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A   ;  5  i 


F  A  N  Y 


•  •  , . 


DIOLOGY 
U 

i 


A  Manual  of 


Botany 


FOR 

INDIAN    FOREST 
STUDENTS 


By  R.  S.  JJOLE,  F.C.H.,  F.L.S..  F.E.s. 

Deputy  Conservator  of  Forests 
and  Forest  Botanist  at 
the  Imperial  Forest  Research 
?  Institute  of  Dehra  Dun  3 


CALCUTTA 

SUPERINTENDENT  GOVERNMENT  PRINTING.  INDIA 
1909 


BIOLOGY 

LIBRARY 

G 


Li*. 


ERRATA. 

Page     11,  lines  14-15  for  are  called  stolons  read  is  called  a  stolon. 

„       27,  line  19,  for  plant  ssuch  read  plants  such. 

„       34,     „    24,/0rlormed  read  formed. 

„       34,     „    25,  for  farge  read  large. 

„       41,     „      8,  after  Dolichandrone  delete  comma. 

„       41,     „      9,  from  below,  for  typica  read  typical. 

„       50,     „       l,for  ligutiform  read  liguliform. 

„       50,  footnote,  for  jurnished  read  furnished. 

„       55,  line  14  from  below,  insert  comma  after  one. 

„       57,  marginal  note,  for  Diagrams  Floral  read  Floral  Diagrams. 

„       58,         „  „       „     S  ed  read  Seed. 

„       63,  line  3,  after  falling  off  insert  or  fading. 

„       70,    „     11  from  below,  for  Alcurene  read  Aleurone. 

„       81,  footnote,  line  4  from  below,  for  —  a  wrinkle)  read(  =  a,  wrinkle). 

„       89,  line  2  from  below, /or  orophyll  read  chlorophyll. 

„       96,  marginal  note,  for  Wtaer  read  Water. 

„     107,  line  4  from  below,  after  side  delete  comma. 

„     109,     „      4  from  below,  for  i.e.  read  e.g. 

„     111,     „     1 5,  for  A  read  As. 

„     111,     „    25,  after  These  insert  are. 

„     111,  marginal  note,  Jor  ertilisation  read  fertilisation. 

„     135,  line  14  from  below,  for  forest  read  forests. 

„     154,  footnote,  for  Physiiogy  read  Physiology. 

„     156,  line  17  from  below,  for  asexua  read  asexual. 

„     180,     „      16,  for  ha  s  read  has. 

„     227,  marginal  note,  for  Plant  read  Plants. 

„     239,  line  3  from  below,  for  mong  read  Among. 

„     244,  marginal  note,  for  Wesern  read  Western. 

„     245,          „  „       „    Forest  read  Forests. 

„     249,  line  23,  for  of  read  or. 
Index,  Page  ii,  after  Bark  internal,  for  80  read  79. 

„          „          after  Bast  internal,  for  80  read  79, 

„          „    X,  after  Internal  bark,  for  80  read  79. 

„          „    xviii,  after  Stomata,  for  48  read  84. 

„          „      „        „      Stool-shoots,  for  71  read  171. 
In  Explanation  of  Figures,   Plate  IV,    last  line,  for  pedately-parted   read 

pedately-di  vided. 
In  Explanation  of  Figures,  Plate  VI,  line   5   from  below,  for   branche  read 

branch. 
In  Explanation  of  Figures,  Plate  XVII,  line  9,  for  wood  parenchyma-bearing 

read  wood-parenchyma  bearing. 

In  Explanation  of  Figures,  Plate  XVII,  line  11,  for  vessel  read  tracheid. 
In  Explanation  of   Figures,  Plate  XVII,  lines  12  and    14,  for  trachea  read 

tracheid. 

lu  Explanation  of  Figures,  Plate  XIX,  line  2,  for    Ureao-spoies  read  Uredo- 
spores. 


S.  G.  P.  J.— 224  I.  G.  F.— 10-1-10— 1,100-  S.  C.  L 


PREFACE. 

1.  THIS  Manual  has  been  pre- 
pared primarily  for  the  use  of  the  students  of  the  Imperial 
Forest  College,  Dehra  Dun.     In  the  absence  of  a  suitable  botani- 
cal text-book  a  great  deal  of  the  time  of  the  instructors  and 
students  at  the  Forest  College  has  hitherto  been  spent  in  dic- 
tating and  copying  lecture  notes,  respectively.     The    publi- 
cation of  the  present  Manual,  which  contains  the  back-bone,  as 
it  were,  of  the  College  course  of  botanical  lectures,  will  enable 
the  instructor  to  profitably  employ  much  of  the  time  formerly 
wasted  on  dictation  in  repeating  parts  of  the  course,  in  further 
explaining  and  illustrating  points  which  are  found  to  give  diffi- 
culty, and  in  frequently  examining  and  explaining  the  character- 
istics of  living  plants  in  the  field  and  in  class.     Although  some 
plants  have  been  mentioned  as  examples  in  the   Manual,  the 
number  of  these  is  necessarily  very  limited.     It  must,  however, 
be  remembered  that  much  may  be  learnt  from  any  individual 
plant  and  that  the  constant  illustration  of  the  facts  mentioned 
in  the  lectures  by  a  reference  to  living  plants  is  essential  if  the 
teaching  is  to  be  made  interesting  and  of  any  practical  value, 
and  if  it  is  to  have  any  result  other  than  the  committal  to 
memory  of  a  mass  of  imperfectly  understood  facts  to  be  repro- 
duced at  an  examination  and  then  forgotten. 

2.  The   object   of  the   present 

Manual  is  to  give  a  general  introduction  to  the  science  of  Botany 
in  accordance  with  modern  knowledge,  in  so  far  as  this  is  pos- 
sible in  the  limited  time  available  for  botanical  instruction  at 
the  Forest  College,  particular  attention  being  paid  to  those 
points  which  at  present  seem  to  be  of  special  importance  for  the 
Forest  Officer  in  India. 

3.  I  have    throughout    endea- 
voured to  keep  in  view  the  needs  of  the  practical  Forest  Officer. 


PREFACE. 

He  should  for  example  possess  a  good  knowledge  of  the  charac- 
ters most  commonly  employed  in  plant  descriptions  and  thus  be 
able  to  identify  plants  from  their  written  descriptions  and  to 
use  a  Flora  with  advantage.  He  should  also  have  an  eye  for 
those  characters  of  bark,  habit,  etc.,  which  enable  one  to  recog- 
nise on  sight  different  species  in  the  forest  at  different  times 
of  year. 

Such  characters  are  dealt  with  in  Part  I  of  the  Manual. 
Those  for  whom  this  book  is  primarily  intended  have  very  little 
time  for  microscopic  work  both  during  their  course  of  training 
and  thereafter.  Anatomy  therefore  is  very  briefly  dealt  with 
in  Part  II  and  only  so  far  as  has  been  considered  necessary  for 
a  fairly  clear  idea  of  the  main  facts  of  Physiology  and  Pathology 
as  given  in  Parts  III  and  V. 

4.  In  accordance  with   the  in- 
structions which  were  received  regarding  the  preparation  of 
this  Manual  systematic  botany  is  treated  very  shortly  in  Part  IV. 
An  attempt  has  there  been  made  to  explain  the   principles  of 
classification,  and  while  avoiding  unnecessary  detail  to  give  a 
fair  idea  of  the  chief  characters  and  general  appearance  of  the 
plants    contained    in    the  main    divisions    of   the  Vegetable 
Kingdom,  as  until  a  student  is  able  to  place  any  plant,  accord- 
ing to  the  aggregate  of  its  characters,  into  at  all  events  its  main 
group  with  fair  accuracy  he  cannot  be  considered  competent  to 
use  a  special  Flora  which  only  deals  in  detail  with  one- or  more 
minor  groups.     The  detailed  classification  of  Dicotyledons  and 
Monocotyledons  has  not  been  dealt  with  but  the  students  at 
the  Forest  College  are  required  to  fully  utilise  every  opportunity 
they  may  have  of  collecting,  examining  and  endeavouring  to 
classify  the  plants  met  with  on  their  tours — a  task  in  which  they 
are  assisted  by  Kanjilal's  most  useful  Forest  Flora  of  the  School 
Circle  which  contains  full  descriptions  of  most  of  the  important 
forest  species  likely  to  be  found,  and  also  by  a  short  course  of 
lectures  delivered  during  the  students'  tours,  giving  the  charac- 
ters of   the  most  important  cohorts   and    natural  orders   of 
together  with  the  names  of  the  chief  plants  of 


PREFACE.  Ill 

economic  importance  which  they  contain.  If  a  future  edition  of 
this  Manual  is  called  for  it  will  probably  be  then  advisable  to 
add  a  section  dealing  with  this  part  of  the  subject. 

5.  It  is  to  be  hoped  that  in  a  few 

years'  time   our   knowledge   of   injurious   fungi   will  be  suffi- 
cient for  the  compilation  of  a  fully  illustrated  hand-book  des- 
cribing the  most  important  forest  fungi  and  paying  particular 
attention  to  characters  which  can  be  recognised  in  the  forests. 
For  the  present,  however,    the    correct   identification  of  any 
particular  fungus  depends  on   skilled  microscopic  work  and 
must  be  left  to  the  expert.     The  paragraphs  of  this  book  which 
deal  with  fungi   may  therefore  be   considered   unnecessarily 
detailed  for  students  who  have  little  opportunity   for  working 
with  the  microscope.     The  facts  regarding  fungi  given  in  Parts 
/Fand  V  and  the  plates  illustrating  the  same,  however,  merely 
aim  at  giving  an  idea  of  the  general  appearance  of  some  typical 
fungi  (the  majority  of  which  also  the  student  will  have  oppor- 
tunities   of  seeing   and  becoming  familiar  with  on  his  tours), 
of  their  life  history  and  injurious  action  on  the  plants  attacked 
and  also  of  the  microscopic   characters  on  which  their  classi- 
fication depends,  with  the  object  of  helping  the  student  to 
recognise  in   the  forest  the  presence  of  an  unknown  injurious 
fungus,  to  select  specimens  of  the  same  necessary  for  its  identi- 
fication by  the  expert  and  to  form  some  idea  as  to  the  best 
remedial  and  protective  measures  to   be  taken  pending  the 
identification  of  the  pest. 

6.  The  question  of  Plant  Dis- 
eases has  perhaps  been  treated  on  rather  broader  lines  than 
is  usual  in  an   elementary  text-book.     The  principle  that  a 
plant's  welfare  depends  on  the  balance  struck  between  the  effects 
of  a  numbei  of  injurious  and  favourable  factors  and  that  a 
disease  is  rarely  due  to  a  single  factor  frequently  appears  to  be 
insufficiently  appreciated,  while,  until  we  know  more  about  the 
normal  physiology  and  life  history  of  our  important  forest  species 
and  about  the  relations  which  exist  between  them  and  other 
organisms,  plants  as  well  as  animals,    very  little  real  progress 


IV  PREFACE. 

in  the  investigation  and  prevention  of  the  diseases  of  ouf 
valuable  forest  plants  and  in  our  efforts  to  favour  the  reproduc- 
tion of  the  same  appears  possible.  An  attempt  has  been  made 
to  emphasize  this  point  of  view  in  Parts  V  and  VI  of  the  Manual. 

7.  While    endeavouring   in   all 

cases  to  avoid  didactic  assertions  on  points  which  have  not  yet 
been  sufficiently  proved.  I  have  not  hesitated  to  indicate  the 
possible  bearing  of  well-established  botanical  facts  on  Indian 
forest  phenomena  with  the  object  of  stimulating  further  obser- 
vation and  research,  and  in  this  respect  it  is  hoped  that  the  book 
may  possibly  be  of  some  use  to  others  besides  those  for  whom 
it  is  primarily  intended. 

With  reference  to  the  plants  which  have  been  quoted  as  ex- 
amples of  the  various  points  considered,  I  have  as  a  rule  selected 
those  which  the  College  students  will  frequently  meet  with  on 
their  tours,  but,  with  the  object  of  making  the  book  more  inter- 
esting to  Forest  Officers  generally,  I  have  also  included  a  few 
well-known  species  which  are  not  found  wild  in  the  neighbour- 
hood of  Dehra  Dun.  In  such  cases,  however,  the  species  in 
question  are  usually  available  for  reference  in  the  College 

Herbarium  or  Museum. 

8.  As  regards  the  illustrations, 

in  the  time  available  for  the  preparation  of  this  Manual  only  a 
limited  number  of  plates  could  be  prepared  and  those  have  been 
selected  which  at  present  seem  most  likely  to  be  useful.  Thus 
no  drawings  of  the  minute  structure  of  plants  for  example  were 
prepared,  there  being  a  good  series  of  anatomical  wall-plates 
available  for  instruction  purposes  in  the  Forest  College. 

9.  My  special  thanks  are  due  to 

Mr.  S.  Eardley  Wilmot,  Inspector  General  of  Forests  to  the 
Government  of  India,  who  very  kindly  entrusted  me  with  the 
present  work  ;  to  Dr.  E.  J.  Butler,  Imperial  Mycologist  of  the 
Pusa  Agricultural  Research  Institute,  who  has  most  generously 
allowed  me  to  use  his  original  work  and  drawings  for  the  purpose 
of  this  book  and  to  whose  help  the  value  of  the  portions  dealing 
with  fungi  is  entirely  due  ;  to  Mr.  H.  H.  Haines  of  the  Imperial 


PREFACE.  V 

Forest  Service  who  has  most  kindly  read  the  proofs  of  Parts  IV, 
Fand  VI  and  given  me  many  valuable  suggestions  and  criticisms 
most  of  which  I  have  been  able  to  give  effect  to,  and  to  Captain 
Gage,  Director  of  the  Botanical  Survey  of  India,  for  his  kind 
help  and  advice. 

10.  During  the  preparation  of 

this  book,  covering  as  it  does  the  main  sub- divisions  of  the  whole 
science  of  botany,  the  number  of  books  referred  to  has  neces- 
sarily been  large  and  only  those  to  which  I  am  most  indebted  can 
be  mentioned  here.  These  are  : — 

(1)  The  various  works  of  the  late  Professor  H.  Marshall 

Ward,  for  whose  inspiring  teaching  and  encourage- 
ment also  the  writer  takes  this  opportunity  of  ex- 
pressing his  deep  sense  of  gratitude. 

(2)  Asa  Gray's  Botanical  Text- Book.,  Volume  I. 

(3)  Dr.  E.  Strasburger's  Text-Book  of  Botany. 

(4)  A  Students'  Text-Book  of  Botany,  by  S.  H.  Vines. 
(5) .  Sachs'  Lectures  on  the  Physiology  of  Plants. 

(6)  Physiology  of  Plants,  by  Dr.  W.  Pfeffer. 

(7)  Species  and  Varieties,  their  origin  by  Mutation,  by  H. 

de  Vries. 

(8)  Diseases  of  Trees,  by  R.  Hartig,  Eng.   Trans.,  edited 

by  H.  Marshall  Ward. 

(9)  Plant  Geography,  by  Dr.  A.  F.  W.  Schimper. 

(10)  Brandis'  Flora  of  North-  West  and  Central  India. 

(11)  Brandis'  Indian  Trees. 

(12)  Gamble's  Manual  of  Indian  Timbers. 

(13)  Indian  Forester. 

(14)  Mr.  P.  H.  Clutterbuck's  Working  Plan  for  the  Forests 

of  Jaunsar-Bawar. 


VI  PREFACE. 

11.  Finally  it  must   be  noted 

that  all  the  figures  for  PUtes  I  to  IX,  XI  to  XIII  and  XX  in- 
elusive,  with  the  only  exception  of  the  diagrams  included  in 
Plates  II  and  IX  have  been  drawn  from  nature  by  my  wife,  to 
whose  work  therefore  this  book  owes  much  of  any  interest  or 
value  it  may  possess. 

R.  8.  HOLE. 
April  1908. 


CONTENTS. 


INTRODUCTION. 

Two  Biological  Sciences  —  Plants  and  Animals  —  Sub-divisions  of  Botany  — 
Principal  Plant  Organs  and  their  Functions  —  Selection  of  Types  for 
Description  ........... 


PAGE. 


1—4 


PART  I- MORPHOLOGY. 

CHAPTER  I.— THE  ROOT. 

Primary,  secondary  and  adventitious  roots — Types  of  Root  Systems — Deve- 
lopment of  Root  precedes  that  of  Shoot — Root  Suckers — Fibrous, 
Tuberous,  Woody,  Subterranean,  Aquatic  and  JSrial  Roota]  .  . 

CHAPTER  II.— THE  STEM. 

Nodes,  Internodes,  Position  of  Leaves  and  Buds — Kinds  of  Stems — Branch- 
ing— Shape  of  Stem — Stem  Structure,  Bark — Leaf-scars,  Lenticels — 
Pith,  Heart- Wood,  Sap- Wood,  Annual  Rings,  Medullary  Rays,  Pores, 
Resin-Canals,  Vascular  Bundles  .  .  .  .  .  •  • 

CHAPTER  III.— THE  LEAF. 

Lamina,  Petiole,  Nerves — Venation— Leaf-margin — Simple  and  Compound 
Leaves— General  Shape  of  Leaves— Polymorphic  JLeaves — Leaves  of 
Young  Plants— Other  Characters  of  Leaves— Colour — Smell — Taste- 
Glands— Characters  of  Petiole — Leaf-Base — Stipules — Metamor- 
phosed Leaves— Phyllotaxy— Buds — Vernation — Homologous,  Ana- 
logous Members  .,....•••• 


6—8 


)— 19 


20—36 


CHAPTER  IV.— THE  INFLORESCENCE  AND  FLOWER. 

Flowering-Shoot — Bracts — Peduncle,  Pedicel,— Types  of  Inflorescence — 
Parts  of  the  Flower,  Perianth,  Stamens,  Pistils,  Torus — Pollination  and 
Fertilisation — Characters  of  Perianth — Position  and  Numbers  of  Floral 
Parts— Distribution  of  Sexual  Organs— Symmetry  of  Flower — Floral 
Parts  are  Leaves — Cohesion  and  Adhesion — Hypogynous,  Perigynous 
and  Epigynous  Flowers— Disc — Descriptive  Terms  for  Petals  and 
Sepals— Estivation— Characters  of  Stamens— of  Pistils— of  Ovules- 
Types  of  Flowers  which  may  cause  Difficulty— Wheat— Pea— Pine— 
Gyinnosperms  and  Angiosperuis— Floral  Diagrams  and  Short-hand 


37—57 


viii  CONTENTS. 

PART  I.— MORPHOLOGY— concld. 

CHAPTER  V.— THE  SEED  AND  FRUIT. 

PAGE. 

The  Seed — Testa,  Tegmen,  Aril — Albuminous  and  Exalbuminous   Seeds — 

Cotyledons,  Radicle  and  Plumule  of  Embryo — Epicotyl — Hypocotyl — 
i  Monocotyledons   and   Dicotyledons — The   Fruit — Different   Types    of 
Fruit — Descriptive  Terms  for  Shape  of  Fruit  and  Seed         .         .         .        68 — 62 

CHAPTER  VI.— GENERAL. 

Duration  of  Plant  Members — Texture  of  Plant  Members — Prickles,  Hairs, 
Scales,  Glands — Habit — Herbs,  Trees,  Shrubs, — Annuals,  Biennials, 
Perennials  —  Gregarious  and  Sporadic  Plants — Evergreen  and  Deci- 
duous Plants  .  .  .  .  . 63—67 

PAET  IL-ANATOMY. 

CHAPTER  I.— CELLS. 

Protoplasm — The  Cell— Cell-contents — Cell-division — Pits  and  Thickening 

of  Cell-wall— Middle  Lamella— Shape  of  Cells 68—72 

CHAPTER  II.— TISSUES. 

Different  kinds  of.  Tissues  and  their  Elements — Tegumentary,  Vascular 
Bundle  and  Fundamental  Tissue  Systems — Structure  and  Develop- 
ment of  Vascular  Bundles  .  .  .  .  .  .  .  .  73 — 76 

CHAPTER  III,— STRUCTURE  AND  DEVELOPMENT  OF  PLANT 

MEMBERS. 

Stem  of  Gymnosperms  and  Dicotyledons — of  Monocotyledons — Root? — 
Abnormal  Development — Periderm  and  Bark — Lenticels — Shedding 
of  Leaves — Development  of  Secondary  Members — Root-cap — Root 
hairs — Shortening  of  Roots — Structure  of  Leaves  ....  77 — 84 

PART  III-PHYSIOLOGY. 

CHAPTER  I.— FUNCTIONS  OF  PLANTS  IN  GENERAL. 

General  Conditions  of  Plant  Life — Osmosis — Turgescence — Transpiration 
— Manufacture  of  Organic  Food  Materials — Respiration  and  Meta- 
bolism— Conditions  of  Existence  in  a  Highly  Organised  Plant  .  .  ^5 — 92 

CHAPTER  II.— FUNCTIONS  OF  THE  HIGHER  PLANTS  IN  DETAIL. 

Plant  Food  Materials — Essential  and  Non-essential  Substances — Avail- 
ability— Absorption  and  Ascent  of  Water  in  Plants — Transpiration — 
Assimilation — En/ymes — Respiration — Growth — Plant  Movements — 


CONTENTS.  IX 

PART   III.— PHYSIOLOGY— concld. 

CHAPTER  II.— FUNCTIONS   OF   THE  HIGHER  PLANTS   IN 

DETAIL— concld. 

PAGE 

Reproduction — Sexual  and  Asexual  Methods — Cross  and  Self-Fertili- 
sation— Anemophilous,  Entomophilous,  Ornithophilous  Plants — Con- 
trivances for  Facilitating  Cross- Fertilisation — Fertilisation  of  t  Flowers 
of  Salvia  lanata  and  Berberis  Lycium — Cleistogamic  Flowers — Dis- 
semination of  Seeds  .........  93 — lift 

PART  IV.-CLASSIFICATION. 

CHAPTER  I.— DEFINITIONS  AND   EXPLANATIONS. 

Necessity  for  Classification — Natural  and  Artificial  Systems — Unit  of  Classi- 
ijcation-^Species,  Sub-species,  Variety,  Race,  Genus  and  larger  Natural 
Groups  —  Nomenclature  —  Practical  Determination  cf  the  Natural 
Groups  . 117—125- 

CHAPTER  II.— PRINCIPAL  SUB-DIVISIONS  OF  THE 
VEGETABLE  KINGDOM. 

Cryptogams  and  Phanerogams — Sub-divisions  of  the  Cryptogams  —  Algae 
— Bacteria — Practical  Importance  of  Bacteria — Fungi — Classification 
of  Fungi — Phycomycetes  —  Ascomycetes  —  Basidiomycetes  —  Ustila- 
ginaceae — Uredinaceaa — Lichens — Liverworts  and  Mosses — Ferns, 
Horsetails  and  Club-Mosses — Phanerogams  divided  into  Gymnos- 
perms  and  Angiosperms — Classification  of  Gymnosperms — Angios- 
perms  divided  into  Monocotyledons  and  Dicotyledons — Synopsis  of 
Principal  Sub-divisions  of  the  Vegetable  Kingdom  ....  126—147 

CHAPTER  III.— THE  ORIGIN  OF  SPECIES. 

Struggle  for  Existence — Importance  of  Sub-species  and  Races — Hybrids — 
Fluctuating  Variability — Mutations — Other  Kinds  of  Variation — 
Heredity — Selection — Origin  of  Species  in  Nature  ....  143  — 161 

CHAPTER  IV.— COLLECTION  AND  PRESERVATION  OF 
SPECIMENS. 

Collection — Preservation — Herbarium  .         .         .         .  .         .     162 — 164 

PART  V- WOUNDS  AND  DISEASES. 

CHAPTER  I.— WOUNDS. 

Healing  of  Wounds  by  means  of  a  Cork  Layer — Callus  Formation  and  heal- 
ing by  Occlusion — Bruises — Pruning — Girdling — Cuttings — Pollard- 
Shoots,  Stool-Shoots,  Root-Suckers — Propagation  by  Layers  and  Guti 
—Grafting— Budding  .....  ...  165—175 


X  CONTENTS. 

PABT  V.- WOUNDS  AND  DISEASES-cow^. 

CHAPTER  II.— DISEASES. 

SECTION  I.— INTRODUCTORY  REMARKS. 

a:      PAGE. 
Duration  of  Plant  Life — Definition  of   Disease — Struggle   for   Existence — 

Factors  influencing  the  Relations  existing  between  Organisms — Sapro- 
phytes— Competitors,  Parasites,  Symbionts — Symptoms  of  Disease — 
Disease  rarely  due  to  one  Factor  alone — Knowledge  required  for 
Investigation  of  Diseases — Sub-division  of  the  Subject  .  ,  .  176—183 

SECTION  II.— INFLUENCES  OF  OTHER  PLANTS  ON  PLANT- 
DEVELOPMENT. 

(a)  PLANT  COMPETITORS. 

Deodar  and  Blue  Pine — Teak  and  Bamboos — Trees  and  Grasses — Trees  and 

Strobilanthes — Climbers — Epiphytes  ...,,,     183 — 186 

(&)  PLANT  PARASITES. 

PhytophtJiora  infestans — PhytophtJiora  omnivora — Fames  annosus — Armil- 
laria  mdlea — Tramcies  Pini — Puccinia  graminis — Mcidium  montanum 
— Orobanche  indica — Cuscuta  reflexa — Loranthus  longiftorus — Parasi- 
tism of  Santalum  album 186 — 202 

(c)  PLANT  SYMBIONTS. 

Lichens — Bacteria      and    Leguminosce — Mycorhizas — Distant    Symbiosis 

between  Green  and  Non-green  Plants — Decay  of  Wood       .         .         .     202 — 205 

SECTION  III.— INFLUENCES  OF  THE  SOIL  ON  PLANT-DEVELOPMENT. 

Necessary  Salts  in  suitable  Condition  and  Quantities — Poisonous  Sub- 
stances— Necessary  Water  and  Oxygen — Accumulation  of  Starch  in 
Leaves  indicates  Root  Trouble 205 — 208 

SECTION  IV.— INFLUENCES  OF  THE  ATMOSPHERE  ON  PLANT- 
DEVELOPMENT. 

Extremes  of  Temperature — Effects  of  Excessive  Heat — Of  Excessive  Cold — 
Effects  of  Light — Of  Excessive  Moisture—Of  Winds — Of  Poisonous 
Substances — Lightning 208 — 216 

SECTION  V.— EFFECT  OF  FIRE. 

Effect  of  Fire — Necessity  of  studying  Indirect  Effect  of  each  Factor     .     .     216—217 


CONTENTS.  Xi 

PART   VI -GEOGRAPHY. 

CHAPTER  I.— FACTORS   INFLUENCING   THE   DISTRIBUTION 

OF  PLANTS. 

PAGB. 

-Available    Water — Soil — Temperature — Light — Air — Existence    of     other 

Plants — Existence  of  Animals — Fire — Action  of  Man — Necessity  of 
avoiding  Hasty  Conclusions  as  to  Factors  responsible  for  Distribution 
—  Power  of  Adaptation — Xerophytes,  Hygrophytes,  Tropophytes  .  218 — 230 

CHAPTER  II.— PRINCIPAL   TYPES  OF  VEGETATION. 

Woodland,  Grassland  and  Desert — Their  Distribution  depends  chiefly  on 
Moisture,  Soil,  and  Action  of  Man — Pioneer  Plants — Succession  of  differ- 
ent Plant  Generations  or  Crops — Action  of  Man  in  altering  Course  of 
Natural  Events  is  limited — Conversion  of  Grassland  into  Forest — Trees 
producing  Root-suckers  favoured  in  the  Struggle  with  Grasses  .  .  231 — 236 

CHAPTER  III.— PRINCIPAL  TYPES  OF,  INDIAN   FORESTS. 

Types  of  Forests— Factors  responsible  for  their  Distribution— Arid-country 
Forests — Deciduous  Forests — Evergreen  Forests — Hill  Forests — Tidal 
or  Littoral  Forests — Riparian  Forests — Distribution  of  important 
Species  ......  ....  237—250 


INTRODUCTION. 

1.  All  living  beings    are  said  to  Two 

belong  either  to  the  Animal   or   Vegetable   Kingdom,  in  other  Biplo8ical 
words  they  are  classed,    respectively,  as    Animals  or  Plants. 
Biology  is  a  term   applied  to  the  study  of  all   living  beings, 

i.e.  of  both  animals  and  plants,  The  two  biological  sciences 
are  Zoology  and  Botany,  the  former  embracing  the  study  of 
animals,  the  latter  that  of  plants. 

2.  So    long   as   we  confine  our  plants  and 

attention    to    those    highly    organised  forms  of   life    which  Animals, 
have    an   elaborate    and    complicated  structure    we   find  no 
difficulty  in    distinguishing   the  members    of  the  two  natural 
kingdoms.     Hence,  in  early  times,  when  the  total  number  of 
plants  and  animals  known   to  scientists    was   comparatively 

small  while  those  which  were  known  belonged  to  the 
higher  forms,  a  definite  line  of  distinction  could  be  easily 
drawn  between  them.  With  the  progress  of  time  knowledge 
has  gradually  extended,  and  with  the  help  of  the  microscope 
we  have  gradually  passed  from  a  consideration  of  the  large 
and  obvious  to  a  study  of  the  minute  and  formerly  invisible 
things  of  life,  and  we  are  now  aware  of  the  existence  of  a 
number  of  simple  forms  of  life  to  which  the  ordinarily  ac- 
cepted notions  of  what  constitutes  an  animal,  or  a  plant,  fail  to 
apply  and  which,  in  consequence,  we  cannot  classify  as  either. 
The  study  of  such  forms  which  stand  at  the  bottom  of  the 
scale  drives  us  to  the  conclusion  that  no  definite  boundary  line 
can  be  drawn  between  the  Animal  and  Vegetable  Kingdoms, 
and  it  should  be  borne  in  mind  that  the  substance  which  con- 
stitutes the  basis  of  life,  viz.  protoplasm,  is  essentially  the 
same  in  both  animals  and  plants.  With  the  exception  of  such 
simple  forms,  however,  there  is  generally  a  great  difference 
between  the  members  of  the  two  kingdoms,  and  it  is  obviously 
convenient  for  the  purpose  of  study  to  keep  them  separate. 
Animals,  then,  as  compared  with  plants,  usually  have  more 
independence  of  action,  while  perhaps  the  mo&b  characteristic 
distinction  of  the  majority  of  plants  consists  in  the  fact  that 
they  possess  a  green  colouring  matter  called  chlorophyll  (a 
word  which  means  leaf -green),  by  means  of  which  they  are 


able  to  manufacture  the  food  materials  necessary  for  the  main- 
tenance of  life  from  the  carbon  dioxide  of  the  air  and  water. 
Animals,  being  unable  to  do  this,  are  obliged  to  maintain  them- 
selves by  consuming  the  food  manufactured  by  plants,  and  if 
plants  were  exterminated  all  life  on  the  earth  would  cease.  Thus 
plants  are  often  called  the  great  food-producers  and  animals 
the  great  food-consumers.  At  the  same  time  it  must  be  noted 
that  a  large  group  of  plants  known  as  the  Fungi  are  excep- 
tional in  this  respect,  for  they  possess  no  chlorophyll  and  like 
\  animals  depend  on  green  plants  for  their  food. 

Sub-divisions  3.  Botany   may  be  sub-divided 

of  Botany.    jn^o  fae  following  divisions  :— 

I.  Vegetable  Morphology. 

II.          ,,        Anatomy. 
IIT.          „        Physiology. 
IV.  Classification  or  Systematic  Botany. 

V.   Vegetable  Pathology. 
VI.  Geographical  Botany. 

The  study  of  the  economic  uses  of  plants  and  of  their  pro- 
ducts is  also  sometimes  treated  as  a  separate  branch  which  is 
then  termed  Economic  Botany. 

Vegetable  Morphology  comprises  the  study  of  the  form  of 
plant  bodies  and  of  the  several  parts  of  such  bodies  as  visible 
to  the  naked  eye. 

Vegetable  Anatomy  comprises  the  study  of  the  minute  struc- 
ture of  plant  bodies  as  seen  by  the  aid  of  the  microscope. 

Vegetable  Physiology  comprises  the  study  of  the  plant  as  a 
living  being.  Just  as  morphology  regards  each  plant  body, 
or  part  of  such  body,  as  a  fact,  as  something  which  exists  and 
possesses  a  certain  shape  and  structure,  so  Physiology  regards 
it  as  a  factor,  as  something  which  is  capable  of  action  and  of 
performing  definite  work.  While  morphology  is  content  with 
the  knowledge  of  the  existence  of  certain  plant  structures, 
Physiology  asks  what  function  these  structures  fulfil  and  how 
they  do  it.  Eegarded  from  a  purely  morphological  stand-point, 
the  various  parts  of  a  plant  body  are  called  members.  When 
such  parts  are  regarded  as  performing  some  particular  kind  of 
work,  i.e.  from  the  physiological  point  of  view,  they  are 
termed  organs.  The  greater  the  number  of  organs,  each  being 
adapted  to  the  performance  of  one  or  more  functions,  possessed 
by  a  plant,  the  more  highly  organised  is  the  plant  said  to  be. 

Although  Part  III  of  this  book  is  devoted  to  Physiology,  a 


few  physiological  details  have  also   been  mentioned,  here  and 
there,  in  the  other  parts  where  this  has  been  thought  advisable. 

Classification  aims  at  arranging  all  plants  in  groups  accord- 
ing to  their  resemblances,  with  the  object  of  facilitating  the 
identification  of  plants  and  of  enabling  anyone  to  rapidly  ac- 
quire a  knowledge  of  the  plants  of  any  particular  region  and  of 
their  correct  names,  as  universally  recognised  by  botanists. 

Vegetable  Pathology  includes  the  study  of  plant  diseases, 
wounds  and  injuries. 

Geographical  Botany  considers  the  distribution  of  plants 
over  the  earth  and  its  causes. 

4.  If  we  take  a  seedling  of  any  of  Principal 
our  common  forest  trees  we  can  at  once  distinguish  : —  anTt 

(a)  The  Root  which  goes  downwards  and  firmly  anchors 
the  plant  in  the  substratum  on  which  it  is  grow- 
ing. 

(6)  The  Shoot  which  goes  upwards  in  a  direction  dia- 
metrically opposite  to  that  of  the  root  and  which 
consists  of  an  axis,  or  stem,  bearing  green  leaves. 

It  will  help  us  to  understand  the  various  forms  assumed  by 
these  two  sets  of  organs  and  their  behaviour  under  different 
circumstances  if  we  realise,  at  the  outset,  that  the  root  ab- 
sorbs from  the  substratum  in  which  it  is  growing  water  with 
mineral  substances  in  solution,  which,  however,  are  as  yet  use- 
less to  the  plant  as  food  and  which  may  be  therefore  called 
raw  food  materials.  This  water  ascends  the  stem  and  passes 
into  the  green  leaves.  The  protoplasm  in  these  leaves,  by  the 
aid  of  the  green  chlorophyll  and  when  it  is  supplied  with 
water  by  the  roots  and  has  air  and  light  in  contact  with  it,  is 
able  to  build  up  a  carbohydrate,  which  usually  first  becomes 
visible  in  the  form  of  starch,  from  the  carbon  dioxide  of  the 
air.  This  starch  contains  the  energy  by  means  of  which  the 
plant  is  able  to  live,  grow  and  perform  its  vital  functions — is 
able  not  only  to  feed  and  keep  alive  its  protoplasm,  but  also  to 
actually  manufacture  additional  living  protoplasm  from  the 
inorganic  substances  brought  up  by  the  roots  and  to  carry  out 
other  work.  The  root  must  thus  grow  and  spread  in  the  dark 
and  moist  substratum,  while  the  shoot  must  come  forth  into 
the  sunlight  and  air  in  order  to  obtain  its  supplies  of  carbon 
dioxide  and  to  expose  its  chlorophyll  to  the  rays  of  sunlight. 

The  shoot  also  eventually  gives  rise,   first,  to  flowers   and 
then,  to  fruits,  the  latter  containing  the  seeds  which  will  once 

B2 


more  develop  into  seedlings  and  new  plants.     All  plant  organs 
may  thus  be  primarily  subdivided  into  :— 

(1)  Vegetative  Organs  which  enable  the   plant   to   exist 

and  maintain  itself  alive,  such  as  the  root,  stem 
and  leaves. 

(2)  Reproductive  Organs  the  function  of  which  is  to  pro- 

duce new  plants,  such  as  flowers. 

The  root  may  be  distinguished  from  the  shoot  by  the  fact 
that  the  former  never  directly  gives  rise  to  leaves,  or  true  re- 
productive organs. 

Selection  5.  Finally,    it    is    necessary    to 

of  Types  for  p0int  out  that  the  forms  of  plants  and  of  their  members 
Description.  ^re  a^mos^  infinitely  various  and  that,  probably,  no  two 
members  of  even  one  and  the  same  plant  are  ever  exactly 
alike,  although  the  differences  between  them  are  frequently 
extremely  minute.  Hence  it  is  obviously  impossible  to 
separately  define  every  form  which  exists.  Generally  speaking, 
therefore,  morphology  aims  at  the  selection  of  broad  types 
for  description,  which  can  be  easily  distinguished  from  one 
another  and  each  of  which  may  be  taken  as  fairly  represent- 
ing the  most  important  characters  of  a  large  number  of 
subordinate  forms  between  which  the  differences  are  very 
slight  and  difficult  to  distinguish.  At  the  same  time,  it  must 
be  noted  that  these  typical  forms  are  generally  connected  by 
intermediate  forms,  through  which  one  type  passes  over  by 
almost  insensible  gradations  into  another,  and  that,  therefore, 
although  the  descriptions  of  the  types  apply  to  a  large  propor- 
tion of  all  existing  forms,  there  are  always  some  forms  for 
which  it  is  not  easy  to  decide  which  typical  description  is  most 
suitable  At  the  same  time  this  is  no  practical  disadvantage, 
for  forms  which  are  precisely  intermediate,  may,  with  equal 
justice,  be  referred  to  either  of  the  two  types  between  which 
they  stand,  while  all  others  are  referred  to  the  types  which 
thev  resemble  most  closelv. 


PART  I.-MORPHOLOGY. 

CHAPTER  I.— THE  EOOT. 

6.  The  first  root  developed   by  Primary, 

a   seedling    is    called    the    primary     root.      All    other    roots  Secondary 
whether   developed   from    the    primary   root,    or    from  other  Adventitious 
parts  of  the  plant,  are  called    secondary.     The   root,    like  the  Roots.  .    ^ 
shoot,   is   capable    of    branching    and   rebranching,    the   new 
roots  being  normally  developed  in  longitudinal  rows,  laterally, 
on   the  parent  root,  in  such  a  way  that  the  youngest  are  always 
nearest  the  growing-apex  of  the    mother-root,   i.e.  they  are 
developed  in  acropetal  succession    and    a    line,    following   the 
course    of   development   of   the   lateral  roots,  passes  from  the 
oldest  ones  at  the  base  of  the  parent-root  to  the  youngest  ones 
at  its  apex. 

The  young  roots  are  developed  from  the  inner  tissues  of 
the  parent-root  and  as  they  grow  they  break  through  the  outer 
tissues.  This  can  be  well  seen  in  the  long  roots  which  hang 
from  the  branches  of  the  Banyan  (Ficus  bengalensis),  the 
young  roots  pushing  through  cracks  in  the  outer  tissues  of  the 
parent  roots.  The  aggregate  of  roots  developed  by  a  plant 
constitutes  its  root-system.  All  roots  which  are  not  branches  of 
the  primary  root,  or  which,  being  such  branches,  are  not 
developed  in  acropetal  succession,  are  said  to  be  adv entitious, 
this  term  being  applied  to  all  parts  of  plants  which  develop 
either  in  abnormal  positions,  or  out  of  their  proper  order. 
Roots,  therefore,  may  first  of  all  be  classified  according  to  their 
mode  of  origin  into : — 

(1)  Primary. 


(2)  Secondary  ]  }?>  t V 

J    {(0)  Adventitious. 

Roots  which  spring  from  the  stem  are  therefore  adventi- 
tious, such  as  are  those  often  seen  on  the  sugarcane  as  shown 
in  Plate  I,  Fig.  4. 

7.  If  we  examine   the   seedling  Types  Of 

of  one   of    our    common  forest    trees,    say    an   Oak.  we  shall  Root- 
find    that    the    primary    root    elongates    and    continues    to  yst<  IM§ 
grow    vigorously  for  a  considerable    period.     Such   a   root    is 
termed  a  tap-root  and  all  the  roots  developed  from  it   are   its 
lateral  roots,  see  Fig.  7,  Plate  I.     If  the   tap-root  is   injured  a 


6 

lateral  root  usually  takes  its  place  and  continues  its  growth,  in 
the  direction  of  the  original  tap-root.  A  root-system  with  a 
tap-root  has  thus  a  distinct  main  axis  and  usually  penetrates  to 
a  considerable  depth  in  the  soil.  A  strong  tap-root  is  deve- 
loped by  many  of  our  forest  trees,  e.g.  Teak  (Tectona  grandis) , 
Sissoo,  (Dalbergia  Sissoo)  and  Anjan  (Hardwickia  binata). 
Such  a  root  of  Sissoo,  or  Anjan,  may  attain  a  length  of  six 
feet  in  the  first  year. 

In  many  plants,  however,  such  as  grasses  and  their  allies,  the 
first  root  sent  out  by  the  seedling  dies  off  after  a  short  time,  or 
does  not  develop  vigorously,  and  stronger  roots  then  spring 
from  the  base  of  the  stem  above  the  primary  root.  These, 
although  they  often  arise  above  the  ground  in  the  air,  may 
ultimately  penetrate  the  soil  and  branch  there,  and  in  many 
plants  the  entire  root-system  consists  of  such  adventitious 
roots.  They  may  be  well  seen  in  the  sugarcane.  For  illustra- 
tions, see  Plate  I,  Figs.  3  and  4. 

The  question  whether  a  plant  develops  a  strong  tap-root, 
or  a  superficial  root-system,  is  of  great  practical  importance. 
Plants  with  the  same  type  of  root-system  compete  with  each 
other  in  the  same  layers  of  soil  for  water  and  raw  food 
materials.  A  tree  with  a  long  tap-root,  like  the  Sissoo,  is  thus 
a  good  nurse  to  protect  tea-bushes  from  the  effects  of  frost, 
the  roots  of  these  two  plants  not  interfering  with  each  other, 
while  the  superficial  roots  of  Toon  (Cedrela  Toona)  and  other 
species,  which  are  often  planted  in  avenues  near  field  crops,  are 
injurious  to  the  latter. 

The  extent  of  the  root-system  developed  by  a  plant  depends 
on  the  amount  of  green  foliage  produced  by  the  shoot.  The 
greater  the  latter,  the  larger  and  more  vigorous  is  the  former. 
Plants  with  leaves  floating  in  water  usually  have  a  small  total 
area  of  leaf  surface,  while  water  also  is  plentiful  close  at  hand, 
and  such  plants  usually  possess  a  small  root-system.  If,  on  the 
other  hand,  we  collect  all  the  roots  of  a  large  forest  tree,  such 
as  the  Sal,  and  place  them  end  to  end  they  would  probably 
cover  a  distance  of  several  miles. 

Development  8.  In  the  case   of    many  of  our 

of  Root        forest   trees  the  development  of  the    root  to    a   great  extent 

itf*A/%nri<vQ  •»  1*1 

precedes  that  of  the  shoot.  The  plant,  as  it  were,  has  to 
make  sure  of  its  foundations  below  ground  before  pro- 
ceeding with  the  superstructure  and  the  development  of  the 
shoot  above  ground  may  often  be  quite  insignificant  for  several 
years,  during  the  early  life  of  the  seedling.  Fig.  1,  Plate  I, 
shows  the  strong  well-developed  tap-root  of  a  young  seedling  of 


ih.eJ$euciOsk(Quercusmcana)  and  the  insignificant  young  shoot. 
In  some  cases  no  true  leaves  at  all  are   developed    during  the 
first  year  but  only  rudimentary  leaves,  small  scaly  structures,  such 
as  are  seen  in  Figs.  I  and  2,  Plate  I.     In  many  cases  the  young 
shoot  of  the    seedling   dies    back   year  after  year  more  or  less 
completely,  while  the  development  of  the  root-system  steadily 
progresses,  such  as  frequently  happens  with  Teak  and  SaL  Even- 
tually, when  a  vigorous  root-system  has  been  developed,  a  strong 
shoot  is  sent  up  which  is  capable  of  ultimately   forming   a   full- 
sized  mature  shoot.     The  advantage  of  this  procedure  is  often 
obvious,  such  as  is  the  case  for  example  with  those   trees   like 
Sal,  Harra  (Terminalia  Chebula),  Jamun  (Eugenia  Jambolana), 
and  Mahua  (Bassia  latifolia),    which,   in  the  plains   of  India, 
come  into  leaf  during  the  hot  dry  season  from   March   to  May, 
when  the  roots  are  obliged  to  send  up  large  quantities  of  water 
to  supply  the  leaves,  and  it  is  therefore  necessary  that  the  roots 
should  be  in  the  deep  moist   subsoil   and  not  in  the  parched 
surface  soil.     In  such  trees,  therefore,  a  strong  deep-going  root- 
system  must    be    formed    before    a    vigorous    shoot    can    be 
developed. 

9.^Although  roots  never  direct-  Root 

ly  give  rise  to  leaves,  they  are  capable,  under  certain Suckera- 
circumstances,  of  bearing  buds  which  may  develop  into 
leafy  shoots  called  root-suckers,  which  are  characteristic  of 
many  of  our  forest  trees  and  shrubs,  such  as  Sissoo,  Tendu 
(Diospyros  tomentosa)  and  many  others.  They  are  often  found 
springing  from  roots  which  have  been  exposed  to  the  light  at 
the  sides  of  a  road- cutting,  or  on  the  banks  of  streams,  for 
example. 

10.  A  typical  root  is  usually  Fibrous  and ' 
cylindrical  in  shape  and  when  fine  and  thread-like  it  is 
termed  fibrous.  Many  plants  manufacture  more  food  material 
than  is  required  for  immediate  consumption,  this  excess 
material  being  stored  until  it  is  wanted.  The  subterranean 
roots  and  the  lower  part  of  the  stem  are  favourite  reservoirs 
for  such  food  and  as  they  are  stuffed  with  the  material  they 
become  much  swollen  and  lose  their  ordinary  shape.  In  some 
plants,  such  as  the  Radish,  Turnip,  Carrot  and  Beet,  the  tap- 
root is  swollen,  in  others,  such  as  the  Dahlia,  the  secondary 
roots  are  stuffed.  Such  swollen  roots  are  said  to  be  tuberous. 
If  the  swollen  root  is  broader  in  the  middle  and  tapers  towards 
both  ends,  like  a  spindle,  it  is  said  to  be  fusiform,  if  it  is  like  a 
turnip,  broader  than  high,  it  is  napiform.  and  if  tapering  re- 
gularly from  a  broad  base  to  the  tip  like  a  carrot,  it  is  conical. 


8 

Woody  11.  In  the    case  of    most  trees 

Boots.  the  old  roots  become  hard  and  woody  as  they  thicken  and 
then  they  no  longer  absorb  water  and  food  materials  from 
the  soil  but  merely  serve  as  conducting  pipes  to  pass  on 
to  the  stem  the  supplies  which  have  been  collected  by  the 
younger  roots.  They  also  anchor  the  tree  more  strongly  in 
the  ground,  the  massive  roots  resisting  the  lever-like  strains 
exerted  by  the  heavy  trunk  with  its  great  crown  of  foliage, 
as  it  sways  in  the  wind.  In  the  case  of  many  Indian  trees  the 
roots  form  woody  supports,  like  buttresses,  at  the  base  of  the 
trunk,  which  are  for  instance  very  characteristic  of  Bombax 
malabaricum,  see  Fig.  4}  Plate  XL 

Subterra-  12.  According     as    roots     are 

nean  developed  in   the   earth,  water,    or  air,    they   may   be  called 

JSdai  Roots,  subterranean,  aquatic  or  cerial.  The  aerial  roots  of  the 
Banyan  on  reaching  the  soil,  branch  and  develop  a  vigorous 
root-system  therein,  while  the  aerial  portions  become  woody 
and  support  the  horizontal  branches  like  columns.  In  Rubus 
lasiocarpus,  the  serial  roots  produced  at  the  tips  of  the  long 
curved  branches  give  rise  to  young  plants,  which  become  inde- 
pendent and  are  separated  from  the  parent  by  the  decay  of  the 
connecting  branches. 

Other  aerial  roots  serve  as  attachment  organs  and  enable 
climbing  stems  to  cling  firmly  to  their  substratum ;  these  are 
iound  in  the  Ivy,  Ficus  scandens,  and  many  others.  These 
roots  are  also  in  some  cases  able  to  absorb  moisture  and  food 
materials,  provided  that  such  are  available.  Thus  some  climbing 
stems  are  able  to  exist  after  they  have  been  cut  and  their 
connection  with  the  subterranean  root  system  severed. 

Sometimes  the  aerial  roots  originating  on  the  stem  grow 
obliquely  downwards  into  the  ground  and  form  remarkable 
stilt-like  supports  which  may  be  seen  in  Screw-Pines  (Panda- 
mis),  Mangroves  (Rhizophora),  and,  in  miniature,  sometimes 
in  the  Maize  (Zea  Mays).  Some  aerial  roots  possess  the  excep- 
tional power  of  being  able  to  absorb  moisture  from  the  air  by 
means  of  a  spongy  outer  sheath  of  tissue  called  the  velamen, 
found  in  many  orchids.  In  a  few  cases  also  the  aerial  rootF 
are  green  m  colour  and  possess  chlorophyll. 


CHAPTER  II.— THE  STEM. 

13.  As  has  been    noted    above,  Nodes, 
the    stem    of   a   plant    bears   the   structures   which    we   call 

•  T  £     i    J.T_    A     j-x. 

leaves.  If  we  examine  an  ordinary  stem  we  rind  tnat  tne  Leaves  and 
leaves  are  situated  at  certain  definite  positions  on  it,  which  Buds, 
are  separated  by  a  more  or  less  considerable  length  of  stem 
which  bears  no  leaves.  The  places  where  the  leaves  are 
borne  are  called  the  nodes,  the  piece  of  stem  lying  between  two 
successive  nodes  is  called  an  internode.  The  nodes  are  often 
swollen  and  are  sometimes  marked  by  a  distinct  line  or  joint, 
as  in  Gnetum.  The  youngest  leaves  are  found  to  be  nearest  the 
growing  apex  of  the  stem,  i.e.  they  are  developed  in  acropetal 
succession.  If  only  two  leaves  occur  on  one  node  they  are  on 
opposite  sides  of  the  stem  and  are  said  to  be  opposite,  if  there 
is  only  one  leaf  at  each  node,  the  leaves  are  said  to  be  alternate,  or 
scattered.  If  more  than  two  leaves  occur  at  a  node,  they  are 
situated  at  equal  distances  apart  and  are  then  said  to  be  in 
whorls  (or  in  verticils)  or  whorled  (or  verticillate).  If  at  each 
node  there  are  two  leaves  which  are  nearly,  but  not  quite, 
opposite,  they  are  said  to  be  sub-opposite.  When  more  than  two 
leaves  occur  at  a  node  which  are  nearly,  but  not  quite,  at  the 
same  level,  they  are  said  to  be  in  false  whorls  (or  verticils).  In 
practical .  descriptive  botany,  however,  the  difference  between 
true  and  false  whorls  (or  verticils)  is  often  not  insisted  on, 
both  being  merely  called  whorls  (or  verticils).  When  pairs  of 
opposite  leaves  are  arranged  on  the  stem  in  such  a  way  that 
the  longitudinal  axis  of  each  pair  is  at  right  angles  to  the  axis 
of  the  pair  above  and  below  it,  the  leaves  are  said  to  be 
decussate,  and  are  then  in  four  vertical  ranks.  When  leaves 
are  arranged  in  two  vertical  ranks  they  are  distichous,  or  bifa- 
rious.  When  two  or  more  leaves  are  inserted  very  close  together 
so  that  they  appear  to  be  in  tufts,  the  leaves  are  said  to  be 
fascicled,  or  in  fascicles,  as  in  the  Deodar.  The  upper  angle 
between  a  leaf  and  the  stem  is  called  the  axil.  On  the  stem, 
in  addition  to  leaves,  we  find  the  structures  which  we  call  buds 
and  which  are  capable  of  growing  out  into  new  stems,  or 
branches.  These  buds  are  normally  found  in  the  axils  of  the 
leaves  when  they  are  called  axillary,  or  at  the  apex  of  the  stem 
when  they  are  said  to  be  terminal.  The  leaf,  in  the  axil  of 
which  a  bud  appears,  is  called  the  subtending  leaf,  while  the  bud, 
having  a  leaf  close  below  it,  is  said  to  be  subtended  by  the  leaf. 


10 

This  fact  that  buds,  and  the  branches  which  develop  from 
them,  as  a  rule  occupy  a  definite  position  as  regards  the  leaves 
is  of  great  importance,  and  we  are  thus  helped  in  doubtful 
cases  to  arrive  at  a  decision  as  to  whether  a  certain  member  is 
to  be  regarded  as  of  the  nature  of  a  branch  or  of  a  leaf.  What- 
ever first  arises  in  the  axil  of  a  leaf  may  usually  be  regarded  as 
a  branch,  and  what'  subtends  a  branch  as  a  leaf.1 

Various  14.  A  stem    which   is   soft  and 

*?S?f  more  or  less  succulent   is   said  to  be  herbaceous,     if  firm,  and 

more  or  less  tough  and  hard,  it  is  said  to  be  woody. 

In  some  plants  there  is  very  little,  if  any,  stem  and  they 
are  said  to  be  stemless,  or  acaulescent,  e.g.  Plicenix  acaulis. 
In  others  there  are  several  stems  of  equal  vigour  ;  such  stems, 
occurring  in  tufts,  are  said  to  be  caespitose,  as  in  many 
Bamboos.^ 
Sterjcwfare  :  — 


erect,  when  ascending  perpendicularly, 

ascending,  when  rising  obliquely, 

decumbent,     when    erect,    but    with    the    basal    portion 

horizontal,  or  nearly  so, 

reclining,  when  the  basal  portion  is  more  or  less  erect 
and  the  upper  portion  curved  downwards  with  the 
apex  trailing  on  the  ground, 

procumbent,  or  prostrate,  when  lying  on  the  ground, 
repent,  or  creeping,  when  prostrate  and   also   rooting   as 

they  grow, 

soandeni,  when  climbing, 

twining,  when  they  climb  by    spirally  coiling  around   a 

support.     Twiners  may  ascend  in  two  directions,  i.e. 

they  may  ascend  from  left  to  right,  as  viewed  from 

outside  the  coil,  in  which  case   they  are  said  to  be 

dextrorse   and   to    move   in     an    anti  —  or    counter- 

clockwise direction,  or  they  may  ascend  from   right 

to  left,  when    they  are  said  to  be   sinistrorse  and  to 

move  in  a  clockwise  direction,  see  Fig.  1,  Plate  II. 

Stems  may  also   climb  by  means    of    special  organs  called 

tendrils.     These   are    slender,    thread-like    bodies,    simple    or 

branched,  which  firmly  attach  themselves  to  a  support  by  coiling 

around  it,  or  by  adhering  to  it,  and  thus  hold  the  stem    secure- 

ly.     Others    climb    by    means    of     hooks    and    spines,     e.g. 

Calamus  tennis. 

The  stems  of  some  plants,  although  not  distinctly  climbing, 
are  wide-spreading  and  may  often  be  found  resting  on,  and 
more  or  less  entangled  with,  the  stems  and  branches  of  other 


11 

plants.  They  are,  as  it  were,  weak,  or  imperfect,  climbers, 
and  are  usually  described  as  rambling,  or  straggling,  e.g. 
Quisqualis  indica  and  Deeringia  celosioides. 

A  large  climber  with  a  woody  stem  is  called  a  liane,  e.g. 
Bauliinia  Vahlu. 

A  stem  rising  directly  from  the  ground,  bearing  flowers, 
but  no  green  leaves,  is  called  a  scape,  e.g.  Orobanche  indica. 

The  peculiar  jointed  stems  of  grasses  and  bamboos  which 
are  hollow  between  the  joints  are  called  culms. 

The  unbranched  columnar  stem  of  palms  and  Tree  Ferns  is 
termed  a  caudex. 

An  a3rial  or  subterranean  branch,  rooting  and  giving  off 
shoots  which  become  independent  plants,  by  the  decay  of  the 
branch  connecting  them  with  the  parent  stem,  are  called 
stolons.  The  Potato  has  subterranean  stolons,  while  the 
rooting  branches  of  Rubus  lasiocarpus  are  a3rial  stolons. 

A  runner  is  a  slender  stolon  with  long  internodes,  well  seen 
in  the  Strawberry.  A  plant  which  produces  a  number  of 
stolons,  or  runners,  is  said  to  be  sarmentose. 

A  rhizome  is  a  stem  of  root-like  appearance  prostrate  on,  or 
buried  under,  the  ground,  giving  off  slender  roots  usually  at  the 
nodes  and  producing  erect  serial  shoots,  or  leaves,  progressively 
from  the  growing  apex,  as  found  in  many  Grasses,  Bamboos  and 
the  common  Bracken  Fern,  Pteris  aquilina. 

Stolons,  runners,  and  rhizomes,  are  sometimes  very  like  roots, 
but  they  may  always  be  distinguished  from  roots  by  the  fact 
that  they  directly  give  rise  to  leaves  in  the  axils  of  which  buds 
appear,  although  such  leaves  are  usually  minute  and  scale-like. 

A  bulb  is  a  short  shoot  with  a  flattened  or  conical  stem, 
provided  with  thick,  fleshy,  scale-like  leaves  and  from  the 
base  of  which  roots  are  developed,  e.g.  the  Onion. 

A  corm  is  like  a  solid  bulb,  the  main  portion  of  the  corm  con- 
sisting of  the  thickened  stem  which  is  naked,  or  with  a  few 
inconspicuous,  scaly,  investing  leaves. 

A  tuber  is  a  short  thickened  shoot,  or  part  of  a  shoot,  bearing 
.  inconspicuous  scaly  leaves,  with  buds  in  their  axils.  In  the 
Potato,  tubers  are  formed  at  the  ends  of  stolons,  the  so-called 
"eyes"  being  the  axillary  buds.  A  tuber  can  be  distinguish- 
ed from  a  tuberous  root  by  the  fact  that  it  bears  scaly  leaves, 
in  the  axils  of  which  the  buds  arise.  Small  bulbs,  corms  and 
tubers  sometimes  appear  on  a3rial  stems  in  the  leaiL-axils  and 
they  are  then  called  bulbils,  which  are  found  in  many  species 
of  Dioscorea.  They  are  capable  of  developing  roots  and 
producing  independent  plants. 


12 

A  cladode  is  a  stem  coloured  green,  with,  or  without, 
inconspicuous  scale  leaves,  which  more  or  less  resembles  a  leaf 
and  performs  the  functions  of  a  leaf,  e.g.  the  stem  of  the 
prickly  pear,  Opuntia  Dillenii. 

Branching.  15.  Branching,    i.e.    the    pro- 

duction of  new  stems  from  the  parent  stem,  takes  place  in  two 
principal  ways  termed,  respectively,  dichotomous  and  lateral 
branching.  In  the  first  the  growing  point  of  the  parent 
stem  divides  into  two  ;  two  branches  of  equal  vigour  are  thus 
formed,  as  in  a  two-pronged  fork,  and  there  is  no  continuous 
main  axis.  This  method  of  branching  is  very  rare. 

In  lateral  branching  the  growing  point  of  the  parent  stem 
does  not  divide  and  new  stems  are  developed  laterally  from  the 
parent.  There  are  two  principal  kinds  of  lateral  branching, 
the  monopodial  and  cymose.  In  the  former  there  is  a  distinct 
and  simple  main  axis,  formed  by  the  elongation  of  the  parent 
stem.  Such  a  main  axis  is  termed  a  monopodium.  Such  a 
system  of  branching  is  well  seen  in  a  Pine  tree  in  which  the 
leader  has  uninterruptedly  developed  from  the  terminal  bud. 

In  cymose  branching  there  is  no  simple  main  axis  formed 
by  the  elongation  of  the  parent  stem  and  the  lateral  branches 
grow  faster,  or  for  a  longer  period,  than  the  parent.  Of  this 
form  of  branching  there  are  two  types,  the  sympodial  and 
falsely  di-  or  tri-chotomous.  In  sympodial  branching  there  is  a 
main  axis  but  this,  instead  of  being  a  simple  axis  consisting  of 
the  parent  stem,  is  built  up  of  the  basal  portions  of  a  number 
of  lateral  branches.  On  each  segment  of  such  an  axis,  one 
lateral  branch  develops  more  vigorously  than  any  other,  i.e. 
there  is  one  leader.  The  axis  so  formed  is  termed  a  sympo- 
dium,  or  false  axis,  and  is  at  first  crooked  but  frequently 
straightens  subsequently,  see  Fig.  3,  Plate  II.  This  may  often 
be  seen  on  the  twigs  of  trees  the  terminal  buds  of  which  have 
died,  or  not  developed  vigorously,  and  the  leaders  have  been 
formed  by  the  axillary  and  therefore  lateral  buds.  In  false- 
ly dichotomous  branching,  such  as  occurs  for  instance  in 
Rhamnus  virgatus,  the  branches  are  forked  and  at  first 
sight  there  appears  to  have  been  a  division  of  the  grow  ing  point 
of  the  parent  stem.  On  closer  inspection  it  appears  that  in 
this  plant  the  growing  point  really  terminates  in  a  spine  and 
that  there  is  no  dichotomy.  In  some  plants  the  terminal 
bud  becomes  a  flower,  and  in  others  it  dies,  or  does  not  develop. 
A  system  of  branches  each  one  of  which  forks  in  this  way 
and  produces  a  pair  of  lateral  branches  of  equal  vigour 


13 

is  called  a  false-dichotomy,  or  a  dichasium.  If  three  lateral 
branches  develop  with  equal  vigour  the  branching  is  falsely  tri- 
chotomous.  In  botanical  descriptions,  however,  no  distinction 
is  as  a  rule  made  between  falsely  and  truly  di-  or  tri-chotom- 
ous,  the  branching  being  merely  described  as  dichotomous,  or 
trichotomous,  respectively.  We  are  accustomed  to  associate  the 
word  branching  principally  with  the  stem  of  plants,  but  it 
must  be  remembered  that  precisely  similar  branching  may  be 
exhibited  by  the  root,  or  leaf,  or  other  members. 

Branches  as  a  rule  arise  normally  in  acropetal  succession  and 
in  the  leaf  axils,  i.e.  they  are  axillary.  Those  which  originate 
otherwise  are  adventitious,  i.e.  they  are  developed  out  of 
their  proper  order,  or  in  altogether  exceptional  positions. 

As  branches  are  usually  axillary  their  arrangement  follows 
that  of  the  leaves,  and  like  leaves  they  may  therefore 
be  opposite,  sub-opposite,  alternate,  whorled  and  so  on. 
In  many  plants  while  some  branches  remain  stunted  and 
short,  others  grow  rapidly,  and  there  are  in  consequence 
elongated  and  dwarf  shoots.  These  occur,  for  instance,  in 
Randia  dumetorum  and  Pyrus  Pashia,  see  Fig.  3,  Plate  II. 
Such  dwarf  shoots  in  many  cases  develop  into  spines  and  at 
first  sight  it  is  not  always  easy  to  decide  whether  such  spines 
are  to  be  regarded  as  altered  (metamorphosed)  leaves,  or  stems, 
but  as  a  rule  they  are  found  in  the  axils  of  leaves,  or  they 
themselves  directly  give  rise  to  leaves,  thus  indicating  that 
they  are  stems,  or  rather  branches.  The  spines  of  the  Bel, 
Aegle  Marmelos,  of  Flacourtia  Cataphracta  and  others  may 
frequently  be  seen  bearing  leaves,  see  Figs.  1  and  2,  Plate  III. 
In  the  Pine  we  find  that  there  are  two  kinds  of  leaves. 
The  first  are  small  and  scale-like,  in  the  axils  of  which  small 
dwarf  shoots  with  brown  scales  at  the  base  and  terminating  in 
two  or  more  long  green  leaves,  or  needles,  are  borne.  On  the 
dwarf-shoots  of  the  Deodar  the  internodes  remain  so  short 
that  the  leaves  (needles)  appear  to  be  in  tufts,  or  fascicles. 

The  tendrils  of  Gouania  leptostachya  do  not  at  first  sight 
appear  to  be  branches  but,  as  in  the  case  of  the  spines  of  the 
Bel,  the  true  nature  of  these  organs  is  indicated  by  the  fact 
that  they  originate  in  the  leaf -axils  and  bear  leaves,  just  as  do 
normal  branches,  see  Fig.  3,  Plate  III. 

In  some  plants  the  stem  remains  simple  and  does  not 
branch,  e.g.,  in  the  majority  of  Palms.  In  a  plant  in  which 
the  stem  repeatedly  branches,  the  branches  thus  produced 
again  branching  and  so  on,  such  as  is  the  case  with  most  of  our 


14 


Shape  of 
Stem. 


Stem 

Structure, 

Bark. 


forest  trees,  the  smallest  branches  which  have  been  produced 
most  recently  are  called  twigs,  the  smaller  branches  branchlets, 
and  the  larger  ones  branches. 

16.  The  shape  of   the  stem  and 

branches  varies  considerably  in  different  plants  and  often  helps 
the  Forester  to  distinguish  his  trees  in  the  forest.  Many  trees 
have  large  buttresses  at  the  base  of  the  trunk,  e.g.  Bombax 
malabaricum,  see  Fig.  4,  Plate  XI.  In  Ougeinia  dalbergioides 
the  stem  -is  usually  short  and  almost  always  crooked.  In 
Acacia  leucophloea  it  is  almost  always  crooked  and  knotty 
(gnarled).  In  the  Hornbeam.  Carpinus  viminea,  it  is  distinctly 
fluted,  i.e.  with  broad,  shallow,  curved  grooves. 

In  many  plants  the  stem  is  cylindrical,  i.e.  terete,  in 
others  the  stem  has  3,  5,  or  more  distinct  ridges  or  angles. 
Sometimes,  in  addition  to  being  angled,  the  stem  is  more  or 
less  deeply  grooved,  or  hollowed  out  between  the  angles,  and 
it  is  said  to  be  channelled.  If  the  stem  is  marked  with  more 
or  less  parallel  furrows  it  is  sulcate,  if  the  furrows  are  very 
minute  and  look  like  mere  lines  the  stem  is  said  to  be  striate. 
The  stems  of  lianes  are  frequently  irregular  in  shape  and  are 
flattened  or  deeply  grooved. 

Such  peculiarities,  although  as  a  rule  not  noticeable  on 
the  old  stems  and  branches  of  trees  and  shrubs,  are  often  very 
characteristic  of  branchlets  and  twigs.  The  branchlets  of 
Flemingia  stricta,  for  instance,  are  3-angled  (triquetrous),  those 
of  Coriaria  nepalensis  are  quadrangular,  those  of  Teak  are 
channelled,  and  those  of  Flemingia  congesta  are  sulcate. 

17.  If  .now     we    cut    through 

a  stem  of  one  of  our  common  forest  trees,  say  the  Teak, 
and  examine  the  cut  surface,  we  find  the  greater  portion 
of  the  interior  of  the  stem  to  be  solid  and  hard,  consisting 
of  the  so-called  wood',  which  is  enveloped  in  a  comparatively 
thin  external  coat  of  softer  substance  of  a  different  colour, 
which  can  be  detached  from,  and  stripped  off,  the  solid  woody 
cylinder  which  it  covers.  This  outer  coat  is  the  so-called 
cortex,  or  bark,  of  the  Forester ;  its  characteristics  vary  greatly 
in  different  species  and  the  Forester  finds  them  very  valuable 
for  the  identification  of  trees  in  the  forest.  In  some  plants, 
e.g.  Palms  and  Bamboos,  there  is  no  true  bark.  The  first 
characteristic  of  the  bark  to  be  noted  is  its  thickness  ;  on  the 
stems  of  young  plants  and  the  twigs  of  old  plants  it  is  very 
thin.  On  the  old  stems  of  trees  and  shrubs  its  thickness  varies 
greatly  in  different  species.  Thus  in  Ougeinia  dalbergioides  it 


15 

is  only  about  one-sixth  of  an  inch  thick,  whereas  in  Sal  its  thick- 
ness is  from  1  to  2  inches. 

There  is  usually  a  considerable  difference  both  in  colour  and 
texture  between  the  outer  and  inner  layers  of  bark,  thus  in 
Phyllanthus  Emblica  the  outer  bark  is  pale  grey  and  the  inner 
substance  is  red  ;  in  most  trees  and  shrubs  also  the  inner  sub- 
stance is  distinctly  moist  while  the  outer  tissue  is  dead  and  dry. 

On  the  stems  of  young  plants  and  twigs  of  old  trees  the 
exterior  of  the  bark  is  usually  smooth,  but  on  the  old  stems  of 
trees  and  shrubs  the  degree  of  roughness  varies  greatly  in 
different  species.  InSterculia  wrens,  for  example,  it  is  very 
smooth  and  affords  a  marked  contrast  to  the  rough  bark  of  say 
an  old  Sal  tree.  Smooth  stems  may,  or  may  not,  be  shining  : 
in  Anogeissus  latifolia  the  smooth  stem  is  not  shining,  while 
in  the  Birch  it  is. 

The  colour  of  the  external  bark  is  not  only  different  in 
different  species  but  varies  on  different  parts  of  one  and  the 
same  plant.  Thus  the  twigs  and  branches  of  Carissa  spinarum 
are  bright  green,  but  the  old  stems  are  grey,  or  yellowish. 

The  twigs  and  branches  of  Berberis  aristata  are  dark  red,  of 
Rubus  lasiocarpus  purple  and  of  Salix  daphnoides  dark  green, 
or  almost  black.  The  colour  of  the  bark  on  the  stems  of  trees 
and  shrubs  is  very  important  for  identification  and  the  follow- 
ing may  be  noted  as  examples  : — 

In  Betula  utilis  it  is  almost  white. 

In  Albizzia  procera  it  is  greenish. 

In  Boswellia  serrata  it  is  yellowish. 

In  Stephegyne  parvifolia  it  is  bluish-grey. 

In  Anogeissus  latifolia  it  is  pale  grey. 

In  Terminalia  Arjuna  it  is  pale  pinkish-grey. 

In  Diospyros  tomentosa  it  is  almost  black. 

In  Sterculia  urens  it  is  sometimes  dark  red. 

The  texture  of  the  bark  also  varies  greatly  in  different 
species :  in  Betula  utilis  it  is  papery,  in  Erythrina  suberosa 
corky,  while  in  Teak  and  Cupressus  torulosa  it  is  fibrous,  i.e. 
it  breaks  up  into  thin  long  threads. 

The  bark  of  old  branches  and  stems  may,  or  may  not,  have 
fissures,  or  cracks,  of  various  kinds.  The  stem  of  Anogeissus 
latifolia  is  smooth  without  characteristic  fissures.  When  there 
are  fissures  they  may  be  shallow  as  in  Teak,  or  deep  as  in 
Pinus  longifolia.  The  way  in  which  such  fissures  run,  whether 
vertically  or  horizontally,  and  the  way  in  which  they  join,  or 
cross,  one  another  are  also  characteristic.  In  Buchanania 


16 


Leaf -Scars 

and 

Lenticels. 


Internal 
structure. 


Pith. 


latifolia  these  fissures  cross  at  right  angles  cutting  the  surface 
of  the  bark  into  small  squares  and  producing  a  tessellated 
appearance  ;  in  other  cases  strips  and  plates,  or  scales,  of  various 
shapes  and  sizes  are  produced. 

As  the  stem  of  a  tree  or  shrub  increases  in  size  the  outer 
layers  of  bark  are  cast  off  and  the  way  in  which  this  is  done  is 
often  characteristic.  In  the  Birch,  the  bark  peels  off,  i.e.  ex- 
foliates, in  rolls,  in  the  Khair  (Acacia  Catechu)  in  long  narrow 
strips  which  remain  for  some  time  on  the  trees  giving  a  ragged 
appearance,  and  in  Gmelina  arborea  in  large,  irregularly  shaped, 
scales,  thus  exposing  patches  of  pale  yellowish  surface  below 
which  contrast  strongly  with  the  grey  external  bark. 

18.  The   bark   on   young   stems, 

on  branchlets  and  on  twigs  often  exhibits  peculiar  and 
characteristic  marks,  the  principal  of  which  are  the 
so-called  leaf-scars  which  mark  the  spots  from  which  the 
leaves  have  fallen,  and  the  lenticels.  The  leaf -scars  of  course 
follow  the  arrangement  of  the  leaves  and  in  deciduous  species 
we  can  at  once  see  from  the  position  of  these  scars  whether 
the  leaves  are  opposite,  alternate,  whorled  and  so  on.  Their 
size  and  shape  vary  greatly  in  different  species.  A  reference 
to  Figs.  1  and  2,  Plate  XI,  will  indicate  how  we  may  readily 
distinguish  two  trees  (Odina  Wodier  and  Hymenodictyon  ex- 
celsum),  which,  when  bare  of  leaves,  are  somewhat  alike  in  the 
forest,  by  an  inspection  of  their  twigs  and  principally  by  the 
shape  and  arrangement  of  their  leaf-scars.  In  some  cases  the 
entire  leaf  is  not  shed  and  the  bases  of  the  leaves  persist  and 
give  the  stem  a  ragged  appearance,  as  in  species  of  Phoenix. 

Lenticels  are  interruptions  in  the  outer  coat  of  bark  which 
allow  air  to  penetrate  to  the  internal  tissues  and  they  usually 
appear  as  raised  corky  spots,  or  lines,  of  varying  shape  and  size. 
In  the  Birch,  for  instance,  they  are  horizontal  lines,  being  very 
large  and  conspicuous,  while  in  Pyrus  Pashia  they  are  small 
spots.  They  are  often  of  a  paler  colour  than  the  surrounding 
bark  and  are  thus  made  conspicuous. 

19.  If  now   we   look  at   the  cut 

surface  of  that   portion  of  our  teak  stem  which  lies  inside  the 
bark  we  find  certain  peculiarities  which  require  notice. 

In  the  centre  there  is  a  small  area  of  soft  tissue  called  the 
pith  which,  in  young  stems,  branches  and  twigs,  is  often  large 
and  conspicuous.  The  size  and  shape  of  the  pith  varies  in 
different  species.  As  a  rule  it  is  small,  but  in  Teak  it  is  rela- 
tively large  and  conspicuous.  In  the  Walnut  and  Prinsepia 
uiilis  it  is  divided  into  characteristic  large  chambers,  In  cross 


17 

section  it  is  quadrangular  in  Teak,  nearly  circular  in  Corylus 
Colurna,  the  shape  of  a  cross  in  Bauhinia  Vahlii,  usually 
oblong  in  Birches  and  pentagonal  in  Oaks. 

The  remainder  of  the  stem  lying  between  the  pith  and  the  Heart-  and 
bark  is  occupied  by  the  wood  proper  and,   unless   our  stem   is  Sap~Wood> 
too  young,  we  shall  find  that  the  central  portion  of  this  woody 
cylinder  is  dark-coloured  and  is  surrounded  by  a  belt  of   paler 
wood.    The  former  is  the  heart-wood  or  duramen  and  the  latter 
the  sap-wood  or  alburnum.     In  some  trees,  eg.    the  Silver   Fir, 
there  is  no   heart- wood,  but  when  this  is  present  its  colour  and 
size  in  comparison  with  those   of  the  sap-wood  are  important 
characters.    In  Teak  the  heart-wood,  when  fresh,  is  dark  golden 
yellow  and  the  sap-wood  is  white. 

In  the  wood,  arranged  around  the  pith,  we  find  numerous  Alinual 
lines  which  divide  the  wood  into  concentric  layers  ;  the  latter  Rmgs' 
are  the  annual  rings,  so-called  because  each  of  them  usually 
represents  the  amount  of  wood  formed  in  one  year  and 
hence  by  counting  these  rings  the  age  of  the  stem  may  be 
calculated.  These  rings  are  due  to  a  difference  in  the  density 
of  adjacent  concentric  layers  of  wood,  the  wood  formed  on  the 
outer  side  of  each  ring  being  denser  than  that  on  the  inner 
side  of  the  next  ring  which  adjoins  it.  In  several  species  no 
annual  rings  are  found,  e.g.  the  Mango  ;  in  others  there  are 
so-called  false  rings,  i.e.  bands  of  tissue  which  do  not  pass 
uninterruptedly  around  the  stem  and  which  often  run  into 
one  another.  These  may  be  seen  in  Pongamia  glabra  and 
Quercus  semecarpifolia. 

Through  the    wood,    passing    from    the    centre   outwards,  Medullary 
towards  the  circumference  of  the  stem,  are  radiating  lines  called  Rays> 
the  medullary  rays.     The  presence  or  absence,  width  and  other 
characters,  of  these  rays  are  important  points  in  the  identi- 
fication of  different  woods.     If  we  split  a  piece  of    the    Teak 
stem  radially  we   find  the   medullary   rays   appearing  on  the 
surface  of  the  radial  section  as  shining  plates,    giving   an   orna- 
mental appearance  to  the  wood  and  constituting  the  so-called 
silver-grain.  Looking  again  at  the  cross  section  of  our  stem  we 
find  a  number  of   minute    pores    which   are   larger    and    more Pores- 
numerous  in  the  inner  part  of   each  annual  ring    than    in   the 
outer  part.     The  size  of  these  pores  and  the  way  in  which  they 
are  distributed  throughout  the  wood  are   important  characters 
for  the  identification  of  trees  from  their  wood. 

