so 
many  other  cherished  traditions  from  the  broad  highway  of  science 
into  the  byways  where  it  still  flourishes  at  the  present  time  and  will 
no  doubt  continue  to  do  so  for  a  long  time  to  come.  Though 
somewhat  more  complicated  in  its  processes,  mediaeval  and  modem 
astrology  is  practically  identical  with  the  form  it  took  on  in  Greece.** 
Not  only  did  Greek  astrology  make  its  way  throughout  the  West  but 
it  spread  also  to  the  East,  for  it  has  been  definitely  ascertained  that 
what  we  find  of  it  in  India  and  even  in  China  is  due  to  the  spread  of 
the  sphere  of  Greek  influence  ;**  and  the  same  holds  good  for  Egypt, 
where  it  begins  to  flourish  with  the  rise  of  Hellenistic  culture.'^ 

"*  Bouche-Leclercq,  /.  c,  Chap.  XVI.,  "  L'AstroIogie  dans  le  Monde  Re- 
main "  and  "  Cumont/'  "  Les  Religions  Orientales  dans  le  Paganisme  Romain  " 
(Paris,  1907),  Chap  VII. 

*•  See  Cumont,  Tlnfluence  religieuse  de  TAstrologie  dans  le  Monde  Ro- 
main (Transactions  of  the  3d  International  Congress  for  the  History  of 
Religions,  II.,  pp.  197-198). 

•*  Bouche-Leclercq,  pp.  624  seq. 

*  Compare  for  example  the  ideas  associated  with  the  planets  in  a  modem 
manual  of  astrology  like  Ellen  H.  Bennett's  "Astrology"  (New  York,  1897), 
PP-  93-100,  with  Bouche-Leclercq's  statement  of  the  Greek  views  ("  UAstrol- 
ogie  Grecque,"  pp.  93-101  and  311-326). 

"Thibaut,  "Astronomic,  Astrologie  und  Mathematik,"  in  Biihler-Kiel- 
horn,  "Grundriss  der  Indo-Arischen  Philologie,"  III.,  9,  p.  15,  and  Kugler, 
"  Kulturhistorische  Bedeutung  der  babylonischen  Astronomic,"  p.  49. 

"  It  is  one  of  the  many  merits  of  Bouche-Leclercq  to  have  demonstrated 


I908.1  IN   BABYLONIA  AND  ASSYRIA.  671 

We  thus  find  the  source  of  all  astrology  in  the  ancient  world 
in  the  system  that  arose  in  the  Euphrates-Valley;  and  in  view 
of  this  it  will  be  admitted  that  the  thorough  study  of  Babylonian- 
Assyrian  astrology  is  a  factor  of  considerable  importance  in 
tracing  the  intellectual  development  of  mankind.  Coming  backj 
therefore,  to  our  immediate  subject  we  have  the  curious  phe- 
nomenon that  about  coincident  with  the  period  when  a  genuine 
science  of  astronomy  takes  a  firm  footing  in  Babylonia,  astrol- 
ogy begins  its  triumphant  march  throughout  the  world.  It  is 
tempting  to  suppose  that  we  have  in  this  phenomenon  the  symp- 
tom of  an  "  exchange  "  of  influences  that,  while  on  the  one  hand 
Babylonia  gave  astrology  to  Greece,  the  contact  with  the  scien- 
tific spirit  of  Greece  resulted  in  giving  an  impetus  to  astronom- 
ical investigations  in  Babylonia.  The  possibility,  indeed,  of  Greek 
influence  on  Babylonian  astronomy  was  suggested  by  Bouche- 
Leclercq  and  is  favored  by  Kugler.*'  Since,  as  now  appears,  the 
credit  for  the  discovery  of  the  precession  of  the  equinoxes  rests  with 
the  Greek  astronomer,  Hipparch,  who  announced  it  c.  130  B.  C.,  and 
since  it  would  indeed  appear  that  in  the  second  century  B.  C.  the 
Babylonians,  according  to  Kugler,  were  still  ignorant  of  this  prin- 
ciple, there  is  certainly  every  reason  to  suppose  that  the  Babylonians 
were  in  this  instance  the  pupils,  and  the  Greeks  the  teachers.  On  the 
other  hand,  the  Greek  astronomers  seem  to  have  obtained  from  the 
Babylonians  the  names  for  the  constellations  of  the  ecliptic  which  we 
still  use  at  the  present  time.  Certainly,  for  the  beginnings  of  their v 
astronomy  the  Babylonians  are  not  indebted  to  the  Greeks  since  \ 
those  beginnings  reach  back  beyond  the  contact  of  Orient  with  j 

in  his  great  work  on  Greek  astrology  the  worthlessness  of  the  traditions 
which  ascribe  Greek  astronomy  and  astrology  to  an  Egyptian  origin.  Sec 
especially  the  important  note  ("  U Astro! ogie  Grecque,"  pp.  51-52)  from  which 
it  appears  that  "  Chaldean  "  and  "  Egyptian  "  are  used  almost  interchangeably 
by  uncritical  Greek  and  Roman  writers  who  hand  down  more  or  less  fanciful 
traditions.  Since  Boll  ("  Sphaera,"  p.  159  seq.)  and  others  have  demon- 
strated the  late  origin  of  the  zodiac  of  Denderah,  the  chief  evidence  for  the 
early  introduction  of  astronomy  in  Egypt  has  fallen  away;  and  there  is  no 
reason  for  assuming  that  astrology  flourished  in  Egypt  before  the  Ptolemaic 
period. 

"  Bouch6-Leclercq,  0.  c,  p.  50  and  Kugler,  "  Kulturhistorische  Bedeutung 
der  babylonischen  Astronomic,"  p.  48. 


672  JASTROW— HEPATOSCOPY  AND  ASTROLOGY        [December 4, 

Occident,  but  that  would  not  preclude  the  possibility  of  influences 
from  the  side  of  Greece  at  a  later  stage  in  the  development  of  astro- 
nomical lore. 

To  account  for  the  point  of  departure  for  the  unfolding  of  a 
g[enuine  science  as  astronomy^  independent  of  merely  empirical  ob- 
servations in  the  interest  of  astrology,  and  which  as  we  saw**  dates 
from  the  sixth  century  B.  C,  we  have  another  factor  entering  into 
Babylonia  about  this  time  that  must  have  exerted  a  profound  in- 
fluence— ^the  appearance  of  Persia  on  the  scene  and  with  it  the 
advanced  form  of  faith  known  as  Zoroastrianism  and  which  by  com- 
parison with  the  emphatically  polytheistic  conceptions  of  the  Baby- 
lonian religion  was  superlatively  rationalistic.  Contact  with  a 
strange  culture  is  always  attended  by  an  intellectual  stimulus,  and 
this  takes  place  whether  the  contest  be  friendly  or  hostile.  Though 
the  Persian  rulers  even  after  Darius  with  whom  the  full  sway 
of  Zoroastrianism  may  be  said  to  beg^n,  maintained  a  conciliatory 
attitude  towards  the  gods  of  Babylonia,  Cyrus  going  so  fan  as  to 
claim  that  his  conquest  of  the  country  was  in  the  interest  of  Mar- 
/duk,^®  nevertheless,  the  presence  of  a  totally  different  religion,  recog- 
/  nized  as  the  oflScial  one  by  the  Persian  rulers  from  the  days  of 
*  Darius  on,  must  have  acted  as  a  disintegrating  element  that  led  to  a 
\  decline  in  the  belief  in  the  Babylonian  gods  and  to  a  corresponding 
\weakening  of  the  hold  that  the  ofiicial  rites  had  on  the  people.  I  ven- 
ture to  think  that  the  influence  of  Zoroastrianism,  bringing  in  its 
wake — as  did  Christianity  and  as  did  Islamism — a  wave  of  intellec- 
tual advance,  is  the  factor  which  accounts  for  the  definite  separation 
of  the  study  of  the  heavenly  phenomena  from  being  merely  an  ad- 
junct to  a  system  of  divination,  to  take  its  position  as  a  genuine  and 
independent  science.  A  further  impetus  to  the  new  science  was  given 
by  the  contact  with  Greek  culture  with  the  further  possibility  of  a 
direct  influence  of  Greek  astronomical  theories  and  methods  on  the 
investigations  of  the  Babylonian  priests. 

The  advance  of  astronomy  must,  however,  have  reacted  also 
on  the  basic  principle  which  we  have  seen  underlay  Babylonian- 
Assyrian  astrology.    Though  even  the  &aru-priests,  while  still  com- 

•  See  above,  p.  667. 

^  Hagen,  Cynis-Texte  in  "  Beitrage  zur  Assyriologie/'  II.,  p.  229. 


,9o8.]  IN   BABYLONIA  AND  ASSYRIA.  673 

pletely  enthralled  by  astrology,  must  have  been  impressed  with  the 
domain  of  law  in  the  movements  and  phenomena  of  the  heavens, 
there  remained  enough  scope  for  caprice  in  the  more  unusual  phe- 
nomena which  the  imperfect  knowledge  placed  outside  of  the  sphere 
of  regularly  working  law.  With  the  gradual  reduction  of  this 
scope  until  through  astronomical  calculations  even  such  phenomena 
as  eclipses  came  within  the  range  of  recognized  law,  the  belief  in  as- 
trology must  have  suffered  a  decline,  at  all  events  in  the  minds  of  the 
better  informed  priests.  Astronomy  and  astrology  presented  a  con- 
trast not  unlike  that  which  in  modern  times  is  frequently  represented 
by  science  and  religion  and  though  no  open  conflict  ensued,  the  / 
growth  of  astronomy  must  have  involved  the  decline  of  astrology. 
If  the  data  of  astrology  are  all  due  to  the  workings  of  inevitable  and 
clearly  recognized  eternal  laws,  there  is  no  room  for  any  spontaneity 
on  the  part  of  the  gods,  so  far  at  least  as  the  stars  manifest  divine 
activity.  Every  advance  in  astronomy,  therefore,  removed  a  stone 
from  the  foundation  on  which  the  structure  of  astrology  was  reared, 
until  the  stability  of  the  entire  structure  was  endangered.    The  last 


three  centuries  before  our  era  represent  in  general  a  period  of  de- 
clining faith  in  the  gods  both  in  Babylonia  as  well  as  in  Greece  and 
elsewhere.  The  old  order  throughout  the  ancient  world  of  cultural 
development  was  passing  away,  and  the  growing  strength  of  astron- 
omy is  in  itself  symptomatic  of  the  new  order  destined  to  take 
the  place  of  the  old.  It  is  no  unusual  phenomenon  to  find  a  great 
civilization  handing  over  to  posterity  as  a  legacy  at  the  period  of  its 
decay — a  superstition  instead  of  a  real  achievement.  "  The  evil 
that  men  do  lives  after  them;  the  good  is  oft  interred  with  their 
bones  "  applies  to  nations  as  to  individuals,  and  so  it  happens  that 
while  the  wholesome  fruits  of  the  Babylonian-Assyrian  civilization 
were  not  entirely  lost,  the  overripe  products  with  the  odor  of  decay 
pervading  them  were  the  first  to  be  exported  to  other  climes. 
What  became  proverbial  among  Greeks  and  Romans  as  "  Chaldaean 
wisdom  "  is  not  the  astronomy  of  Babylonia  but  the  astrology  which, 
after  having  spent  its  force  in  the  soil  in  which  it  arose,  takes  root 
elsewhere  and  soon  flourishes  more  luxuriantly  than  it  ever  did 
in  its  native  heath.  We  have,  however,  also  seen  that  in  the  care 
of  others  the  original  plant  was  modified  through  the  transfer  from 


\ 

/ 


674  JASTROW— HEPATOSCOPY  AND  ASTROLOGY        [December  4. 

/  the  Orient  to  the  Occident.  Astrology  in  Babylonia  declines  as 
astronomy  grows,  for  the  very  reason  that  astronomy  is  an  outgrowth 
of  astrology,  representing  the  evolution  of  a  science,  by  the  break- 
ing away  from  attachment  to  a  religion  and  a  cult.  In  Greece 
astronomy  arises  as  do  other  sciences  through  the  growth  of  the 
spirit  of  investigation.  There  was  so  far  as  we  can  see  no  religious 
tradition  out  of  which  or  in  opposition  to  which  astronomy  took  its 
rise.  There  is  no  antecedent  astrology  from  which  astronomy 
emerges  as  the  butterfly  from  the  chrysalis.  Therefore,  astrology 
coming  to  the  Greeks  as  a  novel  conception,  with  all  the  force  of  an 
apparently  practical  application  of  a  scientific  theory,  suggesting 
the  possibility  of  a  direct  communion  with  the  arbiters  of  human 
fate — the  conscious  goal  or  unconscious  hope  of  all  religions — ^it  was 
capable  of  being  assimilated  to  the  already  firmly  established  astron- 
omy. Astrology  as  further  developed  by  the  Greeks  became  merely 
one  of  the  phases  of  astronomy,  as  is  shown  by  the  synonymity 
of  the  two  terms,  AorpoAoyia  and  turrpovoyuiaJ^ — a  condition  which 
persisted  till  mediaeval  scholasticism,  which  distinguishes  merely 
as  a  matter  of  definition  between  "  natural  astrology  "  or  theoretical 
astronomy  and  "  judicial  astrology  "  or  divination  through  the  stars 
as  the  application  of  the  theory  to  human  life. 

