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CONTENTS 

\ 

INVITED  REVIEW 

EHINGER,  BERNDT  E.  J. 

Retinal  circuitry  and  clinical  ophthalmology    333 

BEHAVIOR 

HlDAKA,  MlCHIO 

Nematocyst  discharge,  histoincompatibility,  and  the  formation  of  swee- 
per tentacles  in  the  coral  Galaxea  fascicularis  350 

STEELE,  CRAIG  W. 

Non-random,  seasonal  oscillations  in  the  orientation  and  locomotor 
activity  of  sea  catfish  (Arius  felis)  in  a  multiple-choice  situation  ....  359 

DEVELOPMENT  AND  REPRODUCTION 

ANDERSON,  SUSAN  L.,  WALLIS  H.  CLARK,  JR.^  AND  ERNEST  S.  CHANG 

Multiple  spawning  and  molt  synchrony  in  a  free  spawning  shrimp  (57- 
cyonia  ingentis:  Penaeoidea) 377 

PETRAITIS,  PETER  S. 

Females  inhibit  males'  propensity  to  develop  into  simultaneous  her- 
maphrodites in  Capitella  species  I  (Polychaeta) 395 

WEIS,  VIRGINIA  M.,  DOUGLAS  R.  KEENE,  AND  LEO  W.  Buss 

Biology  of  hydractiniid  hydroids.  4.  Ultrastructure  of  the  planula  of 
Hydractinia  echinata  403 

ECOLOGY  AND  EVOLUTION 

LONSDALE,  DARCY  J.,  AND  JEFFREY  S.  LEVINTON 

Latitudinal  differentiation  in  embryonic  duration,  egg  size,  and  newborn 
survival  in  a  harpacticoid  copepod  : 419 

. 

PHYSIOLOGY 

COBB,  JAMES  L.  S. 

The  neurobiology  of  the  ectoneural/hyponeural  synaptic  connection  in 

an  echinoderm    432 

SANGER,  JOSEPH  W.,  AND  JEAN  M.  SANGER 

Sarcoplasmic  reticulum  in  the  adductor  muscles  of  a  Bermuda  scallop: 
comparison  of  smooth  versus  cross-striated  portions  447 

SILVERMAN,  BARRY  A.,  RONALD  W.  BERNINGER,  RICHARD  C.  TALAMO,  AND 

FREDERICK  B.  BANG 

The  use  of  the  urn  cell  complexes  of  Sipunculus  nudus  for  the  detection 
of  the  presence  of  mucus  stimulating  substances  in  the  serum  of  rabbits 
with  mucoid  enteritis 


461 


SHORT  REPORTS 


CHOW,  SEINEN,  YASUHIKO  TAKI,  AND  YOSHIMITSU  OGASAWARA 

Cryopreservation  of  spermatophore  of  the  fresh  water  shrimp,  Macro- 
brachium  rosenbergii    471 

JAFFE,  LIONEL  F.,  AND  ANTHONY  E.  WALSBY 

An  investigation  of  extracellular  electrical  currents  around  cyanobacterial 
filaments    476 

ROBINSON,  B.  W.,  AND  R.  W.  DOYLE 

Trade-off  between  male  reproduction  (amplexus)  and  growth  in  the 
amphipod  Gammarus  lawrencianus    • 482 

Index  to  Volume  168  489 


Volume  168  Number  3 

SYMPOSIUM  SUPPLEMENT  TO  THE 

BIOLOGICAL  BULLETIN 


PUBLISHED  BY 
THE  MARINE  BIOLOGICAL  LABORATORY 

C.  B.  Metz,  Editor;  P.  L.  Clapp,  Assistant  Editor 

THE  NAPLES  ZOOLOGICAL  STATION  AND 

THE  MARINE  BIOLOGICAL  LABORATORY: 

ONE  HUNDRED  YEARS  OF  BIOLOGY 

CONTENTS 

GROSS,  PAUL  R. 

Preface     1 

NINETEENTH-CENTURY  BACKGROUND,  AND  PERSONALITIES 

GROEBEN,  CHRISTIANE 

Anton  Dohrn — the  statesman  of  Darwinism    4 

MAIENSCHEIN,  JANE 

Agassiz,  Hyatt,  Whitman,  and  the  birth  of  the  Marine  Biological  Lab- 
oratory         26 

MONROY,  ALBERTO,  AND  CHRISTIANE  GROEBEN 

The  "new"  embryology  at  the  Zoological  Station  and  at  the  Marine 
Biological  Laboratory  35 

REINGOLD,  NATHAN,  AND  JOEL  N.  BODANSKY 

The  sciences,  1850-1900,  a  North  Atlantic  perspective  . 44 

THE  EVOLUTION  OF  DISCIPLINES 

GROSS,  PAUL  R. 

Laying  the  ghost:  embryonic  development,  in  plain  words  62 

EBERT,  JAMES  D. 

Cell  interactions:  the  roots  of  a  century  of  research 80 

RUSSELL-HUNTER,  W.  D. 

An  evolutionary  century  at  Woods  Hole:  instruction  in  invertebrate 
zoology 88 

FANTINI,  BERNARDINO 

The  sea  urchin  and  the  fruit  fly:  cell  biology  and  heredity,  1900-1910       99 

ALLEN,  GARLAND  E. 

Heredity  under  an  embryological  paradigm:  the  case  of  genetics  and 
embryology , 107 

Continued  on  Cover  Two 


GHIRETTI,  FRANCESCO 

Comparative  physiology  and  biochemistry  at  the  Zoological  Station  of 
Naples 122 

COHEN,  SEYMOUR  S. 

Some  struggles  of  Jacques  Loeb,  Albert  Mathews,  and  Ernest  Just  at 

the  Marine  Biological  Laboratory    127 

FLOREY,  ERNST 

The  Zoological  Station  at  Naples  and  the  neuron:  personalities  and 
encounters  in  a  unique  institution .  .  137 

YOUNG,  J.  Z. 

Cephalopods  and  neuroscience    153 

BENNETT,  M.  V.  L. 

Nicked  by  Occam's  razor:  unitarianism  in  the  investigation  of  synaptic 
transmission  . 159 

TOMAS,  CARMELO  R. 

Marine  botany  and  ecology  at  Stazione  Zoologica     168 

PAST  AND  FUTURE:  SHORTER  COMMUNICATIONS  ON  POLICY 

AND  POLITICS 

EBERT,  JAMES  D. 

Carnegie  Institution  of  Washington  and  marine  biology:  Naples,  Woods 
Hole,  and  Tortugas 172 

EBERT,  JAMES  D. 

Evolving  institutional  patterns  for  excellence:  a  brief  comparison  of  the 
organization  and  management  of  the  Cold  Spring  Harbor  Laboratory 
and  the  Marine  Biological  Laboratory 183 

MAIENSCHEIN,  JANE 

First  impressions:  American  biologists  at  Naples   187 

MAIENSCHEIN,  JANE 

Early  struggles  at  the  Marine  Biological  Laboratory  over  mission  and 
money 192 

RUSSELL-HUNTER,  W.  D. 

The  Woods  Hole  laboratory  site:  history  and  future  ecology    197 

RUSSELL-HUNTER,  W.  D. 

From  Woods  Hole  to  the  world:  The  Biological  Bulletin  200 

MIRALTO,  ANTONIO 

What  laboratories  for  what  science? 203 

Index  205 


Reference:  Biol.  Bull  168  (suppl.):  1-3.  (June,  1985) 


LIBRARY 


PREFACE 

The  Stazione  Zoologica  of  Naples  was  founded  in  1 872;  the  Marine  Biological 
Laboratory  received  its  first  students  and  investigators  in  Woods  Hole  during  the 
summer  of  1888.  That  each  has  survived  is  surprising;  that  they  have  survived  these 
tumultuous  hundred  years  of  science  not  merely  intact,  but  with  steady  growth  of 
influence  worldwide,  is  remarkable.  There  were  close  and  cooperative  relationships 
among  leading  scientists  of  the  two  laboratories  during  the  decades  between  1890 
and  the  outbreak  of  the  first  World  War.  Cooperative  relationships  have  continued, 
but  the  years  of  the  last  century's  turn  were  momentous  ones  for  what  became 
"modern"  biology.  What  went  on  at  the  Stazione  and  at  the  MBL  is  thus  of  great 
interest  as  intellectual  history  and  for  the  lessons  to  be  learned  about  the  role  of 
institutions,  as  such,  in  the  advance  of  science. 

In  October  of  1984,  a  meeting  was  held  at  the  Stazione's  ecology  laboratory  on 
the  Island  of  Ischia  (which  laboratory  was  once  a  villa  belonging  to  Anton  Dohrn, 
the  Stazione's  founder).  With  timely  assistance  from  the  Commonwealth  Fund,  the 
Klingenstein  Fund,  and  Italian  agencies  supporting  the  Stazione,  a  group  of 
European  and  American  scholars — historians  of  science  and  biologists  in  about 
equal  numbers — met  to  review  the  events,  the  ideas,  and  the  personalities  of  a 
hundred  years  of  biology  at  the  two  places.  As  is  generally  known,  the  two  ostensibly 
"marine"  laboratories  have  had  impact  on  far  more  than  marine  sciences  in  that 
interval. 

The  participants  could  barely  scratch  the  surface  of  an  immense  subject,  many 
of  whose  source  materials  are  still  unexamined.  But  there  was  general  agreement  on 
its  importance  and  on  the  opportunities  for  new  and  significant  research.  Thus  the 
Ischia  meeting  and  its  contributions  were  meant  to  serve  rather  more  than  the 
normal  purposes  of  a  scholarly  gathering:  they  were  and  are  meant  to  be  a  goad 
and  catalyst  for  new  historical  research,  and  to  act  thus  upon  working  biologists 
with  a  taste  for  history  as  well  as  upon  historians  of  biology  and  medicine. 

Such  efforts  are  rarely  successful:  historians  and  experimental  scientists  (and 
administrators  of  science)  often  fail  to  communicate.  They  have  different  styles  of 
work  and  different  imperatives.  The  specialties  have  diverged  greatly  during  their 
professionalization  of  the  past  twenty-five  years.  The  high  standards  of  one  group 
are  not  necessarily  of  interest  or  importance  to  the  other;  and  what  is  inexcusable 
as  form  for  one  can  be  perfectly  acceptable  to  another  scholar. 

The  Ischia  meeting  was,  nevertheless,  a  success.  It  included  contributions  from 
both  categories  of  participant  that  were  exciting  and  novel  to  the  other:  the  speakers 
did  not  speak  past  one  another.  Catalysis  of  new  projects  was  in  the  air.  The 
auguries  for  new  writing  in  time  for  the  MBL's  1988  Centennial  are  strong. 

This  volume  contains  the  written  version  of  papers  presented  viva  voce  at  Ischia. 
Their  publication  is  essential  for  the  catalytic  purposes  mentioned.  This  posed, 
however,  a  problem  inherent  in  the  different  working  styles  of  historians,  archivists, 
experimental  scientists,  and  administrators.  We  saw  the  choice,  initially,  as  one  of 
forcing  all  the  papers  into  one  or  the  other  mold:  that  of  an  historical  journal  or 
that  of  a  biological  journal  such  as  THE  BIOLOGICAL  BULLETIN.  In  the  end  we 
rejected  both,  for  it  seemed  to  us  more  important  to  communicate  the  events  and 
flavor  of  the  Ischia  meeting  accurately  than  to  dress  up  the  biologists'  presentations 
to  fit  the  style  of  historical  scholarship  or  the  reverse.  The  chapters  of  this  volume 


PREFACE 


Ischia  participants  left  to  right:  J.  D.  Ebert,  J.  Maienschein,  M.  V.  Bennett,  G.  Allen,  C.  Groeben,  P. 
Gross,  N.  Reingold,  J.  Z.  Young,  E.  Florey.  Mrs.  Florey,  W.  D.  Russell-Hunter,  E.  Cohen.  S.  Cohen.  A. 
Monroy,  and  F.  Ghiretti.  Not  shown:  B.  Fantini. 


are,  therefore,  si{i  generis:  they  are  what  went  on  at  Ischia,  with  transduction  only 
from  the  spoken  to  the  written  word. 

The  well-disposed  reader  will  therefore,  I  hope,  tolerate  what  might  drive  a 
proper  editor  to  fury:  as  many  different  forms  of  bibliographic  citation,  nearly,  as 
there  are  papers,  and  some  abrupt  transitions  of  purpose  and  point  of  view.  So  be 
it:  we  believe  that  there  is  to  be  gained  an  invaluable  sense  of  the  importance  of  the 
subject  to  different  biological  interests.  Much  will  be  found  here  that  is  new,  and 
some  well-known  stories,  or  stories  assumed  to  be  well-known,  are  re-told  with  new 
insights.  The  price  of  novelty  is  a  much  greater  intrusion  of  the  personalities  of  the 
writers  than  is  allowed  today  in  most  professional  journals. 

The  papers  as  printed  follow  fairly  closely,  but  not  exactly,  the  order  of 
presentation  at  the  meeting.  The  broad  design,  established  initially  by  S.  S.  Cohen 
and  A.  Monroy,  the  organizers,  was  for  an  opening  session  on  late-nineteenth 
century  science  in  general,  followed  by  developments  in  the  characteristic  disciplines 
of  the  two  laboratories:  zoology,  embryology,  genetics,  biochemistry,  neurophysiology, 
botany,  ecology.  These  sessions  were  succeeded  by  discussions  of  particular  institu- 
tional policies  and  problems,  e.g.,  the  role  of  teaching  and  the  funding  of  research. 
There  were,  finally,  a  number  of  informal  discussions  of  present-day  issues  of 
organization  and  policy.  This  sequence  is  maintained,  on  the  whole,  in  the  volume. 

The  participants  are  grateful  to  the  supporting  agencies,  and  to  the  Naples  and 
Ischia  staff  members  for  their  cooperation  and  characteristic  warm  hospitality.  I  am 
personally  grateful  to  my  colleagues.  Garland  Allen  and  Jane  Maienschein,  for  their 


PREFACE  3 

indispensable  assistance  in  the  substantive — and  hence  serious — editorial  work.  We 
all  hope  that  publication  of  these  proceedings  will  attract  more  of  our  colleagues  to 
the  subject,  and  to  exploring  riches  in  the  archives  of  both  laboratories.  There  are 
powerful  continuities  of  purpose  underlying  the  key  ideas  of  modern  biology,  and 
those  emerge  strongly  from  study  of  the  two  institutions.  Such  continuities  do  not 
necessarily  emerge  from  preoccupation,  however  intense  and  skilled,  with  the  current 
literature  of  biology.  Yet  there  is  reason  to  believe  that  understanding  those  driving 
forces  (and,  yes,  intellectual  prejudices)  can  be  helpful,  perhaps  essential,  for  wise 
decisions  about  tomorrow's  research  directions  in  the  laboratory.  If  the  continuities 
are  communicated  even  to  a  few  readers  of  this  volume,  as  they  have  largely  ceased 
to  be  communicated  in  graduate  biological  education,  then  the  efforts  of  organizing 
the  meeting  and  assembling  this  volume  will  have  been  rewarded. 

Paul  R.  Gross 

Marine  Biological  Laboratory 

March,  1985 


Reference:  Biol.  Bull.  168  (suppl.):  4-25.  (June,  1985) 

ANTON   DOHRN— THE  STATESMAN   OF  DARWINISM 

To  commemorate  the  75th  anniversary  of  the  death  of  Anton   Dohrn 

CHRISTIANE  GROEBEN 
Stazione  Zoologica,  Villa  Comunalc,  80121  Naples.  Italy 

ABSTRACT 

Based  on  personal  accounts  from  the  Archives  of  the  Stazione,  this  paper  traces 
the  development  of  Anton  DohnVs  personality.  His  inner  motivations,  his  personal 
experiences,  and  the  external  factors  and  traditions  that  led  to  the  foundation  of 
the  Naples  Zoological  Station  are  described. 

Dohrn  strove  for  a  greater  degree  of  organization  in  science.  The  paper 
investigates  the  factors  helping  to  achieve  this  goal  and  outlines  the  influence  that 
the  Naples  Station  had  on  the  foundation  of  other  institutions. 

DISCUSSION 

In  youth  we  plume  our  fancy's  wings 

To  flit  from  sun  to  sun; 
At  length  we  fold  them  up.  and  then 

A  little  work  is  done. 
Compared  with  our  imagined  deeds 

How  small  and  poor  it  seems! 
But  then — one  little  act  outweighs 

A  thousand  glorious  dreams. 

Charles  Grant1 

Anton  Dohrn  (1840-1909),  founder  of  the  Stazione  Zoologica  in  Naples,  once 
said:  "To  create,  to  organize,  to  develop — this  is  my  need,  even  passion."2  It  is  on 
these  motivations  that  I  shall  focus  in  the  following  presentation. 

The  Naples  Zoological  Station  was  an  offspring  of  Anton  Dohrn"s  personality. 
Dohrn  has  been  denned  as  "an  independent  pioneer  of  science  politics"  (Heuss),3 
"a  catalyst  in  the  encouragement  and  stimulation  of  creative  ideas"  (I.  Miiller, 
1975,  p.  193),  and  also  as  being  gifted  with  "an  unusually  sure  eye  for  the  significance 
of  the  different  sections  of  our  science,  and  for  the  way  in  which  they  interrelate 
and  complement  each  other""  (Th.  Boveri,  1910,  p.  31).  Dohrn"s  personality 
dominated  and  catalyzed  the  activity  and  the  work  carried  out  at  the  Station,  from 
its  founding  until  long  after  his  death  in  1909. 

In  1975  O.  Skalova,  in  her  sociological  case  study  of  the  Naples  Zoological 
Station,  contacted  2000  guest  scientists  who  had  worked  at  the  Station  during  the 
previous  40  years.  From  the  650  who  replied  it  was  clear  that  one  of  the  most 
highly  rated  factors  was  "the  creative  atmosphere""  prevailing  in  the  laboratory 
(Skalova,  1975,  pp.  26-28).  This  atmosphere  undoubtedly  went  a  long  way  to 
making  the  Zoological  Station  a  "permanent  Congress  of  Zoologists,"'  as  Theodor 
Boveri  called  it  in  his  1910  commemoration  of  Anton  Dohrn  (Th.  Boveri,  1910, 
p.  40). 

I  have  long  attempted  to  discover  the  factors  that  led  to,  or  that  help  to  maintain, 
the  much-discussed  "creative  atmosphere""  of  the  Naples  Zoological  Station.  It  was 


ANTON   DOHRN  5 

a  peculiar  condition  in  which  most  scientists  became  engulfed  but  could  not  explain. 
It  emanated  in  the  first  instance  from  Anton  Dohrn.  The  intriguing  question  is  how 
this  catalyzing  process  actually  worked,  or  better,  how  the  personality  of  Anton 
Dohrn  brought  about  such  a  sense  of  creativity  in  others.  It  is  also  important  to 
analyze  the  external  and  internal  factors,  that  contributed  to  the  foundation  of  a 
research  institute  where,  in  the  words  in  Hans  Driesch  in  1909,  "9/u>  of  all  basic 
work  in  modern  zoology  has  been  done'"  (Driesch,  1909,  p.  514). 

Much  has  been  written  on  the  foundation  and  the  history  of  the  Naples  Station 
(Kofoid,  1910;  Groeben  and  Miiller,  1975;  Miiller,  1975,  1976;  Partsch,  1980; 
Miiller  and  Groeben,  1984).  In  1940,  Theodor  Heuss  published  an  exhaustive 
biography  of  Anton  Dohrn  (Heuss,  1962);  and  the  scientist  Dohrn  and  his  relation 
to  the  zoology  of  his  time  has  been  excellently  described  by  Alfred  Kiihn  (1950). 
Yet,  the  question  of  the  influence  of  Dohrn's  personality  on  the  scientific  work  of 
the  Station  remains  unexplored. 

Anton  Dohrn's  relationship  with  his  father  Carl  August  Dohrn  (1806-1892),  a 
well  known  amateur  entomologist  (Heuss,  1962,  pp.  23-49;  P.  Dohrn,  1983,  pp. 
30-81),  played  an  important  role  in  forming  Anton  Dohrn's  character.  His  search 
for  acknowledgment  among  his  peers  and  his  essential  loneliness  as  an  adult  date 
from  his  childhood,  as  do  his  liberal  views,  his  interest  in  people,  and  his  wide 
knowledge  of  literature,  music,  and  science. 

The  relationship  between  Carl  August  and  Anton  Dohrn  was  difficult  and 
tormented,  ranging  from  heated  discussions,  ruptures,  and  disinheritance  to  recon- 
ciliation and  eventually  mutual  respect.  Dohrn  was  struck  by  his  father's  brilliance 
and  knowledge,  and  his  wide-ranging  correspondence  with  eminent  scientists.  He 
longed  to  live  up  to  his  father's  expectations,  hoping  to  win  his  respect  and  love.  In 
1897,  on  the  occasion  of  the  25th  anniversary  of  the  Naples  Zoological  Station, 
Anton  Dohrn  acknowledged  his  debt  to  his  father,  his  "intellectual  protoplasma," 
but  he  also  noted  that  in  their  family  circle  it  was  more  important  to  remember 
quotations  from  Goethe  or  to  recognize  music  from  Beethoven,  than  to  excel  in 
Greek  or  mathematics  (A.  Dohrn,  1897,  p.  34). 

After  having  received  a  humanistic  education,  Dohrn  studied  medicine  and 
zoology  at  Konigsberg,  Bonn,  Jena,  and  Berlin,  obtaining  his  Ph.D.  at  Breslau  in 
November  1865.  He  became  Privatdozent  at  Jena  in  January  1868.  To  friends  he 
admitted: 

I  have  no  vocation  as  a  Zoologist.  You  see,  to  pass  my  whole  day  observing 
through  glasses  whether  this  crab  has  seven  or  eight  segments,  whether  here  lie 
cells  with  two  or  three  nuclei,  whether  this  tissue  has  grown  thus  or  thus — this  I 
can't  do.4 

He  was  always  prone  to  bigger  thoughts,  to  more  grandiose  schemes. 

Dohrn's  attitude  suddenly  changed,  when  through  the  influence  of  Ernst  Haeckel 
at  Jena  in  1862,  Dohrn  became  acquainted  with  Darwin's  works.  As  he  exclaimed 
in  the  letter  of  1866,  cited  above,  "[I  felt]  a  really  piercing  excitement;"  the  once 
dry  Zoology  was  no  longer  an  end  in  itself,  but  a  new  tool  to  acquire  knowledge. 
The  way  Dohrn  put  that  new  tool  to  use  was  through  the  study  of  morphology: 
that  branch  of  zoology  in  which  fields  such  as  comparative  anatomy,  embryology, 
and  physiology  were  used  as  a  means  of  elucidating  the  phylogenetic  history  of 
various  groups.  Comparative  embryology  became  a  cornerstone  of  morphology, 
based  on  Haeckel's  recapitulation  theory:  that  the  individual  in  its  embryonic 
development  passes  through  the  major  stages  of  its  own  evolutionary  past.  Morphology 
thus  became  one  of  the  major  ways  in  which  zoologists  sought  to  expand  and 


6  C.  GROEBEN 

develop  Darwinian  theory  in  the  last  30  years  of  the  nineteenth  century.  As  a  result 
of  his  conversion  to  evolutionary  theory  in  the  early  1860's,  Dohrn  promised  to 
dedicate  his  whole  life  to  Darwinism — a  promise  he  indeed  kept. 

In  1865,  Dohrn  accompanied  Haeckel  on  his  famous  expedition  to  Helgoland 
where,  for  the  first  time  Dohrn  studied  marine  organisms  (Fig.  1 ).  It  seems  to  have 
been  in  Helgoland,  while  struggling  home  with  buckets  full  of  sea  water,  that  Dohrn 
and  Haeckel  talked  about  a  "Zoological  Station"  (Uschmann,  1959,  p.  65).  Dohrn's 
friendship  with  Haeckel  ended  a  few  years  later  when  the  younger  man,  influenced 
by  Kant  and  F.  A.  Lange's  Geschichte  des  Materialismus  (1866),  could  no  longer 
subscribe  to  HaeckeFs  philosophical  generalizations.  But  familiarity  with  marine 
organisms  as  objects  for  study,  and  the  idea  of  establishing  a  marine  laboratory 
remained  an  important  legacy  from  Haeckel  for  the  rest  of  Dohrn's  life. 

In  1867  (July-September)  and  again  in  1868  Dohrn  went  to  Millport,  Isle  of 
Cumbrae,  Scotland,  to  continue  his  studies  on  Arthropoda.  There  he  lived  as  a 
guest  of  David  Robertson  (1806-1896),  a  self-taught  Scottish  zoologist  and  later 
founder  of  the  Millport  Biological  Station  (1885). 

It  was  during  this  first  trip  to  England  that  Dohrn,  through  an  introduction  by 
his  father  to  the  entomologist  Henry  Tibbats  Stainton  (1822-1892),  met  many 
English  zoologists,  most  importantly  Thomas  Henry  Huxley  (1825-1895),  with 


FIGURE  1.  Excursion  to  Helgoland,  1865.  Standing,  left  to  right:  Anton  Dohrn,  Jena;  Richard 
Greef,  Bonn;  Ernst  Haeckel,  Jena.  Sitting,  left  to  right:  Salverda,  Delft;  Pietro  Marchi,  Florence. 
(Reproduced  with  permission  from:  Uschmann,  1959,  fig.  23.) 


ANTON   DOHRN  7 

whom  a  close  and  warm  friendship  developed.  Huxley  even  suggested  a  visit  to 
Charles  Darwin  (1809-1882),  but  Dohrn  only  sent  Darwin  his  publications  at  that 
time.  This  led  to  an  exchange  of  letters  (Dohrn  did  meet  Darwin  two  years  later,  in 
September  1870;  see  Groeben,  1982).  Dohrn  also  reported  on  his  work  at  the  annual 
meeting  of  the  British  Association,  held  that  year  (1868)  at  Dundee. 

Dohrn's  close  ties  with  British  scientists  were  to  pay  off  in  future  years.  During 
the  difficult  period  (1870-1874)  when  he  was  trying  to  build  the  Zoological  Station, 
Dohrn  received  the  greatest  encouragement  and  help  from  his  British  colleagues. 
Apart  from  the  active  support  of  Darwin  and  Huxley,  Dohrn  found  a  great  source 
of  strength  and  encouragement  for  his  dream  of  a  marine  laboratory  in  the  British 
natural  history  tradition  and  the  readiness  of  the  British  to  accept  adventurous  and 
even  somewhat  eccentric  plans. 

In  the  period  after  1860,  stimulated  particularly  by  French  and  British  exploring 
expeditions,  general  interest  in  the  sea  and  its  riches  and  in  the  potential  of  marine 
organisms  for  systematic  and  morphological  study  had  increased  greatly  throughout 
Europe  (Deacon,  1971;  Rice  and  Wilson,  1980).  In  1870  at  the  annual  meeting  of 
the  British  Association  in  Liverpool,  a  Committee  was  formed  "for  the  purpose  of 
promoting  the  foundation  of  zoological  stations  in  different  parts  of  the  world" 
(Report,  1871;  Dohrn,  1872a;  Rice  and  Wilson,  1980).  Indeed,  it  was  this  committee, 
through  its  many  reports  as  well  as  notes  and  articles  regularly  published  in  Nature, 
that  gave  such  widespread  publicity  in  the  English-speaking  world  to  Dohrn's 
Stazione  Zoological. 

In  Germany  the  general  concepts  of  using  marine  organisms  for  biological 
research  was  first  promoted  by  Johannes  Miiller  (1801-1858).  Through  his  numerous 
visits  to  the  North  Sea  (1843,  1848,  1854,  and  1855)  and  especially  the  Mediterranean 
(in  1849  at  Nice  and  Villefranche,  1850-52  to  Trieste,  in  1853  at  Messina,  1856  at 
Cette  and  Nice),  Miiller  publicized  widely  the  concept  of  marine  biological  research 
as  a  means  of  elucidating  fundamental  biological  concepts.  Miiller  was  a  gifted 
teacher,  with  whom  most  of  Germany's  principal  zoologists  and  physiologists  had 
studied  (Kiihn,  1950).  Thus,  his  interest  and  influence  was  of  considerable  importance. 

After  the  British  Association  meeting  in  1868,  Dohrn  returned  to  Millport, 
where,  together  with  David  Robertson  he  constructed  a  portable  aquarium,  to  take 
with  him  to  Messina,  Sicily,  where  he  planned  to  spend  the  winter  of  1868-69. 

The  Straits  of  Messina  were  famous  at  that  time  for  the  richness  and  variety  of 
their  marine  fauna  and  flora.  In  1788,  Spallanzani  had  studied  the  pelagic  fauna  of 
the  area,  followed  many  years  later  by  the  German  zoologist  August  D.  Krohn 
(1846).  And,  Johannes  Miiller  must  have  conveyed  his  enthusiasm  for  seaside 
studies  there  to  many  of  his  students,  since  Messina  soon  became,  according  to  one 
report,  the  "Mecca  of  the  German  Privat-Dozent/v>  So  favorable  were  collecting 
conditions  at  Messina,  for  example,  that  in  one  year  (1859-60),  Haeckel  (also  one 
of  Miiller's  students  and  inspired  by  him  to  do  marine  investigations),  discovered 
more  than  1 44  new  species  of  Radiolaria  in  the  straits. 

Messina  was  thus  the  obvious  choice  for  Dohrn  to  conduct  marine  studies. 
There  Dohrn  met  up  with  his  friend,  the  Russian  zoologist  Nikolai  N.  Mikloucho- 
Maclay  (1848-1888),  also  a  former  pupil  of  Haeckel,  and  later  to  become  an 
eminent  anthropologist  (Miiller,  1980).  Maclay  introduced  Dohrn  to  the  Russian- 
Polish  family,  de  Baranowski,  whose  eldest  daughter  Marie  (1856-1918)  was  to 
become  Dohrn's  wife  in  1874. 

Dohrn  and  Maclay  rented  two  rooms  at  the  Palazzo  Vitale,  Strada  Garibaldi, 
right  on  the  port,  with  a  breathtaking  view  of  the  Straits  of  Messina  and  the  coast 
of  Calabria.  They  bought  some  chairs  and  had  two  work-benches  made  for  them. 
Commenting  on  their  simple  life,  Dohrn  wrote  in  1868: 


8  C.  GROEBEN 

.  .  .  and  so  we  live,  including  service  and  meals,  for  5  francs  a  day,  much 
cheaper  and  better  than  at  a  Hotel,  where  we  would  have  had  to  climb  3-4  floors 
and  would  also  have  had  difficulties  because  of  our  working  problems.  The 
fishermen  invade  our  flat  and  bring  us  lots  of  animals.6 

The  portable  aquarium  Dohrn  had  built  in  Scotland  proved  to  be  very  valuable 
indeed.  For  the  first  time  he  could  observe  the  breeding  of  crustacean  eggs.  However, 
faced  with  such  difficulties  as  bad  weather,  lack  of  animal  supply,  and  lack  of 
literature,  Dohrn  realized  how  useful  it  would  be  for  scientists  to  arrive  and  find 
the  "table  laid"  for  their  work  (Fig.  2).  That  is,  to  find  on  arrival  instruments, 
rooms,  service,  chemicals,  and  books  available  together  with  records  of  where  and 
when  certain  species  could  be  found  and  useful  information  on  local  conditions, 
etc.  In  this  spirit  he  left  his  equipment  and  diary  at  Messina  where  friends  promised 
to  take  care  of  them.  In  February  1869  Dohrn  could  thus  write  to  his  father:  "The 
Zoological  Station  of  Messina  has  now  been  established.  I  retain  it  an  important 
progress  for  our  Science  should  we  succeed  in  getting  beyond  the  embryonic  stage." 
Dohrn  and  Maclay  had  optimistically  planned  to  cover  the  world  with  a  network 
of  Zoological  Stations.  Although  Dohrn  thought  initially  of  Venice,  Nice,  Gibraltar, 
Portugal,  Ceylon,  Australia,  and  the  Cape  of  Good  Hope8,  in  the  end  Naples  was 
all  he  could  successfully  accomplish. 

The  potential  of  marine  organisms  for  morphological  and  systematic  studies  had 
already  led  several  scientists  to  try  to  establish  study  facilities  near  the  sea,  e.g.,  the 
French  stations  at  Concarneau  (1859),  Arcachon  (1863),  and  Roscoff  (1872). 
However,  these  were  mostly  field  stations,  connected  to  a  University  or  Institute, 
and  were  not  independent  facilities  to  house  a  host  of  different  sorts  of  investigations 
and  a  wide  variety  of  projects. 

Dohrn  always  acknowledged  Vogt  (1817-1895)  as  his  forerunner.  In  1852,  Vogt 
had  tried  at  Villefranche,  and  in  1863  at  Naples,  to  found  a  research  and  observation 
station  (Vogt,  1871;  Dohrn,  1871,  pp.  6-8;  Oppenheimer,  1980).  Neither  had 
materialized,  however. 

Leaving  Messina  in  April  1869,  Dohrn  returned  to  Jena  for  his  summer  term 
lectures  and  also  to  collect  money  for  a  small  building  to  be  constructed  at  Messina 
to  house  the  equipment.  In  so  doing  Dohrn  put  to  work  his  "creative  imagination"9, 
a  quality  he  was  aware  of  possessing  to  an  unusual  degree. 

Dohrn's  enthusiasm  for  Darwinism  did  little  to  remove  his  dissatisfaction  with 
Zoology — research  and  teaching — as  it  stood  at  that  time.  He  felt  that  his  interests 
were  too  wide,  or  Zoology  at  the  University  too  narrow — to  be  able  to  continue  his 
University  career.  Science  in  itself  and  scientific  discoveries  just  didn't  stimulate 
him,  or  as  Th.  Boveri  put  it  (1910,  p.  24): 

he  lacked  the  most  elementary  urge  of  a  scientist,  the  urge  to  observe,  to  discover 
hitherto  unknown  facts,  even  if  they  are  unknown  only  to  the  observer  himself. 
Not  that  he  did  not  acknowledge  the  value  of  new  discoveries.  But  it  did  not 
matter  to  him  to  make  them  himself. 

Dohrn  wanted  to  make  a  contribution  to  advancing  Darwinism  that  would 
exploit  his  complex  personality  and  gifts.  He  once  put  it  like  this: 

Proper  zoological  work  has  elements  that  do  not  appeal  to  me  and  do  not  at  all 
take  into  account  an  inner  need,  which  is  to  occupy  myself  in  a  practical  way, 
to  make  an  impact  in  the  outside  world,  to  be  of  service  to  others.10 

And  again  in  1874,  during  one  of  his  frequent  bouts  of  deep  depression — a  heritage 


ANTON   DOHRN 


FIGURE  2.  N.  N.  Mikloucho-Maclay,  letter  to  Anton  Dohrn,  January  1869,  Messina,  describing 
and  illustrating  the  hardships  he  had  to  cope  with  in  order  to  pursue  his  studies.35  (ASZN,  Dohrn 
Archives,  Ba  735) 


from  his  mother's  family — he  wrote  in  an  unfinished  last  will:  "In  the  center  of  my 
existence  lies,  I  may  say,  a  passion  for  helping  others,  direct  or  indirect." 

The  growing  interest  in  exploring  life  at  sea,  the  need  for  marine  organisms  for 
research  in  morphology  and  embryology,  Dohrn's  own  marine  experiences,  his 
championing  of  Darwinism,  and  his  need  to  prove  himself — all  these  currents 
converged  into  the  creation  of  the  Naples  Zoological  Station.  Dohrn's  attention  had 
turned  to  Naples  because  he  wanted  to  connect  the  Station  with  an  aquarium  open 
to  the  public,  the  entrance  fees  thus  providing  the  means  to  pay  a  permanent 
assistant.  Dohrn  therefore  had  to  choose  a  large  city  that  attracted  many  tourists. 
Naples  at  that  time  was  still  one  of  the  largest  and  most  attractive  cities  of  Europe 
with  more  than  500,000  inhabitants  and  about  30,000  tourists  a  year  (Vogt,  1871). 

In  1870  Dohrn  went  to  Naples;  with  luck  he  overcame  doubts,  ignorance,  and 
misunderstandings  and  persuaded  the  city  authorities  to  give  him,  free-of-charge,  a 
plot  of  land  at  the  sea  edge,  in  the  beautiful  Royal  Park  (today,  the  Villa  Comunale). 
For  his  part  he  promised  to  build  a  Zoological  Station  at  his  own  expense.  Dohrn 
himself  gives  a  very  colorful  description  of  his  experiences  with  the  Neapolitan 
authorities  in  his  "History  of  the  Naples  Zoological  Station,"  which  he  started  to 
compile  in  1895.  (The  unfinished  manuscript  is  kept  in  the  Dohrn  Archives  at  the 
Stazione.) 

In  October,  1871,  Dohrn  installed  himself  with  all  his  equipment  and  books  at 
the  Palazzo  Torlonia,  near  the  small  port  of  Mergellina  in  Naples,  together  with  the 


10 


C.  GROEBHN 


biologist  E.  Ray  Lankester  (1847-1929).  The  two  friends  were  thus  able  to 
immediately  start  their  scientific  research.  The  foundations  of  the  Zoological  Station 
were  laid  in  March,  1872,  while  by  September,  1873,  the  whole  building  was 
finished  (Fig.  3).  Two-thirds  of  the  building  costs  came  out  of  Anton  Dohrn's  and 
his  father's  pockets,  the  remaining  third  was  provided  by  loans  from  friends. 

The  finished  building  contained  pumps,  machines,  store  rooms,  and  sea  water 
tanks  in  the  basement;  a  public  aquarium  on  the  ground  floor;  a  large  laboratory 
for  about  20  scientists  and  the  frescoe  room  with  the  library  on  the  first  floor;  and 
12  smaller  labs  and  living  quarters  for  the  custodian  and  assistants  on  the  second 
floor.  The  first  scientist  to  work  at  the  Station,  the  German  anatomist  Wilhelm 
Waldeyer  (1846-1921),  arrived  in  late  September,  1873. 

The  first  department  was  that  of  Morphology,  then  the  dominant  branch  of 
biology.  In  1876  a  botanical  department  was  created.  Although  there  was  no  room 
initially  in  the  main  building,  physiology  was  added  in  1882  by  renting  a  small 
building  near  the  Station.  The  space  problem  was  alleviated  by  adding  two  new 
sections  to  the  main  station,  one  in  1885-1888,  and  another  in  1903-1906.  A 
department  of  bacteriology  was  added  in  1887,  while  physiology  could  now  expand 
to  include  both  comparative  physiology  and  physiological  chemistry  (Figs.  4,  5).  In 
1906  Dohrn  had  his  son,  Reinhardt,  build  a  summer  home,  the  Villa  Acquario,  on 
his  beloved  island  of  Ischia;  today  the  Villa  Acquario  houses  the  Station's  Ecology 
Department. 

In  order  to  cover  the  running  expenses  of  the  institute  Dohrn  devised  the  so- 
called  "Table-System:"  against  an  annual  fee  the  contract  partner  (universities, 
ministries  of  education,  scientific  institutions,  private  individuals)  had  the  right  to 
nominate  one  scientist  to  use  a  table  for  one  year.  A  "table"  included  lab  space, 
fresh  animal  supply,  chemicals,  and  the  use  of  the  library  and  other  facilities.  All  of 


FIGURE  3.     The  Naples  Zoological  Station  in  1873.  (Dohrn  Archives) 


ANTON   DOHRN 


11 


the  arrangements,  financing,  and  guidance  of  day-to-day  activities,  were  supervised 
by  Dohrn  himself.  To  give  one  example  of  his  successful  managerial  skill:  by  1890, 
all  debt  on  the  original  building  and  the  first  addition  had  virtually  been  paid;  the 
annual  balance  showed  a  remarkable  profit  of  8,000  francs  (out  of  a  total  of  200,000 
francs  income);  and  36  tables  were  rented  annually  by  15  different  countries.  Dohrn 
also  started  a  specimen  supply  program  as  another  source  of  income.  Thanks  to  the 
inventiveness  and  skill  of  preparator  Salvatore  Lobianco  (1860-1910),  who  entered 
the  service  of  the  Station  at  the  age  of  14,  the  Zoological  Station  soon  became 
known  for  the  beauty  and  perfection  of  its  collections  of  preserved  marine  animals. 
These  were  sold  to  museums,  institutes,  and  individuals  all  over  the  world,  and 
many  samples  are  still  on  display  in  the  Station  today. 

Through  lectures,  conferences,  and  written  articles  Dohrn  continuously  tried  to 
make  the  Zoological  Station  known  to  a  wider  public.  He  always  solicited  international 
support  as  a  way  of  insuring  the  scientific  and  political  independence  of  the  Station. 
In  Germany  Dohrn  concentrated  on  interesting  influential  politicians,  scientists, 
industrialists,  dukes,  and  kings,  always  hoping  that  they  would  further  the  interests 
of  the  Station.  The  Naples  Station  soon  became  a  "must"  for  every  aspiring  biologist 
around  the  world  and  for  every  visitor  to  Naples.  In  1877  the  Berlin  Academy  of 
Sciences  and  the  Prussian  Ministry  of  Education  provided  funds  for  the  "Johannes 
Miiller,"  a  5-ton  steam  launch  which  served  for  both  collecting  and  excursion  trips. 
In  fact,  important  guests  to  the  station  were  usually  taken  on  an  excursion  on  the 
"Vaporetto"  to  one  of  the  many  beautiful  places  in  the  Gulf  of  Naples  (Fig.  6). 

In  1879-80  the  Zoological  Station  started  three  different  publications:  the 
Mittheilungen  aus  der  Zoologischen  Station  zu  Neapel  (vol.  1,  1879),  intended  for 


'  -^.  ___ 


_.. 


FIGURE  4.     The  Naples  Zoological  Station,  first  and  second  building.  November   1889.  (Dohrn 
Archives) 


12 


C.  GROEBEN 


-.-.' 


FIGURE  5.    The  Naples  Zoological  Station  today.  (Photo  Lab,  Stazione  Zoologica) 


the  research  results  of  staff  and  guest  scientists;  the  series  of  monographs  Fauna 
und  Flora  des  Golfes  von  Neapel  (vol.1,  1880)  as  an  inventory  of  the  Mediterranean; 
and  finally  the  Zoologischer  Jahreshcbericht  (Vol.  1,  1880),  a  reference  journal  that 
soon  became  famous  for  its  rapid  publication  and  accuracy. 

Although  Darwin  advised  Dohrn  that  establishing  a  library  would  be  too  great 
an  expense  and  consumption  of  time  (Groeben,  1982,  p.  29),  Dohrn  thought  that 
availability  of  all  the  major  published  sources  was  a  necessity  for  his  research 
institute.  He  gave  his  own  large  collection  to  the  Station,  and  got  publishers  and 
scientists — among  them  Darwin — to  donate  their  publications  in  Zoology  and 
related  fields.  The  Naples  Station's  biological  reference  collection  is  still  unrivalled 
in  Europe  today  (Fig.  7). 

Dohrn  was  also  able  to  obtain  some  of  the  latest  equipment  through  donations 
or  at  special  low  prices.  For  example,  Ernst  Abbe  (1840-1905)  of  the  Zeiss  factory, 
one  of  Dohrn's  few  close  friends,  allowed  the  Station  to  purchase  sets  of  Zeiss 
instruments  at  a  significant  discount;  in  return,  workers  at  the  Station  suggested 
ways  in  which  the  equipment  could  be  improved,  and  Zeiss  was  brought  to  the 
attention  of  the  international  scientific  community  (Fig.  8). 

Assistants  and  guests  collaborated  in  improving  section-cutting  and  staining 
methods,  thus  maintaining  the  high  level  of  technical  services  offered  by  the  Station. 
As  C.  O.  Whitman  (1883)  aptly  summarized  it  in  his  article  on  "The  advantages  of 
study  at  the  Naples  Zoological  Station,"  written  after  he  had  worked  at  Naples  in 
1881-82: 


ANTON   DOHRN 


13 


FIGURE  6.  "Johannes  Miiller"  (right)  and  "Frank  Balfour"  (left),  the  two  steam  launches  of  the 
Naples  Zoological  Station,  at  Mergellina.  On  the  left  in  the  background:  the  Naples  Station.  December 
1891.  (Dohrn  Archives) 


[The  Station  is]  a  sort  of  international  depot  for  the  reception  of  discoveries  and 
improvements  made  elsewhere.  The  heterogeneous  material  thus  obtained  is 
sifted,  systematized,  tested,  further  elaborated  and  refined  and  redistributed. 

When  the  25th  anniversary  of  the  foundation  of  the  Zoological  Station  was 
celebrated  in  1897,  nearly  2,000  scientists  presented  a  signed  address  to  Anton 
Dohrn,  saying,"  .  .  .  that  we  are  incapable  of  conceiving  what  the  present  state  of 
biological  science  would  be  without  the  influence  of  the  Zoological  Station"  (Dohrn, 
1897,  p.  13). 

In  1872  Dohrn  had  published  the  article  "Der  gegenwartige  Stand  der  Zoologie 
und  die  Griindung  zoologischer  Stationen"  (On  the  present  state  of  Zoology  and 
the  founding  of  Zoological  Stations)  (Dohrn,  1872b)  in  the  Preussische  Jahrbiicher, 
an  important  cultural  journal  widely  read  by  the  educated  public.  An  Italian 
translation  appeared  a  few  months  later  in  a  similar  type  of  journal.  This  essay  is 
often  quoted  as  one  of  the  classical  programmatical  writings  in  zoology,  for  it  lays 
out  clearly  and  openly  the  importance  of  marine  stations  for  the  future  of  biological 
research — a  future  that  had  been  seen  by  Dohrn  as  early  as  1868. 

Dohrn  explains  that  since  the  time  of  Darwin  zoology  has  entered  a  new  stage, 
leaving  mere  systematics  behind.  Experience  had  shown,  however,  that  all  practical 
efforts  to  promote  the  new  direction,  especially  the  study  of  marine  organisms 
usually  ended  in  a  waste  of  time,  money  and  energy,  because,  in  Dohrn's  view 
[zoology]  "lacks  organization"  (p.  4).  He  argued  that  future  efforts  should  concentrate 


14 


C.  GROEBEN 


FIGURE  7.  The  Naples  Zoological  Station  Library  in  the  frescoe  room.  The  cycle  of  frescos  depicting 
scenes  from  Neapolitan  life,  were  created  in  1873  by  the  German  painter  Hans  von  Marees  and  the 
German  sculptor  Adolf  von  Hildebrand.  1895.  (Dohrn  Archives) 


on  two  topics:  the  struggle  for  existence  and  natural  selection;  and  the  recapitulation 
of  phylogeny  in  the  development  of  the  individual  (i.e.,  embryology).  The  latter  was 
clearly  in  line  with  HaeckeFs  "Biogenetic  Law,"  while  the  former  anticipated  the 


ANTON   DOHRN 


15 


FIGURE  8.     Anton  Dohrn  in  his  office  at  his  Zeiss  microscope.  November  1889.  (Dohrn  Archives) 


study  of  populations  in  their  environment  and  animal  behavior.  In  addition,  new 
positions  in  zoology  (comparative  anatomy  and  embryology)  were  also  required  to 
guarantee  a  new  structure,  and  organization  for  zoology.  Zoological  Stations  would 
serve  a  crucial  function  as  "greenhouses"  for  young  zoologists. 

The  Zoological  Station,  by  providing  such  perfect  working  conditions,  fully 
answered  the  need  for  organization  that  Dohrn  saw  so  lacking  in  zoology  at  the 
time.  Good  organization  contributed  to  saving  money,  time,  and  energy  for  research. 
Dohrn  wanted  to  render  scientific  research  possible,  but  he  had  no  desire  to  interfere 
with  what  was  done  with  the  means  and  tools  he  put  at  the  disposal  of  his  guests. 
The  creation  of  such  a  complex  and  complicated  organism  as  a  research  station 
also  answered  Dohrn's  personal  need  "to  be  of  service  to  others."  To  his  satisfaction, 
he  had  created  something  new,  and  in  his  own  sphere  he  always  felt  like  a  monarch; 
he  compared  himself  often  to  Bismarck,  whom  he  greatly  admired.  Conscious  of 
his  achievements,  Dohrn  liked  to  call  himself  "the  Statesman  of  Darwinism." 

Dohrn  disdained  fame  and  honors,  although  he  wasn't  beyond  using  his  own 
numerous  decorations  if  they  could  be  of  use  in  gaining  further  support.  He  hoped, 
through  the  small  kingdom  he  had  created,  to  gain  power13  and  "to  exert  a  strong 
influence  on  the  outward  conditions  of  Science."  4  For  Dohrn  this  meant  influence 
on  those  who  made  or  represented  science. 

Almost  as  a  Leitmotiv,  Dohrn  in  his  letters  to  his  wife  continuously  quotes  from 
the  prologue  to  Goethe's  Faust:  "In  the  beginning  was  the  deed."  '  While  still  at 
Jena  he  had  decided  to  channel  all  his  efforts  into  doing,  acting,  and  creating,  aware 
that  this  meant  he  would  have  to  forego  such  other  interests  as  socialism,  poetry. 


16  C.  GROEBEN 

philosophy,  and  music.  Several  times  Dohrn  calls  the  Station  a  creation,  a  work  of 
art,  similar  to  sculptures,  poems,  or  paintings. 

At  this  point  we  should  consider  the  role  of  Anton  Dohrn's  own  scientific  work 
and  its  significance,  both  for  him  and  for  the  development  of  the  Station.  From 
1881  until  1907  he  published  25  "Studien  zur  Urgeschichte  des  Wirbelthierkorpers," 
thus  following  the  line  of  research — phylogenetic  studies  (i.e..  morphology) — for 
which  the  Zoological  Station  had  been  built.  While  the  Station  welcomed  new 
scientific  approaches — for  example,  the  Entwicklungsmechnik,  or  "developmental 
mechanics"  of  Roux  and  Driesch — during  the  nineties,  Dohrn  remained  a  mor- 
phologist,  using  the  Station's  facilities  as  did  any  other  table-holder.  Dohirfs  aim 
was  not  just  to  reconstruct  the  phylogenetic  tree  of  vertebrates;  his  ultimate  motive 
or  "main-spring"16  was  to  explain  the  history  of  human  form,  of  the  physical 
structure  of  man.  In  a  letter  to  E.  B.  Wilson  in  1900,  Dohrn  well  characterizes  his 
scientific  approach: 

Phylogeny  is  a  subtle  thing,  it  wants  not  only  the  analytical  powers  of  the 
"Forscher",  but  also  the  constructive  imagination  of  the  "Kiinstler", — and  both 
must  balance  each  other,  which  they  rarely  do, — otherwise  the  thing  does  not 
succeed.17 

With  Dohrn  it  was  very  often  imagination  that  prevailed,  which  was  invaluable 
when  creative  talents  were  called  for,  but  it  was  ill-fitted  for  scientific  research  where 
conclusions  must  be  drawn  from  facts,  and  not  facts  made  to  fit  into  conclusions. 

In  Dohrn's  life,  periods  of  practical  activity  concerning  the  Zoological  Station 
were  intertwined  with  periods  of  intense  scientific  study.  In  his  scientific  work, 
Dohrn  ranged  in  mood  and  attitude  from  vigorous  enthusiasm  for  the  broad  scope 
of  morphological  work  (for  example,  his  own  particular  idea,  the  Annelid-theory  of 
vertebrate  origin),  to  resigned  pessimism  during  which  he  limited  himself  only  to 
establishing  facts  and  details,  without  hope  of  ever  solving  the  larger  problem  of 
vertebrate  origin. 

One  cannot  help  feeling  that  Dohrn's  intense  activity  was  psychologically 
motivated  by  the  desire  to  prove  himself  to  those  who  had  doubted  his  ability  of 
ever  achieving  anything  in  the  way  of  serious  scientific  work.  Principal  among  those 
were  his  father  and  his  teachers  Ernst  Haeckel  and  Carl  Gegenbaur  (1826-1903), 
who  already  at  Jena  had  doubted  his  qualifications  as  a  scientist,  criticizing  his 
superficiality.  Dohrn  said  in  1872: 

I  have  been  ridiculized  [sic],  laughed  at,  they  have  told  me  that  I  had  no 
character,  that  I  were  [sic]  weak, — in  short  all  those  amiable  flatteries  which  can 
bring  a  man  to  despair  or  to  fury.  Then  they  doubted  my  beginnings,  called  me 
adventurer  and  phantast,  disbelieved  my  energy, — now  they  begin  to  give  me 
credit,  to  believe  in  me,  and  by  and  by  they  will  praise  me  and  admire  me.18 

Dohrn  was  flattered  that  E.  B.  Wilson  taught  his  works  and  theories,  especially 
the  Functionswechsel  (change  of  function)19  concept  in  his  student  courses  at 
Columbia  University.  He  was  especially  pleased  that  while  he  was  at  Woods  Hole 
in  August  of  1897,  he  was  asked  to  discuss  his  Vrspnmg  tier  Wirbelthiere.20  Without 
wanting  to  detract  from  the  thoroughness  of  Dohrn's  scientific  work,  it  must  be 
said  that  research  and  theorizing  was  only  one  part  of  his  life.  As  he  put  it  in  a 
letter  to  his  wife  in  1897: 

I  have  told  you  several  times  that  I  make  a  work  of  art  out  of  life  itself  ...  I 
found  chaos  before  me  and  have  created  out  of  that  both  a  practical  organism: 


ANTON   DOHRN 


17 


the  Station,  and  a  theoretical  one,  the  "Urgeschichte  der  Wirbelthiere".  Each 
step  on  the  path  of  these  two  things  I  have  visioned  beforehand,  as  an  artist  first 
sees  the  complete  work  of  art  and  then  starts  to  create  its  parts.21 

Dohrrfs  model  was  Goethe's  ideal  of  harmonic  humanity  (harmonische  Mensch- 
lichkeit)22,  that  is,  the  unity  and  harmony  between  what  one  can  do  and  what  one 
wants  to  do.  That  Dohrn  acquired  this  balance  is  shown  by  the  fact  that,  after 
having  absorbed  the  directive  influences  exerted  on  him  during  his  early  years,  from 
about  1870  on,  he  did  not  start  or  maintain  any  close  friendships.  By  then  he  relied 
only  on  himself.  This  was  true  with  regard  to  the  Station  and  also  with  regard  to 
his  family. 

Anton  and  Marie  Dohrn  had  four  sons  (Boguslav,  called  Bux,  b.  1875;  Wolf,  b. 
1878;  Reinhard,  b.  1880;  Harald,  b.  1885)  and  a  daughter  who  lived  only  one  year 
(1876-1877)  (Fig.  9).  Anton  Dohrn  was  a  loving  and  caring  father,  his  letters  to  his 
wife  are  full  of  plans  and  projects  about  how  to  help  their  sons'  intellectual 
development;  they  too  were  something  to  be  shaped  according  to  his  wishes. 

Although  the  administrative  and  social  duties  of  the  Station  were  a  heavy  burden 
for  him — and  his  health  suffered  in  consequence — Anton  Dohrn  would  not  share 
this  responsibility.  He  was  convinced  that,  having  to  answer  only  to  himself  he 


FIGURE  9.  Anton  Dohrn  and  his  family.  Standing,  left  to  right:  Reinhard  (1880-1962),  Bux  (1875- 
1960),  Wolf  (1878-1914),  and  Harald  (1885-1945)  Dohrn.  Sitting:  Mane  ( 1856-1918)  and  Anton  (1840- 
1909)  Dohrn.  Naples,  ca.,  1905.  (Dohrn  Archives) 


18  C.  GROEBEN 

could  keep  the  Station  free  from  bureaucracy  and  could  give  full  range  to  his 
"creative  imagination."' 

This  also  explains  why  there  were  no  strong  or  outstanding  characters  on  his 
staff.  Hugo  Eisig  (1847-1920),  Paul  Mayer  (1848-1923),  Wilhelm  Giesbrecht  (1854- 
1913),  and  Salvatore  Lobianco  were  all  very  efficient  in  their  specific  areas  and  they 
delivered  good  research,  but  they  weren't  personalities.  Strong  characters  like 
Nikolaus  Kleinenberg  (1842-1897)  (Muller,  1973),  J.  J.  von  Uexkiill  (1864-1944), 
or  Albrecht  Bethe  (1872-1954) — to  name  a  few — did  not  remain  long  at  the  Station, 
possibly  because  they  clashed  with  the  equally  strong-willed  Dohrn. 

Now,  how  did  Anton  Dohrn  actually  formulate,  create,  and  direct  his  Institute 
and  influence  science  in  general? 

First,  it  should  be  stressed  that  Dohrn  was  a  very  generous  and  charming  person 
who  was  unconcerned  by  social  differences  (Fig.  10).  He  was  accepted  by  everyone: 
the  fishermen  at  Mergellina  and  at  the  Station  discussed  their  problems  with  him; 
the  German  Emperor  chatted  politics  with  Dohrn  down  in  the  Aquarium;  and 
Carmen  Sylva,  the  Queen  of  Romania,  almost  fell  in  love  with  him.  Dohrn  put  at 
ease  guests  from  all  nationalities  and  backgrounds.  Hans  Driesch  reported  that 
Dohrn  used  to  say  "Here  we  are  a  family"  (1909,  p.  515);  in  fact,  Driesch, 
representing  a  completely  new  direction  of  scientific  work  (experimental  embryology), 
would  have  had  reason  enough  to  feel  unwanted,  but  he  did  not.  Scientific 
discussion,  along  with  very  heated,  though  enjoyable  arguments,  were  frequent;  but 
they  never  grew  personal. 

Dohrn  was  also  good  company,  taking  excursions,  enjoying  jokes  and  even 
childish  games.  He  even  installed  a  billiard-table  and  bowling-alley  at  the  Station, 
and  also  organized  concerts  and  promoted  literary  discussions  at  his  home.  Guests 
at  the  "Casa  Dohrn"  and  the  Zoological  Station  thus  experienced  science  and  art 
as  the  two  complementary  sides  of  European  culture  (Groeben,  1984)  (Fig.  1 1). 

Twice  Dohrn  was  offered  important  chairs  in  zoology,  in  1879  at  Naples  and 
four  years  later  at  Berlin;  both  times  he  refused.  He  did  not  like  teaching  and 
preferred  to  exert  his  personal  influence  through  organizing  and  administration  of 
science.  With  regard  to  the  Berlin  chair  he  explained  to  his  father:  "Here  at  the 
Station  I  can  better  influence  the  development  of  Zoology,  and  have  in  my  hands 
human  material  that  is  much  riper  than  students."  This  wish  to  mold  human 
material  was  also  reflected  in  the  selection  and  handling  of  his  staff. 

To  get  out  of  people  like  Eisig,  Mayer,  Meyer,  etc.,  what  is  possible,  for  a  great 
purpose,  this  is  the  touchstone  of  the  art  of  handling  people;  to  find  them  boring 
is  a  very  cheap  and  shortsighted  pleasure.24 

What  for  many  of  the  guest  researchers  was  a  perfect,  fruitful,  and  stimulating 
stay  at  Naples,  was  often  directed  and  organized  by  Dohrn;  one  cannot  help 
comparing  him  to  a  puppeteer.  One  example  of  this  is  his  statement: 

And  when  I  see  daily,  with  how  much  elan  and  how  much  joy  all  these  persons 
go  to  work  here,  who  come  from  outside,  and  how  I  succeed  through  the  timely 
interposition  of  excursions  on  the  steam-launch  to  keep  up  the  good  humour 
and  high-tensed  spirit — then  I  feel  deep  joy  about  the  results  of  hard  work  and 
fearful  years.25 

That  Anton  Dohrn  himself  was  a  well-known  and  respected  scientist  only 
contributed  to  the  stimulating  atmosphere.  From  him  guests  could  expect  under- 
standing of,  though  not  necessarily  agreement  with,  their  work.26  As  a  scientist  he 


ANTON   DOHRN 


19 


FIGURE  10.     Anton  Dohrn.  Sketch  by  Johannes  Martini,  a/..  1897.  (Dohrn  Archives) 

behaved  as  a  guest  himself,  not  influencing  what  was  done  at  the  Station,  only  how 
it  was  done.  As  an  impartial  diplomat  Dohrn  would  straighten  out  technical, 
scientific,  and  political  wrangles.  Unobtrusively,  he  dedicated  more  time  to  professors 


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ANTON   DOHRN  21 

than  to  younger  scientists,  because  he  respected  their  position  and  personality.  He 
also  appreciated  their  influence  in  directing  their  students  to  Naples,  and  of  course 
he  realized  that  it  was  good  publicity  to  attract  important  figures  like  August 
Weismann  (who  came  in  1877,  1882,  and  1887),  Carl  Vogt  (who  came  in  1879  and 
1884),  and  Emil  du  Bois-Reymond  (who  came  in  1878). 

Dohrn  was  also  an  excellent  public  relations  man,  constantly  seeking  to  extend 
the  name  of  the  Station  around  the  World.  For  example,  he  was  furious  at  C. 
Mereschkowsky,  of  St.  Petersburg,  because  in  several  communications  of  1882,  the 
latter  never  once  mentioned  that  his  results  had  been  obtained  at  Naples.27  Dohrn 
also  tried  to  promote  the  name  of  the  Station  through  solid  publications;  he  did  not 
approve  of  "preliminary  notes"  that  often  led  to  nothing  more,  and  excluded 
polemic  articles  in  the  Zoologischer  Janesbericht,  making  the  journal  well-respected 
and  a  must  for  all  libraries  and  research  workers  in  morphology. 

Summarizing,  it  may  be  said  that  as  far  as  the  Zoological  Station  was  concerned, 
the  creative  atmosphere  was  something  consciously  created  by  Anton  Dohrn,  who 
in  the  process  proved  himself  to  be  the  perfect  host  or,  the  perfect  statesman  of 
Zoology. 

Two  main  aspects  of  the  Naples  Station  have  inspired  the  foundation  of  similar 
institutes  all  over  the  world. 

The  first  "generation"  of  scientists  was  mostly  impressed  by  the  technical 
perfection  of  instruments  and  service  (animal  supply,  sea  water  tanks,  staining  and 
section-cutting  techniques,  library)  and  the  high  standard  of  research  offered  at 
Naples.  In  the  U.  S.  among  the  first  visitors  to  the  Station  were  E.  B.  Wilson,  C.  O. 
Whitman,  and  Emily  A.  Nunn,  all  of  whom  came  in  1882-1883,  having  learned  of 
the  wonderful  research  potentials  from  European  colleagues.  Wilson  had  applied  in 
January,  1883,  to  visit  Naples  to  work  on  the  development  of  Penatulida  "and  to 
learn  your  beautiful  methods  of  research."'28  After  returning  to  the  laboratory  of 
Rudolf  Leuckart  (1822-1898)  in  Leipzig,  after  his  stay  at  Naples,  C.  O.  Whitman 
had  complained,  "We  have  here  (at  Leipsic)  a  magnificent  building,  but  nothing  to 
work  with.  The  liberal  supply  at  Naples  contrasts  strongly  with  the  supply  of  tables 
here."29  And,  after  returning  to  the  United  States,  both  Whitman  and  Emily  Nunn 
wrote  glowing  accounts  in  Science  of  the  Station's  opportunities.  This  did  much  to 
spread  the  reputation  of  the  Station  to  younger  workers  in  the  United  States,  who 
subsequently  visited  Naples  in  the  1890's  and  early  1900's  (for  example,  T.  H. 
Morgan,  W.  M.  Wheeler,  Nettie  M.  Stevens,  among  others). 

The  idea  of  a  well-equipped  marine  station  for  pure  research  influenced 
Whitman  in  his  project  for  a  marine  station  at  Woods  Hole  that  concentrated  on 
summer  courses.  It  also  guided  Kakichi  Mitsukuri  (1858-1909)  in  founding  the 
Misaki  Marine  Laboratory  (1887)  and  E.  R.  Lankester  in  building  the  MBL  at 
Plymouth  (1888).  And  Spencer  F.  Baird  (1823-1887),  head  of  the  United  States 
Fish  Commission,  asked  Dohrn  for  technical  information  on  the  installation  of  the 
aquarium  and  the  sea  water  supply  in  1875  and  1883,  and  also  for  blueprints  and 
advice  for  the  fisheries  laboratory  he  was  building  at  Woods  Hole.  Anton  Dohrn 
was  delighted  that  Naples  had  become  a  standard  for  others,  but  he  was  also  anxious 
to  safeguard  Naples'  position  as  the  "mother-station." 

The  second  feature  of  the  station  that  so  impressed  and  influenced  visitors  to 
the  Naples  laboratory  was  its  complete  freedom  of  research.  This  freedom  of 
research,  of  being  host  to  the  international  scientific  community,  is  the  aspect  that 
has  also  survived  Anton  Dohrn's  personal  imprint.  Given  his  essentially  German 
background  and  his  independent  character,  the  Naples  Station  could  only  become 
a  "monarchy,"  whereas  significantly  enough  C.  O.  Whitman,  from  the  very 


22  ('•   GROHBliN 

beginning,  tried  to  put  the  MBL  on  a  democratic  basis  (Lillie,  1910,  p.  XXXI). 
Dohrn's  greatness  consisted  in  creating  an  organism,  the  potential  of  which  was 
larger  than  national,  contemporary  and  socio-political  ties. 

In  1888  President  G.  Stanely  Hall  of  Clark  University,  Worcester,  Massachusetts, 
visited  Naples  and  went  to  see  Anton  Dohrn  at  Berlin.  In  his  first  annual  report. 
Hall  stressed  where  Clark  University  had  or  had  not  followed  the  example  of  Naples 
(Clark  University,  1891,  p.  7;  1893,  p.  15). 

President  D.  C.  Oilman  (1831-1908)  of  Johns  Hopkins  University  visited  the 
Station  in  189030  and  again  in  1902  as  President  of  the  Carnegie  Institution,  which 
led  to  the  unexpected  rent  of  two  "Carnegie-Tables"  for  American  scientists  in 
December  1902.31 

The  Kaiser  Wilhelm  Institutes  (today:  Max  Planck  Institutes)  at  Berlin  were 
founded  as  research  institutions  dedicated  to  basic  science,  following  the  example 
set  by  Naples.  Through  the  offices  of  Julian  Huxley  (1887-1975)  the  Naples  Station 
later  served  as  a  model  for  the  constitution  of  UNESCO  research  programs  (Skalova, 
1975,  p.  1). 

Lastly,  one  of  the  most  striking  examples  of  the  influence  that  the  Naples  Station 
has  exerted,  concerns  the  Rockefeller  Institute  for  Medical  Research  in  New  York. 
Simon  Flexner  (1863-1946),  the  newly  appointed  Director  of  the  Institute  in  1901, 
while  on  a  year  long  recruiting  trip  through  Europe,  visited  Naples  in  November 
1903  and  spent  a  few  days  with  Anton  Dohrn  at  the  Station;  here  he  sought  advice, 
and  explained  to  Dohrn  and  others  about  the  Institute  planned  for  New  York.32  A 
few  days  later  Dohrn  asked  E.  B.  Wilson  in  a  letter:  "A  Dr.  Flexner  was  here:  he 
has  to  do  with  the  Rockefeller  Medical  Research  Institute.  What  does  that  mean? 
Is  it  a  serious  and  reliable  thing?"  3  Flexner  on  the  other  hand,  in  a  long  letter  to 
Christian  Herter  (1865-1910),  reported  on  his  impressions  of  Dohrn  and  the  Station. 
This  is  one  of  the  rare  lively,  personal,  immediate  impressions  about  Anton  Dohrn: 
therefore,  I  quote  it  extensively: 

.  .  .  The  most  important  event  of  my  trip  so  far  has  been  the  meeting  of  Dr. 
Dohrn  at  Naples.  Fortunately  he  was  most  courteous  and  permitted  me  to  see  a 
great  deal  of  himself.  Besides  several  meetings  in  his  laboratory  where  he  talked 
most  freely  of  his  plan  of  organization  and  its  results,  he  did  Helen  and  me  the 
courtesy  to  take  us  for  an  afternoon's  sail  on  the  Bay  of  Naples  in  the  Station 
Steamer  "Johannes  Miiller".  The  afternoon  was  perfect,  the  sun  shining  brightly 
on  a  placid  sea  of  the  deepest  azure  blue,  and  the  shores  with  Vesuvius,  Capri 
and  Ischia  in  the  background  were  of  the  loveliest  tints  [?]  and  form.  As  some 
of  the  Station  workers  were  along  I  found  it  easy  to  pass  much  of  the  time 
speaking  with  Dr.  Dohrn  whose  large  and  remarkable  experience  at  Naples 
supplied  an  endless  fund  of  entertaining  descriptions. 

Dohrn  is  now  63,  a  large  hale  man,  of  large  voice  and  pronounced  manners. 
His  love  for  science  is  an  enthusiasm  which  is  probably  greater  today  than  when 
at  30  he  conceived  the  idea  of  the  Zoological  Station — the  first  station  of  the 
kind  in  the  history  of  the  world.  He  is  a  great  man — of  that  I  have  no  doubt.  I 
think  he  impressed  me  more  favorably  and  strongly  than  anyone  I  ever  met 
before.  .  .  . 

He  advised  me  to  begin  in  a  small  way  and  was  delighted  with  the  idea  of 
putting  the  first  force  into  a  small  house  where  the  early  organization  can  be 
completed  and  from  where  a  move  can  readily  be  made  into  the  larger  laboratory. 

But  the  advice  that  he  urged  most  strongly  was  "freedom".  He  has  never 
acted  as  censor  to  his  workers.  They  come  to  him,  everything  is  supplied  them 
for  their  problems,  but  their  results  are  their  own.  Any  discoveries  they  make 
are  theirs;  and  blunders  theirs  too.  "Public  opinion,  not  I,  is  the  censor"  was  his 


ANTON   DOHRN  23 

repeated  statement.  "Men  work  here,"  he  said,  "in  a  dozen  different  branches  of 
biological  science,  can  I  be  authority  on  them  all."  "No,  no,  give  them  perfect 
freedom;  let  them  search  where  and  how  they  will;  help  them  in  every  way  that 
you  can,  but  do  not  pretend  to  be  master  over  them."  It  was  a  remarkable 
pronouncement  and  coming  from  such  an  authority  and  one  of  the  most 
successful  research  leaders  of  the  world,  worthy  of  the  most  thoughtful  consid- 
eration. And  the  more  I  have  thought  over  the  subject  the  more  I  have  come  to 
his  point  of  view.  I  wonder  how  it  impresses  you? 

He  is  putting  up  now  a  third  building — for  Physiology.  Of  late  comparative 
physiology  has  grown  with  such  rapidity  that  a  special  laboratory  is  demanded. 
His  enthusiasm  in  the  subject  is  delightful  and  he  expects  important  returns  from 
this  branch  of  work.  "Dog  and  Cat  physiology"  as  he  called  it  have  about  reached 
their  limit  of  value:  what  we  require  is  a  study  of  simpler  forms  for  the  unravelling 
of  complex  phenomena  and  to  render  their  understanding  possible.  It  was 
splendid  to  hear  him  exclaim  over  "Science"  and  say  again  and  again  in  reference 
to  this  exertion,  or  that  privation,  or  a  specially  striking  illustration,  "But  it  is  all 
for  Science."' 

ACKNOWLEDGMENTS 

1  wish  to  thank  Dr.  Antonietta  Dohrn  and  Dr.  Peter  Dohrn,  Mr.  James  Thomas 
Flexner,  and  the  Rockefeller  University  Archives  for  the  permission  to  quote  from 
Anton   Dohrn's  correspondence,   and  from   Simon   Flexner's  letters  to  Christian 
Herter,  respectively. 

NOTES 

'Charles  Grant  (1841-1889),  Scottish  poet  and  literary  critic  and  close  friend  of  Anton  Dohrn. 
Dohrn  quotes  this  poem  in  a  letter  to  his  wife  as  referring  to  his  own  life.  Anton  Dohrn  to  Marie  Dohrn. 
Aug.  5,  1886,  Naples.  Bd  379.  Numbers  and  letters  (e.g.,  Bd  379.)  indicate  reference  numbers  of  the 
Archives  of  the  Naples  Zoological  Station  (ASZN).  If  not  mentioned  otherwise,  quotations  are  translated 
from  German. 

2  Anton  Dohrn  to  Marie  Dohrn.  Aug.  1883,  Stettin.  Bd  246. 

3  Th.  Heuss.  Bemerkungen  zu  dem  Plan  einer  Anton  Dohrn  Biographic.  1939.  3  pp.,  typescript.  Be. 
1939.  H. 

4  Anton  Dohrn  to  Fanny  Lewald  and  Adolf  Stahr,  April  19.  1866.  Jena.  Ba  1178. 

5  F.  de  Filippe,  quoted  by  Francesco  Todaro  in  his  speech  on  the  occasion  of  the  25th  anniversary 
of  the  foundation  of  the  Naples  Zoological  Station,  April  14,  1897.  Dohrn,  1897,  pp.  5-10:  8. 

6  Anton  Dohrn  to  Fanny  Lewald  and  Adolf  Stahr,  Oct.  17,  1868,  Messina.  Ba  1241. 

7  Anton  Dohrn  to  Carl  August  Dohrn,  Feb.  15.  1869.  Messina.  Be  16. 

8  Anton  Dohrn  to  Thomas  Henry  Huxley,  April  24,  1870,  Naples,  Ba  473,  and  to  Fanny  Lewald 
and  Adolf  Stahr.  July  6,  1870,  Stettin.  £«  1278. 

9  Anton  Dohrn  to  Marie  Dohrn,  Aug.  1,  1886.  Naples.  Bd  372:  and  May  15,  1890,  Naples.  Bd  681. 
111  Anton  Dohrn  to  Fanny  Lewald  and  Adolf  Stahr,  April  15,  1866,  Jena.  Ba  117: 

"  Anton  Dohrn,  unfinished  manuscript,  September  1874,  Hoekendorf,  Bd  0173. 
12  Anton  Dohrn  to  Marie  Dohrn,  July  1,  1884,  Hoekendorf.  Bd  283. 

"Anton  Dohrn  to  Mane  de  Baranowska-Dohrn,  Nov.  8,  1873,  Naples,  Bd  0114:  Oct.  2.  1881. 
Berlin.  Bd  207. 

14  Anton  Dohrn  to  Marie  Dohrn,  June  6,  1884,  Moscow.  Bd  268. 

15  Anton  Dohrn  to  Marie  Dohrn,  Aug.  22,  1888,  Hoekendorf.  Bd  543. 

16  Anton  Dohrn  to  Marie  Dohrn.  Dec.  24,  1889,  Naples.  Bd  648. 

17  Anton  Dohrn  to  E.  B.  Wilson,  Feb.  20,  1900,  Naples.  Bd  846  (in  English). 

'"  Anton  Dohrn  to  Marie  de  Baranowska,  Dec.  31.  1872,  Berlin.  Bd  020  (in  English). 
14  E.  B.  Wilson  to  Anton  Dohrn,  April  13,  1909,  New  York.  G, XXXI. 46. 

20  Anton  Dohrn  to  Marie  Dohrn,  (Aug.)  13,  1897,  Detroit.  Bd  1101. 

21  Anton  Dohrn  to  Marie  Dohrn,  Aug.  1,  1886,  Naples.  Bd  372. 

22  Anton  Dohrn  to  Marie  Dohrn,  Dec.  27,  1884,  Berlin.  Bd  327/28. 

23  Anton  Dohrn  to  Carl  August  Dohrn,  May  17,  1883.  Naples.  Be  33. 

24  Anton  Dohrn  to  Marie  Dohrn.  July  7.  1885,  Naples.  Bd  353. 


24  C.  GROEBEN 

;s  Anton  Dohrn  to  Marie  Dohrn,  April  1  1,  1890,  Naples,  lid  670. 

16  Margret  Boveri  to  Theodor  Heuss,  July  5,  1939,  Stockholm,  copy  for  Reinhard  Dohrn.  Be, 1939, H. 

21  Anton  Dohrn  to  Marie  Dohrn,  July  29/30,  1882,  Stettin.  Bd  223. 

2S  E.  B.  Wilson  to  Anton  Dohrn,  Jan.  29,  1883,  Cambridge.  A,1883,W. 

^C.  O.  Whitman,  May  1882.  This  passage  has  been  copied  by  Paul  Mayer  on  a  letter  from  J. 
Barrois  (original  missing).  A, 1882, B. 

30  D.  C.  Oilman  to  Anton  Dohrn,  Nov.  7,  1894,  Baltimore.  AJ894.G. 

11  From  Anton  Dohrn's  letters  and  from  the  Presidential  Files  of  the  Carnegie  Institution  (personal 
communication,  James  D.  Ebert)  it  shows  that  the  two  "Carnegie-Tables"  were  rented  thanks  to  the 
insistence  of  E.  B.  Wilson. 

32  Simon  Flexner  to  Anton  Dohrn,  Nov.  4,  1903,  Naples.  A.  1903, F 

33  Anton  Dohrn  to  E.  B.  Wilson,  Nov.  11,  1903,  Naples.  Ba  853. 

34  Simon  Flexner  to  Christian  Herter,  Nov.  21,  1903.  Perugia.  Rockefeller  University  Archives  RG 
417  Folder  7a  #5.  Quoted  with  permission  from  the  Rockefeller  Archives  and  J.  T.  Flexner,  New  York. 

J5  Nikolai  Nicklolajewitsch  Mikloucho-Maclay.  Letter  to  Anton  Dohrn,  Jan.  1869.  Messina  BA  735. 
Published  in  I.  Miiller.  1980.  p.  39. 


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Gra:  am  18.  August  1910.  S.  Hirzel,  Leipzig.  43  pp.  Reprinted  in:  Naturwiss.  28  (1940):  787- 

798;  and  in:  Simon,  1980,  pp.  106-149. 

Clark  University,  1891.  Second  Annual  Report  oj  the  President  .  .  .  Worcester,  Massachusetts. 
Clark  University,  1893.  Third  Annual  Report  of  the  President  .  .  .  Worcester,  Massachusetts. 
DEACON,  M.  1971.  Scientists  at  the  Sea  1650-1900.  a  Study  oj  Marine  Science.  Academic  Press,  London, 

New  York.  445  pp. 
DOHRN,  A.  1871.  Kurzer  Abriss  der  Geschichte.  sowie  Gutachten  und  Meinungsausserungen  hervorragender 

Naturjorscher  fiber  die  Griindung  der  Zoologi.se/ien  Stalionen.  Neapel.  8  pp.  Reprinted  in:  Simon, 

1980,  pp.  13-20. 

DOHRN,  A.  1872a.  The  foundation  of  Zoological  Stations.  Nature  5:  277-280;  437-440. 
DOHRN,  A.    1872b.   Der  gegenwartige  Stand  der  Zoologie  und  die  Griindung  zoologischer  Stationen. 

Prems.  Jb..  30:  137-161.  Reprinted  in:  Naturwiss..  19  (1926):  412-424;  and  in:  Simon,  1980. 

pp.  23-46.  Italian  translation  in:  Nuova  Antologia,  Jan.  1873:   1-27.  Reprinted  in:  Boll.  Zool. 

35(1968):  507-531. 
DOHRN,  A.    1897.  Das  25  jdhrige  Jubiliium  der  Zoologischen  Station  :u  Neapel  am   14. April  1897. 

Breitkopf  &  Hartel,  Leipzig.  44  pp.  Reprinted  in:  Simon,  1980,  pp.  61-104. 

DOHRN,  K.  1983.  I 'on  Biirgern  und  \Veltbiirgern.  Fine  Familiengeschichte.  G.  Neske,  Pfullingen.  272  pp. 
DRIESCH,  H.  1909.  Zur  Erinnerung  an  Anton  Dohrn.  Siiddeutsche  Monatshejte  (Nov.):  513-518. 
GROEBEN,  C.,  ed.  1982.  Charles  Darwin — Anton  Dohrn.  Correspondence.  Macchiaroli,  Napoli.  118  pp. 
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(Exhibition  Catalogue).  Naples.  110  pp. 

HEUSS,  T.  1962.  Anton  Dohrn.  Rainer  Wunderlich,  Tiibingen.  448  pp.  (1st  ed.:  1940;  2nd  ed.:  1948). 
KOFOID,  C.  A.   1910.  The  Biological  Stations  of  Europe.  Washington.  360  pp.  (Bull.  U.  S.  Bureau  of 

Education  no.  4). 
KUHN,  A.   1950.  Anton  Dohrn  und  die  Zoologie  seiner  Zeit.  Puhbl.  Sta:.  Zool.  Napoli,  Suppl.   1950. 

205  pp. 
LANCE,  F.  A.   1866.  Geschichte  des  Materialismus.  Iserlohn.  New  edition:  A.  Schmidt,  ed..  Frankfurt, 

1974. 

LILLIE,  F.  R.  1910.  Charles  Otis  Whitman  (1842-1910).  J.  Morphol.  22:  XV-LXXVII. 
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heute — ein  "permanenter  Kongress".  Naturwiss.  Rundschau  37:  429-437. 
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Reference:  Hiol.  Hull   I6S  (suppl.):  26-34.  (June,  1985) 


AGASSIZ,   HYATT,   WHITMAN,   AND  THE   BIRTH   OF 
THE   MARINE   BIOLOGICAL   LABORATORY 

JANE   MAIENSCHEIN 

Department  of 'Philosophy.  Arizona  State  University.  Tempe.  Arizona  85287 

ABSTRACT 

This  paper  establishes  that  the  MBL  began  as  a  self-consciously  American 
marine  laboratory,  following  the  lead  of  its  American  predecessors.  In  particular, 
Louis  Agassiz's  School  of  Natural  History  at  Penikese  Island  and  Alpheus  Hyatt's 
Laboratory  for  instruction  at  Annisquam,  Massachusetts,  directly  inspired  the  MBL. 
Archival  sources  reveal  the  connections  and  the  MBL's  goals.  Teaching  and  research 
were  accepted  as  the  dual  and  compatible  goals  for  the  Laboratory,  and  it  was  left 
to  the  first  director,  Charles  Otis  Whitman,  to  work  out  how  best  to  combine  the 
two.  These  emphases  and  the  clientele  thus  attracted  clearly  distinguished  the  MBL 
from  the  European  laboratories,  such  as  the  Naples  Zoological  Station,  which 
concentrated  on  independent  research.  In  part  the  MBL  achieved  success  because 
both  the  teaching  and  research  focused  on  shared  basic  questions,  namely  develop- 
mental questions  posed  within  a  solid  morphological  tradition.  Epigenesis  and 
preformation,  the  role  of  cells  in  development,  cell  lineage  study  of  early  egg 
organization:  such  themes  ran  through  most  of  the  work  done  at  the  MBL  in  the 
first  year. 

DISCUSSION 

Over  the  years  a  number  of  myths  have  appeared  about  the  early  history  of  the 
MBL,  some  better  than  the  truth — as  is  the  way  with  myths.  One  such  story, 
reported  to  the  New  York  Times,  is  representative.  There  David  Starr  Jordan,  who 
was  then  a  biologist  at  Stanford,  discussed  the  founding  of  the  MBL.  He  reported 
that  "Senator  F.  Baird,  Secretary  of  the  Smithsonian  Institution,  .  .  .  and  his 
associates  met  in  1888  and  formally  organized  a  corporation,  separate  from  the 
Bureau  of  Fisheries,  to  carry  on  the  work"  of  marine  biology  (Jordan,  1926). 
Fortunately  Jordan  was  a  better  biologist  than  he  was  an  historian.  Actually  Spencer 
(not  Senator)  Fullerton  Baird  had  died  in  1887  so  was  unlikely  to  have  done  much 
incorporating  in  1888.  Other  myths  include  the  claim  that  the  MBL  was  simply  a 
copy  of  the  Naples  Zoological  Station  (Lillie,  1944,  pp.  14-15). 

I  shall  examine  more  closely  the  foundation  of  the  MBL,  and  concentrate  on 
two  points:  (1)  The  MBL  was  established  as  a  self-consciously  American  marine 
laboratory,  even  while  it  reflected  influence  from  Naples,  and  (2)  despite  the  declared 
desire  to  include  all  of  biology,  it  was  the  concentration  of  research  and  instruction 
around  shared  concerns,  particularly  developmental  problems,  which  brought  about 
the  MBL's  early — and  continuing — success. 

Beginning  with  the  first  claim,  that  the  MBL  began  as  an  American  effort,  I 
shall  discuss  briefly  the  several  cornerstones  in  the  lab's  foundations.  These  include 
the  influence  of  Louis  Agassiz,  of  Alpheus  Hyatt,  and  especially,  of  Charles  Otis 
Whitman.  Others  then  constructed  a  strong  edifice  on  the  solid  foundation. 

Agassiz's  summer  school  at  Penikese  Island  provided  the  initial  vital  stimulus. 
As  the  MBL's  first  director  Whitman  said  repeatedly,  the  MBL  was  a  lineal 
descendent  of  its  genetic  ancestors,  Agassiz's  Penikese  School  and  Hyatt's  Annisquam 

26 


BIRTH   OF  THE   MBL  27 

laboratory  (Whitman,  1883,  1903).  For  some  time,  Agassiz  had  considered  the 
prospect  of  running  a  summer  school  to  provide  students  of  natural  history  with 
practical  experience.  In  1873,  he  finally  gained  financial  backing  from  a  wealthy 
New  Yorker  and  opened  the  Anderson  School  of  Natural  History  on  Penikese 
Island.  The  clientele  was  to  be  school  teachers  who  sought  field  experience  to  inform 
their  classroom  instruction;  thus  the  lab  was  oriented  toward  instruction  rather  than 
original  research  (Agassiz,  1885,  chapter  25;  Wilder,  1907;  Morse,  1923;  Conklin, 
1927;  Wright  and  Wright,  1950;  Dexter,  1974).  The  school  opened  in  1873  with 
about  fifty  attendees.  According  to  one  report  the  women  were  very  "schoolma'amy" 
and  "the  gentlemen  are  not  a  whit  behind"  ("Penikese  Island,"  1873,  p.  378).  Yet 
the  group  appeared  earnest  and  eager,  the  same  reporter  acknowledged,  and  the 
group  included  four  individuals  of  particular  importance  for  the  MBL:  Alpheus 
Hyatt,  William  Keith  Brooks,  Charles  Otis  Whitman,  and  Cornelia  Clapp.  Hyatt 
was  a  lecturer  rather  than  a  student,  and  it  was  Hyatt  who  was  to  become  the  real 
father  of  the  MBL  (Dexter,  1974,  p.  159). 

Agassiz  was  a  master  of  publicity  and  made  the  first  day  a  real  show — unlike 
the  first  day  at  the  MBL.  The  students  and  a  number  of  guests  met  on  the  dock  in 
New  Bedford  and  went  together  by  steamer  to  Penikese.  There  all  were  treated  to  a 
dinner  in  the  newly  (and  rather  hastily)  constructed  buildings  and  to  an  inspiring 
informal  convocation.  One  student  admitted  that  after  the  guests  had  departed  and 
the  show  was  over  that  the  reality  of  the  island  proved  a  bit  discouraging.  There 
they  were  stuck  on  an  island  about  %  mile  long  and  'A  mile  wide,  which  was 
virtually  barren  of  trees  or  other  accoutrements  (Anonymous,  1895,  p.  21).  The 
student  did  not  have  long  to  fret,  however;  work  began  immediately  the  next 
morning  and  consumed  all  available  time  for  the  duration  of  the  summer — except 
Sundays  when  most  of  the  students  refused  to  carry  out  ungodly  biological  studies. 

Some  popular  accounts  give  the  impression  that  the  students  spent  their  days 
wandering  idly  about  the  island  collecting  things  without  purpose.  It  is  true  that  the 
instruction  was  highly  individualized,  with  each  student  spending  a  good  part  of 
each  day  exploring,  collecting,  observing,  recording,  and  generally  studying  nature 
rather  than  books — as  Agassiz  instructed.  Yet  good  books,  not  mere  repetitive 
textbooks,  did  have  their  place.  So  did  lectures.  Agassiz  invited  a  number  of 
important  biologists  to  lecture  to  the  group  on  a  range  of  natural  history  topics 
(Popular  Science  Monthly,  1874).  In  fact,  each  day  began  with  structured  lectures, 
followed  by  an  hour  or  so  of  dissection.  Afternoons  often  brought  freedom  to  roam 
and  collect,  but  students  spent  most  evenings  attending  lectures,  dissecting  by 
candlelight,  and  writing  up  their  notes  from  the  day's  work.  Such  a  system  obviously 
worked  best  for  those  students  capable  of  framing  their  own  questions  and  following 
through  with  relevant  collecting,  but  Agassiz  and  his  invited  speakers  helped 
articulate  appropriate  problems  as  well. 

Not  everyone  approved  of  Agassiz's  school.  The  highly  respected  British  naturalist 
E.  Ray  Lankester  admitted  that  what  he  called  "the  spasmodic  descent  upon  the 
sea-coast"  offered  a  very  nice  vacation  for  naturalists  who  could  not  otherwise 
afford  such  luxuries.  Such  trips  might  even  result  in  collection  of  a  few  new  species. 
he  admitted.  But,  Lankester  insisted,  "it  is  not  in  this  way  that  the  zoology  of 
to-day  can  be  forwarded"  (Lankester,  1880,  pp.  497-499).  A  naturalist  needs  to 
work  at  settling  "important  questions,"  he  believed.  Even  Whitman  recalled  that  at 
first  he  found  Agassiz's  methods  unproductive,  but  that  he  soon  came  to  admire 
them  and,  indeed,  incorporated  some  into  the  approach  of  the  MBL  (Craig,  1910). 

Agassiz's  Penikese  School  continued  for  a  second  year,  despite  Louis's  death  in 
1873  and  his  son  Alexander's  illness  in  the  second  session  of  1874.  Then  it  closed. 


28  J.   MAIENSCHEIN 

Not  for  lack  of  funds,  as  Jordan  and  others  have  claimed,  but  more  for  lack  of 
anyone's  having  taken  the  initiative  to  keep  it  going  (G.  R.  Agassiz,  1913,  pp. 
129,  131).' 

In  1879  one  of  the  Penikese  students,  Alpheus  Hyatt,  began  another  seaside  lab 
in  Annisquam,  on  Cape  Ann  in  Massachusetts  (Dexter,  1952;  Boston  Society  of 
Natural  History;  Kohlstedt,  1979).  This  laboratory  had  a  purpose  closely  following 
that  of  the  Penikese  School.  Intended  to  provide  opportunities  for  science  teachers 
to  observe  and  study  marine  animals,  the  lab  was  also  the  inspiration  of  the  Boston 
Society  of  Natural  History  and  was  supported  by  the  Woman's  Education  Association 
of  Boston.  Hyatt  served  as  director,  with  the  Boston  Society  Assistant  B.  H.  van 
Vleck  (who  had  been  a  student  at  Penikese)  as  instructor.  After  two  years  in  Hyatt's 
house,  the  lab  moved  to  a  separate  location  nearby  in  1881.  Then,  with  the 
continuing  financial  aid  of  the  Woman's  Education  Association,  the  Annisquam 
Laboratory  operated  as  a  department  of  the  Society  of  Natural  History.  Clearly 
Hyatt's  ideals  helped  direct  the  effort,  but  the  specific  purpose  of  providing 
educational  opportunities  for  instructing  science  teachers  came  from  the  Boston 
Society,  for  which  Hyatt  was  the  Curator.  At  times  the  level  of  the  students' 
commitment  and  preparation  seemed  hopelessly  low.  As  Mrs.  Hyatt  wrote  to 
Alpheus  while  he  was  at  sea  on  an  expedition,  the  group  was  very  uninteresting, 
even  tedious.  They  were  essentially  raw  recruits,  hopelessly  elementary  students  who 
were  beginning  to  drive  van  Vleck  to  despair  (Dexter,  1956-1957).  But  the  school 
did  attract  a  few  men  such  as  Thomas  Hunt  Morgan  who  certainly  became  a 
serious  researcher  and  one  of  the  backbones  of  the  MBL. 

In  1887,  the  Woman's  Education  Association  decided  that  the  project  had 
succeeded  and  that  they  would  withdraw  support  since  they  held  the  goal  of  seeding 
projects  until  they  caught  on,  then  leaving  them  on  their  own  (Hyatt,  1887).  The 
Annisquam  project  seemed  a  success.  But  Hyatt  was  rather  tired  and  wished  to 
develop  an  American  marine  laboratory  on  an  independent  basis:  an  institution 
separate  from  the  Society  of  Natural  History  and  from  himself  as  director.  He  also 
felt  that  a  new  site  would  prove  preferable  to  that  of  Annisquam,  which  was 
becoming  polluted.  Thus  came  the  move  to  Woods  Hole. 

Why  Woods  Hole?  The  answer  lies  largely  with  Spencer  Baird.  For  several  years, 
Baird  had  wanted  his  friend  Hyatt  to  move  the  Annisquam  school  to  Woods  Hole, 
which  had  purer  water,  more  abundant  marine  life,  a  congenial  setting  and,  not 
coincidentally,  was  home  of  the  United  States  Fish  Commission  which  Baird 
headed  (Galtsoff,  1962;  Boston  Society  Minutes,  1888,  pp.  563-564).  Baird  wanted 
to  attract  researchers  and  students  to  form  a  research  community  at  the  Fish 
Commission.  In  some  details  he  seems  to  have  been  influenced  by  the  research 
emphasis  of  the  Naples  Zoological  Station,  opened  in  1872.  At  first  his  efforts 
seemed  to  be  succeeding  (Galtsoff;  1962,  p.  29;  Whitman,  1883,  p.  97;  Parker,  1946, 
p.  136).  But  the  connection  of  the  Fish  Commission  with  the  government  and  its 
mandate  to  investigate  practical  fisheries-related  questions  made  it  very  difficult  for 
him  to  develop  in  the  same  way  as  the  more  independent  Naples  Lab.  Baird  did 
attract  cooperation  from  the  Johns  Hopkins  University,  which  sent  Professor 
William  Keith  Brooks  and  some  students  to  the  Fish  Commission,  and  from 
Princeton  and  Harvard.  Yet  Baird  failed  to  gain  the  necessary  financial  support  to 

1  G.  R.  Agassiz  says  that  Alexander  was  always  against  the  Penikese  lab  and  that  the  financial 
situation  became  impossible  when  Anderson  withdrew  his  support  after  the  second  year.  Letters  from 
Alexander,  May  30  and  June  23,  1888,  indicate  that  he  probably  felt — probably  not  quite  fairly — that  he 
had  tried  to  maintain  the  Penikese  and  other  marine  labs  and  had  received  no  support  from  others. 


BIRTH   OF  THE   MBL  29 

attract  other  researchers  and  to  establish  a  permanent  research  lab  in  the  1880's  in 
Woods  Hole. 

In  1887  as  Hyatt,  the  Woman's  Education  Association,  and  the  Boston  Society 
of  Natural  History  began  to  consider  sites  for  their  laboratory,  they  did  find  Woods 
Hole  attractive.  Baird  had  helped  the  Annisquam  school  by  sending  specimens.  He 
had  urged  a  friend  to  buy  land,  near  the  Fish  Commission,  which  was  held  for  the 
benefit  of  any  educational  institution  that  might  build  there.  He  had  welcomed 
Hyatt  at  Woods  Hole.  When  the  MBL  was  incorporated  in  1888  the  Trustees  chose 
Woods  Hole  as  their  site,  and  looked  to  the  Fish  Commission  for  further  encour- 
agement. 

With  Hyatt  as  president,  the  MBL  trustees  decided  to  hire  Johns  Hopkins 
Professor  of  Zoology  William  Keith  Brooks  as  the  first  director  (MBL  Minutes, 
1888,  pp.  11-13).  Hyatt  knew  Brooks,  and  had  recommended  him  for  his  job  at 
Hopkins  (Oilman  Papers).  Perhaps  Brooks  would  take  the  job  without  pay,  Hyatt 
suggested,  and  perhaps  the  Hopkins  would  lend  financial  support  to  the  laboratory 
effort.  Now,  you  know  that  the  first  director  was  actually  Charles  Otis  Whitman. 
After  all,  there  is  no  Brooks  laboratory  building  at  the  MBL  these  days.  Brooks 
turned  down  the  offer.  Why,  you  may  ask?  Why  would  anyone  turn  down  the 
opportunity  to  become  first  director  of  America's  first  permanent  research  laboratory 
for  marine  biology?  Why  would  anyone  reject  the  chance  to  summer  in  Woods 
Hole? 

The  Trustees  offered  no  salary  at  first,  but  that  alone  probably  would  not  have 
deterred  Brooks.  Who  was  this  man,  then,  who  rejected  his  chance  to  become 
immortalized  at  the  MBL?  Brooks  was,  quite  simply,  the  zoologist  with  the  best  job 
in  America  at  the  time.  He  was  the  only  professor  of  morphology  at  the  American 
research  university.  A  student  of  Agassiz's  and  participant  in  the  Penikese  School, 
he  was  teacher  of  Edmund  Beecher  Wilson,  Thomas  Hunt  Morgan,  Edwin  Grant 
Conklin,  Ross  Granville  Harrison,  and  others  who  assumed  central  importance  for 
MBL  history  and  for  the  history  of  biology  in  general.  He  was  also  founder  and 
director  of  the  most  significant  marine  research  lab  in  America  to  date,  the 
Chesapeake  Zoological  Laboratory,  run  by  the  Johns  Hopkins  (McCullough,  1969; 
Benson,  1979,  1985;  Oilman  Papers).  The  Chesapeake  Laboratory  was  an  informal 
arrangement  each  summer  where  Hopkins  graduate  students,  usually  accompanied 
by  Brooks,  explored  marine  life  in  one  or  another  location,  ranging  from  Beaufort, 
North  Carolina  to  Jamaica  or  Bermuda  (Chesapeake  Zoological  Laboratory  Reports, 
Oilman  Papers).  Brooks's  reports  to  the  Hopkins  President  about  these  sessions 
reveal  his  enthusiasm,  but  clearly  show  that  his  leadership  style  was  best  for  a  very 
few  specially  selected  men  at  the  Chesapeake  Laboratory  (and  one  woman,  once— 
Emily  Nunn,  later  Whitman's  wife).  Brooks  liked  the  summer  research  trips,  and 
he  liked  Woods  Hole  during  his  visits  at  the  Fish  Commission.  But  Brooks  was  one 
of  the  most  unassuming,  retiring,  and  unlikely-to-be-director  sorts  of  men  imaginable. 
Perhaps  he  lacked  vision.  He  did  not  believe  that  Woods  Hole  could  support,  or 
should  support,  two  research  labs  in  marine  biology.  He  chose  rather  to  ally  the 
fate  of  his  Chesapeake  Zoological  Laboratory  with  the  Fish  Commission. 