The  hardness  and  weight  of  wood  are  also  well-known 
characters  for  distinguishing  trees.  With  Teak,  for  example, 
we  may  compare  the  much  softer,  lighter  wood  of  Erythrina 


18 

tuberosa  and  the  considerably  harder  and  heavier  wood  of 
Xylia  dolabriformis. 

Finally  the  characteristic  scent  of  the  Teak  wood  should  be 
noted  and  compared,  for  example,  with  that  of  the  wood  of 
Sandal,  Toon  and  Deodar. 

The  stem  of  our  example,  the  Teak,  of  which  we  have  now 
shortly  considered  the  most  important  and  obvious  characteris- 
tics, may  be  taken  as  typical  of  the  stems  of  the  great  majority 
of  the  trees  and  shrubs  of  our  forests,  and  some  of  the  modi- 
fications of  this  typical  structure  found  in  different  species 
have  also  been  noted.  It  must  be  mentioned,  however,  that  in 
some  species  there  is  a  remarkable  departure  from  the  type  in 
that  there  are  narrow,  more  or  less  concentric,  bands  of  soft, 
bark-like  tissue,  alternating  with  bands  of  woody  tissue 
throughout  the  stem.  These  are  found  for  example  in 
Dalbergia  paniculata  and  Cocculus  laurifolius.  A  similar 
structure  is  often  seen  in  the  woody  stems  of  climbers,  e.g.  in 
Bauhinia  Vahlii,  in  which  the  layers  of  porous  wood  alternate 
with  soft,  red,  bark-like  tissue. 

If  now  we  look  at  a  section  of  a  Deodar  stem  we  find, 
as  before,  a  pith,  distinct  bark,  and  wood  showing  both 
heart-  and  sap-wood,  while  in  the  wood  are  annual  rings  and 
medullary  rays.  There  is,  however,  a  great  difference  from  the 
Teak  wood  in  that  here  we  have  no  pores,  the  annual  rings 
being  marked  by  the  darker  colour  of  the  denser  wood  on 
the  outside  of  each  ring  contrasting  with  the  lighter  colour 
of  that  on  the  inside  of  the  rings.  This  absence  of  pores  is 
characteristic  of  the  wood  of  all  the  trees  and  shrubs  commonly 
known  as  Conifers.  In  some  conifers,  although  there  are  no 
Resin-o.uals.  pores,  there  are  wThat  are  called  resin-canals  or  -ducts,  which  on 
a  transverse  section  may  at  first  be  mistaken  for  true  pores. 
They  may  be  distinguished  by  their  irregular  outline  and  they 
are  usually  sparsely  distributed  through  the  wood.  They  are 
well  seen  on  a  transverse  section  of  Pinus  longifolia. 

If  now  we  look  at  the  section  of  a  Palm,  e.g.  Borassus 
flabellifer,  we  find  that  there  is  no  distinct  bark  which  can  be 
stripped  off  and  there  are  no  medullary  rays  or  annual  rings. 
Scattered  throughout  the  soft  tissue  of  the  stem,  we  find 
numerous  rounded,  hard,  black  areas,  in  the  interior  of  some 
Vascular  of  which  is  a  large  pore.  These  are  the  so-called  vascular 
bundles.  Those  situated  towards  the  exterior  of  the  stem  are 
usually  without  pores  and  are  closer  together  than  those 
towards  the  centre,  and  the  outer  portion  of  the  stem  is  conse- 
quently much  harder  and  denser  than  the  interior.  The  section 


19 

of  the  stem  of  a  Bamboo  is  somewhat  similar,  but  the  vascular 
bundles  are  not  so  conspicuous  and  the  interior  of  the  stem 
is  usually  hollow,  except  at  the  joints.  In  the  stem  of  a  Tree 
Fern  also  there  is  no  distinct  bark,  and  we  find  an  exterior  coat 
consisting  of  the  bases  of  fallen  leaves  and  adventitious  roots 
and  a  central  portion  of  soft  tissue,  which  often  disappears 
and  leaves  a  hollow  in  old  stems,  while  between  these  lies 
a  ring  of  harder  woody  tissue  which  contains  characteristic 
vascular  bundles,  usually  crescent-shaped  and  with  a  dark- 
coloured  border. 


20 


CHAPTEII  III.— THE  LEAF. 

20.  If  we  look  at  the  foliage  leaf 

of    say  the   Teak   tree  we  at  once  recognise   that  it   consists 
of  two  principal  parts  (1)  the  expanded  apical  portion  called 
the  blade  or  lamina   and  (2)    the    stalk,    or   petiole.     In  some 
Nerves.         plants   the   leaves   have   no   petiole   and   they  are   then   said 
to  be  sessile,  whereas  leaves  with  a  distinct  stalk  are   petiolate. 
On  looking  at  the  lamina   we   find  a   framework   of  firm  ribs 
traversing  its   surface    in  all    directions,    between  which   the 
soft    green   tissue   is   stretched,  much    like  the  cloth  over  the 
ribs  of  an  umbrella.  The  largest  and  most  prominent  of  these 
are  usually    distinguished   as  ribs,  nerves,    or   veins,  and    the 
smallest  as  veinlets.     For  the  present   we  may  include   them 
all  under  the    general  term   strands.     These    strands  of   firm 
tissue,    if   followed    up,    will  be    found    to   extend    from   the 
leaf  blade  through  the  petiole   and  stem  down  to  the  root,  the 
strands  in  the  leaf  being,  in  fact,   branches   of  similar  stouter 
strands  in  the  stem,  which   are  the  so-called  vascular  bundles. 
These  give  the  necessary  rigidity  and   strength  to  the    plant 
enabling  the  stem  to   stand   erect  and  the  flat  leaf-blades  to 
remain   extended    in  the  sunlight.     These  cords  of  tissue  also 
contain  minute  pipe-like  structures  which  will  be  considered  in 
detail  later,  but  through  some  of  which,  it  may  now  be   noted, 
water  with   mineral    salts   in  solution  passes  from  the  roots   to 
the  green  leaf,  while  others  bring  back  into  the  stem  the  food 
materials  manufactured  in  the  leaf  by  the    help  of  the  green 
chlorophyll  under  the  influence  of  sunlight,  and  carry    them 
to  those  points  where  they  are   required.     When    leaves    are 
cast  oft',  the  places  where  the  vascular  strands  pass  into    the 
stem  from  the  leaf  are  usually  distinguishable  on  the  resulting 
leaf -scars  as    more  or  less,  evident  dots,  or    rounded  marks. 
The  number,  shape,  and  arrangement  of   these  marks    on   the 
scar  are  often  very  characteristic. 

Veration.  21.  The  way  in  which  the  strands 

are  arranged  in  the  leaf  is  important  and  is  called  the  venation 
of  the  leaf.  Two  main  classes  of  leaves  are  usually  distin- 
guished, viz  :— 

(1)  Parallel -veined. 

(2)  Reticulate-  or  net-veined. 

In  parallel  venation  all  the  strands  which  can  be  easily  seen 


21 

with  the  naked  eye  run  approximately  parallel  to  one  another « 
They  usually  run  from  the  base  of  the  leaf  to  the  apex,  as  in  a 
Grass,  or  Bamboo.  The  term,  however,  is  also  applied  to  cases 
where  there  is  one  main  strand  traversing  the  leaf-blade  from 
base  to  apex,  called  the  midrib,  and  from  which  all  other 
noticeable  strands  run  approximately  parallel  to  each  other 
towards  the  leaf-margin,  as  in  a  Banana  leaf.  In  both  cases  a 
few  minute,  inconspicuous,  and  generally  straight,  strands 
usually  connect  the  main  strands  and  run  at  right  angles  to 
them.  These  are  conspicuous  in  the  leaves  of  Arundinaria 
spathiflora.  In  reticulate  venation  the  strands  are  seen  to 
branch  in  all  directions  and  thus  give  rise  to  an  elaborate  net- 
work, as  in  a  Teak  leaf.  When  there  is  a  distinct  midrib  from 
which  all  the  other  nerves,  directly  or  indirectly,  spring,  the 
nerves  which  spring  from  the  midrib  are  called  lateral  nerves 
and  the  leaf  is  said  to  be  penninerved,  or  pinnately-veined.  In 
such  cases  also  the  midrib  is  said  to  be  the  primary  nerve,  the 
larger  nerves  springing  from  the  midrib  are  called  secondary 
nerves,  and  the  smaller  nerves  springing  from  the  latter  the 
tertiary  nerves. 

In  some  cases  in  this  type  of  leaf,  the  lower  pair  or  two  of 
lateral  nerves  are  much  more  prominent  than  the  remainder, 
as  in  Cinnamomum  Camphora,  see  Fig.  5,  Plate  IV.  Such 
cases  bring  us  by  insensible  gradations  to  the  next  type  of 
venation  in  which  there  is  more  than  one  main  rib,  or  primary 
nerve,  entering  the  base  of  the  blade  and  the  leaf  is  then  said 
to  be  palminerved,  or  palmately-  or  digitately -veined.  Accord- 
ing to  the  number  of  the  strong  basal  nerves  the  leaf  is  said  to 
be  palmately — or  digitately — 5  nerved,  7  nerved  and  so  on. 

Nerves  may  be  straight  or  more  or  less  curved.  Those 
which  are  slightly  bent  in  the  form  of  a  bow  and  run  in  a 
regular  sweeping  curve  are  called  arcuate,  those  which  are 
more  sharply  bent  are  arched.  A  leaf  of  Quercus  incana  may 
be  taken  as  an  example  of  a  penninerved  leaf  with  straight- 
secondary  nerves,  see  Fig.  1,  Plate  IV.  Such  a  leai  differs 
from  a  typical  parallel-veined  leaf,  such  as  that  of  the  Banana, 
in  having  the  secondary  veins  further  apart  and  the  spaces 
between  them  rilled  with  distinctly  reticulated  strands.  A 
leaf  of  Acer  caesium,  Fig.  2,  Plate  IV,  furnishes  an  example 
of  a  palminerved  leaf  with  straight  primary  nerves.  The 
penninerved  leaf  of  Cornus  macropliylla,  Fig.  3,  Plate  IV, 
has  arcuate  secondary  nerves.  The  palminerved  leaf  of 
Smilax  parvifrtia,  Fig.  4,  Plate  IV.  has  arcuate  primary  nerves. 
By  some  botanists  the  term  palminerved  is  restricted  to  cases 


22 

in  which  the  primary  nerves  are  practically  straight;  when 
they  are  curved,  the  leaf  is  merely  described  as  having  5, 
7,  or  more,  basal  nerves.  When  the  secondary  nerves  are 
curved  each  one  frequently  runs  into  and  joins  the  next  secon- 
dary above  it,  the  junction  taking  place  near  the  leaf  margin. 
A  continuous  strong  nerve,  composed  of  the  ends  of  the 
secondary  nerves,  is  thus  formed  close  to  the  leaf  margin. 
This  is  called  a  marginal,  or  intramarginal,  nerve  and  may 
be  seen  in  Ficus  bengalensis,  or  Eugenia  Jambolana.  Such  a 
nerve  prevents  the  leaf  blade  from  being  easily  torn.  A 
Banana  leaf  has  no  such  nerve  and  is  quickly  torn  into  strips 
by  the  wind.  If  this  were  not  so  the  enormous  leaf  blade 
would  offer  great  resistance  to  the  wind  and  the  plant's 
tissues  would  be  subjected  to  a  dangerous  strain.  As  a  rule 
a  midrib  divides  the  leaf  blade  into  two  practically  equal  divi- 
sions. When  these  divisions  are  unequal  the  leaf  is  said  to 
be  oblique,  as  is  the  case  in  Ficus  Cunia.  When  nerves 
continually  fork  and  divide  into  two  branches  of  approxi- 
mately equal  size,  the  venation  is  called  furcate  and  is  charac- 
teristic of  many  Ferns,  see  Fig.  6,  Plate  IV. 

Leaf  margin.  22.  The  margin  of  a  leaf  is  said 

to  be  entire  when  it  is  an  even  line  not  indented  in  any 
way,  serrate  when  with  sharp  teeth  directed  towards  the 
leaf  apex,  bi-serrate  when  each  main  tooth  is  again  serrated, 
serrulate  when  serrate  with  very  small  teeth,  ruminate 
when  serrate  with  the  teeth  directed  backwards,  dentate  or 
toothed  when  the  teeth  are  triangular  and  directed  out- 
wards, crenate  when  with  rounded  teeth,  repand  when  it  is 
a  gently  undulating  line,  sinuate,  or  undulate,  when  the 
undulations  are  more  pronounced.  When  the  margin  is 
fringed  with  fine  and  close-set  hairs  it  is  ciliate,  and  when  it 
is  cut  into  a  number  of  long  narrow  segments  it  is  fimbriate. 
W'hen  the  incisions  are  deeper,  but  do  not  extend  more 
than  half-way  to  the  midrib  in  the  case  of  a  penninerved 
leaf,  or  to  the  base  of  the  blade  in  the  case  of  a  palminerved 
leaf,  the  leaf  is  said  to  be  lobed  or  cleft,  according  as  the  leaf 
divisions,  or  the  spaces  between  them,  are  broad  and  rounded 
or  narrow  and  acute.  WThen  the  incisions  extend  still  deeper, 
the  leaf  is  said  to  be  partite,  or  parted,  and  when  they  reach 
the  midrib  or  base  of  the  blade,  thus  dividing  the  leaf  into 
distinct  parts,  the  leaf  is  said  to  be  divided.  In  this  case  the 
parts  so  divided  off  are  called  segments  and  they  cannot  be 
separated  from  the  midrib  or  petiole  without  the  lamina  being 
torn.  When  each  of  these  divisions  has  a  separate  insertion 


'23 

of  its  own,  there  being  no  trace  of  lamina  at  the  place  of 
insertion,  and  when  each  of  them  can  he  separated  from  the 
common  leaf -stalk  without  tearing,  just  as  the  whole  leaf  may 
be  separated  from  the  stem,  the  leaf  is  known  as  a  compound 
leaf,  in  contra-distinction  to  a  simple  leaf  which  is  not  thus 
divided.  The  parts  of  a  simple  leaf  blade  are  known  as  Simple  and 
segments,  or  lobes,  while  each  separate  division  of  a  com-  Compound 
pound  leaf  is  called  a  leaflet.  The  terms  given  above  for  the 
description  of  a  simple  leaf,  as  to  its  venation,  margin  and  so  on, 
are  equally  applicable  to  each  leaflet  of  a  compound  leaf. 
Each  leaflet  may  be  sessile  or  provided  with  a  stalk,  the  latter 
being  called  a  petiolule.  The  various  shapes  of  the  leaf  blade 
are. caused  by  portions  of  the  leaf  growing  faster  than  others, 
and  we  get  first  an  uneven  margin,  then  a  lobed,  cleft,  parted, 
and  finally  a  divided,  or  compound  leaf.  As  in  such  cases  the 
growth  in  the  direction  of  the  main  nerves  is  usually  more 
vigorous  than  that  of  the  portions  of  the  blade  between  them, 
a  pinnately-nerved  leaf  gives  rise  to  a  pinnately-lobed, — 
cleft, — parted, — divided,  or  compound  leaf,  and  a  palmately- 
nerved  leaf  to  a  palmately, — lobed, — cleft, — parted, — divided,, 
or  compound  leaf.  Pinnately-  and  palmately -cleft  are  synony- 
mous  with,  and  often  replaced  by,  the  terms  pinnatifid  and 
palmatifid  respectively 

Similarly   the     terms    pinnately — and    palmately — divided 
are  synonymous  with  the  terms  pinnatisect  and  palmatisect. 

23.  The  following  terms  are  most  General 

frequently  employed  for  describing  the  general  shape   of  the  shape  of 
blade  of  a  leaf  or  leaflet.     They   are  of    course   applicable   to Leaves- 
any    plane  surface  and    so  may   be  used    for   other  organs 
besides   leaves.     As   they    refer    to    plane   surfaces   with   an 
entire  margin,  in   order  to  describe  the   general  shape  of   a 
leaf  the   margin  of   which  is    not    entire,   it  is  usual   to  con- 
sider the    general   shape   as    defined  by  an   imaginary    line 
passing  through  the    base   and   apex   of  the  leaf-blade    and 
touching  the  summits  of   all   the    principal   teeth,    lobes,   or 
segments,  of  the  lamina. 

1.  Acicular ;    needle-shaped,    very  much    longer  than 

wide  and  tapering  to  a  point,  such  as  a  Pine  leaf. 

2.  Linear ;  several  times  longer  than  wide  with  almost 

parallel  sides. 

3.  Subulate  ;  awl-shaped,  acicular  with  a  broad  base. 

4.  Lanceolate ;  like  a  spear-head.    Several  times  longer 

than  wide  but  tapering   to  both  ends,  with  the 
•'.    -  greatest  width  below  the  centre. 


24 

5.  Oblong ;   two  or  three  times  as  long  as  broad  with 

almost  parallel  sides. 

6.  Elliptical;  oblong  with  a  regularly  curved  outline. 

7.  Oval ;  broadly    elliptic.    The    width  more  than  half 

the  length. 

8.  Ovate ;  like  the  outline   of  an  egg  with  the   broader 

end  at  the  base. 

9.  Orbicular  ;  circular  or  nearly  so. 

10.  Cuneate  ;  wedge-shaped.     Broad  at  apex  and  narrow 

at  base. 

11.  Deltoid  or    triangular  ;    cuneate    with  the  width  at 

the  base. 

12.  Spatulate  ;  spoon-shaped.    Rounded  above,  long  a.nd 

narrow  below. 

13.  Reniform ;  kidney- shaped. 

14.  Falcate  ;  sickle-shaped.     Curved  like  the    blade  of  a 

sickle,  or  scythe. 

For  describing  the  extremity  of  the  leaf,  whether  the 
base  or  apex,  the  following  terms  are  most  commonly  used, 
1  to  9  being  generally  applied  to  the  apex  and  10  to  13  to 
the  base. 

1.  Acuminate  ;  with  a  long  tapering  point. 

2.  Acute  :  ending    in    an  acute   angle,  the  point  not 

being  prolonged. 

3.  Obtuse  :  with  a  blunt  or  rounded  apex. 

4.  Truncate  :  ending  abruptly  as  if  with    the  end  cut 

off  by  a  straight  line. 

5.  Retuse  :  with  a  shallow  notch  in  a  rounded  apex. 

6.  Emarginate  :  with  a  decided  terminal  notch. 

7.  Mucronate  :  ending  abruptly  in  a  short,  stiff,  sharp 

point. 

8.  Cuspidate  :    ending  in   a  long,  tapering,  stiff,  sharp 

point. 

9.  Caudate  :  ending  in  a  long  slender  tail. 

10.  Cordate  :    when    of    two  broad   rounded   lobes  one 

on  either  side  of  a  deep  notch. 

11.  Aurided  :  when  of  tw  o   narrow   rounded  lobes    one 

on  either  side  of  a  deep  notch, 

12.  Sagittate :    when   the    lobes    on    each   side   of   tho 

notch  are  pointed  and  directed   downwards. 

13.  Hastate :  when  the  lobes  on  each  side  of  the   notch 

are  pointed  and  directed  outwards. 


25 

An  ovate  leaf  which  is  cordate  at  the  base  and  pointed  at 
the  apex  is  called  a  cordate  leaf ;  similarly  a  leaf  which  is 
sagittate,  or  hastate,  at  the  base  and  pointed  at  the  apex  is 
called  a  sagittate,  or  hastate,  leaf.  If  in  a  lanceolate,  ovate, 
or  cordate  leaf  we  imagine  the  petiole  to  be  attached  to  the 
apex  instead  of  to  the  base,  we  get  an  ob-lanceolate,  ob-ovate, 
or  ob-cordate  leaf,  respectively. 

A  peltate  leaf  is  one  in  which  the  petiole  instead  of  being 
attached  to  the  margin  of  the  leaf-blade  is  joined  to  some  part 
of  its  under-surface,  so  that  the  leaf-blade  is  more  or  less  at 
right-angles  to  the  petiole,  as  in  Nelumbium  speciosum. 

If  in  a  deeply  cordate  leaf  we  imagine  the  basal  lobes  to 
grow  together  and  join  in  front  of  the  petiole  we  get  a  peltate 
leaf ;  if  now  such  a  cordate  leaf  was  sessile  on  the  stem  and  the 
same  thing  happened,  the  stem  would  appear  to  pass  through 
the  leaf.  Such  a  leaf  is  said  to  be  per/oliate  and  the  base 
of  the  leaf  which  thus  embraces,  or  clasps,  the  stem  is  said 
to  be  amplexicaul.  If  the  base  only  partly  surrounds  the  stem 
it  is  semi-amplexicaul.  Occasionally  the  bases  of  two  opposite 
sessile  leaves  grow  together  and  are  then  said  to  be  connate.  In 
some  cases  the  leaf-blade  is  continued  along  the  stem  below 
the  leaf  insertion  and  the  leaf  which  thus  seems  to  run  down 
the  stem  is  called  decurrent,  as  seen  in  Verbascum  Thapsus. 
Pinnately — and  palmately — compound  leaves  are  usually  known 
as  pinnate  and  palmate  leaves,  respectively. 

The  prolongation  of  the  petiole  of  a  pinnate  leaf,  which 
corresponds  to  the  midrib  of  a  pinnately-veined  simple  leaf,  is 
called  the  rhachis. 

When  the  rhachis  terminates  in  an  odd  leaflet  the  leaf  is 
said  to  be  impari — ,  or  odd — pinnate  and  when  there  is  no  termi- 
nal leaflet  it  is  pari — ,  or  abruptly — pinnate. 

The  leaflets  of  a  pinnate  leaf  are  usually  opposite  to  each 
other  in  pairs,  but  sometimes  they  are  alternate  on  the  rhachis. 
The  impari-pinnate  leaves  of  Sissoo  have  alternate  leaflets. 
When  the  leaflets  of  a  pinnate  leaf  vary  greatly  in  size  the 
leaf  is  interruptedly-pinnate.  When  the  terminal  leaflet  or 
pair  of  leaflets  is  largest,  those  below  it  gradually  decreasing 
in  size  with  the  smallest  at  the  base,  the  leaf  is  lyrately  pinnate, 
as  are  the  leaves  of  Picrasma  quassioides. 

WThen  the  rhachis  of  a  pinnate  leaf,  instead  of  giving  rise 
directly  to  leaflets,  develops  branches  on  which  the  leaflets 
are  borne,  these  branches  are  termed  pinnae  and  the  whole 
leaf  is  said  to  be  bi-pinnate,  such  as  are  the  leaves  of  Khair 
(Acacia  Catechu).  When  pinna3,  instead  of  bearing  the  leaflets 


26 

directly,  also  develop  branches  on  which  the  leaflets  arise,  these 
secondary  branches  are  termed  pinnules  and  the  leaf  is  said 
to  be  tn-pinnate,  as  are  often  the  leaves  of  Moringa  pterygosperma. 
A  compound  leaf  is  usually  described  by  the  number  of  leaflets, 
thus  J}i~,  tri-,  quadri-foliolate  and  so  on. 

It  is  not  always  easy  at  first  sight  to  distinguish  between 
a  pinnately-and  palmately-trifoliolate  leaf,  but  on  close  in- 
spection it  will  be  seen  that  in  the  former  the  leaflets  do  not 
all  spring  from  the  same  point — see  the  pinnately  3-foliolate 
leaf  of  Desmodium  tilicefolium,  Fig.  1,  Plate  F,  in  which  an 
obvious  rhachis  extends  from  the  point  of  insertion  of  the 
pair  of  lateral  leaflets  to  that  of  the  terminal  leaflet.  The 
junction  between  the  rhachis  and  the  base  of  the  terminal 
leaflet,  or  of  its  petiolule,  is  often  marked  by  a  distinct  joint 
or  articulation ;  sometimes  also  the  petiolule  of  the  terminal 
leaflet  is  swollen  and  is  thus  easily  distinguished  from  the 
rhachis.  The  leaf  of  Desmodium  tiliaefolium  may  be  compared 
with  the  palmately  5-foliolate  leaf  of  Holboellia  latifolia  in 
which  all  the  leaflets  spring  from  a  distinct  joint  at  the  apex  of 
the  petiole,  Fig.  2,  Plate  V. 

It  is  not  always  easy  to  at  once  decide  whether  foliar 
structures  are  leaves  or  leaflets.  If  we  look  at  a  branch  of 
Phyllanthus  Emblica  we  find  the  small  leaves  arranged  in  two 
ranks  along  the  twigs  giving  the  latter  the  appearance  of 
pinnate  leaves.  Minute  buds  may,  however,  be  often  found 
in  the  axils  of  these  small  leaves,  thus  indicating  that  they 
are  true  leaves,  no  buds  being  normally  produced  in  the  angle 
between  a  leaflet  and  the  axis  from  which  it  springs.  Moreover 
flowers  are  found  on  these  leaf-bearing  twigs  thus  showing  that 
the  latter  aje  branches  of  the  stem  and  not  petioles,  rhachises, 
or  branches  of  them,  on  which  no  flowers  normally  arise. 

A  leaf  which  is  at  first  palmately  3-nerved  but  in  which 
each  of  the  lateral  nerves  is  forked,  the  outer  branch  of  each 
fork  sometimes  again  forking,  is  said  to  be  pedately -nerved. 
Such  a  leaf  may  be  lobed,  cleft,  parted,  divided  or  compound, 
just  as  is  the  case  with  an  ordinary  palminerved  leaf,  and  in 
the  last  case  we  get  a  pedately -compound  leaf,  which  is  usually 
simply  called  a  pedate  leaf.  See  Fig.  7,  Plate  IV. 

The  principal  terms  which  are  usually  employed  for 
describing  the  general  shape  of  leaves  have  now  been 
enumerated  above,  but  it  must  be  remembered  that  the  forms 
of  leaves  are  infinitely  various  and  that  it  is  neither  possible 
nor  desirable  to  have  separate  names  and  definitions  for  every 
form  that  may  be  met  with.  Here,  as  elsewhere,  morphology 


27 

selects  and  describes  types,  and  we  must  expect  to  find  a 
number  of  forms  intermediate  between  the  types.  For  each 
intermediate  form  we  employ  the  term  which  seems  to  describe 
it  most  correctly,  and  in  many  cases  a  combination  of  terms, 
such  as  ovate-lanceolate,  oblong -lanceolate,  and  so  on,  are  found 
to  be  most  suitable.  The  student  should  collect  leaves  of 
various  plants  and  endeavour  to  frame  terse  descriptions  of 
the  same,  so  that  another  person  from  the  descriptions  may 
be  able  to  picture  to  himself  the  exact  form  of  the  leaves  in 
question.  On  one  and  the  same  plant  all  the  leaves  are  not 
exacthr  the  same  size  or  shape,  and  hence,  when  describing  the 
leaves  of  the  plant,  the  extremes  must  be  given,  such  as 
:'  varying  from  ovate  to  elliptic,  length  of  blade  4-6  inches, 
breadth  2-3  inches." 

24.  When  the    size  and  shape   of  Polymorphic 

the   leaves   on  one    and  the  same  plant  vary  very  consider-  Leayes- 
ably  the  plant  is  said  to  be  heterophyllous  and  the  leaves   are 
said  to  be  polymorphic.      This  is    the    case  in    many  climbing 
plant  ssuch  as  Ficus  pumila,  the  Ivy  and  others,  see  Figs.  1 
and  2,  Plate  VI. 

To  understand  this  phenomenon  we  must  remember  that 
the  function  of  the  green  leaves,  viz.  that  of  manufacturing 
food  for  the  plant,  can  only  be  properly  performed  when  the 
green  leaf-blades  are  exposed  to  the  sunlight  and  cannot  be 
carried  out  in  the  dark,  and  that  the  amount  of  food  which  can 
be  manufactured  depends  on  the  amount  of  green  surface 
exposed  to  the  light.  Creeping  stems  which  are  often  found  in 
shady  places  on  the  ground,  or  on  the  trunks  of  other  trees, 
must  make  the  most  of  the  faint  light  at  their  disposal  and 
must  keep  as  large  a  surface  of  green  leaf-tissue  exposed  to 
the  light  as  possible.  We  thus  find  beautiful  arrangements  of 
leaves  of  very  various  size  and  shape,  small  leaves  occupying 
the  spaces  between  the  large  ones  and  the  lobes  and  segments 
of  some  leaves  fitting  into  the  spaces  between  similar  lobes 
and  segments  of  adjoining  leaves.  Each  leaf  is  thus  arranged 
so  as  to  shade  its  neighbours  as  little  as  possible  and  no  light 
is  allowed  to  pass  through  on  to  the  substratum  below  with- 
out being  utilised. 

Such  arrangements  of  leaves  are  called  leaf-mosaics.  When 
such  creeping  stems  send  up  erect  serial  stems  standing  away 
from  the  substratum  covered  by  the  leaf-mosaic  and  which 
ultimately  give  rise  to  flowers  and  fruit,  the  leaves  borne  by  such 
stems  usually  differ  considerably  from  those  on  the  creeping  stems 
and  are  more  uniform  in  size  and  shape,  see  Fig.  2,  Plate  VI. 


Leaves  of 

Young 

Plants. 


Other 
Characters 
of  Leaves. 


Such  aerial  stems  are  usually  surrounded  by  brighter  light 
than  are  the  creeping  stems  and  their  leaves  are  exposed  not 
only  to  overhead  light,  but  also  to  considerable  lateral 
illumination,  so  that  in  their  case  the  partial  shading  of  the 
lower  leaves  by  the  upper  ones  is  not  so  important,  the  loss  of 
overhead  light  being  compensated  by  the  addition  of  lateral 
light.  Moreover  as  in  such  stems  plenty  of  room  is  available 
to  enable  the  leaves  to  develop  at  different  levels  it  is  not  so 
necessary  that  the  leaves  arising  at  any  one  level  should  fit 
together  closely  and  accurately,  for  the  light  which  passes 
through  the  interstices  of  the  upper  leaf-layers  may  be  received 
and  utilised  by  the  lower  leaves  and  is  not  therefore  lost  to 
the  plant.  Hence  in  these  shoots  leaves  of  such  size  and  shapa 
as  are  necessary  to  form  a  close-fitting,  more  or  less  flat, 
mosaic  are  not  required. 

Although  hete^ophylly  is  one  of  the  expedients  resorted  to 
by  plants  in  order  to  have  as  large  a  leaf  surface  as  possible 
exposed  to  the  light,  it  is  by  no  means  the  only  one,  and  in  other 
plants  the  same  end  may  be  attained  in  a  variety  of  ways. 

25.  The   leaves  of   seedlings  and 

young  plants  usually  differ  considerably  from  those  of  mature 
plants.     Fig.    5,   Plate  XII,  shows   a    seedling    of    Oroxylum 
indicum  and  indicates  the  difference  which  may  exist  between 
the    first   leaves    and    those  subsequently    developed,  for  the 
leaves  there  shown    bear    not  the    least    resemblance  to  the 
enormous    bi-    and    tri-pinnate    leaves    of    the  mature    tree. 
The  leaves  of  young  pfants   of    Quercus   semecarpifolia   have 
spines  and  are    toothed,  while   on   old  trees  they  are  usually 
entire,   and   in  Gardenia   turgida   the  leaves   of  young  plants 
are  quite  unlike  those  of  older  plants.     In  species  of  Phcenix 
the   leaves    of   young  plants    are    usually    entire  while  those 
of  mature  plants  are  pinnate.     Similarly  the  leaves  on  coppice 
shoots    often    differ  considerably    in    size    and    shape    from 
those   on  uninjured  plants.      On    coppice    shoots    of    Pyrus 
Pashia  the  leaves  are  often  lobed  or   cleft,  whereas  they  are 
normally  crenate.     This  point  is  one  of  practical   importance 
for  Foresters,   who,   unless   they   are   able   to  recognise  their 
trees  and  shrubs  at  different  stages  of  their  life-history,  can 
know  very  little  about  the  reproduction  of  their  forests. 

26.  Leaves    may     be    glossy    or 

shining,  like  those  of  Cocculus    laurifolius,   or  dull    and   not 
shining  like  those   of   Euonymus  Hamiltonianus  and  in  this, 
as  in  other  respects,  the  upper  and  lower  surface   of  the  leaf- 
blade   may    differ    considerably.     As    regards  colour,   mature 


29 

leaves  are  usually  of  some  shade  of  green  and  the  lower  sur-  Colour, 
face  is  frequently  paler  than  the  upper.  The  colour  of  young 
leaves  is  sometimes  characteristic  and  is  often  of  some  shade 
of  red  or  purple.  As  examples  we  may  note  the  pinkish,  or 
purplish,  young  leaves  of  Quercus  incana,  the  bright  red  ones 
of  Acer  caesium,  the  beautiful  purple  and  brown  of  those  of 
Mango  and  the  dark  red  brown  of  Cassia  Fistula.  Mature 
leaves  frequently  undergo  a  striking  change  of  colour  before 
they  fall  off  the  plant.  Such  colours  are  usually  called 
autumn  tints  and  often  enable  the  Forester  to  recognise  his 
trees  from  a  long  distance.  As  examples  we  may  note : 

The  yellow  of  Odina  Wodier,  the  pale  leather-brown  of 
Lagerstrcemia  parviflora,  the  brick -red  of  Antidesma  diandrum, 
the  beautiful  red,  purple,  and  orange  of  Sapium  sebiferum,  and 
the  dark  red,  or  bronze,  of  Anogeissus  latifolia.  Some  leaves, 
on  drying,  undergo  a  characteristic  change  of  colour  ;  those  of 
Dalbergia  paniculata  turn  black  and  those  of  many  species  of 
Symplocos  turn  bright  yellow. 

Many  leaves  have  a  strong  and  characteristic  smell  especially  Smell, 
when  they  are  crushed.  As  examples  may  be  noted  the  un- 
pleasant smell  of  leaves  of  Viburnum  foetens,  Premna  latifolia 
and  Solanum  verbasci folium,  the  aromatic  odour  of  those  of 
Skimmia  Laureola  and  Cinnamomum  Tamala,  the  black-cur- 
rant-like smell  of  Pogostemon  plectranthoides  and  the  well-known 
smell  of  TJiymus  Serpyllum. 

The  taste  of  leaves  is  also  often  peculiar.     The  leaves  of Taste- 
Eauhinia    malabarica,   Acacia  pennata   and   Antidesma  dian- 
drum, for  example,  have  a  characteristic  acid  taste. 

Leaves  which  are  strongly  aromatic  frequently  have  the  Glands, 
ethereal  oil,  to  which  the  smell  is  due,  stored  in  small  recep- 
tacles called  glands  in  the  leaf  tissue.  Such  glands,  being 
translucent,  are  easily  seen  when  the  leaf  is  held  up  to  the 
light  as  pale  spots.  Such  leaves  are  said  to  be  gland-dotted. 
As  examples  may  be  taken  a  leaf  of  the  Orange,  or  of  Zan- 
thoxylum  alatum. 

The  texture  of  leaves,  as  well  as  excrescences,  such  as 
hairs,  scales  and  so  on,  which  commonly  appear  on  other  parts 
of  the  plant  besides  the  leaves,  will  be  considered  later,  when 
the  detailed  account  of  each  member  has  been  completed. 

27.  The    principal   points   to  be  p™™jfrs  of 
paid  attention  to  as   regards  the  petiole   of  the  leaf  are  its 
lengthy  which  varies  greatly  in  different  plants,   and  its  gen- 
eral shape.     In  some  plants  it  is  very  slender  and  thread-like 
and   is  said   to  be  filiform,  in    others  it  is  stout.     Like  the 


30 

stem,  the  petiole  may,  or  may  not,  be  terete,  flattened, 
angled,  grooved,  striate,  and  so  on.  Very  frequently  it  is 
channelled,  or  grooved,  on  the  upper  surface  as  in  Viburnum 
cotinifolium,  see  Figs.  8  and  9,  Plate  IX. 

In  some  cases  the  lamina  is  continued  along  the  sides  of 
the  petiole  which  thus  appears  to  be  winged,  as  seen  in  the 
Orange. 

Leaf-Base.  28.  In   some   cases  [the   base   of 

the  leaf  is  more  or  less  clearly  differentiated  from  the  rest 
of  the  leaf  and  in  many  plants  it  becomes  a  swollen  cushion  of 
tissue  called  the  pulvinus,  well  seen  for  example  at  the  base 
of  the  petiole  of  Millettia  auriculata,  and  the  small  pulvinus 
often  found  at  the  base  of  leaflets  is  called  a  pulvinule. 

.  This  differentiation,  however,  is  particularly  remarkable  in 
many  Bamboos  in  which  the  leaf-base  is  large  and  forms  a 
long  sheath  to  the  stem,  the  petiole  of  the  lamina  being 
separated  from  the  sheath  by  a  distinct  joint  or  articulation. 
The  sheaths  which  arise  directly  from  the  Bamboo  culms, 
called  the  culm -sheaths,  in  the  axils  of  which  the  branches 
arise,  usually  develop  no  leaf -blade,  or  only  an  imperfect  one, 
but  all  gradations  between  sheaths  with  no  blade  and  those 
with  a  normal  lamina  may  frequently  be  seen. 

This  sheath  is  very  characteristic  of  Bamboos  and  Grasses 
generally,  as  is  also  the  peculiar  structure  known  as  the  ligule, 
a  membranous  outgrowth  developed  from  the  inner  face  of 
the  leaf -sheath  at  the  junction  of  the  lamina,  or  petiole,  with 
the  leaf-sheath,  and  which  may  be  well  seen  in  Arundinaria 
falcata. 

In  Berberis  Lycium  a  distinct  joint  is  found  between  the 
leaf-base  and  the  lamina  of  the  apparently  simple  leaf,  see  Fig. 
4,  Plate  V.  In  Berberis  nepalensis,  however,  a  plant  which 
in  important  characters  resembles  B.  Lycium  and  which  there- 
fore, being  considered  to  be  fairly  closely  related  to  it,  bears 
the  name  of  Berberis,  the  leaves  are  pinnate  and  there  is  a 
distinct  joint  between  the  terminal  leaflet  and  the  rest  of 
the  leaf  and  at  the  insertion  of  the  lateral  leaflets,  see 
Fig.  3,  Plate  V ;  hence  it  is  concluded  that  the  leaf  of  B. 
Lycium  is  really  a  compound,  i.e.  pinnate,  leaf,  reduced  to 
the  terminal  leaflet.  The  apparently  simple  leaf  of  the 
Orange  is  also  articulated  to  the  petiole  and  is  considered  to  be 
a  unifoliolate  compound  leaf. 

Stipules.  29.  The  leaf -base  frequently   de- 

velops   two    more    or    less  noticeable   lateral  branches  called 


31 

stipules,  there  being  normally  two  stipules  to  each  leaf, 
one  on  each  side  of  the  petiole.  These  are  very  large 
and  leaf-like  in  Albizzia  stipulate  and  the  common  Pea ; 
sometimes  they  are  like  small  bristles  and  are  said  to  be 
setaceous  and  in  Capparis  spinosa  they  are  developed  as 
thorns.  Stipules  frequently  fall  off  shortly  after  their  first 
appearance  and  often  before  the  leaf  to  which  they  belong 
is  fully  developed,  as  in  Holoptelea  integrifolia.  In  falling, 
however,  more  or  less  noticeable  scars  are  left  behind  and  by 
observing  these  it  can  be  known  that  stipules  were  developed 
and  an  idea  formed  of  their  shape  and  size.  The  large  stipules 
of  many  species  of  Ficus  for  instance  leave  ring-like  scars. 

In  Rosa  the  stipules  are  adnate  to  the  petiole.  In  some 
cases  where  the  leaves  are  opposite  a  stipule  of  one  leaf  be- 
comes united  to  the  stipule  which  is  opposite  to  it  and  which 
belongs  to  the  other  leaf  on  the  node,  there  thus  being 
apparently  two  large  stipules  at  each  node,  situated  between 
the  petioles  of  the  leaves.  Such  a  stipule  is  called  an  inter- 
petiolar  stipule  and  is  well  seen  in  Stephegyne  parvifolia,  where 
each  large  stipule  can  be  easily  separated  into  its  component 
stipules  along  their  line  of  junction,  see  Fig.  9,  Plate  VIII. 

In  some  plants  in  which  there  is  only  one  leaf  at  each 
node,  the  stipules  cohering  by  their  outer  edges,  entirely 
embrace  the  stem  and  there  is  apparently  one  stipule  opposite 
the  leaf.  In  some  cases  the  stipules  cohere  both  by  their 
outer  and  inner  margins  and  thus  form  a  complete  tubular 
sheath  around  the  stem,  called  an  ocrea,  as  seen  in  many 
species  of  Polygonum. 

The  leaflets  of  a  compound  leaf  often  have  stipules  which 
are  then  called  stipels.  These  may  be  seen  in  Desmodium 
tiliaefolium,  Fig.  1,  Plate  V. 

Leaves  which  produce  stipules  are  said  to  be  stipulate  and 
those  which  do  not  are  exstipulate.  Similarly  leaflets  which 
produce  stipels  are  described  as  stipellate. 

•  30.  True  leaves,  like  other  organs,  Metamor- 
are  often  found  to  be  metamorphosed,  or  altered  in  form,  in 
order  that  they  may  be  able  to  carry  out  other  work  than 
that  usually  performed  by  leaves.  In  Gloriosa  superba  the 
apex  of  the  leaf-blade  is  prolonged  and  becomes  a  tendril,  in 
the  common  Pea  some  of  the  terminal  leaflets  are  trans- 
formed into  tendrils,  in  Clematis  montana  the  petiole  coils 
round  supports  and  serves  as  a  climbing  organ,  in  Berberis 
Lycium  the  entire  leaf  is  often  transformed  into  a  three- 


32 

pronged  spine  and  on  one  and  the  same  shoot  intermediate 
forms  between  the  typical  spine  and  normal  foliage  leaf  can 
often  be  found,  see  Fig.  3,  Plate  VI. 

That  these  spines  are  really  leaves  is  indicated  by  the  fact 
that  buds  and  branches  arise  in  their  axils  just  as  they  do  in 
the  axils  of  true  leaves  ;  moreover  the  base  of  the  spines  is 
furnished  with  minute  stipules  j  list  as  are  those  of  the  normal 
leaves. 

31.  The  way  in  which  the   leaves 

are  arranged  upon  the  stem  is  called  phyllotaxy,  or  phyttotaxis. 
Some  of  the  arrangements  commonly  found,  such  as  opposite 
leaves,  alternate  leaves,  and  so  on,  have  already  been  noted 
above,  but  it  is  usual  to  distinguish  some  additional  types  of 
the  alternate  arrangement  by  fractions,  such  as  ^,  f,  f  and 
so  on.  To  determine  the  phyllotaxy  of  a  plant  one  leaf  is 
selected  on  the  stem  as  a  starting  point  which  we  may 
call  (a)  and  then,  passing  up  the  stem,  we  note  the  first 
leaf  which  is  situated  exactly  vertically  above  it  which  may 
be  called  (b).  A  spiral  line  is  then  traced  from  the  start- 
ing point  (a),  passing  the  shortest  way  around  the  stem 
through  the  insertions  of  all  the  leaves  situated  between  (a) 
and  (b)  in  consecutive  order  and  ending  at  (b).  If  this  line  is 
found  to  have  passed  twice  around  the  stem  and  to  have 
passed  through  5  leaves,  omitting  (a)  but  including  (6),  the 
phyllotaxy  is  said  to  be  -f ;  if  the  line  has  passed  three  times 
around  the  stem  through  8  leaves  it  is  f  and  so  on.  When 
alternate  leaves  are  distichous,  therefore,  the  phyllotaxy  is  ^ 
and  when  they  are  tristichous,  i.e.  in  three  vertical  ranks,  it  is  |-. 

The  phyllotaxy  occasionally  varies  on  one  and  the  same 
plant ;  thus  on  erect  branches  of  Coriaria  nepalensis  the  leaves 
are  decussate  in  4  vertical  ranks,  whereas  on  horizontal  branches, 
by  a  twisting  of  the  stem,  they  become  distichous  and  the 
leaves  are  also  twisted  so  that  their  blades  are  practically 
horizontal,  i.e.  with  the  leaf-surface  perpendicular  to  the 
direction  of  the  rays  of  light  coming  from  above,  see  Plate 
XIII. 

We  have  noted  already  how  important  it  is  for  plants  to 
secure  a  suitable  degree  of  illumination  for  their  green  leaf- 
surface,  and  this  necessity  is  to  a  great  extent  responsible  for 
the  different  forms  of  leaves  and  for  the  way  in  which  they 
are  arranged  upon  the  stem.  It  must,  however,  always  be 
remembered  that  all  plants  have  not  identical  requirements, 
that,  for  instance,  the  most  suitable  degree  of  illumination  for 
one  plant  may  be  injurious  to  another,  and  secondly  that  the 


33 

same  object  may  be  attained  by  different  plants  in  entirely 
different  ways.  Thus  one  plant  may  obtain  the  best  illumi- 
nation for  its  foliage  b}T  heterophylly,  another  by  adopting 
a  different  arrangement  of  the  leaves  upon  the  stem,  or  by 
varying  the  lengths  of  the  petioles,  another  by  adopting  a 
climbing  habit,  and  so  on. 

32.  Buds  normally  arise  in  the  Eud8- 

leaf  axils,  or  at  the  apex  of  the  stem  or  branches.  Those 
arising  elsewhere  are  adventitious.  Some  buds  contain  only 
rudimentary  foliage  leaves,  others  rudimentary  flowers  and 
some  both  rudimentary  leaves  and  flowers. 

If  we  examine  an  ordinary  foliage  bud,  especially  when 
it  is  bursting  into  vigorous  growth,  we  can  easily  satisfy  our- 
selves that  it  is  merely  a  young  stem,  or  branch,  bearing  the 
as  yet  imperfectly  developed  leaves  and  with  very  short  inter- 
nodes. 

Some  buds  are  covered  with  dry  scales  which  protect  the 
delicate  young  parts  within  them  from  being  dried  up  and 
killed;  these  are  well  seen  in  the  large  buds  of  the  Horse  Chestnut, 
Aesculus  indica.  The  bud  is  often  still  further  protected  by  a 
coating  of  wax,  gum,  or  resin,  while  the  young  parts  inside  the 
scales  are  also  often  protected  by  a  covering  of  wool,  or  hair. 
Other  buds  have  no  protecting  scales  and  are  said  to  be  naked, 
e.g.  those  of  Teak.  If  we  watch  the  development  of  a  scaly 
bud  we  find  that,  shortly  after  the  bud  opens  and  begins  to 
grow  vigorously,  the  scales  which  previously  covered  and  pro- 
tected it  fall  off,  leaving  behind  several  small  scars  close 
together  on  the  stem,  indicating  the  spots  where  they  were 
inserted.  These  can  be  well  seen  in  Fig.  1,  Plate  VII,  which 
shows  a  twig  of  Acer  caesium  from  the  terminal  bud  of  which 
4  leaves  have  developed,  the  bud-scales  having  just  been  shed. 
In  all  trees  and  shrubs  therefore  in  which  the  growth  of  stems 
and  twigs  is  stopped  once  annually,  scaly  buds  being  produced 
at  their  apices  during  the  period  of  rest,  it  is  clear  that  we  . 
can  calculate  their  age,  so  long  as  these  scars  of  the  bud-scales 
are  visible.  In  Fig.  1,  Plate  VII,  for  instance,  the  twig 
shows  the  completed  growth  of  3  years,  while  that  of  the  4th 
year  has  just  commenced.  In  the  spruce,  Picea  Morinda,  the 
bud  scales  do  not  fall  off  singly,  but  cohere,  ac.d  are  carried 
up  like  a  cap  on  the  tip  of  the  young  expanding  shoot. 

In  connection  with  this  question  of  the  age  of  shoots  should 
be  noted  also  the  fact  that  in  Pines  one  whorl  of  lateral 
branches  is  usually  developed  each  year,  so  that  the  age  of 


34 

branches  and  young  trees  can  be  calculated  by  counting  such 
whorls,  allowance  being  of  course  made  in  the  case  of  stems 
for  the  number  of  years  required  before  the  first  whorl  is  formed 
on  the  seedling.  There  are,  however,  exceptions  and  unusual 
conditions  may  cause  the  development  of  more  than  one  whorl 
in  the  year,  but  it  is  important  to  note  this  as  an  example 
of  what  may  be  done  by  carefully  watching  the  development  of 
different  species.  Bud  scales  are  sometimes  metamorphosed 
leaves.  If  we  examine  an  opening  bud  of  Aesculus  indica  we 
find  that,  while  the  outermost  scales  remain  dry  and  fall  off, 
some  of  the  inner  scales  begin  to  grow  and  those  which  grow 
the  most  very  closely  resemble  normal  leaves.  Figs.  1 — 8, 
Plate  VIII,  give  a  series  of  such  scales  removed  from  an  open- 
ing bud  which  clearly  shows  the  bud -scales  to  be  normal  leaves 
which  have  been  arrested  in  their  development  and  made  to 
function  as  a  protective  covering  to  the  other  young  leaves. 
Fig.  2,  Plate  VII,  shows  a  young  shoot  developing  from  the 
bud  of  Carpinus  viminea,  and  in  this  case  the  bud-scales  are 
seen  to  be  stipules,  the  inner  scales  being  situated  one  on 
each  side  of  a  leaf  petiole  just  as  are  normal  stipules.  The 
bud-scales  of  species  of  Ficus  are  also  found  to  be  stipules  and 
in  Stephegyne  parvifolia  the  large  interpetiolar  stipules  act  as 
bud-scales,  see  Fig.  9,  Plate  VIII.  All  the  buds  which  are 
lormed  on  a  plant  do  not  necessarily  develop  ;  in  many  cases  a 
farge  number  remain  dormant  for  long  periods  and  develop  only 
under  exceptional  circumstances,  such  as  when  neighbouring 
buds,  or  young  shoots,  are  destroyed,  such  resting  buds  are 
termed  dormant  buds. 

Usually  there  is  only  one  bud  in  each  leaf  axil,  but  not 
infrequently  there  are  additional  buds  situated  on  each  side  of, 
or  above,  the  true  axillary  bud  ;  these  are  termed  accessory 
buds.  In  some  cases  these  are  obviously  only  the  lowest 
buds  of  a  normal  axillary  shoot,  which,  at  first  sight,  appear 
to  spring  directly  from  the  parent  stem,  owing  to  the  lower 
nodes  of  the  axillary  shoot  remaining  very  short. 

In  Prinsepia  utilis  there  is  frequently  an  accessory  bud 
which  develops  into  a  spine  and  serves  as  a  protection  to  the 
axillary  bud  below  it,  see  Fig.  10,  Plate  VIII.  Buds  which  arise 
outside  of  the  leaf -axil  are  termed  extra-axillary. 

Buds  vary  greatly  in  size,  shape,  colour,  in  the  number  of 
their  external  scales  and  other  characters  which  are  useful  for 
identifying  deciduous  trees  and  shrubs  in  the  forest. 

In  the  Plane,  Platanus,  the  buds  are  found  to  be  complete- 
ly covered  by  the  hollow  base  of  the  petiole  and  only  become 


35 

visible  when  the  leaves  fall  off,  a  characteristic  circular  scar 
around  the  bud  then  showing  where  the  leaf  was  inserted. 

33.  The  way  in  which  the  young  Vernation 

leaves,  or  leaflets,  are  folded  in  the  bud,  which  is  called 
vernation,  varies  in  different  species.  In  some  cases  the  edges  of 
the  leaf  are  rolled  in  towards  the  midrib  on  the  upper  surface  of 
the  leaf  when  the  vernation  is  involute.  In  other  cases  the  edges 
are  similarly  rolled  in  towards  the  midrib  on  the  under  surface, 
the  vernation  being  revolute.  In  others  the  leaf  is  not  rolled 
and  the  longitudinal  halves  of  the  blade,  turning  on  the  midrib 
like  a  hinge,  are  placed  flat  face  to  face  with  their  upper  surfaces 
in  contact,  the  vernation  being  conduplicate.  In  others  the  leaf 
is  so  folded  on  its  longitudinal  axis  that  the  under-surface  is 
outside  and  one  edge  covers  the  other,  the  vernation  being 
convolute.  In  others  the  leaf  is  coiled  inward  from  the  apex, 
the  vernation  being  circinnate.  In  others  the  leaf-blades  are 
folded  between  the  nerves,  the  vernation  being  plicate. 

For  illustrations  see  Plate  IX. 

Finally,  in  some  cases,  there  is  no  obvious  folding,  or 
rolling,  and  the  young  leaves  are  placed  practically  flat,  or 
slightly  curved,  one  against  another. 

A  young  leaf  in  the  early  stages  of  its  development  has  its 
tissues  imperfectly  developed,  and  the  soft  green  tissue  lying 
between  the  nerves  is  particularly  susceptible  to  injury  and 
is  liable  to  be  dried  up  and  destroyed  if  fully  exposed  to  the, 
sun  and  air. 

Some  protection  is  therefore  commonly  provided  for  it, 
sometimes  by  the  folding  of  the  lamina,  sometimes  by  the 
position  assumed  by  the  young  leaves,  and  sometimes  by 
protective  coverings  of  hair,  wax,  gum,  or  resin. 

Thus  in  Viburnum  cotinifolium  we  find  the  young  leaves 
erect  on  the  stem,  each  pair  of  opposite  leaves  having  their 
upper  surfaces  placed  close  together,  while  only  the  under 
surfaces  are  exposed  to  the  light  and  air. 

The  upper  surface  of  each  leaf  is  concave,  the  under 
surface  convex,  and  the  lamina  is  plicately  folded,  the  delicate 
green  tissue  being  thrown  into  deep  folds  between  the  nerves, 
these  folds  projecting  into  the  concavity  of  the  upper  surface. 
Thus  the  green  tissue  is  effectively  protected  during  its  early 
development  by  the  close-set  framework  of  nerves  which  alone 
are  exposed  on  the  exterior  of  the  young  leaves.  As  the  young 
leaves  are  thus  erect  and  closely  adpressed,  we  find  that  tfce 
upper  surface  of  the  petioles  is  grooved,  room  biding  thus 
provided  for  the  terminal  bud  between  the  opposite  leaves. 

2D 


36 

As  eacli  leaf  gets  older  the  folds  of  green  tissue  are  flattened 
out.  the  leaf  is  thrown  outward  and  downward  and  the  lamina 
is  expanded  more  or  less  horizontally,  with  the  upper  surface 
directly  exposed  to  the  sun's  rays.  See  Figs.  7 — 9,  Plate  IX. 
Homologous  34.  Several  cases  of  the  so- 

and  Analo-  ca}iefi  metamorphosis  of  plant  members  have  been  incidentally 
Members.  mentioned  in  this  and  the  previous  chapters  from  which  it  will 
be  seen  that  members  which  are  alike  in  their  mode  and  place 
of  origin  and  general  plan  of  structure  are  regarded  as 
morphologically  the  same,  however  different  they  may  super- 
ficially appear  to  be  in  their  form,  or  function.  Members 
which  are  thus  of  the  same  morphological  value  are  said  to 
be  homologous,  whereas  those  members  which,  although  alike 
in  some  respects,  are  not  morphologically  the  same,  such 
as  the  spines  of  Berberis  and  Bel,  are  called  analogous. 


37 


CHAPTER  IV.-THE  INFLORESCENCE  AND  FLOWER. 

35.  A  shoot  which  bears  a  flower  Flowering- 

is  called  a  flowering-shoot,  and  like  an  ordinary  leafy  shoot  it  shoot> 
may  be  axillary,  or  terminal.    In  some   species   the   flowering 
shoots  spring  from  the  older  branches,  or  stems,  from  which 

the  leaves   have   disappeared,  as  in  Ougeinia  dalbergioides  and 
Ficus  Cunia. 

The  flowering  shoot  frequently  branches  and  produces,  not  inflorescence. 
one,  but  several  flowers.  The  collection  of  flowering  shoots  thus 
formed  is  the  inflorescence.  According  as  the  flowering  shoots 
are  stiff  and  erect,  or  drooping  and  pendulous,  so  is  the 
inflorescence  described  as  erect,  or  pendulous.  The  inflorescence 
varies  greatly  in  size  :  in  the  Teak  it  is  very  extensive  and  is 
then  described  as  ample,  or  merely  large,  whereas  in  other 
cases  it  is  reduced  to  a  few  small  flowers,  or  even  to  a  single 
flower,  in  which  case  the  flowers  are  solitary,  and  the  inflores- 
cence may  be  further  said  to  be  many-  or  few-flowered,  and  much- 
branched  or  little-branched.  Again  if  its  branches  are  close 
together  the  inflorescence  is  compact,  or  dense,  otherwise  it  is 
lax,  or  loose. 

The  first  important  point  to  notice  regarding  the  flowering  Bracts, 
shoots  is  that  the  leaves  which  they  bear,  and  from  the  axils 
of  which  they  spring,  are  usually  smaller  than  the  ordinary 
foliage  leaves,  of*  a  different  shape  and  sometimes  also  of  a 
different  colour.  Such  altered  leaves  are  called  bracts.  Those 
which  are  borne  on  the  ultimate  branches  of  the  flowering 
shoots,  i.e.  on  the  stalks  of  the  individual  flowers,  are  distin- 
guished by  the  name  of  bracteoles.  A  flowering  shoot  on  which 
bracts  are  produced  is  said  to  be  bracteate,  if  no  bracts  are 
produced  it  is  ebracteate. 

The  stalk  of  a  flower  is  the  peduncle  ;  when  the  peduncle 
branches,  that  portion  of  it  from  which  the  lateral  flowering 
branches  arise  is  the  rhachis  and  the  stalks  of  the  single 
flowers  are  then  called  pedicels,  terms  which  will  recall  the 
analogous  ones  of  petiole,  rhachis  and  petiolule  employed  for 
the  leaf.  Stalked  flowers  are  said  to  be  pedunculate,  or 
pedicellate,  as  the  case  may  be,  while  flowers  without  a  stalk 
are  sessile. 

36.  We  have  seen  above  that 

there  are  two  principal    kinds  of  branching,    viz.,  monopodial 
and  cymose,  and  according  to  the  way  in  which   the  flowering 


ceuce 


38 

shoot  branches  two  main  types  of  inflorescence  are  distin- 
guished as  under,  although  here,  as  in  other  cases,  intermedi- 
ate forms  are  found  : 

(1)  Monopodial  or  racemose. — Here  there  is  a  distinct  main 

axis  formed  by   the    continued   elongation   of   the 
parent  flowering  shoot. 

(2)  Cymose. — Here  there  is  no  main  axis  formed   by   the 

elongation  of  the  parent  flowering  shoot  and  lateral 
flowering  shoots    grow  more    vigorously  than  the 

o  <->  c?  */ 

parent  axis. 
The  principal  kinds  of  racemose  inflorescences  are  :— 

(a)  Raceme. — The  main  axis  is  elongated  and  bears 
pedicellate  flowers. 

(6)  Corymb. — This  ia  a  raceme  in  which  the  main  axis 
is  short  and  the  lower  pedicels  are  longer  than 
the  upper  ones,  the  inflorescence  being  more  or 
less  flattened  and  with  a  convex  outline. 

If  in  a  raceme,  or  corymb,  the  main  axis,  instead  of  directly 
producing  pedicels,  develops  lateral  branches  which,  directly 
or  indirectly,  bear  the  pedicels,  we  get  a  compound-raceme  or 
-corymb,  respectively. 

(c)  Spike. — The  main  axis  is  elongated  and  bears  sessile 

flowers.  A  small  spike  is  called  a  spikelet.  A 
spike  having  inconspicuous  unisexual  flowers 
and  which  falls  off  entire  from  the  plant  after 
flowering,  or  fruiting,  is  a  catkin.  It  is  usually 
pendulous. 

(d)  Spadix  is  a  spike  with  a  thick   or    fleshy    axis.     A 

spadix  is  usually  more  or  less  enclosed  in  a  large 
bract  which  is  called  the  spathe.  In  palms  the 
spadix  is  branched  and  has  several  spathes. 

(e)  Umbel. — The  main  axis  bears  a  number  of   pedicel- 

late flowers  at  its  apex.  The  pedicels  which  all 
radiate  from  the  same  point  are  called  the  rays 
of  the  umbel.  At  the  apex  of  the  peduncle 
bracts  are  often  found  forming  a  whorl  below  the 
pedicels,  or  rays,  which  is  called  the  involucre. 
When  the  rays  instead  of  bearing  flowers  bear 
secondary  umbels,  the  latter  are  termed  umbellules 
and  their  involucres  are  involucels,  while  the 
inflorescence  becomes  a  compound  umbel. 
(/)  Head  or  Capitulum. — The  main  axis  bears  a  number 
of  sessile  flowers  at  its  apex.  Here  also  there 


39 

is  usually  an  involucre  of  bracts  below  the  flowers. 
The  apex  of  the  peduncle  on  which  the  flowers  are 
borne  and  which  is  really  the  shortened  rhachis 
is  here  called  the  receptacle  of  the  inflorescence. 
Small  scale -like  bracts  are  often  found  on  the 
receptacle  subtending  the  individual  flowers. 
In  a  typical  raceme  there  is  no  obvious  reason  why  the 
main  axis  should  not  continue  to  grow  and  produce  lateral 
branches  in  acropetal  succession  indefinitely,  whereas  if  it  soon 
produced  a  terminal  flower,  as  is  the  case  in  a  cyme,  its  growth 
is  at  once  checked  and  the  further  development  of  the 
inflorescence  then  devolves  on  the  lateral  branches.  Racemose 
inflorescences  are  therefore  often  called  indeterminate,  or 
indefinite,  and  cymose  inflorescences  determinate,  or  definite. 
Again  if  a  plan  is  drawn  of  a  raceme  the  youngest  flowers  are 
in  the  centre  and  the  oldest  are  outside,  while  a  line  following 
the  course  of  development  from  the  oldest  to  the  youngest 
flowers  of  such  an  inflorescence  passes  from  the  outside  to  the 
centre,  whereas  in  a  cyme  the  reverse  is  the  case,  the  oldest 
flower  being  in  the  centre.  Hence  racemose  inflorescences  are 
called  centripetal  and  cymose  inflorescences  centrifugal.  In  the 
case  of  inflorescences  in  which  the  shoots  are  much  reduced  in 
length  and  the  flowers  are  therefore  brought  together  more  or 
less  at  the  same  level,  it  is  often  not  easy  to  decide  to  which 
type  they  must  be  referred  and  the  relative  position  of  the 
flowers  then  often  helps  us  to  decide  the  question.  In  a  head, 
for  instance,  the  facts  that  the  oldest  flowers  which  open  first 
are  on  the  outside  and  that  the  youngest  which  open  last  are  in 
the  centre,  indicate  that  the  inflorescence  is  racemose.  A 
compound  racemose  inflorescence  also  is  usually  easily 
recognised  by  the  fact  that  there  is  a  distinct  main  axis  with 
the  oldest  and  longest  branches  at  its  base  and  the  youngest 
and  shortest  at  the  apex. 

The  principal  kinds  of  cymose  inflorescences  are  : — 

(a)  Dichotomous  cyme.. — The  main  axis  terminates  in 
a  flower  and  two  vigorous  and  equal  lateral 
branches  develop.  If  there  are  three  equally 
vigorous  lateral  branches  instead  of  two  the 
cyme  becomes  tricJiotomous.  If  this  kind  of 
branching  is  continued  and  each  lateral  branch 
in  its  turn  terminates  in  a  flower  and  in  its  turn 
develops  2  or  3  vigorous  lateral  branches  and 
so  on,the  inflorescence  becomes  a  compound  di- 
or  tri-chotomous  cyme  respectively. 


40 

(6)  Helicoid  cyme. — The  main  axis  here  terminates  in 
a  flower  as  before,  but  the  further  growth  of  the 
inflorescence  is  carried  on,  not  by  2  or  3  equally 
vigorous  lateral  branches,  but  by  one  only,  the 
latter  also  soon  terminates  in  a  flower  and 
produces  another  lateral  branch  and  so  on,  the 
lateral  branches  always  developing  on  the  same 
side  of  the  parent  axis. 

(c)  Scorpioid  cyme. — This  is  similar  to  (b)  but  the  lateral 
branch,  instead  of  always  developing  on  the  same 
side  of  the  parent  axis,  develops  alternately  on 
opposite  sides. 

In  both  (b)  and  (c)  there  is  thus  a  sympodium,  or  false  axis. 
When  this  false  axis  straightens,  as  it  usually  does,  the  inflores- 
cence has  a  strong  superficial  resemblance  to  one  of  the  racemose 
type.  In  such  cases  the  position  of  the  bracts  often  indicates 
at  once  the  true  character  of  the  inflorescence,  for  the  flower 
stalks,  instead  of  springing  from  the  axils  of  the  bracts,  as  they 
would  do  in  a  raceme,  are  opposite  to  the  bracts. 

A  Fascicle  is  a  general  term  for  a  tuft  or  cluster  of 
flowers,  or  of  flowering  shoots,  without  reference  to 
its  being  of  the  racemose  or  cymose  type,  and  is  used 
when  the  true  character  of  the  inflorescence  is  not 
easily  made  out. 

Panicle  is  a  much  used  term  which  is  applied  to  all 
compound  and  much  branched  inflorescences  of  which 
the  first  ramifications  are  racemose.  A  panicle 
which  is  ovate  or  lanceolate  in  outline  is  called  a 
thyrsus. 

A  cyme  which,  in  outline  and  general  appearance,  resem- 
bles a  raceme,  corymb,  or  umbel,  is  called  a  racemiform,  co- 
rymbiform,  or  umbelliform  cyme,  and  similarly  a  panicle  may 
be  racemiform,  corymbiform,  or  umbelliform. 

For  illustrations  of  some  of  the  above  types  of  inflorescen- 
ces see  Figs.  1  tu  10,  Plate  X. 

Finally  there  are  the  more  complicated  so-called  mixed 
inflorescences  in  which  more  than  one  type  is  combined. 
Thus  the  ultimate  ramifications  of  a  panicle,  the  first  branches 
of  which  are  developed  according  to  the  racemose  type,  may 
be  cymes,  and  such  a  panicle  may  be  described  as  a  cymose 
panicle,  or,  perhaps  better,  we  may  describe  the  inflorescence 
as  one  consisting  of  cymes  arranged  in  a  panicle. 

If  the  student  will  examine  for  himself  the  following 
examples  of  typical  inflorescences,  they  will  help  him  to 


41 

realise   how  the   above  terms   may  be  employed  in    practical 
descriptive  botany : — 

(1)  Lax,  long,  pendulous  racemes  of  Cassia  Fistula. 

(2)  Dense,  pedunculate  racemes  of  Abrus  precatorius. 

(3)  Flowers    of  Desmodium    pulchellum  in  small  fascicles 

in  the  axils  of  2-foliolate  bracts  arranged    in  ter- 
minal and  axillary  racemes. 

(4)  Few --flowered  corymb  of  Dolichandrone,  falcata. 

(5)  Spike  of  Terminalia  belerica. 

(6)  Spadix  of  Arisaema  Wallichianum,  the  Cobra  Plant. 

(7)  Fascicled  heads  of  Acacia  Farnesiana. 

(8)  Compound  trichotomous  cymes  of  Eugenia  operculata. 

(9)  Flowers    of    Trewia  nudifiora    fascicled  in    pendulous 

racemes. 

(10)  Many -flowered     umbelliform      cymes     of    Leptadenia 

reticulata. 

(11)  Sessile      flowers    of    Wendlandia    exserla,     in    large 

conical,  or  pyramidal,  panicles. 

(12)  Panicle  of  heads  of  Acacia  caesia. 

(13)  Panicle  of  umbels  of  Heptapleurum  venulosum. 

(14)  Dichotomous    cymes    of    Cornus  macropliylla  in  large 

terminal  panicles. 

(15)  Heads  of  Albizzia    odoratissima    in  compact  corymbs, 

arranged  in  large  panicles. 

(16)  Compact  thyrsus  of  Phlogacanthus  thyrsiflorus. 

37.  In  a  typical   flower  we  can  Parts  of  the 

at  first  sight  distinguish  3  obviously  distinct  kinds  of  organs ;  p^J^ 
the  outer  ones  are  more  or  less  clearly  leaf -like  in  shape,   often  stamens' 
with  a  distinct  venation  and  sometimes  green  in  colour.     These  Pistils, 
may  be  called  the  floral  envelopes  and  together  constitute  the 
perianth.     One  of  the  important  duties  which  the  perianth  has 
to  perform    is  to    envelop  and  protect,    especially    when  they 
are  young,  the  essential  floral  organs  situated  within  them  and 
which  are  the  stamens  and  pistils.     The  former  together  con- 
stitute the  male   portion  of  the  flower,  called  the  androzoium, 
and  the  latter  the  female  part,  called  the  gyncecium.     A  typica 
stamen  consists  of  a  swollen  head  called  the  anther,    situated 
on  a  slender  stalk  called  the  filament.     In  the  anther  is  stored 
a  yellow  powdery  substance  called  pollen,   contained  in  two 
little  bags  which  are  placed  side  by  side  in  the  anther  and  which, 
when  the   anther  is  mature,    open    and    scatter   the    pollen. 
This  opening  is  termed  the  dehiscence  of  the  anther.     A  typical 
single  pistil  consists  of  a  swollen  hollow,  base,  called  the  ovary, 
which   contains  the   small  rounded  bodies  called  ovules  (these 


42 


Torus  or 
sceptacle. 


Pollination 
Fertilisation. 


eventually  developing  into  what  are  known  as  seeds),  an 
elongated  neck  called  the  style,  which  is  frequently  more  or 
less  expanded  at  its  apex  and  there  forms  the  so-called 
stigma.  The  stigma  is  a  specially  differentiated  part  of  the 
style,  usually  with  a  moist  and  sticky  surface,  which  varies 
greatly  in  shape  and  size  ;  sometimes  it  is  a  rounded  knob  ; 
sometimes  it  is  merely  a  single,  or  double,  line  running  along 
the  side  of  the  style,  while  sometimes  the  apex  of  the  style  is 
forked  and  the  sticky  surface  of  the  stigma  (called  the  stig- 
matic  surface)  is  found  on  the  branched  tip  of  the  style.  The 
essential  parts  of  the  pistil  are  the  ovary  and  the  stigma. 
The  style  may  be,  and  often  is,  absent,  the  stigma  then 
being  sessile.  The  ovary  being  the  principal  part  of  a  pistil,  the 
word  ovary  is  often  used  to  signify  the  whole  pistil.  A  flower 
may  contain  one  or  several  pistils. 

The  perianth  leaves,  stamens  and  pistils  are  all  inserted 
cjose  together  on  the  more  or  less  expanded  apex  of  the 
flower  -stalk  and  this  portion  of  the  stalk  from  which  the 
floral  organs  spring  is  called  the  torus,  or  floral  receptacle. 

38>  The  object  of  the  flower  is 

^°  Pr°duce  seeds  which  are  capable  of  developing  new  and 
independent  plants.  No  seeds,  however,  can  arise  in  a  pistil 
unless  some  of  the  powdery  pollen  which  is  contained  in  the 
anthers  is  able  to  reach  the  stigma.  The  small  grains  of  pollen 
which  happen  to  be  caught  by,  and  to  adhere  to,  the  sticky 
stigma  develop  minute  tubes  which  penetrate  the  style  and 
grow  down  into  the  ovary  to  the  ovules,  the  contents  of  each 
pollen  grain  passing  into  its  tube.  When  one  of  these  tubes 
reaches  an  ovule,  a  part  of  its  contents  pass  into  and  fuse 
with  a  part  of  the  contents  of  the  ovule,  which  is  then  said 
to  have  been  fertilised  and  which  thereafter  grows  and 
develops  into  a  seed,  containing  the  embryo  of  a  new  plant. 
The  transference  of  pollen  to  the  stigmas  (pollination)  is 
thus  of  vital  importance,  it  being  essential  for  fertilisation 
and  the  production  of  seeds.  In  some  plants  such  as 
Grasses,  this  is  done  by  the  wind  and  these  plants  have  no 
conspicuously-coloured  flowers  ;  in  other  plants  this  work  is 
done  by  insects  and  birds  who  visit  the  flowers  for  the  sweet 
juice,  or  nectar,  which  they  contain,  get  dusted  by  pollen  in  the 
process,  and  carry  this  away  to  the  stigmas  of  other  flowers. 
Those  flowers  which  can  be  most  easily  seen  and  found  by  birds 
and  insects  obviously  therefore  have  the  best  chance  of  being 
visited  and  of  having  their  ovules  fertilised,  and  this  is  the 
reason  why  so  many  flowers  possess  conspicuous  and  brightly 


43 

•coloured  perianth  leaves  and  attractive  scents.  From  this  we 
see  that,  however  conspicuous  the  perianth  leaves  of  a  flower 
may  be,  these  are  after  all  of  only  subsidiary  importance  and 
that  the  really  essential  organs  of  a  flower  are  the  stamens  and 
pistils,  without  which  no  seed  can  be  formed.  There  are 
indeed  some  flowers  which  only  contain  each  a  single  pistil, 
•or  a  single  stamen,  but  which  nevertheless  must  be  regarded 
as  true  flowers. 

39.  The    perianth    of  a   flower  Perianth' 

may  consist  of  leaves  which  are  all  alike,  or  it  may  consist  of 
two  distinct  sets  of  leaves,  the  outer  of  which  are  usually 
green  and  the  inner  of  more  delicate  texture,  and,  being  white, 
or  of  some  colour  other  than  green,  the  latter  serve  to  make 
the  flower  conspicuous.  The  former  are  called  sepals  and 
together  constitute  the  calyx,  the  latter  are  the  petals  and 
together  constitute  the  corolla.  In  Rosa,  moschata  for  instance, 
we  find  no  difficulty  in  distinguishing  the  inner  5  large  white 
petals  from  the  outer  green  sepals. 

A  flower  which  has  both  sepals  and  petals  is  said  to  be 
dichlamydeous,  one  which  is  without  both  sepals  and  petals, 
i.e.  with  no  perianth,  is  achlamydeous,  while  one  which  has 
only  sepals,  or  only  petals,  is  monochlamydeous.  A  flower 
which  has  no  petals  is  also  said  to  be  apetalous.  When  the 
perianth  leaves  are  all  similar  and  there  is  therefore  no 
obvious  calyx  and  corolla,  the  leaves  are  merely  called  perianth 
leaves.  If  they  are  all  coloured  like  petals  the  perianth  is 
said  to  be  petaloid,  whereas  if  they  all  resemble  sepals  the 
perianth  is  sepaloid.  If  the  perianth,  calyx,  or  corolla,  of  a 
flower  consists  of  distinct  leaves,  which  are  separate  from  one 
another  and  each  of  which  has  a  distinct  insertion  of  its  own 
upon  the  receptacle,  the  perianth,  calyx,  or  corolla,  respectively, 
is  said  to  be  polyphyllous,  polysepalous,  or  polypetalous.  On 
the  other  hand  a  'perianth,  calyx,  or  corolla,  of  united,  or 
connate,  leaves,  is  said  to  be  gamopliyllous,  gamosepalous,  or 
gamopetalous.  According  to  the  degree  of  cohesion  between 
the  respective  perianth  leaves,  sepals,  or  petals,  they  are 
said  to  be  connate  at  the  base,  to  or  beyond  the  middle,  nearly 
to  the  apex,  and  so  on. 

In  cases  of  cohesion,  where  there  is  a  distinct  narrow  base 
formed  by  the  lower  connate  portions  of  the  leaves,  this  is 
called  the  perianth-,  calyx-,  or  coiolla-tube  and  the  upper 
expanded  portion  is  called  the  limb.  The  intermediate  portion, 
usually  slightly  more  expanded  than  the  tube  and  not  so  much 
expanded  as  the  limb,  is  called  the  throat.  The  calyx  or 


44 


Position  and 
Number  of 
Parts  of  the 
Flower. 


corolla  is  also  said  to  be  toothed  when  the  sepals  or  petals 
are  united  nearly  to  the  apex,  deft,  or  lobed,  if  divided  to 
about  the  middle,  and  parted  if  divided  nearly  to  the  base. 
The  same  terms  may  also  be  used  for  the  perianth. 

40.  When     both     calyx     and 

corolla  are  present  the  number  of  sepals  is  very  frequently 
equal  to  the  number  of  petals.  The  organs  of  the  flower  are 
usually  inserted  in  whorls  on  the  receptacle ;  the  internodes 
between  the  whorls  being  as  a  rule  very  short,  the  whorls  are 
all  brought  close  together  and  the  members  of  successive  whorls 
are  seen  to  alternate  with  each  other.  Thus  we  find  an  outer 
whorl  of  say  5  sepals  and  then,  inside  and  above  them  on  the 
receptacle,  a  whorl  of  5  petals  so  arranged  that  each  petal 
stands  in  the  gap  between  two  sepals,  then  inside  the  petals 
a  whorl  of  5  stamens  alternating  with  the  petals  and  each 
stamen  therefore  opposite  a  sepal,  then  another  and  inner 
whorl  of  stamens  alternating  with  the  outer  and  each  of  its- 
stamens  therefore  opposite  a  petal,  and  finally  the  pistil,  or 
pistils.  Some  flowers  have  a  whorl  of  bracteoles  outside  the 
calyx,  which  at  first  sight  may  be  mistaken  for  an  extra  whorl 
of  sepals.  By  carefully  counting  the  obvious  petals  and  sepals, 
however,  their  true  nature  may  usually  be  detected.  Thus  in  a 
flower  of  Hibiscus  Rosa-sinensis  we  find,  outside  the  stamens, 
5  large  red  obvious  petals  and  then  a  gamosepalous  calyx  of  5 
connate  sepals  alternating  with  them.  In  addition  to  these, 
however,  we  find  6  or  7  linear  bracteoles.  If  the  latter  formed 
a  normal  whorl  of  sepals  they  would  be  5  in  number  alternating 
with  the  whorl  above,  which  they  do  not,  but  even  if  they 
were  5  in  number  and  did  correctly  alternate  with  the  sepals 
above  them,  by  including  them  among  the  sepals  the  latter 
would  become  twice  as  numerous  as  the  petals,  which  is  unusual. 
Again  in  the  cotton  plant  (Gossypium)  we  find  an  obvious, 
corolla  outside  the  stamens  composed  of  5  large  yellow  petals 
with  a  purple  centre  and  a  gamosepalous  calyx  of  5  connate 
sepals  and  then  3  large  leafy  bracteoles  j  ust  below  the  calyx. 
One  or  more  whorls  of  bracteoles  which  in  this  way  resemble 
an  outer  calyx  of  a  single  flower  form  what  is  called  an 
epicalyx  and  remind  us  of  the  so-called  involucre  of  bracts  at 
the  base  of  a  head  or  umbel  of  several  flowers. 