Lastly,  if  another  suggestion  be  permitted,  the  "  Chaldaeans  " 
whom  we  encounter  so  frequently  in  Greek  and  Roman  writers 
acting  as  "  diviners  "  on  such  various  occasions,  appear  to  be  indeed 
Babylonian  fedrw-priests  or  the  disciples  of  these  priests  who,  because 
of  the  decline  of  faith  in  astrology  in  the  centers  in  which  it  arose, 
left  their  homes  to  seek  their  fortunes  elsewhere.  As  with  the 
growth  of  astronomical  lore,  the  hold  of  the  old  system  of  astrology 
was  loosened,  the  occupation  of  the  fedrM-priests  was  gone.  Their 
condition  was  not  unlike  that  of  the  Levites  who,  as  the  priests  of  the 
local  sanctuaries  in  Palestine,  were  deprived  of  their  standing  and 
livelihood  with  the  decline  of  these  sanctuaries  through  the  gradual 
concentration  of  Jahweh  worship  in  the  central  sanctuary  of  Jerusa- 
lem. These  Levites  wandered  to  Jerusalem  where,  according  to  the 
Priestly  Code,  provision  was  made  for  them  by  assigning  them  to 
posts  as  assistants  to  the  kohanim — the  legitimate  priests  of  the  cen- 

"See  Bouche-Leclercq,  0.  c,  p.  3,  note  2. 


i9o8,] 


IN   BABYLONIA  AND  ASSYRIA.  675 


tral  sanctuary."  The  Wru-priests  of  Babylonia  in  their  capacity  as 
astrologers  wandered  to  the  West,  there  to  ply  their  trade  for  which 
a  market  was  no  longer  forthcoming  in  their  own  homes.  Baby- 
lonian astrology,  enjoying  the  popularity  in  Greece  and  in  the  Roman 
empire  frequently  granted  to  a  foreign  importation  in  preference 
to  a  home  industry,  became  the  fashion  of  the  Occident  during  the 
centuries  that  marked  the  decline  of  belief  in  the  gods  of  Greece 
and  Rome  and  that  offered  a  hospitable  welcome  to  all  kinds  of 
strange  faith  and  mystic  cults,  until  the  term  "  Chaldaean  "  became 
synonymous  with  "  astrologer."  In  time  it  was  no  doubt  applied 
to  the  one  who  divined  through  the  stars  irrespective  of  his  origin.^** 
Besides  astrology,  hepatoscopy  was  also  practiced  by  these  "  Chal- 
daeans,"^'  but  both  forms  of  divination,  being  derived  from  an 
official  cult  and  practiced  purely  as  a  profession  that  was  presumably 
not  without  profit  suffered,  as  was  inevitable,  a  degeneration,  with 
the  result  that  a  measure  of  reproach  became  attached  to  the  term 
"  Chaldaean,"  which  acquired  almost  the  force  of  trickster  and  de- 
ceiver. It  was  nevertheless  fortunate  that  the  term  survived  as  a 
fingerpost,  directing  us  to  the  land  in  which  the  system  of  divination 
arose  that  after  strange  vicissitudes  has  survived  in  the  form  as 
modified  under  Greek  influences  and  with  some  additions  in  the 
mediaeval  period,  to  our  own  days,  still  finding  many  devotees  in 
circles  where  one  would  hardly  expect  to  encounter  them.^* 

The  degenerating  process  through  which  the  term  "  Chaldaean  " 

"See  e.  g.,  Baudissin,  Geschichte  des  Altestamentlichen  Priesterhums 
(Leipzig,  1889),  p.  287. 

^^  So,  e.  g.,  Teukros,  the  author  of  a  Greek  treatise  on  astrolog>%  is  called 
"  the  Babylonian  "  evidently  in  the  sense  of  "  astrologer."  See  the  fragments 
of  this  treatise  published  by  Boll  ("  Sphaera,"  pp.  16-21)  who  places  Teukros 
in  the  first  century  of  this  era. 

"  See  the  story  told  by  Polyaenus,  "  Strategicon,"  IV.,  20,  of  the  decep- 
tion practised  upon  the  army  of  Attalus  I.  of  Pergamon  by  Soudinos  "  a 
Chaldaean  augur"  who  writing  the  words  "  victory  of  the  king"  ({SaoiXicj^  vIkj]) 
backwards  on  the  palm  of  his  hand,  pressed  the  smooth  side  of  the  liver  of 
a  sacrificial  animal  on  his  hand,  and  then  held  the  liver  with  the  significant 
words  inscribed  on  it  to  the  gaze  of  the  army,  who  regarded  it  as  a  sign 
sent  by  the  gods.     See  also,  above,  p.  650,  note  13. 

^*The  late  Richard  Gamett  is  only  one  of  many  examples  of  men  other- 
wise abreast  with  modem  thought  who  cling  to  the  faith  in  the  revelations 
of  the  stars. 


676  JASTROW— HEPATOSCOPY   AND  ASTROLOGY         [December  4, 

passed  must  not,  however,  lead  us  to  the  conclusion,  which  would  be 
decidedly  false,  that  astrology  when  it  passed  over  to  the  West 
became  wholly  at  the  mercy  of  professional  tricksters.  This  is  but 
one  phase  of  the  subject  which,  seriously  cultivated  by  Greek  physi- 
cists, became  bound  up  as  we  have  seen  with  advanced  forms  of 
astronomy,  mathematics  and  philosophical  speculation.  It  is  the 
old  Babylonian  astrology  directly  imported  by  "  Chaldaeans  "  as 
professional  asfrologers  that  degenerated  into  a  dishonest  trade, 
whereas  the  modification  of  the  Babylonian  system  under  the  in- 
fluence of  the  Greek  scientific  spirit  was  raised  to  the  dignity  of  a 
genuine  science ;  and  belief  in  it  remained  an  integral  part  of  science 
throughout  the  middle  ages.  In  our  days  when  the  new  scientific 
spirit  has  definitely  broken  with  astrology,  we  are  witnessing  a 
process  not  unlike  that  which  set  in  when  faith  in  the  Babylonian 
system  declined  in  the  land  of  its  birth.  Whatever  justifiable  basis 
(if  any)  it  may  have  had  is  entirely  obscured  by  those  who  exploit 
it  as  a  profession.  The  modem  "  astrologers  "  are  not  the  Greek 
astronomers  attaching  to  their  science  a  divinatory  aspect,  but  the 
old  Mfw-priests  in  a  new  garb,  plying  a  trade  that  flourishes  through 
the  readiness  of  people  to  be  deceived — a  readiness  that  amounts 
almost  to  willingness.  Why  then,  it  may  be  asked,  search  out  the 
follies  and  superstitions  of  the  past  ?  Bouche-Leclercq^*  supplies  us 
with  the  answer  when  he  says  "  that  it  is  not  a  waste  of  time  to  find 
out  how  other  people  wasted  theirs." 


TB  II 


UAstrologie  Grecque,"  p.  ix. 


FURTHER  STUDIES   ON   TRANSPLANTATION   OF 

VESSELS  AND   ORGANS.^ 

By  ALEXIS  CARREL. 
(Read,  November  6,  1908.) 

It  is  known  that  tissues  can  be  removed  from  an  animal,  trans- 
planted into  another  animal  and  live  normally  in  the  body  of  their 
new  owner.  The  transplantation  of  anatomical  structures  has 
already  been,  and  will  be  again  in  the  future,  used  in  human  sur- 
gery. For  instance,  an  excellent  method  of  treating  an  aneurism 
of  the  femoral  artery  would  be  the  extirpation  of  the  diseased  part 
and  its  replacement  by  a  piece  of  artery  of  same  caliber.  This 
new  artery  cannot  be  taken  from  an  animal  and  grafted  on  man, 
for  the  serum  of  an  animal  is  toxic  for  the  cells  of  an  animal  of 
different  species.  A  dog's  vessel  transplanted  on  man  could  pos- 
sibly perform  its  arterial  functions,  but  the  histological  structure 
of  its  wall  would  be  deeply  modified  and  accidents  could  occur. 
It  is  probable  that  arteries  from  anthropoid  ape  would  be  of  safer 
use,  because  man  and  ape  are  closely  related  from  a  zoological 
standpoint.  But  this  would  be  exceedingly  expensive  and  not  prac- 
tical. It  will  be  safer  and  simpler  to  graft  on  man  vessels  taken 
from  another  man.  The  vessels  can  be  extirpated  from  an  ampu- 
tated limb  or  from  the  body  of  a  criminal  or  of  a  man  killed  by 
accident.  But  it  is  sure  that  these  cases  will  not  present  themselves 
at  the  time  convenient  for  the  surgeon  and  his  patient.  Therefore, 
it  is  important  to  find  a  method  to  store  human  vessels  during  the 
period  which  will  elapse  between  their  extirpation  and  their  graft 
on  the  patient.  With  this  view,  I  have  attempted  to  preserve 
arteries  in  a  condition  of  latent  life,  in  order  that,  after  having  spent 
several  days  or  several  weeks  outside  of  the  body,  they  can  be  trans- 
planted  successfully. 

*  From  the  Laboratories  of  the  Rockefeller  Institute  for  Medical  Research. 

677 


678  CARREL— FURTHER  STUDIES  ON  [NoYcmbere. 

Before  describing  the  method  which  renders  possible  the  preser- 
vation of  arteries,  I  shall  briefly  summarize  some  of  the  results 
obtained  at  the  Rockefeller  Institute  in  the  transplantation  of  blood 
vessels  and  organs.  These  operations  became  possible  as  soon  as  a 
practical  method  of  uniting  blood  vessels  was  found.  Success  in 
transplanting  organs  is  direct  function  of  the  circulation.  The  cir- 
culation cannot  be  immediately  reestablished  but  by  the  sewing  of 
the  vessels  of  the  organ  to  those  of  the  host.  The  sewing  of  vessels 
is  today  a  very  easy  operation.  Some  years  ago,  while  I  was  work- 
ing at  the  University  of  Lyons,  I  found  a  method  of  uniting  severed 
arteries  or  veins,  which  gave  excellent  results.  This  method  was 
progressively  improved  in  such  a  manner  that  it  is  practically 
always  successful.  The  vessels  heal  very  quickly  and  no  coagula- 
tion of  the  blood  occurs  when  the  operation  is  aseptic  and  the  union 
of  the  vascular  ends  accurate.  The  scar  of  the  severed  vessels  is, 
in  many  cases,  so  small  that  after  a  few  months  it  is  hardly  dis- 
cernible. On  a  renal  vein  examined  a  little  over  two  months  after 
the  sewing,  it  was  impossible  to  localize  exactly  the  position  of  the 
anastomosis.  The  anastomosis  of  the  renal  artery  was  represented 
only  by  an  indistinct  line  crossing  the  intima.  Twelve  months  after 
the  anastomosis  of  a  carotid  artery,  the  anatomical  specimen  was  re- 
moved and  examined.  After  longitudinal  incision  of  the  wall,  no 
scar  was  seen  on  the  intima,  there  was  no  modification  of  the  caliber. 
But,  in  one  small  point,  the  vessel  had  lost  part  of  its  elasticity  and 
it  permitted  to  localize  approximately  the  anastomosis.  The  results 
are  permanent.  Two  and  three  years  after  the  operation,  the  circu- 
lation through  the  anastomosis  remains  normal.  It  must  be  known 
also  that,  if  the  method  is  not  correctly  applied,  or  a  fault  of  tech- 
nique, even  very  slight,  is  made,  thrombosis  may  occur.  Success 
depends  much  less  on  the  way  of  handling  the  needles  or  passing 
the  threads  than  on  the  knowledge  of  the  causes  which  are  able  to 
produce  thrombosis  and  their  removal.  On  human  beings,  this 
method  has  already  been  successfully  used  by  American  and  Euro- 
pean surgeons,  and  on  animals,  it  has  permitted  to  perform  the 
transplantation  of  blood  vessels,  organs  and  limbs. 

The  graft  of  a  segment  of  artery  on  an  artery  of  another  animal 
of  the  same  species  is  ordinarily  successful  when  the  vessels  are  of 


,908.]  TRANSPLANTATION  OF  VESSELS  AND  ORGANS.  679 

sufficient  caliber.  After  a  few  months,  the  transplanted  segment 
assumes  exactly  the  same  appearance  as  the  normal  vessel.  The 
carotid  of  a  dog  was  examined  three  months  after  the  graft  of  an 
arterial  segment.  The  transplanted  segment  was  exactly  similar  to 
the  other  parts  of  the  artery.  There  was  no  modification  of  caliber. 
The  elasticity  was  normal.  The  only  evidence  of  the  operation  was 
two  whitish  transverse  lines  on  the  intima.  The  remote  results  are 
excellent.  A  dog,  into  whose  aorta  a  segment  of  aorta  from 
another  dog  had  been  transplanted,  was  living  and  in  good  health 
nine  months  after  the  operation  and  the  femoral  pulse  was  normal. 
The  transplantation  of  arteries  has  already  been  attempted  in  human 
surgery  by  Pierre  Delbet  in  the  treatment  of  aneurism.  When  a 
large  artery  is  wounded  and  partially  destroyed,  or  when  a  tumor 
adherent  to  the  main  vessels  of  a  limb  renders  necessary  the  extir- 
pation of  these  vessels,  the  substitution  of  a  new  piece  of  artery  to 
the  removed  part  would  -prevent  the  occurrence  of  gangrene. 

The  graft  of  an  artery  of  an  animal  into  an  animal  of  different 
species  is  often  successful  if  the  animals  are  closely  related.  I 
transplanted  several  times  segments  of  dog's  carotid  arteries  on  the 
abdominal  aorta  of  cats  with  excellent  functional  results.  Never- 
theless, these  results  cannot  be  compared  with  those  obtained  in 
transplantation  between  animals  of  same  species.  Sometimes  the 
lumen  becomes  dilated,  or  even  a  fusiform  aneurism  can  be  found. 
Even  when  the  functions  of  the  transplanted  segment  are  perfect, 
its  wall  undergoes  marked  histological  changes.  The  elastic  frame- 
work disappears  and  progressively  the  muscular  fibers  are  resorbed. 
After  a  few  months,  they  have  practically  disappeared.  The  ves- 
sel is  then  composed  mainly  of  connective  tissue. 

Veins  can  easily  be  grafted  on  arteries.  I  performed  several 
times  the  transplantation  of  the  vena  cava  on  the  aorta,  on  dogs  and 
on  cats,  with  excellent  results.  A  segment  of  vein  transplanted 
into  an  artery  undergoes  immediately  very  marked  changes.  The 
wall,  which  is  very  thin,  becomes  thicker  and  stronger.  The  lumen 
is  often  dilated,  but  no  aneurism  has  ever  been  observed.  On  the 
contrary,  the  vein  reacts  against  the  increased  blood  pressure  by 
thickening  its  wall.  The  thickening  is  due  to  an  hyperplasy  of  the 
muscular  cells  and  an  hypertrophy  of  the  adventitia.    There  is  also 


680  CARREL— FURTHER    STUDIES    ON  [November  6. 

a  very  large  increase  of  the  interstitial  connective  tissue  of  the 
media.  The  venous  wall  becomes  as  strong  as  the  arterial  wall. 
The  function  has  created  the  organ.  Therefore,  veins  can  act  as 
a  substitute  for  arteries.  This  is  of  practical  importance  in  human 
surgery,  for  on  the  patient  himself  an  abundant  supply  of  vein  can 
always  be  found. 