After  Baird's  death  in  1887,  the  next  Fish  Commissioner,  Colonel  McDonald, 
wished  to  expand  investigation  at  his  lab.  He  encouraged  Brooks  to  work  there  as 
a  consultant  and  researcher  and  to  bring  a  few  of  his  students  as  well  (Oilman 
Papers).  Brooks  was  happy  with  the  Fish  Commission  and  was  therefore  never 
convinced  that  the  MBL  was  a  good  idea.  He  believed  in  1888  that  McDonald  was 
making  progress  in  improving  the  Fish  Commission  as  a  research  facility.  No  other 


J.   MAIENSCHE1N 

lab  was  needed,  he  felt,  and  especially  not  in  Woods  Hole.  As  he  wrote  of  the  MBL 
idea, 

I  said  all  that  I  could  to  convince  Sedgwick  [one  of  the  MBL  Trustees]  that 
the  Boston  Laboratory  would  be  much  more  valuable  if  some  other  place  than 
Woods  Hole  were  selected,  so  that  naturalists  might  have  the  benefit  of  stations 
at  two  points,  and  if  McDonald  is  able  to  carry  out  his  plans  and  to  open  this 
laboratory  to  investigators  in  future  years,  I  do  not  believe  that  the  other 
laboratory  can  succeed.2 

As  I  said,  perhaps  Brooks  lacked  vision.  Presumably  his  convictions  led  him  to  turn 
down  the  directorship  of  the  MBL.  Perhaps  he  was  also  tired  after  years  of  running 
the  Chesapeake  summer  sessions.  Perhaps  he  did  not  wish  to  take  on  a  lab  with  a 
very  weak  financial  base  and  fight  the  inevitable  battles  for  funding,  with  no  obvious 
general  support.  Evidently  he  felt  uneasy  about  having  women  in  his  biology  classes 
and  laboratories,  and  women  would  be  hard  to  avoid  at  the  MBL  because  of  the 
laboratory's  connection  with  the  Women's  Education  Association.  For  various 
reasons,  then.  Brooks  rejected  the  MBL  offer. 

Immediately  after  receiving  Brooks'  rejection,  the  Trustees  forwarded  an  offer 
to  Charles  Otis  Whitman,  then  director  of  the  Allis  Lake  Laboratory  in  Milwaukee, 
Wisconsin  (MBL  Minutes,  1888,  p.  27).  Hyatt  probably  knew  Whitman  through 
Whitman's  two  summers  at  Agassiz's  Penikese  school  and  also  from  the  years  that 
Whitman  spent  at  the  Museum  of  Comparative  Zoology  at  Harvard.  Though 
certainly  not  as  prestigious  as  the  Johns  Hopkins,  Whitman  headed  the  other 
American  biological  research  laboratory  at  the  time.  Immediately,  Whitman  accepted 
the  MBL  offer.  With  only  vaguely  articulated  goals,  the  Trustees  instructed  Whitman 
to  begin  the  lab  within  a  few  months:  to  open  in  July  of  1888.  They  circulated  an 
announcement  to  solicit  students  and  support. 

The  Women's  Education  Association  donated  the  equipment  from  Annisquam 
to  the  MBL  and  also  helped  the  MBL  Trustees  raise  money  for  the  new  laboratory. 
Van  Vleck  served  as  first  instructor,  as  he  had  at  Annisquam,  so  the  MBL 
maintained  connections  with  its  founders.  Yet  Hyatt  led  the  Trustees  in  making  it 
clear  that  change  was  also  in  order,  that  the  lab  should  offer  both  instruction  and 
individual  investigation,  and  that  as  director.  Whitman  should  develop  the  lab  as 
he  saw  appropriate.  As  Frank  Lillie  later  wrote,  this  decision  worked  well,  for  in 
Whitman  "the  trustees  had  found  a  man  not  only  fitted  to  carry  out  their  purposes 
but  possessing  imagination  adequate  to  transform  their  shadowy  ideas,  the  zeal  and 
determination  required  to  give  them  form  and  substance,  and  the  courage  to  face 
whatever  difficulties  might  arise"  (Lillie,  1944,  p.  36). 

The  first  year  began  inauspiciously.  Cornelia  Clapp,  who  had  also  attended  the 
Penikese  School,  arrived  on  time  for  the  new  session  and  found  the  carpenters  still 
at  work  building  the  lab.  Whitman  had  not  yet  arrived,  reportedly  because  of  family 
illness.  No  equipment  had  arrived;  it  remained  side-tracked  somewhere  along  the 
way.  No  one  had  made  arrangements  for  boarding  or  lodging.  In  short,  there  really 
was  no  lab.  But  Clapp,  buoyed  by  her  enthusiasm  and  by  the  arrival  of  the  other 
attendees — about  half  and  half  male  and  female — stayed  and  waited.  Finally,  the 
equipment  from  Annisquam  arrived.  Whitman  appeared,  the  one  laboratory  building 
was  completed,  and  aside  from  such  troubles  as  tripping  at  night  over  the  many 
boulders  in  the  paths,  that  first  session  of  the  MBL  proceeded  successfully,  if  quietly. 

2  Letters,  Brooks  to  Oilman,  no  date.  Oilman  Collection,  and  Brooks  to  Oilman,  December  1980, 
on  the  need  for  a  summer  lab  in  the  southern  United  States  since  he  did  not  regard  the  MBL  as 
satisfactory.  Alexander  Agassiz,  letter.  May  30,  1888,  shows  his  opposition  to  the  new  laboratory. 


BIRTH   OF  THE   MBL  31 

During  those  first  years,  the  Fish  Commission  proved  very  helpful  in  sharing 
specimens,  providing  sea  water,  a  boat,  nets,  etc.  And  the  Fish  Commission  men 
(for  unlike  the  MBL  group,  they  were  all  men)  visited  and  discussed  projects.  Clapp 
recorded  that  Whitman  taught  basic  techniques  and  how  to  observe  productively 
and  to  get  results  in  morphological  research.  As  she  enthused  about  that  first  year, 
the  year  before  the  appearance  of  Wilson,  Conklin,  or  Morgan,  "The  atmosphere 
of  that  laboratory  was  an  inspiration;  the  days  were  peaceful  and  quiet;  there  were 
no  lectures  nor  anything  else  to  distract  the  attention  from  the  work  at  hand" 
(Clapp,  1927).  That  she  fell  in  love  with  the  MBL  experience  is  clear  from  her  life- 
long active  association  there.  This  remarkable  woman  went  on  to  obtain  a  second 
Ph.D.  degree  with  Whitman  when  he  became  chairman  of  the  new  University  of 
Chicago  biology  department  in  1890  (Rossiter,  1982,  pp.  19-21,  86,  88).  That  the 
MBL  succeeded  in  attracting  such  loyal  and  able  supporters  undoubtedly  contributed 
to  its  early  success. 

The  MBL  had  begun.  At  the  same  time  that  the  MBL  attracted  more  researchers 
and  students,  the  Fish  Commission  began  to  have  problems  with  private  researchers. 
In  effect,  the  government  bureaucracy  wanted  the  Commission  to  emphasize  fisheries 
research  and  did  not  wish  to  allow  private  investigators  (Oilman  Papers).  Brooks' 
predictions  about  the  redundancy  and  failure  of  the  MBL  soon  proved  wrong;  the 
MBL  soon  became  the  preeminent  marine  lab  in  Woods  Hole  and  in  the  United 
States.  With  success,  the  Trustees  and  especially  Whitman  began  to  have  greater 
aspirations  for  the  lab. 

So  far,  it  should  be  clear,  this  lab  had  American  roots.  It  was  clearly  a  biological 
laboratory  for  America — the  first  such  permanent  facility.  The  particular  mix  of 
instruction  and  research,  with  the  resulting  communication  among  the  students, 
young  faculty,  and  established  researchers,  was  peculiarly  American,  possible  only 
in  a  country  which  had  no  established  hierarchy  in  research.3  The  inexperienced 
learned  from  direct  contact  with  the  more  proficient.  Thus,  those  who  taught  the 
courses  at  the  MBL  could  introduce  a  new  generation  to  the  problems  and  methods 
they  saw  as  important.  Many  untrained  American  scientists  received  that  practical 
learning  with  nature  which  Agassiz  had  sought — as  they  could  not  have  at  the 
European  stations.  The  democratic  control  by  a  corporation  of  scientists  overseen 
by  interested  trustees  came  only  after  some  reform  and  struggle  in  1897,  but  the 
organization  was  uniquely  American  and  surprisingly  successful.  The  lab  had 
achieved  truly  national  support.  The  links  between  that  character  and  that  of  the 
Naples  Station  remain  to  be  examined. 

I  come  now  to  my  second  theme:  that  interest  in  development  (broadly 
conceived)  served  as  an  important  unifying  focus  for  the  MBL.  Despite  the  expressed 
goal  of  including  all  of  biology,  the  success  of  the  MBL  depended  on  the  way  the 
shared  problems,  namely  developmental  problems,  brought  the  participants  to  work 
together  and  to  communicate  in  an  exciting,  productive,  and  cooperative  way. 

The  fact  that  developmental  questions  dominated  early  work  at  the  MBL  and 
to  a  lesser  extent  subsequent  work  as  well  is  not  entirely  surprising.  Both  Whitman 
and  Brooks  concentrated  on  developmental  questions,  and  these  two  exerted  the 
greatest  influence  on  the  young  researchers  who  worked  at  the  MBL  in  the  first 
decades.  More  generally,  the  morphological  tradition  had  come  to  regard  marine 
invertebrates  as  particularly  useful  for  revealing  homologies  as  well  as  evolutionary 
histories,  or  phylogenies.  By  1890,  many  MBL  researchers  had  focused  on  the 

3  W.  D.  Russell-Hunter  has  pointed  out  that  the  Milport  laboratory  in  Scotland  offered  an  example 
parallel  in  some  respects;  this  suggestion  calls  for  further  careful  study. 


32  J     MAIENSCHEIN 

question  of  how  the  egg  becomes  fertilized  and  begins  development.  Specifically,  a 
number  of  American  researchers  began  to  ask  whether  development  follows  a 
pattern  which  is  predominantly  inherited  or  which  is  acquired  and  hence  emerges 
only  gradually:  that  is,  whether  preformation  or  epigenesis  predominates.  In  particular. 
Whitman  focused  on  the  question:  to  what  extent  does  the  egg  cell  already  experience 
organization?  (Whitman,  1896;  Maienschein,  1985). 

It  is  not  easy  to  answer  that  question.  What  sorts  of  things  might  even  count  as 
evidence  that  either  preformation  or  epigenesis  occurs?  What  sorts  of  work  should 
be  done  to  attack  the  problem — careful  descriptive  observation  of  prepared  materials 
or  experimental  manipulations  to  acquire  new  sources  of  data?  Such  questions  led 
to  intense  debates  by  the  1890's,  which  I  do  not  have  time  to  discuss  here.  But  the 
intensity  of  debate  and  the  concentration  of  research  around  exciting  problems 
clearly  added  to  the  MBL  atmosphere. 

Whitman  believed  that  some  early  organization  occurs,  that  the  egg  is  not 
simply  a  "blank  slate,"  but  he  left  open  the  question  of  how  much  such  organization 
occurs  and  whether  cytoplasm  or  the  nucleus  is  the  center  of  organization.  He  also 
suggested  how  to  attack  such  questions,  namely  through  cell-lineage  studies  (Whitman, 
1878,  1887).  Cell-lineage  does  just  what  it  sounds  like — traces  the  lineages  of  each 
cell  through  every  cleavage  stage  until  the  investigator  gets  tired  of  the  tedius  effort 
or  until  the  cells  become  too  difficult  to  identify  further. 

In  1890,  Edwin  Grant  Conklin  was  at  the  Fish  Commission  examining  early 
developmental  stages.  He  heard  that  Edmund  Beecher  Wilson,  at  the  MBL,  was 
doing  something  similar.  So  Conklin  walked  across  the  street,  talked  to  Wilson,  and 
both  were  astonished  at  how  closely  their  results  agreed.  As  Conklin  reported, 
"Wilson  was  as  excited  by  those  results  as  I  was  and  he  reported  this  to  Whitman. 
Whitman  at  once  sent  for  me  to  come  over  to  see  him  in  the  office.  .  .  ."  (Conklin, 
1968,  p.  116).  Of  course,  Conklin  rushed  right  over,  and  Whitman  said  he  would 
like  to  publish  Conklin's  work  in  his  journal.  The  Journal  of  Morphology.  Others 
joined  in,  including  Thomas  Hunt  Morgan  and  Ross  Harrison,  though  they  never 
actually  published  their  cell  lineage  work  (Costello,  1967).  As  Whitman's  student 
and  second  director  of  the  MBL,  Lillie,  said,  when  Whitman  told  him  of  the  people 
working  on  cell-lineage  and  of  their  findings,  "I  accepted  his  advice  to  take  up  this 
subject:  and  worked  on  freshwater  Unio,"  for  which  he  had  to  take  the  train  back 
and  forth  to  a  little  pond  in  Falmouth,  Massachusetts,  carrying  his  heavy  wading 
boots  and  a  heavy  bucket  (Lillie,  1926).  Cell-lineage  work  served  as  a  rallying  point 
and  attracted  researchers  to  the  MBL  for  a  specific  purpose  (Maienschein,  1978).4 

By  the  mid  1890's,  cell-lineage  work  had  begun  to  pale.  Several  researchers  at 
the  MBL  turned  to  other  morphological  questions  and  to  problems  of  regeneration, 
and  to  physiology  and  related  problems  (Werdinger,  1980;  Maienschein,  1976; 
Haraway,  unpub.).  Also,  the  experimental  work  of  Jacques  Loeb  and  Charles 
Manning  Child  in  physiology  of  development  and  by  the  German  developmental 
experimentalists  began  to  attract  more  attention  as  a  possibly  productive  method 
for  attacking  those  same  questions  about  epigenesis  and  preformation,  or  whether 
the  egg  is  organized  as  a  mosaic  or  develops  regulatively.  Stimulated  by  successes 
from  Germany  and  Naples,  biologists  became  increasingly  enthusiastic  about  the 
promises  of  experimental  manipulation,  which  seemed  to  many  to  offer  quicker 
and  more  dramatic  results  than  more  traditional  methods  such  as  cell-lineage  work. 

4  Publications  in  journals  edited  by  Whitman,  Biological  Lectures  and  Journal  oj  Morphology,  report 
the  results  of  the  cell-lineage  work. 


BIRTH   OF  THE   MBL  33 

As  Herbert  Spencer  Jennings  later  reflected,  this  led  to  a  mad  rush  toward 
experimentation  by  some.  He  said  of  the  period: 

.  .  .  their  tales  disagreed  radically.  They  tried  for  a  long  time  to  convince  each 
other,  but  failed.  And  the  reason  was  that  there  was  no  way  of  deciding  which, 
if  any,  of  the  tales  were  correct.  But  what  hath  the  man  of  science  of  all  his  labor 
and  of  the  vexation  of  his  heart,  if  it  leads  to  no  general  agreement,  to  nothing 
that  can  be  demonstrated?  And  so,  the  zoologists  gave  it  up;  they  looked  upon 
the  works  that  their  hands  had  wrought,  and  behold  all  was  vanity  and  vexation 
of  spirit.  Henceforth,  they  said,  we  must  so  work  that  our  results  and  conclusions 
can  be  tested;  can  be  verified  or  refuted.  We  must  be  able  to  say:  Such  and  such 
things  happen  under  such  and  such  conditions,  and  if  you  don't  believe  it  you 
may  supply  the  conditions,  you  may  try  it  for  yourself,  and  you  will  find  it  to  be 
true.  But  that  is  precisely  experimentation;  and  so  they  flocked  with  enthusiasm 
to  experimentation.  (Jennings,  1926,  p.  98) 

This  led  to  a  good  deal  of  argument,  with  experimental  evidence  cited  as  proving 
one  or  another  point  of  view.  As  Jennings  later  reflected,  that  period  seemed  a  bit 
like  a  comic  opera  with  everyone  dancing  about  singing  frenetically  "You  are  right 
and  I  am  right  and  he  is  right  and  all  are  right"'  (Jennings,  1926,  p.  99).  Not 
everyone  had  embraced  experimentation,  of  course.  And  the  turn  to  experimentation 
did  not,  in  fact,  solve  all  the  problems  as  some  had  hoped. 

The  rush  to  experimentation  settled  down,  but  the  concern  with  shared  problems 
remained  into  the  twentieth  century.  The  cross-fertilization  of  ideas  and  exchange 
of  methods  really  did  dominate  developmental  work  at  the  MBL,  as  revealed  in  the 
Biological  Lectures  published  from  the  Laboratory.  By  1910,  things  had  begun  to 
change,  as  they  have  continued  to  do  since.  Research  has  steadily  diverged  in 
different  directions  with  resulting  proliferation  of  more  specialized  research  projects, 
courses,  and  publications — which  is  another  story.  The  early  sense  of  shared 
developmental  concerns,  which  provided  such  a  strong  foundation  for  the  first 
permanent  American  marine  laboratory  has  faded,  for  better  or  for  worse.  As 
Conklin  suggested,  such  changes  in  biology  have  not  always  advanced  biological 
understanding.  Biologists,  he  said,  have  become  a  lot  like  squid.  They  have  come 
to  progress  rapidly  backwards  while  excreting  large  quantities  of  ink  (Conklin,  n.d.). 
Squid,  like  ink  and  progress,  have  played  an  important  role  in  the  MBL's  history. 

ACKNOWLEDGMENTS 

I  wish  to  thank  the  MBL  librarians  and  especially  Ruth  Davis  who  offered 
special  help  and  encouragement  at  many  points,  Ann  Blum  at  the  Museum  of 
Comparative  Zoology,  Philip  Pauly  for  identifying  resources,  and  the  archivists  at 
the  Johns  Hopkins  University  Archives  for  their  assistance.  Archival  materials 
quoted  with  permission.  Research  was  supported  by  NSF  grant  #SES-8309388. 

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THE  "NEW"   EMBRYOLOGY   AT  THE  ZOOLOGICAL  STATION 
AND  AT  THE   MARINE   BIOLOGICAL   LABORATORY 

ALBERTO   MONROY   AND  CHRISTIANE  GROEBEN 

Stazione  Zoologica,  80121  Napoli,  Italy 

The  time  will  never  come  when  direct  interchange  of  thought  and  comparison 
of  methods  of  research  will  cease  to  be  of  the  highest  importance  to  the 
biologist. 


:.  O.  Whitman.  1883.  The  advantages  of  study 
at  the  Naples  Zoological  Station.  Science  2: 
93-97 

ABSTRACT 

The  Naples  Zoological  Station  was  one  of  the  main  centers  of  the  revolt  against 
Haeckelian,  phylogenetic  embryology.  On  the  other  hand,  the  founder  of  the  Station, 
Anton  Dohrn,  while  being  a  distinguished  embryologist,  was  an  enthusiastic  follower 
of  HaeckeFs  theories.  The  question  discussed  here  first  is  that  of  the  interactions 
between  Dohrn  and  the  followers  of  the  new  trend  in  embryology,  the  Entwicklungs- 
mechaniker,  among  whom  Herbst,  Driesch,  and  Boveri  were  regular  visitors  to  the 
Station.  While  Dohrn  fully  acknowledged  the  significance  of  the  discoveries  arising 
from  the  new  experimental  approach  to  embryology,  he  remained  faithful  to 
phylogenetic  embryology.  Examining  the  interactions  between  two  American  biol- 
ogists most  involved  in  the  foundation  of  the  MBL,  namely  C.  O.  Whitman  and 
E.  B.  Wilson,  and  the  leaders  of  the  "new"'  embryology,  we  then  discuss  the  effect 
of  these  interactions  on  the  development  of  embryological  research  at  the  MBL. 
We  suggest  that  the  main  effect  was  to  promote  the  new  conceptual,  and  hence 
methodological,  approach  to  the  problems  of  development.  The  Naples  group  saw 
the  egg  as  a  cell  that  could  be  manipulated  in  an  effort  to  answer  questions 
concerning  cell  physiology.  In  contrast,  the  Woods  Hole  group  was  interested  in  the 
egg  as  the  starting  point  of  development.  This  was  reflected  also  in  the  choice  of 
the  experimental  material:  the  sea  urchin  egg  in  the  former  case  and  the  highly 
"determined"  eggs  of  mollusks  and  annelids  in  the  latter. 

DISCUSSION 

The  Naples  Zoological  Station  was  one  of  the  strongholds  of  the  revolt  against 
Haeckelian,  phylogenetic  embryology.  In  fact,  it  was  in  Naples  that  the  advocates 
of  the  new  experimental  approach  to  the  problems  of  development — Entwicklnngs- 
mechanik  and  Entwicklungsphysiologie — made  some  of  their  most  important  dis- 
coveries, thus  starting  a  new  era  in  the  study  of  development.  We  wish  to  discuss 
here  the  interactions  between  the  followers  of  the  new  approach  and  the  founder  of 
the  Zoological  Station,  Anton  Dohrn. 

Dohrn  was  an  embryologist  who  moved  in  the  footsteps  of  Ernst  Haeckel  and 
who  had  immense  admiration  of  HaeckeFs  scientific  achievements.  Dohrn  wrote, 
in  1867,  that  "In  Haeckel's  Gesamte  Morphologic  der  Organismen  lies  the  foundations 
of  a  new  science"  and,  later,  that  "it  is  to  be  considered  an  established  fact  that  the 
development  of  an  animal  in  the  egg  and  in  the  larval  condition  is  a  condensed 


35 


36  A.   MONROY   AND  C.  GROEBEN 

and  sometimes  obscured  image  of  the  development  of  its  genealogic  tree."  One  of 
Dohrn's  immediate  goals  was  to  analyze  in  detail  one  of  Haeckel's  constructions. 
"In  the  same  way  as  linguists  reconstruct  original  languages  .  .  .  the  zoologist 
should  be  able  to  outline  a  comprehensive  picture  of  the  development  of  an  animal 
group  from  a  large  number  of  embryological  data"  and  hence  "to  identify  the 
ancestor  of  the  whole  group"  (Dohrn,  1872).  He  thought  his  "Funktionswechsel," 
which  he  considered  as  probably  his  most  important  intellectual  achievement,  was 
the  key  to  the  changes  underlying  evolution.  The  essence  of  this  principle  is 
epitomized  in  two  passages:  "The  way  of  life  is  the  agent  that  keeps  the  shape  of 
the  larva  until  its  development  has  reached  the  right  size  and  the  tissues  from  which 
the  insect  (Fliige)  will  arise  are  already  there,"  and 

The  succession  of  functions  which  are  carried  out  by  the  same  organ  caused  the 
change  of  the  organ.  Each  function  is  the  resultant  of  many  components,  one  of 
which  is  the  main  or  primary  function,  while  the  others  are  side  or  secondary 
functions.  The  lowering  of  the  primary  function  and  the  building  up  of  a 
secondary  function  alters  the  overall  function — when  the  secondary  function 
becomes  the  primary  one,  the  overall  function  changes  and  the  result  of  the 
whole  process  is  that  the  organ  changes  [Dohrn,  1875]. 

As  examples  he  refers  to  the  anterior  limbs  of  crustaceans  turning  into  chelae, 
to  vertebrate  gills  turning  into  mouth  apparatus. 

Dohrn's  theory  was  the  target  of  a  vehement  attack  by  Carl  Gegenbaur  (1876), 
who  dismissed  it,  in  particular  rejecting  the  speculations  about  the  origin  of 
vertebrates  from  annelids  as  "a  striking  example  of  unscientific  comparative 
anatomy."  Hence,  it  must  have  been  gratifying  for  Dohrn  to  receive  a  sympathetic 
letter  from  Charles  Darwin  (24  May  1875)  in  which  Darwin  cautiously  expressed 
his  interest  in  Dohrn's  ideas  about  the  descent  of  vertebrates.  Another  letter  dated 
2  February  1875,  comments  humorously  on  the  theory  of  the  vertebrate's  descent 
from  annelids:  "I  shall  be  very  sorry  to  give  up  the  ascidians  to  whom  I  feel 
profound  gratitude."  Dohrn's  "Funktionswechsel"  principle  was  also  endorsed  by 
August  Weismann  and  Emil  du  Bois-Raymond. 

Thus,  for  Dohrn  as  for  Haeckel  embryology  was  just  a  tool,  albeit  a  most 
powerful  one,  to  construct  genealogic  trees  of  the  various  animal  groups  and 
eventually  to  identify  the  original  form  from  which  all  groups  branched.  The  embryo 
as  such  was  uninteresting.  Thus,  embryology  was  reduced  to  an  almost  sterile 
mental  exercise.  Saying  this  does  not  imply  that  all  embryological  work  carried  out 
under  Haeckel's  influence  was  worthless.  On  the  contrary,  some  important  discoveries 
even  paved  the  way  for  future  work.  Yet  most  observations,  no  matter  how 
important  in  their  own  right,  were  contorted  in  order  to  fit  hypothetical  and 
unprovable  genealogic  trees. 

Wilhelm  Roux  violently  opposed  this  approach.  The  introduction  to  the  first 
volume  of  his  Archiv  fiir  Entwicklungsmechanik  der  Organismen  (1894)  may  be 
considered  the  "manifesto"  of  the  new  embryology.  His  views  opposed  those  of 
HaeckeKs,  both  theoretically  and  methodologically,  and  led  to  an  entirely  different 
approach  to  the  analysis  of  development.  The  main  claim  was  that  the  study  of  the 
embryo  was  interesting  in  its  own  right.  A  few  excerpts  illustrate  Roux's  point.  For 
example,  the  program  is  outlined  in  the  opening  sentence: 

Developmental  mechanics  or  causal  morphology  of  organisms  ...  is  the  doctrine 
of  the  causes  of  organic  forms,  and  hence  the  doctrine  of  the  causes  of  the  origin, 
maintenance  and  involution  of  these  forms  .  .  .  the  general  problem  of  devel- 
opmental mechanics  [is]  the  ascertainment  of  the  formative  forces  of  energies. 


THE  "NEW"   EMBRYOLOGY  37 

In  so  far,  however,  as  forces  or  energies  are  only  known  to  us  by  their  effects, 
i.e.,  every  kind  of  force  by  its  specific  mode  oj  operating,  the  problem  may  be 
defined  as  the  ascertainment  of  the  formative  modi  operandi. 

From  these  premises  follows  the  conclusion  that  "The  causal  method  of 
investigation  ...  is  experiment.  'Certainty  in  causal  deduction  can  only  come  from 
experiment,  either  from  ^artificial''  or  from  "nature's  experiment,  such  as  variation, 
monstrosity,  or  other  pathological  phenomena,"  and  particularly  important,  "devel- 
opmental mechanics  must,  so  far  as  possible,  seek  to  utilize  for  its  own  ends,  all  the 
ends  and  ways  of  causal  investigation  of  organisms  and  the  results  thereby  attained, 
and  not,  in  foolish  conceit,  cast  aside  any  biological  discipline  as  being  useless."  On 
the  other  hand,  Roux  did  not  disdain  phylogenetic  studies.  Indeed 

...  in  accordance  with  the  double  course  of  development,  viz.  the  phyletic  and 
ontogenetic,  developmental  mechanics  must  look  for  the  causes,  or  modi  operandi, 
of  each  of  these  two  courses;  hence  an  ontogenetic  and  phylogenetic  developmental 
mechanics  are  to  be  perfected.  But  in  consequence  of  the  intimate  causal 
connections  existing  between  the  two.  many  of  the  conclusions  drawn  from  the 
investigation  of  ontogeny  will  also  throw  light  on  phylogenetic  processes  .  .  . 

As  long  as  comparative  anatomy  attempted  to  establish  only  the  main  course 
of  development  in  the  animal  kingdom,  following  in  a  general  way  the  continuous 
development  of  forms  only  through  the  classes  of  each  type,  comparison  of 
different  forms  showed  that  essentially  and  unequivocally  the  same  course  of 
progressive  development  is  followed  by  nearly  all  systems  of  organs.  But  in 
further  approximation  of  a  higher  degree,  viz.,  in  tracing  that  development 
through  the  orders,  families,  genera,  and  species,  even  to  the  individual,  so  many 
incongruities  in  the  development  of  organ  systems  and  organs  made  their 
appearance,  that  comparative  anatomy  has  been  compelled  to  call  in  the 
assistance  of  quite  a  number  of  developmental  mechanical  hypotheses,  for  the 
correctness  of  which  only  experimental  tests  can  give  complete  security. 

And  a  few  lines  later  .  .  .  "it  would  be  encouraging  if  comparative  anatomists 
would  themselves  resort  to  experimentation  for  the  purpose  of  solving  .  .  .  the 
problems  in  which  they  are  interested"  (Roux,  1894).  These  passages,  which  are 
generally  overlooked  or  ignored,  show  that  Roux's  position  was  much  more  open- 
minded  than  that  of  some  of  his  followers.  He  seems  to  have  accepted  phylogenesis 
in  the  realm  of  Entwicklungsmechanik;  and  in  fact  the  last  sentence  is  a  plea  for 
cooperation  with  the  comparative  anatomists.  As  we  shall  see,  quite  a  different 
attitude  from  that  of  Hans  Driesch! 

The  most  strenuous  and,  indeed,  uncompromising  and  arrogant  defender  of  the 
new  movement  was  Hans  Driesch.  It  is  worth  citing  some  passages  from  a  virulent 
and  somewhat  amusing  article  written  by  Driesch  mostly  directed  against  Hugo 
Eisig  and  Edmund  Beecher  Wilson. 

What  do  the  phylogeneticists  want  then?  They  cannot  even  do  consistent 
biological  research!  Yes!  can  they  do  research  at  all?  .  .  .  and  yet  [phylogenesis] 
is  there  but  not  as  a  science  as  it  is  not  even  entitled  to  this  name  .  .  .  We  know 
what  we  can  do  and  what  we  cannot  do  for  the  time  being.  Our  opponents  think 
that  they  know  what  we  don't  even  want  to  know.  With  their  comparisons  they 
deal  with  questions  that  by  their  very  nature  we  have  not  dealt  with  and  which 
in  fact  cannot  be  approached  ...  we  have  started  to  approach  scientifically  a 
very  small  part  of  morphological  problems,  others,  such  as  the  very  important 
problem  of  morphological  diversities,  not  at  all.  We  are  well  aware  of  the  problem 
of  Transformation'  but  we  consider  it  for  the  time  being  an  impregnable  fortress 
and  we  address  ourselves  to  'Developmental  Physiology'  as  here  we  see  the 
possibility  of  obtaining  results  while  our  opponents  represent  us  as  if  we  thought 
that  Developmental  Physiology  were  all  Morphology  [Driesch,  1899]. 


A.   MONROY   AND  C.   GROEBEN 

And  in  a  letter  to  Eisig  (28  August  1898)  he  wrote  as  an  explanation  of  his  attack, 
".  .  .  it  is  you  that  I  have  as  opponent,  and  not  only  you,  but  also  Wilson  who 
with  his  last  publication  has  become  a  renegade  [referring  to  the  1898  Wilson 
paper].  This  is  why  I  have  to  deal  in  the  article  with  persons  whom  I  know  closely 
and  esteem  personally  and  scientifically:  this  is  why  I  had  to  write  it." 

How  did  Anton  Dohrn  feel  about  Driesch  and  the  new  direction  of  embryological 
research  which,  ironically,  had  become  so  centered  at  the  Zoological  Station  that 
the  leaders  (including  such  people  as  Driesch,  Herbst,  Boveri,  Morgan,  and  Wilson) 
were  known  in  Europe  as  "Neapler  Entwicklungsmechanikef!"  There  are  very  few 
testimonies  of  Dohrn's  opinion  of  Driesch  as  a  person  and  as  a  scientist.  In  a  letter 
to  A.  H.  Davis  (10  July  1903  quoted  by  Heuss,  1962)  he  expresses  his  sympathy 
and  understanding  for  Driesch:  "although  I  cannot  follow  the  weight  of  his 
argument,  or  his  point  of  view  and  of  his  conclusions."  And  a  year  later  he  wrote 
to  Wilson: 

If  one  day  he  could  be  persuaded  that  he  ought  not  to  boast,  even  in  a  mild  way, 
of  the  excellence  of  his  intellectual  field  but  to  be  courteous  and  magnanimous 
with  other  pursuits,  he  would  grow  very  much  in  value.  I  hope  he  will  one  day 
be  wise  enough  to  feel  that;  if  not  his  Personlichkeit  will  never  attain  the 
dimensions  of  his  intellect.  I  have  an  intense  interest  in  Driesch,  though 
sometimes  I  wish  him  a  good  licking  [27  July  1904,  A.  Dohrn  to  E.  B.  Wilson]. 

Driesch,  on  the  other  hand,  greatly  admired  the  contribution  to  biology  of  the 
Zoological  Station  and  hence  of  Dohrn.  Indeed,  in  his  memorial  address  for  Dohrn 
he  wrote  that  the  Zoological  Station  was  "the  place  where  most  of  the  cytology  and 
experimental  developmental  physiology  had  originated,  however  in  their  own  right 
and  not  as  sciences  at  the  service  of  the  theory  of  descent,  and  in  fact  in  open 
contrast  with  Phylogenesis"  (Driesch,  1909). 

Driesch  was  certainly  one  of  the  most  interesting  and  influential  personalities  at 
the  Naples  Station  beginning  with  his  first  visit  in  1891  (Driesch,  1951).  For  a 
number  of  scientists  who  worked  at  the  Station  his  influence  was  a  decisive  factor 
in  their  scientific  life.  Also,  it  was  largely  due  to  Driesch  that  researchers  recognized 
the  great  advantage  of  the  sea  urchin  egg  as  an  experimental  tool.  Until  recently,  in 
fact,  90%  of  the  work  on  fertilization  and  a  large  percentage  of  all  embryological 
work  rested  on  experiments  on  the  sea  urchin  egg.  Indeed,  it  had  become  a 
widespread  belief  that  "what  is  true  for  the  sea  urchin  must  be  true  for  all  animals." 
The  discovery  of  the  sea  urchin  egg  as  an  experimental  material  can  be  traced  back 
to  the  work  at  Villefranche  by  the  Hertwigs,  who  followed  in  vivo  the  details  of  the 
formation  of  the  zygote  nucleus  from  the  fusion  of  the  male  and  the  female 
pronuclei  (Hertwig,  1876).  It  was,  however,  due  to  Curt  Herbst  and  Driesch  that 
embryologists  realized  how  well  the  sea  urchin  egg  lent  itself  to  experimental 
manipulations.  One  of  the  earliest,  and  in  fact  one  of  the  most  important  break- 
throughs, was  Herbsf  s  discovery  that  blastomeres  of  the  cleaving  sea  urchin  egg 
would  be  separated  from  one  another  after  a  brief  exposure  to  calcium-free  sea 
water  (Herbst,  1908).  These  experiments  paved  the  way  to  one  of  the  most 
fascinating  areas  of  research  in  embryology,  that  of  cell  interactions.  Almost  at  the 
same  time  Driesch  succeeded  in  separating  the  first  two  blastomeres  and  showing 
that  a  whole  embryo  could  arise  from  each  one  of  them.  This  observation,  which 
at  first  appeared  to  contradict  Roux's  results  on  the  amphibian  egg,  began  the  long 
controversy  on  the  mosaic  versus  regulatory  organization  of  the  egg.  The  aggressive 
method  used  by  Driesch  to  separate  the  blastomeres  by  violently  shaking  the  egg 
(the  embryologists  who  used  Driesch's  method  were  derisively  called  "egg-shakers") 
were  later  superseded  by  the  highly  sophisticated  microsurgical  technique  devised 


THE   "NEW"   EMBRYOLOGY  39 

by  Sven  Horstadius  in  the  early  thirties  (Horstadius,  1973).  To  this  day  Horstadius's 
work  is  one  of  the  points  of  reference  for  sea  urchin  embryology.  In  Theodor 
Boveri's  hands  the  sea  urchin  egg  proved  also  to  be  an  excellent  material  for  the 
study  of  the  nuclear-cytoplasmic  interactions  in  development  through  the  analysis 
of  the  hybrid  combinations,  a  method  further  developed  by  his  student  Fritz  Baltzer 
and  still  widely  used  (Baltzer,  1967).  (The  unfinished  manuscript  of  Boveri's  last 
work  on  the  development  of  merogonic  and  partially  merogonic  sea  urchin  hybrids 
is  preserved  in  the  MBL  Library.)  The  sea  urchin  egg  was  equally  important  for  the 
study  of  physiological  and  biochemical  problems,  not  only  those  specifically  related 
to  development  but  also  those  of  a  more  general  character.  In  this  context.  Otto 
Warburg's  work  has  a  special  prominence.  His  discoveries  of  the  change  in  the 
respiration  of  the  sea  urchin  egg  as  a  result  of  fertilization  started  a  new  field  of 
"chemical  embryology/'  a  research  line  pursued  at  the  Zoological  Station  primarily 
by  John  Runnstrom  and  his  school  (Warburg,  1910).  Runnstrom  was  indeed  a 
frequent  visitor  to  the  Station  almost  to  his  death. 

In  Woods  Hole,  however,  experiments  on  the  sea  urchin  egg  did  not  start  on  a 
large  scale  until  Jaques  Loeb's  discovery  of  artificial  parthenogenesis  (1899),  and 
until  Frank  Lillie  became  interested  in  fertilization  early  in  1900  [even  though 
Wilson  had  shown  how  well-suited  the  sea  urchin  egg  was  for  the  study  of 
fertilization  and  cell  division  (Wilson,  1895)].  This  reflects  the  different  kinds  of 
problems  which  the  embryologists  in  Woods  Hole  were  considering.  In  Naples, 
largely  under  the  influence  of  Driesch  and  Herbst  and  later  Boveri,  interest  focused 
on  what  Driesch  called  "developmental  physiology,"  or  the  study  of  mechanisms 
controlling  the  early  stages  of  development.  Woods  Hole  embryologists,  mostly 
under  the  influence  of  Charles  Otis  Whitman,  were  interested  in  the  "program  of 
development"  and  hence  primarily  in  cell  lineage.  They  were  interested  in  the  egg 
and  the  oocyte  as  the  point  of  departure  for  development.  The  problem  of 
"promorphology,"  which  was  completely  alien  to  the  Naples  group,  was  central  to 
the  Woods  Hole  embryologists. 

The  few  documents  available  suggest  that  Dohrn  was  aware  of  the  revolution  that 
was  taking  place  in  the  field  of  embryology.  He  recognized  the  strength  of  the  tide, 
but  still  felt  lingering  fondness  for  his  "fallen  idol."  He  had  to  "play  the  host,"  and 
as  Director  of  the  Station  it  was  his  duty  to  stay  an  dehors  de  la  melee:  to  take 
sides  might  have  endangered  the  position  of  the  Station.  In  this  connection,  Dohrn 
complained  of  Froriep's  use  (Froriep,  1902)  of  meaningless  and  empty  expressions: 

1  see  in  those  expressions  only  empty  schemes  of  the  kind  that  have  greatly 
damaged  the  already  discredited  phylogenetic  research,  and  which  one  should 
strenuously  avoid,  if  one  want  to  repel  within  due  limits  the  highbrowed  criticism 
that  phylogenesis  receives  from  the  completely  differently  oriented  Entwicklungs- 
mechanik  or  Entwicklungsphysiologie,  and  if  one  want  to  strengthen  its  indepen- 
dent and  fundamental  importance  as  a  historic-biological  discipline  [Dohrn. 
1904]. 

Thus  Boveri  may  not  have  been  fair  when  he  wrote: 

Not  that  he  did  not  recognize  the  value  of  new  discoveries  .  .  .  Isn't  it  surprising 
that  the  rich  mine  that  he  himself  had  opened,  has  had  essentially  no  influence 
on  his  own  work?  The  direction  of  his  research  was  not  determined  by  any  kind 
of  external  stimuli,  but  the  specific  problems  that  originated  in  his  mind 
developed  into  theories  that  he  then  tried  to  verify  with  facts  [Boveri,  1910]. 

Let  us  turn  to  the  question  of  how  and  to  what  extent  the  atmosphere  at  the 
Zoological  Station  and  Dohrn's  personality  influenced  visitors.  We  shall  limit  our 
analysis  to  only  two  of  the  first  American  biologists  to  work  at  the  Station:  Whitman 


40  A.   MONROY   AND  C.   GROEBEN 

and  Wilson.  Given  the  role  played  by  these  two  men  in  establishing  the  new  trend 
of  embryological  research  in  the  United  States,  rather  than  their  own  research 
contributions,  we  feel  that  Whitman  should  be  considered  the  key  figure  and  in  fact 
"the  inspirational  leader"  (Maienschein,  1978).  Whitman  visited  Naples  for  the  first 
time  from  November  1881  to  May  1882;  ten  years  earlier  than  Driesch.  When  he 
went  to  Naples  he  had  already  had  graduate  training  with  Rudolf  Leuckart  in 
Leipzig,  and  hence  he  had  already  been  exposed  not  only  to  the  physiological 
methods  but  also  to  the  anti-Haeckel  wave  already  in  full  swing  in  Germany,  whose 
most  prominent  figure  was  Wilhelm  His.  At  that  time  Whitman  had  already 
published  his  important  paper  on  "The  embryology  of  Clepsine'"  (Whitman, 
1878,  1888). 

These  dates  are  important  as  they  show  that  when  Whitman  went  to  Germany 
he  already  had  certain  well-defined  ideas  and,  more  importantly,  he  had  already 
developed  working  hypotheses  as  to  the  problems  of  development  and  the  method- 
ological approach  to  them.  Whitman  rejected  the  idea  that  gastrulation  was  the  first 
important  event  of  the  development — an  idea  that  carried  much  weight  in  phylo- 
genetic  embryology  and  that  had  been  the  cornerstone  of  the  "Gastrea"  theory. 
Whitman  was  among  the  first  to  herald  the  idea  that  the  study  of  development  had 
to  start  from  the  divided  egg.  In  particular,  he  maintained  that  the  origin  of  the 
germ  layers  (a  topic  of  fundamental  importance  to  the  Haeckelians)  could  not  be 
properly  understood  without  knowledge  of  the  principal  events  of  cleavage,  in 
particular  of  the  eventual  significance  of  the  various  blastomeres.  This  means  that 
the  main  object  of  embryology  was  the  study  of  the  history  of  the  embryo,  from 
the  egg  to  the  adult,  rather  than  from  the  embryo  to  the  ancestors  (Whitman  1894b, 
1895).  This  was  well  in  line  with  the  approach  acclaimed  by  the  new  German 
embryological  school.  What  seems  new  is  that  Whitman  stressed  the  importance  of 
the  organization  of  the  egg — the  promorphology — as  the  basis  to  understanding 
development. 

Whitman's  views  on  promorphology,  organization  and  epigenesis  are  best 
explained  in  his  lecture  "Evolution  and  epigenesis"  (1894a)  where  he  wrote: 

It  has  become  perfectly  clear  .  .  .  that  epigenesis,  as  now  understood,  does 
not  cover  the  whole  field.  Only  the  old  epigenesis  .  .  .  ever  pretended  to  start 
the  development  of  organisms  from  the  level  of  inorganic  matter.  .  .  . 

The  indubitable  fact  on  which  we  now  build  is  not  bit  of  inorganic  homogeneity, 
into  which  organization  is  to  be  sprung  by  a  coagulating  principle,  or  cooked  in 
by  a  calidum  innatum.  or  wrought  out  by  a  spinning  archaeus,  but  the  ready- 
formed,  living  germ,  with  an  organization  cut  directly  from  a  preexisting,  parental 
organization  of  the  same  kind.  .  .  . 

The  essential  thing  here  is,  not  simply  continuity  of  germ  substance  of  the 
same  chemico-physical  constitution,  but  actual  identity  of  germ  organization  with 
stirp-organization  .  .  . 

Let  this  'organization'  stand  for  no  more  than  our  neoepigeneticists  freely 
concede,  namely,  that  original  constitution  of  the  germ,  which  predetermines  its 
type  of  development  and  the  form  which  ultimately  distinguishes  it  from  other 
species  developing  under  like  external  conditions. 


and 


The  question  does  not  now  turn  on  either  of  the  old  hinges,  but  on  what 
factors  determine  the  type  of  development.  Instead  of  asking,  are  all  the  parts 
predelineated?  we  ask,  how  are  they  delineated?  Instead  of  referring  development 
to  a  deus  ex  machind,  or  accident,  we  ask,  what  is  the  mechanism  of  the  germ 
which  enables  it  under  suitable  conditions  to  grow,  divide,  differentiate,  and 
reproduce  all  the  complicated  details  of  its  own  species?  We  see  that  every  form 


THE   "NEW"   EMBRYOLOGY  41 

presented  in  development  issues  as  the  product  of  what  has  gone  before  and  as 
the  foundation  of  what  is  yet  to  come.  Retrospectively,  it  is  a  'determinate,' 
prospectively,  it  is  a  'determinant.' 

Indeed,  Whitman  challenged  the  anti-historical  approach  of  the  Entwicklungs- 
machaniker,  though  not  that  of  Roux.  Organisms  are  the  product  of  a  long  evolution 
and  hence  they  cannot  be  properly  analysed,  let  alone  understood,  without  keeping 
this  fact  well  in  mind.  Whitman  consequently  rejected  the  view  of  embryology  as 
only  "physiology  of  development"  which  was  based  on  the  assumption  that 
evolutionary  history  does  not  play  any  part  in  the  explanation  of  ontogeny.  Hence, 
he  made  a  plea  for  cooperation  between  morphology  and  physiology. 

Without  any  direct  sources  of  information  it  is  impossible  to  say  whether  Dohrn 
and  Whitman  exerted  any  scientific  influence  on  one  another.  Certainly  Whitman's 
1883  article  on  the  Zoological  Station  shows  that  he  was  strongly  impressed  by  its 
organization  and  by  the  atmosphere  that  Dohrn  had  managed  to  create  there  yet, 
as  we  shall  see  shortly,  he  did  not  share  some  of  Dohrn's  principles.  Nor  is  it 
possible  to  make  any  meaningful  inferences  from  Whitman's  papers  after  his  stay 
in  Naples.  It  does  seem  unlikely  that  the  topic  of  evolution  was  never  mentioned 
during  their  conversations.  The  most  that  can  be  said  is  that  daily  contact  with 
Anton  Dohrn  may  have  been  influential  in  mitigating  the  later  anti-phylogenetic 
attitude  of  the  most  fervent  German  Entwicklungsmechanikers.  This  is  why  it  is 
important  that  Whitman  was  in  Naples  before  the  Driesch  era  since  he  developed 
his  own  views  without  being  influenced  by  Driesch's  overwhelming  personality. 

This  experience  at  Naples  was  of  great  importance  when  Whitman  began  the 
Marine  Biological  Laboratory.  Indeed,  he  brought  not  only  the  spirit  of  freedom  he 
had  experienced  at  the  Zoological  Station,  but  also  an  open-minded  attitude  to 
approaches  to  the  problems  of  development.  Nothing  gives  better  evidence  of  the 
atmosphere  that  has  prevailed  at  the  Marine  Biological  Laboratory  since  its 
foundation  than  the  Biological  Lectures.  Contrary  to  Anton  Dohrn  who,  under  the 
banner  of  freedom,  banned  teaching,  lectures,  and  debates  from  the  Station, 
Whitman  stressed  these  aspects.  And  this  is  why  now,  nearly  a  century  later,  we 
have  an  invaluable  testimony  of  not  only  how  biological  research  and  thinking 
evolved,  but  of  the  disagreements  and  personality  clashes  that  occurred  along  the 
way.  For  this  reason  developmental  biologists  owe  Whitman  a  debt  equal  to  that 
which  is  owed  Anton  Dohrn. 

Wilson's  story  is  quite  different.  A  student  of  William  Keith  Brooks,  one  of  the 
staunchest  Haeckelians,  Wilson  soon  became  dissatisfied  with  the  phylogenetic 
approach  to  embryology.  It  is  significant  that  his  work  ripened  during  his  first  stay 
in  Woods  Hole.  There  he  met  Whitman  who  took  great  interest  in  this  research  on 
the  development  of  Nereis.  It  is,  however,  certain  that  Theodor  Boveri  exerted  the 
greatest  influence  on  Wilson,  followed  by  Hans  Driesch  whom  he  met  in  Naples  in 
1892.  It  was  Boveri  who  remained  his  point  of  reference  throughout  his  life,  and  it 
is  significant  that  Wilson's  celebrated  and  classic  book  The  Cell  in  Development  and 
Inheritance  (1896)  is  dedicated  "To  my  friend  Theodor  Boveri."  It  is  no  wonder, 
then,  that  very  early  in  his  scientific  career  Wilson  was  decidedly  oriented  toward 
the  cellular  approach  to  the  study  of  development,  with  major  emphasis  on  the 
nucleus.  In  The  Cell  he  writes: 

The  primary  determining  cause  of  development  is  the  nucleus,  which  operates 
by  setting  up  a  continuous  series  of  specific  metabolic  changes  in  the  cytoplasm. 
This  process  begins  during  ovarian  growth,  establishing  the  external  form  of  the 
egg,  its  primary  polarity,  and  the  distribution  of  substances  within  it.  The 
cytoplasmic  differentiations  thus  set  up  form,  as  it  were,  a  framework  within 


42  A.   MONROY   AND  C.   GROEBEN 

which  the  subsequent  operations  take  place  in  a  course  which  is  more  or  less 
firmly  fixed  in  different  cases. 

Thus  Wilson,  as  did  Whitman,  looked  at  the  events  of  oogenesis  as  the  most 
important  events  for  the  subsequent  development  of  the  embryo.  This  does  not, 
however,  make  him  a  "preformationist"  in  the  usual  meaning  of  the  word. 

A  detailed  account  and  discussion  of  Wilson's  embryological  work,  especially 
with  respect  to  the  problem  of  cell  lineage,  is  provided  by  Maienschein  (1978). 
Whitman's  influence  is  felt  in  Wilson's  position  toward  the  evolutionary  interpretation 
of  development.  Wilson's  views  are  best  expressed  in  the  two  lectures  on  the 
"Embryological  criterion  of  homology"  (1894)  and  "Cell  lineage  and  ancestral 
reminiscence"  (1898a,  b).  Concerning  homology,  which  was  another  cornerstone  of 
phylogenetic  embryology,  his  position  was  indeed  similar  to  Whitman's:  .  .  .  "//  is 
the  prospective  and  not  the  retrospective  aspect  of  development  that  is  decisive  .  .  ." 
This  is  shown  most  clearly  in  the  case  of  the  germ  layers  and  the  cleavage  stages. 
In  the  latter  case  embryonic  origin  and  position  are  utterly  worthless  apart  from 
developmental  destiny.  In  all  these  cases  homology  is  determined  "not  by  origin  but 
by  fate"  A  few  pages  later,  ".  .  .  the  events  of  ontogeny  are  essentially  adaptive, 
and  .  .  .  the  persistence  of  ancestral  reminiscences  in  development  or  of  similarities 
in  the  development  of  homologous  parts  is  in  some  way  connected  with  the  persistence 
of  ancestral  conditions  of  development"  (Wilson,  1894). 

Wilson  was  also  a  close  friend  of  Anton  Dohrn,  but  again  their  correspondence 
reveals  only  their  respect  for  each  other's  point  of  view  and  remarks  concerning 
common  extrascientific  interests.  Although  our  inquiry  has  not  established  whether 
and  to  what  extent  Dohrn  may  have  influenced  scientists  working  at  the  Station,  or 
their  influence  on  him,  it  does  lead  to  some  indications  as  to  why  the  Naples  Station 
and  the  MBL  developed  in  different  directions.  This  divergent  growth  can  be  traced 
to  the  different  roles  their  founders  conceived  for  them.  When  Anton  Dohrn 
founded  the  Zoological  Station,  science  in  Europe  and  particularly  in  Germany, 
was  flourishing  as  never  before  and  it  enjoyed  great  esteem  in  social  and  political 
circles.  The  university  offered  excellent  teaching  facilities,  but  here  were  two  things 
a  university  could  not  provide.  One  was  a  place  where  people  could  work  under 
conditions  of  complete  freedom,  meaning  that  no  demands  should  be  made  on 
them.  The  second  was  the  marine  material  which  was  proving  increasingly  to  offer 
unique  experimental  opportunities.  In  the  U.  S.,  on  the  other  hand,  biology  was  in 
a  backwater  compared  to  Europe,  the  cultural  center  of  the  world.  The  Americans 
felt  isolated.  This  is  indeed  the  leitmotiv  of  the  articles  that  Whitman  (1883)  and 
Morgan  (1896)  wrote  on  the  Zoological  Station.  Their  most  urgent  need  was  to 
educate  a  class  of  high  caliber  scientists  who  could  compete  with  the  Europeans. 
And,  as  said  before,  one  of  Whitman's  greatest  achievements  was  to  create  the 
Marine  Biological  Laboratory  primarily  as  an  educational  center.  Educational 
meaning,  in  the  broadest  sense,  a  place  where  the  exchange  of  ideas  and  collaborations 
were  almost  forced  upon  the  investigators  through  series  of  lectures  and,  above  all, 
as  a  result  of  their  participation  in  various  courses.  The  Embryology  course  was  one 
of  the  earliest  and  most  successful  courses,  judging  from  the  number  of  eminent 
biologists  who  had  their  starts  there.  What  was  the  take-home  lesson  for  the 
American  embryologists  who  worked  in  Naples?  It  seems  to  be  two-fold.  First,  it 
was  the  idea  of  having  a  laboratory  devoted  to  research  where  people  had  the 
opportunity  to  meet  and  exchange  their  ideas,  a  kind  of  "permanent  congress,"  in 
an  atmosphere  of  complete  freedom.  Second,  contact  with  the  leaders  of  the  new 
intellectual  and  methodological  approach  to  the  problems  of  development  gave  the 
Americans  new  impetus  for  a  fresh  approach  to  their  own  problems. 


THE  "NEW"   EMBRYOLOGY  43 

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in  Graz  am  18  Aug.  1910. 
DOHRN,  A.   1867.  Eugerion  Boeckingi  und  die  Genealogie  der  Arthropoden.  Stettin.  Entomol.  Z.  28: 

145-153. 
DOHRN,  A.  1872.  Der  gegenwartige  Stand  der  Zoologie  und  die  Grundung  zoologischer  Stationen.  Preuss. 

Jahrb  30:  137-161. 
DOHRN,  A.  1875.  Die  Ursprung  der  Wirbelthieren  und  das  Prinzip  des  Funktionswechsel.  W.  Englemann, 

Leipzig. 
DOHRN,  A.  1904.  Studien  zur  Urgeschicht  des  Wirbelthierkorper.  24.  Die  Premandibularhole:  Mitt.  Zool. 

Station  Neapel  17:  117-299. 
DRIESCH,  H.  1899.  Von  der  Methode  der  Morphologic.  Kriiische  Erortemngen.  Biol.  Centralblatt  19: 

3-58. 

DRIESCH,  H.  1909.  Zur  Erinnerung  an  Anton  Dohrn.  Siiddt.  Monatsch.  6:  514. 
DRIESCH,  H.  1951.  Lebenserinnerungen.  Reinhardt  Ver.,  Miinchen  &  Basel. 

EISIG,  H.  1898.  Zur  Entwicklungsgeschichet  der  Capitalliden.  Mitt.  Zool.  Station  Neapel  13,  1-292. 
FRORIEP.  A.  1902.  Einige  Bemerkungen  zur  Kopffrage.  Anal.  An:.,  21:  545-553. 
GEGENBAUR,  C.  1876.  Die  Stellung  und  Bedeutung  der  Morphologic.  Morphol.  Jahrb.  1:  1-19. 
GROEBEN,  CH.,  ed.    1982.  Charles  Darwin  (1809-1882)-Anton  Dohrn  (1840-1909)  Correspondence. 

Macchiaroli,  Napoli. 
HERBST,  C.  1908.  Ueber  die  osmotischen  Eigenschaften  und  die  Entsteheung  der  Befruchtungsmembran 

beim  Seeigelei.  Arch.  Entwiddungsmech.  d.  Org.  26:  82-88. 
HERTWIG,  O.  1876.  Beitrage  zur  Kenntniss  der  Bildung,  Befruchtung  und  Theilung  des  thierischen  Eies. 

Morphol.  Jahrb.  1:  347-434. 
HEUSS,  TH.  1962.  Anton  Dohrn  in  Neapel. 

HORSTADIUS,  S.  1973.  Experimental  Embryology  of  Echinodenns.  Clarendon  Press,  Oxford. 
KUHN,  A.   1950.  Anton  Dohrn  und  die  Zoologische  Station  Neapel.  Publ.  Sta:.  Zool.  Napoli,  Suppl. 

1950,  203  pages. 
LOEB,  J.  1899.  On  the  nature  of  the  process  of  fertilization  and  the  artificial  production  of  normal  larvae 

(plutei)  from  the  unfertilized  eggs  of  the  sea  urchin.  Am.  J.  Physio/.  3:  135-138. 
LOEB,  J.  1900.  On  the  Nature  of  the  process  of  fertilization.  Biol.  Led..  Woods  Hole  1:  273-282. 
LOEB.  J.   1908.  Ueber  die  osmotischen  Eigenschaften  und  die  Entsteheung  der  Befruchtungsmembran 

beim  Seeigelie.  Arch.  Entwiddungsmech.  d.  Org.  26:  82-88. 
MAIENSCHEIN,  J.  1978.  Cell  lineage,  ancestral  reminiscence,  and  the  biogenetic  law.  J.  Hist.  Biol.  11: 

129-158. 

MORGAN,  T.  H.  1896.  Impressions  of  the  Naples  Zoological  Station.  Science  53:  16-18. 
Roux,  W.  1894.  Einleitung.  Arch.  Entwiddungsmech.  d.  Org.  Iu42. 
WARBURG,  O.   1910.  Ueber  die  Oxydationen  in  lebenden  Zellen  nach  Versuche  am  Seeigelei.  Ztscli. 

Physiol.  Chemie  66:  305-340. 

WHITMAN,  C.  O.  1878.  The  embryology  of  Clepsine.  Q.  J.  Microsc.  Sci.  18:  252-258. 
WHITMAN,  C.  O.  1883.  The  advantages  of  study  at  the  Naples  Zoological  Station.  Science  2:  93-97. 
WHITMAN,  C.  O.  1888.  A  contribution  to  the  history  of  germ  layers  of  Clepsine.  J.  Morphol.  1:  105-182. 
WHITMAN,  C.  O.  1894a.  Evolution  and  epigenesis.  Biol.  Led.,  Woods  Hole  3:  205-224. 
WHITMAN,  C.  O.  1894b.  The  inadequacy  of  the  cell  theory  of  development.  Biol.  Led..  Woods  Hole  2: 

105-124. 

WILSON,  E.  B.  1984.  The  embryological  criterion  of  homology.  Biol.  Led.,  Woods  Hole  3:  101-124. 
WILSON,  E.  B.  1895.  An  Atlas  of  the  Fertilization  and  Karyokinesis  of  the  Ovum.  Columbia  Univ.  Press. 
WILSON,  E.  B.  1896.  The  Cell  in  Development  and  Inheritance.  Columbia  Univ.  Press. 
WILSON,  E.  B.  1898a.  Cell  lineage  and  ancestral  reminiscence.  Biol.  Led.,  Woods  Hole  6:  21-42. 
WILSON,  E.  B.  1898b.  Contributions  on  cell  lineage  and  ancestral  reminiscence.  Ann.  N.  Y.  Acad.  Sci. 

11:  1-27. 


Reference:  Biol.  Hull.  168  (suppl.):  44-61.  (June,  1985) 


THE  SCIENCES,    1850-1900,   A   NORTH   ATLANTIC   PERSPECTIVE 

NATHAN   REINGOLD  AND  JOEL  N.   BODANSKY 
Henry  Papers,  Smithsonian  Institution,  Washington,  DC  20560 

ABSTRACT 

For  an  overview  of  the  sciences  in  the  last  half  of  the  nineteenth  century,  a 
series  of  tables  are  presented  on  the  official  support  of  the  sciences  for  Germany, 
the  United  Kingdom,  and  the  United  States  at  ten-year  intervals  plus  a  set  of  tables 
with  a  similar  but  more  detailed  breakdown  for  the  United  States.  The  limitations 
of  the  data  are  discussed  and  certain  subdivisions  of  the  tables  are  analyzed  in  terms 
of  both  national  characteristics  and  indications  of  later  trends.  A  review  of  the 
United  States  situation  at  the  turn  of  the  century  follows,  particularly  stressing  a 
few  relative  differences  from  the  other  two  nations.  The  paper  concludes  with  a 
brief  consideration  of  similarities  disclosed  by  the  data. 

DISCUSSION 

To  give  an  overview  of  what  was  happening  in  the  sciences,  1850-1900,  this 
paper  will  present  a  series  of  quantitative  data  of  carefully  limited  scope.  Much  of 
what  follows  will  consist  of  a  discussion  of  the  characteristics  of  the  data,  of  what 
the  data  apparently  indicates,  and  of  some  general  questions  arising  from  these 
findings.  Quantification  is  not  viewed  by  the  authors  as  necessarily  the  road  to 
historical  understanding,  let  alone  as  encompassing  all  significant  aspects  of  the 
past.  Conceptual,  institutional,  and  ideological  (in  a  non-Marxist  sense)  factors  often 
elude  quantitative  history.  Nevertheless,  our  data  represents  another  way  of  slicing 
the  historical  pie,  and  one  producing  results  of  reasonable  validity  and  even  some 
heuristic  values. 

The  original  computations  resulted  from  the  desire  of  the  senior  author  to 
extend  backward  in  time  the  kind  of  information  for  the  United  States  given  by  the 
National  Science  Foundation's  well  known  series  Federal  Funds  for  Science  in  order 
to  test  the  veracity  of  contemporary  and  retrospective  statements  about  the  history 
of  science  in  the  United  States  and  to  compare  the  situations  in  the  North  American 
republic  with  analogous  developments  in  Europe.  Known,  existing  published  Amer- 
ican sources  only  permit  a  limited  extrapolation  back  to  the  late  1930's;  all  known 
earlier  data  for  this  century  are  patchy  and  flawed  to  a  point  of  near  uselessness. 
For  the  last  century,  an  attempt  was  made  to  gauge  the  development  of  a  professional 
scientific  community  using  numbers  derived  from  studies  of  the  historical  bibliography 
of  the  sciences  and  from  the  occupational  data  given  by  the  decennial  U.  S.  Census. 
Although  lacking  a  high  degree  of  precision,  the  findings  provided  useful  orders  of 
magnitude  illuminating  questions  for  further  research.1 

The  present  paper  is  an  extension  of  the  prior  effort  to  the  estimation  of  the 
official  support  of  the  sciences,  1850-1900,  stimulated  by  the  existence  of  two 
similar  attempts  for  the  United  Kingdom  and  Germany:  R.  M.  MacLeod  and 

E.  K.  Andrews'  unpublished  compilation  for  the  U.  K.,  "Selected  Science  Statistics 
Relating  to  Research  Endowment  and  Higher  Education:  1850-1914,"  1976;  and 

F.  R.  Pfetsch's  Zur  Entwicklung  der  Wissenschaftspolitik  in  Deutschland,   1750- 
1914,  (Berlin,   1974).  The  date  span  of  the  paper  derives  from  the  U.  K.  study's 

44 


THE  SCIENCES,    1850-1900  45 

earliest  data  and  the  fact  that  private  sector  funding  became  much  more  consequential 
in  the  United  States  after  1900,  making  the  federal  funds  a  significantly  lesser  aspect 
of  the  whole  than  for  the  preceding  decades,  a  situation  persisting  until  the  scientific 
mobilization  of  World  War  II.  The  figures  for  the  United  States  are  taken  from 
published  reports  of  receipts  and  expenditures  of  the  Department  of  the  Treasury 
supplemented  as  necessary  by  recourse  to  the  published  annual  reports  of  the 
administrative  entities.  All  the  data  of  the  three  nations  are  formally  comparable  as 
to  origins,  a  fact  that  proves  both  a  virtue  and  a  hazard  to  the  historian. 

The  tables  are  in  two  sets,  one  being  the  comparative  accounts,  the  second 
consisting  of  a  more  detailed  breakdown  for  the  United  States;  in  both  sets  results 
are  given  at  ten  year  intervals  (1850,  1860,  1870,  1880,  1890,  1900).  Presumably, 
similar  detailed  breakdowns  are  possible  for  Great  Britain  and  Germany.  The  three- 
country  limitation  is  regrettable  but  not  fatal,  arising  from  the  lack,  to  our 
knowledge,  of  works  comparable  to  Pfetsch  and  to  MacLeod  and  Andrews.  No 
doubt  similar  data  exists  in  the  official  statistics  of  many  nations,  if  not  in 
unpublished  archival  sources.  The  three  countries  are  an  interesting  sample,  consisting 
of  the  two  leading  national  scientific  communites  at  the  turn  of  the  century  plus  a 
significantly  rising  presence  in  the  world's  scientific  economy.  From  other  evidence 
for  specific  disciplines,  notably  physics,  and  for  other  countries,  particularly  France, 
we  believe  that  little,  if  any,  of  the  general  pattern  disclosed  will  require  major 
amendment.  Nevertheless,  we  hope  others  compile  equivalent  data  for  countries 
like  France,  Italy,  Russia,  Japan,  Austro-Hungary,  Sweden,  the  Netherlands,  and 
Switzerland. 

To  maximize  comparability,  the  U.  S.  data  is  given  by  the  categories  and 
standards  of  the  United  Kingdom  and  German  sources  even  though  this  has 
seriously  complicated  our  labors  at  a  number  of  points.  Pfetsch  is  far  less  explicit 
in  explaining  his  data  than  MacLeod  and  Andrews,  but  we  believe  this  has  not 
vitiated  our  basic  premises.  National  differences,  whether  of  current  compilers  or  of 
the  historic  data,  produce  interesting  problems  not  always  amenable  to  easy  solution. 
Both  the  German  and  British  compilations,  for  example,  include  their  respective 
patent  offices  as  a  "scientific"  activity,  something  we  would  have  excluded  and 
suspect  would  not  have  figured  in  any  U.  S.  computation  of  the  late  nineteenth 
century.  We  regard  the  Patent  Office  as  a  property-granting  entity,  not  one  engaged 
in  research  or  in  the  administration  of  a  development  program.  The  resulting 
swelling  of  the  U.  S.  figures  obscures  the  entire  question  of  the  theory /applications 
relationship. 

Concentrating  on  official  support  obviously  evades  the  entire  question  of  the 
private  sector's  role  in  the  support  of  science,  clearly  an  important  matter  in  the 
United  States  and  in  Great  Britain  (to  a  lesser  extent)  in  our  period  and  at  other 
times,  but  of  still  lesser  importance  in  Germany.  Non-official  support  will  be 
discussed  in  a  few  contexts. 

Concentrating  on  governmental  support  produces  two  important  consequences 
for  the  data  presented  in  these  tables.  The  first  is  that  Pfetsch  can  and  must  give 
considerable  information  on  the  German  states  both  before  and  after  the  founding 
of  the  Bismarckian  empire.  Not  only  are  published  sources  readily  available,  but 
the  former  states,  like  today's  Lander,  play  an  important  role  in  the  political 
economy  of  German  science.  In  contrast,  for  the  United  States,  doing  all  of  the 
states  in  existence  1850-1900  is  a  formidable  research  task  and  one  of  doubtful 
desirability  given  the  ample  indications  of  the  modesty  of  state  and  local  support 
relative  to  federal  patronage.  Nevertheless,  some  such  support  will  be  appropriately 
noted  in  the  discussion,  and  readers  should  assume  that  the  U.  S.  figures  are 
understated  to  a  slight  but  not  yet  determined  degree. 


46  N.   REINGOLD  AND  J.   N.   BODANSKY 

Relying  on  official  sources  means  using  line  items  as  given  in  each  nation's 
respective  national  accounts.  These  do  not  exactly  match.  Further,  we  have 
encountered  many  instances  where  research  funding  is  not  differentiated  from 
general  allocations.  In  the  U.  S.  case  a  number  of  instances  have  yielded,  after 
further  research,  to  more  precise  breakdowns.  In  general,  all  three  national  sets  of 
data  are  derived  from  attempts  to  concentrate  on  organizations  with  primary  or 
clear  R  &  D  missions  or  with  an  R  &  D  support  role.  In  the  U.  S.  case  we  have 
excluded  instances  of  routine,  non-R  &  D  uses  of  scientific  and  technical  personnel 
and  made  judgments  of  inclusion  based  on  evidences  of  scientific  sophistication. 
For  clarity  and  comparability,  data  are  reported  by  administrative  units  even  though 
many  units  will  engage  in  diverse  work  in  terms  of  subject  matter.  Since  further 
subdivision  is  often  not  possible  even  with  arduous  and  perhaps  dubious  estimating 
procedures,  the  present  array  is  a  useful  first  approximation  with  the  virtue  of 
informing  us  of  what  the  past  was  like  in  its  own  terms  without  a  spurious  precision 
deployed  anachronistically  in  the  service  of  today's  concepts. 

Because  of  the  existence  of  an  extraordinary  statistical  compilation  for  the 
discipline  of  physics  around  1900,2  for  comparability  we  will  similarly  report  all 
expenditures  in  German  marks.  Throughout  this  period  there  was  a  great  stability 
of  exchange  rates:  4.2M/$1  and  20.6M/£.  All  values  are  current  without  standard- 
ization for  inflation/deflation  nor  have  we  attempted  to  normalize  for  comparative 
costs  of  living.  For  example,  the  physics  data  indicates  that  U.  S.  figures  are 
overstated  in  comparison  to  Western  Europe  because  of  higher  costs  for  salaries, 
supplies,  and  equipment. 

Let  us  consider  specific  categories  in  the  comparative  national  accounts.  Fully 
in  keeping  with  the  pride  of  place  accorded  in  the  German  rhetoric  of  the  past  and 
in  contemporary  historic  accounts  are  the  large  sums  reported  for  the  support  of 
higher  education  by  the  German  states.  Yet  these  impressive  sums  are  far  from 
unequivocal  in  their  meaning.  First,  the  numbers  are  not  limited  to  research 
expenditures  and  include  work  in  humanistic  and  social  science  fields,  not  only  the 
physical  and  biological  sciences.  They  represent  essentially  the  cost  for  the  mainte- 
nance of  a  system  of  higher  education.  In  contrast,  the  "Anglo-Saxons'"  largely 
attempt  to  restrict  the  sums  to  scientific  and  technical  education.  It  is  not  simply 
the  distinction  between  German  "WissenschafT  and  English  "science"  but  differing 
perceptions  of  the  relationship  of  peaks  of  achievement  to  the  broader  cultural 
infrastructure.  We  suspect  that  the  sciences  were  subsumed  under  a  broader  rubric 
like  "Kultur"  with  research  and  education  in  "Kultur"  being  seen  as  co-extensive. 
That  conjecture  is  reinforced  by  a  second,  related  characteristic  of  the  data.  Pfetsch 
asserts  that  the  Imperial  government  did  not  generally  support  "zweckfrei"  research. 
In  his  analysis  what  we  would  designate  as  pure  research  is  located  in  this  educational 
sector;  in  fact  all  of  those  funds  are  designated  "zweckfrei."  As  we  cannot  conceive 
of  that  categorization,  there  is  obviously  a  differing  concept  here  of  the  relationship 
of  pure  research  to  application.  The  two  issues  raised  in  connection  with  higher 
education  crop  up  elsewhere  in  the  tables;  the  British  occupy  a  middle  position, 
perhaps  somewhat  closer  to  the  Germans  than  to  their  American  cousins.  Pfetsch 
and  other  Germans  of  the  past  do  not  consider  the  training  of  individuals  for  the 
professions  as  a  mission  and  tend  to  restrict  "zweck"  largely  to  whatever  effects 
"wirtschaftswachstum."3 

Also  predictable  are  the  large  U.  S.  investments  in  knowledge  and  use  of  natural 
resources  (lands,  forests,  agriculture,  fisheries),  given  the  obvious  drive  to  settle  and 
to  exploit  a  large  continental  mass.  Two  qualifications  are  pertinent.  Given  the 
comparative  areas,  the  German  and  British  sums  are  hardly  disproportionately 


THE  SCIENCES,    1850-1900  47 

small,  especially  if  we  remember  that  the  analagous  costs  for  the  imperial  possessions 
of  Britain  are  not  in  these  tables  but  in  the  separate  governmental  accounts  of 
India,  Canada,  Australia,  Egypt,  etc.  Given  what  is  known  of  similar  French  and 
Russian  activities,  everybody  was  doing  it,  not  simply  the  expanding  U.  S.  republic. 
If  there  is  a  crucial  difference,  it  is  qualitative.  The  Americans  were  particularly 
adroit  here  in  linking  scientific  research  with  efficient  technological  exploitation.  To 
be  more  exact,  a  vast  research  enterprise  arose  with  a  basic  research  component 
supported  for  its  own  sake  and  for  a  belief  in  its  ultimate  utility.  The  Fish 
Commission,  associated  with  the  history  of  MBL,  is  a  modest  example. 

Notably  absent  from  the  U.  S.  federal  figures  are  sums  for  academies  and 
societies.  While  some  state  funds  went  to  these  bodies,  those  organizations  are 
overwhelmingly  part  of  the  private  sector  to  this  day.  European  observers  and  some 
American  scientists  used  to  regard  this  as  a  serious  blemish,  disregarding  that  learned 
and  professional  societies  were  proliferating  in  the  country  from  1850  even  to  this 
day.  Disregarded  also  is  the  obvious  point  that  in  most  western  countries  in  the 
same  period  the  support  and  conduct  of  research  have  increasingly  come  in  the 
hands  of  other  kinds  of  organizations  (government  bureaus,  universities,  research 
laboratories,  industrial  concerns,  etc.).  The  particular  arrangement  in  the  United 
States  has  had  no  obvious  deleterious  effect  on  the  growth  of  the  sciences.  What 
vexed  U.  S.  scientists  from  time  to  time  was  the  sense  of  a  lack  of  an  assured  high 
status  in  a  national  hierarchy. 

Both  III,  Medicine  and  Health,  and  VIII,  Military,  have  interesting  limitations 
to  their  data  and  at  the  same  time  are  unexpectedly  informative.  For  all  three 
countries,  attempts  were  made  to  separate  out  routine  hospital  and  clinical  care  in 
favor  of  medical  research  and  public  health  administration.  Roughly,  a  distinction 
was  made  between  prevention  and  treatment.  As  far  as  the  sources  permit,  our  data 
estimates  the  former.  Completely  omitted  for  the  U.  S.,  for  example,  is  the  hospital 
system  of  the  Marine  Hospital  Service  (the  forerunner  of  the  U.  S.  Public  Health 
Service)  and  the  medical  departments  of  the  Army  and  Navy.  By  analogy  with  the 
British  Local  Government  Health  Board  and  similar  German  bodies,  we  have 
included  the  federally  funded  District  of  Columbia  Health  Department.  As  no 
American  state  and  local  entities  are  reported,  U.  S.  figures  understate  the  support 
for  this  activity.  The  1890  and  1900  numbers  include  a  small  sum  for  the  Marine 
Hospital  Services'  Hygienic  Laboratory,  the  forerunner  of  the  National  Institutes  of 
Health.  We  should  note  here  that  Pfetsch,  like  other  Germans,  classifies  medicine 
as  "zweckfrei,"  again  a  categorization  we  cannot  accept. 

The  military  figures  probably  understate  slightly  the  amount  of  investment  in 
weaponry.  Only  rarely  are  such  sums  given  separately.  We  assume  that  R  &  D  for 
warfare,  to  the  extent  actually  performed  in  that  era,  is  largely  subsumed  under  the 
heading  of  procurement  costs  and  the  like.  Much  of  the  U.  S.  expenditures  are  for 
peaceful  (or  at  least,  semi-warlike)  purposes.  Although  similar  uses  were  made  of 
the  military  in  other  countries,  the  lack  of  a  serious  military  threat  in  the  last 
century  coupled  with  the  general  sense  in  the  last  century  of  a  lack  of  alternate 
institutions  and  a  shortage  of  skilled  personnel  resulted  in  the  pattern  disclosed 
here.  When  the  security  environment  changed  markedly  during  and  after  World 
War  II,  it  was  relatively  easy  for  the  defense  establishment  to  assume,  openly  and 
otherwise,  a  substantial  role  in  the  support  of  the  sciences. 

Finally,  the  U.  K.  support  of  museums  is  eye-catching.  Even  making  allowances 
for  private  sector  support  in  the  United  States,  the  sums  expended  are  quite  high, 
quite  disproportionate  to  our  subjective  sense  of  the  relative  merits  of  contributions 
from  the  three  nations.  We  suspect  that  the  U.  K.  figures  represent,  in  part,  the 


48  N.   REINGOLD  AND  J.   N.   BODANSKY 

enduring  strength  of  a  wide  public  interest  and  participation  in  natural  history  in 
that  country.  We  hesitate  to  speculate  on  how  this  passion  for  museums  aided  or 
hindered  the  growth  of  support  for  experimental  and  quantitative  branches  of 
biology,  not  to  mention  geophysics  with  its  quite  different  approach  to  the  history 
of  the  Earth. 

To  conclude,  we  will  first  look  at  the  United  States  around  the  turn  of  the 
century  and  then  offer  observations  on  a  number  of  characteristics  common  to  the 
three  nations.  From  Forman,  Heilbron,  and  Weart,  we  know  that  in  1900  physics 
in  the  United  States  received  2,990,000  M,  partly  from  state  governments  but  mostly 
from  the  private  sector.  To  date,  we  have  not  succeeded  in  obtaining  any  reliable 
estimate  for  research  support  in  general  in  1900.  Shortly  after  the  turn  of  the 
century,  the  newly  established  Carnegie  Institution  of  Washington  reported  that  the 
funds  specifically  earmarked  for  the  support  of  research  totaled  $2,952,642,  yielding 
an  annual  income  of  $  199, 625. 4  But  that  figure  is  clearly  grossly  on  the  low  side, 
given  both  the  federal  numbers  and  the  data  for  physics.  What  CIW  reported  were 
the  relatively  few  endowments  solely  designated  for  research  or  nearly  so,  omitting 
support  coming  from  other,  more  general  bodies  of  funds.  In  doing  that,  the 
Carnegie  Trustees  not  only  exhibited  a  desire  to  increase  funding  ticketed  exclusively 
for  research  but,  as  we  shall  shortly  note,  acted  in  accordance  with  a  still  persisting 
national  characteristic.  Since  we  know  that  higher  education  then  and  now  loomed 
large  in  U.  S.  science,  let  us  consider  that  class  of  institutions.  In  1900  more  than 
4%  of  the  18-21  age  group  were  enrolled  in  American  colleges  and  universities,  a 
high  percentage  in  terms  of  European  norms  of  that  date.  The  endowments  of  these 
institutions  (overwhelmingly  private)  totaled  $166,193,529,  yielding  an  annual 
income  of  $1 1,995,463.  To  this  must  be  added  the  considerable  sums  received  for 
tuition.  Although  overwhelmingly  devoted  to  instruction,  clearly  some  fraction  of 
higher  education's  resources  went  into  research.5 

Obviously,  by  1900  the  United  States  of  America  was  engaged  in  an  extensive, 
if  uneven,  effort  to  expand  research  and  development.  Certain  peculiarities  are 
worth  noting  with  the  proviso  that  they  are  relative,  not  absolute,  differences.  First, 
the  private  sector  plays  a  greater  role  in  the  U.  S.,  then  and  now.  Second,  we  infer 
from  our  data  and  later  evidence  a  persisting  desire  to  give  priority  to  erecting  a 
supporting  infrastructure,  rather  than  simply  to  target  great  men,  great  ideas, 
intellectual  breakthroughs,  or  the  like.  Even  so-called  "wars"  (on  cancer)  or 
mobilizations  (for  defense  or  space)  were  accompanied  by  a  concern  for  expanding 
facilities  and  the  supply  of  specialized  personnel.  We  conjecture  this  arose  initially 
from  a  sense  of  deficiency  in  the  national  scheme  of  things  and  suspect  that  in 
Europe  a  contrary  view  prevailed  at  many  points  in  time  based  on  a  degree  of 
complacency  buttressed  by  an  unspoken  motive  to  perpetuate  a  desired  hierarchical 
social  order. 

Third,  we  are  impressed  by  the  relative  willingness  of  the  U.  S.  organizations 
and  institutions  to  split  out  research  and  other  activities  from  otherwise  undiffer- 
entiated  wholes;  witness  the  contrasting  treatment  of  the  German  university  and 
the  Carnegie  Institution's  narrow  focus  on  research  endowments.  Partly  this  arises 
because  Europeans  tend  to  see  all  these  activities  as  part  of  a  cultural  whole,  in  turn 
part  of  a  specific  stratified  social  order.  The  U.  S.  tendency,  (manifested  in  such 
seemingly  narrow  technicalities  as  the  wording  of  Circular  A-21  on  research  costs), 
represents  a  traditional  view  of  a  limited  role  for  government  requiring,  therefore, 
a  careful,  often  nit-picking  analytic  differentiation  by  function,  process,  intent, 
origin,  even  ultimate  result.  This  tendency  encourages  attempts  at  precise  differen- 
tiations of  "pure,"  "applied,"  "development,"  and  other  categories.  That  such 


THE  SCIENCES,    1850-1900  49 

analyses  hardly  represent  the  real  world  of  research  and  development  is  quite 
apparent  from  the  disputes  in  the  United  States  since  World  War  II  over  support 
of  university  science,  the  role  of  health  research,  and  the  nature  of  research  and 
development  under  the  auspices  of  the  armed  services.  It  is  not  accidental  that  this 
is  the  case;  all  three — the  university,  the  health  sector,  and  the  defense  establishment — 
are  multi-purpose  continuums  whose  individual  parts  make  only  partial  sense 
isolated  from  the  whole. 

All  three  nations  display  great  overall  similarity  in  their  patterns  of  funding, 
1850-1900.  Even  a  difference  like  the  absence  of  U.  S.  support  for  academies  and 
societies  is  an  artifact  of  data  limited  to  the  federal  sector.  Nor  is  that  surprising  as 
all  three  are  part  of  a  larger  entity — Western  Civilization — and  one  can  assume  an 
overall  tendency  to  emulate,  if  not  to  compete  with,  successful  innovations  anywhere, 
both  intellectual  and  administrative.  We  are  postulating  a  kind  of  steady  state,  a 
situation  now  spread  to  large  portions  of  the  globe.  Each  major  scientific  country 
and  many  lesser  scientific  nations  observe  changes  in  other  national  communities 
and  very  often  adjust  their  national  accounts  accordingly. 

Despite  the  understandable  emphasis  in  the  literature  on  novelties — conceptual, 
factual,  and  applied — what  the  data  shows  in  this  period  (and  similarly  for  later  in 
this  century)  is  not  a  grand  commitment  to  the  generation  of  peaks  of  creativity. 
On  the  contrary,  at  all  times,  in  so  far  as  we  can  judge  from  this  and  other 
admittedly  fragmentary  evidence,  the  preponderance  of  support  is  for  the  routine 
as  judged  in  retrospect  and  even  as  given  in  the  perspective  of  each  historical  epoch. 
"Routine"  as  used  by  us  does  not  necessarily  imply  lack  of  sophistication  nor  are 
we  equating  that  term  with  applied  work.  What  we  mean  is  that  the  funding  is  to 
continue  patterns  of  activities  and  behavior  already  in  place  which  may  or  may  not 
be  changing  in  some  significant  ways,  quantitative  or  qualitative. 

Nor  is  that  necessarily  bad  from  the  perspective  of  intellectual  advance.  The 
funding  pattern,  taken  as  a  whole,  reflects  two  purposes:  the  maintenance  of  a 
culture  of  "science"  or  of  "research,"  and  the  erection  of  a  supporting  infrastructure 
of  organizations,  social  processes,  and  value  judgments  furthering  the  maintenance 
and  growth  of  that  culture.  What  that  indicates  to  us  is  that  current  arguments 
about  R  &  D  funding  in  the  U.  S.  are  couched  in  traditional  terms  selected  hopefully 
for  their  power  to  convince  legislators,  administrators,  and  other  dispensers  of  funds. 
What  is  largely  absent  from  such  arguments  are  the  real  issues:  disputes  about  the 
nature  of  the  culture  of  research  and  disputes  about  protecting  or  expanding 
particular  pieces  of  the  infrastructure.  What  is  important  from  the  perspective  of 
intellectual  advance  as  a  goal,  whatever  the  specifics  at  issue,  is  that  now,  as  in  the 
years  1850-1900,  there  is  general  agreement  that  intellectual  advance  is  important 
for  its  own  sake  and  for  its  possible  applied  consequences. 

ACKNOWLEDGMENTS 

Grateful  acknowledgement  is  made  to  the  National  Endowment  for  the  Human- 
ities and  the  National  Historical  Publications  and  Records  Commission  for  their 
support  in  the  preparation  of  both  the  text  and  the  appendix. 

NOTES  TO  TEXT 

1  REINGOLD,  N.  1976.  "Definitions  and  speculations:  the  professionalization  of  science  in  America  in  the 
nineteenth  century."  Pp.  33-69  in  The  Pursuit  of  Knowledge  in  the  Early  American  Republic 
.  A.  Oleson  and  S.  Brown,  eds.  Baltimore. 


50  N.   REINGOLD  AND  J.   N.   BODANSKY 

2FoRMAN,  PAUL,  JOHN  L.  HEILBRON,  AND  SPENCER  WEART.  1975.  Physics  circa  1900:  Personnel, 
Funding,  and  Productivity  of  the  Academic  Establishment,  vol.  5  of  Historical  Studies  in  the 
Physical  Sciences.  Princeton. 

3  Pfetsch  makes  a  distinction  between  support  for  "Imperializing"  activities  following  German  unification, 

"general"  support  described  as  "mainly  academic  and  medical,"  and  economically  oriented 
research.  To  North  American  eyes  "zweck"  or  mission  related  research  occurs  under  each 
category.  A  notable  example  of  how  concentrating  on  economic  results  organizes  quantitative 
data  bearing  on  the  sciences  is  PETER  LUNDGREEN,  BiUiung  und  Wirtschaftswachstwn  in 
Industrialisierungsprozess  des  19.  Jahrhunderts,  Berlin,  1973. 

4  Carnegie  Institution  of  Washington,  Report  oj  the  Executive  Committee  to  Board  oj  Trustees,  Washington, 

1902,  pp.  247-269. 

5  See:  REINGOLD,  N.  1978.  National  style  in  the  sciences:  the  United  States  case.  Pp.  163-173  in  Human 

Implications  oj  Scientific  Advance,  E.  G.  Forbes,  ed.  Edinburgh. 


I.  COMPARATIVE  TABLES 
A.   1850* 


GERMAN 

USA 

UK             REICH 

STATES 

I.         AGRICULTURE  AND  FISHERIES 

18 

115 

106 

A.   Agriculture  (including 

Forestry) 

18 

— 

B.    Botanical  Gardens 

— 

115 

C.   Fisheries 

— 

— 

II.  GENERAL  SCIENCE,  TECHNOLOGY  1750               163                                                62 

A.  Geophysics  (including 

topographic  surveys)  1247 

B.  Geology  122               163 

C.  Meteorology 

D.  Standards  [8] 

E.  Patents  318 

F.  Other  63                                                                    40 

III.  MEDICINE  AND  HEALTH  205  682 

IV.  ACADEMIES  AND  SOCIETIES  129  252 

V.  OTHER  SCIENTIFIC  ACTIVITIES  385 

VI.  MUSEUMS  84  25 

VII.  HIGHER  EDUCATION  1  40  2507 

VIII.  MILITARY  732  111  245 

A.  Army  206 

1.  Surveys  153 

2.  Education  53 

3.  Other 

B.  Navy  526  111 

1.  Hydrography 

2.  Astronomy  (including  j  405 

nautical  instruments)  1  1  1 

3.  Nautical  Almanac  —  — 


4.  Education 

63 

5.  Other 

58 

TOTAL 

2501               847 

4264 

All  amounts  given  in  1000's  of  German  marks. 


THE   SCIENCES.    1850-1900 


51 


1.  COMPARATIVE  TABLES 
B.   1860* 


GERMAN 

USA 

UK              REICH 

STATES 

I. 

AGRICULTURE  AND  FISHERIES 

201 

552 

173 

A.   Agriculture  (including 

Forestry) 

168 

— 

B.    Botanical  Gardens 

33 

330 

C.   Fisheries 

— 

222 

II. 

GENERAL  SCIENCE.  TECHNOLOGY 

3225 

399 

143 

A.   Geophysics  (including 

topographic  surveys) 

2236 

— 

B.   Geology 

— 

399 

— 

C.   Meteorology 

— 

— 

— 

D.  Standards 

[10] 

— 

— 

E.    Patents 

967 

— 

— 

F.    Other 

22 

— 

143 

III. 

MEDICINE  AND  HEALTH 

3 

124 

759 

IV. 

ACADEMIES  AND  SOCIETIES 

— 

161 

100 

V. 

OTHER  SCIENTIFIC  ACTIVITIES 

630 

674 

81 

VI. 

MUSEUMS 

17 

641 

126 

VII. 

HIGHER  EDUCATION 

— 

257 

3221 

VIII 

MILITARY 

1184 

1246 

195 

A.  Army 

691 

270 

195 

1.  Surveys 

611 

— 

2.  Education 

74 

239 

3.  Other 

6 

31 

B.    Mm- 

493 

976 

1.  Hydrography 

C 

626 

2.  Astronomy  (including 

329 

nautical  instruments) 

1 

222 

3.  Nautical  Almanac 

69 

91 

4.  Education 

39 

— 

5.  Other 

56 

37 

TOT  A  I 

5260 

4054 

4798 

*  All  amounts  given  in  1000's  of  German 

marks. 

I.  COMPARATIVE  TABLES 

C.    1870* 

GERMAN 

USA 

UK              REICH 

STATES 

I. 

AGRICULTURE  AND  FISHERIES 

734 

1184 

569 

A.   Agriculture  (including 

Forestry) 

628 

280 

B.    Botanical  Gardens 

106 

515 

C.   Fisheries 

— 

389 

II.       GENERAL  SCIENCE,  TECHNOLOGY  4642  3435 

A.   Geophysics  (including 

topographic  surveys)  2215  2461 


33 


149 


52 


N.    REINGOLD   AND  J.   N.    BODANSKY 


I.  COMPARATIVE  TABLES  (Continued) 
C.    1870* 


GERMAN 

USA 

UK              REICH 

STATES 

B.   Geology 

83 

700 

( 
\    n 

C.   Meteorology 

[63] 

206 

I    '- 

D.  Standards 

[18] 

33 

5 

E.    Patents 

2340 

—                   — 

— 

F.    Other 

4 

68 

132 

III. 

MEDICINE  AND  HEALTH 

5 

439 

975 

IV. 

ACADEMIES  AND  SOCIETIES 

— 

37 

113 

V. 

OTHER  SCIENTIFIC  ACTIVITIES 

8 

126 

349 

VI. 

MUSEUMS 

17 

1030 

230 

VII. 

HIGHER  EDUCATION 

— 

409 

5244 

VIII 

.  MILITARY 

1939 

1580                744 

382 

A.   Army 

1257 

402                709 

382 

1.  Surveys 

1084 

— 

2.  Education 

110 

257 

3.  Other 

63 

145 

B.   Navy 

682 

1178                  35 



1.  Hydrography 

173 

914 

2.  Astronomy  (including 

nautical  instruments) 

329 

193 

3.  Nautical  Almanac 

96 

50 

4.  Education 

84 

— 

5.  Other 

— 

21 

TOTAL 

7345 

8240                777 

801  1 

*  All  amounts  given  in  1000's  of  German 

marks. 

I.  COMPARATIVE  TABLES 

D.    1880* 

GERMAN 

USA 

UK             REICH 

STATES 

I. 

AGRICULTURE  AND  FISHERIES 

1,560 

1,229               460 

1,455 

A.   Agriculture  (including 

Forestry) 

903 

534 

B.    Botanical  Gardens 

85 

395 

C.   Fisheries 

572 

300 

II. 

GENERAL  SCIENCE,  TECHNOLOGY 

5,018 

4,384             1,516 

679 

A.  Geophysics  (including 

topographic  surveys) 

2,379 

2,853 

B.   Geology 

448 

1,008 

1  £.  A 

C.   Meteorology 

[1,575] 

299 

\  164 

D.  Standards 

27 

33                100 

5 

E.    Patents 

2,118 

43               624 

— 

F.    Other 

46 

148               792 

510 

III. 

MEDICINE  AND  HEALTH 

960 

1,597                128 

1.318 

IV. 

ACADEMIES  AND  SOCIETIES 



42 

281 

THE   SCIENCES,    1850-1900 


53 


I.  COMPARATIVE  TABLES  (Continued) 
D.   1880* 


GERMAN 

USA 

UK 

REICH 

STATES 

V. 

OTHER  SCIENTIFIC  ACTIVITIES 

85 

38 

— 

380 

VI. 

MUSEUMS 

926 

3,820 

87 

326 

VII. 

HIGHER  EDUCATION 

10 

521 

400 

14.604 

VIII. 

MILITARY 

3,106 

1,777 

1,553 

— 

A.  Army 

2,176 

294 

949 

— 

1.  Surveys 

416 

— 

2.  Education 

122 

231 

3.  Other 

1,638 

63 

B.    Navy 

930 

1,483 

604 

— 

1.  Hydrography 

251 

961 

2.  Astronomy  (including 

nautical  instruments) 

287 

274 

3.  Nautical  Almanac 

109 

67 

4.  Education 

90 

175 

5.  Other 

193 

6 

TOT  A  I 

11,665 

13,408 

4.144 

19,043 

*  All  amounts  given  in  1000's  of  German 

marks. 

I.  COMPARATIVE  TABLES 

E.    1890* 

GERMAN 

USA 

UK 

REICH 

STATES 

I. 

AGRICULTURE  AND  FISHERIES 

8,120 

1,269 

40 

1,455 

A.   Agriculture  (including 

Forestry) 

6,718 

463 

B.    Botanical  Gardens 

96 

429 

C.   Fisheries 

1,306 

377 

II. 

GENERAL  SCIENCE,  TECHNOLOGY 

9,503 

7,164 

1,833 

1,478 

A.   Geophysics  (including 

topographic  surveys) 

1,888 

4.445 

B.   Geology 

3,220 

900 

— 

1     199 

C.    Meteorology 

[830] 

315 

— 

I 

D.  Standards 

16 

87 

125 

5 

E.    Patents 

3,455 

1,205 

1.159 

— 

F.    Other 

924 

212 

549 

1,274 

III. 

MEDICINE  AND  HEALTH 

468 

1.294 

199 

2,505 

IV. 

ACADEMIES  AND  SOCIETIES 

— 

119 

— 

304 

V. 

OTHER  SCIENTIFIC  ACTIVITIES 

43 

21 

3 

1,351 

VI. 

MUSEUMS 

757 

2,425 

63 

815 

VII. 

HIGHER  EDUCATION 

21 

1,041 

400 

18,224 

VIII 

.  MILITARY 

3,009 

1,965 

2,294 

— 

A.   Army 

1,165 

398 

1,855 

— 

1.  Surveys 

14 

— 

2.  Education 

150 

93 

3.  Other 

1,001 

305 

54 


N.    RKINGOLD   AND  .1.    N.    BODANSKY 


1.  COMPARATIVE  TABLETS  (Continued) 
E.    1890* 


GERMAN 

USA 

UK             REICH 

STATES 

B.    Navy 

1,844 

1,567               439 

1.  Hydrography 

408 

564 

2.  Astronomy  (including 

nautical  instruments) 

842 

344 

3.  Nautical  Almanac 

125 

74 

4.  Education 

105 

208 

5.  Other 

364 

377 

TOTAL 

21,921 

15,298            4,832 

26,132 

*  All  amounts  given  in  1000's  of  German 

marks. 

I.  COMPARATIVE  TABLES 

F.    1900* 

GERMAN 

USA 

UK              REICH 

STATES 

I.         AGRICULTURE  AND  FISHERIES 

13,375 

2,100               528 

2.452 

A.   Agriculture  (including 

Forestry) 

11,072 

1  .  1  54 

B.    Botanical  Gardens 

109 

615 

C.   Fisheries 

2,194 

331 

II.       GENERAL  SCIENCE,  TECHNOLOGY 

13.874 

7,432             3,629 

2,519 

A.   Geophysics  (including 

topographic  surveys) 

2,243 

4,342 

B.   Geology 

2,841 

829 

f    ,,A 

C.   Meteorology 

4.157 

315               1    3 

j     436 

D.  Standards 

44 

80               436 

8 

E.    Patents 

4,122 

1,405             2,484 

— 

F.    Other 

467 

461                706 

2,075 

III.      MEDICINE  AND  HEALTH 

1,860 

1,967                605 

2,938 

IV.      ACADEMIES  AND  SOCIETIES 

— 

152 

413 

V.       OTHER  SCIENTIFIC  ACTIVITIES 

2,409 

240 

3,508 

VI.      MUSEUMS 

935 

2,926                100 

1,031 

VII.    HIGHER  EDUCATION 

5,082 

2,073                400 

27,839 

VIII.  MILITARY 

2,614 

1,978             2,761 



A.   Army 

893 

180             2,366 



1.  Surveys 

155 

— 

2.  Education 

189 

124 

3.  Other 

549 

56 

B.   Navy 

1,721 

1,798                394 



1.  Hydrography 

602 

740 

2.  Astronomy  (including 

nautical  instruments) 

359 

459 

3.  Nautical  Almanac 

117 

82 

4.  Education 

126 

154 

5.  Other 

517 

363 

TOTAL 

40,149 

18,868             8,023 

40,700 

*  All  amounts  given  in  1000's  of  German  marks. 