What  has  been  just  said  must  of  course  not  be  taken  to 
mean  that  in  a  flower,  in  which  both  calyx  and  corolla  are 
present,  the  number  of  sepals  must  equal  the  number  of  petals 
and  the  members  of  each  whorl  must  alternate  with  those  of  the 
next,  as  there  are  obviously  several  flowers  in  which  this  is  not 


45 

so,  such  as  those  of  the  Poppy,  in  which  there  are  4  petals  and 
2  sepals.  The  fact  is  only  here  emphasized  that  this  is  the  case 
in  a  very  large  number  of  plants  and  the  fact  will  often  be 
found  useful  in  helping  us  to  readily  diagnose  flowers  in  the  field. 

If  now  we  look  at  a  flower  of  Berberis  Lycium  we  find, 
outside  the  6  stamens,  12  perianth  leaves,  the  inner  6  of  which 
are  all  the  same  size  and  colour  and  at  first  sight  appear  to  be  in 
one  whorl  with  a  stamen  in  front  of  each  of  them.  The  outer  6 
perianth  leaves,  however,  are  distinctly  arranged  in  two  whorls, 
each  of  3  leaves,  the  outer  3  being  greenish  and  sepal-like 
alternating  with  the  3  inner  leaves  which  are  larger,  brighter 
yellow,  and  obviously  more  like  petals.  If  we  examine  the 
flower  more  carefully  and  remember  that  as  a  rule  the  number 
of  sepals  and  petals  is  equal,  we  find  that  it  is  really  quite 
normal  with  2  whorls  each  of  3  sepals,  2  whorls  each  of  3 
petals  and  2  whorls  each  of  3  stamens,  the  members  of  each 
whorl  alternating  with  those  of  the  next.  In  this  case  the 
outermost  3  leaves  are  more  or  less  obviously  sepal-like, 
but  the  3  leaves  in  the  next  whorl  are  undoubtedly  more  like 
typical  petals,  and  this  indicates  that  the  only  real  difference 
between  sepals  and  petals  is  one  of  position,  the  former  being 
the  outer  and  the  latter  the  inner  set.  Outside  the  outer  whorl 
of  sepals  there  are  usually  also  2  or  3  small  bracteoles  which 
closely  resemble  the  outer  sepals,  and  we  see  that,  so  far  as  mere 
appearance  goes,  it  is  not  always  possible  to  readily  distinguish 
bracteoles  from  sepals.  A  flower  which  has  its  parts  in  whorls 
of  2,  3,  4,  5,  or  6  respectively  is  said  to  be  di-,  tri-,  tetra-, 
penta-  or  hexa-  merous. 

According  as  the  number  of  stamens  in  a  flower  is  one,  two, 
three,  or  many,  it  is  said  to  be  mon-,  di-,  tri-,  or  poly- 
androus. 

41.  A    flower  which     contains  Distribution 

both  stamens  and  pistil,  or  pistils,  is  said  to  be  hermaphrodite, 
or  bisexual  ;  a  flower  which  either  has  no  stamens,  or  no  pistil, 
is  unisexual ;  a  unisexual  flower  with  stamens  is  a  male,  or 
staminate,  flower,  and  is  usually  shortly  written  thus —  $  flower, 
while  one  with  pistils  only  is  a  female,  or  pistillate,  flower  and 
is  written  ?  flower.  When  the  flowers  of  a  plant  are  unisexual 
but  both  $  and  s  flowers  occur  on  the  same  individual  plant,  the 
flowers  are  monoecious  and  the  plant  itself  may  be  described  as 
moncecious.  When  the  flowers  of  a  plant  are  unisexual  and  the  <? 
and  ?  are  found  on  different  individuals  the  flowers  and  plant 
are  said  to  be  dioecious.  When  the  flowers  of  a  plant  are  both 
unisexual  and  hermaphrodite,  which  may,  or  may  not,  occur 


46 

on  the  same  individual,  the  plant  and  flowers  are  said  to  be 
polygamous.  A  flower  which  produces  no  seed  is  said  to  be 
sterile  ;  this  may  be  because  it  possesses  no  pistil,  or  because  its 
pistils  fail  to  produce  seed. 

Symmetry  of  42.  A  flower  is  normally  borne 

in  the  axil  of  a  bract  which  is  said  to  subtend  the  flower, 
just  as  an  ordinary  foliage  leaf  subtends  its  axillary  branch. 
That  part  of  the  flower  which  faces  the  flowering  shoot  from 
which  it  springs  laterally  is  called  the  upper,  superior,  or 
posterior,  while  the  opposite  part  facing  the  subtending  bract  is 
the  lower,  inferior,  or  anterior.  The  plane  which  passes  through 
the  posterior  and  anterior  part  of  the  flower,  and  therefore 
through  the  parent  flowering  axis  and  the  middle  of  the 
subtending  bract,  is  the  median  plane,  or  section,  of  the  flower,, 
while  a  vertical  plane  passing  through  the  flower  at  right 
angles  to  the  median  plane  is  called  the  lateral  plane  or  section, 
and  any  other  plane  which  passes  through  the  intersection  of 
these  may  be  called  the  oblique  plane  or  section. 

A  radial  longitudinal  section  of  a  flower  is  one  which  is 
parallel  to  its  longitudinal  axis  and  passes  through  the  centre 
of  the  flower,  while  a  transverse  section  is  one  at  right  angles  to 
its  longitudinal  axis. 

If  in  a  flower  the  members  of  each  whorl  of  floral  organs  are 
all  alike  in  size  and  shape,  the  flower  is  regular.  If  not  it  is 
irregular.  The  regularity  or  otherwise  of  a  flower  is  naturally 
most  noticeable  in  the  perianth  leaves  and  especially  in  the 
corolla,  these  being  as  a  rule  the  most  conspicuous  part  of  the 
flower,  and  hence  when  in  descriptive  botany  a  flower  is 
spoken  of  as  regular,  or  irregular,  these  terms  may  as  a  rule  be 
taken  to  refer  to  the  corolla,  or  perianth  leaves. 

A  flower  which  is  divisible  in  two  or  more  radial  longitudinal 
planes  into  similar  halves,  so  that  in  each  section  one  half  is  to 
the  other  as  an  object  is  to  its  image  seen  in  a  mirror  and  the 
halves  in  one  section  are  similar  to  those  in  the  other  sections, 
the  flower  is  actinomorphic.  When  there  is  only  one  radial  longi- 
tudinal section  in  which  a  flower  can  be  divided  into  similar 
halves  it  is  said  to  be  zygomorphic,  and  if  there  is  no  plane  in 
which  it  can  be  divided  in  this  way  it  is  asymmetric. 

The  p»rts  of  43.  The  flower  is  really  a  short- 

ate  LeavJ  enec^  shoot  and  its  parts,  which  compose  the  calyx, 
corolla,  androecium  and  gynoecium  are  really  nothing 
but  leaves.  The  flower  bud  like  the  bud  of  an  ordinary  leafy 
shoot  is  either  terminal,  or  it  arises  in  the  axil  of  a  leaf,  the 
latter  being  a  normal  green  leaf,  or  one  which  has  only  been 


47 

slightly  altered  in  character  in  which  case  it  is  called  a  bract. 
The  green  sepals  are  often  large  and  obviously  leaf-like ;  the 
petals  also  in  shape  more  or  less  resemble  leaves  and  they  often 
have  a  distinct  venation.  Colour  alone  is  a  characteristic  of 
very  little  importance  in  enabling  us  to  decide  on  the  true 
nature  of  a  member,  as  is  shown  by  the  fact  that  the  bracts  of 
several  plants,  although  they  are  obviously  leaves,  are 
brilliantly  coloured  like  petals,  e.g.  the  bracts  of  Bougainvillea. 
The  remarkably  conspicuous  pure  white,  or  coloured,  calyx 
lobe  of  Mussaenda  should  also  be  noted.  Stamens  it  is  true  do 
not  at  first  sight  at  all  remind  us  of  leaves,  but  in  some 
flowers,  especially  in  those  which  have  numerous  petals,  as  in 
the  Water  Lily  (Nymphaea),  we  find  that  the  petals  gradually 
pass  into  the  stamens  through  a  series  of  intermediate  forms, 
which  clearly  indicate  how  one  and  the  same  organ  may,  by 
a  number  of  slight  modifications,  become  a  normal,  more  or 
less  leaf-like,  petal,  or  a  perfect  stamen. 

The  pistil  also  does  not  at  first  sight  resemble  a  leaf,  but  if 
we  take  a  simple  pistil  such  as  a  young  pea-pod,  and  compare 
it  with  an  involutely-folded  leaf,  the  margins  of  which  are 
brought  together  and  made  to  cohere,  we  see  that  very  slight 
modifications  would,  after  all,  suffice  to  convert  what  would 
ordinarily  develop  into  a  normal  leaf  into  a  pistil. 

Occasionally  also,  e.g.  in  Roses,  examples  may  be  found  of 
so-called  "green-flowers,"  in  which  the  different  members, 
instead  of  assuming  their  ordinary  shape  and  colour,  appear  as 
more  or  less  perfectly  developed  green  leaves.  The  facts  noted 
above  which  indicate  the  true  nature  of  the  floral  members  are 
found  to  be  confirmed  by  a  microscopic  examination  of  their 
mode  of  origin,  minute  structure  and  development  during  the 
early  stages  of  their  growth,  when  they  are  found  to  be 
indistinguishable  from  true  leaves.  The  floral  parts  arise  on 
the  flower  stalk  just  as  leaves  do,  on  an  ordinary  shoot.  Occa- 
sionally they  are  in  spirals,  but  as  a  rule  they  are  arranged  in 
whorls.  The  internodes  in  the  flower  usually  remain  very  short, 
and  all  the  floral  parts  are  therefore  crowded  close  together  on 
the  portion  of  the  shoot  from  which  they  spring,  i.e.  on  the 
so-called  receptacle,  or  torus.  This,  however,  is  not  always  the 
case,  and  in  Capparis  there  is  a  distinct  internode  between  the 
stamens  and  the  pistil,  the  latter  appearing  to  have  a  stalk 
which  is  called  the  gynophore ;  internodes  are  also  occasionally 
developed  between  other  floral  parts,  e.g.  between  the  petals  and 
stamens  in  species  of  Grewia,  and  a  stalk  which  thus  appears 
to  bear  both  the  stamens  and  the  pistil  is  called  a  gonophore. 


48 


Cohesion. 


Flowers. 


Although  the  apex  of  the  floral  axis  as  a  rule  ceases  to  grow 
at  a  very  early  stage  in  the  development  of  the  flower,  excep- 
tional cases  are  known  in  which  this  axis  has  continued  to 
grow,  and  we  find  a  green  leafy  shoot  growing  apparently  out 
of  the  centre  of  the  flower.  Very  rare  cases  are  also  known 
in  which  buds  have  been  developed  in  the  axils  of  the  floral 
members,  just  as  they  are  usually  developed  in  the  axils  of 
normal  leaves. 

44.  Owing  to  the  very  limited 

growth  of  the  floral  axis  and  to  the  non-development  of  its 
internodes  the  floral  parts  are,  as  has  already  been  noted, 
crowded  close  together  and  in  many  cases  this  crowding 
results  in  the  young  members,  which  originate  separately  on 
the  receptacle,  coalescing  at  their  bases,  and  they  then  grow 
up  together  instead  of  separately.  This  is  how  a  gamosepalous 
calyx,  or  a  gamopetalous  corolla,  for  instance,  arises.  When 
members  belonging  to  the  same  whorl  become  thus  fused 
together  the  phenomenon  is  termed  cohesion  and  the  members 
are  said  to  be  connate. 

Members  belonging  to  adjacent  whorls  also  often  become 
fused  together  and  we  find,  for  example,  stamens  fused  with 
petals,  the  stamens  looking  as  if  they  had  grown  out  of  the 
corolla.  In  such  a  case  we  say  there  is  adhesion.  Members 
which  are  thus  adnate  appear  to  originate  not  directly  from 
the  receptacle  in  the  usual  way  but  at  some  distance  from  it, 
their  apparent  point  of  insertion  having  been  carried,  as  it 
were,  away  from  the  receptacle.  Cohesion  then  is  the  union 
of  members  belonging  to  the  same  whorl,  adhesion  the  union 
of  those  developed  at  different  levels,  while  members  which 
are  neither  connate  nor  adnate  are  said  to  be  free. 
,  45.  When  there  is  no  adhesion 

an(l  the  sePals>  petals  and  stamens  all  arise  directly  from  the 
receptacle,  these  members  obviously  originate  directly  below 
the  gyno3cium.  The  sepals,  petals  and  stamens  are  then 
said  to  be  hypogynous,  or  inferior,  and  the  gyncecium  is  free  and 
superior.  The  entire  flower  may  also  be  described  as  hypogynous. 
When,  however,  there  is  some  degree  of  adhesion  and  the 
petals  and  stamens,  instead  of  springing  from  the  receptacle, 
appear  to  spring  from  the  calyx-tube,  the  apparent  insertions 
of  the  sepals,  petals  and  stamens  are  no  longer  directly 
below  the  gynoecium,  but  are  carried  out  away  from  the 
receptacle  and  form  a  circle  around  the  gynoecium.  In 
this  case  the  sepals,  petals  and  stamens  are  per  igy  nous 
and  the  flower  itself  is  said  to  be  perigynous.  Here,  however, 


49 

there  is  no  adhesion  between  the  gynoeciuin  and  the  other 
floral  members,  the  gynoscium  is  still  quite  free  from  the 
latter  and  is  superior.  Finally,  there  are  cases  in  which 
the  other  floral  members  are  united  with  the  gynoecium  and 
in  which  the  adhesion  is  carried  right  up  to  the  top  of  the 
gynoecium,  so  that  the  latter  appears  to  be  completely  enclosed 
and  covered  in,  while  the  sepals,  petals,  and  stamens  appear 
to  arise  on  the  top  of,  or  above,  the  gynoecium.  The  sepals,, 
petals  and  stamens  are  here  said  to  be  epigynous,  or  superior,, 
the  flower  itself  is  epigynous  and  the  gynoecium  is  inferior.  For 
illustrations  see  Figs.  11 — 14,  Plate  X.  The  typical  cases  noted 
above  are  linked  together  by  a  number  of  intermediate  forms ; 
thus  in  some  flowers  there  is  adhesion  between  the  gynoecium 
and  the  other  members,  but  this  adhesion  only  extends  about 
half-way  up  the  gynoecium.  Such  flowers  are  best  considered 
as  imperfectly  epigynous,  and  the  gynoecium  may  be  described 
as  half -inferior.  In  the  case  of  other  flowers,  where  the 
insert  ons  of  the  petals  and  stamens  are  only  very  slightly  re- 
moved from  the  receptacle,  it  is  often  difficult  to  decide  if 
they  should  be  classed  as  hypogynous  or  perigynous.  The 
fact  that,  sometimes,  the  so-called  calyx-tube  is  found  to  bear 
rudimentary  leaves,  indicates  that  this  is',  in  many  cases,  really 
an  outgrowth  of  the  floral  axis,  or  receptacle,  and  that  it  does 
not  consist  merely  of  the  coalescent  bases  of  the  sepals  and 
other  floral  leaves. 

46.  The  torus,  or   floral   recep-  The  Diso. 
tacle,    frequently   grows   out     and     forms   what    is     called    a 

disc,  this  being  merely  an  expansion  of  the  receptacle. 
It  is  usually  a  fleshy,  swollen  ring  of  tissue  which  may, 
or  may  not,  be  lobed,  or  divided.  When  the  disc  is 
divided  into  distinct  portions  the  separate  parts  of  it  are  termed 
glands.  The  disc  and  glands  frequently  excrete  a  sweet, 
sugary  fluid  commonly  known  as  nectar  and  each  is  then  called 
a  nectary.  The  disc  is  usually  developed  between  the  stamens 
and  the  gynoecium,  but  sometimes  also  between  the  petals  and 
stamens,  and  sometimes  it  bears  the  insertions  of  the  petals 
and  stamens.  According  to  its  apparent  place  of  origin,  the 
disc  may  be  hypogynous,  perigynous,  or  epigynous. 

47.  In  addition    to    the   terms  Descriptive 

which    have    been    already    given    for   ordinary    leaves,   the  Terms  for 
following    are  often   used  for  describing  the  shape    cf  sepals  and 

and  petals.    If  they  have   a  narrow  base  supporting  a  broad 
expanded  blade,  the  former  is  termed  the  claw  and  the  latter 

the    blade,    or   lamina.       If     strap-shapsd    they   are     termed 


50 

liguliform.*  Sometimes  there  is  an  outgrowth  from  the 
inner  face  of  the  petals,  usually  at  the  junction  of  the  blade 
and  claw,  or  at  the  throat  of  a  gamopetalous  corolla,  which 
is  often  scale-like  and  reminds  one  of  the  ligule  of  Grasses. 
Such  outgrowths  in  a  flower,  collectively,  constitute  the  so- 
•called  corona. 

The  calyx  or  corolla  is  said  to  be  :— 

Bi-labiate,  if  divided  into  two  more  or  less  distinct  lips. 

Rotate,  or  wheel-shaped,  when  with  a  spreading  limb 
and  very  short  tube. 

Crateriform,  or  cup-shaped,  when  like  rotate  but  with 
the  limb  curving  upwards. 

Hypocrateriform,  or  salver -shaped,  when  with  a  long 
narrow  tube,  abruptly  expanding  into  a  broad 
limb. 

Trumpet-shaped,  if  salver-shaped  with  the  tube 
expanding  near  the  top. 

Infundibuliform,  or  funnel-shaped,  when  the  tube 
expands  gradually  from  the  base. 

Campanulate,  or  bell-shaped,  when  the  tube  is  expand- 
ed from  the  base,  with  its  length  not  more 
than  i/wice  its  breadth  and  with  the  sides 
spreading  outwards  at  the  top. 

Urceolate,  or  urn-shaped,  when  the  tube  is  swollen 
in  the  middle  and  contracted  at  the  top  and  base. 

Tubular,  when  there  is  a  conspicuous  nearly  cylin- 
drical tube  and  a  relatively  inconspicuous  limb. 

Gibbous,  when  with  a  small  pouch-like  swelling  near 
the  base. 

Ventricose,  when  swollen  on  one  side,  like  the  corolla 
of  several  species  of  Strobilanthes. 

Saccate,  when  swollen  at  the  base  like  a  bag. 
Spurred,  when  with  a  slender,  hollow  prolongation. 

The  limb  or  lobes  of  the  calyx  are  sometimes  reduced  to 
hairs  or  bristles  which  are  often  branched  and  feathery  and 
which  constitute  the  so-called  pappus.  The  calyx  in  such  a 
case  is  said  to  be  pappose.  The  pappus  is  often  more  or  less 
invisible  in  the  flower  and  only  becomes  fully  developed  when 
the  fruit  ripens,  as  is  the  case  in  Valeriana  Wallichu. 
.  48.  The  way  in  which  the 

parts   of     the    flower   are    arranged   in    the    bud     is   called 

*  The    term    ligulate    is    sometimes    used  in    th;s    sense    but.   it    also    means 
furnished  with  a  ligule. 


51 

-aestivation.     The  aestivation  of  the  sepals  and  petals  is : — 

crumpled,  when  they  are  irregularly  folded  ; 

valvate,  when  ..their  margins  are  in  contact  and  there 
is  no  overlapping.  When  the  margins,  besides 
being  in  contact,  are  turned  straight  inwards  it 
is  valvate-induplicate,  and  when  the  margins  are 
involute  it  is  valvate -involute  ; 

imbricate,  when  their  margins  overlap  and  at  least 
one  of  each  whorl  has  both  margins  covered ; 

contorted,  when  every  one  of  the  same  whorl  has  one 
margin  covered  and  one  margin  uncovered.  Assum  • 
ing  that  the  observer  stands  outside  the  whorl  and 
therefore  behind  the  sepals,  or  petals,  as  the  case 
may  be,  when  the  right-hand  margin  of  each  is  un- 
covered they  are  said  to  overlap  to  the  right  and 
when  the  left-hand  margin  of  each  is  uncovered  to 
overlap  to  the  left. 

In  addition  to  overlapping  the  leaves  may  be  twisted, 
and  this  twisting  may,  or  may  not,  be  in  the  same 
direction  as  the  overlap. 

plaited,  or  plicate,  when  they  are  in  longitudinal 
folds. 

For  illustrations  see  Figs.  15 — 19,  Plate  X. 

49.  As  already  noted  a  normal  characters 

.-stamen   consists  of  a  swollen  head  (the  anther)    on    a    stalk  of  Stamens, 
(the  filament). 

A  typical  anther  consists  of  two  longitudinal  halves,  or 
lobes,  called  thecae,  each  of  which,  at  least  when  young,  contains 
2  cells,  or  chambers.  These  cells  contain  the  pollen  and  are 
called  the  pollen-sacs.  When  mature  the  anther  opens  (i.e., 
dehisces)  and  allows  the  pollen  to  escape.  Each  theca  com- 
monly opens  by  a  longitudinal  slit  in  such  a  way  that  one  slit 
opens  at  once  into  both  sacs,  each  theca  thus  appearing  to  be 
one-celled.  The  two  sacs  in  each  theca  also  are  sometimes 
confluent  before  dehiscence.  In  a  few  cases  the  anther  has  only 
one  cell.  According  to  the  number  of  cells  it  is  uni-,  bi-,  or 
quadri-locular.  The  anthers  sometimes  dehisce  by  means  of 
small  holes  (pores)  and  sometimes  by  valves.  That  portion  of 
the  filament  which  runs  up  between  the  anther-lobes  is  called 
the  connective.  The  filament  is  sometimes  wanting  and  then 
the  anther  is  said  to  be  sessile.  Sometimes  the  anther  is  not 
developed  and  sir.ch  an  imperfect  stamen  is  called  a  siaminodium 

f)T  staminode. 

E  2 


52 

The  stamens  are  said  to  be  :— 

monaddphous,  when  their  filaments  are  all  united  into  - 

a  tube ; 

diadelphous,  when  in  two  sets,  or  bundles  ; 
polyadelphous,  when  in  several  sets ; 
syngenesious,  when  the  anthers  are  united  in  a  ring ; 
didynamous,  when  there  are  4  stamen s;    two  of  them 

long  and  two  of  them  short ; 
tetradynamous,  when  there  are  6  stamens,  4  of   them 

long  and  two  of  them  short. 

If  the  stamens  are  inserted  on  the  petals  they  are  cpi- 
peialous,  and  if  opposite  the  petals  they  are  antipetalous. 

If  the  stamens  are  the  same  number  as  the  sepals  and 
petals,  the  flower  is  isostemonous,  if  the  stamens  are  double  the 
number,  the  flower  is  diplostemonous,  and  if  they  are  double 
the  number  and  they  do  not  alternate  correctly  with  the 
petals,  the  flower  is  obdiplostemonous. 

Stamens  which  protrude  beyond  the  corolla  are  said  to  be* 
exserted ;  if  hidden  by  the  corolla  they  are  included. 

The  anther  is  said  to  be :— 

introrse,  when  the  thecse  are  turned  inwards  towards  the 
centre  of  the  flower,  i.e.  when  the  thecaB  are  on 
that  side  of  the  anther  which  faces  the  floral  axis 
(ventral  side) ; 

',  «.  extrorse,  when  the  theca3  are  turned  outwards,  facing 
away  from  the  floral  axis,  i.e.  when  the  thecse 
are  on  that  side  of  the  anther  which  is  turned 
away  from  the  axis  (dorsal  side) ; 

innate,  when  the  filament  appears  to  be  fixed  in  the 
centre  of  the  base  of  the  anther,  the  thecaa  not 
facing  inwards  or  outwards.  The  anther  is  also 
sometimes  said  to  be  basi fixed  in  this  case  ; 

adnate,  when  the  filament  appears  to  run  up  at  the  back, 
or  front,  of  the  anther,  the  thecae  facing  inward s,. 
or  outwards ; 

versatile,  when  it  swings  freely  at  the  end  of  the  filament ; 

dorsifzed,  when  the  filament  appears  to  be  inserted  in 
the  back  of  the  anther ; 

The  stamens  sometimes  branch,  e.g.  in  the  Castor  Oil  Plant. 
This  is  often  indicated  by  the  anthers  being  only  one-celled, 
i  e.  each  is  really  half  a  stamen,  although  in  some  cases  this  may 
be  due  to  the  non-development  of  half  the  stamen  Anther:? 


53 

may  be  provided  with  bristles,  points,  spur-or  horn -like  pro- 
cesses. The  connective  is  also  sometimes  prolonged '  into  a 
point  beyond  the  anthers.  The  filament  is  usually  slender,  but 
varies  in  shape  and  is  sometimes  provided  with  wings. 

Pollen  is  usually  a  powdery  substance,  which,  when  magni- 
fied, is  seen  to  consist  of  distinct  grains.  The  shape  and  size  of 
the  grains  varies  in  different  plants  and  the  surface  of  the 
grains  may,  or  may  not,  be  provided  with  points,  knobs, 
ridges,  or  other  markings.  In  those  plants,  the  pollen  of  which 
is  distributed  by  the  wind,  the  grains  are  smooth  and  in  others, 
the  pollen  of  which  is  distributed  by  insects,  or  birds,  the 
grains  are  usually  sticky.  In  some  plants  the  grains  adhere 
together  in  masses,  the  latter  being  called  pollinia. 

50.  As  has  been  already  pointed  characters 

out,  a  simple  pistil  really  consists  of  a  single  infolded  of 
leaf  which  bears  the  ovules  on  its  inner  margins.  Such 
an  ovuliferous  pistil-leaf  is  called  a  carpel.  A  pistil  con- 
sisting of  one  carpel  is  said  to  be  simple,  if  of  more  than 
one  carpel  it  is  compound.  A  gynoecium  consisting  of  simple 
pistils  is  called  apocarpous,  the  carpels  being  distinct,  a  gyn- 
cecium  consisting  of  a  compound  pistil  is  syncarpous,  the 
carpels  being  combined.  In  a  simple  pistil  the  lower 
surface,  or  back,  of  the  carpel  is  represented  by  the  exterior 
of  the  pistil,  the  upper  surface,  or  front,  of  the  carpel  forms 
the  interior  of  the  pistil.  The  united  margins  of  the  carpel 
which  bear  the  ovules  look  towards  the  centre  of  the  flower 
(the  axis)  and  form  what  is  called  the  ventral  suture  or  seam. 
The  opposite  ridge,  along  the  back  of  the  pistil,  representing 
the  midrib  of  the  carpel,  forms  the  dorsal  suture.  In  a  com- 
pound pistil  the  carpels  may  be  joined  by  their  margins  only, 
or  these  may  grow  inwards  and  be  united  in  the  centre  of 
the  pistil.  In  the  former  case  the  pistil  is  one -celled  and  in 
the  latter  it  is  divided  by  partitions  called  dissepiments  into 
several  cells,  there  being  as  a  rule  as  many  cells  as  there  are 
carpels.  (Sometimes  this  is  not  the  case,  there  being  so-caQed 
false-partitions  which  grow  in  from  the  back  of  the  carpels.) 

The  surface  on  which  the  ovules  are  borne,  which  is  often 
more  or  less  enlarged,  is  called  the  placenta  and  the  way  in 
which  the  placentae  are  arranged  is  called  the  placentation.  In 
the  case  of  a  compound  pistil  in  which  the  carpels  cohere  by 
their  margins  only,  the  placentae  are  on  the  walls  of  the  pistil 
and  the  placentation  is  called  parietal ;  when  the  margins  are 
brought  together  and  cohere  in  the  axis  of  the  pistil,  the 
placentae  are  in  the  inner  angles  of  the  cells,  and  the  placenta- 


54 

tion  is  called  axile.  In  some  cases  the  ovules  are  borne  on  a 
central  axis  which  is  not  connected  by  dissepiments  with  the 
walls  of  the  ovary  and  the  pistil  is  consequently  one -celled. 
The  placentation  in  such  cases  is  called  free-central. 

As  a  rule  the  ovules  are  borne  on  the  margins  of  the 
carpels,  but  in  some  cases  they  are  developed  all  over  the 
internal  face  of  the  carpels  and  the  placentation  is  then 
called  superficial. 

In  a  compound  pistil  the  styles  may,  or  may  not,  be  more 
or  less  completely  combined  and  the  number  of  styles,  or  of 
the  divisions  of  the  style,  or  stigma,  often  indicates  the 
number  of  carpels  in  the  pistil. 

Owing  to  one  side  of  the  ovary  growing  more  rapidly 
than  another,  the  style  sometimes  becomes  displaced  to  one 
side  and  appears  to  arise  from  the  base  instead  of  from  the 
apex  of  the  ovary.  Such  a  style  is  said  to  be  qynobasic. 

In  some  cases  the  floral  receptacle,  or  torus,   is  prolonged 
upwards  between  the  carpels  forming  a  kind  of  beak  to  which 
the  ovaries  and  styles  adhere  and  from   which   they   separate 
when  the  fruit  ripens.     Such  a  beak  is  called  a  carpophore. 
Characters  51.   The    rounded    out-growths 

of  Ovules.  Of  -f/kg  carpeis  wnich  are  destined  to  become  the  seeds 
are  called  ovules.  These  may  be  sessile,  or  they  may  be 
provided  with  a  distinct  stalk  called  the  funicle.  A 
typical  ovule  consists  of  a  central  portion  of  soft  tissue 
called  the  nucellus  which  is  invested  with  one  or  two  coats 
called  the  integuments.  The  latter  do  not  completely  cover  in 
the  nucellus,  a  narrow  hole  being  left  at  the  apex  which  is 
called  the  micropyle.  The  spot  from  which  the  integuments 
originate  at  the  base  of  the  ovule  is  called  the  chalaza. 

Ovules  are : — 

erect,  when  they  rise  from  the  base  of  the  ovary ; 
ascending,  when  they  rise  obliquely  and  are  attached  near 

the  bottom  of  the  ovary ; 
horizontal,   when   they   are  borne    on    the  side    of  the 

ovary  with  their  long  axis  horizontal ; 
suspended,  when  hanging  from  the  apex    of  the  ovary; 
pendulous,  when  hanging  obliquely  from  near  the  apex 

of  the  ovary; 
orthotropous,  when  they  are    straight,  with  the  chalaza 

and  micropyle  in  one   straight  line    and  the  chalaza 

at  the  apex  of  the  funicle , 
campylotropous,    when    the     line    joining    the    chalaza 

and  micropyle  is  curved,  the  micropyle  being  brought 


55 

round  and  down  near  the  chalaza.  This  is  caused 
by  one  side  of  the  ovule  growing  faster  than  the 
other ; 

anatropous,  when  they  are  completely  inverted.  The 
chalaza  and  the  micropyle  are  in  the  same  straight 
line,  but  the  upper  part  of  the  funicle  is  apparently 
adherent  to  the  side  of  the  ovule  and  forms  a  ridge 

O 

called  the  raphe. 

Intermediate  forms  between  the  above  types  of  course 
occur. 

The  spot  where  the  end  of  the  funicle  is  joined  to  the  ovule 
is  called  the  hilum.  When  the  ripe  seed  separates  from  its  stalk 
a  scar  marks  the  position  of  the  hilum.  In  orthotropous  and 
campylotropous  ovules  the  hilum  and  chalaza  coincide ;  in 
anatropous  ovules  they  are  at  opposite  ends  of  the  ovule. 
When  the  raphe  is  turned  towards  the  ventral  suture  of  the 
carpel  it  is  said  to  be  ventral  and  when  turned  towards  the 
dorsal  suture  it  is  dorsal.  In  the  nucellus  of  the  ovule  there  is 
a  large  cell  called  the  embryo-sac,  containing  protoplasm  which 
is  more  or  less  divided  up  into  separate  da  lighter- cells  which 
may,  or  may  not,  be  provided  with  delicate  cell-walls.  When 
a  ripe  pollen-grain  reaches  the  sticky,  surface  of  the  stigma  it 
germinates.  Its  outer  coat  is  burst  and  a  long  delicate  tube  is 
sent  out,  called  the  pollen-tube  into  which  pass  the  proto- 
plasmic contents  of  the  pollen-grain.  This  tube  penetrates 
the  tissue  of  the  stigma,  and  of  the  style  if  there  is  one,  usually 
enters  the  micropyle  and  grows  until  its  tip  comes  in  contact 
with  the  apex  of  the  embryo-sac.  A  portion  of  the  protoplasm 
then  passes  out  from  the  pollen-tube  and  unites  with  a  part 
of  the  protoplasm  in  the  embryo-sac,  and  with  the  fusion  of 
these  separate  masses  of  protoplasm  into  one  fertilisation  is 
completed.  The  lump  of  protoplasm  surrounds  itself  with  a 
cell-wall,  begins  to  divide,  and  develops  into  the  so-called 
embryo,  or  young  plant,  the  ovule  now  having  become  the 
seed. 

52.    Having      completed      our  Types  Or 

examination   of  the   parts   of  the   flower   a  few  points  which  Flowers 
may   cause  difficulty    remain  to  be  noted. 

In  cases  where  some  of  the  floral  parts  are  suppressed  Difficulty, 
and  not  developed,  it  is  not  always  easy  to  decide  on  the  true 
nature  of  the  parts  which  are  present.  Thus  in  the  flower  of 
Clematis,  there  are  usually  4  conspicuous  leaves  which  look 
like  petals,  but  which  are  really  sepals,  no  petals  being  develop- 
ed. This  is  indicated  by  the  fact  that  rudimentary  petals  are 


56 

sometimes  found  in  their  proper  place  inside  the  petal-like 
sepals.  The  presence  of  rudiments  is  thus  seen  to  be  of  im- 
portance, and  in  doubtful  cases  a  careful  comparison  of  allied 
plants  often  helps  us  to  arrive  at  a  decision. 

The  student  should  carefully  examine  the  flowers  of  the 
following  plants  as  they  may  cause  some  difficulty.  They  are 
fairly  typical  of  the  flowers  of  a  large  number  of  important 
plants  and  a  few  special  terms  are  necessary  for  their  des- 
cription. 

wheat,  (a)  Wheat. — The  flowers  are  in  spikelets,  and  each  spikelet 

contains  from  3 — 5  flowers.  Each  flower  consists  of  3  stamens 
and  one  pistil  with  two  feathery  stigmas,  while  the  perianth  is 
represented  by  two  small  hypogynous  scales  called  lodicules. 
Each  flower  is  enclosed  between  2  scaly  bracts,  the  outer  of 
which  is  the  flowering -glume,  while  the  inner  and  smaller  is  the 
palea.  Finally  outside  of  all,  at  the  base  of  the  spikelet  there 
are  two  more  dry,  scaly  bracts,  which  are  called  the  outer- 
glumes. 

The  flowers  of  grasses  RW!   I  am  boos  closely  resemble  this 
type. 

Pea-  (b)  Common  Pea,  Bean,  Butea  frondosa,  or  a  Dalbergia. — 

The  striking  peculiarity  here  is  in  the  corolla,  which  is 
called  papilionaceous  from  some  supposed  resemblance  to 
a  butterfly.  The  two  lower  anterior  petals,  which  are  inside 
the  others,  are  narrow  and  fit  close  together,  forming  the  so- 
called  keel  (carina).  Outside  the  keel  are  two  more  narrow 
petals  called  the  wings  (alae),  one  on  each  side  of  the  keel, 
while  finally,  outside  the  wings,  there  is  a  large  posterior 
upper  petal  called  the  standard  (vexillum).  It  should  also 
be  noted  that  the  filaments  of  the  stamens  are  more  or  less 
united  into  a  tube. 

Pine.  (c)  Pine    (Pinus). — The   flowers  are    unisexual.     The  male 

flower  consists  of  an  axis  around  which  are  spirally  arranged 
the  numerous  scale -like  stamens,  each  stamen  bearing  two 
pollen-sacs  on  its  under  surface. 

The  female  flower  also  consists  of  an  axis  with  numerous 
spirally-arranged  scales  which  represent  the  carpels.  In  the 
axil  of  each  carpel  there  are  two  ovules,  lying  on  another  scale 
called  the  placental  scale.  As  the  fruit  ripens,  the  placental 
scales  become  woody  and  form  the  familiar  Pine  cone.  In 
this  nase  the  flowers  are  of  very  simple  construction  and  there 
is  no  perianth.  Both  the  stamens  and  carpels  are  scale-like, 
and  finally  the  carpels  do  not  form  an  enclosed  ovary,  the 
ovules  merely  lying  exposed  on  their  upper  surfaces.  Plants 


57 

like   this  which  have  their  ovules  uncovered  are  called  Gymno- 
sperms  (naked  seeded  plants),    while  other  plants  having  their  and  Anglos 
ovules  enclosed  in  an  ovary  are  distinguished  as  Angiosperms.  P6™8- 

53.  When  studying  flowers    it 

is  convenient     to   be    able     to   shortly   and    clearly   express  Di,grarrs 
on     paper    the     number,      relative      positions      and     other  Floral 
characteristics      of     the     floral     parts,      instead     of     giving  and  Short- 
long     written    descriptions.       This    can    be    done   by    floral  hand- 
diagrams   and  formulae.     A   floral   diagram   is  a  plan    of   the 
flower  in  which  the  position    of  the  various  floral  parts  is  in- 
dicated   by    a    diagramatic    cross-section.      To    indicate  the 
posterior  and  anterior  sides  of  the   flower,  the  position  of  the 
bract,  in  the  axil  of  which   the  flower  is  borne,  must  be    shown 
.and  of  the  main  axis  from  which  the  flower  springs. 

In  floral  formulae  the  following  equivalents  are  used:  — 

K=calyx  (sepals). 
C=  corolla  (petals). 
P  =  perianth. 

A=andro3cium  (stamens). 
G  =  gynoacium    (carpels). 

The    following    examples    will    indicate  how  formulae    are 
;<written  :  — 


G  (3).  —  The  flower  has  a  polyphyllous  perianth 
of  6  leaves,  in  two  whorls,  each  of  3  leaves. 
There  are  6  free  stamens,  also  in  two 
whorls,  each  of  3,  and  a  syncarpous, 
infeiior  gynoecium  of  3  carpels.  Nothing 
being  said  to  the  contrary  the  flower  is 
regular. 

€5  A  <»  Goo.  —  There   are  here  5  free  sepals  and  5  free 
petals,  while  the  stamens  and  carpels   are 
too    numerous    for    easy    counting.     The 
stamens  are  free  and  the  gyncecium  apocar- 
pous and  superior. 

A  €5  A(s  +  5)  GI.  —  Here  there  is  a  gamosepalous  calyx 
of  5  sepals,  a  zygomorphic  corolla  of  5 
free  petals,  10  stamens  with  their  fila- 
ments united  into  a  tube  and  a  gyncecium 
of  one  superior  simple  pistil. 


58 


S  ed. 


Testa, 
Tegmen  Aril. 


Albuminous 
Ex  albumin- 
ous seeds. 


Cotyledons 
Radicle  and 
Plumule  of 
Embryo. 


CHAPTER  V.— THE  'SEED  AND  FRUIT. 

54.  The  normal  seed  is  en- 
veloped in  one  or  two  outer  coats  which  correspond  to 
the  integuments  of  the  ovule.  The  outer  is  usually  firm 
and  often  hard  and  is  called  the  testa  ;  the  inner,  when  it  is 
present,  is  thin  and  delicate  and  is  called  the  tegmen.  In  some 
cases  there  is  an  additional  covering  to  the  seed  developed 
after  fertilisation  has  taken  place  and  which  is  accordingly  not 
visible  on  the  ovule.  This  covering  usually  originates  from 
the  funicle  or  placenta  but  occasionally  also  from  the  micro - 
pyle.  It  is  known  as  an  aril,  e.g.  the  pulp  of  the  Lichi. 

An  outgrowth  morphologically  similar  to  an  aril  but 
which  is  smaller  and  may  develop  from  various  parts  of  the 
seed  is  called  a  caruncle,  or  sfrophiole. 

The  testa  of  the  seed  sometimes  grows  out  into  a 
membranous  wing,  as  in  Oroxylum  indicum,  see  Fig.  1,  Plate 
XII,  and  sometimes  is  provided  with  long  soft  hairs,  as  in  the 
Cotton  plant,  these  structures  aiding  the  distribution  of  the 
seed  by  wind. 

Inside  the  seed- coats  we  find  what  is  popularly  called  the 
kernel ;  this  may  consist  only  of  the  embryo,  or  the  latter  may 
occupy  only  a  portion  of  it,  the  remainder  of  the  kernel 
consisting  of  the  so-called  albumen.  (This  is  cellular  tissue 
densely  packed  with  food-materials,  such  as  starch,  or  other 
substances,  which  are  to  serve  as  food  for  the  young  embryo.) 
In  the  former  case  the  seed  is  said  to  be  exalbuminous  and 
in  the  latter  albuminous.  As  examples  of  the  former  we  may 
take  those  of  the  Oak  and  Oroxylum  indicum,  see  Figs.  1  and  2, 
Plates  I  and  XII,  and  of  the  latter  the  seed  of  the  Pine  (Pinus). 

If  the  albumen  has  originated  inside  the   embrvo-sac  it  is 

o  v 

called  endosperm,  if  it  is  a  part  of  the  tissue  of  the  nucellus  it 
is  called  perisperm. 

Albumen  may  be  mealy  (when  it  is  easily  broken  into 
powder),  oily,  fleshy,  or  hard,  and  even  bony.  If  the  outer 
surface  of  the  albumen  is  crumpled,  or  puckered,  into  narrow 
folds  it  is  said  to  be  ruminate. 

In  a  well-developed  embryo  wre  can  distinguish  an  axis 
bearing  one  or  more  minute  leaves ;  these  leaves  are  called 
cotyledons.  That  part  of  the  axis  situated  below  the  insertion 
of  the  cotyledons  and  from  which  the  primary  root  of  the 
seedling  will  be  developed  is  called  the  radicle,  while  the  part 
of  the  axis  above  the  cotyledons,  which  is  the  end  of  the 


59 

minute  shoot,  is  the  plumule.  The  radicle  of  the  embryo 
always  points  towards  the  micropyle.  The  position  of  the 
embryo  in  the  albumen  is  often  characteristic ;  it  may  be 
straight  or  curved,  it  may  be  in  the  centre  of  the  albumen 
(axial),  eccentric,  or  external  to  it  as  in  Grasses.  The  cotyledons 
may  be  placed  straight  in  the  seed  or  variously  folded.  The 
cotyledons  are  :— 

incumbent,   when   the   radicle  is  laid  along  the  back  of 

one  cotyledon  ; 

accumbent,  when  the   radicle   is  laid  along  the  edge  of 
the  cotyledons. 

In  exalbuminous  seeds  all  the  food  material  is  stored  in 
the  tissue  of  the  embryo  itself,  and  usually  in  the  cotyledons, 
which  are  then  thick  and  fleshy.  In  other  cases  the  cotyledons 
are  thin  and  more  or  less  foliaceous.  On  germination  taking 
place  the  cotyledons  may  remain  below  the  ground,  as  in  the 
Oak,  see  Figs.  1  and  2,  Plate  I,  or  be  raised  up  on  the  growing 
stem  into  the  light  and  air,  when  they  usually  become  green 
and  more  or  less  like  ordinary  leaves,  as  in  Oroxylum  indicum, 
see  Plate  XII.  The  portion  of  the  stem  of  a  seedling  situated 
below  the  insertion  of  the  cotyledons  is  called  the  hypocotyl  Hypocotyi. 
while  that  part  of  the  stem  lying  between  the  insertion  of  the  EPlcotyi- 
cotyledons  and  that  of  the  first  foliage  leaf  or  leaves  is  the 
epicotyl. 

The  plants  termed  Angiosperms,   which   have  been  men-  Monocotyie- 
tioned  above,  are  sub-divided    into  two  great  groups  known 
as  the  Dicotyledons  and    Monocotyledons  respectively.     In  the  dons, 
former,  which  includes    most  of  our  important  forest    species 
the  embryo  has  typically    two   opposite    cotyledons';  in    th3 
latter,  which  includes  such  plants   as  the  Grasses,  Bamboos 
and  Palms,  the  embryo  has  typically  only  one  cotyledon. 

55.  The  mature  gyncecium  of  a  Fruit. 

flower  containing  the  seeds,  with  everything  which  may  be 
joined  to  it,  constitutes  the  fruit.  The  fruit  may  consist 
of  a  single  ripe  pistil,  in  which  case  it  is  a  simple  fruit, 
or  of  the  collection  of  separate  pistils  belonging  to  one 
flower  when  it  is  an  aggregate  fruit,  or  of  the  pistils  belonging 
to  several  flowers  united  together,  when  it  is  a  multiple  fruit, 
or  infructescence.  If  it  is  provided  with  a  stalk  the  fruit  is 
said  to  be  stipitate,  if  not,  it  is  sessile. 

The  ovary  wall  becomes  the  wall  of  the  fruit  which  is" 
called  the  pericarp  and  this  immediately  surrounds  the  seeds. 
The  position  occupied  by  the  embryo  with  reference  to  the 
rest  of  the  fruit  is  important,  and  the  radicle  is  said  to  be 


60 

superior  when  it  points  upwards  towards  the  apex  of  the  fruit 
and  inferior  when  it  is  directed  downwards  towards  the  base. 
The  pericarp  may,  or  may  not,  be  differentiated  into  definite 
layers  and  sometimes  as  many  as  3  can  be  distinguished,  in 
which  case  the  outer  layer  is  called  the  epicarp,  the  inner 
layer  the  endocarp,  and  the  middle  layer  the  mesocarp. 

In  many  cases  a  conspicuous  part  of  the  fruit  is  not  deve- 
loped from  the  pistils,  but  from  some  other  part  of  the  flower. 
Thus  in  the  Strawberry  the  succulent  portion,  which  forms  the 
greater  part  of  the  fruit,  is  developed  from  the  receptacle,  in 
the  Sal  the  calyx  persists  and  develops  into  large  veined 
wings,  in  Semecarpus  Anacardium  the  base  of  the  calyx 
and  the  receptacle  form  a  conspicuous  fleshy  base  to  the 
fruit,  while  in  the  Oak  the  "cup"'  of  the  acorn  is  formed  of  bracts. 
Fruits  which,  when  ripe,  open  to  let  the  seeds  escape  are 
said  to  be  dehiscent,  those  which  do  not  do  so  are  indehiscent. 
Types  of  56.  The  principal  types  of  fruits 

Fmit. 


A.  —  DRY   AND   DEHISCENT. 

(1)  Follicle,  consisting  of  one  carpel  and   dehiscing  by  one 
suture. 

(2)  Legume,  consisting  of  one  carpel  and  dehiscing  both  by 
the  dorsal  and  ventral  sutures.     A  legume  which  is  much  con- 
stricted between  the    seeds   is  called  a  lomentum.     This  often 
breaks  up  into  one-seeded  joints  when  ripe. 

(3)  Capsule,  arising  from  a  compound  pistil.     May  be  one 
or    many    celled.     If   a    capsule    splits    open    along    the  dis- 
sepiments its  dehiscence  is  septicidal,  if  it  splits  open  through 
the  back  of  each  carpel  the  dehiscence  is  loculicidal.     In   both 
cases  the   dissepiments    separate  from   the  axis.     When  the 
dissepiments  remain  attached  to  the  axis  and  the  valves  merely 
separate    from  the  ends    of   the    dissepiments,   as    in  Cedrela 
Toona,  the  dehiscence  is  septijragal.     In  some  cases  the  dehis> 
cence  may  be  partly  according  to  one  and  partly  according  to 
another  of  these  types.    In  septicidal  dehiscence,  for  instance, 
if  the  dissepiments  remain  attached  to  the  axis,    the  valves  of 
the  capsule  may  separate  from  the  ends  of  the  dissepiments  as 
in  septifragal  dehiscence. 

(4)  Schizocarp,  arising  from  a  compound  pistil    and  break- 
ing up  at  maturity  into  distinct  portions,   each  of  which  is 
usually    indehiscent    and  looks  like  a  separate    fruit,    being 
called  a  coccus.     When  a  coccus  is  small  and  one  -seeded  it  is 
called  a  nutlet. 


61 

B. — DRY  AND  INDEHISCENT. 

(5)  Nut,  a  one-seeded  fruit  with  a    hard,    dry,    pericarp 
such  as  the  acorn  of  the  Oak. 

(6)  Achene,  a  one-seeded    fruit    with    a    thin,  leathery, 
pericarp. 

(7)  Caryopsis,    similar  to  an  achene    but  the  pericarp  is 
closely  adherent  to  the  seed,  the  latter  completely  filling  the 
cell,  whereas  in  the  achene  this  is  not  so. 

(8)  Samara,  any  dry  indehiscent  fruit  which  is  provided 
with  a  wing  developed  from  the  pericarp. 

C. — SUCCULENT. 

(9)  Berry,  the  whole  pericarp  with  the  exception    of    the 
outer  skin  (epicarp)  is  succulent  and  the  seeds  are  immersed 
in  the  pulp. 

(If))  Drupe,  the  pericarp  is  differentiated  into  three  layers; 
the  inner,  or  endocarp,  is  hard  and  forms  the  so-called  stone 
of  the  fruit,  containing  the  seed,  or  seeds,  and  which  may  be 
one  or  more  celled.  The  stone  is  immersed  in  the  mesocarp 
which  is  generally  succulent,  while  the  epicarp  forms  a  skin 
over  the  mesocarp.  The  succulent  mesocarp  is  sometimes 
called  the  sarcocarp  and  the  hard  endocarp  the  putamen. 
In  the  Mango  the  portion  eaten  is  the  sarcocarp  and  the 
endocarp  is  fibrous. 

When  a  drupe  is  formed  from  a  gyncecium  composed  of 
more  than  one  carpel  and  the  carpels  separate  and  give  rise 
each  to  a  distinct  endocarp,  so  that  in  the  ripe  drupe  there  is 
more  than  one  stone,  each  of  these  stones  is  called  a  pyrene, 
as  in  Grewia. 

51,    For  describing   the  shape  General 

of  fruits,  or  seeds,  the  terms  which  have  been  noted  above  in  Terms  for 
connection  with   other    solid  parts  of    plants    may  be  used,  thrshape  of 
such  as  fusiform,  conical  and  so  on.     The  terms  given  for  flat  Seeds  and 
surfaces  are  also  employed  for  describing  the  outline  (as  seen 
in    elevation)   of  solid    parts  and  in  addition    the    following 
terms  are  frequently  used  : 

cylindrical,    elongated    with    circular    cross    section. 

spherical,  round ; 

globose,  or  spheroidal,  nearly  spherical ; 

ovoid,  an  egg-shaped  solid ; 

ellipsoid,   an  elliptical  solid ; 

discoid,  circular,  and  flat,  or  depressed,  in  centre  ; 

compressed,    flattened    sideways ; 


62 

depressed,  flattened  from  above  ; 

clavate,  club-shaped,  thickened  above  and  slender  below  ; 

turbinate,  top-shaped,  ob-conical ; 

pyriform,  pear-shaped  ; 

didymous,    slightly   2-lobed  ; 

plano-convex,  flat  on  one  side,  convex  on  the  other; 

concavo-convex,  concave  on  one  side,  convex  on  the  other  ; 

moniliform,  cylindrical,  but  constricted  at  intervals  like 

a  string  of  beads  ; 
torulose,  slightly  moniliform. 


63 

CHAPTER  VI.— GENERAL. 

58.  According   to  their  duration  Duration  of 
pirts  of  plants  are  said  to  be :—  Sant, 

iii  ) 

fugacious,    falling    off   very    early,  almost   as  soon     as 
they   are   developed,    as    the  petals  of    Linseed,    or 
Reinwardtia  trigyna ; 
caducous,  falling  off  early,  e.g.  petals  falling  before  the 

flower  is  fertilized ; 

deciduous,  falling  at  the  usual  season,  e.g.  petals  which 
fall  soon  after  the  fertilization  of  the  flower  has  taken 
place  ; 

persistent,  remaining  attached  to  their  support  beyond 
the  usual  season,  e.g.  petals,  or  sepals,  which  remain 
attached  to  the  fruit. 

Parts  which  wither,  but  still  remain  attached  to  their 
support,  are  said  to  be  marcescent ;  parts  which  persist  and 
increase  in  size  are  said  to  be  accrescent,  such  as  the  sepals  of 
Sal  which  become  enlarged  in  the  fruit. 

59.  According  to    their  texture  Texture  of 
parts  of  plants  are  said  to  be  : —  Members. 

osseous,  bony,  e.g.  the  stone  of  many  fruits  ; 
corneous,  like  horn,  e.g.  albumen  of  Date  Palm ; 
cartilaginous,    hard     and    tough  like    parchment,    e.g. 
endocarp  of  an  apple  which  surrounds  the  "  pips,"  or 
seeds. 
chartaceous,  thin,   like   paper,  e.g.  outer  bark  of  young 

Birch  stems  ; 

coriaceous,  firm  and  tough  like  leather,  e.g.  leaf  of  Mango  ; 
•sub- coriaceous,  thin  and  pliable  like  leaves  of  Berchemia 

floribunda  ; 

membranous,  very  thin  and  pliable  and  somewhat  trans- 
parent like  skin.  e.g.  leaves  of  Staphylea  Emodi. 
If  membranous  but  dry  and  more  or  less  colourless, 
the  texture  is  scarious  ; 

-paleaceous,  like  chaff ;   scarious  but  rather  stiff ; 
-fleshy,  thick  and  soft,  like  leaves  of  Saxifraga  ligulata  ; 
succulent,  fleshy  and  juicy,  like  leaves  of  Fedum  rosulatum; 
crustaceous,  hard  and  brittle,   like   the  r'nd,    or  epicarp; 

of    fruit   of  Grewia  pilosa. 

The  texture  often  varies  according  to  the  age  of  the 
member,  thus  leaves  which  are  membranous  when  young 
may  become  coriaceous  when  mature. 

60.     The  outer  skin  or  surface 
of  plant   mem  bers  is  often    provided   with    pricldes.     These  Gland's. 


64 

may  be  distinguished  from  spines,  which  are  really  branches, 
i.e.  continuations  of  the  internal  tissues,  by  the  fact  that  they 
are  superficial  structures  which  are  easily  broken  off,  and  if 
the  bark  is  stripped  off  they  often  come  away  with  it  such  as 
do  those  of  Roses  and  Rubus. 

A  few  of  the  terms  used  for  describing  the  surface  of 
plant  members  have  been  given  above  and  in  addition  to  them 
the  following  are  in  common  use  :-— 

echinate,  with  sharp  prickles,  like  a  hedgehog; 
verrucose,  or  tuberculate,  warty,  with  knobby  excrescen- 
ces; 

scabrous,  rough  to  the  touch ; 
rugose,  wrinkled. 

The  surface  is  also  often  provided  with  hairs,  or  wax,  and 
said  to  be  : — 

glabrous,  if  without  hairs ; 

glabrescent,  almost  glabrous,  with  very  few  hairs ; 
pubescent,  with  short,  soft,  straight  hairs;  if  the  hairs 
are  very  close  together  and  the  surface  feels  like 
velvet  it  is  said  to  be  pilose,  if  the  hairs  are  very- 
short  and  only  just  perceptible  to  the  touch  the 
surface  is  puberulous ; 

hirsute,  pubescent  but  the  hairs  are  longer  and  stiffer  ; 
villous,  shaggy.     The  hairs  are  long  and  weak   and  not 

matted  ; 
tomentose,  densely  pubescent,  but  the  hairs  are  matted 

and    the    surface  feels  softer  and  more  woolly  ; 
woolly,  tomentose  with  long  hairs,  looking  like  wool ; 
floccose,  woolly  but  the  hairs  are  easily  detached,  e.g. 
those  on  the  undersurf  ace  of  leaves  of  Pyrus  lanata  ; 
mealy,  hairs  are  very  short  and  are  easily  rubbed  off  like 
powder,    e.g.    those   on    young   leaves  of  Loranthus 
pulverulentus ; 

hoary,  or  canescent,  with  a  greyish-white  appearance,  due 
to    very    minute  hairs,  too  small  to  be  easily  distin- 
guished ; 
strigose,  with  pointed,  straight,  stiff  hairs,    lying  along 

the  surface ; 

silky,  with  closely  adpressed,  soft,  fine  hairs  ; 
hispid,  rough  with  rigid  hairs ; 
glaucous,  pale  bluish  green  in  colour,  often  due  to  a  thin 

coating  of  wax,  or  so-called  bloom, 
pruinose,    with    a   covering   of   wax,     or    bloom,    e.g. 
branches  of  Rubus  lasiocarpus. 


65 

The  surface  is  sometimes  covered  with  scales,  as  in 
Elaeagnus,  when  it  is  said  to  be  lepidote  and  in  other 
plants  with  a  sticky  secretion  when  it  is  said  to  be  viscid. 
Hairs  which  have  branches  radiating  from  one  point  are 
called  stellate,  and  those  with  a  swollen  head  excreting  a 
sticky  ^  substance  are  called  glandular,  e.g.  the  hairs  on 
Roses  and  on  the  petiole  of  Corylus  Colurna.  Structures 
known  as  glands  are  often  found  on  the  surface  of  plants. 
These  are  more  or  less  prominent,  somewhat  fleshy,  swellings 
which  may,  or  may  not,  secrete  sticky  substances.  They 
are  common  on  the  leaf-stalks  of  Acacias.  Other  kinds  of 
glands  have  been  described  in  previous  paragraphs. 

61.  The  general   appearance  of  Habit. 

a  plant    is  called    its  habit.     This    depends  largely    on    the 
stem,    whether     it     is     erect,    prostrate,    climbing    and     so 
on,    also    on    whether    the    stem  is    branched  or    not.     The 
simple    columnar    stem  of  most    Palms  is  thus  readily  dis- 
tinguished   from  those  of  the  majority  of    our    forest  trees 
which  are    much    branched.     Habit    also     depends    on    the 
position,  arrangement,  and  number  of  the  branches-,  branchlets, 
and  twigs,  and  on  their  size  and  mode  of  growth.     Most  trees 
for  instance  are    bare  of  branches  for  some    distance  from 
the  ground,  whereas  the  stems  of  shrubs  branch  close  to  the 
ground.    Bombax  malabaricum  also  is  easily   recognised  in   the 
forest  by  the  fact  that  its  branches  are  arranged  in  whorls,   or 
false  whorls,  see  Fig.  3,  Plate  XI.    In  some  trees  .the  branchlets 
and  twigs  are  thick  and  few  in  number,  in  many  of  the    fleshy 
Euphorbias  and  some  Sterculias,  for  instance  :  in    others  there 
is  a  large  number  of  small  branchlets  and  twigs,  as  in  Hard- 
wickia  binata.     In  old  Deodar  the  branches  are  almost  horizon- 
tal and  form  terraces  of  foliage,  while  in  the   spruce,    from    the 
more  or  less  horizontal  branches,  the  branchlets  and  twigs  hang 
downwards.     In  Ailanthus  excelsa  the  branches  are  more  or  less 
decumbent.     In  many  trees,  the  branches  or  twigs  are  pendulous 
or 'drooping,  such  as  are  the  terminal  shoots  of  young  Deodar, 
the   branchlets  of  Anogeissus  acuminata  and   Anjan,  and  the 
branches    of    Salix     babylonica.     Branches    which    leave   the 
parent-stem  or  branch  at  a  wide  angle  are  said  to  be  divaricate, 
such  as  those  of  Combretum  ovalifolium,  or  Hamiltonia  suaveo- 
lens.  On  the  branches  also  to  a  great  extent  depends  the  general 
shape  of  the   crown   which   is   often   very   characteristic ;   in 
Albizzia  stipulata  the  crown  is  broad  and  flat-topped,  in  che  Silver 
Fir  it  is  more  or  less  cylindrical,  in  the  spruce  conical,  and  in 
the  mahua,  Bassia  latifolia,  usually  rounded. 


66 

Herbs,  Trees,  ^  plant  the  serial  stem  of  which  is  herbaceous  and  dies 
down  to  the  ground  annually  is  called  a  herb.  There  are 
some  plants  the  stem  of  which,  although  it,  dies  down 
annually,  is  firm  and  more  or  less  woody,  e.g.  Psoralea 
corylifolia.  They  are  also  usually  classed  as  herbs.  Plants 
of  which  the  serial  stems,  or  the  greater  part  of  them,  per- 
sist for  more  than  one  year  and  which  are  more  or  less  woody* 
are  classed  as  shrubs  and  trees  respectively. 

A  shrub  is  distinguished  from  a  tree  by  its  smaller  size 
and  by  the  fact  that  it  usually  has  branches  near  the  base. 

According  to  their  size,  shrubs  andtrees  are  again  usually 
sub-divided  into  shrubs,  large  shrubs,  small  trees,  medium- 
sized  trees,  large  and  very  large  trees. 

An  undershrub  is  a  plant,  of  the  aerial  stem  of  which  a 
very  small  portion  persists  for  more  than  one  year.  Such  a 
plant  is  also  sometimes  said  to  be  suffruticose,  or  suffru- 
tescent. 

Large  herbs  which  in  size  and  general  appearance  re- 
semble true  shrubs  are  usually  described  as  shrubby,  e.g. 
Sesbania  aculeata. 

Climbing  plants  are  termed  herbaceous  if  their  aerial  stems 
are  herbaceous  and  die  down  annually,  and  woody  if  the 
stems  persist  for  more  than  one  year  and  are  more  or  less 
woody. 

Biennials,  A  plant,  or  part  of  a  plant,  which  only  lives  for  one  year 

Perennials,     is  called  an  annual,  one  which  lives  for  2  years  a  biennial  and 

one  which  lives  for  more  than  2  years    a   perennial.      Annual 

plants   are    always  herbs,    biennial   plants   are  usually  herbs, 

while  a  perennial  plant  may  be  a  herb,  shrub,  or  tree. 

The  persistent  basal  portion  of  the  stem  of  an  undershrub 
and  the  persistent  base  of  an  herbaceous  perennial,  lying  at, 
or  just  below,  the  ground  surface,  and  usually  including  a 
small  portion  of  the  bases  of  the  aerial  stems  and  the  upper 
thickened  portion  of  the  roots  and  from  which  new  herbaceous 
stems  are  annually  produced,  is  called  a  stock,  or  sometimes, 
also,  a  root-stock.\ 

Such  root -stocks  gradually    increase   in    size   and  often 

*  In  a  few  cases  stems  which  persist  for  more  than  one  year  are  distinctly 
herbaceous,  e.g.  those  of  the  Banana. 

|  A  rhizome  is  also  called  a  root-stock  by  some  botanists. 


67 

form  large  woody  masses  at,  or  close  below,  the  ground  sur- 
face.    Examples  are  :— 

Clerodendron    serratum,    Careya    lierbacea    and  Combretum 
nanum. 

A  plant  which  is  usually  found  growing  in  company  with  Gregarious 
many  other  plants  of  the  same  species  is  said  to  be  gregarious,  p"ant8p° 
such  as  the  Sal,  many  Bamboos  and  species   of   Strobilanthes. 

A  plant  the  individuals  of  which  are  widely  scattered  from 
each  other  is  sporadic. 

Although  the  above  distinctions  are  useful  for  describing 
plants  it  must  be  understood  that  here,  as  elsewhere  in  mor- 
phology, the  distinctions  are  not  absolute.  Intermediate  forms 
between  the  types  selected  are  frequently  found  while  in  one 
and  the  same  plant  the  distinctions  cannot  always  be  in- 
sisted on.  In  our  Indian  forests  we  have  many  examples 
of  plants  which  in  some  localities  are  climbers  while  in 
others  they  are  erect  shrubs,  or  trees,  e.g.  Acacia  pennata 
and  Carissa  spinarum.  Again  in  some  localities  Leea  aspera 
and  Clerodendron  serratum  are  perennial  herbs,  in  others 
undershrubs,  and  in  others  large  shrubs.  Trees  which  are 
continually  browsed  by  cattle  when  young  may  be  prevented 
from  developing  normally  and  be  temporarily  reduced  to 
shrubs  ;  in  other  cases  their  serial  stems  may  be  more  or 
less  completely  killed  down  to  the  ground  annually  by  fires, 
or  frost,  and  they  may  thus  be  reduced  temporarily  to 
undershrubs,  or  perennial  herbs,  new  shoots  being  sent  up 
yearly  from  the  stock. 

The  common  arhar,  Cajanus  indicus,  is  usually  an  annual, 
but  is  sometimes  biennial.  By  artificially  preventing  the 
flowering  of  some  plants,  also,  it  has  been  found  possible  to 
convert  an  annual  into  a  perennial.  In  annuals  the  whole 
plant  dies  annually,  and  although  in  perennials  this  does  not 
happen,  still  some  portion  of  the  plant  usually  dies  and 
is  shed  annually.  In  many  trees  and  shrubs  the  leaves  do 
not  live  for  12  months  and  the  plants  are  consequently 
bare  of  green  foliage  during  some  part  of  the  year.  They  are 
called  deciduous,  while  those  plants  which  have  green  foliage  ^ 

, ,  ,  . ,  T  TIL     Evergreen 

throughout  the  year  are  evergreen.     In  some  trees  and  shrub?  Deciduous 
small  twigs   die  and  are  thrown  off,    as  in  Pines,  Phyllanthus  Plants. 
Emblica  and  Strobilanthes  Wallichii,  while  in    the    American 
Swamp   Cypress,    Taxodium  distichum,   large    branches     are 
periodically   shed. 


68 


PAET  II.-ANATOMY, 

1 

Hf;  CHAPTER  I.— CELLS. 

Protopksm.  62.  The  bodies  of  plants  and  of 

animals  consist  primarily  of  protoplasm  ;  this  is  a  colourless 
substance,  the  consistency  of  which  varies  with  the  quantity 
of  water  it  contains  from  that  of  a  viscid  fluid  to  that  of  a 
firm  substance  almost  like  wax.  Protoplasm  always  contains 
carbon,  hydrogen,  oxygen,  nitrogen,  sulphur,  phosphorus 
and  iron.  When  heated  strongly  protoplasm  gives  off  water 
and  then  it  blackens  and  gives  off  ammonia.  If  heated  to 
red  heat  an  ash  is  left  consisting  of  small  quantities  of  mineral 
substances.  Protoplasm,  however,  is  a  living  substance  which 
can  be  easily  killed  by  too  high  a  temperature,  by  small 
quantities  of  poisons  and  by  other  factors.  It  must  not  be 
regarded  as  a  simple  chemical  compound,  and  we  cannot 
produce  protoplasm  artificially  by  collecting  and  mixing  to- 
gether all  the  elements  which  appear  to  enter  into  its  compo- 
sition. 

Some  plants  exist  which  consist  of  only  a  naked  mass  of 
protoplasm.     Such  a  mass  of  living  protoplasm  is  found  to  be 
capable  of  creeping  movements,    portions    of  the   protoplasm 
being  pushed  out  which  draw  the  remainder  of  the  protoplasmic 
body  after  them.     It  is  also  capable  of  taking  up  and  absorb- 
ing bodies   containing  nourishing    materials    with    which    it 
comes  in  contact,  of  building  up  new  protoplasm   and   thus   of 
growing  and  increasing  in  size,  of  respiration,  i.e.  of  absorbing 
oxygen  from  the  air  and  giving  off  carbon  dioxide,  and  finally 
of  throwing  off  pieces  of  itself  which  are  capable  of   an   inde- 
pendent existence.     In  addition  to  these  powers  of  movement, 
nutrition,  growth,    respiration  and  reproduction,    living   pro- 
toplasm, in  common  with    all    living  matter,    possesses  the 
remarkable  characteristic  of  irritability,  i.e.  it  is  sensitive  to, 
and  capable  of    re-acting  in  response    to,  a    stimulus.     The 
quality  is  most  easily  recognised  in  cases  where  the  response 
is  greatly  out  of  proportion  to  the  stimulus.     If  the     leaves  of 
the  sensitive  plant,  Mimosa  pudica,   are  lightly    touched,    the 
leaflets  and  pinnae  all    close   together    and    the    whole   leaf 
sinks  downwards.     In  this  case  contact    with    another    body 
has  supplied  the  stimulus  to  which  the  living  protoplasm  has 


69 

re-acted  or  responded  in  a  particular  way,  the  final  result  of 
such  response  being  manifested  in  the  obvious  movements  of 
the  leaf  and  leaflets. 

63.  The      protoplasm      in      the  The  Cell. 

majority  of  plants  does  not  occur  in  naked  masses    but    sur- 
rounds itself  with  an  elastic,  membranous  wall,  of  a  substance 
called  cellulose,  inside  of  which    it  lives.     Such  an  enclosed 
piece  of  protoplasm  is  called  a  cell  and  its  outer  envelope  the 
cell-waU.     Many  plants  are  known,  each  of  which  consists  of 

only  a  single  cell,  but  the  body  of  the  more  highly  organised 
plants  with  which  we  are  chiefly  concerned  is  made  up  of  a 
multitude  of  such  cells  which  live  together  in  intimate  contact, 
although  every  such  plant  originally  began  life  as  a  single 
minute  cell. 

64.  If  a  thin   section    from   the  Ceil 

growing  apex  of  such  a  plant  is  examined  with  a  microscope,  it  Contents. 
will  be  found  to  consist  of  a  number  of  nearly  cubical  cells, 

each  of  them  being  provided  with  a  delicate  cell-wall  and  all  of 
them  fitting  closely  together.     Each  cell  is  full  of  protoplasm 
and  contains  in  the  centre  a  rounded   body  called  the  nucleus 
which  is  separated  by  a  definite  boundary  from  the  rest  of 
the  protoplasm.     If  the  protoplasm    is  treated  with  a  suitable 
stain,   the    nucleus   is  seen  to   consist   of   a  number   of   fine 
twisted    threads,   or    filaments,    which   become  more   deeply 
coloured  than  the  rest  of  the  protoplasm.     The  latter,  which 
fills  all  the  rest  of  the  cell  outside    the    nucleus    and    which 
usually  contains  distinct  granules,  is  distinguished  as  cytoplasm. 
As  such  a  cell  increases  in  size  the    protoplasm    is    no    longer 
able  to  completely  fill  its  cavity  and    spaces,     the    so-called 
vacuoles,  arise  in  the  protoplasm  which   usually     occupy    the 
greater  part  of  the  mature  cell,  the  protoplasm  being  confined 
to  a  layer  around  the  cell-wall,  with  perhaps   a    few   connecting 
threads  across  the  cell  cavity.      However    large   the    vacuoles 
may  be,    there    is    always    a  continuous   layer  of  cytoplasm 
surrounding  the  nucleus  and  lining  the  inside  of  the  cell- wall. 
The  vacuoles  contain  a  watery  fluid  termed  cell-sap  which  con- 
tains a  variety  of  substances  in  solution,  some  intended  as  food 
for  the  protoplasm  and  others  having    been    excreted  by  the 
protoplasm  as  waste  products.     The  cell-sap    also    frequently 
contains  a  red,  or  blue,  colouring  matter  called   antliocyanin, 
to  which  the  colour  of  many  flowers,  fruits  and  young  leaves 
is  due.    The  cytoplasm  in  such  a  cell  is  often  seen  to  be  in  active 
movement,  the  latter  being  indicated  by  the  granules  contained 
in  it,  which  are  seen  to  be  carried    along   by  the  living    pro- 


70 

toplasm  like  grains  of  sand  in  a  stream.  If  the  living  cell 
examined  is  in  a  part  of  the  plant  exposed  to  the  light,  such 
as  a  leaf,  it  will  be  found  to  contain,  in  addition  to  the.  nucleus 
and  cytoplasm,  the  so-called  chlorophyll  corpuscles.  These 
are  green  bodies,  generally  ellipsoidal  in  shape,  consisting 
of  dense  protoplasm  and  which  owe  their  colour  to  the  green 
pigment  called  chlorophyll  which  they  contain.  This  pigment 
is  soluble  in  alcohol  and  if  leaves  are  boiled  and  placed 
in  alcohol  the  green  colour  can  be  extracted  in  the 
form  of  a  solution.  These  chlorophyll  corpuscles  possess  the 
power  of  making  starch  from  water  and  the  carbon  dioxide 
of  the  air  under  the  influence  of  light.  The  small  starch  grains, 
when  formed,  are  dissolved  and  go  to  feed  the  protoplasm, 
or  to  be  accumulated  and  stored  in  certain  cells  as  reserve 
food  material  until  wanted.  In  parts  of  plants  not  exposed 
to  light,  protoplasmic  corpuscles  essentially  similar  to  the 
chlorophyll  corpuscles  are^formed  but  without  the  characteris- 
tic green  pigment  and  sometimes,  as  in  flowers  and  fruits,  the 
green  pigment  is  replaced  by  a  red,  or  yellow,  one. 

The  living  contents  of  the  cell  thus  comprise : — 

The  Nucleus 
„     Cytoplasm 
„     Chlorophyll  Corpuscles 

for  all  of  which,  collectively,  the  general  term,  protoplasm  is 
commonly  employed.  Among  the  non-living  substances  fre- 
quently found  in  living  cells  the  most  important  are  : — 

Starch  grains. — Large  grains  are  only  found  in  cells  where 
a  store  of  reserve  starch    is  being  accumulated.     Such   grains 
usually  exhibit  a  distinct  stratification,  their  substance   being- 
arranged    in  layers.      They   turn  blue  when  treated  with  a 
solution  of  iodine. 

Alcurene  or  Proteid  grains, — These  consist  chiefly  of  albu- 
minous substances  and  turn  yellow-brown  when  treated  with 
a  solution  of  iodine.  They  frequently  contain  albumen  crystals 
which  can  be  distinguished  from  crystals  of  inorganic  sub- 
stances by  the  fact  that  they  absorb  stains  and  swell  when 
treated  with  water.  They  also  usually  contain  rounded,  or  crys- 
talline, masses  of  mineral  matter.  They  are  common  in  seeds. 

Albuminous  substances  are  the  most  complex  bodies 
found  in  the  plant,  with  the  exception  of  the  living  proto- 
plasm itself,  and  they  contain  carbon,  oxygen,  hydrogen,, 
nitrogen,  sulphur,  and  phosphorus. 


71 

Other  substances  are  tannins,  fats,  ethereal  oils,  resin, 
caoutchouc,  and  mineral  crystals  the  latter  usually  consisting 
of  Calcium  Oxalate 

65.  So  far  as  is  known  at  pre-  ceil 
sent,  every  cell  originates  from  a  pre-existing  cell  and  this 
is  usually  brought  about  by  normal  cell-division.  The  whole 
cell  about  to  divide  increases  in  size,  the  filaments  contained 
in  the  nucleus  contract  and  become  converted  into  a  few 
thick  threads  of  equal  length.  The  nuclear  membrane  en- 
closing the  nucleus  then  disappears,  while  each  of  the  threads 
divides  into  two  longitudinal  halves,  the  latter  separating  and 
passing  to  opposite  ends  of  the  cell.  The  halves  collected  at 
each  end  again  coil  up  and  each  mass  forms  a  new  nucleus 
provided  with  a  membrane,  while  between  them  the  forma- 
tion of  a  cell-wall  completes  the  division  into  2  cells.  In  this 
way  an  accurate  division  of  the  nuclear  substance  contained 
in  the  cell  is  insured. 

66.  The  cell-wall  is  at  first  very  Thickening  of 

thin,  but  when  the  cell  has  attained  its  full  size  the  wall  is  *J®  j^."^*11 
more  or  less  thickened,  the  thickening  substance  being  applied  tion  of  pits, 
in  layers  to  the  original  membrane.  Such  thickened  walls 
are  therefore,  as  a  rule,  distinctly  stratified.  Parts  of  the 
original  cell- wall,  however,  usually  remain  unthickenecl.  Some- 
times very  small  areas  are  thus  left  and  narrow  channels 
through  the  thickening  layers  consequently  arise,  forming  the 
so-called  pits.  The  pit  in  the  wall  of  one  cell  is  usually 
continued  through  the  thickening  layers  of  the  wall  of  the 
adjacent  cell,  so  that  the  cells  are  connected  as  it  were  by 
a  narrow  channel  which  is,  however,  blocked  in  the  middle 
by  the  membranous  primary  wall.  A  bordered-pit  is  a  modi- 
fication of  the  simple  pit,  the  channel  of  which  is  wide  in 
the  centre  and  narrow  at  the  ends,  the  centre  of  the  original 
w^all  which  blocks  the  middle  of  the  channel  being  thickened 
and  forming  the  so-called  torus,  exactly  facing  the  narrow  en- 
trances to  the  channel.  In  this  case  the  thickening  layers  are 
built  out  into  a  dome,  instead  of  being  closely  adpressed  to 
the  primary  wall  as  is  the  case  with  a  simple  pit.  The 
entrance  to  the  bordered-pit  is  at  the  apex  of  the  dome. 