The  organs,  kidneys,  spleen,  or  thyroid  gland,  for  instance,  can 
be  transplanted  from  an  animal  to  another  animal  and  their  circu- 
lation immediately  reestablished  by  suture  of  the  blood  vessels  to 
those  of  their  host.  Two  methods  are  used — ^the  simple  trans- 
plantation, and  the  transplantation  in  mass.  The  simple  trans- 
plantation consists  of  dissecting  the  organ,  cutting  its  vessels,  and 
uniting  these  vessels  directly  to  those  of  the  host.  In  the  trans- 
plantation in  mass,  the  organ  is  extirpated,  together  with  the  sur- 
rounding tissues  and  organs,  its  nerves,  vessels  and  the  main  vessels 
of  the  region.  After  transplantation,  the  anastomoses  are  not  made 
on  the  vessels  of  the  organ  themselves,  but  on  the  main  vessels  of 
the  anatomical  region.  The  transplantation  in  mass  of  the  kidneys 
has  been  performed  on  cats.  It  consists  of  extirpating  from  a  first 
animal  both  kidneys,  Jtheir  vessels  and  the  corresponding  segments 
of  the  aorta  and  vena  cava,  their  nerves  and  nervous  ganglia,  their 
ureters  and  the  corresponding  part  of  the  bladder ;  of  placing  these 
anatomic  specimens  into  the  abdominal  cavity  of  a  second  animal 
whose  kidneys  have  been  previously  resected  and  the  aorta  and 
vena  c^va  cut  transversely ;  and  of  suturing  the  vascular  segments 
between  the  ends  of  the  aorta  and  vena  cava,  and  of  grafting  the 
flap  of  bladder  onto  the  bladder  of  the  host.  In  every  case  the 
reestablishment  of  the  renal  functions  was  observed.  These  func- 
tions were  determined  by  the  character  of  the  urine  and  the  general 
condition  of  the  animals. 

The  secretion  of  urine  often  begins  as  soon  as  the  arterial  circu- 
lation is  reestablished.  In  some  cases  the  amount  of  urine  during 
the  first  twenty-four  hpurs  was  more  than  loo  c.c.  However,  a  cat 
urinated  only  25  c.c.  during  the  first  twenty- four  hours ;  the  second 
day  the  amount  of  urine  passed  was  only  16  c.c. ;  this  urine  was 
highly  concentrated  and  contained  much  urea.  Every  cat  urinated 
abundantly  every  day,  but  the  animals  presented  sooner  or  later 


,9o8.J  TRANSPLANTATION   OF  VESSELS  AND  ORGANS.  681 

some  complication,  which  modified  in  some  measure  the  renal  func- 
tions. As  is  to  be  expected  after  an  operation  as  complex  as  the 
transplantation  in  mass,  various  accidents  occurred ;  hydronephrosis, 
intestinal  compression  by  peritoneal  adhesions,  volvulus,  phlegmon, 
puerperal  infection,  compression  of  the  renal  veins  by  organized 
hematoma  of  the  connective  tissue,  which  were  the  direct  or  indirect 
causes  of  death  in  these  animals.  However,  in  two  experiments  the 
functions  of  the  kidneys  seem  to  have  been  for  a  certain  time  almost 
completely  normal.  The  color  of  the  urine  was  yellow,  generally, 
or  often  less  dark  than  the  normal  urine  of  the  cat.  Its  reaction 
was  acid.  Its  quantity  for  twenty-four  hours  oscillated  between 
1 20  and  160  c.c,  but  it  might  be,  exceptionally,  25  and  even  15  cc, 
or  in  another  case,  215  or  255  cc.  for  twenty- four  hours.  The 
density  was  very  far  from  constant;  generally  it  oscillated  between 
1.018  and  1.03Q,  going  sometimes  as  high  as  1.035  ^^^  1-051. 
Among  the  abnormal  constituents  of  the  urine  the  presence  of  albu- 
min only  has  been  looked  for.  In  some  cases  there  was  a  little 
albumin  during  the  first  days,  ranging  from  0.50  to  0.25  for  1,000 
cc.  In  other  cases  the  albumin  disappeared  about  one  week  after 
the  operation. 

The  general  condition  of  the  animal  can  be  used,  in  some  meas- 
ure, to  indicate  the  perfection  of  the  urinary  elimination.  As  long 
as  no  complications  were  present  the  animals  lived  as  normal  cats 
do,  without  presenting  any  symptoms  which  could  be  considered  as 
produced  by  renal  insufficiency.  When  general  complications  oc- 
curred the  cats  reacted  against  them  in  normal  ways.  In  one  case, 
the  animal  was  in  apparently  normal  condition  four  days  after  the 
operation.  She  walked  about  the  room,  played  and  ate  a  great  deal 
of  raw  meat.  Her  condition  remained  excellent  for  several  weeks. 
Twenty  days  after  the  operation  she  was  in  good  health,  had  glossy 
hair,  was  very  fat,  ate  with  appetite  all  kinds  of  food  and  urinated 
normally.  There  was,  however,  albumin  in  the  urine,  and  slow  and 
progressive  enlargement  of  the  kidneys  took  place,  which  showed 
that  she  was  not  in  an  entirely  normal  condition.  It  remained  in 
excellent  health  until  the  twenty-ninth  day  after  the  operation. 
Then  gastro-intestinal  symptoms  appeared,  and  death  occurred  on 
the  thirty-first  day  after  the  operation. 

PROC.  AMER.  PHIL.  SOC,  XLVII.  I90  RR,  PRINTED  FEBRUARY  9,  I909. 


682  CARREI^FURTHER    STUDIES    ON  [November  6, 

In  another  experiment  the  animal  was  a  female  cat  which  had 
lived  in  the  laboratory  for  several  months.  She  was  in  excellent 
condition  when  she  was  operated  on  and  recovered  very  quickly 
from  the  operation.  Her  life  went  on  just  the  same  as  before.  The 
kidneys  were  movable  and  small.  She  looked  in  excellent  health 
and  lived  as  a  normal  cat.  On  the  eighteenth  day  after  the  trans- 
plantation albumin  appeared  in  the  urine  and  a  direct  examination 
of  the  kidneys  was  made  to  ascertain  the  cause.  The  general  con- 
dition was  little  affected  by  the  operation  and  the  albumin  disap- 
peared on  the  twenty-first  day,  but  reappeared  again  a  little  later. 
On  the  thirty-fifth  day  the  animal  was  very  weak  and  emaciated. 
She  died  on  the  thirty-sixth  day  of  acute  calcification  of  the  arteries. 

These  results  show  that  the  functions  of  the  kidneys  reestab- 
lished themselves  after  the  transplantation.  Since  an  animal  can 
live  in  an  apparently  prosperous  condition  of  health  fifteen  or 
twenty-five  days  and  more,  after  a  double  nephrectomy,  and  elim- 
inate each  twenty- four  hours  from  120  to  160  c.c.  of  urine  through 
the  new  kidneys,  it  is  certain  that  the  functions  of  the  transplanted 
organs  are  efficient. 

The  "  simple  transplantation  "  of  the  kidneys  consists  of  dissect- 
ing a  kidney,  cutting  the  renal  vessels  and  ureter  a  few  centimeters 
below  the  hilus,  implanting  the  organ  on  the  same  or  another  ani- 
mal, and  of  anastomosing  its  vessels  to  the  renal  vessels  of  the  host 
I  performed  the  double  nephrectomy  and  the  replantation  of  one 
kidney  in  five  dogs.  The  secretion  of  the  urine  remained  normal 
as  long  as  no  ureteral  complication  occurred.  The  conditions  of  the 
kidneys  were  excellent.  A  little  more  than  two  months  after  the 
operation,  the  location  of  the  anastomoses  of  the  renal  vein  could 
not  be  detected.  The  anastomosis  of  the  renal  artery  was  seen  as  a 
small  and  indistinct  line  on  the  intima. 

The  remote  results  of  this  operation  are  excellent.  On  February 
6,  1908,  the  left  kidney  of  a  middle-sized  bitch  was  extirpated,  per- 
fused with  Locke's  solution  and  put  into  a  jar  of  Locke's  solution 
at  the  temperature  of  the  laboratory.  The  ends  of  the  vessel  were 
prepared  for  anastomoses,  and  afterward  the  kidney  was  replaced 
into  the  abdominal  cavity.  The  circulation  was  reestablished  after 
suture  of  the  vessels  and  the  ends  of  the  ureter  united.    The  animal 


ijoB.]  TRANSPLANTATION   OF  VESSELS   AND  ORGANS.  683 

made  an  uneventful  recovery.  Fifteen  days  afterward  the  right 
kidney  was  extirpated.  The  animal  remained  in  perfect  health. 
The  urine  did  not  contain  any  albumin.  It  is  generally  of  low 
density.     Today  the  animal  is  in  perfect  condition.     (Fig.  i.) 


Fia  I.     The  dog,  who  is  jumping,  underwent  nine  months  ago  a  double 
nephrectomy  and  replantation  of  one  kidney. 

This  observation  demonstrated  definitely  that  an  animal  can  live 
in  normal  condition  after  both  kidneys  have  been  extirpated  and  one 
replaced.  It  removes  also,  without  need  of  further  discussion,  the 
objections  of  the  experimenters  who  claim  that  the  section  of  the 
renal  nerves,  the  temporary  suppression  of  the  renal  circulation  or 
the  perfusion  of  the  kidneys  produce  necessarily  dangerous  and  even 
fatal  lesions  of  this  organ. 


(>84  CARREL— FURTHER  STUDIES  ON  [November  6, 

By  using  the  method  of  transplantation  in  mass  it  becomes  pos- 
sible to  perform  the  transplantation  of  a  whole  anatomic  region, 
with  its  main  artery  and  vein.  From  a  first  dog,  the  right  part  of 
the  scalp  and  the  auricle  were  extirpated  in  one  mass  wtih  the  car- 
tilaginous portion  of  the  auditory  canal  cut  close  to  the  skull,  the 
connective  tissue  and  the  glands  of  the  retro-maxillaris  space,  the 
tissues  of  the  carotid  region,  and  the  upper  portions  of  the  external 
jugular  vein  and  of  the  common  carotid  artery.  On  a  second  dog 
the  auricle  and  a  portion  of  the  scalp  was  extirpated  and  the  right 
part  of  the  neck  opened  through  a  longitudinal  incision.  The  ana- 
tomic specimen  was  then  placed  close  to  the  wound,  and  the  periph- 
eral end  of  the  carotid  artery  and  of  the  jugular  vein  united  to  the 
central  end  of  the  corresponding  vessels  of  the  host,  at  the  level  of 
the  middle  part  of  the  neck.  The  circulation  was  then  reestablished. 
Then  the  neck  was  closed  by  two  rows  of  suture.  A  few  minutes 
after  the  establishment  of  the  circulation  the  ear  and  the  scalp 
assumed  their  normal  appearance.  The  new  ear  was  fixed  by  cir- 
cular suture  of  its  cartilaginous  canal  to  the  cartilaginous  canal  of 
the  host.  The  auricular  muscles  were  sutured  and  the  operation 
completed  by  continuous  catgut  suture  of  the  skin  without  drainage. 

Three  weeks  after  the  operation  the  auricle  and  the  transplanted 
tissues  were  in  normal  condition.  The  temperature  of  both  auricles, 
normal  and  transplanted,  )vere  about  the  same.  The  transplanted 
ear  was  as  thin  and  glossy  as  the  normal  one.  Except  for  the  dif- 
ference of  color,  it  could  not  have  been  seen  that  the  ear  did  not 
belong  to  the  dog. 

The  transplantation  of  a  limb  from  one  animal  to  another  of  the 
same  species  is  a  problem  very  much  simpler  than  the  transplanta- 
tion of  a  gland.  In  April,  1907,  I  found  that  a  thigh,  extirpated 
from  the  fresh  cadaver  of  a  dog,  and  transplanted  onto  another  dog, 
could  begin  to  heal  in  a  very  satisfactory  manner.  One  year  after, 
by  using  more  careful  asepsis  in  the  transplantation  of  the  1^  from 
one  fox  terrier  to  another,  I  observed  union  by  first  intention  of  the 
new  leg  to  its  host. 

A  white,  middle-aged  male  fox  terrier  was  etherized  and  the  left 
leg  cut  just  below  the  knee.  The  limb  was  perfused  with  Locke's 
solution,  wrapped  in  a  greased  silk  towel  and  kept  on  a  table  at  the 


,9o8]  TRANSPLANTATION   OF  VESSELS  AND  ORGANS.  686 

temperature  of  the  laboratory.  A  white,  young  female  fox  terrier 
was  etherized.  She  was  of  the  same  size  and  shape  as  the  first  dog. 
Her  nails  and  bones  were  very  slightly  smaller.  The  leg  was  ampu- 
tated circularly  just  below  the  knee.  The  new  leg  was  immediately 
fixed  to  the  central  end  of  the  tibia  of  the  host  by  an  Elsberg's  alu- 
minum splint.  The  muscles,  nerves  and  femoral  vessels  were  united 
to  the  corresponding  parts  of  the  host,  and  the  circulation  reestab- 
lished. A  small  exploratory  incision  was  made  between  the  second 
and  third  toes.  Hemorrhage  of  red  blood  occurred.  The  animal 
recovered  quickly  and  remained  in  normal  condition.  The  tempera- 
ture of  the  new  foot  was  at  first  higher  than  that  of  the  normal  one. 
It  was  a;lso  edematous.  After  a  few  days  the  edema  disappeared 
and  the  foot  had  exactly  the  same  appearance  as  the  normal  one. 
The  temperature  went  slightly  down.  There  was  only  a  difference 
of  one  tenth  of  a  degree  centigrade  between  the  normal  and  the 
new  foot. 