THE  SCIENCES.    1850-1900 


55 


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2.  Naval  Observatory 
Nautical  Instruments36 
Other  Astronomy37 

3.  Nautical  Almanac38 
4.  Naval  Academy  —  science39 

5.  Meteorology40 

n  d\ 

surveys 
Torpedo  Corps42 
Other  Ordnance  and  Steam  Engineering  Experiments43 
Miscellaneous44 

*  Estimate. 
1  The  forerunner  of  the  U.  S.  Department  of  Agriculture,  the  Pate 
:nce  in  the  Government.  In  1862,  the  work  of  the  division  was  transfi 
2  Unlike  its  counterpart  in  the  twentieth  century,  with  its  heavy  emp 
Agriculture  was  primarily  a  center  for  the  promotion  of  agricultural  s< 
luded  research  in  biology,  botany,  chemistry,  zoology  and  entymology 
purchase  and  distribution  of  seeds.  Because  its  orientation  in  this  pe 
ept  that  for  1900  the  expenses  of  the  Weather  Bureau,  which  though  i 

3  Though  a  private  botanical  garden,  under  the  sponsorship  of  the 
ction,  the  first  direct  appropriation  for  a  national  Botanical  Garden  ca 
[he  Wilkes  Expedition  on  the  site  of  the  by  then  defunct  Columbian  1 
4  The  United  States  Fish  Commission  was  established  by  Congress 
:arch  in  icthyology  and  marine  biology. 
5  The  survey  of  the  coasts  of  the  United  States  was  first  authorized 
vever.  Congress  repealed  its  original  authorization,  and  the  Coast  Sur 
War  and  Navy  Departments.  In  1832,  Congress  reestablished  the  Coa 
partment  at  the  beginning  of  this  century,  save  for  the  period  1834-1 
Coast  and  Geodetic  Survey  in  recognition  of  the  geodetic  activities  wi 

IIUI1  IU  lla  lUJJUgiapiiiv.,  nyuivjgia^iiiiv.,  anvj  gv.uvjv.iiv.  juivv-^ing  u(jv.i 

:ion,  and  even  a  little  meteorology,  the  great  preponderance  of  its 
860  —  Minnesota  boundary,  Texas  boundary. 
870  —  California-Oregon  boundary,  Nebraska  boundary.  Nebrask; 
880  —  Colorado-Utah  boundary. 
900  —  Idaho-Montana  boundary.  Locating  the  98th  meridian. 
850  —  U.  S.-Mexico  boundary,  U.  S.  -British  provinces  northeastei 

860  —  U.  S.-Mexico  boundary. 
Jsted  in  expenditures  as  "Completing  geological  surveys  of  Michi 

1|* 

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58 


N.   REINGOLD  AND  J.   N.   BODANSKY 


F.  V.  Hayden,  originated  in  a  Congressional 
though  the  survey  included  work  in  natural 
(  1  )  its  mission  was  primarily  geological  and 
stence  in  1879. 
ical  Survey  of  the  Territories  (Hayden)  and 
and  the  Geographical  Surveys  West  of  the 
(King),  was  also  antecedent  to  the  creation 
e  9.  For  King  Survey,  see  footnote  28.  For 
itures  for  either  1870  or  1880. 
rritories". 

nditures  for  this  establishment  are  therefore 
y  —  Meteorology,  but  in  brackets  to  indicate 
ig  to  farmers,  a  new  Weather  Bureau  was 
870,  some  meteorological  observations  had 

1f*tf*r\rr\\r\n\/\  tVia  \/fo^,'^il  r*r*A  CT«^ 

was  under  the  direction  of  the  Smithsonian 
:eorological  work  was  supported  from  the 
n  1873,  the  meteorological  activities  of  the 

line  appropriation  for  this  purpose  appears 
oast  Survey  expenditures,  but  which  it  has 
are  used  to  indicate  that  this  is  the  second 

designated  the  Office  of  Standard  Weights 
National  Bureau  of  Standards. 

;s  of  expenditures,  in  the  case  of  the  Patent 
or  National  Museum,  for  example,  where 
hie  most  imposing  structures  in  Washington 
jres  of  the  Patent  Office  by  including  these 
and  1870. 

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TABLE  II  (Continued) 

jeographical  Survey  of  the  Territories,  or  Hayden  Survey,  under  the  direction  of  geologist 
logical  survey  of  Nebraska,  by  which  label  it  is  still  identified  in  the  1870  expenditures.  Al 
ly,  along  with  its  strictly  geological  activities,  it  has  been  included  under  Geology  because 
the  U.  S.  Geological  Survey,  under  which  it  was  subsumed  when  the  latter  came  into  exi 
yey  was  established  in  1879  by  the  merger  and  expansion  of  the  Geological  and  Geograph 
1  Surveys  of  the  Rocky  Mountain  Region  (Powell),  both  under  the  Interior  Department, 
•  the  War  Department.  A  fourth  survey,  the  Geological  Exploration  of  the  Fortieth  Parallel 
hough  it  had  actually  terminated  its  work  a  year  earlier.  For  Hayden  Survey,  see  footnot 
.  The  Powell  Survey,  which  was  active  from  1874  to  1879  does  not  appear  in  the  expend 
e  statistical  information  concering  the  gold  and  silver  mines  of  the  western  States  and  Te 
n  the  Territories. 

» 
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lie  weather  service  was  established  in  the  Signal  Corps  of  the  U.  S.  Army  in  1870.  The  expe 
although  for  convenience  they  have  also  been  shown  under  General  Science  and  Technolog 
uded  elsewhere.  In  1891,  in  recognition  of  the  importance  of  reliable  weather  forecastil 
Agriculture,  and  the  meteorological  work  of  the  Signal  Corps  transferred  to  it.  Prior  to  1 
Government  deoartments.  including  the  Hvrlrnpranhir  Offirp  nf  thp  Naw  r«pp  Naw  \. 

ie  General  Land  Office.  The  largest  meteorological  establishment  prior  to  1870,  however, 
19,000  M  to  support  its  network  of  weather  observations.  But  the  Smithsonian's  mel 
not  Congressional  appropriations,  and  thus  cannot  be  counted  as  Government  science.  I 
e  Signal  Corps. 

dard  weights  and  measures  was  established  in  the  Coast  Survey  in  1830,  the  first  specific 
for  1850,  1860,  and  1870,  therefore,  represent  funds  which  are  included  in  the  general  C 
devoted  to  salaries  for  the  Survey's  weights  and  measures  activities.  Again,  the  brackets 

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al  expenditures  on  buildings  for  scientific  organizations  have  been  included  in  the  estimat< 
been  made.  In  contrast  to  the  buildings  for  the  Botanic  Gardens.  Naval  Observatory, 
instructed  specifically  suited  to  the  needs  of  the  institution,  the  Patent  Building  was  one  oft 
the  activities  it  housed.  For  this  reason,  it  was  thought  undesirable  to  inflate  the  expenditl 
refore  been  deleted  from  the  totals  for  the  years  where  it  is  relevant,  namely  1850,  1860 

jnses  of  the  Patent  Office's  Agricultural  Division  have  been  separated  out  and  placed  unc 
ring  Expedition,  under  the  direction  of  Lieutenant  Charles  Wilkes,  was  confined  to  the 

ollections,  etc.,  continued  for  many  years,  and  it  was  for  this  purpose  that  the  expenditui 
>ark  was  authorized  by  Congress  in  1889.  The  unusually  large  expenditures  for  1890  pres 

is  report;  Smithsonian:  protection  against  fire. 

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Reference:  Bio/  Hull.  168  (suppl.):  62-79.  (June,  1985) 


LAYING  THE  GHOST:   EMBRYONIC   DEVELOPMENT, 

IN   PLAIN  WORDS 

PAUL   R.  GROSS 

Marine  Biological  Laboratory,  Woods  Hole,  Massachusetts  02543 

—Met  him  what?  he  asked. 

—Here,  she  said.  What  does  that  mean? 

—He  leaned  downward  and  read  near  her  polished  thumbnail. 

—Metempsychosis? 

—Yes.  Who's  he  when  he's  at  home? 

—Metempsychosis,  he  said,  frowning.  It's  Greek:  from  the  Greek.  That  means 

the  transmigration  of  souls. 
— O  rocks!  she  said.  Tell  us  in  plain  words. 

— James  Joyce,  Ulysses.  II. 

CREATIVE  GHOSTS'- 

The  battle  between  Preformationism  and  Epigenesis  is  more  than  its  recurrent 
skirmishes  on  the  field  of  embryology.  It  is  among  the  oldest  of  philosophical  battles. 
In  the  seventeenth  century,  Thomas  Hobbes,  from  whom  a  materialist,  epigeneticist 
position  might  have  been  expected,3  nevertheless  made  the  pure  preformationist 
case,  in  plain  words:  "Nothing  taketh  a  beginning  from  itself/'  (The  epigeneticist 
response  could  have  been:  "Creation  occurs;  complexity  obviously  arises  from 
simplicity.")  On  this  issue  philosophers  and  scientists  have  spoken  inconsistently 
more  often  than  on  most  others,  and  great  biologists  have  been  no  exception. 

Epigenetics  of  the  nineteenth  century,  when  the  Stazione  Zoologica  in  Naples 
was  a  beacon  and  the  MBL  in  Woods  Hole  was  gathering  strength,  did  not  require 
Creation  to  follow  Archbishop  Ussher's  scenario — the  world  established  by  successive 
strokes  of  God  a  few  thousand  years  earlier.  It  could  be  more  subtle.  Moreover, 
questions  of  meaning  and  "explanation"  were  in  the  air,  and  there  was  an 
exhilaration  with  change.  Lecturing  to  the  MBL  community  in  1895  on  mind  and 
matter,  A.  E.  Dolbear  said: 

.  .  .  the  past  thirty  years  shows  that  in  the  field  of  natural  history  everybody 
has  changed  from  a  creationist  to  an  evolutionist.  But  this  has  been  only  the 
beginning  of  the  change,  for  if  that  science  was  true  that  made  such  a  change 
necessary,  it  is  also  true  that  the  same  science  will  make  needful  other  changes 
in  men's  conceptions  of  what  kind  of  a  universe  they  live  in,  how  it  works,  and 
how  they  came  to  be  and  think  as  they  do;  and  not  unlikely  that  what  we  please 
to  call  evolution  will  have  to  be  explained  and  restated  in  very  different  terms 
than  those  in  vogue  now.4 

So  spoke  Dolbear  of  Tufts  College,  and  he  could  with  confidence  dismiss  all 
antecedent  theology  and  metaphysics  as  irrelevant,  specifically  to  the  issue  of  origins. 
But  then  in  the  end  he,  too,  foundered  on  the  seeming  plain  sense  of  preformation: 
you  can't  get  something  from  nothing.  In  the  end  he  argued  that  the  universal  ether, 
at  least,  must  have  been  created.  Having  first  laid  to  rest  an  earlier  ghost,  he  raised 
a  different  one.  The  battle  continues,  as  it  has  since  then — if  not  since  Aristotle— 
with  the  forms  of  argument  changing  as  the  foci  of  scientific  theory  change.  By  the 
nineteenth  century,  serious  fighting  about  the  origins  of  complexity  had  fallen  to 
science,  and  there  it  remains.  Like  all  arguments  of  principle  and  common  sense,  it 

62 


LAYING  THE  GHOST  63 

is  fought  not  just  with  reason,  mathematics,  observation,  or  measurement:  but 
sometimes  with  feelings,  alliances,  power,  and  politics,  with  all  the  trappings  of 
society  and  culture,  which  some  contemporary  historians  insist  are  the  primary 
drives  of  scientific  inquiry,  rather  than  a  subset  of  them.  I  reject  the  insistence;  but 
that  the  trappings  are  there  is  undeniable. 

Preformationism  versus  Epigenesis  is  not  merely  a  diversion  for  embryologists; 
nor  for  cosmologists.  It  is  difficult  for  a  sober  mind  to  imagine  how  order  might 
arise  from  disorder,  long-range  structure  from  randomness.  The  discoveries  of 
nineteenth  century  physics,  especially  of  thermodynamics,  put  an  end  to  the  most 
primitive  possibility — that  organized  macroscopic  structure  may  appear  spontaneously 
in  a  structureless  or  truly  homogeneous  system.  In  the  absence  of  appropriate  work 
done  from  the  outside,  this  does  not  happen.5  But  embryologists  of  the  late 
nineteenth  century  and  early  twentieth,  although  many  were  impressed  by  the 
power  of  physics,  seem  not  to  have  been  overly  concerned  with  thermodynamics. 
And  in  modern  physics  the  issue  becomes  clouded  again,  at  least  in  the  sense  that 
a  vacuum  is  no  longer  what  we  had  thought  it  to  be:  in  the  quantum  universe 
material  particles  can  and  do  arise  spontaneously;  not  from  "nothing,"  but  because 
it  is  necessary  to  re-define  "nothing."6 


THE  MIND  IN  THE  EMBRYO 

The  search  for  an  explanation  of  development — that  most  awe-inspiring  of  all 
processes  of  the  biological  world,  in  which  a  complex  and  functional  being  emerges 
from  a  zygote  that  is  seemingly  structureless  under  the  microscope — has  included  a 
long  succession  of  creative  ghosts.  Those  have  been  the  agencies  or  blueprints  by 
which  the  form  of  the  embryo  and  adult  is  built.  They  have  changed  in  character 
from  the  ridiculous — the  homunculi  of  ovists  and  animalculists  in  the  seventeenth 
and  eighteenth  centuries — to  the  sublime — the  originally  naked  and  self-replicating 
polynucleotide,  from  which  everything  else  takes  its  guidance  and  form.  But  the 
effort  to  exorcise  the  ghosts,  material  or  immaterial,  continues  nevertheless,  and  not 
only  from  the  embryo:  from  an  anthropic  cosmos  no  less  than  from  consciousness 
(whose  ghost  is  an  entity  called  "mind"). 

Toward  the  turn  of  the  last  century,  before  the  Proud  Tower  of  Europe  began 
to  lean,7  but  when  the  strains  were  already  felt  and  influential  in  the  styles  of 
scientists,  the  ghost  in  the  embryo  assumed  a  new  and  unexpected  form.  For 
obscure  and  to  some  extent  accidental  reasons,  the  creative  ghost  became,  under 
the  influence  of  Haeckel,  the  old  Scala  Naturae  in  a  new  dress:  the  embryo  retracing 
faithfully  and  by  necessity  the  evolution  of  its  ancestors.  The  inevitable  opposition, 
when  it  arose,  did  not  so  much  refute  the  broad  claim  (although  it  was  refuted  in 
detail)  as  bypass  it  and  deem  it  irrelevant.  This  reaction  began  in  Europe,  notably 
with  Wilhelm  His,  but  the  arguments  were  heaviest  and  most  influential  among 
others,  at  the  Stazione  Zoologica;  curiously  so,  given  the  commitment  of  the 
founder,  Anton  Dohrn,  to  Haeckel's  genealogical  apparition.  By  this  time,  there 
were  able  young  Americans  who  could  commit  themselves  fully  to  fundamental 
research  in  biology.  Those  who  later  became  founders  of  the  Marine  Biological 
Laboratory  in  Woods  Hole  were  aware  of,  and  later  in  the  thick  of,  the  Naples 
arguments.  They  did  not  fail  to  carry  them  to  Woods  Hole.  The  concentration  of 
intellect  and  skills  at  the  MBL  led  to  newer  and  more  powerful  arguments;  to 
important  discoveries  of  observation  and  experiment;  and  in  the  end  to  laying  to 
rest  the  ghost,  for  at  least  the  interval  from  then  until  now. 


64  P.   R.  GROSS 

Whether  it  has  been  exorcised  for  good  I  do  not  venture  to  guess;  but  allow  me 
my  doubts.  Mind/Brain  dualism  remains  in  good  health,  even  among  respectable 
psychologists;  and  as  it  becomes  clear  to  most  biologists  that  there  are  not  enough 
genes  to  specify  all  the  connections,  one  by  one,  among  neural  elements,  let  alone 
the  emergent  operating  systems  of  the  brain,  the  ghost  may  walk  again.  This  too  is, 
after  all,  a  problem  of  biological  development.  In  any  case,  the  foundations  of 
several  branches  of  modern  biology  were  laid  in  the  course  of  early  embryological 
arguments  at  Woods  Hole,  not  the  least  of  those  branches  being  experimental 
embryology,  general  physiology  in  the  new  forms  that  led  to  what  we  now  identify 
as  "cell  biology"  and  "biophysics,"  and  modern  genetics.  It  is  a  tangled  story  which 
needs  still-unwritten  books  in  order  to  be  told  properly. 

HAECKEL'S  SYNTHESIS 

Ernst  Heinrich  Haeckel  grew  up  in  the  conservative  atmosphere  appropriate  to 
his  father's  position  as  a  higher  civil  servant  (Regierungsraf)  in  the  best  Prussian 
mold.  Haeckel  was  a  gifted  student  and  studied  under  the  best  teachers:  botany 
under  Schleiden  (Jena);  anatomy  and  physiology  at  Wiirzburg  under  Kolliker; 
pathology  under  Virchow;  and  physiology  under  J.  Miiller  in  Berlin,  where  he 
acquired  his  devotion  to  marine  research.  He  published  an  important  work  of 
comparative  morphology,  Die  Radiolarien,  after  extensive  marine  collections  at 
Messina,  and  became  Professor  of  Zoology  at  Jena  in  1862,  where  he  remained 
until  1909.  Afterward  he  continued  writing  until  his  death,  at  the  age  of  75, 
in  1919. 

It  was  a  life  full  and  productive,  and  enormously  influential.  The  influence  upon 
embryology  was  one  among  many,  but  the  embryological  one  lasted  longer  than 
did  the  others.  This  influence  owed  much  to  the  particular  ingenuity  with  which 
Haeckel  made  an  unique  amalgam  of  the  three  great  but  disparate  influences  on 
his  own  thought:  Darwin,  Goethe,  and  Lamarck.  How  this  unlikely  Trinity  came 
to  be  is  in  essence  simple,  but  the  essence  does  poor  justice  to  HaeckeFs  energy  in 
applying  it  to  a  whole  system  of  thought  about  life  on  earth,  and  in  triumphing,  by 
polemic,  over  its  many  detractors. 

Haeckel  was  for  a  long  time  a  good  boy;  conscientious,  devoted  to  his  parents 
and  their  ideals,  such  as  German  unification.  Right-thinking  came  easily  to 
him.  Thus,  Nordenskiold,  referring  to  Haeckel's  letters  written  when  he  was  away 
from  home:8 

True,  he  could  cause  his  parents  anxiety  on  account  of  his  dislike  for 
medicine  and  his  propensity  for  unpractical  dreaming,  but,  on  the  other  hand, 
he  was  always  ready,  with  a  somewhat  rhetorical  and  precocious  eloquence,  to 
confess  his  weaknesses  to  his  old  parents  and  to  promise  to  make  them  happy  in 
the  future.  The  most  striking  feature  of  these  letters  is  their  Christian  piety, 
which  contrasts  strongly  with  the  hatred  that  Haeckel  felt  for  Christianity  in 
later  years.  .  .  . 

He  was  for  a  time  an  ardent  German  nationalist,  and  a  passionate  denouncer 
of  Germany's  enemies,  within  and  without.9  But  he  was  also  an  observer.  The 
enthusiast  of  twenty-five  became  a  skeptic  at  thirty-five.  A  new,  imperial  Germany 
did  not  broaden  the  rights  of  man,  nor  loosen  the  bonds  of  religion  allied  with 
reactionary  elements  of  the  state;  quite  the  reverse.  Freedom  of  thought,  which  is 
important  to  professors,  did  not  increase  with  material  progress  and  national  power: 
it  was  decreased,  often  brutally  so.  Haeckel  made  early  the  discovery  that  noble 


LAYING   THE  GHOST  65 

thoughts,  translated  into  political  reality,  do  not  necessarily  pass  directly  into  good 
actions.  They  are  just  as  likely  to  pass  to  the  reverse. 

All  the  while,  his  professional  work,  molded  strongly  by  the  standards  of 
Wiirzburg  (where  Karl  Gegenbaur  now  held  sway),  was  devoted  to  the  good  German 
purpose  of  creating  a  Weltanschauung,  a  worldview  and  an  ordered  system,  through 
which  living  nature,  and  man  within  it,  might  be  fully  comprehended  under  a 
liberal  ideal  of  progress.  The  central  tool  for  its  construction,  as  already  established 
by  Gegenbaur  and  others,  would  be  comparative  morphology;  the  structure,  an 
objective  and  self-consistent  phylogeny,  which  would  explain  what  and  why  the 
living  world  is  as  it  is,  and  in  what  directions  it  would  go. 

Disappointed  in  his  hopes  for  reform  and  progress  in  the  spheres  of  politics  and 
general  culture,  he  fashioned  a  system  of  thought  within  his  professional  discipline 
that  would  substitute;  that  would,  at  one  and  the  same  time,  deny  the  significance 
of  princes,  politicians,  and  the  priests  who  supported  them  in  oppression,  and  hold 
out  to  mankind  a  vision  of  harmony  and  progressive  change  in  the  forms  and 
conditions  of  life.  He  found  within  his  experience  three  guides.  The  first  was 
Darwinism,  enthusiasm  for  which,  in  the  form  of  comparative  morphology,  he  had 
already  acquired  as  a  student  and  a  correspondent  with,  and  collaborator  of, 
Gegenbauer.10  It  was  the  evidence  of  common  descent  that  gave  Haeckel  the  key  to 
his  Weltanschauung.  Common  descent  immediately  defies  special  creation:  thus  it 
defied  the  establishment;  common  descent  bespeaks  flux,  change,  relatedness  among 
all  things  living,  and  by  a  very  small  (and  then  still  allowable)  step,  it  implies 
progressiveness.  The  second  guide  was  Goethe.  It  was  not  merely  that  the  poet  and 
polymath  of  Weimar  had  been  a  natural  philosopher,  the  definer  of  "morphology," 
but  that  he  had  been  a  champion  of  evolution  before  Darwin.  He  had  also,  however, 
been  an  honored  Minister  of  State  in  liberal  Weimar.  He  had  rhapsodized  change 
and  gotten  away  with  it.  And  withal,  Goethe  had  lived  to  a  ripe  old  age:  a  survivor, 
with  authority.  This  example  before  him,  Haeckel,  who  insisted  upon  the  importance 
of  a  mechanical  basis  for  his  new  system,  took  on  in  fact  Goethe's  unmechanical, 
romantic  idealism,  in  a  form  that  had  already  been  expunged  from  the  physical 
sciences  and  from  physiology." 

And  finally  Lamarck;  to  whom  Haeckel  gave  credit  for  having  understood  the 
mechanism  of  living  transformations  over  time,  whereas  Darwin  he  seems  to  have 
credited  as  more  or  less  the  demonstrator  of  their  occurrence.  Throughout  the 
whole  ran  that  strong  tendency  of  the  time  toward  viewing  history  as  the  determiner 
of  events — a  tendency  in  no  way  limited  to  biology:  a  description  of  how  things 
had  come  about  was  accepted  as  the  explanation  of  why  things  are  as  they  are. 

HaeckePs  greatest  work  was  the  Generelle  Morphologic,  published  in  1866,  with 
its  latter  and  most  important  part  modestly  dedicated  to  Darwin,  Goethe,  and 
Lamarck.  For  embryology,  however,  it  is  not  this  impressive  tome,  but  the  later 
Anthropogenic  oder  Entwicklungsgeschichte  des  Menschen  (1874)  that  was  to  be 
determining  for  decades  to  come.  This  book  had  no  less  a  purpose  than  a  complete 
description  of  the  biological  origins  of  the  human  race,  as  reconstructed  from  the 
facts  of  paleontology,  embryology,  and  comparative  anatomy. 

In  it  Haeckel  presents  fully-formed  what  came  to  be  called  the  "Biogenetic 
Law/'  i.e.,  that  "ontogeny  is  a  brief  and  rapid  recapitulation  of  phylogeny."  The 
general  idea  was  not  new:  it  had  been  proposed  earlier  by  others,  including,  notably, 
Meckel  and  Fritz  Miiller;  but  in  Haeckel's  hands  it  becomes  the  motto  and  the 
program  for  biological  research,  especially  in  embryology.  The  purpose  of  embryology 
is  to  trace  out  genealogy — to  identify,  by  means  of  close  observation  of  embryonic 
form,  the  details  of  phylogenetic  relationships  among  animals,  the  history  of  their 


66  P.  R.  GROSS 

descent.  The  history  will  be  the  explanation;  for  the  fact  of  organic  evolution  is  not 
in  doubt,  and  its  mechanisms  seem  close  to  being  known.  For  this  program  of 
embryological  research  there  was  ready  to  hand  a  most  fortunate  tool:  the  morpho- 
genesis of  germ  layers,  whose  universal  homology  seemed  to  Haeckel  perfectly 
obvious. 

The  facts  of  embryonic  development,  in  short,  he  saw  as  the  strongest  support 
of  common  descent,  and  at  the  same  time  the  still  unexplored  territory  by  which 
its  details — hence  the  detailed  history  of  life — are  to  be  wrested  from  nature.  The 
germ  layers  arise  at  gastrulation:  hence  the  development  of  form  after  that  stage  of 
embryogenesis  is  to  be  studied.  From  such  studies  are  to  be  reconstructed  the 
genealogies  that  will  establish  not  only  the  tree  of  life,  but  also  the  sap  flowing  in 
it.  For  twenty-five  or  thirty  years  this  program  was  the  preoccupation  of  embryology, 
which  was  in  turn  perceived  as  the  strong  right  arm  of  general  morphology,  and,  of 
course,  of  Darwinism.  Haeckel,  exorcising  the  ghost  of  special  creation  from 
embryonic  development,  substituted  another  one:  phylogeny  as  the  directing  intel- 
ligence, phylogeny  not  merely  the  result,  but  the  mechanical  cause  of  development. 
(Why  bother,  then,  with  physiology,  or  with  cleavage,  which  was  a  mere  subdivision 
of  the  bulky  egg  into  better-deployable,  small  units?12) 

Reaction  was  not  slow  in  developing,  although  thirty  years  were  required  for  it 
to  affect  the  work  of  most  biologists.  Haeckel  gathered  round  himself  a  group  of 
enthusiastic  followers,  many  of  whom  associated  themselves  willingly  with  a  political 
radicalism  they  conceived  as  matching  their  scientific  stance.  But  not  expectedly, 
for  every  accomplished  scientist  who  espoused  the  radical  program  entire,  there  was 
an  equally  distinguished  one  who,  for  reasons  of  his  own,  opposed  it.  The  first 
attacks  upon  Haeckel  had  other  than  purely  scientific  motives.  For  example: 
Haeckel's  system  included  (naturally)  a  program  for  educational  reform:  this  was 
attacked  by  no  less  a  figure  than  Haeckel's  former  teacher,  Rudolf  Virchow,  who  in 
so  doing  placed  himself  in  the  uncharacteristic  position  of  arguing  that  the  teaching 
of  hypotheses  should  be  banned. 

These  arguments,  emotional  as  they  were,  attracted  less  attention  than  desired 
by  their  proponents.  Militarists  and  socialists  had  other  fish  to  fry;  and  the  arguments 
of  zoologists  were  in  any  case  incomprehensible  to  most  policy-makers,  let  alone  to 
the  masses.  Haeckel's  program  for  embryological  and  morphological  research 
survived  such  arguments  by  many  years,  as  it  did  the  other  philosophical  quarrels 
that  marked  the  death-throes  of  natural  philosophy.  The  destruction  of  Haeckelian 
embryology  came  from  another  quarter  entirely:  that  of  cytology,  with  its  remarkable 
discoveries  of  cell  structure,  growth,  and  division  (Strasburger's  monumental  Zell- 
bildimg  und  Zelltheilung  was  published  in  1874  and  became  truly  influential  a 
decade  later);  of  fertilization  and  its  early  consequences  (Fol,  1879);  and  from  the 
new  experimental,  i.e.,  manipulative  and  invasive,  embryology,  championed  by 
Wilhelm  His,  by  Roux,  Boveri,  the  Hertwigs,  and  in  the  end  by  a  whole  school  of 
Naples  investigators  whose  most  aggressive — and  abrasive — voice  was  that  of  Hans 
Driesch. 

The  reaction  at  Naples  is  critical  to  the  unfolding  story  of  Haeckel's  overthrow 
and  the  rise  of  modern  embryology;  but  I  shall  not  address  it  here.  It  is  the  subject 
of  a  chapter  elsewhere  in  this  volume.  Suffice  it  to  say  that  the  Americans  were 
aware  of  it  and  involved  in  it,  and  that  without  such  involvement  the  MBL  would 
not  have  come  to  be  what  it  is.  American  biology  would  have  been  something  less. 
The  MBL  story,  insofar  as  it  can  be  represented  here,  must  begin  not  with  outspoken 
anti-phylogenesis — for  that  came  later — but  with  the  transitional  work  of  its 
remarkable  founding  Director,  Charles  Otis  Whitman. 


LAYING  THE  GHOST  67 

WHITMAN  SPURS,  AND  CONTROLS,  THE  REVOLUTION  IN  AMERICA 

If  Haeckel  caused  a  revolution  in  embryology,  then  the  mechanist-experimental 
wave  was  its  counter-revolution.  Whitman  helped,  despite  himself,  to  bring  it  about, 
by  demonstrating  the  power  and  value  of  cell  lineage  research;  and  yet  he  was  able 
to  control  the  inevitable  after-shocks  (which  included  a  rejection  even  of  cell  lineage) 
by  the  power  of  persuasion,  by  the  breadth  of  his  mind,  and  by  his  skill  as  a 
scientific  administrator.  It  is  true  that  the  name  "cell  lineage"  was  applied  first  by 
E.  B.  Wilson  to  "the  study  of  the  cell-by-cell  origin  of  body  regions  and  organs 
characteristic  of  the  development  of  annelids,  most  mollusks,  many  Crustacea,  the 
tunicates.  and  a  few  other  animals,  in  which  the  cleavage  of  the  egg  is  determinate."13 
But  Whitman  must  be  counted  among  the  actual  inventors  of  the  methodology,  the 
first  to  demonstrate  its  power. 

Writing  in  The  Proud  Tower  of  Thomas  B.  Reed,  Republican  of  Maine  and 
elected  speaker  of  the  U.  S.  House  of  Representatives  in  1890,  Barbara  Tuchman 
uses  words  that  might  apply  equally  well  to  the  contemporary  Whitman: 

...  in  character,  intellect  and  a  kind  of  brutal  independence  .  .  .  (he)  .  .  . 
represented  the  best  that  America  could  put  into  politics  in  his  time.  He  was 
sprung  from  a  rib  of  that  hard  northern  corner  of  New  England  with  the 
uncompromising  monosyllabic  name.  .  .  .  The  sons  of  Portland  families  went 
to  Bowdoin,  not  to  satisfy  social  custom,  but  to  gain  a  serious  education. 

So  too  Maine-born  Whitman,  who  studied  the  classics  at  Bowdoin,  displayed 
throughout  his  life  a  certain  brutal  independence,  supported  by  an  agile  intellect 
and  an  unwillingness  to  submit  to  authority  unless  its  sources  had  survived  his  own 
examination.  The  United  States  had  neither  a  landed  aristocracy  nor  an  intellectual 
one,  except  for  the  small  cluster  of  thinkers  who  survived  in  the  vicinity  of  Boston. 
Those  did  not  exert  much  influence  upon  the  culture  outside  a  few  universities. 
The  nation  was  a  German  liberal's  dream  come  true:  abdication  of  the  rich  from 
politics;  government  loose,  flexible,  manipulable — and  self-serving;  growth  in  wealth 
and  power  but  without  the  restrictions  of  landed  inheritance  and  hereditary  privilege; 
the  poor  under  control.  A  capitalist's  heaven,  and  one  in  which  as  much  honor  was 
given  to  financial  achievers  (even  if  achievement  was  by  dubious  means)  as  was 
denied  to  politicians. 

To  be  sure,  not  all  was  peace  and  tranquility.  Healing  the  Civil  War's  wounds 
was  the  healing  of  a  large  adolescent,  growing  like  a  weed,  feeling  the  desires  of 
adolescence.  The  inevitable  conflict  erupted  in  the  Spanish-American  war  of  1898- 
a  conflict  between  those  hungry  to  use  their  strength  to  seize  external  riches,  and 
those  who  saw  in  the  emulation  of  European  imperialism  the  death  of  the  American 
social  ideal.  Leading  the  former  were  those  two  "degenerate  sons  of  Harvard"  (as 
Harvard's  President  Eliot  named  them),  Theodore  Roosevelt  and  Senator  Lodge, 
supported  by  the  popular  press;  and  leading  conscience  were  a  few  of  the  Bostonians 
(including  the  same  Eliot),  a  few  journalists,  and  some  honest  politicans  (including 
Reed).  Conscience  lost  and  Jingoism  won.  But  the  Caribbean,  and  more  so  the 
Pacific  islands,  were  sufficiently  remote  from  the  great  land  between  two  oceans  so 
that  they  had  small  effect.  It  was  an  expanding  and  confident  society,  unencumbered 
by  ancient  power-structure  and  innocent,  after  Appomattox,  of  war  on  its  own  soil. 
Able  men  of  conscience  who  did  politics  did  it,  and  were  lonely.  Able  men  of 
science  or  the  arts  did  those;  and  they  did  not  bother  much  about  politics. 

Like  Thomas  Reed,  Whitman  became  a  school-teacher  upon  graduation  from 
Bowdoin:  at  age  twenty-six  he  became  principal  of  the  Westford  Academy  in 


68  P.   R.  GROSS 

Westford,  Massachusetts;  and  in  1872,  at  the  age  of  thirty,  he  joined  the  faculty  of 
the  English  High  School  in  Boston.  It  was  here  that  his  interest  turned  to  natural 
history.  He  attended  the  two  sessions  of  Louis  Agassiz's  Anderson  School  on 
Penikese  Island  in  1873  and  1874.  In  1875,  having  decided  to  stake  his  future  upon 
a  career  in  science,  he  did  what  it  was  necessary  to  do  in  order  to  obtain  the  best 
training:  he  sailed  for  Germany  with  the  intention  of  studying  natural  history  and 
earning  a  doctorate  there  (in  the  laboratory  of  Rudolf  Leuckart,  at  Leipzig).  In 
1878,  a  man  no  longer  young  to  be  starting  in  science,  he  received  the  doctorate 
and  published  a  dissertation  on  the  embryology  of  leeches  of  the  genus  Clepsine.14 

Just  as  Whitman  himself  emerged  as  an  unique  transitional  figure  in  scientific 
thought  and  administration  in  America — aware  of  all  that  was  new,  and  receptive 
to  it,  but  unwilling  to  sacrifice  the  values  of  the  old  simply  for  the  sake  of  novelty— 
so,  too,  is  this  hundred-page  memoir  of  Whitman's  transitional.  Its  length  was  not 
unusual  at  the  time,  nor  for  some  time  longer:  witness  the  extraordinary  length  and 
detail  of  monographic  papers  published  by  such  followers  of  Whitman  as  Wilson 
and  Conklin.  Nor  is  there  any  direct  or  obvious  rejection  of  the  genealogical 
concerns  that  so  characterized  contemporary  works  of  embryology  or  comparative 
morphology.  It  is  simply  that  these  concerns  are,  in  Whitman's  work,  mere 
obeisances,  by  comparison  with  the  extraordinary  detail  and  attention  devoted  to 
the  three  central  sections.  These  deal,  in  order,  with  the  cytology  of  maturation  and 
of  the  uncleaved  zygote — employing  every  sophistication  of  method  recently  intro- 
duced by  the  cytologists  ("die  aniline-barber,"  of  whom  other  embryologists  were 
scornful);  with  an  exact  description  of  cleavage  itself  and  of  the  fates  of  blastomeres; 
and  finally  with  the  morphogenetic  movements  of  gastrulation  and  neurulation, 
which — despite  Whitman's  delicacy  in  not  making  a  great  point  of  it — threw  doubt 
upon  the  prevailing  notions  of  germ  layer  homology. 

This  paper  was  not  simply  a  doctoral  student's  labor  in  animal  morphology, 
although  that  is  how,  under  Leuckart's  guidance,  the  work  had  begun.  It  was  a 
canonical  work  of  morphology,  but  it  was  also  a  beginning  of  cell  lineage  research, 
which  was  to  become  the  first  of  those  subdisciplines  to  be  uniquely  associated  with 
Woods  Hole.  Close  observation  of  the  earliest  processes  of  embryogenesis,  conducted 
with  every  technical  advantage  to  be  gained  from  contemporary  advances  in 
microscopy,  largely  free  of  preoccupation  with  phylogeny  and  the  intractable  issues 
of  origins,  produced  evidence  of  the  superficiality  of  Haeckelian  embryology  and  a 
new  means  for  the  analysis  of  emerging  form  in  the  embryo.  No  matter  that  this 
disposal  of  the  phylogenetic  ghost  would  create  a  substitute  one:  heredity  borne  by 
substances  or  particles  of  the  egg.  The  fight  about  that  would  come  later. 

Whitman  returned  in  due  course  to  the  United  States,  as  independent  a 
personality  as  he  had  left  it;  but  he  returned  with  scientific  skills  and  habits  of 
reasoning  that  incorporated  most  of  the  good,  and  little  of  the  bad,  of  German 
science.  He  had  begun  early,  and  continued  later,  exorcism  of  the  wraith  that 
Haeckel  had  imposed  upon  embryology.  But  in  so  doing  he  installed  what  others 
in  Woods  Hole  were  to  see  as  equally  irrelevant  for  an  explanation:  the  notion  of 
"promorphology,"  or  "predetermination"  of  the  egg,  in  consequence  of  which  the 
patterns  of  maturation  and  cleavage  were  irrevocably  fixed.  Thus  fixed,  the  behavior 
of  blastomeres  would  inevitably  lead  to  a  reinforcement  of  the  original  polarity;  to 
the  acquisition  of  symmetries  more  complex  than  spherical  or  radial,  and  to  the 
programmed  emergence  of  adult  form.  But  the  question  could  then  rightly  be  asked, 
as  it  was:  "Whence  does  the  'promorphology'  arise;  and  why  do  we  need  it  at  all?" 

Whitman  would  reply  with  cogency  and  reason,  although  not  sufficient  to 
convince  some  of  the  younger  and  newly  self-confident  investigators  who  came  to 


LAYING  THE  GHOST  69 

Woods  Hole.  But  his  replies  were  for  all  that  cogent  and  in  advance  of  their  time; 
the  former  Bowdoin  scholar,  like  the  rapier-witted  Thomas  Reed  in  the  Congress, 
gave  as  good  as,  or  better  than,  he  got.  In  the  optimistic,  civilized,  relatively 
unpolitical  atmosphere  of  the  MBL  (as  in  the  two  university  departments — Clark 
and  Chicago — that  Whitman  ran  so  autocratically),  the  Director  could  and  did 
encourage  alternative  views.  He  could  and  did  enlarge  the  scope  of  teaching  and 
research,  even  beyond  his  own  abilities,  as  changes  in  biology  made  such  enlargement 
necessary;  and  he  modified  his  own  views,  but  without  retreating  from  the  first 
position  that  the  hereditary  constitution  of  the  zygote  is  the  primary  source  of 
information  for  its  early  development,  and  hence  for  the  morphology  finally 
achieved. 

STEMMING  THE  TIDE 

Few  were  happy  with  the  new  ghost:  heredity,  or  species-specific  determinants, 
residing  in  the  egg  and  directing  its  development,  replacing  a  still  vaguer  sort  of 
phylogenetic  director.  In  Europe,  the  entwicklungsmechanik  of  Roux  moved  irre- 
sistibly toward  physical  and  chemical  interference  with  development  as  the  investi- 
gative device,  and  hence  toward  the  explanation  of  developmental  events  in  physical 
and  chemical  terms.  In  America,  especially  at  Woods  Hole,  there  were  at  first 
murmurs  and  later,  when  T.  H.  Morgan  joined  forces  with  the  German-American 
Jacques  Loeb,  well-spoken  criticisms.  They  identified  Whitman's  hereditary  deter- 
miners— and  soon  those  implied  by  the  work  of  Wilson,  Conklin,  Lillie,  and 
others — as  a  new  category  of  homunculi.  To  explain  development  as  due  to  the 
action  of  "plasms"  or  "idiophores"  in  the  protoplasm  was  for  these  critics  no  better 
than  to  explain  it  as  the  outcome  of  an  "idea."  For  them,  it  was  no  explanation 
at  all. 

Whitman  fought  back,  while  at  the  same  time  encouraging  the  work  of  his 
opponents.  He  was  no  compulsive  speaker  and  publisher,  but  his  lectures  and 
papers  were  strong,  and  his  pen  could,  as  Conklin  remarked,  be  dipped  in  gall.15 
Thus,  in  1894,  six  years  after  the  founding  of  the  MBL,  he  delivered  a  lecture  upon 
"Evolution  and  Epigenesis,"  which  was  devoted  to  a  scholarly  attack  on  the  gathering 
opposition. 

.  .  .  The  possibility — not  to  say  probability — that  the  egg  is  from  the 
beginning  of  its  existence  as  an  individual  cell  definitely  oriented,  has  received 
but  little  attention.  Many  difficult  questions  are  involved  which  can  only  be 
settled  after  the  most  exhaustive  analysis  of  its  structure  and  the  most  careful 
examination  of  its  entire  history.  It  is  not  enough  to  catch  a  fact  here  and  there, 
in  this  or  that  species;  the  whole  series  of  phenomena  must  be  studied  genetically, 
and  in  as  many  forms  as  possible.  It  often  happens  that  we  have  to  snatch  facts 
as  opportunity  brings  them  within  reach,  regardless  perhaps  of  their  connections; 
but  so  long  as  they  stand  isolated,  they  are  unsafe  pegs  to  hang  theories  upon. 
Examples  abound  on  this  one  question  of  the  orientation  of  the  egg,  and  the 
mention  of  'isotropisirf  will  recall  more  than  one  windfall  of  premature  specu- 
lations."16 

The  windfall  Whitman  referred  to  was  the  growing  body  of  evidence,  derived 
especially  from  experiments  on  regulative  eggs  such  as  that  of  the  sea  urchin,  which 
led  the  investigators  to  propose  that  the  egg  must  be  isotropic — i.e.,  regionally 
homogeneous:  and  therefore  not  predetermined.  That  style  of  research  to  which 
Whitman  referred  with  favor,  i.e.,  the  "most  exhaustive  analysis  of  structure,"  was 
of  course  the  style  he  had  himself  employed  and  to  some  extent  pioneered  in 
embryology.  For  its  execution  there  came  to  the  MBL  a  group  of  exceptional 


70  P.   R.  GROSS 

scientific  talents,  some  of  them  Whitman's  own  students,  others  independent.  So 
"exhaustive"  in  fact  were  their  productions  that  they  stemmed  the  tide  of  experi- 
mentalism  for  a  time,  and — much  more  importantly — because  the  investigators 
were  young  enough  to  incorporate  what  was  useful  in  the  new  experimental 
embryology,  they  became  in  due  course  experimenters  themselves.  At  first  the  work 
was  unabashed  observation,  but  later  most  of  them  became  observer-experimenters, 
and  in  the  end  it  was  one  of  them — E.  B.  Wilson — who  provided  the  commanding 
synthesis  of  observational  and  experimental  embryology.  It  was  he  who  brought  the 
revolution  against  nineteenth  century  natural  philosophy,  and  the  old  spirits  and 
specters,  to  a  triumphant  conclusion.17 

Wilson,  E.  G.  Conklin,  and  F.  R.  Lillie  were  the  leaders  of  this  school — at  first 
observers  and  tracers  of  cell  lineage,  later  experimenters,  finally  synthesizers.  I  may 
not  in  this  essay  examine,  as  it  has  been  necessary  to  some  extent  to  do  for  Haeckel 
and  Whitman,  the  personalities  and  backgrounds  of  these  men,  however  individual 
and  fascinatingly  different  they  were.  To  do  so  here  would  soften  the  focus  of  the 
argument — for  it  did,  indeed,  gather  focus  toward  the  turn  of  the  century.  Suffice  it 
to  say  that  these  were  Americans,  who  shared  the  personal  styles  of  their  American 
contemporaries  more  than  they  did  the  styles  of  the  Europeans  who  came  to  Naples. 

Thus  Wilson,  an  accomplished  cellist  (whose  close  friendship  with  Anton  Dohrn 
began  with  their  mutual  delight  in  a  Schumann  string  quartet)18  and  a  profound 
scholar,  seems  not  to  have  been  caught  up  in  the  politics  of  culture  in  the  way  the 
Europeans  were.  Much  less,  even,  were  the  other  two.  Pragmatists,  deeply  committed 
to  their  work  and  to  the  immediate  concerns  of  academic  life,  they  and  their 
contemporaries  created  in  the  short  course  of  ten  years  a  practice  of  embryology 
which  was  not  merely  American,  but  was  in  fact  the  vanguard  of  embryology. 

Wilson  was  the  first  to  carry  out  a  complete  cell  lineage  study  at  Woods  Hole: 
this  was  his  splendid  work  of  development  of  the  annelid.  Nereis. l9  Conklin's 
descriptive  work  on  Crepidula  was  perhaps  even  more  elegant.20  Though  it  did  not 
excite  the  admiration  of  his  doctoral  supervisor,  W.  K.  Brooks,  it  ranks  still  among 
the  defining  examples  of  cell  lineage  research.  Lillie,  under  Whitman's  direction, 
carried  out  a  similar  study  on  the  lamellibranch,  Unio.21  Lillie  records  Wilson's 
remark  to  him  in  1891:  "I  believe  I  am  going  to  destroy  the  germ  layer  theory  of 
development!"22 

And  so  he,  and  the  others,  did,  in  the  following  sense:  that,  contrary  to  the 
convictions  of  two  decades  before,  to  the  effect  that  the  process  of  cleavage  was 
indifferent,  a  mere  subdivision  of  the  egg  preparatory  to  germ  layer  formation,  and 
the  germ  layers  thereafter  strictly  homologous  species  to  species;  the  cleavages  were 
in  fact  (at  least  in  the  forms  they  studied),  determined  as  to  plane,  cell  size,  and 
timing.  They  showed  that  the  germ  layers  arise  by  the  early  segregation  of  regionally 
different  cytoplasms  in  consequence  of  the  cleavages;  and  that  the  resultant  germ- 
layers  are  not  homologous  in  the  way  that  was  earlier  believed. 

A  decade  later,  Wilson  and  Conklin  had  become  experimenters  as  well  as 
observers.  Wilson  published  an  impressive  synthesis  of  the  facts  and  arguments 
derived  from  both  kinds  of  research  in  two  monograph-length  papers,  on  Dentalium 
and  Patella2"1  Conklin's  comprehensive  study  of  "Organization  and  cell  lineage  of 
the  ascidian  egg"  was  published,  with  its  many  exquisitely  drawn  plates,  by  the 
Philadelphia  Academy  of  Natural  Sciences.24  And  by  that  time  they  had  both  taken 
full  account  of  the  issues — such  as  capacity  for  regulation  after  injury  or  blastomere 
displacement — raised  by  the  experimentalists.  The  cell  lineage  chapter  became  the 
initiating  chapter  of  all  modern  treatments  of  embryonic  development. 


LAYING  THE  GHOST  71 

But  the  later  synthesis  takes  me  ahead  of  my  particular  story.  In  the  early 
1890's,  cell  lineage  was  observation;  the  opposition  was  devoted,  on  the  other  hand, 
to  experimental  interference  with  the  early  embryo,  under  the  rationales  advanced 
originally  by  Roux  and  later  by  Hertwig,  Boveri,  and  Driesch.  They  had  two 
complaints:  first,  that  observation  alone,  however  painstaking,  could  never  lead  to 
explanation,  since  explanation  should  account  for  biological  events  in  terms  of 
chemistry  and  physics,  not  through  the  naming  of  unknown  and  unknowable 
"plasms."  Second,  that  their  own  experiments  showed,  at  least  in  the  eggs  they 
studied  (which  were,  of  course,  rarely  the  same  ones  as  the  lineagists  studied),  that 
ordinary  processes  of  early  cleavage  may  to  a  very  large  extent  be  disrupted  without 
loss  of  the  embryo's  capacity  to  form  a  normal-appearing  larva.  The  campaign  to 
exorcise  the  unknowable  from  a  directive  role  in  embryogenesis  was  in  full  career. 
Epigenesis  was  to  be  established  upon  a  firm  basis  in  the  physical  sciences,  as  an 
objective  account  of  the  egg's  responses  to  environmental  stimuli.  That  this  thrust 
caused  Morgan — who  would  later  lead  the  emergence  of  a  new  science  of  genetics— 
to  reject  the  idea  of  hereditary  determiners  of  development,  does  not  matter:  Mendel 
was  not  yet  rediscovered,  nor  the  virtues  of  Drosophila  melanogaster. 

EPIGENETICS  UNCOMPROMISING:  ISOTROPISM 

Jacques  Loeb  was  a  scientific  materialist,  a  cosmopolitan  European  educated 
amid  the  cultural  tensions  that  formed  the  various  camps  of  European  science. 
Unlike  many  of  his  contemporaries  in  biology,  he  was  solidly  trained,  not  only  in 
mechanistic  physiology  (including,  of  course,  the  physics  and  chemistry  of  the  time), 
but  also  in  philosophy.  Commenting  upon  Loeb's  background,  Jeffrey  Werdinger 
says:25  "Like  his  teachers,  his  goal  was  to  analyze  and  explain  organic  phenomena 
on  the  basis  of  physico-chemical  laws  and  principles.  Indeed,  he  came  to  regard 
classical  nineteenth  century  physics  as  the  very  model  of  science  properly  understood." 

Arrived  at  the  Stazione  Zoologica  in  the  winter  of  1889-90,  he  began  a  lifelong 
course  of  research  on  regeneration,  tropisms,  and  animal  behavior,  seeking  to 
explain  them  all  under  a  simple,  but  comprehensive  set  of  physical  principles.  At 
Naples  he  met  several  of  the  young  Americans  who  had  begun  to  visit  regularly, 
and  with  their  urging  and  assistance,  emigrated  to  America,  where  he  obtained  a 
teaching  post  at  Bryn  Mawr  College.  Another  young  biologist  was  recruited  to  the 
faculty  that  year  (1891):  Thomas  Hunt  Morgan.  Whitman,  ever  on  the  alert  for 
new  ideas  backed  up  by  solid  work,  was  much  taken  with  Loeb,  and  brought  him 
eventually  to  the  department  at  Chicago,  where  Whitman  had  gone  as  Chairman. 
Loeb  came  with  Whitman  to  Woods  Hole,  there  to  found  the  course  in  General 
Physiology  that  has  continued,  with  high  distinction,  to  be  taught  ever  since. 

Except  in  these  matters  of  origin  and  prior  training,  Loeb  was  not  different  from 
the  other  young  men  of  his  generation  at  the  MBL.  He  was,  perhaps,  more 
passionate  about  the  broad  implications  of  arguments  that  were  in  progress:  he  had 
brought  with  him,  along  with  the  attitudes  of  the  new  physical  biology,  much 
positivist,  social-progressive  freight;  and  he  applied  it  in  his  activities,  his  writings, 
and  his  lectures,  exerting  considerable  influence  in  American  scientific  circles  (but 
not,  perhaps,  as  much  as  he  might  have  done  in  Germany).  A  greater  influence 
came,  however,  from  the  brash  simplicity  of  his  experiments,  which  were  at  the 
same  time  highly  reproducible.  Loeb  saw  no  reason  why  explanations  of  development 
should  assign  cause  to  unknown,  inherited  factors  of  the  cytoplasm  or  nucleus,  or 
even  worse,  deny  altogether  the  value  of  investigating  cause.  For  him,  the  program 


P.   R.  GROSS 

of  embryology,  indeed  of  all  biology,  was  to  discover  causes — which  were  certain  to 
he  ordinary,  immediate,  physical  and  chemical  processes.  "Explanation"  would 
consist  in  the  identification  of  such  processes. 

On  that  basis,  his  criticisms  of  observational  cell  lineage  research  can  be 
understood.  His  experimental  accomplishments  with  embryos  were  regular  and 
abundant.  It  is  true  that  the  first  popular  evidence  of  non-determinacy  in  cleavage 
(popular  in  the  sense  that  it  was  widely  discussed)  was  not  Loeb's  but  that  of  Hans 
Driesch,  working  at  Naples.26  It  is  true  that  the  capacity  of  some  eggs  to  "regulate" 
development,  i.e.,  to  compensate  for  loss  or  damage  to  blastomeres  in  early  cleavage, 
was  to  some  extent  accepted  even  by  the  descriptive  morphologists,  although  they 
contrived  to  believe  for  a  while  in  its  irrelevance.  But  Loeb,  in  America,  went  much 
further  than  Driesch.  His  was  a  most  radical  epigenesis,  backed  by  simple  experiments 
whose  results  were  very  difficult  to  explain  if  one  believed,  with  Whitman,  that  the 
egg  is  always  "predetermined,  if  not  predelineated." 

Loeb's  later  work  on  artificial  parthenogenesis,  and  its  implications  for  the 
argument  here  described,  is  too  well-known  to  describe  again.  A  single  additional 
example  of  his  work  and  the  critique  implicit  in  it  must  suffice,  and  it  is  best  that 
it  be  from  the  early  1 890's,  the  period  of  our  interest.  In  the  Journal  of  Morphology, 
that  year,  Loeb  published  a  paper  as  remarkable  for  its  brevity,  and  the  boldness  of 
its  conclusions,  as  were  the  papers  of  the  other  side  for  their  length  and  complexity. 
The  third  in  a  series  with  the  general  title,  "Investigations  in  physiological  mor- 
phology," this  one  was  subtitled,  simply,  "Experiments  on  cleavage."27 

The  experiments  described  are  a  series  of  manipulations  of  the  concentration 
and  composition  of  salts  in  the  sea  water  in  which  sea  urchin  (Arbacia)  eggs  were 
fertilized  and  allowed  to  develop.  With  careful  observation,  the  consequences  of 
these  treatments  for  karyokinesis,  cytokinesis,  and  the  pattern  of  cleavage  were 
recorded.  Loeb  made  the  discovery  that  nuclear  division  could  take  place  without 
cleavage  of  the  egg  cytoplasm.  When  such  undivided,  multinucleate  embryos  were 
replaced  in  normal  sea  water,  they  cleaved,  not  by  the  normal  stepwise  process,  but 
all  at  once  into  many  cells  (as  many,  Loeb  thought,  as  there  were  nuclei).  These 
experiments  were  very  carefully  controlled  against  the  possibility  of  polyspermy, 
which  was  known  to  cause  multipolar  division. 

Loeb  argued: 

.  .  .  the  segmentation  of  the  nucleus  proceeds,  although  more  slowly  than 
under  normal  conditions,  whilst  no  segmentation  of  the  protoplasm  is  possible. 
The  fact  in  itself  is  of  some  technical  value,  as  it  enables  us  to  separate  two 
processes  which  nature  generally  produces  together.  ...  In  regard  to  our 
knowledge  of  segmentation,  we  see  from  this  that  the  physiological  conditions 
for  segmentation  of  the  nucleus  are  very  different  from  the  physiological 
conditions  of  the  segmentation  of  the  protoplasm.  .  .  .  But  these  experiments 
allow  us  to  go  one  step  farther  and  make  clear  one  element  in  the  complex  called 
segmentation,  namely  the  physiological  cause  for  the  segmentation  of  the 
protoplasm. 

There  follows  a  short  digression  on  the  water  content  of  cells,  the  effects  of 
different  ions,  and  "irritability;"  then  by  a  chain  of  argument  appropriate  to  the 
state  of  knowledge  of  the  time,  the  conclusion:  ".  .  .  The  segmentation  of  the 
protoplasm  is  the  effect  of  a  stimulus  which  the  nucleus  applies  to  the  protoplasm, 
and  which  makes  the  protoplasm  close  around  the  nucleus." 

The  implication  is  not  merely  that  a  simple  nuclear  signal  is  the  cause  of 
cleavage,  but  that  the  form  of  the  resulting  blastomere  is  a  direct  result  of  it.  It  will 
be  evident  that  such  experiments  and  conclusions  were  very  remote  from  the 


LAYING  THE  GHOST  73 

observations  and  conclusions  of  cell  lineage  research.  They  were,  in  some  degree, 
offensive  to  that  group  of  investigators — although  never,  apparently,  taken  personally. 
It  is  impossible  to  do  justice  by  any  summary  to  the  gadfly-quality  of  Loeb's  results 
in  this  period.  The  best  that  can  be  done  is  to  say  that,  save  for  a  few  cases  of  over- 
interpretation,  his  productions  did  show  that  there  is  little  fixity  ( 1 )  of  cleavage 
pattern,  or  (2)  of  cell  size  required  during  very  early  development  for  the  eventual 
production  of  a  normal  larva  of  the  sea  urchin;  moreover,  that  such  rules  of  cleavage 
as  could  be  discerned  implied  a  great  capacity  of  the  embryo  for  repair  and 
"regulation,"  provided  that  the  physical  and  chemical  environment  were  appropriate. 
The  emerging  position,  from  this  body  of  work  and  the  work  of  others — Morgan 
and  Ross  Harrison  in  America,  Theordor  Boveri,  Oskar  Hertwig,  Hans  Driesch  in 
Europe,  was  precisely  the  one  that  a  physicist,  comfortable  with  simplifying 
assumptions  and  absolutely  committed  to  causal  analysis,  would  take:  that  the  egg 
(note  "the  egg,"  i.e.,  every  egg)  is  isotropic;  regionally  homogeneous  and  undiffer- 
entiated;  that  it  is  through  a  series  of  physical  stimuli  and  responses,  the  normal 
environment  acting  on  a  tabula  rasa,  that  all  the  events  of  development  are  initiated 
and  controlled.  This  was  truly  a  Radical  Epigenetics,  originated  by  Driesch's  "egg- 
shaking." 

In  the  end,  that  original  egg-shaker,  faced  with  the  inconsistencies  of  his  own 
radical  epigenetics  alongside  its  truths,  and  with  the  truth  of  determinate  cleavage 
in  other  eggs  than  the  sea  urchin,  along  with  its  uncertainties,  would  opt  out  of  it 
and  embrace  a  vitalism  more  reminiscent  of  Plato  than  of  Schelling  and  Goethe. 
But  Loeb  did  not.  One  can  find  in  Loeb's  lecture  of  1912,  delivered  to  the  First 
International  Congress  of  Monists  at  Hamburg,  that  same  dedication  to  a  purely 
mechanistic  interpretation  of  life — but  with  better  experimental  justification — that 
is  evident  in  his  1894  lecture  at  the  MBL  "On  the  limits  of  divisibility  of  living 
matter,"28  which  was  an  open  challenge  to  the  claim  of  an  "organized,"  anisotropic 
egg  cytoplasm.  The  difference  between  the  two  lectures  is  that  for  the  Monists,29 
Loeb  felt  free  to  express  his  lifelong  ethical  convictions,  and  could  add  some  of  the 
new  knowledge  of  mutations  to  them: 

Economic,  social,  and  political  conditions  of  ignorance  and  superstition  may 
warp  and  inhibit  the  inherited  instincts  and  thus  create  a  civilization  with  a 
faulty  or  low  development  of  ethics.  Individual  mutants  may  arise  in  which  one 
or  the  other  desirable  instinct  is  lost,  just  as  individual  mutants  without  pigment 
may  arise  in  animals;  and  the  offspring  of  such  mutants  may,  if  numerous 
enough,  lower  the  ethical  status  of  a  community.  Not  only  is  the  mechanistic 
conception  of  life  compatible  with  ethics;  it  seems  the  only  conception  of  life 
which  can  lead  to  an  understanding  of  the  source  of  ethics.30 

Nor  did  Thomas  Hunt  Morgan  ever  abandon  physical  mechanism  and  epigenetics. 
Indeed,  his  insistent  rejection  of  predetermination,  and  of  undefined  hereditary 
elements  directing  the  course  of  development,  was  surely  one  reason  for  his 
continued  indifference  to  genes  in  development,  after  the  rediscovery  of  Mendel's 
work;  even  for  a  time  after  the  chromosome  theory  of  heredity  had  blossomed  so 
suddenly  at  Woods  Hole.  His  entry  into  genetics,  and  his  transformation  of  it  at 
Columbia  and  in  the  summers  at  Woods  Hole,  had  to  await  a  simple  and  explicit 
experimental  device  by  which  he  could  analyse  genes  and  chromosomes  at  the  same 
time:  the  breeding  of  fruit-flies. 

Earlier,  Morgan  did  the  same  sort  of  embryological  work  as  Loeb,  but  he 
brought  to  it  stronger  morphological  and  observational  equipment  (having  done  his 
doctoral  research  with  Brooks  at  Johns  Hopkins).  In  close  contact  with  Driesch  as 


P.   R.  GROSS 

well  as  with  Loeb,  he  complemented  the  work  of  those  two  with  egg-shaking 
experiments  of  an  unprecedented  precision  and  thoroughness. 

Morgan's  progress,  first  as  critic  of  predetermination  and  cell  lineage,  later  as 
synthesizer  of  data  from  both  sides,  is  well  illustrated  by  experiments  he  conducted 
over  a  period  of  fifteen  years  at  Naples  and  Woods  Hole,  employing  two  different 
egg-shaking  techniques.  The  first  was  egg-shaking  pure  and  simple,  a  la  Driesch, 
but  enhanced  by  a  refinement  of  existing  methods  for  obtaining  egg  fragments  and 
isolated  blastomeres  from  early  embryos.  (He  shook  the  eggs  with  tiny  shards  of 
glass!)  The  second,  following  Boveri  but  again  with  innovations  of  technique  (e.g., 
in  the  use  of  the  centrifuge),  was  analysis  of  development  in  embryos  in  which  there 
were  forced  alterations  of  the  cleavage  planes.  These  analyses  were  carried  out  with 
expert  cytologic  technique,  which  made  the  conclusions  much  more  detailed  and 
less  bold  than  they  would  otherwise  have  been.  Cytologic  detail  then,  like  the  best 
ultrastructural  work  today,  makes  the  experienced  observer  cautious.  But  Morgan's 
conclusions  stand  far  better,  in  the  light  of  subsequent  discovery,  than  do  many  of 
Loeb's,  and  better  than  any  of  Driesch's. 

The  following  example  will  serve.  Loeb  had  reported,  in  the  summer  of  1894,  a 
series  of  startling  experiments  on  "extra-ovates,"31  masses  of  egg-substance  extruded 
in  the  course  of  a  brief  osmotic  shock  caused  by  immersing  the  fertilized  eggs  in 
distilled  water.  (It  is  important  to  note  here  that,  however  troubled  this  work  might 
have  been  by  over-interpretation,  it  anticipated  by  many  years  the  ligation  experiments 
on  amphibian  eggs  which  brought  Hans  Spemann  so  much  fame,  and,  in  part,  his 
Nobel  Prize.)  Loeb  observed  that  Driesch,  in  shaking  apart  early  sea  urchin  embryos, 
was  able  to  obtain  normal  development  from  any  of  the  first  four  blastomeres,  but 
never  from  blastomeres  of  the  eight  cell  stage  or  beyond.  This  was  not,  Loeb  argued, 
the  best  test  of  the  minimum  size  of  a  mass  of  "protoplasm"  capable  of  developing 
into  a  whole  larva.  A  better  one  would  be  somehow  to  divide  the  uncleaved  egg 
into  very  small  pieces,  and  to  determine  what  kind  of  development  those  might  be 
capable  of. 

The  experiments  were  done  by  causing  osmotic  swelling  of  fertilized  eggs  in 
mass  culture,  producing  small,  spherical  extrusions  through  the  fertilization  mem- 
brane; returning  the  culture  to  normal  sea  water,  allowing  it  to  develop,  and  then 
determining  what  minimum  size  of  extruded  (or  original,  i.e.,  still  within  the 
fertilization  membrane)  fragment  could  form  a  pluteus.  The  conclusion  was  that  a 
nucleated  volume  of  material  as  little  as  one-eighth  that  of  the  whole  egg  could 
produce  a  pluteus  larva.  Moreover  it  did  not  seem  to  matter  which  part  of  the  egg 
it  was:  Loeb  thought  that  any  part  would  do.  Hence  the  egg  is  isotropic,  and  the 
limitation  upon  full  differentiation  is  quantitative,  not  qualitative  as  the  argument 
from  cell  lineage  research  on  other  species  required. 

There  were,  or  should  have  been,  quantitative  questions  about  this  work,  the 
most  obvious  of  which  arises  from  the  fact  that  Loeb  had  no  way  of  deciding  what 
fraction  of  all  possible  egg  fragments  behaved  in  this  way,  and  could  not  therefore 
be  certain  that  any  part  of  an  egg,  one-eighth  the  original  volume,  would  do.  Nor 
did  he  provide  the  detailed  cytological  information  that  one  would  demand  in  order 
to  be  convinced  that  the  eighth-volume  objects  were  indeed  "plutei,"  rather  than 
asymmetric,  multicellular  masses.  Nevertheless  this  work,  coming  after  Driesch's 
(whose  conclusions  were  similar),  was  a  shock  for  those  who  thought  they  had 
reason  to  believe  that  the  egg  is  not  only  polar,  but  regionally  differentiated,  with 
the  parts  of  the  larva  laid  down,  before  cleavage  begins. 

Morgan  took  up  this  line  of  work,  but  carried  it  out  with  even  greater  care  than 
Loeb  had  done.  His  measurements  were  rigidly  quantitative  and  they  were  supported 


LAYING  THE  GHOST  75 

by  thorough  microscopical  work  on  the  fixed  and  stained  products  of  the  experiments. 
In  1895  he  published  the  results  of  studies  done  at  Naples,  on  "partial"  larvae  of 
the  sea  urchin  Sphaerechinus.31  Introducing  them,  he  wrote: 

Up  to  the  present  time  however  some  important  questions  in  regard  to  these 
("partial")  larvae  remain  unanswered.  For  instance,  we  do  not  yet  know  the 
smallest  size  possible  for  the  larvae,  or  whether  this  size  varies  according  to  the 
means  first  employed  to  get  the  fragment.  We  do  not  know  whether  the  small 
larvae  produce  the  same  number  of  cells  as  do  the  normal,  or  whether  they  can 
assume  the  definitive  form  with  fewer  cells.  The  number  of  karyokinetic  divisions 
that  take  place,  and  the  resulting  sizes  of  nuclei  and  cells  are  also  unknown. 

All  these  unknowns  he  undertook  to  make  known,  and  to  a  large  extent 
succeeded  in  doing  so,  although  the  central  objection  we  would  now  apply  to  Loeb's 
work  applied  here  as  well — that  there  was  no  proof  that  any  small  piece  whatsoever 
of  an  egg  can  give  rise  to  a  complete  larva.  That  test  had  to  wait  until  the  brilliant 
microsurgical  work  of  Horstadius  was  done,  forty  years  later;  and  its  outcome  was 
the  both  Loeb  and  Morgan  were  wrong. 

Nevertheless:  Morgan's  very  careful  study  confirmed  the  main  observations,  if 
not  the  broadest  conclusions,  of  his  predecessors,  and  added  more  to  shock  the 
predeterminists.  He  minced  no  words  in  the  conclusion  to  this  very  complete  paper: 

The  conclusion  is  forced  upon  us — and  I  see  no  escape  from  it — that  the 
formation  of  the  embryo  is  not  controlled  by  the  form  of  cleavage.  The  plastic 
forces  heed  no  cell  boundaries  but  mould  the  germ-mass  regardless  of  the  way  it 
is  cut  up  into  cells.  That  the  forms  assumed  by  the  embryo  in  successive  stages 
are  not  dependent  on  cell-division  may  be  demonstrated  in  almost  any  egg. 
Whitman's  conclusions  receive  I  think  strong  support  from  the  results  of  the 
experiments  recorded  in  the  preceding  pages. 

"Whitman's  conclusions"  are  those  offered  in  an  earlier  lecture  at  Woods  Hole 
("On  the  inadequacy  of  the  cell  theory").  And  how  Whitman  must  have  simmered, 
to  discover  his  thoughts  on  the  matter,  oflfered  originally  in  a  different  context  and 
in  aid  of  quite  different  conclusions,  here  quoted  in  support  of  radical  epigenesis! 

It  is  instructive  to  follow  Morgan's  continued  work  on  this  problem,  especially 
as  described  in  papers  published  six  and  fifteen  years  later.  The  radicalism  disappears 
as  the  breadth  and  incisiveness  of  the  experiments  increases.  Thus  the  conclusion 
of  the  1901  paper  is  already  much  softer:  the  reality  of  determinate  cleavage  is  at 
least  recognized,  and  the  extent  to  which  fragments  of  regulative  eggs  may  develop 
is  no  longer  overstated:32 

We  see  then  that  even  in  cases  with  a  perfectly  definite  type  of  cleavage  that 
give  rise  to  embryos  having  definite  relations  to  this  cleavage  there  is  no  absolutely 
necessary  relation  between  the  two,  for  if  the  conditions  are  changed  the  relation 
may  also  be  altered  or  at  least  parts  of  the  egg  may  invaginate  that  do  not  do  so 
under  ordinary  circumstances.  Nevertheless  the  pre-existing  protoplasmic  relations 
in  the  segmenting  egg  appear  to  have  an  important  influence  on  the  formation  of 
the  normal  embryo  .  .  .  [my  emphasis]. 

The  last  paper  of  this  group,  published  in  1910,33  is  a  sophisticated  analysis 
whose  important  conclusion,  for  our  purpose  here,  is  implicit  in  its  title:  "The 
effects  of  altering  the  position  of  the  cleavage  planes  in  eggs  with  precocious 
specification."  The  recognition  is  now  an  open  one;  that  some  degree  of  specification 
exists  in  all  uncleaved  eggs;  in  effect,  a  high  degree — "precocious  specification"  —in 
mosaic  development,  and  a  low  one  in  regulative.  As  well  the  development  of  eggs 


76  P.   R.  GROSS 

such  as  that  of  the  sea  urchin  might  have  been  characterized  as  showing  "low-but- 
not-zero"  specification.  Morgan's  progress  was  the  conversion  of  a  radical  epigenesis 
to  epigenesis  without  isotropy;  while  the  later  research  of  Wilson  and  Conklin  was 
a  conversion  of  rigid  predeterminism  to  a  relative  determinacy  of  cleavage,  with 
recognition  of  the  capacity  for  regulation  and  the  importance  of  cell-cell  interactions 
in  guiding  the  course  of  embryo-formation,  even  in  mosaic  forms. 

With  the  MBL's  first  decade  behind  it,  its  scope  of  scientific  interests  already 
greatly  broadened  and  America  still  innocent  of  the  drumbeat  for  war  reverberating 
throughout  Europe,  these  embryologists  could  look  with  satisfaction  on  their 
accomplishments,  although  few  were  yet  of  an  age  to  look  backward,  and  none 
rested  on  their  laurels.  In  a  nation  that  had  so  recently  been  a  biological,  if  not 
wholly  a  scientific  backwater,  there  were  now  not  one  or  two,  but  a  growing  number 
of  research  universities,  with  departments  whose  biological  faculties  matched  and, 
in  some  cases,  surpassed  their  equivalents  in  the  ancient  institutions  of  Europe. 
Here  was  the  MBL,  where  every  summer  there  gathered  a  group  of  biologists  who, 
because  of  the  war  in  Europe  and  its  attendant  dislocations,  surpassed  in  productivity 
and  would  soon  surpass  in  influence  the  group  that  gathered  at  Naples.  The  MBL 
offered,  moreover,  advanced  instruction  for  research  students,  under  the  guidance 
of  that  group,  and  hence  under  a  teaching  faculty  that  could  nowhere  be  matched. 
This  insured  that  the  enterprise  of  fundamental  research  in  biology  would  not 
merely  survive  in  the  U.S.A.,  but  that  by  the  law  of  compound  interest  (the  mean 
number  of  trainees  per  trainer  being  greater  than  one)  it  would  grow  exponentially 
in  the  future. 

Most  important,  embryology,  jewel  in  the  crown  of  evolution  a  decade  earlier, 
had  been  made  over  into  something  else,  something  more  important.  Wilson  had 
been  right:  cell  lineage  toppled  the  germ  layer  theory  of  development,  and  with  it 
had  fallen  the  Haeckelian  paradigm  of  embryological  research.  The  embryo  and  its 
doings  came  to  center-stage  in  their  own  right,  as  a  biological  subject  worthy  of  the 
most  intense  study  because  its  secrets  were  clearly  at  the  heart  of  multicellular  life. 
Recapitulation  was  an  irrelevancy:  the  stages  of  vertebrate  embryogenesis  (only) 
showed  resemblances,  and  only  resemblances,  among  genera,  orders,  and  classes; 
but  certainly  not  as  between  embryos  of  one  and  adults  of  an  other.  And  the 
reasons  for  such  resemblance  had  lost  all  value  as  explanation  of  development. 
Explanations  would  have  to  be  sought  within  the  cell  and  even  within  the  molecules 
of  the  cell.  The  genealogical  ghost  had  been  driven  out. 

In  the  effort  to  rout  remaining  ghosts,  the  radical  epigeneticists  had  attempted 
to  find  explanations  for  development  solely  in  the  physics  and  chemistry  of 
"protoplasm,"  and  in  the  stimuli  and  responses  of  the  surrounding  medium  and 
the  zygote,  respectively.  They  failed,  by  and  large,  to  convince.  The  response  to 
them  might  well  have  been  the  reasonable  one  that,  since  the  eggs  of  two  different 
species,  developing  side-by-side  in  the  same  culture  dish,  produce  two  quite  different 
embryos,  and  by  two  different  patterns  of  cleavage  and  morphogenetic  movement, 
there  must  be  hereditary  determinants  of  some  kind  directing  development.  And  of 
course  it  would  soon  become  apparent  that  genes  on  chromosomes  were  the  likeliest 
although  not  the  only  candidates.  For  the  radical  epigeneticists,  not  yet  quite 
reconciled,  this  argument  retained  a  smell  of  preformationism,  but  it  was  at  least  a 
preformationism  of  matter,  not  of  spirit.  In  some  sense  it  was  a  return  to  the 
eighteenth  century,  and  yet  means  by  which  the  chemistry  and  physics  of  the 
postulated  determinants  might  be  investigated  were  already  coming  into  being — 
hence  Loeb's  increasing  concern  with  turning  "colloid  chemistry"  and  the  "proto- 
plasm" concept  into  a  proper  physical  chemistry  of  proteins.  That  concern  would 


LAYING  THE  GHOST 

spawn,  among  Loeb"s  students  and  descendants,  an  important  part  of  modern 
protein  chemistry  and  enzymology. 

Morgan,  a  hard-headed  experimentalist  and  logician  to  the  last,  would  continue 
a  little  too  long  to  shave  with  Occam's  Razor  (hence,  perhaps,  his  somewhat  untidy 
beard);  but  his  energy  and  powerful  intelligence  would  turn  him  at  the  last — the 
genetics  and  cytogenetics  of  Drosophila  having  shown  what  it  could  reveal — to 
attempt  a  synthesis  of  genetic  predetermination  and  epigenetic  biochemistry.  He 
did  not  solve  the  problem,  but  he  was  able  to  state  it,  and  thereby  to  summarize 
outcomes  of  those  exorcisms  I  have  touched  upon  here,  as  a  part  of  his  Nobel 
address  of  1934:34 

...  it  is  conceivable  that  different  batteries  of  genes  come  into  action  one 
after  the  other,  as  the  embryo  passes  through  the  stages  of  its  development.  This 
sequence  might  be  assumed  to  be  an  automatic  property  of  the  chain  of  genes. 
Such  an  assumption  would,  without  proof,  beg  the  whole  question  of  embryonic 
development,  and  could  not  be  regarded  as  a  satisfactory  solution. 

But  it  might  be  that  in  different  regions  of  the  egg  there  is  a  reaction  between 
the  kind  of  protoplasm  present  in  those  regions  and  in  specific  genes  in  the 
nuclei;  certain  being  affected  in  one  region  of  the  egg,  other  genes  in  other 
regions.  Such  a  view  might  give  also  a  purely  formal  hypothesis  to  account  for 
the  differentiation  of  the  cells  in  the  embryo.  The  initial  steps  would  be  given  in 
the  regional  constitution  of  the  egg. 

The  first  responsive  output  of  the  genes  would  then  be  supposed  to  affect  the 
protoplasm  of  the  cells  in  which  they  lie.  The  changed  protoplasm  would  now 
act  reciprocally  on  the  genes,  bringing  into  activity  additional  or  other  batteries 
of  genes.  If  true,  this  would  give  a  pleasing  picture  of  the  developmental  process. 

EXODOS 

Realization  of  the  broad  possibilities  offered  by  Morgan  in  that  lecture  was  not 
to  be  until,  I  should  judge,  the  early  1970's.  A  small  volume  devoted  to  the  question 
of  gene  action  and  the  control  of  embryogenesis — not  the  first  and  not  necessarily 
the  most  complete  of  several  near-contemporary  volumes — was  published  by  Max 
Hamburgh  in  197 1.35  I  recommend  it,  for  its  brevity,  to  the  non-embryologist  reader 
who  would  gain  an  impression  of  what  had  been  done  in  the  intervening  forty  years 
about  "Theories  of  Differentiation."  To  summarize  the  book,  which  is  itself  a 
summary:  a  productive  science  of  developmental  genetics,  imagined  only  dimly  by 
Morgan,  emerged,  and  while  its  main  achievements  were  at  first  in  identifying  genes 
involved  in  later  morphogenesis  of  various  animals,  it  left  little  doubt  that  morpho- 
genesis is  under  the  direct  control  of  genes.  All  the  new  insights  into  pathways  and 
mechanisms  of  intermediary  metabolism  had  been  applied  to  embryos,  and  the 
energetics  of  development  had  come  in  a  general  way  to  be  understood. 

Curiously,  it  was  through  biochemistry  rather  than  formal  genetics,  that  the  first 
coordinated  attacks  upon  genetic  control  of  early  development,  hence  of  determi- 
nation, were  mounted.  Protein  synthesis,  now  understood  to  be  the  second  major 
step  in  gene  action  (transcription  being  the  first),  was  shown  to  be  continuous  and 
necessary  throughout  development  and  from  the  very  start:  hence  there  were  no 
"silent"  periods  for  genetic  control  of  development.  And  a  large  part  of  that 
synthesis  was  shown  to  be  under  control  of  genetic  messengers  prepared  earlier- 
during  oogenesis;  thus  providing  an  explanation  for  the  apparent  independence  of 
some  steps  in  pre-gastrula  development  from  the  zygotic  genome,  and  an  important 
set  of  candidates — now  in  definable  chemical  form — for  those  morphogenetic 
determinants  upon  whose  existence  the  lineagists  had  staked  their  reputations.36  3 


78  P.   R.   GROSS 

How  appropriate  it  is  that  advanced  embryological  research  should  again  be 
preoccupied,  today,  with  cell  lineage;  of  systems  in  which  the  differential  distribution 
of  specific  and  identified  gene  products  can  be  traced,  and  in  which  the  mechanisms 
by  which  those  genes  are  activated  or  repressed  differentially  can  be  studied!  I  know 
no  better  general  reference  for  the  state  of  these  issues  at  the  time  I  write  this — and 
for  the  embryo's  ghostly  intelligence,  if  any — than  the  volume  of  lectures  given  in 
the  MBL's  embryology  course  the  summer  of  1983,38  ninety  years  after  its  first 
offering  in  a  small  wooden  building  at  Woods  Hole. 

NOTES  AND   REFERENCES 

1  I  am  much  in  debt  to  the  following  for  insights  into  the  work  and  personalities  of  founders  of  the  MBL: 

Garland  Allen,  for  his  Life  Science  in  the  Twentieth  Century  (Cambridge  University  Press. 
1978),  for  several  important  papers  on  T.  H.  Morgan,  and  for  his  biography  Thomas  Hunt 
Morgan:  The  Man  and  his  Science  (Princeton  University  Press,  1978);  Jane  Maienschein,  for 
her  paper  on  Cell  Lineage,  Ancestral  Reminiscence,  and  the  Biogenetic  Law  (J.  Hist.  Biol.  II: 
129-158,  1978)  and  for  lectures  delivered  in  1984  at  the  MBL  and  at  the  Ischia  meeting;  and 
Jeffrey  Werdinger,  whose  doctoral  dissertation  Embryology  at  Woods  Hole:  The  Emergence  of  a 
New  American  Biology  (1980),  is  available  from  University  Microfilms,  Ann  Arbor,  Michigan. 
A  recent  work  of  European  history  in  the  period  of  interest  has  been  helpful:  Norman  Stone, 
Europe  Transformed:  1878-1919  (Harvard  University  Press,  1984).  The  first  edition  of  Eric 
Davidson's  Gene  Activity  in  Early  Development  (Academic  Press,  New  York,  1968)  is  a  valuable 
source  for  cell  lineage  research  in  relation  to  experimental  embryology. 

2  "Ghost"  is  no  hyperbole.  The  common  definition — the  disembodied  soul  of  a  dead  person — is  only  the 

second.  The  first,  (e.g.,  in  Webster's  New  Collegiate  Dictionary,  1981)  is  "The  seat  of  life  or 
intelligence."  That  one,  applied  to  the  entities  imagined  by  natural  philosophers  to  direct 
embryonic  development,  fits  well.  At  worst,  it  is  a  metonymy. 

3  RUSSELL,  BERTRAND.  1945.  A  History  of  Western  Philosophy,  fourteenth  printing.  Simon  and  Schuster, 

New  York.  Pp.  546  if.  The  particular  plain  words  between  quotation  marks  were  chosen  for  use 
in  an  editorial  introduction  to  a  series  of  papers  on  "The  Mind,"  in  The  Wilson  Quarterly  8(5): 
47  (1984). 

4  Amos  Emerson  Dolbear  (1837-1910);  physicist  and  inventor.  Professor  of  Physics  at  Tufts  College.  The 

quoted  passage  is  from  the  sixth  of  the  MBL's  Biological  Lectures  of  1895,  but  the  reluctant 
conclusion  that  creation  of  the  ether  must  have  taken  place  is  reached  in  the  fifth.  See  Biological 
Lectures.  Ginn  &  Company,  Boston,  1896,  pp.  63-82;  83-99. 

5  Sadi  Carnot  laid  the  foundation  for  the  First  Law  in   1 824.  The  grand  structure  was  completed  by 

William  Thomson,  later  Lord  Kelvin,  24  years  later.  The  Second  Law,  which  identifies  the 
restricting  principle  in  processes  whose  forms  of  energy  include  heat,  was  established  by  Clausius 
and  Lord  Kelvin  in  the  1850's.  The  next  half-century  saw  the  rise  of  a  statistical,  kinetic  theory 
of  matter,  and  a  probabilistic  definition  of  entropy.  Toward  the  end  of  that  time,  a  thoughtful 
physicist  would  have  judged  the  appearance  of  form  in  a  formless  macroscopic  object,  without 
appropriate  external  work  done  on  it,  to  be  about  as  probable  as  the  spontaneous  boiling  of 
water  in  a  kettle  held  in  a  deep-freeze.  See  R.  A.  MILLIKAN,  D.  ROLLER,  AND  E.  C.  WILSON. 
1937.  Mechanics,  Molecular  Physics,  Heat  &  Sound.  Ginn  &  Co.  (M.I.T.  Press  edition,  1965). 

6  Virtual  particles  in  a  quantum  vacuum  are  individually  undetectable,  but  they  exist  because  their 

macroscopic  consequences,  such  as  the  Casimir  effect,  exist.  Moreover,  if  the  "Big  Bang"  took 
place  in  an  interval  like  the  Planck  time,  10  41  s,  that  might  be  considered  an  event  of  creation 
(were  not  there  a  "creationism"  with  which  I  reject  the  remotest  connection).  See  JOHN  D. 
BARROW  AND  JOSEPH  SILK.  1983.  The  Left  Hand  of  Creation.  Basic  Books,  Inc.,  New  York, 
for  a  modern  cosmologist's  view. 

7  The  reference  is  to  BARBARA  W.  TUCHMAN.  1966.  The  Proud  Tower.  Macmillan,  New  York. 

8  NORDENSKIOLD,  ERIK.  1928.  The  History  of  Biology.  Translated  from  the  Swedish  by  L.  E.  Eyre,  Tudor 

Publishing  Co.,  New  York. 

9  Some  dates:  Bismarck's  "blood  and  iron"  statement  was  made  in   1848;  three  wars  followed  in  the 

unification  of  Germany  under  Prussian  hegemony.  The  last,  the  Franco-Prussian  war,  ended  in 
disaster  for  France.  With  the  loss  of  Alsace-Lorraine,  the  humiliating  capture  of  Napoleon  III, 
and  the  indemnity  of  a  billion  dollars  France  was  forced  to  pay,  there  were  planted  the  seeds  of 
the  Great  War  of  1914.  The  German  Empire  under  Wilhelm  I  was  proclaimed  formally  on 
January  18,  1871,  at  Versailles. 

10  Although  Haeckel  and  Gegenbaur  were  collaborators,  and  Haeckel  and  Anton  Dohrn  united  for  a  time 

in  friendship  and  in  their  views,  Gegenbaur  and  Dohrn  were  mortal  enemies.  The  former, 
especially,  never  lost  an  opportunity  to  malign  the  latter. 


LAYING  THE  GHOST  79 

"  NORDENSKIOLD,  E.  Op.  cit.,  for  Haeckel's  antecedents;  but  see,  for  a  devastating  judgment  of  Goethe 
as  scientist,  the  essay  of  SIR  CHARLES  SHERRINGTON.  1949.  Goethe  on  Nature  and  on  Science. 
2nd  ed.  Cambridge  University  Press. 

12  See,  inter  alia,  J.  WERDINGER,  Op.  cit.,  and  STEPHEN  JAY  GOULD.  1977.  Ontogeny  and  Phytogeny. 

Belknap  Press  of  Harvard  University,  Cambridge,  Massachusetts. 

13  LILLIE,  F.  R.   1944.  The  Woods  Hole  Marine  Biological  Laboratory,  University  of  Chicago  Press.  P. 

123. 

14  WHITMAN,  C.  O.  1878.  The  embryology  of  Clepsine.  Q.  J.  Microsc.  Sci.  18:  215-314. 

15  LILLIE,  F.  R.  1944.  Op.  cit.  P.  160. 

16  WHITMAN,  C.  O.  1895.  Evolution  and  epigenesis.  In  Biological  Lectures.  Ginn  &  Co.,  Boston. 

17  WILSON,  E.  B.  The  Cell  in  Development  and  Heredity,  Macmillan,  New  York,  edition  of  1925. 
18GROEBEN,  C.  AND  I.  MiJLLER.  1975.  Exhibition  Catalogue:  The  Naples  Station  at  the  Time  of  Anton 

Dohrn.  Pp.  89-90.  The  quartet  was  the  A  major,  composed  by  Schumann  in  1842.  Elsewhere 

in  this  volume  is  reproduced  a  letter  from  Wilson  to  Dohrn  (7  June  1899),  signed  with  a  cello 

device  and  quoting  the  cello  line,  starting  at  the  12th  measure,  of  Beethoven's  C-sharp  minor 

string  quartet.  Op.  131. 
w  WILSON,  E.  B.  1892.  The  cell-lineage  of  Neresis:  a  contribution  to  the  cytogeny  of  the  annelid  body.  /. 

Morphol.  6:  361-480. 

2(ICONKLIN,  E.  G.  1897.  The  embryology  of  Crepidnla.  J.  Morphol.  13:  1-226. 
21  LILLIE,  F.  R.  1895.  The  embryology  of  Unionidae.  J.  Morphol.  10:  1-100. 
!2  LILLIE,  F.  R.  1944.  Op.  cit.  P.  124. 
21  WILSON,  E.  B.  1904.  Experimental  studies  on  germinal  localization.  I.  The  germ-regions  of  the  egg  of 

Dentalium.  AND  //.  Experiments  on  the  cleavage-mosaic  in  Patella  and  Dentalium.  J.  Exp. 

Zool.  1:  1-72;  197-268. 
24CONKLIN,  E.  G.  1905.  Organization  and  cell-lineage  of  the  ascidian  egg.  /  Acad.  Natl.  Sci.  Philadelphia 

13:  1-119. 
15  WERDINGER,  J.  Op.  cit.  P.  266. 

26  "Every  historian  is  aware  that  the  'revolutionary  moment'  is  rather  apt  to  be  the  time  in  which  certain 

previously  known  ideas,  or  theories,  or  doctrines  receive  a  new  turn  that  brings  them  forcibly 
to  the  minds  of  everyone,  or  are  given  a  sudden  incisiveness  by  new  experiments  .  .  .  made  in 
so  striking  a  way  that  no  one  can  escape  considering  them." 

This  passage  is  from  I.  Bernard  Cohen,  in  his  Introduction  to  Margaret  G.  Foley's  translation 
of  Galvani's  De  Mribus  Electricitatis  .  .  .,  issued  by  the  Burndy  Library,  Norwalk,  Connecticut 
(1953). 

27  LOEB,  JACQUES.   1892.  Investigations  in  physiological  morphology.  III.  Experiments  on  cleavage.  J. 

Morphol.  7:  253-263. 

28  LOEB,  JACQUES.  1895.  On  the  limits  of  divisibility  of  living  matter.  Pp.  55-65  in  Biological  Lectures. 

Ginn  &  Co.,  Boston. 

29  Monism  can  be  traced  back  at  least  to  Parmenides.  It  is  the  doctrine  that  all  manifestations  of  the 

world  are  properties  of  a  single,  material  unity.  The  Greek  atomists  were  monists:  so  was 
Spinoza.  Curiously,  Haeckel  was  a  monist.  Ernst  Mach  (1838-1916),  physicist-turned-philosopher, 
was  the  leading  nineteenth  century  exponent.  He  was  also  a  vitalist  (though  he  would  have 
denied  it).  Loeb's  affinities  for  a  doctrine  that  holds  all  phenomena,  perception  as  well  as  external 
action,  to  be  manifestations  of  an  underlying  material  reality,  can  readily  be  understood. 

30  LOEB,  JACQUES.  1912  (January).  The  mechanistic  conception  of  life.  Pp.  5-21  in  Pop.  Sci.  Monthly. 

31  See  ref.  28  for  Loeb,  and  for  Morgan:  T.  H.  MORGAN.  1895.  The  'partial'  larvae  of  Sphaer echinus. 

Archiv.  f.  Entwicklungsmech.  2:  81-126. 

32  MORGAN,  T.  H.  1901.  The  proportionate  development  of  partial  embryos.  Archiv.  f.  Entwicklungsmech. 

13:416-635. 

33  MORGAN,  T.  H.  1910.  The  effects  of  altering  the  position  of  the  cleavage  planes  in  eggs  with  precocious 

specification.  Archiv.  f.  Entwicklungsmech.  29:  205-224. 

34  This  speech  was  published  in  Scientific  Monthly,  41:  5-18  (1935).  Reproduced  here  is  the  quotation 

from:  IAN  SHINE  AND  SYLVIA  WROBEL.  1976.  Thomas  Hunt  Morgan:  Pioneer  of  Genetics.  The 
University  Press  of  Kentucky,  Lexington. 

35  HAMBURGH,  MAX.  1971.  Theories  of  Differentiation.  American  Elsevier  Pub.  Co.,  New  York. 

36  GROSS,  P.  R.  1968.  Biochemistry  of  differentiation.  Ann.  Rev.  Biochem.  37:  631-660. 

37  See  also  two  other  reviews,  published  fifteen  years  apart:  P.  R.  GROSS.  1967.  Curr.  Top.  Dev.  Biol.  2: 

1-46;  and  E.  H.  DAVIDSON  et  al.  1982.  Science  217:  17-26. 

38JEFFERY,  W.  R.  AND  R.  A.  RAFF,  eds.  1983.  Time,  Space,  and  Pattern  in  Embryonic  Development. 
MBL  Lectures  in  Biology,  Vol.  2.  Alan  R.  Liss,  Inc..  New  York. 


Addenda  to  "Laying  the  ghost:  embryonic  development,  in  plain  words,"  by  Paul  R.  Gross. 


PLATE  I.  Top:  Old  Main,  the  MBL's  first  teaching  and  research  laboratory,  on  the  site  of  which  the 
Loeb  Laboratory  stands  today.  Photograph  made  by  Baldwin  Coolidge  in  1893.  Bottom:  the  first 
Embryology  class,  1893.  Coolidge  photograph.  C.  O.  Whitman  at  center,  standing.  F.  R.  Lillie  at  his  far 
right.  The  four  women  students,  left  to  right,  are:  S.  Emma  Keith;  Elizabeth  E.  Bickford;  Bertha  M. 
Brown;  Marie  L.  Minor.  The  blackboard  inscription  is  "Isotropism."  Others  of  the  class  are  identified 
opposite  page  91  in  Lillie.  1944  (13).  This  and  all  other  photographs  courtesy  of  the  Archives,  MBL 
Library. 


PLATE  II.  Top  left:  C.  O.  Whitman,  1908.  Photograph  by  R.  M.  Strong.  Top  right:  E.  B.  Wilson. 
Origin  of  photograph  unknown.  Bottom  left:  E.  G.  Conklin,  1922.  Bottom  right:  F.  R.  Lillie,  1921. 
Photos  of  Conklin  and  Lillie,  and  of  Loeb  in  Plate  III,  were  taken  by  the  legendary  "Wireless  Pete"  of 
Woods  Hole. 


PLATE  III.  Left:  Jacques  Loeb  in  1922.  Right:  Thomas  Hunt  Morgan  at  age  25,   1891.  From  the 
Johns  Hopkins  University  Yearbook. 


Reference:  Biol.  Hull.  168  (suppl.):  80-87.  (June,  1985) 


CELL  INTERACTIONS:   THE   ROOTS  OF  A  CENTURY  OF  RESEARCH 

JAMES   D.   EBERT 

Carnegie  Institution  oj  Washington,  1530  P  Street,  N.  W.,  Washington.  DC  20005 

ABSTRACT 

Only  rarely  have  marine  organisms  provided  experimental  systems  par  excellence 
for  concerted  and  continuing  analyses  of  mechanisms  of  cell  interactions,  in  either 
of  the  two  main  categories  I  consider:  inductive  and  morphogenetic.  Although  there 
were  significant  findings  with  marine  organisms  from  the  beginning,  they  were 
frequently  overshadowed,  possibly  because  the  seasonal  character  of  research  on 
marine  organisms  resulted  in  a  focus  on  experiments  that  did  not  require  continuity, 
and  on  comparative,  rather  than  on  mechanistic,  analyses. 

The  roots  of  the  study  of  inductive  interactions — contemporary  with  the 
establishment  of  the  Marine  Biological  Laboratory  (MBL) — are  found  in  the  work 
of  Chabry  and  Roux  on  the  development  of  isolated  blastomeres  of  an  ascidian  and 
a  frog,  respectively,  after  destruction  of  neighboring  blastomeres.  Roux's  findings 
had  a  larger  impact  than  Chabry's,  in  part  because  Roux  cast  his  findings  in  larger 
terms  than  did  Chabry,  but  also  because  of  the  differences  in  experimental 
approaches  that  emerged.  The  fact  that  the  blastomeres  of  many  marine  embryos 
could  be  dissociated  readily,  lead  to  repeated  comparative  studies  of  the  capacity 
for  development  of  isolated  blastomeres.  The  use  of  amphibian  systems  produced  a 
drive  to  understand  the  failure  of  independent  development;  and  the  use  of 
transplantation  techniques  resulted  in  the  discovery  of  the  "organizer,"  that  set  in 
train  fifteen  years  of  intensive  and  largely  fruitless  research  on  its  chemical  nature, 
punctuated  by  Lester  Earth's  demonstration  of  "neural  differentiation  without 
organizer." 

The  impact  of  studies  of  marine  forms  on  the  emergence  of  today's  ideas  on 
morphogenetic  interactions — cell  adhesion  molecules,  etc. — is  less  clear.  The  con- 
tributions of  Herbst  in  dissociating  embryos  in  calcium-free  sea  water,  and  of  H.  V. 
Wilson,  and  later  Galtsoff,  in  re-aggregating  sponge  cells  had  no  significant  effect 
until  after  Holtfreter's  far  reaching  studies  of  "tissue  affinity." 