In  other  cases  when  the  greater  part  of  the  original  wall 
remains  unthickened,  the  thickening  portions  take  the  form 
of  narrow  bands  which  sometimes  anastomose  and  form  a  net- 
work. The  cell- wall  consists  chiefly  of  cellulose,  a  carbohydrate 
which  after  treatment  with  sulphuric  acid  turns  an  iodine 
solution  blue,  but  as  it  thickens  it  is  usually  more  or  less 


72 

changed  by  the  addition  of  various  chemical  substances  and 
is  often  converted  into  wood,  i.e.  is  lignified,  or  into  cork, 
i.e.  is  suberised.  The  communication  between  the  living 
protoplasm  of  adjacent  cells  is,  however,  not  cut  off  by  the 
thickening  of  the  wall,  for  exceedingly  fine  connecting  threads 
of  protoplasm  are  found  to  pass  chiefly  through  the  thin 
membranes  of  the  pits  but  sometimes  also  through  the  thick- 
ened wall. 

When  a  cell-wall  separating  two  adjacent  cells  has  become 
thickened,  its  central  portion  which  occupies  the  position  of 
Middle  the  original  primary  wall  is  called  the  middle  lamella,  and  this 
Lamella.  differs  more  or  less  markedly  from  the  rest  of  the  wall.  In 
lignified  cell- walls  it  is,  as  a  rule,  more  highly  lignified  than 
the  rest  of  the  wall,  and  if  a  piece  of  wood  is  treated  with  a 
solution,  such  as  nitric  acid  with  chlorate  of  potash,  which  is 
capable  of  dissolving  woody  substance  (lignin),  the  individual 
cells  which  compose  the  woody  mass  will  be  isolated  by  the 
dissolution  of  the  middle  lamella  and  the  structure  of  the 
wood  is  consequently  destroyed.  In  this  case  only  the  outer 
thickening  layers  of  the  walls  will  persist  which  consist  mainly 
of  cellulose  and  contain  less  lignin  than  the  middle  lamella. 
In  some  cases  the  latter  dissolves  easily  in  boiling  water  and 
some  cells  can  thus  be  dissociated  by  being  placed  in  hot 
water. 

Shape  of  67.  Young  cells  such  as  those  at 

Cells-  the  growing  points  of  plants,  are  at  first  more  or  less  cubical, 

or  rectangular,  in  shape,  but  as  they  grow  older  and  increase 
in  size,  as  they  do  just  behind  the  growing  points,  they  may 
undergo  a  great  change  of  form,  and  may  also  lose  their 
living  protoplasm,  in  which  case  they  are  often  termed  dead 
cells,  although  really  they  are  then  only  cell  cavities  bounded 
by  the  cell  walls. 


73 


CHAPTER  II.— TISSUES. 

68.  A  single  cell  which  continues  Different 

to  grow  and  divide   will  result  in  producing  an  aggregate  of  K.inds  of 
cells   all  of  which  are  coherent  from  the  time  of   their  origin;  and^heii 
such  an  aggregate  of  cells  which  are  more  or  less  similar,  which  Elements, 
erow  in  the  same  way  and  which  have  similar   duties  to   per- 
form is  called  a  tissue.     As  the  cells  of  a  tissue  grow  older  their 
walls   frequently  separate    somewhat  from  one  another    and 
spaces  arise  between  them  which  are  called  intercellular  spaces. 
The  tissues  of  plants  may  be  first  classified  as  : — 

(A)  Meristematic,  or  embryonic. — Those  consisting  of  cells 

still  capable  of  growing  and  dividing  and  thus  of 
producing  new  cells. 

(B)  Permanent. — Those  consisting  of  cells  which  have  com- 

pleted their  growth  and  have  ceased  to  divide. 

The  cells  of  permanent  tissues  vary  greatly  in  size,  shape, 
thickness  of  their  walls,  and  in  other  particulars.  They  may 
remain  more  or  less  rectangular,  or  cubical,  in  shape,  and  a 
tissue  consisting  of  such  cells  is  named  parenchyma.  The  cell 
walls  are  usually  thin  and  the  cells  usually  contain  protoplasm. 
Sometimes  the  cells  have  their  walls  thickened,  but  the  thick- 
ening substance  is  situated  mainly  at  the  corners  of  the  cells, 
in  which  case  the  tissue  is  called  collenchyma.  In  other  cases 
the  cells  become  much  elongated  in  proportion  to  their  width 
and  their  ends  are  pointed.  They  then  usually  contain  no 
protoplasm  but  only  air  and  water,  while  their  walls  are 
generally  thickened  and  provided  with  simple  pits.  Such  cells 
are  called  fibres.  Cells,  whether  parenchymatous,  or  fibrous, 
which  have  their  walls  very  much  thickened  and  in  which  the 
cell-cavity  is  consequently  very  small  and  in  some  cases  almost 
obliterated,  are  said  to  be  sclerenchymatous,  and  together  form 
the  tissue  known  as  sclerencltyma.  Cells  similar  to  fibres  but 
which  are  somewhat  shorter  and  wider,  with  blunter  ends,  are 
called  tracheids. 

By  the  absorption  of  their  transverse  partitions,  also,  rows  of 
cells  may  become  converted  into  long  tubes  which  are  termed 
vessels,  or  tracheae.  Both  vessels  and  tracheids  frequently 
have  typical  bordered  pits  on  their  walls  and  they  are  also 
distinguished  as  annular,  spiral,  or  reticulate,  when  the 
thickening  bands  on  their  walls  form  distinct  rings,  spirals,  or 
a  net  work,  respectively.  When  the  bordered  pits  of  a  vessel 


74 

are  transversely  elongated  and  are  situated  close  together,  one- 
above  the  other,  the  thickened  portion  of  the  wall  alternating 
regularly  with  the  pit  apertures,  they  look  like  the  rounds  o±  a 
ladder  and  the  vessel  is  then  described  as  scalariform. 

The  essential  difference  between  a  vessel  and  a  tracheid 
consists  in  the  fact  that  the  latter  is  a  single  cell  and  the  former 
.  is  produced  by  the  fusion  of  several  cells.  The  tr  cheid  corre- 
sponds, as  it  were,  to  a  single  joint  of  a  vessel.  Vessels,  however, 
frequently  attain  a  much  larger  diameter  than  do  tracheids 
and  they  are  often  clearly  visible  to  the  naked  eye  as  pores 
on  the  transverse  section  of  a  stem.  Neither  tracheids 
nor  vessels  contain  protoplasm,  they  are  tubes  whose  special 
duty  is  the  conveyance  of  water  from  the  roots  to  the  leaves. 
Intermediate  forms  between  these  typical  elements  often  occur. 

The  so-called  sieve-tubes  are  formed,  like  vessels,  by  the 
fusion  of  a  number  of  cells,  but  in  this  case  the  transverse 
partitions,  instead  of  being  completely  absorbed,  are  penetrated 
by  a  number  of  small  holes,  such  perforated  partitions 
being  called  sieve-plates.  The  walls  of  sieve-tubes  are  never 
lignified  and  are  usually  thin,  with  an  interior  lining  of  living 
protoplasm.  They  are  the  principal  conductors  of  organic 
food  substances  from  the  leaves  to  the  places  where  they  are 
required. 

Many  plants  contain  so-called  laticiferous -tubes  or  -vessels. 
Sometimes  these  arise  by  the  continued  growth  and  elongation 
of  a  single  cell  and  sometimes  by  the  fusion  of  several  cells,  as 
in  ordinary  vessels. 

Laticiferous  tubes  are  usually  branched,  their  walls  have  a 
thin  lining  of  protoplasm  and  are  usually  thin,  while  their  sap 
consists  of  a  milky,  usually  white,  fluid.  Such  tubes  are 
common  in  Figs,  Euphorbia,  and  many  other  plants,  and  their 
latex  often  contains  valuable  caoutchouc  and  gutta-percha. 
The  so-called  resin-canals  of  Conifers  are  really  intercellular 
spaces  into  which  resin  is  excreted  from  the  neighbouring 
cells.  The  oil  glands  in  the  leaves  of  the  Orange,  Lime  and 
other  plants  are  formed  by  the  disorganisation  of  a  group  of 
cells,  the  cell  walls  being  absorbed  and  the  cavity  so  formed 
-being  more  or  less  filled  with  ethereal  oil. 

Jyssstems.  69.  The    tissues    of    the    higher 

plants  are  usually  classified  in  three  groups,  or    systems,   as 
follows  : — 

(1)  Tegumentary  Tissue  System. 

(2)  Vascular  Bundle  Tissue  System. 

(3)  Fundamental  Tissue  System. 


75 

The  Tegumentary  System  comprises  the  epidermis,  or  entire  Tegu 

i  •  -c    J.-L          i       j.          v   i,  n  -_t         £  -i     System. 

outer  skin   01   the  plant,   which  usually  consists  ot   a  single  " 

continuous  layer  of  cells.     These  fit  close  together  and,  in  the 

serial  parts  of  plants  at  all   events,  have   their   external  walls 

more  or  less  thickened  and  cuticularised,  i.e.    converted  into 

cutin,  a  substance  resembling  cork  but  which  is  more  resistant 

to  the  action  of    caustic  potash  and  strong  sulphuric   acid. 

When  such  cuticularised  external  walls  are   very  thick  their 

outermost  layers  are  more    cuticularised   than   the  rest   and 

form   the   so-called  cuticle,  which  can  often  be  detached  as  a 

separate   membrane   from  the  other   thickening  layers.     The 

epidermis  is  also  frequently  provided  with  an  additional  external 

covering  in  the  shape  of  deposits  of  wax,  or  various  forms  of 

hairs,  the  latter  being  outgrowths  of  the  epidermal  cells.     The 

cells  of  the  epidermis  do  not  usually  contain  chlorophyll.     It  vascular 

was  pointed  out,   in  Part  I   above,   that  the    firm  nerves,  to  Bundle 

which  the  venation   of  the  leaves    is  due,  pass  into  the  stem,   ys 

where  they  join  on  to  stouter  similar  strands  which  are   again 

continued  into  the   root,  and    also    that   these  strands    were 

called  vascular  bundles.     In  some  Balsams,  the  stems  of  which 

are  more    or  less  transparent,  these  strands  passing  down  the 

stem  can  be  seen  with  the  naked  eye  and  in  other  cases  they 

may  often  be  isolated  from  young  stems  which   have    been 

allowed  to  rot  in  water  by  carefully  washing  away  the  softer 

tissue.    Vascular    bundle    skeletons  of    leaves,  also,  are  often 

naturally  produced  as  the  leaves  slowly  decay.     These  vascular 

bundles  together  constitute  the  Vascular  Bundle  Tissue  System. 

A.11  tissue  which  does  not  belong  either  to  the  Tegumentary  Fundamental 

or   Vascular  Bundle  System  is  classed  as  Fundamental,   and  system. 

consists  usually  of  parenchyma,  but  also  often  contains  collen- 

chyma  and  sclerenchyma. 

70.  A  vascular  bundle  consists  structure  o£ 

of  two  portions,  the  Xylem,  or  Wood  portion  and  the  Phloem,  Bundles* 
or  Bast  portion.  The  former  consists  chiefly  of  vessels  and 
tracheids,  or  sometimes  of  tracheids  alone,  with  some  paren- 
chyma, while  the  phloem  portion  contains  sieve-tubes  and 
parenchyma.  Fibres  may  also  be  present  in  bothxylem  and 
phloem.  Most  commonly  the  xylem  and  phloem  are  in  contact 
on  one  side  only  and  the  bundle  is  said  to  be  collateral.  In 
the  stem  the  xylem  and  phloem  portions  of  a  collateral  bundle 
are  found  on  the  same  radius  of  the  stem,  the  xylem  being 
nearest  the  centre  of  the  stem  and  the  .phloem  nearest  the 
circumference.  In  some  cases  the  xylem  has  phloem  not  only 
on  the  outside,  but  also  on  the  inside,  the  xylem  being  placed 


76 

between  two  groups  of  phloem  and  in  this  case  the  bundle 
is  bi-collateral.  In  cases  where  either  the  xylem  or  the 
phloem  is  entirely  surrounded  by  the  other,  the  bundles 
are  called  concentric.  In  some  cases  all  the  cells  of  a  vascular 
bundle  develop  into  either  xylem  or  phloem  elements,  but 
in  others  certain  cells  remain  meristematic,  i.e.  retain  their 
power  of  growth  and  division.  In  the  former  case  the  bundle 
is  said  to  be  closed  and  in  the  latter  open,  the  meristematic  tissue 
being  called  the  cambium.  On  the  outside  of  the  vascular  bundle 
sclerenchymatous  fibres  are  usually  found,  which  often  form  a 
more  or  less  complete  sheath  to  the  bundle, 

Devei    m  ^'  ^  vascular  bundle  does  not  of 

of  Vascuia?n  course  suddenly  arise  in  the  plant  tissues  with  all  its  parts  corn- 
Bundles,  plete,  but  is  developed  gradually,  like  all  other  parts  of  the 
plant,  from  cells  which  at  first  are  homogeneous.  In  the  very 
young  plant  embryo  in  the  seed  the  vascular  bundles  are  not 
yet  recognisable.  As  the  embryo  develops,  however,  certain  rows 
of  cells  grow  longer  than  their  neighbours  and,  dividing  tangen- 
tially,  give  rise  to  strands  of  narrow  elongated  cells,  which  are 
destined  to  become  the  vascular  bundles.  The  form  of  these 
cells  gradually  changes  as  they  complete  their  growth,  some 
elongate  to  form  fibres  and  tracheids,  the  width  of  others 
increases  to  form  vessels,  their  walls  become  variously  thickened 
and  pitted  and  their  chemical  composition  is  altered,  many 
cells  lose  their  protoplasmic  contents,  while  in  other  cases  the 
cross  walls  are  more  or  less  absorbed  to  form  vessels  and  sieve- 
tubes.  This  differentiation  of  a  collateral  vascular  bundle 
proceeds  from  the  exterior  of  the  bundle  on  both  sides  inwards 
and  the  first  formed  elements  of  the  xylem  and  phloem,  which 
are  thus  in  this  case  the  outermost,  are  called  protoxylem  and 
protophloem,  respectively.  In  the  protoxylem  annular  or  spiral 
vessels  or  tracheids  are  found  which  do  not  occur  in  fche  rest 
of  the  xylem. 

The  cells  at  the  growing  apices  of  the  shoots  and  roots  of 
older  plants  are  also  practically  homogeneous  and  meriste- 
matic, just  as  they  are  in  the  plant  embryo,  and  it  is  only 
at  some  distance  behind  the  so-called  growing  points  that  we 
find  the  vascular  bundles  fully  developed. 


77 


CHAPTER  III.  -STRUCTURE   AND    DEVELOPMENT   OF 
PLANT    MEMBERS. 

72.  If    the    young    stem    of    a  s,tem  of 

Gymnosperm,  or   Dicotyledon,    is*  cut  across  after  the  colla-  pfrmg03" 
teral  vascular  bundles  have  become  differentiated,  the  latter  and  Dicoty-- 
will  be  found  to  be  in  a  circle  around  the  stem,  the  xylem  of  Ied°ns. 
each  bundle  being  nearest  the  centre     of  the  stem  and  the 
phloem  nearest  the  circumference.     The  fundamental    tissue 
occupying  the  centre  of  the  stem,  inside  the  circle  of  vascular 
bundles,  forms  the  pith,  while  that  between  the  bundles  con-^  "  ary 
stitutes    the    primary    medullary   rays.     In    perennial    stems, 
and  sometimes  also  in  annual  stems,  after  the  differentiation 
of    the  primary  vascular  bundles,  a  further  process  sets  in 
termed  secondary  growth  in  thickness.     This  is  accomplished 
by  the  activity  of  the  cambium  of  the  vascular  bundles  which 
is  a  layer  of  elongated,  delicate-walled  cells,  full  of  protoplasm, 
situated  between  the  xylem  and   phloem,  and  by  the  forma- 
tion and  growth  of  a    similar    thin  layer  of  cambium  in  the 
primary  medullary  rays  between  the   bundles,  which,   uniting 
with  the    cambium  of  the  bundles,   forms  a  complete    ring 
around    the    stem.      From    this     cambium     new    cells    are 
continually    being    formed,    through    the      division    of    the 

tt 

cambium  cells  by  tangential  walls,  both  on  the  inside  and 
outside  of  the  ring,  towards  the  centre  and  circumference 
of  the  stem  respectively ;  the  former  develop  into  xylem  and 
the  latter  into  phloem  elements.  The  primary  medullary  rays 
are  thus  broken  up  by  the  formation  of  new  xylem  and  phloem 
elements  from  the  cambium  between  the  bundles.  These 
elements,  however,  follow  a  more  or  less  sinuous  course  longi- 
tudinally leaving  a  net-work  of  narrow  elongated  openings 
which  are  filled  by  rows  of  cells  of  fundamental  tissue,  chiefly 
parenchyma,  running  horizontally  from  the  centre  of  the  stem 
towards  the  circumference.  These  cells  constitute  the  second- 
ary medullary  rays  and  as  the  cambium  cells  opposite  these 
rays  continually  form  new  medullary  ray  cells  opposite  thorn, 
both  on  the  inside  and  outside  of  the  cambium  ring,  a  medul- 
lary ray,  once  started,  is  kept  open  as  the  stem  increases  in 
thickness  and  extends  both  through  xylem  and  phloem  elements. 
As  the  circumference  of  the  stem  gradually  enlarges,  now 
medullary  rays  originate  from  the  cambium  between  those  al- 
readv  formed.  By  mean0  of  the  medullary  ravs  communication 

tf  w  •  ,       • 


Annual 
Rings. 


Heart- 
Wood 

Wood 
Elements 


Bast 
Elements. 


Stem  of 
Monocoty- 
ledons. 


is  insured  between  all  the  lining  cells  of  the  stem.  The  rays 
first  formed  extend  from  the  pith  to  the  circumference,  while 
those  arising  later  originate  at  some  distance  from  the  pith. 

In  plants  the  growth  of  which  exhibits  periods  of  activity 
alternating  with  periods  of  rest,  there  is  usually  a  more  or  less 
marked  difference  between  the  wood  formed  at  the  beginning 
and  end  of  the  period  of  activity  respectively.  In  the  former 
the  vessels  are  often  larger,  or  more  numerous,  or,  in  the 
absence  of  vessels,  the  other  elements  are  usually  wider  and 
with  thinner  walls,  and  the  wood  of  which  they  constitute  a  part 
contrasts  strongly  with  that  which  it  adjoins  formed  at  the  end 
of  the  previous  season's  period  of  activity,  the  elements  of  which 
are  denser.  More  or  less  obvious  concentric  rings  thus  arise  in 
the  wood,  which  are  the  so-called  annual  rings.  What  is  known 
as  heart-wood  consists  of  dead  cells,  the  cavities  of  which  are,  as  a 
rule,  blocked  up  by  gums,  or  other  substances,  and  which,  being 
more  or  less  saturated  with  tannins,  is  usually  of  a  dark  colour. 

The  wood  of  Conifers  consists  almost  entirely  of  tracheids 
which  have  bordered  pits  chiefly  on  their  radial  walls,  with  some 
parenchyma,  and  also  sometimes  with  scattered  resin  ducts. 

The  wood  of  Dicotyledons  consists  of  vessels,  tracheids, 
parenchyma,  and  fibres. 

The  phloem,  or  bast,  of  both  Gymnosperms  and  Dicoty- 
ledons consists  of  sieve-tubes,  parenchyma,  and  long,  thick- 
walled,  fibres  ;  the  latter,  unlike  the  fibres  of  the  wood,  usually 
have  their  walls  very  slightly,  or  not  at  all,  lignified,  and  they 
are  therefore  tough  and  flexible.  It  is  owing  to  this  fact  that 
the  bast  of  many  species  is  of  commercial  value  for  the  manu- 
facture of  ropes  and  cordage. 

The  above  elements  occur  in  varying  proportions,  the  wood 
or  bast  of  a  particular  species  having  more  of  a  certain  element 
than  that  of  another  species  while  in  some  species  certain 
elements  may  not  be  represented. 

73.  In  the  young  stem  of  a  Mono- 
cotyledon the  primary  collateral  vascular  bundles,  instead  of 
being  arranged  in  a  circle,  are  often  irregularly  scattered 
throughout  the  fundamental  tissue  of  the  stem.  The  xylem 
of  each  bundle  is  turned  towards  the  centre  of  the  stem  as 
before,  but  the  bundles  are  all  closed,  i.e.  no  meristematic 
tissue  remains  in  them  to  produce  secondary  growth  in 
thickness.  In  this  case  no  distinction  can  be  made  between 
the  pith  and  primary  medullary  rays.  In  a  few  Mono- 
cotyledons, e.g.  some  Palms,  secondary  growth  in  thickness 
does  occur,  a  cambium  ring  arising  in  the  fundamental  tissue 
outside  the  vascular  bundles,  but  this,  instead  of  forming 


79 

'continuous  wood  on  the  inside  of  the  ring,  produces  only  new 
areas  of  fundamental  tissue  with  isolated  vascular  bundles. 

74.  In  the  young  roots  of  Mono-  Roats. 

cotyledons,  Dicotyledons,  and  Gymnosperms  the  arrangement 
of  the  primary  vascular  bundles  is  different  from  that  found  in 
the  stem.     The   xylem   and  phloem   strands   of  each  bundle 
separate  from  each  other  as   they   pass  into  the  root  and  are 
there  arranged  side  by   side   on   different  radii  of  the  root  and 
the  bundles   are  then  said    to    be   radial.     The  xylem  strands 
also  become   twisted  on  themselves   so   that  the   protoxylem, 
instead    of    being    internal,    as    in    the    stem,    is    external. 
Sometimes  the  xylem  strands   meet  in  the  centre   of  the   root 
and   sometimes   they  do   not,    there     then   being    a   central 
pith.     These  separate  strands  of  xylem  and  phloem  follow    a 
straight  longitudinal  course  in   the  root   instead  of  curving  as 
do  the  bundles  of  the  stem.     In  the  roots  of  Dicotyledons  and 
Gymnosperms   secondary  growth    in   thickness   occurs    simul- 
taneously with  the  similar  growth  in  the  stem.      In  the  roots 
cambium  layers  first  appear  on  the  inside  of  the  phloem  strands 
and,  extending  thence  laterally  on  both  sides,  they  eventually 
coalesce  opposite  the  xylem  strands,  and  thus  form  a  complete 
-cambium  ring.     This   jing  gives  rise  to  xylem  elements  on  the 
inside  and  phloem  elements  on  the    outside,    as    in  the   stem, 
broad  medullary  rays  being  usually  formed  opposite  the  primary 
-xylem  strands.     The  wood  of  an  old  root  is    usually  more 
porous  than  that  of  the  stem  but  otherwise  they  very  closely 
Tesemble  each.~oth.er  in  the  possession  of  annual  rings  and  other 
characters.     The  cambium  of  the  root  is  a  continuation   of 
that  of  the  stem,  there  thus  being  an  uninterrupted    cambial 
layer  throughout  the  stem  and  root  and  their  branches. 

In  those  Monocotj'ledons  which  exhibit  secondary  growth 
a,  cambial  layer  may  arise  in  the  root  outside  the  primary 
hurdles  but  it  only  produces  closed  vascular  bundle  strands 
scattered  in  fundamental  tissue,  as  in  the  stem. 

75.  In    addition  to    the    typical  Abnormal 

mode  of  secondary  growth  above  described,  cases  of  abnormal  Deveiop- 
•development   are   not   uncommon     among   Dicotyledons   andmentt 
Gymncsperms.    In  Cocculus  laurifolius,  Bauhinia  Vahlii,  Cycas, 
and  several  other  plants,  the  cambium  ring  first  formed  ceases 
to  grow  alter  a  time  and  then  a  second  cambium  ring  arises 
-outside  the  bast  formed  by  the  first.     This  also  ceases  to  grow 
after  a  limited  time  and  a   third  ring  arises,   and  so  on,   the 
stem  in  consequence   exhibiting  very  characteristic,  more  or 
less  concentric,  bands  of  wood  and  soft  bark-like  tissue,  which 
have  been  already  mentioned  on  page  18. 


80 

In  other  cases  the  cambium  forms  much  more  wood  or  bast 
at  certain  points  than  at  others,  and,   on  a  cross  section.,  radial 
masses  of  wood  and  bast  are  formed  alternating  with  each  other, 
the  cambium    forming  an  undulating  layer  instead  of  a  circle. 
Periderm  76.  In    those    Dicotyledons    and 

Gymnosperms   in    which  the  interior    of    the    stem   is    con- 
tinually increasing    in  thickness    through    the  activity  of  the 
cambium  ring,  the   fundamental  tissue   and  epidermis,  which 
at    first    surrounded    the  primary  vascular   bundles,   is   sub- 
jected to    pressure  and  tension.     In  some  cases  the  cells  com- 
posing  these  tissues   continue    to  grow  and   divide  and   thus 
keeping   pace   with   the    internal    development   are    able    to* 
persist.     This  occurs  in  most  annual   stems  and   in  perennial 
stems  during  the  first  year   of  their  development   and   occa- 
sionally for  a  longer  period.      In  the   majority   of  perennial 
stems,  to  wards  the  close  of  the  first  year's  growth,  the  epidermis 
being  no  longer  able  to  withstand  the  strain  becomes  ruptured 
and   falls  off.     Before  this  happens,  however,  a   layer    of  cells 
has  been  produced  close  below  the  epidermis  which  acts  like  a 
secondary  cambium  and  is  called  the  Phellogen  9  or  Cork  Cambium. 
By  the  active  growth  and   division  of  this  layer  new  cells    are 
added  both  towards  the  exterior  and  interior  of  the  stem.    The 
former  lose  their  protoplasm  and  become  converted  into  cork, 
forming  the  so-called  Phellem.,  while  the    latter    retain    their 
protoplasm  and  usually  contain    chlorophyll,    constituting  the 
Phelloderm.    The  term  Fender m  includes  the  Phellem,  Phellogen 
and  Phelloderm.     The  cork  cells  of  the  phellem  form  an  elastic 
protective  covering  to  the  stem  which  is  almost    impermeable 
to  air  and  water  and  which  effectually   takes  the  place  of  the 
ruptured  epidermis.     It  is  owing  to  the  formation  of  this  layer 
of  cork  that  the  colour  of  young  twigs  usually  changes    from 
green  to  some  shade  of  brown,  the  chlorophyll  in  the  cells    of 
the  fundamental  tissue  beneath  it  being  no  longer  visible    as 
was  the  case  when  these  cells  were  covered  by  the  thin  epider- 
mis.    In  some  cases  the  periderm,  instead  of  originating   close 
below  the  epidermis,  is  formed    deeper   down   in    the    funda- 
mental tissue,  and  sometimes  close  to  the  vascular  bundles.     In 
such  cases,  a?  all  the  living  cells  outside  it  are  prevented  from 
obtaining  the  supply  of  water    and    food   necessary    for   their 
existence,  on  account  of  the  impermeable  cork  layer,  they  diy 
np    and   die,    forming  the   rough   outer  bark  which  is  sooner 
or  later  exfoliated.     In  some  trees,   e.g.    the   Birch,    the    first 
formed,  or  primary,  phellogen  remains  active  for  several  years 
and  in  some  species  it  persists  through  the   entire    life   of   the 
tree,  new  cork  cells  being  added  below  as  the  outer  ones  are- 


81 

exfoliated.  In  this  way  a  thick  corky  bark  may  be  formed. 
As  a  rule,  however,  its  activity  ceases  after  a  short  time  and 
another,  or  secondary,  phellogen  arises  deeper  in  the  stem 
beneath  the  first  and  nearer  the  actively  growing  cambium 
layer.  In  this  case  again  all  the  living  cells  outside  the  second- 
ary phellogen  die  and  are  added  to  the  dead  outer  bark  just 
as  happens  when  the  primary  phellogen  develops  deep  in  the 
fundamental  tissue.  In  this  way  a  number  of  successive  peri- 
derms  may  be  formed,  succeeding  each  other  at  short  intervals. 
It  will  thus  be  seen  that  the  outer  dead  bark  may  consist  of 
cork  cells  only  or.  in  addition,  of  dead  elements  which  originally 
formed  a  part  of  the  primary  fundamental  tissue,  or  of  successive 
periderms,  or  even  of  the  phloem  developed  from  the  cambium. 
Taking  also  the  word  cortex,  or  bark,  in  the  usual  sense  as- 
including  all  the  tissues  outside  the  cambium  ring,  it  will  be 
seen  that  this  consists  of  an  outer  portion  of  dead  elements  and 
an  inner  part  which  is  largely  composed  of  living  cells  containing 
protoplasm.  According  as  the  periderm  forms  a  regular  circle 
around  the  stem,  or  only  arcs  of  a  circle,  so  do  the  outer  layers 
of  dead  bark  exfoliate  as  rings  or  scales.* 

Dead  bark  which  does  not  exfoliate  rapidly  becomes  deeply 
cracked  and  fissured. 

In  the  roots  of  Dicotyledons  and  Gymnosperms  which 
exhibit  secondary  growth  the  primary  phellogen  usually  origi- 
nates close  to  the  primary  phloem,  all  the  tissue  external  to  it 
then  peeling  off,  while  subsequent  phellogens  arise  as  in  the  stem. 
In  the  stems  and  roots  of  those  Monocotyledons  which  possess 
a  cambium  ring  and  which  therefore  increase  in  thickness^ 
periderm  is  also  formed  outside  the  cambium  ring  in  the  way 
just  described  only  in  this  case  the  cambium,  instead  oi 
producing  bast  continuously  on  the  outside,  gives  rise  to  new 
fundamental  tissue. 

77.  At   certain    points    in    both 
roots  and   stems  the  phellogen,  instead  of  producing  cork  cells 

*  In  this  book  the  word  '•'  bark  "  is  used  in  its  ordinary  sense  and  denotes  all 
tissues  situated  outside  the  cambium.  In  some  botanical  books  it  is  applied  in  a  res- 
tricted sense  to  the  dead  tissues  situated  outside  the  phellogen.  For  the  latter,  however, 
the  word  rhytidomc  seems  preferable. 

The  word  rJiytidome  appears  to  have  been  first  employed  in  Indian  Botanical 
literature  by  W.  R.  Fisher  in  his  Morphological  Botany  (Roorkee  1888),  but  it  has 
not  as  yet  been  generally  adopted  by  botanists.  In  that  book  Mr,  Fisher  has 
spelt  the  word  in  two  ways  rfiytiderm  (page  76)  and  rJiytidome  (page  12C).  In  his 

Tree*,  8f,rub?,  and  Wcody  Climbers  of  the  Bombay  Presidency,  2nd  edition,  page  x, 
Mr.  W.  A.  Talbot  has  adopted  the  word  rJiytidome.  The  word  is  believed  to  be 
derived  from  Greek  pvn$  =  a  wrinkle)  and  Zepua.  ( =  skin,  hide)  and  thus  signi- 
fies wrinkled,  or  shrivelled,  skin.  pvn$  has  already  given  rise  to  words  which  are 
generally  accepted  in  botanical  literature,  such  as  rytidocarpus  (=  wrinkled  fruit), 

so  that  a  better  spelling  of  this  useful  term  would  appear  to  be  rytiderm. 

a 


82 


Shedding  of 
Leaves. 


Develop- 
ment of 
Secondary 
Members. 


Develop- 
ment  of 
the  Root. 
Root- cap. 


which  fit  close  together  without  intercellular  spaces,  gives  rise 
externally  to  a  number  of  loosely  united  cells  which  often 
protrude  considerably  beyond  the  surface.  These  areas  of 
loose  cork  cells  are  the  lenticels,  and  they  allow  the  necessary 
interchange  of  gases  to  take  place  between  the  intercellular 
spaces  in  the  interior  of  the  plant  and  the  outer  air. 

78.  When    leaves    are  about    to 

be  shed   a    layer  of    cork,  continuous    with    the  periderm  of 
the  stem,    is  formed   across  the  base    of  the  petiole,  which  is, 
however,  penetrated   by  the   vascular  strands   which  descend 
from   the,  leaf.     Shortly   after  its  formation  a  layer  of  cells, 
a   little   distance  above  it  in  the  petiole,  becomes  absorbed, 
resulting  in   the   entire    separation   of  the  tissues  of  the  leaf 
from  those  of  the  stem  and  the  leaf  accordingly  falls  off.     The 

<_>   •/ 

cork  layer  below  the  surface  of  the  scar  now  protects  the 
inner  tissues  of  the  stem  from  injury  and  this  protection  is 
completed  by  the  blocking  up  of  the  cavities  of  the  vessels 
and  sieve-tubes  with  gums  or  cork. 

79.  Young  leaves   and   branches 

first    appear   on  the  stem  as  minute   humps    caused  by  the 
growth  and  division  of  a  group  of  cells  situated  close  below  the 
epidermis.     As  these  cells  grow  the  epidermis  increases  in  area 
accordingly  but  not  in  thickness,  and  it  thus  remains  as  a  con- 
tinuous layer  over  the  leaves  and  young  branches  and  stems. 
Young  roots,  on  the  other  hand,  originate  deep  in  the  funda- 
mental tissue  of  the   parent    root,    just  outside  the    vascular 
bundles,    and  as  they   develop  they  force  their  way    through 
and   rupture   the  tissue  lying  between  them  and  the  exterior 
of  the  root,  so  that  they  emerge  through  slits  in  the  tissue  of 
the  parent.     Each  young  root  also,  as  a  rule,   originates  just 
outside  a  xylem  strand  of  the  parent,  and  they  are  therefore 
found  in  straight  longitudinal  rows,    there  being    usually    as 
many  rows  as  there  are    xylem  strands    in    the   parent  root. 
These  lateral   roots  also  always  first  appear  at  some  distance 
behind   the    growing    apex    of   the    parent   root,   where   the 
tissues  have  already  become  differentiated 

80.  The  growing  point  of  a  root, 

unlike   that   of  a   stem,   is  protected    by   a    mass   of    tissue 
called  the  root-cap    formed    by    the  growth   and    division    of 
the   external   layer  of  cells  at  the    apex.     The   outer  cells    of 
this  cap    are    continually   exfoliated  while  new  cells  are  con- 
tinually added  at  the  base.    The  actually  elongating  portion 
of  a  growing  root   comprises  a  relatively  small  area,  situated 
immediately  behind  the  apex,  and  as  this  elongates  it  pushes 
the  root  apex,  protected  by  its   slippery  conical  cap,  in  front  of 
it   into  the   earth   like   a    shield.     If   the   roots    of    a  young 


83 

plant,  e.g.  a  wheat  plant  such  as  is  shown  in  Plate  7,  Fig. 
3,  are  carefully  removed  from  the  soil  and  examined,  their 
lower  portions  will  be  found  projecting  from  a  mass  of 
soil  particles  like  naked  white  threads.  Closer  inspection  shows 
that  these  soil  particles  are  clinging  to  a  number  of  minute 
hairs  which  look  like  very  fine  shining  lines.  These  are  the 
so-called  root-hairs  which  attach  themselves  so  firmly  to  the  Root-hairs.: 
soil  particles  that  the  latter  can  only  with  difficulty,  if  at  all, 
be  separated  from  them.  When  looked  at  with  a  microscope, 
each  of  these  hairs  is  seen  to  be  a  tubular  outgrowth  of  a  single 
epidermal  cell,  into  which  it  opens  at  the  base,  with  a  thin  cell 
wall  of  cellulose  and  containing  living  protoplasm.  It  is  prin- 
cipally by  means  of  these  hairs  that  roots  absorb  their 
necessary  supplies  of  water  and  substances  in  solution  from 
the  soil,  the  older  parts  of  the  roots  taking  no  part  in  this  work 
of  absorption.  If  these  delicate  hairs  were  formed  on  the 
portion  of  the  root  which  is  rapidly  elongating  they  would  be 
rubbed  off  as  the  root  pushes  downwards  into  the  soil,  and 
hence  the  elongating  region  is  destitute  of  these  hairs.  The 
root -hairs  as  a  rule  only  live  for  a  few  days  and  then  die  off,  so 
that  they  are  absent  from  the  older  parts  of  the  root,  but  as 
new  hairs  are  continually  being  formed  behind  the  elongating 
region  a  zone  of  active  living  hairs  is  constantly  maintained 
there.  Owing  to  the  strong  adhesion  of  the  root-hairs  to  the 
soil  particles,  the  root  is  firmly  held  in  the  soil  and  the  elonga- 
tion of  the  area  immediately  behind  the  apex  must  drive  the 
tip  of  the  root  down  into  the  soil,  as  it  cannot  force  up  the 
firmly  anchored  part  of  the  root  behind  it.  To  enable  the  root 
to  obtain  as  much  water  and  mineral  salts  as  possible  from 
the  soil,  it  is  obviously  an  advantage  for  it  to  have  a  large  ab- 
sorbing surface  which  shall  come  in  contact  with  as  many  soil 
particles  as  possible.  Accordingly,  the  tip  of  the  root  as  it  glows 
downwards,  usually  follows  an  irregular  spiral  course,  and  the 
multitudes  of  root-hairs  give  it  a  very  large  absorbing  surface. 
The  root-hairs  having  obtained  all  the  supplies  they  can  from 
the  soil  within  their  reach  die  off,  while  the  lateral  roots  then 
tap  those  layers  of  soil  which  were  beyond  the  reach  of  the 
hairs  of  their  parent  root. 

The  upper  older  portion   of  the  young  roots   of  seedlings  Shortening 
frequently   exhibits    the    peculiar  property  of  contracting,  or of  Roots- 
shortening.     A  pull   is  thus  exercised   on  both    the   root  and 
stem,  and  as  the  former  is  strongly  anchored  the  lower  portion 
of  the  stem  is  pulled  down  into  the  soil  and  the  young  plant 
is  thus  firmly  established.     The  rooting  tips  of  the  branches 
of   Rubus  lasiocarpus  may  thus  be  seen  to  be  drawn  into  the 
ground  and  similar  contraction  i«  often  seen  in  the  older  roots 

Q  tf 


84 

of  mature  plants,  especially  of  those  plants  which  have  rosettes- 
of  leaves  close  to  the  ground.  In  such  plants  the  short  stem 
increases  slightly  in  length  each  year  and  the  new  leaves  arise 
higher  and  higher  on  the  stem  but,  notwithstanding  this,  the 
new  leaves  are  always  situated  close  to  the  ground.  Trans- 
verse wrinkles,  or  folds,  at  the  base  of  the  root,  where  the 
tissues  have  shrunk,  are  often  clearly  visible  where  such 
contraction  has  taken  place. 

Structure  of  gl.  The  leaf  is  composed  of  funda- 

mental tissue  traversed  by  vascular  bundles  and  covered  by 
an  epidermis,  all  of  which  are  continuous  with  the  corresponding 
tissues  of  the  stem.  The  xylem  of  each  bundle  is  nearest  the 
upper  surface  of  the  leaf  and  the  phloem  nearest  to  the  lower 
surface.  The  fundamental  tissue  of  the  leaf  may  be  uniform, 
but  is  usually  differentiated  into,  the  so-called  palisade  tissue, 
situated  under  the  upper  epidermis,  and  the  spongy  parenchyma, 
which  extends  from  the  palisade  tissue  to  the  lower  epidermis. 
The  palisade  tissue  consists  of  cylindrical  cells,  rich  in  chloro- 
phyll, situated  close  together,  with  their  long  axes  perpendicular 
to  the  leaf  surface.  The  spongy  parenchyma  consists  of  irre- 
gularly shaped,  often  stellate,  cells,  containing  less  chlorophyll, 
and  between  which  there  are  large  intercellular  spaces.  In  the 
leaf,  as  in  other  parts  of  the  aerial  shoot  of  the  plant,  the  cells 
of  the  epidermis  fit  close  together  except  at  the  spots  where  the 
openings,  termed  stomata,  occur.  These  stomata  are  particularly 
numerous  on  the  leaves  and  in  the  leaves  of  land  plants  they 
are  found  almost  exclusively  on  the  undersurface.  A  stoma 
is  formed  by  the  division  of  an  epidermal  cell  into  two 
daughter  cells,  which  then  separate  from  one  another  by  the 
splitting  of  the  wall  between  them,  an  opening  into  the  tissues 
of  the  leaf  thus  being  formed.  These  two  cells  contain 
chlorophyll,  are  sausage-shaped,  and  are  called  the  guard-cells. 
Immediately  below  the  guard-cells  is  a  large  intercellular 
space  which  is  in  communication  with  the  other  intercellular 
spaces  of  the  leaf,  so  that  the  internal  tissues  of  the  leaf  are 
placed  in  communication  with  the  air  which  can  thus  obtain 
access  to  the  cells,  while  the  escape  of  gases  and  water  vapour 
from  the  leaf  is  also  provided  for.  According  to  the  quantity 
of  water  in  the  guard-cells  their  free  walls  which  adjoin  the 
stomatal  opening,  approach,  or  recede  from,  each  other,  thus 
closing,  or  opening,  the  stoma.  When  the  guard-cells  are 
turgid  and  full  of  water  their  free  walls  are  drawn  apart  and  the 
stoma  is  opened,  while  if  the  guard-cells  lose  water  and  their 
walls  are  no  longer  tightly  stretched,  as  they  are  in  turgescence. 
their  free  walls  are  pushed  together  and  close  the  stoma. 


PART  IIL-PHYSIOLOGY. 

fp4 
CHAPTER  I.— FUNCTIONS  OF  PLANTS  IN  GENERAL. 

82.  It  has  been  noted  in  Part  II  General 

•that  the  essential  living  substance  of  plants  consists    of  pro-  Conditions ' 
toplasm  and  that  this  protoplasm  is  capable  of  doing   various01  >lantLlfe- 
kinds  of  work,  of  performing  various  functions,  such  as  those  of 
nutrition,  respiration,  growth,  movement  and  reproduction.  See 
page  68. 

Protoplasm  can,  however,  only  perform  these  functions 
provided  that  the  external  conditions  are  favourable.  If 
no  water  is  available  protoplasm  at  once  becomes  inactive 
and  may  die,  while  it  is  easily  destroyed  by  too  high  or  too 
low  a  temperature. 

The  protoplasm  contained  in  the  cell  from  the  growth 
and  development  of  which  a  Teak  tree  is  ultimately  produced 
must  clearly  have  a  very  different  structure  from  that  contained 
in  the  cell  which  eventually  develops  into  a  Sal  tree :  with  the 
bost  microscopes  at  present  available,  however,  no  such 
difference  can  be  detected.  Still  we  must  not  on  that  account 
expect  the  protoplasm  of  different  plants  to  always  behave 
in  exactly  the  same  way  when  the  external  conditions  are 
the  same,  and,  as  a  matter  of  fact,  we  know  that  different 
plants  have  different  needs  and  that  the  effect  of  one  and 
the  same  factor  on  different  plants  may  be  very  various.  The 
amount  of  light,  for  instance,  which  is  absolutely  essential 
for  the  healthy  development  of  some  plants  (ligkt-demanders) 
is  often  injurious  to  those  plants  which  live  in  shady  places 
(shade  bearers). 

In  all  ordinary  green  plants,  however,  a  suitable  supply 
of  water  and  mineral  salts,  a  suitable  amount  of  light  and  heat, 
as  well  as  of  free  oxygen  and  carbon  dioxide,  are  essential 
for  their  existence  and  for  the  vital  activity  of  their  protoplasm. 

In  Physiology,  then,  we  must  consider,  on  the  one  hand, 
the  living,  irritable,  protoplasm,  and  on  the  other  hand,  the 
non-living  factors  of  the  plant's  environment  which  are  capable 
of  acting  as  stimuli  to  the  protoplasm,  of  inducing  it  to  behave 
in  a  certain  way  and  to  perform  definite  work,  while,  under 
certain  circumstances,  they  may  be  able  to  altogether  prevent 
its  activity  and  even  to  kill  it. 


86 

For  every  such  external  factor  there  is  in  fact  for  each  in- 
dividual plant  a  minimum.,  maximum,  and  optimum,  degree  of 
intensity  with  reierence  to  its  effect  on  any  particular  function, 
the  first  two  being  the  extremes  beyond  which  the  function 
ceases  to  be  performed  and  the  last  being  that  degree  of  inten- 
sity under  the  effect  of  which  the  protoplasm  is  best  able  to 
exercise  the  function  in  question.  Green  plants,  for  instance, 
cannot  manufacture  their  organic  carbonaceous  food  from  inor- 
ganic materials  without  light,  while  if  the  light  is  too  intense 
the  green  chlorophyll,  by  means  of  which  the  manufacture  is 
made  possible,  is  destroyed,  so  that  the  manufacture  of  food 
is  only  possible  provided  that  the  intensity  of  the  light  is 
suitable.  Again  if  the  temperature  of  the  soil  falls  below 
a  certain  minimum,  or  rises  above  a  certain  maximum,  the 
protoplasm  in  the  root-hairs  becomes  inactive  and  the  roots 
can  no  longer  provide  the  plant  with  the  necessary  water 
and  salts  from  the  soil. 

83.  The    entire    body    of    some 

minute  green  plants  which  live  in  water  and  are  called  alga3 
consists  of  a  single  cell,  such  as  has  been  described  in  Part  II, 
in  which  more  or  less  of  the  cavity  of  the  cell  is  occupied  by 
the  so-called  vacuolea,  filled  with  cell-sap  and  surrounded  by 
a  layer  of  protoplasm  which  lines  the  thin  outer  wall  of 
cellulose  and  in  which  are  imbedded  the  nucleus  and  chlorophyll 
corpuscles.  Such  a  minute  plant  is  just  as  capable  of  exercis- 
ing the  functions  of  nutrition,  growth,  respiration,  and  repro- 
duction, as  are  plants  the  bodies  of  which  consist  of  enormous 
numbers  of  such  cells  and  exhibit  an  elaborate  differentiation 
into  ^rgans.  Such  a  cell,  for  instance,  is  able  to  absorb 
fro  n  the  water  in  which  it  lives  water  with  traces  of  mineral 
salts  in  solution.  This  is  effected  by  osmosis  or  diosmosis, 
a  phenomenon  by  means  of  which  two  different  liquids,  or  gases, 
which  are  capable  of  mixing  and  which  are  separated  by  a 
thin  membrane  permeable  to  both  of  them,  are  able  to  diffuse 
through  the  membrane  until  a  condition  of  equilibrium  is 
attained,  the  liquid  or  gas  on  each  side  of  the  membrane  then 
being  of  the  same  density  and  composition.  If  the  membrane 
is  more  easily  permeated  by  one  solution  than  by  the  other  a 
larger  quantity  of  the  former  will  pass  through  it  and  the 
latter  will  consequently  increase  while  the  former  decreases  in 
volume.  The  current  which  flows  towards  the  solution  which 
increases  in  volume  is  distinguished  as  endosmose  and  the  cur- 
rent in  the  opposite  direction  is  exosmose.  Endosmose  usually 
takes  place  towards  the  denser  liquid  and  if  a  pig's  bladder  is- 


87 

filled  with  a  strong  solution  of  common  salt,  or  some  other 
solution  denser  than  water,  and  is  then  immersed  in  pure 
water  the  quantity  of  water  passing  into  the  bladder  is  con- 
siderably greater  than  that  of  the  solution  which  passes  out  of 
it.  The  more  concentrated  the  solution  also,  the  more  rapid 
is  the  endosmotic  current.  The  wall  of  a  plant  cell,  like  other 
organised  substances,  is  capable  of  imbibing  water  and  swelling. 
Protoplasm  is  in  this  way  able  to  imbibe  water  to  the  extent  of 
90  %  of  its  entire  weight,  although  it  then  becomes  almost  fluid. 
That  great  pressures  may  be  created  by  this  property  of  im- 
bibition is  shown  by  the  fact  that  the  swelling  of  wooden 
wedges,  which  have  been  moistened  with  water,  will  suffice 
to  split  granite  rocks,  thus  indicating  how  strong  this 
attraction  for  water  is.  The  walls  of  living  active  cells  are 
always  saturated  with  the  water  they  have  imbibed  and  in 
this  condition  they  are  permeable  to  solids  and  gases,  pro- 
vided these  are  in  a  state  of  solution,  and  in  this  state  these 
substances  are  able  to  diosmose  freely  through  the  cell  walls. 

In  the  living  cell,  however,  another  factor  besides  the  cell- 
wall  enters  into  the  case  which  is  able  to  a  certain  extent  to 
control  and  regulate  this  purely  physical  phenomenon  of  dios- 
mosis  and  that  is  the  living  protoplasm  which  lines  the  interior 
of  the  cell-wall.  It  by  no  means  follows,  for  example,  that 
substances  which  are  able  to  readily  pass  through  the  cell-wall 
will  be  able  to  pass  through  the  protoplasmic  lining  and  thus 
penetrate  to  the  cell-sap.  The  protoplasm  in  fact  decides  what 
shall,  and  what  shall  not,  pass  respectively  from  the  cell-sap  to 
the  exterior  of  the  cell  and  from  the  exterior  into  the  cell-sap. 
Seeing  that  the  protoplasm  of  one  plant  differs  more  or  less 
fundamentally  in  character  from  that  of  another  plant  and 
that  the  protoplasm  of  each  individual  thus  has  its  own  parti- 
cular requirements  to  satisfy,  we  should  natural!}'  expect 
that  the  materials  absorbed  should  differ  in  different  cases, 
and  that  while  one  plant  takes  up  large  quantities  of  a  particular 
substance  another  may  take  up  less,  and  this  is  actually  the  case. 
Moreover,  plants  on  account  of  this  selective  power  are  able 
to  collect  and  accumulate  large  amounts  of  substances  which 
exist  only  in  very  small  quantities  in  their  surrounding  medium, 
diffusion  from  outside  continually  replacing  the  small  quantities 
of  any  particular  substance  which  the  plant  may  absorb  from  its 
environment. 

84.  The  controlling  power  of  the  Turgescence. 
living   protoplasm    is   also   responsible    for  the   phenomenon 
called  turgescence.     In    the  cell-sap  of    the    living    alga,    for 


88 

iastance,  strong  solutions  of  sugars  and  other  substances 
exercising  a  powerful  attraction  for  water  are  formed  by  the 
protoplasm  and  are  held  by  it  in  the  cell,  so  that  a 
current  of  water  is  drawn  rapidly  into  the  cell  by  endos- 
mosis  just  as  happens  in  the  case  of  the  pig's  bladder.  Any 
Rubstances  also  which  are  in  solution  in  the  water  surround- 
ing the  cell  which  are  either  absent  from  the  cell-sap,  or  of  which 
a  smaller  proportion  exists  in  the  cell-sap  than  is  present 
in  the  surrounding  water,  will  also  pass  into  the  cell-sap  with 
the  water  current,  provided  that  the  protoplasm  permits  them 
to  pass. 

Now,  although  the  protoplasm  in  this  way  allows  a  solution 
of  particular  materials  to  pass  through  its  substance  into  the 
cell-sap,  it  does  not  permit  it  to  filter  back  again  out  of  the  cell, 
but  holds  it  fast.  As  water  continues  to  pass  into  the  cell 
the  protoplasmic  lining  is  pressed  outwards  against  the  cell- 
wall  which  in  its  turn  becomes  distended,  but  which,  by  its 
resistance  to  the  pressure,  develops  a  state  of  tension  in  the 
cell.  A  cell  in  this  state  becomes  stiff  just  like  a  bladder 
distended  by  air  and  is  said  to  be  turgid,  or  in  a  state  of  tur- 
gescence. 

The  young  growing  leaves  and  shoots  of  the  higher  plants, 
whioh  consist  almost  entirely  of  living  cells,  each  one  of  which 
closely  resembles  our  algal  cell,  are  enabled  to  stand  erect 
and  to  exhibit  considerable  rigidity,  owing  to  a  number  of  the 
cells  being  turgid  and  stiff.  If  a  young  shoot  is  cut  and  kept 
without  water  we  know  that  it  soon  droops  and  becomes 
flaccid,  the  cells  composing  it  having  lost  water  and  being 
no  longer  able  to  remain  turbid. 

Again,  if  a  living  cell  is  brought  into  contact  with  a  con- 
centrated solution  of  a  substance  which  has  a  stronger  attraction 
for  water  than  have  the  substances  contained  in  the  cell-sap 
water  may  be  drawn  out  of  the  cell  to  such  an  extent  that  it 
is  no  longer  able  to  remain  turgid  and  the  protoplasm  becomes 
inactive  and  unable  to  perform  its  functions.  It  is  on  this 
account  that  plants  are,  as  a  rule,  unable  to  obtain  their  supplies 
of  water  and  hence  also  of  dissolved  mineral  salts  from  concen- 
trated solutions. 

Transpira-  35 .  As  the  mineral  salts   required 

as  raw  food  materials  must  thus  enter  the  living  turgid  cell 
as  very  dilute  solutions,  in  order  that  the  cell  may  obtain  a 
sufficiency  of  salts  a  large  quantity  of  water  must  continually 
be  got  rid  of,  and  the  protoplasm,  while  not  allowing  the  salts 
which  it  requires  to  pass  out  of  the  cell  and  be  lost,  does 


^continually  permit  the  water  which  is  in  excess  of  that  re- 
-  quired  to  maintain  the  cell  in  a  state  of  turgidity  to  escape. 
In  terrestrial  plants  this  excess  water  passes  off  in  the  form 
of  water- vapour  and  the  phenomenon  is  termed  transpiration. 
Room  is  thus  made  in  the  cell  for  more  water  and  a  con- 
tinual fresh  supply  of  water  and  salts  is  provided  for. 

86.  Not     Only     solids,      but      also  Manufacture 

gases,  in  solution  are  able  to  enter  the  cell  by  osmosis  andpoj[gan 
oxygen  and  carbon  dioxide,  which  are  dissolved  in  the  water  Material, 
surrounding  it,  are  thus  able  to  penetrate  into  the  algal 
cell.  Now,  provided  that  the  temperature  is  suitable  and 
there  is  sufficient  light,  the  protoplasm  is  able,  by  the  help 
-of  the  green  chlorophyll,  to  decompose  the  water  and  car- 
bon dioxide  contained  in  the  cell  and  to  build  up  from 
them  an  organic  carbohydrate  which  is  usually  starch,  oxygen 
being  given  off.  This  is  merely  a  preliminary  process  consist- 
ing in  the  building  up  of  a  food  material  which  in  this  case 
precedes,  and  is  quite  independent  of,  nutrition  proper,  just 
as  the  collection,  or  cooking,  of  food  may  precede  actual 
feeding.  Some  plants  indeed  exist  which,  as  we  shall  see 
later,  can  dispense  altogether  with  this  preliminary  process, 
although  they  are  composed  of  living  protoplasm  which 
requires  feeding  just  as  does  that  of  green  plants.  Such  plants 
possess  no  chlorophyll,  but  by  various  means  they  are  able  to 
obtain  their  organic  food  materials  ready-made  for  them  by 
-green  plants. 

87.  Living    protoplasm,    as    has  Respiration 

been  already  noticed,  respires,  and  plants,  like  animals,  absorb  Metabolism. 
oxygen  and  give  off  carbon  dioxide.      During    this    process 

some  of  the  substance  of  the  protoplasm  itself  appears  to  be 
destroyed  and  broken  down  into  simple  compounds,  energy 
being  evolved  which  is  in  great  part  dissipated  in  the 
form  of  heat.  If  therefore  the  protoplasm  is  to  be  main- 
tained alive  and  enabled  to  grow,  there  must  be  a  continual 
supply  of  energy  available  by  means  of  which  the  very  com- 
•plex  protoplasm  may  be  again  built  up  from  simpler  sub- 
stances, and  further  in  order  that  the  protoplasm  should 
actually  increase  in  bulk  and  grow,  it  is  obvious  that  not 
only  must  those  molecules  be  replaced  which  have  been  des- 
troyed but  additional  molecules  must  be  constructed.  This 
constant  source  of  fresh  energy  is,  in  green  plants,  supplied  by 
the  starch  which  is  being  continually  manufactured  in  the 
orophyll  corpuscles,  and  a  considerable  proportion  of 
which  is  a#ain  being  continually  broken  down  into  carbon 


90 

dioxide  and  water  in  respiration.  By  the  breaking  up  of  a 
molecule  of  starch  energy  is  liberated  and  this  energy  can  be 
employed  by  the  protoplasm  in  so  working  on  the  various 
mineral  salts  and  other  substances  in  the  cell  that  they,  or 
some  of  them,  are  bound  together  into  protoplasm  which  is 
thus  continually  nourished  and  enabled  to  grow,  while  surplus 
energy  is  also  available  for  various  kinds  of  work,  such  as  the 
building  up  of  substances  having  a  strong  attraction  for  water 
and  which  are  thus  able  to  cause  osmotic  currents.  One 
and  the  same  substance,  however,  may  be  dealt  with  by  the 
living  protoplasm  in  various  ways  according  to  its  needs 
at  the  time.  The  starch  which  has  been  manufactured  in  the 
chlorophyll  corpuscles,  for  instance,  may  be  stored  as  reserve 
material  and  put  aside  in  the  form  of  starch  grains,  as  not 
being  at  present  needed,  it  may  be  converted  into  cellulose 
and  built  into  the  cell- wall,  it  may  be  used  as  food  and  built  up 
into  protoplasm,  or  it  may  be  at  once  broken  up  in  respiration, 
made  to  give  up  its  contained  energy  and  converted  again 
into  carbon  dioxide  and  water. 

When  it  is  considered  also  that  there  is  a  variety  of 
substances  in  the  living  cell,  both  mineral  salts,  organic 
substances  and  gases,  and  that  they  all  may  be  dealt  with 
in  a  variety  of  ways,  it  is  obvious  that  the  actual  processes 
going  on  in  the  cell  are  exceedingly  complex  and  as  a 
matter  of  fact  they  are  very  little  understood.  The  main 
fact  known  about  them  is,  however,  that  they  may  all  be 
placed  in  two  great  groups  —  (1)  those  which  consist  in 
the  breaking  down  of  complex  to  simple  substances  result- 
ing in  the  liberation  of  energy,  some  of  which  passes  off  as 
heat  but  some  of  which  is  always  available  for  work  in  the 
cell,  and  (2)  those  which  consist  in  the  building  up  of  complex 
from  more  simple  compounds,  the  ultimate,  or  end,  product 
being  the  exceedingly  complex  protoplasm  itself.  The  latter 
processes  collectively  are  termed  assimilation,  or  anabolism, 
and  the  former  disassimilation,  or  Jcatabolism,  while  all  these 
processes  together  are  termed  metabolism,  which  thus  means  the 
sum  total  of  all  the  chemical  changes  which  go  on  in  a  living 
cell.  The  building  up  of  starch  by  green  plants  is  thus  a  process 
of  assimilation.  A  large  number  of  the  substances  which  are 
formed  in  the  cell  during  these  various  changes  are  of  further 
use  to  the  plant  and  can  again  be  used  in  anabolism,  or  kata- 
bolism,  but  some  are  of  no  further  use  and  these  so-called 
waste -products  are  got  rid  of  by  the  protoplasm  and  if  possible 
are  excreted,  i.e.  passed  out  of  the  cell. 


91 

88.  If  now,  instead   of  consider-  Conditions  of 

ing  a  single  cell,  we  turn  our  attention  to  a  large  plant  like  a  f  Highly! 1E 
tree  which  has,  as  we  have  already  seen.,  a  very   complicated  Organised 
structure,  we  begin  to  realise  the  necessitv  for  its  variously plant' 

o  **  «/ 

differentiated  parts.  If  the  body  of  such  a  plant  were  com- 
posed of  an  aggregate  of  thin-walled  cells,  it  is  obvious  that 
turgidity  alone  would  not  suffice  to  enable  it  to  stand  erect  and 
to  resist  the  enormous  strains  to  which  it  is  subjected.  Many 
cells  therefore  are  deprived  of  their  protoplasm  and  are  converted 
into  thick-walled  elements  such  as  fibres,  which  serve  to  strength- 
en and  support  the  plant  body.  Further,  although  food 
materials  in  solution  could  diffuse  through  such  a  large  plant 
body  if  it  were  composed  entirely  of  thin- walled  living  cells,  by 
osmosis  from  cell  to  cell,  the  currents  thus  produced  would  be  too 
slow  to  satisfy  the  plant's  requirements  and  hence  we  find 
that  some  cells  are  converted  into  conduction  pipes,  such 
as  the  vessels  and  tracheids,  which  are  able  to  conduct  water 
and  salts  rapidly  from  the  roots  to  the  leaves,  and  the  sieve- 
tubes  which  conduct  the  organic  food  from  the  leaves  to  the 
places  where  it  is  required.  By  means  of  a  system  of  inter- 
cellular spaces,  also,  which  communicate  with  the  outer  air, 
fresh  currents  of  carbon  dioxide  and  oxygen  are  continually 
brought  into  contact  with  the  walls  of  the  living  cells,  while 
the  water  vapour  exhaled  by  the  latter  is  able  to  quickly  pass  off 
into  the  outer  air.  That  water  is  actually  absorbed  by  the 
roots  and  is  given  off  by  the  leaves  of  a  highly-organised 
plant  can  be  easily  seen,  if  the  uninjured  roots  of  a  young 
plant  are  put  in  an  air-tight  vessel  of  water,  the  whole 
being  placed  in  a  balance.  .The  amount  of  water  then  taken 
up  by  the  roots  can  be  directly  measured,  while  the  amount 
given  off  by  the  leaves  is  given  by  the  loss  of  weight  indicated 
by  the  balance. 

If  the  external  conditions  remain  constant,  the  amount  of 
water  absorbed  by  the  roots  is  equal  to  the  amount  given  off 
by  the  leaves. 

The  fact  that  green  plants  can  grow  and  develop 
normally  in  pure  sand,  or  distilled  water,  to  which  certain 
mineral  salts  have  been  added,  but  which  contains  no  carbon, 
shows  that  the  carbon  required  for  their  organic  food  is  not 
obtained  from  the  soil.  The  further  fact  that  starch  is  actually 
formed  by  .the  green  leaves  is  also  easily  shown  by  the  follow- 
ing experiment. 

Part  of  a  healthy,  living  leaf  on  a  plant  which  has  been  kept 
in  the  dark  for  some  hours  is  covered  with  a  piece  of  tinfoil, 


92 

or  other  substance,  which  does  not  allow  light  to  pass  through  it, 
and  the  plant  with  its  partly  covered  leaf  is  then  placed  in 
bright  sunshine  for  some  hours.  The  leaf  is  then  cut  off 
and  killed  by  being  placed  a  short  time  in  boiling  water. 
It  is  then  placed  in  warm  alcohol  and  thus  decolourised,  the 
chlorophyll  passing  into  the  alcohol  in  solution.  The  leaf  is 
then  placed  in  a  solution  of  iodine  and  finally  washed  with  water, 
when  all  parts  which  had  been  exposed  to  the  light  will  be  found 
to  have  turned  blue-black,  the  colour  indicating  the  presence  of 
starch,  the  manufacture  of  which  was  only  possible  in  those 
parts  where  the  chlorophyll  corpuscles  were  exposed  to  the 
light. 


CHAPTER  II.— FUNCTIONS  OF  THE  HIGHER  PLANTS  IN 

DETAIL. 

89.  It  has  been  already  pointed  SaffcF,ood 

-,  •    i  t  ^          i      ^i     •  i-  <•        •  ,  Materials. 

out  that  the  higher  plants  absorb  their  supplies  of  mineral 
salts  and  water  mainly  by  means  of  the  root-hairs,  each  of 
which  is  merely  the  prolongation  of  a  single,  living,  epidermal 
cell,  provided  with  a  thin  wall  of  cellulose  and  containing 
protoplasm,  a  nucleus,  and  cell-sap.  In  all  essential  details,  in 
fact,  it  resembles  our  algal  cell  except  that  the  green  chlorophyll 
corpuscles  are  absent.  In  such  a  root-hair,  therefore,  the 
assimilation  of  a  carbohydrate  from  inorganic  materials  cannot 
take  place,  but  respiration  and  the  absorption  of  mineral 
substances  in  solution  can  take  place,  as  in  the  alga. 

The  chemical  constituents  of  the  plant  body  can  be  found 
by  analysis.  Water  in  considerable  quantities  always  occurs 
In  a  living  plant,  and  in  turgid  herbaceous  plants  water 
may  amount  to  as  much  as  90%  of  the  entire  weight  of 
the  plant.  About  half  the  dry  weight  of  a  plant  is  made  up  of 
carbon. 

In  Part  II  we  have  seen  that  a  large  number  of  additional 
substances  enter  into  the  composition  of  the  protoplasm,  but 
other  substances  also  occur  and  practically  all  known  elements 
have,  at  one  time  or  another,  been  detected  in  plants. 

In    order  to  discover   which    of   all   these   substances  are  Essential 
absolutely  essential  and  in  what  quantities  they  are  required  Substancss. 
for  the  growth  and  development  of  various  plants,  the  latter 
must  be  grown  in  pure  water  to    which  only  known  quantities 
of  known  substances  are  added.     By  such  water-culture    ex- 
periments it  has  been  discovered  that  the  following  are  indis- 
pensable for  the  higher  plants : — 

Carbon.  Phosphorus. 

Hydrogen-.  Potassium. 

Oxygen.  Magnesium . 

Nitrogen.  Calcium. 

Sulphur.  Iron. 

All    of    these    substances,    excepting    potassium,  magnesium 

and    calcium,    are    believed   to  enter  into  the  composition  of 

the  protoplasm    itself.     The    calcium    appears    to    be    often 

useful  in  combining  with  poisonous  substances,    e.g.    oxalic 

acid,  which  are  then  deposited    in   the  plant    in   a    harmless 

form.     Of  some  of  these  essential  substances    only  very  small  Quantities 

quantities  are   required,  but  if  that   small  quantity  is  absent  teiu!l'ei1- 


94 


plant  growth  and  development  are  impossible.  Thus  very 
little  iron  is  required,  but  unless  the  necessary  minimum 
is  present  no  chlorophyll  is  developed.  Of  these  various 
elements  the  carbon  is  obtained  from  the  carbon  dioxide 
of  the  air,  oxygen  is  obtained  partly  from  the  air  and 
partly  from  water,  hydrogen  from  water  and  the  others  are  all 
Availably,  absorbed  in  solution  by  the  roots  from  the  sc-il.  Large  quanti- 
ties of  these  necessary  substances,  however,  may  be  present  in 
the  soil  and  yet  not  be  available  for  the  plant,  because  they 
occur  in  a  form  in  which  the  plant  can  make  no  use  of  them. 
As  only  substances  in  solution  can  penetrate  the  cell-wall  and 
thus  enter  the  absorbing  root-hairs,  only  such  substances  as 
are  soluble  can  be  utilised  by  the  plant.  Although  it  appears 
that  some  of  the  higher  plants  can  directly  utilise  ammonium 
salts,  most  of  them  can  only  obtain  their  nitrogen  readily 
.Soil—  when  it  occurs  in  the  form  of  nitrates.  It  a  solution  of  organic 

Absorption.  an(j  morganjc  substances  is  poured  on  ordinary  soil,  it  will 
be  found  that  the  water  which  filters  through  is  usually 
different  in  composition  to  that  originally  added.  This  is  due 
to  the  power  possessed  by  the  soil  of  absorbing  and  retaining 
various  substances,  partly  on  account  of  chemical  combi- 
nations taking  place,  and  partly  on  account  of  the  attraction 
exercised  by  the  particles  of  soil.  Soils  vary  considerably  with 
regard  to  their  absorptive  power  ;  those  containing  humus 
absorb  the  most  and  are  thus  able  to  store  up  large 
quantities  of  valuable  plant  food-materials,  while  sandy  soils 
have  very  little  absorptive  power.  All  substances,  however, 
are  not  retained  with  equal  vigour ;  ordinary  soils  allow  the 
valuable  nitrates  to  filter  through  and  pass  away  in  drainage 
water,  while  phosphates,  potassium  salts  and  ammonium  are 
usually  strongly  held.  Such  absorbed  substances  can,  however, 
be  gradually  washed  out  again  if  sufficient  water  is  continually 
added  to  the  soil,  so  that  from  the  stored  substances  the 
dilute  solutions  required  by  plants  are  thus  made  available 
in  a  moist  soil. 

Living  active  root-hairs  are  continually  excreting  carbon 
dioxide  as  a  product  of  their  respiration,  and  this,  passing  into 
the  water  which  surrounds  them,  aids  materially  in  dissolving 
and  making  available  for  the  plant  useful  substances  in  the 
soil.  Calcium  carbonate,  for  example,  is  readily  soluble  in 
water  containing  carbon  dioxide  and,  if  a  plant  is  allowed  to 
grow  with  its  roots  touching  a  polished  marble  plate,  the 
latter  will  be  corroded  and  etched  where  the  roots  have  come 
in  contact  with  it. 


Power  of 
Plants  to 
obtain 

Necessary 
-Substances. 


95 

Plants,  however,  differ  considerably  in  their  power  of  extract- 
ing their  supplies  of  necessary  substances  from  the  soil  and  those 
which  can  tap  supplies  which  are  not  available  for  other  plants 
may  therefore  develop  normally  on  soils  where  the  latter  cannot 
exist.  As  a  general  rule,  the  substances  which  the  higher  plants 
seem  to  find  most  difficult  to  obtain  in  sufficient  quantities  are 
nitrogen,  phosphorus,  and  potassium,  and  these  substances  in 
suitable  combinations  are,  generally  speaking,  the  most  valu- 
able manures  for  field  crops,  the  soil  being  able  to  supply 
a  sufficiency  of  the  other  substances.  Those  plants  which 
have  special  difficulty  in  obtaining  any  one  of  these  substances 
from  ordinary  soil  may  therefore,  if  grown  continually  on  the 
same  ground,  be  unable  after  a  few  years  to  obtain  a  sufficient 
supply  for  their  vigorous  development  and  the  yield  of  agricul- 
tural crops  may  thus  sink  to  Jrd  the  average  produce. 

To  show  how  plants  vary  in  this  respect,  however,  it  should  be 
mentioned  that,  according  to  experience  in  Europe,  although 
turnips  remove  large  quantities  of  nitrogen  and  potassium  from 
the  soil,  they  as  a  rule  only  require  a  manure  containing  phos- 
phorus, whereas  barley  and  many  other  field  crops  usually 
require  an  additional  supply  of  nitrogen  in  the  form  of  manure, 
if  a  good  crop  is  to  be  produced.  Again,  experiments  extending 
over  50  consecutive  years  have  shown  that,  when  barley  :s 
grown  on  one  and  the  same  soil  continuously,  only  nitrogen 
and  phosphorus  being  given  as  manure  and  no  potassium, 
the  crop  will  still  be  nearly  normal  and  equal  to  that  produc- 
ed when  a  full  manure  of  potassium,  nitrogen  and  phospho- 
rus is  given.  If  grown  continuously  for  the  same  period  in 
the  same  soil  with  no  manure  at  all,  the  crop  will  sink  to  nearly 
^rd  the  normal,  while  if  nitrogen  is  omitted  from  the  manure 
the  yield  sinks  to  less  than  \  the  normal. 

Very  little  is  known  regarding  the  peculiarities  of  our  forest 
trees  in  this  respect,  but  it  is  clear  that  we  have  to  consider 
not  only  the  quantity  of  essential  substances  contained  in  the 
soil  and  actually  removed  by  the  plants,  but  also  the  difficulties 
which  each  species  in  nature  experiences  in  availing  itself  of 
the  substances  which  are  present  in  the  soil,  as  on  this 
may  depend  the  question  of  whether,  or  not,  a  certain  species 
can  be  grown  continuously  for  long  periods  in  the  same  soil. 

The  root  systems  of  different  plants  also  develop  in  different 
layers  of  soil,  and  on  such  facts  in  great  measure  depends  the 
theory  of  rotation  of  crops  and  the  growing  of  mixed  crops. 

Many  plants  are  in  nature  found  to  take  up  large  quantities  of 
substances  which,  so  far  as  we  can  judge  from  water- culture  Substances. 


96 


Absorption 
and  Ascent 
of  Wtaer  in 
Plarts. 


Root- 
Pressure. 


experiments,  are  not  essential  for  their  growth  and  de\  elopment. 
These  substances,  however,  may  often  be  indirectly  useful ;. 
thus  silica  is  often  taken  up  in  large  quantities  and  in  many 
plants  is  deposited  in  the  epidermal  cell-walls.  This  increases 
the  hardness  and  rigidity  of  the  plant  and  may  aid  it 
considerably  in  the  struggle  for  existence  by  rendering  it  nn~ 
palatable  to  grazing  animals.  In  this  way  the  existence  of 
a  plant,  in  its  natural  surroundings,  may  depend  on  the 
presence  of  substances  which  appear,  at  first  sight,  to  be  un- 
necessary for  its  healthy  development. 

90.  Substances  in    solution   pass 

into  the  root-hairs  by  osmosis  as  in  the  case  of  the  algal 
cell,  but,  unlike  the  latter,  an  individual  root-hair  does  not 
remain  in  a  continual  state  of  turgidity.  The  protoplasm 
of  the  turgid  root-hair  after  a  time  appears  to  relax  its 
resistance  to  the  outward  pressure  of  the  water  and  sub- 
stances in  solution  which  have  accumulated  in  the  cell-sap 
and  this  absorbed  water  with  substances  in  solution  conse- 
quently filters  back  out  of  the  cell  under  a  pressure  corre- 
sponding to  that  previously  exercised  by  the  protoplasm  sup- 
ported by  the  cell- wall  by  means  of  which  the  state  of  tur- 
gescence  was  attained.  That  this  pressure  can  be  very  consi- 
derable is  indicated  by  the  fact  that,  when  a  young  root  pene- 
trates a  crack  in  a  rock,  or  masonry  wall,  as  the  root  increases 
in  size,  the  pressure  exercised  by  its  cells,  in  their  efforts  to 
become  turgid  and  grow,  may  suffice  to  split  the  rock,  or  wall, 
as  the  case  may  be. 

The  water,  however,  which  is  thus  pressed  out  of  the  root- 
hairs,  instead  of  passing  out  into  the  soil  and  thus  being  lost  to 
the  plant,  finds  its  way  from  cell  to  cell  of  the  fundamental 
tissue  of  the  root  and  eventually  passes  into  the  vessels  and 
tracheids  of  the  vascular  bundles. 

That  water  with  salts  in  solution  is  thus  pressed  by  the 
roots  into  the  vessels  and  tracheids,  and  often  with  consider- 
able force,  can  be  seen  by  felling  a  tree  when  the  so-called 
"sap"  is  rising,  i.e.  just  before  the  leaves  appear  and  when  the 
roots  have  commenced  to  actively  absorb  water  from  the  soil. 
The  water  can  be  plainly  seen  to  exude  from  the  vessels  and 
tracheids  on  the  cut  section  and  if  analysed  it  is  found  to  contain 
various  substances  in  solution.  By  attaching  a  bent  tube  con- 
taining mercury  to  the  rooted  stumps  of  cut  plants  the  pres- 
sure with  which  this  water  is  pumped  up  can  be  measured,  and 
it  is  found  to  frequently  exceed  that  of  one  atmosphere. 


97 

Such  "  bleeding "  may  continue  for  several  days,  or  even 
weeks,  and  if  the  roots  are  kept  warm  and  well  supplied  with 
water  the  quantity  of  fluid  exuded  may  be  large,  and  often 
considerably  exceeds  in  volume  the  total  volume  of  the  root 
system. 

That  this  so-called  root-pressure,  however,  in  itself  does  not 
suffice  to  explain  the  ascent  of  the  water  current  in  high  trees  is 
shown  by  the  following  facts.  If  the  total  quantity  of  water 
given  off  by  such  bleeding  stems  is  measured  it  is  found  to  be 
very  much  less  than  the  amount  actually  required  and  given 
off  by  the  transpiring  leaves.  Again  if  an  actively  transpiring 
plant  in  full  leaf  is  cut,  the  stump,  instead  of  exuding  water, 
will  at  first  actively  absorb  it,  and  water  placed  on  the  cut  section 
will  be  taken  up  by  the  vessels  and  tracheids,  thus  showing 
that,  instead  of  a  positive  root-pressure  existing  in  the  latter, 
the  existing  pressure  is  less  than  that  of  one  atmosphere.  It 
has  been  ascertained  also  that  when  transpiration  is  actively 
proceeding  the  water  does  not  occur  in  continuous  columns 
in  the  conducting  elements,  but  as  short  columns,  alternating 
with  bubbles  of  air,  and  that  the  pressure  in  them  generally 
decreases  towards  the  top  of  the  tree. 