Fifteen  days  after  the  operation  the  new  leg  was  perfectly  healed 
by  first  intention,  but  the  bones  were  not  very  strongly  united.  The 
Elsberg  splint  had  broken  and  the  tibia  was  a  little  incurved.  The 
exploratory  incision  of  the  foot,  although  having  been  slightly  in- 
fected, was  completely  cicatrized.  The  new  leg  had  the  same 
appearance  as  the  normal  one.  The  animal  was  in  good  condition, 
but  coughed  a  little.  At  this  time  several  other  dogs  died  of 
broncho-pneumonia.  The  animal  became  sick.  Twenty  days  after 
the  operation  her  condition  became  worse  and  a  marked  dyspnea 
appeared.  The  dog  died  on  the  twenty-second  day  after  the  opera- 
tion. Postmortem  examination  showed  a  double  diffuse  broncho- 
pneumonia. The  new  leg  was  perfectly  healed ;  with  linear  cutane- 
ous scars.  Its  appearance  was  exactly  the  same  as  the  normal  leg. 
The  bones  were  strongly  united  by  a  fibrous  callus.  The  explora- 
tory incision  of  the  foot  had  liealed  without  visible  scar. 

This  experiment  is  the  first  example  of  successful  grafting  of 
a  new  limb  on  an  animal.  It  demonstrates  that  the  leg,  in  spite  of 
the  change  of  owner,  remains  normal.  If  further  experiments  show 
that  the  functions  of  the  transplanted  limb  are  normally  reestab- 
lished, it  will  be  permissible  to  try  on  man  the  transplantation  of 


686  CARREL— FURTHER    STUDIES    ON  [November  6. 

limbs,  or  segments  of  limbs,  taken  from  an  amputated  limb,  or  from 
the  body  of  a  man  killed  by  accident. 

All  these  experiments  show  that  the  remote  results  of  the  trans- 
plantation of  fresh  vessels  can  be  perfect,  that  transplanted  kidneys 
functionate,  that  an  animal  having  undergone  a  double  nephrectomy 
and  the  transplantation  of  both  kidneys  from  another  animal  can 
live  normally  for  a  few  weeks,  and  that  an  animal  which  has  under- 
gone a  double  nephrectomy  and  the  graft  of  one  of  his  own  kidneys 
can  recover  completely  and  live  in  perfect  health.  Finally,  it  has 
been  demonstrated  that  a  leg  extirpated  from  a  dog  and  substituted 
for  the  corresponding  1^  of  another  dog  heals  normally. 

Since  the  experimental  transplantation  of  arteries  are  perma- 
nently successful,  it  is  permissible  to  use  this  method  in  human  sur- 
gery ;  for  instance,  in  treating  aneurisms  as  it  has  been  already  tried 
by  Delbet  in  Paris.  The  era  of  these  operations  being  opened,  the 
attempt  of  preserving  blood  vessels  outside  of  the  body  in  a  condi- 
tion of  latent  life  was  made  with  the  view  of  rendering  these  opera- 
tions more  practicable. 

The  length  of  the  period  which  elapses  between  the  extirpation 
of  a  tissue,  and  the  reestablishment  of  its  circulation  after  trans- 
plantation, is  an  important  factor  of  success  or  failure.  The  result 
of  the  graft  depends  entirely  on  the  condition  of  the  tissues  at  the 
time  of  the  reestablishment  of  the  circulation.  They  must  still  be 
alive;  although  apparently  dead.  If  the  tissues  are  really  dead,  the 
graft  is  completely  unsuccessful.  There  are  two  kinds  of  death, 
general  death  or  death  of  the  whole  organism,  and  elemental  death 
or  death  of  the  tissues  and  organs.  It  is  impossible  to  give  a  defi- 
nition of  general  death.  Everybody  understands  what  it  means. 
Nevertheless,  we  are  as  ignorant  about  it  as  about  life.  General 
death  can  occur  suddenly,  while  elemental  death  is  a  slow  process. 
A  man,  for  instance,  is  stabbed  through  the  heart  and  killed.  His 
personality  has  disappeared.  He  is  dead.  However,  all  the  organs 
and  tissues,  which  compose  the  body,  are  still  living.  The  life  of 
every  tissue  and  organ  of  the  body  could  go  on  if  a  proper  circula- 
tion was  given  back  to  them.  If  it  were  possible  to  transplant  imme- 
diately after  death  the  tissues  and  organs,  which  compose  this  body, 
into  other  human  organisms,  no  elemental  death  would  occur,  and 


X908J  TRANSPLANTATION  OF  VESSELS  AND  ORGANS.  687 

all  the  constituent  parts  of  the  body  would  continue  to  live.  The 
man,  however,  would  be  dead,  for  his  personality  would  have  dis- 
appeared. In  this  case,  general  death  can  be  defined  as  the  rupture 
of  the  contract  of  association  between  the  tissues  and  organs  of  the 
organism  by  failure  of  one  of  the  partners,  the  heart  Therefore, 
general  death  is  very  diflferent  from  elemental  death.  It  is  merely 
the  starting  point  of  the  disintegrative  phenomena  which  lead  to 
elemental  death. 

Immediately  after  general  death,  elemental  death  begins.  It  is 
a  complex  and  slow  process  which  progressively  destroys  the  living 
matter.  We  cannot  know  directly  whether  or  not  a  tissue  is  living 
and  by  what  chemical  or  physical  peculiarities  a  living  being  differs 
from  its  corpse.  There  is  no  reagent  of  life.  Living  matter,  in  a 
condition  of  non-manifested  life,  is  apparently  similar  to  non-living 
matter.  We  perceive  life  only  through  its  manifestations.  Our 
ignorance  renders  for  us  unmanifested  life  similar  to  death.  If 
seeds  or  microbes  are  placed  in  physico-chemical  conditions,  where 
manifested  life  is  impossible,  living  matter  canot  be  distinguished 
from  dead  matter.  What  is  the  difference  between  a  dead  seed  and 
the  seed  which  will  produce  a  large  tree?  We  do  not  know.  Be- 
tween a  vessel  which  will  live  normally  after  transplantation,  and 
another  one  which  will  undergo  deep  microscopical  lesions,  there  is 
no  morphological  difference.  We  know  merely  that,  immediately 
after  general  death,  the  tissues  are  still  alive,  because  they  manifest 
life  if  they  are  given  back  their  normal  circulation.  We  know  ako 
that  some  time  after  general  death  they  die,  because  they  are  not 
able  to  manifest  life  again,  even  when  replaced  in  normal  physio- 
logical condition.  Between  the  death  of  the  organism  and  the  ele- 
mental death  there  is  a  period  where  the  tissues  are  progressively 
invaded  by  cadaveric  disintegration.  At  the  beginning,  the  cadaveric 
changes  are  slight,  and  the  tissues  can  recover  if  placed  back  into 
normal  condition.  Later,  irreversible  changes  take  place  and  ele- 
mental death,  that  is,  destruction  of  the  living  matter,  occurs. 

The  duration  of  this  period  intermediate  between  death  of  the 
organism  and  elemental  death  is  longer  or  shorter,  according  to  the 
nature  of  the  tissue.  The  cerebral  substance  disintegrates  so  quickly 
that,  after  a  few  minutes  of  complete  anemia,  irreparable  lesions 


688  CARREL-FURTHER   STUDIES  ON  [November  6, 

take  place.  The  spleen,  liver  and  kidneys  are  also  rapidly  destroyed. 
On  the  contrary,  the  anatomical  structures  which  compose  a  limb 
are  very  strong  and  can  overcome  for  a  long  time  the  cadaveric 
processes.  The  different  parts  of  the  same  organ  do  not  present 
similar  resistancy  to  cadaveric  disintegration.  Among  the  anatom- 
ical components  of  renal  substance,  the  cells  of  the  secretory  tubules 
are  extremely  delicate  and  may  present  marked  morphological 
changes  a  short  time  after  death.  The  celk  of  the  excretory  tubuli 
are  stronger.  The  glomeruli  are  still  more  resistant.  It  may  hap- 
pen that  the  epithelial  cells  are  already  dead,  while  the  glomeruli 
and  the  vessels  are  still  living.  The  vascular  endothelium  seems  to 
be  the  "  ultimum  moriens  "  of  the  organ,  according  to  Wells.  The 
vessels,  which  are  the  necessary  condition  of  life  of  organs,  are  also 
the  part  of  the  organs  which  resists  longer  the  disintegrative  proc- 
esses. The  elements  which  compose  the  wall  of  an  artery  differ 
widely  in  resistancy.  The  muscular  fibers  die  first.  Immediately 
after  the  stopping  of  the  circulation,  all  the  elements  of  the  vascular 
wall  are  alive.  If  the  transplantation  is  performed  at  this  moment, 
the  artery  lives  in  the  body  of  its  host  and  keeps  its  normal  consti- 
tution. If  the  transplantation  is  performed  a  little  later,  when  the 
muscular  fibers  are  already  dead,  the  wall  of  the  artery  will  be  com- 
posed mainly  of  connective  and  elastic  tissue,  and  the  muscular  fibers 
will  disappear.  If  the  artery  is  completely  dead  when  the  trans- 
plantation is  made,  its  wall  will  be  composed  of  amorphous  sub- 
stance, around  which  the  organism  will  create  an  envelope  of  dense 
connective  tissue. 

Elemental  death  is  brought  about  by  microbian  and  autolytic 
enzymes.  Immediately  after  general  death,  the  microorganisms 
from  the  digestive  tract  diffuse  through  the  body  and  their  ferments 
begin  to  destroy  the  tissues.  At  the  same  time,  the  autolytic  fer- 
ments, which  are  not  any  longer  held  in  check  by  the  serum,  "con- 
tribute also  to  the  disintegration  of  the  organs.  This  destructive 
process  is  increased  or  retarded  by  the  causes  which  activate  or 
retard  the  enzymotic  actions,  and  the  multiplication  of  the  micro- 
organisms. For  instance,  the  rate  of  cadaveric  disint^ration,  which 
is  very  rapid  at  35®  or  40**  C,  becomes  very  slow  at  +  i®  or  -f-  2**  C. 
It  is  completely  stopped  by  desiccation  of  the  tissues.    The  preser- 


i9o8.]  TRANSPLANTATION  OF  VESSELS  AND  ORGANS.  689 

vation  of  the  tissues  in  the  serum  of  the  same  animal  will  also  retard 
very  much  the  organic  destruction. 

The  occurrence  o#  cadaveric  changes  in  tissues,  which  will  be 
used  for  transplantation,  must  be  prevented.  This  can  be  attained 
in  two  different  manners:  by  stopping  completely  the  chemical 
activities  of  the  tissue,  or  merely  by  retarding  so  much  the  evolution 
of  autolytic  disintegration  that,  after  a  few  days  or  a  few  weeks, 
the  lesions  are  so  small  that  they  are  not  dangerous. 

The  first  method  would  be  ideal.  The  tissue,  being  in  a  condi- 
tion of  chemical  indifference,  could  be  preserved  theoretically  for  an 
indefinite  period.  There  are  many  instances  of  this  form  of  latent 
life  in  the  animal  kingdom.  Two  centuries  ago,  Loevenhoeck 
obtained  the  resurrection  of  Milnesium  tardigradum,  which  had  been 
completely  dried  for  a  long  time,  by  moistening  it  with  water.  In 
1840,  Doyere  studied  also  the  peculiarities  of  latent  life  of  Milnesium 
tardigradum.  He  dried  completely  a  few  of  these  animals,  heated 
them  at  a  temperature  of  100®  C,  and,  after  having  humidified 
them,  observed  that  they  lived  again.  These  observations  are  very 
important  because  Milnesium  tardigradum  is  highly  organized  and 
contains  muscular  fibers,  nerves,  nervous  ganglia,  etc.  Paul  Bert, 
in  several  famous  experiments,  attempted  to  preserve  tissues  of 
mammals  in  a  condition  of  latent  life.  One  of  those  experiments 
consisted  of  cutting  the  tail  of  a  rat,  drying  it  in  vaccum,  and  sub- 
mitting it  to  a  temperature  of  +  100**  C.  The  tail  was  afterwards 
transplanted  onto  another  rat.  It  was  observed  that  the  dimensions 
of  the  tail  grew  larger,  that  its  vessels  united  to  the  vessels  of  the 
host  and  that  the  bone  marrow  underwent  fibrous  degeneration.  It 
showed  that  the  heated  and  dried  tail  could  live  again.  I  attempted 
to  preserve  arteries  in  latent  life  by  a  similar  method.  Carotid 
arteries  from  dogs  were  extirpated  and  placed  in  sealed  glass  tubes, 
part  of  which  were  filled  with  calcium  chloride.  Within  a  few 
hours,  the  arteries  became  yellow  brown,  shrank  and  looked  like 
pieces  of  catgut.  One  tube  was  heated  for  twelve  minutes  at 
-f-  100**.  When,  after  several  days,  the  dried  vessels  were  put  into 
Locke's  solution,  they  took  back  their  water  and  assumed  again  their 
normal  color,  size  and  consistency.  Two  of  them  were  transplanted 
onto  the  carotid  arteries  of  dogs.     It  was  found  that  they  could 


690  CARREL— FURTHER   STUDIES  ON  [November  6, 

perform  normally  their  functions.  Two  weeks  after  the  operation, 
one  of  the  vessels  was  examined.  The  circulation  was  normal. 
The  transplanted  segment  looked  very  much  4ike  the  other  parts  of 
the  carotid.  It  was  covered  by  a  normal  connective  tissue  sheath. 
The  wall  was  of  same  color  and  thickness  as  the  wall  of  the  normal 
carotid.  Its  consistency  was  a  little  harder.  Nevertheless,  it  was 
found,  by  microscopical  examination,  that  this  wall  was  composed 
of  an  elastic  framework  and  amorphous  material  surrounded  by  a 
new  wall  of  connective  tissue.  The  vessel  was  dead.  The  death 
of  the  vessel  was  perhaps  due  more  to  the  way  in  which  the  desicca- 
tion was  done  than  to  the  desiccation  itself.  With  a  better  tech- 
nique, results  similar  to  those  of  Paul  Bert  could  possibly  be  ob- 
tained. Actually,  this  method  is  dangerous  because  the  artery  is 
not  any  longer  a  living  structure,  but  merely  a  foreign  body,  as  a 
piece  of  rubber  tubing  or  an  artery  preserved  in  formalin  or  killed 
by  heating. 