INTRODUCTION 

When  I  entered  the  Johns  Hopkins  University  as  a  graduate  student  in  1946, 
destined  to  work  with  Professor  B.  H.  Willier,  the  field  of  embryology,  as  it  was 
then  called  (for  the  phrase  "developmental  biology"  did  not  become  popular  until 
the  early  1950s),  was  in  the  doldrums.  There  were,  to  be  sure,  a  few  active 
laboratories,  with  extraordinary  leaders;  in  addition  to  Willier,  Paul  Weiss,  then  at 
Chicago,  Viktor  Hamburger,  at  Washington  University,  Johannes  Holtfreter,  at 
Rochester,  and  Victor  Twitty  at  Stanford,  come  to  mind.  With  such  luminaries 
working  in  the  United  States,  and  at  least  an  equal  number  in  Europe — Horstadius 
and  Runnstrom;  Brachet,  Dalcq  and  Pasteels;  Woerdemann  and  ten  Cate;  and 
Waddington  and  Abercrombie,  why  do  I  say  the  field  was  in  the  doldrums?  My 
answer  is  this:  each  of  the  leading  centers  had  its  own  special  interests,  its  own 
special  preserve.  Each  was  tackling  important  problems,  with  the  technology  of  the 
day.  Still,  there  was  no  major  thrust,  no  "true  imperative"  in  the  field.  With  the 

80 


EARLY   STUDIES  OF  CELL   INTERACTIONS  81 

exception  of  Hamburger  and  Willier  in  the  United  States,  and  Brachet  and 
Waddington  in  Europe,  most  of  the  leaders  in  the  field  paid  little  attention  to 
genetics.  The  embryology  textbooks  of  the  day  commonly  contained,  at  most,  a 
single  chapter  on  developmental  genetics,  despite  the  contributions  of  Dunn, 
Landauer,  Wright,  Hadorn,  and  others.  (Willier  el  ai.  1955,  reflects  the  state  of  the 
field  after  World  War  II). 

The  seeds  of  change,  that  would  lead  to  a  spirited  renaissance  of  the  field,  had 
been  sown  in  the  1930s  and  1940s,  by  Ephrussi  and  Beadle,  by  Beadle  and  Tatum, 
by  Caspersson,  and  by  Brachet,  to  name  only  a  few  of  the  pioneers  who  pointed 
the  way  to  the  intellectual  wedding  of  developmental  biology  and  genetics.  But  they 
had  only  begun  to  influence  the  field. 

Less  heralded  at  the  time  were  the  strides  taken  by  Holtfreter,  and  later  by 
Moscona  and  Abercrombie,  in  analyzing  what  Holtfreter  called  "tissue  affinity," 
studies  that  were  the  forerunners  of  today's  widespread  interest  in  "cell  adhesion 
molecules."  I  shall  return  to  these  studies  later  under  the  (over  simplified)  heading 
"Cell  Surface  Interactions  in  Morphogenesis." 

Let  me  return  to  the  embryology  of  the  1940s.  I  observed  that  the  field  lacked 
a  primary  focus.  It  had  not  always  been  so.  In  fact,  for  nearly  two  decades,  attention 
was  focused,  more  in  Europe  than  in  the  United  States,  but  to  a  considerable  degree 
throughout  the  world,  on  what  came  to  be  called  "inductive  cell  interactions."  Cell- 
cell  interaction  is  a  necessary  condition  for  the  formation  of  the  cellular  architecture 
of  organs  as  well  as  for  their  organization  and  interconnection.  The  first  demonstration 
of  embryonic  induction,  by  Spemann  (190  la,  1907)  and  W.  H.  Lewis  (1904,  1907a, 
b)  provides  the  classic  example.  During  normal  development  the  optic  vesicle  grows 
out  to  contact  the  epidermis.  A  short  time  later  the  vesicle  invaginates  to  form  the 
optic  cup;  at  the  same  time  the  epidermis  at  the  point  of  contact  thickens  and  sinks 
beneath  the  surface,  following  the  retinal  surface  of  the  cup.  Becoming  detached 
from  the  epidermis,  this  group  of  cells  rounds  up  and  differentiates  to  form  the 
lens,  generally  as  a  consequence  of  the  influence  of  (or  induction  by)  the  optic 
vesicle. 

The  nature  of  induction  was  then,  and  remains  today,  a  fascinating  problem.  In 
the  1940s  however  embryologists  were  groping — there  is  no  better  word — for  a  new 
approach,  following  the  failure  of  then  existing  techniques  to  solve  a  problem  whose 
solution  had  seemed,  for  a  time,  to  be  within  their  grasp. 

Let  us  return  to  the  roots  of  the  problem. 

INDUCTIVE  CELL  INTERACTIONS 

The  roots  of  the  study  of  inductive  interactions — contemporary  with  the 
establishment  of  the  Marine  Biological  Laboratory — are  found  in  the  work  of 
Chabry  (1887;  see  Morgan,  1927),  and  Roux  (1888)  on  the  development  of  isolated 
blastomeres  of  an  ascidian  and  a  frog,  respectively,  after  destruction  of  neighboring 
blastomeres.  Roux's  findings  had  a  larger  impact  than  Chabry's,  in  part  because 
Roux  cast  his  findings  in  larger  terms  than  did  Chabry.  The  questions  addressed  in 
these  works  are  the  following:  Is  the  differentiation  of  a  given  blastomere  a  function 
of  its  position  in  the  whole,  i.e.,  is  it  dependent  upon  contact  or  influences  of 
neighboring  cells,  or  is  its  development  an  act  of  "self-differentiation,"  which  implies 
that,  at  a  definite  moment,  it  contains  all  the  specific  conditions  for  further 
differentiation?  Isolation  and  defect  experiments  were  widely  used  in  analyzing  this 
problem. 

The  blastomeres  of  the  developing  frog's  egg  are  so  closely  united  that  their 
complete  isolation  is  difficult.  At  the  2-cell  or  4-cell  stage  of  Rana  esculenta,  Roux 


J.    D.   EBERT 

pricked  one  blastomere  with  a  heated  needle  in  order  to  exclude  it  from  further 
development.  This  method  is  far  less  satisfactory  than  complete  isolation  because  it 
is  difficult  to  determine  the  extent  of  injury  to  the  pricked  blastomere  and  the  effect 
of  the  punctured  blastomere  on  the  surviving  blastomere.  However,  the  following 
results  were  obtained:  ( 1 )  by  pricking  one  blastomere  at  the  2-cell  stage,  either  a 
lateral  or  an  anterior  hemi-embryo  was  produced,  depending  on  the  direction  of 
the  first  cleavage  plane;  and  (2)  by  pricking  one  blastomere  at  the  4-cell  stage, 
"three-quarter"  embryos  were  produced.  From  these  results  Roux  concluded  that 
from  the  4-cell  stage,  development  is  a  mosaic  of  four  essentially  "self-differentiating" 
pieces. 

In  many  cases  the  half-embryo  restored  the  missing  half.  According  to  Roux 
this  process,  which  he  called  "post-generation,"  was  accomplished  by  utilization 
("organization")  of  the  protoplasm  of  the  punctured  blastomere  by  the  half-embryo. 
It  appears  more  likely,  however,  that  the  hot  needle  merely  retarded  development, 
and  did  not  exclude  the  punctured  blastomere  from  further  development. 

It  was  shortly  thereafter  that  Driesch  presented  his  now  famous  studies  of  the 
production  of  whole  embryos  from  isolated  blastomeres  of  the  sea  urchin.  This 
separation  was  accomplished  by  shaking  fertilized  eggs  in  artificial  sea  water  from 
which  calcium  had  been  removed,  following  the  method  of  Herbst. 

These  contrasting  observations  of  Roux  and  Driesch  had  far  reaching  conse- 
quences. Indeed,  from  them  emerged  two  separate  trails  of  research,  each  with  its 
own  philosophy.  In  his  book  Experimental  Embryology,  Morgan  (1927)  devoted 
five  successive  chapters  to  comparative  studies  of  the  capacity  for  development  of 
isolated  blastomeres,  and  the  fate  of  cells  and  their  location — the  study  of  cell 
lineage.  The  fact  that  the  blastomeres  of  many  marine  embryos  could  be  dissociated 
readily  led  to  an  emphasis  on  comparative  studies  of  the  development  of  isolated 
blastomeres.  As  Morgan  reported,  "It  has  been  found  that  the  isolated  blastomeres 
of  sea  urchins,  of  certain  hydroids,  of  nemerteans,  of  Amphioxns,  offish,  of  Triton 
and  of  the  frog  give  rise  to  whole  embryos;  while  the  isolated  blastomeres  of 
ctenophores,  molluscs  and  ascidians  give  rise  to  half-embryos."  In  contrast,  the  use 
of  amphibian  systems,  stemming  from  the  observations  by  Roux,  produced  a  drive 
to  understand  the  failure  of  independent  development,  that  is  to  identify  that  part 
of  an  embryo  so  crucial  to  the  developmental  process  that  its  absence  resulted  in  a 
defective  embryo.  It  was  this  drive  that  promoted  the  study  of  cell  interactions  and 
resulted  in  the  discovery  of  embryonic  induction.  It  is  this  trail  that  I  shall  follow, 
pausing  only  to  reflect  briefly  on  the  origins  of  these  divergent  trails  of  research.  At 
this  meeting.  Gross  has  addressed  the  philosophic  differences  between  the  morphol- 
ogists  (cell  lineage)  and  the  physiologists  (Entwicklungsmechanik).  I  would  add  only 
one  other  perception,  based  on  E.  G.  Conklin's  lectures  and  personal  communications 
from  B.  H.  Willier. 

At  the  end  of  his  career,  Conklin  lectured  each  summer  in  the  Embryology 
Course  at  the  Marine  Biological  Laboratory.  In  his  lecture  in  the  summer  of  1946, 
he  dwelt  on  the  difficulties  presented  by  "seasonal  research"  and  spoke  plaintively 
of  his  longing  for  a  year-round  supply  of  marine  eggs  and  embryos,  so  that  research 
could  be  undertaken  on  materials  selected,  not  for  their  ease  or  convenience,  but 
for  their  importance. 

My  own  teacher,  B.  H.  Willier,  who  was  a  student  of  Frank  Lillie,  (possibly 
Lillie's  most  successful  student)  spent  several  summers  in  Woods  Hole,  and  was  in 
fact  a  Trustee  of  the  Marine  Biological  Laboratory.  In  the  early  1960s,  when  I  was 
considering  an  invitation  to  head  the  Laboratory's  Embryology  course,  Willier 
advised  me  to  accept — but  with  the  caveat  that  I  must  not  allow  my  summers  at 


EARLY   STUDIES  OF  CELL  INTERACTIONS  83 

the  MBL  to  dilute  my  ongoing  year-round  research  program.  He  went  on  to  tell  us 
(Clement  Markert  and  me)  that  he  was,  in  fact,  repeating  Frank  Lillie's  advice  to 
him.  According  to  Willier,  Lillie,  in  his  later  years,  often  wondered  aloud  about  the 
shortcomings  of  the  MBL  as  a  center  for  research  (in  contrast  to  "intellectual 
rejuvenation")  and  especially  about  the  decisions  on  research  direction  imposed  by 
the  summer  season,  which  tended  to  favor  descriptive  over  experimental  studies. 
Lillie  was  said  to  have  observed,  "Willier,  that's  why  there  are  no  Harrisons  at  the 
Laboratory." 

Now,  let  me  return  to  cell  interactions.  Our  trail  takes  us  largely  to  Europe, 
although  as  we  have  already  observed,  the  contribution  of  Warren  Lewis,  a 
contemporary  and  colleague  of  Ross  G.  Harrison  in  Baltimore,  was  pivotal  in 
the  field. 

We  pick  up  the  trail  at  Spemann's  laboratory  at  the  turn  of  the  century,  as 
revealed  in  articles  he  published  in  1901(b),  1902,  and  1903.  There  can  be  little 
doubt  that  these  observations  had  their  origin  in  comparative  studies  typical  of  their 
time.  Spemann  was  examining  the  potencies  of  isolated  blastomeres  of  Triton. 
When  can  restriction  of  prospective  potency  to  prospective  value  be  detected? 

The  method  used  in  these  experiments  was  constriction  of  eggs  or  embryos  with 
a  fine  hair  loop,  as  devised  by  O.  Hertwig.  This  technique  makes  possible  complete 
separation  of  the  first  two  blastomeres,  halves  of  blastulae  and  gastrulae.  [A  slight 
modification  of  the  technique,  in  which  a  small  cytoplasmic  "bridge"  is  left,  was 
used  in  Spemann's  later,  also  classic  studies  of  twinning  combined  with  delayed 
nucleation  (Spemann,  1914,  1918;  see  Weiss,  1939.).] 

In  one-third  to  one-fourth  of  the  embryos  studied  the  first  cleavage  coincides 
with  the  axis  of  symmetry  of  the  gastrula.  However,  in  the  majority  of  embryos, 
the  second  cleavage  coincides  with  this  plane  of  symmetry. 

Development  subsequent  to  the  application  of  firm  constriction  in  the  plane  of 
the  first  cleavage  produces  two  kinds  of  results:  ( 1 )  one  blastomere  produces  a 
normal  embryo  and  the  other  forms  a  "Bauchstuck"  and  (2)  each  of  the  two 
blastomeres  produces  a  normal  embryo.  Subsequent  observations  revealed  that  in 
the  former,  the  blastopore  formed  in  and  was  restricted  to  the  descendants  of  one 
blastomere  only.  (Spemann  referred  to  this  as  the  "dorsal"  blastomere  and  to  its 
product  as  the  "dorsal-embryo-half.")  This  dorsal-embryo-half  develops  into  a 
normal  embryo,  the  ventral-embryo-half  forming  the  Bauchstiick. 

When  two  embryos  were  formed,  the  lip  of  the  blastopore  was  bisected  by  the 
ligature.  In  this  case  Spemann  assumed  that  the  constriction  was  median  and 
complete.  (With  "incomplete"  constriction,  double-headed  monsters  are  produced.) 

Spemann  concluded  that  the  first  two  blastomeres  are  of  equal  potency  only 
when  they  contain  equal  portions  of  the  blastopore. 

Eggs  constricted  in  the  blastula  stage  gave  similar  results  leading  him  to  conclude 
that  even  before  the  appearance  of  the  blastopore,  an  area  is  present  in  the  egg  the 
equal  division  of  which  imparts  equal  potencies  to  both  halves  of  the  blastulae. 

Thus,  early  in  the  first  decade  of  the  new  century,  Spemann  took  the  first  crucial 
step  toward  identifying  the  need  for  cell-cell  interaction  in  the  developing  amphibian 
embryo. 

Nor  should  we  lose  sight  of  the  fact  that  Spemann  himself,  and  Lewis  in 
Baltimore,  were,  in  those  very  years,  establishing  the  need  for  "induction"  in  the 
development  of  the  lens. 

There  followed  a  series  of  experiments  using  both  isolation  and  recombination 
(transplantation)  techniques  in  which  Spemann  (1918)  analyzed  the  process  of 
progressive  determination  in  the  urodele  gastrula.  The  experiments  carried  out  are. 


84  J.   D.   EBERT 

in  principle,  as  follows:  pieces  were  removed  from  stated  areas  of  two  Triton 
gastrulae,  and  reciprocally  transplanted;  and  the  subsequent  development  of  the 
transplant  was  followed.  All  changes  were  homoplastic.  The  results  may  be  sum- 
marized as  follows: 

I.  Exchange   of  pieces   from   presumptive    medullary   plate   and   presumptive 
epidermis: 

A.  Early  gastrulae:  presumptive  epidermis,  when  transplanted  to  the  future 
medullary   plate  area,   develops  into  brain;   presumptive   medullary   plate,   when 
transplanted  to  the   future  epidermal   region,   becomes  epidermis — development 
according  to  position. 

B.  Late  gastrulae:  presumptive  epidermis,  when  transplanted  to  the  future 
medullary  plate  area,  develops  into  epidermis;  presumptive  medullary  plate,  in 
future   epidermal    region,   becomes   medullary   plate — development   according   to 
origin. 

C.  Exchanges  between  one  early  and  one  yolk-plug  stage  gastrula:  regardless 
of  whether  the  "older"  tissue  transplanted  was  presumptive  medullary  plate  or 
epidermis,  development  was  according  to  position.  However  in  every  case  the  older 
tissue  retains  its  "advantge"  for  several  days. 

II.  Exchange  of  presumptive  epidermis  and  the  region  immediately  dorsal  to 
the  blastopore  (early  gastrulae):  presumptive  epidermis,  transplanted  to  the  area  of 
the  dorsal  lip,   produces  brain;  the  region  just  dorsal   to  the  blastopore,   when 
transplanted  to  the  future  epidermal  region,  "becomes  brain  and  notochord." 

In  1918  Spemann  concluded  that  the  region  above  the  blastopore  is  presumptive 
medullary  plate;  both  brain  and  notochord  develop  from  donor  tissue;  therefore  the 
region  above  the  blastopore  is  fixed  in  its  fate  (medullary  plate)  earlier  than  the  rest 
of  the  medullary  plate  region. 

This  conclusion  was  surely  incorrect,  yet  Spemann  himself  did  not  (in  1918) 
reveal  serious  concern  about  it  in  print.  Nevertheless,  he  moved  immediately  to  try 
to  confirm  and  extend  the  1918  study,  repeating  the  experiments,  but  using 
heteroplastic  grafts  between  two  differently  pigmented  species  of  Triton.  The 
findings,  published  in  1921,  differed  from  those  reported  earlier,  but  not  conclusively. 
Spemann  again  observed  that  the  region  of  the  blastopore  became  determined 
earlier  than  regions  more  distant  from  it  and  that  there  appeared  to  be  an  additional 
"organizing  action"  of  the  dorsal  lip  region  when  transplanted  to  another  embryo. 
These  suggestive  findings  set  the  stage  for  the  definitive  experiments  by  Spemann 
and  Mangold  (1924)  which  proved  that  a  piece  of  the  dorsal  lip  of  the  blastopore, 
in  the  process  of  gastrulation,  exerts  an  organizational  effect  upon  surrounding 
tissues  in  such  a  manner  that  it  causes  a  secondary  embryo  to  be  formed  if 
implanted  in  an  indifferent  place  on  another  embryo.  The  grafted  piece  of  the 
blastopore  was  therefore  designated  as  an  "organizer." 

Following  the  discovery  of  the  organizing  power  of  the  dorsal  lip,  and  the 
demonstration  by  Spemann  and  Hilde  Mangold  that  this  capacity  was  not  specific 
between  closely  related  species,  Spemann  wondered  how  far  this  non-specificity 
might  extend.  At  his  suggestion  Geinitz  (1925),  using  both  Spemann's  transplantation 
method  and  O.  Mangold's  "Einsteck"  technique,  tested  less  closely  related  species. 

Results  of  transplantation  of  "organizers"  from  Pleurodeles  waltli,  Amblystoma 
mexicanum,  Rana  temporaria,  R.  esculenta,  and  Bombinator  pachypus  to  Triton 
taeniatus,  T.  cristatus,  or  T.  alpestris  hosts  showed  that  the  action  of  the  organizer 
is  non-specific  between  closely  related  genera,  between  families,  and  even  between 
different  orders  (or  sub-classes).  The  most  successful  case  of  the  last  mentioned  type 


EARLY   STUDIES  OF  CELL  INTERACTIONS  85 

is  described  in  considerable  detail.  Using  Spemann's  method,  a  piece  of  Bombinator 
"organizer"  was  placed  in  the  ventral  side  of  a  young  T.  tacniatus  gastrula.  It 
became  invaginated  near  the  host  blastopore  and  eventually  gave  rise  to  a  notochord, 
undifferentiated  mesoderm,  and  somites.  In  addition  it  induced  from  the  host  tissue 
somites  and  a  neural  tube.  Several  similar  but  less  striking  cases  were  also  mentioned. 
Transplants  from  Bufo  vulgaris  and  Hyla  arborea  to  T.  taenialus  were  unsuccessful. 
In  the  case  of  transplantation  in  the  reverse  direction  (urodele  to  anuran)  no 
induction  occurred. 

A  complementary  series  of  experiments  dealt  with  the  capacity  of  grafted 
Bombinator  presumptive  epidermis  to  be  influenced  by  its  position  in  the  host. 
When  stuck  into  the  blastocoel  it  remained  undifferentiated;  if  implanted  in  the 
surface  of  the  T.  taeniatus  host  it  developed  according  to  position,  becoming 
epidermis  or  being  invaginated  and  subsequently  becoming  mesoderm  in  the  cases 
reported. 

Geinitz  observed  that  the  more  distant  the  phylogenetic  relationship  between 
graft  and  host,  the  less  the  tendency  of  the  cells  to  mutually  contribute  to  organs. 
He  suggested  that  the  earlier  the  stages  used,  the  more  successful  the  grafts,  possibly 
because  these  may  be  antecedent  to  processes  chemically  differentiating  the  species. 
Most  important,  these  experiments  illustrate  the  lack  of  species-specificity  of  the 
induction  process. 

Geinitz's  contribution  was  but  the  first  in  the  long  series  of  studies  of  the  non- 
specificity  of  the  induction  process,  further  demonstrated  in  the  series  of  articles  by 
Holtfreter,  of  which  his  1935  article  is  fully  representative. 

The  discovery  of  the  "organizer"  and  the  revelation  that  it  was  not  species 
specific  set  in  train  fifteen  years  of  intensive  and  largely  fruitless  research  on  its 
chemical  nature,  by  Spemann  himself,  especially  in  collaboration  with  Fischer  and 
Wehmeier  (1933),  by  Joseph  Needham  and  C.  H.  Waddington,  and  their  colleagues, 
by  the  leading  Dutch  investigators,  especially  Woerdeman  and  Raven  and  by  E.  J. 
Boell  and  collaborators.  As  described  by  Brachet  (1950),  at  various  times  attention 
was  focused  on  glycogens,  fatty  acids,  sterols,  and  nucleoproteins.  Fischer  and 
colleagues,  influenced  by  Holtfreter,  finally  concluded  that  no  specific  chemical,  or 
group  of  chemicals,  could  be  stated  to  be  the  organizer.  On  the  other  hand  Needham, 
Waddington,  and  associates  postulated  that  the  active  "evocator"  is  a  sterol  type 
substance,  normally  bound  in  a  protein-glycogen-sterol  complex  and  released 
normally  during  gastrulation,  or  under  experimental  conditions  by  cytolyzing  agents. 

It  was  Holtfreter,  and  especially  Lester  Barth,  whose  observations  brought  the 
"quest  for  the  organizer"  to  a  halt,  just  as  World  War  II  clouded  the  horizon.  The 
MBL  provided  a  platform  in  1939  for  Barth  to  first  present  his  observations, 
subsequently  published  in  1941  as  "Neural  differentiation  without  organizer." 
Earth's  discovery,  that  the  course  of  differentiation  of  small  aggregates  of  cells 
prepared  from  explants  of  ventral  ectoderm  of  frog  gastrulae  depends  upon  the 
composition  of  the  solution  in  which  they  are  cultured  lent  weight  to  Holtfreter1  s 
ideas  on  the  release  of  effectors  by  "mild  and  reversible  cytolysis." 

Following  World  War  II  the  "quest  for  the  organizer"  continued,  turning  for  a 
time  to  the  nucleic  acids,  especially  the  ribonucleic  acids  as  effectors  (Brachet,  1950). 
While,  in  principle,  cell-cell  interaction  involving  the  exchange  of  "information"  at 
the  point  of  impact  by  the  exchange  of  vesicular  material  or  by  cytoplasmic  bridges 
cannot  be  excluded,  attention  is  now  centered  on  the  exchange  of  interactants  via 
cytoplasmic  bridges,  specialized  junctions  or  other  mechanisms,  and  on  signals 
generated  at  the  membrane,  transmitted  via  specific  receptor  sites  and  intracellular 
mediators  like  Ca++  and  cyclic  AMP  to  specific  genes. 


86  J.   D.   EBERT 

Indeed,  the  works  of  Barth  presaged  a  quiet  revolution  that  emphasized  the 
possible  importance  of  internal  release  and  redistribution  of  inorganic  ions,  leading 
Barth  and  Barth  to  a  series  of  contributions  (1959-1974)  in  which  they  attempted 
to  formulate  a  general  theory  of  ionic  regulation  of  normal  embryonic  induction. 

But  immediately  after  World  War  II,  the  revolution  was  quiet  indeed,  as  students 
of  embryology  were  swept  up  in  the  wave  of  extraordinary  advances  generated  by 
the  one  gene-one  enzyme  hypothesis,  leading  to  the  emergence  of  nucleic  acid 
chemistry  and  molecular  genetics  as  focal  fields  of  research. 

CELL  SURFACE  INTERACTIONS  IN  MORPHOGENESIS 

I  have  already  remarked  that  the  first  decade  of  the  twentieth  century  saw 
embryology  come  of  age  as  an  experimental  science.  Indeed,  it  was  probably 
embryology's  first  "golden  age,"  with  advocates  of  both  cell  lineage  and  Entwick- 
lungsmechanik  contributing  one  discovery  after  another.  The  decade  was  capped  by 
what  Michael  Abercrombie  called  "an  astonishing  stride  forward  in  the  history  of 
biology,"  Ross  G.  Harrison's  (1907,  1910)  introduction  of  the  technique  of  tissue 
culture  in  the  study  of  nerve  outgrowth.  Less  heralded  at  the  time  and  less 
provocative  in  immediately  opening  up  new  vistas  of  experiments  to  be  done  were 
the  observations  of  Harrison's  contemporary,  H.  V.  Wilson  (1907,  see  Morgan, 
1927),  on  the  species  specific  aggregation  of  dissociated  sponge  cells.  Neither  Wilson's 
pioneering  observations,  nor  the  later  extension  of  the  work  by  Galtsoff,  had  a 
significant  impact  on  concepts  of  morphogenesis  for  three  decades.  Nor  did  the 
contributions  of  Herbst  at  the  turn  of  the  century  in  dissociating  embryos  in 
calcium-free  sea  water  contribute  significantly  in  the  conceptual  sense.  Indeed,  these 
earlier  observations  came  to  the  fore  only  after  Holtfreter  had  published  his  classic 
article  "Tissue  affinity,  a  means  of  embryonic  morphogenesis"  in  which  he  developed 
the  concept  of  selective  affinities  between  embryonic  cells  and  tissues.  He  showed 
that  when  tissues  of  amphibian  embryos  are  exposed  to  solutions  of  high  pH  they 
dissociate.  Upon  being  returned  to  saline  at  a  physiological  pH,  they  reconstruct 
the  tissue  of  origin.  Only  then  did  the  concepts  of  adhesive  selectivity  in  cell 
interactions,  of  "cell  linking  macromolecules"  and  "cell  adhesion  molecules" 
gradually  emerge. 

LITERATURE   CITED 

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BARTH,  L.  G.  1966.  The  role  of  sodium  chloride  in  sequential  induction  of  the  presumptive  epidermis  of 

Rana  pipiens  gastrulae.  Biol.  Bull.  131:  415-426. 
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unconditioned  medium.  J  Embryo/.  Exp.  Morphol.  7:  210-222. 
BARTH,  L.  G.,  AND  L.  J.  BARTH.  1962.  Further  investigations  of  the  differentiation  in  vitro  of  presumptive 

epidermis  cells  of  the  Rana  pipiens  gastrula.  J.  Morphol.  110:  347-373. 
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HOLTFRETER,  J.  1939.  Tissue  affinity,  a  means  of  embryonic  morphogenesis.  Pp.  186-225  reprinted  in 
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Org.  48:  533-570. 

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Reference:  Biol.  Hull.  168  (suppl.):  88-98.  (June,  1985) 


AN   EVOLUTIONARY   CENTURY   AT  WOODS   HOLE:   INSTRUCTION 

IN   INVERTEBRATE  ZOOLOGY 

W.   D.   RUSSELL-HUNTER 

Marine  Biological  Laboratory.  Woods  Hole,  Massachusetts,  02543,  and  Department  of  Biology, 
Syracuse  University,  Syracuse,  New  York,  13210 

ABSTRACT 

Whitman,  Lillie,  and  their  successors  always  regarded  research  and  instruction 
as  complementary  functions  for  the  Marine  Biological  Laboratory.  An  invertebrate 
zoology  course  was  taught  each  summer  at  Woods  Hole  for  90  years.  It  is  suggested 
that  the  strengths  of  this  course  and  its  capacity  to  evolve  came  from  the  re-sorting 
of  eight  or  nine  instructors  with  diverse  research  interests  every  five  years  or  less, 
within  the  constraints  of  a  stable  and  highly  structured  instructional  environment. 
A  chronological  summary  of  instructional  staffs  is  followed  by  a  brief  survey  of  the 
invertebrate  materials  covered  and  the  conceptual  approaches  used  in  instruction. 
The  course  was  never  a  comprehensive  systematic  coverage  of  all  marine  invertebrates, 
while  the  teaching  laboratory  never  provided  the  best  conditions  for  teaching 
comparative  anatomy,  but  was  a  much  better  place  to  investigate  feeding  mechanisms, 
locomotory  patterns,  or  reproductive  behavior  in  a  reductionist  fashion.  Each  group 
of  eight  or  nine  instructors,  attracted  by  opportunities  for  personal  research,  almost 
inevitably  included  both  mechanistic-physiologists  and  population-naturalists  and 
the  dialectic  which  resulted  had  both  educational  and  research-generating  value. 
The  corporate  body  of  active  investigators  at  the  MBL  was  sustained  in  part  by  the 
conditions  of  recruitment  for  both  course  instructors  and  post-course  research 
students  in  invertebrate  zoology. 


INTRODUCTION 

An  invertebrate  zoology  course  was  taught  each  summer  at  Woods  Hole  for  90 
years,  for  81  of  them  in  the  same  teaching  laboratory  on  the  lower  floor  of  the 
south  wing  of  Old  Main.  However,  this  was  far  from  representing  an  evolutionary 
stasis.  To  a  physiological  ecologist  turned  Whig  amateur  recorder  of  its  history, 
there  was  a  continuing  dynamic  equilibrium.  Doubtless  professional  historians  given 
to  the  dialectic  method  (including  Allen,  1979,  1981)  would  detect  a  long  sustained 
struggle  between  the  phyletic  and  the  experimental  students  of  the  invertebrates, 
while  a  Namierian  historian  could  readily  compile  proportionate  statistics  of  the 
research  publications  and  institutional  connections  of  the  instructors  and  plot  the 
switches  (sometimes  acute,  sometimes  dampened)  of  interest  in  matters  evolutionary. 
In  this  brief  survey  I  shall  try  to  go  beyond  my  natural  Whig  celebration  or 
encomium,  and  utilize  a  little  from  such  alternative  approaches.  Two  preliminary 
matters  concern  the  mechanics  of  staffing  the  course  and  the  logistics  of  its 
instructional  laboratory.  Both  stem  in  part  from  the  course's  origins  and  antedate 
the  establishment  of  it  and  of  the  Marine  Biological  Laboratory  in  1888. 


88 


INVERTEBRATE  INSTRUCTION  AT  WOODS  HOLE  89 

HISTORY 

Staffing  and  laboratory  logistics 

"Whitman  regarded  research  and  instruction  as  co-ordinate  functions  with  a 
single  aim,"  noted  Lillie  (1944).  Instruction  at  Penikese  in  1873  and  1874,  and  at 
Annisquam  in  1881-1886  (see  Lillie,  1944;  Maienschein,  1985)  provided  models 
for  the  course  in  marine  invertebrate  zoology  which  began  with  the  founding  of  the 
MBL  in  1888.  Both  Whitman  and  Lillie  noted  that  instructors  from  a  variety  of 
home  institutions  could  be  recruited  for  each  summer  by  the  incentive  of  opportunities 
for  research.  In  later  years,  in  addition  to  a  laboratory  with  healthy  marine  organisms 
and  running  sea  water,  these  opportunities  included  an  open  and  magnificent 
research  library,  contacts  with  an  international  group  of  peers,  and  (in  some  periods) 
access  to  technical  equipment  not  available  in  many  colleges.  One  result  of  this 
mode  of  recruiting  noted  in  a  later  report  on  educational  policy  (Buck  et  a/.,  1963) 
was  that  it  has  always  been  impossible  to  staff  the  invertebrate  zoology  course  with 
a  set  of  specialists  in  each  group  of  marine  invertebrates  because  it  has  always  been 
more  important  to  insist  that  all  instructors  show  research  distinction  in  experimental 
science. 

Two  other  aspects  of  staffing  are  important.  First,  the  content  and  conduct  of 
the  courses  have  always  been  the  exclusive  responsibility  of  the  course  instructors. 
The  Director  and  the  standing  committee  on  instruction  become  directly  involved 
only  with  the  selection  of  each  successive  instructor-in-charge  and  with  no  other 
aspect  of  any  ongoing  instructional  program.  Secondly,  we  have  the  practice,  which 
began  early  and  soon  became  formalized  for  the  invertebrate,  embryology,  and 
physiology  courses,  that  the  term  as  instructor-in-charge  of  an  established  course 
should  not  exceed  five  years. 

Some  other  constraints  arise  from  the  fact  noted  by  Lillie  that  the  students  in 
invertebrate  zoology  included  undergraduates  and  were  generally  the  youngest  group 
in  the  summer  institution.  In  fact,  academic  and  calendar  ages  varied  greatly,  with 
one  or  two  holders  of  Ph.D.  or  M.D.  degrees  in  most  classes.  Over  at  least  six  of 
the  decades,  a  majority  of  each  class  consisted  of  seniors  and  beginning  graduate 
students  (a  fact  which  may  confuse  historians  using  our  archives  since  many 
invertebrate  course  students  would  have  different  home  institutions  listed  at  spring 
acceptance  and  at  completion  of  the  summer  course).  The  number  of  students  was 
also  larger  than  in  the  other  formal  courses,  occasionally  being  over  50:  in  later 
years  it  was  usually  35  to  45,  from  whom  10-12  would  be  selected  for  post-course 
research.  In  turn  this  dictated  a  larger  number  of  instructors  than  in  the  other 
formal  courses,  usually  eight,  but  ranging  from  seven  to  ten. 

Another  determinant  of  the  evolution  of  invertebrate  instruction  at  the  MBL 
came  from  the  architecture  and  furnishings  of  the  teaching  laboratory.  Whitman 
and  Lillie  believed  that  students  should  give  their  full  time  during  the  six  weeks  or 
so  of  a  formal  summer  course.  Each  student  was  thus  to  be  assigned  a  private  work 
space,  which  could  be  occupied  at  any  time  of  the  day  or  night.  For  8 1  years,  right 
up  to  the  last  summer  (1968)  before  the  opening  of  the  Loeb  laboratories,  this  was 
in  the  ground-floor  laboratory  of  the  south,  or  original,  wing  of  the  Old  Main 
building  and  each  individual  space  was  actually  a  quarter  of  a  large,  wide  workbench 
like  a  "Naples"  table  (Groeben,  1975)  with  two  students  facing  two  others.  For 
most  of  the  period,  a  long  suite  of  sea-tables  was  placed  down  the  middle  of  the 
laboratory  between  seven  workbenches  on  each  side,  and  these  housed  the  students' 
study  materials,  the  supply  of  healthy  marine  invertebrates. 


90  W.   D.   RUSSELL-HUNTER 

The  theme  of  this  paper  is  one  of  structural  continuity  underlying  functional 
change.  By  most  appropriate  measures,  the  MBL's  invertebrate  course  made  a 
major  contribution  to  evolutionary  biology  during  the  last  century.  My  thesis  is  that 
the  strengths  of  this  contribution  came  largely  from  the  resorting  of  eight  or  nine 
instructors  every  five  years  or  less  (like  chiasma-mandated  recombination)  within 
the  constraints  of  a  stable  and  highly  structured  instructional  environment  (with  all 
the  faunal  diversity  that  can  imply). 

Instructors  and  invertebrates 

It  is  possible  to  summarize,  chronologically  and  briefly,  the  instructional  staffing 
and  organization  of  the  course  and  then,  systematically  but  even  more  briefly,  the 
kinds  of  invertebrate  material  covered.  In  parts,  these  summaries  will  be  obviously 
eclectic. 

In  the  beginning  most  of  the  principal  investigators  (and  founding  fathers)  also 
took  part  in  instruction.  In  the  years  1888  through  1895,  summer  students  of 
zoology  heard  more  than  merely  "highlight"  lectures  from  E.  G.  Conklin,  Cornelia 
M.  Clapp,  and  C.  O.  Whitman  himself.  In  the  first  year,  B.  H.  Van  Vleck  who  had 
assisted  Alpheus  Hyatt  at  Annisquam  was  listed  as  instructor.  By  1896-1897,  two 
younger  instructors  taking  part  were  C.  M.  Child  and  F.  R.  Lillie.  It  may  be 
significant  that  by  the  tenth  session  (1897),  Whitman  was  referring  to  a  Department 
of  Investigation  in  Zoology  and  a  Department  of  Instruction  in  Zoology,  with  the 
only  three  other  individuals  listed  under  "Officers  of  Instruction"  forming  a 
Department  of  Botany.  A  year  later  (1898),  we  find  Zoology  [printers  or  directors/ 
editors  seemed  to  use  the  diaeresis  in  alternate  years]  divided  into  departments  of 
Investigation,  Embryology  (F.  R.  Lillie,  Head  Instructor),  and  Anatomy  (James  I. 
Peck,  Head  Instructor),  with  separate  departments  of  Physiology  (Jacques  Loeb, 
Head  Instructor)  and  of  Botany  (Bradley  M.  Davis,  Head  Instructor).  In  other 
printed  announcements  for  1898,  the  "Anatomy"  course  was  described  (more 
correctly  in  present  usage)  as  a  course  on  Morphology  of  Marine  Invertebrates,  and 
we  can  note  that  the  students  received  additional  lectures  from  Whitman,  E.  G. 
Gardiner,  and  V.  L.  Kellogg.  [With  the  subsequent  separation  of  the  Embryology 
course  from  the  Zoology  group,  the  traditional  four  MBL  courses — Botany,  Inver- 
tebrate Zoology,  Embryology,  and  Physiology — had  been  established  by  the  beginning 
of  the  20th  century.] 

In  1897,  James  I.  Peck  of  Williams  College,  who  was  also  acting  as  Assistant 
Director  to  Whitman,  was  instructor-in-charge  of  invertebrate  zoology,  and  recorded 
staff  discussion  about  course  content  (see  below).  He  was  succeeded  in  1901  by 
Gilman  A.  Drew  of  the  University  of  Maine,  who  subsequently  became  instructor- 
in-charge  of  the  embryology  course  in  1908,  and  then  continued  as  Assistant 
Director  under  Lillie  for  many  years  thereafter.  Drew's  last  summer  heading  the 
invertebrate  course  was  1907,  when  assisting  him  as  a  regular  instructor  was  Otto 
C.  Glaser;  the  class  heard  lectures  from  Morgan,  Conklin,  Lillie,  E.  B.  Wilson,  and 
possibly  Whitman.  A  student  in  the  course  that  summer  from  Syracuse  University 
was  Charles  Packard,  and  another  student  from  Columbia  University  was  Ethel  N. 
Browne. 

In  invertebrate  zoology,  Drew  was  succeeded  by  Winterton  C.  Curtis,  then  of 
the  University  of  Missouri  (but  see  below),  who  headed  the  course  1907-191 1.  By 
1916-1918,  Caswell  Grave,  W.  C.  Allee,  W.  H.  Taliaferro,  and  T.  W.  Painter  were 
now  instructors  in  the  course,  and  in  1918-1919,  Libbie  H.  Hyman  was  doing  work 
on  invertebrate  respiration  under  Child's  direction,  but  stemming  directly  from 


INVERTEBRATE   INSTRUCTION   AT  WOODS   HOLE  91 

Jacques  LoetTs  earlier  investigations.  Allee  was  in  charge  in  1919-1920  when  a 
separate  course  on  Protozoology  under  Gary  N.  Calkins  was  set  up.  By  1925,  B.  H. 
Willier  and  H.  B.  Baker  were  instructors  and  by  1928,  B.  H.  Grave  and  Elbert  C. 
Cole.  The  last  fourteen  instructors  named  include  both  experimentalists  and 
naturalists.  Cole  was  instructor-in-charge  in  1932-1936,  and  was  succeeded  for 
1937-1941  by  T.  H.  Bissonette,  under  whom  P.  S.  Crowell  and  J.  S.  Rankin  were 
instructors. 

Subsequent  instructors-in-charge  were:  A.  J.  Waterman  of  Williams  College 
( 1 942),  John  B.  Buck  ( 1 943- 1 944),  F.  A.  Brown  Jr.  ( 1 945- 1 949),  Lewis  H.  Kleinholz 
(1950-1954),  Theodore  H.  Bullock  (1955-1957),  Grover  C.  Stephens  (1958-1960), 
Clark  P.  Read  (1961-1963),  W.  D.  Russell-Hunter  (1964-1968),  James  F.  Case 
(1969-1971),  Robert  K.  Josephson  (1972-1974),  and  Michael  J.  Greenberg  (1975- 
1977).  The  last  ten  named  could  all  be  described  as  comparative  physiologists,  and 
broadly  the  majority  of  their  staffs  could  also  be  so  described,  although  in  fact  they 
ranged  in  their  own  research  activities  from  comparative  biochemistry  and  biophysics 
through  functional  morphology  to  the  physiological  ecology  of  invertebrates,  with 
only  a  few  being  additionally  systematists.  A  few  staff  lists  can  illustrate  this:  for 
1947  we  have  F.  A.  Brown,  Jr.  (in  charge),  W.  D.  Burbanck,  C.  G.  Goodchild,  John 
H.  Lochhead,  Madelene  E.  Pierce,  W.  M.  Reid,  Mary  D.  Rogick,  and  Talbot  H. 
Waterman;  for  1954:  Lewis  H.  Kleinholz  (in  charge),  John  H.  Lochhead,  Norman 
A.  Meinkoth,  Grover  C.  Stephens,  John  M.  Anderson,  Muriel  Sandeen,  L.  M. 
Passano,  and  Morris  Rockstein;  for  1967:  W.  D.  Russell-Hunter  (in  charge),  George 
G.  Holz,  Jr.,  Norman  Millott,  Eric  L.  Mills,  James  F.  Case,  Frank  M.  Fisher,  Jr., 
Robert  K.  Josephson,  Jonathan  P.  Green,  Meredith  L.  Jones,  and  Hugh  Y.  Elder; 
and  for  1971:  James  F.  Case  (in  charge).  Garth  Chapman,  Alan  Gelperin,  David  C. 
Grant,  Michael  J.  Greenberg,  Joseph  B.  Jennings,  Charlotte  P.  Mangum,  James  G. 
Morin,  and  Dorothy  M.  Skinner.  In  the  period  1969-1977,  the  course  was  called 
Experimental  Invertebrate  Zoology,  perhaps  echoing  a  period  in  the  thirties  when 
the  embryology  course  was  named  Experimental  Embryology.  In  1978,  the  inver- 
tebrate course  was  replaced  by  one  entitled  Neural  Systems  and  Behavior  under  the 
direction  of  Alan  Gelperin  and  subsequently  of  Ronald  R.  Hoy. 

Potential  material  on  invertebrates  is  enormous.  Over  the  9 1  years,  the  invertebrate 
zoology  course  attempted  to  increase  understanding  of  the  biology  of  > 300,000 
diverse  species  classified  in  32  animal  phyla  (including  two  "new"  phyla  Pogonophora 
and  Gnathostomulida  which  were  added  later  in  the  20th  century,  and  several 
redefinitions  during  the  period).  For  most  of  the  nine  decades,  Protozoa  were  not 
included  (except,  for  example,  from  1963  to  1969),  and  most  aspects  of  the  biology 
of  insects  and  of  other  land  arthropods  were  deliberately  excluded.  The  material 
was  generally  made  more  manageable  by  placing  greater  emphasis  on  the  nine  or 
so  "major"  phyla,  designated  as  such,  not  merely  as  encompassing  larger  numbers 
of  species  or  of  individuals,  but  also  in  recent  years  by  quantifiable  ecological 
measures.  To  illustrate  these,  of  the  solar  energy  incorporated  into  green  plants,  a 
disproportionately  large  share  flows  through  representatives  of  such  major  phyla  as 
the  Arthropoda  and  the  Mollusca,  whereas  the  energy  flow  through  representatives 
of  the  minor  phylum  Entoprocta  in  any  ecosystem  is  normally  several  orders  of 
magnitude  smaller.  For  the  77  years  from  Drew  to  Greenberg  at  least,  the  biology 
of  Cnidaria,  Annelida,  Arthropoda,  Mollusca,  and  Echinodermata  always  received 
considerable  attention,  with  the  extent  of  work  on  sponges,  flatworms,  nematodes, 
and  invertebrate  chordates  varying  from  one  instructional  group  to  another.  The 
other  twenty-two  minor  phyla  were  usually  neglected — the  course  was  never  a 
comprehensive  systematic  coverage  of  all  kinds  of  marine  invertebrates.  Certain 


92  W.   D.   RUSSELL-HUNTER 

naturalists  among  the  course  staffs  were  always  dissatisfied  by  this  (see  below)  as,  in 
the  early  decades,  were  certain  institutions  which  financially  supported  students  for 
the  invertebrate  course. 

Although  this  strategy  was  consistent  for  9 1  years,  staff  debate  on  the  tactics  of 
appropriate  neglect  continued  for  at  least  70  of  them.  In  preparing  to  write  this 
paper,  I  was  surprised  to  read  about  a  staff  (and  student)  discussion  (Peck,  1896) 
centering  on  "rapidly  going  over  many  forms,  versus  doing  a  few  forms  more 
thoroughly"  reported  for  4  July  1896  by  James  I.  Peck,  a  discussion  paralleled  in 
1966  by  my  course  colleagues.  We  concluded,  as  Peck's  associates  apparently  did, 
that  with  a  heterogeneous  group  of  students,  a  diversity  of  material  should  be 
available,  but  that  levels  of  investigation  were  better  paced  individually  by  each 
student's  needs.  Of  course,  we  were  only  echoing  the  training  policy  established 
even  earlier  for  the  Annisquam  Laboratory  by  Alpheus  Hyatt.  Concentration  upon 
"representative  types"  of  invertebrates,  an  educational  legacy  of  mixed  value  derived 
from  Louis  Agassiz,  ebbed  and  flowed  in  the  invertebrate  course,  reaching  a  later 
apogee  in  1945-1949,  and  being  as  deliberately  eschewed  in  1961-1974. 

This  somewhat  arbitary  limitation  of  invertebrate  groups  to  be  studied  was 
paralleled  by  some  restriction  of  conceptual  approach.  Again  we  find  considerable 
continuity  over  the  91  years  in  what  could  be  a  dynamic  synthesis  continually 
refashioned.  The  treatment  of  the  invertebrates  presented  to  most  students  was 
intermediate  in  regard  to  both  grade  of  biological  organization  and  level  of  concept; 
that  is,  it  was  concerned  with  whole  animals  considered  mainly  at  the  mechanistic- 
physiological  and  adaptive-functional  levels  of  explanation.  The  structural-descriptive 
level  was  rarely  emphasized,  and  the  evolutionary-historical  level  only  explored 
from  time  to  time.  It  is  possible  that  the  tissue  and  cell  grades  of  organization  were 
more  often  added  to  the  whole-animal  and  organ  studies  in  the  first  three  decades 
of  the  course,  and  that  population  and  community  interactions  were  considered 
more  frequently  after  1920,  but  whole-animal  investigations  remained  central. 
Research  themes  recurring  over  the  years,  although  with  increasing  sophistication 
of  study  techniques,  include  patterns  of  feeding,  mechanics  of  locomotion,  and 
reproductive  behavior. 

Value  of  MBL  courses 

The  mechanistic  functional  morphology  basic  to  this  conceptual  approach 
requires  surprisingly  little  instruction  in  formal  systematics,  and  little  had  ever  been 
given  in  the  Woods  Hole  invertebrate  courses.  In  the  last  years  of  the  Invertebrate 
Zoology  course  as  such,  the  argument  was  being  used  that  there  were  many  other 
courses  being  taught  elsewhere  in  the  United  States  which  dealt  with  the  systematics 
and  natural  history  of  marine  invertebrates  at  the  senior  undergraduate  level,  and 
therefore  it  was  unnecessary  to  do  this  at  the  MBL.  This  argument  ignored  the  fact 
that  many  of  these  courses  elsewhere  had  been  founded  by  "refugees"  who  believed, 
in  part  correctly,  that  these  kinds  of  studies  had  always  been  neglected  in  MBL 
courses.  I  am  sure  that  Drs.  John  M.  Anderson,  John  M.  Kingsbury,  John  S. 
Rankin,  and  Donald  J.  Zinn  would  claim  that  this  educational  mission  was  taken 
up  by  the  newer  institutions  with  which  they  were  connected  largely  because  it  had 
been  neglected  for  so  long  at  the  MBL.  Clearly,  this  interpretation  was  not  known 
to  other  internal  critics  concerned  with  other  training  programs  at  the  MBL.  I 
remember  arguing  with  the  late  Harry  Grundfest  in  the  middle  sixties  that  the  MBL 
needed  an  invertebrate  zoology  course  largely  because  American  biology  still  needed 
an  experimentalist  counter  to  such  natural  history  courses,  even  if  only  to  assure 
an  appropriate  diversity  of  marine  invertebrate  systems  to  be  used  in  future  studies 


INVERTEBRATE   INSTRUCTION   AT  WOODS   HOLE  93 

in  comparative  physiology  and  physiological  ecology.  In  making  any  comparisons 
between  the  MBL's  invertebrate  course  and  the  marine  invertebrate  courses  currently 
offered  at  the  Isles  of  Shoals,  at  Duke's  Beaufort  Laboratory,  at  Avery  Point,  at 
Lewes,  at  Pacific  Grove,  and  elsewhere,  I  would  make  an  exception  of  the  courses 
offered  by  the  Friday  Harbor  Laboratories  of  the  University  of  Washington,  which 
have  been  closely  similar  to  those  at  the  MBL,  and  have  had  a  similar  output  of 
investigators  and  publishable  investigations. 

Measures  of  the  contribution  of  MBL  courses,  including  invertebrate  zoology, 
to  both  evolutionary  and  physiological  biology  in  North  America  are  varied.  One, 
in  some  ways  similar  to  contemporary  use  of  citation  indices  by  sociologists  of 
science  and  by  college  administrators,  was  first  used  by  G.  A.  Drew  in  1923.  He 
tabulated  the  number  of  students  from  the  summers  of  1908  to  1917  inclusive 
whose  names  had  appeared  in  the  1922  edition  of  American  Men  of  Science.  By 
this  measure  18%  of  all  students  and  12%  of  those  from  the  invertebrate  courses 
were  regarded  as  successful  investigators  and  college-level  teachers.  A  larger  sample 
(1918-1931)  surveyed  by  Charles  Packard  in  1939  (Packard,  1940)  yielded  29.6% 
of  all  students  (49.1%  of  males)  and  25.4%  of  invertebrate  zoology  students  (47%  of 
males).  When,  for  federal  training  grant  applications  in  1964  and  1968,  I  surveyed 
the  invertebrate  classes  from  the  early  1950's  my  figures  were  somewhat  higher  than 
Packard's.  In  any  such  survey,  individual  names  may  have  consequence  beyond 
such  statistics.  As  a  single  example,  significant  contributions  to  our  understanding 
of  evolutionary  processes,  using  distinct  conceptual  approaches  and  widely  diverse 
invertebrate  data  bases,  have  been  made  by  four  students  from  this  period:  J.  O. 
Corliss,  M.  J.  West-Eberhard,  J.  S.  Farris,  and  L.  B.  Slobodkin.  Thus,  ultimate  as 
well  as  proximate  biological  causalities  have  been  profitably  explored  by  the 
invertebrate-trained  cohorts.  However,  speculative  phylogeny  has  never  been  prom- 
inent in  invertebrate  instruction  at  the  MBL.  At  least  from  Drew's  time  onward, 
the  instructors  concerned  with  evolutionary  (ultimate)  questions  or  the  neo-Darwin- 
ians,  although  usually  outnumbered  by  those  concerned  with  physiological  (proximate) 
questions  or  the  Loebians,  have  been  population-statistical  investigators  with  stochastic 
yet  testable  hypotheses  rather  than  idealist  morphologists. 

One  of  the  investigators  at  the  MBL  in  the  first  years  of  the  century  does 
exemplify  the  worst  aspects  of  the  speculative  schools  of  post-Haeckel  morphology. 
He  was  William  Patten  who  published  metabiological  reports  on  comparative 
anatomy  in  the  Journal  of  Morphology  and  elsewhere,  which  postulated  an  origin 
of  vertebrates  in  arachnids  like  Limiilm.  This  represents  a  comparatively  late 
derivative  of  the  Naturphilosophen  of  J.  W.  Goethe,  and  is  perhaps  closest  in 
concept  to  the  publications  of  Lorenz  Oken  in  the  first  half  of  the  nineteenth 
century.  Patten's  reports  can  be  contrasted  with  contemporary  MBL  work  by  Otto 
C.  Glaser  who  was  then  an  instructor  in  the  zoology  course.  In  the  Journal  of 
Experimental  Zoology  in  1907,  Glaser  published  a  paper  whose  title  seems  to 
anticipate  the  Neural  Systems  and  Behavior  course  under  Alan  Gelperin  which 
formally  replaced  the  invertebrate  zoology  course  in  1978.  It  was  entitled  "On 
movement  and  problem  solving  in  Ophiura" 

Instructional  material  from  the  invertebrate  course  continued  to  affect  more 
formal  pedagogy  elsewhere.  Many  textbooks  had  origins  at  Woods  Hole.  Among 
others,  Winterton  C.  Curtis,  Chauncey  G.  Goodchild,  Douglas  A.  Marsland,  and 
Libbie  H.  Hyman  were  involved  in  early  undergraduate  texts.  The  only  advanced 
invertebrate  text  in  English  remains  uncompleted  after  Libbie  H.  Hyman  prepared 
the  first  five  magnificent  volumes  (1940-1954),  each  as  a  real  pandect.  Further, 
Woods  Hole  instructional  work  stimulated  the  first  English  coverage  of  comparative 
animal  physiology  in  1961  by  C.  Ladd  Prosser  and  Frank  A.  Brown,  Jr.  Somewhat 


94  W.   D.   RUSSELL-HUNTER 

earlier,  the  course  instructors,  along  with  a  few  associates  under  the  editorship  of 
Frank  Brown,  had  produced  the  useful  volume,  Selected  Invertebrate  Types  (1950). 
This  was  a  worthy  successor  to  older  German  Prakticum,  as  a  summary  of 
comparative  invertebrate  morphology  for  a  restricted  series  of  type  invertebrates.  It 
emerged  in  conversation  with  some  of  the  authors,  that  they  did  not  regard  it  as 
representative  of  what  they  should  teach  at  Woods  Hole,  but  rather  as  background 
material  they  would  have  liked  all  students  to  have  known.  More  recently,  symbiotic 
instructional  activities  in  the  invertebrate  course  are  acknowledged  as  part  of  the 
origins  for  a  laboratory  guide  to  the  invertebrates  in  1970  by  I.  W.  Sherman  and 
V.  G.  Sherman,  for  two  of  my  paperbacks  (1968  and  1969),  and  for  my  larger 
textbook,  A  Life  of  Invertebrates  (1979).  Of  course,  primary  research  literature  was 
continually  produced.  Over  the  last  30  years  of  the  course,  5-9  research  papers 
resulted  each  year  from  the  summer  investigations  of  instructors  and  postcourse 
students.  In  general,  many  distinguished  investigators,  many  new  research  topics, 
and  a  wide  variety  of  publications  were  produced  for  American  science  by  the 
invertebrate  zoology  course  at  the  MBL. 

CONSTRAINTS  AND  PURPOSES 
The  dual  mission  of  the  MBL 

For  the  invertebrate  course,  as  for  the  MBL  as  a  whole,  the  dual  mission  has 
always  been  to  produce  new  biological  investigators  as  well  as  new  biological 
investigations.  Successive  directors  (and  boards  of  trustees)  have  held  firm  to  that 
mission,  despite  recurrent  pressures  to  cut  back  on  instruction  (or,  in  earlier  years, 
to  place  it  under  the  administration  of  one  or  two  colleges).  They  have  also 
deliberately  opposed  any  proposals  to  decouple  instruction  from  investigation  at 
Woods  Hole.  In  1978,  the  first  annual  report  of  Paul  R.  Gross  quotes  Whitman, 
"Other  things  being  equal,  the  investigator  is  always  the  best  instructor."  Charles 
Packard's  annual  report  for  1940  quotes  as  the  first  point  promulgated  by  the  then 
committee  on  instruction,  "The  instructorships  are  to  be  regarded  as  aids  to 
research.'"  As  noted  in  this  symposium  and  elsewhere  (Lillie,  1944;  Maienschein, 
1985),  the  prehistory  of  the  MBL  was  responsible  for  the  initial  importance  of 
instruction.  This  was  unusual  among  the  early  marine  laboratories;  initially  Con- 
carneau,  Naples,  Plymouth,  and  Monaco  had  no  great  instructional  committments, 
although  provision  of  aquariums  for  the  general  public  was  important. 

One  other  institution,  the  laboratory  of  the  Scottish  Marine  Biological  Association 
at  Millport  in  Scotland,  had  mixed  origins  like  the  MBL  (Yonge,  1972;  Currie, 
1983)  and  an  early  instructional  mission.  It  was  founded  as  a  floating  laboratory, 
the  Ark,  by  John  Murray  (of  Challenger  fame)  at  Granton  in  1884  and  was  towed 
to  Millport  (1885)  on  the  Isle  of  Cumbrae  in  the  Clyde  Sea  Area  to  join  forces  with 
Dr.  David  Robertson  (earlier  an  associate  and  advisor  of  Anton  Dohrn,  see  Groeben, 
1984),  and  a  group  of  Glasgow  naturalists  (several  involved  with  adult  education). 
The  Association  was  incorporated  as  a  non-profit  company  to  promote  research 
and  education  in  marine  biology,  in  a  close  parallel  to  the  MBL.  For  nearly  seven 
decades,  investigation  and  instruction  were  symbiotic  at  Millport  as  at  Woods  Hole. 
However,  in  1968-1970  almost  all  research  activities  were  transferred  from  Millport 
to  a  new  set  of  laboratory  buildings  at  Dunstaffnage  near  Oban  on  the  west  coast 
of  mainland  Scotland.  The  Millport  laboratory  has  been  left  to  continue  teaching 
and  the  supply  of  marine  material  for  instruction  elsewhere,  as  the  University 
Marine  Biological  Station,  administered  jointly  by  London  and  Glasgow  Universities. 
It  is  to  the  credit  of  Norman  Millott  and  John  A.  Allen,  successive  directors  of  the 
"rump"  station,  that  some  worthy  research  has  continued  at  Millport.  Within  fifteen 


INVERTEBRATE   INSTRUCTION   AT  WOODS   HOLE  95 

years,  the  Dunstaffnage  laboratory  has  grown  much  more  like  the  Woods  Hole 
Oceanographic  Institution  than  the  MBL,  and  it  is  interesting  that  they  are  now 
similarly  trying  to  re-establish  tenuous  links  with  graduate  education.  The  dual 
committment  to  research  and  training  at  the  MBL  should  never  be  compromised. 

Given  that  the  double  mission  has  helped  create  the  rich  MBL  summer 
environment  of  collective  curiosity  and  scientific  debate,  what  was  uniquely  important 
about  the  MBL's  invertebrate  course?  My  thesis  suggests  that  both  the  microenvi- 
ronment  of  the  instructional  setting,  and  the  numbers,  recruiting,  and  turnover  of 
the  instructional  staff  were  unusual,  possibly  unique.  Both  features  may  merit  a 
further  gloss. 

The  instructional  theater 

The  part  played  by  the  invertebrate  teaching  laboratory  as  the  theater  in  which 
instruction  in  invertebrate  zoology  evolved  over  nearly  a  century,  can  hardly  be 
exaggerated.  The  availability  of  healthy  marine  invertebrates  in  the  tanks  and  dishes 
of  the  sea-tables  almost  mandated  that  over  nine  decades  some  students  in  every 
summer  would  pursue  studies  on  whole  animal  physiology  or  behavior.  The 
organization  of  this  proscenium  arch  (along  with  the  conditions  of  recruitment  for 
the  actors)  created  certain  limitations  as  well  as  opportunities  which  remained 
consistent  throughout  the  years.  It  is  important  to  realize  that  the  teaching  laboratory 
in  Old  Main  never  provided  the  best  conditions  for  teaching  comparative  anatomy, 
as  successive  instructors  who  had  inclinations  toward  such  classical  morphology 
(such  as  Drew,  or  forty  years  later  Bullock,  or  sixty  years  later  Schopf)  complained 
from  time  to  time.  It  was  always  a  much  better  place  to  study  feeding  mechanisms 
or  locomotory  patterns  or  reproductive  behavior — in  other  words  to  investigate 
problems  of  whole  animal  physiology  in  a  mechanistic,  reductionist  fashion.  Thus 
it  was  entirely  apt  that  the  successor  teaching  laboratory,  partly  designed  on  the 
basis  of  the  Old  Main  one,  should  be  housed  in  a  training  building  named  after 
Jacques  Loeb. 

The  case  of  Winterton  C.  Curtis  provides  a  revealing  excursus  on  student  work- 
spaces. He  was  a  student  in  the  Zoology  Course  in  1 896,  an  independent  investigator 
in  the  laboratory  in  the  last  years  of  the  century,  and  later  instructor  and  then 
instructor-in-charge  for  four  years  ending  in  1911.  He  went  on  to  head  the  first 
National  Research  Committee  on  the  biological  effects  of  radiation,  and  as  such 
was  responsible  for  obtaining  early  X-ray  equipment  both  for  H.  J.  Muller  and  for 
the  MBL.  As  every  historian  knows,  he  was  one  of  the  expert  witnesses  used  so 
effectively  by  Clarence  S.  Darrow  as  defense  attorney  in  the  Scopes  trial  in 
Tennessee.  I  first  encountered  Dr.  Curtis,  then  a  spry  gentleman  in  his  mid-eighties, 
when  he  came  into  the  invertebrate  teaching  laboratory  in  Old  Main  to  ascertain 
who  was  sitting  in  his  old  seat.  He  returned  within  an  hour  to  present  the  student 
with  some  suitably  inscribed  books.  On  inquiry,  I  found  he  had  been  doing  this  for 
decades  and,  since  the  seating  plan  of  Old  Main  did  not  change,  the  student  honored 
each  year  always  had  a  surname  beginning  with  the  letter  'B\  'O,  or  *D\  He  always 
talked  for  a  while  with  the  student  (carefully  avoiding  any  interference  with  class 
work).  It  was  all  very  discreet  and  I  may  know  more  about  the  matter  than  most 
instructors,  simply  because  one  letter  'B\  Stephen  C.  Brown,  was  subsequently  a 
research  associate  of  mine.  Dr.  Curtis  continued  his  annual  visits  to  Old  Main  until 
the  summer  of  1964;  he  died  in  1966. 

The  importance  of  each  student  being  assigned  an  individual  working  place, 
which  may  be  occupied  at  any  time  of  day  or  night,  was  established  by  Whitman 
and  is  confirmed  as  policy  at  several  places  in  Lillie's  account.  As  Curtis  knew. 


96  W.   D.   RUSSELL-HUNTER 

alphabetic  assignment  of  places  had  persisted  for  at  least  six  decades.  What  Lillie 
(1944)  noted  in  his  retrospect  of  forty  years  was  still  true  in  the  middle  sixties,  and 
lights  remained  on  in  the  invertebrate  laboratory  of  Old  Main  long  into  most  nights. 
Of  course,  the  Old  Main  community  was  never  totally  asocial;  many  human 
interactions  took  place  across  and  around  the  sea-tables.  Temporary,  and  more 
lasting,  partnerships  seemed  to  be  exogamous.  Therefore,  a  hypothesis  to  be  tested 
by  a  curious  sociologist  working  our  archives  is  that  subsequent  marriages  involved 
alphabetically  distant  pairs. 

The  setting  also  provided  an  open  consulting  clinic  on  invertebrate  biology  for 
other  investigators  at  MBL.  If  some  physiologist's  research  animals  became  parasitized, 
there  was  always  a  possibility  that  someone  (staff  or  student)  in  the  teaching 
laboratory  could  identify  the  parasite.  If  a  comparative  biochemist  wanted  different 
sources  of  cartilage  in  invertebrates  or  of  elastin  fibers,  information  (and  possibly 
living  examples)  could  be  available.  A  truly  synergic  exchange  of  knowledge  between 
the  collectors  of  the  supply  department  and  the  instructors  in  the  invertebrate 
laboratory  extended  over  both  the  early  decades  and  the  last  twenty  years  of  the 
course.  This  occasionally  took  on  unusually  systematic  aspects,  with  instructors 
providing  diagnostic  keys  for  collectors  dealing  with  congeneric  species,  or  collectors 
providing  habitat  details  for  less  frequent  forms.  I  have  been  told  that,  over  a  period 
of  two  intermediate  decades,  this  commensal  relationship  went  sour,  with  certain 
collectors  being  secretive  not  only  about  specific  localities  but  also  about  general 
habitat  conditions.  In  general,  however,  the  workers  in  both  the  invertebrate  teaching 
laboratory  and  the  supply  department  provided  cognate  and  complementary  resources 
on  invertebrate  diversity  to  the  rest  of  the  Woods  Hole  community.  A  restored 
invertebrate  zoology  course  could  complement  the  functioning  of  the  proposed 
Marine  Resources  Center  in  a  similar  way. 

The  prevalence  of  mechanistic  investigations  of  feeding  and  locomotion  over 
the  years  has  already  been  noted,  and  was  clearly  stimulated  by  the  availability  of 
healthy  animals  in  the  sea-tables  of  the  teaching  laboratory.  As  early  as  the  tenth 
session  in  1897,  a  great  deal  of  "live  histology"  was  going  on,  making  use  of  the 
translucency  of  tissues  in  certain  invertebrates  to  observe  microscopically  subcellular 
processes.  Productive  variants  of  these  techniques  continued  for  eighty  years,  and 
contribute  to  ongoing  work  on  the  visualization  of  microtubules.  Indirectly,  the 
transference  of  such  techniques  used  in  Woods  Hole  to  living  mammalian  tissues 
in  the  development  of  the  rabbit's  ear  chamber  by  Eliot  R.  Clark  in  1929  led,  as 
was  documented  by  W.  E.  LeGros  Clark  (1958),  to  the  modernization  of  much 
vertebrate  histology  and  to  some  continuing  techniques  of  tissue  culture. 

Research  projects  in  the  open  teaching  laboratory  were  always  carried  out  amidst 
a  critical  public.  While  on  occasion  an  instructor  or  post-course  student  would  feel 
somewhat  over-advised,  at  least  there  was  no  danger  that  any  experimental  design 
would  be  deficient  in  controls  or  involve  pseudoreplication.  Finally,  the  significance 
of  the  living  invertebrates  in  the  sea-tables  can  be  illustrated  by  two  prohibitions 
which  were  among  the  very  few  laboratory  rules  for  students  in  the  course.  First, 
histological  fixatives  were  banned  from  the  students'  work-benches  and  from  the 
sea-table  area.  Secondly,  students  were  strongly  discouraged  from  making  personal 
collections  of  preserved  animals  or  of  molluscan  shells.  Keeping  a  diversity  of  living 
material  available  for  all  students  was  regarded  as  paramount. 

Instructor  turnover  and  dialectic 

Great  evolutionary  strength  resulted  from  the  MBL  policy  that  the  term  as 
instructor-in-charge  of  an  established  course  should  not  exceed  five  years.  This 


INVERTEBRATE   INSTRUCTION   AT   WOODS   HOLE  97 

allowed  conceptual  shifts  as  well  as  technological  ones  to  occur  in  the  invertebrate 
course,  the  one  continuous  feature  for  more  than  80  years  being  the  teaching 
laboratory  with  its  healthy  living  marine  invertebrates  in  the  running-water  sea- 
tables.  It  is  clear  that  over  part  of  this  time,  instruction  in  the  parallel  course  in 
marine  botany  hardly  evolved  at  all.  A  single  instructor-in-charge  ran  that  course 
for  18  years  and,  after  a  3-year  interval,  his  designated  successors  ran  a  largely 
unchanged  course  for  a  further  15  years.  This  relative  stasis  in  instruction  has 
created  major  planning  difficulties  in  marine  botany  for  the  Director  and  instruction 
committee  of  the  MBL  to  this  day. 

Not  only  the  limited  term  for  each  course  head,  but  also  the  larger  number  of 
instructors  to  be  recruited,  maintained  the  dynamic  strength  of  the  invertebrate 
course.  All  instructors  were  investigators.  To  put  it  crudely,  the  instructor-in-charge 
of  any  MBL  course  dispensed  patronage  in  the  form  of  opportunities  for  research. 
The  single  characteristic  shared  by  all  chosen  as  instructors  has  always  been  their 
predictable  research  yield.  This  is  why  I  believe  the  larger  size  of  the  invertebrate 
course  staff  was  important.  Each  group  of  eight  or  nine  instructors,  recruited  as 
having  research  interests  in  different  invertebrates,  almost  inevitably  included  both 
mechanistic-physiologists  and  population-naturalists,  with  experimentalists  largely 
but  not  exclusively  in  the  former  group.  Each  year  some  aspects  of  a  thesis-antithesis 
(Allen,  1979,  1981;  Mayr,  1982)  would  be  publicly  renewed  to  the  great  benefit  of 
the  student  group.  When  in  the  invertebrate  classroom  headed  by  Clark  P.  Read  24 
years  ago,  I  challenged  the  biochemical  generalizations  on  invertebrate  aging  put 
forward  by  Bernard  L.  Strehler,  using  criticisms  drawn  from  demographic  statistics 
for  life-cycles,  I  did  not  then  realize  my  part  in  continuing  an  educational  process 
of  long  standing.  If,  in  similar  debates,  pejorative  labels  were  used  such  as  "tissue- 
grinding"  versus  "dickybird-listing,"  they  more  often  referred  to  contrasting  meth- 
odologies rather  than  to  conceptual  paradigms  (as  clearly  perceived  in  other  cases 
by  our  historians,  see  Allen,  1981;  Maeinschein,  1981),  and  obviously  sounded  both 
more  significant  and  more  polemic  to  outside  observers.  There  was  often  greater 
tolerance  of  the  broader  paradigms,  and  most  instructors  recognized  (when  they 
bothered  at  all)  that  both  the  evolutionary  (population)  and  the  physiological  levels 
of  explanation  were  of  legitimate  interest  to  biologists.  In  discussing  students  of 
ultimate  (evolutionary)  and  proximate  (physiological)  causations,  Mayr  (1982)  has 
even  retrodicted  the  polarity  back  to  the  late  sixteenth  century  and  to  the  contrast 
between  herbalist-naturalists  and  physician-physiologists. 

Aside  from  their  importance  in  an  educational  process,  the  continuously  renewed 
debates  among  invertebrate  course  instructors  generated  or  reconstituted  a  number 
of  active  research  areas.  A  few  from  the  last  thirty  years  can  be  mentioned.  Debate 
and  research  on  the  occurrence  and  nature  of  biological  "clocks"  spread  from 
invertebrate  course  instructors.  Revival  of  interest  in  August  Putter's  theories  on 
direct  uptake  of  organic  solutes  by  invertebrates  had  a  similar  history.  Research 
controversies  involving  the  hormones  and  neurohormones  of  arthropods  moved 
into  and  out  from  the  course.  In  several  periods,  dissension  about  invertebrate 
orientation  and  navigation  gave  rise  to  fruitful  research.  Debate  on  a  heart-like 
structure  in  sea  urchins  was  speculative  and  proved  to  be  functionally  ill-based,  but 
gave  rise  to  investigations  of  the  role  of  the  axial  organ  in  responses  to  infective 
microorganisms  in  echinoderms.  It  is  possible  that  a  majority  of  such  research 
questions  would  not  have  been  debated  and  developed  as  active  projects,  but  for 
the  concurrence  as  instructors  of  mechanistic-physiologists  and  population-naturalists. 

On  the  broader  scale,  invertebrate  zoology  and  the  other  formal  courses  have 
provided  essential  recruitment  (along  with  hierarchical  selective  processes)  to  renew 
the  corporate  body  of  active  investigators  at  the  MBL.  Over  the  years,  successive 


98  W.   D.   RUSSELL-HUNTER 

directors  of  the  MBL  resisted  any  proposals  to  decouple  instruction  from  investigation 
(thus  avoiding  the  fate  of  the  Scottish  marine  station  at  Millport),  or  to  modify  for 
the  embryology,  physiology,  and  invertebrate  zoology  courses  the  policy  that  the 
term  of  each  instructor-in-charge  should  not  exceed  five  years  (thus  avoiding  the 
evolutionary  difficulties  of  the  marine  botany  course).  Continuing  and  evolving 
strength  of  the  MBL  as  a  research  community  depends  upon  its  demography  being 
dynamic.  In  turn,  this  requires  both  intracohort  competition  (no  matter  how  research 
space  is  assigned,  accounted,  or  funded)  and  effective  continuing  recruitment.  The 
latter  was  well  served  by  the  "research  patronage"  mode  of  recruitment  for  both 
course  instructors  and  postcourse  students. 

ACKNOWLEDGMENTS 

I  am  truly  indebted  (along  with  all  the  other  participants  in  the  Ischia  symposium) 
to  Paul  R.  Gross  and  Alberto  Monroy  for  creating  it,  and  to  Seymour  S.  Cohen 
and  Garland  E.  Allen  for  helping  them  organize  the  integrated  program.  It  is 
obvious  that  my  own  charge,  concerning  invertebrate  teaching  in  the  MBL's 
evolution,  has  been  treated  rather  literally  in  this  paper.  As  a  historical  contribution, 
its  biases  are  obvious,  and  it  may  tend  to  resemble  the  eulogies  of  a  regimental 
history.  However  it  should  have  some  value  not  only  as  a  participant's  recall,  but 
also  as  an  ecologist's  view  of  a  population-habitat  interaction.  Perhaps  physiological 
ecologists,  like  the  above  developmental  biologists  turned  historians,  are  preadapted 
as  amateurs  of  history  by  their  concern  with  the  synoptic  analysis  of  dynamic 
processes  through  time.  I  must  also  thank  Jane  Fessenden  (MBL  Librarian),  Ruth 
Davis  (MBL  Archivist),  Garland  E.  Allen,  and  Peregrine  D.  Russell-Hunter  for  help 
with  reference  materials.  Once  again,  this  paper  has  required  help  at  all  stages  of  its 
production  from  my  wife,  Myra  Russell-Hunter,  to  and  for  whom  I  continue  always 
thankful. 

LITERATURE  CITED 

ALLEN,  G.  E.  1979.  Naturalists  and  experimentalists:  the  genotype  and  the  phenotype.  Stud.  Hist.  Biol. 

3:  179-209. 

ALLEN,  G.  E.  1981.  Morphology  and  twentieth-century  biology:  a  response.  J.  Hist.  Biol.  14:  159-176. 
BUCK,  J.  B.,  A.  LAZAROW,  T.  HAYASHI,  B.  KETCHUM,  AND  J.  W.  GREEN.  1963.  Report  of  the  instruction 

committee  on  educational  policy.  Biol.  Bull.  125:  50-53. 
CURRIE,  R.  I.  1983.  Marine  science  (Two  hundred  years  of  the  biological  sciences  in  Scotland).  Proc.  R 

Soc.  Edinburgh  (B)  84:  231-250. 

DREW,  G.  A.  1923.  Data  on  course  students.  Reproduced  in  Packard  (1940)  below. 
GROEBEN,  C.  1975.  The  Naples  Zoological  Station  at  the  Time  of  Anton  Dohrn:  Exhibition  and  Catalogue. 

English  translation,  published  by  the  Goethe-Institut  (German  Cultural  Center),  Paris.  1 10  pp. 
GROEBEN,  C.  1984.  The  Naples  Zoological  Station  and  Woods  Hole.  Oceanus  27:  60-69. 
LEGROS  CLARK,  W.  E.  1958.  The  Tissues  oj <  the  Body,  4th  ed.  Oxford  University  Press,  Oxford.  415  pp. 
LILLIE,  F.  R.  1944.  The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press,  Chicago. 

284  pp. 
MAIENSCHEIN,  J.  1981.  Shifting  assumptions  in  American  biology:  embryology,  1890-1910.  J.  Hist.  Biol. 

14:  89-113. 
MAIENSCHEIN,  J.  1985.  Agassiz,  Hyatt,  Whitman  and  the  birth  of  the  Marine  Biological  Laboratory.  Biol. 

Bull.  168:(suppl.):  26-34. 
MAYR,  E.  1982.  The  Growth  of  Biological  Thought:  Diversity,  Evolution  and  Inheritance.  Belknap  Press, 

Harvard  University  Press,  Cambridge,  974  pp. 
PACKARD,  C.  1940.  The  scientific  record  of  students  in  courses  at  the  Marine  Biological  Laboratory.  Biol. 

Bull.  79:  25-27. 
PECK,   J.    I.    1896-1898.    Manuscript   notes   on   courses,    bound   with   marked   copies   of  the   annual 

announcements  on  summer  instruction.  Archives,  Marine  Biological  Laboratory,  Woods  Hole. 
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Reference:  Biol.  Bull.  168  (suppl.):  99-106.  (June,  1985) 


THE  SEA   URCHIN   AND  THE   FRUIT  FLY: 
CELL   BIOLOGY  AND  HEREDITY,    1900-1910 

BERNARDINO   FANTINI 

Dipartimento  di  Genet ica  e  Biologia  Molecolare,  Universita  degli  Sludi  di  Roma  "La  Sapienza.  "  and 
Gruppo  di  Storia  delle  Science  Biologiche,  Staiione  Zoologica  di  Napoli 

ABSTRACT 

The  choice  of  an  experimental  subject  particularly  suitable  for  a  specific  research 
field  opened  the  way  for  new  discoveries  and  new  theories.  The  sea  urchin  and  the 
fruit  fly,  the  material  of  choice  for  embryological  and  genetic  research,  symbolize 
two  different  research  traditions. 

Knowledge  regarding  these  animals  was  vast  but  largely  separate,  with  little  cross 
fertilization,  so  that  by  the  '30s  little  was  known  about  the  genetic  system  of  the  sea 
urchin,  and  embryological  studies  of  Drosophila  were  just  beginning. 

This  paper  illustrates  the  reason  for  this  disciplinary  distinction.  Embryologists 
concentrated  on  the  relationships  between  nucleus  and  cytoplasm  while  geneticists 
of  the  '"Drosophila  Group"  concentrated  on  the  nucleus,  focusing  exclusively  on 
the  transmission  aspects  of  heredity.  T.  H.  Morgan's  work  forms  a  central  focus,  as 
he  moved  from  an  embryological  concern  to  build  a  new  scientific  program  of 
genetic  transmission  work. 

DISCUSSION 

Many  science  historians  have  stressed  the  importance  of  laboratory  materials 
for  the  development  of  research  in  particular  fields.  Such  materials  include:  the 
chicken  egg  for  von  Baer's  morphological  theory  of  embryonic  development, 
Salamandra  maculata  for  Flemming's  cytological  investigation  (because  of  the  large 
size  of  its  cells  and  nuclei),  Pisum  sativum  for  Mendel's  early  research  on  the 
hereditary  laws  (as  opposed  to  the  difficult  second  choice  Hieracium),  Triton  for 
Spemann's  experiments  on  the  organizer,  Neurospora  for  chemical  genetics,  and 
Escherichia  coli  and  the  T-phages  for  molecular  biology. 

The  sometimes  deliberate  and  sometimes  serendipitous  choice  of  an  experimental 
subject  suitable  for  a  specific  field  has  opened  the  way  for  advancement  of  research 
and  theories.  Much  progress  has  depended  upon  the  fortuitous  discovery  of 
organisms  that  clearly  illustrate  a  process  or  a  structure,  or  that  lend  themselves  to 
convenient  experimentation.  For  instance,  Flemming's  choice  of  organisms  with 
long  chromosomes  as  his  research  material  undoubtedly  helped  him  to  elucidate 
the  main  features  of  mitosis.  At  other  times  the  choice  has  been  less  fortunate.  For 
example,  Mendel's  decision  to  test  his  theory  of  inheritance  in  Hieraciiim  in  order 
to  generalize  it,  (as  suggested  by  Nageli),  was  certainly  one  of  the  reasons  for  the 
neglect  of  his  work.  In  this  genus,  as  we  now  know,  parthenogenesis  is  common, 
which  led  Mendel  to  results  incompatible  with  his  original  theory.  De  Vries's 
selection  of  Oenothera  lamarkiana  as  experimental  support  of  his  own  Mutation- 
theorie  provides  another  such  negative  example. 

The  sea  urchin  and  the  fruit  fly,  Paracentrotus  lividus  and  Drosophila  melano- 
gaster,  the  materials  of  choice  for  embryological  and  genetic  research,  positively 
symbolize  two  different  research  traditions.  As  a  laboratory  organism,  the  sea  urchin 

99 


100  B.   FANTINI 

was  first  introduced  by  Hertwig  in  the  spring  of  1875,  during  his  research  on  the 
process  of  fertilization.  The  egg  in  this  species  is  small,  has  little  yolk,  and  remains 
transparent  even  at  high  magnifications.  Moreover  both  egg  and  sperm  are  easy  to 
preserve.  This  material  is  therefore  very  suitable  for  embryological  studies.  The  sea 
urchin  was  the  organism  on  which  Driesch  performed  his  famous  experiments,  in 
1891,  separating  the  two  first  blastomeres  and  thereby  challenging  W.  Roux's  mosaic 
theory  of  embryonic  development.  In  this  way  it  became  the  material  of  choice  for 
that  part  of  embryology  concerned  with  cell  physiology  and  the  mechanism  of 
determination  ("physiological  morphology"). 

In  contrast,  Drosophila  was  introduced  as  an  experimental  organism  for  hereditary 
studies  in  W.  E.  Castle's  laboratory  around  1901.  In  1906  Castle  described  some 
Mendelian  characters  in  Drosophila,  and  T.  H.  Morgan  started  to  use  the  fly  by 
1908  to  induce  de  Vriesian  mutations  by  exposing  the  larvae  to  radium.  The 
following  years  he  studied  breeding  in  Drosophila  in  connection  with  evolutionary 
studies  and  found  this  animal  a  "wonderful  material."0  However,  only  after  1910 
did  he  carry  out  the  experiments  which  founded  a  new  discipline,  the  chromosome 
theory  of  the  gene. 

Embryologists  and  geneticists  studied  their  respective  experimental  objects  in 
impressive  depth,  yet  knowlege  of  these  animals  remained  distinct,  with  little  cross 
fertilization.  Between  1906  and  1907  Stevens  explored  Drosophila  for  cytological 
studies,  but  the  fruit  fly  did  not  enter  the  realm  of  "cytological  animals."  In  the 
'30s,  then,  after  three  decades  of  Mendelism,  little  was  known  about  the  genetic 
system  of  the  sea  urchin  and  the  embryological  studies  of  Drosophila  were  just 
beginning.  Why  for  many  decades,  notwithstanding  the  growing  importance  of 
research  performed  on  these  organisms,  did  these  subjects  of  study  and  the  questions 
asked  of  them  remain  so  distinct? 

By  the  1890s  experimental  embryologists  stressed  that  development  was  the 
result  of  factors  internal  to  the  organism  itself,  even  though  they  differed  on  how 
the  internal  mechanisms  operated.  That  mechanism  relates  to  the  notion  of  heredity, 
the  transmission  of  some  internal  organizing  factors  from  parents  to  offspring 
through  the  zygote.  In  particular,  some  embryologists  thought  that  the  entire  process 
was  determined  by  the  organization  of  the  egg  cytoplasm  prior  to  fertilization.  They 
asked:  what  kind  of  factor  is  actually  inherited?  How  do  they  control  embryonic 
differentiation?  In  such  a  way  the  problems  of  heredity  and  those  of  embryogenesis 
seem  to  be  neatly  connected.  Yet,  the  two  fields,  embryology  and  genetics,  remained 
separated  for  many  decades.  Why?  To  answer  such  questions,  it  is  necessary  to 
study  the  historical  developments  of  both  disciplines.  In  particular,  Morgan's  shift 
from  embryology  to  genetics  and  back  to  embryology,  well  known  thanks  to 
Garland  Allen's  book,  provides  a  useful  case-study  for  some  general  remarks  about 
disciplinary  relationships.1 

Embryological  work  includes  a  variety  of  distinct  research  programs.  Entwick- 
lungsmechanik  dominated  the  embryological  community,  at  least  in  Europe,  but 
this  "sea  urchin  tradition"  was  in  a  morphological  phase,  even  if  it  was  a 
physiological  morphology,  looking  for  a  causality  of  morphogenesis.  Cell  lineage 
was  a  different  tradition  and  many  of  its  problems  were  common  to  cytology,  the 
morphological  study  of  cell  structure.  Within  the  cytological  tradition  the  main 
concern  was  the  multiplication  of  cells  and  nuclei  and  in  particular  the  role  and 
behavior  of  chromosomes.  Already  in  1888  Boveri  had  demonstrated  that  the 
members  of  each  pair  of  homologous  chromosomes  are  qualitatively  different  in 
their  hereditary  determinants  from  members  of  all  other  pairs.  Thus  each  chromosome 
pair  was  unique,  having  individuality,  even  when  it  disappeared  during  some  phases 


CELL   BIOLOGY   AND  HEREDITY  101 

of  cell  division.  Another  research  program,  Mendelian  genetics,  was  derived  from  a 
tradition  different  from  chromosome  studies,  both  theoretically  and  institutionally, 
and  scientific  communications  between  the  two  fields  remained  rare.  Study  of 
heredity  was  not  concerned  with  transmission  genetics  but  with  corpuscular  theories 
of  heredity  and  embryology.  The  close  relationship  between  the  study  of  heredity 
and  embryology  prior  to  Mendel's  rediscovery  disappeared  in  the  early  XXth 
century. 

The  main  point  of  contact  between  embryology  and  heredity  was  the  problem 
of  sex  determination.  This  problem  was  well  studied  by  the  cell-lineage  school,  but 
it  was  less  interesting  for  the  typical  Entwicklungmechanicker,  who  was  more 
concerned  with  the  very  early  phases  of  embryonic  development,  especially  the  first 
cell  divisions.  This  problem  lay  at  the  boundary  of  the  two  separate  disciplines:  it 
was  studied  by  cytologists,  like  E.  B.  Wilson,  and  by  geneticists,  like  W.  Bateson,  L. 
Cuenot,  and  W.  E.  Castle,  with  very  different  methods  and  theories. 

The  various  theories  of  sex  determination  current  in  the  decade  1900-1910  fell 
into  two  general  groups.  One  group  maintained  that  external  conditions  were  the 
primary  factors  in  determining  sex;  the  opposing  group  maintained  that  sex  was 
determined  only  by  internal  factors.  Cytologists  generally  held  the  first  theory  and 
geneticists  the  second,  presenting  a  different  form  of  the  old  dichotomy  between 
epigenesis  and  preformation. 

In  1902,  C.  E.  McClung  presented  the  first  suggestion  that  the  "accessory"" 
chromosome,  described  by  Henking  in  1891,  could  be  associated  with  sex  determi- 
nation in  adults,  on  the  basis  of  the  existence  of  two  classes  of  sperm,  one  with  an 
accessory  chromosome.  Shortly  afterward,  in  1902,  Boveri  and  Sutton  pointed  out 
the  extraordinary  similarity  between  the  cytologically  observed  separation  of  members 
of  each  chromosome  pair  during  gamete  production  and  Mendel's  postulated 
segregation  of  independent  factors.  That  was  evidence  of  the  possible  link  between 
chromosomes  and  Mendelian  factors.  Boveri's  evidence  was  conclusive,  however, 
only  for  people  already  convinced  of  the  chromosome  theory,  for  it  indicates,  as 
Morgan  wrote  in  1910  "that  individual  chromosomes  do  not  in  any  sense  contain 
either  preformed  germs  or  determinants,  or  unit  characters,  or  even  stand  for  the 
production  of  particular  organs  in  any  sense."2 

In  1905  Wilson's  and  Stevens'  evidence  that  the  accessory  chromosome  was  the 
sex-determining  element  confirmed  a  close  parallel  between  cytological  observations 
and  the  determination  of  sex.  In  spite  of  these  results,  even  among  the  supporters 
of  the  theory  of  chromosomal  determination  of  sex,  the  conclusive  evidence  of 
these  observations  was  weak.  Wilson  concluded  that  "great  if  not  insuperable 
difficulties  are  encountered  by  any  form  of  the  assumption  that  these  chromosomes 
are  specifically  male  or  female  sex  determinants." 

Richard  Hertwig's  work  in  1906-1907,  studying  cross  fertilization  between  frogs 
of  different  geographic  races,  furnished  additional  evidence  that  chromosomes  really 
did  have  a  significant  role  in  sex  determination.  Those  experiments  showed  that  if 
one  used  the  sperm  of  a  single  male  for  fertilizing  different  females,  the  normal  sex 
ratio  among  the  offspring  would  result.  In  contrast,  fertilizing  a  female  with  sperm 
from  different  males  produces  quite  variable  results.  It  could  be  concluded  that  the 
sperm  was  really  the  sex-determining  element.  Morgan  was  highly  impressed  by 
these  experiments  and  thereafter  considered  Wilson's  interpretations  more  favorably. 
Nevertheless,  Morgan  was  clearly  opposed  to  the  idea  that  chromosomes  contain 
specific  hereditary  units.  The  chromosomes  can  have  an  important  role  in  the 
determination  of  characters,  but  not  necessarily  as  the  carriers  of  genetic  factors. 
The  two  roles  remain  quite  distinct.  There  are  two  different  kinds  of  problems:  (1) 


102  B.   FANTINI 

the  chromosomal  determination  of  sex  and  (2)  sex  as  a  Mendelian  factor.  One 
could  accept  the  first  without  accepting  the  second.  The  epigenetic  embryological 
tradition  was  ready  to  accept  the  causal  connection  between  chromosomes  and 
characters,  but  not  the  concept  of  the  chromosomes  as  specific  Mendelian  factors. 
Wilson  eventually  concluded  that  "sex  production  stands  in  some  relation  with  the 
chromosomes  and  can  be  treated  from  the  stand  point  of  Mendelian  phenomena, 
as  interpreted  by  the  Sutton-Boveri  chromosome  theory."4  But  this  was  more  an 
act  of  faith,  the  endorsement  of  a  new  theory,  than  the  result  of  empirical 
demonstration.  Wilson  considered  the  chromosome  as  the  Mendelian  factor,  "sex 
determination  being  a  matter  of  Mendelian  dominance,  more  specifically  of  chro- 
mosome dominance." 

In  contrast,  Morgan  rejected  the  simple  version  of  chromosome  theory  as  naive 
given  the  small  number  of  chromosomes  compared  to  the  large  number  of  inherited 
traits.  He  generalized  his  attack  against  the  assumption  that  the  nucleus  must  be 
the  bearer  of  the  hereditary  qualities  of  the  male,  arguing  that  "the  protoplasm  may 
account  for  the  results."  For  Morgan,  sex  was  determined  in  some  way  more 
complex  than  the  action  of  a  single  element  like  the  X-chromosome,  in  particular 
by  the  action  of  the  cytoplasm  on  the  hereditary  material  or  by  the  relationship 
between  nucleus  and  cytoplasm.  In  the  period  between  1894  and  1906  Morgan  had 
often  invoked  a  gradient  or  polarity  concept  of  substance  in  the  formative  phenomena, 
in  particular  for  regeneration.  According  to  this  theory,  the  fate  of  tissues  was 
determined  by  the  concentration  of  some  particular  substances  in  the  cytoplasm. 

Morgan's  critiques  of  chromosomal  determination  of  sex  were  two-fold.  First 
the  evidence  was  contradictory.  Wilson's  report  that  the  spermatozoon  having  the 
extra  chromosome  produces  a  female  every  time  and  that  the  one  without  it 
produces  a  male  was  exactly  the  opposite  of  McClung's  conclusion.  Wilson  and 
Stevens  had  studied  mostly  insects  and  concluded  that  males  were  heterozygous 
(XY)  and  females  homozygous  (XX)  for  the  accessory  chromosome.  Punnett  and 
Raynor  had  worked  primarily  with  moths  and  chickens  and  had  found  just  the 
opposite.  Second,  for  Morgan  the  supposition  was  contradictory  in  itself  because  it 
suggested  that  the  same  chromosome  will  be  female-determining  in  one  generation 
and  male-determining  in  the  next. 

As  a  consequence  Morgan  criticized  "this  modern  way  of  referring  everything 
to  the  chromosomes".6  He  decided  in  favor  of  the  cytoplasmic  determination  of 
hereditary  phenomena.  He  argued  that  chromosomes  might  fuse  completely  during 
the  process  of  intertwining  (synapsis),  and  thought  that  the  union  of  two  homologous 
chromosomes  during  synapsis  was  as  homogeneous  as  when  two  drops  of  water  fuse 
into  one.7  In  other  words,  for  Morgan  the  chromosomes  do  not  contain  hereditary 
information  for  individual  characters  of  the  embryo,  but  an  embryo's  hereditary 
traits  result  from  the  interactions  of  materials  produced  by  the  "entire  constellation 
of  chromosomes."8  At  this  moment  Morgan  again  heralded  an  epigenetic  view,  and 
stressed  the  importance  of  process  as  opposed  to  form.  For  him  the  chromosome 
theory  and  Mendelian  genetics  demanded  "a  morphological  basis  in  the  germ  for 
the  minutest  phase  (factor)  of  a  definitive  character."  It  is  essentially  a  morphological 
conception  with  but  a  trace  of  functional  concern.  Mendelism  "utilizes  not  a  single 
finding  of  the  science  of  biochemistsry  .  .  .  With  an  eye  seeing  only  particles  and  a 
speech  only  symbolizing  them,  there  is  no  such  thing  as  the  study  of  process 
possible".9  As  E.  S.  Russell  pointed  out  in  1930: 

epigenesis  and  preformation  represent  two  different  attitudes  to  the  problem  of 
development,  arise  from  two  fundamentally  different  philosophies.  The  epigenetic 


CELL   BIOLOGY   AND   HEREDITY  103 

view  is  dynamic,  vitalist,  physiological;  the  preformationist  is  static,  deterministic, 
and  morphological.  The  one  stresses  time  or  process,  the  other  space  and 
momentary  state — the  one  emphasizes  function,  the  other  concentrates  on  form.1" 

Usually  Morgan's  attitude  toward  Mendelism  prior  to  1910  has  been  presented 
as  a  reaction  against  the  use  of  metaphysical  concepts  such  as  determinants, 
idioplasm,  biophores,  micelles,  etc.  However,  Morgan  was  also  trying  to  generalize 
the  de  Vriesian  theory  of  mutation  which  was  based  on  the  concept  of  pangenes, 
hereditary  particles  "in  many  respects  analogous  to  the  molecules  of  chemists/'12 
Yet  this  kind  of  theorizing  remained  in  the  background  because  de  Vries  and 
Morgan  were  trying  to  deduce  hereditary  and  evolutionary  laws  by  indirect  methods, 
such  as  breeding  and  selection  of  varieties.  As  an  embryologist  by  training,  Morgan 
was  quite  accustomed  to  the  concept  of  material  hereditary  particles,  the  idea  that 
the  nucleus  and  the  chromosomes  in  particular  are  directly  involved  in  hereditary 
transmission.  From  this  point  of  view,  he  was  looking  for  a  material  basis  of 
evolutionary  jumps,  suitable  for  experimental  investigation.  Therefore,  his  negative 
attitude  towards  both  Mendelian  genetics  and  the  chromosomal  theory  can  be 
attributed  to  the  lack  of  scientific  coherence  between  them.  Only  after  the  elucidation 
of  the  phenomenon  of  chiasmatype  could  the  connection  between  genetical  and 
cytological  evidence  be  established  and  the  idea  of  material  factors  governing 
heredity  and  embryogenesis  be  openly  accepted. 

The  turning  point  and  the  beginning  of  the  fruit  fly  model  was  Morgan's  own 
studies  on  the  "sex-limited"  inheritance  on  Drosophila  melanogaster.  The  appearance 
of  the  white-eyed  mutant  suddenly  shifted  Morgan's  interest  from  evolutionary 
studies  to  heredity.  The  important  fact  was  that  this  particular  character,  produced 
by  a  mutation  totally  different  from  a  de  Vriesean  mutation,  not  only  appeared  to 
be  a  pure  Mendelian  character  but  also  seemed  clearly  connected  with  sex  inheritance. 
In  explaining  these  results  Morgan  could  accept  the  basic  structure  of  Mendelian 
explanation  and  Wilson's  and  Stevens'  idea  that  determination  of  sex  has  a 
chromosomal  basis.  Even  in  1909,  before  the  discovery  of  the  fly  with  white  eyes, 
Morgan  assumed  the  role  of  a  combination  of  hereditary  factors.  In  order  to  explain 
the  deviations  from  Mendelian  ratios  observed  in  1905  with  yellow  mice,  Cuenot 
had  assumed  the  idea  of  selective  fertilization  proposed  by  Castle.  In  such  a  way  no 
yellow  mouse  would  be  obtained  by  breeding  heterozygotes  since  yellow  factors 
from  the  sperm  do  not  combine  with  yellow  factors  from  the  egg. 

From  his  epigenetic  point  of  view,  Morgan  could  not  accept  that  such  trivial 
differences  as  hair  color  could  prevent  the  fertilization  between  gametes  carrying 
those  colors.  Morgan  was  thus  skeptical  about  the  Mendelian  concept  of  purity  of 
gametes  and  their  absolute  segregation  during  cell  divisions.  In  consequence,  he 
explained  Cuenot's  results  by  assuming  that  certain  alleles  always  remained  together 
rather  than  segregating  in  a  random  fashion.  His  complex  mechanism  assumed  that 
yellow  and  gray  "unit  characters"  stayed  together  in  50%  of  the  offspring.  One 
major  point  of  his  chromosomal  theory,  namely  linkage,  was  already  present  as  a 
formal  concept  before  the  discovery  of  the  cytological  phenomenon  of  chiasmata. 
But  it  was  used  against  the  concept  of  the  purity  of  gametes.  Morgan  at  first  rejected 
both  segregation  and  purity  of  gametes,  accepting  both  only  after  his  own  results 
on  sex-linked  characters  and  cytological  evidence. 