Now,  if  for  any  reason  there  exists  a  demand  for  a  particular 
substance  in  a  certain  cell,  that  substance  will  continue  to 
be  attracted  to  the  cell  until  the  demand  is  satisfied.  If. 
for  instance,  owing  to  the  presence  of  a  concentrated  solution 
of  a  substance  having  a  strong  attraction  for  water,  an  osmotic 
current  into  the  cell  is  created,  and  further,  if  this  water  is  driven 
off  in  the  form  of  vapour  as  fast  as  it  enters  the  cell,  the  inward 
current  will  continue  to  flow,  since  the  needs  of  the  cell  in  res- 
pect of  water  remain  unchanged.  If,  on  the  other  hand, 
the  water  is  allowed  to  dilute  the  solution  and  to  continually 
carry  off  some  of  the  same  to  neighbouring  cells  by  exosmosis, 
a  condition  of  equilibrium  will  sooner  or  later  be  reached, 
the  concentration  of  the  solution  in  all  the  cells  being  the 
same.  Glucose,  or  grape-sugar,  is  a  soluble  substance  derived 
from  starch  by  the  addition  of  a  molecule  of  water,  and  if  we 
suppose  that  a  dilute  solution  of  glucose  is  flowing  into  a  cell 
by  osmosis  and  that  on  reaching  the  cell  the  glucose  becomes 
converted  into  insoluble  starch,  which  is  deposited  as  starch 
grains,  the  inward  flow  of  glucose  will  continue,  the  cell's  needs 
in  respect  of  glucose  remaining  unsatisfied.  It  is  in  this  way 
that  large  quantities  of  the  so-called  reserve  food-materials 
are  accumulated  and  stored  in  certain  cells  until  they  are 
required.  Similarly,  although  currents  of  water  with  salts 


98 


Sucking- 
Force  in 
Leaves. 


Transpira- 
tion. 


in  solution  are  continually  flowing  by  osmosis  into  the  living 
cells  of  the  leaf,  each  of  which,  like  the  algal  cell,  is  busily 
engaged  in  the  sunlight  in  manufacturing  starch  and  in  building 
up  protoplasm  to  replace  that  lost  by  respiration,  yet,  as  the 
water  which  enters  is  again  continually  exhaled  in  transpiration 
and  as  the  salts  which  entered  with  the  water  are  continually 
being  seized  on  by  the  protoplasm,  made  to  combine  with  other 
substances  and  built  up  into  protoplasm,  or  otherwise  changed, 
the  attraction  exercised  by  the  cells  for    water    and   these   par- 
ticular salts  in  solution  remains  unchanged,  so  long  as  the  vital 
processes    mentioned    continue     actively,     and     water    and 
salts    are   in    consequence    continually   withdrawn   by  the  living 
cells  of  the  leaves  from  the  vessels  and  tracheids  with  which  they 
are  in  contact.    In  this  way  a  backwardly-acting  sucking  force   is 
developed  which,  in  some  way  not  yet  clearly  understood,  extends  to 
the  roots  and  causes  fresh  supplies  of  water  and  salts   to  continually 
pass    into    the    xylem    elements  of    the  vascular  bundles  from 
the  root-hairs.    So  much,  however,  is  certain  that   there    is  no 
enormous  pressure  acting  from  below  which  forces  the  water  up 
to  the  summit  of  high  trees,   and  that,  although    the   suction 
force  acting  from  above  is  considerable,  it  also  by  itself  is  not 
sufficient    to    account    for    the    transpiration    current.     The 
most  recent  researches  show  that,  if  the  living  cells  in  the  wood 
are  killed,  the  latter  soon  loses  its  power  of  conducting  water, 
and  it  is  thought  that  the  living  cells  of  the  medullary  rays 
and  wood  parenchyma  which  are  in  contact  with  the   vessels 
and  tracheids  assist  in  helping  on  the  flow  from  point  to  point. 
It  has,  however,  been  ascertained  that  the  water  current  ascends 
mainly  in  the  cavities  of  the  vessels  and  tracheids  and  not  in 
their  walls,  and  by  compressing  transpiring  shoots  and  thus 
diminishing  the  sectional  area  of  the  cavities  of  their  elements 
the  water  current  can  be  greatly  reduced.     It  has  also  been 
ascertained    that  the  water  current  ascends  in  the  younger 
layers  of  wood  and  that  when  heart- wood  is  present  it  takes 
no   part   in   the    conduction   of   water. 

91.  In    large   thin    leaves    the 

living  cells  are  not  shut  off  from  the  outer  air  so 
effectually  as  are  those  in  the  stem  and  branches  which 
are  soon  protected  by  an  outer  coat,  several  layers  of 
cells  in  thickness,  of  cork,  or  other  more  or  less  impervious 
material,  and  hence,  although  the  outer  walls  of  the  epidermal 
cells  of  such  leaves  are  usually  more  or  less  thickened,  a  certain 
quantity  of  water  is  continually  abstracted  from  the  leaf  by 
evaporation.  This  process  also  is  largely  promoted  owing  to 


99 

the  fact  that,  by  means  of  the  numerous  stomata  and  large 
intercellular  spaces  in  the  leaf  tissue,  the  atmospheric  air  is 
•able  to  actually  come  in  contact  with  the  thin  walls  of  the 
living  turgid  cells  in  the  interior  of  the  leaf  which  are  saturat- 
ed with  water.  Room  is  thus  made  in  these  cells  for  more 
water  which  accordingly  flows  into  them  from  the  vessels 
and  tracheids  of  the  leaf  vascular  strands  and  continually 
brings  with  it  a  fresh  supply  of  the  necessary  salts.  This 
evaporation  of  water  from  the  leaves  is  called  transpiration. 
Within  certain  limits  plants  are  able  to  regulate  and  control  Regulation  of 
the  amount  of  water  which  shall  thus  escape  from  them  byTranspiratioiu 
closing  the  stomata,  by  developing  a  thick  coating  of  hairs,  or 
other  protective  covering,  from  the  epidermis,  or  by  other 
devices,  and  consequently  the  quantity  of  water  transpired 
from  a  given  surface  of  living  leaves  is  always  less  than  that 
which  would  have  evaporated  from  an  equal  surface  of  water 
while  a  dead  leaf,  also,  loses  water  by  evaporation  quicker  than 
a  living  leaf.  With  this  proviso,  however,  the  factors  on  which 
transpiration  depends  are  essentially  the  same  as  those  which  re- 
gulate ordinary  evaporation,  and  the  amount  of  water  transpired 
therefore  depends  on  the  temperature  of,  and  amount  of 
water  contained  in,  the  surrounding  air,  the  presence  or  absence 
of  air  currents,  and  the  area  of  the  surface  exposed  to  eva- 
poration. As  a  general  rule,  the  rapid  removal  of  water 
from  the  actively  assimilating  cells  of  the  leaf  is  very  necessary 
to  enable  them  to  obtain  the  necessary  quantity  of  salts  they 
require  from  the  very  dilute  solution  absorbed  by  the  roots. 
There  are,  however,  obvious  cases  in  which  rapid  transpiration 
may  be  highly  injurious,  such  as  when  the  roots  can  obtain 
very  little  water  from  a  dry  soil,  and  it  is  essential  that  plants 
should  be  able  to  regulate  this  loss  of  water.  That  this  can 
only  be  done  within  definite  limits  is  indicated  by  the  fact 
that,  during  a  hot  dry  day,  plants  may  frequently  be  seen  to 
droop  and  become  flaccid,  the  amount  of  water  which  has  been 
lost  by  transpiration,  in  such  cases,  being  so  great  that  the 
living  cells  have  been  unable  to  obtain  sufficient  water  from  the 
roots  to  enable  them  to  remain  turgid.  They  have  accordingly 
become  inactive  and  unable  to  perform  their  vital  functions. 
Unless  the  loss  of  water  has  been  too  great  such  plants  usually 
recover  in  the  night,  owing  to  the  diminished  transpiration. 
In  some  cases  when  little  or  no  evaporation  can  take  place,, 
owing  to  the  saturation  of  the  atmosphere,  or  other  cause, 
plants  may  become  filled  with  an  excess  of  water  and  salts 
in  solution  which  is  pumped  into  them  by  the  active  roots. 

H  2 


100 

Water-Pores  ^his  excess  water  is  often  got  rid  of  by  exudation  from  the 
ordinary  stomata,  or  from  special  openings  in  the  leaves  called 
water- pores,  such  as  are  found  for  instance  at  the  tips  of  the 
leaves  of  Colocasia  antiquorum.  Such  pores  are  usually 
larger  than  ordinary  stomata,  and,  as  their  guard-cells  contain 
no  living  protoplasm  and  cannot  therefore  alter  their  shape, 
the  pores  are  always  open.  In  the  early  morning  drops  of  water 
may  often  be  seen  exuding  from  the  leaves  of  grasses  and  other 
plants  which  have  become  surcharged  with  water  by  the 
actively  absorbing  roots  during  the  night.  In  such  cases 
the  escaping  water  carries  with  it  salts  in  solution,  whereas 
in  transpiration  only  the  water  escapes  in  the  form  of  vapour, 
the  salts  remaining  behind.  As  has  already  been  noted,  plants 
can  to  a  certain  extent  control  transpiration  by  opening  or 
closing  their  stomata  and  this  is  effected  mainly  by  the  guard- 
cells  ;  when  the  latter  are  turgid  the  stomata  open  and  vice 
versa.  The  guard-cells  become  turgid  when  active  transpira- 
tion is  advantageous,  in  strong  sunlight  when  plenty  of  water 
is  available,  or  when  the  surrounding  air  is  moist,  while  they 
become  flaccid  and  thus  close  the  stomata  when  active  trans- 
piration is  likely  to  be  injurious  owing  to  the  failure  of  the 
water  supply,  or  other  cause.  In  dry  localities  where  roots 
can  obtain  very  little  water  from  the  soil,  only  those  plants 
can  exist  which  possess  very  efficient  means  of  preventing 
rapid  transpiration,  or  which  get  over  the  difficulty  in  some 
other  way.  Thus  in  desert  plants  the  total  area  of  the  leaf 
surface  is  usually  greatly  reduced  and  leaves  in  many  species 
may  be  entirely  absent,  the  living,  chlorophyll- containing 
cells  being  situated  in  the  thick  stems  and  protected  by  a 
more  or  less  impervious  epidermis,  instead  of  being  arranged 
in  large  thin  leaves  provided  with  numerous  stomata. 

In  many  localities  the  water  difficulty  only  becomes  acute 
•  during  certain  seasons  of  the  year  and  plants  are  able  to  exist 
provided  they  can  complete  their  development  during  the 
period  when  the  soil  around  their  roots  is  sufficiently  moist 
to  supply  their  needs.  Thus  in  the  plains  of  India,  most 
grasses  and  herbs  complete  their  development  during  the  rainy 
season  and  die  down  in  the  hot  weather.  Deciduous  trees 
also  lose  their  leaves  and  remain  inactive  during  the  seasons 
when  #n  account  of  either  the  low  temperature  of  the  soil, 
or  the  scarcity  of  water,  the  roots  would  find  difficulty  in 
obtaining  their  supplies.  Evergreen  trees  on  the  other  hand 
may  be  able  to  exist  through  such  seasons  owing  to  the 
effective  checks  on  transpiration  possessed  by  them  in  the 


101 

way  of  a  reduced  leaf  area,  protective  coverings  to  the  leaf, 
and  so  on.  That  such  arrangements  may  be  very  effective  is 
indicated  by  experiments  carried  out  in  Europe  which  have 
shown  that  the  amount  of  water  transpired  by  evergreen 
conifers  is  frequently  only  iVth  of  the  amount  transpired  by 
deciduous  dicotyledons,  a  result  which  is  not  surprising 
when  we  consider  the  small  area  of  leaf-surface  exposed  to  the 
air  in  the  narrow  coniferous  needles,  the  strongly  thickened 
cuticle,  the  stomata  sunk  below  the  surface  of  the  needles  and 
thus  protected  from  air-currents,  and  the  presence  of  scler- 
enchymatous  tissue  beneath  the  epidermis  which,  as  a  rule, 
surrounds  and  protects  the  thin- walled  chlorophyll- contain- 
ing parenchyma  of  such  needles.  Again,  owing  to  their  deep- 
going  root-systems  some  trees  are  able  to  tap  a  perennial 
water  supply  and  can  thus  produce  fresh  foliage  and  transpire 
actively  in  the  hottest,  dryest  season  of  the  year. 

92.  That  a    carbohydrate,  which  Assimilation. 
is  usually  starch,  is  formed  in  the    green    leaves   of    a  healthy 
plant  exposed  to   sunlight,  from  water  and  the  carbon  dioxide 
of  the  air  has  already  been  pointed  out.     During  this  process 
oxygen  is  evolved,    the    volume    of    oxygen  given-  off  being 
approximately  equal  to  that  of  the  carbon  dioxide  absorbed. 
If  a  green  plant  is  placed  in    an   air-tight    jar    in   an    atmos- 
phere   from   which    the   carbon    dioxide  has  been    removed, 
no   starch   is    formed  in   the   leaves,     thus     indicating   that 
the   carbon    dioxide    of    the    atmosphere    is  essential  for  the 
process.     If  the  cut  stem  of  a  green  aquatic  plant  is  placed 
in  a  cylinder  of  water  in  the  sunlight,  bubbles  of  gas  will  be 
found  to  escape  from  the  intercellular  spaces  at  the  cut  surface 
which  can  be  easily  collected  and  proved  to  be  nearly  pure 
oxygen.     By  counting  the  number  of  bubbles  given  off  per 
minute,  an  idea  can  be  formed  of  the  rate  at  which  carbon 
dioxide  is   being   absorbed   and   starch   manufactured.     That 
light  is  essential  for  this  assimilation  is  shown  by  the  fact 
that  no  starch  is  formed  in  that  part  of  a  green  leaf  which  is 
covered  with  tinfoil,  or  otherwise  protected  from  the  light. 
On  the  other  hand  it  can  be  shown  by  experiment  that  all  the  Rays  of  Light 
rays  of  light  are  not  of  the  same  value  for  this  process.     If,  for  re^uired« 
instance,  a  bright  solar  spectrum  is  projected   on  a  green  leaf 
for  several  hours,  the  leaf  then  being  decolourised  with  alcohol 
and  treated  with  a  solution  of  iodine  as  before  described,  the 
blue-black  colour,  indicating  the  presence  of  starch,  will  be 
found  to  be  most  intense  in  that  part  of  the  leaf  which  was  ex- 
posed to  the  red  and  orange  rays.     The  same  thing  can  also 


102 

be  shown  by  means  of  the  green  water  p^ant  mentioned  in  the 
previous  experiment,  for  if  it  is  placed  in  a  cylinder  of  red 
glass  the  bubbles  will  be  evolved  almost  as  quickly  as  in  ordinary 
sunlight,  whereas  in  a  cylinder  of  blue  glass,  which  intercepts 
and  cuts  off  the  red  and  orange  rays,  the  evolution  of  bubbles 
Light  absorb-  almost  ceases.  If  the  light  which  has  passed  through  a 
e<*  by  chlor°  solution  of  chlorophyll  is  decomposed  by  a  prism,  a  dark  band 
will  be  found  in  the  red  and  orange  indicating  that  these 
rays  have  been  absorbed  by  the  chlorophyll.  The,  latter, 
the  presence  of  which  is  essential  for  the  assimilation  of  carbon 
in  the  higher  plants,  is  thus  seen  to  absorb  energy  from  the 
sun  in  the  shape  of  certain  rays  of  light.  Assimilation  is, 
however,  a  vital  function  performed  by  the  living  protoplasm 
which,  in  some  way  which  is  not  yet  understood,  utilises 
the  energy  absorbed  by  the  chlorophyll  in  decomposing  the 
Formation  of  wa-ter  and  carbon  dioxide  and  in  building  up  from  them  a  carbo- 
Starch.  hydrate,  such  as  starch.  The  chlorophyll  is  thus  merely 
a  part  of  the  apparatus  which  the  protoplasm  employs  in  this 
work  of  assimilation.  As  might  be  expected,  it 'will  be  found 
that  a  leaf  exposed  to  the  light  which  has  passed  through 
fi  solution  of  chlorophyll,  or  through  another  green  leaf,  is 
unable  to  assimilate  carbon  in  the  way  described,  the  effective 
light  rays  being  no  longer  available  for  the  process. 

Among  the  factors  which  directly  influence  assimilation 
are  a  suitable  supply  of  iron  and  carbon  dioxide,  the  tem- 
perature, and  light,  for  each  of  which  there  is  an  optimum 
degree  of  intensity.  It  has  been  already  noted  that  no  chloro- 
phyll is  formed  unless  the  necessary  quantity  of  iron  is  available 
and  that  without  carbon  dioxide  the  assimilation  of  carbon 
becomes  impossible.  The  amount  of  carbon  dioxide  in  the 
air  is  as  a  rule  very  small,  about  "04  %  and  provided 
that  the  light  is  sufficiently  intense  assimilation  may  be 
increased  by  increasing  the  amount  of  carbon  dioxide  in  the 
air  up  to  a  certain  extent.  More  than  10  %  car- 
bon dioxide,  however,  has  a  decidedly  injurious  effect 
and  many  plants  die  if  continually  exposed  to  air  containing 
as  little  as  4  %.  Temperature  has  a  marked  effect  on  assi- 
milation, as  well  as  on  all  other  vital  functions,  and  if  the 
temperature  is  below  a  certain  minimum  the  protoplasm 
forms  no  chlorophyll.  In  the  absence  of  light  assimilation 
ceases  at  once,  while  it  increases  in  proportion  to  the  intensity 
of  the  light  until  the  optimum  degree  of  intensity  is  reached, 
after  which  it  again  decreases.  If  the  intensity  of  the  light 
continues  to  increase  the  chlorophyll  is  ultimately  destroyed. 


103 

93.  The  starch  formed  in  the  leaves 
is  not  allowed  to  accumulate  there,  but  is  continually  being 
transported  to  other  parts  of  the  plant  where  it  is  required  for  use, 
or  storage.  This  transport  goes  on  both  day  and  night,  but  as, 
during  the  day,  the  amount  of  starch  manufactured  is  consider- 
ably in  excess  of  that  removed,  starch  does  accumulate  in  the 
leaves  to  a  certain  extent,  which ,  however,  is  usually  removed 
during  the  night.  Thus  leaves  as  a  rule  contain  more  starch  at 
nightfall  than  in  the  early  morning.  Only  those  substances  which 
are  soluble  are  able  to  traverse  the  cell-walls  and  they  can  thus 
be  transported  easily  and  quickly  from  cell  to  cell  in  a  plant. 
Starch,  however,  is  not  soluble  in  water  at  the  ordinary  tem- 
perature, and  hence,  for  the  purposes  of  transport  in  the  plant,  it 
is  converted  into  a  soluble  sugar,  such  as  glucose,  or  maltose,  and 
this  is  effected  by  means  of  the  substances  called  enzymes.  The 
latter  are  chemical  substances  formed  by  the  protoplasm 
during  metabolism  which  possess  the  power  of  more  or  less 
altering  various  compounds  without  being  themselves  effected 
and  many  are  able  to  convert  insoluble  into  soluble  substances. 
Diastase  is  a  general  term  for  a  group  of  enzymes  which  are 
frequently  found  in  plants,  some  of  which  can  turn  starch 
into  glucose,  or  fruit-sugar,  while  others  turn  it  into  maltose, 
or  cane-sugar.  In  addition  to  this  power  of  enabling  valuable 
food  materials  to  be  easily  transferred  from  place  to  place, 
enzymes  are  also  frequently  employed  by  plants  for  converting 
compounds  into  substances  which  can  be  readily  assimilated 
and  food  materials  stored  in  seeds  are  often  thus  made  available 
for  assimilation  by  the  young  plant, 

94.  All  plants,  like  animals,  K*sPiration- 
require  to  breathe,  and.  like  animals,  plants  absorb  oxygen 
and  give  off  carbon  dioxide,  heat  being  evolved  in  the 
process  which  is  known  as  respiration.  In  the  day  time 
respiration  is,  as  a  rule,  not  obvious  in  green  plants  growing 
in  the  sunlight  and  which  are  found  to  continually  enrich 
the  air  with  oxygen.  This  is  due  to  the  fact  that  the  quantity 
of  oxygen  evolved  in  carbon-assimilation  is  as  a  rule  much 
in  excess  of  that  required  for  respiration  and  that  nearly  all 
the  carbon  dioxide  produced  during  respiration  is  reassimilated 
in  the  cells  containing  the  green  chlorophyll.  If  a  green 
plant,  however,  is  placed  in  an  air-tight  jar  and  covered  with 
black  cloth,  or  otherwise  protected  from  the  sunlight,  it  will 
be  found  to  absorb  oxygen  from  the  air  in  the  jar  and  to  give 
off  an  equal  volume  of  carbon  dioxide.  Again,  if  a  quantity 
of  germinating  seeds  are  placed  in  an  air-tight  jar  and  after 


104 

some  hours  a  lighted  taper  is  introduced  into  the  jar,  the 
taper  may  be  extinguished  owing  to  the  exhaustion  of  the 
oxygen  and  the  accumulation  of  carbon  dioxide.  In  such  seeds 
which  contain  no  chlorophyll  there  is  no  carbon-assimilation, 
but  respiration,  nutrition,  and  growth  are  still  actively  going 
on  at  the  expense  of  the  food  materials  stored  in  the  seeds. 
The  fact  that  seeds  can  germinate  and  that  seedlings  can  develop 
and  attain  considerable  dimensions  in  the  dark,  when  no  carbon- 
assimilation  is  possible,  clearly  shows  that  nutrition  is  by  no 
means  the  same  thing  as  carbon-assimilation.  In  the  case 
of  such  seedlings  which  have  developed  in  the  dark,  nutri- 
tion and  growth  have  taken  place  at  the  expense  of  the  organic 
food  materials  contained  in  the  seed  and  the  dry  weight  of  such 
a  seedling  is  found  to  be  actually  less  than  that  of  the  seed 
from  which  it  sprang.  These  substances  have  in  fact  been  lost 
in  respiration,  the  process  which  has  supplied  the  energy 
necessary  for  nutrition  and  growth.  If  on  the  other  hand 
a  seedling  is  allowed  to  develop  normally  in  the  sunlight,  green 
leaves  are  produced,  carbon  is  assimilated,  and  the  dry  weight 
of  the  plant  is  soon  considerably  in  excess  of  that  of  the  seed 
from  which  it  sprang,  owing  to  the  fact  that  more  organic 
substance  has  been  manufactured  in  the  leaves  than  has  been 
required  for  respiration. 

Growth.  95.  During  the  growth  of  cellular 

plants  the  actual  enlargement  of  the  cells  is  effected  more  by 
increasing  the  actual  surface-area  of  the  cell- walls  than  by  in- 
creasing the  substance  of  the  protoplasm,  although  the  latter 
does  of  course  take  place.  This  enlargement  of  the  cell-walls 
can  only  take  place  provided  that  the  cells  are  in  a  state  of 
turgidity  and  hence  a  liberal  supply  of  water  is  essential  for  the 
growth  of  such  plants.  Like  other  vital  phenomena,  growth  is 
only  possible  within  a  definite  range  of  temperature.  When  the 
latter  falls  below  0°  C,  or  rises  above  40°  or  50°  C,  growth 
as  a  rule  becomes  impossible,  while  the  optimum  tem- 
perature for  growth  usually  lies  between  22°  and  37°  C. 

Light  retards  growth  and  plants  which  have  developed 
in  the  dark  as  a  rule  have  unusually  long  internodes,  while 
their  tissues  contain  more  water  and  their  cell-walls  are  thinner 
than  would  have  been  the  case  if  the  plants  had  been  exposed 
to  the  sunlight.  As  carbon-assimilation  is  impossible  in  the 

Ligkt. 6C  y  dark,  even  if  leaves  were  developed  they  would  be  unable 
to  perform  their  essential  functions  and  plants  which  have 
developed  in  the  dark  as  a  rule  form  no  green  chlorophyll. 
Moreover  in  such  cases  material  and  energy  are  not 

C?*' 


105 

wasted  on  the  construction  of  leaves  of  normal  shape  and  size 
and  the  leaves  which  are  formed  are  as  a  rule  unusually 
small  or  thin,  the  plant's  efforts  being  mainly  devoted  to 
increasing  the  length  of  the  stem,  with  the  object  of  enabling 
it  to  reach  the  sunlight  where  green  leaves  can  once  more 
assimilate  carbon  from  the  air. 

Provided  that  other  important  factors,  such  as  the  amount 
of  available  water  and  the  temperature,  remain  constant, 
plants,  as  might  be  expected,  usually  grow  more  at  night 
than  in  the  day. 

The  rays  of  light  which  have  most  effect  on  growth  are  the 
so-called  ' '  chemical  rays  ' :  which  are  situated  at  the  blue- 
violet  end  of  the  spectrum.  In  plants  which  have  developed 
in  a  red  light,  therefore,  the  retarding  effects  of  light  which 
prevent  excessive  growth  are  not  seen,  and  although  in  such 
plants  chlorophyll  is  formed,  and  carbon-assimilation  enables 
them  to  increase  in  weight,  their  growth  in  other  respects 
resembles  that  of  plants  grown  in  deep  shade. 

96.  Owing     to    its     remarkable  Plant  Move- 

power  of  irritability  the  living  protoplasm   of  plants  is,  as  it m 
were,  able    to  perceive  the  existence   of  external   factors  and 
to  regulate  its  actions  accordingly,  a  fact  which  is  often  made 
manifest  by  the  obvious  movements  of  plant  organs. 

Thus  the  protoplasm  in  the  roots  and  stems  of  the  higher 
plants  is  sensitive  to  the  force  of  gravity  and,  guided  by  the 
direction  in  which  this  force  acts,  it  is  able  to  direct  the  growth 
of  the  elongating  portions  of  these  organs  in  such  a  way  that 
they  assume  the  positions  best  suited  for  the  performance  of 
their  functions.  Thus  if  a  healthy  seedling  is  placed  with  its 
primary  shoot  and  root  in  a  horizontal  position,  the  growing 
portions  of  these  organs  will  be  found  to  curve  in  such  a  way 
that  the  tip  of  the  root  and  stem  respectively  point  directly 
downwards  towards  the  centre  of  the  earth  and  vertically  up- 
wards in  a  directly  contrary  direction,  growth  then  being 
continued  in  these  directions  without  further  curvature.  The 
curvatures  to  which  these  movements  are  due  can  only  take  place 
in  those  parts  of  the  root  and  stem  which  are  still  growing  and 
they  are  caused  by  the  unequal  growth  of  the  opposite  sides  of 
the  organ  concerned.  In  the  root  the  upper  side  grows  more 
rapidly  and  in  the  stem  the  opposite  occurs.  This  phenomenon 
of  movement  executed  in  response  to  the  force  of  gravity  is  called 
geotropism.  The  root  which  grows  towards  the  centre  of  the 
earth,  i.e.  in  the  direction  along  which  the  force  of  gravity  is 
acting,  is  said  to  be  positively  geotropic,  while  the  stem  which 


106 

grows  in  the   opposite  direction,  away  from  the  centre  of  the 
earth,  is  negatively  geotropic.     Those  parts  of  plants  which  grow 
horizontally,  i.e.  in   a  direction  at  right  angles  to  that  along 
which  gravity  acts,  are  said  to  be  dia-geotropic,  such  as  are 
often   the  primary  branches  of  the  stem  and  root.     It  is  also 
found  that  if  the  seedling,  after  being  placed  in  a  horizontal 
position,   is  continuously   and  evenly  rotated  so  that  no  side 
of  the  root  or  stem  is  allowed  to  remain  stationary,  no  geotropic 
curvature  takes  place,   inasmuch  as  no  sooner  does  any  part  of 
the  root  or  stem  receive  a  stimulus  to  grow  faster  than  the  side 
opposite  to  it  than  it  almost  immediately  receives  a  contrary 
stimulus  which  neutralises  it.      In  these  movements,  as  in  the 
case  of  those  of  the  leaves  of  Mimosa  pudica,  we  see  that  the 
response  of  the  plant  is  out  of  proportion  to  the  stimulus  applied 
and  the  result  is   quite  different  to  that  obtained  when  the 
same  stimulus  is  applied  to  lifeless  matter.     Thus  in  this  case 
the  shoot  moves  in  a  direction  exactly  opposite  to  that  followed 
by  a  lifeless  body  acted  on  by  gravity,  and  which  falls  down- 
wards towards  the  centre  of  the    earth  merely  by  its  own 
weight.     Moreover  a  primary  root  growing  geotropically  down- 
wards is  able  to  penetrate  and  force  its  way  into  mercury  which 
is  specifically  much  heavier  than  itself,  which  would  not  be  the 
case  if  the  root  was  composed  of  lifeless  substance  and  was  acted 
on  by  gravity.     The  irritability  of  a  plant  organ  may  vary  at 
different  periods  of  its  existence,  as  is  shown  by  the  fact  of  a 
dia-geotropic  lateral  root  becoming  positively  geotropic  and 
taking  the  place  of  a  tap-root  which  has  been  cut  off,  or  injured, 
and  the  same  thing   occurs  when  a  lateral  branch   replaces  a 
damaged  leading  shoot.      This  question  of  the  movements  of 
plant  organs  is  complicated   by  the  fact  that  in  nature  one 
and  the  same  organ  is  frequently  exposed  to  the  influence  of 
several  factors,  or  stimuli,  to  each  one  of  which  individually  it 
may  be  able  to  respond  in  a  particular  way.     Thus  the  move- 
ments of  such  an  organ  would  be  the  result  of  the  combined 
influence  of  all  the  stimuli  affecting  it  at  one  and  the  same  time. 
Hydrotro-      Thus  roots  are  also  found  to  be  sensitive  to  the  presence  of  water 
and  are  able  to   regulate   their   growth  with  reference  to  the 
source  of  moisture,   i.e.  they   are  hydrotropic.     If  a   primary 
root  is  growing  in  soil  with  plenty  of  moisture  available  on  all 
sides,    it    will    proceed    downwards,   following  the  direction 
suggested  by  gravity,  but  if  it  is  situated  in  a  very  dry  soil,  in 
which  moist  patches  occasionally  occur,  the  root  will  be  found 
to  grow  towards  the  moist  areas,  it  thus  being  positively  hy- 
drotropic, irrespective  of  whether  these  areas  lie  directly  below  it, 


J07 

or  in  any  other  position,  the  hydrotropism  of  the  root  in  this  case  Heiiotropism. 
overpowering  its  geotropism.  The  aerial  roots  of  some  climbing 
stems  are  also  found  to  be  sensitive  to  the  direction  of  the  rays 
of  light  which  fall  upon  them  and  they  respond  by  growing 
away  from  the  source  of  light,  i.e.  they  are  negatively 
heliotropic.  Such  roots  being  also  positively  hydro  tropic  and 
very  slightly,  if  at  all,  positively  geotropic,  turn  towards  the 
moist  dark  crevices  of  their  support  and  not  downwards 
towards  the  centre  of  the  earth. 

The  primary  stem,  in  addition  to  being  negatively  geotropic, 
is  also  sensitive  to  the  direction  of  the  rays  of  light  falling  on  it 
and  responds  by  turning  its  apex  towards  the  source  of  light,  i.e. 
it  is  positively  heliotropic.  In  this  case  heliotropism  is  usually 
more  powerful  than  geotropism  and  if  the  stem  is  exposed  to  il- 
lumination on  one  side  it  will  grow  obliquely  towards  the  light 
rather  than  vertically  upwards.  Most  leaves  are  also  sensitive 
both  to  gravity  and  also  to  light,  they  being  both  dia-geotropic 
and  dia-heliotropic,  but  their  position  in  nature  depends  mainly 
on  the  direction  of  the  light  rays.  As  a  rule  they  place  them- 
selves at  right  angles  to  the  incident  rays  of  light  with  their  upper 
surface  turned  directly  towards  the  source  of  light.  Plate  XIII 
for  instance  shows  how  the  leaves  of  Coriaria  nepalensis  secure 
this  position  on  erect  and  horizontal  branches,  respectively.  In  sleep- 
addition  to  this  sensitiveness  to  the  direction  of  the  light  rays,  themc 
floral  envelopes  and  also  the  foliage  leaves  of  many  plants  are 
found  to  be  sensitive  to  variations  in  the  intensity  of  light  and 
also  to  variations  of  temperature.  Thus,  with  a  rising  tempera- 
ture and  light  becoming  more  intense,  many  flowers  open,  while 
they  close  with  a  fall  of  temperature  and  a  diminished  intensity 
of  light.  The  so-called  sleep-movements  of  some  foliage  leaves 
are  due  to  the  same  cause,  the  leaves  in  this  case  assuming  a 
different  position  at  niotfit  to  that  taken  up  in  the  day,  and  as  the 
temperature  falls,  arid  the  intensity  of  the  light  decreases,  in  the 
evening,  these  leaves,  or  leaflets,  are  found  to  close  together  and 
to  usuallyexpose  only  their  edges  to  the  zenith.  In  some  cases  it 
appears  that  these  movements  are  of  service  to  the  plant  in  pre- 
venting excessive  loss  of  heat  by  radiation.  The  so-called  night- 
position  may  also  be  taken  up  in  some  cases  in  the  daytime, 
owing  to  the  light  being  too  intense,  or  the  temperature  too  great. 
Many  tendrils  are  found  to  be  sensitive  to  contact  with 
rough  solid  bodies,  as  a  result  of  which  the  side,  in  contact  with 
the  substance  grows  slower  than  the  opposite  side,  and  the 
tendril  accordingly  coils  around  the  obstructing  object,  (unless 
the  latter  is  too  large)  and  supports  the  plant  stem. 


108 


The  movements  hitherto  considered  are  those  which  are 
brought  about  chiefly  by  the  unequal  growth  of  opposite  sides 
of  the  organ  concerned  and  hence  they  are  only  possible  in  parts 
which  are  still  growing.  Those  organs  which  have  completed 
their  growth  therefore  have  become  fixed  in  position,  and  this 
position  isthe  one  which  is  onthe  whole  the  best  suited  for  the 
performance  of  the  functions  of  each  individual  organ,  having 
regard  to  the  conditions  of  its  environment.  Thus  a  leaf  which 
is  no  longer  able  to  move  according  as  the  direction  of  the  light 
rays  falling  on  it  changes,  takes  up  a  fixed  position  in  which 
it  receives  the  full  benefit  of  the  greatest  quantity  of  the 
most  suitable  light  rays. 

Movements  In  addition,  however,  to  the  movements  which  are  brought 
a^>out  by  a  difference  in  the  rate  of  growth  of  different  parts  of 
organs,  plants  are  also  in  some  cases  able  to  effect  the  movement 
of  organs  after  the  latter  have  completed  their  growth,  in  res- 
ponse to  contact,  light,  temperature,  and  other  stimuli.  In  these 
cases  the  movement  is  brought  about  by  an  alteration  in  the 
turgidity  of  the  cells  on  opposite,  sides  of  the  organ  concerned. 
In  foliage  leaves  the  pulvinus  and  pulvinule  are  organs  especially 
adapted  for  effecting  movements  of  this  description,  which  can 
be  well  seen  in  the  leaves  of  Mimosa  pudica.  If  the  cells  on 
the  upper  side  of  the  pulvinus  are  turgid,  while  those  on  the 
lower  side  are  flaccid,  the  leaf  will  move  downwards,  and  vice 
versa,  the  leaf  in  each  case  turning  on  the  pulvinus  like  a  hinge. 
As  cells  are  only  capable  of  becoming  turgid  when  the  cell  walls 
are  elastic  and  not  rigid,  we  find  that  the  greater  part  of  the 
pulvinus  consists  of  parenchymatous  cells  with  non-lignified 
elastic  cell-walls,  the  vascular  strands  and  strengthening 
tissue  being  united  in  a  central  strand  where  it  offers  least- 
resistance  to  bending,  instead  of  being  distributed  nearer  the 
circumference. 

Eeproduc-  97.  There  are  two  modes  of  repro- 

SS'  AseS1  Auction  m  tne  Vegetable  Kingdom  known  as  the  Asexual,  or 
Methods.  Vegetative)  and  the  Sexual,  respectively.  In  the  former  a  portion 
of  the  protoplasm  of  the  parent  plant,  which  may  be  a  single 
cell,  or  a  multicellular  structure  such  as  a  bud,  separates  from 
the  parent  and,  either  at  once,  or  after  further  growth,  consti- 
tutes a  new  individual  plant.  In  sexual  reproduction  a  new 
individual  plant  can  only  be  produced  after  the  union  of  two 
pieces  of  protoplasm  has  taken  place  which  have  been  developed 
on  different  plants,  or  on  different  parts  of  the  same  plant. 
Several  plants  exist  which  propagate  themselves  only  by  the 
asexual  method,  others  are  found  to  do  so  only  by  the  sexual 


109 

method,  while  others  again  employ  both  methods.  In  the 
higher  plants  asexual  reproduction  is  often  brought  about  by 
means  of  stolons  and  runners,  as  in  the  Potato,  Strawberry, 
and  Rubus  lasiocarpus.  The  young  plants  developed  become 
separated  from,  and  independent  of,  their  parent  by  the  decay 
of  those  portions  of  the  stolons  and  runners  which  lie  between 
the  young  plants  and  their  parent.  The  same  thing  apparently 
often  occurs  in  the  case  of  root-suckers  which  become  indepen- 
dent by  the  decay  of  the  connecting  roots.  Asexual  reproduction 
is  also  effected  by  means  of  tubers,  bulbs,  corms,  bulbils,  and 
tuberous  roots. 

Sexual  reproduction  in  the  higher  plants  is  effected  by  means 
of  seeds.  From  the  fact  that  two  methods  of  reproduction  occur 
and  that  some  plants  which  depend  only  on  the  asexual  method, 
and  some  which  depend  only  on  the  sexual  method,  are  able  to 
exist  and  successfully  maintain  themselves  in  different  parts  of 
the  earth,  we  should  naturally  infer  that  each  method  possesses 
certain  advantages.  There  seems  to  be  no  doubt  that  this  is 
the  case  and  that,  under  the  conditions  of  existence  to  which 
plants  are  exposed  in  nature,  at  one  time  one  method,  and  at 
another  time  the  other  method,  may  prove  most  advantageous. 
Compared  with  the  structures  by  means  of  which  asexual  re- 
production is  effected,  such  as  bulbs,  corms  and  tubers,  seeds  are 
usually  smaller  in  size,  are  usually  produced  in  larger  numbers 
and  generally  possess  more  efficient  means  for  their  wide  dis- 
tribution. In  the  higher  plants,  therefore,  sexual  reproduction 
tends  to  result  in  the  establishment  of  a  large  number  of  young 
plants  scattered  at  a  considerable  distance  both  from  their 
parent  and  from  each  other.  To  a  great  extent,  therefore,  the 
roots  of  each  young  plant  are  able  to  develop  in  layers  of  soil 
which  have  not  been  exhausted  by  the  roots  of  its  parent,  and 
each  young  plant  is  to  a  great  extent  freed  from  a  severe  com- 
petition for  its  necessaries  of  life  with  other  young  plants  of  its 
own  species,  which  have  the  same  needs  and  requirements. 

With  asexual  reproduction,  on  the  other  hand,  it  is  found  that 
the  young  plants,  although  they  are  not  so  effectually  separated 
from  their  parent  and  each  other,  grow,  at  all  events  at  first,  more 
vigorously  and  attain  large  dimensions  quicker  than  do  those 
developed  from  seed.  This  power  of  vigorous  growth  during  early 
youth  may  often  be  of  vital  importance  for  the  existence  of  the 
plant,  i.e.  in  forest  areas  which  are  covered  with  a  heavy  growth 
of  grass  in  which  the  slow-growing  young  seedlings  of  forest 
trees  are  often  smothered  and  killed,  whereas  strong- growing 
root-suckers  may  be  able  to  successfully  establish  themselves. 


110 

As  regards  sexual  reproduction,  it  is  clear  that  in  an  herma- 
phrodite flower  it  is  possible  (1)  for  the  pistil  to  be  fertilised  with 
pollen  developed  in  the  same  flower,  in  which  case  the  flower 
is  said  to  be  self -fertilised  and  (2)  for  the  pistil  to  be  fertilised  by 
pollen  developed  in  another  flower  occurring  on  the  same  plant 
or  in  a  flower  belonging  to  a  separate  plant,  in  which  case  the 
flower  is  said  to  be  cross-fertilised.  Now,  a  young  plant 
arising  from  a  seed  resulting  from  the  second  kind  of 
cross-fertilisation  must  inherit  something  from  each  of  its 
parents,  and  it  therefore  tends  to  vary  and  to  exhibit 
characters  which  were  not  found  in  the  mother-plant.  It 
appears  that  this  tendency  to  differ  from  the  mother-plant  also 
exists  to  a  certain  extent  in  plants  arising  from  seed  produced  by 
self -fertilisation,  and  all  plants  raised  from  seed  tend  to  differ 
more  or  less  considerably  from  the  mother-plant.  In  asexual 
reproduction  this  is  not  the  case  and  the  young  plants  are 
invariably  found  to  resemble  their  mother-plant  very  closely. 
In  the  case  of  the  young  plants  arising  from  seed,  a  large  pro- 
portion of  which  frequently  find  themselves  at  a  long  distance 
from  their  mother  and  exposed  to  conditions  of  existence  more 
or  less  fundamentally  different  from  those  under  which  their 
mother-plant  developed,  the  possession  of  slightly  different 
characteristics  may  obviously  be  of  the  utmost  use  in  enabling 
them  to  develop  successfully.  On  the  other  hand  considerable 
variation  in  the  case  of  asexually  produced  offspring,  established 
in  the  immediate  neighbourhood  of  their  parent,  would  as  a  rule 
be  a  disadvantage. 

98.  As    regards  the  respective 

Seif-Fertiiisa-  advantages     of    cross     and     self-fertilisation,    it     has     been 
tion.  proved  by  actual  experiment   in  the  case  of  several  plants  in 

Europe  that  cross-fertilisation  results  in  the  production  of 
more  seeds,  which  in  their  turn  are  able  to  produce  more 
vigorous  plants,  than  is  the  case  with  self-fertilisation.  Even 
if  this  were  universally  the  case,  which  has  not  been  proved, 
it  must  always  be  remembered  that  cross-fertilisation  is  effected 
with  considerable  difficulty  and  is  therefore  far  more  uncertain 
than  self-fertilisation,  inasmuch  as  fewer  flowers  are  likely  to  be 
fertilised.  It  would  therefore  be  quite  possible  for  a  plant 
which  uniformly  produces  a  large  number  of  young  plants  as 
the  result  of  self -fertilisation  to  be  equally,  even  if  not  more, 
successful  in  the  struggle  for  existence,  in  comparison  with  a 
plant  which  habitually  produces  very  few  offspring  as  the  result 
of  cross-fertilisation,  even  if  the  offspring  of  the  latter  were 
slightly  more  vigorous  than  those  of  the  former.  It  is  in  fact 


Ill 

probable  that,  as  is  the  case  with  sexual  and  asexual  methods, 
according  to  the  conditions  under  which  plants  exist,  sometimes 
self-  fertilisation,  and  sometimes  cross-fertilisation,  may  be  most 
advantageous.  That  cross-fertilisation,  however,  is,  on  the 
whole,  most  advantageous  in  the  majority  of  cases,  is  indicated 
by  the  fact  that  a  very  large  number  of  contrivances  exist  which 
appear  to  aim  at  on  the  one  hand  the  prevention,  or  at  least  the 
postponement,  of  self  -fertilisation,  and  on  the  other  hand  at 
facilitating  cross  -fertilisation. 

99.   For    the     transference     of 

pollen  from  the  stamens  to  the  stigmas  the  principal  agencies  pilous,  Omi 
employed  by  plants  are  wind,  insects,  and  birds.  Plants  which  thophii 
are  pollinated  by  the  wind  are  called  anemophilous  and  usually 
have  small  inconspicuous  flowers,  with  no  brightly-coloured 
perianth,  and  with  no  sweet  nectar  or  attractive  odour.  A 
the  pollen  is  distributed  in  all  directions  by  the  wind,  it  is 
usually  produced  in  very  large  quantities  to  insure  some  of  it 
reaching  the  female  organs,  as  is  the  case  for  instance  in  Pines 
and  Firs.  In  Grasses  wind  -pollination  is  aided  by  the  fact  that 
the  large  anthers  are  versatile  and  swing  freely  in  the  wind, 
while  the  feathered  stigmas  offer  a  large  catchment  surface  for 
the  pollen.  Anemophilous  plants  are  also  frequently  gregarious, 
such  as  are  Pines  and  many  Grasses. 

Plants  which  are  pollinated  by  insects  are  termed  entomo- 
philous  and*  those  pollinated  by  birds  are  ornithophilous.  These 
as  a  rule  characterised  by  the  possession  of  conspicuously- 
coloured  floral  envelopes  with  often  also  nectar  and  attractive 
scents.  The  insects,  or  birds,  attracted  to  the  flowers  obtain 
nectar,  or  pollen,  or  both,  as  food  and  in  their  visits  carry  out 
for  the  plant  the  desired  transference  of  some  of  the  pollen 
to  the  stigmas. 


100.   The  chief  contrivances  by 

means  of  which  plants  endeavour  to  bring    about  cross-fertili-  ing  Cross 
sation  are:— 

(1)  The  separation  of  the  sexes.  —  This  is  effectually  secured 
in  dioecious  and  monoecious  plants  and  to  a  certain 
extent  also  in  polygamous  plants.  This  is  also 
effected  by  dichogamy,  i.e.  although  the  stamens 
and  pistil  occur  in  the  same  flower  they  mature  at 
different  times.  Flowers  in  which  anthers  mature 
fost  are  protandrous,  those  in  which  stigmas  mature 
first  and  become  ready  for  the  pollen  before  the 
stamens  dehisce  are  protogynous. 


112 

(2)  Heterostyly. — In  some  plants  it  is  found  that  all  the 
flowers  on  some  individuals  have  stamens  and  styles 
of  different  lengths  from  those  in  the  flowers  of  other 
individuals  of  the  same  species.  These  plants  and 
flowers  are  .said  to  be  heterostyled.  In  species  of 
Primula  the  flowers  are  dimorphic,  or  of  two  forms. 
The  flowers  on  some  individuals  have  short  styles 
with  the  anthers  situated  above  them  at  the  throat 
of  the  corolla,  and  those  on  other  plants  have  long 
styles  with  the  stigma  at  the  throat  of  the  corolla 
and  the  anthers  below  them,  deep  in  the  corolla 
tube.  Other  plants  exist  which  have  trimorphic 
flowers,  some  with  long,  some  with  short,  and 
others  with  medium-sized  styles,  with  the  anthers 
also  arranged  at  corresponding  heights  in  different 
flowers,  the  stigma  and  anthers  of  the  same  height 
never  occurring  on  one  and  the  same  plant. 
In  such  plants  it  is  found  that  full  fertility  is  only  obtained 
when  the  stigma  is  fertilised  by  the  pollen  taken 
from  anthers  standing  at  a  corresponding  level 
(i.e.  by  pollen  taken  from  the  flowers  of  another 
plant).  All  other  crosses  are  termed  illegitimate 
and  are  found  to  be  more  or  less  sterile,  i.e.  very  few 
seeds  are  produced  as  a  result  of  them  and  if  seeds 
are  produced  there  is  less  chance  of  healthy  plants 
developing  from  them  than  in  the  case  of  a  legiti- 
mate cross.  As  the  anthers  and  stigmas  standing 
at  the  same  level  in  different  flowers  must  come  in 
contact  with  the  same  part  of  the  bodies  of  the  insects 
which  visit  them,  cross-fertilisation  is  more  or  less 
frequently  insured,  while  self -fertilisation  is  not 
prevented.  Several  heterostyled  plants  occur  in 
India,  e.g.  Reinwardtia  trigyna  and  Woodfordia  ftori- 
bunda. 

(3)  Mechanical  arrangements. — As  illustrations  of  the  truly 
remarkable  mechanisms  found  in  various  flowers  two 
plants  have  been  selected,  viz.  Salvia  lanata  and 
Berberis  Lycium,  both  of  which  are  common  in 
Jaunsar.  For  illustrations  to  compare  with  the 
following  accounts  see  Plate  XIV. 
Fertilisation  101.  In  Salvia  lanata  the 

corolla  is  divided  into  two  obvious  lips,  the  lower  lip 
forming  a  convenient  landing-place  for  insects,  while  the 
upper,  helmet-shaped  lip  rises  above  it  like  a  hood.  Nectar 


113 

is  excreted  by  the  yellow  disc  at  the  base  of  the  ovary 
and  collects  in  the  corolla  tube.  The  flowers  are  much 
\isited  by  bees  and  a  bee  alighting  on  the  landing-stage 
and  wishing  to  get  at  the  nectar  has  to  pass  forward  under  the 
hooded  upper  lip  of  the  corolla  and  it  then  finds  the  entrance 
into  the  nectar-containing  tube  effectually  blocked  by  an  in- 
genious piece  of  mechanism  consisting  of  the  two  stamens  which 
is  shown  in  Fig.  1,  dissected  from  the  flower.  Each  stamen  is 
attached  to  the  corolla  by  a  short  filament  on  which  the  long 
connective  swings  like  a  lever  on  its  fulcrum.  The  upper  longer 
arm  of  the  connective  is  slender  and,  together  with  the  pollen- 
bearing  anther-lobe  which  it  carries  at  its  apex,  is  concealed  in 
the  hooded  upper  corolla  lip.  The  lower  thickened  connective 
arm  is  joined  to  that  of  the  other  stamen  which  is  placed  side  by 
side  with  it  in  the  flower,  the  metamorphosed  lower  anther- 
lobes  borne  at  their  extremities  coalescing  to  form  a  little  pouch- 
like  structure  which  effectually  blocks  the  entrance  to  the  corolla- 
tube,  like  a  trap-door.  See  Fig.  2.  The  anthers  dehisce  by  a 
longitudinal  slit  which  directly  faces  the  landing 'stage.  A  bee 
wishing  to  get  at  the  nectar  and  pushing  against  the  obstructing 
door  will  move  the  latter  upwards  and  backwards,  the  path  to 
the  nectar  thus  being  opened,  while  the  upper  connective  arms 
descend  towards  the  landing  stage  and  bring  their  pollen-covered 
anthers  down  on  the  bee's  back.  See  Fig.  3.  So  soon  as  the 
bee  withdraws,  the  stamens  swing  back  into  their  former  posi- 
tion. The  flowers  of  this  plant  are  protandrous,  i.e.  the  anthers 
mature  and  begin  to  shed  their  pollen  before  the  stigma  is 
ready  for  pollination."  In  a  young  flower  the  stigma  occupies 
the  position  shown  in  Fig.  4  (a),  the  stigmatic  surfaces  being  close 
together  and  well  out  of  the  way  of  an  insect  entering  the  flower. 
As  the  flower  gets  older  the  style  becomes  depressed  towards 
the  landing  stage  while  the  stigmatic  lobes  separate  and  become 
recurved,  Fig.  4  (6).  A  bee,  before  it  can  enter  the  flower  at  all, 
must  first  bring  its  back,  which  has  probably  been  dusted  with 
the  pollen  of  a  separate  younger  flower,  in  contact  with  the 
stigmatic  surface  and  cross-fertilisation  is  thus  effected. 

102.  In   the     case  of   Berberis  Fertilisation 

Lycium,  if  we  look  at  a  flower-bud  we  find  the  stamens  stand-  of  Flowers  of 
ing  erect  in  the  centre  of  the  flower  with  their  anthers  close  to 
the  stigma,  but  the  anther-valves  are  then  unopened  as  shown  in 
Fig.  5  (a),  Plate  XIV.  As  the  flower  expands  and  the  petals 
open  out  the  stamens  are  bent  back  with  them,  the  back  of  the 
filaments  being  closely  adpressed  to  the  upper  surface  of  the 
petals.  As  the  flower  opens,  also,  the  anthers  dehisce.  During 


114 

dehiscence  the  valves  remain  attached  only  at  one  point  at  the 
top  of  the  connective,  and,  hinged  on  this  point,  each  valve 
gradually  rises  up,  pulling  out,  as  it  does  so,  most  of  the  pollen, 
which  remains  attached  to  its  inner  surface.  Fig.  5  (b).  Each 
valve  after  rising  to  an  almost  horizontal  position  turns  its  inner 
surface  which  is  covered  with  pollen  inwards  towards  the  centre 
of  the  flower.  Fig.  5  (b)  and  (c). 

In  the  open  flower  these  pollen-covered  valves  are  protected 
from  the  rain  and  weather  by  means  of  the  shelter  afforded  by 
the  curled-in  tips  of  the  petals,  so  that  in  looking  into  the  ex- 
panded flower  the  valves  are  hidden  from  sight.  Fig.  6.  Look- 
ing again  at  the  open  flower  we  see  that  each  staminal  filament 
widens  out  at  its  base  so  as  to  come  in  contact  with  the  neigh- 
bouring filament  on  each  side  of  it.  Also  a  little  above  the  base 
of  the  filament,  on  each  side  of  it,  there  is  an  oval,  orange- 
coloured,  gland  which  excretes  nectar,  two  of  these  so-called 
nectaries  being  situated  near  the  base  of  each  petal  on  its  upper 
surface.  Figs.  6  and  7.  Each  staminal  filament  fits  in  closely 
between  two  nectaries  and,  as  already  noted,  lies  with  its  back 
closely  pressed  against  the  surface  of  the  petal.  The  nectar 
excreted  by  the  nectaries  flows  over  the  base  of  the  filaments 
and  forms  a  glistening  ring  around  the  base  of  the  ovary.  The 
bell-shaped  flowers  of  this  plant  are  either  horizontal,  or  they 
hang  downwards,  and  the  sepals  and  petals  effectually  prevent 
rain  from  entering  the  flower  and  damaging  the  nectar,  or 
pollen.  It  will  also  be  noted  that  as  the  petals  have  their  tips 
curled  in  to  protect  the  pollen  they  are  not  so  conspicuous  as 
they  would  be  if  they  were  fully  expanded,  but  to  compensate 
for  this  and  to  make  the  flower  more  noticeable,  the  three  inner 
sepals  are  much  enlarged  and  coloured  bright  yellow.  Fig.  6, 
If  now  the  base  of  one  of  the  filaments  in  the  expanded  flower  is 
touched  with  a  pointed  instrument  such  as  a  thin  pencil,  the 
stamen,  with  a  sudden  spring,  flies  up  from  its  sheltering  petal 
and  resumes  the  erect  position  it  occupied  in  the  unexpanded 
bud.  The  pollen- covered  valves  are  thus  brought  into  violent 
contact  with  the  pencil,  but  when  the  latter  is  removed  it  will 
be  seen  that  the  valves  are  not  near  enough  to  the  centre  of  the 
flower  to  actually  touch  the  stigma.  Fig.  8. 

A  bee,  searching  for  nectar  with  its  proboscis  and  touching 
the  irritable  base  of  one  of  the  filaments,  will  thus  cause  the 
stamen  to  spring  inwards  and  dust  with  pollen  that  side  of  its 
head  which  is  turned  away  from  the  stigma.  The  insect  being 
struck  by  the  stamen  will  often  fly  away  at  once  and  after 
visiting  a  few  flowers  its  head  will  be  dusted  all  over  with 


115 

pollen,  so  that,  on  visiting  any  other  flower,  it  must  inevitable 
rub  off  some  of  this  pollen  on  to  the  edge  of  the  stigmatic  disc 
and  may  thus  effect  cross-fertilisation.  The  flowers  of  this 
species  are  often  visited  by  small  ants  and  beetles  which 
consume  both  the  pollen  and  the  nectar.  Such  insects  only 
occasionally  spring  a  stamen  and  then,  as  they  do  not  leave  the 
flower  at  once,  but  continue  to  crawl  about  in  search  of  food, 
they  must  at  all  events  occasionally  bring  about  the  pollination 
of  the  stigma  with  the  flower's  own  pollen.  Moreover,  as  the 
pollen- covered  valves  of  a  sprung  stamen  are  practically  on  the 
same  level  as  the  stigmatic  surface,  it  seems  certain  that  some 
of  the  flower's  own  pollen  must,  sometimes  at  least,  reach  its 
stigma.  Thus  while  self -fertilisation  is  not  prevented,  more 
or  less  frequent  cross-fertilisation  is  insured. 

103.  It  must  always  be  re- 
membered  that,  although  cross-fertilisation  may  be,  and 
apparently  is  as  a  rule,  preferable  to  self -fertilisation,  it  is 
easy  for  a  plant  to,  as  it  were,  over-reach  itself,  if  in  endeavour- 
ing to  secure  the  former  it  makes  the  latter  absolutely 
impossible ;  for  in  the  event  of  cross-fertilisation  not  being 
effected,  which  is  in  many  cases  a  possible  contingency, 
no  seeds  at  all  would  be  formed  and  sexual  reproduction 
would  absolutely  fail.  Thus  many  plants  appear  to  find  it 
safest  to  at  all  events  insure  self -fertilisation  in  the  event  of  the 
failure  of  their  efforts  to  secure  cross-fertilisation.  This  is 
perhaps  most  clearly  seen  in  plants  which,  like  many  violets, 
possess  two  kinds  of  flowers,  (1)  large  conspicuous  flowers 
adapted  for  cross-fertilisation,  and  (2)  small,  inconspicuous 
closed  flowers  adapted  solely  for  self-fertilisation  (  —  cieisto- 
gamie flowers],  the  latter  insuring  a  supply  of  seed  in  the 
event  of  the  ordinary  flowers  not  being  fertilised. 

On  the  other  hand  there  are  certain  species  which  trust  en- 
tirely to  cross-fertilisation.  The  pollen  of  some  of  these  plants 
not  only  fails  to  effect  the  fertilisation  of  its  own  flower  but 
acts  as  a  poison,  and  if  applied  to  the  stigmas  results  in  the 
death  of  the  flower.  Finally  it  must  be  noted  that  if  a  stigma 
has  been  pollinated  with  pollen  from  its  own  flower,  self -fertilisa- 
tion may  be  prevented  and  cross-fertilisation  effected,  even  if 
the  stigma  is  pollinated  after  a  considerable  interval  with 
pollen  formed  in  another  flower  of  the  same  species,  owing  to 
the  latter  pollen  being  prepotent,  i.e.  able  to  effect  fertilisation 
quicker  than  the  flower's  own  pollen. 

104.  To  insure  the  wide  distribu-  insemina- 
tion of  seeds   various  devices   are   employed  by  plants.     In tiou  of  Seeds- 

i  % 


116 

some  cases  the  seed  or  fruit  is  provided  with  outgrowths 
which  tend  to  make  it  buoyant  and  adapted  for  dispersal 
by  wind,  such  as  the  hairs  on  the  seed  of  Holarrhena 
antidysenterica  and  the  wing  of  the  seed  of  Oroxylum  indicum 
(Plate  XII).  In  other  cases  the  seed  or  fruit  is  adapted 
for  conveyance  by  water,  such  as  are  the  seeds  of  Sissoo  which 
remain  enclosed  in  the  light  pod,  the  latter  serving  as  a  float. 
The  wing-like  outgrowths  also  of  many  seeds  and  fruits  appear 
to  serve  equally  well  for  transport  by  water  or  wind.  The 
winged  Sal  fruit  for  instance  is  often  carried  considerable  dis- 
tances by  water.  In  other  cases  the  seeds  are  distributed  by 
animals  which  eat  the  fruit  and  excrete  the  undigested  seeds. 
The  seeds  of  species  of  Zizyphus  appear  to  be  widely  distri- 
buted in  this  way  by  jackals.  The  seeds  of  species  of  Loran- 
ihus  are  largely  distributed  by  birds  which  eat  the  pulp  of  the 
fruit  and  rub  off  the  seeds  on  the  branches  when  wiping  their 
bills. 

Sometimes  the  seeds  are  forcibly  expelled  to  a  considerable 
distance  by  the  bursting  of  the  ripe  fruit,  as  in  the  case  of  the 
capsule  of  species  of  Impatiens. 

Finally,  fruits,  and  rarely  also  the  seeds,  may  be  provided 
with  spines,  hooks,  or  bristles,  by  means  of  which  they  adhere 
to  clothing,  the  hair  or  fur  of  animals,  etc.,  and  are  thus  carried 
long  distances. 


117 


PART  IV.-CLASSIFICATION. 

CHAPTER  I.— DEFINITIONS  AND  EXPLANATIONS. 

105.  The    fact    that    there    are  Necessity  for 

numerous  distinct  kinds  of  plants  has  been  recognised  from  Classifica- 
very  early  days,  and  the  uneducated  native  of  our  forests tlon' 
has  required  no  botanical  training  to  teach  him,  for  instance, 
that  the  Sal  tree  (Shorea  robusta]  is  distinct  from  the  Sam 
(Terminalia  tomentosa)  or  to  enable  him  to  recognise  these 
two  kinds,  or  species,  of  tree  in  the  forest.  In  early  days, 
when  the  total  number  of  plants  known  to  any  single 
individual  was  insignificant  the  want  of  an  elaborate  system 
of  classification  was  not  felt,  for  one  could  readily  acquire 
and  retain  in  the  memory  an  intimate  knowledge  of  the 
characteristics  of  the  known  plants,  which  explains  how  it  is 
that  the  aboriginal  is  often  found  to  be  well  acquainted  with, 
and  able  to  recognise  at  sight,  the  majority  of  the  plants 
growing  in  the  jungle  near  his  village.  It,  however,  soon 
became  evident  that,  if  any  individual  desired  to  con- 
siderably extend  his  knowledge  of  plants  and  to  be  able 
to  become  quickly  acquainted  with  those  of  foreign  countries, 
some  system  of  classification  was  essential,  under  which  all 
plants  could  be  shortly  and  concisely  described  (only  essential 
points  of  difference  being  noted)  and  grouped  in  such  a  way  as 
would  enable  one  to  quickly  refer  to  its  group  any  plant,  where 
its  brief  description  and  correct  name  would  then  be  obtained. 
Once  the  botanical  name  of  a  plant  has  been  ascertained,  all 
the  information  that  has  ever  been  placed  on  record  regarding 
it  can  then  be  obtained  by  referring  to  the  necessary  books. 
Botanical  classification  thus  aims  at  placing  all  plants  in  groups 
under  groups,  according  to  their  resemblances,  the  smallest  groups 
being  combined  to  form  larger  groups  and  the  latter  again 
formed  into  still  larger  groups  and  so  on,  the  plants  included 
in  the  smallest  of  these  groups  being  all  very  much  alike, 
while  those  included  in  the  largest  groups  have  fewer  points  of 
resemblance.  The  peculiarities  which  enable  us  to  distinguish 
one  plant  from  another  are  called  characters, 

In  a  Flora,  therefore,  which  is  a  book  containing  the  descrip- 
tions of  all  the  plants  of  any  given  country  or  district,  the 
essential  characters  which  enable  us  to  distinguish  the  plants 


iis 

in  any  one  of  the  largest  groups  from  those  in  all  the  others  are 
first  given  ;  then  under  each  of  the  largest  groups  follow  the 
essential  characters  distinguishing  the  next  lower  plant  groups 
from  one  another  and  so  on,  so  that,  if  we  compare  the  characters 
of  any  given  plant  with  those  given  in  the  Flora,  we  are  led 
quickly  on  from  group  to  group  until  ultimately  we  arrive  at  the 
smallest  group  and  the  correct  name  of  our  plant,  without  wast- 
ing time  or  getting  confused  in  reading  and  comparing  long 
descriptions  of  unimportant  characters. 

Natural  106.  It  has  been  recognised  from 

a  very  earlyPeriod  tnat  "  h^e  begets  like,"  that,  with  plants 
as  with  animals,  the  offspring  resemble  their  parents,  and  hence, 

tion.  also,  it  has  been  accepted  as  a  fact  from  early  times  that  all 

organisms  which  closely  resemble  each  other  must  be  nearly 
related. 

When  botanical  classification  was  commenced,  it  became 
necessary,  first,  to  analyse  and  clearly  define  the  essential  points 
of  difference  between  individual  plants,  i.e.  their  characters, 
and,  secondly,  to  group  together  those  plants  with  similar 
characters.  It  was  then  found  that,  if  only  one  character  was 
relied  on,  plants  were  often  placed  in  entirely  different  groups, 
although,  as  regards  all  their  other  characters,  they  were  very 
much  alike  and  were  in  consequence  held  to  be  very  closely 
related.  A  system  of  classification,  therefore,  which  only  relied 
on  single  characters,  regardless  as  to  whether,  or  not,  the  plants 
which  were  most  nearly  related  were  thus  kept  together,  was 
known  as  an  Artificial  System,  whereas  a  system,  according  to 
which  all  characters  were  taken  into  account  and  which  resulted 
in  placing  the  most  closely  allied  plants  in  the  same  group, 
was  known  as  the  Natural  System. 

The  best  known  example  of  an  artificial  system  is  the 
so-called  "sexual  system"  of  Linnaeus.  Under  this  system 
only  the  characters  of  the  stamens  and  the  distribution  of  the 
sexual  organs  in  different  flowers  were  taken  into  consideration, 
and  all  known  plants  were  accordingly  classified  under  24  groups. 
As  an  instance  of  the  artificial  nature  of  this  scheme  of  classifica- 
tion it  may  be  noted  that  Class  XXI  Moncecia  includes  two  such 
distantly  related  plants  as  the  Maize  (Zea  Mays),  a  Monocoty- 
ledon, and  the  Oak  (Quercus),  a  Dicotyledon. 

A  good  artificial  system  which  does  not  concern  itself  with 
the  fact  as  to  whether  nearly  related  plants  are,  or  are  not,  kept 
together  in  the  groups  which  it  defines,  but  which  merely  refers 
to  isolated  and  easily  recognised  characters,  often  enatyes  us  to 


119 

quickly  assign  any  given  plant  to  the  group  where  its  description 
is  to  be  found,  and  thus  in  many  cases  facilitates  the  work  of 
identification,  for  which  reason  it  is  still  utilised  to  some  extent 
in  modern  Floras  in  the  compilation  of  keys  to  the  larger 
natural  groups. 

Striking  characters,  however,  are  often  inconstant  and  by 
themselves  are  not  always  trustworthy  guides  in  classification, 
while,  apart  from  the  fact  that  reliance  on  a  single  character 
is  apt  to  lead  us  wrong,  it  must  be  remembered  that,  if  we 
depend  on  one  or  two  characters  only,  we  are  quite  unable  to 
classify  any  plant  unless  the  individual  in  question  exhibits 
those  characters  at  the  particular  moment  of  its  life- history 
when  we  happen  to  see  it.  If,  for  instance,  we  rely  only  on 
the  number  of  cotyledons  for  the  definition  of  primary  groups 
and  the  cotyledons  are  not  to  be  found  on  our  plant,  we  are 
at  once  brought  to  a  standstill  and  are  unable  to  proceed  with 
the  identification.  We  are  then  driven  to  the  conclusion  that, 
as  a  rule,  in  botanical  classification,  an  aggregate  of  characters 
is  of  more  value  than  one  or  two  striking  points  of  difference, 
both  in  indicating  general  resemblance,  and  therefore  relation- 
ship, and  in  the  work  of  identification.  At  the  present  day, 
botanists  endeavour  to  make  all  permanent  classification  as 
natural  as  possible  throughout,  and  hence,  before  deciding  to 
which  group  any  plant  belongs,  all  characters  are  as  far  as 
possible  taken  into  consideration  and  we  then  decide  which 
group  it,  on  the  whole,  appears  to  resemble  most,  the  plants 
in  any  one  natural  group  resembling,  in  the  sum  of  theiv 
characters,  each  other  more  than  any  other  plant. 

107.  Although,    as    has     been  The  Unit  of 

pointed  out  above,  it  has  been  recognised  from  a  remote  9assifica' 
period  that  the  offspring  of  any  given  plant  closely  resem- 
bles its  parents,  yet  close  observation  showed  that,  in 
reality,  no  two  individual  plants  were  ever  exactly  alike.  If 
then  we  accept  similarity  of  form  and  structure  alone  as 
the  basis  of  classification,  we  must  adopt  the  individual  plant 
as  the  unit.  Since,  however,  every  individual  plant  has 
only  a  limited  period  of  existence  and  sooner  or  later  dies, 
such  a  procedure  would  lead  to  no  practical  results.  With 
the  individual  as  the  unit  we  should,  for  instance,  require  an 
unlimited  number  of  names  for  all  the  individuals  which 
now  exist  or  which  will  arise  in  the  future  from  existing  forms 
by  reproduction  ;  we  should  only  be  able  to  catalogue  and  de- 
scribe a  very  small  number  of  all  the  individuals  ever  existing 
on  the  earth  at  one  and  the  same  time  ;  while  the  descriptions, 


120 

after  the  death  of  the  individuals  to  which  they  refer,  would  be 
useless  for  the  purpose  of  identifying  living  plants  and  would, 
therefore,  be  of  little  value  to  us  or  our  successors.     It  has, 
however,  long  been  recognised    that    although  the  immediate 
offspring   of   any  individual  usually  differ  slightly  from  their 
parents  and  each  other,  they,  on  the  whole,  invariably  resemble 
their  parents  and  each  other  very  closely,  and  this  undoubted 
fact  that,    within    certain    limits,    all    organisms    breed    true, 
affords  the  only  basis  for  a  natural  history  classification  which 
shall   be    of    practical    value.     By    discovering    within    what 
limits  each  different  kind  of  plant  breeds  true,  i.e.  *by  determin- 
ing  which    characters    are    always    transmitted    truly    to  its 
immediate  offspring,  we  are  able  to  obtain  a  unit  which,  so  far 
as  we   can  see,  is  permanent,  the  marks  which  distinguish  the 
individuals  belonging    to  this  unit  from  all  other  plants  being 
always    transmitted  unchanged     from    parents    to    offspring 
through     successive     generations.     Such   a  unit    can    conse- 
quently be  recognised  and   studied  by  our'  successors,  while,  by 
only  giving  a  separate   name  to  each    such  unit  instead  of  to 
each   individual,     the    number    of   names  which    will   be  re- 
quired is  enormously  reduced.     Such  considerations  have  led  to 
the  selection  of  the  so-called  species  as  the  unit  of  classification. 
It  will  be  seen  also  that  the  morphological  characters  which 
are    most  important  in    classification   are    those    which  are 
always  transmitted  unchanged  from  parents  to  their    offspring 
and  which  therefore  indicate  genetic  relationship. 

Species,  (  108    With  these  preliminary  remarks 

Variety,0*  '  tne  following  definitions  are  now  given  :— 

Genus.  DEFINITION  1. — A  SPECIES  is  the  smallest  group  of  plants 

existing  wild  in  nature  which  can  be  readily  distin- 
guished from  all  other  groups  owing  to  the  fact  that  the 
individuals  composing  it  all  possess  in  common  certain 
well  marked  characters  (=•  specific  characters)  by 
which  they  can  be  distinguished  from  all  other  plants. 
The  individuals  also  which  compose  the  species  are, 
when  developed  normally  in  a  state  of  nature,  always 
able  to  transmit  their  specific  characters  unchanged  to 
the  majority  of  their  immediate  offspring. 

DEFINITION  2. — A  SUB-SPECIES  is  a  group  essentially 
similar  to  a  species  but  subordinate  to  it.  The 
differences  separating  any  two  individuals  belonging  to 
different  sub-species  not  being  so  great  as  those  which 
separate  individuals  belonging  to  different  species. 


121 

DEFINITION  3. — A  VAKIETY  is  a  group  of  plants  subordinate 

to  a  species.  The  differences  between  any  two  varieties 

{  of  the  same  species  are  not  constant,  i.e.  they  are  not 

always  transmitted  unchanged  from  the  parent  to  the 

majority  of  its  immediate  offspring. 

DEFINITION  4. — A  RACE  is  a  variety  of  considerable  fixity. 
The  characters  distinguishing  the  individuals  which 
compose  it  from  those  constituting  the  rest  of  the 
species  are  frequently  (e.g.  in  certain  localities  or 
under  certain  conditions  of  existence),  but  not  always, 
transmitted  from  the  parent  to  the  majority  of  its 
immediate  offspring. 

DEFINITION  5. — A  number  of  species  which  closely  resem- 
ble one  another  with  respect  to  their  important 
morphological  characters  are  combined  into  a  higher 
group  termed  a  GENUS. 

Every  species  in  the  same  genus  bears  the  same  name,  Nomencla- 
known  as  the  generic  name,  while,  to  distinguish  the  various ture- 
species  included  in  a  genus,  each  one  is  given  an  additional 
name  known  as  the  specific  name.  The  name  of  every  species, 
therefore,  consists  of  two  words.  Thus  all  plants  belonging  to 
the  oak  genus  bear  the  common  generic  name  of  Quercus, 
while  the  various  species  are  distinguished  by  their  specific 
names,  thus  we  have  Quercus  incana,  Quercus  glauca,  and  so 
on.  The  same  specific  name  cannot  be  used  for  more  than  one 
species  in  the  same  genus,  but  may,  of  course,  be  used  in  another 
genus. 

In  the  majority  of  modern  Floras  no  distinction  is  drawn 
between  sub-species,  races  and  varieties,  all  sub-divisions  of 
the  species  being  indiscriminately  termed  varieties.  Of  such 
varieties  those  which  are  held  to  be  most  important  are 
given  separate  names,  the  varietal  name  following  the  specific 
name,  thus  Cedrus  Libani  var.  Deodara.  Unimportant  varieties 
are  merely  noted  below  the  description  of  the  species  in  which 
they  are  included  and  are  designated  by  numbers  or  letters. 
As  different  botanists  have  sometimes  given  different  names 
to  one  and  the  same  plant  and  as  different  plants  have  some- 
times received  the  same  name,  it  is  necessary,  in  order  to  avoid 
confusion,  to  write  after  the  name  of  the  plant  the  name,  in 
full  or  abbreviated,  of  the  author  who  first  gave  it  that  name : 
thus  Rhus  Wallichii  Hook  f.  means  that  J.  D.  Hooker  was  the 
botanist  who  first  gave  the  plant  this  name.  A  name  of  a 
plant  which  has  been  superseded  by  another,  owing  to 
its  having  been  considered  incorrect  for  some  reason,  is 


Natural 
Order,  and 
Class. 


Principal 
Natural 
Groups 
of  Plants. 


122 

known  as  a  synonym.  Thus  Rhus  Wallichii  Hook  f.  syn  R. 
vernicifera  Brandis  means  that  the  plant,  the  correct  name  of 
which  is  considered  to  be  R.  Wallichii  given  by  J.  D. 
Hooker,  is  the  same  as  that  which  had  been  also  named 
R.  vernicifera  by  Sir  Dietrich  Brandis.  The  author,  whose 
name,  or  abbreviated  name,  is  attached  to  each  species,  is  the 
person  who  first  put  the  plant  into  the  correct  genus  and  who 
therefore  is  not  necessarily  the  person  who  was  the  first  to 
describe  the  plant  or  to  give  it  a  name. 

Just  as  species  are  combined  into  genera,  so  are  a  number 
of  genera  which  closely  resemble  one  another  in  their  impor- 
tant morphological  characters  combined  into  larger  groups 
known  as  NATURAL  ORDERS,  and  the  latter  are  similarly 
combined  into  still  higher  groups  termed  CLASSES. 

109.  The     principal     groups    of 

plants  therefore  which  are  the  most  frequently  met  with,  and 
which  form  the  groundwork  of  all  systems  of  classification 
are  as  follows,  commencing  with  the  largest : — 

Class 
Order 
Genus 
Species 
Variety 

These  groups,  however,  are  not  sufficient  to  indicate  all 
the  degrees  of  resemblance  which  are  found  to  exist,  and  hence 
they  are  often  sub -divided,  while  additional  groups  are  also, 
if  necessary,  created.  The  following  is  a  list  of  the  names 
most  frequently  employed  for  such  sub-divisions  of  the 
Vegetable  Kingdom,  given  in  sequence  commencing  with  the 
largest.  It  must,  however,  be  noted  that  several  of  these 
terms  have  been  applied  to  groups  of  somewhat  different  value 
by  different  botanists  :•— 

Sub -Kingdom 
Division 

Class  . 

Sub-class 
Series 
Cohort 
Order* 
Sub -order 


*  According  to  the  Rules  adopted  by  the  International  Botanical  Congress, 
held  at  Vienna  in  1905,  the  groups  usually  known  hitherto  in  English  literature 
as  cohort  and  order  respectively  should  be  designated  order  and  family. 


Tribe 
Genus 
Section 
Species 
Sub-species 
Race 
Variety. 

The  main  object  of  classification  is  to  enable  us  to  rapidly 
become  acquainted  with  the  principal  groups  of  plants 
indigenous  in  various  countries,  and  it  must  be  remembered 
that  the  above  definitions  refer  to  groups  of  plants  as  they 
exist  growing  wild  in  a  state  of  nature,  and  that  there  are  many 
plants  which,  in  the  garden,  breed  true  and  give  the  impression 
of  being  constant  forms  and  of  constituting  good  species  which 
are  not  found  as  wild  species  in  a  natural  state,  owing  to 
their  being  unable  to  survive  in  the  struggle  for  existence 
or  to  other  causes.  Hence,  if  terms  similar  to  those  given 
above  are  used  for  analogous  groups  of  cultivated  plants,  it 
should  invariably  be  stated  that  plants  under  cultivation 
are  referred  to. 

110.  Proof  that  a  particular  group  Practical 

of  individuals  has  descended    from  another  group    is    not  in  Determina- 
itself    sufficient   reason    for    combining  them  together  as  one  ^ of  the 
species,  for,  in  the  course  of  time,  the  intermediate  forms  which  Groups, 
once  united  the  two  groups  may  have  disappeared,    causing 
the  two  groups  to  occur  in  nature  as  distinct  species  separ- 
ated by  well-marked  and  constant  differences. 

A  species  is  a  group  of  plants  which  actually  exists  in 
nature,  the  recognition  and  correct  definition  of  which  are 
independent  of  the  way  in  which  the  group  originated. 