The  second  method  of  preserving  arteries,  outside  of  the  body, 
consists  in  lowering  the  power  of  the  microbian  and  autol)rtic  en- 
zymes, by  keeping  the  tissues  at  a  low  temperature.  This  method 
cannot  suspend,  for  an  indefinite  time,  the  occurrence  of  elemental 
death.  It  increases  only  the  length  of  the  period  during  which  the 
cadaveric  changes  are  slight  and  not  able  to  interfere  with  a  com- 
plete,  or  almost  complete,  recovery  of  the  artery  after  transplanta- 
tion. If  a  vessel  is  extirpated  aseptically,  placed  in  a  sterilized 
sealed  tube  and  kept  in  a  refrigerator  just  above  the  freezing  point, 
it  can  be  preserved  for  a  long  time  in  good  condition.  From  a 
surgical  standpoint,  it  is  sufficient  that  the  vessels  are  kept  safely 
for  a  few  days  outside  of  the  body  before  being  transplanted. 
Nevertheless,  it  is  far  from  perfect.  The  ideal  method  would  be 
certainly  to  place  the  tissues  in  a  condition  of  latent  life,  as  is  pos- 
sible for  Milnesiunt  tardigradum  and  other  organisms. 

The  technique  that  I  use  is  very  far  from  being  original.  The 
vessels  are  merely  preserved  in  cold  storage  as  are  commonly  eggs, 
or  chickens,  or  vegetables.  They  are  removed  from  a  living  or  a 
dead  animal  soon  after  death,  perfused  and  washed  with  Locke's 
solution  and  placed  in  sterilized  glass  tubes,  the  atmosphere  of  which 
is  moistened  with  a  few  drops  of  water.    The  tubes  are  immediately 


PnoctEDmas  Am.  Phium.  Soc.  Vol,  XLVII.  No.  190 


SegineTit  of  ulery  preserved  in  ■  sealed  sterilized  lube. 

Sqrment  of  artery  preserved  for  Iwenty  two  days  in  cold  storage.     Six  months 

after  Iransplantatron. 


I908.]  TRANSPLANTATION   OF  VESSELS  AND  ORGANS.  091 

sealed.  (Plate  VII,  Fig.  i.)  Sometimes,  the  arteries  are  put  in 
a  fluid.  A  few  vessels  have  been  preserved  in  isotonic  sodium 
chloride  solution.  The  result  was  unsatisfactory;  for  the  muscular 
fibers  of  the  artery  were  killed  in  twenty-four  hours.  The  results 
obtained  with  Locke's  solution  were  much  better.  However,  a  still 
better  method  would  consist  in  keeping  the  vessels  in  serum  of  an 
animal  of  the  same  species  or  in  inactivated  serum  of  an  animal  of 
different  species.  The  serum  is  more  exactly  isotonic  for  the  tis- 
sues than  Locke's  solution ;  it  is  slightly  bactericidal,  and  it  contains 
antibodies  for  the  autolytic  ferments  of  the  cells.  I  performed 
once  only  the  transplantation  of  a  segment  of  dog's  carotid,  pre- 
served in  dog's  serum  for  forty-eight  hours.  Fifteen  days  after 
the  transplantation,  the  vessel  was  examined  and  found  in  a  perfect 
microscopical  condition. 

The  sealed  tubes  containing  the  arterial  segments  are  put  into  a 
thick-walled  ice-box,  the  temperature  of  which  remains  constantly 
between  o  and  -}- 1**  C.  The  temperature  must  not  go  down  below 
o°  C.  When  the  vessels  have  been  frozen,  the  wall  presents  soon 
after  the  transplantation  marked  microscopical  lesions.  If  the  tem- 
perature is  too  high,  and  the  operation  not  thoroughly  aseptic, 
microbian  colonies  may  settle  in  the  wall  of  the  vessels.  Oblitera- 
tion or  <levelopment  of  fusiform  aneurism  are  the  consequence  of 
these  faults  of  technique.  When  the  operation  has  been  correctly 
performed,  the  artery  keeps  its  normal  appearance  for  a  long  time. 
After  several  weeks,  its  color  and  consistency  are  generally  normal. 
The  wall  is  a  little  softer  and  the  vessel  flattens  itself  more  easily. 
After  six,  seven  and  even  ten  months,  the  macroscopical  appearance 
of  the  vessel  is  not  markedly  modified.  Sometimes  it  looks  com- 
pletely normal.  From  a  microscopical  standpoint,  the  condition  of 
the  arteries  is  very  variable.  In  some  cases,  the  nuclei  of  the  mus- 
cular fibers  are  modified.  In  other  cases  they  are  absolutely  normal. 
A  section  of  a  pig's  carotid  artery,  preserved  in  a  sealed  tube  with 
a  few  drops  of  Locke's  solution  from  April  to  November,  1908,  was 
entirely  normal.  It  looked  as  if  it  had  been  extirpated  from  the 
animal  a  few  moments  before  being  fixed  in  Zenker's  fluid,  while  it 
had  been  preserved  for  six  months  outside  of  the  body. 

A  few  minutes  before  the  transplantation,  the  tube  is  removed 


692  CARREL— FURTHER    STUDIES    ON  [November  6. 

* 

from  the  ice-box  and  broken.  The  vessel  is  removed  from  the  tube, 
put  in  a  jar  of  Locke's  solution  at  the  temperature  of  the  laboratory, 
thoroughly  washed  and  placed  in  warm  vaseline.  Afterward,  the 
vaseline  is  expressed  from  its  lumen,  and  the  segment  grafted  onto 
the  artery  of  the  host.  As  soon  as  the  circulation  is  established 
through  the  artery  of  the  host,  the  transplanted  s^ment,  which  is 
white,  takes  back  immediately  its  normal  color  and  becomes  almost 
similar  to  the  other  parts  of  the  artery.  Sometimes  the  small  ves- 
sels of  the  adventitia  appear  neatly  injected  with  Jjlood.  In  seg- 
ments of  carotid  artery,  preserved  for  eight  and  eleven  months  in 
cold  storage  and  grafted  on  the  carotid  of  a  dog,  the  vasa  vasorum 
were  seen  full  of  blood  as  soon  as  the  circulation  was  reestablished. 
The  results  of  the  transplantation  of  arteries,  preserved  in  cold 
storage,  are  generally  excellent  from  a  functional  standpoint,  even 
if  the  vessel  has  been  kept  for  one  or  two  months  outside  of  the 
body.  But,  from  an  anatomical  standpoint,  the  microscopical  con- 
stitution of  the  vessel  is  markedly  modified  when  it  has  spent  a  long 
time  in  cold  storage.  The  duration  of  the  period  during  which  a 
vessel  can  be  preserved  without  occurrence  of  any  lesion,  is  not 
exactly  determined.  However,  it  seems  that  an  artery,  preserved 
for  more  than  eight  days  in  cold  storage,  undergoes  always,  after 
transplantation,  a  degeneration  of  its  muscular  fibers,  while  the 
other  parts  of  the  vessel  seem  to  remain  normal.  Several  times  a 
perfect  histological  condition  of  the  transplanted  artery  was  ob- 
served. A  piece  of  carotid  artery  from  a  dog  was  put  in  a  sealed 
tube  with  a  few  drops  of  Locke's  solution  and,  two  days  afterward, 
transplanted  onto  the  carotid  artery  of  another  dog.  Two  weeks 
after  the  operation,  the  neck  of  the  dog  was  reopened.  The  circu- 
lation through  the  carotid  was  normal.  The  transplanted  segment 
looked  like  the  other  parts  of  the  carotid.  It  was  resected  and 
examined  histologically.  The  adventitia  was  thickened  and  con- 
tained several  small  vessels.  The  media  was  normal.  The  nuclei 
of  the  muscular  fibers  were  found  entirely  similar  to  those  of  a  nor- 
mal artery.  The  intima  was  well  preserved  and  slightly  thickened. 
This  observation  shows,  evidently,  that  a  vessel  can  be  preserevd  in 
cold  storage  and  live  again  normally  when  transplanted.  It  is  not 
a  dead,  but  a  living  artery,  with  all  its  normal  anatomical  elements. 


i9o8.]  TRANSPLANTATION  OF  VESSELS  AND  ORGAl>fs.  693 

Thus,  the  vessel,  while  in  cold  storage,  was  in  a  condition  of  unmani- 
fested  life. 

The  behavior  of  a  vessel,  transplanted  after  having  been  killed, 
by  formalin  or  by  heating  at  80®  C,  is  different.  Often  its  appear- 
ance is  normal,  from  a  gross  anatomical  standpoint.  Nevertheless, 
a  few  days  after  transplantation,  its  microscopical  constitution  is 
deeply  modified.  Its  wall  is  composed  of  an  amorphous  material 
where  no  nuclei  can  be  observed,  but  where  the  elastic  framework 
still  is  visible,  although  very  modified  in  its  shape.  The  wall  is 
surrounded  by  a  layer  of  connective  tissue  produced  doubtless  by 
the  host.  A  dead  vessel  is  merely  a  foreign  body,  which  would  pro- 
gressively be  resorbed  and  replaced  by  connective  tissue.  Throm- 
bosis frequently  occurs  after  this  kind  of  transplantation  and  its 
use  is  dangerous  from  a  clinical  standpoint.  On  the  contrary,  a 
vessel,  preserved  for  a  few  days  in  a  condition  of  latent  life,  is  still 
a  living  structure  when  it  is  transplanted.  Its  use  is  as  safe  as  that 
of  a  fresh  artery. 

In  all  the  cases  where  the  vessels  spent  more  than  eight  days  in 
the  ice-box,  the  muscular  fibers  of  the  media  disappeared  a  few 
days  after  transplantation.  Nevertheless,  the  anatomical  results 
were  often  so  perfect  that,  after  a  few  months,  the  location  of  the 
transplanted  segment  on  the  artery  of  the  host  was  hardly  discerni- 
ble. On  April  2,  1908,  a  piece  of  carotid,  preserved  for  twenty-two 
days  in  cold  storage,  was  transplanted  on  the  carotid  of  a  dog.  On 
October  15,  1908,  the  neck  was  opened  and  the  carotid  dissected. 
It  was  not  possible  to  find  the  location  of  the  transplanted  segment. 
After  longitudinal  opening  of  the  carotids,  the  location  of  the  anas- 
tomoses could  be  determined.  (Plate  VII,  Fig.  2.)  The  result  of 
the  graft  of  a  vessel  which  had  spent  seventy  days  in  cold  storage 
was  as  satisfactory.  Six  months  after  the  operation  a  section  was 
made  through  the  middle  part  of  the  transplanted  segment.  The 
adventitia  was  normal  and  the  intima  thickened.  The  media  was 
composed  of  elastic  fibers  which  had  retained  their  ordinary  wavy 
appearance.     All  the  muscular  fibers  had  been  destroyed. 

The  actual  method  failed  to  give  positive  results  in  the  trans- 
plantation of  arteries  after  several  months  in  cold  storage.  Graft 
of  arteries  which  had  spent  eight  months  outside  of  the  body  was 


694  CARREL— FURTHER   STUDIES   ON  [N»«Bib«6, 

attempted  in  two  cases.  Thrombosis  occurred.  The  vessels  were 
dead,  and,  in  spite  of  their  almost  normal  appearance,  markedly 
disintegrated. 

The  remote  results  of  the  transplantations  of  preserved  vessels 
are  very  satisfactory   from  a  clinical  standpoint.     In   November, 


Ftc,  2.     Cat  in  which  a  seg:ment  of  the  abdominal  aorta  was  replaced  by  a 
piece  of  dog's  carotid. 

1906,  a  segment  of  the  abdominal  aorta  of  a  cat  was  extirpated  and 
replaced  by  a  piece  of  dog's  carotid  preserved  in  cold  storage  for 
twenty  days.  The  animal  remained  in  excellent  health.  After  a 
few  weeks,  the  abdomen  was  reopened  and  the  transplanted  artery 


ijBi.]  TRANSPLANTATION  OF  VESSELS   AND  ORGANS.  695 

examined.  The  circulation  through  the  new  artery  was  excellent, 
and  its  caliber  normal.  The  abdomen  was  closed.  The  cat  spent 
the  years  1907  and  1908  at  the  Rockefeller  Institute  in  excellent 
health.  The  femoral  pulse  was  normal.  The  condition  of  the 
femoral  pulse  is  an  indication  of  the  condition  of  the  circulation 
through  the  abdominal  aorta.  Partial  or  complete  occlusion  of  the 
aorta  produces  diminution  or  disappearance  of  the  pulse  of  the 
femoral  arteries.  To-day,  twenty-five  months  have  elapsed  since  the 
operation,  the  cat  is  in  good  condition  (Fig.  2)  and  the  femoral 
pulse  normal' 


Fig.  3.     Dog  in  which  a  segment  of  the  abdominal  aorta  was  replaced  by  a 
piece  of  human  popliteal  artery. 

In  May,  1907,  a  short  portion  of  the  abdominal  aorta  of  a  small 
bitch  was  extirpated.  Between  its  cut  ends  was  grafted  a  segment 
of  popliteal  artery  from  a  young  man's  leg  amputated  at  the  Pres- 
byterian Hospital  by  Dr.  Ellsworth  Eliot.  Before  being  trans- 
planted, the  popliteal  artery  had  been  preserved   for  twenty-four 

'This  cat  was  presented  before  the  American  Physiological  Society,  De- 
cember  igo6. 


696  CARREL— TRANSPLANTATION   OF  ORGANS.        [November  6, 

days  in  cold  storage.  The  femoral  pulse  remained  normal.  A  few 
months  after  the  operation,  the  abdomen  was  reopened  and  the  cir- 
culation through  the  new  artery  found  normal.  There  was  no  modi- 
fication of  its  caliber.  The  animal  remained  in  good  health.  Dur- 
ing the  years  1907  and  1908,  no  modification  of  the  femoral  pulse 
occurred.  It  is  still  normal  to-day,  one  year  and  a  half  after  the 
operation,  and  the  animal  is  in  excellent  condition  (Fig.  3). 

These  experiments  demonstrate  that  the  clinical  results  of  the 
transplantation  of  preserved  vessels  can  remain  satisfactory  for  a 
long  time.  However,  in  both  cases,  the  operation  was  performed 
under  unfavorable  circumstances.  The  grafted  arteries  belonged  to 
an  animal  of  different  species  and  the  method  of  preservation  used 
in  both  cases  was  imperfect.  The  wall  of  these  vessels  underwent 
certainly  marked  histological  changes.  Nevertheless  they  are  still 
able  to  perform  normally  their  functions. 

Conclusions. 

The  results  of  the  experiments  of  preservation  of  arteries  in  cold 
storage  must  be  considered  from  both  the  anatomical  and  the  prac- 
tical standpoint. 