If  adult  characters  are  determined  by  hereditary  particles  on  chromosomes  then 
a  large  number  of  traits  must  be  inherited  together,  coupled.  Coupling  had  already 
been  observed  by  Bateson,  who  explained  it  first  with  the  theory  of  attraction  and 
repulsion  and  later  with  the  theory  of  reduplication.  The  idea  of  "crossing-over" 


104  B.   FANTINI 

based  on  Janssens'  chiasmatype  was  an  alternative  to  the  complex  cytological 
mechanism  introduced  by  Bateson.  Morgan  assumed  at  the  same  time  that  "the 
materials  that  represent  these  factors  are  contained  in  the  chromosomes"  and  that 
these  factors  that  "couple"  lie  near  each  other  in  a  linear  series.  These  couplings 
can  be  separated  by  crossing  over.  The  difference  in  the  strength  of  the  coupling 
depends  on  "the  linear  distance  apart  of  the  chromosomal  materials  that  represents 
the  factors."  From  this  assumption  it  follows  that  the  observed  hereditary  phenomena 
"are  a  simple  mechanical  result  of  the  location  of  the  materials  on  the  chromosome, 
and  of  the  method  of  union  of  homologous  chromosomes."  Mendelism  is  no  longer 
a  logical  construction  or  a  numerical  system  but  the  result  of  the  location  of  the 
factors  on  the  chromosomes. 

Thus  Morgan  abandoned  his  agnosticism.  For  embryologists,  the  "sea  urchin 
people,"  the  most  important  question  was  not  so  much  how  hereditary  information 
was  transmitted  from  one  generation  to  the  following,  but  rather  how  that  information 
was  translated  into  adult  characters.  For  an  embryologist,  heredity  is  not  a  problem 
but  a  prerequisite.  Morgan  abandoned  that  previous  problem — how  inherited 
information  is  transformed  into  adult  characters — to  study  genetics  or  the  transmis- 
sion of  information.  He  moved  from  the  sea  urchin  tradition  to  a  new  Drosophila 
tradition. 

Two  comments  are  necessary.  The  first  is  that  Morgan's  attitude  against 
theoretical  supposition,  which  he  often  used  to  reject  the  ideas  of  others,  was  freely 
abandoned  to  accept  a  very  powerful  and  innovative  hypothesis.  The  second 
comment  is  that  the  bridge  between  cytology  and  genetics  which  Morgan  stressed 
remained  only  a  hope.  The  Drosophila  group  grasped  the  powerful  idea  of  a  linear 
disposition  of  factors  on  the  chromosome  introduced  by  Morgan  and  developed  by 
Sturtevant.  The  group,  especially  the  younger  members  who  pushed  Morgan  in  this 
direction,  considered  the  chromosomes  as  black  boxes,  or  perhaps  as  a  string  of 
black  pearls. 

Embryology  for  its  part  followed  its  own  tradition  without  caring  too  much 
about  the  success  and  the  appeal  of  the  new  research  program  in  genetics.  In  the 
'30s  Morgan  returned  to  his  first  and  greatest  love,  embryology.  He  felt  the  need 
for  a  general  text  on  the  relationships  between  genetics  and  embryology,  but  openly 
confessed  to  critics  that  he  had  discussed  embryology  and  genetics,  not  their 
relationships.13  Indeed  no  evidence  was  given  by  the  Drosophila  group  about  the 
physiological  nature  of  genes  or  the  way  in  which  they  interact  during  embryonic 
development.  In  the  first  decade  of  Mendelism,  many  scientists  described  factors  in 
biochemical  and  physiological  terms,  like  Bateson's  theory  of  presence-absence, 
Castle's  model  of  factor  interaction,  and  G.  S.  Shull's  explanation  of  the  multiple 
characters  phenomenon.  However,  the  Drosophila  group  pushed  this  point  into  the 
background,  as  a  problem  too  difficult  to  explain  at  the  moment. 

Embryology  could  not  be  interested  in  that  kind  of  theory.  Strong  epigenetic 
attitudes  and  physiological  models  were  directing  embryology  towards  other  con- 
cerns— that  is  organizer  theories  and  chemical  embryology,  new  forms  of  physiological 
morphology.  Chemical  embryology  was  in  fact  looking  for  a  chemical  explanation 
of  morphogenetic  movements,  using  may  different  substances  as  stimuli.  Embryol- 
ogists were  more  concerned  with  the  larger  changes  in  the  whole  organism  than 
with  the  lesser  qualities  known  to  be  associated  with  genetic  action.  As  E.  E.  Just14 
said,  embryologists  were  much  more  interested  in  the  back  than  in  the  bristles  on 
the  back  and  more  in  how  the  eyes  are  built  than  in  how  they  acquire  their  color. 

In  the  decades  of  triumph  of  the  chromosome  theory,  embryology  was  in  a 
period  of  depression  and  the  great  expectations  of  the  1 890's  and  the  first  decade  of 


CELL   BIOLOGY   AND   HEREDITY  105 

the  twentieth  century  were  replaced  by  a  sense  of  impotence  towards  the  difficulties 
of  understanding  development. 

There  was  a  time  of  discouragement.  .  .  .  The  fertility  of  the  soil  seemed  to  have 
suddenly  run  out  and  tillage  no  longer  worth  while.  What,  more  human,  then, 
than  the  gold  rush  to  genetics  and  general  physiology?15 

Many  embryological  experiments  seemed  to  stress  the  role  of  the  cytoplasm  in 
differentiation.  The  destruction  of  the  cytoplasm  brought  about  serious  disturbances 
of  normal  development,  but  the  nucleus  could  be  transplanted  without  such 
disturbance,  at  least  in  the  earliest  stages.  The  role  of  the  nucleus  seemed  to  be 
secondary  for  early  cleavage  and  in  the  metabolism  of  the  embryo,  becoming 
important  only  for  morphogenetic  movements.  Embryology,  according  to  its  own 
tradition,  concentrated  on  both  the  nucleus  and  the  cytoplasm;  and  "The  prestige 
of  success  enjoyed  by  the  gene  theory  might  easily  become  a  hindrance  to  the 
understanding  of  development  by  directing  our  attention  solely  to  the  genome, 
whereas  cell  movements,  differentiation  and  in  fact  all  developmental  processes  are 
actually  effected  by  the  cytoplasm/'16 

In  contrast,  the  Drosophila  group  notwithstanding  Morgan's  earlier  emphasis, 
concentrated  on  the  nucleus,  and  more  precisely  on  the  chromosomes,  focusing 
exclusively  on  the  transmissional  aspects  of  heredity.  The  cell,  with  its  biochemistry 
and  physiology,  was  considered  a  black  box.  Study  of  the  egg  produced  no  scientific 
success  and  therefore  was  considered  non-scientific  for  the  Drosophila  group.  Even 
if  the  problem  of  a  biochemical  and  physiological  description  of  the  transformation 
of  the  egg  into  an  adult  is  very  important  and,  "there  can  be  no  question  of  the 
paramount  importance  of  finding  out  what  takes  place  during  development,"  we 
cannot  tell  how  much  this  information  may  lead  to  a  better  understanding  of  the 
chromosome  theory.  "For  a  knowledge  of  the  chemistry  of  all  the  pigments  in  an 
animal  or  plant  might  still  be  very  far  removed  from  an  understanding  of  the 
chemical  constitution  of  the  hereditary  factors  by  whose  activities  these  pigments 
are  ultimately  produced."17 

The  central  problem  of  embryology  in  genetic  terms  was  how  embryonic  cells 
arising  from  the  same  zygote  and  therefore  sharing  the  same  genetic  constitutions 
develop  along  different  lines  during  tissue  differentiation.  Even  in  1935  this  problem, 
the  triggering  mechanism  of  the  expression  of  the  genes,  was  felt  to  be  inaccessible 
to  scientific  research.  Many  alternatives  remained  possible  and  Morgan  in  his  Nobel 
Lecture  confessed  "that  we  must  wait  until  experiments  can  be  devised  to  help  us 
to  discriminate  between  these  different  possibilities."18  Morgan,  in  order  to  reach  a 
scientific  theory  of  genetic  transmission,  had  to  abandon,  for  a  while,  his  own 
tradition  and  interests  to  build  a  new  tradition.  Thus  he  abandoned  the  sea  urchin 
in  favor  of  the  fruit  fly.19 

LITERATURE  CITED 

0  Morgan  to  Dreisch.  23  November  1910.  Quoted  in  ALLEN.  1978.  Thomas  Hunt  Morgan.  The  Man  and 

His  Science.  Princeton  University  Press,  Princeton.  P.  153. 

1  ALLEN,  GARLAND.  1978.  Thomas  Hunt  Morgan.  The  Man  and  His  Science.  Princeton  University  Press, 

Princeton. 

2  MORGAN,  T.  H.  1910.  Chromosomes  and  heredity.  Am.  Nat.  44:  449-496:461. 

3  WILSON,  E.  B.  1905.  The  chromosomes  in  relation  to  the  determination  of  sex  in  insects.  Science  22: 

500-502:502. 

4  WILSON,  E.  B.  1907.  Sex  determination  in  relation  to  fertilization  and  parthenogenesis.  Science  25:  376- 

379:376. 

5  Morgan  to  Driesch,  23  October  1905,  quoted  in  G.  ALLEN  (footnote  1 ),  p.  137. 


106  B.   FANTINI 

6  Morgan  to  Driesch  (footnote  5). 

7  T.  H.  MORGAN  (footnote  2). 
s  T.  H.  MORGAN  (footnote  2). 

9  MORGAN,  T.  H.  1909.  Recent  experiments  on  the  inheritance  of  coat  colors  in  mice.  Am.  Nat.  43:  449- 

510:510. 

10  RUSSELL,  E.  S.   1930.  The  Interpretation  of  Development  and  Heredity.  Clarendon  Press,  Oxford.  P. 

121. 

11  MORGAN,  T.  H.  1907.  Sex  determining  factors  in  animals.  Science  25:  382-384:384. 

12  DE  VRIES,  H.  Intracellulere  Pangenesis.  Gustav  Fischer,  Jena  1897  (Eng.  Transl.  Intracellulur  Pangenesis, 

Open  Court,  Chicago  1910). 

13  Quoted  by  G.  ALLEN  (footnote  1),  p.  300. 

14  JUST,  quoted  by  R.  G.  HARRISON.  1937.  Embryology  and  its  relations.  Science  85:  369-374:372. 

15  R.  G.  HARRISON,  manuscript  1925,  quoted  by  Harrison  (footnote  14),  p.  370. 
15  R.  G.  HARRISON,  (footnote  14),  p.  372. 

17  MORGAN,  T.  H.,  A.  H.  STURTEVANT,  H.  J.  MULLER,  AND  C.  B.  BRIDGES.  1915.  The  Mechanism  of 

Mendelian  Heredity.  Henry  Holt,  New  York.  P.  227. 

18  MORGAN,  T.  H.  1935.  The  relation  of  genetics  to  physiology  and  medicine.  Nobel  Lecture,  Sci.  Monthly 

41:  5-18. 

19  Footnote  17,  p.  227. 


Reference:  Biol.  Bull.  168  (suppl.):  107-121.  (June,  1985) 


HEREDITY   UNDER  AN   EMBRYOLOGICAL   PARADIGM:   THE  CASE 
OF  GENETICS  AND  EMBRYOLOGY 

GARLAND   E.   ALLEN 
Department  of  Biology,  Washington  University.  St.  Louis,  Missouri  63130 

INTRODUCTION 

In  examining  the  history  of  genetics  at  the  Marine  Biological  Laboratory  (MBL), 
a  curious  paradox  emerges:  although  the  leader  of  the  classical  school  of  Mendelian 
heredity  in  the  United  States,  Thomas  Hunt  Morgan  (1866-1945),  was  a  long- 
standing summer  investigator  at  the  MBL  and  brought  his  Drosophila  research 
group  to  Woods  Hole  every  summer  from  1912  through  1940s,  genetics  as  such 
never  became  incorporated  into  official  MBL  work.  There  were  few  other  geneticists 
regularly  at  the  Marine  Biological  Laboratory  other  than  Morgan  and  his  students; 
and,  genetics  did  not  get  incorporated  into  the  courses  of  instruction  at  the 
Laboratory  (it  still  is  not  a  separate  course  today).  At  the  same  time,  embryology, 
so  closely  related  conceptually  and  historically  to  the  study  of  heredity,  was  from 
the  beginning,  and  still  remains  today  an  official  mainstay  of  the  MBL's  research 
and  instructional  program.  Why  should  embryology  have  remained  such  a  strong 
part  of  MBL  work  while  genetics  never  really  achieved  the  same  status?  Was  this  a 
local  phenomenon  peculiar  to  the  MBL,  or  was  it  a  reflection  of  some  more  general, 
intellectual  and/or  social  trend  within  biology  in  the  period  after  the  rediscovery  of 
Mendel  in  1900?  In  attempting  to  answer  these  questions  I  was  led  to  consider  the 
larger  issue  of  the  historical  relationship  between  genetics  and  embryology  in  the 
United  States  (and,  to  some  extent,  internationally  as  well)  in  the  early  decades  of 
the  twentieth  century.  What  begins  as  an  issue  in  the  specific  history  of  the  Marine 
Biological  Laboratory,  broadens  into  the  larger  question  of  the  history  of  twentieth 
century  biology. 

In  the  present  paper  I  will  explore  (1)  the  early  history  of  the  relationship 
between  genetics  and  embryology  from  1890  to  1910;  (2)  the  growing  divergence 
between  the  two  fields  during  the  period  to  1940,  and  (3)  the  role  that  T.  H.  Morgan 
himself  played  in  creating  the  divergence  between  genetics  and  embryology;  and 
finally  (4)  how  the  various  factors  may  have  been  influential  in  causing  Morgan  to 
shift  his  position  so  profoundly.  At  the  end  of  the  paper,  I  will  come  back  to  the 
original  paradox  with  regard  to  the  MBL  and  suggest  why,  on  an  institutional  level, 
the  relationship  between  genetics  and  embryology  seemed  to  mirror  what  was 
occurring  on  the  national  level. 

Focussing  on  the  role  of  Thomas  Hunt  Morgan  in  this  historical  analysis  has 
several  advantages.  First,  Morgan's  career  spanned  the  period  during  which  embryol- 
ogy and  heredity  went  from  the  holistic  and  unified,  to  the  more  restricted  and 
limited  view.  Second,  Morgan  was,  himself,  a  major  architect  of  the  split  that 
occurred  between  genetics  and  embryology.  Although  he  began  his  scientific  career 
as  an  embryologist,  and  to  some  extent  always  remained  one,  he  was  willing  to 
separate  the  study  of  genetics  from  that  of  embryology  during  a  major  segment  of 
his  career.  Third,  focusing  on  the  work  of  a  single  individual  allows  a  more  in- 
depth  analysis  and  the  emergence  of  a  more  clear  picture  of  the  interrelationships 
of  historical,  social,  and  intellectual  factors  than  would  be  possible,  in  the  confines 

107 


108  G.   E.   ALLEN 

of  a  single  paper,  if  I  included  a  large  number  of  investigators,  or  a  wider  range  of 
problems. 

THE  UNIFIED  VIEW  OF  HEREDITY:  1 890- 1910 

Following  the  lead  of  Charles  Darwin  and  others,  many  late  19th  and  early  20th 
century  biologists  chose  to  construct  synthetic  theories  that  related  the  processes  of 
genetic  transmission  to  those  of  embryonic  differentiation  and  development.  Theo- 
rizers  such  as  Ernst  Haeckel  (1834-1919)  and  August  Weismann  (1834-1914)  saw 
the  vertical  process  of  "transmission"  from  parent  to  offspring  and  the  horizontal 
process  of  "translation"  of  heredity  potential  into  adult  traits  as  part  of  the  same 
fundamental  process.  At  the  same  time,  they  made  no  distinction  between  what  we 
would  call  today  genotype  and  phenotype;  rather  they  saw  the  two  as  inseparable, 
a  division  between  potentiality  and  actuality  as  meaningless.  They  also  saw  cell 
nucleus  and  cytoplasm  as  an  integrated  whole  as  one  constantly  interacting  system, 
each  component  inconceivable  without  the  other.  The  theories  of  Haeckel  and 
Weismann  were  stimulating  in  their  synthetic  power,  and  recognized  a  reality — 
namely  that  the  processes  of  genetic  transmission,  embryonic  development,  and 
evolution  of  species  were  interrelated,  an  interrelationship  all  biologists,  young  and 
old,  recognized. 

However,  as  many  younger  biologists  around  the  turn  of  the  century  complained, 
the  methods  of  Weismann,  Haeckel,  and  others  generated  theories  which  were  non- 
testable,  and  thus  could  never  be  verified  or  disproven.  As  long  as  biologists  indulged 
in  this  kind  of  theory-making,  some  younger  investigators  argued,  biology  would 
never  gain  a  solid  base  as  a  hard  science  such  as  physics  or  chemistry.  As  I  have 
shown  elsewhere  (Allen,  1978b)  T.  H.  Morgan  was  among  those  younger  biologists 
who  sought  more  experimental  and  rigorous  methods  for  pursuing  biological 
problems. 

While  Morgan  and  his  contemporaries  rejected  the  speculative  methods  of  his 
predecessors  (including  his  teacher  at  Johns  Hopkins,  W.  K.  Brooks)  they  retained 
from  them  the  unified  view  of  heredity — a  process  embodying  both  genetic  trans- 
mission and  embryonic  development.  In  1910  Morgan  wrote: 

We  have  come  to  look  upon  the  problem  of  heredity  as  identical  with  the 
problem  of  development.  The  word  heredity  stands  for  those  properties  of  the 
germ  cells  that  find  their  expression  in  the  developing  and  developed  organism. 
When  we  speak  of  the  transmission  of  characters  from  parent  to  offspring,  we 
are  speaking  metaphorically;  for  we  now  realize  that  it  is  not  characters  that  are 
transmitted  to  the  child  from  the  body  of  the  parent  but  that  the  parent  carries 
over  the  material,  to  both  parent  and  offspring  [Morgan,  1910a:  p.  449]. 

His  friend  from  both  Hopkins  and  MBL  days,  Edwin  Grant  Conklin  (1863-1952), 
held  similar  views.  Indeed,  Conklin  felt  strongly  that  the  problem  of  heredity  was 
the  central  issue  of  biology,  as  he  stated  clearly  in  1908: 

Indeed,  heredity  is  not  a  peculiar  or  unique  principle  for  it  is  only  similarity 
of  growth  and  differentiation  in  successive  generations.  ...  In  fact  the  whole 
process  of  development  is  one  of  growth  and  differentiation,  and  similarity  of 
these  in  parents  and  offspring  constitutes  hereditary  likeliness.  The  causes  of 
heredity  are  thus  reduced  to  the  causes  of  successive  differentiation  of  development, 
and  the  mechanism  of  heredity  is  merely  the  mechanism  of  differentiation 
[Conklin,  1908:  pp.  89-90]. 


GENETICS  AND  EMBRYOLOGY  109 

Conklin  introduces  his  discussion  by  claiming: 

Heredity  is  today  the  central  problem  of  biology.  This  problem  may  be 
approached  from  many  sides — that  of  the  breeder,  the  experimenter,  the  statistician, 
the  physiologist,  the  embryologist,  the  cytologist — but  the  mechanism  of  heredity 
can  be  studied  best  by  the  investigation  of  the  germ  cells  and  their  development 
[Conklin,  1908:  pp.  89-90]. 

It  is  clear  that  for  Morgan  and  Conklin,  of  all  the  approaches  to  heredity,  the 
embryological  conception  was  the  most  important  and  fruitful. 

As  part  of  his  adherance  to  the  unified  concept  of  heredity,  Morgan  attacked 
the  Mendelian  and  chromosome  theories  of  heredity  as  isolating  one  component  of 
the  hereditary  process  from  the  other,  and  thus  smacking  of  artificiality.  In  1909 
Morgan  attacked  the  Mendelian  theory  for  being  preformationist,  overly  concerned 
with  particles,  and  not  with  the  actual  process  by  which  hereditary  traits  are 
manifested  in  the  adult.  As  he  wrote: 

The  nature  of  Mendelian  interpretation  and  description  inextricably  commits 
to  the  'doctrine  of  particles'  in  the  germ  and  elsewhere.  It  demands  a  'morpho- 
logical' basis  in  the  germ  for  the  minutest  phase  (factor)  of  a  definitive  character. 
It  is  essentially  a  morphological  conception  with  but  a  trace  of  functional  feature. 
With  an  eye  seeing  only  particles  and  a  speech  only  symbolizing  them,  there  is 
no  such  thing  as  a  study  of  a  process  possible.  ...  It  has  been  possible.  I  think, 
to  show  by  means  of  what  we  know  of  the  genesis  of  these  color  characters  that 
the  Mendelian  description — of  color  inheritance  at  least — has  strayed  very  wide 
of  the  facts;  it  has  put  factors  in  the  germ  cells  that  it  is  now  quite  certainly  our 
privilege  to  remove;  it  is  declared  a  discontinuity  where  there  is  now  evident 
epigenesis  [Morgan,  1909:  p.  509]. 

In  the  following  year  he  criticized  the  chromosome  theory  in  much  the  same  terms: 

It  may  be  said  in  general  that  the  particulate  theory  is  the  more  picturesque 
or  artistic  conception  of  the  developmental  process.  As  a  theory  it  has  in  the  past 
dealt  largely  in  symbolism  and  is  inclined  to  make  hard  and  fast  distinctions.  It 
seems  to  better  satisfy  a  class  of  type  of  mind  that  asks  for  a  finalistic  solution, 
even  though  the  solution  be  truly  formal.  But  the  very  intellectual  security  that 
follows  in  the  train  of  such  theories  seems  to  me  less  stimulating  for  further 
research  than  does  the  restlessness  of  spirit  that  is  associated  with  the  alternative 
[that  is  epigenetic  or  embryological]  conception  [Morgan,  1910,  pp.  451-452]. 

What  was  telling  to  any  embryologist,  according  to  Morgan,  was  the  propensity  for 
those  adhering  to  particulate  theories  to  speak  of  adult  traits  as  if  they  actually 
resided,  in  miniature  within  the  fertilized  egg.  This  was  preformationism  at 
its  worst. 

Just  as  Morgan  opposed  localizing  the  adult  trait  in  a  particle,  hypothetical  or 
real,  he  also  opposed  focusing  largely  on  the  cell  nucleus  as  the  seat  of  all  genetic 
and  developmental  events.  Those  who  looked  to  the  nucleus  alone,  were  ignoring 
the  cytoplasm  which  "was  the  seat  of  the  really  interesting  events  [in  embryogenesis]" 
(Morgan,  1897:  p.  121).  In  these  words  Morgan  was  rejecting  the  recent  studies  by 
Boveri,  E.  B.  Wilson,  and  many  others  which  claimed  that  it  was  the  nucleus,  and 
specifically  the  chromosomes,  which  were  the  major  determiners  of  the  hereditary 
process. 

Thus,  to  Morgan,  as  an  embryologist  in  the  early  twentieth  century,  the  problems 
of  genetic  transmission  and  embryonic  development  were  inseparable.  Learning 
about  transmission  of  information  between  parents  and  offspring  was  of  no  value 


1  10  G.   E.  ALLEN 

without  also  learning  about  the  development  of  the  trait  into  its  ultimate  adult 
form.  Quite  literally,  as  Scott  Gilbert  has  emphasized  (1978),  the  Mendelian  theory 
developed  by  the  Morgan  school  began  with  solid  roots  in  the  field  of  embryology. 
After  1910,  however,  Morgan  began  to  reverse  his  position,  and  by  1926  had 
come  to  accept  (indeed,  in  many  respects,  create)  the  rigorous  separation  of  the 
problems  of  hereditary  transmission  from  those  of  embryonic  development.  In  his 
book.  The  Theory  of  the  Gene  (1926),  Morgan's  new  position  is  clear: 

Between  the  characters,  that  furnish  the  data  for  the  theory  [that  is,  Mendelian 
theory]  and  the  postulated  genes,  to  which  the  characters  are  referred,  lies  the 
whole  field  of  embryonic  development.  The  theory  of  the  gene,  as  here  formulated, 
states  nothing  with  respect  to  the  way  in  which  the  genes  are  connected  with  the 
end-product,  or  character.  The  absence  of  information  relating  to  this  interval 
does  not  mean  that  the  process  of  embryonic  development  is  not  of  interest  for 
genetics  .  .  .  but  the  fact  remains  that  the  sorting  out  of  the  characters  in 
successive  generations  can  be  explained  at  present  without  reference  to  the  way 
in  which  the  gene  affects  the  developmental  process  [Morgan,  1926:  p.  26]. 

Morgan  continued  to  believe  in  and  even  give  acknowledgment  to,  the  relationships 
between  genetic  transmission  and  embryonic  development.  But  he  did  not  know 
how  to  study  this  relationship  in  a  concrete  and  experimental  way.  Hence,  he  settled 
for  a  distinct  separation  between  the  two.  Although  Morgan  himself  attempted  to 
synthesize  the  two  fields  in  Embryology  and  Genetics  (1934),  he  was  remarkably 
unsuccessful.  But  at  least  he  admitted  the  difficulty  of  affecting  a  real  synthesis. 
When  a  colleague  told  him  of  his  disappointment  in  not  seeing  a  synthesis  between 
embryology  and  genetics  in  the  book,  Morgan  is  reported  to  have  said:  "Well,  what 
did  you  expect?  I  did  exactly  what  I  said  I  would  do  in  the  title:  I  discussed 
embryology  and  I  discussed  genetics." 

We  now  turn  to  the  question  of  what  factors  affected  Morgan's  change  from  the 
more  synthetic  and  holistic  to  the  more  analytical  and  restricted  notion  of  heredity 
between  1910  and  1925. 

FACTORS  INFLUENCING  MORGAN'S  SEPARATION  OF  GENETICS 
FROM  EMBRYOLOGY,  1910-1925 

Several  factors,  both  specific  and  general,  contributed  to  Morgan's  redefinition 
of  the  concept  of  heredity,  and  thus  his  separation  of  the  study  of  transmission 
(genetics)  from  that  of  translation  (embryology).  Among  the  most  prominent  specific 
factors  were  the  success  of  the  Mendelian  and  chromosome  theories  in  explaining 
his  own  work  with  Drosophila  (after  1910),  and  his  introduction  to  the  genotype- 
phenotype  conception  of  Wilhelm  Johannsen  in  1911.  Among  the  general  factors 
were  Morgan's  committment  to  mechanistic,  materialistic  (and  physico-chemical) 
philosophy,  his  desire  to  establish  a  new  field  of  biology  with  definite  aims  and 
boundaries,  and,  finally,  the  agricultural  revolution  in  the  United  States  that  made 
funds  available  for  genetic  (as  opposed  to  embryological)  work,  thus  giving  direction 
to  research  in  transmission  that  went  far  and  above  that  available  for  work  in  other 
areas  of  biology  such  as  embryology.  Let  us  see  how  each  of  these  factors  contributed 
to  Morgan's  change  of  position. 

First,  the  specific  factors.  When  Morgan  published  his  first  paper  on  the  white- 
eyed  mutant  Drosophila  (1910b),  he  made  a  clear  interpretation  of  its  inheritance 
pattern  in  Mendelian  terms.  He  refrained,  however,  from  associating  the  "factor" 
for  eye  color  directly  with  the  sex  (X)  chromosome.  In  the  ensuing  year,  however, 
as  more  mutants  were  discovered  and  their  patterns  of  transmission  established. 


GENETICS  AND   EMBRYOLOGY  1  1  1 

Morgan  quickly  embraced  the  chromosome  theory  as  well.  The  work  of  the  Morgan 
group  from  191 1  onward,  especially  the  study  of  sex-linked  traits  and  chromosome 
mapping,  provided  the  sort  of  material  basis  for  the  Mendelian  theory  that  had 
been  lacking,  at  least  in  Morgan's  view,  previously.  Moreover,  the  association  of 
Mendelian  breeding  results  with  the  cytological  studies  of  chromosome  behavior 
and  structure,  had  yielded  what  Lindley  Darden  and  Nancy  Maull  termed  an 
"interfield  theory"  of  great  predictive  power  (Darden  and  Maull,  1977).  Moreover, 
by  admitting  that  the  material  basis  of  heredity  resides  in  the  chromosomes,  Morgan 
was  forced  to  give  the  nucleus  a  greater  role  in  the  hereditary  life  of  the  cell  than 
he  had  previously  been  willing  to  do.  In  fact,  so  complete  was  Morgan's  change  of 
view  on  the  relative  roles  of  nucleus  and  cytoplasm  in  governing  the  process  of 
"heredity,"  that  by  1919  he  could  write  in  a  letter  to  his  friend  Jacques  Loeb, 
that  he  wanted  to  dispel  particularly  the  still-prevalent  notion  of  "cytoplasmic  in- 
heritance:" 

It  is  this  point  [cytoplasmic  inheritance]  that  I  am  anxious  to  go  for,  because 
of  its  widespread  belief  among  biologists  in  general  for  which  I  can  find  absolutely 
no  real  basis  except  an  emotional  one.  It  is  for  this  reason  mainly  that  I  have 
not  hesitated  to  hold  up  as  examples  two  of  my  best  friends  and  a  very  famous 
German  investigator  [Morgan  to  Loeb,  14  May  1919;  Loeb  Papers,  Library  of 
Congress]. 

In  the  face  of  the  extraordinary  work  on  Drosophila,  the  idea  of  cytoplasmic 
influence  on  determination  of  any  adult  traits  seemed  to  lose  all  force. 

A  second  specific  influence  that  seems  to  have  been  important  in  Morgan's 
development  was  his  explicit  recognition  of  the  genotype-phenotype  distinction  as 
examined  by  Wilhelm  Johannsen  in  his  book  of  1909,  but  especially  in  his  paper 
for  the  American  Naturalist  of  191 1.  The  content  and  polymical  nature  of  Johannsen's 
paper,  as  well  as  its  historical  significance,  have  been  well  analyzed  by  two  recent 
scholars,  Fred  Churchill  (1974)  and  Jan  Sapp  (1984).  Churchill  was  the  first  to  point 
out  that  Johannsen  made  a  distinction  between  the  horizontal  and  vertical  concepts 
of  heredity,  while  Sapp  was  the  first  to  emphasize  that  Johannsen's  paper  was  a 
specific  polemic  against  the  phenotype  conception,  which  he  claimed  was  not 
amenable  to  rigorous  experimental  or  mathematical  analysis.  To  Johannsen,  the 
phenotype  was  a  horizontal  concept,  which,  in  terms  of  Ernst  Haeckel's  biogenetic 
law,  was  essentially  an  historical  view.  That  is,  the  phenotype  was  the  result  of  a 
long  evolutionary  process  in  which  the  ontogenetic  development  of  the  individual, 
interacting  with  its  environment,  produced  the  final  appearance  of  the  adult  form. 
However,  what  was  important  to  Johannsen  was  the  underlying  mechanism  by 
which  the  phenotype  was  determined  irrespective  of  environmental  differences.  The 
genotype  concept  was  thus  a  vertical,  and  purposely  ahistorical,  view.  To  Johannsen, 
trained  as  a  chemist,  Mendelian  genes  had  to  be  regarded  as  unchangeable  entities, 
analogous  to  the  atoms  of  chemistry.  Just  as  atoms  combine  and  recombine  into 
various  molecules,  but  nonetheless  retain  their  individual  properties,  so  too  could 
genes  combine  and  recombine  in  different  genotypes  in  successive  generations,  and 
yet  retain  their  own  individuality.  A  gene  for  white  eye  remained  a  gene  for  white 
eye  even  when  masked  for  a  generation  or  two  with  a  dominant  gene  for  red  eye. 
To  Johannsen,  "the  fundamental  nature  of  heredity  lay  hidden  deep  within  the 
gamete."  Only  by  the  use  of  analytical  and  experimental  methods  could  the  nature 
of  heredity  be  discovered,  and  the  laws  used  for  predictive  purposes.  Johannsen 
made  a  clear  and  explicit  separation  between  the  field  of  heredity  and  that  of 
embryology.  Heredity  passes  through  the  germ  line  only,  in  accordance  with 


112  G.   E.   ALLI-N 

Weismann's  germ-plasm  theory,  and  deals  only  with  the  genotype.  Embryology,  on 
the  other  hand,  embodies  the  influence  of  environment  as  well  as  the  whole  past 
history  of  the  species,  and  thus  deals  only  with  the  phenotype.  To  make  heredity 
amenable  to  experimental,  analytical,  and  mathematical  methods,  Johannsen  argued 
that  the  proper  sphere  of  study  was  the  genes  within  the  gametes,  and  not  the  final 
expressed  potential  of  the  adult  phenotype. 

It  is  clear  that  Morgan  must  have  learned  about  Johannsen's  genotype-phenotype 
distinction  by  late  1910  or  early  1911.  Both  men  were  present  at  the  Princeton 
meeting  of  the  American  Society  of  Naturalists  on  19  December  1910,  where  they 
participated  in  a  symposium  entitled  "Study  of  Pure  Lines  of  Genotypes/'  Morgan, 
Johannsen,  and  a  number  of  other  biologists  addressed  the  genotype  conception  of 
heredity  and  evaluated  this  notion  for  the  present  study  of  genetics.  At  this 
conference,  Johannsen  gave  his  now-famous  paper,  "The  Genotype  Conception  of 
Heredity,"  (Johannsen,  1911),  and  Morgan  spoke  on  sex-limited  inheritance  and 
sexual  dimorphism  (Morgan,  1911).  Furthermore,  Johannsen  was  invited  to  spend 
part  of  the  winter  term  of  191 1  at  Columbia,  where  it  is  highly  unlikely  that  he  and 
Morgan  would  not  have  met  and  discussed  their  mutual  interests  in  the  problems 
of  the  emerging  new  field  of  genetics.  As  early  as  1913  Morgan  makes  reference  to 
Johannsen's  work  in  Heredity  in  Sex  (Morgan,  1913)  and  again  similar  references 
appear  in  Mechanism  of  Me ndelian  Heredity  (Morgan  el  al,  1915).  Although 
Morgan  does  not  say  it  in  so  many  words,  I  think  it  is  safe  to  infer  that  Johannsen's 
phenotype-genotype  distinction  provided  an  important  conceptual  foundation  for 
Morgan's  growing  awareness  of  the  distinction  that  could  be,  and  needed  to  be 
made,  between  genetics  and  embryology. 

Among  the  more  general  factors  influencing  Morgan's  separation  of  genetics 
and  embryology  was  his  long-standing  committment  to  the  philosophy  of  mechanistic 
materialism.  Morgan  had  enthusiastically  embraced  the  new  mechanistic  biology 
beginning  as  early  as  1891  upon  first  meeting  the  German-born  mechanistic 
philosopher  Jacques  Loeb  when  they  both  joined  the  faculty  of  the  biology 
department  at  Bryn  Mawr  College  that  year.  Later,  as  a  result  of  his  visits  to  the 
Stazione  Zoologica  in  Naples,  and  particularly  his  association  there  with  another 
German,  Hans  Driesch,  Morgan's  mechanistic  views  were  renewed  and  strengthened 
with  new  experimental  embryology.  To  Morgan,  the  mechanistic  materialist  philos- 
ophy meant  belief  in  the  material  existence  of  the  world,  its  structure  in  terms  of 
separate  and  separable  components,  and  the  method  of  analysis,  in  which  complex 
processes  could  be  broken  down  into  their  simpler  components  and  studied 
independently,  under  controlled  conditions.  This  meant,  of  course,  using  rigorous 
experimental  methods,  and,  at  least  in  its  early  twentieth-century  form,  a  committ- 
ment to  a  kind  of  physico-chemical  reductionism.  Morgan  was  not  as  highly 
reductionist  as  Loeb,  for  he  always  had  a  feel  for  the  whole  organism  that  prevented 
him  from  embracing  naive  views  equating  the  organism  with  a  machine,  or  a  "bag 
of  enzymes."  The  key  element  in  Morgan's  mechanistic  materialist  philosophy,  at 
least  for  understanding  his  willingness  to  go  against  his  own  earlier  beliefs  and 
separate  genetics  and  embryology,  is  analysis.  To  Morgan,  experiments  only  made 
sense  in  an  analytical  framework.  If  conditions  could  not  be  controlled,  and  the 
effects  of  complex  systems  could  not  be  studied  in  isolation,  then  rigorous  biology 
was  not  possible.  It  was  essential  to  take  complex  processes  and  break  them  down 
into  their  component  parts.  The  older,  more  inclusive  concept  of  heredity  was  thus 
not  amenable  to  mechanistic  analysis.  No  satisfactory  experimental  techniques  had 
been  developed  to  isolate  and  study  the  process  of  embryonic  differentiation — how 
it  was  controlled  at  the  cellular,  tissue,  and  organ-system  level.  However,  with  the 


GENETICS  AND   EMBRYOLOGY  113 

advent  of  the  Drosophila  work,  at  least  one  component  of  the  broad  view  of 
heredity,  namely,  transmission  from  parent  to  offspring,  suddenly  became  amenable 
to  experimental  and  analytical  methods.  Though  Morgan  may  have  been  predisposed 
to  retain  the  broader  definition  of  heredity,  the  newer  techniques,  compatible  as 
they  were  with  his  basic  philosophical  orientation  toward  experimentation,  dictated 
separating  the  broad  definition  into  its  simpler  parts.  The  result  was  that  Morgan 
focused  on  the  transmission  side  of  heredity  for  the  next  fifteen  years. 

Another  general  factor  stimulating  the  rapid  growth  of  the  Mendelian  chromosome 
theory,  as  opposed  to  embryology,  was  the  American  agricultural  revolution  taking 
place  around  the  turn  of  the  century.  In  the  post-Civil  War  period,  a  number  of 
changes  had  occurred  in  United  States'  economic  and  social  organization  which 
directly  affected  agricultural  production.  The  most  important  was  the  rapid  devel- 
opment of  industry,  which  led  to  migration  of  workers  from  the  farms  and  rural 
areas  to  the  great  centers  of  industrial  production.  The  resulting  urbanization  created 
a  great  demand  for  food,  but  at  the  same  time  meant  there  were  now  fewer  hands 
working  the  land.  By  the  turn  of  the  century,  major  financiers  had  turned  their  eyes 
toward  the  profitability  of  managed,  large-scale  agriculture.  This  meant,  among 
other  things,  the  development  of  new,  faster-growing  or  higher-yielding  crops. 
"Scientific  agriculture"  was  not  new  at  this  time,  but  the  form  in  which  science  was 
applied  to  agriculture  had  begun  to  change. 

In  the  latter  half  of  the  nineteenth  century,  from  Justes  von  Liebig  onward, 
much  research  had  been  put  into  what  was  loosely  called  "agricultural  chemistry." 
This  field  involved  the  scientific  study  of  soils,  the  development  of  natural  and 
chemical  fertilizers,  the  production  of  animal  feeds,  attention  to  animal  and  plant 
nutrition,  etc.  While  these  efforts  had  yielded  some  major  increases  in  productivity, 
by  the  late  nineteenth  century  they  had  reached  a  limit  in  both  extent  and 
profitability.  Fertilizers  or  feed  additives,  for  instance,  had  to  be  added  continually 
to  keep  up  production  levels.  This  was  not  so,  however,  with  scientific  breeding. 
The  results  of  good  breeding  had  a  very  different  economic  potential.  In  1910  U.  S. 
Secretary  of  Agriculture,  James  Wilson,  argued  openly  for  the  study  of  heredity  as 
a  way  of  developing  a  more  economically  profitable  agriculture.  Writing  in  the 
opening  pages  of  the  newly  founded  American  Breeders  Magazine,  Wilson  noted 
that  both  fertilizers  and  animal  nutritive  feeds  must  be  reapplied  year  after  year  to 
have  the  desired  effects,  whereas  the  hereditary  effects  obtained  through  scientific 
breeding  were  self  perpetuating:  "Heredity  is  a  force  more  subtle  and  more 
marvelous  than  electricity.  Once  generated  it  needs  no  additional  force  to  sustain 
it.  Once  new  breeding  values  are  created  they  continue  as  permanent  economic 
forces."  (Wilson,  1910:  p.  5).  Wilson  then  goes  on  to  make  his  point  more  explicit. 

But  the  cost  of  improvements  through  breeding  usually  represents  only  a 
small  fraction  of  the  added  values.  The  increase  of  products  secured  pays  the 
price  in  a  short  time,  and,  since  there  is  no  further  expense,  the  annual  increase 
afterward  is  clear  profit.  The  farmer  will  be  able  to  retain  a  part  of  the  larger 
production  in  the  form  of  added  profit  and  part  will  help  to  reduce  the  cost  of 
living  to  those  in  the  cities.  Larger  production  on  the  farm  will  also  give  increased 
business  to  the  transportation  company,  the  manufacturer,  and  the  merchant, 
and  will  provide  the  nation  the  larger  product  with  which  to  hold  our  balance  of 
trade.  [Ibid.] 

Wilson's  enthusiasm  was  not  mere  political  rhetoric.  There  was  a  widespread 
belief,  partly  catalyzed  by  the  rediscovery  of  Mendel's  laws  in  1 900,  that  the  science 
of  breeding  was  off  to  a  new  and  momentous  start.  As  Charles  Rosenberg  has 


1  14  G.   E.   ALLEN 

shown,  the  new  genetics  found  an  especially  warm  welcome  in  many  state  agricultural 
experiment  stations,  and  agricultural  schools  (Rosenberg,  1976a,  b).  And,  though 
the  payoffs  were  not  always  as  dramatic  as  Secretary  Wilson  and  others  initially 
imagined,  the  new  genetics  did  contribute  profoundly  to  the  development  of 
agriculture  in  the  United  States.  The  work  of  investigators  such  as  Donald  Jones 
(Connecticut  Agricultural  Station),  E.  M.  East  and  later  Paul  Mangelsdorf  (Harvard's 
Bussey  Institution,  devoted  to  practical  and  ornamental  breeding),  Charles  Zeleny 
(University  of  Illinois),  L.  J.  Stadler  and  Barbara  McClintock  (University  of 
Missouri),  and  R.  A.  Emerson  at  Cornell  (initially  a  state  agricultural  college)  was 
all  carried  out  in  a  specific  agricultural  context,  and  in  many  cases  led  directly  to 
some  practical  agricultural  gains  (hybrid  corn  being  one  of  the  most  notable). 

That  all  of  this  was  not  lost  on  genetics,  and  even  Morgan  in  particular,  is 
evidenced  by  the  fact  that  in  1918  E.  B.  Babcock  and  R.  E.  Clausen  published  one 
of  the  first  and  most  widely  used  applied  genetics  texts,  Genetics  in  Relation  to 
Agriculture,  dedicated,  appropriately,  to  Morgan.  In  the  Introduction  to  that  book 
Babcock  and  Clausen  state  clearly  the  economic  importance  of  applying  known 
genetic  principles  to  agricultural  breeding: 

Of  all  the  sciences  that  contribute  to  the  great  .  .  .  composite  which  is  known 
as  agriculture  none  is  more  important  economically  than  genetics.  .  .  .  Without 
doubt  vast  possibilities  await  realization  through  the  more  thorough  and  systematic 
development  of  our  living  economic  resources.  Such  development  is  directly 
dependent  on  the  successful  utilization  of  genetic  principles  in  plant  and  animal 
breeding.  The  science  of  genetics  is  still  very  young,  but  it  is  firmly  established 
and  is  developing  rapidly.  It  claims  the  attention  of  the  producer  of  today  and 
invites  the  most  serious  study  of  the  agriculturists  of  tomorrow  [Babcock  and 
Clausen,  1918:  pp.  vii]. 

That  Morgan  agreed  with  these  principles  is  clearly  demonstrated  by  a  memorandum 
that  he  sent  to  various  geneticists  around  this  same  time,  outlining  plans  for  a 
department  of  genetics  at  Columbia  University.  In  the  opening  paragraphs  of  that 
outline,  Morgan  stated: 

Until  within  recent  years  scientific  agriculture  has  to  do  almost  solely  with 
the  feeding  of  plants  and  animals.  This  condition  arose  from  the  fact  that  the 
persons  who  first  became  interested  in  developing  a  science  of  agriculture  were 
chemists.  To  increase  the  productiveness  of  domesticated  plants  and  animals  by 
the  use  of  fertilizers  and  properly  proportion  rations  has  been  the  goal  of  the 
great  bulk  of  scientific  work  in  agriculture.  It  is  now  evident,  however,  that  this 
is  only  one  side,  and  fundamentally  the  least  important  side,  of  the  matter.  What 
an  animal  or  a  plant  produces  is  fundamentally  determined  by  what  that  plant 
or  animal  is.  The  innate  hereditary  constitution  of  the  individual  and  the  race  is 
the  basis  on  which  all  improved  feeding  and  fertilizing  must  end.  The  science  of 
genetics  (or  breeding)  is  fundamental  for  all  agriculture  [From  Morgan  Folder, 
Raymond  Pearl  Papers,  American  Philosophical  Society  Library;  no  date  on 
manuscript]. 

It  must  be  remembered  that  Morgan  was  also  one  of  the  early  members  of  the 
American  Breeders  Association,  and  attended  at  least  two  of  their  meetings  (St. 
Louis,  Missouri,  December,  1903,  and  Columbia,  Missouri,  January,  1909)  (Kim- 
melman,  1983:  p.  194). 

Funding  patterns  in  the  first  decade  of  the  twentieth  century  suggest  that 
considerably  more  money  was  becoming  available  for  agriculture-related  research 


GENETICS  AND   EMBRYOLOGY  115 

than  had  been  true  in  previous  years.  For  example,  the  United  States  Department 
of  Agriculture  and  the  State  Agricultural  Experiment  Station  were  both  centers  for 
Mendelian  research  after  1901.  In  addition,  many  states  were  developing  agricultural 
experiment  stations  of  their  own  where  none  had  existed  before  (Rosenberg,  1976a, 
b).  And,  private  foundations  turned  toward  agriculture.  In  setting  funding  priorities 
for  the  newly  established  Carnegie  Institution  of  Washington  (1902),  Andrew 
Carnegie  was  particularly  interested  in  supporting  the  work  of  Luther  Burbank.  His 
reasons  were  explicitly  that  Burbank's  much  publicized  new  methods  of  grafting 
would  have  important  economic  results.  At  almost  the  same  time  the  Carnegie 
Institution  also  funded  Charles  B.  Davenport's  Station  for  the  Experimental  Study 
of  Evolution  at  Cold  Spring  Harbor,  founded  in  1904  with  an  initial  grant  of 
$34,250.  By  1918  the  annual  budget  for  the  Station  had  risen  to  $60,000,  and  by 
1935  to  $115,000.  While  all  of  the  research  carried  out  at  the  Station  was  not  all 
directly  agricultural,  Davenport's  initial  goal  and  continued  purpose  always  remained 
to  study  the  factors  influencing  heredity  and  breeding  in  a  variety  of  animals  and 
plants.  And,  after  1915,  Morgan's  work  with  Drosophila  was  funded  by  the  Carnegie 
Institution  of  Washington  (starting  with  annual  grants  of  $3,600  between  1915  and 
1919,  and  jumping  to  an  annual  average  of  $12,000  from  1920  through  1934). 

The  funding  for  embryology,  in  contrast,  was  significantly  less.  The  Carnegie 
Institution  of  Washington  made  its  first  grant  for  embryology  in  1913,  to  F.  P.  Mall 
at  the  Anatomical  Institute  of  Johns  Hopkins  University,  for  a  total  of  $15,000. 
Perhaps  more  than  anything  else  this  comparison  suggests  why  pursuit  of  the  study 
of  hereditary  transmission  had  a  certain  immediacy  to  it  which  the  study  of 
embryology  lacked. 

I  suggest,  therefore,  that  around  the  turn  of  the  century  there  was  more  than 
passing  academic  or  intellectual  interest  in  the  study  of  heredity.  Hereditary 
transmission  had  a  practical,  economic  imperative  which  perhaps  encouraged  its 
isolation  from  other  related  problems  such  as  embryonic  development.  It  was  this 
atmosphere  that  may  have  given  some  impetus,  even  indirectly,  to  the  redefinition 
of  heredity  effected  by  Morgan  and  his  followers  after  1910.  Let  me  emphasize 
clearly,  however,  that  I  am  not  suggesting  that  Morgan  or  other  Mendelian  geneticists 
raced  toward  the  study  of  genetic  transmission  merely  for  financial  gain.  I  am 
suggesting  that  the  availability  of  funds  does  influence  the  direction  for  research 
programs  when  the  programs  can  be  carried  out  with  new  techniques  and  concepts. 
The  important  point  I  am  trying  to  raise  here  is  that  the  context  for  determining 
the  availability  of  funds  was  not  merely  the  academic  and  intellectual  interest 
inherent  in  one  or  another  field.  It  was  more  related  to  the  social  and  economic 
imperatives  present  in  the  society  at  large,  which  encouraged  funding  in  certain 
areas  to  a  far  greater  extent  than  others. 

At  a  more  sociological  level,  the  splitting  of  transmission  genetics  from  embryonic 
development  had  an  important  practical  consequence  within  the  scientific  community. 
It  allowed  Morgan,  his  immediate  followers,  and  others  who  took  up  the  Mendelian- 
chromosome  theory,  to  define  a  new  and  separate  field  of  investigation.  Between 
1910  and  1925  the  Mendelian-chromosome  theory  became  what  Irmre  Lakatos 
calls  a  full-fledged  research  program  (Lakatos,  1970).  Lakatos  has  emphasized  that 
research  programs  not  only  consist  of  concepts  but  also  of  methods  of  research  (in 
this  case,  for  example,  breeding  coupled  with  cytological  observations),  standard 
protocols  (for  example  the  use  of  pure  strains  for  breeding,  or  the  correlation 
between  breeding  and  cytological  data),  and  philosophical  methods,  including  the 
notion  of  what  is  proper  explanation  in  a  field  (for  example,  mechanistic  vs.  holistic 


116  G.   E.   ALLEN 

interpretation;  or  the  role  of  quantitative  and  mathematical  thinking  in  scientific 
explanation).  By  establishing  a  research  program,  Lakatos  points  out,  scientific 
workers  define  their  fields  and  problems,  thus  influencing  the  direction  of  future 
research  and  the  development  of  field-wide  methods  for  dealing  with  challenges  to 
accepted  orthodoxy.  For  example,  Morgan  and  his  group  not  only  rigorously  defined 
the  problems  which  were  to  become  the  future  focus  of  genetic  research,  but  also 
established  an  orthodoxy  which  focused  almost  exclusively  on  the  cell  nucleus  as 
the  center  of  heredity.  Any  attempts  to  discuss  cytoplasmic  inheritance  (or  what 
was  sometimes  referred  to  as  "maternal  effects")  were  strongly  discouraged.  Jan 
Sapp  has  studied  the  history  of  this  subject  exhaustively,  and  has  shown  how  those 
who  sought  to  publish  on  cytoplasmic  inheritance,  such  as  Tracy  Sonnenborn  and 
later  his  student,  David  Nanney,  at  first  found  their  papers  rejected  by  orthodox 
genetic  journals  (Sonneborn,  1978). 

Sapp  applied  the  idea  of  struggle  for  authority  among  competing  fields,  as 
developed  by  French  sociologist  Pierre  Bourdieu  (1975),  specifically  to  the  case  of 
nuclear  versus  cytoplasmic  genetics  in  the  early  twentieth  century  (Sapp,  1984). 
Bourdieu's  idea  is  that  scientific  fields,  or  research  programs  in  the  Lakatosian 
sense,  are  in  competition  with  other  fields  for  money,  students,  and  the  opportunity 
to  control  academic  or  research  positions.  The  competition  is  most  keen  among 
closely  related  fields,  but  exists  to  one  degree  or  another  between  all  fields.  In 
Bourdieu's  model,  scientific  authority,  or  competence,  is  understood  as  the  socially 
recognized  legitimacy  of  the  individual  to  speak  and  act  on  scientific  matters.  The 
content  of  scientific  ideas  is  thus  seen  as  related  to  the  social  reality  of  establishing 
a  professional  niche,  that  is,  a  research  program.  Thus,  Bourdieu  sees  the  choice  for 
pursuing  certain  theories  over  others,  as  well  as  the  manner  in  which  the  theories 
are  put  forward,  as  an  integral  part  of  the  social  context  among  competing  fields. 

Applied  to  the  development  of  genetics  and  its  separation  from  embryology, 
Bourdieu's  idea  suggests  the  following  scenario:  as  Mendelian  genetics  began  to  have 
some  success  dealing  with  the  process  of  transmission,  it  became  advantageous  to 
begin  determining  the  boundaries  of  the  new  field — that  is,  to  establish  its  problems 
and  its  scope.  This  meant  for  Morgan  and  his  group  that  it  became  increasingly 
advantageous  to  eliminate  from  the  study  of  heredity  itself  the  knotty  problems  of 
embryonic  development  with  which  Morgan  himself  (and  others)  had  had  little 
experimental  success.  By  so  doing,  Morgan  was  able  to  outline  what  appeared  to  be 
a  successful  and  easily  approachable  field  of  scientific  endeavor.  Had  he  insisted  on 
working  simultaneously  with  the  problems  of  the  development  of  phenotype  and 
transmission  of  the  genotype,  it  is  doubtful  that  the  field  could  have  developed  in 
any  clear-cut  way.  Although  he  himself  never  renounced  either  his  interest  in 
embryology,  or  his  belief  that  the  Mendelian  gene  ultimately  had  to  be  interpreted 
in  embryological  terms,  Morgan  was  pragmatic  enough  to  see  the  advantage  of 
pushing  embryology  aside  for  the  time  being.  Developing  the  new  field  of  Mendelian 
genetics  with  a  strong  central  focus  (transmission,  assortment,  and  recombination), 
a  set  of  research  techniques,  and  most  importantly  some  clear  and  immediate 
results,  Morgan  was  able  to  attract  attention,  students,  and  (ultimately)  research 
money  in  a  way  that  would  have  been  impossible  had  he  insisted  on  studying 
heredity  in  the  older,  more  holistic  way.  Morgan  thus  drew  a  boundary  between  the 
new  field  of  genetics  and  the  old  field  of  embryonic  development.  Everything  within 
the  boundary  was  included  in  the  new  research  program;  those  who  would  try  to 
force  upon  the  Mendelian-chromosome  theory  the  burden  of  explaining  embryonic 
development  were  told  to  become  (or  remain)  "embryologists." 


GENETICS  AND   EMBRYOLOGY  117 

In  putting  forth  this  analysis  I  do  not  wish  to  suggest  that  Morgan  or  his  group 
made  such  choices  consciously,  or  were  acting  in  a  particularly  ruthless  or  opportunist 
way  to  exclude  certain  topics  from  their  new  research  program.  I  do  want  to  suggest, 
however,  that  the  conscious  and  subconscious  aspects  of  what  it  means  to  establish 
a  new  field  of  research  and  to  gain  the  recognition,  money,  and  students  which  can 
result,  is  not  a  negligible  factor  in  what  constitutes  the  formation  of  a  scientific 
research  program.  After  all,  research  programs  are  more  than  merely  good  ideas. 
They  involve  techniques,  equipment,  laboratories,  people,  the  desire  for  individual 
recognition,  and  money,  all  of  which  have  some  direct  influence  on  the  content 
and  direction  of  the  scientific  ideas  themselves. 


ANALYSIS  AND  CONCLUSION 

To  return  to  the  beginning  of  our  paper,  the  question  still  remains:  why  did 
genetics  not  enter  the  mainstream  of  MBL  courses  or  research?  If  the  above  analysis 
is  correct — namely,  that  genetics  was  highly  favored  by  funding  patterns  and  by  its 
adherence  to  mechanistic  and  experimental  and  analytical  lines — it  would  have 
seemed  all  the  more  likely  that  it  would  have  played  a  prominent  role  at  the  MBL. 
Here,  I  think,  institutional  factors  peculiar  at  the  MBL  in  the  1920s  and  '30s  played 
a  major  part. 

From  its  inception,  the  MBL  had  been  a  bastion  of  embryology  and  the 
attendant  holistic  definition  of  heredity  inhertied  from  the  nineteenth  century. 
C.  O.  Whitman,  founder  and  first  Director  of  the  MBL,  was  a  morphologist  and 
Haeckelian  while  pioneering  the  newer  methods  of  cell  lineage  and  experimentation. 
In  addition,  several  of  Morgan's  contemporaries  at  the  MBL  objected  to  the  new 
"genotypic"  definition  of  heredity  which  Morgan  was  championing.  In  1916  Jacques 
Loeb  pointed  out  the  difficult  implications  for  embryology  of  the  new  work  in 
heredity. 

The  difficulties  besetting  the  biologist  in  this  problem  [harmonious  interaction 
of  parts  of  an  organism]  have  been  rather  increased  than  diminished  by  the 
discovery  of  Mendelian  heredity,  according  to  which  each  character  is  transmitted 
independently  of  any  other  character.  Since  the  number  of  Mendelian  characters 
in  each  organism  is  large,  the  posibility  must  be  faced  that  the  organism  is  merely 
a  mosaic  of  independent  hereditary  characters.  If  this  be  the  case  the  question 
arises:  What  moulds  these  independent  characters  into  a  harmonious  whole? 
[Loeb,  1916:  pp.  v-vi] 

A  decade  later  F.  R.  Lillie,  then  Director  of  the  MBL,  clearly  pointed  out  his 
objections  to  the  new  Mendelian  conception  of  the  gene  with  respect  to  embryological 
processes: 

I  do  not  know  of  any  sustained  attempt  to  apply  the  modern  theory  of  the 
gene  to  the  problem  of  embryonic  segregation.  As  the  matter  stands,  this  is  one 
of  the  most  serious  limitations  of  the  theory  of  the  gene  considered  as  a  theory 
of  the  organism.  We  should,  of  course,  be  careful  to  avoid  the  implication  that 
in  its  future  development  the  theory  of  the  gene  may  not  be  able  to  advance  into 
this  unconquered  territory.  But  I  do  not  see  any  expectation  that  this  will  be 
possible,  even  in  principle,  as  long  as  the  theory  of  the  integrity  of  the  entire 
gene  system  [i.e.,  that  all  genes  are  present,  or  at  least  active]  in  all  cells  is 
maintained.  If  this  is  a  necessary  part  of  the  gene  theory,  the  phenomena  of 


118  G.   E.   ALLEN 

embryonic  segregation  must,  I  think,  lie  beyond  the  range  of  genetics  [Lillie, 
1927:' p.  366]. 

Lillie  even  went  on  to  claim  that  he  did  not  foresee  a  future  synthesis  possible — at 
least  in  the  late  nineteenth  century  (by  which  he  meant  Weismannian)  sense — given 
the  new  view  of  geneticists  about  what  constituted  heredity  (Lillie,  1927:  p.  367). 
Another  critic  of  the  Morgan  school  was  Morgan's  old  friend  and  colleague,  both 
from  Woods  Hole  and  Johns  Hopkins  days,  Ross  G.  Harrison.  Harrison  wrote  in 
1937  that  the  new  gene  theory,  as  prestigious  as  it  seemed  to  be,  was  much  too 
one-sided.  It  focused  only  on  the  problem  of  what  hereditary  potentialities  were 
passed  from  parent  to  offspring,  and  failed  to  deal  in  any  significant  way  with  the 
embryological  issues  (Harrison,  1973:  p.  372).  Beyond  this,  Harrison  probed  more 
deeply  at  the  philosophical  foundations  of  the  split.  He  argued  that  geneticists  were 
too  atomistic,  while  embryologists  sought  a  more  holistic  interpretation  of  the 
hereditary  process: 

The  embryologist,  however,  is  concerned  more  with  the  larger  changes  in  the 
whole  organism  and  its  primitive  systems  of  organs  than  with  the  lesser  qualities 
known  to  be  associated  with  gene  actions.  As  Just  remarked  ...  he  is  more 
interested  in  the  back  [of  a  fruit  fly]  than  in  the  bristles  on  the  back,  and  more 
in  the  eyes  than  the  eye  color  [Harrison  1937:  p.  372]. 

In  addition  to  classical  embryologists  such  as  Lillie  and  Harrison,  another  group, 
namely  those  concerned  with  the  problem  of  cytoplasmic  inheritance  reacted  against 
the  Morgan  definition  of  heredity.  As  Jan  Sapp  pointed  out,  both  Herbert  Spencer 
Jennings  and  more  particularly  his  student  Tracy  Sonneborn  found  the  definition 
of  heredity  associated  with  the  Morgan  school  far  too  limiting.  In  his  autobiography 
Sonneborn  described  himself  as  a  "lifelong  critic  of  what  seemed  to  be  a  blind  and 
erroneous  faith  in  the  gene  as  a  source  of  all  heredity"  (Sonneborn,  1978:  p.  1).  In 
the  same  paragraph  he  referred  to  the  transmission  conception  of  heredity  as  "a 
stifling  dogma."  Focusing  as  much  attention  as  it  did  on  the  nucleus,  and  particularly 
the  chromosomes,  the  Mendelian-chromosome  theory,  as  defined  by  Morgan,  left 
little  room  for  nucleo-cytoplasmic  interaction.  Viktor  Hamburger  has  pointed  out 
that  as  late  as  1951  the  effects  of  this  rigid  separation  (between  genotype  and 
phenotype  and  between  nucleus  and  cytoplasm)  were  felt  by  Belgian  embryologist 
Albert  Dalcq.  Arguing  that  classical  genetics  ignored  the  important  role  that 
cytoplasm  as  "an  organized  system"  played  in  the  process  of  differentiation,  he 
wrote, 

This  notion  [of  pattern  in  the  cytoplasm  and  of  the  importance  of  the  whole], 
so  intimately  tied  to  a  pattern,  is  lacking  in  the  system  of  concepts  used  by 
geneticists  .  .  .  [These]  are  based  on  a  particularistic,  atomistic  viewpoint  which 
neglects,  despite  everything,  this  other  factor  which  resides  in  the  totality  of  the 
organization  [Dalcq  (1951):  p.  135;  translated  by  Hamburger]. 

With  this  sort  of  a  general  objection  widely  felt  among  embryologists  it  is  no 
wonder  that  the  MBL  was  not  a  likely  atmosphere  for  the  development  of  genetics 
as  a  new  field.  Add  to  this  the  fact  that  Morgan  did  not  like  to  bother  himself  with 
teaching,  and  was  undoubtedly  making  no  efforts  to  develop  a  genetics  course  on 
his  own  at  MBL,  the  atmosphere  at  the  Laboratory  was  simply  not  conducive  to 
the  new  Mendelian  genetics  as  an  official  Woods  Hole  research  or  teaching  program. 
In  conclusion,  then,  what  I  want  to  emphasize  is  that  the  development  of 
genetics  as  a  field  separate  from  the  older,  more  inclusive  notion  of  heredity,  ended 


GENETICS   AND  EMBRYOLOGY  119 

up  separating  the  study  of  genetics  from  the  study  of  embryology.  To  summarize, 
the  separation  was  the  result  of  several  interacting  factors: 

( 1 )  A  pervasive  commitment  among  biologists — especially  T.  H.  Morgan  and 
his  school — to  mechanistic  materialism  and  its  associated  analytical  methods  by 
which  complex  problems  and/or  processes  are  broken  down  into  their  simpler 
components; 

(2)  the  conscious  awareness  of  biologists  of  the   useful  distinction   Wilhelm 
Johannsen  had  made  between  the  genotype  and  the  phenotype,  corresponding  as  it 
did  to  the  distinction  between  genetics  and  embryology; 

(3)  the  rapid  and  exciting  development  of  the  Mendelian  work  with  the  fruit  fly 
Dmsophila  melganogaster;  Drosophila  was  an  extremely  favorable  organism  for  the 
study  of  Mendelian  and  chromosome  transmission; 

(4)  the  competition  between  fields  which  makes  the  delineation  of  separate 
disciplines  advantageous,  especially  to  a  new  field  trying  to  establish  its  own  identity, 
its  own  areas  of  research  focus,  its  own  funding,  and  its  own  students; 

(5)  and  last  and  perhaps  most  important,  the  agricultural  context  in  which  the 
study  of  how  (and  in  what  pattern)  traits  are  passed  on  from  one  generation  to 
another  seeing  as  being  greater  economic — and  I  mean  by  that  consciously  profit — 
gain  is  viewed  as  more  important  than  how  traits  develop  from  fertilized  egg  to 
adult.   It  was  this  agricultural   imperative  that  translated   not  only  into  greater 
enthusiasm  and  optimism  about  the  potential  which  genetics  held,  but  also  into 
greater  financial  and  institutional  support. 

Thus,  what  started  as  a  paradox  about  the  specific  institutional  history  of  the 
MBL,  now  can  be  seen  as  part  of  a  larger  development  both  inside  and  outside  the 
biological  community.  The  widening  gap  between  genetics  and  embryology,  both 
conceptually  and  in  terms  of  rapidity  of  new  advances  in  the  1920s  and  "30s,  was 
thus  a  result  of  a  number  of  converging  factors  during  the  first  several  decades  of 
the  century.  T.  H.  Morgan,  an  important  figure  at  the  MBL  during  this  period, 
played  a  major  role  in  creating  that  widening  gap.  That  his  own  most  exciting  work 
did  not  find  a  more  hospitable  a  home  in  the  MBL  is  due  to  both  institutional  and 
wider  intellectual/philosophical  factors — to  the  difference  in  the  holistic  world  view 
of  the  embryologists  and  the  mechanistic,  analytical  world  view  of  the  geneticists. 
Such  is  the  stuff  of  which  scientific  history  is  made. 

ACKNOWLEDGMENTS 

An  earlier  version  of  this  paper  was  prepared  for  the  British  Society  for 
Developmental  Biology's  annual  meeting  at  Nottingham,  England,  in  April,  1983. 
I  am  much  indebted  to  the  British  Society,  and  to  Dr.  Timothy  Horder,  for  the 
opportunity  to  put  these  ideas  together  for  the  first  time.  In  addition,  a  number  of 
people  have  contributed  substantially  to  my  own  thinking,  either  directly  or 
indirectly,  during  the  preparation  of  the  paper:  Viktor  Hamburger,  Jan  Sapp,  Scott 
Gilbert,  and  Barbara  Kimmelman. 

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Reference:  Biol.  Bull.  168  (suppl.):  122-126.  (June,  1985) 


COMPARATIVE   PHYSIOLOGY   AND   BIOCHEMISTRY   AT  THE 
ZOOLOGICAL  STATION   OF  NAPLES 

FRANCESCO  GHIRETTI 

Department  of  Biology,  University  of  Padova,  Italy 

The  Zoological  Station  of  Naples  was  planned  and  realized  in  1872  as  a  research 
institute  for  zoology  and  morphology.  Its  founder  and  director  Anton  Dohrn  was  a 
zoologist  and  morphologist;  the  first  assistants  were  zoologists  Nicholaus  Kleinenberg 
and  Hugo  Eisig.  Zoologists  and  morphologists  were  the  first  guests  of  the  new 
Institute:  von  Waldeyer-Hartz,  Francis  Balfour,  Ray  Lankester,  August  Weismann, 
Giovanni  Battista  Grassi,  Antonio  Delia  Valle,  Oscar  Schmidt,  Willem  Hiibrecht, 
and  others.  The  three  publications  issued  by  the  Station,  Mittheilungen  der  Zoolo- 
gischen  Station  in  Neapel,  Zoologischer  Jahresbericht,  and  Fauna  und  Flora  des 
Golfes  von  Neapel  dealt  with  zoology  and  botany.  In  1876  Anton  Dohrn  added  a 
section  of  Botany,  and  botanists  also  were  among  the  first  guests. 

Research  projects  reflected  the  origin  of  the  scientists  themselves,  but  morpho- 
logical studies  dominated.  Dohrn's  original  interest  was  morphology  of  the  inverte- 
brates and  comparative  embryology.  In  1875  he  published  a  book,  Urspriing  der 
Wirbeltiere  und  das  Prinzip  des  Functionswechsel  which  was  the  basis  for  his  25 
subsequent  publications  under  the  general  title:  Studies  on  the  Origin  of  the 
Vertebrate  Body.  Later  his  main  research  interest  centered  around  the  problem  of 
explaining  the  structure  of  the  vertebrate  head. 

Although  a  morphologist  until  the  end  of  his  life,  Dohrn  had  an  unusual  feeling 
for  the  significance  of  new  scientific  currents,  especially  the  developing  branches  of 
physiology,  comparative  physiology,  and  physiological  chemistry.  Taking  account 
of  the  rapid  development  of  these  disciplines  which  had  sprouted  in  the  early  1800's 
under  the  leadership  of  Jean  Baptiste  Dumas  and  Jean  Baptiste  Boussingault  in 
France,  and  Johannes  Miiller  and  Justus  von  Liebig  in  Germany;  fearing  the 
competition  of  other  countries,  which  already  had  more  up-to-date  marine  stations; 
and  hoping  to  attract  and  link  new  scientists  to  the  Naples  Institute;  Dohrn  built, 
in  1888,  another  building  and  connected  it  to  the  older  one  with  a  foot-bridge.  He 
called  the  new  building  the  Department  of  Physiology.  Soon,  however,  the  new 
facilities  failed  to  meet  the  requirements  of  the  ever-expanding  field  of  comparative 
physiology,  and  he  took  the  third  step  of  his  unique  enterprise  by  adding  a  larger 
building,  attached  to  the  older  one.  This  was  finished  in  1906.  The  Zoological 
Station  maintained  its  well-known  appearance  until  1965,  when  the  empty  space 
spanned  by  the  foot-bridge  was  filled  with  the  new  library.  As  Dohrn  used  to  say: 
the  three  buildings  give  the  impression  of  a  railway  or  of  a  tramway,  with  an  engine 
and  two  wagons.  In  the  new  Department,  two  sub-departments  were  established: 
one  for  Physiological  Chemistry  and  the  other  for  Animal  and  Comparative 
Physiology. 

Dohrn  died  in  1909  and  was  unable  to  appreciate  fully  the  tremendous  impetus 
he  had  given  to  scientific  activities  of  the  Institute  with  the  realization  of  these  new 
sub-departments.  When  the  history  of  the  scientific  activities  of  the  Station  is 
written,  as  I  hope  it  will  be,  chronologically  and  with  all  the  studies  which  have 
been  carried  out  properly  described,  the  scale  will  tip  definitely  towards  physiology. 
There  is,  perhaps,  no  aspect  of  cell  physiology  or  of  comparative  physiology  and 

122 


COMPARATIVE   PHYSIOLOGY   AND   BIOCHEMISTRY  123 

biochemistry  which  has  not  been  for  a  shorter  or  longer  time  the  subject  of 
important  research  at  the  Institute.  It  is  a  long  line  of  flashing  lights,  some  very 
luminous,  some  less  so,  which  glows  almost  uninterrupted  to  the  present  time.  I 
shall  not  even  try  to  recall  all  of  them,  but  I  cannot  avoid  mentioning  a  few 
scientists  who,  by  having  worked  at  the  Station  for  considerable  periods,  left  an 
imprint  on  the  scientific  history  of  the  Institute. 

From  1892  to  1902,  von  Uexkiill  was  head  of  the  Physiology  Department,  and 
for  ten  years  he  carried  out  original  and  ingenious  experiments  on  the  nervous  and 
muscular  systems  of  marine  animals.  Uexkiill  was  a  pioneer  of  modern  behavioral 
biology.  He  investigated  the  environmental  relations,  the  "sphere  of  function"  as  he 
called  it,  which  connect  the  individual  to  its  environment. 

During  the  same  period  Oscar  Hertwig  and  Theodor  Boveri  laid  the  foundations 
of  the  mechanism  of  fertilization.  Especially  after  the  investigations  of  Boveri,  the 
egg  and  the  larvae  of  echinoderms  became  classic  objects  of  cell  physiology. 
Cytophysiological  studies  of  the  unfertilized  and  fertilized  sea  urchin  egg  cover  a 
large  part  of  the  scientific  history  of  the  Station;  this  field  has  been  elucidated  by 
Monroy  and  by  other  participants  of  this  meeting  more  expert  than  myself. 

Active  "customers"  of  the  Physiology  Department  at  the  end  of  the  last  century 
and  the  beginning  of  this  one  were  Silvestro  Baglioni  and  Filippo  Bottazzi,  who,  as 
human  physiologists  themselves,  were  well  aware  of  the  great  importance  of  the 
expanding  new  disciplines  of  comparative  physiology  and  physiological  chemistry. 
From  1905  to  1909,  Baglioni  worked  mostly  on  the  sensory  organs  of  cephalopods 
and  fishes;  Bottazzi  carried  out  his  well-known  experiments  on  osmoregulation  in 
marine  animals.  This  line  of  research  originated  from  the  concept  of  the  milieu 
interieur,  formulated  years  before  in  France  by  Claude  Bernard.  The  results  obtained 
caused  Bottazzi  to  distinguish  aquatic  animals  as  homeosmotic  and  pecilosmotic,  a 
distinction  which  is  maintained  today  with  the  modern,  if  not  more  transparent 
words,  osmoconformers  and  osmoregulators.  Baglioni  and  Bottazzi's  work  at  the 
Station  can  be  examined  in  their  extensive  chapters  in  the  Handbuch  der  Verglei- 
chende  Physiologic,  that  monumental  treatise  edited  by  Winterstein  between  1914 
and  1925. 

Martin  Henze,  who  came  to  Naples  in  1902,  was  in  charge  of  the  sub-department 
of  Physiological  Chemistry  for  ten  years,  until  1914.  He  made  many  important 
contributions  to  the  chemistry  of  hemocyanin,  the  oxygen-carrying  pigment  which 
had  been  discovered  by  Leon  Fredericq  while  working  at  the  marine  station  of 
Roscoff.  By  his  discovery  of  tyramine  and  other  amines  in  the  salivary  glands  of 
cephalopods,  Henze  was  the  initiator  of  a  fruitful  line  of  biochemical  research  which 
developed  into  several  other  biochemical  and  physiological  fields,  such  as  the 
biotoxins  of  marine  animals  and  their  humoral  and  nervous  correlations.  All  these 
have  been  pursued  at  the  Station  until  very  recently  by  many  scientists. 

From  1908  to  1914,  Otto  Warburg  spent  several  periods  at  the  Station,  where 
he  carried  out  his  first  major  independent  work  on  the  oxygen  consumption  which 
occurs  when  a  sea  urchin  egg  begins  to  develop  after  fertilization.  He  made  the 
classic  discovery  that  upon  fertilization  the  rate  of  respiration  rises  as  much  as  six- 
fold. These  results,  which  provided  the  basis  for  his  future  work  on  the  metabolism 
of  tumor  cells,  are  described  in  three  of  his  very  first  papers  (in  Hoppe  Seyler's 
Zeitschrift  fur  Physiologische  Chemie)  completed  before  obtaining  his  M.D.  degree 
in  1911  at  Heidelberg. 

Warburg  had  exceptional  skill  in  selecting  the  right  kind  of  material  for  solving 
a  specific  problem.  He  chose  the  sea  urchin  egg  (and  not,  for  instance,  the  frog's 
egg,  which  was  available  in  Heidelberg)  because  the  amount  of  living  matter  is  large 


124  F.  GHIRETTI 

in  relation  to  the  yolk  mass,  and  because  development  of  the  fertilized  egg  is  very 
rapid,  so  that  the  changes  he  was  looking  for  would  take  place  in  a  short  time.  His 
aim  was  to  demonstrate  that  in  the  course  of  growth  chemical  work  must  be  done, 
and  therefore  the  rate  of  energy  supply  must  increase.  This  is  self-evident  to  any 
student  who  studies  biology  today,  but  80  years  ago  the  experiments  of  earlier 
investigators  of  this  problem  had  been  inconclusive.  More  astonishing  is  that 
Warburg  carried  out  his  oxygen  determinations  not  manometrically,  but  with  the 
titrimetric  method  developed  by  Winkler,  which  he  had  improved  and  checked 
carefully  for  all  possible  causes  of  errors.  The  theory  and  practice  of  manometry 
were  perfected  by  Warburg  later,  in  1920,  and  they  were  the  key  techniques  of  his 
later  discoveries. 

After  1922,  the  Belgian  physiologists  Henry  Fredericq  (son  of  Leon  Fredericq) 
and  Zeno  Bacq  became  regular  guests  of  the  Station.  Here  they  worked  mainly  on 
the  physiology  of  the  autonomic  nervous  system  of  cephalopods;  the  presence  and 
release  of  noradrenaline  and  acetylcholine  from  the  visceral  nerves  of  invertebrates 
was  the  main  object  of  their  work.  It  was  during  these  studies  that  Bacq  discovered 
nemertine  and  amphiporine  from  marine  annelids,  a  discovery  which  can  be  placed 
in  another  line  of  research,  i.e.,  biotoxins  from  marine  animals.  This  original  line 
goes  back  to  Martin  Henze,  and  includes  among  its  exploiters  Vittorio  Erspamer, 
with  his  discovery  of  murexine  and  of  eledoisin.  Such  research  is  pursued  today  by 
some  members  of  the  staff  of  the  Station  including  Lucio  Cariello. 

In  1935,  Bacq  and  Francesco  Paolo  Mazza  demonstrated  the  presence  of 
acetylcholine  in  the  optical  ganglia  of  the  octopus,  and  identified  the  substance 
chemically.  Earlier,  in  Germany,  acetylcholine  had  been  identified  as  the  "vagusstojf" 
by  Otto  Loewi.  The  work  of  Bacq  and  Mazza  is  a  keystone  in  the  history  of 
chemical  nervous  transmission,  and  it  was  the  first  direct  demonstration  of  its 
existence  in  nervous  tissue. 

In  the  same  line  of  research  belongs  the  work  of  Enrico  Sereni,  who  was  in 
charge  of  the  Department  of  Physiology  for  several  years.  Sereni  was  a  brilliant 
physiologist,  and  among  many  other  studies  I  like  to  remember  are  his  ingenious 
experiments  on  nervous  and  humoral  correlations  of  the  activity  of  the  chromato- 
phores,  and  the  peripheral  nervous  system  in  cephalopods,  in  collaboration  with  the 
young  J.  Z.  Young. 

Young  founded  another  line  of  research,  followed  for  many  years,  almost 
uninterrupted  at  the  Station.  His  name  is  written  in  capital  letters  in  the  scientific 
history  of  the  Institute.  This  afternoon  we  shall  listen  to  a  recollection  by  Young 
himself  of  his  work  on  the  neurophysiology  of  the  squid  and  the  octopus. 

I  close  this  sketch  by  asking  once  again  that  experts  in  zoology,  botany, 
embryology,  physiology,  biochemistry,  etc.,  who  have  worked  at  the  Station,  write 
the  scientific  history  of  the  Institute.  As  a  multifaceted  crystal,  the  Zoological  Station 
of  Naples  will  reflect  a  hundred  years  of  the  history  of  Biology. 

It  has  been  said  that  we  cannot  conceive  what  the  present  state  of  biological 
sciences  would  be  without  the  influence  of  the  Zoological  Station.  This  is  pure 
rhetoric:  too  many  rhetorical  statements  have  been  made  about  the  Station. 

I  want  to  ask  a  simple  and  much  less  global  question:  what  has  been  the 
influence  of  the  Station  on  the  development  of  Biology  in  Italy!  After  all,  it  was 
founded  and  developed  in  this  country,  and  it  is  not  irrational  to  suppose  that 
exchanges  occurred  between  this  scientific  institute  and  the  rest  of  the  Italian 
scientific  community. 


COMPARATIVE   PHYSIOLOGY   AND   BIOCHEMISTRY  125 

In  fact,  the  impact  of  the  Station  on  Italian  zoology  is  enormous.  As  Giovanni 
Battista  Grassi  said,  in  the  last  century  the  Italian  zoologists  and  botanists  were 
more  Linnaeans  than  Linnaeus  himself.  Only  in  later  years  were  they  influenced  by 
the  great  revolution  of  the  so-called  scientific  zoology,  which  had  arisen  in  Germany 
in  1848  with  the  foundation  of  the  Zeitschrift  fur  wissenschaftliche  Zoologie.  Dohrn 
was  a  crusader  of  this  revolution.  The  Italian  zoologist  Antonio  Delia  Valle  (to 
mention  only  one),  who  worked  at  the  Station  from  1878  to  1899,  contributed 
greatly  to  the  development  of  zoology  in  this  country. 

But  the  same  did  not  occur  in  the  physiological  and  biochemical  sciences.  This 
is  surprising  because,  as  I  have  recalled,  the  Station  had  developed  more  space  for 
physiology  and  biochemistry  than  for  zoology  and  morphological  sciences.  In  my 
opinion  there  are  several  reasons  for  this,  which  deserve  careful  analysis,  especially 
in  consideration  of  the  present  status  and  of  the  future  prospects  of  the  Station. 

Physiological  and  biochemical  work  accomplished  at  the  Station  remained 
almost  unknown  to  Italian  physiologists  and  biochemists.  This  was  due  primarily 
to  the  indolence  and  the  rather  restricted  preparation  of  the  Italian  scientists.  No 
physiologist  in  Italy,  not  even  one  of  his  pupils,  was  the  heir  of  Bottazzi.  Even 
today  comparative  physiology  in  this  country  is  considered  an  eccentric  discipline 
by  traditional  physiologists. 

This  is  on  one  side.  On  the  other  side,  the  Station  always  refused  to  create 
something  which  would  have  even  the  appearance  of  a  school  of  physiology.  Yet 
the  Institute  had  its  own  scientific  staff;  it  had  heads  of  Departments,  and  Assistants. 
And  several  of  them,  as  I  have  recalled,  succeeded  in  establishing  productive  lines 
of  research  in  the  Department  of  Physiology — in  spite  of  the  politics  of  the  Institute, 
I  should  add. 

Reinhard  Dohrn  used  to  say:  "there  are  no  positions  at  the  Z.S.,  but  only 
functions."  By  this  rather  obscure  statement  he  meant  that  we  were  not  really  Head 
of  the  Department  or  Assistant,  but  we  had  to  behave  as  though  we  were.  If,  in 
addition,  we  wanted  to  carry  out  scientific  work,  that  was  our  business. 

The  fear  that  somebody  on  the  staff  might  become  the  scientific  head  of  a 
Department  is  the  original  sin  at  the  Zoological  Station.  At  the  time  of  Anton 
Dohrn,  Uexkiill  was  fired  because  he  wanted  to  be  the  literal  leader  of  the  Physiology 
Department.  "Discord  developed  between  Dohrn  and  Uexkiill  because  the  latter,  as 
head  of  the  Physiology  Department,  expected  special  treatment."  This  is  reported 
in  the  Archives  of  the  Station. 

This  is  also  the  reason  why  the  Station  never  had  its  own  research  programs.  Of 
course  that  is  not  precisely  true.  During  Anton  Dohrn's  lifetime  the  Station  did 
carry  out  an  ambitious  program  on  its  own.  It  was  in  fact  the  personal  scientific 
program  of  the  Director.  All  the  Assistants  (and  I  want  to  remember  among  them, 
particularly,  Salvatore  Lo  Bianco),  most  of  the  zoologists  and  morphologists, 
foreigners  as  well  as  Italians,  who  crowded  the  Institute,  even  in  disparate  fields, 
centered  their  interests  around  a  pillar,  which  was  the  original  scientific  program  of 
the  Station.  But  the  situation  changed  suddenly  with  appearance  of  the  Departments 
of  Physiology  and  of  Physiological  Chemistry,  since  Dohrn  was  neither  a  physiologist 
nor  a  biochemist,  and  it  was  radicalized  later  when  the  Director  was  not  even  a 
scientist. 

The  Station  never  made  a  sincere  effort  to  introduce  itself  to  the  Italian  scientific 
community,  however  reluctant  and  narrow-minded  were  the  Italian  physiologists 
and  biochemists.  Today  we  are  proud  to  recall  the  Nobel  prizes  of  famous  scientists 
who  worked  at  the  Station:  Otto  Loewi,  Albrecht  Kossel,  Otto  Warburg,  Otto 


126  F.  GHIRETTI 

Meyerhof,  Theodor  Svedberg,  Albert  Szent-Gyorgyi.  How  many  excellent  occasions 
those  imply  for  shaking  the  sloth  of  the  teachers,  and  for  attracting  young  people 
from  the  Italian  universities! 

With  the  exception  perhaps  of  Neapolitan  students  who  knew  the  building  from 
the  outside,  nobody  for  years,  in  the  Italian  universities,  ever  heard  about  the 
existence  of  a  Zoological  Station  in  Naples.  Postgraduate  students  were  admitted 
for  the  first  time  in  1947,  when  Giuseppe  Reverberi  obtained  support  via  short-term 
grants  from  the  Italian  National  Research  Council.  I  had  the  good  luck  to  be  among 
them,  and  I  wonder  how  welcome  it  was  to  have  around  a  dozen  restless,  noisy 
young  people,  not  used  to  the  atmosphere  of  such  an  old  and  traditional  Institute. 

When  Monroy  last  year  told  me  of  his  plan  to  celebrate  the  hundred  years  of 
scientific  activity  at  the  Zoological  Station  of  Naples  and  at  the  Marine  Biological 
Laboratory  in  Woods  Hole,  I  realized  suddenly  how  different  these  two  institutions 
have  approached  their  intentions  and  purpose:  the  former  aristocratic  and  exclusive; 
the  other  accessible  and  educational.  We  shall  discuss  further  the  biological  future 
and  the  functional  perspectives  of  the  Zoological  Station  of  Naples.  I  hope  that  the 
history  of  scientific  activity  at  the  Station,  which  we  are  sketching  at  this  meeting, 
will  not  remain  a  mere  academic  celebration,  but  will  rather  help  to  define  the 
place  of  this  Institute  in  the  Italian  scientific  community  of  today;  what  sort  of 
relationship  it  can  have  (or  must  have)  with  the  Italian  universities;  and  with  other 
scientific  institutions.  History,  said  the  Romans,  is  the  master  of  life.  But  history 
never  repeats  itself. 


Reference:  Biol.  Bull.  168  (suppi.):  127-136.  (June,  1985) 


SOME   STRUGGLES  OF  JACQUES   LOEB,   ALBERT   MATHEWS,   AND 
ERNEST  JUST  AT  THE   MARINE   BIOLOGICAL   LABORATORY 

SEYMOUR   S.   COHEN 

Depart  men!  oj  Pharmacological  Sciences,  State  University  of  New  York 
at  Stony  Brook,  Stony  Brook,  New  York  1 1 794 

ABSTRACT 

Jacques  Loeb  led  the  Department  of  Physiology  at  the  Marine  Biological 
Laboratory  (MBL)  in  1892,  four  years  after  the  opening  of  the  Laboratory.  In  that 
year  he  was  also  the  first  to  study  the  development  of  fertilized  sea  urchin  (Arbacia 
punctulata)  eggs  and  detected  a  selective  effect  of  hypertonic  sea  water  in  eliciting 
nuclear  cleavage.  By  1 899  he  had  discovered  artificial  parthenogenesis.  His  priority 
in  this  discovery  was  challenged  by  an  assistant  in  his  Department,  A.  P.  Mathews, 
who  criticized  other  aspects  of  Loeb's  work  both  from  the  MBL  and  from  their 
common  University.  Mathews  also  leaked  Loeb's  progress  to  the  press.  Loeb 
attempted  to  have  Mathews  fired,  but  Mathews  remained  as  head  of  the  course  at 
the  MBL  and  also  became  Professor  at  the  University  of  Chicago  after  Loeb  left. 
Loeb  worked  for  over  a  decade  to  prove  Mathews  incorrect  in  his  claims  and 
criticism.  Loeb's  work  eventually  helped  to  destroy  a  colloidal  theory  which  Mathews 
upheld  in  his  book,  the  first  American  text  of  biochemistry.  For  over  two  decades, 
Mathews  taught,  advised,  and  encouraged  E.  E.  Just,  who  in  support  of  the 
embryological  work  of  another  mentor,  F.  R.  Lillie,  publicly  attacked  Loeb's  work. 
Loeb's  antipathy  to  Just  has  been  attributed  to  racism,  but  possibly  resulted  in  large 
part  from  his  scientific  differences  with  Mathews,  Lillie,  and  Just. 

INTRODUCTION 

For  at  least  the  past  thirty  years,  there  has  been  a  poker  game  in  Woods  Hole 
each  Thursday  night  of  every  summer.  Although  the  participants  have  all  been 
eminent  practitioners  of  a  reductionist  quantitative  biology  at  the  MBL,  the  "poker 
games"  have  consisted  of  a  variety  of  different  card  games  in  which  winning  has 
depended  on  large  components  of  luck  coupled  with  concentrated  study  by  the 
players.  Almost  uniformly  the  games  have  included  "wild  cards"  or  "jokers",  i.e., 
either  the  cards  can  be  whatever  the  players  would  like  or  the  rules  have  been 
altered  to  permit  hidden  surprises.  In  maximizing  chance  the  participants  have 
relaxed  by  deliberately  changing  their  habits  of  work,  and  have  appealed  to  the 
power  of  the  unknown. 

Reductionism,  the  usual  methodology  of  these  gamblers,  has  often  incorrectly 
carried  the  implication  of  an  arrogant  presumption  of  knowledge.  Although  a  large 
majority  of  my  colleagues  hold  the  view,  as  did  Jacques  Loeb,  that  phenomena  of 
biology  obey  the  laws  of  physics  and  chemistry,  and  attempt  to  analyze  many  of 
these  phenomena  in  these  terms,  I  do  not  know  any  who  believe  that  physiological 
phenomena  are  entirely  interpretable  with  our  presently  available  knowledge. 
However,  to  many  of  Loeb's  contemporaries,  descriptive  morphologists  at  the  end 
of  the  nineteenth  century,  the  search  for  manipulable  simple  systems  as  an  approach 
to  generalizing  biological  laws  may  have  seemed  an  affront  to  the  then  obvious 
complexity  of  the  biological  world. 

127 


128  S.   S.   COHEN 

At  present  the  practitioners  of  this  methodology  will  say  only  that  they  are 
following  both  the  biologists,  who,  in  the  words  of  F.  G.  Hopkins  "know  best  the 
lay  of  the  land"  and  the  chemists  (and  physicists)  who  have  devised  ever  more 
penetrating  exploitative  techniques.  Despite  an  interest  in  performing  ever  more 
sophisticated  and  elegantly  controlled  experiments,  most  investigators,  even  today, 
are  enthused  by  the  possibility  of  the  unexpected,  i.e.,  a  serendipitous  discovery.  In 
this  sense,  the  poker  game,  replete  with  wild  cards,  may  be  merely  the  reflection  of 
the  aspirations  of  some  well-known  experimentalists. 

In  my  recent  examination  of  the  early  history  of  physiology,  embryology,  and 
biochemistry  at  the  MBL,  I  have  also  turned  up  some  unsuspected  wild  cards, 
which  appear  to  challenge  some  current  notions  about  our  predecessors. 

George  Sarton  has  told  us  that  a  historian  of  science  should  know  both  history 
and  science,  and  this  ideal  has  proven  to  be  quite  difficult  to  realize.  In  the 
occasional  mixed  symposia  of  historians  and  scientists,  these  groups  have  tended  to 
focus  on  different  aspects  of  a  problem  under  consideration  and  most  often  have 
talked  past  each  other.  In  addition  the  two  groups  have  developed  a  certain  tension 
between  them.  The  historians  are  suspicious  of  the  personal  involvement  of  the 
practitioners  with  the  subject  matter  and  other  possibly  related  long  forgotten  events, 
while  the  scientists  point  to  the  neglect  or  lack  of  understanding  of  the  scientific 
qualities  of  the  problems  by  the  historians.  In  my  recent  studies,  it  has  appeared  to 
me  that  an  interplay  of  scientific  and  historical  knowledge  is  essential  in  permitting 
a  realistic  look  at  the  evolution  of  physiology  at  the  MBL.  It  is  not  really  possible 
to  understand  the  growth  of  the  discipline  at  that  institution  without  a  close  study 
of  almost  all  of  the  actors  and  their  scientific  problems,  as  well  as  their  interactions 
on  matters  of  personal,  scientific,  and  institutional  concern. 

WHITMAN,  LOEB,  AND  PHYSIOLOGY 

In  examining  the  records  of  the  MBL,  the  name  of  Jacques  Loeb  appears  very 
early  in  the  introduction  of  physiology  at  the  Laboratory.  A  course  of  work  in 
physiology  as  such  was  not  introduced  "for  lack  of  space"  in  1888,  the  first  year  of 
the  opening  of  the  laboratory  (Whitman  1892).  Whitman  called  attention  to  this 
deficiency  in  1891  and  acted  to  correct  it  in  1892.  In  his  Annual  Report  of  1892, 
he  stated  that 

Morphology  and  physiology  are  two  quite  distinct  sides  of  biology,  each  with 
definite  and  constant  peculiarities  of  method  and  aim;  but  these  two  sides  are 
only  the  statical  and  the  dynamical  aspect  of  one  and  the  same  thing;  one 
presents  Ihe  feature  the  other  the  expression.  It  is  only  as  a  matter  of  convenience 
that  these  two  aspects  are  dealt  with  separately;  they  are  complemental  and  have 
their  full  meaning  only  when  united. 

Whitman  continued  by  insisting  that  the  two  sides  be  "kept  in  working  contact." 
Separation  has  kept  "physiology  too  exclusively  in  the  service  of  Medicine"  .  .  . 
"The  biological  economy  of  organisms  must  become  an  integral  part  of  physiology." 
Both  branches  relate  to  problems  of  both  evolution  and  development,  i.e.,  to 
paleontology  and  embryology,  and  "the  history  of  morphology  and  physiology  is 
one  continuous  illustration  of  their  interdependence,"  as  exemplified  by  Harvey's 
discovery  of  the  circulation  of  the  blood.  There  has  been  a  "lack  of  interest  in 
general  physiology"  and  in  marine  biology.  "It  has  been  our  good  fortune  to  draw 
into  connection  with  the  Laboratory  Dr.  Loeb,  whose  enthusiasm,  zeal,  and 
accomplishments  in  general  physiology,  make  him  a  fitting  director  of  this 
department." 

After  a  year  at  Bryn  Mawr  in  1891,  Loeb  was  recruited  by  Whitman  to  the 
University  of  Chicago  in  1892.  Whitman's  plan  for  the  growth  of  an  academic 


THE  LOEB,   MATHEWS,   JUST  CONNECTION  129 

biology  that  reflected  the  natural  world  rather  than  that  of  medicine  called  for  a 
man  of  Loeb's  broader  interests  at  both  his  University  Department  and  the  MBL. 

In  the  summer  of  1892  Loeb  began  in  a  new  building  at  the  MBL,  and  worked 
with  three  postdoctoral  associates  and  a  number  of  students  in  a  course  of  lectures 
and  experimental  studies.  He  soon  became  interested  in  embryological  studies  and 
reportedly  was  the  first  to  perform  experiments  on  the  eggs  of  the  sea  urchin, 
Arbacia  punctulata  (Harvey,  1956).  His  paper  described  effects  of  hypertonic  sea 
water  on  nuclear  division  without  cell  division  (Loeb,  1892).  T.  H.  Morgan,  also 
working  at  the  MBL,  published  on  the  same  subject  a  year  later  (Morgan,  1893).  It 
is  of  interest  that  at  the  MBL  both  of  these  men,  who  had  had  some  administrative 
differences  as  fellow  faculty  members  at  Bryn  Mawr,  were  pursuing  similar  problems 
in  physiological  morphology,  i.e.,  embryological  development,  on  the  same  biological 
system.  Although  frequently  differing  in  their  conclusions,  these  men  did  respect 
each  other's  work,  and  had  similar  orientations  to  the  major  biological  problems  of 
the  day.  Both  eventually  left  this  complex  but  apparently  confusing  embryological 
system  for  more  manageable  materials,  and  as  we  know,  both  made  fundamental 
contributions  to  quantitative  biology.  Morgan  became  an  innovator  of  genetics  and 
the  theory  of  the  gene,  while  Loeb  can  be  considered  to  be  a  founding  father  of  the 
physical  chemistry  of  proteins.  An  important  book  on  the  latter  subject  (Cohn  and 
Edsall,  1943)  is  dedicated  to  the  memory  of  Jacques  Loeb  (1859-1924),  in  addition 
to  tributes  to  Sir  William  Bate  Hardy  (1864-1934),  Thomas  Burr  Osborne  (1859- 
1929),  and  S0ren  Peter  Lauritz  S0rensen  (1868-1939).  The  inscription  in  this 
volume  of  Cohn  and  Edsall  reads  "Their  investigations  laid  the  foundations  for  the 
physical  chemistry  of  proteins." 

LOOKING  AT  LOEB  AND  His  ASSOCIATES 

Any  working  biochemist  trained  in  the  193CTs  and  194CTs  would  have  many  ties 
to  Loeb.  Taking  courses  in  General  Physiology  and  Colloid  Chemistry  in  the  3(Ts, 
one  would  have  learned  that,  as  a  result  of  Loeb's  work,  the  alleged  distinctions 
between  crystalloids  and  colloids,  as  applied  to  protoplasmic  constituents,  were 
spurious.  He  had  shown  that  proteins  combine  stoichiometrically  with  acids  and 
alkalis  as  a  function  of  the  hydrogen  ion  concentration  (pH)  and  reactive  ionizable 
groups  on  the  proteins  (Loeb,  1924).  As  a  result,  proteins  may  be  soluble  at  one 
pH  and  aggregated  at  another.  This  manipulation  was  fundamental  in  the  extraor- 
dinary isolation  and  crystallization  of  many  proteins  and  enzymes  in  the  late 
twenties,  thirties,  and  forties  by  John  Northrop  and  Moses  Kunitz,  both  of  whom 
had  been  close  associates  of  Loeb  at  the  Rockefeller  Institute.  Also,  in  another 
important  branch  of  protein  chemistry,  the  productive  laboratory  of  E.  J.  Cohn 
during  World  War  II  used  the  physical  chemical  methods  pioneered  by  Loeb  to 
prepare  many  medically  useful  fractions  of  blood  plasma.  Loeb  is  an  important 
figure  to  a  biochemist  of  my  period,  and  it  was  fascinating  to  learn  that  his 
biochemical  discoveries  began  at  the  MBL  with  experiments  on  sea  urchin  eggs  and 
hypertonic  sea  water. 