With  a  few  exceptions,  the  individuals  of  one  and  the 
same  species  cross  readily  and  produce  fertile  offspring  while 
individuals  belonging  to  distinct  species  very  frequently  do 
not  do  so.  At  the  same  time  fertility  cannot  be  accepted  as 
an  infallible  criterion  of  species,  for  illegitimate  unions  between 
the  different  forms  of  flowers  which  occur  in  one  and  the  same 
species,  in  plants  with  dimorphic  and  trimorphic  flowers, 
produce  very  little  fertile  seed  and  the  plants  raised  from  such 
seed  are  sterile  inter  se,  just  as  is  frequently  the  case  with 
the  illegitimate  unions  between  distinct  species.  The  male  and 
female  forms  of  some  organisms  differ  widely  from  each  other 
in  many  important  characters,  while  an  organism  may  also 
exhibit  an  entirely  different  appearance  at  different  periods 


124 

of  its  life-history.  However  great  such  differences  might  "be 
they  would  obviously  not  justify  our  classing  different  forms 
of  one  and  the  same  organism  as  distinct  species.  When 
defining  species,  therefore,  care  must  be  taken  not  only  to 
observe  what  characters  are  likely  to  vary  in  the  immediate 
offspring  of  one  and  the  same  individual  plant,  but  also  to 
note  how  the  characters  of  the  same  plant  vary  at  different 
periods  of  its  life-history.  An  imperfect  knowledge  of  life- 
histories  has  led  to  mistakes  being  made  in  the  classification 
of  some  fungi,  different  stages  in  the  life-history  of  one  and 
the  same  individual  having  been  sometimes  described  as  entirely 
distinct  species.  The  words  "  always  able  to  transmit"  in 
definition  1,  while  indicating  constancy  under  varying  condi- 
tions of  existence,  also  imply  that,  while  plants  capable 
of  both  sexual  and  asexual  reproduction  cannot  constitute  a 
species  if  they  only  transmit  their  essential  characters  truly  by 
asexual  reproduction,  organisms  only  capable  of  asexual  repro- 
duction are  not  thereby  precluded  from  forming  a  true  species. 

The  characters  which  are  used  to  define  species  must 
not  only  be  constant,  but,  also,  in  order  to  facilitate  identi- 
fication, they  should,  as  far  as  possible,  be  such  as  can  be 
easily  recognised,  and  further,  to  be  of  use  in  written  descrip- 
tions, they  must  be  such  as  can  be  easily  described  in  words. 

Among  the  higher  flowering  plants,  with  which  we  are 
chiefly  concerned,  characters  of  the  floral  structure  are  as  a 
rule  most  constant  and  important  and,  hence,  special  atten- 
tion is  paid  to  them  in  most  Floras,  but  these,  owing  to  their 
minuteness,  are  often  difficult  to  recognise,  while  flowers  are 
also  only  available  at  certain  seasons ;  just  as,  however,  the 
savage,  who  pays  no  attention  to  such  minute  characters, 
is  still  able  to  recognise  the  different  trees  in  the  forest,  so  can 
also  the  expert  forester,  by  utilising  characters  referring  to 
the  buds,  leaf-scars,  kind  of  bark  on  young  and  old  stems, 
the  method  of  branching,  the  colour  of  the  foliage  at  different 
seasons,  and  others,  which,  as  a  rule,  are  not  included  in 
botanical  books,  owing  to  their  not  being  easily  described  in 
words,  to  their  not  being  easily  recognisable  in  herbarium 
specimens,  or  to  their  not  being  sufficiently  constant  over 
large  areas  and  in  different  localities.  For  this  reason,  also, 
a  botanical  classification  based  on  a  single  character  is  not 
necessarily  artificial,  for  such  a  character  may  be  correlated 
with  several  others,  the  latter  having  been  omitted  from  the 
scheme  of  classification  owing  to  their  being  difficult  to  de- 
scribe or  to  distinguish.  ^ 


125 

In  order  to  determine  whether  particular  groups  of  plants 
found  wild  are  to  be  considered  as  species,  sub-species,  races, 
or  varieties,  a  knowledge  of  the  life-history,  as  well  as  of  the 
appearance  of  the  seasonal  forms  of  individual  plants,  and  of 
the  kind  and  amount  of  variation  which  may  occur  among  the 
immediate  offspring  of  one  and  the  same  individual,  growing 
under  different  conditions,  is  essential.  It  is,  however,  as  a 
rule,  impossible  to  ascertain  with  certainty  the  parent  of  a 
plant  found  wild,  and  very  few  species  have  been  experiment- 
ally cultivated  with  the  object  of  recording  the  variation 
exhibited  by  them.  The  systematic  botanist,  therefore,  must 
rely  for  his  determinations  mainly  on  morphological  char- 
acters, coupled  with  his  knowledge  of  the  development  and 
variation  of  the  few  forms  which  have  be*en  studied. 

His   conclusions    are,  consequently,   liable  to  error  and   to 
correction  in  the  light  of  subsequent  research,  while,  in  the 
present  state  of  our  knowledge,  there  is  obviously  room  for 
considerable  difference  of  opinion  as  to  the  kind  of  characters 
and  the  amount  of  variation  which  should  be  held  sufficiently 
important  to  constitute  an  inter-specific  gap,  in  consequence 
of    which  plants  which  are  regarded  by  some  botanists  as 
distinct   species   are    considered   to   be   varieties    by   others. 
The  discovery  of  intermediate  forms,   also,   sometimes  leads 
to  two  groups  of  plants,  hitherto  considered  distinct,  being 
combined  as  one  species,  it  being  possible,  in  such  a  case,  to 
arrange  a  complete  series  of  individuals  between  which  there  are 
no  marked  differences,  the  slight  differences  noticed  being  such 
as  might  be  expected  to  occur  in  the  offspring  of  one  individual. 
In  defining  the  limits  of  the  larger  natural  groups  and 
in  deciding   which   species   are   to  be   included   in   the   same 
genus,    which    genera   in   the    same    order,    and  so  on.   there 
is  even  more  room  for  difference  of  opinion,  and  it  must  be 
remembered  that  the  difference  is  not  so  much  as  regards 
facts,  such  as,  for  instance,  the  existence  of  certain  characters, 
but  rather  as  regards  decisions  on  such  questions  as  to  whether 
the  possession  of  certain  characters  in  common  by  several 
plants,  may,  or  may  not,  be  considered  to  prove  a  relationship 
between  them,  and  as  to  the  degree  of  such  relationship.     It 
will  thus  be  seen  that  the  Natural  System  of  Classification  is, 
as  yet,  by  no  means  perfect,  and  is  liable  to  modification  and 
alteration  as  our  knowledge  extends. 


126 


CHAPTER     II.— PRINCIPAL     SUB-DIVISIONS     OF    THE 
VEGETABLE   KINGDOM. 

.  .  111.  All    plants  are  first  divided 

-Division    •    ,         .-I  i    i  •        i  i-i-i 

of  the  into    the    two  great  groups,  or  sub -kingdoms,    called  respec- 

Vegetabie      tively  Cryptogams  and  Phanerogams.      The    term  Cryptogam 
is  derived  from   two    Greek   words   signifying  "  hidden   mar- 
Cryptogams  riage  "  and  was  originally  applied  to  all  plants  whose  reproduc- 
and  tive  organs  were  minute  and  inconspicuous. 

Phanero- 
gams. From    the    absence  of  what  are  usually  called  flowers,  or 

structures  containing  stamens,  or  pistil,  or  both,  this  great 
group  of  Cryptogams  is  sometimes  referred  to  as  the  one 
containing  "  flowerless,"  or  "  non-flowering,"  plants,  while  the 
group  of  Phanerogams  is  similarly  termed  that  which  includes 
the  "  flowering  "  plants. 

The  essential  difference  between  Cryptogams  and  Phanero- 
gams, however,  consists  in  the  fact  that  the  latter  produce 
true  seeds,  i.e.  many-celled  bodies  containing  the  multi- 
cellular  embryo  or  young  plant,  whereas  Cryptogams,  in  place 
of  seeds,  produce  spores  which  are  unicellular  structures. 
These  spores  may  be  developed  asexually,  or  they  may  arise 
as  the  result  of  a  sexual  process,  the  latter  being  manifest  in 
the  mingling  together  of  the  protoplasmic  contents  of  distinct 
cells.  When  the  sexual  cells  are  externally  similar,  each  is 
termed  a  gamete,  when  one  is  smaller  and  more  active  than  the 
other,  the  small  active  cell  is  termed  the  spermatozoid,  and 
the  large  passive  cell  the  egg-cell,  or  oosphere.  Spores  which 
are  capable  of  remaining  dormant  for  considerable  periods  are 
distinguished  from  those  capable  of  germinating  immediately 
after  their  formation  by  the  name  of  resting  spores. 

the  112'    These  tw°    great    gr°UPS  are 

Cryptogams,  further  sub-divided  as  follows  : — 

CRYPTOGAMS. 

I. — THALLOPHYTA.  ")  Cellular  plants  with  no   true   vas- 
II. — BRYOPHYTA.      j      cular  bundles. 

III. — PTERIDOPHYTA.    Vascular    plants    with   true    vas- 
cular bundles. 

Thaiio-  THALLOPHYTA. — The  plants  in  this  group  show  no  distinct 

phytes.         differentiation  into  stem    and  leaves  and  the  cellular  body 


127 

which  serves  such  plants  for  stem  and  leaves  is  called  a  thallus. 
For  our  purposes  the  plants  in  this  group  may  be  sub-divided 
as  follows  :  — 

1.  Algae      .  Plants   containing   chlorophyll. 

3*  ^Ta]°olourleSS   plants   without   chlorophyll. 


1  13.  Algae.     This  group  is  of  very 

little  importance  to  the  Forest  Officer  and  contains  plants  of 
simple  structure  which  live,  chiefly,  in  fresh  or  salt  water,  but 
also  on  a  damp  substratum,  such  as  moist  soil,  the  bark  of 
trees  in  shady  forests,  etc.  Many  are  coloured  green  ;  others 
are  blue-green,  brown,  red  or  purple,  in  such  cases  some 
other  pigment  more  or  less  masking  the  green  chlorophyll. 
Many  are  unicellular,  or  consist  of  minute  filaments,  while 
some  of  the  so-called  "  seaweeds  v  are  more  elaborate  in 
structure,  some  possessing  a  root-like  attachment  organ  and 
with  a  floating  thallus  which  may  attain  a  length  of  800 
feet,  or  even  more. 

114.  Bacteria.     This    group     in-  Bacteria. 

eludes  the  smallest  of  all  known  plants.  They  are  exceed-  Account 
ingly  minute  and  of  very  simple  structure,  being  unicellular 
or  filamentous,  with  no  indications  of  any  specialised  tissue. 
Many  are  always  present  in  the  air  and  soil,  others  live  in 
water,  and  others  in  dead  or  living  organic  matter.  Many 
are  well  known  in  connection  with  animal  diseases  and  are 
popularly  spoken  of  as  "  germs,"  or  "  microbes." 

Regarding  the  size  of  bacteria,  we  may  take,  as  a  fairly 
typical  example,  Bacterium  aceti,  the  little  individual  rods  of 
which  have  a  length  and  breadth  of  about  —  ^  of  an  inch 
and  —OQ  of  an  inch,  respectively.  Many  are  capable  of  active 
movement  by  means  of  exceedingly  fine,  hair-like  processes 
termed  cilia  or  flagella. 

The  presence  or  absence  of  these  flagella  and  the  shape  of 
the  individual  cells  are  important  characters  for  the  classi- 
fication of  bacteria.  The  following  forms,  for  instance,  are  dis- 
tinguished by  the  characters  given  :  — 

(a)  Cocci,  minute  spherical  cells. 

(b)  Bacteria,  rod-like  cells  with  no  flagella. 

(c)  Bacilli,  rod-like  cells  with   flagella  scattered   over  the 

whole  surface  of  the  cell. 

(d)  Spirally  curved  forms  (Vibrio  closely  wound,  usually 

with  one   polar  flagella,  Spirillum    closely   wound 


128 

with  polar  tufts  of  flagella,  Spirochaete  loosely 
wound,  long  and  filamentous). 

The  form  of  one  and  the  same  individual,  however,  may 
vary  considerably  at  different  stages  of  its  life-history  or  ac- 
cording to  the  medium  in  which  it  is  living.  The  individual 
bacterial  cells  are  thus  sometimes  united  into  chains,  or 
masses  of  varying  form,  and  owing  to  the  partial  dissolution 
and  swelling  of  the  outer  layers  of  the  cell  walls  of  the  indivi- 
dual cells  such  masses  are  often  distinctly  mucilaginous  or 
gelatinous. 

Plants  which  are  thus  capable  of  assuming  several  different 
forms  are  termed  polymorphic. 

So  long  as  conditions  are  favourable,  bacteria  grow  and 
multiply  very  rapidly.  Each  tiny  bacterial  cell,  whether 
spherical,  rod-shaped  or  curved,  having  grown  to  a  certain 
size,  becomes  divided  by  a  cell- wall  into  two  equal  portions, 
and,  these  segments  separating  from  one  another,  two  distinct 
individuals  are  formed,  each  of  which  then  continues  to  grow 
until  it  has  attained  the  normal  size,  when  it  also  divides  into 
two,  and  so  on.  This  mode  of  multiplication  by  cell-division 
and  the  separation  of  the  segments  is  very  characteristic  of 
the  bacteria  which  are  in  consequence  often  called  the  Fission — 
or  Splitting-Fungi  (Schizomycetes).  When,  from  the  exhaus- 
tion of  the  substratum  or  some  other  reason,  conditions  become 
unfavourable  for  growth,  the  preservation  of  the  plant  is 
insured  by  the  formation  of  resting  spores  which  can  exist  for 
long  periods  without  further  development.  In  the  formation 
of  spores  the  protoplasm  of  the  individual  cells  rounds  itself  oil 
and  becomes  surrounded  by  a  specially  thickened  membrane, 
each  cell  thus  becoming  a  spore.  On  germination,  the  protec- 
tive coat  disintegrates  and  the  cell  proceeds  to  grow  and  form 
new  individuals  by  division  as  before.  All  species  are  not  known 
to  form  spores.  There  is  no  sexual  reproduction  and,  when 
spores  are  formed,  no  special  spore-bearing  organ  is  developed. 
Many  bacteria  which  can  be  destroyed  by  boiling  after  ger- 
mination are  able  to  resist  high  temperatures  in  the  spore 
condition.  Hence  boiling  a  substance  or  liquid  once  is  by 
no  means  always  sufficient  to  sterilise  it,  i.e.  to  absolutely 
destroy  all  life  in  it.  The  boiling  should  be  repeated  several 
times,  an  interval  of  a  day  being  allowed  to  elapse  between 
each  operation,  so  that  some  of  the  spores  which  might  have 
survived  the  previous  operations  may  have  time  to  germinate. 
Direct  sunlight  is,  as  a  rule,  prejudicial  to  the  growth  and 
development  of  bacteria. 


129 

115.  Bacteria  are  of  great  practical  Practical 
importance  for  the  following  reasons  :- 

(1)  Many   species  possess   the  power    of    more  or  less  de- 

composing, and  altering  the  composition  of,  the 
material  from  which  they  obtain  their  food. 

Bacterium  acidi  lactici  breaks  up  milk  causing  it  to 
become  sour. 

Bacterium  aceti,  the  Vinegar  Bacterium,  converts  alco- 
hol into  acetic  acid. 

Bacillus  vulgaris  is  the  most  common  cause  of  the 
decomposition  of  meat. 

Such  decomposing  actions  are  sometimes  spoken  of  as 
decay  or  rotting,  sometimes  as  fermentation,  and 
sometimes,  owing  to  the  evolution  of  evil-smelling 
gases,  as  putrefaction. 

(2)  Many  species  cause  very  virulent  and  dangerous  animal 

diseases. 

Bacillus  tetani  is  found  in  the  soil  and  causes  tetanus. 
Bacillus  typlii  causes  typhoid  fever. 
Vibrio  cholerae  causes  cholera. 

(3)  Many  species  help  other  plants    to   obtain   the    supply 

of  valuable  nitrogenous  materials  which  are  neces- 
sary for  their  existence. 

Bacillus  radicicola  lives  in  tubercles  on  the  roots  of  vari- 
ous plants  and  is  able  to  fix  the  free  nitrogen 
of  the  air  and  to  pass  it  on  in  a  form  in  which  it 
can  be  utilised  by  these  plants,  this  source  of 
nitrogen  being  otherwise  not  available  for  them. 

The  so-called  Nitrifying  Bacteria  live  in  the  soil  and  con- 
vert ammonia  to  nitrous  and  nitric  acid.  Nitrates  are  thus 
eventually  formed  which  are  the  most  valuable  source  of 
nitrogen  for  the  majority  of  the  higher  plants. 

It  will  be  seen  that,  although  bacteria  are  responsible 
for  many  of  the  most  virulent  animal  diseases,  they  seldom 
affect  other  plants  injuriously,  this  being  partly  due  to  the 
difficulty  which  such  minute  organisms  find  in  passing 
through  the  cell  walls  of  plant  tissues. 

116.  Fungi.    In  this  group  are  in-  Fun?!- 

eluded  a  great  variety  of  plants  which  are  popularly  known  by 
various   names,  such   as   moulds,   rusts,   toadstools,  mushrooms, 

etc.  They  are  of  special  importance  to  the  Forester  on 
account  of  their  being  responsible  for  the  majority  of  the 
most  destructive  of  known  plant  diseases.  Some  fungi  are 


130 

very  minute  and  with  a  structure  scarcely  more  elaborate 
than  that  of  the  bacteria.  The  common  Brewers'- 
Yeast  Plant,  for  instance,  consists  of  single,  oval 
cells,  each  cell  having  a  diameter  of  about  ~^  of  an  inch. 
It  can,  however,  be  at  once  distinguished  from  the  bacteria 
by,  among  other  things,  its  considerably  larger  size,  its 
peculiar  mode  of  multiplication  by  what  is  known  as  budding, 
or  sprouting.,  and '  by  the  way  in  which  the  spores  are  formed 
as  will  be  clear  from  the  life-history  given  below.  The 
majority  of  fungi,  however,  have  a  much  more  elaborate 
structure  than  that  of  the  simple  Yeast  Plant,  and,  although 
they  never  show  traces  of  leaves,  there  is  commonly  a  more 
or  less  clear  differentiation  into  root-like  and  shoot-like  portions. 
Special  spore-bearing  organs  are  commonly  developed  and 
sexual  reproduction  is  often  met  with.  Many  of  the  simpler 
forms  resemble  colourless  alga3.  Some  fungi  then  are  unicellular, 
but  the  majority  consist  of  slender,  thin- walled,  more  or  less 
branched  filaments,  or  tubes,  called  hyphae,  which  are  only 
capable  of  growing  in  length  at  their  apices.  These  hyphae 
may  possess  cross- walls  which  thus  divide  them  into  segments, 
in  which  case  they  are  said  to  be  septate,  or  they  may  be 
aseptate,  i.e.  undivided.  The  whole  vegetative  body  of  the 
fungus  consists  of  these  hyphae  and  is  called  the  mycelium. 
This  ramifies  in  the  substance  from  which  the  fungus  derives 
its  nourishment  and  absorbs  the  necessary  food  materials 
from  it  and,  like  the  roots  of  higher  plants,  usually  remains 
out  of  sight.  Fungi  are,  therefore,  best  known  by  their 
reproductive  organs,  which  are  developed  at  the  surface  of  the 
substance  on  which  the  fungus  is  growing,  or  on  aerial  bran- 
ches of  the  hyphae.  Sometimes  the  mycelium  is  superfi- 
cial and  only  specially  developed  short  branches  of  the  hyphae, 
termed  haustoria,  or  suckers,  penetrate  the  substance  on  which 
the  fungus  is  growing  and  absorb  the  necessary  food 
materials. 

The  mycelium  may  consist  of  a  few  delicate  hyphae  and 
thus  be  invisible  to  the  naked  eye,  or  the  hyphae  may  be 
collected  into  dense  masses  and  become  visible  in  the  form 
of  papery  or  skin-like  membranes,  or  thick,  felt-like  sheets 
and  masses.  In  some  species  the  slender  thin-walled  hyphae 
are  united  into  bundles,  which  are  provided  with  a  hard, 
dark-coloured,  protective  coat.  Such  strands  may  some- 
times be  mistaken  for  the  fine  roots  of  higher  plants,  which 
they  resemble  in  general  appearance,  and  they  are,  in  conse- 
quence, termed  rhizomorphs.  or  root-like  structures.  Pro- 


131 

tected  in  these  strands,  the  tender  hyphae  are  able  to  spread 
through  areas  which  are  unsuitable  for  their  growth  and 
development,  the  hyphae  being  again  extruded  to  absorb 
nourishment  when  favourable  conditions  are  met  with. 

The  fungi  may  be  divided  into  the   following  groups  :  —  Classifica- 

tion of 

I.  —  Phy  corny  cetes.  Fungi. 

II.  —  Ascomycetes. 

III.  —  Basidiomy  cetes. 

IV.  —  UstUaginaceae. 
V.  —  Uredinaceae. 


117.  Phi/corny  cetes.  The  hyphae  are 
generally  aseptate.  Mycelium  does  not  form  a  compact  m-7ce 
mass.  Sexual  reproduction  is  common.  The  asexual  spores 
may  be  developed  in  spore-cases  (—  sporangia),  the  protoplasmic 
contents  of  which  divide  up  to  form  the  spores,  or  the  end  cf 
a  hypha  may  swell  up,  the  swollen  tip  eventually  separating 
from  the  hypha  to  form  a  spore,  which,  in  such  a  case,  is 
called  a  conidium,  •  and  the  portion  of  the  hypha  on  which  it 
is  borne  is  called  a  conidiophore. 

Phytophthora  infestans  causes  the  potato  disease.  This 
has  been  selected  as  a  typical  example  of  this 
group  and  its  life-history  is  given  in  detail  in  Part 
V  below. 

Other  species  of  Phytophthora  and  species  of  Pythium  are 
very  destructive  to  seedlings  and  to  palm  trees. 

118.  Ascomycetes.   Hyphse  are  septate. 

Spores  are  produced  in  a  special  kind  of  elongated  sporangium 
termed  an  ascus.  The  number  of  spores  produced  in  an  ascus 
is  almost  always  definite,  usually  eight,  whereas  in  a  sporan- 
gium the  number  of  spores  is  usually  indefinite.  In  an  ascus, 
also,  the  protoplasmic  contents  are  not  all  used  up  in  the  for- 
mation of  the  spores,  as  is  the  case  in  a  sporangium.  These 
asci  may  be  without  any  covering,  or  may  be  grouped  in 
special  structures,  called  the  ascocarps,  thus  forming  the  ascus- 
fruits.  A  completely  closed  ascocarp  is  called  a  cleistothe- 
cium,  one  with  a  small  aperture  at  the  apex  a  perithecium,  and 
one  which  is  open,  saucer-shaped,  or  hat-shaped,  an  apothecium. 
Sexual  reproduction  sometimes  occurs. 

Among     the    interesting     fungi     included    in    this    group  }; 

are  the   minute,   unicellular  plants   known  collectively  as   the 

Yeast  Fungi,   or  the  Saccharomy  cetes,   which   are   capable  of 

causing    the    alcoholic  fermentation    of  sugar  solutions.     \Ve 

Jmow  that,  if  the  sweet  juice  extracted    from    the   sugarcane 


132 

is  not  at  once  boiled  and  is  left  exposed  to  the  air,  the  process 
known  as  fermentation  rapidly  sets  in  and  that  if  this  is 
allowed  to  continue  we  ultimately  obtain  in  place  of  the  sugar 
solution  one  which  contains  alcohol,  carbon  dioxide  gas  being 
evolved  during  the  process.  Yeast  Fungi  are  therefore  of 
great  commercial  importance  in  the  manufacture  of  wine 
from  the  sugar  in  grapes,  in  the  preparation  of  spirit  from  the 
sweet-tasting  mahua  (Bassia  latifolia)  flowers  and  in  the  manu- 
facture of  wine,  beer  and  spirits,  generally.  If  the  minute  oval, 
or  spherical,  cells  of  a  Yeast  Fungus  are  placed  in  a  suitable 
nutrient  sugar  solution  they  are  found  to  grow  and  multiply 
with  great  rapidity,  the  peculiar  method  of  multiplication 
being  termed  budding.  The  cell  wall  of  the  Yeast  Plant 
bulges  out  and  a  protuberance  is  formed  which  gradually 
increases  in  size,  the  neck  connecting  it  with  the  mother-celt 
remaining  narrow.  A  cell-  wall  is  then  formed  across  the 
narrow  neck  at  the  point  of  union  and  the  swollen  protu- 
berance, separating  from  the  mother-cell,  becomes  a  separate 
cell  which  then  behaves  in  the  same  way.  When  all  the  sugar 
has  become  converted  into  alcohol  and  the  food  material  is 
consequently  exhausted,  the  yeast  cells  can  no  longer  continue 
growing  and  budding  and  they  proceed  to  form  spores.  The 
protoplasm  in  each  cell  divides  into  four  little  blocks,  each 
block  becoming  a  spore,  which  is  eventually  set  free  by  the 
disintegration  of  the  wall  of  the  mother-cell.  Under  favour- 
able conditions  each  of  these  spores  germinates  and  at  once 
proceeds  to  grow  and  bud  off  new  cells  as  described. 
Ferment  It  has  been  noted  above  that  bacteria  are  capable 

Qf  causmg  various  kinds  of  fermentation  but  they  do  not 
produce  the  alcoholic  fermentation  so  characteristic  of  the 
Yeasts.  Plants,  such  as  bacteria  and  fungi,  which  are  able 
to  produce  fermentation  are  known  as  Ferment  Organisms 
(sometimes  also  as  Organised  Ferments).  Such  plants  appear 
to  owe  their  power  of  producing  fermentation  mainly  to  the 
chemical  substances  known  as  Enzymes  which  they  containr 
and  which,  of  course,  are  not  living  organisms.  The  action 
of  the  Yeast  on  the  sugar  solution  will  be  understood  from 
the  following  equation  :  — 


(grape   sugar)         (alcohol)         (carbon   dioxide) 

and  it  should  be  noted  that,  although  the  Yeast  derives  its 
food  f-rom  the  sugar,  yet  only  a  very  small  proportion  of  the 
solution  is  actually  utilised  as  food,  the  remainder  being 


133 

decomposed  and  broken  up  into  simpler  compounds.  Thus 
only  5  per  cent,  of  the  sugar  may  be  actually  used  as  food 
while  95  per  cent,  is  reduced  to  alcohol  and  carbon  dioxide. 

Other  interesting  plants  belonging  to  this  group  are 
the  species  of  Meliola  which  form  sooty  black  patches  of  mould 
on  the  leaves  of  various  trees,  e.g.  the  Sal,  Mango^and  Orange. 
The  mycelium  is  entirely  superficial  and  does  not  penetrate 
the  leaves,  the  fungi  living  on  the  sweet  juices  excreted  by 
aphides,  scale  and  other  insects.  These  fungi,  unless  excep- 
tionally numerous,  do  very  little  harm  to  the  trees.  Preven- 
tive measures  should  aim  at  destroying  the  insects. 

Rosellinia  bunodes  is  a  species  which  has  been  found  to 
be  very  destructive  to  forest  trees,  such  as  Litsaea  angusti folia, 
Schleichera  trijuga  and  others,  in  Mysore  and  Assam. 

The  roots  are  first  attacked  and  the  fungus  is  best  recog- 
nised by  the  clusters  of  small,  round,  black  perithecia,  con- 
taining the  asci,  which  appear  at  the  base  of  the  attacked  stems 
just  above  the  ground  surface.  These  perithecia  have  a  car- 
bonaceous structure  and  appearance  and  can  be  crushed  in 
the  fingers  like  fragments  of  coal. 

119.  Basidiomyceles. — This'  group 
includes  the  most  highly  developed  forms  of  fungi  among  mycetes. 
which  are  those  commonly  known  as  Mushrooms  and  Toad- 
stools. The  hyphse  are  septate.  There  is  no  sexual  repro- 
duction. The  asexual  spores  are  developed  on  elongated, 
club-shaped,  terminal  cells  of  the  hyphse,  which  are  called 
basidia.  These  basidia  are  usually  placed  close  together,  side 
by  side,  and  form  the  hymenium  or  hymenial  layer.  At  the 
top  of  each  basidium  are  situated  four  spores,  each  on  a  little 
stalk.  This  hymenial  layer  may  be  smooth  and  flat,  it  may 
cover  the  sides  of  thin  lamellae  or  gills,  it  may  line  the 
interior  of  pores  or  tubes,  or  it  may  cover  the  surface  of 
raised  spikes,  knobs,  or  irregular  folds.  A  large  number  of 
species  are  known  to  be  injurious  to  Indian  trees,  of  which 
only  a  few  can  be  mentioned. 

Armillaria  mellea  (perhaps  better  known  as  Agaricus  mel- 
leus)  is  a  very  destructive  species  in  Europe  and  is  believed 
to  occur  in  India. 

In  Europe  it  is  particularly  destructive  to  coniferous  trees. 
It  is  one  of  the  few  species  characterised  by  the  development 
of  rhizomorphs  from  which  in  this  case  the  mushroom-like 
spore-bearing  organs,  or  sporophores,  arise.  The  sporophore 
consists  of  two  principal  parts,  the  stalk,  or  stipe,  and  the 
umbrella-shaped  cap,  or  pileus.  On  the  under-surface  of  the 


pileus  are  a  number  of  radiating  lamellae,  or  gills,  the  sides  of 
which  are  covered  with  the  hymenial  layer,  on  which  the  spores 
are  produced.  The  sporophores  are  usually  found  in  clumps 
near  the  base  of  the  attacked  tree — the  pileus  is  yellowish,  or 
brownish,  in  colour,  with  dark  scales,  and  the  stipe  bears  a  mem- 
branous, collarlike  ring.  The  sporophores  are  edible.  The 
mycelium  forms  firm  white  sheets  between  the  bark  and  the 
wood  on  the  roots,  or  at  the  base  of  the  trunk.  The  fungus 
spreads  by  means  of  its  spores,  which  are  disseminated  in 
myriads  as  a  fine  white  powder  and  which  are  capable  of 
producing  a  new  mycelium,  and  also  by  means  of  the 
rhizomorphs,  which  spread  through  the  soil,  bore  into  sound 
roots  and  produce  a  vigorously  growing  mycelium  in  them. 

Fomes  annosus  causes  the  well-known  Deodar  root-disease. 

Trametes  Pini  has  been  found  destroying  Pinus  excelsa. 

The  life  histories  of  these  two  species  is  given  in  detail  in 
Part  V  below. 

Much  of  the  so-called  "  dry  rot "  of  timber  which  has  been 
used  in  construction  is,  in  Europe,  caused  by  a  fungus  named 
Merulius  lacrymans,  which  is  believed  to  occur  also  in  India. 

Fomes  Pappianus  is  destructive  to  babul  (Acacia  arabica). 

120.   Ustilaginaceae.—Kyphdd  are 

septate.  No  sexual  reproduction.  The  mycelium  produces 
dark-brown,  or  black,  resting  spores.  Each  of  these  on 
germination  produces  a  short  tube  from  which  numerous 
small  conidia  called  sporidia  are  abstricted.  If  supplied 
with  sufficient  nourishment,  as  would  be  the  case  in  a 
field  which  has  been  manured,  these  conidia  are  capable  of  very 
rapid  multiplication,  by  budding,  or  sprouting,  after  the  manner 
of  yeast  cells.  As  the  nourishment  becomes  exhausted,  each 
conidium  develops  a  hypha,  and  proceeds  to  form  the  mycelium 
which  produces  the  resting-spores.  These  fungi  are  particularly 
destructive  to  cereal  crops,  such  as  wheat,  oats,  maize,  and 
others,  and  are  also  found  on  many  wild  grasses,  the  mycelium 
living  in  the  tissues  of  these  plants  and  ultimately  developing 
masses  of  dark-coloured  resting  spores.  These  characteristic 
spores,  resembling  as  they  do  a  sooty  powder,  have  caused  the 
fungi  in  this  group  to  be  characterised  by  the  popular  name  of 
smuts.  Several  fungi  in  this  group,  e.g.  one  of  the  commonest 
species  occurring  on  oats,  are  characterised  by  the  fact  that  the 
mycelium  is  only  able  to  enter,  and  infect,  the  plants  attacked 
when  the  tissues  of  the  latter  are  very  young  and  tender,  i.e. 
shortly  after  the  germination  of  the  seed.  Having  once  gained 
an  entrance,  the  mycelium  of  the  fungus  spreads  through  the 


135 

tissues  of  the  plant  attacked,  and  as  the  latter  grows,  the 
mycelium  grows  along  with  it,  without,  however,  so  far  as  cne 
can  see,  injuring  the  plant  in  any  way.  The  presence  oi  the 
fungus  is  indeed  only  made  manifest  to  the  ordinary  observer 
shortly  before  the  time  of  harvest.  The  hyphae  of  the  fungus 
entering  the  flowers  feed  on  and  destroy  the  substances  which 
should  have  been  devoted  to  the  development  of  the  grain,  and 
eventually,  instead  of  the  ripe  corn,  we  find  the  soot-like  masses 
of  resting  spores. 

Ustilago  Maydis  is  common  on  maize  in  India  and  is  easily 
recognised  by  the  boil-like  swellings,  or  blisters,  which  may  occur 
on  leaves,  stem,  flowers,  or  fruit  heads.  These  blisters,  .which 
may  attain  the  size  of  one's  fist,  contain  the  spores  and,  when  the 
latter  are  ripe,  the  blisters  burst  and  the  dark-coloured  spores 
escape.  The  maize  heads  are  usually  attacked  and  the  grain, 
is  consequently  destroyed. 

121.    Uredinaceae.    Hyphse    are,.    ,. 

septate.  Sexual  reproduction  doubtful.  The  fungi  included  in 
this  group  are  characterised  by  their  polymorphism.  Several 
kinds  of  spores  are  usually  produced  by  them  and  their  com- 
plicated life-history  will  be  best  understood  from  the  detailed 
account  of  Puccinia  graminis  given  below  in  Part  V. 

Owing  to  the  frequent  development  of  masses  of  orange- 
yellow,  or  rust-coloured,  spores,  which  give  the  appearance  of 
rusty  streaks,  or  patches,  on  the  leaves  of  the  plants 
attacked  by  them,  these  fungi,  in  contradistinction  to  those  of 
the  previous  group,  are  popularly  known  as  rusts.  A  very 
large  number  of  these  fungi  occur  on  the  leaves  of  our  Indian 
forest  trees  and  shrubs,  but,  as  a  rule,  they  are  not  very 
injurious  and  are,  consequently,  not  of  great  importance.  Only 
a  few  of  those  which  are  most  noticeable  in  the  forest  of  the 
North- West  Himalaya  will  be  mentioned  here. 

One  of  the  most  remarkable  is  that  known  as  Bardaydla 
deformans  (formerly  named  Aecidium  Thomsoni)  which  causes 
the  orange-red  tassels  so  frequently  seen  in  the  forests  of 
Jaunsar-Bawar  on  the  spruce,  Picea  Morinda  (see  Plate  XV, 
Fig.  1  (a)  and  (b)  ).  Only  the  current  year's  shoots  are  attacked, 
these  being  "stunted,  thickened  and  densely  covered  with 
curved  needles."  Every  needle  on  the  attacked  shoot  is  affected 
and  they  afford  a  striking  contrast  to  the  straight-,  healthy 
needles.  The  masses  of  orange-red  spores  (teleuto-spores) 
form  "  two  continuous  flattened  beds  on  the  upper  surface  " 
and  two  smaller  beds  on  the  under-surface  of  the  needles. 
When  young,  the  tassels  emit  a  disagreeable  odour.  The 


136 

attacked  shoots  eventually  shrivel  up  and  turn  black.  As  a 
rule,  the  damage  done  is  not  great,  the  leading  shoot  not  being 
often  attacked.  The  teleuto-spores  may,  also,  be  sometimes 
found  on  the  cone  scales  (see  Fig.  1  (b) ). 

The  needles  of  Pinus  longifolia,  the  chir,  are  often  seen  to 
produce  prominent,  flattened,  reddish-yellow  blisters,  usually 
about  one-fifth  of  an  inch  long  and  one-tenth  of  an  inch  high. 

Those  contain  the  spores  (secidiospores)  of  the  fungus 
Peridermium  complanatum.  The  mycelium  lives  in  the 
needles,  but  not  much  harm  results.  Sometimes  the  orange 
blisters  may  be  seen  on  the  stem,  or  young  branches,  and  as,  in 
such  cases,  the  mycelium  destroys  the  cambium  and  young 
cortex  the  damage  is  more  severe.  This  form  on  the 
stem  is  considered  to  be  a  variety  of  the  above  fungus  and 
is  distinguished  by  the  name  of  corticola. 

Peridermium  brevius  (see  Plate  XV,  Fig.  2)  is  a  fungus  closely 
resembling  the  last  which  attacks  the  needles  of  Pinus  excelsa. 
The  blisters  are  smaller  than  those  caused  by  P.  complanatum. 
Chrysomyxa  Himalense  causes  the  very  noticeable,  orange -red 
blisters  often  seen  on  the  petioles  of  Rhododendron  arboreum  and 
R.  campanulatum. 

Gymnosporangium  Cunninghamianum  (see  Plate  XV,  Figs. 
3  and  4)  is  an  interesting  example,  inasmuch  as  it  is  one 
of  the  many  species  known  in  this  group,  which,  for  its 
full  development  and  completion  of  its  life-history,  requires  to 
live  on  two  distinct  species  of  plants.  Part  of  its  life  is  spent  on 
the  cypress,  Cupressus  torulosa,  and  the  remainder  on  the  pear, 
Pyrus  Pashia.  On  the  small  branches  and  twigs  of  the  cypress 
homisperical,  or  elongated,  dark  brown  spore  masses  (teleuto- 
spores)  arise,  which,  during  moist  weather,  swell  up  enormously 
into  gelatinous  masses.  These  soon  become  yellowish  in  colour 
owing  to  the  germination  of  the  teleuto-spores  and  the  forma- 
tion of  small,  orange-red  conidia,  called  sporidia,  which  are 
abstricted  from  the  teleuto-spore  germ-tube  as  described  in  the 
case  of  the  resting  spores  of  the  Vstilaginaceae.  These 
conidia  are  capable  of  developing  a  mycelium  in  the  leaves  of 
the  pear.  These  leaves  ultimately  produce  conspicuous, 
thickened  patches  which  may  be  |"  in  diameter,  orange-red 
above  and  yellowish  below.  On  the  lower  surface  of  these 
patches,  little  tubular  structures  are  found,  1 — 2-mm.  long, 
which  are  called  the  aecidia  and  in  which  the  aecidiospores  are 
produced.  These  spores  on  germination  infect  the  cypress. 
This  fungus  does  a  lot  of  harm  to  young  cypress  seedlings  and 
plants  in  Jaunsar.  It  is  thought  that  the  eradication  of  the 


137 

pear  near  the  cypress  plantations  would  lead  to  the  disappear- 
ance of  the  pest,  the  fungus  being  then  unable  to  complete  its 
normal  life-cycle. 

Perhaps  the  most  remarkable  rust  of  all  is  that  called  Gam- 
bleola  cornuta  which  is  often  seen  on  Berberis  nepalensis  in 
Jaunsar  and  near  Mussoorie.  It  produces  clusters  of  long, 
black,  wavy  hairs  on  the  under -surface  of  the  leaves  and  some- 
times on  the  twigs.  Each  hair  consists  of  chains  of  teleuto- 
spores  which  adhere  closely  to  one  another  (see  Plate  XV,  Fig.  5). 

122.  Before    leaving   the     great  Lichens, 
group     of     the    Thallophytes,      we    must      shortly    consider 

the  curious  plants  called  Lichens.  The  general 
appearance  of  a  lichen  is  well  known,  viz.  that  of  a  crust  on 
stones,  earth,  trees,  etc.,  of  an  irregularly  shaped,  foliaceous 
structure  adhering  loosely  to  the  substratum  ;  or  again,  that 
of  a  tufted,  shrubby  mass  of  elongated  branching  processes, 
such  as  is  often  seen  festooning  the  stems  and  branches  of  trees  • 
in  cold  damp  forests  of  the  Himalayas ;  while  others  form 
gelatinous,  jelly-like  masses.  A  lichen  consists  of  two  distinct 
plants  which  live  together,  in  partnership  so  to  speak,  one  of 
these  partners  being  a  fungus,  usually  belonging  to  the 
Ascomycetes,  and  the  other  a  minute  alga.  The  green, 
chlorophyll-containing  cells  of  the  latter  are  enmeshed  in 
the  closely  woven  hyphae  of  the  fungus,  the  latter  being  res- 
ponsible for  the  general  form  and  outline  of  the  lichen.  The 
lichen  is  attached  to  the  sub-stratum  by  root-like  hairs,  or 
attachment  organs,  developed  by  the  hyphae  of  the  fungus. 
The  fructification  of  the  lichen  is  developed  by  the  fungus,  but 
the  germinating  fungus  spores  usually  perish  if  unable  to  find  a 
suitable  alga  with  which  to  enter  into  partnership.  Lichens  are 
commonly  reproduced  by  minute  buds,  each  of  which  contains  a 
few  algal  cells  entwined  with  hyphse.  These  are  distributed  by 
the  wind  and  grow  into  new  lichens.  The  gelatinous  lichens 
are  usually  dark-brown,  or  olive-green,  in  colour,  while  others 
;are  usually  greyish-green,  or  yellowish. 

123.  We    now    pass    on   to    the  Bryophytes. 
second  great  sub-division  of  cryptogams,  viz.  the  BRYOPHYTA. 

The  plants  in  this  sub-division  are  divided  into  two  groups  :— 

(1)  The  Liverworts  (Hepaticae). 

(2)  The  Mosses  (Musci). 

Liverworts  are  minute  plants  of  no  forest  importance  which  Liverworts, 
may  be  found  growing  in  damp  soil,  on  rocks,  tree-trunks  and 
even   in   the  water.     Many  of  them  show  no  differentiation 
into  stem  and  leaves  and  possess  a  lobed  thallus,  thus   resem- 


138 

bling  the  Algae.  They  are  included  in  the  Bryophyta  mainly 
on  account  of  their  life-history  and  the  structure  of  their 
reproductive  organs.  Those  which  possess  leaves  usually  have 
the  leaves  inserted  in  two  rows. 

Mosses.  The  mosses  are  small  herbaceous  plants,  showing  a  distinct 

differentiation  into  a  slender,  often  wiry,  stem  and  small 
green  leaves,  which  are  commonly  found  growing  in  damp 
shady  places,  on  the  ground,  on  trees,  rocks,  etc. 

The  leaves  are  usually  arranged  spirally  and  not  in  two  rows 
as  in  the  Liverworts.  They  possess  no  true  vascular  bundles, 
and  consequently  the  leaves,  which  have  a  very  simple 
structure  and  are  usually  only  one  cell  thick,  exhibit  no  distinct 
venation,  although  a  rudimentary  midrib  may  be  present. 
The  roots  of  these  plants  do  not  possess  the  elaborate  structure 
of  true  roots,  but  are  very  simple  organs,  consisting  of  single 
rows  of  cells  and  are  known  as  rhizoids.  The  characteristic 
fruit  of  a  moss  is  a  stalked  capsule,  called  the  sporogonium, 
which  contains  microscopic  spores.  These,  on  germination, 
develop  a  growth  of  minute  green  filaments  called  the  protonema, 
on  which  buds  arise.  From  these  buds  are  produced  what  we 
know  as  the  moss  plant.  The  microscopic  reproductive  organs 
arise  in  groups  at  the  apex  of  the  shoot,  or  in  the  leaf  axils. 
The  male  organs,  termed  antheridia,  are  stalked  and  club- 
shaped  and  contain  a  number  of  small  cells.  When  mature  the 
antheridium  ruptures  at  the  apex  and  expels  these  cells. 
Each  of  the  latter  then  liberates  a  minute  twisted  filament 
called  the  spermatozoid  which  is  provided  with  two  long 
cilia.  The  female  organs,  termed  the  archegonia,  are  flask-shaped 
with  a  slender  neck.  The  dilated  basal  portion  contains  a 
,  naked  mass  of  protoplasm  called  the  egg-cell,  or  oosphere,  which 
lies  in  the  bottom  of  the  archegonium  like  jelly  in  a  flask.  The 
spermatozoids,  swimming  by  means  of  their  cilia,  in  the  dew  or 
rain  water  with  which  the  sponge-like  moss  tufts  are  so  often 
saturated,  reach  the  archegonium,  pass  down  its  neck  and  one  of 
them  penetrates  the  oosphere.  With  the  fusion  of  the  proto- 
plasm of  these  two  bodies  fertilisation  is  accomplished  and  the 
fertilised  oosphere,  which  is  now  called  the  oospore,  surrounds 
itself  with  a  cell-wall  and  at  once  begins  to  grow  and  to  divide. 
After  the  first  cell-division  has  taken  place,  the  body  is  no  longer 
called  the  oospore  but  the  embryo.  By  further  growth  and 
division  this  embryo  eventually  develops  into  the  sporogonium, 
bearing  the  well-known  moss-capsule  at  its  apex  and  with  its 
foot  sunk  in  the  tissues  of  the  moss  plant.  The  capsule  itself 
is  called  the  them  and  it  is  often  provided  with  a  long  stalk  called 


139 

the  seta.  In  this  capsule  are  produced  the  asexual  spores.  As 
the  embryo  develops  in  the  archegonium,  the  wall  of  the  latter 
is  stretched  and  eventually  ruptures,  a  portion  of  the  torn 
investment  being  carried  up  on  the  capsule,  as  the  seta 
elongates,  and  thus  forms  the  cap,  or  calyptra.  The  capsule  is 
usually  closed,  until  ripe,  by  a  lid  called  the  operculum  and  a  row 
of  minute  teeth,  called  the  peristome,  is  often  found  fringing  the 
upper  edge  of  the  capsule  inside,  just  below  the  operculum. 
We  thus  see  that  there  are  two  distinct  stages  in  the  life -history 
of  a  moss,  the  general  form  and  appearance  of  the  plant  varying 
considerably  in  the  two  stages.  There  is  thus  the  sexual  stage,, 
the  ordinary  moss -pi  ant,  which  arises  from  the  asexual  spore 
and  produces  the  sexual  reproductive  organs.  As  a  result 
of  the  sexual  process  of  fertilisation,  the  oospore  is  formed,  and 
with  this  commences  the  asexual  stage  which  ends  with  the 
production  of  asexual  spores.  Each  of  these  stages  is  called  a 
generation,  and  when  they  alternate  regularly,  as  in  the  moss^ 
the  phenomenon  is  termed  an  alternation  of  generations.  The 
sexual  generation  is  called  the  Oophyte,  or  Egg-plant,  and  the 
asexual  the  Sporophyte,  or  spore-plant.  The  former  arises  from 
a  spore  produced  asexually,  the  latter  from  a  spore  produced 
sexually.  The  life-history  of  the  moss  may  be  shortly  expressed 
as  follows : — 

Sporophyte  Oophyte 

Spore —  —  ^-Protonema 

I 
Moss  Plant 


c?  antheridia          $  archegonia 

!  I 

spermatozoid     =     oosphere 

! 

Sporogonium^—  — -oospore 

124.  We  now  come  to  the   third  Pterido- 

and  last  great  group  of  cryptogams,  viz.  the  PTERIDOPHYTA.  P^*63- 
The   plants    in   this   group  exhibit  a    distinct   differentiation 
into  root,  stem  and  leaves  and  possess  true  vascular  bundles, 
while  in  the    case    of    these    plants,    as    in    the    Bryophyta,  Ferns, 
there  is  a  distinct    alternation  of  generations.      The    plants 
of  most  forest  importance   included   in  this  group  are  those 
known  as  the  Ferns  (Filices).     The  leaves    of  ferns  are  called 
fronds,  their   venation  is  usually  furcate  and   their   vernation 
circinnate.      The  young   stem    and    leaf-stalks    are    usually 


140 

provided  with  remarkable,  brown,  scaly  hairs  called  ramenta. 
The  primary  root  does  not  persist  and  form  a  tap  root,  but 
soon  dies  back,  its  place  being  taken  by  adventitious  roots 
springing  from  the  stem  or  leaf-stalks.  Most  ferns  are  her- 
baceous with  a  creeping  rhizome  of  which  the  common  bracken 
(Pteris  aquilina)  is  a  good  example  and  which  sends  up  each 
year  one  or  more  large  fronds.  Others,  however,  are  known  as 
Tree  Ferns  and  attain  a  height  of  some  50  feet  and  a  diameter 
of  1  foot  with  fronds  12  feet  in  length.  In  their  general 
appearance  these  Tree  Ferns  resemble  some  species  of  Palms 
with  their  unbranched  stems  and  terminal  rosettes  of  large 
pinnate  leaves.  The  venation  and  vernation  of  the  leaves, 
however,  is  usually  very  characteristic,  while  the  stem 
exhibits  a  different  structure  from  those  of  other  woody  plants. 
The  stems  of  the  Pteridophytes  contain  no  permanent  cam- 
bium and  hence  there  is  no  secondary  growth  in  thickness. 
The  closed  vascular  bundles  are  usually  concentric  and 
arranged  more  or  less  in  a  circle.  A  section  across  the  stem 
of  a  Tree  Fern  therefore  shows  :— - 

(1)  A  central  portion  of  cellular  tissue  which  often  decays 

and   leaves    a    hollow   in    old    stems. 

(2)  An  outer  mass  composed  of  the  bases  of  the  fallen 

leaves    and    adventitious    roots. 

(3)  An  intermediate  zone  between  (1)  and  (2)  containing 

the  closed  vascular  bundles,  each  of  which  is 
usually  crescent-shaped  with  a  dark-coloured 
border. 

On  the  fern  fronds  and  usually  on  their  under-surface 
are  produced  the  little  capsule-like  spore-cases,  or  sporangia 
which  to  the  unaided  eye  look  like  little  granules.  Those 
leaves  which  bear  the  sporangia  are  distinguished  from  the 
barren  fronds  by  the  term  sporophylls  (  — spore-bearing  leaves). 
The  sporangia  are  usually  collected  in  groups,  each  group  being 
termed  a  sorus,  and  in  many  cases  they  are  protected  when 
young  by  an  outgrowth  of  the  leaf-tissue  called  the 
indusium.  The  rupturing  of  the  sporangium  results  in  the 
dispersal  of  the  microscopic  spores,  each  of  which,  on  germina- 
tion, is  capable  of  producing  a  small,  flat,  usually  heart-shaped 
structure  called  the  prothallium,  which  resembles  a  minute 
cordate  leaf  and  is  usually  considerably  less  than  an  inch  in 
diameter.  This  usually  contains  chlorophyll,  and,  developing 
numerous  root-hairs  from  its  under-surface,  becomes  free 
from  the  spore.  On  the  under-surface  of  the  prothallium 
arise  the  microscopic  reproductive  organs,  termed  antheridia 


141 

and  archegonia,  which  are,  in  the  main,  similar  to  those 
described  above  for  the  mosses,  only  of  somewhat  simpler 
structure.  The  oosphere  contained  in  the  archegonium,  on 
being  fertilised  by  one  of  the  spermatozoids  liberated  by  the 
antheridia,  becomes  an  oospore,  surrounds  itself  with  a  cell 
wall  and  proceeds  at  once  by  growth  and  cell-division  to  develop 
into  the  young  fern  plant.  The  latter  remains  attached  to- 
the  pro  thallium  by  a  special  organ  termed  the  foot,  by  means 
of  which  it  obtains  its  nourishment  from  the  prothallium 
until,  with  the  development  of  its  own  leaves  and  roots,  it 
becomes  independent  and  then  the  prothallium  withers  and 
dies.  The  oophyte  in  the  fern,  therefore,  is  represented  by 
the  prothallium,  while  the  fern  plant  itself  is  the  sporophyte. 
The  prothallium  of  the  Pteridophyta  is  consequently  homo- 
logous with  the  protonema  and  what  we  call  the  '  '  plant  '  '  of 
the  Bryophyta,  while  the  '  '  plant  '  :  of  the  Pteridophyta  is 
homologous  with  the  sporogonium  of  the  Bryophyta.  The 
life  history  of  the  fern  may  be  shortly  expressed  as  follows  :  — 

Sporophyte  Oophyte 

Spore  _  Froth  allium 


d"  antheridia  ?  archegonia 

spermatozoid     =       oosphere 

Fern  Plant<—  —oospore 

It  should  be  noted  that  in  both  the  Bryophyta  and 
Pteridophyta  water  is  necessary  for  fertilisation,  to  enable  the 
spermatozoids,  which  swim  in  the  wrater,  to  reach  the  arche- 
gonia. 

The  Pteridophyta  are  usually  sub-divided  into  the  follow- 
ing three  groups  :  — 

1.  Ferns     proper.      Plants     with     well-developed     large  classification 

leaves.     Sporophylls  are    usually    similar    to    the  of  Pterido- 
foliage  leaves  and  are  not  aggregated  into  cone-  Phytes- 
like    structures.     Each  sporophyll    usually     bears 
numerous  sporangia. 

2.  Horsetails.     Plants    with    well-developed    stems    with  Ferns, 

distinct  nodes  and   internodes,    more  or  less  ribbed  •horsetails, 
and  with  small  leaves  in  whorls,  forming  a  sheath 
at  each  node.     The   small  peltate  sporophylls  are 
aggregated  into  a   cone-like  flower   at   the  apex  of 
each  fertile  shoot. 


142 

3.  Club-mosses.  Plants  with  numerous  small  scattered 
leaves.  Sporophylls  resemble  foliage  leaves  but  are 
sometimes  aggregated  into  conelike  flowers. 
Sporangia  are  borne  singly  either  on  the  upper 
surface  of  the  sporophyll,  or  on  the  stem. 

Now,  in  the  true  ferns,  the  asexual  spores  are  all  alike  and 
one  and  the  same  prothallium  bears  both  antheridia  and 
archegonia.  In  some  Pteridophytes,  however,  the  spores 
are  of  two  kinds,  the  larger  ones  being  termed  macrospores 
and  the  smaller  ones  microspores.  These  are  contained  in 
separate  sporangia  which  are  called,  respectively,  macrospor- 
angia  and  microsporangia ;  the  leaves  bearing  the  former 
being  the  female  sporophylls  and  the  leaves  bearing  the  latter 
the  male  sporophylls.  From  the  macrospore  is  developed  a 
female  prothallium  which  produces  only  archegonia  and 
from  the  microspore  a  male  prothallium  producing  only  antheri- 
dia. Moreover,  in  such  plants,  the  prothallium  is  in  every 
case  pulled  further  back  into  the  spore,  from  which  it  now 
never  becomes  independent,  as  we  have  seen  to  be  the  case 
in  the  ferns.  The  male  prothallium  in  particular  becomes 
very  much  reduced  and  consists  of  only  a  few  cells.  Moreover, 
in  the  Horsetails  and  some  Club  mosses  the  sporophylls  are 
aggregated  into  small,  cone-like  flowers.  These  plants  thus 
lead  us  up  to  the  great  group  of  the  most  highly  developed 
plants,  known  as  the  :— 

125.   Phanerogams. — This     contains 

all  the  plants  which  are  not  included  in  the  groups  already 
enumerated  and  briefly  described  above,  and  hence  it 
includes  practically  all  our  forest  trees  and  shrubs.  These 
plants  exhibit  a  distinct  differentiation  into  true  root,  stem 
and  leaves,  and  possess  true  vascular  bundles.  They  are 
characterised  by  the  production  of  true  seeds.,  and  the 
majority  possess  flowers  containing  structures  which  we  at 
once  recognise  as  stamens  and  carpels.  In  Phanerogams, 
as  in  the  higher  Cryptogams,  there  is  an  alternation  of 
generations,  although  at  first  sight  this  is  not  obvious, 
owing  to  the  great  reduction  in  the  oophyte,  that  which 
we  see  and  recognise  as  the  ' '  plant ' :  being  the  sporo- 
phyte.  The  various  organs  and  members  of  phanerogamic 
plants  were  described  and  received  definite  names  when  the 
life  histories  of  most  of  the  cryptogams  were  very  imperfectly 
known.  This  led  to  different  names  being  given  to  organs 
which  are  really  homologous  and  this  has  resulted  in  further 
obscuring  the  resemblances  between  the  plants  of  these  two 


143 

great  groups.     Thus    we    find    that    the    male  and    female 
sporophylls    of   phanerogams  are  called  stamens   and  carpels 
respectively.     The  sporophylls  with  the  portion  of  the  shoot 
on  which  they  are  borne  constitute  the  flower,  while  in  many 
cases  other  leaves  surround  the  sporophylls  and  form  a  perianth. 
The  microsporangia  are  the   pollen-sacs.,  the  macrosporangia, 
the    ovules.     The    microspores     are   the   pollen     grains,    the 
macrospore  is  the  embryo-sac  ;  both  kinds  of  spores  containing 
as  usual  a  single  nucleus,  the  division  of   this  nucleus   in  each 
case  is   the  first   step  towards   the   formation    of  a  prothal- 
lium    (i.e.   the   oophyte).     The    male   prothallium   is    always 
minute  and  consists  of  only    a  few  cells,  the  most  noticeable 
portion  of  it  being  the  so-called  pollen-tube.      Spermatozoids 
are  rarely  formed  and  are  usually   represented  by  two  small, 
naked,  nucleated,   male  cells  which   travel    down  the   pollen 
tube  and  eventually  pass  out  at  the  apex.     The   cells  arising 
from  the  repeated  nuclear  division  in  the  macrospore    consti- 
tute the  female  prothallium,  which    is  sometimes   large   and 
forms  the  tissue  known   as    endosperm.      In    the   prothallium 
an  oosphere  arises,  either    as   a   naked   cell,  or  in  an  arche- 
gonium.     Fertilisation  is  accomplished  with  the  fusion  of  the 
nucleus  of  a  small  male  cell  with  that  of  the  oosphere  ;   while, 
from  the  resulting  oospore,  the  embryo,  or  young  sporophyte, 
is  produced.     Here  then  the  prothallium  never    has  an  in- 
dependent existence  apart  from   the   spore    which    has   pro- 
duced it.     The  female    prothallium  remains  altogether   inside 
the   macrospore,   the     latter  remains   inside    the    macrospo- 
rangium, and  the  latter  remains  for  a   long  time  attached  to 
the    parent    plant    or   sporophyte.      Eventually     the    whole 
macrosporangium  with   the  macrospore  and  contained  embryo 
separates   from   the   parent    plant    as    the    seed.     Thus    the 
oophyte  does  not  form  a  distinct  and  separate  stage  in  the 
life  history.     The  embryo    during  its  growth  and  development 
frequently   absorbs    the   rest    of   the    tissue     formed   in   the 
macrosporangium  and  consequently  the  kernel  of  the  ripe  seed 
consists  of  the  embryo  alone.     In  such  cases   the  seed   is  said 
to  be  eocalbuminous.     In  other  cases  some  of  the   tissue  of  the 
macrosporangium  persists  in  the  ripe  seed  and  the  latter  is 
said  to  be  albuminous.   Such  persistent  tissue  is  distinguished  as 
endosperm,  or  perisperm,  according  as  it  has   developed  within, 
or   without,  the  macrospore.    See   also  p.   58.     Phanerogams 
are  divided  into  the  two  following  main  divisions  : — 
I.  Gymnosperms. 
II.  Angiosperms. 


144 


Gymno- 
sperms. 


Cycadace*. 


126.  Gymnosperms.     The   ovules  are 

naked   and   not   enclosed  in    an    ovary.      The  pollen    grains 
are  thus  able  to  come  directly  in  contact  with  the  micropyle 
of  the  ovule.     The  oophyte  developed  by   the  macrospore  is 
represented  by  a   mass  of   tissue  which    completely  fills   the 
macrospore    (embryo    sac)    and  is    called    endosperm.      The 
oosphere     is     contained     in  an  archegonium.      There    is    no 
distinct  perianth    to  the   flower.     The  flowers   are  unisexual. 
Cotyledons,  two  or  more,  usually  appearing  above    ground 
on  germination,    are   perennial  trees  and  shrubs,  usually  resin- 
ous.     Leaves  are  often  acicular.     The  primary  root  persists 
and    forms  a  tap-root.     The  vascular  bundles  are  open,  col- 
lateral and  arranged  in  a  circle   round   the   pith.     Secondary 
growth  in  thickness  takes  place  by  means  of  a  normal  cambium 
ring  and  the  stem  consequentlv  exhibits  a  central   pith,   with 
distinct  bark  and  wood,  while  it  increases  in  size   by   rings   of 
new  growth  added  at  the   outside   of   the   wood-cylinder   and 
immediately  beneath  the  bark.     True  vessels  are   not   present 
in  the  wood  except  in   that   of  Gnetaceae.     Gymnosperms  are 
sub-divided  into  the  following  three  orders  :— 

Cycadacese. 

Conifers. 

Gnetacea3. 

127.  Cycadaceae.     In  this  order  are 

included   the     plants    which     are     often     seen    in      Indian 
gardens  and  are  sometimes  wrongly  called  the  Sago  Palms. 
The       columnar,      usually     unbranched,      stem      with     the 
terminal  rosette   of     large   pinnate     leaves   of    these   plants 
gives     them     a     superficial    resemblance      to     the     Palms. 
In  India  they  rarely  attain  a  greater    height    than    30    feet. 
The  wood  is    characteristic  owing  to  its  possessing  alternate 
layers  of  woody  and  bast  tissue  with  no  true  vessels,  and    the 
pith  is  usually  large.     The  flowers  have   no  perianth  and  are 
dioecious.     They  are    situated  at,  or  near,  the  apex  of  the 
stem,  the   male  flower  being  a  cone  with  a  number  of  thick 
scales,  on  the  under-surface  of  each  of  which  are    situated 
numerous  pollen-sacs.     In  the  female  Cycas  tree  the    charac- 
teristic   macrosporophylls,   or  carpels,   arise  on  the  apex     of 
the  main  stem,  taking  the  place  as  it  were  of  the  foliage  leaves. 
Although    smaller   and   not   green,    these    carpels   somewhat 
resemble  the  foliage  leaves.     The  upper  portion  of  the  carpel 
is   pinnate,  the  ovules,   wrhich  may  attain  the  size  of  small 
plums,  being  borne  laterally  on  the  basal  portion  in  the  posi- 
tion of  leaflets.      Cycas  Rumphii,  and  C.  pectinata  are  well- 


145 

known  Indian  trees,  from  the  stems  of  which  sago  or  starch 
is  obtained. 

128.  Coniferae.      These    include  Conifer®. 

much  branched  woody  plants  with  undivided,  usually  acicuiar, 
leaves.  The  wood  does  not  contain  true  vessels  but  often 
possesses  resin  ducts.  The  flowers  have  no  perianth  and  are 
usually  cones,  i.e.  shoots  carrying  scale-like  sporophylls,  the 
latter  bearing  the  ovules  and  pollen-sacs.  In  this  group 
are  included  very  many  important  forest-trees  such  as  the 
Pines,  Firs,  Spruces,  Cedars,  Larches,  Cypresses,  Junipers 
and  others. 

129.  Gnetaceae.  The  leaves   are  un-  Gnetaceae. 
divided,    often  broad   and   usually   opposite.     The  stem   and 
branches  are  jointed  at  the  nodes.    The   flowers  are  not  cones, 

they  have  a  rudimentary  perianth  and  are  usually  arranged  in 
slender  spikes.  The  wood  contains  true  vessels. 

Ephedra  Gerardiana  is  a  small,  apparently  leafless,  shrub, 
fairly  common  in  Jaunsar. 

Several  species  of  Gnetum  occur  in  India.  They  are 
generally  climbers  with  broad,  well  developed,  opposite  leaves. 

130.  Angiosperms.     The  macrospo-  Angiosper 
rophylls,  or  carpels,  form  closed  cavities  called    ovaries,  within 
which  are   contained  the  ovules.     The  pollen  is   received   not 

on  the  micropyle  of  the  ovule,  but  on  a  specialised  portion  of 
the  carpel  called  the  stigma.  The  oophyte  developed  by  the 
macrospore  is  small  and  consists  of  only  a  few  cells  which  are 
usually  without  cell-walls,  one  of  these  naked  cells  being 
the  oosphere.  After  fertilisation  has  taken  place, 
however,  the  cavity  of  the  spore  becomes  filled  with 
tissue  which  has  also  been  called  endosperm,  but  which  is 
clearly  not  the  same,  morphologically,  as  the  endosperm  of 
Gymnosperms  which  is  developed  before  fertilisation.  The 
flowers  are,  typically,  hermaphrodite  with  a  perianth.  The 
Angiosperms  are  sub-divided  into  two  groups  : — 

Monocotyledons. 

Dicotyledons. 

131.  Monocotyledons.  The  embryo  Monocoty- 

has  typically  one  cotyledon  which  usually  remains  below  the ledons- 
ground.  These  plants  are  usually  herbaceous  and  are 
rarely  shrubs  (species  of  Smilax  for  example),  or  trees 
(Palms  and  Bamboos).  The  flowers  have  their  parts  typically 
in  whorls  of  three.  The  leaves  usually  have  parallel 
venation.  Species  of  Smilax  and  Dioscorea  are  the  most 
important  exceptions  with  reticulate  venation.  The  leaves 


146 

are  usually  without  stipules,  with  a  well-developed  leaf  base 
and  not  disarticulating  readily  from  the  parent  axis.  The 
primary  root  soon  dies  and  its  place  is  taken  by  adven- 
titious roots  which  arise  successively  higher  and  higher  on 
the  stem.  The  collateral,  closed  vascular  bundles  are  scat- 
tered irregularly  through  the  stem.  Secondary  growth 
in  thickness  rarely  occurs,  but  when  it  does  and  a  cambium 
ring  is  present,  the  latter  does  not  form  a  continuous  ring 
of  wood  on  the  inside,  but  produces  closed  vascular  bundles 
scattered  in  fundamental  tissue.  Perennial  stems  usually 
have  no  distinct  bark,  no  annual  rings,  and  no  medullary 
rays,  but  possess  true  vessels. 

Dicotyle-  132.  Dicotyledons.    Embryo     has 

dons.  typically  two  opposite   cotyledons   which   may  remain  within 

the  seed,  or  become  green  and  appear  above  the  ground  in 
germination.  These  plants  are  herbs,  shrubs  or  trees.  The 
ilowers  have  their  parts  usually  in  whorls  of  5  or  4.  The 
leaves  generally  have  reticulate  venation.  The  leaves  often 
have  stipules,  rarely  have  a  conspicuous  sheathing  leaf-base, 
and  usually  disarticulate  freely  from  the  parent  axis.  The 
primary  root  usually  persists  to  form  a  tap  root.  The  collateral, 
open  vascular  bundles  are  arranged  in  a  circle.  Secondary 
growth  in  thickness  takes  place  in  perennial  stems  by  means 
of  a  complete  cambium  ring  which  continually  forms  woody 
tissue  on  the  inside  and  cortical  tissue  outside.  Perennial 
stems  usually  possess  distinct  bark,  annual  rings,  medullary 
rays  and  numerous  vessels. 

Synopsis  of  133.  The  following  Key  will  now 

theSub-divi-  help  us  to  bring  into  focus  the  principal  groups  of  the  Vegetable 
which  we  have  considered  above  : — 


Kingdom. 

CRYPTOGAMS.     Plants  not  forming  true  seeds. 

("  Algae  .         .       Plants   contain- 

THALLOPHYTA.  ing      chloro- 

phyll, chiefly 
aquatic. 
Cellular  plants  with  no  differentia- -^    Bacteria.  -•  Very^ 


tion  into  stem  and  leaves.     An 


altogether  wanting  or  irregular. 


minute      plants  i    Colourless  plants 


alternation    of    generations    is          multiplying    by  L      without 


splitting.  i        chlorophyll. 

L   Fungi.  J 


BRYOPHYTA. 


f  Mosses         .         .~]  Plants  contain- 
|  Liverworts. —  i      ing       chloro- 

•Cellular  plants  usually    showing  a  |       ^mute      plants  |      phyll. 


differentiation  into  stem  and  j 
leaves,  and  with  a  distinct  alter-  | 
nation  of  generations,  the  t 
oophyte  constituting  "the  I 
plant." 

PTERIDOPHYTA. 


"Plants  showing  a  differentiation 
into  stem  and  leaves.  Possess 
true  vascular  bundles,  and 
with  a  distinct  alternation  of- 
generations,  the  sporophyte  con- 
stituting the  "  plant."  Chiefly 
herbaceous.  Only  a  few  of  the 
Ferns  attain  the  dimensions  of 
trees. 


often     with    no  f-  Chiefly     terres- 
|      d  i  S  e  rentiation  I      trial. 
i      into   stem    and  I 
L     leaves  J 

f  Ferns. — Plants  with  relatively  large 

leaves.       Sporophylls      are      not 

aggregated  into  cones. 
Horsetails. — Plants     with     distinct 

nodes  and   internodes  and   small 

leaves  in  whorls.     Peltate  sporo- 

phylls  aggregated    into   cone -like 

flowers. 
Club-mosses. — Plants      with     small 

scattered      leaves.       Sporophylls 


resemble  foliage  leaves,  but  are 
sometimes  aggregated  into 
cone -like  flowers. 


PHANEROGAMS.     Plants  forming  true  seeds. 


GYMNOSPERMS. 

Ovules  not  enclosed  in  an 
Trees  and  shrubs. 


ANGIOSPERMS. 


or 
no 


fCycadaceae. — "  Sago      Palms 
Cycads.         Flowers     have 
perianth  and  are  usually  cones. 
Ccniferae. — Freely  branching    trees 
ovary.  •{      and  shrubs.     Flowers  cones  with 

no  perianth. 

Gnetaceae. — Flowers   are    not  cones 
and       have       a        rudimentary 
^     perianth. 

r  Monocotyledons. — Embryo  with  one 
J      cotyledon. 

Ovules     enclosed    in    an    ovary.  ]  Dicotyledons. — Embryo     with     two 
Trees,  shrubs  and  herbs.  L     opposite  cotyledons. 


L2 


148 


CHAPTER  III.— THE  ORIGIN  OF  SPECIES. 

struggle  for  134.  On  all  sides  around  us,   in 

existence,      nature,  the    great    battle  of   life    is    always    going    on,     or, 
as   it    is   often   termed,     the   struggle  for    existence.      On  all 
sides   we    find    organisms    directly   destroying    others,  carni- 
vorous animals  preying  on  other  animals,  some  plants,   such 
as  fungi,   killing  and  feeding  on  other  plants,   while  every- 
where animals  are    found    injuring    and    destroying   plants. 
Not    so    obvious,    but    no    less    severe,    is    the    competition 
between   similar  organisms  for  the   same  kind  of    food.     This 
is  usually  most  severe   between  organisms  which  require  the 
same    food    materials    and    similar    external    conditions    for 
their  existence.     To  realise  this  we  have  only  to  watch   the 
development    of    crowded    seedlings    in    a    nursery     seed-bed, 
or  in  the  forest.     The  roots  of  all  are  ramifying  in  the   same 
layers  of  soil  and  are  competing  with  each  other  for  the   same 
moisture  and  the  same  solutions  of  mineral  salts — the    stems 
of  all  are  competing  in  their  endeavours  to  expand  their  green 
leaves  in  the  same  light  and  air.     Some  individuals  are    soon 
seen  to  be  growing  more  slowly  than  their  vigorous    neigh- 
bours— their  leaves   turn   pale,   their   steins    grow   weak   and 
finally  they  die,  while  the  seedlings  around  soon  cover   over 
the   gaps   caused   by   their   disappearance,    leaving   no    trace 
of  those  which  have  been  starved  and  killed.     The   Forester, 
therefore,   in   order  to  secure  the   best   development  of    the 
individuals  belonging  to  his   valuable   species,   is   obliged    to 
interfere   in  this   struggle   between  plant   and   plant    and    to 
transplant   his    seedlings    and    to    clean    and   thin    his  young 
woods.     Realising,  also,  that  the  competition  between  plants 
belonging  to  different  species  is  often  scarcely  less  severe,  he 
takes     measures    to    prevent     undesirable    species    interfering 
with  the    healthy  development  of    his  valuable  trees,   while 
the    agriculturist    carries    out    essentially   similar    operations 
in  his  fields,  by   eradicating  the  "weeds"   which,  year  after 
year,  appear,  in  addition  to  those  plants  which  alone  he  wishes 
to  cultivate.     In  order  to  maintain  itself,  also,  every  organism 
is  obliged  not  only  to  fight  against   other  living   organisms, 
but,     in    addition,     to    struggle     against    the    unfavourable 
factors  of  its  non-living  environment,  e.g.  climatic    influences, 
great  heat  and  cold,  drought,   excessive   moisture,   and   so  on. 
The  importance  and  universality  of   this   natural  phenomenon 
of  the  struggle    for   existence  is  perhaps  best  realised  by   a 


consideration  of  the  great  powers  of  increase  possessed  by  all 
living  organisms.  It  has  been  calculated,  in  the  case  of  a  small 
herbaceous  plant,  about  two  feet  in  height,  which  produces 
on  an  average  10,000  seeds  annually,  that,  if  all  the  seeds 
developed  and  produced  fertile  plants,  each  of  which  in  their 
turn  ripened  a  similar  number  of  seeds  and  so  on,  all  the 
dry  land  on  the  earth  would  be  completely  occupied 
by  such  plants  at  the  end  of  five  years.  Many  plants,  also, 
produce  a  greater  number  of  seeds  than  the  example  given, 
and  an  ordinary  Tobacco  plant  is  calculated  to  produce,  on 
an  average,  360,000  seeds  annually.  That  no  single  species 
is  able  to  actually  monopolise  the  earth  in  this  way  is  due 
to  the  struggle  for  existence. 

"  While  the  offspring  always  exceed  the  parents  in  num- 
ber, generally  to  an  enormous  extent,  yet  the  total  number 
of  living  organisms  in  the  world  does  not,  and  cannot,  increase 
year  by  year.  Consequently  every  year,  on  the  average,  as 
many  die  as  are  born,  plants  as  well  as  animals ;  and  the 
majority  die  premature  deaths.  *  *  There  is  thus  a 
perpetual  struggle  among  them  which  shall  live  and  which 
shall  die  ;  and  this  struggle  is  tremendously  severe,  because  so 
few  can  possibly  remain  alive."* 

135.  It  has  been  noted  above  that  importance 

in  many   Floras  all    sub -divisions   of  the  species  are  indiscri-  of  s?b~ 
minately  termed  varieties.     Obviously  such  varieties  are  by  no 
means  all  of  the  same  value.     We  may  find  that  the  distinguish- 
ing   characters    of  a   particular  variety    are    always,    under 
varying  conditions  of  existence,   transmitted  unchanged  from 
the  parent  to  its  immediate   offspring,   that  the  variety  is, 
in    fact,    a    true    sub-species.     Since    the    characters    which 
distinguish  a  particular  sub-species  from  the  rest  of  its  species 
may  include  the    power  of  producing  a  valuable  commercial 
product,    immunity    from    particular    forms    of     injury    and 
disease  caused  by  fungi,  insects  or  other  injurious  factors  such 
as    frost,    and    so    on,    a    knowledge    of  such  sub-species  is 
obviously  of  great  importance  for  the  Forester,  as  well  as  for 
the  Gardener  and  Farmer.     Many  races,  again,  the  distinguish- 
ing   characters    of    which   only    remain    constant    under  cer- 
tain   conditions  of  existence  and  which  are  therefore  of  sub- 
ordinate importance  in  nature,  are  of  considerable   importance 
in  cultivation  where  the  conditions  of  their  existence  can  be 
regulated  so    as    to    keep    their    characters    constant.     The 
selection  of  the  most  suitable  sub-species   and  races  for  culti- 

*  Darwinism — by  A.  R.  Wallace,  page  11. 


150 

vation,  therefore,  plays  a  very  important  part  in  modern 
agriculture  and  horticulture.  In  a  single  field  of  wheat,  or 
other  cereal,  several  different  sub-species  may,  and  usually  do, 
co-exist.  The  most  desirable  of  these  may  be  observed  to 
constitute  a  very  small  proportion  of  the  crop  when  culti- 
vated without  special  care.  The  reason  for  this  is  usually 
to  be  sought  in  the  struggle  for  existence,  the  conditions  being 
more  favourable  to  the  less  valuable  sub-species  and  thus 
enabling  them  to  overcome  and  kill  out  their  less  vigorous 
neighbours,  just  as  we  have  seen  to  be  the  case  in  a  crowded 
seed-bed,  and  this  is  especially  the  case  in  unfavourable 
seasons.  The  Farmer,  therefore,  carefully  isolates  and  saves 
the  seeds  of  the  desirable  sub-species,  say  those  with  the 
largest  ears  and  the  biggest,  most  numerous  grains,  and  cul- 
tivates it  separately,  treating  all  other  kinds  of  the  cereal 
as  weeds,  and  endeavouring,  as  far  as  possible,  to  eliminate 
them  from  his  fields. 

Again  the  distinguishing  characters  of  some  races  of  plants 
do  not  remain  constant  when  they  are  grown  in  the  vicinity 
of  other  plants  by  which  they  are  easily  fertilised.  This 
does  not  prevent  such  plants  being  of  great  value  to  the  horti- 
culturist who  endeavours  to  isolate  such  races  and  to  pre- 
vent, as  far  as  possible,  intercrossing  with  allied  plants.  The 
beds  in  which  such  plants  are  grown,  however,  can  rarely 
be  separated  sufficiently  to  prevent  occasional  crossing  by 
bees,  which  carry  the  pollen  from  one  to  the  other. 

Frequently,  therefore,  the  seed  of  some  of  our  best  garden 
flowers,  as  sold  in  the  market,  cannot  be  guaranteed  as  pure, 
some  of  the  resulting  seedlings  invariably  showing  aberrant 
characters,  owing  to  their  parents  having  been  crossed  by 
allied  forms. 