From  an  anatomical  standpoint,  they  show  that  an  artery  from 
an  animal  can  be  kept  outside  of  the  body  for  two  days  at  least, 
transplanted  onto  another  animal  of  the  same  species,  and  live  again 
without  presenting  any  change  of  its  constituent  elements.  The 
transplantation  of  vessels  killed  by  drying,  heating  or  fixation  in 
formalin  is  followed  by  degeneration  of  the  wall  and  replacement 
by  connective  tissue  from  the  host.  When  the  vessel  is  kept  in  cold 
storage  for  a  longer  period  of  time,  all  the  muscular  fibers  of  the 
media  disappear  a  few  days  after  transplantation.  If  the  period 
spent  in  cold  storage  is  still  longer,  eight  months  for  instance,  throm- 
bosis occurs. 

From  a  practical  standpoint,  these  experiments  demonstrate  that 
the  preserved  vessels,  even  if  their  muscular  fibers  are  completely 
resorbed,  are  an  excellent  substitute  for  arteries  and  perform  nor- 
mally their  functions  for  months  and  years. 


,9o8.]  MINUTES.  697 

Stated  Meeting  November  20,  1908.  . 

President  Keen  in  the  Chair. 

The  decease  was  announced  of  Prof.  William  Keith  Brooks,  at 
Baltimore,  on  November  12,  1908,  set.  60. 

The  following  papers  were  read : 

"  The  Early  History  of  the  American  Philosophical  Society,"  by 
Mr.  Joseph  G.  Rosengarten. 

"The  Recapitulation  Theory  of  Embryologists,"  by  Prof. 
Thomas  H.  Montgomery,  Jr.,  which  was  discussed  by  President 
Keen,  Prof.  Kraemer,  Prof.  Doolittle,  Prof.  Pratt  and  Prof. 
Montgomery. 

Stated  Meeting  December  4,  1908, 

President  Keen  in  the  Chair. 

Prof.  Edwin  G.  Conklin  read  an  obituary  notice  of  Prof.  William 
Keith  Brooks  (see  page  iii). 

The  following  papers  were  read : 

"  Astrology  in  Ancient  Babylonia,"  by  Prof.  Morris  Jastrow,  Jr. 
(see  page  646),  which  was  discussed  by  President  Keen  and  Mr. 
Goodwin. 

"  On  the  Effect  of  a  Radio-Active  Mineral  on  Plant  Growth," 
by  Mr.  Joseph  Willcox. 

Dr.  John  L.  Shober  exhibited  some  photographs  made  by  radia- 
tions from  radium  and  uraninite. 

Stated  Meeting  December  18,  1908, 
President   Keen   in  the   Chair. 

Professor  Herbert  Weir  Smyth,  a  newly-elected  member,  was 
presented  to  the  chair  and  took  his  seat  in  the  Society. 

A  letter  was  read  from  the  Geological  Society  of  Glasgow  stating 
that  it  would  celebrate  its  jubilee  on  January  28,  1909,  and  inviting 
the  Society  to  be  represented  thereat.  The  invitation  was  accepted 
and  Sir  William  Turner,  K.C.B.,  was  appointed  the  Society's 
representative. 

PROC.  AMER.  PHIL.  SOC.,  XLVII.  I90  SS,  PRINTED  FEBRUARY  8,  I909. 


698  MINUTES.  [December  ,8, 

The  decease  was  announced  of 
Dr.  Ernest  T.  Hamy,  at  Paris,  on  November  i8,  1908,  act.  65. 
Prof.  Oliver  Wolcott  Gibbs,  at  Newport,  R.  I.,  on  December 

9,  1908,  set  86. 
Professor  Herbert  Weir   Smyth   read  a  paper  on   "Ancient 
Greek  Conceptions  of  the  Future  Life  "  which  was  discussed  by 
Professor  Lamberton,  Professor  Newbold  and  Professor  Smyth. 

Special  Meeting  December  21,  ipo8. 

President  Keen  in  the  Chair. 

The  President  introduced  Professor  Gugliebno  Ferrero,  who 
read  a  paper  on  "Antony  and  Cleopatra." 


INDEX. 


Absorption  spectra  of  solutions,  17, 
276 

Alaska  Boundary,  15,  87 

,  University  of  Pennsylvania  ex- 
pedition to,  1907,  13 

Algebraic  equations  in  infinite  series, 
16,  III 

American  Institute  of  Electrical  En- 
gineers, invitation  from,  2 

American  Philosophical  Society,  early 
history  of,  697 

Andaman  Islander,  train  of,  14,  51 

Anderson  and  Jones,  absorption 
spectra  of  solutions,  17,  276 

Antony  and  Cleopatra,  698 

Art  and  ethnology,  14,  30 

B 

Babylonia,  hepatoscopy  and  astrology 
in  ancient,  646,  697 

Balch,  E.  S.,  art  and  ethnology.  14, 30 

^  T.  W.,  law  of  Oresme,  Coper- 
nicus and  Gresham,  14,  18 

Barnard,  photographs  of  Daniel's 
Comet,  16 

Bates,  Greek  vases  in  the  Museum 
of  Science  and  Art,  University  of 
Pennsylvania,  17 

Bauer,  ocean  magnetic  work  of  the 
Carnegie  Institution,  16 

Bermuda  sand  dune  plants,  15,  97 

Bloomfield,  a  Vedic  Concordance,  17 

Bombsl^ell  ore,  135,  136 

Brain,  comparison  of  that  of  man 
with  albino  rat,  14 

Brains  of  natives  of  the  Andaman 
and  Nicobar  Islands,  14,  51 

Brashear,  astronomical  photography, 
16 

Brooks,  William  Keith,  obituary  no- 
tice of,  301,  iii 

Brown,  completion  of  the  lunar 
theory,  17 

Brown  and  Reichert',  crystallographic 
study  of  the  Hemoglobins,  14,  298 

C 

Carnegie,  delegate  to  American  In- 
stitute of  Electrical  Engineers,  2 


Carnegie  Institution,  ocean  magnetic 
work  of  the,  16 

Carrel,  further  studies  on  transplan- 
tation of  vessels  and  organs,  645, 

Cassandre  in  the  Oresteia  of  -^schy- 

lus,  14 
Cetacea,  classification  of  the,  15,  385 
Chance,  origin  of  bombshell  ore,  135, 

136 
Chilian  copper  minerals,  15,  7p 
College    of    Physicians,    laying    of 

conor-stone  of  new  building,  13, 14 
Congres  International  de  Botanique 

(3d),  invitation  from  the,  135 
Congress  of  chemistry  and  physics,  2 
Coniclin,    obituary    notice    of    Prof. 

William  Keith  Brooks,  iii 
Cytomorphosis,  14 


Daniel's  Comet,  16 

Darwin,  commemoration  of  the  cen- 
tenary of  the  birth  of,  644 

Davenport,  determination  of  domi- 
nance   in    Mendelian    inheritance, 

15,  59 
Death  penalty  by  electricity,  14,  39 

Descent,  Australian  laws  of,  134 

Donaldson,  comparison  of  the  albino 

rat  with  man  in  respect  to  brain 

and  spinal  cord,  14 
Doolittle,    Eric,    personal    error    in 

double  star  measures  which  depend 

on  position  angle,  16 
Double  star  measures,  16 

E 

Earth,  physics  of  the,  15,  157 
Election  of  members,  16 

Officers  and  Councillors,  i,  2 

Electricity,  death  penalty  by,  14,  39 
Ethnology,  art  and,  14,  30 
Excretory  organs  of  Metazoa,  15,  547 

F 

Ferrero,  Antony  and  Cleopatra,  698 
Francke,  mediaeval  German  sculpture 
in  the  Germanic  Museum  of  Har- 
vard University,  17,  636 
Fungi  of  Pennsylvania;   gasteromy- 
cetes,  15 


699 


700 


INDEX. 


Gasteromycetes ;  fungi  of  Pennsyl- 
vania, 15       . 

Geological  Society  of  Glasgow,  invi- 
tation to  jubilee  of,  697 

Goethe's  private  library  as  an  index 
of  his  literary  interests,  14 

Goodspeed  and  Richards,  recent  ad- 
vances in  color  photography,  12 

Gordon,  University  of  Pennsylvania 
expedition  to  Alaska,  1907,  13 

Greek  vases,  notes  on,  17 

H 

Hale,  telescopes  for  solar  research, 

Harshbcrger,  leaf  structure  of  the 
Bermuda  sand  dune  plants,  15,  97 

Hart,  artificial  refrigeration,  12 

Hartzell,  photographs  by  the  Lu- 
miere  process,  12 

Haupt,  lost  tribes  of  Israel  and  the 
Aryan  ancestry  of  Jesus  and  His 
first  disciples,  17 

Hemoglobins,  crystallographic  study 
of  the,  14,  298 

Hepatoscopy  and  astrology  in  Baby- 
lonia and  Assyria,  646,  697 

Heredity,  variation  and  evolution  in 
Protozoa,  15,  393 

Hewett,  Goethe  s  private  library  as 
an  index  of  his  literary  interests, 

U 

Holland,  delegate  to  College  of  Phy- 
sicians, 14 

Hovey,  contribution  to  history  of 
Mount  Pelee,  645 


Ingen,  stratigraphic  observations  in 
vicinity  of  Susquehanna  Gap,  15 

Inheritance,  Mendelian,  15,  59 

International  Archaeological  Con- 
gress (second),  13 

Congress      of      Mathematics 

(fourth).  12,  13 

Israel,  lost  tribes  of,  and  Aryan  an- 
cestry of  Jesus  and  His  first  disci- 
ples, 17 


Jastrow,  hepatoscopy  and  astrology 
in  Babylonia  and  Assyria,  646,  697 

sign    and   name   for   planet   in 

Babylon,  17,  141 

Jennings,    inheritance    in    Protozoa, 

15,  393 
Jones     and     Anderson,     absorption 

spectra  of  solutions,  17,  276 


Keller,  Chilian  copper  minerals,  15, 

79. 
Kelvin,  Lord,  memorial  in  honor  of,  2 

Kennelly  and  Upson,  the  humming 
telephone,  329,  644 

Kraemer,  influence  of  heat  and  chem- 
icals on  the  starch  grain,  15 


Lambert,  algebraic  equations  in  infi- 
nite series,  16,  iii 

Lamberton,  dramatic  function  of 
Cassandra  in  the  Oresteia  of 
yEschylus,  14 

Le^on  sur  Tintegration  des  equations, 
etc.,  12 

Life,  ancient  Greek  conceptions  of 
the  future,  697 

Loeb,  tumor  growth  and  tissue 
growth,  2,  3 

Lost  tribes  of  Israel  and  Aryan  an- 
cestry of  Jesus  and  His  first  disci- 
ples, 17 

Lovett,  integrable  oases  of  the  prob- 
lem of  those  bodies  in  which  the 
force  function  is  a  function  only 
of  the  mutual  distances,  12 

Legon    sur    Tintegration    des 

equations,  etc.,  12 

problems   of   three   bodies    on 

surfaces,  17 

Lumiere  process,  photographs  by  the, 

12 
Lunar  theory,  17 

M 

Mason,  explosion  of  the  Saratoga 
septic  tank,  14 

Mathews,  notes  on  Australian  laws 
of  descent,  134 

Mediaeval  German  sculpture  in  Mu- 
seum of  Harvard  University,  17 

Meeting,  General,  13 

Stated,    I,   2,    12,    13,   134,  644, 

645.  697,  698 

Members,  deceased: 

Becquerel,  Antoine  Henri,  644 
Brooks,  William  Keith,  697,  iii 
Qeveland,  Grover,  644 
Davenport,  Sir  Samuel,  13 
Gibbs,  Oliver  Wolcott,  698 
Gilman,  Daniel  Coit,  676 
Hamy,  Ernest  T.,  698 
Mascart,  E.,  644 
Mason,  Otis  T.,  645 
Meltzel,  Hugo  von,  644 
Rohrig,  F.  L.  Otto,  644 
Sellers,  Coleman,  i 


INDEX. 


701 


Members,  deceased — continued 
Seymour,  Thomas  Day,  i 
Spofford,  Ainsworth  Rand,  644 
Young,  Charles  Augustus,  2 

elected: 

Brumbauffh,    Martin    Grove,    16 
Cannon,  Walter  Bradford,  16 
Christie,  Jame?,  16 
Hallock,  William,  16 
Hopkins,  Edward  Washburn,  16 
Nys,  Ernest,  16 
Pearson,  Learned,  16 
Penck,  Albrecht  F.  K.,  16 
Royce,  Josiah,  16 
Schurman,  Jacob  G.,  16 
Smyth,  Charles  Henry,  16 
Smjrth,  Herbert  Weir,  16 
Spangler,  Henry  Wilson,  16 
Spitzka,  Edward  Anthony,  16 
Sterrett,  John  Robert  Sitlington, 

16 
Tucker,  Richard  Hawley,  16 
Wood,  Robert  Williams,  16 

presented,  134,  644,  697 

Membership  accepted,  134,  644 
Mendeleef,  memorial  in  honor  of,  2 
Mendelian  inheritance,  15,  59 
Metabolism,    effect    of    certain   pre- 
servatives upon,  17 
Metozoa,  excretory  organs  of  the,  15, 

Michelson,     elected     Vice-President, 

134,  135 
Milk,  production  and  distribution  of, 

Minot,  cytomorphosiff,  14 

Mont  Pelee,  contribution  to  history 

of.  645 
Montgomery,    excretory    organs    of 

the  Metazoa,  15,  547 
,  recapitulation  theory  of  embry- 

ologists,  697 
Moore,  a  living  representation  of  the 

ancestors  of  the  plant  kingdom,  17, 

91 

N 

Newcomb,  delegate  to  Fourth  Inter- 
national Congress  of  Mathematics, 

Nicobar  Islander,  bram  of,  14,  51 
Nipher,  effect  of  an  angle  in  a  wire 
conductor  in  spark  discharge,  17 


Oases,  integrable,  12 

Obituary   notice    of   William    Keith 

Brooks,  301,  iii 
Officers  and  Council,  election  of,  1,2 
Ore,  bombshell,  135,  136 


Oresme,    Copernicus    and    Gresham, 

law  of,  14,  18 
Organs,  transplantation  of,  677 
Osborn,   appointed   to   represent    So- 
ciety at  Cambridge,  celebration  of 
birth  of  Darwin,  644 


Pearson,  production  and  distribution 
of  milk,  13 

Photographs  by  the  Lumiere  process, 
12 

Photography,  astronomical,  16 

color,  recent  advances  in,  12 

Physico- Medical  Society  at  Erlangen 
sends  thanks  for  congratulatory 
address,  644 

Planet,  sign  and  name  for,  in  Baby- 
lon, 17,  141 

Plant  growth,  effect  of  a  radio-active 
mineral  on,  697 

kingdom,   representation  of  the 

ancestors  of  the,  17,  91 

Protozoa,  inheritance  in,  15,  393 

Preservatives,  influence  of,  upon 
health  and  metabolism,  302 

Pupin,  delegate  to  American  Insti- 
tute of  Electrical  Engineers,  2 

Pylc  and  Titchener,  after-images  of 
subliminally  colored  stimuli,  366, 
644 

R 

Rabies,  pathology  of,  14 
Radio-active    mineral,    effect    of,    on 

plant  growth,  697 
Rat,  albmo,  comparison  of,  with  man 

in  respect  to  brain  and  spinal  cprd, 

Ravenel,  pathology  of  rabies,  14 

Recapitulation  theory  of  embryolo- 
gists,  697 

Refrigeration,  artificial,  12 

Reichert  and  Brown,  crystallographic 
study  of  hemoglobins,  14,  2^ 

Rhinochimsera,  brain  of,  14,  37 

Richards  and  Goodspeed,  recent  ad- 
vances in  color  photography,  12 

Rosengarten,  early  history  of  Amer- 
ican Philosophical  Society.  697 


Sand  dune  plants  of  Bermuda,  15,  97 

Santa  Cruz  tjrpotheria,  15,  64 

See,  further  researches  on  the  physics 

of  the  earth,  15,  157 
Septic  tank,   explosion   of,  at   Sara- 
toga. 14 
Sinclair,  Santa  Cruz  typotheria,  15, 64 
Smyth,  ancient  Greek  conceptions  of 
t\\e  future  life,  697 


702 


INDEX. 