LOEB'S  EARLY  RESEARCH  AT  THE  MBL 

Several  historians  have  outlined  Loeb's  early  life  as  student  and  post-doctoral 
research  associate  in  Germany  (Fleming  1973;  Pauly,  1980).  His  initial  interest  in 
the  localization  of  brain  function  raised  questions  on  the  control  of  locomotion  and 
animal  behavior,  subjects  which  led  to  studies  of  various  tropisms.  He  then  opposed 
the  concept  of  instinctual  behavior  and  objected  to  the  use  of  psychological  terms 
in  describing  biological  reality.  In  the  late  1880's  Loeb  had  discussed  problems  of 


130  S.   S.   COHEN 

plant  motion  with  Julius  Sachs  who  had  wished  "to  reduce  to  chemistry  and  physics 
those  functions  amenable  to  such  explanations"  and  had  pointed  to  the  effects  of 
osmotic  pressure  and  salts  on  plants  (Pauly,  1980).  As  described  by  Pauly,  Loeb 
saw  science,  and  particularly  biological  science,  as  part  of  a  human  effort  to  cope 
more  effectively  with  the  environment.  Going  to  the  Stazione  Zoologica  in  Naples 
in  1889  and  1890,  he  began  to  study  regeneration  and  the  creation  of  new  animal 
forms  in  marine  organisms. 

Loeb's  marriage  to  an  American  in  1890,  his  evaluation  of  his  potential  career 
in  medically  oriented  Germany,  and  his  friendship  in  Naples  with  Americans,  such 
as  Christian  Herter,  eased  his  decision  to  emigrate  and  to  accept  the  offer  of  a 
position  at  Bryn  Mawr.  His  move  to  the  larger  world  of  the  newly  established 
University  of  Chicago  and  an  association  with  the  enlightened  and  dynamic 
Whitman  permitted  him  to  become  a  virtually  instantaneous  leader  of  physiology 
in  an  America  in  which  the  discipline  scarcely  existed.  It  must  be  noted  that 
Darwinian  evolution  in  this  pre-Mendelian  era  could  offer  no  scientific  proof  of  the 
mechanism  of  the  origin  of  species  and  variation,  and  Loeb,  as  well  as  Morgan, 
were  far  from  convinced  at  this  time  of  the  validity  of  "evolutionism."  It  has  been 
suggested  by  Pauly  that  Whitman  hoped  research  by  Loeb  and  the  physiologists 
would  contribute  to  knowledge  of  the  evolution  of  physiological  function,  whereas 
Loeb  was  interested  primarily  in  the  control  of  known  function. 

In  any  case,  Loeb's  research  began  at  the  MBL  with  the  effects  of  hypertonic 
media  on  fertilized  sea  urchin  eggs,  and  he  believed  he  had  detected  nuclear  cleavage 
without  cellular  cleavage  (Loeb,  1892),  a  result  which  was  challenged  (Morgan, 
1893).  With  the  cytological  aid  of  W.  W.  Norman  of  Texas,  an  associate  in  the 
Physiology  Course  at  the  MBL  and  previously  at  the  Naples  Laboratory  (Norman, 
1896),  Loeb  was  proved  correct  and  proceeded  to  exploit  the  observation  of  Morgan 
who  had  seen  that  salts  produced  mitosis-like  effects  in  unfertilized  eggs.  By  1899 
Loeb  had  discovered  and  reported  "the  artificial  producton  of  normal  larvae  (plutei) 
from  unfertilized  eggs  of  the  sea  urchin"  (Loeb,  1899).  Loeb  had  indeed  begun  to 
"control"  biological  systems.  In  the  summer  of  1900,  this  result  was  confirmed  by 
many  workers  and  the  cytology  of  parthenogenetic  development  of  these  eggs  had 
been  clarified  (Wilson,  1901). 

It  should  be  evident  that  this  surprising  and  dramatic  feat  was  the  consequence 
of  many  lines  of  work.  Loeb,  Sachs,  and  others  had  rejected  the  medical  direction 
of  German  biology  and  had  sought  ever  more  controllable  model  systems.  The 
Naples  laboratory  had  helped  to  educate  Whitman,  Loeb,  Morgan,  and  Norman  in 
the  importance  of  marine  systems.  Whitman's  vision,  energies,  and  administrative 
skills  had  built  the  MBL  and  the  Department  at  Chicago,  and  had  recruited  Loeb. 
The  personal  qualities  and  interplay  of  all  of  these  men  had  also  contributed 
crucially.  Nevertheless  the  induction  of  nuclear  cleavage  to  artificial  parthenogenesis 
had  taken  seven  years  of  determined  effort  to  answer  Morgan  and  to  go  on  from 
there.  The  need  of  a  working  scientist  to  define  the  nature  of  a  possible  mistake 
and  to  correct  it,  if  necessary,  can  be  a  powerful  driving  force,  and  perhaps  has  not 
been  considered  sufficiently  in  analyzing  the  mainsprings  of  discovery.  In  looking 
at  Loeb's  career,  we  shall  see  that  Loeb  worked  very  hard  and  long  to  answer 
criticisms  of  his  scientific  results  and  conclusions.  The  nature  of  that  need  is  not 
easily  clarified. 

A.  P.  MATHEWS,  THE  WILD  CARD 

In  1895,  E.  B.  Wilson  and  Albert  Prescott  Mathews,  the  latter  a  student  at 
Columbia  University,  made  an  interesting  contribution  to  the  study  of  the  devel- 
opment of  the  sea  urchin  egg  (Wilson  and  Mathews,  1895).  Mathews  spent  a  year 


THE   LOEB,   MATHEWS,  JUST  CONNECTION  131 

in  Germany  with  Albrecht  Kossel  on  the  chemistry  of  sperm  and  in  1898  he 
completed  his  doctoral  dissertation  at  Columbia  on  "The  Physiology  of  Secretion" 
(Harvey,  1958).  Having  studied  the  chemistry  of  staining  and  the  structure  of  cells 
of  the  pancreas,  and  with  biochemical,  cytological,  and  embryological  training, 
Mathews,  now  Assistant  Professor  of  Physiology  at  Tuft's  College,  joined  the  MBL 
faculty  of  the  Department  of  General  and  Comparative  Physiology  in  1899  with 
Loeb  and  E.  P.  Lyon,  Instructor  in  Biology  of  the  Bradley  Institute  of  Technology 
of  Peoria,  Illinois.  Norman  appears  to  drop  from  the  course  after  1898.  This  is  the 
important  summer  of  Loeb's  great  discovery,  but  in  1900  Mathews  announced  that 
quite  independently  he  had  caused  cell  division  in  unfertilized  Arhacia  eggs  by 
anaerobiosis,  heat  and  ether,  alcohol,  and  chloroform.  He  also  added  that  Morgan 
had  conducted  the  first  fruitful  experiments  on  chemical  ferilization  in  1898,  and 
that  Loeb  had  confirmed  and  extended  Morgan's  work  in  1899.  According  to 
Mathews,  Loeb  had  been  incorrect  initially  in  his  interpretations,  but  had  now 
modified  his  views. 

In  1901  (Mathews,  1901)  Mathews  published  from  Harvard  on  secretion,  and 
on  the  effects  of  salts  in  the  conductivity  of  the  nerve.  In  another  paper  we  learn 
more  of  artificial  parthenogenesis,  in  this  instance  of  starfish  eggs,  produced  by 
mechanical  agitation.  This  paper  also  included  a  confirmation  attributed  to  Morgan, 
as  well  as  unpublished  data  by  Loeb  on  Chaetoptems  and  Nereis.  It  may  be 
mentioned  that  embryologists,  including  E.  E.  Just,  thought  that  this  effect  of 
shaking  was  due  to  the  carbon  dioxide  or  acidity  generated  in  unduly  concentrated 
suspensions  of  eggs. 

In  1901  Mathews  came  to  the  University  of  Chicago  in  the  Department  of 
Physiology.  He  informed  his  brother,  a  newspaper  reporter,  of  exciting  unpublished 
work  from  Loeb's  laboratory,  which  was  then  sensationalized  in  the  press  (Pauly, 
1980;  Kohler,  1982).  Loeb  was  upset,  and  writing  to  President  Harper,  stated  that 
Mathews  should  not  have  confiscated  Loeb's  unpublished  work  or  have  claimed 
that  of  Loeb's  students,  that  Mathews'  work  was  unsound  "and  of  such  a  character 
as  to  sooner  or  later  injure  the  reputation  of  the  University"  (Kohler,  1982).  Kohler 
has  also  recorded  some  independent  opinions  casting  aspersions  on  Mathews' 
scientific  judgement.  The  resulting  brouhaha  led  to  the  development  of  a  "Code  of 
Scientific  Ethics"  by  the  faculty  of  the  University  of  Chicago  (Pauly,  1980).  By  1903 
Loeb  had  left  the  University  of  Chicago  for  the  University  of  California,  but 
Mathews  remained  in  Chicago  and  developed  work  in  chemical  biology,  rising  to 
the  rank  of  full  professor  by  1905  (Kohler,  1982).  He  left  Chicago  only  to  go  to  the 
University  of  Cincinnati  where  he  began  a  program  in  clinical  chemistry. 

In  long  papers  in  Science  on  nerve  stimulation,  Mathews  turned  his  attention 
to  physical  chemistry,  attempted  to  correct  some  of  his  own  initial  mistakes  in 
attributing  salt  effects  to  the  production  of  hydroxyl  ions,  and  then  enlarged  upon 
new  results  correcting  errors  assigned  to  Loeb  (Mathews,  1902).  When  Loeb 
responded  criticizing  some  of  Mathew's  notions,  such  as  those  of  stimulation  and 
inhibition  being  due  to  a  precipitation  and  resolution  of  nerve  colloids  respectively, 
Mathews  continued  at  length,  and  introduced  new  speculations  on  his  results  with 
eggs,  kidneys,  and  central  nervous  system  (Mathews,  1903). 

Loeb  now  had  a  large  number  of  criticisms  and  claims  to  which  to  respond.  In 
fact,  although  far  more  circumspect  than  Mathews  in  referring  to  his  antagonist  by 
name,  Loeb  did  continue  to  explore  the  matters  raised  by  Mathews  and  eventually 
responded  to  many  of  them.  For  example  in  1899  Loeb  had  reported  that  calcium 
ion  inhibited  muscle  twitching  caused  by  NaCl  and  had  suggested  that  stimulation 
by  citrate,  oxalate,  fluoride,  and  phosphate  might  be  due  to  precipitation  or  other 
binding  of  calcium.  On  the  other  hand  Mathews  expressed  the  belief  in  1903  that 


132  S.  S.  COHEN 

only  unions  are  stimulating  and  that  cations  are  depressing.  Exploring  the  effects  of 
salts  of  ammonium  and  tetraethyl  ammonium  ions  and  other  amines,  and  a  large 
array  of  other  salts  Loeb  and  Ewald  showed  in  1916  that  calcium  "does  not  inhibit 
the  efficiency  of  the  stimulating  salt  by  depressing  the  irritability  of  the  nerve." 
Many  other  facts  are  adduced  and  in  conclusion  "All  these  facts  contradict  the 
hypothesis  of  Mathews.  .  .  ."  (Loeb  and  Ewald,  1916). 

Loeb's  continuing  concern  with  Mathews1  counter  hypotheses  can  be  found  in 
some  of  his  later  papers  (Loeb,  1914a).  It  may  be  asked  if  his  almost  exclusive 
concentration  on  salt  effects  and  ion  binding  in  his  later  experimental  years  did  not 
result  in  significant  measure  from  the  knowledge  he  had  developed  in  responding 
to  these  old  criticisms.  Loeb's  initial  formulation  of  the  idea  that  ion  effects  on 
protoplasm  were  interpretable  by  the  laws  of  chemistry  and  physics  was  stated  as 
early  as  1904,  and  his  criticisms  of  the  proposed  differences  between  colloidal  and 
crystalloidal  proteins  sharpened  as  his  studies  on  ion  binding  became  more  rigorous. 

The  existence  of  this  deep  personal  and  professional  antagonism  between  Loeb 
and  Mathews  raises  many  questions  concerning  Mathews'  position  at  Chicago  and 
at  the  MBL.  How  did  he  manage  to  remain  at  Chicago?  Mathews  led  the  MBL 
physiology  course  after  Loeb's  departure  for  the  University  of  California  in  1903, 
and  remained  associated  with  it  until  1919.  Indeed  he  was  elected  a  Trustee  of  the 
MBL  in  1906  and  remained  a  Trustee,  even  if  emeritus,  until  his  death  in  1957 
(Harvey,  1958).  When  Loeb  left  California  for  the  Rockefeller  Institute  in  1910,  he 
spent  his  summers  at  the  Institute  laboratory  at  the  MBL,  with  associates  among 
whom  were  Northrop  and  Kunitz.  He  also  was  a  Trustee  until  his  death  in  1924.  It 
might  be  imagined  that  Mathews  and  Loeb  might  have  had  quite  divergent  opinions 
on  the  organization  of  the  physiology  course,  and  might  well  have  expressed  these. 
However  there  is  no  significant  mention  of  Mathews  in  the  minutes  of  the  Trustees 
and  of  the  Executive  Committee  of  the  Trustees.  The  minutes  and  formal  reports 
are  models  of  tact,  obscuring  the  tensions  of  the  times. 

THE  MANNING-JUST-LOEB  CONNECTION 

As  indicated  earlier,  a  career  in  biochemistry  and  cell  physiology  does  sensitize 
one  to  the  name  of  Jacques  Loeb.  In  addition  a  literary  event  in  1983  has  most 
forcefully  brought  Loeb  to  our  attention.  The  biography  of  the  Negro  biologist, 
Ernest  Everett  Just,  written  by  the  black  historian,  K.  R.  Manning  (Manning,  1983), 
was  widely  and  favorably  reviewed  this  past  year.  The  book  describes  the  social  and 
institutional  difficulties  of  a  black  biologist  in  America  from  1910  through  1940. 
Woods  Hole  and  the  MBL  are  portrayed  as  a  somewhat  racist  community  and 
institution  respectively,  in  which  the  apprentice  Just  earned  his  keep  by  menial 
labor  and  was  trained  to  do  research  by  the  Director,  F.  R.  Lillie.  Just  eventually 
obtained  a  doctoral  degree  under  Lillie  at  the  University  of  Chicago  and  worked 
independently  at  the  MBL  on  problems  of  embryological  development.  Although 
Just  had  risen  to  a  Professorship  at  Howard  University,  had  published  extensively 
with  Lillie,  and  was  recognized  as  quite  knowledgeable  in  his  field,  he  was  unable 
to  obtain  a  position  in  a  major  white  University,  and  indeed  he  had  been  warned 
of  this  on  many  occasions  by  his  teacher  and  adviser,  Lillie.  He  had  indeed  failed 
to  "break  out"  of  Howard  to  a  more  respected  position,  and  was  unable  to  make  a 
"lasting  psychological  adjustment"  to  his  predicament.  Although  recognized  as  a 
great  Negro  biologist,  he  wished  desperately  to  be  recognized  as  a  great  biologist.  In 
describing  Just's  failures.  Manning's  book  imputes  racist  behavior  to  the  Jew,  Loeb, 
dubs  Abraham  Flexner,  a  "paternalistic  segregationist,"  and  dwells  gratuitously  on 
the  unpleasant  personal  reputations  of  some  Jewish  physiologists.  Just  is  quoted  as 


THE   LOEB,   MATHEWS,  JUST  CONNECTION  133 

having  written  in  an  appeal  to  Lady  Astor  in  the  1930's  that  he  had  been  "put  in 
the  bad  book  by  Jews  generally." 

However  throughout  the  book  one  finds  that  Loeb  had  helped  Just  considerably 
on  several  occasions  early  in  his  professional  career,  that  Flexner  had  assisted  Just 
often  through  various  philanthropic  agencies,  and  that  Just  had  obtained  warm, 
even  caring  personal  support  from  some  Jewish  colleagues  and  friends.  Nevertheless 
one  is  left  with  the  claim  that  Loeb  was  a  major  factor  in  the  tragedy  of  Just. 

It  is  appropriate  to  consider  the  evidence  that  Loeb  was  a  racist  because 
Manning  has  not  discussed  the  succession  of  events  in  the  context  of  the  interactions 
of  scientists  who  disagreed  publicly  as  well  as  privately  on  matters  of  science.  Some 
of  the  weaknesses  of  Manning's  arguments  may  be  summarized  as  follows: 

(1)  Since  1904,  Loeb  had  abandoned  the  mystique  of  colloidal  chemistry  which 
was  retained  by  Just,  Mathews,  and  close  colleagues  of  Just  such  as  L.  V.  Heilbrunn 
through  the  late  1930s.  This  neglect  of  scientific  differences  will  be  considered  in 
more  detail  below. 

(2)  The  twenty-year-old  relations  of  Just  to  his  teacher  and  adviser  Mathews 
are  discussed  without  reference  to  the  unfriendly  relations  of  Mathews  and  Loeb 
described  above. 

(3)  Loeb's  antiracist  publications  in  1914  (Loeb,  1914b)  are  not  quoted,  but  he 
is  rebuked  for  not  having  made  speeches  at  NAACP  meetings. 

(4)  In  a  letter  to  a  Flexner,  Loeb  is  known  to  have  objected  to  Just  as  a  possible 
colleague  at  the  Rockefeller  Institute.  Just  is  stated  to  be  "incompetent."  The 
referencing  is  confused,  and  the  exact  quotations  are  distorted  (Manning,  1983).  In 
any  case,  Loeb  believed  that  Just  "was  one  of  the  men  who  were  making  Woods 
Hole  an  impossible  place  for  a  decent  scientist  to  live  in"  (Pauly,  1980).  If  read 
carefully,  this  might  be  interpreted  to  indicate  that  Just  is  only  one  of  numerous 
miscreants  at  the  MBL,  and  this  could  not  be  an  argument  on  racial  grounds.  The 
suggestion  that  Just  would  be  better  off  as  a  high  school  teacher  is  also  suggested  to 
be  racially  motivated.  None  of  these  inferences  are  discussed  in  the  context  that 
Just  had  been  denouncing  Loeb's  science  in  print  and  at  meetings  for  some  years. 

(5)  Despite  such  attacks  by  Just  against  Loeb,  the  latter  is  expected  to  have 
supported  Just's  appointment  at  the  Rockefeller  Institute,  as  "symbolic  for  the 
whole  black  race"  (Manning,  1983).  Manning,  the  historian,  is  calling  for  "affirmative 
action"  in  1923  from  Loeb,  who  has  been  publicly  criticized  by  Just. 

(6)  Commenting  on  Loeb's  scientific  disagreements  with  Lillie,  the  most  impor- 
tant of  Just's  teachers,  advisers  and  collaborators,  Manning  suggests  that  Loeb 
founded  the  Journal  of  General  Physiology  in  part  to  block  Lillie  from  publishing 
(Manning,  1983).  John  Northrop's  memory  of  Loeb's  impatience  in  publishing  in 
previously  existing  journals  seems  far  more  convincing  (Pauly,  1980). 

There  does  not  appear  to  be  direct  evidence  of  racism  on  Loeb's  part.  It  should 
also  be  noted  that  Just's  alienation  from  American  society  continued  and  exacerbated 
long  after  Loeb's  death  in  1924. 

THE  MATHEWS-JUST  CONNECTION 

The  point  is  made  by  Manning  that  Just's  attacks  on  Loeb  are  often  attributable 
to  Just's  loyalty  to  his  teacher  and  adviser,  Lillie,  whose  scientific  outlook  and 
analyses  differed  from  those  of  Loeb.  It  is  relevant  to  note  that  A.  P.  Mathews  was 
also  teacher,  friend,  and  scientific  adviser  to  Just  and  that  Loeb's  attitude  to  Just 
may  also  have  been  influenced  by  the  other  protracted  war  between  Loeb  and 
Mathews  described  earlier. 


134  S.   S.  COHEN 

When  Just  studied  at  the  University  of  Chicgo  in  the  academic  year  of  1914-15 
to  fulfill  his  residency  requirement  for  the  doctorate,  he  took  a  course  in  cell 
chemistry  with  Mathews  (Manning,  1983).  Mathews  has  been  quoted  to  the  effect 
that  Just  was  "better  than  any  other  student  in  the  class,"  and  likely  to  become 
"one  of  the  most  original  and  creative  men  in  zoology  in  the  United  States."  Their 
association  continued  at  the  MBL  and  professionally  through  the  late  1930's.  In 
1935,  Mathews  encouraged  Just  to  express  his  controversial  opinions  challenging 
current  genetic  theory  at  a  meeting  of  the  American  Society  of  Zoologists,  and  to 
emulate  Lavoisier's  revolution  in  18th  century  chemistry  (Manning,  1983). 

A.  P.  Mathews  is  credited  with  having  written  the  first  American  textbook  in 
biochemistry  in  1915.  It  was  written  for  medical  students,  was  quite  popular,  and 
went  through  six  editions,  of  which  the  last  was  in  1936  (Mathews,  1936).  It  was 
said  to  have  irritated  colleagues  with  personal  asides,  among  other  non-standard 
components  (Kohler,  1982).  Mathews'  style  was  apparently  consistent.  The  1915 
book  has  been  described  (Chittenden,  1930)  as  comprehensive,  critical  and  valuable. 
Chittenden's  evaluation  of  Mathews'  contributions  comments  favorably  on  a  discovery 
of  the  spontaneous  oxidation  of  sugars  and  cysteine  in  alkaline  solution  but  notes 
his  work  on  protein  precipitation  by  ions  without  other  comment. 

However  Florkin  did  comment  unfavorably  on  the  1924  edition  of  Mathews' 
book  in  a  chapter  on  "The  Dark  Age  of  Biocolloidology"  (Florkin,  1972).  Mathews 
was  quoted  as  follows: 

It  is  by  means  of  the  colloids  of  a  protein,  lipoid  or  carbohydrate  nature, 
which  make  up  the  substratum  of  the  cell  that  this  localization  of  chemical 
reactions  is  produced;  the  colloids  furnish  the  basis  for  the  organization  or 
machinery  of  the  cell;  and  in  their  absence  there  could  be  nothing  more  than  a 
homogeneous  conglomeration  of  reactions.  .  .  .  The  colloids  localize  the  cell 
reactions  and  furnish  the  physical  basis  of  its  physiology;  they  form  the  "cell 
machinery." 

In  1936,  long  after  the  important  work  of  Loeb,  Adair,  and  Svedberg  in  1924, 
Mathews'  text  was  still  describing  as  uncertain  the  molecularity  of  hemoglobin  and 
other  defined  proteins  (Mathews,  1936).  Unfortunately,  Just's  orientation  to  cell 
chemistry  in  1939,  the  year  of  publication  of  his  book  The  Biology  of  the  Cell 
Surface  (Just,  1939a)  which  was  to  make  his  final  statement,  was  essentially  still 
that  of  the  colloidal  chemistry  of  his  teacher  and  even  recent  adviser,  A.  P.  Mathews. 
It  was  unfortunate  that  Just,  focussing  on  the  potential  roles  of  cell  membrane  and 
the  underlying  cytoplasm,  which  he  termed  "ectoplasm,"  died  in  1941,  before  the 
power  of  electron  microscopy  became  known  and  available  and  before  the  underlying 
nucleoprotein  particles  in  cytoplasm  had  been  described  by  Caspersson  and  isolated 
and  analyzed  by  Brachet  and  Claude.  Just's  book,  published  just  before  World  War 
II,  could  not  have  received  active  attention  by  either  the  Europeans  at  war,  or  the 
increasingly  distracted  Americans. 

WAS  LOEB  WRONG  ABOUT  JUST? 

Loeb  and  others  considered  Just  to  be  arrogant,  conceited,  and  boastful  in  his 
aggressive  claims  for  recognition  of  achievement  (Manning,  1983).  However  it  is 
probable  that  Just  felt  it  necessary  to  assert  himself  more  forthrightly  than  is 
customary  in  obtaining  recognition  by  his  peers.  What  is  peer  review  for  a  black 
biologist? 

Of  the  three  groups  with  which  Just  had  been  associated  at  the  MBL,  all  of 
whom  had  studied  the  embryological  development  of  marine  eggs,  only  Just  had 


THE   LOEB.   MATHEWS.  JUST  CONNECTION  135 

stayed  with  the  problem  and  had  come  to  a  sense  of  the  importance  of  cytoplasm. 
T.  H.  Morgan,  after  confusing  starts  and  shifts,  had  dropped  the  problem  and  had 
focused  on  mutations  in  Drosophila;  his  work  in  genetics  culminated  in  a  Nobel 
prize.  Loeb  also  had  dropped  the  problem  and  in  attempting  to  understand  his 
earliest  work  had  become  both  a  founding  father  of  protein  chemistry  and  a  legend, 
depicted  in  part  in  "Arrowsmith."  Lillie,  with  whose  research  work  Just  had  been 
associated  for  years,  had  become  President  of  the  National  Academy  of  Sciences. 
Meanwhile  embryological  work  had  demonstrated  that  egg  fragments  lacking  nuclei 
and  genes  could  develop  after  fertilization,  or  even  divide  after  stimulation  without 
fertilization.  Just  emphasized  the  importance  of  the  cytoplasm,  as  had  Morgan 
earlier,  but  finally  had  overreached  himself  (Just,  1939b)  with  unnecessary  attacks 
on  genetics  (Manning,  1983)  and  extreme  theories  on  the  nature  of  the  cellular 
division  of  labor. 

However  it  may  be  noted  that  in  November  1936  Just  wrote  to  Lillie  from 
Howard  University,  described  his  most  recent  work,  and  requested  that  Lillie  write 
to  support  another  request  for  financial  help. 

"Whilst  as  you  would  appreciate  my  interest  is  in  the  biology,  I  have  had  to  go 
into  the  chemical  end  of  the  nucleoproteins.  In  this  I  have  had  great  help  from  Dr. 
A.  P.  Mathews.  He  has  also  written  a  very  strong  letter  in  my  behalf.  .  .  ."  (Just, 
1936).  Later  in  Just's  "Status  of  my  research  program  in  embryology  .  .  ."  to  Lillie. 
apparently  written  in  1939,  he  states  "as  a  law  that  differentiation  during  development 
never  appears  without  attendant  progressive  synthesis  of  nucleoprotein"  (Just, 
1939b).  Believing  the  increase  of  nuclear  substance  to  be  effected  in  the  cytoplasm 
he  proposes 

a  more  exact  study  of  nucleo-protein  synthesis  to  embrace  as  many  different 
types  of  eggs  as  possible.  .  .  .  My  measurements,  up  to  now  as  rough  as  those 
of  others  who  had  made  such,  I  can  refine.  The  aim  here  is  to  follow  by  careful 
observation  and  experiment  the  moment  to  moment  changes  in  the  cytoplasm 
as  the  nuclei  are  built  up,  to  correlate  them  with  nuclear  synthesis  and  thus  to 
derive  a  biological  law  of  general  validity  for  all  cells  undergoing  differentiation. 

Could  the  aging  biologist.  Just,  have  learned  these  techniques?  Could  he  have 
determined  that  DNA  is  not  synthesized  in  the  cytoplasm  and  then  have  learned 
more  about  the  division  of  labor  within  cells?  Could  he,  like  Loeb  and  Morgan, 
have  achieved  genuinely  important  contributions  to  our  understanding  of  cellular 
biology  relatively  late  in  life?  We  cannot  know,  but  understand  that  his  thread  of 
an  idea  might  have  become  the  rope  on  which  he  might  have  climbed  into  the 
modern  cellular  science  of  the  post-war  world. 

CLOSING  REMARKS 

A  look  at  the  newly  assembled  Archives  at  the  MBL  reveals  old  data  and  new 
questions.  How  did  these  perpetually  quibbling,  quarreling,  or  warring  scientists  live 
together,  experiment,  and  survive  and  grow  professionally?  How  had  Woods  Hole 
in  1923  become  an  impossible  place  for  a  decent  scientist  to  live  in?  And  what  was 
a  decent  scientist?  Will  we  be  able  to  develop  historians  of  biology  who  will  know 
enough  history  and  enough  science,  as  well  as  a  few  other  essential  disciplines? 

ACKNOWLEDGMENTS 

My  inquiries  into  our  Archives  and  other  sources  in  our  Library  were  greatly 
helped  by  the  assistance  given  to  me  by  Mrs.  Ruth  Davis  and  Ms.  Carol  Horgan, 
Archivists,  Ms.  Jane  Fessenden  and  staff  of  the  MBL  Library,  and  by  Homer  Smith, 


136  S.   S.  COHEN 

the  interested  General  Manager  of  the  MBL.  I  wish  also  to  thank  the  MBL  for  its 
Hospitality  and  permission  to  publish  the  Just-Lillie  items  from  its  Archives.  It  is  a 
great  pleasure  to  acknowledge  weekly  discussions  with  Dr.  Garland  Allen  and  his 
associates  of  the  MBL  History  of  Biology  Seminar,  whose  research  had  impinged  at 
every  turn  on  my  own  studies.  I  also  with  to  thank  Mrs.  Rita  Krant  for  secretarial 
assistance. 

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THE  ZOOLOGICAL  STATION  AT  NAPLES  AND  THE   NEURON: 
PERSONALITIES  AND  ENCOUNTERS   IN  A   UNIQUE  INSTITUTION 

ERNST  FLOREY 

Fakultat  j'iir  Biologic  der  Universitdt  Konstanz,  D-775  Konstan:,  FRG. 

INTRODUCTION 

When  the  young  Dozent  of  Comparative  Anatomy  at  the  University  of  Jena, 
Dr.  Anton  Dohrn,  became  obsessed  with  the  idea  of  building  a  Zoological  Laboratory 
on  the  shores  of  the  Mediterranean  Sea  at  Naples,  his  enthusiasm  was  riding  the 
crest  of  a  wave  of  excitement  that  had  spread  throughout  Europe  and  eventually 
reached  the  North  American  continent.  It  was  the  excitement  generated  by  Darwin's 
new  theory  of  evolution  which  was  so  infectious  because  it  reinforced  the  new 
tendency  towards  a  rational  and  mechanistic  explanation  of  life  phenomena  that 
had  already  been  in  full  development. 

The  science  of  Zoology  had  only  recently  been  established.  Only  five  years 
earlier,  in  1865,  Ernst  Haeckel  had  been  appointed  professor  of  Zoology  at  the 
University  of  Jena  on  the  recommendation  of  the  great  Karl  Gegenbaur,  then 
professor  of  Anatomy  and  Zoology  at  the  same  university,  and  was  provided  with 
a  new  Zoological  Institute.  This  was  the  year  in  which  Anton  Dohrn  received  his 
Ph.D.  degree  at  the  University  of  Breslau  under  Eduard  Adolf  Grube  (who  had 
been  appointed  Professor  of  Zoology  there  in  1857).  Dohrn  had  studied  under 
Haeckel  in  1862,  after  Haeckel  had  just  become  Extraordinarius  of  Comparative 
Anatomy  in  Gegenbaur's  institute.  In  1868,  Dohrn  became  Privatdozent  of  Zoology 
in  Jena,  but  later,  in  the  same  year  we  find  him  already  in  England  and  Scotland, 
and  in  1869  at  Messina  in  southern  Italy,  carrying  out  embryological  studies  on 
marine  organisms.  What  a  restless  character  he  must  have  been:  while  a  student,  he 
changed  universities  five  times!  From  the  University  of  Konigsberg  he  went  to 
Bonn,  then  to  Jena,  moved  to  Berlin,  and  finally  received  his  degree  at  Breslau.  He 
was  indeed  a  man  of  action:  within  a  span  of  only  three  years  after  his  return  from 
Messina  he  had  established  the  Zoological  Station  at  Naples  which  opened  its  doors 
in  1873!  It  must  be  remembered,  however,  that  at  that  time  the  Zoological  Station 
was  not  yet  the  international  institution  it  was  to  become.  The  Stazione  Zoologica 
was  built  with  Dohrn's  private  funds.  Indeed,  the  contract  with  the  city  of  Naples 
which  granted  him  the  right  to  use  the  land  on  which  he  had  built  his  institute  was 
signed,  but  two  years  later.  All  along  it  had  been  Dohrn's  intention,  however,  to 
offer  the  services  of  his  institute  to  scientists  from  all  countries.  To  make  this 
possible  he  sought,  and  obtained,  on  the  recommendations  of  some  of  the  most 
important  scientists  of  his  time,  funds  from  scientific  institutions  and  governments 
of  many  countries  to  whom  he  "rented"  research  facilities:  the  so-called  "tables." 

From  the  beginning  Dohrn  regarded  Zoology  as  an  experimental  science  and 
saw  the  main  thrust  in  the  development  of  morphology,  embryology,  and  physiology. 
He  seized  the  newly  won  status  of  zoology  which  he  described  so  emphatically  in 
his  programmatic  paper  on  "the  present  state  of  Zoology  and  the  founding  of 
zoological  stations"  which  appeared  in  the  Preussische  Jahrbiicher  (vol.  30)  in  1872: 

When  thus  Zoology  with  all  its  branches  has  acquired  new  stature  and  importance, 
it  is  not  surprising  that  in  zoological  circles  everyone  labors  with  redoubled 

137 


138  E.   FLOREY 

energy.  As  after  a  great  victory  the  members  of  the  victorious  nation  appear 
among  the  other  nations  with  elated  selfconfidence  and  are — albeit  grudgingly- 
regarded  by  them  with  increased  respect,  so  appear  the  Zoologists  in  the  midst 
of  the  other  scholars  in  the  full  consciousness  that  it  is  their  science  that  has 
developed  and  brought  to  maturity  the  greatest  concept  of  modern  research,  and 
that  it  is  their  task  to  nurse  and  extend  it,  and  that  the  other  sciences  must 
receive  it  and  must  be  fertilized  and  reformed  by  it. 

In  this  important  publication,  Dohrn  spelled  out  the  new  goals  of  Zoology:  to  study 
the  basis  of  natural  selection  (naturliche  Ziichtung)  and  to  investigate  the  evolutionary 
origins  of  animal  adaptation  to  the  environment.  Dohrn  envisions  an  ecologically 
oriented  comparative  physiology  and  emphasizes  the  need  to  explain  organ  functions 
on  the  basis  of  adaptation  and  evolutionary  history.  The  foundation  of  zoological 
stations  should  provide  the  ideal  places  where  such  a  science  could  be  developed. 

Dohrn  saw,  however,  another  important  task  in  the  foundation  of  marine 
stations:  the  opportunities  they  offer  for  the  rising  generation  of  zoologists.  He 
wanted  to  provide  promising  young  zoologists  with  the  freedom  to  carry  out  research 
in  the  most  conducive  environment  a  zoologist  can  find,  and  to  free  them  from 
financial  worries  for  the  time  of  their  stay  at  the  zoological  station. 

FRIDTJOF  NANSEN:  THE  STRUCTURE  OF  THE  NERVOUS  SYSTEM 

The  motivation  to  encourage  and  support  promising  young  zoologists  must  have 
prompted  Dohrn  to  accept,  in  1886,  a  young  Norwegian  zoologist  by  the  name  of 
Fridtjof  Nansen  as  a  guest  of  the  Stazione  Zoologica,  at  a  time  when  neither 
Norway,  nor  any  of  the  other  Scandinavian  countries  had  reached  financial 
agreements  with  the  Zoological  Station.  Nansen  was  keenly  interested  in  a  problem 
that  engaged  both  physiologists  and  histologists  in  heated  debates:  the  relationship 
between  ganglion  cells  and  nerve  fibers,  the  nature  of  the  nervous  impulse  and  the 
cellular  basis  of  the  functioning  of  the  brain.  Nansen  had  been  led  to  these  questions 
through  his  investigation  of  a  class  of  parasitic  annelids,  the  myzostomids.  When 
he  studied  the  histology  of  their  nervous  system  he  found  himself  confronted  with 
this  fundamental  issue  and  had  to  discover  that  the  existing  research  literature  could 
not  help  to  resolve  it.  He  felt  impelled  to  carry  out  comparative  studies. 

Nansen  has  become  famous  not  for  his  work  in  zoology,  or  neurohistology,  but 
for  his  exploration  of  the  arctic.  With  Sverdrup  he  crossed  Greenland  (two  years 
after  his  stay  at  Naples),  and  from  1890  to  1896  he  carried  out  his  exploration  of 
the  north  pole.  As  high  commissioner  of  the  League  of  Nations  he  introduced  (in 
1921)  the  famous  Nansen-passport,  and  in  1922  he  received  the  Nobel  peace  prize. 
His  earlier  accomplishments  as  a  scientist  and  explorer  have  been  recorded  in  a 
very  readable  biography  by  W.  C.  Broegger  and  Nordahl  Rolfsen:  "Fridtjof  Nansen 
1861-1893"  (1896).  This  book  contains  a  22-page  chapter  entitled  "In  Naples" 
which  describes,  partly  in  Nansen's  own  words  (quoted  from  his  1887  article  in 
Naturen),  the  Zoological  Station  and  the  great  impression  Anton  Dohrn  made  upon 
the  receptive  mind  of  the  young  Nansen. 

Nansen,  then  Curator  at  the  Museum  of  Natural  History  at  Bergen,  Norway, 
was  only  25  years  old  when  he  came  to  Naples.  He  was  dissatisfied  with  the 
prevalent  histological  techniques  then  available  for  the  study  of  the  structure  of  the 
nervous  system.  He  was  open-minded  enough  to  recognize  the  great  potential  of 
the  new  staining  method  invented  and  developed  by  Camillo  Golgi  at  Pavia.  A 
born  explorer,  he  immediately  set  out  to  travel  to  Pavia  to  get  first-hand  knowledge 
of  the  new  method.  In  1885  Nansen  had  won  the  Joachim  Friele  gold  medal  for 


THE  ZOOLOGICAL  STATION   AT  NAPLES 


139 


JOH.  v.  d.  FEHR 


BERGEN. 


FIGURE  1.  Fridtjof  Nansen  sent  this  portrait  to  Anton  Dohrn.  On  its  back  he  wrote  the  following 
dedication:  "Dem  Herrn  Prof.  Dr.  A.  Dohrn  mit  vorziiglicher  Hochachtung  zur  freundlichen  Erinnerung 
von  Fridjof  Nansen  Bergen  3.  April  1887"  (Private  archive  of  the  Dohrn  Family). 


140  E.   FLOREY 

his  work  on  that  peculiar  class  of  annelid  worms,  the  Myzostoma  (today  known  as 
Myzostomida).  He  accepted  the  medal  in  copper  and  used  the  value  of  the  gold  for 
his  traveling  expenses.  After  a  short  stay  at  Pavia  he  continued  to  Naples  where  he 
was  assigned  a  working  space  in  a  large  upstairs  laboratory  already  occupied  by  five 
other  scientists.  The  Zoological  Station  had  already  been  open  for  a  dozen  years. 
Numerous  countries  supported  it  in  exchange  for  the  right  to  one  or  more  "research 
tables"  for  their  respective  scientists.  Norway  at  that  time  did  not  yet  participate, 
and  it  was  due  to  the  generosity  of  the  director  of  the  Statione  Zoologica  that 
Nansen  was  given  the  opportunity  to  spend  two  months  at  this  institution.  Upon 
his  return  to  Norway,  Nansen  published  an  article  on  the  Zoological  Station  in  the 
Norwegian  popular  science  magazine  Naturen.  Excerpts  can  be  found,  in  English 
translation,  in  Brogger  and  Rolfsen's  Nansen  Biography  of  1 896.  Nansen's  enthusiasm 
for  Anton  Dohrn's  work  and  achievements  can  be  gleaned  from  these  quotations: 

The  whole  basement  of  the  great  building  is  fitted  up  as  an  aquarium  for  the 
general  public;  an  aquarium  which  it  would  certainly  be  difficult  to  rival.  This 
great  room,  with  its  many  tanks,  is  soberly  decorated,  with  a  complete  avoidance 
of  all  humbug  [sic]  or  fantastic  ornament,  which  would  only  serve  to  distract  the 
attention  from  its  essential  purposes.  It  has  a  great  attraction  not  only  for  the 
ordinary  traveller,  but  for  the  scientific  student  as  well.  Down  here  he  is  able  to 
pass  hours  in  communion  with  nature,  and  face  to  face  with  the  rarest  of  marine 
organisms,  and  in  a  comparatively  brief  time  he  may  learn  more  of  the  life  of 
the  world  than  he  could  by  long  grubbing  in  volumes  of  printed  wisdom,  or 
rooting  through  the  dead  treasures  of  museums.  He  will  contract  the  habit  of 
using  his  eyes  and  of  his  powers  of  observation  upon  living  nature,  and  learn  to 
regard  life  as  the  essential  object  of  research. 

Acquaintance  with  the  Station,  for  the  majority  of  tourists,  does  not  extend 
beyond  this  room.  Far  more  important  to  science,  however,  are  the  laboratories 
situated  in  the  upper  stories  of  the  building.  Here  naturalists  from  almost  all 
European  countries  are  at  work,  here  they  have  everything  they  can  possibly 
require  for  their  studies.  They  can  come  to  the  Station,  sit  down  at  the  work- 
table  assigned  to  them,  tell  the  Curator,  Salvatore  Lo  Bianco,  what  particular 
animals  they  want,  and  presently  the  animals  are  brought  alive  to  their  very 
tables,  where  they  can  study  them  at  leisure,  with  no  need  to  stir  from  their 
places  except  for  meals  and  sleep.  Instruments,  smaller  tanks  in  which  to  keep 
the  animals  alive,  and  an  excellent  library,  are  all  just  at  hand.  This  concentration 
of  appliances  is  the  novel  and  important  feature  of  the  institution.  ...  If  the 
workers  are  tired  of  the  laboratory,  they  are  free  to  go  out  in  the  vessels  belonging 
to  the  Station,  and  watch  the  gathering  in  of  fresh  specimens.  Beside  several 
fishing  boats,  the  Station  owns  two  small  steamers.  .  .  .  These  steamers  and 
boats  are  equipped  for  dredging,  trawling,  net-fishing,  surface-fishing,  and  so 
forth.  They  are  also  supplied  with  diving  apparatus,  so  that  in  this  way,  too,  you 
can  fetch  up  whatever  you  want. 

What  was  it  that  Nansen  wanted  to  accomplish  at  Naples;  what  actually  did  he 
accomplish?  When  his  biographer  W.  C.  Brogger  inquired  of  Anton  Dohrn  what  he 
remembered  of  Nansen's  stay  in  his  laboratory,  Dohrn  felt  somewhat  embarrassed 
that  he  could  not  be  of  much  assistance;  all  he  could  recall  was  that  Nansen  was 
working  "mainly  on  Amphioxus  and  on  Selachians,  making  use  of  the  new  Golgi 
method."  Dohrn  recalled,  however,  that  Nansen  was  "a  smart  dancer,  who  certainly 
did  not  disdain  the  company  of  lively  ladies.  I  believe  not  to  err  when  I  report  to 
you  that  a  beautiful  Scottswoman  competed  dangerously  with  his  studies  and 
presumably  was  the  cause  of  his  exchanging  Naples  for  Rome  earlier  than  he  had 
originally  intended." 


THE  ZOOLOGICAL  STATION   AT  NAPLES  141 

In  actual  fact,  Nansen  had  used  his  time  very  well  indeed.  Of  course,  the 
research  he  did  at  Naples  was  only  a  part  of  a  wider  ranging  endeavor.  He  continued 
his  comparative  studies  on  the  nervous  system  in  Bergen  where  he  also  received 
from  Naples  more  specimens  of  A  mph ioxus,  "most  excellently  prepared  in  different 
ways  by  Salvatore  Lo'Bianco."  Already  at  Naples  he  had  immersed  himself  in  a 
thorough  study  of  the  research  literature.  It  seems  incredible  that  already  in  1887, 
only  one  year  after  his  return  from  Naples,  where,  after  all,  he  had  spent  only  two 
months,  Nansen  published  a  214-page  monograph  with  11  plates,  entitled  The 
Structure  and  Combination  of  the  Histological  Elements  of  the  Central  Nervous 
System.  Fully  80  pages  of  the  monograph  are  devoted  to  the  history  of  the  subject. 
His  literature  list  is  respectable  and  amounts  to  no  less  than  21  pages.  In  this  very 
illuminating  account  of  the  research  of  others  and  of  his  own,  Nansen  reaches 
conclusions  about  the  general  structure  and  function  of  nervous  systems.  Because 
he  investigates  ganglion  cells  mostly  in  invertebrates,  and  studies  only  the  spinal 
ganglion  cells  of  vertebrates,  he  assumes  that  ganglion  cells  have  mainly  a  nutritive 
function.  He  correctly  interprets  the  nature  of  what  Franz  Leydig  had  called  the 
"Punktsubstanz"  and  what  later  became  to  be  known  as  the  neuropil,  as  being 
composed  of  innumerable  fine  nerve  branches.  He  also  correctly  interprets  the 
course  of  nervous  excitation  during  reflex  actions.  As  to  the  connections  between 
ganglion  cells  and  nerve  fibers  he  is  somewhat  ambiguous  but  favors  the  view  that 
nerve  fibers  are  outgrowths  of  ganglion  cells. 

It  may  seem  surprising  that  Nansen  does  not  once  mention  the  work  and  ideas 
of  Anton  Dohrn  on  the  subject,  but  Dohrn's  research  on  the  nature  of  nerve  cells 
and  ganglion  cells  had  not  yet  advanced  to  the  stage  of  publication,  and  the  subject 
may  never  have  come  up  in  conversation  between  these  two  men.  There  was, 
however,  an  Hungarian  neurohistologist  working  at  the  Zoological  Station  who 
certainly  influenced  greatly  the  thinking  of  Anton  Dohrn,  and  who  also  was  well 
acquainted  with  Fridtjof  Nansen.  He  was  Stephan  von  Apathy  from  Koloszvar. 
Apathy  had  developed  a  staining  method  involving  gold-chloride,  formic  acid, 
methylene  blue,  and  hematein,  which  permitted  the  demonstration  of  a  "Fibrillen- 
gitter"  ganglion  cells  and  nerve  fibers  were  shown  to  contain  fibrils.  Apathy's 
preparations  seemed  to  indicate  that  these  fibrils  formed  a  continuous  network 
throughout  the  nervous  system.  The  fibrils  were  seen  to  cross  from  one  cell  process 
to  the  next  at  their  contact  points.  Apathy  considered  these  fibrils  to  be  the  true 
conducting  structures  of  the  nervous  system.  Apathy  was  indeed  an  excellent 
histologist,  and  his  superb  preparations  convinced  and  influenced  many  of  the 
leading  histologists,  among  them  Wilhelm  His,  Franz  Nissl,  Max  Bielschowsky,  and 
Hans  Held. 

That  it  should  have  been  possible  to  subscribe  to  the  notion  that  the  neurofibrils 
(to  use  a  modern  term)  are  the  "conducting  elements"  of  the  nervous  system  is 
almost  incredible  when  one  remembers  that  electrophysiologists  (Du  Bois-Reymond, 
Hermann,  Bernstein)  had  long  before  established  the  electrical  nature  of  the  nerve 
impulse:  it  would  have  been  inconceivable  to  the  biophysically  trained  physiologists 
that  separate  action  currents  can  be  conducted  within  a  given  nerve  cell  or  even 
within  a  nerve  process.  Evidently,  electrophysiology  (today  this  would  be  called 
biophysics)  was  not  taken  that  seriously  by  the  histologists,  and  even  a  physiologist 
like  Albrecht  Bethe  became  so  impressed  by  Apathy's  results  that  he  was  willing  to 
accept  the  hypothesis  that  the  fibrils  and  not  the  nerve  fibers  are  the  true  elements 
(or  'units'  as  we  would  say  today)  of  the  nervous  system. 

In  their  Nansen  biography,  Brogger  and  Rolfsen  (1896)  quote  a  letter  evidently 
written  to  them  by  Apathy,  who  reminisces  about  the  life  he  and  others  shared  with 


142  E.   FLOREY 

Nansen  while  they  were  at  Naples.  "One  of  these  friends,  a  Hungarian  scientist," 
they  state, 

writes  to  us:  'He  was  the  life  of  all  our  little  festivities.  Most  of  the  students  then 
working  at  the  Station  were  in  the  habit  of  meeting  at  the  Cafe  Basta  on  the 
Corso  Vittoriao  Emmanuele;  every  evening  at  suppertime  there  was  a  little  feast 
here,  a  musical  gathering,  light-hearted  and  refreshing  in  the  highest  degree. 
Nansen  contributed  greatly  to  the  prevailing  gaiety.  It  some  times  happened  that 
we  devotees  of  science  became  so  enlivened  with  wine  and  music,  that  we 
proceeded  to  dance  a  quadrille;  and  on  these  occasions  Nansen  was  Master  of 
Ceremonies. 

'Once  we  chartered  a  carriage  to  drive  to  Castellamare  and  Sorrento  by  the 
famous  coast  road.  On  the  way,  another  carriage  with  two  ladies  came  up  behind 
us.  The  ladies  amused  themselves  by  racing  us  and  laughing  at  us  as  they  shot 
past;  whereupon  Nansen  sprang  out  of  the  carriage  and  ran  by  the  side  of  the 
horse  a  long  stretch  of  the  way.  Thus  we  overtook  the  ladies  again,  to  the 
unbounded  merriment  of  both  parties. 

'In  Sorrento  Nansen  met  some  Norwegian  ladies.  I  was  very  tired  and  went 
to  bed;  but  the  Norwegian  ladies  wanted  to  get  up  a  dance,  and  as  there  was  a 
scarcity  of  partners,  my  presence  was  required.  Nansen  declined  to  give  a 
moment's  peace  till  I  got  up  and  dressed  myself.  Then  he  dragged  me  into  the 
drawing-room,  where  we  were  greeted  with  loud  applause  by  the  ladies,  who 
were  quite  alive  to  the  situation. 

'At  other  times  he  would  be  quiet  and  absorbed,  and  would  sit  by  the  hour 
without  uttering  a  word.  I  have  seen  him  at  the  foot  of  Vesuvius,  among  the 
ruins  of  San  Sebastiano,  and  on  the  melancholy  lava-wastes.  San  Sebastiano  was 
devasted  by  the  eruption  of  1874;  nothing  was  left  but  a  church.  I  have  seen  him 
sitting  on  a  block  of  lava  there  by  the  church,  hour  after  hour  without  stirring; 
he  simply  sat  and  gazed  out  into  the  distance.  Time  after  time  we  others  tried  to 
make  a  start,  and  called  to  him — he  never  moved.  Afterwards,  on  the  way  home, 
as  he  and  I  walked  together,  arm  in  arm,  I  tried  to  make  him  talk,  but  found 
him  absolutely  mute — there  was  not  a  word  to  be  got  out  of  him.' 

ANTON  DOHRN:  THE  NERVE  FIBER 

Before  continuing  with  Stephan  von  Apathy  and  his  impact  through  the 
Zoological  Station  at  Naples,  let  me  return  once  again  to  Anton  Dohrn.  With  an 
immense  effort,  Dohrn  occupied  himself  with  his  "Studien  zur  Urgeschichte  des 
Wirbelthierkorpers"  (Studies  on  the  early  history  of  the  vertebrate  body).  As  by  a 
magnet  he  was  drawn  to  the  problem  of  the  structure  of  the  nervous  system  which 
he  tried  to  resolve  by  studying  lower  vertebrates  and  their  embryological  development. 
It  was  a  heroic  effort  of  trying  to  understand  the  central  nervous  system  of  higher 
vertebrates  on  the  basis  of  Haeckel's  "biogenetic  law,"  the  notion  that  ontogeny 
recapitulates  phylogeny  and  that  any  structure  found  in  higher  organisms  must  have 
its  primitive  counterpart  in  the  earlier  stages  of  evolution  as  represented  in  primitive 
animals.  The  question  was,  in  Dohrn's  words,  "is  the  nerve  fiber  an  outgrowth  of 
the  ganglion  cell?  or  is  it  composed  of  numerous  cells,  the  exponent  of  which  can 
be  regarded  the  Schwann  nuclei?"  Dohrn  called  the  decision  of  this  question  "the 
foundation  of  our  ideas  about  the  nervous  system."  In  the  16th  of  his  "studies"  he 
comes  to  the  definite  conclusion  that  the  nerve  fiber  arises  through  the  fusion  of 
Schwann  cells  and  that  "the  central  ganglion  cells  have  nothing  to  do  with  the 
beginning  of  the  axis  cylinder  or  the  entire  formation  of  the  nerve  fiber."  Ganglion 
cell  and  nerve  fiber  are  connected  by  contact,  they  are  not  genetically  related.  In 
his  views,  Dohrn  finds  himself  supported  by  Apathy  whom  he  quotes  extensively. 


THE   ZOOLOGICAL  STATION   AT  NAPLES  143 

Shortly  after  publication  of  his  16th  "Studie"  in  1891,  Dohrn  was  plagued  by 
doubts  about  the  correctness  of  his  interpretations.  He  rushed  into  print  a  retraction 
in  the  Anatomischer  Anzeiger  (vol.  7,  p.  348).  The  famous  Anatomist  Albert 
Koelliker  had  announced  a  Lecture  to  be  given  at  the  Congress  of  Anatomists  to 
be  held  in  Munich  with  the  ominous  title  "Ueber  die  Entwicklung  der  Elemente 
des  Nervensystems,  contra  BEARD  and  DOHRN."  Because  of  Dohrn's  retraction, 
the  actual  lecture  had  the  simpler  title  ".  .  .  contra  BEARD."  Ten  years  later, 
Dohrn  decided  that  he  should  not  have  retracted  his  views.  His  doubts,  he  said  in 
his  20th  "Studie,"  had  been  mainly  "of  subjective  origin,  and  were  due  to  a  nervous 
depression  caused  by  overwork,  climatic  and  other  influences,  the  like  of  which  I 
unfortunately  had  to  suffer  repeatedly  due  to  the  abrasive  work  and  the  complicated 
conditions  of  life  connected  with  my  position  as  director  of  the  Zoological  Station" 
(p.  139,  vol.  15  of  the  Mitth.  Zool.  Stat.  Neapel,  1901). 

It  would  be  unfair  to  judge  Anton  Dohrn  by  his,  by  present  standards,  misguided 
endeavor  to  solve  once  and  for  all  the  enigma  of  the  cellular  relationship  between 
ganglion  cell  and  nerve  fiber.  Dohrn's  vision  was  far  more  wide  ranging,  and  truly 
important.  He  clearly  saw  the  need  to  encourage  the  development  of  comparative 
physiology  and  biological  chemistry,  and  he  was  able  to  generate  interest  in  the 
enormous  potential  of  a  marine  laboratory  for  the  expansion  of  knowledge  in  this 
important  area  of  experimental  biology.  Many  of  the  important  physiologists  were 
attracted  to  the  Zoological  Station,  among  them  Max  Verworn,  Sigmund  Exner, 
Jaques  Loeb,  Willem  v.  Einthoven,  and  Jacob  von  Ueuxkiill. 

In  1906  Dohrn  created,  with  newly  acquired  funds,  departments  for  physiology 
and  for  physiological  chemistry  in  the  Zoological  Station;  he  employed  H.  Burian 
and  M.  Henze  as  heads  of  these  departments.  Dohrn  motivated  Otto  von  Fiirth  to 
write  the  influential  I'erg/eichende  chemische  Physiologic  niederer  Tiere  (Jena, 
1903),  probably  the  first  text  of  biological  chemistry.  There  can  be  no  doubt  that  it 
was  due  to  the  influence  of  Anton  Dohrn  and  his  Zoological  Station  that  Winterstein 
was  induced  to  publish  the  monumental  Handbuch  der  vergleichenden  Physiologic 
(published  in  eight  volumes  between  1911  and  1925,  together  no  less  than  9321 
pages!):  almost  all  of  the  authors  had  been  working  at  the  Zoological  Station.  Like 
this  Station,  the  Handbuch  was  an  international  affair:  the  contributing  authors 
hailed  from  eight  different  countries. 

Already  in  the  late  1890"s,  Hermann  J.  Jordan  had  been  Dohrn's  private 
assistant.  Later,  Jordan  was  to  become  one  of  the  most  influential  comparative 
physiologists.  On  the  recommendation  of  Jordan,  T.  H.  Morgan  appointed  two  of 
Jordan's  pupils  to  important  posts  at  the  California  Institute  of  Technology: 
C.  A.  G.  Wiersma,  and  A.  van  Harreveld.  The  work  of  these  great  neurobiologists 
thus  reflects  the  heritage  of  Anton  Dohrn. 

ALBRECHT  BETHE:  GANGLION  CELL  AND  REFLEX 

The  neuron  doctrine,  enunciated  by  Wilhelm  Waldeyer  in  his  famous  paper  in 
the  Deutsche  Medizinische  Wochenschrift  of  1891,  was  welcomed  by  physiologists. 
But  neurohistologists  like  Held,  Nissl,  and  others,  continued  either  to  oppose  it  or 
to  regard  it  with  utmost  caution.  In  1896,  Albrecht  Bethe,  a  pupil  of  Goltz  in 
StaOburg,  later  professor  of  Physiology  at  Frankfurt,  came  to  Naples  to  study  the 
nervous  system  of  the  shore  crab,  Carcinus.  His  goal  was  a  complete  histological 
and  physiological  description  of  the  neurons  in  what  he  then  considered  to  be  a 
"simple  nervous  system."  His  encounter  with  Apathy  forced  him  to  completely 
revise  his  ideas  about  structure  and  function  of  nervous  systems.  In  his  important 


144  E.   FLOREY 

book  Allgcmeine  Anatomic  und  Physiologic  des  Nervensystems,  published  in  1903, 
Bethe  writes  that  he  had  been  skeptical  at  first  of  Apathy's  papers: 

because  of  the  aprioristic  form  in  which  they  were  written,  and  because  of  the 
peculiarity  of  the  result  which  were  suported  either  with  no  pictures  at  all,  or 
with  only  schematic  illustrations.  My  doubts  disappeared  rapidly,  however,  when 
Mr.  von  Apathy,  on  the  occasion  of  our  meeting  at  the  Zoological  Station  at 
Naples  in  the  Fall  of  1896,  had  the  kindness  to  show  me  his  preparations.  On 
the  evening  before  this  memorable  day  I  had  still  told  him  that  I  considered 
what  he  had  published  outside  of  all  possibilities,  and  that  it  must  be  due  to  self- 
deception  when  he  thought  that  he  could  follow  such  fine  fibrils  individually  for 
millimeters.  What  has  been  shown  to  me  then,  however,  was  of  such  convincing 
clarity,  that  I  was  forced,  after  some  pretended  objections,  to  relinquish  my 
opposition.  What  happened  to  me  was  experienced  by  many  others,  and  nobody 
of  normal  vision  can  elude  the  convincing  impression  of  Apathy's  preparations 
unless  his  eye  is  beclouded  with  envy  or  injured  vanity. 

As  a  consequence  of  his  "conversion,"  Bethe  looked  at  the  nervous  system  as  a 
syncytium.  The  ganglion  cells  were  unnecessary  to  explain  reflex  actions,  as  he 
demonstrated  by  an  experiment  that  henceforth  would  be  known  in  the  literature 
and  in  physiology  texts  as  the  "Bethe  experiment."  In  Naples,  working  with 
Carcinus,  Bethe  did  the  following  experiment:  he  removed  the  ganglion  cells  that 
surround  the  neuropil  of  the  second  antennae,  severed  the  connections  between  this 
neuropil  and  the  rest  of  the  nervous  system,  and  cut  the  esophageal  commisures. 
He  noted  that  the  antennae  still  maintained  their  tonus  and  that  they  were  held 
stiffly  in  their  normal  raised  position.  When  mechanically  stimulated  the  antennae 
were  retracted,  but  afterwards  were  once  again  extended.  From  this  experiment 
Bethe  concluded  that  ganglion  cells  are  either  unnecessary  for  these  reflexes  to 
occur,  or  the  reflex  arcs  do  not  go  through  the  ganglion  cells.  As  he  was  convinced 
now  that  it  is  the  fibrils  which  serve  as  the  connecting  elements,  he  concluded  that 
the  "Primitivfibrillen"  (the  elementary  fibrils  of  which  the  composite  fibrils  are 
composed)  are  the  true  conducting  elements  of  the  nervous  system. 

Bethe's  experiment  was  widely  quoted  by  those  opposing  the  neuron  doctrine. 
Indeed,  Bethe  himself  declared  "we  must  stop  considering  the  neuron  as  a  physio- 
logical unit  and  must  admit  that  one  and  the  same  neuron  is  capable  of  many 
diverse  actions,  depending  on  which  fibrillar  tract  is  in  operation." 

The  Zoological  Station  at  Naples  was  indeed  a  cross-roads  of  the  biological 
sciences.  The  meeting  there  between  Apathy  and  Dohrn  who  became  close  friends, 
gave  Dohrn  the  needed  confidence  that  he  was  on  the  right  track,  that  he  had  solved 
the  riddle  of  the  fundamental  cellular  nature  of  the  vertebrate  nervous  system. 
Dohrn's  work  in  turn  gave  affirmation,  as  did  Apathy's  work,  to  the  many 
histologists  (and  physiologists)  who  opposed  the  neuron  doctrine.  The  meeting  of 
Apathy  and  Bethe  led  Bethe  to  completely  reconsider  his  ideas  about  the  structure 
and  function  of  the  nervous  system.  In  1893,  Bethe  had  set  out  to  completely 
describe  the  nervous  system  of  what  he  considered  to  be  a  simple  animal  using 
neurohistological  techniques  as  well  as  physiological  experiments.  He  went  to  Naples 
with  the  express  idea  of  "mapping  the  nervous  system,"  of  describing  all  its  neurons 
and  their  interconnections — a  task  which  has  not  been  tried  again  until  C.  A.  G. 
Wiersma  began  his  pioneering  studies  of  the  central  nervous  system  of  crayfish  in 
the  late  1950's  at  the  California  Institute  of  Technology  (see  Identified  Neurons  and 
Behavior  of  Arthropods,  edited  by  G.  Hoyle,  1977).  Bethe  could  have  accomplished 
much  of  what  neurobiologists  started  to  do  six  decades  later,  had  he  not  been 
discouraged  by  the  histological  findings  of  Apathy  which  he  was  able  to  confirm  in 


THE  ZOOLOGICAL  STATION   AT  NAPLES  145 

his  own  work  on  Carcinm  carried  out  at  Naples.  It  is  touching  to  read  the  final 
passage  of  his  third  report  on  his  Carcinus  experiments  which  was  published 
in  1998: 

When  I  began  this  work  three  years  ago,  I  expected  to  advance  with  my  knowledge 
of  the  anatomical  structure  of  the  Nervous  system  of  Carcinus  to  a  point  where 
I  could  describe  about  all  the  nervous  elements  and  their  branches.  After 
preliminary  studies  I  considered  their  interconnections  not  to  be  too  complicated, 
and  I  believed  I  should  be  able  to  clearly  reveal  the  significance  of  each  by 
physiological  experiment.  At  that  time  it  seemed  that  if  this  was  achieved  we 
would  be  very  much  closer  to  an  understanding  of  the  nervous  system.  The 
epochal  work  of  Apathy  has  shaken  this  hope  in  its  foundations. 

It  would  now  be  necessary  to  discover  the  course  of  each  single  fibril — "and  this 
is  unthinkable.  As  I  now  overlook  my  whole  work,  I  reach  the  sad  conclusion  that 
nothing  has  been  gained  from  it  for  our  factual  knowledge.  Were  there  not 
satisfaction  in  the  search  for  knowledge,  one  would  have  to  say  in  resignation:  it  is 
too  difficult  for  us  humans," 

The  "Bethe  experiment'"  on  Carcinus  has  been  a  stumbling  block  for  the  general 
acceptance  of  the  neuron  doctrine  which  holds  that  the  axon  is  an  outgrowth  of  a 
ganglion  cell,  that  all  nerve  cells  are  ganglion  cells  (while  Schwann  cells  like  most 
other  cells  of  the  nervous  system,  are  glia  cells),  and  that  each  neuron  is  a  separate 
unit  which  does  not  fuse  anywhere  with  another  cell,  contacts  being  only  in  the 
nature  of  synapses.  In  1909  the  famous  Otto  Langendorff,  writing  in  Nagel's 
authoritative  Handbuch  der  Physiologic  des  Menschen,  debates  the  Bethe  experiment. 
He  accepts  Bethe's  conclusions  as  valid  for  crustaceans  but  expresses  the  opinion 
that  "reflexes  of  invertebrates  are  perhaps  of  a  lower  level,"  hence  the  neuron  theory 
can  still  be  valid  for  vertebrates.  As  late  as  1927,  another  well  known  German 
physiologist,  Emil  Abderhalden,  states  in  his  textbook  of  Physiology  (Lehrbuch  der 
Physiologic,  1927,  p.  115)  that  it  is  "an  established  fact"  that  the  nervous  system  is 
constituted  of  cells  which  are  interconnected  by  strands  of  fibrils.  Bethe's  experiment 
on  Carcinus  plays  an  important  role  in  Abderhalden's  arguments. 

STEPHAN  VON  APATHY:  NEUROFRIBRILS 

In  the  history  of  neurohistology,  Stephan  von  Apathy  has  been  of  great 
importance.  The  major  breakthrough  was  the  development  of  new  staining  methods 
while  he  was  working  at  the  Stazione  Zoologica  at  Naples.  For  three  years  (1886- 
1889)  Apathy  occupied  the  Hungarian  "research  table"  at  Naples.  Anton  Dohrn 
assigned  to  him  the  task  of  writing  a  monograph  on  the  Hirudinea  for  the  now 
famous  Fauna  und  Flora  des  Golfes  von  Neapel.  Both  men  became  close  friends. 
Apathy  based  much  of  his  later  speculations  on  the  fine-structure  of  the  nervous 
system  on  whole-mount  preparations  of  the  gastrointestinal  tract  of  the  marine 
leech  Pantobdella  muricata.  His  publications,  especially  his  paper  "Das  leitende 
Element  des  Nervensystems  und  seine  topographischen  Beziehungen  zu  den  Zellen," 
published  in  1897  in  the  Zoological  Station's  "house  journal,"  the  then  prestigious 
Mittheilungen  aus  der  Zoologischen  Station  zu  Neapel  (vol.  12),  generated  excitement 
and  heated  debate  among  neurohistologists. 

In  1870,  Apathy  was  appointed  to  the  chair  of  Zoology  at  the  University  of 
Kolozsvar.  He  was  27  years  old.  A  few  years  later  he  was  also  in  charge  of  the 
chairs  of  histology  and  embryology  (see  the  biography  by  A.  Abraham,  1963). 
Kolozsvar  is  the  Hungarian  name  of  the  former  capital  of  Transylvania,  an  old 
Hungarian  settlement  which,  mostly  in  the  13th  century,  received  German  settlers 


146  E.   FLOREY 

and  was  named  Klausenberg.  It  became  part  of  the  Austrian  empire  in  1691,  and 
was  claimed  and  occupied  by  Hungary  in  1848.  As  Albrecht  Bethe  remembers  in 
his  obituary  of  Stephan  von  Apathy,  who  died  in  1922,  von  Apathy  returned 
unopened  any  letter  addressed  to  him  at  "Klausenburg."  His  Hungarian  nationalism 
was  so  strong  that  he  refused  to  travel  through  Austria,  an  attitude  which  made  it 
a  matter  of  some  complexity  to  reach  the  Zoological  Station  at  Naples.  Stimulated 
by  the  example  of  Anton  Dohrn's  accomplishments  at  Naples,  Apathy  instituted  a 
"table  system"  at  his  institute  and  provided  foreign  scientists  with  laboratory  space 
where  they  could  carry  out  histological  studies  and  get  acquainted  with  his  widely 
acclaimed  techniques.  Among  his  guests  were  Albrecht  Bethe,  Wilhelm  Waldeyer, 
and  a  Dutch  histologist,  J.  Boeke,  who  later  became  a  famous  neurohistologist  and 
extended  Apathy's  studies  to  the  mammalian  autonomic  nervous  system.  Boeke 
remained  critical  of  the  neuron  theory  and  maintained  that  autonomic  nerve  fibers 
terminated  in  a  terminal  reticulum  composed  of  (neuro)fibrils.  Typical  of  his  point 
of  view  is  his  paper  of  1949  "The  sympathetic  end  formation,  its  synaptology,  the 
interstitial  cells,  the  periterminal  network,  and  its  bearing  on  the  neuron  theory" 
(Acta  Anatomica  8:  18-61).  Like  Anton  Dohrn,  Apathy  traveled  widely  to  many 
European  universities  and  became  personally  acquainted  with  the  best  scientists  of 
his  day.  He  put  his  whole  effort  into  establishing  a  new  "Zoological  Station"  at 
Koloszvar,  and  in  1909  his  new  Zoological  Institute,  one  of  the  finest  in  all  of 
Europe,  was  opened — complete  with  loggias  and  extensive  facilities  for  the  mainte- 
nance of  freshwater  and  marine  animals.  As  in  Naples,  the  public  facilities  (the 
aquaria,  museum,  lecture  rooms),  were  on  the  lower  floor,  the  two  upper  floors 
contained  the  research  laboratories,  the  administration,  and  the  library. 

How  Apathy  would  have  liked  to  show  off  his  accomplishment  to  his  friend 
Dohrn!  Fate  decided  otherwise:  on  3  October  1909,  Apathy  attended  the  funeral 
service  for  Anton  Dohrn  at  Jena. 

With  the  end  of  the  first  world  war,  the  golden  era  of  Apathy's  institute  and  of 
his  scientific  career  came  to  a  sudden  end.  Apathy  had  become  a  politician  but 
could  not  prevent  the  take-over  of  Transylvania  by  Rumania.  When  he  was  released 
from  prison,  he  accepted  a  position  at  the  Hungarian  University  at  Szeged  and  tried 
once  again  to  create  a  new  zoological  institute.  He  no  longer  had  the  strength;  he 
died  two  years  later. 

J.  C.  ALEXANDROWICZ:  STRETCH  RECEPTOR  NEURONS 

Stephen  von  Apathy  had  survived  Anton  Dohrn  by  eleven  years.  In  the 
meantime,  the  directorship  of  the  Zoological  Station  at  Naples  had  been  transferred 
to  Anton  Dohrn's  son  Reinhard  Dohrn,  who,  except  for  his  years  of  exile  during 
1915-1924,  conducted  the  affairs  of  the  Zoological  Station  at  Naples,  guiding  this 
prestigious  and  precious  institution  through  the  political  turmoils  of  a  nationalistic 
era  and  the  so-difficult  war  and  post-war  years  until  he  relinquished  his  leadership 
to  his  son  Pietro  Dohrn  in  1954.  (The  accomplishments  of  Pietro  Dohrn  and  the 
later  history  of  the  Zoological  Station  have  been  critically  reviewed  in  leading 
articles  in  Science,  1969,  and  in  Nature,  1983). 

Reinhard  Dohrn's  diplomatic  activities  succeeded  in  1924  to  reach  an  agreement 
with  Polish  authorities  to  establish  a  Polish  research  table  at  the  Stazione  Zoologica. 
One  of  the  faithful  scientists  using  this  table  was  J.  C.  Alexandrowicz,  professor  of 
ophthalmology,  and  from  1937  Undersecretary  of  State  in  the  Polish  Ministry  of 
Education.  Alexandrowicz's  meticulous  neuro-anatomical  studies  of  crustaceans  and 
cephalopods  using  methylene-blue  staining  techniques  have  earned  him  a  sepcial 


THE  ZOOLOGICAL  STATION   AT  NAPLES  147 

place  in  neurobiology.  Perhaps  his  most  important  discovery,  rivaled  only  by  J.  Z. 
Young's  discovery  of  the  giant  axons  of  squid,  are  the  stretch  receptor  organs  of 
crustaceans.  Although  this  discovery  was  made  at  Naples  just  before  the  second 
world  war,  it  became  known  to  the  scientific  world  only  several  years  after  the  end 
of  the  war:  when  the  war  started,  Alexandrowicz  became  an  officer  of  the  Polish 
military  medical  corps.  After  the  defeat  of  Poland  when  this  country  was  divided 
up  between  Germany  and  the  Soviet  Union,  Alexandrowicz  was  taken  prisoner  by 
the  Russian  army,  and  then  was  sent  with  the  Polish  expeditionary  force,  known  as 
the  Anders  Army,  to  North  Africa  to  help  the  British  defeat  the  Germans.  The 
contingent,  in  which  he  served  as  education  officer,  never  saw  action.  When  the 
war  ended,  Alexandrowicz  was  taken  to  England  to  become  a  farm  laborer.  It  was 
Reinhard  Dohrn  who  traced  him  with  the  aid  of  the  Red  Cross,  and,  through  his 
connections  with  members  of  the  Royal  Society,  initiated  the  establishment  of  a 
special  professorship  for  Alexandrowicz  at  the  Marine  Laboratory  at  Plymouth. 
Alexandrowicz  had  lost  all  his  valuable  preparations.  He  now  repeated  his  investi- 
gations and  in  1952  began  his  series  of  publications  on  the  structure  and  histology 
of  crustacean  stretch  receptors  that  have  become  classics. 

Alexandrowicz's  investigations  are  the  basis  of  important  physiological  work 
that  was  begun  almost  immediately  after  their  publication  and  led  to  those 
discoveries  (e.g.,  Kuffier  and  Eyzaguirre,  1955)  that  have  become  the  key  to  our 
understanding  of  how  sensory  neurons  translate  a  stimulus  into  a  series  of  nerve 
impulses,  how  they  encode  stimulus  strength  into  an  impulse  frequency.  Alexan- 
drowicz described  an  efferent  innervation  of  the  stretch  receptor  neurons  which  was 
shown  later  to  be  purely  inhibitory  (Eyzaguirre  and  Kuffler,  1955).  The  crustacean 
stretch  receptor  preparation  thus  became  an  important  tool  in  the  investigation  of 
inhibitory  synaptic  transmission.  It  was  in  these  stretch  receptor  neurons  that  the 
first  evidence  was  obtained  that  7-aminobutyric  acid  (GABA)  is  the  transmitter 
substance  of  inhibitory  neurons,  and  that  the  transmitter  action  can  be  blocked  by 
picrotoxin  (Florey,  1953;  Bazemore  et  ai,  1957).  Indeed,  it  can  be  said  without 
exaggeration  that  the  stretch  receptor  neurons  discovered  by  Alexandrowicz  at  the 
Zoological  Station  at  Naples  have  been  a  cornerstone  in  the  development  of 
neurophysiology.  To  mention  only  two  of  the  key  findings:  it  was  on  stretch  receptor 
preparations  that  the  Japanese  physiologist  K.  Uchizono  showed  for  the  first  time 
that  inhibitory  nerve  terminals  are  characterized  by  clear  oval  synaptic  vesicles — in 
contrast  to  cholinergic  terminals  which  always  contain  clear  round  vesicles.  The 
work  of  the  Swedish  physiologist  D.  Ottoson  on  isolated  stretch  receptor  neurons 
provided  the  first  clear  proof  that  the  site  of  initiation  of  the  nerve  impulse  is  not 
the  soma  of  the  nerve  cell,  but  the  initial  segment  of  its  axon. 

ERNST  SCHARER:  NEUROSECRETION 

The  Zoological  Station  has  been  instrumental  in  yet  another  important  advance 
in  the  field  of  neurobiology:  the  discovery  of  what  has  become  known  as  "neurose- 
cretion,"  the  elaboration  and  secretion  of  hormones  by  nerve  cells.  In  1928  Ernst 
Scharrer,  then  Assistent  under  Karl  von  Frisch  at  the  Zoological  Institute  of  the 
University  of  Munich,  was  granted  the  use  of  a  research  table  at  the  Zoological 
Station  at  Naples.  As  he  stated  in  a  letter  to  Reinhard  Dohrn,  he  wanted  to  fix  the 
brain  of  many  species  of  fish,  and  "if  possible  to  investigate,  with  the  aid  of 
methylene  blue  staining,  the  innervation  of  the  epiphysis."  He  discovered  that 
certain  neurons  in  the  midbrain  show  evidence  of  secretion,  confirming  earlier 
findings  of  Carl  Speidel  (1919).  With  material  from  Naples,  Scharrer  continued  his 


148  E.   FLOREY 

studies  on  these  "neuroglandular"  cells  of  fishes  and  extended  these  studies  to  higher 
vertebrates  where  the  same  cell  type  was  found.  After  he  had  married  another  pupil 
of  von  Frisch,  Berta  Scharrer,  the  inseparable  couple  continued  to  explore  the 
comparative  aspects  of  "neurosecretion,"  as  the  phenomenon  was  soon  to  be  called: 
Berta  Scharrer  in  invertebrates,  Ernst  Scharrer  in  vertebrates.  Their  association  with 
Reinhard  Dohrn  and  the  Zoological  Station  became  important  not  only  for  the 
development  of  this  important  field  of  neurobiology;  the  political  situation  in 
Germany  made  life  intolerable  for  the  Scharrers,  and  it  was  Reinhard  Dohrn  who 
helped  with  their  emigration  to  the  United  States. 

Already  before  the  end  of  the  second  world  war,  in  1944,  American  scientists 
like  R.  E.  Cooker  (Chapel  Hill),  A.  R.  Moore  (Oregon),  and  Ernst  Scharrer,  urged 
the  president  of  the  National  Research  Council  (the  precursor  of  the  National 
Science  Foundation)  and  Italian  authorities  to  reopen  the  Statione  Zoologica  and 
to  support  the  directorship  of  Reinhard  Dohrn.  Thanks  to  the  untiring  effort  of 
Ernst  Scharrer,  who  had  moved  from  Ohio  to  the  Department  of  Anatomy  at 
Denver/Colordao,  the  National  Research  Council,  as  well  as  Columbia  University 
(Wilson  Fund),  the  American  Association  for  University  Women,  and  the  American 
Society  of  Zoologists  were  persuaded  to  contribute  funds  in  support  of  the  Naples 
Zoological  Station.  In  the  summer  of  1946,  the  president  of  the  National  Research 
Council,  Ross  Granville  Harrison  (whose  activities  at  Woods  Hole  are  discussed  in 
other  contributions  of  this  Symposium)  set  up  a  Committee,  chaired  by  Ernst 
Scharrer,  to  aid  the  Zoological  Station  at  Naples.  The  Committee,  which  included 
as  its  members  E.  G.  Conklin,  Mrs.  E.  B.  Harvey,  R.  G.  Harrison,  S.  Hecht,  L.  H. 
Kleinholz,  A.  R.  Moore,  and  H.  H.  Plough,  met  for  the  first  time  on  14  August 
1946  at  Woods  Hole  to  decide  on  a  program-in-aid  to  assist  in  the  re-establishment 
of  the  Zoological  Station  at  Naples  as  an  international  center  of  biological  research. 
The  program  envisioned  the  establishment  of  additional  American  research  tables, 
a  fund-drive  to  improve  and  enlarge  the  library,  and  a  shipment  of  food  for  the 
Mensa.  Further  plans  concerned  the  modernization  of  the  laboratory  facilities  and 
research  equipment.  The  activities  of  this  committee  led  to  contributions  by  the 
Rockefeller  Foundations  and  the  UNESCO.  The  committee  advised  the  Zoological 
Station  in  matters  of  library  acquisitions  and  the  purchase  of  research  equipment. 
After  1950  the  newly  established  National  Science  Foundation,  the  Lilly  Endowment, 
Inc.,  and  the  Rockefeller  Foundation  increased  the  American  engagement  in  the 
affairs  of  the  Zoological  Station  enormously — but  this  is  not  the  place  to  record  the 
international  ramifications  of  the  Zoological  Station  and  the  history  of  the  material 
support  it  has  received  from  the  international  scientific  community  (which,  after 
all,  includes  members  of  other  nations  that  have  made  substantial  contributions  to 
the  development  of  the  Zoological  Station).  We  return,  therefore,  to  the  topic  of 
the  role  of  the  Zoological  Station  at  Naples  in  the  development  of  neurobiology. 

On  the  initiative  of  Ernst  Scharrer  and  Wolfgang  Bargmann,  the  Zoological 
Station  hosted,  in  1953,  the  first  International  Symposium  on  Neurosecretion  (this 
was  the  fourth  international  symposium  held  at  the  Zoological  Station).  This  event 
has  a  special  place  in  the  history  of  biology  because  it  was  this  symposium  that 
established  the  concept  that  neurons  produce  hormones  and  that  neurosecretion  is 
an  essential  feature  of  the  chemical  control  of  animal  development  and  function. 

J.  Z.  YOUNG:  GIANT  AXONS,  LEARNING  AND  MEMORY 

It  is  impossible  to  review  the  relationship  between  the  Zoological  Station  and 
the  neuron  without  mentioning  the  research  on  the  giant  axons  of  squid,  discovered 


THE  ZOOLOGICAL  STATION   AT   NAPLES  149 

by  J.  Z.  Young  in  1936  when  he  worked  at  this  institution.  Since  Prof.  Young  will 
review  the  history  of  this  discovery  in  his  lecture,  it  will  be  sufficient  to  restrict  the 
discussion  here  to  some  further  development  of  research  made  possible  by  the 
incredibly  large  size  of  the  "giant  synapses"  between  second-  and  third-order  giant 
axons.  In  1966  Berhard  Katz  and  Riccardo  Miledi  from  the  University  College  in 
London  came  to  the  Stazione  Zoologica  to  investigate  the  relationship  between 
calcium  and  transmitter  release.  Their  experiments  have  become  classics;  they  prove 
that  extracellular  calcium  is  essential  for  transmitter  release  to  occur  and  that 
calcium  ions  enter  the  nerve  terminal  when  this  becomes  depolarized  by  the 
incoming  presynaptic  action  potential. 

The  year-round  availability  of  Octopus  at  Naples,  and  the  recognition  of  the 
advanced  development  of  the  brain  of  these  animals  has  prompted  Y.  Z.  Young  to 
embark,  at  the  Statione  Zoologica,  on  a  study  of  learning  and  memory  in  these 
creatures.  This  was  made  possible  by  a  large  grant  to  the  Stazione  Zoologica  for  the 
establishment  of  a  large  "cephalopod  facility"  which,  in  its  best  days,  included  more 
than  two  hundred  tanks  in  which  as  many  octopuses  could  be  individually  housed 
and  maintained.  Together  with  several  collaborators,  especially  Bryan  Boycott, 
Martin  Wells,  and  John  Messenger,  he  mapped  the  neuronal  circuits  of  the  octopus 
brain  and  through  ingenious  training  experiments  he  explored  the  learning  ability 
of  these  animals.  These  studies  led  to  new  concepts  of  the  neuronal  mechanisms 
underlying  memory.  Several  important  monographs  resulted  from  this  research: 
M.  J.  Wells:  Brain  and  Behaviour  in  Cephalopods.  1962;  J.  Z.  Young:  A  Model  of 
the  Brain,  1964;  J.  Z.  Young:  The  Anatomy  of  the  Nervous  System  of  Octopus 
vulgaris,  1971;  and  J.  Y.  Young:  Programs  of  the  Brain,  1978. 

CONCLUSION 

By  providing  research  facilities  near  the  sea  where  marine  animals  can  be  readily 
obtained  and  maintained,  and,  more  importantly  perhaps,  by  providing  the  intellectual 
atmosphere  conducive  to  intensive  research  and  stimulating  interaction  with  other 
scientists,  the  Stazione  Zoologica  has  permitted  major  advances  in  neurobiology  to 
occur.  As  long  as  this  institution  was  able  to  pursue  the  goals  envisioned  by  its 
founder,  Anton  Dohrn,  it  was  eminently  successful.  But  such  simple  words  cannot 
explain  the  impact  the  Zoological  Station  at  Naples  had  on  biologists  all  over  the 
world.  Intentionally  I  use  the  word  'biologists'  and  not  the  abstract  form  'biology.' 
The  Stazione  Zoologica  has  been  dominated  by  the  spirit  of  its  former  directors,  by 
the  immense  human  dimension  of  its  founder  Anton  Dohrn,  by  that  great  European, 
Reinhard  Dohrn  ...  all  this  has  been  attested  to  by  so  many  public  statements, 
that  no  further  emphasis  is  needed.  The  most  recent  eulogies  were  presented 
(typically,  in  four  languages)  by  dignitaries  from  many  countries  on  the  occasion  of 
the  celebration  of  the  100th  birthday  of  Reinhard  Dohrn  on  13  March  1980  in  the 
Vila  Pignatelli  in  Naples  (Reinhard  Dohrn  1880-1962,  edited  by  C.  Groeben,  1983). 

And  yet,  neither  the  location  nor  the  personalities  of  the  directors  can  explain 
the  miracle  of  the  "Naples  experience"  (to  quote  Maurice  Wilkins)  or  the  affection 
all  those  great  scientists  felt,  and  still  feel,  for  the  Zoological  Station.  Good  research 
institutes  can  be  found  in  many  places  in  many  countries,  and  the  Statione  Zoologica 
is  certainly  not  among  the  best  equipped  laboratories — perhaps  it  never  was!  Nor 
has  it  been  attractive  because  it  harbored  a  local  scientific  genius  at  whose  feet  it 
was  desirable  to  sit,  in  whose  laboratory  it  was  essential  to  learn  methods  unattainable 
anywhere  else.  The  idea  of  an  international  home  for  an  international  science,  nay, 
for  the  unfettered  pursuit  of  the  highest  ideal  of  science,  this  original  idea  of  Anton 


150  E.   FLOREY 

Dohrn  was  so  infectious,  that,  from  the  start,  it  caught  the  imagination  of  all  those 
great  minds  who  came  in  contact  with  it:  Charles  Darwin,  Thomas  Henry  Huxley, 
Thomas  Hunt  Morgan,  Edward  Beecher  Wilson,  Hermann  von  Helmholtz,  Emil 
Du-Bois  Reymond,  Filippo  Botazzi,  Silvestro  Baglioni — and  Fridjof  Nansen,  Stephan 
von  Apathy,  Albrecht  Bethe,  J.  C.  Alexandrowicz,  Ernst  Scharrer,  J.  Z.  Young,  but 
also  of  Benedetto  Croce,  Theodor  Heuss,  .  .  .  the  list  is  endless. 

The  Zoological  Station  became  a  place  of  the  mind,  an  ideal  jointly  possessed 
and  cherished  by  all  those  who  experienced  it.  This  is  the  reason  why  the  organism 
of  the  Zoological  Station  was  able  to  survive:  not  because  it  was  itself  strong  enough 
to  surmount  all  the  adversities  it  encountered  in  its  long  history,  but  because  it  was 
revived  from  the  outside  on  the  strength  of  the  idea  carried  by  the  community  of 
all  those  who  kept  this  idea  alive  and  found  ways  to  revitalize  it  both  spiritually 
and  materially.  Science  is  not  an  abstraction  but  an  immensely  human  activity.  It 
is  lived,  not  written,  thus  it  needs  a  true  home,  not  only  a  laboratory  or  an  office. 
The  Zoological  Station  at  Naples  has  been  such  a  home  and  has  been,  and  is  being 
regarded  with  that  special  kind  of  nostalgia  accorded  only  to  those  special  places  in 
which  the  true  spirit  of  man  is  recognized.  Thomas  Hunt  Morgan  called  it  a  "holy 
city."  Our  present  age  would  do  well  to  live  up  to  its  tradition. 

The  brief  histories  given  here  are  the  stories  of  important  scientists  and  of  their 
important  discoveries.  But  they  are  also  memorials  of  great  ideals,  passions,  and 
sacrifices,  and  they  bear  witness  to  the  importance  of  a  great  dream  which  happened 
to  come  to  life  at  Naples  in  the  Zoological  Station,  the  creation  of  that  remarkable 
man,  Anton  Dohrn. 

ACKNOWLEDGMENTS 

The  passages  quoted  from  the  writings  of  Anton  Dohrn,  Albrecht  Bethe,  and 
Ernst  Scharrer,  have  been  translated  into  English  by  myself.  I  am  grateful  to  the 
librarian  of  the  Stazione  Zoologica,  Walter  Groeben,  for  making  available  to  me 
the  original  sources  of  the  scientific  literature  referred  to.  I  am  especially  grateful  to 
Christiane  Groeben  for  providing  access  to  the  Dohrn  Archives  which  she  has  so 
impressively  filed  and  organized,  and  for  providing  copies  of  relevant  documents 
and  correspondence.  I  am  indebted  to  Professor  John  Edwards  of  the  University  of 
Washington  for  introducing  me  to  the  Nansen  biography  of  Brogger  and  Rolfsen 
(1896),  and  for  lending  me  his  copy  of  Fridjof  Nansen's  so  important,  and  unjustly 
forgotten  work  The  Structure  and  Combination  of  the  Histological  Elements  of  the 
Central  Nervous  System  (1886).  Much  of  the  biographical  and  historical  information 
on  the  history  of  the  Zoological  Station  stems  from  the  invaluable  recent  work  by 
Karl  Josef  Partsch  Die  Zoologische  Station  in  Neapel,  Model!  internationaler 
Wissenschaftszusammenarbeit  which  provides  extensive  documentation. 

Important  sources  were  the  Geschichte  der  Mikroskopie  edited  by  H.  Freund 
and  A.  Berg  (vols.  1  and  2,  1963,  1964),  Alfred  Kiihn's  important  work  Anton 
Dohrn  und  die  Zoologie  seiner  Zeit  (1950),  Theodor  Heuss'  biography  of  Anton 
Dohrn  (2nd  edition,  1962),  and  the  Festschrift  Reinhard  Dohrn  1880-1962  edited 
by  Christiane  Groeben  in  collaboration  with  Antonie  and  Pietro  Dohrn  (1983). 

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YOUNG,  JOHN  Z.   1971.  The  Anatomy  of  the  Nervous  System  of  Octopus  vulgaris.  Clarendon  Press, 

Oxford.  690  pp. 
YOUNG,  JOHN  Z.  1978.  Programs  of  the  Brain.  Oxford  University  Press,  Oxford.  325  pp. 


Reference:  Biol.  Bull.  168  (suppl.):  153-158.  (June,  1985) 


CEPHALOPODS  AND  NEUROSCIENCE 

J.  Z.  YOUNG 

The  Wellcome  Institute  for  the  History  of  Medicine,  183  Euston  Road,  London  NW1  2BP,  Great  Britain 

INTRODUCTION 

Study  of  cephalopods  at  marine  laboratories  has  provided  material  for  some  of 
the  outstanding  discoveries  of  neuroscience  in  this  century.  The  giant  nerve  fibers 
are  the  most  conspicuous  example,  but  studies  of  photoreceptors  and  the  memory 
mechanisms  of  the  brain  have  been  very  fruitful,  as  has  work  on  chromatophores 
and  many  other  topics.  It  would  be  impossible  to  summarize  all  this  work  but  it 
may  be  interesting  to  show  the  sequence  in  which  some  of  it  has  developed  at 
Naples,  Plymouth,  and  Woods  Hole,  in  much  of  which  I  have  been  concerned. 

EYES 

Perhaps  the  earliest  contribution  of  cephalopods  to  fundamental  neural  processes 
was  the  discovery  of  the  electroretinogram  by  Frohlich  at  Naples  in  1914.  The 
curious  electrical  phenomena  in  the  rhabdomes  are  still  only  partly  understood  and 
have  been  the  subject  of  many  later  investigations.  Outstanding  has  been  the 
demonstration  by  Hagins  and  McGaughty  (see  Messenger,  1981)  that  the  opening 
of  channels  to  produce  generator  potentials  takes  place  locally,  near  the  site  of 
photon  absorbtion  in  a  rhabdome.  Speaking  of  the  retinal  pigments  will  remind  us 
of  the  use  made  by  Hubbard  and  Wald  at  Woods  Hole  of  cephalopod  eyes  to 
provide  the  rhadopsin  for  their  fundamental  research.  Indeed  the  history  of  the  use 
of  these  eyes  for  neuroscience  merits  a  symposium  of  its  own.  Cephalopods  appear 
to  have  no  color  vision,  in  spite  of  their  own  colorful  displays  (Messenger,  1981). 
But  the  capacity  to  detect  the  plane  of  reflected  polarized  light,  first  suggested  by 
the  geometry  of  the  rhabdomes  (Young,  1960;  Saibil,  1982)  and  proved  experimentally 
at  Naples  by  Moody  and  Parris  (1961)  and  by  electrical  recordings,  may  provide  a 
sort  of  substitute  for  color  vision  (Saidel  et  al.,  1983). 

STATOCYSTS 

The  statocyst  is  an  organ,  one  of  whose  major  functions  is  stabilization  of  the 
visual  image.  It  has  been  investigated  at  Naples  by  Young  (1960)  and  Wells  (1960). 
Thorough  studies  have  been  made  by  Budelmann  at  Regensburg  using  large  numbers 
of  octopuses  and  cuttlefishes  carried  alive  from  Naples.  He  and  I  have  analyzed  the 
oculomotor  control  system  (Budelmann  and  Young,  1985).  Recently  he  has  discov- 
ered that  an  octopus  monitors  its  fast  and  slow  movements  separately  by  a  unique 
system  of  large  and  small  cupulae  (Budelmann  and  Williamson,  1985). 

Another  recent  development  has  been  the  discovery  that  the  peduncle  lobe  and 
basal  lobes  of  the  brain  contain  systems  of  small  cells  with  parallel  fibers.  These 
resemble  the  vertebrate  cerebellum  and  like  that  organ  are  involved  in  the  optomotor 
reflexes. 

Measurement  of  the  statocysts  of  many  species  collected  from  Plymouth,  Naples, 
Woods  Hole,  Miami,  and  Hawaii  have  shown  a  system  similar  to  the  semicircular 
canals  of  vertebrates  (Stephens  and  Young,  1975;  Maddock  and  Young,  1984).  The 

153 


154  J.  Z.   YOUNG 

canals  are  formed  by  a  series  of  knobs,  the  anticristse.  They  are  best  developed  in 
the  rapidly  moving  loliginids  and  ommastrephids.  In  the  slowly  moving  neutrally 
buoyant  forms,  the  statocyst  is  large  and  empty. 

EXTRAOCULAR  PHOTORECEPTORS 

What  may  be  called  the  modern  epoque  of  cephalopod  research  at  Naples  was 
begun  by  Enrico  Sereni  who  made  many  experiments  on  the  chromatophores  and 
salivary  secretion,  summarized  in  a  long  article  in  1930.  At  the  time  of  his  early 
death  he  and  I  were  collaborating  in  a  study  of  regeneration  of  the  stellar  nerves, 
which  was  completed  after  his  death  (Sereni  and  Young,  1932).  In  the  course  of 
this  study  I  was  attracted  to  a  small  orange  spot  at  the  hind  end  of  the  stellate 
ganglion  of  Eledone.  I  cut  sections  of  it,  with  no  hypothesis  other  than  curiosity.  It 
proved  to  be  a  hollow  vesicle  into  which  passed  a  number  of  projections,  apparently 
of  nerve  cells.  After  discussion  with  Sereni  it  was  named  the  epistellar  body  (Young, 
1929).  I  was  interested  at  the  time  in  the  vertebrate  adrenals  and  made  the  hypothesis 
that  these  projections  into  the  epistellar  cavity  were  secretory.  Ernst  Scharrer  eagerly 
seized  upon  this  as  one  of  the  earliest  examples  of  neurosecretion.  But  he  and  I 
were  sadly  mistaken.  Forty  years  later  Howard  Bern,  himself  an  endocrinologist, 
thought  it  time  to  study  this  organ  properly.  The  E.M.  quickly  showed  that  the 
processes  inside  the  epistellar  body  are  rhabdomes  (Nishioka  et  al.,  1966).  It  is  not 
a  gland  at  all  but  a  photoreceptor,  though  without  any  lens  or  other  dioptric 
apparatus.  Alex  Mauro  working  at  Naples  and  Ischia  confirmed  that  it  produces  its 
own  minielectroretinogram  (Mauro  and  Baumann,  1968).  What  can  this  photore- 
ceptor be  doing  inside  the  mantle?  The  epistellar  body  is  especially  large  in  deep- 
sea  octopods,  which  are  transparent.  One  hypothesis  is  that  it  serves  to  detect  the 
presence  of  a  mass  of  luminous  material  in  the  mantle,  which  would  attract  a 
predator.  The  oesophagus  is  deeply  pigmented,  presumably  for  the  same  reason. 
However  Houck  (1982),  working  at  Hawaii,  has  recently  shown  that  in  octopuses 
with  the  optic  nerves  cut  diurnal  rhythms  can  still  be  entrained  by  light,  perhaps 
detected  by  the  epistellar  bodies. 

The  extra-ocular  photoreceptors  in  decapods  are  in  the  head,  not  on  the  stellate 
ganglion  (Thore,  1939;  Boycott  and  Young,  1956).  They  have  been  thoroughly 
studied  by  R.  E.  Young  (1978)  at  Hawaii  in  many  species  of  squid.  In  some 
mesopelagic  forms  such  as  Abraliopsis  they  serve  to  monitor  the  downward 
illumination  emitted  by  photophores  for  countershading.  For  this  light  to  be  effective 
in  making  the  squid  invisible  from  below  it  must  match  the  downwelling  light.  This 
match  is  ensured  by  the  photosensitive  vesicles  which  are  in  two  sets,  one  looking 
up  to  the  surface  and  the  other  towards  the  animal's  own  luminous  organs  (Young 
and  Roper,  1976).  The  system  even  ensures  an  appropriate  match  to  the  wave 
length,  if  necessary,  in  moonlight! 