136.  Plants  raised   from  the  seed 

Hybrids,  produced  by  the  crossing  of  two  plants  belonging  to  distinct 
species,  or  varieties,  are  called  hybrids,  or  bastards,  and 
are  usually  distinguished  as  species-hybrids  and  variety- 
hybrids.  Very  commonly  the  crossing  of  individuals  belong- 
ing to  distinct  species  is  accompanied  by  entire  or 
partial  sterility,  i.e.  there  is  less  chance  of  fertile  seed 
resulting  from  the  cross  and  of  fertile  individuals  being  deve- 
loped from  such  good  seed  as  may  have  been  produced, 
than  is  the  case  in  normal  fertilisation.  Such  sterility  varies 
greatly  in  degree,  for  while  crosses  between  very  nearly  related 
species,  or  between  varieties  of  the  same  species,  may  be, 
and  usually  are,  perfectly  fertile,  crosses  between  specieSv. 


151 

which  are  not  closely  allied  are  characterised  by  a  consider- 
able degree  of  sterility  and  are  often  entirely  sterile.  As 
fertilisation  consists  essentially  in  the  union  of  two  sexual 
cells,  a  portion  of  the  protoplasm  of  each  cell  entering  into 
the  constitution  of  the  embryo  which  results  from  such  a 
union,  we  should  naturally  expect  the  distinguishing  charac- 
teristics of  each  parent  to  become  more  or  less  apparent 
in  the  hybrid.  This  is  the  case  and  the  hybrid  is  found  either 
to  be  intermediate  between  the  parents,  the  characters  of 
both  parents  being  combined  in  it,  or  else  to  resemble  one 
parent  more  than  the  other.  Occasionally  also  it  may  pre- 
sent entirely  new  characters.  In  some  cases  the  result  of 
hybridisation  is  different  according  to  which  parent  is  chosen 
as  the  father  or  mother  respectively.  Thus  a  cross  may 
only  be  possible  when  an  individual  of  a  certain  species  is 
always  chosen  as  the  father,  the  reciprocal  cross  with  the 
same  individual  as  the  mother  being  sterile. 

Again,  the  hybrid  in  some  cases  is  always  found  to  re- 
semble one  parent,  e.g.  the  father,  more  than  the  other.  Fertile 
hybrids  are  those  which,  when  fertilised  by  their  own 
pollen  or  by  that  of  another  individual  of  the  same  hybrid 
cross,  produce  fertile  seed  from  which  fertile  plants  are 
developed.  Fertile  hybrids  may  be  constant,  as  regards  their 
distinguishing  characters,  from  the  first,  but  in  the  majority 
of  cases  they  are  found  to  split,  i.e.  to  revert  to  the  parent 
forms  in  subsequent  generations.  In  such  cases  of  reversion, 
however,  a  few  individuals  are  often  found  which  remain 
constant  hybrids,  with  characters  distinguishing  them  from 
both  parents.  A  hybrid  may  be  successfully  crossed  not 
only  with  another  hybrid  but  with  one  of  its  parents ;  if  this 
is  repeated,  the  derivative  hybrid  obtained  from  such  a  cross 
being  again  crossed  with  the  same  parent  and  so  on,  the  off- 
spring are  found  to  resemble  that  parent  more  and  more, 
and  finally  to  completely  revert  to  it.  The  hybrid,  instead 
of  being  crossed  with  one  of  its  parents,  may  be  crossed  with 
an  individual  belonging  to  an  entirely  distinct  species  and 
this  operation  may  also  be  repeated.  In  this  way  it  has 
been  found  possible  to  obtain  a  derivative  hybrid  in  which 
the  characters  of  six,  or  even  more,  different  species  are  com- 
bined. Hybrids  occur  in  nature  but  not  very  frequently ; 
probably  this  is  partly  due  to  the  fact  that,  in  the  majority 
of  cases,  where  the  flowers  of  a  plant  are  simultaneously  pol- 
linated by  different  plants,  the  pollen  o£  an  individual  belong- 
ing to  the  same  species  is  prepotent,  i.e.  it  is  able  to  accom- 


152 

plish  fertilisation  quicker  than  the  pollen  of  plants  belonging 
to  different  species.  The  action  of  such  foreign  pollen  being 
thus  excluded,  hybridisation  is  prevented.  In  other  cases 
the  plants  resulting  from  hybridisation  may  be  unable  to 
establish  themselves  in  nature,  owing  to  the  severity  of  the 
struggle  for  existence  with  other  plants.  Among  the  char- 
acters which  are  frequently,  possessed  by  hybrids  and  which 
serve  to  distinguish  them  from  their  parents  are  an  increased 
tendency  to  variation,  a  more  luxuriant  and  vigorous  vege- 
tative growth,  and  larger,  more  brilliantly  coloured  flowers, 
which  also  often  tend  to  become  double.  For  this  reason 
alone  hybrids  are  important  in  horticulture.  The  great 
importance  of  hybridisation,  however,  depends  chiefly  on  the 
fact  that  it  affords  a  means  of  combining  in  one  plant  the 
valuable  attributes  of  several,  and  a  species  yielding  a  valuable 
commercial  product  which  is  liable  to  damage  by  frost,  or  to  a 
particular  form  of  disease,  may  thus  be  rendered  hardy  by 
crossing  it  with  an  allied,  but  hardy,  species,  the  hybrid  off- 
spring, while  yielding  the  valuable  product  of  one  parent, 
possessing  also  the  hardy  attributes  of  the  other.  If  such 
a  desirable  hybrid  is  obtained  it  would  of  course  be  carefully 
isolated  and  cultivated  in  such  a  way  as  to  prevent,  as  far 
as  possible,  intercrossing  with  other  plants.  At  the  same 
time  it  must  be  remembered  that  the  practical  attainment 
of  such  a  result  is  attended  with  great  difficulties,  the  greatest 
obstacles  to  successful  work  in  this  direction  being  the 
inconstancy  of  many  hybrids  and  the  comparative  sterility 
of  others.  For  successful  hybridisation  the  flowers  of  the 
plant  chosen  as  the  mother  (if  hermaphrodite)  should  be 
deprived  of  their  stamens  before  the  latter  are  mature,  the 
flowers  then  being  covered  with  paper  caps  to  prevent  the 
access  of  insects  and  possible  fertilisation  by  the  pollen  of 
other  plants.  With  a  clean  camel-hair  brush  the  pollen  is 
then  taken  from  the  plant  selected  as  the  father  and  gently 
and  lightly  applied  to  the  stigmas  of  the  protected  mother 
flowers.  The  caps  are  then  replaced  on  the  latter.  If  the 
cross  has  been  successful  the  flower  will  quickly  wither,  and 
if  this  does  not  occur  the  cross  should  be  repeated. 

137-  If  we  examine  a  large  number 
Variability,  of  adult  plants  all  raised  from  the  seed  of  a  single 
individual,  we  shall  find  a  few  exceptionally  tall,  and 
a  few  exceptionally  short,  specimens,  while  the  great 
majority  will  be  seen  to  be  intermediate  between  these  two 
extremes,  of  moderate,  or  medium  height,  and  differing  very 


153 

slightly  from  each  other  in  this  respect.  A  large  number 
of  adult  men  placed  in  a  row  would  give  a  similar  picture 
•of  variability.  Again  if  we  examine  and  carefully  measure 
the  leaves  on  a  tree,  we  find  a  few  unusually  large,  and  a  few 
unusually  small,  specimens,  while  the  great  majority  are 
intermediate  and  of  moderate  dimensions.  The-same  thing 
•occurs  in  the  shape  and  size  of  seeds  and  fruits.  Numbers 
also  vary  in  the  same  way,  as  is  seen  in  the  number  of  lateral 
veins  in  leaves  and  leaflets.  Small  numbers,  however,  are 
usually  more  constant  than  large  numbers  and  a  considerable 
degree  of  constancy  usually  characterises  the  numbers  of  the 
different  floral  organs.  Such  characters  as  the  percentage 
of  sugar  in  the  sugarcane,  or  of  starch  in  potatoes,  as  well  as 
the  power  of  resistance  to  frost,  and  so  on,  obey  the  same  rule, 
and  we  may  say  that  this  phenomenon  of  Fluctuating  Vari- 
ability is  almost  universal.  This  type  of  variation  is  obviously 
•directly  dependent  to  a  great  extent  on  nutrition,  and  therefore, 
indirectly,  depends  on  the  factors  which  together  constitute  the 
plant's  environment.  On  vigorous,  well-nourished  coppice 
shoots  the  leaves  are  as  a  rule  considerably  larger  than  on 
normal  branches,  while  gardeners  know  that,  by  diminishing 
the  total  number  of  flowers  or  fruits  on  a  plant,  and  by  thus 
increasing  the  amount  of  food  available  for  the  remainder,  the 
size  of  the  latter  can  be  materially  increased.  The  various 
factors  of  the  environment  also  Have  a  marked  effect.  A  plant 
accustomed  to  a  damp  locality,  on  being  transferred  to  a  dry 
one,  may  be  so  changed  in  appearance  as  to  resemble  another 
species.  In  such  cases  the  size  of  the  leaves  and  the 
growth  in  length  of  all  shoots  are  often  greatly  reduced 
and  thorns  and  spines  are  frequently  developed.  The  common 
Brinjal  (Solanum  Mdongena),  for  instance,  when  cultivated  in  a 
well-watered  garden,  has  large  leaves  which  are  often  almost 
unarmed.  It  may,  however,  be  often  seen  as  an  escape  from 
cultivation,  in  dry  rocky  places,  with  small  and  intensely 
prickly  leaves.  In  India  also  we  have  several  examples  of 
plants  which,  in  dry  localities,  are  erect  shrubs,  or  small  trees, 
and  in  damp  situations  climbers — such  as  Toddalia  aculeata  and 
Alangium  Lamarckii.  The  nature  of  the  soil  also  frequently 
appears  to  affect  plant  characters,  and  individuals  growing  in 
soil  containing  large  quantities  of  lime,  or  other  substances, 
may  differ  very  considerably  from  individuals  of  the  same 
species  growing  in  ordinary  soil.  Light  is  also  a  factor  affect- 
ing the  form  of  plants,  although  it  is  sometimes  difficult  to 
•decide  how  far  an  observed  result  is  to  be  ascribed  to  any  one 


factor,  such  as  light,  and  to  what  extent  other  factors,  such  as- 
available  moisture,  may  have  contributed  to  it.  In  shady  situ- 
ations, in  the  case  of  one  and  the  same  species,  not  only  may 
leaves  be  found  to  be  thinner,  and  often  of  a  different  colour,, 
consistency,  size,  and  shape,  than  they  are  in  places  exposed  to 
bright  sunlight,  but  their  anatomical  structure  may  also  be 
fundamental  i y  changed,  and  we  find  sun-leaves  with  an  epidermis 
devoid  of  chlorophyll,  provided  with  a  thick  cuticle,  and  with 
well-developed  palisade  tissue,  while  shade-leaves  often  have 
no  palisade  tissue  and  an  epidermis  with  very  thin  walls  and 
provided  with  chlorophyll.  Anatomical  variations  also  may 
be  found  in  the  petioles,  stems  and  roots.  Cases  are  known 
of  so-called  dimorphic  species  which  are  able  to  maintain 
themselves  on  high  mountains  and  also  in  the  lowlands,  or 
on  dry  land,  as  well  as  under  the  water,  and  variability  is 
thus  seen  to  be  a  very  useful  character,  inasmuch  _  as  vari- 
able plants  are  thus  enabled  to  adapt  themselves  to  varying 
conditions  of  existence  and  to  widely  extend  their  area  of 
distribution.  Variations  of  the  kind  now  under  discussion 
appear  to  us  to  be  temporary  and  inconstant,  and  individuals 
belonging  to  the  alpine — or  aquatic — form  of  a  dimorphic 
species,  after  existing  for  many  generations  on  the  moun- 
tains or  under  water,  are  found  to  assume  the  characters 
of  the  lowland — or  terrestrial — form,  respectively,  when  cul- 
tivated in  the  plains  or  on  dry  land.  At  the  same  time  cases 
are  known  in  which  plants  .under  the  influence  of  certain  con- 
ditions have  undergone  a  slow  but  progressive  change  and 
have  thus  acquired  definite  characteristics  which  have 
become  constant  and  hereditary.  With  regard  to  this  point 
the  study  of  bacteria  is  of  special  value  seeing  that,  in  their 
case,  "  experiments  can  be  readily  extended 

over  a  far  greater  number  of  generations  than  in  the  case  of 
flowering  plants,  for  a  bacterium  which  divides  once  an  hour 
passes  through  as  many  generations  in  ten  days  as  an  annual 
plant  does  in  240  years."*  By  continued  cultivation  under 
special  conditions,  it  has  been  found  possible  to  permanently 
eliminate  in  certain  species  of  bacteria  the  power  of  producing 
poisons,  pigments,  and  even  the  important  character  of  the 
power  of  spore  production.  Such  changes  do  not  occur  sud- 
denly, but  only  gradually  become  fixed  and  hereditary  and 
if,  after  short  exposure  to  the  special  conditions,  the  bacteria 
are  returned  to  their  original  environment,  the  power  of  pro- 

*  Physiiogy  of  Plants,  by  Dr  W.  Pfeffer,  Volume  11.  pages  192—193. 


155 

ducing  poison,  pigment,  or  spores,  returns.  Attention  has 
hitherto  been  confined  to  the  so-called  fluctuating  variability, 
and  it  must  be  remembered  that,  although  in  a  series  of  such 
variations,  the  two  extreme  forms  may  be  widely  dissimilar, 
these  are  always  linked  together  by  a  number  of  intermediate 
forms  which  differ  very  slightly  from  each  other;  there  is 
no  gap  in  the  chain  and  no  sudden  and  considerable  variation, 
only  a  number  of  very  slight  differences.  We  have  also  seen, 
that  such  variations  are  inconstant,  although  in  some  cases 
at  least,  under  suitable  conditions,  they  can  be  gradually  fixed 
and  rendered  hereditary. 

In  contrast  with  such  variations  are  those  known  as  spon-  Spontaneous 
taneous  variations,  sports,  or  mutations.  Our  knowledge  of  Variations, 
these  and  of  their  mode  of  occurrence  in  nature  depends 
chiefly  on  the  experiments  carried  out  by  Professor  Hugo  De 
Vries.  He  has  demonstrated  that  plants  of  the  species  Oenothera 
Lamarckiana  in  addition  to  producing  normal  offspring  bearing 
the  characteristic  marks  of  their  parents,  produced  also  con- 
siderable numbers  of  individuals  which  possessed  definite  and 
appreciable  characters  not  seen  in  their  parents.  Such  indivi- 
duals are  termed  mutants ;  they  suddenly  come  into  existence, 
there  is  no  progressive  change,  and  no  intermediate  forms  are 
found  linking  them  with  the  species  from  which  they  spring, 
they  are  due  in  fact  to  a  sudden  variation,  i.e.  a  mutation.  A 
mutant,  when  isolated  and  fertilised  with  its  own  pollen,  is 
found  to  be  constant  from  the  time  of  its  origin  and  to  transmit 
its  essential  characters  truly  to  its  offspring.  This  constancy 
sharply  distinguishes  mutations  from  the  inconstant  fluctuating 
variations.  The  species  mentioned  was  found  to  produce  several 
different  types  of  mutants,  differing  in  various  ways  from  the 
original  form,  several  individuals  of  each  type  usually  appeared 
simultaneously,  while  each  type  was  liable  to  be  reproduced  at 
intervals  by  the  parent  species.  The  causes  of  mutations  are  at 
present  unknown.  The  great  majority  of  species  in  nature 
are  fixed  and  constant,  but  from  the  case  of  this  Oenothera 
it  appears  possible  that  all  species  at  some  period  of  their 
existence  throw  off  mutants  in  considerable  numbers.  Such 
a  mutating  period  probably  forms  only  a  small  part  of  the 
total  life  of  a  species.  Such  a  period  moreover  does  not  neces- 
sarily involve  the  death  of  the  parent  species  which,  as  seen 
in  this  Oenothera,  in  addition  to  producing  mutants,  ma}' 
continue  to  transmit  its  specific  characters  truly  to  the  majority 
of  its  offspring.  Mutations  are  not  entirely  confined  to  such 
mutating  periods;  occasional  mutants  may  arise  at  any  time, 


156 


Variations 


but  so  far  as  is  at  present  known,  they  are  rare.  The  occur- 
rence of  double  flowers  in  species  with  usually  single  flowers, 
of  regular  flowers  in  species  with  usually  irregular  flowers, 
of  divided  leaves  in  species  with  entire  leaves,  of  a  weeping, 
or  fastigiate,  habit  in  trees  and  shrubs  with  an  erect,  or  spread- 
ing, crown,  respectively,  are  among  the  most  common  examples 
of  mutations  at  present  known. 

Monstrosities.  Very  striking  variations  which  give  the  impression  of  an 
altogether  abnormal  structure  are  called  monstrosities.  They  are 
often  termed  malformations  and  frequently  give  rise  to  diseased 
conditions.  They  are  of  rare  occurrence  in  nature  ;  the 
cauliflowers  and  turnips  of  cultivation  are  believed  to  have 
arisen  from  monstrosities.  Finally  it  must  be  noted  that 
remarkable  variations  in  the  form  of  plants  may  be  caused  by 

Mutilation.  insects  and  fungi,  which  may,  for  instance,  be  responsible 
for  the  peculiar  structures  known  as  "  galls  "  and  "  witches' 
brooms."  Mutilation  also  may  exercise  a  far-reaching  effect 
such  as  is  seen  in  plants  which  are  continually  browsed  cr 
grazed  by  animals,  while  some  annual  species  may  be  made 
perennial  by  removing  the  flower  buds  and  thus  preventing 
their  reproduction. 

Heredity.  138.  By  heredity  is  understood  the 

transmission  of  characters  from  a  parent  to  its  offspring. 
This  transmission  is  rendered  possible  in  plants  by  means 
of  their  powers  of  sexual  and  asexual  reproduction.  Many 
variations  can  only  be  transmitted  truly  to  the  offspring 
by  asexua  propagation,  and  for  this  reason  individuals 
which  exhibit  a  desirable  variation  are  usually  propagated 
in  horticulture  by  asexual  methods.  This  is  particularly  the 
case  with  hybrids  which  are  frequently  very  inconstant  if 
propagated  by  seed.  The  younger  a  plant  or  organ  is,  the  greater 
will  be  the  effect  of  any  factor  which  is  capable  of  influencing  its 
development,  and  the  early  stages  of  the  growth  of  the  embryo 
constitute  the  most  susceptible  period.  If  this  period  could 
be  cut  out,  as  it  were,  from  a  plant's  life  history  we  should 
expect  it  to  be  less  subject  to  variation  and  to  resemble 
its  parent  more  closely  than  would  be  the  case  if  the  individual 
were  raised  from  seed,  and  this  is  precisely  what  is  effected  by 
asexual  propagation.  It  must,  however,  never  be  forgotten 
that  in  sexual  reproduction  effected  by  the  crossing  of  two 
distinct  individuals  a  portion  of  the  protoplasm  of  each  parent 
enters  into  the  composition  of  the  young  plant,  which  must 
therefore  inherit  something  from  its  father  and  something  from 


157 

its  mother.  When  both  parents  possess  similar  characters  the 
offspring  will  naturally  tend  to  strongly  resemble  the  mother 
plant,  although  this  depends  to  some  extent  on  the  conditions 
under  which  the  young  plant  develops,  but  if  this  is  not  the 
case  the  results  of  the  cross  may  be  very  various,  as  has  been 
noted  in  the  remarks  on  hybrids.  We  have  also  seen  that, 
when  a  hybrid  form  is  repeatedly  crossed  with  one  of  its 
parents,  it  tends  to  quickly  revert  to  that  parent  form,  and  in 
this  way  sexual  reproduction  may  help  to  eliminate  new  vari- 
ations and  to  keep  species  constant  in  nature.  A  character 
which  may  appear  to  have  been  lost  is  frequently  found  to  be 
really  only  dormant,  or  latent,  and  although  it  may  be 
inherited  in  this  state  during  many  generations  it  is  always 
likely  to  reappear  and  again  become  active.  This  phenomenon 
of  the  reappearance  in  a  plant  of  an  ancestral  quality  which 
has  been  latent  in  its  parents  is  termed  atavism. 

139.  We  have  seen  above  that  a  Selection. 

large  proportion  of  all  living  organisms  which  come  into 
existence  must  perish  prematurely  and  that  in  nature  those 
which  are  best  adapted  to  the  conditions  under  which 
they  exist  are  selected,  or  chosen,  as  it  were,  by  nature, 
and  survive,  while  the  remainder  are  rejected  and  perish.  This 
Survival  of  the  Fittest,  therefore,  is  seen  to  be  the  result  of  the 
so-called  Natural  Selection.  Selection  has  been  well  likened  to  a 
sieve,  only  those  individuals  which  possess  the  necessary 
qualifications  are  able  to  remain  and  are  selected,  while  the 
remainder  pass  through  the  sieve  and  are  rejected.  It  is  advis- 
able to  distinguish  two  kinds  of  selection,  viz.  that  which 
selects  a  particular  sub-species,  or  race,  from  among  other  sub- 
species and  races,  and  that  which  selects  particular  individuals 
within  one  and  the  same  sub-species,  or  race.  An  example  of 
the  first  occurs  when,  in  a  field  of  wheat,  a  single  sub-species  is 
selected  from  among  the  others  growing  with  it  and  is  then 
isolated  and  separately  propagated.  The  same  thing  occurs  in 
nature  when,  from  a  number  of  mutations,  only  those  which  are 
best  adapted  to  the  conditions  of  life  are  selected  and  survive, 
while  the  others  perish.  In  addition  to  this,  however,  we  know 
that,  within  one  and  the  same  sub-species,  or  race,  there  is 
always  fluctuating  variability — that  no  two  individuals  are  ever 
exactly  alike — and  that  the  same  quality  or  character  is 
exhibited  in  different  degrees  by  different  individuals.  Hence 
it  might  be  inferred  that  breeding  entirely  from  those  indivi- 
duals which  exhibit  a  desirable  character  in  a  marked  degree 
would  result  in  the  production  of  similarly  characterised 


158 

offspring ;  and  that  a  race  could,  in  this  way,  be  improved  and 
•confined,  as  it  were,  to  a  few  of  the  best  individuals,  the  large 
number  of  moderate,  or  inferior,  individuals  usually  occurring 
in  the  race  being  thus  excluded.     To   a  great  extent  this  is 
found  to  be  the  case,  and  modern  horticulture  and  agriculture 
depend  largely  on  these  two  principles  of  (1)  the  selection  of  the 
most  suitable  race  or  sub-species  and  (2)  the  improvement  of  the 
selected  strain  by  breeding  only  from  the  best  individuals.     In 
cases  of  fluctuating  variability  the  greater  the  number  of  indivi- 
duals examined  the  more  chance  there  is  of  finding  an  indivi- 
dual which  exhibits  a  particular  character  in  an  extreme  degree 
and  for  this  reason  large  cultures  are  often  resorted  to  in  horti- 
culture.    Professor  Hugo  De  Vries  records    a    case   in    which 
40,000  plants  were    cultivated,  only  a  single  individual   being 
finally  selected  from  this  multitude  for  further  propagation,  all 
others  being  destroyed.     Such  an  individual,  when  found,  can 
only  be  propagated  with  certainty  by    asexual   reproduction. 
Many  plants,  however,  cannot  easily  be  propagated  in  this  way 
and  such  large  cultures  are  very  inconvenient.      In  addition   to 
this  the  asexual  method  of  reproduction   affords   no   means  of 
effecting  further  considerable  improvement  of  the  race.     It  is, 
however,  found  'that   such   desirable    variations    can  also    be 
obtained  and  to  a  certain  extent  be  propagated  by   sexual 
methods.      Experience  shows  that  if  individuals  exhibiting  a 
certain  character  in  a  marked  degree  are  chosen   as    parents, 
although  the  average  of  the  offspring  raised  from  their  seed  do 
not  exhibit  this  character  in  so  high  a  degree  as  their  parents, 
they  do  show  a  distinct  improvement   in   this  respect  and  are 
superior  to  the  average  of  the  race  from  which  they  have  arisen. 
By  continuing  this  method   for  some  generations    the    same 
degree  of  improvement  can  be  .obtained  with  comparatively  few 
individuals,  which  could   only  be  obtained   in   one  generation 
by  cultivating  a  very  much  larger  number  of  plants.     Moreover 
the  sexual  method  possesses  the  great  advantage  of  enabling  us 
to  continue  the  work  of  improvement.     This  principle  is  a  very 
important  one  in  modern  agriculture.     The  fact  that   not   one, 
but   several,   characters   have   almost  invariably   to  be  taken 
into    consideration   is  one  of  the  great  difficulties  in   the   way 
•of  successful  practical  work  in  this  line.     If,  when  selecting 
certain   individuals   in   a   race  of  wheat,  we  were  only  to  pay 
attention  to  the  quantity  or  quality  of  grain  yielded,  we  might 
select  those  plants  which  are  also  very  susceptible  to  disease  or 
climatic  influences,   and   the   selection   would   thus  effect  no 
practical  improvement.     It  .is  generally  recognised  that  the 


159 

best  results  are,  as  a  rule,  obtained  by  selecting  not  those 
individuals  which  exhibit  the  best  visible  characters,  but  those 
plants  which  are  found  to  produce  the  most  desirable  offspring ; 
in  other  words,  the  value  of  a  plant  is  gauged  not  by  its  visible 
attributes  but  by  its  power  to  transmit  desirable  qualities  to  its 
descendants.  For  this  purpose  the  seeds  of  each  individual  are 
sown  separately  and,  to  obtain  an  average  value  for  the  offspring 
of  each  individual,  100  seedlings  are  carefully  examined  and  their 
qualifications  considered.  Only  those  groups  which  contain  the 
greatest  number  of  desirable  individuals  are  then  selected  for 
further  cultivation.  This  method  is  based  on  the  idea  that  a 
single  plant,  or  even  a  few  individuals,  may  owe  their  qualifica- 
tions not  to  any  inherited  attribute  but  to  extraordinarily 
favourable  conditions  to  which  they  have  been  subjected  during 
their  own  life-time.  Although  great  improvements  have  been 
wrought  on  the  lines  indicated  above,  no  permanent  improve- 
ment has,  up  to  date,  been  effected  and  no  new  and  constant 
race  has  been  produced.  Experience  hitherto  gained  has  shown 
that  whatever  standard  of  selection  is  adopted,  a  limit  to  the 
improvement  effected  is  soon  reached  beyond  which  it  is  impos- 
sible to  go,  and,  further,  that  when  a  desirable  degree  of 
improvement  has  been  reached,  this  can  only  be  maintained  by 
continued  selection,  for  if  selection  ceases,  the  race  soon  reverts 
to  its  original  type. 

140.  Linnaeus,  the  most  famous  of  Origin  of 
the  early  botanists  who  worked  at  classification,  thought  t 
each  species  had  been  separately  created  at  the  beginning  of  the 
world  and  that  the  individuals  of  a  species  were  only  capable  of 
producing  other  individuals  like  themselves,  i.e.  that  existing 
species  had  remained  unchanged  from  the  creation.  Charles 
Darwin,  however,  subsequently  proved  thatthis  was  not  the  case 
and  that,  from  one  species,  other  and  totally  distinct  species 
could  arise  in  the  progress  of  time.  The  theory  of  evolution  in 
the  animal  and  vegetable  kingdoms  is  now  universally  accepted 
as  correct,  as  is  also  the  fact  that  this  evolution  is  mainly 
dependent  on  two  factors,  which  are : — 

(1)  the  variability  of  living  organisms,  and 

(2)  the  struggle  for  existence,  owing  to  which  only  those 

organisms  are  able  to  survive  which  are  most  per- 
fectly adapted  to  the  conditions  under  which  they 
exist,  which  are  best  able  to  withstand  unfavourable 
conditions  of  soil,  or  climate  and  which  can  best 
hold  their  own  against  other  injurious  organisms. 


160 

Such  a  view  also  satisfactorily  accounts  for  the  fact 
that  everywhere  in  nature  we  find  animals  and 
plants  exhibiting  the  most  beautiful  adaptations  to 
the  conditions  of  their  environment,  by  means  of 
which  they  are  favoured  in  the  struggle  for 
existence. 

Opinions,  however,  differ  as  to  which  type  of  variation  has 
played  the  most  important  part  in  the  history  of  species.  The 
variations  which  are  believed  to  be  most  important  from  this 
point  of  view  may  be  divided  into  three  main  classes  as 
under : — 

(1)  Variations  due  to   the   crossing   of  different 

forms,  i.e.  to  hybridisation. 

(2)  Fluctuating  variations. 

(3)  Mutations. 

It  may  also  be  premised  that,  in  order  to  form  the  beginning 
of  a  new  species,  a  plant  must  be  able  to  maintain  itself  under 
the  conditions  of  existence  to  which  it  is  exposed  in  nature,  it 
must  be  able  to  transmit  its  essential  characters  truly  to  its  off- 
spring, it  must  occur  in  considerable  numbers  and  be  sufficiently 
fertile  to  produce  numerous  offspring.  Such  a  plant  may  possibly 
result  from  hybridisation  alone.  As  regards  fluctuating  vari- 
ability, experience  with  bacteria  indicates  that,  if  the  conditions, 
to  the  stimulating  action  of  which  the  variation  must  be 
ascribed,  are  kept  constant,  in  some  cases,  at  least,  a  new  and 
constant  form  may  be  gradually  evolved  which  may  also  be 
able  to  establish  itself  as  a  new  species.  In  the  case  of  plants 
reproducing  sexually,  such  a  progressive  change  would  be 
aided  by  natural  selection,  for  all  individuals  not  exhibiting  the 
variation,  which,  under  the  circumstances,  is  favourable  to 
their  existence,  would  perish,  while  only  those  possessing  the 
variation  in  a  high  degree  would  remain  to  cross  and  produce 
offspring.  We  have  seen  also  that  new,  fertile,  and  constant, 
forms  in  considerable  numbers  may  arise  by  mutation,  and 
those  which  are  able  to  survive  in  the  struggle  for  existence 
may  form  the  beginning  of  new  species.  Just  as  the  small 
fluctuating  variations  may  be  gradually  accumulated  and 
increased  in  the  same  direction  by  natural  selection,  so 
may  be  also  mutations,  for,  providing  the  conditions  remain 
constant,  of  all  the  mutants  produced  during  successive 
mutating  periods  only  those  mutants  which  vary  in  the 
same  direction  will  be  able  to  survive,  and  thus  evolution 
will  continue  along  the  same  lines,  just  as  with  fluctuating 


161 

variations,  until  those  marked  differences  are  produced  which 
distinguish  the  actually  existing  species  in  nature.  Valuable 
results,  however,  in  the  way  of  producing  new  forms  have  been 
obtained  in  horticulture  and  agriculture  by  a  combination  of 
the  operations  of  selecting  the  best  strains,  of  crossing  distinct 
strains,  and  of  selecting  for  propagation  those  individuals  which 
exhibit  the  most  desirable  fluctuating  variations,  and  we  must, 
therefore,  be  careful  not  to  hastily  ascribe  to  any  one  type  of 
variation,  or  to  any  single  factor,  pre-eminent  importance  in  the 
evolution  of  species  in  nature. 


162 


CHAPTER  IV.— COLLECTION   AND    PRESERVATION  OF 

SPECIMENS. 

• 

Collection  141.  It  is  of  the  first  importance 

of  Speci-       that  specimens  should  be  gathered  which  exhibit  those  parts 
mens.  an(j    organs,    the  characters  of    which    are   included   in   the 

descriptions  and  keys  of  our  Floras.  So  far  as  possible  the 
specimens  selected  should  indicate  the  average  condition  and 
the  range  of  variability  of  the  important  organs  found  on  one 
and  the  same  individual  and  on  the  different  individuals  of  the 
same  species  found  growing  together,  and  in  any  case  should 
not  be  confined  to  those  parts,  or  individuals,  which  show 
extreme  variations,  or  abnormal  development.  The  specimens 
must  not  be  allowed  to  wither  before  they  are  laid  in  the  drying 
papers  and  are  usually  placed,  immediately  after  they  are 
gathered,  in  a  tin  case  which  should  be  kept  as  cool  as  possible. 
The  specimens  having  been  laid  in  the  drying  papers  are 
usually  placed  between  two  boards  with  a  weight  on  the  top. 
For  camp  perhaps  the  most  portable  and  useful  press  is  one 
consisting  of  two  pieces  of  strong  wire  lattice  of  convenient 
size  which  can  be  firmly  bound  together  with  straps  and 
buckles,  or  with  a  piece  of  thin  rope. 

The  specimens  are  kept  in  this  until  dry,  the  drying  papers 
being  of  course  changed  and  dried  when  necessary,  and  they  can 
then  be  packed  in  boxes  in  layers  between  sheets  of  paper,  care 
being  taken  to  keep  them  from  damp.  A  small  notebook 
should  always  be  carried  in  which  sunh  details  as  the  vernacular 
name,  dimensions,  habitat,  habit,  and  general  appearance  of 
the  living  plant  should  be  invariably  noted,  on  the  spot,  and 
not  left  to  memory.  To  each  specimen,  before  being  placed  in 
the  drying  papers,  should  be  fastened  a  small  label  with  a 
number,  and  great  care  should  be  taken  to  see  that  these  labels 
do  not  become  subsequently  detached.  A  journal  should  also 
be  kept  up  in  which  the  specimens  collected  should  be  entered 
daily,  serially,  under  their  respective  numbers,  with  the  date. 
Here  also  should  be  recorded  the  details  noted  in  the  field,  the 
name  of  the  plant  and  natural  order  (if  determined),  the  results 
of  the  inspection  of  the  specimens  with  diagrams  or  sketches  of 
the  parts  examined,  the  locality  where  collected,  and  careful 
details  of  all  characters  observed  which  are  not  exhibited  by 
che  specimens  themselves,  such  as  the  colour  of  the  fresh 
flowers.  It  is  important  that  all  specimens  should,  so  far  as 
possibly  preserve  their  natural  appearance  and  colour,  and  for 


.     163 

this  purpose  they  must  be  dried  as  quickly  as  possible. 
The  drying  papers  should  be  changed  at  least  once  a  day  for  the 
first  few  days  and  the  following  note  may  well  be  borne  in  mind : 
"  Two  or  tlyee  changes  of  the  driers  during  the  first  24  hours 
will  accomplish  more  than  a  dozen  changes  after  the  lapse  of 
several  days.  The  most  perfect  preservation  of  the  beautiful 
colours  of  some  orchids  has  been  effected  by  heating  the  driers 
and  changing  them  every  two  hours  during  the  first  day."* 
In  order  to  hasten  the  drying  of  succulent  plants  they  should 
be  dipped  (with  the  exception  of  the  flowers)  in  boiling  water, 
and  this  is  also  a  good  plan  to  adopt  in  the  case  of  those 
specimens  which  are  apt  to  lose  their  leaves  in  drying.  To 
reduce  the  thickness  of  some  specimens,  such  as  hard  fruits  and 
so  on,  they  may  be  thinned  by  cutting  away  the  underside, 
care  being  taken  to  see  that  the  original  shape  can  be  made 
out. 

142.    The   specimens    when  com-  Preservation 

pletely  dried  should  be  mounted,  and  this  is  perhaps  best  done  of  Speci- 
by  sewing  them  on  to  the  sheets  with  a  needle  and  thread,  never  m 
more  than  one  species  being  fastened  to  the  same  sheet. 
Fragile  specimens  must,  be  glued  to  the  sheets,  care  being 
taken  not  to  use  more  glue  than  necessary,  especially  on  the 
flowers.  Before  being  placed  in  the  herbarium  all  speci- 
mens should  be  poisoned  to  protect  them  from  mould, 
insects  and  vermin.  Corrosive  sublimate  (bichloride  of  mercury) 
is  the  poison  usually  employed  which  should  be  dissolved  in 
spirits  of  wine  in  the  proportion  of  about  \  oz.  of  the  poison  to 
one  pint  of  spirit.  This  should  be  applied  to  the  specimens  with 
a  large  soft  brush.  The  solution  should  not  be  allowed  to  come 
in  contact  with  metal  or  discoloration  may  result.  Leaves  and 
other  parts  of  the  specimens  which  become  detached  should  be 
placed  in  small  envelopes,  the  latter  then  being  fastened  to  the 
mounted  sheets.  At  the  foot  of  each  mounted  sheet  should  be 
carefully  noted  the  natural  order,  scientific  and  vernacular 
name  of  the  plant,  locality  where  found,  date  of  finding,  name 
of  collector  and  details  of  useful  characters  not  exhibited  by 
the  specimen  itself,  such  as  colour  of  the  fresh  flowers,  habit  oi 
the  plant,  and  so  on.  All  the  sheets  of  the  species  belonging  to 
one  and  the  same  genus  should  be  placed  in  one  folded  sheet  of 
strong  paper,  on  the  outside  of  which  should  be  written  the 
natural  order,  the  generic  name  and  the  names  of  the  contained 
species.  A  collection  of  dried  specimens  is  termed  a  herbarium  Herbarium, 
and  should  be  stored  in  cabinets,  or  small  almirahs,  provided 

*Asa  Gray's  Botanical  Text  Book,  Volume  I,  page  377. 

M2- 


164 

with  close-fitting  doors  and  fitted  inside  with  pigeon-hole  com- 
partments in  which  the  genus  covers  are  placed  and  are  thus 
readily  accessible  for  reference.  The  name  of  the  natural  order 
is  placed  above  the  pigeon-holes  containing  the  sheets  belonging 
to  that  order,  while  it  is  sometimes  convenient  to  affix  an 
index  of  the  genera  in  each  pigeon-hole  on  the  inside  of  the 
cabinet  doors,  or  a  list  of  the  genera  may  be  placed  on  the 
top  of  the  sheets  in  each  compartment. 

Before  examining  or  preparing  sections  of  dried  flowers  and 
fruits  they  should  be  placed  in  cold  water  and  then  gradually 
heated  until  sufficiently  soft 


165 


PART  V.-WOUND3  AND  DISEASES. 

CHAPTER  I.— WOUNDS. 

143.  If  a  mass  of  living  cells  is  cut  Healing  of 

through  with  a  sharp  knife,  the  exposed  surface  at  first  turns  ^JJJjfjf1^ 
brown  and  then  becomes  covered  with  a    protecting    coat    of  Cork  Layer, 
corky  tissue.    The  cells  actually  cut  through  die  at  once,  and  in 
consequence  of  the  access  of  air  some  of  their  contents  become 
oxidised,  this  process  causing  the  brown  discolouration.     The 

layer  of  cork  is  formed  by  the  uninjured  cells  immediately 
below  the  cut  surface  which,  by  their  growth  and  division, 
give  rise  to  layers  of  flat  tabular  cells.  These  soon  die  and, 
their  cell  walls  having  become  converted  into  cork,  form  a  pro- 
tective coat  several  layers  of  cells  in  thickness  which,  although 
elastic,  is  very  impervious  to  air  and  water,  and  consequently 
the  rapid  drying  up  and  destruction  of  the  .living  cells  beneath 
it  is  effectually  prevented.  This  process  may  be  well  observed 
on  the  cut  surface  of  a  potato,  although  the  wounded  surface 
of  any  mass  of  living  cells,  in  the  root,  stem,  leaf,  or  elsewhere, 
will  present  essentially  the  same  phenomenon. 

144.  If  the   cells  below   the   cut  Callus- 
surf  ace  are  growing  and  dividing  when  the  cut  is  made,  the  first  ^^^lin 
thing  which  happens  is  the  dying  of  the  actually  cut  cells    and  of  Wounds 
then  the  external  corky  coat  is  formed  as  before.     Now,    this  by  Ooclu- 
thin  layer  of  cork  being  very  elastic,  the  pressure  exerted  by  it Slon> 

on  the  living  growing  tissues  beneath  it  is  very  much  less  than 
was  caused  by  the  original  external  tissues  which  have  been  re- 
moved by  the  cut.  These  living  tissues  are  consequently 
able  to  grow  far  more  vigorously  than  before  and  they  give 
rise  to  a  juicy  cushion  of  thin-walled  cells  which  is  called  a 
callus.  This  at  first  consists  of  a  mass  of  embryonic  cells ; 
some  of  these  remain  capable  of  growing  and  forming  new 
cells,  thus  retaining  their  character  as  embryonic  tissue,  while 
the  remainder  become  gradually  differentiated,  very  much  as 
is  the  case  in  normally  growing  tissues.  This  process  may  be 
well  seen  on  the  vigorously  growing  stem  of  a  young  sapling. 
Suppose,  for  instance,  that  we  cut  away  a  longitudinal  strip 
of  the  external  tissues  from  such  a  stem,  the  cut  extending 
down  to  the  wood  which  is  thus  left  exposed.  The  wound  at 
first  gapes  owing  to  the  contraction  of  the  tissues  of  the  cor- 


166 

tical  jacket,  which,  before    the  wound  was  made,  was  tightly 
stretched    and    exerted    considerable    pressure  on    the  tissues 
below  it.     The  living  cells  of  the  cortex  and  the  cambium  at 
the  edges  of  the  wound  at  once  protect  themselves  with  a 
layer  of  cork    and    then    rapidly    develop  a  callus  as  above 
described,  which    pushes    out    towards    the    surface    of    the 
wound,    i.e.     in    the     direction    where    there    is    the    least 
resistance,  and  thus  forms  a  thick  pad-like   rim   around  the 
wound.     A  layer    of   cells  in  this  callus,  continuous  with  the 
cambium  of  the  stem,  retains  its  embryonic  character  and  acts 
as  a  cambium,  .the   tissue   formed  on  the  inside  of   the   layer, 
abutting  on  the  old  wood,   becoming  differentiated  into  wood 
and  that    outside    the  cambial  layer  developing  into  cortical 
tissue.     This  wood  formed  in  the  callus  is  called   wound-wood 
and,  owing  to  the  diminished  pressure  and  abnormal  conditions 
under  which  it  is  developed,   it  usually  differs  considerably 
from   normal    wood    in    its    structure.     The    cortex    of    the 
callus    usually    remains  thin  and  exerts   far  less  pressure  on 
the   tissues   beneath    it    than    old    cortex  would    do.     These 
callus    cushions    may    consequently    continue    to  grow  very 
rapidly  for  several  years,  and  the  whole  surface  of  the  wound 
may    thus    become    quickly    and    completely   covered    over, 
owing    to    the    various    cushions    coming    in     contact  with 
one  another  and    coalescing.     In  many  cases  no  thick,    outer 
bark    consisting    of      layers    of     dead     elements    has    been 
formed  on  the  callus  pads  when  thev  thus  come  in  contact, 
and,  as  they  continue  to  grow  and  press  against  one  another, 
the  thin    cork  layers    lying  between  them  are   ruptured  and 
squeezed  out,  while  the  cambium  and  living  cortical  tissues 
of  the  adjacent  lips  come  into  direct  contact  and  grow  together. 
The   cambium   of  the   various  cushions  thus   uniting,   a   con- 
tinuous cambial  layer  is  formed  over  the  whole  surface  of  the 
wound,  and  then  further  growth  in  thickness  continues  in  the 
normal  way  as  if    nothing  had  occurred,  and  very  soon  no 
external  sign  of  the  injury  can  be  seen.    If  thick  layers  of  dead 
bark  have  been  formed  on  the  callus  when  the  various  cushions 
come  into  contact,   the    complete  coalescence    of    the  latter  is 
rendered  much  more  difficult  and  may  be  much  delayed.     In 
some  species  the  living  cells  of   the  medullary  rays  on  the  sur- 
face of  the  exposed  wood  also  take  part  in  this  healing  process 
and   form    callus  pads,  in  addition  to  those  developed  at  the 
margins*  of  the  wound.     A  wound  which  is  healed  in  the  man- 
ner described  is  said  to  have  been   occluded,   and   this   healing 
process  is  termed  occlusion. 


167 

It  must  be  noted  that  no  growing  together,  or  intimate 
union,  is  possible  between  the  living  cells  of  the  callus  and 
the  dead  elements  of  the  wood  which  were  exposed  on  the 
wound  surface,  and,  although  the  callus  grows  over  and  is 
closely  adpressed  to  the  wood,  the  junction  between  them 
always  remains  as  a  distinct  line  of  separation."  This  is  often 
well  seen  in  the  case  of  hammer-marks  stamped  on  the 
exposed  wood  of  a  blaze,  the  latter  having  been  subsequently 
completely  healed  over  as  described  above.  Years  afterwards 
such  a  mark  may  be  clearly .  distinguished  deeply  embedded 
in  the  trunk,  the  letters  and  figures  having  been  merely  covered 
over  by  the  new  wood  and  hence  not  obliterated. 

145.  If,  in  the   above   example,  Bruises, 
the   outer  tissues  of  the  sapling  had  been  crushed  and  bruised 

by  a  blow  from  a  hammer,  or  other  means,  instead  of  being 
removed  by  a  clean  cut,  rapid  healing  in  the  manner  describ- 
ed would  have  been  impossible.  The  bruised  cambium  and 
cortical  tissues  die,  contract,  and  remain  adhering  to  the  wood 
below,  thus  obstructing  the  formation  of  a  callus  by  the  living 
tissues  bordering  the  wounded  patch.  Such  wounds  in  conse- 
quence do  not  heal  easily. 

146.  Now,  instead  of  wounding  Wounds 

the    stem    itself,    we  will  suppose    a    branch    to     have    been  °aused  b 
cut    off.   If  the  cut  has  been  made  just  above  one    or  more 

leafy  shoots,  the  latter  will  keep  the  living  tissues  of  the 
stump  alive  and  the  severed  end  will  eventually  be  healed 
over  by  callus  tissue.  If  the  stump  carries  buds*  the 
wound  may  stimulate  the  development  of  these  into  leafy 
shoots  and  the  healing  process  will  be  as  before.  If,  on  the 
contrary,  such  a  branch  stump  bears  no  buds,  or  leaves,  the 
tissues  will  dry  up  and  die.  The  living  cambium  and  cortical 
tissues  of  the  stem,  at  the  base  of  the  dead  stump,  now  develop 
a  ring  of  callus  tissue  around  the  stump,  but  the  formation 
of  this  callus  and  its  further  extension  up  the  stump  are  much 
impeded  by  the  dead  cortical  tissues  of  the  latter,  which  remain 
adhering  to  the  dead  wood  and  under  which  the  callus  has  to 
force  its  way.  The  healing  is  consequently  much  delayed  and 
the  dead  tissues  of  the  stump  frequently  rot  before  the'  process 
is  complete,  the  decay  then  spreading  into  the  sound  wood  of 
the  trunk  and  causing  a  hollow  stem.  Hence,  in  all  cases  of 
pruning  branches,  the  latter  should  be  cut  off  close  to  the 
stem,  the  cut  surface  being  parallel  to  the  surface  of  the  stem, 
the  healing  process  then  being  similar  to  that  described  for  a 
simple  stem  wound,  In  order  to  prevent  decay  spreading  in 


168 

the  wood  exposed  on  the  surface  of  the  wound,  it  is  important 
to  see  that  rain  water  is  not  allowed  to  lodge  on  the  wound 
and  that  air  is  excluded  as  far  as  possible.  In  Conifers  the 
wound  is  often  well  protected  by  a  copious  exudation  of  resin 
and  in  many  Dicotyledons  gums,  to  a  certain  extent,  serve 
the  same  purpose,  but  it  is  always  best  to  cover  the  cut 
surface  with  wood  tar,  or  some  other  substance,  which  will 
not  interfere  with  the  process  of  occlusion  and  will  prevent  air, 
water,  and  fungus  spores  obtaining  access  to  the  wound  until 
the  healing  is  complete.  In  connection  with  artificial  pruning 
may  be  mentioned  the  case  of  the  lower  branches  of  trees 
growing  in  a  dense  wood,  which  are  killed  by  the  heavy  shade. 
If  such  branches  are  .small  their  wood  is  usually  soft,  which 
soon  becomes  quite  rotten.  Such  branches  in  consequence 
quickly  fall  or  are  broken  off  close  to  the  stem  and,  the 
wound  soon  healing,  little  harm  is  done.  In  the  case  of  larger 
branches,  however,  which  often  contain  well  marked  heart-wood, 
the  dead  stumps  persist  for  several  years  and,  as  the  stem  in- 
creases in  thickness,  these  gradually  become  enclosed  in  the 
wood  of  the  stem.  No  intimate  union  being  possible  between 
the  dead  stumps  and  the  living  tissues  of  the  stem  which 
gradually  envelop  them,  such  stumps  form  knots  in  the  wood 
which  may  fall  out  after  conversion  and  cause  holes.  In 
addition  to  being  a  source  of  weakness  such  stumps  are  usually 
more  or  less  decayed  and  may  cause  wide-spreading  decay  in 
the  sound  wood  of  the  stem. 

Girdling  or  147.  Killing  trees  by  girdling   or 

Ringing.  ringing  is  a  common  forest  operation,  e.g.  in  the  case 
of  Teak  in  Burma.  If  a  deep  circular  cut  is  made  in  a 
Teak  tree,  down  to,  and  into,  the  heart-wood,  thus  com- 
pletely severing  the  connection  between  the  layers  of 
cortical  tissue  and  sap-wood  situated  respectively  above 
and  below  the  wound,  the  tree  will  quickly  die.  This  is  due 
to  the  fact  that  the  heart-wood  is  incapable  of  water  conduc- 
tion, and  hence  the  transpiring  crown  of  the  tree  situated 
above  the  girdle  no  longer  receives,  from  the  soil,  the  ne- 
cessary supplies  of  water  to  replace  that  lost  by  transpiration 
and  the  ultimate  death  and  drying  up  of  all  parts  above  the 
ring  is  consequently  assured.  This,  generally  speaking,  holds 
good  for  all  trees  which  possess  a  well-marked  heart-wood. 
Trees,  however,  which  possess  no  distinct  heart-wood  can  also  be 
killed  by  girdling,  e.g.  the  Himalayan  Spruce  and  Silver  Fir,  but 
in  the  case  of  such  trees  the  effect  of  girdling  is  frequently  very 
slow,  the  trees  often  remaining  alive  for  several  years.  In  such 


169 

cases,  it  is  not  possible  to  ascribe  the  death  of  the  tree  to 
any  one  single  factor.  It  is  known  that,  as  a  general  rule,  the 
water-current  from  the  roots  passes  chiefly  through  the  younger 
external  layers  of  wood,  but  it  must  be  remembered  that  all 
living  tissues  are  more  or  less  capable  of  water  conduction,  and 
that  the  ascending  water-current  under  ordinary  circumstances 
only  makes  use  of  the  young  wood  layers  because  it  can  pass 
through  them  more  rapidly  than  it  could  through  any  other 
tissue.  If  a  ring- wound  through  the  young  wood  now  interrupts 
these  rapidly  conducting  channels  the  water- current  is  compelled 
to  make  use  of  some  other  and  less  satisfactory  channels. 
Girdling  may  thus  result  in  the  water-current  being  carried 
through  the  inner  and  older  wood-rings  which,  under  ordinary 
circumstances,  would  not  have  been  utilized,  and  although 
this  interferes  with  the  rapidity  of  the  current  and  is  thus 
more  or  less  injurious,  the  death  of  the  tree  may  not  result  from 
this  cause  for  a  considerable  period.  It  should  also  be  noted 
that  the  longer  the  distance  which  the  water-current  has  to  tra- 
verse in  slowly  conducting  channels,  the  greater  will  be  the 
interference  with  the  rapidity  of  the  current  and,  consequently, 
the  wider  the  ring,  the  more  likely  it  is  to  be  effective  in  causing 
the  death  of  the  tree.  We  must,  however,  consider  not  only 
the  effect  of  the  wound  on  the  upward  water-current,  but  also  its 
effect  on  the  supply  of  food  materials  descending  the  stem  from 
the  leaves.  These  food  substances  are  mainly  carried  through 
the  sieve-tubes  of  the  cortex.  That  the  downward  passage  of 
these  materials  is  interrupted  by  the  ring- wound  is  indicated 
by  the  fact  that  a  callus  often  begins  to  form  only  along  the 
upper  edge  of  the  ring,  the  severed  tissues  along  the  lower 
edge,  which  now  receive  no  food  supplies  from  the  crown, 
being  unable  to  produce  a  callus.  Now  if  the  passage  of  the  food 
materials  from  the  crown  to  the  base  of  the  stem  and  the  roots 
is  thus  prevented,  it  is  obvious  that  although  the  reserve  mate- 
rials in  the  tissues  of  the  roots  and  the  base  of  the  stem  may 
for  a  time  suffice  for  the  nourishment  of  the  cambium,  for  the 
extended  growth  of  the  roots,  and  for  the  production  of  new  root 
hairs  and  water  conducting  layers  of  wood ;  this  cannot  continue 
indefinitely,  and  if  no  other  source  of  food  materials  is  made 
available,  the  supply  of  water  obtained  from  the  soil  by  the 
roots  will  gradually  decrease,  and  the  death  of  the  tree  eventu- 
ally result  from  this  cause  alone.  That  the  reserve  materials 
in  the  roots  are  thus  sometimes  exhausted  is  indicated  by 
the  fact  that,  in  some  cases,  no  coppice  shoots  arise  from  the 
stumps  of  trees  which  have  been  killed  by  girdling,  there 


170 

being  no  reserves  in  the  tissues  of  the  stump  and  roots  avail- 
able for  the  production  of  such  shoots.  If  the  severed  living 
tissues  on  the  lower  edge  of  the  ring  receive  sufficient 
nourishment  from  leafy  shoots  situated  below  the  ring,  they 
will  also  proceed  to  form  a  callus,  which  may  meet  and  coalesce 
with  that  developed  from  the  upper  lip,  the  wound  being  thus 
healed  over  and  the  normal  continuity  of  the  tissues  re-estab- 
lished, although,  here  again,  the  wider  the  ring,  the  longer 
will  the  healing  be  delayed  and  the  more  injurious  will  be  the 
wound.  Cases  will  be  occasionally  met  with  which  do  not 
appear  to  admit  of  an  explanation  according  to  the  above 
facts,  and  such  exceptions  will  be  found  due  to  the  existence  of 
special  conducting  tissue  in  the  interior  of  the  stem,  wood  which 
has  retained  its  power  of  conductivity,  internal  phloem,  etc. 
As  a  general  rule,  however,  the  non-success  of  girdling  is  due 
to  the  operation  not  having  been  thoroughly  performed, 
complete  interruption  of  the  ordinary  conducting  channels  not 
having  been  secured.  In  some  cases  it  is  possible  that  the  failure 
of  the  operation  is  due  to  the  roots  of  the  girdled  tree  being  in 
intimate  connection  with  the  roots  of  neighbouring  trees,  and  to 
their  thus  being  able  to  obtain  their  necessary  food  materials 
from  such  trees. 

-.     .  148.  Now    the    callus   which,    as 

we  have  seen  above,  so  frequently  results  from  a  wound, 
consists,  at  all  events  in  the  early  stages  of  its  development, 
largely  of  embryonic  cells,  i.e.,  cells  which  must  have  the 
capacity  of  producing  any  part  of  the  plant  body  in  which  they 
occur,  for  the  whole  plant  has  been  built  up  by  the  activity 
of  such  cells.  Hence  it  is  not  surprising  that  the  callus  is  often 
found  to  give  rise  to  roots,  and  buds  capable  of  developing  into 
shoots.  Such  roots  and  buds  are  termed  adventitious,  as  are  all 
structures  which  occur  in  places  where  under  normal  conditions 
they  would  not  have  been  developed.  This  may  be  well  seen 
in  the  case  of  cuttings,  these  being  shoots  which  have  been 
separated  from  the  parent  plant  by  a  clean  knife  cut.  The  base 
of  such  a  cutting,  if  kept  moist  by  being  placed  in  soil  or  damp 
air,  soon  becomes  covered  with  a  callus  developed  from  the 
living  growing  tissues  adjoining  the  cut  surface,  in  the  manner 
described  above.  Roots  then  spring  from  this  callus  and  through 
the  cortex  at  the  base  of  the  cutting,  the  latter  thus  becoming 
an  independent  plant.  It  is  important  to  restrict  evaporation 
from  the  cutting  as  much  as  possible  until  the  new  roots  have 
been  developed,  and  the  cuttings  are  often  first  laid  sloping- 
wise  in  the  ground  and  entirelv  covered  with  earth  with  the 


171 

exception  of  the  upper  buds,  until  they  strike,  i.e.  develop  roots. 
The  removal  of  the  transpiring  leaves  from  the  cutting  also,  to 
some  extent,  serves  the  same  purpose.  In  India  large  posts 
of  Boswellia  serrata,  Erythrina  suberosa  and  other  species  which 
have  been  placed  in  village  fences  may  often  be  seen  which  have, 
after  the  manner  of  cuttings,  developed  roots  from  the  base  and 
shoots  from  the  upper  end  and  thus  become  independent  trees. 

149.   In  the  case  of  many  species, 

the  felling  of  the  tree  results  in  a  more  or  less  vigorous  develop-  Poiiard- 
ment  of  callus  from  the  living  tissues  on  the  wounded  shoots, 
surface  in  which  adventitious  buds  may  arise,  while  the  wound  an 
also  often  stimulates  the  cambium  to  form  adventitious  budssuckers. 
elsewhere  on  the  stump  or  on  the  roots.  If  now  there  is  a  suffi- 
cient stock  of  food  materials  in  the  stump,  or  roots,  or  both, 
a  vigorous  development  of  shoots  from  the  adventitious  buds 
in  the  callus,  or  from  the  adventitious  or  dormant  buds 
elsewhere  on  the  stump,  or  from  the  adventitious  buds 
on  the  roots,  may  result.  When  the  young  shoots  arise 
on,  or  close  to,  the  cut  surface,  the  latter  is  usually  quickly 
covered  over  by  the  healthy  tissue  at  the  base  of  the  vigorous 
young  shoots  and,  the  access  of  air  and  water  being  thus 
obstructed,  the  spread  of  decay  into  the  stump  and  root 
system  is  prevented.  When  the  stump  of  the  tree  is  high 
and  the  shoots  arise  at  some  distance  from  the  ground 
they  are  usually  known  as  pollard- shoots,  whereas  if  the  tree  is 
cut  low  and  the  shoots  arise  close  to  the  ground  they  are  called 
stool -shoots,  or  coppice- shoots,  while  the  shoots  springing  from  the 
roots  are  termed  root-suckers.  Root- suckers  usually  develop 
young  roots  of  their  own,  and  this  often  occurs  also  with  stool- 
shoots  which  arise  at,  or  near,  the  surface  of  the  ground.  By 
directly  wounding  the  roots  themselves  and  by  thus  stimulating 
the  development  of  callus,  and  of  adventitious  root-and-shoot- 
buds  on,  and  near,  it,  the  production  of  root-suckers  may  often 
be  increased.  When  considering  the  phenomena  here  alluded 
to,  care  must  be  taken  to  distinguish  those  cases  in  which 
the  individual  plant  is  merely  trying  to  recover  from  the 
severe  injury  which  it  has  received  from  those  in  which  a  more 
or  less  complete  process  of  reproduction  may  be  recognised, 
consisting  in  a  division  of  the  mother-plant  and  the  establishment 
of  new  and  independent  individuals.  The  latter,  for  instance, 
occurs  when  root-suckers  become  independent  of  the  mother- 
tree  and  are  furnished  with  vigorous  root -systems  of  their  own. 
The  same  thing  also  appears  to  occur  in  cases  where  stool-shoots 
produce  roots  of  their  own  which,  owing  to  the  rapid  disinte- 


172 

gration  of  the  old  stool,  are  able  to  develop  vigorously  and  with- 
out obstruction  in  all  directions.  In  the  case,  however,  of  pollard- 
shoots  and  frequently  also  in  the  case  of  stool-shoots,  there 
is,  correctly  speaking,  no  true  reproduction.  Here  the  normal 
development  of  the  individual  tree  has  received  a  very  severe 
check,  the  individual  has  been  badly  wounded  and  injured  by  the 
removal  of  its  crown  of  foliage,  and  the  plant  sets  about  repairing 
the  injury  as  well  as  it  may,  just  as  we  have  seen  is  done  in  the 
case  of  other  wounds.  Thus  the  tree  at  once  endeavours  to 
replace  the  crown  of  foliage  which  has  been  removed  by  a 
vigorous  growth  of  pollard  or  stool- shoots.  As  a  general  rule, 
it  appears  that  the  closer  the  resemblance  between  the  new 
crown  of  foliage  formed  by  the  young  shoots  and  the  old  crown 
of  which  the  tree  has  been  deprived,  the  better  will  the  new 
crown  be  able  to  fulfil  its  duty  in  providing  for  the  healthy 
maintenance  and  extended  growth  of  the  old  root-system,  i.e. 
the  more  perfect  will  be  the  tree's  recovery.  In  other  cases  the 
process  of  recovery  is  slower  and  appears  to  resemble  in  a 
general  way  that  exhibited  by  a  tree,  the  crown  of  which  has 
been  severely  damaged  by  drought  or  frost.  In  such  a  case, 
if  the  injury  has  not  been  too  severe,  young  shoots  appear  on 
those  portions  of  the  stem  and  branches  which  are  still  alive. 
These  grow  and  gradually  take  the  place  of  the  dead  branches 
which  ultimately  fall  off,  and  in  a  few  years  the  recovery  may 
be  so  complete  that  we  can  see  no  signs  of  the  damage  remain- 
ing. A  very  similar  process  often  takes  place  in  the  case  of  a  tree 
which  has  been  pollarded  or  coppiced  ;  the  young  crown  of 
foliage  being  unable  to  supply  sufficient  food  material  to  the 
old  roots,  many  of  the  latter  begin  to  die  back  from  their  tips, 
while  a  crop  of  young  roots  is  adventitiously  developed  from 
those  parts  of  the  old  roots  which  still  remain  alive.  The 
tree,  as  it  were,  appears  to  be  trying  to  start  life  again  with 
a  new  crop  of  shoots  and  young  roots.  Just  as  a  tree  may 
completely  recover  from  severe  damage  by  frost,  drought,  or 
other  injuries,  so  may  it  also  satisfactorily  recover  from  the 
injury  inflicted  by  coppicing  or  pollarding,  but  it  should  be 
remembered  that  any  demand  made  on  the  powers  possessed 
by  a  plant  of  recovering  from  an  injury  is  usually  very  harm- 
ful if  repeated,  and  in  Europe  experience  has  shown  that  if  ash 
or  maple  are  repeatedly  cut  over  they  often  die  after  the 
second  or  third  operation. 

Propagation  150.  Propagation    by     means  of 

by  means  of  layers  and  by  guti  are  operations  often  resorted  to  by  gardeners 
GutiT  *n  India  m  preference  to  propagation  by  cuttings,  they  being  as 


173 

a  rule  more  certain  of  success.  Layering  is  described  as  fol- 
lows by  Firminger :  "  Select  a  branch  of  ripened  wood  of  the 
plant  to  be  layered,  that  will  bear  being  bent  down  to  the  earth 
without  breaking.  Cut  the  branch  half  through  with  a  sharp 
knife  just  under  one  of  the  leaf  buds  towards  its  extremity 
and  then  pass  the  knife  upwards,  so  as  to  slit  the  branch  about 
an  inch  or  two  up.  The  slit-piece,  with  the  leaf-bud  at  its  ex- 
tremity, called  the  '  tongue, '  should  be  kept  open  by  inserting  a 
small  piece  of  tile.  Remove  the  earth  to  the  depth  of  two  or 
three  inches  from,  or  place  a  flower-pot  over,  the  spot  just 
where  the  tongue  falls  on  the  branch  being  bent  down  ; 
then  carefully  bend  the  tongued  part  of  the  branch  into  the 
earth,  or  into  the  flower-pot ;  secure  it  in  that  position  by  a  peg, 
and  cover  it  over  with  earth,  which  should  be  pressed  down  and 
watered.*  "  A  callus  is  developed  from  the  cut  surface  of  the 
tongue  from  which  roots  arise  just  as  in  the  case  of  cuttings,  but 
the  layer  differs  from  a  cutting  in  that  it  receives  supplies  of 
water  and  salts  in  solution  through  the  wood  of  the  unsevered 
portion  of  the  branch.  So  soon  as  the  layer  has  developed 
its  own  roots  and  thus  become  independent  of  this  source  of 
supply,  it  may  be  completely  severed  from  the  parent  plant. 
The  operation  may  be  hastened  by  making  a  half-ring  wound 
on  the  unsevered  half  of  the  branch  down  to  the  wood.  The 
food  materials  descending  from  the  leaves  of  the  layer  which 
would  have  passed  down  .to  the  parent  plant  along  this  half 
of  the  branch  are  thus  intercepted  and  are  also  devoted  to  the 
production  of  callus  and  new  roots. 

The  operation  known  as  guti  consists  in  completely  ring- 
ing a  branch  down  to  the  wood,  the  latter  being  carefully 
scraped  to  insure  the  removal  of  all  the  cambium  and  cortical 
tissues.  The  wound  is  then  covered  with  a  lump  of  adhesive 
earth  or  moss  which  is  maintained  in  position  by  a  bandage 
and  kept  moist.  Here  the  passage  of  the  water  current  along 
the  young  wood  of  the  branch  is  not  interrupted  and  the 
leaves  are  enabled  to  manufacture  food  materials  vigorously ; 
these  materials  passing  down  the  sieve  tubes  in  the  cortex  are 
intercepted  in  their  course  at  the  upper  edge  of  the  ring 
where  they  are  devoted  to  the  production  of  a  callus  and  new 
roots.  So  soon  as  the  latter  are  well  developed,  the  branch 
may  be  completely  severed  from  the  parent  and  an  indepen- 
dent plant  is  produced.  Ficus  elastica  is  frequently  propagat- 
ed in  this  way. 

*  A    Ms.mial  of   Gardening    for   Bengal     and     Upper   India     by  Thomas     A. 
Fumkvger,  3rd  edition,  1874,  page  81. 


174 

Grafting.  151.  When  describing  above  the 

occlusion  of  wounds  it   was    noted    that    the    cambium     and 
living  tissues  of  one  callus  cushion,     coming    into     contact 
with     those     of     another     cushion,     coalesced     with,     and 
became  intimately   united   to    them.     This    power    possessed 
by  living    tissues  of  thus  growing    firmly  together    is    taken 
practical   advantage    of  in   several    ways    in    the    operations 
known   collectively    as   grafting.     Ordinary    grafting     consists 
in  bringing  the     cambium  and   living    tissues   of   one    plant 
into    close   contact  with    similar    tissues    of    another    plant. 
These  ultimately  coalesce,  and  from   the    union    of   parts    of 
two  distinct  plants    we  thus    get    a  single    plant ;  one  of  the 
partners,   called   the   stock,   which   was   provided   with   roots, 
supplying  water  and  mineral  salts  from  the  soil,   the  other 
partner  called  the  scion,  which  possessed  shoot  buds,  supply- 
ing food  materials  which  it  manufactures  in  its  leaves.     Such 
a  union  is,  however,  as  a  rule,  only  possible  between  closely 
related  plants.     In  the  majority  of  cases  of  successful  graft- 
ing   each    partner    is  found    to    preserve    its   own   individual 
characteristics ;  if  one  of  them  naturally  grows  faster  than  the 
other  it  continues  to  do  so  after  grafting  has  taken  place  and 
a  distinct  line  of  demarcation  between  the  fast-and  slow-grow- 
ing tissues  of  the  partners  is  visible  in  the  stem  of  the  plant 
resulting   from   the   grafting ;    similarly   an   Alphonse   mango 
grafted  on  an  ordinary  country-mango  stock,  continues  to  pro- 
duce Alphonse  mangoes  as  before  and  not  the  unpalatable  coun- 
try fruit.     At  the  same  time  rare  cases  are  known  where  graft- 
ing has  resulted  in  more  or  less  altering  the  characteristics  of 
the  partners.   The  method  of  grafting  usually  adopted  in  India 
is  that  known   as   inarching.     Firminger   describes   the   oper- 
ation   as    follows :  "  Procure  a  seedling  of  about  one  or  two 
years  old,  of  the  plant  to  be  inarched,  or  where  a  seedling  is 
not  to  be  obtained,  a  rooted  cutting  of  the  same  age,  of  the 
plant  that  is  to  supply  the  stock.     Put  it  in  a  pot,  and  when 
it  is  well  established    it  will  be  ready  to  be  operated  upon. 
Slice  away  from  one  side  of  the  young  stem  a  piece  of  bark, 
with  a  thin  layer  of  the  wood  beneath  it,  about  two  inches  long ; 
do  the  same  to  a  young  stem  of  the  plant  to  be  inarched  from, 
and  then  bring  together  the  two  stems  that  have  thus  been 
operated  upon  so  that  the  cut  parts  lie  close  in  contact  face  to 
face,   and   bandage   them   with    cotton-twist.     In   course    of 
time,  when  the  parts  have  united,  head  down  the  stook  and 
dissever  the  scion  from  the  parent  plant  by  cutting  it  through 
below   the  bandage.     The  grafted   plant  must  then   be  put 


175 

somewhere  in  a  shaded  place  and  not  removed  from  its  pot 
till  it  has  made  a  vigorous  growth,  and  stock  and  scion  have 
become  thoroughly  incorporated."  * 

This  operation  of  inarching  is  often  performed  naturally, 
and  it  is  not  uncommon  to  find  branches  of  trees  which,  after 
pressing  and  rubbing  against  the  stems  and  branches  of  neigh- 
bouring trees,  have  become  joined  to  them  at  the- points  of 
contact.  This  is  even  more  frequently  the  case  with  roots, 
and  such  natural  root  grafting  probably  explains  some  cases  of 
failure  in  ringing  trees  and  instances  of  stumps  producing 
coppice  shoots  which,  as  a  rule,  are  unable  to  do  so. 

152.  Budding  is  another  variety  Budding. 

of  grafting  which  is  commonly  practised.  This  must  be 
carried  out  when  the  sap  is  up  and  is  best  done  just 
before  the  period  of  vigorous  growth.  A  T-shaped  cut  is 
made  in  the  cortex  of  the  stock  down  to  the  wood.  A  piece 
of  cortex,  called  the  "  shield,"  bearing  a  healthy  stout  bud 
is  detached  from  the  wood  of  the  scion  and  inserted  in  the 
T  cut,  the  latter  being  then  bound  round  tightly  with 
cotton-twist  or  other  bandage.  The  inside  of  the  shield  thus 
lies  closely  against  the  wood  of  the  stock  and  the  bud  projects 
through  the  slit  in  its  cortex.  One  or  several  buds  may  be 
thus  grafted  on  the  stock,  and  when  these  have  developed 
vigorous  shoots,  the  branches  of  the  stock  are  cut  back 
close  above  the  budded  shoots,  so  that  the  water-current  from 
the  roots  may  not  be  diverted  from  the  latter. 

*  Op.  cit.,  pages  84-85. 


176 

CHAPTER  II.— DISEASES. 
SECTION  I. — INTRODUCTORY  REMARKS. 

Duration  153.   So  far  as  we   know  at  pre- 

of  Plant  sent,  all  the  embryonic  tissue  of  plants,  such  as  the  cambium, 
which  has  hot  become  converted  into  specialised  tissue  or 
organs,  is  capable  of  unlimited  life,  providing  the  external 
conditions  are  such  as  to  allow  of  its  growth.  The  continued 
existence  of  plant  life  upon  the  earth  in  fact  depends  on 
this  property  of  embryonic  cells.  Thus  bacteria,  the  cells 
of  which  all  retain  their  embryonic  character  and  power 
of  continued  growth  and  of  producing  new  individuals,  possess 
within  themselves  the  power  of  unlimited  life.  In  annual 
plants  the  death  of  the  plant  ensues  on  the  production  of 
the  seed.  The  seeds,  however,  contain  embryonic  cells,  and 
the  continued  existence  of  the  latter  renders  the  mainten- 
ance of  the  species  possible.  Trees,  on  the  other  hand, 
owing  to  the  presence  of  the  embryonic  cambium,  are 
often  capable  of  living  for  very  long  periods  and  may  reach 
an  age  of  more  than  1,000  years.  In  a  tree  considerable 
portions  continually  die  in  the  ordinary  course  of  natural 
development.  Thus  the  root-hairs  only  live  for  a  short  time, 
large  quantities  of  wood  are  killed  to  form  heart-wood,  masses 
of  external  tissue  die  and  become  dead  bark,  the  leaves  live  for 
a  few  years  only  at  the  longest,  sepals,  petals  and  stamens  have 
a  very  brief  life,  so  that  in  an  old  tree  the  actually  living  tissue 
which  has  arisen  from  the  embryonic  cambium  and  become 
converted  into  special  tissue,  or  organs  with  a  particular  func- 
tion to  perform,  is  at  most  but  a  few  years  old.  Not  only  in 
annuals,  but  in  some  plants  which  do  not  flower  for  several 
years,  such  as  some  species  of  Strobilanthes,  Bamboos  and  some 
Palms,  the  production  of  flowers  and  seed  causes  such 
exhaustion  of  the  plant's  vitality  that  death  ensues.  In  some 
cases  it  has  been  found  possible  to  prolong  a  plant's  existence 
by  artificially  preventing  the  production  of  flowers,  and  cases 
are  known  .in  which  annuals  have  thus  been  made  perennial. 
Whether  on  account  of  internal  or  external  causes,  however, 
so  far  as  we  know  at  present,  every  individual  plant  does 
sooner  or  later  die. 

Definition  of  1^^:-  ^n  nature   the  development 

Disease.         of  every  organism  depends  on  the  number  and  intensity  of  the 

injurious  and  beneficial  influences,  not  only  of   other  organisms, 

but  also  of  the  non-living  environment,  which  affect  its  develop- 


177 

ment.  Only  when  all  the  factors  which  influence  the  develop- 
ment  of  an  organism  combine  to  act  in  the  most  favourable 
way  can  an  ideally  healthy  development  become  possible. 
In  nature,  however,  this  is  probably  never  realized,  except 
perhaps  for  very  limited  periods  in  the  life  of  any  particular 
organism,  and,  with  the  gradually  increasing  predominance 
of  injurious  over  beneficial  factors,  we  get  at  first  a  condi- 
tion which  we  recognise  as  disease  and  eventually  death. 
Seeing,  however,  that  an  ideally  healthy  development,  for  even 
limited  periods  in  the  life-history  of  any  particular  organism, 
is  rarely,  if  ever,  possible,  it  is  obviously  very  difficult 
to  define  exactly  what  we  mean  by  disease.  We  have  seen 
that  the  ordinary  course  of  the  normal  development  of  a 
plant  in  itself  necessitates  the  death  of  considerable  portions 
of  the  plant-body,  or  even  of  the  entire  plant  in  cases  where 
death  naturally  follows  the  production  of  flowers  and  seed, 
yet  here  we  can  hardly  speak  of  disease.  For  practical  pur- 
poses, however,  we  may  regard  as  diseased  any  condition  of 
the  plant,  or  of  any  part  of  it,  which,  unless  ameliorated,  will 
lead  to  the  obviously  premature  death  of  the  plant,  or  of  some 
part  of  it. 

155.  In   Part   IV   above,  it   has  Strugg'e  for 

been  noted  that  what  we  call  the  struggle  for  existence Existence- 
is  responsible  for  the  premature  death  of  organisms,  and 
that  ,every  organism,  in  order  to  maintain  itself,  has  to 
struggle  with  other  living  organisms  and  to  fight  against, 
such  factors  as  unfavourable  climate,  excessive  heat  and 
cold,  drought  and  so  on,  see  p.  148.  When  it  is  considered 
that  these  factors  affect  various  organisms  in  very  different 
ways,  we  begin  to  realize  how  complex  the  relations 
are  which  bind  together  the  organic  world.  Hence  when 
studying  the  diseases  of  a  particular  plant,  w3  must 
not  only  consider  the  effect  exercised  upon  it  directly  by  such 
factors  as  soil,  moisture,  temperature,  light  and  such  like, 
but  we  must  also  discover  how  other  organisms,  plants  as 
well  as  animals,  affect  it,  what  factors  influence  the  develop- 
ment of  all  such  correlated  organisms,  and  in  what  way.  To 
illustrate  how  two  organisms,  between  which  at  first  sight  we 
should  say  there  was  absolutely  no  connection,  may  be  de- 
pendent on  each  other  for  their  existence,  we  cannot  do 
better  than  take  the  following  well-known  example  given  by 
Darwin : — 

"  Humble-bees  alone  visit  red    clover,  as  other  bees  cannot 
reach  the  nectar.     *  *     Hence  we  may  infer  as  highly 


178 


Factors 
which     in- 
fluence the 
Relations 
existing 
between 
Organisms 
in  Nature. 


probable  that,  if  the  whole  genus  of  humble-bees  became 
extinct  or  very  rare  in  England,  the  *  red  clover  would 
become  very  rare,  or  wholly  disappear.  The  number  of 
humble-bees  in  any  district  depends  in  a  great  measure  upon 
the  number  of  field-mice,  which  destroy  their  combs  and 
nests.  *  *  Now  the  number  of  mice  is  largely  de- 

pendent, as  everyone  knows,  on  the  number  of  cats.  * 
Hence  it  is  quite  credible  that  the  presence  of  a  feline 
animal  in  large  numbers  in  a  district  might  determine,  through 
the  intervention  first  of  mice  and  then  of  bees,  the  frequency 
of  certain  flowers  in  that  district !  "  *  We  frequently  do  not 
realize  the  severity  of  this  great  struggle  for  in  nature  a  bal- 
ance is  generally  maintained  between  the  conflicting  forces, 
a  position  of  more  or  less  stable  equilibrium  being  eventually 
arrived  at,  which,  however,  a  very  trifling  circumstance  may 
entirely  upset.  The  plant  known  as  Lantana  aculeata  is  a 
native  of  America  and  was  introduced  into  Ceylon  about  1824. 
Since  then  it  has  spread  with  extraordinary  rapidity  over  the 
Peninsula  of  India,  where  it  "  now  covers,  with  a  dense  network 
of  intertwined  branches,  large  areas  of  country,  almost  to 
the  complete  exclusion  of  other  vegetation."  f  This  well 
illustrates  how  a  plant,  the  development  of  which  is  favoured 
more  than  is  that  of  other  species  by  the  factors  of  the  locality, 
soon  succeeds  in  eliminating  all  other  vegetation.  Again 
man,  by  forming  pure  forests  of  a  single  species  of  tree,  has 
favoured  the  development  of  the  insects  and  fungi  which  prey 
upon  it,  by  collecting  together  and  making  easily  accessible 
for  them  masses  of  their  favourite  food,  a  procedure  which 
more  than  once  has  led  to  the  total  destruction  of  extensive 
forests  in  Europe,  while  in  the  teak  forests  of  India  we  clearly 
see  how  much  more  damage  is  done  in  pure  forests  than  in 
mixed  by  defoliators,  such  as  Hyblae:i  puera  and  Pyrausta 
machoeralis. 