Spitzka,  brains  of  natives  of  the  An- 
daman and  Nicobar  Islands,  14,  51 

infliction  of  the  death  penalty 

by  electricity,  14,  39 

Starch  grain,  influence  of  heat  and 
chemicals  on  the,  15 

Stimuli,  after-images  of  subliminally 
colored,  366,  644 

Sumstine,  fungi  of  Pennsylvania; 
Gasteromycetes,  15 

Susquehanna  Gap,  stratigraphic  ob- 
servations in  vicinity  of,  15 


Telephone,  the  humming,  329,  644 
Telescopes  for  solar  research,  17 
Titchener  and  Pyle,  after-images  of 
subliminally    colored    stimuli,    366, 
644 
Tittmann,  Alaska  boundary,  15,  87 
Torricelli  ter-centenary,  644 
Trelease,  appointed  delegate  to  Uni- 
versity of  Missouri,  645 
True,  classification  of  the  cetacea,  15, 

385 
Tumor  growth  and  tissue  growth,  2, 3 

Turner,  Sir  William,  to  represent 
Society  at  Jubilee  of  Geological 
Society  of  Glasgow,  697 

Typotheria,  Santa  Cruz,  15,  64 


University  of  Cambridge,  invitation 
from,  to  Darwin  centenary,  644 

University  of  Missouri,  invitation  to 
installation  of  President  of,  645 

University  of  Pennsylvania  expedi- 
tion to  Alaska,  1907,  13 

Upson  and  Kennelly,  the  humming 
telephone,  329,  644 


Vedic  Concordance,  17 

Vessels  and  organs,  transplantation 
of,  645,  677 

Vice-President  elected  to  fill  unex- 
pired term  of  Professor  Barker, 
resigned,  134,  135 

W 

Wilder,  brain  of  rhinochimaera,  14,  yj 
Wiley,  effect  of  certain  preservatives 

upon  metabolism,  17 
Willcox,    effect    of    a     radio-active 

mineral  on  plant  growth,  697 
Wilson,  photographs  by  the  Lumiere 

process,  12 
Wire  conductor,  effect  of  an  angle 

in  a,  on  spark  discharge,  17 


OBITUARY  NOTICES 

OF 

MEMBERS  DECEASED 


William  Keith  Brooks. 

William  Keith  Brooks  was  bom  at  Cleveland,  O.,  March  25, 
1848,  and  died  at  his  home,  "  Brightside,"  near  Baltimore,  November 
12,  1908.  His  parents  were  born  in  Vermont,  but  their  ancestors 
had  lived  for  many  generations  at  or  near  Concord,  Mass.,  the  first 
of  the  name  having  come  to  America  from  England  prior  to  1634. 
Young  Brooks  receive^  his  early  education  in  the  public  schools  of 
Qeveland,  and  he  afterward  entered  Hobart  College,  Geneva,  N.  Y., 
where,  he  says,  "  I  learned  to  study,  and,  I  hope,  to  profit  by  but 
not  to  blindly  follow,  the  writings  of  that  great  thinker  on  the  prin- 
ciples of  science,  George  Berkeley."  He  spent  two  years  at  Hobart, 
where  he  took  high  honors,  and  then  entered  the  junior  class  at 
Williams  College.  Here  he  distinguished  himself  as  a  thorough  and 
independent  scholar,  and  is  said  to  have  been  one  of  the  most  bril- 
liant students  in  mathematics  Williams  had  ever  known.  In  1870  he 
received  the  degree  of  bachelor  of  arts  and  was  elected  to  Phi  Beta 
Kappa. 

After  his  graduation  his  father  took  him  into  mercantile  business- 
with  himself,  intending  that  he  should  become  his  successor,  but 
such  work  was  distasteful  to  young  Brooks  and  he  soon  abandoned 
it  and  became  a  teacher  in  a  boys  school  at  Niagara,  N.  Y.  When 
he  left  collie  he  was  undecided  whether  to  devote  himself  to 
mathematics,  to  Greek,  or  to  biology,  for  he  was  unusually  proficient 
in  all  of  these  subjects.  He  was  an  enthusiastic  naturalist;  even 
as  a  boy  he  had  given  much  attention  to  fresh-water  aquaria  and 
to  the  habits  of  animals,  and  he  had  published  some  of  his  observa- 
tions ;  with  one  of  his  friends  he  had  constructed  a  microscope  and 
with  other  associates  he  had  organized  a  class  in  natural  history; 
he  had  also  read  many  books  on  natural  history  and  was  intensely 
interested  in  evolution  and  Darwinism.  He  finally  decided  to  de- 
vote himself  to  biology,  largely  influenced,  we  may  imagine,  by  the 

philosophical  importance  of  this  subject. 

... 

lU 


IV  OBITUARY   NOllCES  OF  MEMBERS  DECEASED. 

At  Harvard  Louis  Agassiz  was  at  the  climax  of  his  wonderful 
career,  and  thither  flocked  many  young  men,  who  afterward  became 
leaders  in  biological  science,  to  study  under  this  great  master; 
among  these  was  Brooks.  In  the  summer  of  1873  he  was  a  student 
at  Agassiz's  laboratory  at  Penikese,  and  from  that  time  until  his 
death  he  remained  a  student  of  marine  life.  The  sea  with  its 
teeming  multitudes  of  living  things  always  had  a  particular  charm 
for  him,  not  merely  because  of  the  interest  and  variety  of  its  forms 
of  life,  but  also  because  it  was  the  scene  of  the  earliest  acts  in  the 
drama  of  evolution. 

In  1875  h^  received  the  d^ree  of  Ph.D.  from  Harvard  Uni- 
versity and  was  appointed  assistant  in  the  museum  of  the  Boston 
Society  of  Natural  History.  On  the  founding  of  the  Johns  Hop- 
kins University  in  1876  Brooks  applied  for  and  obtained  one  of 
their  twenty  famous  fellowships,  which  have  done  so  much  to 
change  the  character  of  university  work  and  ideals  in  this  country. 
Before  he  entered  upon  his  fellowship  his  abilities  as  a  teacher  were 
recognized  and  he  was  appointed  associate  in  biology.  In  1883 
he  was  appointed  associate  professor  of  morphology  and  in  1889 
professor  in  that  subject.  On  the  retirement  of  Professor  H. 
Newell  Martin  from  the  headship  of  the  Biological  Department  in 
1894,  Professor  Brooks  became  head  of  the  department  and  con- 
tinued in  that  position  until  his  death.  His  active  scientific  life 
was  therefore  coextensive  with  that  of  the  Johns  Hopkins  Uni- 
versity, and  his  love  of  the  Biological  Department  and  his  loyalty 
to  his  University  were  among  his  strong  characteristics. 

Although  his  publications  were  numerous  and  important  I  think 
that  his  influence  was  greatest  and  most  far  reaching  in  his  work 
as  a  teacher  and  scientific  director.  To  few  biologists,  perhaps  to 
no  other  in  the  history  of  this  country,  has  it  been  given  to  direct 
the  work  and  shape  the  scientific  ideals  of  so  large  and  influential 
a  body  of  young  men.  Among  those  who  took  their  doctor's  de- 
grees under  him  are  more  than  a  score  of  the  leading  zoologists 
of  this  country,  while  many  other  distinguished  scholars  of  this 
and  foreign  lands  were  his  pupils. 

Although  Professor  Brooks  would  present  a  subject  in  his  lec- 
tures in  a  most  clear  and  entertaining  manner,  he  rarely  if  ever 


OBITUARY   NOTICES  OF  MEMBERS   DECEASED.  V 

attempted  to  smooth  the  path  of  the  investigator;  the  latter  was  to 
a  very  large  extent  thrown  upon  his  own  resources.  He  believed 
so  thoroughly  in  the  law  of  natural  selection,  as  he  once  told  me, 
that  he  thought  it  was  best  for  a  student  to  find  out  for  himself, 
as  soon  as  possible,  whether  he  was  fitted  for  independent  investi- 
gation or  not,  and  by  this  rigid  discipline  the  unfit  were  weeded 
out  from  the  fit.  This  was  certainly  no  school  for  weaklings,  but 
it  afforded  magnificent  training  for  those  who  had  ability  and 
determination.  For  those  who  endured  this  ordeal  he  maintained 
the  warmest  regard,  and  his  interest  and  pride  in  the  work  of  his 
students  was  as  marked  as  it  was  stimulating. 

In  connection  with  his  work  as  teacher  and  director  must  be 
mentioned  the  establishment  by  him  of  the  Chesapeake  Zoological 
Laboratory  in  1878.  This  was  the  second  "marine  laboratory  in 
this  country  founded  for  advanced  work  in  pure  zoology.  The 
first  was  established  by  Louis  Agassiz  on  the  island  of  Penikese  in 
Buzzards  Bay  in  1871.  The  Qiesapeake  Laboratory,  unlike  the  one 
at  Penikese,  was  not  limited  to  one  place,  it  consisted  neither  of 
buildings  nor  equipment,  but  of  men  and  ideas.  For  the  first  few 
years  of  its  existence  it  was  located  at  several  different  points  in 
Chesapeake  Bay ;  afterwards  it  was  located  at  Beaufort,  N.  C,  then 
at  different  places  in  the  Bahama  Islands,  and  finally  in  Jamaica. 
In  the  various  expeditions  of  Brooks  and  his  students  to  these 
different  places  they  made  not  only  a  thorough  biological  survey 
of  each  region,  but  they  did  work  of  most  fundamental  and  far 
reaching  importance  on  the  various  groups  of  animals  found.  Out 
of  these  expeditions  has  grown  the  beautiful  and  permanent  sta- 
tion of  the  U.  S.  Fisheries  Bureau  at  Beaufort,  N.  C,  in  which 
Brooks  took  great  interest  and  pride. 

The  "  Scientific  Results  of  the  Sessions  of  the  Chesapeake 
Zoological  Laboratory  "  were  at  first  published  as  a  separate  journal 
of  which  Brooks  was  the  founder  and  editor,  later  this  was  incor- 
porated in  the  "  Studies  from  the  Biological  Laboratory  "  of  which 
he  was  joint  editor  with  H.  Newell  Martin.  He  subsequently 
established  and  edited  "  Memoirs  from  the  Biological  Laboratory," 
a  large  quarto  for  the  publication  of  important  monographs.      He 


VI  OBITUARY   NOTICES  OF  MEMBERS  DECEASED. 

was  also  one  of  the  editors  of  the  "Journal  of  Experimental 
Zoology." 

As  a  scientific  investigator  Brooks  showed  sound  judgment,  depth 
of  insight,  and  untiring  industry  and  enthusiasm.  In  his  research  he 
did  not  attempt  to  cover  the  whole  field  of  zoology,  but  he  did  attempt 
to  do  thoroughly  and  well  all  that  he  undertook.  His  work  began  at 
a  time  when  descriptive  embryology  was  the  newest  and  most  promis- 
ing branch  of  zoology  and  much  of  his  earlier  work  was  devoted 
to  this  field.  His  first  important  paper  was  on  the  "  Development 
of  Salpa,'*  and  many  of  his  later  works,  some  of  than  monumental 
monographs,  were  devoted  to  the  anatomy,  embryology  and  evolu- 
tion of  this  interesting  group  of  ascidians.  Indeed  his  latest  work 
which  was  left  in  manuscript  and  for  which  he  had  prepared  hun- 
dreds of  beautiful  'drawings,  was  a  continuation  of  his  great 
"  Monograph  on  the  Genus  Salpa,**  Among  other  important  re- 
searches may  be  mentioned  his  studies  on  the  "  Lucayan  Indians," 
"  Development  of  Marine  Prosobranchiate  Gasteropods,"  "  Early 
Stages  in  the  Development  of  Fresh  Water  Fulminates,"  "The 
Development  of  Lingula  and  the  Systematic  Position  of  the  Brachi- 
opoda,"  "  The  Relationships  of  MoUusca  and  MoUuscoidea,"  "  The 
Life  History  of  the  Hydromedusae,"  "The  Stomatopoda  of  the 
Challenger  Expedition,"  "  Lucifer :  A  Study  in  Morphology,"  "  The 
Embryology  and  Metamorphosis  of  the  Macroura"  (with  F.  H. 
Herrick),  and  a  "Monograph  of  the  Genus  Doliolum," 

His  studies  on  the  development  of  moUusks  led  him  to  an  ex- 
amination of  the  life  history  and  habits  of  the  oyster  and  this  was 
followed  by  a  consideration  of  the  best  methods  of  propagating  and 
cultivating  oysters.  His  work  on  this  subject  was  embodied  in  a 
book  called  "  The  Oyster,"  which  has  recently  appeared  in  a  second 
edition.  Because  of  its  economic  importance.  Brooks  has  been 
more  widely  known  through  this  work  than  through  any  other. 
He  was  made  chairman  of  the  Maryland  Oyster  Commission  and  did 
much  to  improve  this  industry  by  a  scientific  treatment  of  the  subject. 