The  extra-ocular  photoreceptors  are  even  larger  in  the  bathypelagic  squids,  such 
as  the  cranchiids,  many  of  which  proceed  to  depths  beyond  the  range  of  daylight, 
especially  for  reproduction.  Here  the  photoreceptors  must  have  another  function.  I 
suggest  that  they  monitor  the  depth  at  which  to  spawn.  They  provide  huge  irregular 
masses  of  photosensitive  material  and  their  nerves  connect  with  the  peduncle  lobe 
of  the  brain,  which  is  probably  concerned  with  movement  in  the  vertical  plane.  It 
may  be  that  the  squids  continue  to  proceed  deeper  and  deeper  until  no  photons  are 
captured  even  by  these  large  masses  of  pigment.  When  the  light  goes  out  it  is  dark 
enough  to  breed!  Conversely  the  photoreceptors  prevent  rising  into  the  dangerous 
lighted  zone. 


CEPHALOPODS  AND  NEUROSCIENCE  155 

GIANT  FIBERS 

These  discoveries  are  all  exciting  but  even  greater  developments  have  flowed 
from  my  original  curiosity  about  the  epistellar  body.  Having  found  it  in  octopods  I 
naturally  also  made  sections  of  the  stellate  ganglion  of  decapods.  No  epistellar  body 
was  there  but  instead  I  found  the  giant  nerve  fibers.  I  am  often  asked  at  what  date 
this  discovery  was  made  but  can  give  no  clear  answer.  Sections  of  the  ganglia  of 
Loligo  were  made  at  Naples  in  1929  but  at  first  I  thought  these  large  spaces  were 
veins.  The  axoplasm  does  indeed  look  quite  like  blood  in  some  sections.  Then  I 
followed  them  towards  the  hind  end  of  the  ganglion  where,  as  we  now  know,  they 
originate  by  the  fusion  of  the  axons  of  many  cells.  This  seemed  to  me,  as  a  faithful 
Oxford  follower  of  Sherrington,  to  be  so  unlikely  that  it  took  some  years  to  persuade 
myself  of  it.  However  in  the  collection  there  is  a  slide,  labeled  in  my  handwriting 
"1  •  5  •  30,"  which  is  a  thick  section  clearly  showing  the  axons  dividing  and  passing 
to  many  cells  of  the  giant  fiber  lobe.  So  I  must  have  "known"  their  anatomy  at 
that  date,  but  was  not  sure  enough  to  publish. 

During  the  early  1930s  I  worked  at  Plymouth,  mostly  with  Sepia,  where  the 
giant  fibers  are  smaller  and  there  is  no  giant  fiber  lobe.  The  first  publication  was 
therefore  a  note  in  the  Journal  of  Physiology  in  1934  claiming  that  the  axoplasm  is 
fluid,  which  we  now  know  to  be  an  error.  Then  in  1936  there  was  a  fuller  account 
suggesting  that  the  epistellar  body  had  been  derived  from  the  giant  fiber  lobe  of  an 
ancestor,  which  is  probably  another  error. 

However  by  now  I  was  fully  convinced  that  they  were  nerve  fibers,  and  in  1936 
was  able  to  prove  this  by  simple  experiments  at  Woods  Hole  (Young,  1938).  Several 
others  then  joined  me:  Bronk,  Gerard,  and  Hartline  all  tried  to  show  action 
potentials  but  the  primitive  oscilloscopes  of  those  days  worked  poorly  and  my 
distinguished  colleagues  could  not  show  reliable  action  potentials  by  electrical 
stimulation.  One  day  Keffer  Hartline  and  I  hooked  a  fiber  to  an  amplifier  and 
speaker  and  put  a  crystal  of  oxalate  on  the  end;  out  came  a  wonderful  buzz — the 
first  giant  fiber  impulses. 

K.  C.  Cole  and  Curtis  were  soon  studying  the  electrical  properties  of  the 
membrane  and  Frank  Schmitt  and  Richard  Bear  showed  me  how  to  study 
biophysical  structure  properly  (Bear  et  al.,  1939).  Material  collected  that  summer  at 
Woods  Hole  provided  the  basis  for  a  full  study  of  the  giant  fiber  system  of  Loligo 
(Young,  1939).  It  was  only  at  this  time  that  I  discovered  that  the  first  order  giant 
cells  in  the  brain  had  been  illustrated  by  Williams  (1909).  His  excellent  monograph 
was  published  in  Holland  and  so  far  as  I  can  discover  the  giant  fiber  system  was 
never  mentioned  throughout  the  succeeding  years.  Williams  followed  the  large  fibers 
into  the  stellate  ganglion  and  stellar  nerves  but  he  seems  to  have  supposed  that  they 
ran  through  the  ganglion  without  synapse.  He  gave  no  figure  of  them. 

The  next  phase  of  work  on  the  giant  fibers  was  mainly  at  Plymouth.  Pumphrey 
and  I  showed  (1938)  that  the  conduction  velocity  follows  the  square  root  of  the 
diameter.  Rapid  conduction  by  giant  fibers  is  an  expensive  luxury  for  a  species.  In 
these  experiments  we  were  helped  by  Alan  Hodgkin,  then  a  student  at  Cambridge; 
this  was  the  first  introduction  of  the  Cambridge  team  to  the  squid  fibers. 

The  axons  provided  the  material  for  the  first  direct  measurements  of  the  internal 
potassium  concentration  of  protoplasm,  made  independently  by  Bear  et  al.  (1939) 
and  Webb  and  Young  (1940). 

The  full  development  of  the  potentialities  of  the  giant  fibers  occurred  after  the 
war  and  I  shall  not  try  to  follow  the  details.  Outstanding  achievements  were  the 
placing  of  an  internal  electrode  and  the  emptying  and  refilling  of  the  axon  by 


156  J.  Z.   YOUNG 

Hodgkin  and  Huxley.  These  investigations  provided  the  data  that  enabled  them  to 
deduce  the  equations  of  the  ionic  exchanges  that  are  involved  (Hodgkin  and  Huxley, 
1952). 

The  special  usefulness  of  the  fibers  is  that  they  allow  monitoring  by  electrodes 
on  both  sides  of  the  membrane.  Numerous  workers  have  used  this  property  for 
studies  of  membrane  transport  at  Woods  Hole,  Plymouth,  and  Naples,  continuing 
to  the  present  day  at  Plymouth  with  the  work  of  Keynes  and  Baker  and  Haydon, 
to  mention  only  three  out  of  many.  Miledi  and  Katz  and  other  groups  have  been 
able  to  study  the  two  sides  of  the  synapse  at  Naples.  The  masses  of  axoplasm  and 
sheets  of  membrane  have  provided  opportunities  for  the  work  of  thousands  of 
physiologists,  biochemists,  and  biophysicists  and  will  continue  to  do  so  in  the  future. 
As  new  problems  and  techniques  appear  these  fibers  will  provide  the  material  of 
choice  for  testing  them.  It  is  curious  to  think  how  different  neuroscience  would 
have  been  had  I  not  made  sections  of  a  yellow  spot — out  of  simple  curiosity.  It  is 
easy  to  say  that  the  fibers  would  have  been  discovered  by  someone  else  soon.  But 
would  they  in  the  present  climate?  Who  would  write  a  grant  application  to  study 
the  possible  structure  of  an  unknown  organ?  It  is  an  example  of  the  need  to  allow 
people  to  pursue  whatever  curious  subject  may  interest  them. 

MEMORY 

The  sections  of  the  brains  of  squids  and  cuttlefishes  that  were  made  to  study 
the  giant  fibers  showed  me  many  other  wonderful  things.  The  supraoesophageal 
lobes  include  a  dozen  distinct  lobules,  each  with  a  different  pattern  of  cells  and 
neuropil.  Surely  these  would  provide  a  good  opportunity  to  study  higher  nervous 
activities,  such  as  memory.  I  felt  that  this  was  an  opening  even  more  important 
than  was  offered  by  the  nerve  fibers.  Already  in  the  1930s  there  was  a  moderately 
good  idea  of  how  nerve  impulses  are  conducted.  Hodgkin  and  Huxley  were  able  to 
carry  this  much  further  and  the  giant  synapse  provided  great  opportunities.  But  the 
really  mysterious  problems  of  neuroscience  were  hidden  there  in  the  neuropils  of 
the  higher  centers.  Biophysics  was  not  ready  to  attack  them,  and  still  cannot  do  so 
even  in  1984. 

However  it  seemed  to  me  that  a  start  should  be  made,  and  Sanders  and  I  were 
able  to  show  at  Plymouth  that  the  learning  power  of  Sepia  is  indeed  dependent  on 
the  vertical  lobe  (1940).  After  a  long  interval  in  the  war,  while  studying  nerve 
regeneration  in  mammals  and  men,  I  returned  to  the  problem  of  memory  at  Naples. 
Octopus  provides  even  better  opportunities  than  Sepia  and  proved  to  be  a  splendid 
learner.  The  supply  at  Naples  seems  to  be  inexhaustable.  The  Posillipo  fishermen 
have  been  able  to  bring  in  20  or  more  octopuses  a  day  in  excellent  condition  and 
the  Stazione  has  generously  provided  space  for  special  tanks  to  be  built  with  funds 
from  the  British  Science  Research  Council.  These  facilities  are  still  available. 

With  the  cooperation  of  Boycott,  Wells,  Sutherland,  and  many  others,  the  two 
memory  systems  of  the  octopus,  visual  and  tactile,  have  been  thoroughly  explored 
(see  Wells,  1966;  Young,  1983).  Lesions  have  shown  that  various  lobes  are  involved 
in  learning,  each  in  a  different  way.  The  visual  and  tactile  memory  systems  each 
includes  four  lobes  with  distinctive  structure  and  function.  Unfortunately  for  some 
reason  it  is  difficult  to  record  the  electrical  activities  of  octopus  neurons.  The 
afferent  fibers  proceeding  from  the  retina  and  statocyst  have  been  thoroughly 
investigated,  but  little  is  known  about  activities  within  the  brain.  There  have  been 
many  investigations  of  the  transmitters  involved  since  the  classical  demonstration 
by  Bacq  (1937)  of  the  huge  amounts  of  acetylcholine  in  the  optic  lobes.  Among 


CEPHALOPODS   AND  NEUROSCIENCE  157 

many  others  Juorio  (1971),  Juorio  and  Barlow  (1976),  and  Tansey  (1979)  have 
shown  the  distribution  of  amines  in  the  brain,  mostly  using  material  obtained  at 
Naples. 

Studies  of  cephalopods  have  not  revealed  all  the  secrets  of  the  mechanism  of 
learning  but  they  have  shown  much,  and  may  show  more.  It  may  be  claimed  that 
we  already  know  from  work  with  large-celled  gastropods,  such  as  that  of  Kandel, 
that  memory  involves  changes  in  synaptic  conduction.  This  is  a  great  advance  but 
does  not  tell  us  how  representations  stored  in  the  brain  enable  an  animal  or  man 
to  recognize  a  rectangle.  There  are  properties  of  aggregates  of  neurons  and  we  still 
require  brains  such  as  those  of  octopuses  that  are  suitable  for  studies  of  them.  It 
will  need  special  methods  that  cannot  yet  be  seen,  and  I  doubt  whether  multiple 
electrodes  will  serve.  Some  methods  must  be  devised  that  can  show  how  numerous 
neurons  interact.  The  various  neuropils  of  an  octopus  may  provide  the  material 
that  is  needed,  just  as  the  giant  fibers  of  the  squid  will  allow  testing  of  new  methods 
for  the  study  of  membranes. 

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NICKED  BY  OCCAM'S   RAZOR:   UNITARIANISM   IN  THE 
INVESTIGATION  OF  SYNAPTIC  TRANSMISSION 

M.   V.   L.   BENNETT 

Division  of  Cellular  Neurobiology,  Department  of  Neuroscience,  Albert  Einstein  College  of  Medicine, 
Bronx,  New  York  10461,  and  Marine  Biological  Laboratory,  Woods  Hole,  Massachusetts  02543 

INTRODUCTION 

Before  the  modes  of  synaptic  transmission  were  well  enough  understood  to 
decide  the  matter,  chemical  and  electrical  mediation  were  widely  accepted  as  the 
two  possibilities.  Neurophysiologists  in  general  thought  that  transmission  was 
electrical,  because  axonal  conduction  was  electrical  and  they  saw  no  need  for  an 
additional  mechanism.  Furthermore  chemical  transmission  seemed  too  slow.  On 
the  other  side,  pharmacologists  favored  chemical  transmission,  because  they  worked 
with  drugs  and  had  a  great  deal  of  evidence  for  chemical  sensitivity  (and  some  for 
chemical  transmission)  at  peripheral  sites  including  viscera  and  skeletal  muscle. 

The  motif  of  this  article  comes  from  a  story  Harry  Grundfest  used  to  tell.  In  his 
1947  paper  in  Annual  Review  of  Physiology  he  inclined  toward  the  view  that 
interneuronal  transmission  is  electrically  mediated,  as  is  axonal  conduction,  and  in 
his  letter  of  submission  with  the  manuscript,  he  said  that  in  spite  of  evidence  for 
chemical  transmission  he  remained  an  enlightened  Unitarian.  By  the  time  of  his 
1957  review  he,  and  others,  were  convinced  that  synaptic  transmission  was  chemically 
mediated.  He  recalled  the  comment  written  on  the  end  of  the  manuscript  by  an 
editor  who  remembered  the  earlier  letter:  "Enlightened  unitarianism  grows  dim." 

Although  chemical  transmission  was  established  and  unity  of  synaptic  transmission 
and  axonal  conduction  defeated,  unitarianism  lived  and  still  does.  Occam's  razor, 
the  mode  of  inference  in  which  the  simplest  explanation  is  favored,  remains  in  use. 
I  shall  give  several  illustrations  from  the  controversy  over  synaptic  transmission  of 
Occam's  razor  cutting  both  ways,  instances  where  the  simplest  explanation  was 
incorrect  or  incomplete,  and  the  investigator  was  misled  by  his  preference  for  a 
Unitarian  view.  Biological  reality  does  have  many  unifying  principles,  but  life  is 
complicated  and  also  historical.  Nervous  systems  in  their  evolution  have  come  to 
use  many  different  mechanisms. 

Both  electrical  and  chemical  transmission  were  proposed  for  the  neuromuscular 
junction  in  the  1800s  (cf..  Brazier,  1959).  In  1904  Elliot  observed  the  similarity 
between  the  actions  of  sympathetic  nerves  and  adrenalin.  Recognizing  the  homology 
between  adrenal  glands  and  sympathetic  ganglia,  he  suggested  that  the  nerves  acted 
by  releasing  adrenalin.  This  proposal  was  made  in  a  proceedings  note  in  the  Journal 
of  Physiology,  but  for  reasons  unknown  to  me  he  failed  to  mention  the  idea  again 
in  the  detailed  publication  of  the  data  appearing  the  following  year  (Elliot,  1905). 

Loewi  is  commonly  given  credit  for  establishing  chemical  transmission  with  his 
1921  experiment  on  the  Vagusstoff.  Indeed,  of  the  two  essential  criteria  for 
identifying  a  transmitter,  evoked  release  of  the  putative  transmitter  and  identity  of 
action  of  the  transmitter  and  the  neurally  released  material,  Loewi's  experiment  was 
the  first  to  demonstrate  release.  In  his  1933  Harvey  Lecture  Loewi  wrote  "In  the 
year  1921  I  was  able  to  prove  without  doubt  the  correctness  of  the  fundamental 
idea"  that  sympathetic  and  parasympathetic  nerves  liberate  a  chemical  substance 
that  produces  the  postsynaptic  action.  But  he  did  not  overgeneralize.  He  raised  the 

159 


160  M.   V.   L.   BENNETT 

possibility  of  chemical  transmission  in  the  central  nervous  system,  seeing  that  it 
would  provide  a  nice  mechanism  for  temporal  summation,  but  noted  (p.  228)  "the 
experimental  proof  may  be  rather  difficult  to  obtain/"  He  continues  (p.  232) 

A  priori  one  can  imagine  that  in  cases  where  the  effect  of  stimulation  takes 
place  in  an  almost  immeasurable  space  of  time  and  ceases  practically  instantly 
after  the  stimulus  is  over,  as  for  instance  in  the  case  of  nervous  stimulation  of 
striated  muscle,  there  is  hardly  enough  time  for  a  substance  to  be  formed  and 
nearly  simultaneously  rendered  ineffective.  But  the  time  factor  is  not  a  decisive 
proof  for  or  against  a  humoral  mechanism  in  the  case  of  spinal  nerves.  Personally 
I  do  not  believe  in  a  humoral  mechanism  existing  in  the  case  of  striated  muscle. 

Although  Loewi's  work  was  important  in  establishing  the  possibility  of  chemical 
transmission,  comparable  experiments  were  done  for  electrical  transmission  much 
earlier  by  Matteuci,  who  showed  that  the  electricity  produced  by  a  contracting 
muscle  would  stimulate  a  nerve  (see  Brazier,  1959).  A  nerve  generates  electricity, 
but  one  can  question  whether  it  is  in  the  right  amount,  at  the  right  time  and  in  the 
right  place  to  account  for  synaptic  transmission.  The  same  question  applies  to 
Loewi's  experiments.  Occam's  razor  suggested  that  the  neurally  released  transmitter 
with  the  same  action  as  caused  by  nerve  stimulation  was  indeed  released  appropriately 
in  terms  of  amount,  time,  and  place,  but  a  definitive  demonstration  of  all  these 
properties  has  still  not  been  made,  at  least  at  any  synapse  where  transmission  is 
rapid.  Other  electrophysiological  criteria  turned  out  to  provide  the  evidence  that 
ultimately  convinced  the  doubting  electrophysiologists  (such  as  Grundfest).  And  in 
my  view  it  was  Occam's  razor,  correctly  used,  that  convinced  the  pharmacologists 
before  all  the  data  were  in.* 

J.  C.  ECCLES  AND  ENLIGHTENMENT 

Eccles  figured  prominently  in  what  was  often  termed  the  sparks  and  soup 
controversy,  and  his  history  of  the  subject  is  insightful  as  well  as  first  hand  (Eccles, 
1982).  He  was  convinced  of  chemical  transmission  in  respect  to  sympathetic 
innervation  prior  to  the  second  world  war.  By  1948  he  also  found  Kuffler's 
experiments  in  1942  and  those  of  himself  and  collaborators  to  be  a  valid  demon- 
stration of  chemical  transmission  at  the  (frog)  neuromuscular  junction.  The  prolon- 
gation of  the  PSP  by  eserine  without  change  in  the  brief  nerve  response  was  indeed 
highly  suggestive.  But  in  the  absence  of  really  knowing  the  anatomy  and  the 
transmembrane  potentials,  one  can  still  regard  the  case  as  somewhat  open.  Indeed 
in  a  1948  review  of  most  of  these  experiments  Kuffler  writes  a  rather  ambiguously 
worded  passage  indicating  uncertainty,  although  it  could  be  construed  as  implying 
chemical.  He  really  did  mean  uncertainty  for  his  note  added  in  proof  cites  an  Eccles 
review  (1948)  stating  that  chemical  transmission  was  definitely  established.  It  is 
interesting  that  Kuffler's  work  is  frequently  referred  to  in  that  Eccles  review. 
Although  the  section  discussing  neuromuscular  transmission  is  unclear  as  to  mode, 
a  direct  statement  is  found  on  p.  112  "the  electrical  theory  of  transmission  provides 
a  satisfactory  answer  for  the  spinal  cord,  which  contrasts  with  the  neuromuscular 
junction  where  transmission  appears  to  be  exclusively  cholinergic."  Also  on  p.  107 
"In  every  respect  the  observations  of  Bullock  on  the  squid  giant  synapse  conform 

*  Ernst  Florey  in  the  discussion  of  the  oral  presentation  of  this  paper  told  of  a  visit  to  Loewi  in  New 
York  after  World  War  II.  Loewi  reported  to  Florey  that  he  had  recently  had  an  unpleasant  experience 
while  attending  a  Rubinstein  concert.  He  was  so  distracted  by  the  evident  impossibility  of  such  rapid  and 
complex  motor  acts  being  mediated  chemically  that  he  could  not  enjoy  the  music. 


OCCAM'S  RAZOR  161 

with  the  predictions  of  the  electrical  theory,"  observations  that  electrophysiologically 
were  very  similar  to  those  at  the  neuromuscular  junction.  Chemical  transmission  in 
muscle  was  accepted,  but  electrical  transmission  between  neurons  was  being 
conserved. 

In  respect  to  Eccles  and  Kuffler  on  the  neuromuscular  junction,  I  recall  Paul 
Cranefield  referring  to  a  paper  where  the  coauthors  disagreed  strongly  about  the 
significance  of  their  findings.  In  writing  the  discussion  they  finally  compromised  on 
the  wording  "one  of  the  authors  is  forced  to  conclude  that  .  .  ."  The  investigation 
of  synaptic  transmission  produced  many  inferences  about  whose  compelling  nature 
there  were  strong  differences  of  opinion.  Intracellular  recording  clarified  the  situation 
greatly. 

Eccles  and  co-workers  in  their  landmark  paper  (Brock  et  ai,  1952)  (justifiably) 
stated  that  their  data  established  chemical  transmission  for  neuronal  inhibition.  The 
demonstration  did  not  involve  identification  of  the  transmitter  or  measurement  of 
its  action  and  release,  but  rather  intracellular  recording  of  activity  that  could  not 
(reasonably)  be  mediated  electrically.  They  were  suitably  careful  about  excitatory 
transmission  and  only  said  that  it  was  probably  chemically  mediated.  Fatt  and  Katz 
(1951)  had  already  applied  intercellular  recording  to  the  neuromuscular  junction, 
and  characterized  chemically  mediated  excitation  to  an  extent  that  had  not  been 
possible  previously.  They  were  also  studying  transmission  in  a  crustacean  muscle 
where  inhibition  resulted  from  a  conductance  increase  that  could  not  be  electrically 
transmitted  from  the  presynaptic  fiber  (Fatt  and  Katz,  1953).  Of  course  this  was  at 
a  crustacean  neuromuscular  junction  not  known  to  be  a  model  for  mammalian 
CNS.  Fatt  subsequently  joined  Eccles  for  a  collaboration  in  which  the  electrophys- 
iological  insights  of  Fatt  and  Katz  were  applied  to  both  excitatory  and  inhibitory 
synapses  in  the  cat  spinal  cord.  Eccles,  in  '57,  wrote:  "It  can  now  be  taken  as 
established  that  transmission  across  synapses  occurs  not  by  the  spread  of  electrical 
currents,  but  by  the  specific  chemical  substances  which  impulses  cause  to  be 
liberated  from  the  presynaptic  membranes1"  (p.  67). 

Eccles  conversion  was  an  impressive  mental  feat  in  that  he  was  able  to  let  go 
rapidly  of  an  idea  he  had  strongly  fought  for.  To  be  sure,  the  influence  of  Karl 
Popper  had  led  him  to  push  the  idea  of  electrical  transmission  perhaps  harder  than 
justified  until  it  was  finally  falsified  (for  cat  motoneurons).  The  pendulum  swung  in 
terms  of  the  body  scientific  as  well,  and  my  impression  is  that  at  this  time  almost 
everyone  believed  that  synaptic  transmission  was  chemically  mediated.  However, 
Furshpan  and  Potter  (1959)  were  studying  the  crayfish  giant  motor  synapse.  Wiersma 
had  previously  shown  that  it  transmitted  impulses  in  only  one  direction  and  from 
sensitivity  to  block  by  pharmacological  agents  he  inferred  transmission  to  be 
chemically  mediated.  It  was  an  attractive  synapse  because  both  pre-  and  postsynaptic 
axons  were  large  enough  that  it  appeared  (and  was)  possible  to  penetrate  both 
elements  close  to  the  synapse.  Furshpan  and  Potter  found  that  here  transmission 
was,  after  all,  electrical,  and  unidirectional  transmission  of  impulses  was  due  to 
rectification  by  the  membranes  in  the  synaptic  region  which  were  known  to  be 
closely  apposed.  Eccles  easily  accepted  these  new  findings  and  in  his  1961  review 
had  a  section  on  electrical  transmission.  He  also  proposed  that  the  junctions  between 
segments  of  septate  axons  found  widely  in  annelids  and  arthropods  "should  not  be 
considered  synapses"'  because  they  conduct  in  both  directions,  but  if  they  were 
considered  synapses  "they  would  be  examples  of  "the  simplest  type""  of  electrical 
synapse"  (p.  363). 

Only  later  did  he  come  to  accept  the  septal  junctions  as  synapses  which  are,  by 
his  definition,  close  appositions  "specialized  for  the  transmission  of  excitation  or 


162  M.   V.   L.   BENNETT 

inhibition"  (Eccles,  1964  p.  VI).  [For  the  record,  Watanabe  (1958)  independently 
discovered  electrical  coupling  between  cardiac  ganglion  cells  in  an  arthropod, 
although  in  the  absence  of  morphological  data  (still  absent  in  many  instances  in 
arthropods  including  this  one)  he  thought  the  cells  were  syncytial.]  Perhaps  because 
electric  transmission  was  initially  found  in  invertebrates  Eccles  (1961,  p.  366)  was 
led  to  write 

It  seems  probable  that  many  examples  of  electrical  [sic]  transmitting  synapse 
may  be  discovered  when  invertebrate  nervous  systems  are  intensively  investigated. 
With  the  vertebrate  nervous  system  the  invariable  presence  of  a  considerable 
synaptic  delay  would  exclude  electrical  transmission  as  significantly  contributing 
to  any  of  the  synaptic  transmissions  that  have  been  investigated  by  intracellular 
recording. 

For  the  record,  there  are  many  instances  in  mammals  and  lower  forms  where 
synaptic  delays  at  chemical  and  electrical  synapses  are  difficult  to  distinguish  (cf., 
Bennett,  1977). 

At  a  meeting  on  the  thalamus  in  1965,  I  presented  a  paper  on  how  electrical 
synapses  synchronized  neural  firing  in  fishes  and  suggested  these  systems  as  a 
possible  model  of  mammalian  CNS  synchronization  (Bennett,  1966).  Afterwards 
Eccles  was  heard  to  say  in  effect  that  it  was  all  very  well  for  fishes  but  Bennett 
would  never  find  electrical  transmission  in  mammals  (D.  P.  Purpura,  pers.  comm.). 
I  admit  that  my  presence  at  the  meeting  was  more  a  result  of  my  being  in  the 
institution  of  the  organizers  than  of  their  conviction  that  the  fishes'  physiology  was 
pertinent  to  the  mammalian  thalamus. 

Others  did  go  on  to  find  electrical  transmission  in  the  mammal  (reviewed  in 
Bennett,  1977,  and  Korn  and  Faber,  1979,  and  new  examples  continue  to  be 
discovered).  Eccles  still  writes  somewhat  perjoratively  in  1982  that  electrical  synapses 
"are  relatively  rare  in  the  mammalian  brain  and  it  has  yet  to  be  shown  that  they 
are  functionally  important  in  the  brain1'  (p.  337).  For  the  mammalian  brain  he 
prefers  there  to  be  only  a  single  important  mode  of  transmission.  I  have  often 
argued  that  there  are  many  synapses  where  it  is  difficult  to  see  a  relative  advantage 
of  excitatory  chemical  over  electrical  transmission  (e.g.,  Bennett,  1977),  but  in  this 
area  of  course  I  am  not  disinterested  and  my  arguments  can  be  seen  as  serving  to 
increase  the  importance  of  my  own  research.  Nevertheless,  in  the  absence  of 
experimental  data  it  is  difficult  to  conclude  that  a  distinct  group  of  synapses  are 
unimportant  in  the  operation  of  the  central  nervous  system. 

H.  GRUNDFEST  AND  NEO-UNITARIANISM 

Let  us  now  return  to  the  195CTs  and  Harry  Grundfest's  1957  review.  Grundfest 
proposed  that  subsynaptic  membrane  was  electrically  inexcitable  and  that  this 
property  was  essential  in  its  operation,  and  thus  all  synaptic  transmission  was 
chemical.  (I  came  to  work  in  Harry's  lab  at  this  time,  shortly  after  finishing  graduate 
school.)  The  originality  of  the  proposal  of  electrical  inexcitability  is  arguable  and 
Fatt  and  Katz  had  previously  stated  that  ACh  action  at  the  neuromuscular  junction 
is  unlike  that  of  an  electric  field  which  causes  generation  of  an  action  potential. 
Eccles  (1957,  p.  52)  wrote  independently  that  by  "analogy  with  the  subsynaptic 
membrane  of  the  neuromuscular  junction  [and  other  examples]  .  .  .  it  is  possible 
that  the  subsynaptic  areas  [of  motoneurons]  are  incapable  of  responding  by 
impulses."  Grundfest  did  bring  a  large  amount  of  data  together  and  in  my  view  his 
major  contribution  in  this  regard  was  in  emphasizing  the  importance  of  a  general 
physiology  that  included  all  synapses  and  not  just  the  favored  few. 


OCCAM'S   RAZOR  163 

Given  that  Grundfest  had  adopted  electrical  inexcitability  and  chemical  trans- 
mission as  universals,  a  grand  generalization,  he  was  presented  with  a  problem 
when  electrical  transmission  was  definitively  established.  He  (with  Kao)  wrote  in 
1957  "The  term  synapse  assumes  more  meaning  than  merely  the  physical  apposition 
of  two  adjacent  excitable  cells.  Synaptic  transmissional  excitation  must  therefore  be 
initiated  by  a  process  of  specialized,  secretory  activity  at  the  presynaptic  terminals"" 
(p.  569). 

It  was  arguable  that  a  necessary  property  of  a  synapse  is  unidirectional  action 
(as  Eccles  did  briefly)  and  Kao  and  Grundfest  saw  the  septa  not  as  synapses,  but  as 
"ephapses  without  synaptic  function,  demarcating  ontogenetic  cellular  boundaries.'" 
The  term  ephapse  was  coined  by  Arvanitaki  (1942)  to  denote  artificial  junctions 
made  by  placing  two  axons  together;  she  proposed  these  junctions  as  a  model  for 
(real)  synapses.  The  rectifying  electrotonic  synapse  described  by  Furshpan  and  Potter 
was  not  so  easily  excluded  because  it  exhibited  polarized  or  unidirectional  conduction. 
Kao  and  Grundfest  wrote  "It  remains  to  be  seen  whether  this  'electrically  excitable 
synapse"  is  a  condition  found  at  other  junctions  or  whether  it  represents  an 
abandoned  evolutionary  variant"  (p.  570).  (One  sees  more  vertebrate  chauvinism 
here.  Arthropods  represent  the  peak  of  another  branch  of  evolution.) 

Grundfest"s  later  solution  to  the  problem  of  electrical  synapses  was  to  change 
the  name.  In  his  1959  chapter,  all  electrical  synapses  have  become  ephapses. 
Grundfest  recognized  that  the  crayfish  "junction  meets  the  criteria  of  anatomical 
discontinuity  and  transmissional  polarization'"  (p.  192),  (now  we  know  there  is 
cytoplasmic  continuity  for  small  molecules)  but  discusses  the  junction's  profound 
differences  from  chemically  transmitting  synapses  with  electrically  inexcitable  sub- 
synaptic  membrane. 

He  also  recognized  that  special  geometric  properties  alone  can  lead  to  polarized 
transmission  (at  electrical  synapses  or  ephapses,  p.  190),  but  now  to  him  "the  crucial 
distinction  is  whether  current  flow  in  a  presynaptic  terminal  can  excite"  the 
postsynaptic  element.  Since  at  chemical  synapses  the  presynaptic  current  is  "far  too 
small  to  excite  the  postsynaptic  cell,"  the  electrical  synapses  must  be  something 
else;  he  chose  ephapses.  There  is  a  clearly  illogical  step  here.  All  synapses  are 
unidirectional;  chemical  synapses  are  unidirectional  and  not  electrical.  Thus  all 
synapses  are  chemical  and  not  electrical  (this  is  the  mistake),  and  if  one  finds  a 
junction  that  is  unidirectional  and  electrical,  it  can't  be  a  synapse.  I  do  not  believe 
that  this  was  the  way  in  which  Grundfest  arrived  at  the  description  of  electrically 
transmitting  junctions  between  neurons  as  ephapses.  Rather  the  constellation  of 
properties  which  he  viewed  as  integral  to  synaptic  transmission  appeared  to  derive 
from  chemical  mediation.  If  some  synapses  were  electrical,  the  thesis  would  have 
required  major  revision,  and  it  was  simpler  to  call  electrical  synapses  by  a  dif- 
ferent name. 

I  leave  the  question  of  electrical  inexcitability  with  a  few  further  comments. 
Many  of  the  properties  of  chemical  transmission  that  Grundfest  cites  can  arise  in 
other  ways,  and  electrical  synapses  can  exhibit  virtually  all  of  them  (except  PSP 
reversal).  I  never  found  it  compelling  that  subsynaptic  membrane  should  be 
inexcitable,  because  it  always  seemed  possible  that  a  transmitter  could  act  on  an 
electrically  excitable  channel.  K  ions  released  by  activity  depolarize  adjacent  cells 
by  changing  the  driving  force  for  both  excitable  and  inexcitable  channels,  and  there 
are  many  examples  of  K  release  by  neurons  affecting  other  neurons.  The  K  effects 
might  be  considered  ephaptic  rather  than  synaptic  (although  chemically  mediated), 
but  the  action  can  still  be  excitatory  on  electrically  excitable  membrane.  Recently 
many  modulatory  synapses  have  been  described  where  transmitters  act  to  alter  an 


164  M.   V.   L.   BENNETT 

electrically  excitable  conductance  (Tsien  and  Siegelbaum,  1983).  To  be  sure  most 
of  these  actions  are  probably  mediated  by  a  second  messenger  that  is  intracellular, 
and  the  subsynaptic  membrane  where  the  transmitter  acts  is  not  directly  involved 
in  impulse  generation. 

In  current  terms  we  know  many  channel  macromolecules  that  are  affected  by 
transmitters,  other  chemical  agents  and  electric  fields  (cf.  Bennett  et  a/.,  1984).  It  is 
reasonable  that  for  an  intramembrane  protein  undergoing  a  conformational  change 
there  would  be  a  dipole  moment  change  that  would  confer  some  degree  of 
electrical  sensitivity.  This  sensitivity  might  be  greater  as  in  the  usual  channels 
involved  in  impulse  generation  or  lesser  as  in  most  transmitter  evoked  changes  at 
subsynaptic  membranes  of  chemical  synapses.  But  the  general  prediction  is  that  any 
chemically  evoked  change  will  also  exhibit  some  degree  of  electrical  sensitivity. 
Furthermore,  channels  primarily  sensitive  to  potential  may  also  be  sensitive  to 
regulatory  molecules,  pharmacological  agents,  and  toxins.  I  do  subscribe  to  Grundfest's 
provocative  view  that  electrical  inexcitability  as  determined  electrophysiologically 
provided  the  only  "direct"  evidence  for  chemical  transmission.  Many  synapses  are 
accepted  as  chemical  on  electrophysiological  grounds  with  no  knowledge  of  the 
transmitter  (let  alone  evoked  release  and  identity  of  action,  cf.,  Brock  et  a/.,  1952). 
However,  one  man's  direct  evidence  is  another  man's  tortured  reasoning,  and 
chemical  transmission  would  not  be  nearly  so  convincingly  demonstrated  by 
electrical  inexcitability  if  there  were  not  all  those  data  on  transmitter  action  and 
transmitter  release. 

My  exposition  of  where  Grundfest  went  slightly  astray  should  be  evaluated  in 
light  of  my  own  agenda.  While  my  recollection  is  that  I  did  not  accept  Grundfest's 
constellation  of  properties  at  the  time,  the  real  difficulty  came  later  when  my 
collaborators  and  I  began  to  find  electrical  synapses  in  modest  profusion.  It  was  not 
in  my  self  interest  to  be  working  on  a  lower  class  of  interneuronal  junction,  with  a 
connotation  of  artificiality,  regardless  of  whether  they  could  exhibit  most  of 
Grundfest's  properties.  There  was  no  doubt  another  factor  that  I  have  heard  called 
Feldberg's  dictum,  that  is  that  a  scientist  would  rather  use  another  scientist's 
toothbrush  than  his  [or  her]  terminology. 

There  were  several  important  precursors  to  electrical  transmission  as  demonstrated 
by  Furshpan  and  Potter.  As  noted  above,  Kao  and  Grundfest  (1959)  saw  the  septa 
as  providing  little  hindrance  to  local  circuit,  electrical  propagation  although  the 
septa  were  described  as  not  synaptic.  Bullock  in  1945  wrote  (p.  70): 

The  high  speed  of  conduction  and  its  unpolarized  character  are  significant  in 
view  of  the  apparently  synaptic  nature  of  the  system  as  demonstrated  histologically. 
These  properties  are  compatible  with  the  [supposition]  that  the  synapse  is  not 
inherently  polarized  nor  delaying  but  is  only  so  as  a  result  of  the  particular 
anatomical  relations  prevalent  in  vertebrates  and  that  these  properties  should  not 
be  a  part  of  the  definition  of  the  synapse. 

The  suggestion  that  septal  synapses  are  like  all  other  synapses  is  again  overinclusive, 
but  the  description  accurately  applies  to  many  electrical  synapses  in  vertebrates  as 
well  as  invertebrates. 

Not  everyone  took  a  Unitarian  view  during  these  developments.  Fatt  wrote 
in  1954: 

it  is  probable  that  electrical  transmission  occurs  at  certain  other  junctions.  One 
possible  arrangement,  which  may  be  envisioned  to  give  a  high  degree  of  electrical 
interaction  is  for  two  fibers  [of  about  the  same  dimensions]  to  be  actually 
touching  and  for  the  membrane  in  contact  to  have  a  low  electric  resistance 


OCCAM'S   RAZOR  165 

compared  with  that  in  neighboring  parts  of  the  fiber.  The  synapse  would  then 
serve  to  direct  current  between  the  interior  of  the  two  fibers,  while  active 
membrane  changes  would  occur  in  the  neighboring  regions  [p.  204]. 

Here  is  an  accurate  description  of  transmission  at  many  electrical  synapses;  many 
more  if  one  ignores  the  qualification  as  to  similarity  of  size  of  the  fibers.  As  an 
aside  there  is  an  implication  of  absence  of  excitability  in  the  connecting  membrane. 
He  went  on: 

A  case  in  which  there  can  be  little  doubt  that  electrical  transmission  operates  is 
in  the  nervous  system  of  the  crayfish,  where  successive  giant  nerve  cells,  each 
extending  along  one  segment,  .  .  .  form  the  lateral  giant  nerve  fibers.  Transmission 
takes  place  in  either  direction  and  such  chains  of  nerve  cells  [form]  synapses 
where  transmission  occurs  electrically  [p.  705]. 

Thus,  the  stage  was  thoroughly  set  for  electrical  transmission.  I  find  it  admirable 
that  in  the  laboratory  that  many  consider  the  primary  source  of  chemical  transmission, 
there  was  recognition  that  in  some  cases  transmission  was  very  probably  electrically 
mediated.  Fatt  thought,  for  reasonable  cause,  that  transmission  at  the  motor  giant 
synapse  of  crayfish  was  chemical,  but  the  basic  understanding  that  led  to  Furshpan's 
and  Potter's  important  findings  were  there. 

NACHMANSOHN  AND  SINGLE-MINDED  UNITARIANISM 

David  Nachmansohn  will  be  used  to  provide  a  brief  coda  to  the  longer 
discussions  above.  He  held  with  an  amazing  perseverence  to  a  unified  theory  of 
chemical  mediation  of  action  potentials,  all  action  potentials  everywhere,  but 
believed  that  synaptic  transmission  was  electrical.  His  thinking  came  to  be  a 
somewhat  distorted  mirror  image  of  the  majority  view. 

The  cholinergic  system  was  an  enduring  concern  of  Nachmansohn,  and  he  made 
many  truly  major  contributions  in  respect  to  the  properties  of  acetylcholine  esterase 
and  the  discovery  of  choline  acetylase  (or  choline  acetyltransferase)  and  coenzyme 
A.  Early  on  he  and  his  collaborators,  in  studying  the  electric  eel,  found  that  the 
concentration  of  AChE  per  unit  length  of  electric  organ  was  quite  linearly  related 
to  the  voltage  developed  per  unit  length.  The  current  hypothesis  to  explain  this 
finding  would  be  that  voltage  is  proportional  to  the  number  of  cells  in  series,  and 
the  number  of  cells  per  unit  length  is  greater  in  anterior  regions.  The  AChE  is 
largely  found  on  the  innervated  face  of  cells,  so  the  amount  of  AChE  is  proportional 
to  the  number  of  cells,  hence  voltage  and  AChE  are  related.  At  the  time  of 
Nachmansohn's  finding  the  series  summation  in  electric  organs  of  essentially 
ordinary  membrane  potentials  had  not  been  established  and  the  inference  of  ACh's 
direct  involvement  in  potential  generation  was  a  tenable  hypothesis.  The  Nachman- 
sohn group  proceeded  to  find  a  great  deal  more  data  of  a  pharmacological  kind  that 
supported  the  involvement  of  the  cholinergeric  system.  Also,  negative  evidence  such 
as  the  failure  of  polar  cholinergic  agents  to  act  on  ordinary  axons  was  accounted 
for  by  postulation  of  permeability  barriers  that  protected  the  actual  sites  of  impulse 
generation.  During  these  developments  there  were  questions  raised  by  others  about 
specificity  or  reproducibility  of  some  of  the  results  in  agreement  with  the  theory. 
Moreover  as  data  accumulated  it  became  necessary  to  postulate  permeability  barriers 
on  the  inside  as  well  as  outside  of  the  active  membrane.  The  theory,  although 
increasingly  ornate,  was  now  extremely  difficult  to  disprove,  certainly  by  Nachman- 
sohn's  standards.  In  the  early  seventies  he  elaborated  a  highly  specific  version  of 
how  ACh  was  involved  in  the  permeability  changes  underlying  action  potential 
generation.  (He  did  not  deny  the  permeability  changes  of  ion  fluxes.) 


166  M.   V.   L.   BENNETT 

His  view  of  synaptic  transmission  was  surprising  given  that  he  required  a 
chemical  step  in  axonal  conduction.  In  emphasizing  the  biochemical  unity  of  life, 
"Nature  has  shown  little  imagination  in  modifying  chemical  mechanisms  associated 
with  given  functions"  (Nachmansohn  and  Neumann,  1975),  he  wanted  transmission 
to  be  electrical.  In  1961  he  wrote 

The  action  of  acetylcholine  with  a  specific  receptor  protein  is  essential  for  the 
conductance  changes  observed  [in  the  axon].  The  agent  that  propagates  impulses 
in  the  axon  and  across  the  synapse  is  the  electric  current  (ion  movements)  but 
the  ion  movements  require  the  trigger  action  of  acetylcholine  [p.  241  of 
Nachmansohn  and  Neumann,  1975]. 

and 

Electric  fields  and  manifestations  must  be  greatly  influenced  by  the  complexity 
of  this  organization  [of  the  synapse  revealed  by  electron  microscopy].  But  there 
is  not  a  single  fact  to  support  the  view  that  the  role  of  acetylcholine  system 
present  in  both  pre-  and  postsynaptic  membranes  differs  fundamentally  in  axonal 
conduction  and  synaptic  transmission.  All  data  available  are  consistent  with  the 
unified  concept  [p.  256]. 

Even  in  1975  his  views  about  the  synapses  were  essentially  unchanged.  "Special 
structural  arrangements  may  make  small  eddy  currents  at  junctions  quite  efficient 
in  initiating  the  chemical  reactions  responsible  for  the  changes  in  the  postsynaptic 
membrane  for  either  hyper-  or  depolarization"  (p.  197). 

I  have  never  understood  why  Nachmansohn  was  so  resistant  to  the  idea  of 
chemical  transmission,  which  would  have  been  permitted  by  very  minor  modifications 
of  his  view  of  the  role  of  acetylcholine.  Perhaps  some  of  the  evidence  for  its  role  in 
axonal  conduction  would  have  been  undermined  if  interpreted  as  associated  with 
chemical  transmission.  Alternately  his  tenacity  in  support  of  electrical  transmission 
may  have  been  simply  another  facet  of  the  personality  trait  that  led  him  to  hold  an 
unpopular  and  finally  indefensible  theory  of  axonal  conduction. 

Many  other  scientists  have  held  on  to  their  theories  for  longer  than  they  were 
tenable.  In  spite  of  Occam's  razor  a  pet  theory  can  often  be  nurtured  by  subsidiary 
hypotheses  far  beyond  the  point  where  the  proposer  would  have  been  likely  to 
formulate  the  theory  ab  initio.  Simplicity  is  a  useful  guide  in  formulating  theories, 
but  once  a  theory  becomes  a  member  of  one's  inner  family,  the  drive  towards 
simplicity  has  a  way  of  losing  its  strength. 

ACKNOWLEDGMENT 
Supported  in  part  by  NIH  grants  NS-07512  and  HD-04248. 

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MARINE   BOTANY  AND  ECOLOGY   AT  STAZIONE  ZOOLOGICA 

CARMELO   R.  TOMAS 

Marine  Botany  Laboratory,  Stazione  Zoologica,  80121  Naples,  Italy 

ABSTRACT 

For  108  years  marine  botany  research  has  been  an  important  component  of  the 
research  conducted  at  the  Stazione  Zoologica  of  Naples.  The  first  researchers  at  the 
Naples  Institute  were  German  guests  who  proceeded  with  descriptive  ecological  and 
taxonomic  studies  and  provided  a  foundation  for  the  later  physiological,  cytological, 
life  cycle,  and  biochemical  studies  conducted  at  the  Stazione.  During  a  major  period 
(47  yrs.)  Prof.  G.  Funk  contributed  ecological  work  giving  the  botanical  research  at 
Naples  a  continuity  which  extended  into  the  late  5(Ts.  From  1960  onwards,  the 
marine  botany  laboratory  assumed  a  different  direction  and  recently  has  returned 
to  a  physiological-ecological  orientation.  The  lasting  impact  of  marine  botany  has 
been  the  contribution  of  an  ecological  dimension  from  which  other  studies  grew 
and  found  support. 

DISCUSSION 

Studies  of  marine  botany  in  the  Gulf  of  Naples  predate  the  founding  of  the 
Stazione  Zoologica  by  nearly  fifty  years.  The  studies  of  Delle  Chiaje  (1823)  and 
Costa  (1838)  describe  species  of  the  rich  flora  found  in  the  waters  of  the  kingdom 
of  Naples.  This  flora  and  the  equally  abundant  and  varied  fauna  were  factors 
influencing  Anton  Dohrn  in  establishing  the  Naples  Institute.  Within  four  years  of 
the  opening  of  the  Stazione  Zoologica,  marine  botany  research  was  begun  in  earnest. 
For  the  past  108  years,  research  in  this  discipline  has  continued  to  contribute  to  the 
overall  scientific  effort.  Marine  botany  and  ecology  are  natural  synonyms  for  the 
Stazione  Zoologica,  since  from  the  beginning  the  ecological  approach  to  the  study 
of  marine  algae  predominated,  resulting  in  quantification  and  evaluation  of  the 
evolution  of  Neapolitan  coastal  waters.  In  addition,  algal  studies  relating  to  cytology, 
physiology,  anatomy,  and  aspects  of  biochemistry  were  pursued.  Both  micro  and 
macro  algae  were  studied  although  the  major  emphasis  was  placed  on  macroscopic 
thallate  forms.  The  activity  of  marine  botany  research  was  greatly  influenced  by  the 
perturbations  imposed  by  the  two  world  wars  as  well  as  the  natural  rhythms  of  the 
Institution's  growth. 

The  earliest  visitors  (1873-1900)  were  almost  exclusively  German  researchers 
encouraged  by  Anton  Dohrn  to  visit  and  work  at  his  station.  Among  these,  J. 
Reinke,  P.  Falkenberg,  G.  Berthold,  R.  Valiante,  and  C.  Sauvageau  were  the  first 
to  extensively  study  the  benthic  algae  of  the  Gulf.  Armed  with  the  modern  elements 
of  taxonomy  and  physiology,  these  early  workers  (Reinke,  1878a,  b;  Falkenberg, 
1879,  1901;  Berthold,  1882a,  b;  Valiante,  1883;  Sauvageau,  1892)  established  vital 
species  lists  as  well  as  distribution  in  the  Naples  area.  In  addition,  their  observations 
on  gametes  of  brown  algae,  cellular  composition  including  ions,  chromoplasts, 
vacuoles,  and  associated  membranes  further  added  to  the  general  knowledge  of 
algae.  The  first  decades  of  the  20th  century  had  macroalgologists  including  A. 
Vickers,  F.  Tobler,  E.  Leick,  and  G.  Funk  as  further  contributors  to  our  understanding 
of  algal  species  distribution  as  related  to  ecological  factors.  Among  these,  the  most 

168 


MARINE   BOTANY   AND   ECOLOGY  169 

prominent  and  one  who  had  the  greatest  impact  on  marine  botany  in  Naples  was 
G.  Funk.  Professor  Funk's  monographs  (1927,  1955)  with  extensive  descriptions  of 
algal  associations,  reproductive  cycles  in  nature,  and  distribution  remained  a 
benchmark  for  algal  research  in  the  Mediterranean.  Funk's  observations  also  served 
as  a  basis  for  quantifying  changes  in  natural  populations  in  the  Gulf  of  Naples 
where  increased  urbanization  and  industrial  development  was  strongly  affecting 
coastal  waters.  An  important  aspect  of  his  research  was  the  sustained  effort  over  a 
47-year  period  of  research  at  the  Stazione  which  established  a  strong  ecological 
perspective  in  the  study  of  marine  algae. 

Microalgal  studies  concomitant  with  those  mentioned  above  resulted  in  the 
establishment  of  new  species  lists  for  diatoms,  dinoflagellates,  and  other  flagellates. 
Castracane  (1889),  Shiitt  (1891,  1892),  Schroder  (1901),  Karsten  (1925),  and 
Balsomo  (1903)  substantially  added  to  the  microalgal  species  discovered  in  the  Gulf 
of  Naples.  Subsequent  works  of  Lindemann  (1924,  1925),  Zimmerman  (1930), 
Schussnig  (1930),  and  Schwarz  (1932)  added  further  understanding  of  dinoflagellates 
and  microflagellates  of  this  area. 

An  understandable  decline  in  activity  preceeded  and  followed  the  wars  but  the 
decade  prior  to  World  War  II  marked  a  period  of  intense  research  with  macro  algae. 
Physiological  studies  dealing  with  temperature  (Biebl,  1939),  growth  substances 
(Weij,  1933),  osmotic  relationships  (Hofler,  1930,  1931,  1932),  ion  permeability 
(Brooks-Moldenhaur,  1932;  Magdefrau,  1933;  Ullrich,  1933,  1934,  1936,  1939), 
and  pigment  composition  (Rodio,  1926,  1929,  1939)  were  actively  pursued.  Life 
cycle  studies,  primarily  with  brown  algae  were  reported  by  Carter  (1927),  Hoyt 
(1928),  Knight  (1929),  Pantanelli  (1923),  and  Ubisch  (1928,  1931),  and  observations 
on  sexual  cycles  and  structures  were  published  by  Hartmann  (1925,  1934,  1937), 
Jollos  (1926),  Foyn  (1934a,  b),  and  Moewus  (1938).  During  this  period  both 
Hammerling  (1934a,  b)  and  Schulze  (1939)  pursued  studies  of  Acetabularia  species 
present  in  the  local  waters.  These  studies  were  part  of  the  pioneering  work  which 
was  pursued  in  their  native  Germany  establishing  Acetabularia  as  an  important 
physiological  model  and  tool  in  the  studies  of  cell  biology. 

The  years  following  the  second  world  war  saw  few  or  no  botanists  at  the 
Stazione.  It  was  not  until  the  early  50's  that  botanical  studies  resumed  full  activity 
in  research,  progressing  with  ecological  studies  of  algal  distribution,  algal  cultivation, 
and  work  dealing  with  various  aspects  of  Acetabularia  metabolism.  In  the  late 
1950s,  a  marked  change  occurred  in  the  marine  botany  laboratory  with  the 
permanent  assignment  of  Dr.  Kurt  Beth,  of  the  Max  Planck  Institute,  as  head  of 
the  algal  laboratory  in  Naples.  As  a  cell  physiologist.  Dr.  Beth  was  primarily 
interested  in  Acetabularia  research  (Beth,  1958;  Thimann  and  Beth,  1959),  ephiphy- 
tism  (Beth  and  Merola,  1960),  and  reproductive  cycles  in  Halimeda  tuna  (Beth, 
1962).  In  1963,  Dr.  Beth  organized  and  hosted  the  First  International  Algal 
Conference  at  Naples. 

With  less  emphasis  on  ecology,  the  botany  efforts  during  the  decade  between 
1960  and  1970  left  open  pressing  questions  and  concerns  regarding  the  study  of  the 
environment.  As  a  result  of  this  and  other  factors,  the  Benthic  Ecology  and  Biological 
Oceanography  laboratories  were  formed  and  pursued  topics  no  longer  conducted  in 
marine  botanical  research.  Ecological  studies  in  the  broad  sense,  including  research 
on  the  physical-chemical  factors  as  well  as  plant  and  animal  communities,  are  now 
being  conducted  by  the  three  laboratories. 

Today  marine  botany  has  regained  activity  in  both  macro  and  micro  forms 
combining  a  physiological-ecological  approach  to  the  study  of  algae.  Population 
dynamics  of  micro  algae,  distribution  and  abundance  of  toxic  and  noxious  forms, 


170  C.   R.  TOM  AS 

and  physiological  requirements  of  open  ocean  species  are  but  a  few  of  the  research 
interests  added  to  the  macro  algal  distribution  and  abundance  studies.  An  active 
herbarium,  consisting  of  3000+  specimens  dating  from  1881  to  the  present  continues 
to  serve  as  an  important  reference  for  taxonomic  and  systematics  studies.  Activities 
involving  international  collaboration  have  resumed  and  expanded.  Advanced-level 
international  courses  are  currently  being  organized. 

A  lasting  impact  of  the  marine  botanical  activities  during  a  century  of  research 
at  the  Stazione  has  been  a  continuum  of  ecologically  oriented  studies  which  served 
as  a  backbone  for  other  research  activities.  This,  primarily  attributed  to  the  efforts 
of  Professor  Funk,  has  served  as  a  basis  of  what  we  find  today  as  botany-ecology  at 
the  Stazione  Zoologica. 

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CARNEGIE   INSTITUTION   OF  WASHINGTON   AND  MARINE   BIOLOGY: 
NAPLES,   WOODS  HOLE,  AND  TORTUGAS 

JAMES  D.   EBERT 
Carnegie  Institution  of  Washington,  1530  P  Street,  N.  W.,  Washington.  DC  20005 

INTRODUCTION 

At  2:45  p.m.  on  29  January  1902  the  Board  of  Trustees  of  Carnegie  Institution 
of  Washington  met  at  the  State  Department  in  Washington,  DC  under  the  temporary 
chairmanship  of  the  honorable  John  Hay,  with  Charles  D.  Walcott  serving  as 
temporary  secretary.  Andrew  Carnegie,  who  was  introduced  by  the  chairman, 
presented  his  deed,  creating  a  trust  for  the  benefit  of  the  Carnegie  Institution  of 
Washington,  DC.  After  adoption  of  the  by-laws  officers  were  elected,  including  the 
honorable  Abram  S.  Hewitt  as  chairman,  Dr.  John  S.  Billings  as  vice-chairman, 
Walcott  as  secretary,  and  as  the  Institution's  first  president.  Dr.  Daniel  C.  Gilman, 
former  president  of  the  Johns  Hopkins  University. 

The  second  meeting  of  the  Board  was  held  the  following  day  at  the  New  Willard 
Hotel  in  Washington.  At  that  meeting  an  Executive  Committee  was  elected,  which 
was  charged  with  "preparing  a  report  upon  the  work  which  should  be  undertaken 
by  the  Carnegie  Institution  in  the  near  future,  such  report  to  be  submitted  to  the 
Board  of  Trustees  at  its  next  meeting — ."  To  that  end  the  Executive  Committee 
appointed  eighteen  advisory  committees,  whose  roles  were  defined  in  a  letter  from 
President  Gilman  to  each  advisor  on  1 1  March  1902.  These  fifty  individuals  were 
"invited  to  act  as  one  of  these  advisors  until  the  Annual  Meeting  of  the  Trustees, 
in  November  next."  The  charge  to  the  committees  was  "to  prepare,  in  the  course 
of  the  summer,  a  plan  of  procedure,  and  in  the  meantime  to  engage  in  preliminary 
studies  of  the  problems  committed  to  them,  by  consultation  with  acknowledged 
authorities  at  home  and  abroad." 

Among  the  committees  was  the  Committee  on  Zoology.  Its  chairman  was  Henry 
F.  Osborn,  the  other  members  being  Alexander  Agassiz,  W.  K.  Brooks,  C.  Hart 
Merriam,  and  E.  B.  Wilson. 

The  several  Committees  reported  to  the  Trustees  on  25  November  1902.  Those 
reports  appear  in  Carnegie  Institution  of  Washington  Yearbook,  Volume  1 . 

The  reports  of  several  of  the  advisory  committees  are  considered  "classics"  in 
their  respective  fields,  perhaps  the  most  far-reaching  being  a  report  of  the  Advisory 
Committee  on  Astronomy,  which  produced  a  veritable  charter  for  work  in  astronomy 
over  several  decades  and  resulted  immediately  in  the  establishment  of  Mount 
Wilson  Observatory.  The  report  of  the  Committee  on  Zoology  cannot  be  classified 
among  the  great  reports  of  the  group,  in  part  because  only  three  of  the  five  members 
of  the  Committee  took  an  active  part  in  the  deliberations.  Agassiz  withdrew  from 
the  Committee  before  it  completed  its  deliberations,  and  Merriam  did  not  attend 
the  final  critical  meeting.  Merriam  was  one  of  the  first  to  argue  that  Carnegie 
Institution  should  be  an  operating,  not  a  granting  institution.  He  was  opposed  to 
any  plan  that  would  result  in  scattering  the  work  and  funds  of  the  Institution.  He 
believed  that  "existing  institutions  should  be  allowed  to  continue  their  work  without 
aid  or  interference  from  the  Carnegie  Institution."  He  was  "fully  convinced  that  the 
Carnegie  Institution  should  carry  on  its  own  work,  under  its  own  name,  and  should 
publish  the  results  in  its  own  series  of  publications." 

172 


CARNEGIE  AND   MARINE   BIOLOGY  173 

The  Zoology  report  was  signed  only  by  Osborn,  Wilson,  and  Brooks.  It  must  be 
noted  that  all  were  active  members  of  the  Corporation  of  the  Marine  Biological 
Laboratory.  Thus  it  is  not  surprising  that  the  Zoology  report  emphasized  marine 
biology,  treating  it  under  four  different  headings. 

In  a  section  entitled  "Permanent  Advisory  Committee"  it  was  proposed  that  a 
permanent  Advisory  Committee  on  Zoology  be  established,  on  the  rotation  system, 
to  act  as  advisors  in  connection  with  the  Marine  Biological  Laboratory  and 
Experimental  Station,  the  encouragement  of  research,  expeditions — and  so  forth. 
The  reader  will  note  that  the  title  "Marine  Biological  Laboratory  and  Experimental 
Station"  is  capitalized. 

Under  a  further  heading,  "Marine  and  Experimental  Stations"  the  Committee 
strongly  endorsed  "the  establishment  of  a  permanent  biological  laboratory  as  a 
central  station  for  marine  biology  in  general,  with  branches  at  such  other  points  as 
may  seem  desirable;  also  affiliated  or  independent  experimental  stations  for  the 
study  of  physiological  zoology  and  problems  relating  to  heredity,  evolution,  etc." 

Under  the  heading  "Subsidies,"  the  Committee  concluded  that  "the  Zoological 
Station  at  Naples  will  in  all  probability  be  one  of  the  most  important  centers  for 
special  research  work.  ...  It  is  therefore  desirable,  and  this  Committee  strongly 
recommends,  that  the  Carnegie  Institution  subscribe  annually  for  a  table  at  Naples 
to  the  value  of  $500.  .  .  ." 

Finally,  under  "Supplementary  Notes"  is  found  a  minority  report,  a  "note  by 
E.  B.  Wilson"  who  argued  that  the  Institution  should  support  regularly  at  least  two 
tables  at  Naples.  Wilson  wrote,  "The  advantages  derived  by  American  biology  as  a 
whole  from  the  Naples  station  in  the  past  has  been  of  incalculable  value.  .  .  ." 

Before  proceeding,  let  me  call  attention  to  the  striking  differences  in  referring  to 
marine  biology  under  two  headings,  just  a  page  apart.  Under  one  heading  the 
Committee  spoke  of  "the  Marine  Biological  Laboratory  and  Experimental  Station," 
while  under  the  other  it  wrote  more  generally  of  the  establishment  of  a  permanent 
laboratory  "as  a  central  station  for  marine  biology  in  general.  .  .  ."  This  disparity 
has  special  significance,  and  the  reasons  for  it  will  be  made  clear.  It  is  indeed  the 
central  part  of  our  story.  First,  however,  let  me  take  up  briefly  the  history  of 
interactions  between  Carnegie  Institution  and  the  Stazione  Zoologica. 

CARNEGIE  INSTITUTION  AND  STAZIONE  ZOOLOGICA,  1902-1924 

E.  B.  Wilson's  minority  report  prevailed  with  the  Carnegie  Executive  Committee, 
which  on  27  October  1902  recommended  to  the  Trustees  that  the  Institution 
subscribe  to  two  tables,  at  a  total  annual  cost  of  $1000.  The  Trustees  approved  and 
grant  number  55  provided  for  two  tables  for  the  year  1903.  One  of  the  tables  was 
occupied  for  three  months  during  the  spring  by  E.  B.  Wilson,  and  the  other  by 
H.  S.  Jennings  (then  at  Michigan).  The  remainder  of  the  year  the  tables  were  to  be 
"open  to  whomever  the  director  of  the  laboratory  might  wish  to  assign  to  them." 

On  20  December  1902,  Anton  Dohrn  wrote  to  President  Oilman,  thanking  him 
for  his  "most  desired  Christmas  present,"  and  asking  whether  the  two  tables  "will 
be  a  permanent  establishment?"  and  whether  Oilman  wanted  "a  contract."  To  this. 
Oilman  replied,  on  5  January  1903,  "I  can  only  say  that  all  our  appropriations  are 
made  annually,  and  I  have  no  authority  to  commit  the  Trustees  beyond  the  present 
year.  At  the  same  time  I  can  see  no  reason  why  they  should  not  continue  this 
appropriation  for  a  term  of  years." 

In  fact,  the  Institution  provided  for  two  tables,  without  interruption,  until  1915. 
In  1903,  Wilson  proposed  that  the  Institution  take  a  third  table,  but  that  request 
was  denied  and  the  level  remained  at  two  tables. 


174  J    D.   EBERT 

The  Institution's  final  check  for  $1000  was  sent  to  the  Stazione  on  28  January 
1915,  and  was  acknowledged  by  Reinhard  Dohrn,  then  Director,  on  15  February. 
The  Carnegie  files  thereafter  are  not  extensive.  During  World  War  I  the  Royal 
Italian  Government  assumed  the  "temporary  and  extraordinary  administration"  of 
the  Stazione,  and  at  the  same  time  Carnegie  Institution  was  involved  deeply  in 
science  in  the  service  of  the  United  States. 

Correspondence  was  resumed  in  1919-1920.  Inquiries  were  received  from  B. 
Harvey  Carroll,  the  American  Consul  in  Naples,  from  E.  B.  Wilson,  and  from 
William  Treadwell,  among  others.  The  correspondence  in  the  Carnegie  Files  reveals 
very  little  of  the  ferment  in  Naples  at  that  time.  In  response  to  all  of  these  inquiries, 
President  Woodward  declined  to  recommend  to  the  Trustees  that  the  Institution  re- 
establish relations  with  the  Stazione.  On  27  March  1920,  Woodward  wrote  to 
Wilson,  "One  of  the  rules  we  have  followed  since  the  foundation  of  the  Institution 
is  not  to  give  funds  to  governments.  .  .  ."  This  was  to  be  the  "established"  or 
formal  explanation  for  Woodward's  decision.  As  Woodward  wrote  to  Wilson, 
"Some  months  ago  I  informed  the  Italian  authorities  that  the  Institution  would  not 
be  likely  to  give  any  aid  to  the  Station  so  long  as  it  is  maintained  as  a  governmental 
establishment." 

The  correspondence  does  reveal  that  Woodward  as  an  individual  was  sympathetic 
to  Reinhard  Dohrn.  The  Institution's  files  contain  no  direct  correspondence  with 
the  then  Director  of  the  Station,  Professor  Monticelli. 

The  question  remained  dormant  until  1924  when  further  inquiries  were  received, 
including  one  from  C.  B.  Davenport,  Director  of  the  Institution's  own  Department 
of  Genetics.  By  this  time  John  C.  Merriam  had  succeeded  Woodward  as  President. 
On  24  July  1924  Merriam's  administrative  secretary  wrote,  "I  am  sure  that  Dr. 
Merriam  is  desirous  of  cooperating  in  such  a  project  if  it  proves  possible  to  do  so, 
but  for  the  present  the  Executive  Committee  of  the  Institution  has  not  considered 
that  there  are  available  funds  for  this  purpose." 

Here  this  story  ends,  but  the  ending  should  come  as  no  surprise.  The  Institution 
was  supporting  its  own  struggling  Department  of  Marine  Biology,  and  financial 
exigencies  were  already  pressing  President  John  C.  Merriam  and  the  Trustees  toward 
C.  Hart  Merriam's  position  that  the  Institution's  "strength  and  influence  should  not 
be  weakened  by  diluting  and  scattering  its  resources,  but  husbanded  for  uses  in 
keeping  with  the  promise  and  scope  of  the  Institution." 

CARNEGIE  INSTITUTION  AND  THE  MARINE  BIOLOGICAL  LABORATORY 

We  pick  up  our  story  again  in  1901,  not  in  Washington  but  in  Woods  Hole  (or 
Woods  Holl  as  it  was  then  called).  The  Marine  Biological  Laboratory,  then  in  its 
thirteenth  year,  was  in  grave  financial  difficulty.  In  the  words  of  a  committee  chaired 
by  Frank  R.  Lillie  (and  including  C.  M.  Clapp,  E.  G.  Gardiner,  C.  O.  Whitman 
and  E.  B.  Wilson),  reporting  to  the  Corporation  of  the  Laboratory  in  the  summer 
of  1902, 

For  several  years  the  financial  needs  of  the  Laboratory  have  been  growing  without 
any  corresponding  increase  of  income  until  the  conditions  became  alarming.  The 
Trustees  had  frequently  been  told  by  those  to  whom  appeals  for  support  were 
made  that  the  defects  of  our  business  organization  were  deterrent  to  those  who 
might  otherwise  contribute  to  its  material  support.  This  condition  was  brought 
to  a  crisis  by  an  offer  received  by  the  Trustees  before  the  last  Annual  Meeting  of 
the  Corporation,  under  which  generous  financial  support  was  guaranteed  provided 
a  suitable  business  organization  could  be  effected.  It  was  to  attain  this  end  that 


CARNEGIE   AND   MARINE   BIOLOGY  175 

the  Trustees  asked  at  the  last  Annual  Meeting  of  the  Corporation  that  the  power 
of  modifying  the  by-laws  be  entrusted  to  them.  This  power  was  given  by  a 
modification  of  the  by-laws  adopted  at  the  Annual  Meeting  in  August,  1901. 

The  plan  under  consideration  at  that  time  involved  the  transfer  of  the 
property  of  the  Laboratory  to  a  small  section  of  the  present  Board,  composed  of 
business  men  who  were  to  assume  full  financial  control  and  management,  while 
the  scientific  members  of  the  Board  were  to  continue  as  an  advisory  scientific 
council,  with  general  supervision  of  the  work  of  the  Laboratory.  It  was  thus 
hoped  to  secure  an  efficient  financial  administration  as  well  as  large  financial 
support,  without  sacrificing  the  independence  of  the  Laboratory  or  the  cooperative 
principle  which  has  been  so  potent  a  factor  in  its  success  in  the  past. 

This  plan  was  adopted  unanimously  at  a  meeting  of  the  Trustees  of  the  Marine 
Biological  Laboratory,  held  in  Chicago,  2  January  1902,  and  it  appeared  that  a 
fundamental  change  in  the  organization  of  the  Laboratory  would  be  effected  that 
very  year,  requiring  only  ratification  by  the  Trustees  at  a  special  meeting  to  be  held 
in  the  state  of  Massachusetts. 

But,  as  the  Lillie  Committee  wrote,  ".  .  .  before  ratification  of  this  action,  the 
announcement  of  the  Carnegie  Institution  suggested  the  possibility  that  the  great 
resources  of  this  endowment  might  be  made  available  for  the  support  of  the 
Laboratory." 

This  plan,  which  was  never  ratified,  is  described  fully  by  F.  R.  Lillie  as  the 
'"Chicago  Plan"  in  his  book  The  Woods  Hole  Marine  Biological  Laboratory  (1944). 
It  must  be  described  here  briefly  because  the  controversy  surrounding  it  helped  to 
shape  the  environment  in  which  the  "Carnegie  Plan"  was  debated. 

On  2  August  1901  President  W.  R.  Harper  of  the  University  of  Chicago  had 
written  to  the  Laboratory's  Director,  C.  O.  Whitman  (also  a  Professor  at  the 
University)  to  say  that  "a  company  of  gentlemen,  including  Mr.  A.  C.  Bartlett  of 
Chicago,  Mr.  Charles  Coolidge  of  Boston,  Mr.  C.  R.  Crane  of  Chicago,  and  Mr. 
L.  L.  Nunn  of  Telluride,  Colorado"  were  prepared  to  become  the  Laboratory's 
Trustees,  assuming  full  financial  control  and  providing  management  (including  a 
guarantee  of  ten  thousand  dollars  in  1902),  with  the  existing  scientific  Board 
assuming  the  new  role  of  scientific  advisors. 

Why  did  the  Chicago  Plan  fail?  The  projected  special  meeting  of  the  Trustees 
was  held,  but  only  seven  Trustees  (barely  a  quorum)  appeared,  and  the  Director 
was  absent. 

As  Lillie  described  it,  there  was  hope  of  securing  aid  from  Carnegie  Institution. 
However  there  was  apprehension  as  well.  After  all  the  plan  emanated  from,  or  was 
conveyed  by,  President  Harper,  and  two  of  the  key  "players"  were  Whitman  and 
Lillie,  also  at  the  University  of  Chicago.  The  Trustees  clearly  feared  control  by  a 
single  university.  These  fears  were  exacerbated  by  the  fact  that  the  new  lay  Trustees 
would  include  "in-laws"  of  both  Whitman  and  Lillie. 

Thus  it  is  not  surprising  that  the  Laboratory's  Trustees  turned  readily  toward 
the  fledgling  Carnegie  Institution,  but  they  were  not  alone  among  marine  biologists 
in  their  dream  of  tapping  the  Carnegie  wealth.  David  Starr  Jordan  submitted  a  plan 
for  studying  the  fish  of  the  Pacific  Ocean.  Alexander  Agassiz,  then  President  of  the 
National  Academy  of  Sciences,  proposed  an  expedition  to  the  Pacific  to  study 
marine  life.  The  Governor  of  Bermuda  sought  support  for  the  marine  station  there. 
The  Marine  Biological  Laboratory  had  one  unique  advantage,  however,  in  the 
person  of  E.  B.  Wilson,  a  devoted  "MBLer,"  and  a  member  of  Carnegie's  Committee 
on  Zoology,  a  staunch  advocate  of  the  establishment  of  a  "permanent  biological 
laboratory  as  a  central  station  for  marine  biology  in  general."  Moreover,  Wilson 


176  J.   D.   EBERT 

was  not  alone  in  his  support  of  the  Marine  Biological  Laboratory,  for  as  we  have 
already  observed,  both  Brooks  and  Osborn  served  both  the  MBL  and  Carnegie. 

Events  moved  quickly.  Overtures  were  made  by  MBL  Trustees  to  their  counter- 
parts at  Carnegie,  and  a  formal  application  for  aid  was  sent  to  President  Oilman. 
On  11  March  1902,  on  Walcorfs  motion,  the  Carnegie  Executive  Committee 
resolved  that  Dr.  J.  S.  Billings  be  appointed  a  special  committee  of  one  to  investigate 
and  report  upon  the  desirability  of  the  Institution  making  a  grant  for  the  maintenance 
of  the  Marine  Biological  Laboratory.  Billings  conferred  almost  at  once  with  Brooks, 
Osborn,  Whitman,  and  Wilson,  and  as  a  result  of  that  conference,  stated  his 
willingness  to  report  favorably  on  the  application.  It  should  be  emphasized  that 
Billings  made  his  own  position  clear  at  the  outset:  Carnegie  would  be  more  than  a 
granting  institution.  He  believed  that  the  Institution  should  place  the  Laboratory 
on  a  permanent  basis,  purchase  land,  erect  and  equip  a  new  laboratory,  and  make 
suitable  provision  for  its  maintenance.  Moreover,  if  this  maintenance  were  to  be  of 
a  permanent  character,  he  argued,  Carnegie  should  be  placed  in  full  financial  control 
of  the  property  of  the  Laboratory. 

Moreover,  the  level  of  support  envisioned  by  Professor  Wilson  and  other 
members  of  the  MBL  Board  of  Trustees  was  made  clear  to  Billings,  who  wrote  to 
President  Oilman  on  24  March  1902,  "This  will  involve  an  expenditure  of  about 
$80,000  within  the  next  three  or  four  years  .  .  .  ,  and  also  an  annual  expenditure 
of  about  $30,000  for  current  expenses." 

Finally,  the  Carnegie  perspective  on  teaching  was  made  clear  as  early  as  13 
March  1902,  when  Billings  wrote  to  Wilson  ".  .  .  it  should  be  understood  that  the 
primary  object  of  the  Laboratory  is  to  promote  original  research  and  to  give  to 
competent  persons  an  opportunity  to  make  such  research,  and  that  it  is  not  of  the 
nature  of  a  school  for  teaching  ordinary  students?" 

As  a  result  of  the  conference  between  Billings  and  the  MBL  group,  a  special 
meeting  of  the  Board  of  Trustees  of  the  Marine  Biological  Laboratory  was  called. 
At  that  meeting,  held  in  New  York,  22  March  1902,  the  Trustees  approved  the 
incorporation  of  the  Marine  Biological  Laboratory  by  the  Carnegie  Institution  "on 
the  lines  indicated  in  the  letter  of  Dr.  Billings,"  and  called  for  the  formation  of  a 
committee  to  work  with  the  Executive  Committee  of  Carnegie  Institution.  Carnegie 
responded  almost  immediately,  following  a  meeting  of  its  Executive  Committee  on 
25  March  1902.  It  was  resolved  that  Carnegie  would  acquire  the  Marine  Biological 
Laboratory,  with  the  understanding  "that  Trustees  of  the  Laboratory  are  willing  to 
turn  over  its  plant  to  the  Institution,  provided  the  latter  will  undertake  the 
maintenance  and  support  of  the  Laboratory."  The  Carnegie  Trustees  envisioned 
that  the  MBL  would  become  the  Institution's  "Department  of  Marine  Zoology."  A 
special  committee  was  formed  by  Carnegie  to  work  with  an  MBL  Committee  to 
prepare  a  detailed  plan  for  the  organization  of  the  new  department.  Moreover,  to 
help  the  MBL  through  its  difficult  times,  the  Executive  Committee  resolved  to 
provide  the  sum  of  $4,000  on  or  after  1  August  1 902  as  a  first  contribution  toward 
the  expenses  of  the  Laboratory — providing  satisfactory  evidence  was  furnished  "that 
the  Trustees  of  the  Marine  Biological  Laboratory  have  full  power  to  transfer  the 
property  of  the  Laboratory  to  the  Institution,  and  have  agreed  to  do  so." 

Two  joint  meetings  of  the  Conference  Committee  were  held  in  April  and  May. 
Now,  second  thoughts  began  to  emerge. 

The  MBL  was  in  a  state  of  ferment — in  crisis — and  looked  to  Carnegie  (as  it 
had  looked  to  the  Chicago  group)  as  its  financial  savior.  I  emphasize  the  word 
"financial."  The  financial  crisis  was  real,  yet  Whitman  played  only  a  small  part  in 
the  early  discussions  with  Carnegie.  It  was  Wilson  to  whom  most  of  the  Carnegie 


CARNEGIE  AND  MARINE   BIOLOGY  177 

correspondence  was  directed.  Wilson  appears  to  have  understood  and  to  have 
accepted  the  Carnegie  position.  Whitman  surely  understood  it — but  he  did  not 
accept  it;  still  he  did  not  oppose  it  openly.  On  10  May  1902,  Wilson  wrote  to 
Walcott  as  follows: 

In  reply  to  your  letter  of  May  3rd:  I  shall  be  glad  to  make  a  number  of  suggestions 
regarding  marine  biological  research  in  case  the  Wood's  Hole  plan  is  not  carried 
out,  but  I  hope  that  there  is  no  danger  of  this  plan  failing.  The  only  difficulty 
with  the  plan  seems  to  be  that  Professor  Whitman  is  reluctant  to  take  any  steps 
which  will  not  unite  the  support  of  the  Carnegie  Institution  with  that  of  Messrs. 
Crane,  Nunn  and  others.  The  rest  of  us  feel  that  the  plan  embodied  in  the  sketch 
of  by-laws  that  has  been  drawn  up  in  consultation  with  Messrs.  Billings  and 
Hewitt  ought  to  insure  this  and  probably  will,  and  I  trust  there  will  be  no 
difficulty  in  making  the  transfer  of  the  property. 

Carnegie,  too,  was  in  ferment,  but  its  ferment  was  of  a  different  kind.  Andrew 
Carnegie  had  decided  not  to  found  a  national  university,  but  to  establish  an 
institution  devoted  to  pioneering  research  and  research  training,  not  college  or 
university  education.  Its  Trustees  were  of  several  minds,  however,  as  to  the  way 
Mr.  Carnegie's  mandate  should  be  followed.  Should  all  of  the  Institution's  funds  be 
devoted  to  its  own  operating  departments,  for  which  there  were  large  demands  from 
the  very  beginning?  Should  the  Institution  be  primarily  a  granting  agency,  and  if 
so,  should  the  recipients  of  grants  be  established  institutions,  or  individuals?  It  was 
many  years  before  the  question  was  fully  resolved,  with  the  Institution  focusing 
entirely  on  operating  its  own  departments.  In  1902,  however  two  trends  had  already 
begun  to  emerge,  the  formation  of  new  operating  departments,  and  grants  to 
exceptional  individuals.  The  idea  of  providing  substantial  grants  on  a  long-term 
basis  to  existing  organizations  found  less  favor.  The  proposal  by  the  Marine 
Biological  Laboratory  provided  the  first  great  test  for  the  Carnegie  Trustees  on  this 
question. 

The  test  began  with  a  proposal  from  the  Laboratory  that  a  joint  Board  of 
Trustees  be  established,  equally  representing  Carnegie  and  the  existing  Board  of  the 
Marine  Biological  Laboratory.  This  proposition  failed,  with  the  Carnegie  represen- 
tatives stating,  in  substance,  that  it  was  not  the  Institution's  policy  to  enter  into 
alliance  with  existing  institutions  in  such  a  manner  as  to  involve  divided  control. 
The  Carnegie  group  made  clear  that  the  Institution  might,  from  time  to  time,  make 
special  grants  to  the  Marine  Biological  Laboratory  (as  it  might  to  other  institutions 
or  individuals)  but  that  permanent  and  continuous  support  could  only  be  promised 
on  condition  of  a  definite  and  complete  transfer  of  property  to  the  Institution,  so 
that  it  might  assume  full  financial  control  and  responsibility.  They  stated  positively 
at  that  time,  and  subsequently,  that  it  was  the  wish  and  intention  of  Carnegie 
Institution  upon  assuming  control,  to  give  "the  managers"  the  fullest  possible 
scientific  independence  and  freedom. 

In  due  course,  the  Conference  Committee  finally  agreed  to  amend  the  by-laws 
of  the  Marine  Biological  Laboratory  to  permit  the  transfer  of  the  Laboratory  to  the 
Institution.  The  new  by-laws  were  conveyed  to  the  MBL  Trustees  on  19  July  1902. 

The  MBL  would  now  constitute  the  Department  of  Marine  Biology  of  Carnegie 
Institution.  It  would  be  under  the  general  charge  of  a  "Board  of  Managers,"  to  be 
elected  by  the  Executive  Committee  of  Carnegie  Institution.  At  the  outset  it  was 
proposed  that  the  Board  of  Managers  would  be  composed  of  the  then  existing  Board 
of  Trustees  at  MBL.  The  Board  of  Managers  would  have  immediate  charge  of  the 
Laboratory,  it  would  appoint  the  director  of  the  Laboratory  and  in  general  would 
undertake  all  those  responsibilities  normally  undertaken  by  the  previous  Board  of 


178  J-   D.   EBERT 

Trustees.  However,  changes  in  the  by-laws  had  to  be  approved  by  the  Executive 
Committee  of  the  Trustees  of  Carnegie  Institution. 

On  12  August  the  Annual  Meeting  of  the  members  of  the  Corporation  of  the 
Laboratory  took  what  proponents  of  the  transfer  of  the  Laboratory  to  Carnegie 
might  have  regarded  as  final  and  conclusive  action.  The  proposed  by-laws  to  be 
enacted  by  Carnegie  were  read  and  explained  to  the  Corporation.  The  Corporation 
was  apprised  that  at  the  MBL  Trustees  meeting  on  19  July  a  deed  was  approved 
that  would  convey  to  Carnegie  the  land  and  other  holdings  of  the  MBL  Corporation. 
At  the  meeting  of  the  Corporation,  by  a  majority  of  over  60,  it  was  voted  that  the 
Treasurer  be  authorized  to  execute,  acknowledge,  and  deliver,  in  the  name  and  in 
behalf  of  the  Corporation  the  deed  conveying  all  title  to  the  Institution.  Only  three 
negative  votes  were  cast,  including  one  by  an  individual  who  would  submit  a 
competing  proposal  to  the  Institution,  C.  B.  Davenport.  All  that  remained  then  was 
for  the  Marine  Biological  Laboratory  to  report  upon  its  needs  and  plans.  Still 
another  committee  was  empanelled,  this  time  chaired  by  Whitman.  Although 
Whitman  had  voted  for  the  transfer  at  the  meeting  of  the  Corporation,  he  had  let 
it  be  known  that  he  favored  an  alternate  course,  namely  of  getting  as  large  a  grant 
as  possible  from  the  Institution,  but  remaining  independent  of  it.  Indeed,  as  Wilson 
indicated  in  his  letter  of  10  May,  Whitman  wanted  monies  from  both  the  Carnegie 
and  Chicago  sources,  but  above  all,  he  wanted  the  Laboratory's  independence, 
which  he  then  set  out  to  ensure.  Both  he  and  J.  McKeen  Cattell  (who  had  cast  one 
of  the  three  negative  votes)  published  articles  in  Science  calling  the  Corporation's 
action  into  question.  Whitman's  article,  "The  impending  crisis  in  the  history  of  the 
Marine  Biological  Laboratory"  (1902)  is  especially  noteworthy.  Moreover  his 
Committee  report,  presented  as  a  "Report  of  the  Trustees  of  the  Marine  Biological 
Laboratory  to  the  Trustees  of  the  Carnegie  Institution"  called  for  an  effort  far 
beyond  the  scope  proposed  by  Carnegie  at  the  outset.  Moreover,  surprising  especially 
to  those  who  have  known  the  Marine  Biological  Laboratory  in  recent  decades,  when 
a  premium  was  placed  on  working  with  marine  organisms,  is  the  statement  that 
"Biological  Station"  would  better  express  the  character  and  aim  then  did  the  name, 
Marine  Biological  Laboratory.  The  report  states  in  fact  that  the  word  "marine  is 
there  for  a  somewhat  misleading  reminiscence  of  an  early  stage  of  development, 
when  sea  forms  alone  occupied  attention." 

The  report  called  for  not  only  a  "central  station  at  Woods  Hole"  but  also  for 
secondary  stations  on  the  Maine  coast  and  in  the  West  Indies.  It  was  stated  further 
that  "fresh  water  ponds"  would  be  required  as  well.  Finally,  it  was  argued  that  a 
Biological  Farm  would  be  needed  for  studies  of  heredity  and  evolution  (possibly  to 
accommodate  the  needs  of  Davenport). 

Charles  Coolidge,  a  Boston  architect,  provided  an  estimate  on  the  cost  of 
buildings,  land  and  equipment,  which  far  exceeded  anything  discussed  previously. 
According  to  this  proposal  the  initial  cost  for  a  wharf,  steam  launch,  buildings, 
ponds,  apparatus  etc.  would  amount  to  over  $450,000.  Maintenance  costs  were 
estimated  at  $30,000  for  1903,  $75,000  for  1904,  and  $100,000  for  1905. 

It  is  difficult  to  evaluate  this  Report  without  indulging  in  "psychohistory."  There 
can  be  no  doubt  that  it  was  prepared  hurriedly,  but  even  considering  that,  it  is  a 
rambling,  poorly  documented  statement,  not  up  to  Whitman's  usual  standard — 
especially  the  long  argument  for  a  Biological  Farm.  The  Committee  had  to  have 
known  that  they  were  calling  for  far  more  than  Carnegie  intended  to  provide,  that 
the  proposal  was  unrealistic  and  doomed  to  fail.  A  "psychohistorian"  could  argue 
that  Whitman  had  decided  that  if  MBL  were  to  lose  its  independence,  it  would  be 
at  a  very  high  price.  This  report  was  prepared  after  1 2  August  and  before  4  October 


CARNEGIE  AND   MARINE   BIOLOGY  179 

1902,  for  on  the  latter  date  the  Carnegie  Executive  Committee  considered  it  along 
with  other  information  received,  from  MBL  Trustees  and  others.  By  then,  it  was 
clear  to  the  Carnegie  Executive  Committee  that  despite  the  MBL  Corporation's 
definitive  action,  there  was  still  substantial  unrest  at  the  Laboratory.  The  Carnegie 
Executive  Committee  felt  constrained  to  inform  MBL  of  the  general  principles  that 
would  govern  the  Executive  Committee  in  its  recommendations  to  the  Carnegie 
Board  in  November. 

The  Carnegie  Executive  Committee  reiterated  that  it  would  have  to  be  distinctly 
understood  that  in  the  case  of  difference  of  opinion  as  to  expenditures  to  be  made 
or  liabilities  to  be  incurred,  or  as  to  the  policy  to  be  pursued  in  the  conduct  of  the 
Laboratory,  the  decision  of  Carnegie  Institution,  after  proper  hearing  of  the  views 
of  the  managers  to  the  Laboratory,  shall  be  final  and  conclusive. 

Moreover,  the  Executive  Committee  reiterated  Billing's  statement  of  24  March 
1902,  that  the  Laboratory  should  be  used  for  research  and  not  for  teaching  and  that 
no  instruction  shall  be  given  except  such  as  may  be  furnished  by  investigators  to 
their  assistants.  The  object  should  be  to  provide  "competent  investigators  with 
facilities  for  making  researches." 

Moreover,  the  scheme  proposed  by  the  MBL  Trustees  of  making  the  MBL  an 
institution  for  the  investigation  of  problems  of  evolution,  heredity,  etc.,  including  a 
Biological  Farm,  would  not  be  approved. 

Finally,  the  Executive  Committee  reiterated  that  it  would  not  recommend  more 
than  $80,000  for  land,  buildings,  etc.,  spread  over  the  first  two  years;  moreover,  it 
would  recommend  $10.000  a  year  for  maintenance.  Finally,  in  a  crucial  statement, 
the  Executive  Committee  concluded 

if,  after  considering  these  statements,  the  Board  of  Trustees  (of  the  Marine 
Biological  Laboratory)  is  of  the  opinion  that  it  would  prefer  to  retain  the 
independence  of  the  Laboratory,  as  urged  by  the  present  Director  (Whitman), 
and  not  turn  over  its  property  to  the  Carnegie  Institution,  the  Executive 
Committee  is  prepared  to  consider  a  proposition  for  granting  aid  to  the  Laboratory 
to  the  amount  of  $10,000  a  year  for  the  next  three  years,  on  condition  that 
twenty  research  tables  be  placed  at  the  disposal  of  Carnegie  Institution,  the 
occupant  of  each  table  to  be  furnished  with  supplies  and  material  substantially 
as  is  done  by  the  Naples  laboratory. 

The  Executive  Committee  asked  the  MBL  Trustees  to  respond  by  25  October  1902. 

The  Carnegie  offer  of  a  significant  grant  for  three  years  was  exactly  what 
Whitman  had  been  seeking.  He  was  quick  to  move,  convening  a  meeting  of  the 
Executive  Committee  of  the  Marine  Biological  Laboratory  to  reply  to  the  Carnegie 
Executive  Committee's  resolutions.  Two  reports  were  submitted  by  the  MBL 
Executive  Committee,  a  majority  report,  signed  by  three  members  of  the  committee. 
Whitman,  Lillie,  and  Jacob  Reighard;  and  a  minority  report  signed  by  E.  B.  Wilson 
and  T.  H.  Morgan.  The  majority  report  stated  that  in  its  view  the  general  principles 
stated  by  the  Carnegie  Executive  Committee  were  "in  some  essential  respects  so 
different  from  anything  that  has  been  hitherto  considered  by  the  Corporation  and 
Trustees  of  the  Marine  Biological  Laboratory,  that  they  would  not  feel  justified  in 
authorizing  the  transfer  of  the  property  .  .  .  without  adequate  reconsideration  by 
that  body  and  the  Trustees."  They  therefore  stated  their  opinion  that  the  MBL 
should  retain  its  independence,  and  requested  a  grant  of  $10,000  a  year  for  a  period 
of  three  years. 

The  minority  report  agreed  with  the  majority  of  the  committee  that  the  transfer 
of  the  Laboratory  to  the  Institution  "is  for  the  present  inexpedient."  They  were 


180  J.   D.   EBERT 

very  explicit  however  in  stating  that  they  held  this  opinion  not  because  they  regarded 
the  Carnegie  recommendations  as  essentially  different  from  the  earlier  plan,  but 
"because  the  Director  of  the  Laboratory  has  come  to  doubt  the  desirability  of  that 
plan."  They  stated  further  "we  do  not  agree  with  the  Director's  view,  but  consider 
the  transfer  of  the  Laboratory  inexpedient  unless  practically  unanimous  action  can 
be  taken."  Thus  the  minority  view  argued  that  the  wisest  course  is  "to  recommend 
application  for  a  grant  of  $10,000  for  one  or  more  years  .  .  .  until  the  situation 
may  become  more  clearly  denned." 

Wilson  and  Morgan  went  on  to  express  the  hope  that  Carnegie  Institution  may 
"without  detriment  to  the  interests  of  the  work  at  Woods  Hole,  establish  or  support 
a  marine  station  or  stations,  devoted  to  pure  research,  at  such  points  as  may  seem 
desirable." 

Neither  minutes  nor  correspondence  reveal  disappointment  on  the  Carnegie 
side,  even  though  the  record  clearly  shows  that  the  final  Carnegie  position  differed 
hardly  at  all  from  the  recommendations  advanced  by  Billings  six  months  earlier. 
MBL  knew  what  Carnegie  had  in  mind  (Wilson's  letter  of  10  May  to  Walcott),  but 
Whitman,  undoubtedly  stung  by  the  earlier  criticisms  he  had  suffered  because  of 
the  "Chicago  plan,"  held  stubbornly  to  his  vision  of  independence,  and  outmaneu- 
vered  (or  perhaps  better,  outlasted)  Wilson.  The  Carnegie  position  was  clear:  its 
Executive  Committee  had  offered  the  MBL  two  propositions  and  one  had  been 
accepted.  Nothing  more  need  be  said. 

It  is  clear  that  on  the  MBL  side  not  only  Wilson  and  Morgan,  but  others  were 
disappointed.  Davenport  was  not.  Carnegie  provided,  as  agreed,  $10,000  annually 
for  three  years.  In  1905  Frank  Lillie  applied  for  continuation  of  aid  for  the 
laboratory  for  the  next  ten  years,  or  indefinitely.  On  23  January  1906,  President 
Woodward  responded,  "I  regret  to  state  that  after  careful  consideration  it  was 
decided  that  your  petition  may  not  be  granted." 

Carnegie  Institution's  Year  Book  Number  3  (1904)  announced  the  establishment 
of  a  Department  of  Experimental  Biology,  including  a  Station  for  Experimental 
Evolution  at  Cold  Spring  Harbor,  New  York,  to  be  directed  by  Charles  B.  Davenport, 
and  a  Marine  Biological  Laboratory  at  the  Dry  Tortugas,  Florida,  directed  by  Alfred 
G.  Mayer. 

The  Station  for  Experimental  Evolution,  combined  with  Carnegie's  Eugenics 
Record  Office,  differentiated  gradually  into  the  Department  of  Genetics,  which 
provided  some  of  the  most  glorious  chapters  in  the  history  of  that  subject,  capped 
by  a  period  in  which  the  principal  scientists  in  residence  included  Hershey, 
McClintock,  Demerec,  Kaufmann,  and  Streisinger,  among  others. 

Although  Mayer  himself  was  a  man  of  exceptional  promise,  whose  contributions 
loomed  large  at  the  time,  the  history  of  the  Laboratory  at  Dry  Tortugas  was  less 
glorious.  It  is  the  subject  of  our  final  chapter. 

CARNEGIE  INSTITUTION'S  DEPARTMENT  OF  MARINE  BIOLOGY 

The  Board  of  Trustees  of  Carnegie  Institution,  encouraged  further  by  the 
Committee  on  Zoology,  continued  to  give  marine  biology  high  priority.  Undaunted 
by  the  failure  of  negotiations  with  the  Marine  Biological  Laboratory  (and  much  to 
the  relief  of  some  of  their  advisors,  and  some  members  of  the  Board  itself)  they 
turned  in  a  new  direction.  Andrew  Carnegie,  and  the  Trustees,  had  emphasized  that 
the  Institution  should  seek  out  and  encourage  the  exceptional  individual.  An  early 
"model"  of  the  development  of  a  truly  outstanding  department,  arising  from  the 
brilliance  and  indomitable  energy  of  one  man,  was  the  Department  of  Astronomy, 


CARNEGIE  AND   MARINE   BIOLOGY  181 

forged  by  George  Ellery  Hale.  Encouraged  by  this  model,  the  Institution  turned  to 
Alfred  G.  Mayer,  a  marine  biologist  and  biological  oceanographer,  then  just  36 
years  old.  By  all  accounts,  Mayer  was  indeed  an  attractive  individual.  The  son  of  a 
distinguished  experimental  physicist,  Mayer  had  early  on  forsaken  his  father's  field 
to  study  biology,  encouraged  by  Alexander  Agassiz,  with  whom  he  had  collaborated. 
Mayer,  then  a  curator  at  the  Brooklyn  Institute  of  Arts  and  Sciences,  was  already 
an  established  investigator  and  prolific  author,  and  he  remained  highly  productive 
for  the  next  eighteen  years  of  his  brief  life.  He  was  a  naturalist,  systematist,  and 
comparative  physiologist,  as  well  as  a  superb  sailor  and  a  gifted  amateur  engineer 
and  artist.  Using  the  Laboratory's  yachts,  Physalia  and  Anton  Dohrn,  he  studied  the 
fauna  along  the  Atlantic  coast,  and  in  other  vessels  made  expeditions  to  the  Pacific. 
He  wrote  about  the  growth  of  coral.  He  was  deeply  interested  in  the  phenomenon 
of  rhythmical  pulsation  in  marine  organisms — in  the  medusa,  in  the  branchial  arms 
of  the  barnacle,  in  the  heart  of  Salpa,  and  of  the  embryo  Logger  Head  turtle.  He 
had  a  continuing  interest  in  the  swarming  of  the  Atlantic  Palolo  worm.  Despite  the 
breadth  of  his  interests,  he  explored  a  number  of  problems  in  such  depth  that  he 
was  elected  to  the  National  Academy  of  Sciences  at  the  age  of  48. 

But  if  the  Trustees  selected  a  brilliant  man  to  head  the  fledgling  department, 
they  allowed  him  to  select  a  poor  site  for  the  Laboratory — Logger  Head  Key, 
Tortugas,  Florida.  From  the  beginning  the  Laboratory  was  seen  to  have  two 
functions,  to  provide  a  base  for  Mayer's  own  expeditions  and  experimental  studies, 
and  to  provide  a  setting  for  intensive  research  during  the  summer  by  university 
scientists.  Unfortunately,  the  "season"  was  brief,  from  May  through  July  when  the 
hurricane  season  began.  Tortugas,  and  the  Laboratory  as  Mayer  developed  it,  did 
not  support  families  and  a  diverse  summer  community  like  that  at  the  Marine 
Biological  Laboratory.  The  number  of  visiting  investigators  was  small,  but  the 
quality  was  high.  In  1905,  for  example,  the  roster  included  E.  G.  Conklin,  H.  S. 
Jennings,  William  K.  Brooks,  R.  P.  Cowles,  and  Jacob  Reighard,  with  Davenport 
Hooker  as  scientific  collector.  In  the  first  five  years  of  operation,  Mayer  reported  a 
total  of  29  visiting  scientists  in  residence. 

From  the  beginning  however,  the  Laboratory  was  beset  by  problems:  great 
hurricanes,  the  necessity  of  bringing  supplies,  including  potable  water,  from  Key 
West,  and  the  inability  of  the  Laboratory  to  accommodate  wives  and  children. 

Thus,  Mayer  was  lead  to  propose  transferring  the  Laboratory  to  the  Bahamas, 
to  Maine,  or  to  Jamaica.  He  had  emphasized  at  the  outset  that  he  had  erected 
portable  laboratories  suggesting  that  he  never  viewed  Tortugas  as  a  permanent  site. 
None  of  Mayer's  pleas  engendered  a  favorable  response  from  Washington. 

World  War  I  brought  further  difficulties.  The  government  converted  the  Anton 
Dohrn  into  a  patrol  boat  and  Mayer  taught  navigation  and  seamanship.  And  as 
Frank  Portugal  has  written  (unpubl.  ms.), 

this  was  an  equally  unhappy  period  for  Mayer.  Strong  anti-German  sentiment  in 
America  made  him  ashamed  of  his  German  name.  An  embarrassed  Mayer  had 
such  difficulties  in  getting  his  passport  renewed  that  he  had  to  ask  Woodward  for 
letters  of  recommendation  attesting  to  his  loyalty  as  an  American  citizen.  Later, 
when  he  arrived  back  in  America,  he  was  subject  to  an  intensive  search  by 
immigration  officials,  who  labeled  him  a  "suspicious  character"  and  suggested 
that  he  carried  wireless  equipment  for  secret  transmissions  to  the  enemy.  He  had 
no  choice.  Writing  Woodward,  he  announced,  "my  name  has  been  legally 
changed  from  Mayer  (a  Hun  name)  to  Mayor  .  .  .  the  old  form  makes  me 
bristle  whenever  I  look  at  it. 


182  J    D    EBERT 

Problems  continued  to  mount.  Mayor  had  tuberculosis,  and  another  hurricane 
in  1919  severely  damaged  the  facility.  Mayor's  death  in  1922  from  tuberculosis 
sounded  the  death  knell  for  the  Laboratory  itself,  although  the  Laboratory's  death 
was  "lingering."  John  C.  Merriam,  who  had  replaced  Woodward  as  President  of  the 
Institution,  had  little  interest  in  the  Laboratory  himself  and  found  no  support  for  it 
among  other  leading  biologists  in  the  Institution.  The  Trustees  considered  successors 
to  Mayor  including  Alfred  Redfield,  then  an  assistant  professor  in  the  Harvard 
Medical  School,  but  finally  decided  to  perpetuate  the  Laboratory  only  as  a  modest 
center  for  the  work  of  investigators  for  other  institutions.  Thus,  instead  of  naming 
a  new  director,  they  named  William  H.  Longley  of  Goucher  College  as  Administrative 
Officer.  All  through  the  1930s  Merriam  and  the  Trustees  vacillated  about  closing 
the  Laboratory.  When  Longley  died  in  1937  David  Tennent  of  Bryn  Mawr  was 
asked  to  succeed  him  on  an  interim  basis. 

Vannevar  Bush,  who  succeeded  Merriam  as  President  in  1939,  settled  the  issue 
quickly.  He  closed  the  Laboratory  and  repaired  the  Anton  Dohrn  and  transported  it 
to  the  Woods  Hole  Oceanographic  Institution. 

Thus  the  dreams  of  Alfred  G.  Mayor,  and  of  the  Institution's  1902  Committee 
on  Zoology  were  never  fully  realized,  for  marine  biology  was  one  of  the  few  fields 
upon  which  the  Institution  embarked  in  which  its  efforts  did  not  truly  "make  a 
lasting  difference." 

LITERATURE  CITED1 

LILLIE,  F.  R.  1944.  The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press.  284  pp. 
WHITMAN,  C.  O.  1902.  The  impending  crisis  in  the  history  of  the  Marine  Biological  Laboratory.  Science 
16:  529-533. 

1  I  have  drawn  heavily  on  Carnegie  Institution  of  Washington  Year  Books,  and  on  the  Institution's 
Archives. 


Reference:  Biol.  Bull.  168  (suppl.):  183-186.  (June,  1985) 


EVOLVING   INSTITUTIONAL   PATTERNS   FOR   EXCELLENCE:   A   BRIEF 

COMPARISON   OF  THE  ORGANIZATION   AND   MANAGEMENT  OF 

THE  COLD  SPRING   HARBOR   LABORATORY   AND 

THE   MARINE   BIOLOGICAL   LABORATORY 

JAMES   D.   EBERT 

Carnegie  Institution  of  Washington,  15 30  P  Street,  N.  W.  Washington.  DC  20005 

INTRODUCTION 

I  was  asked  to  compare  the  organization  and  management  of  the  Cold  Spring 
Harbor  Laboratory  and  the  Marine  Biological  Laboratory.  I  began  with  the  MBL, 
which  was  better  known  to  the  participants  in  the  symposium. 

MARINE  BIOLOGICAL  LABORATORY 

The  crucial  element  in  the  organization  of  the  Marine  Biological  Laboratory  is 
the  relationship  between  the  Trustees  and  the  members  who  constitute  the  Corpo- 
ration, for,  in  F.  R.  Lillie's  words  "this  has  been  a  controlling  factor  in  the  history 
of  the  Marine  Biological  Laboratory.  The  members  constituting  the  Corporation 
proper  elect — usually  from  their  own  membership — the  Trustees,  who,  in  their  turn, 
have  the  exclusive  right  to  elect  the  members  of  the  Corporation."  (Lillie,  1944). 

When  the  Laboratory  was  incorporated  in  March,  1888,  seven  Trustees  were 
elected,  one  of  whom,  Alpheus  Hyatt,  was  the  first  President.  There  were  in  addition 
a  Secretary,  a  Treasurer,  and  a  Clerk.  Immediately  after  they  were  elected,  the  first 
Trustees  elected  forty-seven  additional  members  of  the  Corporation.  For  several 
years  thereafter  it  was  the  custom  to  elect  to  the  Corporation  all  workers  at  the 
Laboratory  who  signified  their  willingness  to  become  members.  By  1894,  according 
to  Lillie,  there  were  304  regular  members  and  52  life  members. 

Gradually,  over  the  years,  the  number  of  Trustees  was  increased — to  eleven  in 
1889,  to  nineteen  in  1891  (plus  two  members  ex-officio),  and  in  1897  to  twenty- 
four,  divided  into  four  classes  of  six  each,  with  three  members  ex-officio,  the 
Director,  Assistant  Director,  and  Clerk. 

By  the  1940s,  the  number  of  Trustees  had  grown  to  thirty-two,  four  classes  of 
eight,  plus  five  members  ex-officio,  the  President  and  Vice-President  of  the  Corpo- 
ration, the  Director,  Treasurer,  and  Clerk.  With  rare  exceptions,  all  of  the  Officers 
and  Trustees  were  scientists. 

This  pattern  did  not  change  significantly  until  the  1960s.  In  1963  the  by-laws 
were  amended,  adding  a  Chairman  of  the  Board  of  Trustees  to  the  already  existing 
President  and  Director.  The  responsibilities  of  the  Chairman  and  President  were 
sharply  defined,  with  the  Chairman  (a  non-scientist)  being  responsible  for  business 
and  external  affairs,  while  the  President,  a  scientist,  would  provide  oversight,  along 
with  the  Director,  over  scientific  matters.  The  first  Chairman  was  Gerard  Swope, 
Jr.,  and  the  President  and  Director  at  that  time  were  Arthur  Parpart  and  Philip 
Armstrong,  respectively.  Later  in  the  1960s  the  number  of  Trustees  was  increased 
to  thirty-six,  with  the  additional  Trustee  in  each  class  being  a  non-scientist,  or  "lay" 
Trustee. 


183 


184  J.   D.   EBERT 

In  the  1970s,  as  the  Laboratory  moved  further  toward  functioning  as  a  year 
round  scientific  center,  further  changes  in  governance  were  required,  and  the 
composition  of  the  Board  of  Trustees  was  changed  again,  evolving  to  the  pattern  of 
the  current  Board,  twenty-four  scientific  Trustees,  six  in  each  class,  elected  by  the 
Corporation,  plus  twelve  "lay"  or  "Board"  Trustees,  three  in  each  class,  elected  by 
the  Board  of  Trustees. 

Since  1970,  the  roles  of  President  and  Director  have  always  been  filled  by  the 
same  individual,  except  for  1975-1976  when  K.  R.  Porter  was  Director,  with  J.  D. 
Ebert  serving  as  President. 

Authority  to  act  for  the  Board  of  Trustees  between  meetings  is  vested  in  the 
Executive  Committee  of  the  Board  of  Trustees,  composed  of  the  officers  of  the 
Board  plus  six  members  elected  by  the  Board  of  Trustees. 

COLD  SPRING  HARBOR  LABORATORY 

I  turn  now  to  the  governance  and  major  affiliations  of  the  Cold  Spring  Harbor 
Laboratory. 

It  may  be  instructive,  before  examining  the  present  organization,  to  take  up 
briefly  the  origins  of  the  Laboratory. 

In  the  mid- 19th  century,  John  H.  Jones  built  a  dock  to  facilitate  the  outfitting 
of  whale-ships  on  the  east  side  of  the  inner  harbor  of  the  Long  Island  village  of 
Cold  Spring  Harbor.  His  son,  John  D.  Jones,  inherited  the  family  homestead  and 
adjoining  grounds.  John  D.  Jones  was  an  uncommon  man  of  his  day,  who  developed 
a  keen  interest  in  science.  In  the  words  of  his  brother,  W.  R.  T.  Jones  (1904), 

the  Brooklyn  Institute  [of  Arts  and  Sciences]  desiring  a  place  to  establish  a  school 
of  biology,  he  [John  D.  Jones]  put  up  for  that  Institute  a  building  suitable  for  its 
purpose,  and  the  school,  under  charge  of  able  professors,  has  been  a  success, 
doing  original  work  which  has  been  a  credit  to  Long  Island,  and  acknowledged 
as  such  by  similar  foreign  institutions.  He  also  leased  to  the  State  of  New  York 
grounds  for  a  fish  hatchery,  which  is  now  turning  out  each  year  several  hundred 
thousand  trout  and  salmon  to  stock  the  inland  waters  of  the  State. 

Thus  science  at  Cold  Spring  Harbor  sprung  in  large  part  from  the  interest  of  an 
exceptional  individual. 

Seeing  the  need  of  an  organization  to  perpetuate  the  management  and  care  of 
the  grounds  and  property  devoted  by  him  to  scientific  research,  John  D.  Jones 
incorporated  the  Wawepex  Society,  the  name  Wawepex  being  taken  from  an  old 
Indian  name  of  the  harbor.  John  D.  Jones  was  the  first  governor  of  the  Society, 
continuing  in  that  office  until  his  death  in  1895. 

The  next  significant  change  at  Cold  Spring  Harbor  was  the  opening  of  Carnegie 
Institution  of  Washington's  Station  for  Experimental  Evolution  on  11  June  1904, 
under  the  leadership  of  C.  B.  Davenport.  The  Wawepex  Society  offered  about  ten 
acres  of  land,  which  would  be  leased  for  fifty  years  to  Carnegie  Institution  of 
Washington  for  a  nominal  sum.  For  twenty  years  the  Carnegie  Department  and  the 
Biological  Station  of  the  Brooklyn  Institute  of  Arts  and  Sciences  lived  side  by  side. 
The  Station  for  Experimental  Evolution  was  combined  with  Carnegie's  Eugenics 
Record  Office  in  1921  to  form  a  Department  of  Genetics.  Davenport  remained 
Director  until  1934,  when  he  was  succeeded  by  A.  F.  Blakeslee,  who  served  in  that 
role  until  1941. 

In  1924  the  Brooklyn  Institute  withdrew  its  support  of  the  Biological  Station, 
and  community  leaders  in  Cold  Spring  Harbor  organized  their  own  association,  the 


MBL  AND  COLD  SPRING   HARBOR  COMPARED  185 

Long  Island  Biological  Association,  that  actually  administered  the  Laboratory  until 
its  reorganization  as  an  independent  unit  in  1962. 

Although  significant  work  had  been  accomplished  at  Cold  Spring  Harbor  from 
the  very  beginning,  the  first  "truly  great"  period  in  research  there  began  with  the 
arrival  of  Milislav  Demerec  at  the  Carnegie  Department  of  Genetics  in  1941.  The 
two  decades,  1942-1962,  were  extraordinarily  rich  in  both  the  Carnegie  Department 
of  Genetics  and  the  Biological  Laboratory,  managed  by  LIBA.  The  two  organizations 
collaborated  closely,  and  for  a  considerable  period  Demerec  served  as  Director  of 
both  organizations. 

In  1962  the  Carnegie  Department  of  Genetics  was  terminated  as  a  separate 
administrative  unit  of  the  Institution.  Those  of  its  staff  who  remained  at  Cold 
Spring  Harbor  in  the  "Genetics  Research  Unit"  cooperated  with  the  new  Cold 
Spring  Harbor  Laboratory  that  emerged  from  the  Biological  Laboratory  in  that  year. 
As  described  in  the  Annual  Report  of  the  Cold  Spring  Harbor  Laboratory,  LIBA 
remains  a  non-profit  organization,  which  represents  a  growing  constituency  of 
"friends  of  the  Laboratory." 

The  current  Board  of  Trustees  of  the  Cold  Spring  Harbor  Laboratory  is  divided 
between  individual  Trustees  and  "institutional"  Trustees.  Institutional  Trustees— 
currently  there  are  representatives  of  twelve  institutions,  including  LIBA  and  the 
Wawepex  Society — were  brought  on  to  the  Board  during  the  1960s  when  the 
Laboratory  was  undergoing  reorganization.  The  current  Annual  Report  of  the 
Laboratory  states  that 

in  addition  to  supplying  scientific  leadership  to  the  governing  body,  participating 
institutions  also  provided  emergency  funds  to  help  keep  the  Laboratory  afloat 
during  this  crucial  phase  of  development.  Although  participating  institutions  now 
give  only  token  financial  support,  their  Trustees  continue  to  help  steer  the  course 
of  the  laboratory's  scientific  and  administrative  policies. 

There  are  in  addition  thirteen  individual  Trustees;  thus  all  told  there  are  twenty- 
five  Trustees,  who  meet  three  or  four  times  a  year.  Again,  authority  to  act  for  the 
Board  between  meetings  is  vested  in  the  Executive  Committee  of  the  Board,  which 
is  composed  of  officers  of  the  Board  plus  members  elected  to  the  Executive 
Committee  by  the  Board  of  Trustees. 

CRUCIAL  DIFFERENCES 

There  are  two  crucial  differences  in  the  organizations  of  the  Cold  Spring  Harbor 
Laboratory  and  the  Marine  Biological  Laboratory.  They  are  found  in  the  existence 
of  the  MBL  Corporation,  and  in  the  substantial  majority  of  scientific  Trustees 
elected  by  the  Corporation  at  MBL.  As  Gross  has  written  (1984),  these  are  "working 
members  of  the  faculty;  dedicated  scholars.  They  are  specifically  not  outsiders.  They 
have  a  high  personal  stake  in  the  day  to  day  operations  of  the  place." 

At  Cold  Spring  Harbor,  other  than  in  the  legal  sense,  there  is  nothing  equivalent 
to  the  MBL  Corporation.  Control  of  the  Laboratory  is  vested  in  the  Board  of 
Trustees,  with  the  individual  Trustees  having  a  slight  majority.  However,  that 
majority  appears  to  have  little  significance,  for  the  institutional  Trustees  nominated 
by  the  participating  institutions  and  who  rarely  have  a  personal  stake  in  the 
Laboratory's  day  to  day  operations,  tend  to  act  in  concert  with  the  individual 
Trustees. 

From  these  significant  differences  in  organization  spring  further  differences  in 
management,  in  both  policy  and  style,  expressed  in  the  expectations  of  many  MBL 


186  J.   D.   EBERT 

Corporation  members  and  Trustees  to  preserve  and  enhance  "their  own  piece  of 
the  action."  The  uniqueness  of  the  MBL's  organization  has  not  prevented  the 
evolution  of  the  Laboratory  into  a  financially  viable  and  creative  modern  center  for 
research  and  advanced  teaching,  but  at  times  it  has  slowed  the  process  significantly. 

LITERATURE  CITED 

GROSS,  P.  1984.  Report  of  the  Director  of  the  Marine  Biological  Laboratory.  Biol.  Bull.  167:  34-45. 
JONES,  W.  R.  T.  1904.  Address  of  Presentation  by  the  Governor  of  the  Wawepex  Society.  Carnegie  Insl. 

Wash.  Year  Book  3:  34-36. 
LILLIE,  F.  R.  1944.  The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press.  284  pp. 


Reference:  Biol.  Bull.  168  (suppl.):  187-191.  (June.  1985) 


FIRST   IMPRESSIONS:   AMERICAN   BIOLOGISTS  AT  NAPLES 

JANE   MAIENSCHEIN 
Department  of  Philosophy,  Arizona  State  University,  Tempe.  Arizona  85287 

ABSTRACT 

This  paper  examines  reactions  of  American  biologists  who  traveled  to  the 
Stazione  Zoologica  in  Naples  during  the  1880s  and  1890s.  The  1890s  took  a  number 
of  Americans  to  Naples  despite  the  development  of  research  resources  in  their  own 
country.  In  part  this  can  be  attributed  to  the  continued  support  of  the  Stazione  by 
those  Americans  who  had  first  gone,  particularly  by  Whitman,  Wilson,  and  Morgan. 
The  Naples  Station  continued  to  exert  an  important  influence  on  American 
biologists  into  the  first  years  of  the  twentieth  century. 

DISCUSSION 

In  1881,  Professor  William  Keith  Brooks  at  Johns  Hopkins  reported  that  E.  B. 
Wilson's  dissertation  work  would  make  just  as  "valuable  and  handsome  a  paper  as 
those  from  Dohrn's  laboratory,"  if  only  a  place  could  be  found  to  publish  it  (Brooks, 
4  June  1881).  This  statement  revealed  two  phenomena:  first,  that  the  United  States 
did  not  offer  acceptable  vehicles  for  publication  of  detailed  biological  work  and 
second,  that  work  at  Dohrn's  laboratory  set  a  standard  for  biology  by  1881.  I  wish 
to  focus  here  on  the  second  of  these  points.  By  1881,  the  Naples  Station  had 
achieved  a  reputation  in  the  United  States  for  publication  and  research,  but  no 
American  student  had  yet  worked  there.  That  situation  changed  when  Charles  Otis 
Whitman  arrived  in  November  1881. 

Presumably  Whitman  had  heard  about  the  Station  while  he  was  a  graduate 
student  studying  under  Rudolf  Leuckart  in  Leipzig,  but  his  first  opportunity  to  visit 
Naples  came  after  his  three  years  of  teaching  at  the  University  of  Tokyo  (Lillie, 
1911;  Morse,  1912).  On  his  way  back  to  the  United  States,  he  stopped  in  Naples, 
hoping  to  stay  a  few  months.  Since  he  had  worked  in  Germany  and  since  no 
institution  from  the  United  States  had  subscribed  to  a  table  at  the  Naples  Station, 
Anton  Dohrn  welcomed  Whitman  as  his  guest  from  November  until  May  1882 
(Lillie,  1911,  p.  xxiv).  Whitman  wrote  to  a  friend  that  he  was  "having  a  delightful 
time  at  work  in  the  station,'"  and  that  he  found  there  "greater  advantages  than  are 
to  be  found  anywhere  else  in  Europe"  (Whitman,  23  February  1892).  While  he  had 
concentrated  on  leeches  in  Leipzig  and  Japan,  in  Naples  he  turned  to  the  parasitic 
dicyemids  (Whitman,  1883).  Whitman  recorded  his  positive  reaction  to  Naples  and 
Dohrn's  Station  in  an  article  in  Science,  concluding  that  the  international  character 
of  the  Station  had  made  it  the  "Mecca  of  biologists,  and  a  seat  of  unprecedented 
prolific  activity."  Naples  was  the  place  to  learn  methods.  Whitman  wrote,  and 

This  one  but  all-important  matter,  to  say  nothing  of  the  many  other  advantages 
that  must  accrue  to  an  occupant  of  a  table  at  the  station, — such  as  social 
intercourse,  direct  knowledge  of  a  very  important  fauna,  and  opportunities  of 
acquiring  a  knowledge  of  the  four  languages  with  which  every  naturalist  must 
now  be  familiar, — makes  it  very  desirable,  particularly  for  our  younger  naturalists, 
to  spend  some  time  at  Naples  [Whitman,  1883,  pp.  94,  95;  1882]. 

He  continued  to  call  for  American  support  of  the  Station,  arguing  that  even  if  the 

187 


188  J.   MAIKNSCHEIN 

United  States  had  its  own  laboratory  (namely  the  United  States  Fish  Commission), 
Naples  offered  special  opportunities. 

Another  American  visited  Naples  very  shortly  after  Whitman  left,  as  revealed 
by  the  records  at  Naples.  Christiane  Groeben  has  compiled  a  list  of  those  Americans 
who  visited  Naples  and  has  identified  a  number  of  items  in  the  Archives  at  Naples 
for  some  of  those  individuals.  Groeben  reports  that  Emily  Nunn  was  the  second  to 
visit  the  Naples  Station.  She  had  been  studying  in  England  and  worked  at  an 
English  table.  She  later  married  Whitman,  though  they  did  not  meet  in  Naples. 
Emily  Nunn  recorded  her  favorable  impressions  of  the  Station,  but  evidently  was 
advised  to  gain  more  experience  in  independent  research  before  spending  more  time 
at  Naples  so  did  not  stay  as  long  as  she  had  intended  (Nunn,  1883;  Groeben,  1984). 
Whitman  was  certainly  the  more  important  American  visitor  as  far  as  Dohrn  was 
concerned. 

That  Whitman  left  a  very  favorable  impression  on  Dohrn  is  clear  from  Dohrn's 
letter  supporting  Whitman's  application  for  a  position  at  Columbia  University.  As 
Dohrn  wrote,  "The  half  year  you  worked  in  the  Zoological  Station  has  given  me 
the  highest  opinion  of  what  you  will  be  able  to  accomplish  in  the  right  situation; 
and  if  my  word  can  have  the  least  influence  with  the  authorities  of  the  College,  it 
will  go  thoroughly  in  your  favor"  (Dohrn,  25  June,  1885).  Columbia  did  not,  in 
fact,  hire  Whitman  for  that  position  but  instead  eventually  chose  Edmund  Beecher 
Wilson  for  a  similar  job.  And  Wilson  was  the  first  man  to  work  at  the  Naples 
Station  officially,  that  is  at  a  properly  subscribed  table  (Osborn,  1895; 
Groeben,  1984). 

At  first  Wilson  encountered  difficulties  in  receiving  acceptance  to  work  at  the 
Station.  He  had  expected  to  work  at  one  of  the  English  tables  but  found  them  full 
when  he  arrived  in  Naples  in  March  1883.  Also  he  learned  that  "it  is  considered 
unfair  to  admit  men  to  the  station  unless  regularly  provided  for  by  a  subscription 
from  their  own  country."  Even  the  prospect  of  a  subscription  would  allow  Dohrn 
to  admit  Wilson,  but  acting  otherwise  would  be  unfair  to  those  of  other  countries 
whom  Dohrn  had  had  to  turn  away.  Naturally,  Wilson  felt  keenly  disappointed,  for 
he  believed  that  "the  station  has  now  become  practically  the  headquarters  from 
which  most  of  the  leading  European  laboratories  derive  their  best  methods,  and 
where,  indeed,  much  of  their  most  telling  work  is  done."  Naples  remained  a 
"tempting  treasure"  to  be  anticipated  (Wilson,  9  March  1883).  Fortunately,  by 
April  Wilson  was  in  place  at  a  table  subscribed  by  Williams  College  in  an 
arrangement  made  by  Wilson's  cousin  Samuel  Clarke,  who  went  to  Naples  the  next 
year  while  Wilson  taught  at  Williams  for  him  (Morgan,  1940,  p.  126;  1941,  pp. 
319-320;  1942,  p.  240). 

In  the  meantime  President  Gilman  of  Hopkins  had  offered  to  take  a  table  for 
Wilson,  and  Wilson,  "as  a  good  Hopkins  man,"  regretted  that  Hopkins  had  not 
been  the  first  American  institution  to  do  so.  In  part  hoping  to  elicit  Oilman's 
further  support,  Wilson  enthused  about  the  opportunities  at  Naples: 

It  is  in  every  respect  the  best  laboratory  I  have  seen  and  my  high  expectations 
have  been  fully  met. 

Two  things  especially  strike  me  as  characteristic  of  this  laboratory.  The  first 
is  the  perfection  of  the  technical  methods  of  research.  It  is  now  almost  proverbial 
for  zoologists  to  say:  "For  methods  go  to  Naples"  and  in  the  same  breath  is 
usually  added  "A  good  method  is  half  the  battle."  Certain  it  is  that  many  of  the 
best  modes  of  work  now  used  at  Leipsic,  Cambridge  and  elsewhere  have 
originated  here.  The  secret  of  this  is  simply  that  fifteen  or  twenty  zoologists  are 
usually  at  work,  who  come  from  laboratories  in  all  parts  of  the  world  and  bring 


AMERICAN   BIOLOGISTS  AT   NAPLES  189 

their  experience  to  a  focus  here.  They  are  all  experimenting  and  comparing 
results  and  new  methods  can  thus  be  very  thoroughly  tested  .  .  .  [Wilson,  13 
April  1883]. 

Wilson  expected  to  stay  awhile,  possibly  as  long  as  a  year  in  Naples,  though  he 
was  not  sure  that  he  could  afford  such  a  lengthy  visit.  Yet  Dohrn  reportedly  soon 
invited  Wilson  to  stay  for  three  years  and  to  publish  a  lengthy  study  of  the  local 
Rcnilla  which  would  complement  Wilson's  study  of  American  Renilla  (Brooks, 
15  July  1882).  Wilson  loved  the  Station,  the  colorful  setting,  and  especially  the 
music  which  played  an  important  part  in  his  life.  For  family  and  career  reasons,  he 
nonetheless  reluctantly  declined  Dohrn's  offer  and  returned  to  the  United  States. 
As  he  said  later,  Naples  had  made  "a  deep  and  lasting  impression"  on  him  (Morgan, 
1941,  pp.  319-320).  In  discussing  future  plans  for  the  Marine  Biological  Laboratory 
in  later  years,  it  is  clear  that  Wilson  retained  fond  admiration  for  the  Naples  Station 
and  in  some  respects  wished  to  make  the  MBL  more  like  Naples  (Lillie  Papers). 

A  decade  later,  in  the  1890's,  Wilson  returned  for  a  second  visit,  along  with  an 
increasing  number  of  other  Americans.  Undoubtedly  the  enthusiasm  expressed  by 
Whitman  and  Wilson  during  the  summers  at  the  MBL,  as  well  as  the  exciting  new 
work  issuing  from  the  Station  stimulated  a  number  of  American  biologists,  including 
George  Howard  Parker,  William  Morton  Wheeler,  Thomas  Hunt  Morgan,  and 
eventually  Edwin  Grant  Conklin,  to  visit  Naples  themselves.  The  Naples  experience 
influenced  each  of  these  men,  as  each  directly  responded  to  the  dominant  questions 
and  research  of  Naples  in  his  own  work.  Each  of  these  men  recorded  the  impressions 
which  the  Naples  experience  left  on  him. 

Parker  arrived  first,  in  the  spring  of  1893.  He  had  spent  three  summers  in 
Woods  Hole  at  the  U.  S.  Fish  Commission  and  the  MBL,  and  then  had  gone  to 
Europe  to  study  the  origins  of  the  nervous  system.  He  resolved  to  spend  a  half  year 
each  in  Leipzig,  Berlin,  and  Freiburg,  then  to  go  on  to  Naples,  for  as  he  said, 
"Every  young  zoologist  of  my  generation  was  desirous,  as  part  of  his  early  training, 
to  work  at  the  Naples  Zoological  Station"  (Parker,  1946,  pp.  136,  82-83,  91). 
Though  he  did  not  say  much  about  his  scientific  work  at  Naples,  it  seems  clear  that 
the  opportunity  to  work  with  a  variety  of  different  organisms  and  to  discuss  results 
with  a  diverse  group  of  researchers  helped  him  to  some  of  his  generalizations  about 
nervous  structure  and  function.  In  leaving  he  concluded  that  "of  all  places  to  spend 
the  opening  half  of  the  year  Naples  stood  at  the  forefront"  (Parker,  1946,  p.  107). 

Wheeler  arrived  in  Naples  at  the  turn  of  the  new  year,  after  a  stay  with  Theodore 
Boveri  in  Wiirzburg.  In  a  postcard  to  a  friend  at  Chicago,  Wheeler  wrote: 

Have  at  last  reached  the  "Mecca"  and  hope  to  go  to  work  tomorrow,  when  my 
place  in  the  lab  will  be  ready  for  me.  Naples  is  even  more  beautiful  than  1  had 
imagined  it  to  be.  There  is  plenty  of  foliage  on  the  trees  and  the  weather  is 
heavenly  compared  with  what  you  are  probably  having  in  Chicago  .  .  .  Regards 
to  the  boys  and  to  Professor  Whitman  [Evans  and  Evans,  1970,  p.  89]. 

Yet  Wheeler  found  the  poverty  and  Neapolitan  lifestyle  appalling,  even  while  he 
found  the  scenery  so  attractive.  Though  later  known  for  his  outstanding  work  on 
ants.  Wheeler  spent  his  time  at  Naples  on  developmental  studies  of  various 
invertebrates,  including  the  Myzostoma,  which  he  found  begin  as  small  young  males 
and  mature  into  females.  Clearly  stimulated  by  the  dominant  concerns  with 
development  at  the  MBL  and  at  the  University  of  Chicago  under  Whitman, 
Wheeler's  early  researches  followed  the  pattern  of  many  of  the  young  American 
biologists.  His  firm  grounding  in  the  methods  and  problems  of  cytology  and 
developmental  morphology  provided  the  foundation  for  later  work  (Evans  and 


190  J     MAIENSCHEIN 

Evans,  1970,  pp.  89-98).  And  he  pursued  that  grounding  during  three  and  a  half 
months  of  embryological  study  at  Naples.  Presumably  it  was  at  Naples  that  Wheeler 
became  interested  in  the  debates  about  development  stimulated  by  the  half  embryo 
experiments  of  Roux  and  Driesch.  Yet  though  he  went  on  to  translate  Roux's 
manifesto  for  Entwickelungsmechanik  for  an  evening  lecture  at  the  MBL,  Wheeler 
always  maintained  a  traditional  morphological  focus  on  the  whole  organism.  He 
resisted  the  rush  exemplified  by  Entwickelungsmechanik  to  cut  up  organisms  and 
to  manipulate  them  with  experimentation.  Leaving  Europe  in  July  to  return  to 
Chicago,  Wheeler  met  Morgan,  who  was  then  on  his  way  to  Naples  (Wheeler,  1895; 
Evans  and  Evans,  1970,  pp.  104,  234-235). 

Morgan  clearly  received  a  particularly  strong  stimulus  to  his  work  at  Naples. 
Biographers  have  emphasized  the  contact  there  with  Hans  Driesch  and  Driesch's 
impact  on  Morgan  (Allen,  1978).  Clearly  Morgan  did  respond  to  the  debates  in 
progress  among  Driesch,  Roux,  and  others  convened  at  Naples — debates  about  the 
extent  to  which  an  embryo  experiences  any  preformation  or  predetermination 
because  of  its  inheritance  or  early  structure.  Of  the  stimulating  setting,  Morgan 
wrote  that  "No  one  can  fail  to  be  impressed  and  to  learn  much  in  the  clash  of 
thought  and  criticism  that  must  be  present  where  such  diverse  elements  come 
together"  (Morgan,  1896).  In  fact  Morgan's  early  work  generally  followed  closely 
the  interests  of  those  around  him  or  responded  to  problems  which  seemed  particularly 
exciting  at  the  time,  so  the  stimulus  at  the  MBL,  then  Naples  directed  him  (Brooks, 
21  June  1891).  The  years  just  prior  to  his  Naples  visit  reveal  his  existing  interest  in 
the  experimental  work  of  Pfliiger,  Born,  Roux,  Chabry,  Driesch,  and  Hertwig  on 
early  development:  whether  the  concrescence  theory  works  for  teleosts  and  frogs 
and  whether  the  echinoderm  egg  is  isotropic  dominated  Morgan's  work,  with  general 
questions  about  whether  preformation  or  epigenesis  best  characterizes  development. 
Much  of  Morgan's  work  began  with  a  review  and  often  the  repetition  of  other 
results,  then  moved  to  Morgan's  own  related  experiments.  At  Naples,  where  Morgan 
became  friends  with  Driesch,  his  attention  turned  very  directly  to  Driesch's  work 
on  fragmentation  and  partial  embryos  and  their  impact  on  interpretations  of 
development.  In  the  heated  debate  about  preformation  and  epigenesis,  about 
Weismann's  and  Roux's  mosaic  or  Driesch's  regulative  views  of  development, 
Morgan  himself  maintained  a  moderate  position,  sympathetic  to  Driesch  but  closer 
to  Whitman's  emphasis  on  "organic  continuity"  to  explain  development  (Sturtevant, 
1959;  Allen,  1978,  pp.  55-60,  78-84). 

Edwin  Grant  Conklin  similarly  rejected  the  developmental  interpretations  of 
Driesch  or  Weismann  and  Roux.  His  detailed  cell  lineage  work  on  ascidian 
development  led  him  to  conclude  that  cleavage  in  some  forms  is  determinate,  in 
others  indeterminate  with  respect  to  later  development.  This  conclusion  led  him 
into  direct  disagreement  with  Driesch,  who  maintained  that  cleavage  remains 
indeterminate.  Yet  when  Driesch  traveled  to  the  United  States  he  visited  Conklin 
in  Princeton  and  established  friendly  relations.  It  was  not  until  1910,  when  he 
attended  a  conference  at  Graz  that  Conklin  visited  Naples  and  followed  up  some  of 
his  disagreements  with  Driesch.  Specifically,  he  examined  at  Naples  the  same 
organism  that  Driesch  had  studied  there  (Phallusia  mamillata)  and  established  that 
that  form  develops  determinately  as  does  Cynthia  or  Amphioxus,  which  Conklin 
had  studied  in  detail  earlier  (Butler,  1952;  Harvey,  1958,  p.  65).  He  did  not  change 
his  mind  or  his  research  program  because  of  his  work  at  Naples;  rather  his  studies 
allowed  him  to  discredit  Driesch's  alternative  interpretations.  And  Conklin  acquired 
further  material  to  support  his  conclusion  about  the  central  role  of  cytoplasmic 
factors  in  development. 


AMERICAN   BIOLOGISTS   AT  NAPLES  191 

After  the  189CTs  the  American  situation  had  substantially  improved,  and  yet 
Americans  still  visited  Naples.  The  MBL  and  the  Journal  of  Morphology  provided 
a  laboratory  and  a  publication  outlet  for  Americans.  And  successful  graduate 
programs  provided  places  to  pursue  degrees.  Yet  the  Americans  went  to  Naples  to 
learn  theories  and  methods,  to  experience  the  special  international  exchange  of 
ideas  that  took  place  at  that  "Mecca,"  and  to  examine  organisms  native  to  the  area. 
The  MBL  did  not  replace  the  Naples  experience  for  Americans  and  was  not 
intended  to  do  so,  but  complemented  it.  Positive  first  impressions  stimulated  a  long 
tradition  of  American  expeditions  to  the  Naples  Station. 

ACKNOWLEDGMENTS 

I  wish  to  thank  Ruth  Davis  at  the  MBL,  Ann  Blum  at  the  Museum  of 
Comparative  Zoology,  Christiane  Groeben  at  Naples,  and  the  archivists  at  the 
University  of  Chicago,  the  Johns  Hopkins  University  Archives,  and  the  Johns 
Hopkins  University  Manuscripts  Collections  for  their  assistance.  All  passages  quoted 
with  permission. 

LITERATURE   CITED 

ALLEN,  GARLAND.  1978.  Thomas  Hunt  Morgan.  Princeton  University  Press,  Princeton. 

BROOKS,  WILLIAM  KEITH,  to  President  Oilman,  letters.  Oilman  Papers,  Johns  Hopkins  University 

Manuscripts. 
BUTLER,  ELMER  GRIMSHAW.  1952.  Edwin  Grant  Conklin  (1863-1952).  Am.  Phil.  Soc.  Yearbook  1952: 

5-12. 
DOHRN,  ANTON,  to  Whitman,  letter,  Agassiz  Collection,  Museum  of  Comparative  Zoology  Archives, 

Harvard  University. 
EVANS,  MARY  ALICE,  AND  HOWARD  ENSIGN  EVANS.  1970.  William  Morton  Wheeler.  Harvard  University 

Press,  Cambridge. 

GROEBEN,  CHRISTIANE.  1984.  List  of  Americans  who  visited  Naples.  Naples  and  MBL  Archives. 
HARVEY,  E.  NEWTON.  1958.  Edwin  Grant  Conklin.  Nat.  Acad.  Sci.  Biog.  Mem.  31:  54-91. 
LILLIE,  FRANK  RATTRAY.  1911.  Charles  Otis  Whitman.  J.  Morphol.  22:  xv-Lxxvii. 
Lillie  Papers,  MBL  Archives. 

MORGAN,  THOMAS  HUNT.  1896.  Impressions  of  the  Naples  Zoological  Station.  Science  3:  16-18. 
MORGAN,  THOMAS  HUNT.  1940.  Edmund  Beecher  Wilson.  Obit.  Notice  Fellows  of  R.  Soc.  3:  123-138 

(p.  126). 

MORGAN,  THOMAS  HUNT.  1941.  Edmund  Beecher  Wilson.  Nail.  Acad.  Sci.  Biog.  Mem.  21:  315-342. 
MORGAN,  THOMAS  HUNT.  1942.  Edmund  Beecher  Wilson.  Science  96:  239-242. 
MORSE,  EDWARDS.  1912.  Charles  Otis  Whitman  1842-1910.  Nat  I.  Acad.  Sci.  Biog.  Mem.  7:  269-288. 
NUNN,  EMILY.  1883.  The  Naples  Zoological  Station.  Science  1:  479-481,  507-510. 
OSBORN,  HENRY  FAIRFIELD.  1895.  American  students  at  the  Naples  Zoological  Station.  Science  1:  238- 

239. 

PARKER,  GEORGE  HOWARD.  1946.  The  World  Expands.  Harvard  University  Press,  Cambridge. 
STURTEVANT,  A.  H.  1959.  Thomas  Hunt  Morgan.  Natl.  Acad.  Sci.  Biog.  Mem.  33:  283-299. 
WHEELER,  WILLIAM  MORTON.  1895.  The  problems,  methods  and  scope  of  developmental  mechanics. 

Biological  Lectures  1894:  149-189. 
WHITMAN,  CHARLES  OTIS.  1882.  Methods  of  microscopical  research  in  the  Zoological  Station  in  Naples. 

Am.  Nat.  16:  697-706,  722-785. 
WHITMAN,  CHARLES  OTIS.  1883.  The  advantages  of  study  of  the  Naples  Zoological  Station.  Science  2: 

93-97. 

WHITMAN,  CHARLES  OTIS.  1892.  Letter,  Whitman  Papers,  University  of  Chicago  Archives. 
WILSON,  EDMUND  BEECHER.  Student  file,  Johns  Hopkins  University  Archives. 


RdiTcncc:  liiol.  Hull.  168  (suppl.):  192-196.  (June,  19X5) 


EARLY   STRUGGLES  AT  THE   MARINE   BIOLOGICAL   LABORATORY 

OVER   MISSION   AND   MONEY 

JANE   MAIENSCHEIN 

Department  of  Philosophy,  Arizona  State  University,  I'einpe,  Arizona  H5287 

ABSTRACT 

In  its  first  decades,  the  MBL  Trustees  and  their  Director,  Charles  Otis  Whitman, 
often  disagreed  over  the  proper  goals  and  justified  expenditures  for  the  MBL.  This 
paper  examines  the  nature  of  those  struggles  and  the  attempts  at  resolution,  leading 
ultimately  to  Whitman's  disappointment  and  resignation. 

DISCUSSION 

In  1884,  before  the  MBL  began,  Edmund  Beecher  Wilson  had  spent  a  year  in 
Europe  after  finishing  his  degree  at  Johns  Hopkins.  He  wrote  to  President  Oilman 
at  Hopkins  that  a  number  of  Europeans  expressed  surprise  at  the  lack  of  interest  in 
biology  in  America.  So  many  organisms  to  be  explored,  so  many  resources  to  be 
developed.  As  Wilson  wrote,  "American  zoology  seems  to  me  a  good  example  of  a 
prophet  without  honor  in  his  own  country"  (Wilson,  13  April  1883).  He  continued 
to  find  it  embarrassing  that  the  United  States  lacked  a  permanent  research  facility. 
Baird's  efforts  at  the  United  States  Fish  Commission  had  not  succeeded  in  establishing 
it  as  a  research  center.  There  was  room  for  an  American  laboratory,  Wilson  felt, 
and  a  need  for  one. 

Naples  offered  a  fine  example  of  a  biological  research  station,  and  for  Wilson  its 
superiority  lay  clearly  in  one  simple  fact:  that  "money  has  not  been  wanting,  so 
that  the  management  has  been  able  to  offer  good  facilities  for  work  and  has  thus 
attracted  the  best  workers"  (Wilson,  13  April  1883).  Of  course,  Dohrn  deserves 
credit  for  supplying  most  of  the  money,  and  he  worked  continually  to  insure  an 
adequate  income  at  Naples.  No  one  connected  with  the  MBL  offered  as  much  as 
Dohrn  did  at  Naples.  Money  remained  a  constant  problem  in  the  first  decades  of 
the  MBL. 

With  only  $10,000  for  the  first  year,  the  MBL  opened  in  the  rather  shoestring 
manner  described  by  Cornelia  Clapp  (Clapp,  1927).  The  Trustees  expected  their 
money  to  prove  sufficient  for  four  years,  then  they  planned  to  secure  a  permanent 
endowment  (MBL  Minutes,  1888).  Over  the  first  few  years,  they  reluctantly 
authorized  adding  modest  new  buildings  as  the  demand  grew.  Still,  by  the  years 
1892-1894  the  MBL  experienced  a  balanced  budget,  with  the  help  of  careful 
planning  and  cutting  of  extras,  but  1894-1895  brought  the  beginning  of  serious 
crises  (MBL  Minutes,  Annual  Reports). 

These  financial  troubles  reflected  more  fundamental  disagreements  as  well.  The 
Trustees  had  not  specified  the  goals  of  the  MBL,  for  example,  preferring  instead  to 
leave  the  definition  to  the  first  director.  But  they  disagreed  even  about  who  should 
be  that  first  director.  Those  Trustees  from  the  Women's  Education  Association  in 
Boston,  who  had  supported  the  Annisquam  precursor  of  the  MBL,  revealed  their 
expectations  from  the  beginning.  Presumably  they  envisioned  the  MBL  as  following 
ak  r  more  or  less  the  lines  that  the  Annisquam  Laboratory  had  pursued,  concen- 
trati.  mainly  on  fairly  introductory  teaching.  Thus  they  held  that  the  director  of 

192 


EARLY  STRUGGLES  193 

the  MBL  should  be  B.  H.  van  Vleck,  who  had  served  as  assistant  at  Annisquam. 
Other  Trustees,  including  the  Annisquam  director  Alpheus  Hyatt,  saw  the  advantages 
of  selecting  a  nationally  recognized  figure  like  William  Keith  Brooks  or  Charles  Otis 
Whitman,  either  of  whom  would  bring  major  changes  to  the  lab.  In  the  end  the 
Trustees  chose  Brooks,  then  Whitman.  They  left  unspecified  such  major  decisions 
as  the  relative  roles  they  expected  teaching  and  research  to  play,  though  the  intention 
was  always  to  pursue  a  balance  of  both  unless  the  Fish  Commission  established  a 
successful  research  center,  in  which  case  the  MBL  would  focus  on  teaching  classes 
(MBL  Minutes,  1888,  p.  38). 

Beyond  this  basic  agreement  to  pursue  both  teaching  and  research,  however,  the 
Trustees  evidently  never  reached  an  agreement  about  the  ways  and  degree  to  which 
they  expected  the  MBL  to  expand.  Thus,  when  Whitman  insisted  on  making  the 
expenditures  he  regarded  as  necessary  to  supply  a  legitimate  laboratory  for  a  growing 
number  of  people,  the  Trustees  felt  that  Whitman  spent  too  much.  They  sought  to 
limit  his  spending  (Whitman  to  Conklin,  MBL  Minutes,  Annual  Reports).  Obviously, 
such  disagreement  led  to  fundamental  struggles  over  who  would  control  the  MBL. 

Whitman's  contributions  to  the  Annual  Reports  reflect  his  ambitions.  The  goal 
at  the  MBL  should  be  "to  organize  one  of  the  strongest  and  most  productive 
biological  stations  in  the  world"  which  would  allow  the  United  States  to  make  a 
good  showing  when  compared  with  such  successful  places  as  the  Naples  Station 
(Whitman,  Annual  Report,  1890,  p.  22).  The  MBL  should  remain  a  laboratory  for 
all  biology,  including  morphology  and  physiology,  zoology  and  botany,  marine  and 
more  general  biology,  and  it  should  be  a  place  for  both  independent  research  and 
teaching.  For  Whitman,  the  MBL  promised  to  become  the  premier  American 
biological  laboratory.  Yet  he  recognized  the  accuracy  of  Wilson's  emphasis  on 
money.  He  saw  the  need  to  obtain  an  endowment  for  the  lab,  ideally  a  half-million 
dollar  endowment  to  establish  a  full-time,  year-round  biological  station.  His  great 
hopes  met  continued  obstacles,  and  the  ideal  of  a  permanent  center  for  both  research 
and  teaching  remained  unrealized  (Whitman,  1891,  1893a,  b,  1894,  1898). 

Success  amplified  the  problems  beginning  in  1894  and  1895.  Attendance  had 
increased  dramatically,  from  15  in  1888  to  199  by  1895,  and  financing  had  become 
ever  more  difficult.  To  Whitman  expansion  seemed  desirable,  both  for  students  and 
for  researchers.  He  urged  more  building  in  1895  to  provide  more  room.  And  he 
continued  to  urge  the  need  for  solid  financial  backing.  It  looked,  in  1895,  as  though 
that  backing  might  materialize.  Miss  Helen  Culver  of  Chicago  evidently  intended 
to  give  a  half-million  dollars  each  to  the  MBL  and  to  the  University  of  Chicago's 
biology  program.  Whitman  headed  both,  and  he  worked  hard  to  obtain  the  dual 
gift.  A  letter  from  Whitman  to  Miss  Culver  clearly  indicates  that  part  of  the  gift 
was  originally  intended  for  the  MBL: 

The  Marine  Biological  Laboratory  has  already  become  an  intercollegiate 
centre  for  research  and  instruction.  Some  over  twenty  colleges  and  universities 
are  now  contributing  to  the  support  of  the  Laboratory  by  subscriptions  to  rooms 
and  tables,  and  no  less  than  eighty-five  institutions  were  represented  in  our 
membership  last  summer.  The  national  character  of  the  Laboratory  is  the  chief 
glory  and  that  I  am  sure  will  be  wisely  guarded  in  the  foundation  you  have 
bestowed. 

Instead,  for  unknown  reasons,  Miss  Culver  gave  the  entire  million  to  the  University 
and  none  to  the  MBL  (Whitman,  to  Miss  Helen  Culver,  20  December  1895).  The 
goals  for  the  MBL  remained  elusive. 

Whitman  continued  to  support  growth  and  expansion,  but  in  1896  the  Trustees 
as  a  whole  said,  in  effect,  "no  more."  Yet  the  Executive  Committee  of  the  Trustees 


194  J.   MAIENSCHEIN 

approved  the  expansion  and  expenditures.  With  Whitman's  personal  financial 
backing,  the  group  finally  achieved  agreement,  but  the  seed  of  crisis  had  been  sown 
(Lillie,  1944,  pp.  43-44;  Trustees,  1897;  Clarke  et  ai,  1897).  The  disagreement 
brought  the  first  confrontation,  with  a  major  split  in  the  Board  of  Trustees.  In  late 
1896  and  early  1897,  the  Trustees  met  and  agreed  to  keep  the  lab  open  only  if  they 
could  raise  $2000  to  cover  costs.  They  met  in  Boston,  and  they  did  not  even  consult 
Whitman.  Further,  they  postponed  announcing  the  1897  session  until  they  had  met 
and  made  financial  decisions  about  whether  to  continue;  the  uncertainty  cut  into 
that  summer's  attendance.  The  annual  meeting  at  Woods  Hole  in  August  1897  was 
very  tense,  with  major  disagreements  about  how  to  run  the  lab.  The  meeting 
brought  new  by-laws  and  election  of  new  Trustees,  with  only  two  of  the  original 
Boston  Trustees  remaining  on  the  Board.  As  Whitman  reported  to  Dohrn,  he  was 
particularly  delighted  to  have  gotten  rid  of  the  "old  maids"  from  the  Women's 
Educational  Association  who  had  no  real  connections  with  biology  (Whitman, 
Naples,  2  September  1897).  The  new  Board  reflected  greater  national  representation 
and  greater  biological  commitment. 

Yet  financial  troubles  continued  despite  the  greater  ideological  support  from  the 
Trustees.  In  1898,  Whitman  wrote  to  his  friend  and  supporter  Edwin  Grant  Conklin 
that  "I  am  having  many  sleepless  hours  over  the  lack  of  funds  to  pay  bills  this  year. 
I  have  about  resolved  to  take  from  my  own  poor  pocket  to  settle  the  $600  unsettled 
salaries.  I  have  reason  to  hesitate  to  do  this,  for  I  do  not  see  the  way  out  of  it.  Were 
it  not  for  the  many  good  hearts  behind  me,  I  should  feel  decidedly  blue"  (Whitman 
to  Conklin,  1  November  1898).  Indeed,  a  friend  reported  that  Whitman  had  suffered 
very  deeply  from  the  troubles  since  the  MBL  was  "the  very  apple  of  his  eye" 
(Whitman,  Chicago,  20  May  1898). 

Whitman  articulated  ever  more  strongly  that  the  United  States  needed  a 
biological  laboratory  and  that  it  should  be  a  permanent  station  with  a  full 
endowment.  Such  a  station  must  have  national  cooperation  and  must  therefore 
remain  financially  and  ideologically  independent  of  any  one  group.  Permanence, 
national  support,  cooperation,  and  independence — these  became  recurrent  themes 
for  Whitman  (Whitman,  Annual  Report,  1890,  pp.  22-23;  Annual  Report,  1892, 
pp.  29-36;  1893;  1901;  to  Morgan  or  Wilson,  1902).  Without  money,  those  ideals 
remained  out  of  reach.  At  one  point,  however,  the  financial  goal  seemed  nearly 
attainable. 

In  1900  came  a  concerted  effort  to  achieve  wider  national  support  from  colleges 
to  underwrite  at  least  the  cost  of  operations.  Following  the  Naples  model,  the  MBL 
did  not  try,  in  the  early  years,  to  extract  more  money  from  the  individual  researchers 
but  appealed  instead  to  institutions  for  more  permanent  support.  Some,  such  as 
Alexander  Agassiz,  who  had  supported  Naples  and  other  efforts  felt  they  had 
"thrown  away  enough  money  on  seaside  Laboratories"  and  balked  at  donating 
money  to  yet  another  attempt  to  build  an  American  laboratory  (Agassiz,  30  May 
1888).  Other  colleges  and  institutions  did  continue  to  provide  support.  Finally  in 
1901  and  1902  two  major  offers  came  to  relieve  the  Trustees  of  the  bulk  of  their 
financial  problems.  The  first  came  from  four  wealthy  businessmen  and  was  presented 
through  President  Harper  of  the  University  of  Chicago.  Whitman  strongly  supported 
their  proposal  and  felt  that  it  would  secure  the  laboratory's  financial  independence 
and  make  realistic  the  establishment  of  a  permanent  research  station  (Whitman  to 
Conklin,  1901  and  1902).  Yet,  though  only  two  of  the  four  men  lived  in  Chicago, 
the  Trustees  felt  that  accepting  the  offer  would  give  too  much  power  to  one 
university  and  that  the  lab  would  therefore  lose  the  very  independence  and  national 
character  it  sought.  Whitman  felt  unsupported  in  the  ensuing  sometimes  bitter 


EARLY   STRUGGLES  195 

struggles,  and  he  lamented  to  Conklin  that  though  he  felt  confident  that  things 
would  work  out,  presumably  in  favor  of  the  plan,  "I  often  regret  that  there  has  been 
such  a  strong  sectional  feeling  in  the  East.  It  is  not  very  pleasant  to  have  ones 
motives  impugned,  and  I  confess,  at  times,  to  have  found  the  suspicion  against 
Chicago  University  and  its  men  almost  beyond  endurance"  (Whitman  to  Conklin, 
2  March  1902).  The  fate  of  the  Chicago  plan  remained  unclear  until  a  second  offer 
came  shortly  thereafter  leading  to  the  rejection  of  the  first  proposal  as  such. 

The  second  offer  came  from  the  Carnegie  Institution  and  went  through  numerous 
revisions.  The  group  most  strongly  supporting  the  Carnegie  proposal  included 
Wilson,  who  saw  this  as  the  chance  to  secure  financial  stability  and  to  make  over 
the  MBL  into  a  research  lab  more  like  his  old  ideal  at  Naples  (Cattell,  1902,  pp. 
529-533;  Lillie,  1944,  p.  57;  Whitman,  MBL-Lillie,  8  and  13  October  1902).  Wilson 
came  into  direct  conflict  with  Whitman  on  this  issue.  It  became  clear  to  Whitman 
that  while  some  of  the  Trustees,  such  as  Conklin,  remained  firmly  behind  him, 
most  had  never  really  fully  accepted  his  ideas  for  the  lab.  As  Frank  Lillie  recorded 
later  in  his  history.  Whitman  found  it  disillusioning  to  realize  that  there  were  "few 
who  held  with  anything  like  equal  intensity  his  belief  that  the  ideals  of  organization 
for  which  he  had  fought  were  of  value  far  superior  to  any  degree  of  financial 
security"'  (Lillie,  1944,  p.  60).  Whitman  feared  that  the  Carnegie  people  would 
absorb  the  MBL  as  just  another  one  of  their  own  departments.  He  feared  the 
proposed  move  to  solely  research  and  the  loss  of  instruction  which  Wilson 
applauded.  He  suggested  somewhat  facetiously  to  Conklin  that  "Perhaps  we  had 
better  abandon  every  class  at  Woods"  Holl,  and  all  compensation  for  services,  and 
revert  to  the  old-time  ideal  of  a  pure  research  station.  I  feel  half  inclined  to  do  this, 
and  so  let  everyone  see  by  actual  experience  the  result"  (Whitman  to  Conklin,  7 
January  1904).  Obviously,  he  did  not  expect  happy  results. 

Eventually,  after  much  discussion  and  revision,  a  plan  was  developed  by  which 
the  Carnegie  Foundation  supported  the  MBL  for  three  years  with  $10,000  per  year, 
matched  by  a  gift  from  three  of  the  Trustees — including  some  of  the  support 
offered  in  the  first  Chicago  proposal — but  the  MBL  Trustees  and  Corporation 
retained  control  of  the  lab.  As  an  admirer  put  it  later.  Whitman  had  held  firm  for 
independence. 

Reportedly  has  Woods  Hole  declined  riches  when  by  its  acceptance  there  is 
only  the  remotest  possibility  of  interference  with  this  indispensible  independence. 
All  of  the  proffers  fell  upon  deaf  ears.  The  nightingale  may  be  captured,  but  it 
can  never  be  made  to  breed  by  the  huntsman  nor  be  made  to  sing  in  confinement. 
It  must  live  in  its  own  peculiar  habitat,  and  this  is  found  for  scientists  in  Woods 
Hole.  In  this  country  we  are  searching  for  heroes  in  productive  science,  but  "the 
birds  that  may  sing  may  seem  to  avoid  the  golden  cage"  [The  Resignation,  1908, 
p.  382]. 

Nonetheless,  Whitman  felt  defeated  and  exhausted  and  withdrew  from  the  MBL. 
His  assistant,  Frank  Lillie,  took  over  the  directorship  (MBL  Annual  Report,  1908, 
pp.  8-13). 

Whitman  looked  upon  the  struggles  of  1902  as  growing  pains  of  sorts,  in  which 
the  impatience  of  some  had  led  to  near  disaster.  As  he  put  it,  the  MBL  had  begun 
like  an  organism,  with  only  seventeen  "ids  in  its  protoplasmic  body — two  instructors, 
eight  students,  and  seven  investigators  (all  beginners).  The  two  instructors  could  be 
likened,  with  no  great  stretch  of  the  imagination,  to  two  polar  corpuscles,  signifying 
little  more  than  that  the  germ  was  a  fertile  one,  and  prepared  to  begin  its  preordained 
course  of  development."'  The  original  incorporators.  Whitman  said,  served  as 


196  J.   MAIENSCHEIN 

sponsors  and  left  the  group  of  ids  to  follow  its  own  course  of  development.  The 
germ  thus  underwent  various  cleavages  and  took  shape.  Founded  on  the  principles 
of  cooperation  and  independence,  it  sought  to  embrace  all  of  biology.  It  even  grew 
and  advanced  to  the  tadpole  stage.  Whitman  reported.  But  some  members  wanted 
to  shed  their  tails  and  become  frogs,  and  to  undergo  that  change  immediately.  They 
forgot  the  golden  motto  of  development,  proceed  slowly.  Fortunately,  the  supporters 
made  it  possible  for  the  tadpoles  to  advance  without  loss  of  their  heads  as  well  as 
their  tails.  But  do  not  forget  the  lesson.  Whitman  urged,  for  "There  is  a  work  before 
you  of  far  greater  magnitude  and  importance  than  perhaps  any  of  us  can  now 
realize,  waiting  only  for  the  energy  and  means  to  grapple  with  it.  In  everything  that 
stands  for  the  upbuilding  of  this  laboratory,  let  us  have  cooperation  with  soul  and 
zeal  to  make  it  effective  and  triumphant"  (Whitman,  address,  11  August  1903). 
Clearly,  Whitman  did  not  fully  agree  with  Wilson  that  the  success  of  the  Naples 
Station  lay  "simply"  with  money. 

Permanence,  national  support,  cooperation,  and  independence  have  been  achieved 
at  the  MBL  to  a  remarkable  extent.  Yet  as  Whitman  stressed,  both  money  and  a 
great  deal  of  dedicated  work  by  a  large  number  of  people  remain  necessary  to 
maintain  such  enduring  marine  laboratories  as  the  MBL  and  the  Naples  Station. 

ACKNOWLEDGMENTS 

I  wish  to  thank  Ruth  Davis  at  the  MBL,  Ann  Blum  at  the  Museum  of 
Comparative  Zoology,  Christiane  Groeben  at  Naples,  and  the  archivists  at  the  Johns 
Hopkins  University  Archives  and  the  University  of  Chicago  for  their  assistance.  All 
archival  material  quoted  with  permission. 

LITERATURE  CITED 

AGASSIZ,  ALEXANDER.  Letter,  30  May  1888,  Agassiz  Papers.  Museum  of  Comparative  Zoology  Archives, 

Harvard  University. 
CLAPP,  CORNELIA.  1927.  Some  recollections  of  the  first  summer  at  Woods  Hole.  1888.  Collecting  Net 

2(4):  3,  10. 

CLARKE,  GARDINER,  AND  MCMURRICH.  1897.  Reply.  Science  6:  475-476. 
LILLIE,  FRANK.    1944.   The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press, 

Chicago. 

MBL  Annual  Reports. 

MBL  Minutes  of  the  Trustees,  MBL  Archives. 
MBL  Board  of  Trustees.  1897.  A  statement  concerning  the  Marine  Biological  Laboratory.  Science  (1897): 

529-534. 
The  resignation  of  Prof.  Whitman  as  Director  of  the  Marine  Biological  Laboratory  at  Woods  Hole,  Mass. 

Anal.  Rec.  (1908)  2:  380-382. 

WHITMAN,  CHARLES  OTIS.  1890,  1892.  Report  of  the  Director.  Annual  Reports. 
WHITMAN,  CHARLES  OTIS.  1891.  Specialization  and  organization.  Biological  Lectures.  1890:  1-26. 
WHITMAN,  CHARLES  OTIS.  1893.  A  Marine  Biological  Observatory.  Pop.  Sci.  Mo.  42:  459-571. 
WHITMAN,  CHARLES  OTIS.  1893.  General  physiology  and  its  relation  to  morphology.  Am.  Nat.  27:  802- 

807. 
WHITMAN,  CHARLES  OTIS.  1894.  The  work  and  the  aims  of  the  Marine  Biological  Laboratory.  Biological 

Lectures  1893:  235-242. 
WHITMAN,  CHARLES  OTIS.  1898.  Some  of  the  functions  and  features  of  a  biological  station.  Science  7: 

37-44. 
WHITMAN,  CHARLES  OTIS.  Letters  to  Conklin,  Morgan,  Wilson,  Miss  Helen  Culver,  Whitman  Papers. 

MBL  Archives. 

WHITMAN,  CHARLES  OTIS.  Letters  to  Lillie,  Lillie  Papers.  MBL  Archives. 
WHITMAN,  CHARLES  OTIS.  20  May  1898.  Whitman  Papers,  University  of  Chicago  Archives. 
WHITMAN,  CHARLES  OTIS.  Address  to  the  Corporation,  1901.  Whitman  Papers.  MBL  Archives. 
WHITMAN,  CHARLES  OTIS.  Address  to  the  Corporation,  1903.  Whitman  Papers.  MBL  Archives. 
WHITMAN,   from   Dohrn,  2  September   1897.   Whitman   Papers.   MBL  Archives.  Original   in  Naples 

Zoological  Station  archives  and  provided  for  MBL  by  Christiane  Groeben. 


Reference:  Biol.  Bull.  168  (suppl.):  197-199.  (June,  1985) 


THE  WOODS   HOLE   LABORATORY   SITE: 
HISTORY  AND  FUTURE   ECOLOGY 

W.   D.   RUSSELL-HUNTER 

Marine  Biological  Laboratory,  Woods  Hole.  Massachusetts,  02543,  and  Department  of  Biology, 
Syracuse  University,  Syracuse,  New  York,  13210 

ABSTRACT 

The  Woods  Hole  site  of  the  Marine  Biological  Laboratory  is  adjacent  to  that 
chosen  for  work  of  the  U.  S.  Fish  Commission  by  Spencer  F.  Baird  in  1871.  In 
most  ecological  respects,  it  remains  an  excellent  locus  for  marine  research,  with 
strong  tidal  currents  ensuring  a  supply  of  pure  sea  water  and  with  a  reasonably 
diverse  fauna  and  flora.  Despite  local  environmental  risks,  there  are  grounds  for 
cautious  optimism  that  both  Woods  Hole  and  Naples  (Ischia  laboratory)  can 
continue  to  provide  opportunities  for  all  kinds  of  biological  research  utilizing  healthy 
marine  organisms. 

INTRODUCTION  AND  HISTORY 

The  site  of  the  Marine  Biological  Laboratory  at  Woods  Hole  is,  in  most 
ecological  respects,  a  good  one  well-buffered  by  flowing  sea  water.  In  particular,  it 
is  excellently  placed  in  relation  both  to  the  major  oceanic  circulation  (Lillie,  1944) 
and  to  local  tidal  currents.  It  was  chosen  by  Spencer  F.  Baird  (possibly  as  early  as 
1863)  as  a  site  for  work  of  the  U.  S.  Fish  Commission,  and  he  began  studies  there 
in  1871  and  1875  (Conklin,  1944).  The  fisheries  laboratory  was  built  in  1885,  and 
the  adjacent  MBL  in  1888.  On  a  narrow  promontory  between  Buzzards  Bay  and 
Vineyard  Sound,  placed  at  an  angle  of  the  larger  peninsula  of  Cape  Cod,  there  are 
no  large  rivers  near  by,  and  Baird  noted  the  purity  of  the  local  sea  water,  free  from 
sediment  or  sewage  contamination  (Conklin,  1944). 

FAUNAL  ASPECTS 

The  waters  off  Cape  Cod  are  affected  by  the  Labrador  Current  to  the  north  and 
the  Gulf  Stream  to  the  south,  and  there  are  corresponding  differences  in  at  least 
25%  of  the  benthic  fauna  and  rather  more  of  the  macrophytic  algae.  Thus  we  have 
both  a  cold-water  Strongylocentrotus  and  a  warm- water  Arbacia  available,  providing 
ripe  eggs  in  spring  and  in  summer,  respectively.  In  one  respect,  the  intertidal  and 
sublittoral  faunas  are  somewhat  impoverished.  Since  Cape  Cod  is  entirely  made  up 
of  glacial  moraine  there  are  no  extensive  rock  outcrops,  and  so  we  lack  Octopus, 
rock-boring  bivalves,  and  certain  species  of  hydroids,  serpulids,  barnacles,  and 
opisthobranchs  (including  most  species  both  of  the  beautiful  nudibranchs  and  of 
the  larger  aplysiomorphs  used  in  neural  systems  investigations).  Parenthetically,  it 
is  clear  that  the  different  cephalopods  available  at  the  Stazione  Zoologica  (Octopus) 
and  at  the  MBL  (Loligo)  led  to  markedly  different  emphases  in  neurobiological 
research  over  the  last  45  years.  The  neural  basis  for  capacity  to  learn  or  be  trained, 
including  controls  utilizing  split-brain  techniques,  were  investigated  at  Naples,  while 
the  biophysics  of  neural  membranes  and  chemical  properties  of  the  giant  axon  were 
investigated  at  Woods  Hole. 

197 


198  W.   D.   RUSSELL-HUNTER 

Periodically,  around  Cape  Cod,  local  grounds  for  squid,  sea  urchins,  and  certain 
fish  and  bivalves  have  been  over-collected,  but  no  faunal  extinction  can  be  attributed 
to  MBL  collectors.  Chaetopterus  may  be  the  one  locally  collector-endangered  genus 
at  present  (Valois,  1984).  Recently,  there  has  been  some  competition  from  commercial 
fishermen  for  squid,  Spisula,  and  a  few  other  forms.  Lack  of  detailed  knowledge  of 
natural  population  dynamics  (even  demography)  for  these  species  makes  it  difficult 
to  predict  the  effects  of  such  sustained  commercial  cropping.  Over  the  last  century, 
the  most  extensive  ecological  changes  in  the  area  were  the  disease-caused  decline  of 
eelgrass  (Zostera)  beds  between  1930  and  1960,  and  their  recovery  during  the  last 
20  years.  It  seems  probable  that  both  a  fungus,  Ophiobolus  halimus,  and  a  slime- 
mold,  Labyrinthula  macrocystis,  were  important  in  this  epidemic  on  both  sides  of 
the  North  Atlantic,  although  higher  water  temperatures  may  also  have  been  involved. 
An  excellent  summary  of  the  literature  along  with  substantive  data  from  Danish 
waters  is  provided  by  Rasmussen  (1973).  At  Woods  Hole  as  elsewhere,  associated 
communities  of  invertebrates,  including  Cumingia  (Russell-Hunter  and  Tashiro, 
1985),  became  very  rare  for  three  decades  and  then  required  a  further  20  years  to 
approach  their  pre-1930  levels  of  abundance.  No  human  pollution  or  other 
environmental  modification  has  ever  caused  such  widespread  and  long-lasting 
ecological  changes. 

WATER  QUALITY 

Tidal  currents  run  through  the  Woods  Hole  passage  (between  the  Cape  mainland 
and  the  Elizabeth  Islands)  reaching  average  maximum  velocities  four  times  daily, 
westwards  of  3.6  knots  and  eastwards  of  4.5  knots  (White,  1984),  and  running  at 
<0.8  knots  for  only  2.8  hours  in  each  day  [1  knot  =  1.853  km/h].  Practicable 
extensions  of  the  present  sea  water  intake  system  of  MBL  into  this  tidal  flow  could 
protect  pure  sea  water  supply  for  the  foreseeable  future.  A  similar  continuous  supply 
of  unpolluted  sea  water  is  ensured  at  the  ecological  laboratory  (Bacci,  1969)  of  the 
Stazione  Zoologica  at  Punta  San  Pietro  on  Ischia.  Despite  increasing  residential 
building  on  Cape  Cod,  sewage  contamination  of  MBL  sea  water  is  not  a  problem 
and  the  1984-1986  sewer  development  for  the  Falmouth  area  further  reduces  the 
risk.  In  the  near  future,  a  more  likely  difficulty  at  Woods  Hole  could  be  ensuring 
an  adequate  supply  of  unpolluted  fresh  water  (as  has  long  been  the  case  on  Ischia). 

Historically,  both  Woods  Hole  and  Naples  have  been  good  sites  for  research  on 
healthy  marine  organisms.  The  majority  of  the  invertebrate  species  surveyed  in 
1871  by  Spencer  Baird's  associate  A.  E.  Verrill  in  the  Vineyard  Sound  area  are  still 
there  today.  Similarly,  Tomas  (1984)  could  state  that  the  flora  and  fauna  of  the 
sublittoral  around  Ischia  were  relatively  unchanged  from  the  time  of  Anton  Dohrn. 
These  historical  continuities  in  natural  populations  provide  grounds  for  cautious 
optimism  regarding  the  future  ecology  of  both  areas.  Despite  this,  we  must  increase 
our  knowledge  of  natural  life-cycles,  demography,  and  ecology  in  the  species  we 
collect,  and  be  prepared  (for  example,  in  the  proposed  Marine  Resources  Center  at 
MBL)  to  move  toward  mariculture  where  appropriate.  For  the  Cape  Cod  area,  the 
only  long-term  environmental  risk  lies  in  the  exploration  for,  and  possible  devel- 
opment of,  offshore  oil  deposits  on  Georges  Bank,  but  Woods  Hole  is  somewhat 
protected  by  position  and  prevailing  winds.  Oil  spills  from  vessels,  even  small  ones, 
could  affect  many  fragile  habitats  such  as  high  salt  marshes,  but  supertankers  do 
not  use  Vineyard  Sound  or  Buzzards  Bay.  Both  cautious  optimism  for  the  future, 
and  a  sense  of  gratitude  to  Spencer  F.  Baird,  Alpheus  Hyatt,  C.  O.  Whitman,  and 
the  others  for  picking  the  site,  seem  appropriate  at  this  time. 


SITE  ECOLOGY,  WOODS  HOLE  199 

ACKNOWLEDGMENTS 

Along  with  my  main  charge  to  survey  the  history  of  invertebrate  teaching  at 
Woods  Hole,  Paul  R.  Gross  and  Seymour  S.  Cohen  suggested  that  I  prepare  for 
discussion  at  the  Ischia  symposium  this  brief  statement  on  the  ecological  setting 
and  future  of  the  MBL.  I  am  grateful  both  to  them  and  to  Carmelo  R.  Tomas,  my 
fellow-discussant  from  the  Stazione  Zoologica.  Other  acknowledgments  are  set  out 
in  my  main  paper  (p.  88  of  this  issue). 

LITERATURE  CITED 

BACCI,  G.  1 969.  A  future  for  ecological  research  at  the  Zoological  Station  of  Naples.  Puhhl.  Staz.  Zool. 

Napoli  31:  7-15. 

CONKLIN,  E.  G.  1944.  III.  The  United  States  Bureau  of  Fisheries.  Pp.  24-26  in  Lillie  (1944). 
LILLIE,  F.  R.  1944.  The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press,  Chicago. 

284  pp. 
RASMUSSEN,  E.  1973.  Systematics  and  ecology  of  the  Isefjord  marine  fauna  (Denmark)  with  a  survey  of 

the  eelgrass  (Zostera)  vegetation  and  its  communities.  Ophelia  11:  1-495. 
RUSSELL-HUNTER,  W.  D.  AND  J.  S.  TASHIRO.    1985.  Life-habits  and  infaunal  posture  of  Cumingia 

tellinoides  (TELLINACEA,  Semelidae):  an  example  of  evolutionary  parallelism.  Veliger  27:  253- 

260. 

TOMAS,  C.  R.  1984.  Reply  during  discussion.  Ischia  Symposium,  8-12  October  1984. 
VALOIS,  J.  J.   1984.  Personal  communication  to  author.  Marine  Biological  Laboratory,   Woods  Hole, 

August,  1984. 
WHITE,  R.  E.   1984.  Eldridge  Tide  and  Pilot  Book,   1984.  Robert  Eldridge  White,  Publisher,  Boston. 

272  pp. 


Reference:  Bin/.  Bull.  168  (suppl.):  200-202.  (June,  1985) 


FROM   WOODS   HOLE  TO  THE  WORLD: 
THE  BIOLOGICAL  BULLETIN 

W.   D.   RUSSELL-HUNTER 

Marine  Biological  Laboratory,  Woods  Hole,  Massachusetts,  02543,  and  Department  of  Biology, 
Syracuse  University,  Syracuse,  New  York.  13210 

ABSTRACT 

The  Marine  Biological  Laboratory,  Woods  Hole,  publishes  The  Biological 
Bulletin:  a  general  research  journal  with  an  international  circulation.  It  has  been 
published  continuously  since  1902  (although  its  predecessors  date  from  1897),  and 
there  have  been  only  eight  editors.  Characterized  by  its  regularity  of  publication, 
diversity  of  contents,  and  editorial  independence,  it  has  not  been  a  "house  journal" 
for  at  least  55  years.  Changes  in  its  editorial  policies  have  historically  been  minor. 
The  MBL  library  has  continued  to  benefit  from  approximately  650  serials  received 
as  "free"  exchanges  for  the  journal.  For  future  historical  research,  an  extensive 
archive  of  editorial  correspondence  will  be  provided. 

INTRODUCTION 

The  journal,  The  Biological  Bulletin,  is  a  little  younger  than  the  laboratory  at 
Woods  Hole,  but  both  are  owned  by  the  corporate  membership  of  the  Marine 
Biological  Laboratory.  To  many  members,  unable  to  work  at  the  laboratory,  the 
conservatively  printed  grey  cover  contains  the  only  tangible  return  for  their 
continuing  membership  dues.  Since  1902,  with  only  minor  changes  of  format  but 
with  a  great  diversity  of  contents,  two  volumes  or  six  issues  per  year  have  appeared 
regularly.  This  continuity  as  a  nonspecialist  research  journal  has  sustained  library 
subscriptions  both  domestic  and  foreign,  and  also  journal  exchanges  with  learned 
societies  and  institutions  throughout  the  world.  Two  examples  of  our  cosmopolitan 
circulation  are  revealed  by  our  inclusion  in  the  citation  and  abstracting  services  run 
by  the  Polish  Academy  of  Sciences  in  Warsaw,  and  the  Indian  Agricultural  Research 
Council  in  New  Delhi,  both  based  on  a  precirculated  contents  proof  for  each  issue. 
The  age  of  the  journal  and  its  regularity  of  publication  contribute  to  its  worldwide 
circulation. 

HISTORY 

The  general  history  of  the  MBL  by  Lillie  (1944)  is  surprisingly  unspecific  about 
the  journal,  but  Redfield  (1941)  provides  an  excellent  review  of  its  early  days.  Two 
volumes  per  year  have  been  published  continuously  for  82  years  (since  the 
resumption  of  publication  with  volume  3  in  Fall,  1902);  although  its  predecessors 
date  from  1897.  Edited  by  the  Director  (C.  O.  Whitman)  and  members  of  the  staff, 
two  volumes  of  the  Zoological  Bulletin  were  published  in  1897  and  1898,  and  the 
first  two  volumes  of  its  successor  The  Biological  Bulletin  in  1900  and  early  1901. 
Frank  R.  Lillie  became  Managing  Editor  with  volume  3  in  Fall  1902,  and  his 
editorial  staff  was  listed  as  Conklin,  Loeb,  Morgan,  Wheeler,  Whitman,  and  Wilson. 
Lillie  remained  editor  until  1927,  and  was  succeeded  in  turn  by:  Carl  R.  Moore 
(1927-1930),  Alfred  C.  Redfield  (1930-1942),  H.  Burr  Steinbach  (1942-1950), 

200 


BIOLOGICAL  BULLETIN  HISTORY  201 

Donald  P.  Costello  (1951-1968),  W.  D.  Russell-Hunter  (1968-1980),  and  Charles 
B.  Metz  (1980-  ).  There  have  been  no  supplementary  publications,  and  the 
history  of  publication  of  a  single  general  journal  from  the  MBL  obviously  differs 
from  that  of  journal  publishing  at  the  Stazione  Zoologica. 

At  Naples,  three  different  serials  were  begun  in  1879-80:  an  abstracting  annual, 
a  house  journal,  and  a  series  of  systematic  monographs.  Publication  of  the 
Zoologischer  Jahresbericht  helped  the  library  of  the  Stazione,  which  received  all  the 
literature  sent  in  for  review.  The  journal  publishing  in-house  work  was  the 
Mittheilungen  aus  der  Zoologischen  Station  zu  Neapel  continued  after  1916  as 
Pubblicazioni  delta  Stazione  Zoologica  di  Napoli,  which  has  been  split  into  two 
since  1979;  Marine  Ecology  (Pubbl.  Staz.  Zool.  Napoli  I)  and  History  and  Philosophy 
of  the  Life  Sciences  (Pubbl.  Staz.  Zool.  Napoli  II).  The  magnificent  monographic 
series,  Fauna  und  Flora  des  Golfes  von  Neapel,  has  no  Woods  Hole  equivalent. 

SCOPE  AND  POLICY 

The  Biological  Bulletin  has  always  been  a  general  rather  than  a  specialist  journal, 
publishing  original  research  reports  of  intermediate  length.  It  was  first  intended  to 
be  a  companion  journal  to  the  Journal  of  Morphology  providing  for  relatively  rapid 
publication  of  snorter  papers  with  simpler  illustrations.  From  1912  to  1929,  it 
steadily  became  less  of  a  "house  journal."  Since  1930,  its  editorial  policies  and 
reviewing  procedures  have  been  deliberately  kept  separate  from  both  the  adminis- 
tration and  the  elected  trustees  of  MBL.  Up  to  1980,  the  managing  editor  was 
unpaid,  and  there  was  usually  a  single  full-time  editorial  assistant  (and  some  part- 
time  help  managing  subscriptions).  Home  institutions  of  editors  tolerated  the 
demands  (both  in  space  and  time)  generated  by  their  work  as  midwives  for  the 
scholarship  of  others. 

To  librarians  and  exchanging  institutions,  it  remained  associated  with  that  select 
group  of  general  journals  of  certain  national  academies  and  royal  societies,  charac- 
terized by  Vannevar  Bush  as  of  greatest  significance  in  the  "invisible  university"  of 
world  science.  Frequently,  new  authors  were  surprised  by  contacts  generated  through 
its  readership  in  Eastern  Europe  and  in  Asia.  The  MBL  library  has  always  benefitted 
from  the  650  or  so  serials  generated  as  "free"  exchanges,  and  continuity  both  of 
these  and  of  regular  subscriptions  has  been  influenced  by  regularity  of  publication 
and  diversity  of  contents.  Maintaining  this  diversity  was  the  declared  strategy  of  five 
of  the  editors  and,  in  retrospect,  changes  in  editorial  policy  were  relatively  trivial. 
A  few  good  systematic  papers  were  published  in  the  periods  1930-1942  and  1968- 
1980,  while  taxonomy  and  systematics  were  deliberately  excluded  in  1912-1930 
and  in  1951-1968.  Similarly,  review  papers  were  excluded  from  1925-1980  although 
some  were  published  before  1925  and  after  1980.  Abstracts  of  each  summer's 
general  meetings  at  MBL  have  been  published  for  over  50  years,  but  the  regular 
papers  have  more  widespread  origins.  In  the  period  1950-1980,  19%  of  the  papers 
published  reported  work  done  wholly  or  partly  at  MBL,  58%  reported  research 
conducted  elsewhere  in  the  U.  S.,  and  23%  had  other  international  origins. 

ARCHIVES 

For  future  historians,  an  extensive  archive  will  be  provided  by  the  files  of 
completed  editorial  correspondence  (including  those  for  rejected  papers)  which  have 
been  preserved  nearly  completely  from  1945,  and  partially  from  1930.  We  have 
proposed  that  these  files  remain  in  their  normal  restricted  use  for  reference  in  the 


202  W.   D.   RUSSELL-HUNTER 

editorial  office  for  6  years,  and  then  be  "closed"  for  a  further  20  years,  before 
becoming  available  for  approved  historical  research. 

ACKNOWLEDGMENT 

Along  with  my  main  charge  to  survey  the  history  of  invertebrate  teaching  at 
Woods  Hole,  Seymour  S.  Cohen  suggested  I  prepare  a  brief  historical  statement  on 
The  Biological  Bulletin  for  discussion  at  the  Ischia  symposium.  I  am  grateful  to 
him.  Other  acknowledgments  are  set  out  in  my  main  paper  (p.  88  of  this  issue). 

LITERATURE   CITED 

LILLIE,  F.  R.  1944.  The  Woods  Hole  Marine  Biological  Laboratory.  University  of  Chicago  Press,  Chicago. 

284  pp. 
REDFIELD,  A.  C.   1941.  The  report  of  the  managing  editor  of  the  Biological  Bulletin.  Biol.  Bull.  81: 

12-17. 


Reference:  Biol.  Bull.  168  (suppl.):  203-204.  (June,  1985) 


WHAT   LABORATORIES   FOR   WHAT  SCIENCE? 

ANTONIO   MIRALTO 

Stazione  Zoologica.  I'illa  Comunale,  1-80121,  Naples.  Italy 

Having  examined  the  roles  that  the  Zoological  Station  and  the  Marine  Biological 
Laboratory  have  played  in  shaping  ideas  in  biology  since  the  turn  of  the  century,  it 
now  seems  appropriate  to  address  the  question  of  the  future  of  marine  biological 
laboratories,  and,  in  fact,  of  biological  research  institutes  in  general.  However,  we 
should  first  determine  the  kind  of  science  we  want  for  the  future.  I  believe  that 
science  should  fit  within  a  broad,  non-restrictive,  non-deterministic  cultural  frame- 
work. Indeed,  science  should  become  the  focal  point  of  a  process  of  cultural  renewal; 
a  point  of  reference  for  future  generations;  and  a  driving  force  in  all  fields,  scientific 
and  non. 

From  many  views,  scientists  of  the  16th,  17th,  and  18th  centuries  had  more 
freedom  than  present-day  scientists.  Today  researchers  are  no  longer  restricted  by 
Church  dogma,  however  governments,  financial  trusts,  and  military  and  economic 
demands  impose  more  sophisticated  and  subtle  pressures,  which  are  totally  unrelated 
to  the  needs  of  humanity. 

This  is  not  a  plea  to  turn  back  the  clock,  but  we  must  study  the  past  to 
understand  where  we  came  from  and  how  important  achievements  evolved  to 
discover  the  roots  that  may  nourish  our  future.  The  past  can  also  help  us  to  avoid 
repeating  mistakes  and  to  outline  a  future  in  which  we  are  aware  of  the  difficulties 
and  responsibilities  it  entails. 

There  is  obviously  no  easy  remedy  for  all  the  problems  that  will  arise.  However, 
any  new  solutions  or  ideas  will  probably  come  from  institutions  that  promote  the 
growth  of  knowledge,  that  is,  our  schools  and  universities. 

In  the  past,  universities  provided  a  broad,  philosophical,  humanistic,  and 
scientific  education,  and  were  not  as  specialized  as  today.  Specialization  on  the 
whole  is  acceptable,  but  it  should  not  confine  a  person  to  a  narrow  corridor.  New 
generations  should  be  educated  to  be  creative  in  solving  the  problems  that  will 
confront  them.  New  researchers  should  be  trained  to  have  a  dialectical  attitude. 
After  all,  nature  itself  is  dialectical. 

Schools  and  universities  must  return  to  being  forums  where  ideas  are  questioned 
and  discussed,  and  they  must  produce  people  capable  of  using  the  most  important 
tools  available  to  mankind:  culture  and  knowledge.  I  believe  in  a  science  that  does 
not  bind  its  fate  to  technology,  but  to  the  growth  of  knowledge. 

After  having  presented  an  idea  of  what  I  believe  is  the  cultural  framework  within 
which  research  should  operate,  I  shall  now  turn  to  the  organization  of  scientific 
institutes. 

First  of  all,  laboratories  should  not  become  isolated  and  closed  structures;  this 
would  be  detrimental  to  the  quality  of  research  and  to  researchers.  This  means 
encouraging  international  exchange  between  research  laboratories  and  other  cultural 
institutions. 

I  envisage  research  institutes  as  helping  to  define  the  cultural  and  material 
requirements  necessary  to  improve  the  quality  of  life.  Laboratories  should  not  be 
limited  to  research  but  should  stimulate  the  discussion  of  scientific  and  cultural 
topics  in  general.  It  is  no  exaggeration  to  say  that  many  researchers  have  excellent 

203 


204  A.   MIRALTO 

technical  know-how,  but  lack  a  solid  cultural  background.  Hence,  the  importance 
of  non-specialized  cultural  activities  such  as  symposia,  workshops,  and  courses.  By 
expanding  the  researcher's  frontiers  of  knowledge,  the  quality  of  his  research  will 
improve. 

Another  aspect  to  be  considered  is  the  organization  of  research  laboratories.  To 
be  efficient,  an  organization  requires  adequate  funds  and  personnel,  and  well-defined 
programs.  Furthermore,  research  groups  consisting  of  scientists  with  different 
working  experience  should  be  encouraged  so  that  scientific  problems  can  be 
approached  as  impartially  as  possible.  Scientists  must  also  be  morally,  ethically,  and 
financially  stimulated. 

We  should  also  favor  the  development  of  science  world-wide  to  counterbalance 
the  tendency  to  reduce  personnel.  This  brings  me  to  funding  for  science.  At  present, 
financial  backing  is  insufficient;  scientists  should  demand  that  government  funds 
now  invested  in  armaments  aimed  at  destroying  life  be  employed  for  research  aimed 
at  improving  the  quality  of  life. 

Lastly,  research  institutes  should  be  known  for  their  eagerness  to  learn,  their 
creativity,  and  their  ability  to  exploit  all  available  intellectual  energy.  All  this  in 
turn  should  be  directed  towards  meeting  the  various  needs  of  humanity. 

My  comments  so  far  concern  not  only  the  better  functioning  of  research 
laboratories,  but  also  offer  guidelines  aimed  at  improving  the  quality  of  life.  It  is 
my  hope  that  science  will  contribute  to  arresting  the  advance  of  what  I  consider  the 
barbaric  tendency  of  focusing  on  material  wealth  and  consumer  items,  rather  than 
on  the  cultural  and  moral  problems  of  modern  man. 

As  in  the  past,  science  can  determine  many  aspects  of  the  future  of  humanity. 
Men  of  science  must  look  beyond  the  limits  of  their  own  research  activities  and,  by 
their  culture  and  enlightenment,  become  intellectual  leaders. 


INDEX 


"New"  embryology  at  the  Zoological  Station  and 
at  the  Marine  Biological  Laboratory,  The,  35 

AGASSIZ,  Louis,  26,  88 

Aggassiz,  Hyatt,  Whitman,  and  the  birth  of  the 
Marine  Biological  Laboratory,  26 

ALEXANDROWICZ,  J.  C,  137 

ALLEN,  GARLAND  E.,  Heredity  under  an  embryo- 
logical  paradigm:  the  case  of  genetics  and 
embryology,  107 

American  biologists  at  Naples,  187 

Anton  Dohrn — the  statesman  of  Darwinism,  4 

VON  APATHY,  STEPHAN,  137 

Archives,  MBL,  200 


B 


BAIRD,  SPENCER  F.,  26,  197 

BENNETT,  M.  V.  L.,  Nicked  by  Occam's  razor: 

unitarianism  in  the  investigation  of  synaptic 

transmission,  159 
BETHE,  ALBRECHT,  137 
Biochemistry,  Naples,  122 
Biological  Bulletin,  The,  history,  200 
Birth  of  the  MBL,  26 
BODANSKY,  JOEL  N.,  see  Nathan  Reingold,  44 


Carnegie  Department  of  Marine  Biology,  Tortugas, 
172 

Carnegie  Institution  of  Washington  and  marine 
biology:  Naples.  Woods  Hole,  and  Tortugas, 
172 

Carnegie  and  Naples,  172 

Carnegie  and  Tortugas,  172 

Carnegie  and  Woods  Hole,  172 

Carnegie  Institution,  172,  192 

Cell  biology  and  heredity,  99 

Cell  interactions:  the  roots  of  a  century  of  research, 
80 

Cell  lineage,  26,  35,  62 

Cell  structure,  127 

Cephalopods  and  neuroscience,  153 

Chemical  transmission,  159 

CLAPP,  CORNELIA,  26,  88,  172,  192 

COHEN,  SEYMOUR  S.,  Some  struggles  of  Jacques 
Loeb,  Albert  Mathews,  and  Ernest  Just  at  the 
Marine  Biological  Laboratory,  127 

Cold  Spring  Harbor  Laboratory,  management,  ori- 
gin, and  organization  of,  183 

Colloid  chemistry,  127 

Comparative  physiology  and  biochemistry  at  the 
Zoological  Station  of  Naples,  122 

CONK.LIN,  EDWARD  GRANT,  26,  62,  80,  107,  172, 
187,  192,  200 

CURTIS,  WINTERTON  C.,  88 


D 

Darwinism,  35,  62 

DAVENPORT,  CHARLES  B..  172,  183 

DOHRN,  ANTON,  1,  4,  35,  122,  137,  172,  187,  197 

DREW,  GILMAN  A.,  88 

DRIESCH,  HANS,  26,  35,  107,  187 

Drosophila,  99,  107,  127 


E 

Early  struggles  at  the  Marine  Biological  Laboratory 
over  mission  and  money,  192 

Early  studies  of  cell  interactions,  80 

EBERT,  JAMES  D..  Carnegie  Institution  of  Washing- 
ton and  marine  biology:  Naples,  Woods  Hole, 
and  Tortugas,  172 

EBERT,  JAMES  D.,  Cell  interactions:  the  roots  of  a 
century  of  research,  80 

EBERT.  JAMES  D.,  Evolving  institutional  patterns 
for  excellence:  a  brief  comparison  of  the  or- 
ganization and  management  of  the  Cold  Spring 
Harbor  Laboratory  and  the  Marine  Biological 
Laboratory,  183 

ECCLES,  J.  C.,  159 

Ecology 

at  Naples,  168 
at  the  MBL,  197 

Eelgrass  disease,  197 

Electrical  transmission,  159 

Embryology,  35,  80,  99,  107,  127 

Embryology  at  Naples  and  the  MBL,  35 

Embryonic  development,  62 

Ephase,  159 

Epigenesis,  26,  35,  62 

Evolutionary  century  at  Woods  Hole:  instruction 
in  invertebrate  zoology.  An,  88 

Evolving  institutional  patterns  for  excellence:  a 
brief  comparison  of  the  organization  and  man- 
agement of  the  Cold  Spring  Harbor  Laboratory 
and  the  Marine  Biological  Laboratory,  183 

Eyes,  153 


FANTINI,  BERNARDINO,  The  sea  urchin  and  the 

fruit  fly:  cell  biology  and  heredity  1900-1910, 

99 
First  impressions:  American  biologists  at  Naples, 

187 
FLOREY,  ERNST,  The  Zoological  Station  at  Naples 

and  the  neuron:  personalities  and  encounters 

in  a  unique  institution,  137 
From  Woods  Hole  to  the  world:  The  Biological 

Bulletin,  200 
Fruit  fly,  see  Drosophila 
Future  ecology,  MBL  site,  197 


205 


206 


INDEX   TO   VOLUME    168  (SUPPLEMENT) 


Ganglion  cell,  1 37 

Genetics  and  embryology,  107 

Germany,  research  funding,  44 

GHIRETTI,  FRANCESCO,  Comparative  physiology 
and  biochemistry  at  the  Zoological  Station  of 
Naples,  122 

Giant  axons,  137 

Giant  fibers.  153 

GRANT,  CHARLES,  4 

GROEBEN,  CHRISTIANE,  Anton  Dohrn — the  states- 
man of  Darwinism,  4 

GROEBEN,  CHRISTIANE,  see  Alberto  Monroy,  35 

GROSS,  PAUL  R.,  1 

GROSS,  PAUL  R.,  Laying  the  ghost:  embryonic 
development,  in  plain  words,  62 

GRUDFEST,  H.,  159 


H 

HAECKEL,  ERNST  HEINRICH,  35,  62,  107,  137 

Heredity,  99 

Heredity   under  an   embryological   paradigm:   the 

case  of  genetics  and  embryology,  107 
History  of  instruction,  MBL,  88 
History,  MBL  site,  197 
History,  botany,  168 
HYATT,  ALPHEUS,  26,  183,  192,  197 


Instruction,  history  at  MBL,  88 

Interactions,  cell,  80 

Interactions,  cell  surface,  80 

Interactions,  inductive,  80 

Invertebrate  instruction  at  Woods  Hole,  88 


Biological  Laboratory  over  mission  and  money, 

192 
MAIENSCHEIN,  JANE,  First  impressions:  American 

biologists  at  Naples,  187 
Marine  Biological  Laboratory,   1,  26,  62,  80,  88, 

107,  127,  172,  183,  187,  192,  197,  200,  203 
Marine   Biological    Laboratory,   organization   and 

management  of,  183 
Marine  botany  and  ecology  at  Stazione  Zoologica, 

168 

MATHEWS,  ALBERT  P.,  127 
MAYER  (Mayor),  ALFRED  G.,  172 
Mechanistic  materialism,  107 
Memory,  153 
Mendelism,  99,  107 
MIRALTO,  ANTONIO,  What  laboratories  for  what 

science?  203 
MONROY,  ALBERTO,  AND  CHRISTIANE  GROEBEN, 

The  "new"  embryology  at  the  Zoological  Sta- 
tion and  at  the  Marine  Biological  Laboratory, 

35 
MORGAN,  THOMAS  HUNT,  26,  62,  99,  107,  127, 

137,  187,  200 
Morphology,  26,  35 

N 

NACHMANSOHN,  O.,  159 
NANSEN,  FRIDTJOF,  137 
Nerve  fiber,  137 
Neurofibrils,  137 
Neuron,  137 
Neurosecretion,  137 

Nicked  by  Occam's  razor:  unitarianism  in  the  in- 
vestigation of  synaptic  transmission,  159 


O 


Occam's  razor,  62,  159 


JONES,  JOHN  H.,  183 

JUST,  ERNEST  EVERETT,  99,  127 


Laying  the  ghost:  embryonic  development,  in  plain 

words,  62 
Library,  MBL,  200 
LILLIE,  FRANK,  26,  62,  80,  88,  107,  127,  183,  192, 

200 

LOEB,  JACQUES,  62,  107,  127 
LOEWI,  O.,  159 


M 

MBL  and  Cold  Spring  Harbor  compared,  183 
MAIENSCHEIN,   JANE,   Agassiz,   Hyatt,   Whitman, 
and  the  birth  of  the  Marine  Biological  Labo- 
ratory, 26 
MAIENSCHEIN,  JANE,  Early  struggles  at  the  Marine 


Penikese  Island,  26 
Photoreceptors,  extraocular,  153 
Physiology,  122,  127 
Preformationism,  26,  35,  62 
Proteins,  127 
Publications 

MBL,  200 

Naples,  200 

R 

Racism,  127 

REDFIELD,  A.  C.,  200 

REINGOLD,  NATHAN,  AND  JOEL  N.  BODANSKY, 

The  sciences,    1850-1900,   a  north  Atlantic 

perspective,  44 
Research   funding,   USA,   United   Kingdom,   and 

Germany,  44 
RUSSELL-HUNTER,  W.  D.,  An  evolutionary  century 


INDEX   TO  VOLUME    168   (SUPPLEMENT) 


207 


at  Woods  Hole:  instruction  in  invertebrate 
zoology,  88 

RUSSELL-HUNTER,  W.  D.,  From  Woods  Hole  to 
the  world:  The  Biological  Bulletin.  200 

RUSSELL-HUNTER,  W.  D.,  The  Woods  Hole  Lab- 
oratory site:  history  and  future  ecology,  197 


u 

USA,  research  funding,  44 

Unitarianism,  159 

United  Kingdom,  research  funding,  44 

United  States  Fish  Commission,  26,  187,  192,  197 


SCHARER,  ERNST,  137 

Science,  problems  relating  to,  203 

Science,  public  support,  44 

Science,  role  of,  203 

Sciences,  1850-1900,  a  north  Atlantic  perspective. 
The,  44 

Sea  urchin  and  the  fruit  fly:  cell  biology  and 
heredity  1900-1910,  The,  99 

Some  struggles  of  Jacques  Loeb,  Albert  Mathews. 
and  Ernest  Just  at  the  Marine  Biological  Lab- 
oratory, 127 

Statocysts,  153 

Stazione  Zoologica  of  Naples,  1,  4,  62,  107,  122, 
137,  168,  172.  187.  192,  197 

Stretch  receptor  neurons,  137 

Synapse,  159 


TOMAS,  CARMELO  R.,  Marine  botany  and  ecology 
at  Stazione  Zoologica,  168 


W 

What  laboratories  for  what  science?  203 
WHITMAN,  CHARLES  OTIS,  26,  35,  88,  127,  172, 

187,  192,  197,  200,  203 
WILSON,  EDMUND  BEECHER,  26,  35,  62,  99,  127, 

137,  172,  187,  192,  200 
Women's  Education  Association,  26,  192 
Woods  Hole  Laboratory  site:  history  and  future 

ecology.  The,  197 


YOUNG,  J.  Z.,  137 

YOUNG,  J.  Z.,  Cephalopods  and  neuroscience,  153 


Zoological  Station  at  Naples  and  the  neuron:  per- 
sonalities and  encounters  in  a  unique  institu- 
tion, !37 

Zoology,  invertebrate  at  MBL,  88 


999#'  027 


ACME 

in  co..  INC. 


MAR  28  1986 


STREET 
CHARLESTOWN,  MASS., 


UH    1B57    R