He  wrote  but  one  text-book,  his  "  Handbook  of  Invertebrate 
Zoology"  (1882)  but  this  was  so  excellent  that  it  still  remains  a 
model,  and  in  some  respects  has  not  been  excelled,  if  equalled,  by 
any  later  book  on  that  subject. 


OBITUARY   NOTICES  OF  MEMBERS   DECEASED.  Vll 

His  chief  interest  was  always  in  the  philosophical  side  of  biology 
and  into  this  he  put  the  larger  part  of  his  life  work.  Even  the 
special  researches,  some  of  which  have  been  named  above,  were 
permeated  by  philosophical  inquiry,  and  most  of  his  books  and  later 
contributions  were  devoted  to  the  deeper  philosophical  meanings 
of  vital  phenomena. 

As  a  boy  he  had  read  the  works  of  Darwin  and  had  been  im- 
mensely impressed  by  them  and  to  the  last  he  yielded  to  no  one  in 
his  admiration  and  reverence  for  that  'great  master.  Probably  no 
other  disciple  of  Darwin  was  more  thoroughly  acquainted  with  his 
works,  and  very  frequently  when  criticisms  of  Darwinism  appeared 
he  would  point  out  the  fact  that  the  critic  did  not  understand  what 
Darwinism  is,  or  that  Darwin  had  already  met  and  answered  the 
objections  raised. 

In  1884  he  published  a  book  entitled  "  The  Law  of  Heredity," 
which  in  some  respects  anticipated  the  theories  of  Weismann,  and^ 
which  won  the  highest  commendation  from  Huxley  and  other 
leaders  of  biology.  But  probaHy  the  book  by  which  he  will  be 
longest  remembered  is  the  series  of  lectures  delivered  at  Columbia 
Uhiversity  and  published  in  the  Biological  Series  of  that  institution 
under  the  title  "The  Foundations  of  Zoology"  (1899).  I"  this 
book  he  deals  with  many  subjects  fundamental  not  only  to  zoology, 
but  to  science  and  philosophy  in  general.  Among  these  may  be 
mentioned  "  Nature  and  Nurture,"  "  Zoology  and  the  Philosophy 
of  Evolution,"  "  Natural  Selection  and  the  Antiquity  of  Life," 
*'  Natural  Selection  and  Natural  Theology,"  '^  Paley  and  the  Argu- 
ment from  Contrivance,"  "  The  Mechanism  of  Nature,"  **  Louis 
Agassiz  and  George  Berkeley,"  etc.  On  the  whole  his  chief  points  of 
view  may  be  summarized  in  his  oft-quoted  remark  of  Aristotle  that 
the  "  essence  of  a  living  thing  is  not  what  it  is  made  of  nor  what  it 
does,  but  why  it  does  it,"  or  as  he  expresses  it  elsewhere,  "the  essence 
of  a  living  thing  is  not  protoplasm  but  purpose  " ;  and  in  the  further 
statements  which  he  draws  from  Berkeley,  that  "  nature  is  a  lan- 
guage," that  "  phenomena  are  appearances,"  and  that  "  natural  laws 
are  not  arbitrary  nor  necessary,  but  natural,  i.  e.,  neither  less  nor 
more  than  one  who  has  the  data  has  every  reason  to  expect." 

On  March  25,  1898,  sixty  of  his  former  students  united  in  pre- 


•  •  • 


Vlll  OBITUARY   NOTICES  OF  MEMBERS  DECEASED. 

senting  to  him  an  oil  portrait  of  himself  together  with  a  congratu- 
latory address,  and  at  the  end  of  his  book  on  the  "  Foundations  of 
Zoology,"  he  added  on  this  date,  the  following  note : 

"For  you  who  have,  at  this  time,  for  my  encouragement,  called  your- 
selves my  students,  I  have  written  this  book  which  has  been  my  own  so 
long  that  I  should  part  with  it  with  regret,  did  I  not  hope  that,  as  you  study 
the  great  works  to  which  I  have  directed  you,  you  may  still  call  me  teacher. 
...  If  you  are  indeed  my  students,  you  are  not  afraid  of  hard  work,  so 
in  this  day  of  light  literature,  when  even  learning  must  be  made  easy,  you 
must  be  my  readers,  and  you  must  do  double  duty;  for  I  take  the  liberty 
of  a  teacher  with  his  pupils,  and  ask  that,  after  you  have  read  the  book,  you 
will  some  day  read  it  again;  since  I  hope  that  what  may  seem  obscure,  may, 
on  review,  be  found  consistent  and  intelligible." 

David  Starr  Jordan  review  this  book  in  Science  under  the 
caption  "  A  sage  in  biology."  Whatever  one  may  be  inclined  to 
say  of  his  conclusions  and  theories,  it  cannot  be  denied  that  in  an 
age  when  biological  investigators  have  been  content  with  discovering 
phenomena,  he  has  attempted  to  go  back  of  phenomena  to  their 
real  meaning  and  significance  and  to  point  out  the  relationship  of 
these  newly  discovered  phenomena  to  the  great  current  of  philoso- 
phy which  has  flowed  down  to  us  from  the  remote  past  , 

In  his  philosophical  writing  he  was  most  deeply  influenced  by 
Aristotle,  Berkeley  and  Huxley.  Much  that  he  has  written  still 
seems  to  me  obscure,  although  I  have  read  it  more  than  once,  but  I 
bear  in  mind  his  parting  request,  and  in  the  meantime  profit  by  that 
which  I  do  understand  and  am  charmed  by  the  classical  and  almost 
poetical  diction  in  which  it  is  written. 

His  abilities  received  early  and  generous  recognition.  Apart 
from  his  university  advancement  he  received  many  honors.  He 
received  the  honorary  degree  of  LL.D.  from  Williams  College  in 
1893,  from  Hobart  College  in  1899,  and  from  the  University  of 
Pennsylvania  at  the  Franklin  Bicentennary  in  1906.  In  1884,  at 
the  age  of  thirty-six,  he  was  elected  a  member  of  the  National 
Academy  of  Sciences;  he  was  chosen  a  member  of  the  American 
Philosophical  Society  in  1886;  of  the  Academy  of  Natural  Sciences 
of  Philadelphia  in  1887;  he  was  also  a  member  of  the  Boston  So- 
ciety of  Natural  History,  the  American  Academy  of  Arts  and 
Sciences,  of  the  Maryland  Academy  of  Arts  and  Sciences,  and  of 


OBITUARY   NOTICES  OF  MEMBERS   DECEASED.  IX 

the  American  Society  of  Zoologists ;  he  was  a  fellow  of  the  Amer- 
ican Association  for  the  Advancement  of  Science,  and  also  a  fellow 
of  the  Royal  Microscopical  Society.  For  his  work  on  the  oyster 
he  received  the  medal  of  the  Societe  d'Acclimatation  of  Paris ;  for 
his  work  on  the  scientific  results  of  the  Challenger  Expedition  he 
was  given  a  Challenger  Medal;  and  he  received  a  medal  at  the  St. 
Louis  Exposition  of  1904,  where  he  gave  an  address.  He  was 
Lowell  Lecturer  in  Boston  in  1901,  and  he  gave  one  of  the  principal 
addresses  before  the  International  Zoological  Congress  in  1907. 

These  honors  he  highly  prized,  and  perhaps  none  of  them  more 
than  his  membership  in  this  society.  Whenever  he  was  able,  he 
attended  the  general  meetings  of  the  society,  and  usually  presented  a 
paper  on  some  philosophical  subject.  He  served  as  a  counsellor 
of  the  society  and  frequently  spoke  to  me  of  its  purposes  and 
policies.  He  greatly  enjoyed  coming  into  this  historic  hall,  rich  in 
its  associations  with  great  men  of  the  past,  and  on  one  occasion 
when  I  spoke  to  him  of  the  plan  to  provide  a  larger  home  for  the 
society  in  a  more  central  part  of  the  city,  he  said  to  me,  "  Do  you 
think  you  have  any  right  to  move  the  home  of  the  society?  It 
seems  to  me  that  you  are  only  trustees  of  a  historic  institution, 
executors  of  an  ancient  trust,  and  that  you  have  no  right  to  remove 
this  monument  from  its  historic  site." 

In  personal  character  Professor  Brooks  was  simple  and  child- 
like, unconventional  in  manners,  dress  and  speech.  With  him  talking 
meant  expressing  ideas,  not  merely  passing  the  time,  and  if  he  had 
no  answer  ready  when  a  question  was  asked  him,  he  usually  gave 
no  answer  until  he  was  ready.  These  characteristics  made  him 
appear  somewhat  unique  and  picturesque,  and  gave  rise  to  many 
charming  anecdotes  about  him  which  his  students  and  friends  relate 
with  merriment,  but  real  affection.  He  was  kind  and  gentle;  and 
neither  in  his  publications  nor  in  his  relations  with  his  students 
did  he  ever  deal  in  scorn,  irony,  nor  invective.  President  Remsen 
said  of  him  that  he  had  been  called  the  most  lovable  man  in  the 
faculty.  His  interest  in  his  former  students  was  genuine  and  hearty 
though  he  rarely  expressed  it  directly  to  the  person  concerned.  He 
was  modest  and  dignified;  sincerity  itself;  loyal  to  his  friends,  his 
university,  and  his  ideals;  independent  in  thought  and  action,  and 


X  OBITUARY  NOTICES  OF  MEMBERS  DECEASED. 

not  easily  moved  from  a  position  he  had  once  taken.  He  was  a 
man  of  wide  culture;  he  loved  the  best  literature,  music  and  art. 
When  I  last  saw  him  at  his  home  we  spent  the  entire  evening  until 
after  midnight  pla3dng,  on  his  automatic  piano,  great  compositions 
of  Beethoven,  Mozart,  Wagfter  and  other  masters  of  harmony. 

In  his  home  life  he  was  most  happy  and  devoted.  He  married 
in  June,  1878,  Amelia  Schultz,  of  Baltimore,  by  whom  he  had  two 
children,  Chas.  E.  Brooks,  Ph.D.,  of  Elizabeth,  N.  J.,  and  Menetta 
W.  Brooks,  A.B.,  who,  after  the  death  of  Mrs.  Brooks  in  1901,  took 
charge  of  his  home. 

Professor  Brooks  once  told  me  that  he  proposed  to  retire  from 
his  professorship  when  he  had  reached  the  age  of  sixty  and  there- 
after devote  himself  entirely  to  philosophical  and  scientific  work. 
He  reached  the  age  of  sixty  last  March,  but  how  different  was  his 
realization  from  his  plan.  His  retirement  was  not  to  the  scholarly 
leisure  for  which  he  longed,  but  to  pain,  weakness  and  mortal 
sickness.  For  nine  months  he  struggled  against  a  complication  of 
organic  heart  trouble  and  kidney  disease  and  at  sunrise  on  Thurs- 
day, November  twelfth,  he  breathed  his  last. 

In  his  death  this  society  has  lost  a  worthy  and  devoted  member, 
the  world  of  scholars  a  man  of  rare  ability  and  accomplishments, 
and  his  friends  and  associates  a  noble  and  lovable  companion. 
Peace  to  his  ashes,  honor  and  reverence  to  his  memory! 

Edwin  G.  Conklin. 

Princeton  University. 


MAGELLANIC    PREMIUM 

Founded  in  1786  by  John  Hyacinth  de  Magellan,  of  London 

1909 

THE  AMERICAN  PHILOSOPHICAL  SOCIETY 

Held  at  Philadelphia,  for  Promoting  Useful  Knowledge 

ANNOUNCES  THAT  IN 

DECEMBER,  1909 

IT   WILL  AWARD    ITS 

MAGELLANIC  GOLD    MEDAL 

TO  THE  AUTHOR   OF   THE   BEST    DISCOVERY,    OR     MOST     USEFUL    INVENTION,    RK 
LATING    TO    NAVIGATION,    ASTRONOMY,   OR      NATURAL     PHILOSOPHY    (mERE. 
NATURAL    HISTORY  ONLY   EXCEPTED)  UNDER  THE   FOLLOWING  CONDITIONS    : 

1.  The  candidate  shall,  on  or  before  November  i,  1909,  deliver  free  of  postage 
or  other  charges,  his  discovery,  invention  or  improvement,  addressed  to  the 
President  of  the  American  Philosophical  Society,  No.  104  South  Fifth  Street, 
Philadelphia,  U.  S.  A.,  and  shall  distinguish  his  performance  by  some  motto, 
device,  or  other  signature.  With  his  discovery,  invention,  or  improvement,  he 
shall  also  send  a  sealed  letter  containing  the  same  motto,  device,  or  other  sig- 
nature, and  subscribed  with  the  real  name  and  place  of  residence  of  the  author. 

2.  Persons  of  any  nation,  sect  or  denomination  whatever,  shall  be  ad- 
mitted as  ciindidates  for  this  premium. 

3.  No  discovery,  invention  or  improvement  shall  be  entitled  to  this  premium 
which  hath  been  already  published,  or  for  which  the  author  hath  been  publicly 
rewarded  elsewhere. 

4.  The  candidate  shall  communicate  his  discovery,  invention  or  improvement, 
either  in  the  English,  French,  German,  or  Latin  language. 

5.  A  full  account  of  the  crowned  subject  shall  be  published  by  the  Society, 
as  soon  as  may  be  after  the  adjudication,  either  in  a  separate  publication,  or  in 
the  next  succeeding  volume  of  their  Transactions,  or  in  both. 

6.  The  premium  shall  consist  of  an  oval  plate  of  solid  standard  gold  of  the 
value  of  ten  guineas,  suitably  inscribed,  with  the  seal  of  the  Society  annexed  to 
the  medal  bv  a  ribbon. 

All  correspondence  in  relation  hereto  should  be  addressed 

To  THE  Secretaries  of  the 

AMERICAN   PHILOSOPHICAL   SOCIETY 

No.  104  South  Fifth  Street 

PHILADELPHIA,  U.  S.  A.