156.  Let    us  now   try  to  realize 

the  factors  which  induce  one  organism  to  injure  or  help 
another  and  which  enable  it  to  do  so,  as  well  as  the  conditions 
which  enable  life  to  still  continue  on  the  earth  amidst  so  much 
destruction. 

Complex  carbon  compounds  generally  known  as  organic 
materials  form  an  essential  part  of  the  food  of  all  living 
organisms.  Green  plants  alone  among  all  living  organisms 


*  Origin  of  Species.     By  Charles  Darwin,  6th  Ed.,  pages  53 — 54. 
t  A  Manual  of  Indian  Timbers.     By  J.  S.  Gamble,  1902,  page  524. 


179 

• 

(with  a  few  rare  and  unimportant  exceptions)  are  able  to  manu- 
facture their  organic  food  from  the  carbon  dioxide  of  the  air ; 
all  other  plants  and  animals  must  obtain  their  carbonaceous 
food  from  materials  which  directly,  or  indirectly,  have  been 
manufactured  for  them  by  green  plants.     Without  green  plants 
then  all   life   on   the   earth   would    eventually   cense.     These 
organic  compounds  contain  energy,  or  in  other  words  capacity 
for  doing  work,  and  they  can  be  made  to  yield  up  this  energy 
which  is  stored  within  them  for  the  use  of  living  organisms, 
in  various  ways.   Some  of  them,  for  instance,  such  as  wood  and 
coal,  when  burnt  in  a  fire  are  made  to  give  up  their  energy  in 
the  form  of  light  and  heat.     Others  can  be  used   as   food  and 
made  to  give  up  their  energy  in  the  process  known  as  respira- 
tion. Whether  these  substances  are  slowly  burnt  off  in  living  or- 
ganisms as  food,  whether  they  are  quickly  consumed  in  a  fire, 
or  whether  they  are  slowly  oxidised  and  disintegrated  under  the 
chemical  and  physical  actions  of  air  and  water,  the  end  product 
of  the  decomposition  is  carbonic  dioxide.  Now  when  green  plants 
first  construct  these  organic  materials  large  quantities  of  car- 
bon dioxide  are  removed  from  the  air ;  a  certain  amount   of 
this  is  again  returned  by  mankind  and  animals  by  utilising 
organic  matter  as  food  or  fuel.     Large  quantities  of  carbon, 
however,  still  remain  locked  up  in  the  dead  bodies  of  plants 
and  animals  which  have  not  been  utilised  in  this  way,  and  if 
these  dead  bodies  were  to  remain  unchanged,  the  available 
supply  of  carbon  dioxide  in  the  air  would  eventually  become 
exhausted  and  all  living  beings  would  be  driven  out  of  exist- 
ence.    We  know,  however,  that  this  is  not  the  case,  but  that 
all  such  dead  remains  rot,  decay,  or  putrify,  and  are  eventually 
broken  up  into  the  elements  of  which  they  were  formed,  car- 
bon dioxide  being  again  set   free.     Ordinary    oxidation,    as 
already  mentioned,  is  responsible  for  a  certain  amount  of  this 
decomposition,  but  we  now  know  that  living  organisms  (espe- 
cially   bacteria    and    fungi)  which    derive  their  carbonaceous 
food  from  these  dead  bodies  take  an  exceedingly  active  part 
in  this  work  and  are  hence  indispensable  for  the  maintenance 
of  life.     Plants   which   thus   derive   their  carbonaceous   food 
from  the  dead  bodies  of  other  organisms,  or  from  substances 
manufactured  by  other  organisms,  are  termed  saprophytes.  They  Saprophytes, 
do  not  obtain  their  supplies  directly  from  the  living  tissues  of  an 
organism.     Green  plants  then,  we  see,  are  continually  building 
up  organic  matter  and  thus  not  only  provide  a  continual  sup- 
ply of  food  for  animals  and  other  plants  but  also   prevent  the 
accumulation  of  poisonous  quantities  of  carbon  dioxide  in  the 


180 

i 

air ;  non-green  plants  and  animals,  on  the  other  hand,  are  con- 
tinually consuming  organic  matter  and  by  returning  carbon 
dioxide  to  the  air  keep  up  the  supply  of  raw  material  necessary 
for  the  manufacture  of  fresh  organic  material  by  green  plants. 
From  the  fact  that  the  average  percentage  of  carbon  dioxide 
in  the  air  remains  practically  constant,  we  see  that  a  balance 
is  maintained  in  nature  between  this  work  of  construction  on 
the  one  hand  and  of  destruction  on  the  other,  a  continuance 
of  life  being  thus  assured. 

We  may  then  divide  all  living  organisms  into  the  following 
two  great  classes  :— 

(1)  Green  Plants  obtaining  their  carbonaceous  food  from 

the  air. 

(2)  Non-green  Plants  and  Animals  obtaining  their  carbon- 

aceous food  directly  or  indirectly  from  green  plants. 
Now  although,  as  ha  s  been  shown   above,  each  of  these 
great  classes  as  a  whole  depends  upon  the  other  for  its  exis- 
tence, yet  we  know  that  in  nature  individual  organisms,  owing 
to  their  great  powers  of  increase  and  to  the  limited  quantity  of 
food   materials  available,  are  continually   being  injured   and 
destroyed    in  the  struggle  for  .  existence  by  other   organisms. 
Regarding  their  relations  then  to  any  particular  organism   or 
group  of  organisms,  all  others  may  be  classed  as  follows  :— 
Competitors.         (i)  Competitors.     Those  which  are  indirectly  injurious  owing 
to  their  competition  for  the  same  necessaries  of  life. 

Parasite?.  (2)  Parasites.     Those  which  are  directly  injurious  owing 

to  their  obtaining  some  or  all  of  their  necessaries 
of  life  directly  from  the  living  bodies  of  organisms. 
The  organism  from  which  a  parasite  obtains  its 
supplies  is  called  a  host. 

Symbionta.  ^3)  Symbionts.     Those  which  not  only  are  not  injurious  but 

are  actually  beneficial  and  are  hence  more  or  less 
necessary  for  the  existence  of  the  organisms  bene- 
fited. 

The  phenomenon  of  two  organisms  living  together  with 
benefit  to  one  or  both  of  them  is  termed  symbiosis.  If  both  or- 
ganisms are  benefited  it  is  called  reciprocal  symbiosis  ;  if  only 
one  organism  is  benefited  it  is  called  antagonistic  symbiosis,  as 
in  the  case  of  a  parasite  and  its  host.  Symbiosis  also  may  be 
close  when  the  organisms  live  in  intimate  connection  with 
each  other,  or  distant  when  there  is  no  direct  or  fixed  union 
between  them.  It  is  of  course  often  impossible  to  insist  on 
these  distinctions ;  organisms,  which  at  one  period  of  their 
life  may  be  beneficial  (symbionts),  at  other  times  may  be 


181 

directly  injurious  (parasites),  and  in  cases  of  close  symbiosis  it 
is  often  difficult  to  decide  whether  or  not  one  organism  is  slight- 
ly injured  by  the  other,  that  is,  whether  they  are  to  be  regarded 
as  reciprocal  symbionts,  or  as  parasite  and  host. 

157.  The  symptoms   which    indi-  Symptoms 

catea    diseased     condition     are     of     course     very     various01 
according    to    the    different    factors     at     work      and     may 
consist    of   unusual    pallor    of   the    leaves,  discoloured  spots 

or  blotches  on  the  leaves,  premature  death  of  leaves,  or 
branches,  and  so  on.  At  the  same  time  such  symptoms  in 
themselves  by  no  means  necessarily  indicate  the  factors  which 
are  responsible  for  the  disease.  Premature  death  of  twigs 
and  branches  may,  for  instance,  be  due  to  the  effects  of  fire, 
frost,  drought,  an  unfavourable  condition  of  the  soil  or  the 
attacks  of  scale  insects. 

158.  In     some    cases   the    direct  One  Factor 

connection  between  the  injurious  factor  at  work  and  the  diseased  alc"ie  ls 
condition  of  the  plant  produced  by  it  is  obvious.   We  may,  for  p^n/biiTfor 
example,  see  a  larva  directly  feeding   on   and    destroying    the  a  Disease. 
leaves,  or  we  may  notice  that  the  death  of  our  tree  follows 
immediately  after  an  unusually  severe  frost,  and  in  such  cases  it 

is  natural  to  conclude  that  in  the  absence  of  the  larva  or  frost  the 
disease  would  have  been  avoided.  At  the  same  time  from  what 
has  been  said  above  regarding  the  struggle  for  existence  it  will 
be  realized  that  one  factor  alone  is  rarely  responsible  for  a 
disease  and  this  may  frequently  be  the  case  even  in  such  appar- 
ently obvious  examples  as  the  above.  The  destructive  larva, 
for  instance,  might  not  have  harmed  our  trees  if  the  weather 
or  some  other  factor  had  not  favoured  its  development  to  an 
unusual  extent,  the  frost  might  not  have  injured  our  trees  if 
they  had  not  been  growing  in  a  damp  valley,  their  twigs  in 
consequence  having  remained  soft  and  full  of  watery  sap  in- 
stead of  becoming  lignified  and  matured  like  those  of  the  trees 
on  the  hills  around.  The  discovery  of  one  obviously  injurious 
agent  at  work  is  thus  by  no  means  necessarily  a  satisfactory 
conclusion  to  the  investigation  of  any  particular  disease.  In 
cases  where  we  have  not  ourselves  been  able  to  note  the  first 
appearance  of  a  disease,  it  is  of  course  most  important  to 
ascertain,  as  far  as  possible,  the  character  of  the  first  visible 
symptoms,  the  time  when  they  were  seen,  and  whether  the 
existen.ce  of  any  unusual  external  influence,  such  as  severe 
drought,  the  occurrence  of  a  fire,  etc.,  was  noticed  when  the 
disease  first  appeared. 


182 


Knowledge 
necessary 
for  a  suc- 
cessful In- 
vestigation 
of  Plant 
Diseases. 


Sub-divi- 
sion of  the 
Subject. 


159.  In  order  that  our  study  of 

tne  diseases  of  our  forest  plants  shall  be  of   practical  value, 
we   must  by  careful  observation  and  study  :— 

(1)  Be  able  to  recognise  the  first  symptoms  of  a  diseased 

condition.  This  among  other  things  necessitates 
a  knowledge  of  the  life-history  of  our  plants.  We 
should,  for  instance,  be  inclined  to  ascribe  such 
phenomena  as  the  annual  dying  back  of  the  seed- 
lings of  bamboos  and  other  species,  or  the  shed- 
ding of  shoots  of  some  species  of  Strobilanthes, 
to  a  disease,  if  we  did  not  know  that  these  are 
necessary  in  the  ordinary  course  of  the  plant's 
development. 

(2)  Be.  able  from  these  symptoms  to   at  once   form   an 

approximate  idea  as  to  the  nature  of,  at  all  events, 
some  of  the  principal  factors  causing  the  disease. 

(3)  Be  acquainted  with,  and  know  the  mode  of  action  of, 

all  important  factors  which  either  beneficially  or 
injuriously  affect  the  normal  development  of  our 
plants.  This  will  then  enable  us  to  discover  in 
any  given  case  of  disease  any  unusual  absence  of 
beneficial  factors  or  presence  of  injurious  influ- 
ences and,  while  thus  indicating  the  principal  factors 
responsible  for  the  disease,  will  help  us  in  deciding 
on  the  most  advisable  remedial  and  preventive 
measures. 

(4)  Be  acquainted,  as  far  as  possible,  with  the  life-histories 

of  plants  and  animals  which  are  capable  of  bene- 
fiting, or  of  injuring,  our  forest  plants  and  with 
the  factors  which  influence  the  development  of 
the  same,  so  that  we  may  be  able  to  take  measures 
for  their  destruction,  if  injurious,  or  for  their 
advantage,  if  beneficial. 

160.  A  detailed    account    of    the 

life 'histories  of  the  various  animals  which  influence  the  develop- 
ment of  forest  plants,    their  mode  of  action,    the  symptoms 
by   which    the    diseased    conditions     caused   by   them     may 
be  recognised,   together  with   an    account    of    the    measures 
to    be     taken    for     their    destruction,     or   advantage,     form 
the   subject   of   Forest  Zoology.     It    is,    however,   interesting 
to  note  that  several   cases  of  symbiosis  between  plants  and 
animals  are  known.     Insects,  for   instance,   effect  the  pollina- 
tion   of     flowers     in   return   for    nectar  or    pollen     received 

the  flowers,  birds  and  other  animals  disseminate  seeds  in 


183 

return  for  food  in  the  shape  of  fruit,  etc.,  while  some  ants  in 
return  for  food  provided  by  the  plant  protect  the  latter  against 
other  injurious  insects. 

The  following  sub-divisions  of  the  subject  will  now  be  con- 
sidered in  detail  :— 

Influences  of  other  Plants  on  plant  development. 

Influences  of  the  Soil  on  plant  development. 

Influences  of  the  Atmosphere  on  plant  development. 

Effect  of  fire  on  plant  development. 

SECTION  II. — INFLUENCES   OF  OTHER    PLANTS  ON  PLANT  DE- 

VELOFMENT. 

161.  Following  the     classification 

proposed  above,  we  will  consider,  in  order,  the  effect  exercised 
by  plants    in  their   capacity  as  (a)  Competitors,  (b)  Parasites, 
and  (c)  Symbionts. 

(a)  Competitors, 

162.  We  have    to    consider  here 

not  only  the  competition  among  roots  for  the  water  and  Plant  Com- 
mineral  salts  in  the  soil,  but  also  that  among  shoots  forPetlt°rs- 
the  sun-light  and  air.  In  so  far  as  competition  with  the 
higher  plants  above  ground  goes,  their  insignificant  de- 
velopment in  the  air  renders  the  Cryptogams  of  little  or 
no  importance,  but  in  the  soil  their  competition  may  be 
injurious.  The  mycelia  of  many  saprophytic  fungi,  for 
example,  are  known  to  compete  actively  with  the  root  hairs  of 
higher  plants  in  the  soil  by  taking  up  large  quantities  of  valuable 
mineral  salts,  especially  of  phosphorus  and  potassium.  The 
most  injurious  competitors  with  the  higher  plants,  however,  are 
undoubtedly  to  be  found  in  the  ranks  of  the  higher  plants 
themselves,  i.e.  among  the  Phanerogams.  This  subject  has 
already  been  alluded  to  in  Part  IV  above,  see  p.  148,  and  only  a 
few  striking  examples  of  competition  can  be  mentioned  here. 

In  the  mixed   forest   of   Deodar    and   Blue   Pine    in  the  Deodar  and 
Western  Himalayas,  the  latter  has  continually  to   be  girdled  Blue  Pme 
to  prevent  its   suppressing    and    ousting   the    more   valuable 
Deodar.     A    similar    state    of  things    exists   in  all  our  Indian 
mixed  forests  and  the  forester  has  to  girdle,  or  fell,  trees  which, 
as  yet,  are  of  no  value  to    prevent  their  vanquishing  the  better 
kinds    in    this     competition     for     the     necessaries     of     life. 
Another  well-known   case  of    competition  is    afforded  by   the  Teak  and 
relations  existing  between  Teak  and  various  gregarious  species 
of  bamboos.     In  Burma,  for  instance,  the  dense  shade  afforded 
by   the   gregarious    Bambusa     poiymorpha    often     effectually 


184 


Trees  and 

Grasses. 


Trees  and 

Strobil- 

anthes, 


Climbers 


prevents  the  establishment  of  any  Teak  reproduction  beneath 
them.  Even  when  such  seedlings  have  become  established 
during  the  gregarious  seeding  and  dying  back  of  the  bamboos 
the  young  trees  may  still  be  caught  up  and  suppressed  by 
the  young  bamboos  resulting  from  the  general  seeding. 

In  areas  which  have  been  cleared  for  cultivation  in  our 
forests  and  then  abandoned,  the  dense  growth  of  grasses 
which  quickly  takes  possession  of  the  ground  often  prevents 
for  many  years  the  re- establishment  of  tree-species.  The 
mass  of  grass  to  some  extent  prevents  the  tree  seeds  from 
reaching  the  ground,  the  dense  network  of  grass  roots  in- 
tersecting the  superficial  layers  of  soil  in  all  directions  have  got 
the  start  in  the  competition  for  water  and  mineral  salts 
and  compete  with  the  delicate  roots  of  the  few  seedling  trees 
which  may  have  germinated,  while  the  heavy  growth  of  grass 
above  the  ground  effectually  prevents  the  struggling  plants  from 
obtaining  the  light  and  air  indispensable  for  their  development. 

Similar  in  many  respects,  with  regard  to  their  effect  on  the 
development  of  other  plants,  are  some  species  of  Strobilanthes. 
The  stems  of  the  gregarious  S.  Wallichii  in  the  Western  Himala- 
yas, for  instance,  "  form  a  dense  matted  covering  to  the  soil,  and 
prevent  the  seeds  of  the  forest  trees,  chiefly  oaks  like  Quercus 
dilatata  and  semecarpifolia,  and  firs  like  Picea  Morinda  and  Abies 
Pindrow,  from  reaching  the  ground,  or  if  they  do  reach  the 
ground,  obtaining  sufficient  light  fojr  germination  and  growth."* 

163.  Climbing    plants    again   are 

often  very  injurious  competitors  of  which  our  forests 
contain  many  examples,  the  most  striking  perhaps  being 
the  well-known  Bauhinia  Vahlii.  The  damage  done  by  such 
plants  consists  chiefly  in  the  injurious  competition  for 
light  between  their  foliage  and  the  crowns  of  the  trees  on  which 
they  climb.  The  species  mentioned,  for  instance,  climbing  to  the 
tops  of  the  highest  trees,  eventually  envelops  them  with  its 
mantle  of  enormous  leaves  which  sometimes  measure  as  much  as 
18  inches  across.  The  dense  curtains  of  its  foliage  effectually 
prevent  the  access  of  light  to  the  tree  branches  covered  by  them, 
and  the  leaves  on  the  latter  being  unable  to  manufacture  food, 
the  tree  is  ultimately  starved.  The  roots  of  such  climbers  also 
*ire  more  or  less  injurious  by  competing  in  the  soil  with  the 
roots  of  the  plants  on  which  they  climb  for  water  and  mineral 
salts.  Moreover,  the  pressure  exerted  by  the  coils  of  woody 
climbers  on  the  thickening  stems  and  branches  of  the  plants 


Manual  of  Indian  Timbers  by  J.  S.  Gamble,  1902,  p.  519. 


185 

supporting  them  more  or  less  interferes  with  the  descent  of  food 
materials  from  the  leaves  in  the  tissues  of  the  cortex  and  conse- 
quently with  the  nourishment  and  growth  of  the  cambium.  The 
latter  may  be  thus  killed  in  places  and  the  stems  or  branches 
become  deformed  and  irregular,  often  assuming  a  corkscrew 
shape  owing  to  the  growth  in  thickness  taking  place  in  a  spiral 
direction.  It  is,  indeed,  not  uncommon  to  find  dead  stems  which 
have  been  effectually  girdled  by  such  climbers. 

164.  This  brings  us  to  a  considera-  Epiphytes, 
tion  of  the  effect  of  so-called  epiphytes  on  other  plants.     The 
student  must  be  careful  not  to  confuse  the  term  epiphyte  with 
that  of  parasite.     An   epiphyte    is  a  plant    which    grows  on 
another  plant,   the  term  having  reference    merely  to   position 
and  thus  serving  to  distinguish  plants    which  grow  on   other 
plants,   from  those   which  grow    inside    other    plants   (endo- 
phytes),  or  on  the  earth  (terrestrial),  or  in  the  water  (aquatic).  The 
word  parasite,  on  the  other  hand,  refers  essentially  to  the  manner 
in  which  the  plant  obtains  its  food,  and  an  epiphyte  in  conse- 
quence may  or  may  not  be  also  a  parasite.     We  often  see  a 
young  Banyan  (Ficus  bengalensis),  or  Pipal  (Ficus    religiosa), 
growing  on  the  top  of  a  large  Mango  or  other  tree.      Year    by 
year  the  fig  grows  bigger  and,   as  its   dense  crown  of  foliage 
develops,  it  shades  the  Mango  leaves  more  and  more   from   the 
sunlight.     The  Mango  then  begins  to  languish,   and  ultimately 
dies.     After  several  years  an  enormous  fig   tree  alone   is   to   be 
found  on  the  spot  and  no    sign  remains  of  the  Mango  on  whose 
branches  it  first  started  life.  If  we  carefully  watch  the  develop- 
ment of  such  a  fig  we  find  that  the  young  plant  sends    out    its 
so-called  serial  roots  which   rapidly    spread    downwards  in  all 
directions  along  the  branches  and   stem  of  the  tree  attacked 
towards  the  soil.     These  become  woody  and  frequently  become 
grafted  together  at  points  where  they  come  in   contact  with 
one    another,   thus   forming  a  tightly-fitting,   latticed,   woody 
mantle  over  the  attacked  stems.     These  roots  also  produce  from 
their  under-surface  small  absorption-roots  which,  being  nega- 
tively heliotropic  and  positively  hydrotropic,  cling  tightly  to 
the  bark  and  penetrate  into  its  dark  cracks  and  fissures  where 
moisture  and  small   quantities  of  dust  and  humus  accumulate. 
The  latticed  roots  themselves  also  provide  lodgment  places  for 
dust  and  organic  debris,  and  from  this  alone  do  such  epiphytic 
roots   derive  their  nourishment,  for  they  do  not  penetrate  the 
living  tissues  of  the  stem  on  which  they  grow.   If  such  roots  are 
torn  away  from  the  stem  enveloped  by  them,  it  is  surprising  what 
a  large  quantity  of  dust  and  earthy  matter  may  often  be  found 


186 

adhering  to  them.  When  these  serial  roots  eventually  reach 
the  ground  they  branch  therein  and  rapidly  develop  a 
wide-spreading  root-system  in  the  soil.  If  by  this  time 
the  death  of  the  tree  attacked  has  not  been  caused  by  the  com- 
petition of  the  fig's  crown  of  foliage,  the  competition  of  the 
fig  roots  in  the  soil  now  help  in  effecting  this  result,  and  the 
fig  with  its  own  root-system  established  in  the  soil  no  longer 
requires  the  help  of  its  first  support.  Many  epiphytic  species  of 
Ficus  are  thus  capable  of  destroying  our  forest  trees,  and  it  is 
interesting  to  note  as  an  historical  fact  that  the  famous  Banyan 
"  in  the  Calcutta  Botanic  Gardens  began  life  as  an  epiphyte  on  a 
wild  date-tree  of  which  all  trace  has  long  disappeared."*  This 
tree  is  said  to  have  been  about  100  years  old  in  1886,  with  232 
supporting  trunks  developed  from  serial  roots.  The  circumference 
of  its  crown  was  then  857  feet,  and  it  was  growing  vigorously. 

Although  they  are  not  competitors  in  the  sense  here  ascribed 
to  the  term,  the  fungi  belonging  to  the  genus  Meliola  deserve 
mention  here  as  injurious  epiphytes.  As  has  already  been  men- 
tioned in  Part  IV  above,  these  minute  plants  growing  on  the 
surface  of  the  leaves  of  trees  and  shrubs,  and  feeding  on  the  sweet 
juices  excreted  by  insects  cover  the  leaves  with  black  incrusta- 
tions which,  by  preventing  the  access  of  light  to  the  green  chloro- 
phyll, are  more  or  less  injurious,  see  p.  133. 

Here  also  may  be  mentioned  the  Lichens  which  are  so  often 
found  growing  on  the  stems  and  branches  of  trees.  A  luxuriant 
growth  of  lichens  is  primarily  an  indication  of  a  moist  atmos- 
phere and  also  frequently  of  very  slow  growth  in  the  trees  on 
which  they  are  found,  for  quick  growth  combined  with  the  rapid 
ex-foliation  of  the  outer  layers  of  bark  prevent  the  establish- 
ment of  a  vigorous  growth  of  lichens.  In  so  far  as  direct  injury 
"is  concerned  the  latter  are  usually  of  very  little  importance, 
their  injurious  action,  being  apparently  confined  to  inter- 
ference with  the  access  of  oxygen  to  the  living  tissues  through 
the  lenticels. 

(b)  Parasites. 

165.  The    most     injurious    plant 

parasites  are  undoubtedly  to  be  found  among  the  Fungi,  of 
which  only  a  few  typical  examples  can  here  be  considered  in 
detail,  viz.  : — 

(1)  Phytophthora  infestans. 

(2)  Fomes  annosus. 

(3)  Trametes  Pini. 

(4)  Puccinia  graminis. 

*  Flora  Simlensis  by  Colonel  Sir  Henry  CoJlett,  page  459. 


187 

166.  This  fungus     belonging     to  (l)  phytoPh- 

the  great  group  of  the  PJiycomycetes  causes   the  most   virulent  \nfestans 
disease  from  which  the  potato  plant    (Solanum   tuberosum)   is  De  Bary. 
known  to  suffer,  and  in   countries  where   potatoes    are   exten-  Species 
sively  grown  the  damage  done  by  it  not  infrequently  amounts  attacked< 
to   a    national    calamity.     In  Ireland,     for    instance,    one     of 
the   worst    famines   of  modern    times    has    been    caused    by 
this  disease,    and  again    in    1879   the  loss   occasioned    by    it 
in  that  country  was  estimated  at   nearly  £6,000,000.      Up  to 
the  present  the  disease  in  India  has  been  chiefly  confined  to  the 
moister  localities  such  as  Assam,  the  Eastern  Himalayas  and  the 
Nilgiris,  and  experience  in  Europe  has  shown  that  an  exposure 
of  4  or  5  hours  to  a  dry  heat  of  104°  F.  is  fatal  to  the  furigus. 
From  the  fact,  however,  that   the  disease  has  of  recent  years 
established  itself  also  in  the  plains  of  Bengal  it  appears  probable 
that  it  will  gradually  spread  throughout  the   country  wherever 
potatoes  are  cultivated.    This  fungus  is  also  known  to  attack  the 
tomato  (Lycopersicum  esculentum)  and  a  few  other  plants  belong- 
ing   to  the  Solanaceae  and  Scrophidariaceae. 

The  presence  of  the  disease  is  usually  first  indicated  by  the  Signs  of 
occurrence  of  brown  patches  on  the  leaves  of  the  potato  plants. the  Dlsease- 
These  increase  in  size  and  depth  of  colour,  coalesce  with  neigh- 
bouring patches,  and  gradually  extend  over  the  whole  leaf 
surface.  Similar  patches  often  occur  also  on  the  petioles  and 
stem.  The  attacked  leaves  eventually  shrivel  up  and  drop  off 
leaving  the  bare  stalks  standing,  or  the  leaves  and  stalks  are  com- 
pletely converted  into  a  rotten  mass  which  emits  a  characteristic 
and  disagreeable  odour.  This  latter  is  chiefly  due  to  the  decom- 
posing action  of  bacteria  on  the  tissues  which  have  been  killed 
by  the  fungus.  The  extension  of  the  disease  is  particularly 
rapid  in  damp  warm  weather. 

If  one  of  the  attacked  leaflets  is  now  turned  over  and  one  of  Life  History 
the  characteristic  brown  patches  on  the  under  surface  examin-  and  Damase 
ed  with  a  lens,  a  number  of  pale  silky  threads  will  be  seen 
standing  up  from  the  leaf  surface  at  the  edges  of  the  dark  spot, 
thus  producing  a  characteristic  pale  mouldy  margin  to  the  patch 
which  is  particularly  noticeable  in  damp  weather.  See 
Plate  XVI  (1).  These  more  or  less  erect  threads  are  the 
serial  hyphae,  or  conidiophores,  of  the  fungus  which  have  been 
sent  out  singly  or  in  tufts  through  the  stomata  on  the  under 
surface  of  the  leaf  for  the  purpose  of  bearing  the  conidia,  or 
asexual  reproductive  organs.  These  fine  tubular  conidiophores 
are  branched,  and  at  the  tip  of  the  main  filament  and  of  each  o{ 
its  branches  the  minute  egg-shaped  conidia  are  produced 


188 

which,  if  sufficiently  numerous,  appear  as  a  fine  white  powder 
through  the  lens.  The  conidia  usually  fall  off  or  are  blown  off  by 
the  wind  almost  as  soon  as  they  are  formed,  small  swellings  in 
the  hyphse  indicating  the  places  where  they  originated.  See 
Plate  XVI  (2).  The  mycelium  or  main  body  of  the  fungus 
consists  of  a  branched  net- work  of  fine  tubular  hyphse 
which  spread  and  branch  in  all  directions  in  the  intercellular 
spaces  of  the  leaf-tissue.  If  a  single  hypha  is  examined  with 
the  microscope  it  will  be  seen  to  consist  of  a  long  tube,  with  a 
transparent  thin  wall  of  cellulose,  containing  water,  protoplasm 
and  other  substances.  The  hyphse  are,  as  a  rule,  not  septate, 
but  transverse  partition  walls  are  occasionally  developed  at 
irregular  intervals  to  separate  the  parts  of  the  hyphse  which 
are  still  growing  and  full  of  protoplasm  from  those  which  have 
completed  their  growth  and  are  empty.  These  hyphae  push 
their  way  between  the  cells  of  the  leaf,  and  as  a  rule  do  not 
directly  enter  the  cells.  Occasionally,  however,  they  develop 
small  branches  which  dissolve  their  way  through  the  neighbouring 
cell-walls  by  means  of  an  enzyme  which  they  secrete,  and  which, 
acting  as  absorbing  organs,  are  called  haustoria,  or  suckers. 

Whether  the  hyphse  enter  the  cells  or  not,  however,  the 
result  is  the  same ;  they  consume  a  quantity  of  the  oxygen  and 
water  which  is  required  by  the  living  leaf -cells,  they  directly  ab- 
sorb the  products  of  assimilation  which  would  ordinarily  have 
been  devoted  to  the  development  of  the  tubers,  and  finally  they 
destroy  the  living  protoplasm  and  kill  all  the  living  cells  with 
which  they  come  in  contact.  The  cell-walls  turn  brown  wherever 
the  hyphae  touch  them,  the  colour  extending  for  some  distance 
along  the  walls  beyond  the  actual  point  of  contact.  Here  and 
there  the  ends  of  the  hyphse  emerge  from  the  leaf  tissues  and 
develop  the  branched  conidiophores  as  above  described.  The 
conidia  being  very  minute  and  falling  rapidly  are  blown  about 
and  distributed  by  the  wind  like  fine  powder,  thus  enabling  the 
fungus  to  spread  rapidly  and  infect  fresh  leaves  and  plants. 

The  conidia  may  germinate  in  one  of  the  three  following 
ways : — 

(1)  If  the  conidium  falls  into  a  drop  of  rain  or  dew  the  proto- 
plasm divides  into  from  6-16  parts,  the  tip  of  the 
conidium  dissolves,  and  these  little  masses  of  proto- 
plasm escape.  These  are  called  zoospores,  and  each 
is  provided  with  two  very  fine  hair-like  appendages 
called  cilia  which  lash  the  water  and  enable  the  zoos- 
pore  to  swim  and  move  about  in  the  water.  In  a 
quarter  of  an  hour  or  so  these  zoospores  lose  their 


189 

cilia  and  come  to  rest.  Each  then  develops  a  very 
fine  tube  which  grows  out  into  a  hypha.  The  latter 
dissolves  its  way  into  the  cells  of  the  leaf,  or  stalk, 
among  which  it  then  grows  and  spreads  and  soon 
reproduces  the  mycelium  above  described.  See 
Plate  XVI  (3)-(6). 

(2)  The  conidium  instead  of  forming  zoospores   itself  puts 

out  a  germ-tube  which  directly  enters  the  plant  tjs- 
sues  and  forms  the  mycelium.  See  Plate  XVI  (7). 

(3)  The  conidium  forms  a   germ-tube  as   before,    but   this 

instead  of  entering  the  plant  tissues  forms  a  secondary 
conidium  at  its  apex  which  in  its  turn  falls  off  and 
gives  rise  to  zoospores  or  to  another  germ-tube  which 
directly  enters  the  plant-tissues. 

The  first  of  these  processes  is  the  most  frequent  especially 
with  conidia  which  come  in  contact  with  water  shortly  after 
falling.  Conidia  lose  their  power  of  germinating  in  about  three 
weeks,  and  dry  weather  is  unfavourable  to  their  production  and 
germination.  Conidia  which  are  washed  by  rain  upon  tubers  can 
infect  the  latter  directly,  the  germ-tube  penetrating  the  tissue 
and  forming  a  mycelium  there.  The  mycelium  may  also  pass 
down  the  potato  stalks  and  infect  the  tubers  in  this  way.  The 
mycelium  having  reached  the  tubers  may  proceed  to  destroy 
them  in  the  same  way  as  the  leaves,  the  numerous  bacteria  in  the 
soil  assisting  in  the  decomposition  of  the  diseased  tissues,  or  the 
mycelium  may  remain  in  the  tubers  in  a  dormant  condition. 
Such  infected  tubers  on  being  cut  open  usually  exhibit  dark 
patches  and  when  stored  frequently  rot.  If  infected  tubers  are 
planted  out  the  next  season  the  mycelium  again  becomes  active 
and  growing  along  with  the  young  shoots  sent  out  by  the  tuber 
reproduces  the  disease. 

The  plants  may  be  sprayed  with  Bordeaux  mixture  which  Preventive 
has  been  described  as  "  the  most  effective  and    cheapest  fungi-  Measures- 
cide  known  ".     This  does  not  poison  the  tubers  or  make  them 
unfit  for  consumption,  but  it  kills  the  conidia  and  zoospores  and 
prevents  their  germination.  In  addition  to  its  action  on  the  fun- 
gus this  mixture  has  actually  proved  beneficial  to  the  potato 
plant  and  results  in  an  increased  yield  of    tubers.     The  copper 
in  the  mixture  in  some  way   which  is  not   clearly   understood 
increases  the  amount  of  chlorophyll  in  the  leaves  and   conse- 
quently the  assimilating  powers  of  the  foliage.* 

Care  must  be  taken  not  to  use  infected  tubers  for  seed. 

*  For  instructions  regarding  the  preparation  and  application  of  Bordeaux 
mixture  see  Potato  Diseases  of  India  by  Dr.  E.  J.  Butler,  published  in  the  Agr  - 
cultural  Ledger,  1903,  No.  4,  pp.  108-109. 


omnivora. 


190 

Some  varieties  of  potatoes  have  been  found  to  resist 
the  disease  better  than  others.  Tubers,  for  instance,  with 
exceptionally  thick  skins  offer  more  resistance  to  the  entrance 
of  the  hyphse  than  do  others  with  thin  skins.  In  order  to 
keep  up  the  power  of  resistance  of  the  potato  plant  to  the 
disease,  care  must  be  taken  to  always  select  the  most  resistant 
tubers  for  continued  cultivation,  and  to  raise  new  and  stronger 
varieties  from  the  seed  obtained  by  crossing  good,  selected, 

«/  c?      o 

varieties.  Diseased  stalks,  leaves  and  tubers  should  be  burnt 
and,  to  prevent  risk  of  infecting  sound  tubers  from  living 
conidia  lying  on  the  ground,  the  tubers  should  not  be  raised 
for  three  weeks  after  the  complete  dying  down  of  the  plants. 

As  moisture  favours  the   development   and  spread  of    the 
fungus,  wet  localities  should  be    avoided  as  far  as  possible. 
Irrigation  of  the  crop  should  be  carried  out  with  caution,  and 
the  tubers  should  be  stored  in  a  dry  well-ventilated  place. 
Phytoph-  167.  Very    closely    allied    to  the 

above  fungus  is  another  named  Phytophthora    omnivora    De 
Bary,   which    in  Europe  has  proved  very  destructive  to  the 
seedlings  of  forest  trees.     It  is  probable  that  the  "  damping 
off "  of    seedlings  in  many  of    our  forest   nurseries   in    India 
will  be  found  to   be  caused  by  it  or  by   some  other   nearly 
related    fungus   with  an    almost    identical     life-history    and 
mode  of    attack-     The  first  signs  of  attack  are  usually  dark 
spots  on  the  stern,  cotyledons,  or  first  leaves  of  the  seedling, 
and  if    the  weather  is  warm  and  damp  the   young    plant    is 
soon  destroyed.     Frequently  only  the  "  collar  '     is    attacked 
and  the  seedlings  then  fall  over.     If  only  the  upper  leaves  are 
attacked  the  seedlings   may  recover.     The   damage  gradually 
spreads     from   one    seedling    to    those    around    it,    and    the 
circle  of  diseased  plants    gradually    spreads     outwards    from 
the  centre.     The    growth    and    injurious    action    of  the    my- 
celium of  this  fungus  in  the    tissues    of    the    attacked    plant 
are  almost  precisely  similar  to  those  of  P.  „  infestans,  except 
that    the    hyphse     more     commonly    absorb    their    nourish- 
ment by  means  of  haustoria.     The  most  important  point  of 
difference  between  them  consists  in  the  fact  that  this  species 
produces  sexual  resting  spores  (oospores)  in  the  tissues  of  the 
attacked  plant.     These  spores  have  the  power  of  remaining 
dormant  for  several  weeks  or  months  before  germinating  and 
hence  their  name  of  resting  spores.     In  the  formation  of  these 
spores  the  end  of  a  hypha  swells  up  into  a  rounded  knob,  and 
the  protoplasm  inside  it  forms  into  a  ball  thus  producing  what 
is  called  the  oosphere  or  egg-cell.     The  end  of  another  hypha 
then  coming  in  contact  with  the  swelling,  sends  out  a  small 


191 

tube  which  pierces  the  cell  wall  and  penetrates  to  the  oosphere. 
the  protoplasm  of  the  hypha  then  passing  into  and  mixing 
with  that  of  the  oosphere,  through  the  short  connecting  tube. 
The  oosphere  is  then  said  to  be  fertilized  and  is  called  the 
oospore.  These  spherical  oospores  soon  develop  a  hard  pro- 
tective cell-wall  and  are  capable  of  retaining  thejr  vitality 
and  power  of  germinating  for  a  considerable  period.  These 
spores  find  their  way  from  the  decaying  tissues  of  the  seedlings 
into  the  soil  and,  if  seed  is  sown  in  the  same  soil,  the  new 
crop  of  seedlings  is  almost  invariably  severely  attacked  by 
the  disease,  the  oospores  in  the  soil  germinating  and  producing 
hyphse  which  then  proceed  to  branch  and  form  conidia  as 
before.  This  fungus  is  as  a  rule  only  destructive  to  seedlings, 
young  plants  one  year  old  and  upwards  being  rarely  injured. 
It  is  also  capable  of  living  in  the  soil  for  a  long  time  as  a 
saprophyte  and  thus  can  exist  independently  of  seedlings, 
although  it  is  always  ready  to  resume  its  parasitic  mode  of  life 
on  any  suitable  seedlings  it  may  meet  with. 

As  in  P.  infestans  moisture  favours  the  spread  and  deve- 
lopment of  the  pest;  consequently  shading  the  seed  beds  is 
inadvisable  as  this  would  prevent  the  rapid  evaporation  of 
moisture.  Spraying  with  Bordeaux  mixture  will  aid  in  pre- 
venting the  spread  of  the  disease.  Isolated  specimens  of 
diseased  seedlings  should  be  carefully  removed  and  burnt, 
care  being  taken  not  to  infect  fresh  seedlings  by  shaking  of? 
the  conidia.  If  a  considerable  number  are  affected  the  best 
way  of  preventing  the  dispersal  of  the  conidia  is  to  cover 
the  patch  carefully  with  soil.  People  passing  through  the 
seed-beds  may  convey  the  conidia  on  their  clothes,  or  boots, 
to  healthy  individuals  which  should  be  avoided  as  far  as  pos- 
sible. Seed-beds  must  be  frequently  inspected  so  that  timely 
measures  may  be  taken  to  check  the  disease.  A  seed-bed 
which  has  been  attacked  should  not  be  used  again  for  seed- 
lings of  the  same  species.  In  some  cases  good  may  be  done 
by  burning  rubbish  on  the  infected  seed-bed,  sufficient  heat 
being  produced  to  kill  the  spores  in  the  soil. 

168.  This  fungus    is    also   known  (2)  Fames 

by  Hartig'  s  name  of  Tmmetes  radiciperda.     In  Europe  it  has  «f  »°*«* 
been  found  to  attack  Pinus,  Picea,  Abies,  Juniperus,  Thuja,  the  species 
Beech,  and  other  dicotyledonous  trees.     It  is  most  destructive  to  attacked, 
conifers,  and  in  India  it  is  at  present  best  known  on  account 
of  the  damage  done  by  it  to  the  Deodar,  Cedrus  Libani  var. 
Deodara,  which  it  attacks  and  kills.     The  fungus  is  frequently 
seen  in   the   forests   of  Jaunsar. 


192 

Signs  of  the  Young  Deodar  trees  6-15  feet  in  height  are  chiefly  attacked. 
One  tree  generally  dies  first  and  then  the  disease  spreads  to 
others  near  it,  gaps  being  formed  which  quickly  extend  cen- 
trifugally.  Resin  at  first  begins  to  e%ude  from  different 

o        «/  o 

points  along  the  stem  and  then  the  needles  turn  yellow  and 
fall  off.     Stems  usually  take  two  years  to  die  completely. 

The  sporophores  or  fruiting  organs  are  developed  at  the  base 
Life-History  of  the  Deodar  stems  and  will  also  probably  be  found 
and  Damage  On  the  roots  below  the  ground  surface.  They  arise  from 
done.  cushions  of  mycelium  which  force  their  way  to  the 

surface  of  the  stem  or  root  at  these  points,  where  the  sporo- 
phores first  appear  as  small,  rounded,  chocolate -brown 
nodules.  These  increase  in  number  and  run  together  forming 
a  brown  incrustation  broken  by  numerous  ledges  and  tuber- 
cles. On  the  younger  portion  of  the  sporophore  a  series  of 
imbricate  bracket-like  protrusions  are  developed.  The  lower 
surface  of  these  brackets  is  white  and  contains  numerous 
minute  pores,  on  the  walls  of  which  the  spores  are  developed, 
while  the  upper  surface  is  rough,  brown,  and  covered  with 
tubercles  and  concentric  ridges.  It  is  probable  that  these 
spores  are  often  carried  in  the  ground  on  the  fur  of  mice  and 
other  burrowing  animals  and  are  rubbed  off  on  healthy  roots. 
On  germination  they  give  rise  to  very  delicate  hyphse  which 
penetrate  between  the  bark  scales,  force  their  way  into  the 
wood  and  give  rise  to  characteristic  white  sheets  and  bands 
of  mycelium,  between  and  under  the  bark  scales.  The  roots 
are  not  the  only  parts  affected.  Broad  bands  of  the 
mycelium  invade  the  collar,  ascending  the  stem  and  forcing 
their  way  along  the  cambial  layer,  in  feathery  strands  of 
snow-white  tissue.  These  completely  destroy  the  cambial 
layer.  In  addition  to  this  the  wood  is  directly  attacked 
and  destroyed  by  the  minute  hypha3  which  bore  their  way 
from  cell  to  cell  by  means  of  a  cellulose-dissolving  enzyme 
which  they  exude.  The  lignin  in  the  cell-walls  of  the 
attacked  wood  is  dissolved  out  first  by  the  hyphae  and 
the  middle  lamella  is  destroyed,  so  that  the  wood 
elements  fall  apart.  As  a  result  of  this  action  white  areas 
of  diseased  wood  arise  which  are  very  characteristic  and  which 
are  usually  elongated  in  the  direction  of  the  long  axis 
of  the  stem.  They  consist  of  dissociated  cells  of  prac- 
tically pure  cellulose,  which,  being  themselves  ultimately 
decomposed,  leave  small  hollows  in  the  wood.  Before  de- 
composition is  completed  the  cell-walls  show  a  fine  striation 
and,  later,  large  spiral  cracks.  Living  cells  in  the  wood  paren- 


193 

chyma  and  medullary  rays  attacked  by  the  hyphse  become 
filled  with  a  brown  substance,  probably  caused  by  the 
decomposition  of  the  protoplasm.  This  discoloration  is  more 
noticeable  in  the  roots  than  in  the  stems  of  the  Deodar.  A 
remarkable  characteristic  of  this  fungus  is  the  possession  of 
rhizomorphs.  These  are  branched,  cylindrical,  or~  flattened, 
bodies  composed  of  an  outer,  black  and  somewhat  brittle, 
cortex,  formed  of  interwoven  tough  brown  hyphse  and  of  a 
central,  tough,  flexible,  whitish  portion  composed  of  fine 
hyphal  filaments.  In  general  appearance  these  resemble  the 
black  petioles  of  some  ferns  and  are  not  unlike  the  ordinary 
fine  Deodar  roots.  They  are,  however,  blacker,  of  a  different 
consistency,  and  do  not  taper  regularly  towards  the  apex. 
They  can  be  distinguished  by  splitting  them  longitudinally 
when  the  silky  white  medulla  can  be  teased  out  with  the 
fingers.  These  are  organs  of  propagation  possessing  the 
power  of  apical  growth.  Springing  from  the  mycelium  in 
the  roots  of  diseased  trees,  they  travel  below  the  surface 
of  the  ground  and,  penetrating  below  the  cortex  of  any  healthy 
Deodar  roots  which  they  may  encounter,  develop  in  them 
the  ordinary  mycelium,  and  thus  spread  the  disease  from  tree 
to  tree.  Trees  growing  close  together  in  the  forest  often  have 
their  roots  in  close  contact  and  are  frequently  found  grafted 
together.  It  is  probable  that  infection  often  takes  place  by 
the  mycelium  thus  growing  directly  from  one  root  into 
another  which  is  in  contact  with  it.  For  illustrations,  see 
Plate  XVII. 

The  only  measures  likely  to  be  effective  are  the  removal  of 
attacked  trees  to  prevent  the  formation  of  sporophores  and  the 
isolation  of  infected  areas  by  trenches  dug  a  few  feet  deep 
and  sufficiently  far  from  the  centre  of  infection  to  include 
all  diseased  roots  and  rhizomorphs.  It  is  doubtful  if  this 
could  be  done  profitably  on  a  large  scale. 

169.    In    its    life-history     and 

action  on  the  plant  attacked,  Armillaria  mellea,  which  has  been 
briefly  noted  in  Part  IV  above,  has  several  points  of  resem- 
blance with  the  fungus  just  described,  see  page  133.  Armillaria, 
however,  is  particularly  interesting  as  being  one  of  those  f ungi 
which,  although  living  at  times  in  a  purely  parasitic  manner, 
are  also  able  to  exist  as  saprophytes.  This  fungus  may 
accordingly  sometimes  be  found  thriving  on  wood  used  in 
construction,  e.g.  in  bridges,  and  also  on  dead  roots  and 
stumps,  and  at  others  as  a  virulent  parasite  attacking 
and  killing  living  trees.  From  cases  such  as  these  we 


194 

/ 

conclude  that  no  hard  and  fast  line  can  be  drawn  between 
these  classes  of  saprophytes  and  parasites,  and  there  is  every 
reason  to  believe  that  the  one  mode  of  life  has  led  to  the  other. 
At  the  same  time  it  must  be  remembered  that  a  hypha,  which 
is  able  to  pierce  the  cell- walls  of  a  plant's  tissues,  is  not  by 
any  means  necessarily  able  to  kill  the  living  protoplasm  in 
the  cells. 

ni  17°'    In    India    thiS    destmctive 

Species"        fungus  has  been  found  on  Pinus   excelsa    near   Simla,    and 
attacked.      it  is  well  known  on  various  conifers  in  Europe. 

Signs  of  the  The  characteristic  sporophores  usually  first  indicate  the 
Disease.  presence  of  the  fungus.  These  appear  on  the  stem  usually  near 
the  stump  of  a  dead  branch,  and  are  hard,  bracket-like,  masses 
often  triangular  in  section.  The  upper  surface  of  the  sporo- 
phore  is  rough,  blackish-brown  in  colour  and  with  concentric 
grooves,  the  fawn-coloured  lower  surface  is  covered  with  the 
minute  pores  in  which  the  spores  are  developed.  See  Plate 
XVIII  (a). 

L'!TT  iRtory         ID  nianv  respects  the  damage  done  by  this  pest  resembles 

and  Damage  •  v  J  £       v 

done.  that    described     above    m    the    case     ot     tomes     annosus, 

but  it  is  important  to  note  that,  whereas  the  latter 
fungus  can  attack  sound  roots,  Trametes  Pini  can  only 
attack  a  tree  through  an  existing  wound,  and  it  is  consequently 
termed  a  "  wound  parasite".  The  spores,  alighting  on  a 
wound  which  is  not  covered  with  resin  or  other  protective 
substance,  on  germination  produce  hyphse  which  penetrate 
into  the  stem.  The  hyphar  bore  through  the  cell-walls  and 
extract  the  lignin  in  them  so  that  practically  pure  cellulose 
is  left  behind.  Characteristic  white  spots,  which  eventually 
become  holes,  then  appear  in  the  attacked  wood,  see  Plate 
XVIII  (6).  The  growth  of  the  mycelium  may  also  cause  cup- 
shake,  or  ring-shake.  The  timber  is  thus  rendered  useless,  and 
if  the  tree  attacked  cannot  oppose  the  invading  mycelium  by 
a  copious  flow  of  resin  the  hyphse  may  spread  to  the  cortex 
and  kill  the  tree.  Wounds  caused  by  breaking  or  cutting  off 
old  branches  which  contain  heart-wood  are  favourite  points 
of  attack,  for  little  or  no  turpentine  exudes  from  the  central 
portion  of  such  wounds,  into  which  in  consequence  the  hypha3 
readily  penetrate.  For  the  same  reason  such  wounds  also 
afford  a  convenient  passage  free  of  resin  by  means  of  which 
the  mycelium  can  pass  from  the  interior  of  the  stem  to  the 
outside  for  the  purpose  of  producing  sporophores,  the  latter 
frequently  appearing  at  such  places.  Sporophores  are  not 
as  a  rule  produced  until  a  luxuriant  growth  of  mycelium  has 


195 

established  itself  in  the  interior  of  the  stem,  and  if  one  sporo- 
phore  is  removed  another  as  a  rule  soon    takes    its    place. 

Diseased    trees     must    be    removed    as    quickly  as   pos-  Preventive 
sible.     Wounding  and  injuring  living  trees  must  be  avoided,  and  Measures. 
cut  surfaces   should  be  covered  with  tar  or  other  protective  sub- 
stance.   So   far   as    we    can,  we  must  help   the  cambium    to 
cover  over  as  quickly  as  possible  all  wounds  which  may  arise. 

171.    It    has    been    estimated  (4)  Pyccmia 

that  the   annual  financial  loss   in   India,  on  account  of  fhegraminis 
damage  done  to  cereal  crops  by  the  fungoid  diseases  popularly  ^ers>. 
known  as  Rusts,   is  probably   not   less  than   Us.  4,00,00,000.  attacked. 
The   best   known  rust  in  India  is  Puccinia  graminis  which  is 
very  destructive  to  wheat,  and  which  we  may  also  take  as  a 
type  of  the  great  group  of  parasitic  fungi  to  which  the  rusts 
belong,  viz.  the  Uredinaceae. 

The  first  sign  of  the  disease   consists    in  the  appearance  of  Signs  of  the 
orange-red    streaks    and    patches  on  the  green  wheat  leaves,  Dlsease- 
culms  and  even  on   the   ears,  which  in  Northern  India  usually 
happens   in  January — February.      These  rust  pustules    swell 
and,    bursting    through    the    epidermal    tissue  of   the    wheat 
plant,    scatter   spores    which  look    like    reddish  dust  and  are 
called  uredospores. 

If  the  tissues  of  the  wheat  plant  are  examined  with  Life  History 
the  microscope  the  mycelial  threads  of  the  fungus  will  beand  Damage 
found  ramifying  in  all  directions  between  its  cells.  This done> 
mycelium  derives  its  nourishment  from  the  green  cells 
of  the  wheat  plant  and  consumes  the  food  materials 
which  should  go  to  form  the  grain ;  consequently  the  yield 
of  the  latter  is  enormously  reduced.  After  growing  and 
spreading  in  the  tissues  of  the  wheat  plant  for  two  or  three  weeks 
the  hyphse  turn  towards  the  epidermis,  and  the  orange,  oval, 
one-celled  uredospores  are  budded  off  from  their  tips ;  the 
gradual  accumulation  of  these  hyphal  branches  and  the  spores 
which  they  shed  result  in  the  swelling  of  the  pustule  and 
ultimate  rupture  of  the  epidermis  with  the  scattering  of  the 
spores.  If  these  spores  fall  into  water  they  germinate  and 
send  out  two  or  more  germ  filaments  [see  Plate  XIX  (2)],  one  of 
which  outstrips  the  rest  and  either  dies  in  the  absence  of  a 
suitable  host  or,  in  the  event  of  its  having  alighted  on  a  wheat 
plant  in  a  drop  of  rain  or  dew  (which  would  often  happen  in 
a  wheat  field),  finds  its  way  into  a  stoma  and  rapidly  gives 
rise  to  a  new  mycelium,  which  in  due  course  produces  a  fresh 
crop  of  uredospores.  The  function  of  these  spores  is  thus 
to  spread  the  disease  rapidly  from  plant  to  plant,  wind  aiding 


196 

their  distribution.  Now  as  the  season  progresses,  the  myce- 
lium which,  for  some  weeks,  has  only  produced  uredospores, 
now  begins  to  form  a  totally  different  kind  of  spore,  which 
are  known  as  teleutospores.  These,  instead  of  being  orange  or 
reddish  in  colour,  are  dark  purple -brown  or  black,  and  their 
formation  can  be  recognized  by  the  appearance  of  grey  streaks 
and  patches  among  the  orange  ones  which  spread  and  get  darker 
in  colour,  until  ultimately  the  black  pustules  entirely  replace  the 
rust- coloured  ones  which  preceded  them  [see  Plate  XIX  (1)]. 
These  teleuto -spores  are  formed  in  the  same  manner  as  the  uredo- 
spores and  by  the  same  mycelium,  the  latter  in  the  beginning 
of  the  season  producing  only  uredospores,  then  later  a  few 
teleutospores  among  the  uredospores,  and  ultimately  nothing  but 
teleutospores.  These  spores  which  are  formed  by  the  fungus 
in  the  later  stage  of  its  life-history,  besides  being  quite  distinct 
in  appearance  from  the  uredospores,  have  a  different  function, 
they  being  virtually  resting  spores.  Besides  the  difference  in 
colour,  each  teleutospore  instead  of  being  one-celled  is  divided 
across  the  middle  into  two  cells  or  chambers.  They  are  more 
elongated  and  club-like  in  shape  and  have  a  thicker  wall.  They 
are  also  as  a  rule  not  shed  so  quickly  as  the  uredospores  but 
remain  firmly  attached  to  the  wheat  straw.  They  are  generally 
incapable  of  germinating  for  some  months  after  their  formation. 
When  they  do  germinate  each  cell  sends  out  a  short  hypha 
which  then  becomes  divided  by  cross  partition  walls  into 
segments.  From  each  segment  a  short  lateral  stalk  (sterigma) 
is  produced,  the  tip  of  which  swells  and  then  falls  off — the 
minute,  ovoid,  spore-like  bodies  so  formed  being  called  sporidia. 
The  short  hyphal  filament  developed  from  the  teleutospore  is 
known  as  the  promycelium.  See  Plate  XIX  (3). 

Now  in  Europe  it  was  found  that  these  sporidia  could  only 
develop  if  placed  on  a  new  host-plant,  a  species  of  Berberis,  i.e. 
an  altogether  distinct  species  from  that  on  which  the  teleuto- 
spores were  formed.  Germinating  then  on  the  leaf  of  a  suitable 
host-plant,  these  sporidia  send  out  their  germ  tubes  which 
directly  pierce  the  leaf -tissue  of  the  Berberis  and  develop  a 
parasitic  mycelium  within  it,  the  mycelium  deriving  its  nourish- 
ment from  the  cells  of  the  Berberis  tissue.  Although  the 
Berberis  plant  is  thus  more  or  less  injured,  the  damage  done  is 
usually  nob  severe,  and  the  great  practical  importance  of  this 
fungus  depends  on  the  injury  done  to  the  wheat  plant,  which 
resubs  in  an  enormous  decrease  in  the  yield  of  grain.  The 
presence  of  the  mycelium  in  the  Berberis  induces  swellings  in  the 
leaf  tissue  on  which  two  different  kinds  of  spores  are  produced. 


197 

The  first  to  appear  called  spermatia  arise  in  small  cup-like  recep* 
tacles  on  the  upper  surface  of  the  leaf.  They  are  very  minute, 
and  their  function  is  at  present  unknown.  On  the  under-surface 
of  the  leaf,  larger,  golden-yellow,  cup -like  receptacles  arise,  which 
are  clearly  visible  to  the  naked  eye  and  are  often,  known  by 
the  popular  term  of  "cluster-cups".  These  cups  have  fringed, 
white  margins  and  are  rilled  with  golden-yellow  spores.  They 
arise  first  beneath  the  epidermis  as  hollow  balls,  the  membranous 
walls  of  which  are  formed  by  the  fungal  hyphse  and  constitute 
what  is  known  as  the  peridium.  The  base  of  these  balls  is 
formed  by  a  number  of  short  hyphae  with  their  ends  directed 
towards  the  apex  of  the  ball.  From  the  ends  of  these  hyphae 
spores  are  ab jointed  in  succession  from  the  apex  downwards, 
long  chains  of  spores  being  thus  produced.  As  the  formation 
of  spores  proceeds,  the  ball  swells  and  ultimately  the  epidermis 
of  the  leaf  and  the  peridium  covering  are  ruptured  and  the  ball 
becomes  a  cup-shaped  receptacle  projecting  from  the  leaf  sur- 
face. As  the  spores  escape  from  these  receptacles  and  are  dis- 
tributed by  the  wind,  fresh  ones  are  developed  from  the  hyphae 
at  the  base  to  replace  them.  See  Plate  XIX  (4),  (5)  and  (6).  This 
form  of  the  fungus  developed  upon  the  Berberis  was  at  first 
thought  to  belong  to  a  distinct  genus  from  that  which  included 
the  form  occurring  on  the  wheat ;  it  consequently  received  the 
name  of  Aecidium  Berberidis,  and  it  was  long  before  the  connec- 
tion between  the  two  was  established.  The  cluster  cups  are 
called  aecidia  and  the  spores  developed  in  them  aecidiospores. 
These  spores  are  very  like  the  uredospores  except  that  they  are 
more  golden-yellow  in  colour  and  are  often  somewhat  polygonal 
in  form,  owing  to  the  pressure  which  they  have  undergone. 
These  spores,  alighting  on  a  damp  wheat  plant,  send  out  germ 
tubes  which,  finding  their  way  through  the  stomata,  give  rise  to 
the  mycelium  in  the  plant  tissue  from  which  uredo-  and  teleuto- 
spores  are  ultimately  produced  as  before.  Aecidium  Berberidis 
is  found  on  Berberis  Lycium  near  Simla  and  also  probably  occurs 
on  other  Indian  species  of  Berberis. 

We  thus  see  that  the  complete  life-history  of  this  fungus 
comprises  two  distinct  stages  which  may  be  called  the  Pucci- 
nial  and  Aecidial  respectively,  and  these  stages  may  be  sub- 
divided into  four  as  follows  :— 

C     (1)  The  parasitic    mycelium  in  the  wheat 
Puccinial  I  plant    developed    from     aecidiospores     and 

,  "!  giving  rise  to  uredospores. 

i      (2)  The  formation  of   teleutospores   from 

Lthe  same  mycelium. 


198 

f     (3)  The  germination  of  the  teleutospores 
|  giving  rise  to  a  promycelium  and  sporidia. 
Aecidial       J  This  is  a  non-parasitic  stage, 
stage.  (4)  The   germination  of    the    sporidia  to 

J  form  a  parasitic  mycelium  which  ultimately 
(^produces  secidiospores. 

In  our  typical  example  the  puccinial  and  aecidial  stages 
are  passed  on  different  host-plants ;  such  a  fungus  is  said  to  be 
heteroecious,  and  the  host  which  nourishes  the  aecidial  stage  is 
in  this  case  called  an  intermediate  host.  Other  allied  fungi 
pass  both  the  puccinial  and  the  aecidial  stage  on  the  same 
host-plant  andare  consequently  called  autoecious. 

Now  in  the  central  areas  of  India  Puccinia  graminis  is  one 
of  the  commonest  wheat  rusts,  and  it  has  been  found  widely  spread 
in  places  over  600  miles  from  any  species  of  Berberis  which 
in  India  are  confined  to  the  high  hill  ranges.  In  Europe  also 
it  has  been  found  that  wind-blown  secidiospores  can  ordinarily 
only  infect  plants  within  a  radius  of  25  yards  from  the  bush 
on  which  they  arise,  and  hence  it  is  improbable  that  wind- 
blown a3cidiospores  could  in  such  cases  be  responsible  for  the 
disease.  This  rust  also  is  common  in  parts  of  Australia, 
e.g.  in  Victoria,  where  no  Berberis  grows  wild  and  very  few  are 
cultivated.  Again,  if  the  secidial  stage  on  the  Berberis  is 
always  necsssary  for  the  continued  existence  of  the  fungus, 
the  complete  destruction  of  all  species  of  Berberis  should  suffice 
to  eradicate  the  disease  in  the  areas  treated.  This,  however,  has 
not  been  found  to  be  the  case.  Finally  Aecidium  Berberidis  is, 
in  India,  restricted  to  a  portion  of  the  Himalayan  Eange,  and 
it  is  precisely  in  this  area  that  Puccinia  graminis  is  extremely 
rare  on  cereals.  We  are  therefore  driven  to  the  conclusion 
that  the  fungus  can,  and  often  does,  dispense  with  the  a^cidial 
stage  and  continues  to  exist  on  the  wheat,  year  after  year,  in 
the  puccinial  stage.  How  this  is  managed  is  not  yet  clearly 
understood.  It  has  been  suggested  that  the  uredospores  may 
pass  from  the  wheat  to  some  wild  perennial  grass  and  that  the 
fungus  may  thus  continue  to  maintain  itself  on  the  latter  in  the 
puccinial  stage  until  the  season  for  the  next  wheat  crop  comes 
round  again.  In  Europe  this  fungus  has  been  found  on  150 
different  species  of  grasses,  including  oats,  wheat,  barley  and 
rye,  these  being  known  as  collateral  hosts.  It  has,  however,  been 
found  that  the  form  on  wheat  is  exclusively  limited  to  the  wheat 
and  cannot  impart  the  disease  to  any  other  collateral  host, 
with  the  sole  exception  of  rye  and  barley,  which  in  rare  cases 


199 

may  be  infected.  On  the  other  hand,  uredospores  from  other 
collateral  hosts  fail  to  infect  the  wheat,  except  in  rare  cases 
under  very  favourable  conditions.  It  has  in  fact  been  clearly 
established  that  this  wheat  rust  is  a  highly  specialized  form  of 
the  fungus  which  possesses  very  little  capability  of  directly 
passing  on  to  and  infecting  other  nearly  allied"  species  of 
plants.*  Another  suggestion  is  that  the  uredospores  may  live  in 
the  soil  during  the  period  when  wheat  is  out  of  the  ground 
(between  April  and  November)  and  be  able  to  infect  the  young 
crop  in  December — January.  Experiments,  however,  with 
allied  fungi  have  shown  that  the  uredospores  soon  lose  their 
power  of  germination,  especially  when  they  are  exposed  to  a 
high  temperature,  as  they  would  be  in  the  hot  season  in  the 
plains  of  India. 

These  explanations  therefore  can  hardly  account  for  the  per- 
sistence of  the  disease.  Again  it  has  been  suggested  that  the 
sporidia  may  be  able  to  give  rise  to  a  mycelium  in  the  wheat 
plant  capable  of  producing  uredo-  and  teleuto-spores.  This  is 
improbable  and  there  is  no  evidence  to  support  it.  The  most 
recent  researches  on  the  subject  indicate  that  the  rust  on  wheat 
may  be  a  truly  hereditary  disease.  The  protoplasm  of  the 
fungus  is  said  to  be  able  to  exist  in  intimate  union  with  the 
protoplasm  of  the  wheat  plant,  and  its  presence  then  can  only 
be  detected  with  great  difficulty.  This  state  of  things  may 
continue  for  months,  or  even  years,  the  disease  germs  passing 
on  from  generation  to  generation  in  the  seed,  and  it  is  only 
under  certain  favourable  conditions  of  moisture,  heat,  and  of  the 
state  of  the  host-plant  itself,  that  the  fungus  becomes  distinctly 
visible  in  the  form  of  a  mycelium,  which  takes  place  just  before 
the  rust  pustules  begin  to  appear. 

The  only  practical  measures  at  present  available  for  com-  Preventive 
bating  this  fungus  consist  in  selecting  the  most  rust-resistant  Measur^s> 
varieties  of  wheat  for  continued  cultivation  and  in  endeavour- 
ing to  produce  new  and  improved  varieties  by  inter-crossing. 


*  Seeing  that  the  various  forms  of  this  fungus,  occurring  on  different  collateral 
hosts,  are  each  more  or  less  confined  to  some  special  species  of  plant  we  cannot 
doubt  that  in  some  way  they  differ  essentially  from  one  another.  At  the  same 
time,  so  far  as  visible  microscopic  characters  are  concerned,  these  forms  cannot  be 
distinguished  and  hence  they  are  all  grouped  under  the  species  Puccinia  graminis. 
Here  we  appear  to  have  an  example  of  the  power  possessed  by  some  plants  of 
adapting  themselves  to  changed  conditions,  and  we  see  that,  having  become 
accustomed  to  one  set  of  conditions,  they  do  not  readily  adapt  themselves  to 
another,  although  hi  time  a  specialized  form  may  arise  which  can  do  so. 


200 


Aecidium 
mnntanum. 


Parasitic 
Phanero 
gams. 


172.  The  life-history  of  several 

other  rust-fungi  exhibit  the  two  stages  described  above,  viz.  the 
Puccinial  and  Aecidial,  but,  as  our  selected  example  clearly 
shows,  these  two  stages  are  not  always  necessary.  Thus  there 
are  species  of  Puccinia  of  which  no  Aecidial  form  is  known 
and  vice  versa.  As  an  example  of  an  Aecidium  of  which  no 
puccinial  stage  is  known,  we  may  take  the  "  cluster-cups  ' '  so 
often  seen  on  Berberis  Lycium,  B.  coriaria,  and  B.  aristata,  in 
Jaunsar  and  near  Mussoorie.  This  species  has  been  named 
Aecidium  montanum.  The  mycelium  of  this  species  is  peren- 
nial, the  hyphae  running  in  the  cell  walls  and  intercellular  spaces 
of  the  stem  and  leaves  and  obtaining  food  by  means  of  little 
finger-like  haustoria  pushed  through  the  walls.  Its  presence 
in  the  tissues  of  the  Berberis  causes  the  formation  of  very 
characteristic  witches'  brooms.  The  affected  shoots  become 
dwarfed,  bear  malformed  leaves  and  grow  vertically  upwards, 
the  long,  yellow,  aecidial  cups  occurring  in  masses  on  the  under- 
side of  the  leaves  and  also  scattered  on  the  shoots  and 
sometimes  on  the  peduncles,  from  which  the  clouds  of  powdery 
orange  aecidiospores  are  shed  and  disseminated  by  the  wind. 
Sometimes  more  than  half  the  bush  may  be  affected  in  this  way. 
A  healthy  bush  on  first  infection  shows  large,  reddish,  or  scarlet, 
patches  on  the  upper  surface  and  numbers  of  the  long  aecidial 
cups  below.  In  this  case  there  is  no  deformity  beyond  a  pucker- 
ing of  the  leaves,  and  it  is  uncertain  whether  first  infection  takes 
place  by  aecidiospores  or  by  sporidia.  If  by  the  latter,  the 
puccinial  form  bearing  the  teleutospores  is  still  unknown. 
This  species  is  distinguished  from  A.  Berberidis  by  the  witches' 
brooms,  by  the  greater  length  of  the  aecidial  cups,  which  may 
amount  to  4  mm.,  and  by  the  greater  size  of  the  patches  seen 
on  newly  affected  normal  leaves,  which  ranges  from  J  to  f  of 
an  inch  in  diameter. 

173.    The     following     may  be 
taken  as  typical   examples  of  parasitic  phanerogams  : — 


(1)  Orobanche  indica. 

(2)  Cuscuta  reflexa. 

(3)  Loranihus  longiflorus. 


(1)  Orobanche 
indica. 


174.  The    handsome   spikes  of 

pale  purple,  or  blue,  flowers  of  this  plant  are  often  seen 
in  the  mustard  fields  of  Northern  India,  and  the  total 
absence  of  green  leaves  at  once  strikes  us  as  peculiar. 


201 

This  species  being  thus  unable  to  manufacture  for  itself  the 
carbonaceous  organic  food  materials  which  it  requires  is  obliged 
to  obtain  them  ready-made  from  other  plants.  If  the  soil  is 
carefully  removed  from  the  roots  of  an  Orobanche  plant  these  are 
found  to  be  joined  to,  and  in  intimate  connection  with, 
the  roots  of  the  neighbouring  Jmustard  plants  from~which  they 
take  their  necessary  food  materials. 

175.  This    is    the    well-known  (2)  Cuscvta 

leafless  Dodder,  masses  of  whose  wire-like  stems  may  often  be  seen  reftexa' 
enveloping  trees  and    shrubs  with  a    yellow,     or     greenish, 
mantle.      Although    the   plant    contains  a   little    chlorophyll, 

this  is  quite  insufficient  for  the  manufacture  of  the  food 
it  requires.  If  we  examine  the  twining  stem  we  find  that, 
where  it  comes  in  contact  with  the  stem  or  twigs  of  the 
host,  small  wart-like  protuberances  are  developed  which 
pass  into  and  are  in  intimate  connection  with  the  tissues  of 
the  host  from  which  it  derives  its  water  and  nourishment. 
These  suckers,  or  haustoria,  are  able  to  force  their  way  into  the 
tissues  of  the  host  without  difficulty,  partly  owing  to  the  action 
of  dissolving  enzymes  which  their  filaments  excrete  after  the 
manner  of  fungal  hyphae.  The  seed  of  this  species  germinates 
on  the  ground,  but  the  rootlet  which  is  developed  soon  dies  off, 
and  if  the  filamentous  stem  fails  to  encounter  a  suitable  host 
the  seedling  perishes.  If  allowed  to  develop  undisturbed 
the  Dodder  may  kill  the  plant  attacked,  and  gaps  in  garden 
hedges  caused  by  it  may  often  be  seen. 

176.  This    very    common   plant  (3)L<"anthu3 

.  ,  f   .  ,  •      i         P     ji  •       longiftorus. 

may  be  taken  as  iairly  typical  ot  the  numerous  species 
of  Loranthus  and  Viscwn  which  occur  in  Indian  forests. 
Their  chief  peculiarity  consists  in  the  fact  that  they  pos- 
sess a  considerable  supply  of  chlorophyll,  many  of  them,  like 
our  example,  being  provided  with  well-developed  green  leaves. 
This  plant  appears  to  be  chiefly  distributed  by  birds  who 
deposit  the  seeds  on  the  branches  and  stems  of  trees  where 
they  germinate  and  grow.  The  young  radicle,  piercing  through 
the  cortex  of  the  branch  or  stem  attacked,  penetrates  to  the 
young  wood,  in  the  tissues  of  which  it  spreads  and  develops. 
This  plant  obtains  its  supplies  of  water  and  mineral  salts 
directly  from  the  wood  elements  of  its  host  and  the  portion  of 
the  branch  or  stem  above  the  point  of  attachment  of  the  para* 
site,  being  deprived  of  its  water-supply,  dies  and  we  get  stag- 
headed  trees.  The  attacked  branches  become  swollen  and  dis- 
figured while  the  parasitic  roots,  being  embedded  in  the  wood, 


202 


render  the  latter  more  or  less  worthless  for  construction.  It 
is  probable  that  this  plant  transfers  some  of  its  carbonaceous 
food  materials  to  its  host,  but  it  is  not  known  whether  it  absorbs 
any  organic  food  substances  from  the  host  in  addition  to  the 
supplies  of  v,Tater  and  mineral  salts.  The  chief  practical 
method  oi  combating  this  pest  consists  in  cutting  off  and  de- 
stroying the  attacked  branches. 

177.  Among  parasitic  phane- 
rogams which  are  furnished  with  an  abundance  of  chlo- 
rophyll the  well-known  sandal  tree  (Santalum  album,)  re- 
quires brief  notice.  It  is  now  known  that  this  tree  is 
dependent  to  a  very  large  extent  on  the  roots  of  the  plants 
near  which  it  grows  for  its  supplies  of  water  and  mineral  salts, 
the  Sandal  roots  developing  numerous  haustoria  which 
penetrate  the  root  tissues  of  the  host-plants  and  draw  from 
them  the  necessary  supplies  of  food  materials.  No  definite 
information  is  at  present  available  as  to  the  extent  to 
which  the  host-plants  are  damaged  by  this  parasitism,  and  it 
appears  probable  that  they  may  benefit  to  some  extent  by  a 
supply  of  carbonaceous  food  which  has  been  manufactured  in 
the  green  sandal  leaves,  in  which  case  the  partnership  would 
be  more  or  less  a  reciprocally-symbiotic  one.  Numerous  costly 
attempts  have  been  made  to  artificially  propagate  sandal  in  pure 
plantations  which,  owing  to  the  absence  of  other  trees  and  shrubs, 
were  of  course  failures,  and  although  this  tree  is  of  such  great 
economic  importance  it  is  a  remarkable  fact  that  its  para- 
sitism was  until  quite  recently  regarded  as  uncertain.  It 
is,  therefore,  not  a  matter  for  surprise  that  the  artificial  pro- 
pagation of  this  tree  was  frequently  a  failure,  or  that  the 
notorious  "  spike  "  disease  from  which  it  suffered  had  baffled 
all  attempts  at  explanation  and  prevention.  Until  we  know 
the  principal  conditions  essential  for  the  healthy  development 
of  our  important  species,  it  is  obvious  that  we  -cannot  expect 
to  propagate  them  successfully  or  to  discover  the  cause  of, 
or  remedy  for,  any  disease  which  may  attack  them.  Hence 
we  must  recognize  the  paramount  importance  of  studying 
the  life-histories  of  our  forest  plants  and  the  relations  which 
exist  not  only  between  them  and  their  non-living  environ- 
ment but  also  between  them  and  other  living  organisms. 

Leaving  parasites   we   now   pass   on   to :— 


Lichens. 


example     of 


(c)  Symbionts. 

178.    Perhaps  the  most    typical 
symbiosis     among     plants      is     afforded     by 


203 

the  lichens.  As  has  been  seen  above,  a  lichen  con- 
sists of  two  distinct  plants,  viz.  an  alga  and  a  fungus,  see  page 
137.  The  fungus  receives  carbohydrates  from  the  alga  in 
return  for  which  it  protects  the  alga  from  drought  and  pro- 
vides it  with  water  and  mineral  salts. 

179.  Another  well-known  exam-  Bacteriaand 

pie  is  furnished  by  the  bacteria  which,  as  noted  in  Part  IV  Leguminosae. 
above,  see  page  129,  live  in  the  tubercles  often  found  on  the  roots 
of  the  Pea,  Bean  and  allied  plants  belonging  to  the  Natural 
Order  Leguminosae.  The  best  known  of  these  bacteria  is  named 
Bacillus  radicicola  which,  passing  through  the  root  hairs, 
gives  rise  to  the  peculiar  tubercles  in  the  root  cortex  of  the 
infected  plants.  The  bacteria  are  localized  in  these  tubercles 
and  do  not  spread  to  neighbouring  tissues.  While  the  bac- 
teria live  on  the  carbohydrates  supplied  by  the  host  they, 
in  their  turn,  are  able  to  provide  the  host  with  valuable 
nitrogenous  food  materials,  owing  to  the  power  possessed  by 
them  of  fixing  the  free  nitrogen  of  the  air  and  of  manufac- 
turing from  it  a  compound  which  can  be  utilized  as  food  by 
higher  plants.  Owing  to  this  remarkable  symbiosis  by  means 
of  which  most  leguminous  plants  can  accumulate  nitrogenous 
organic  material  in  their  tissues,  a  crop  of  such  plants  can 
actually  enrich  a  soil  poor  in  nitrates.  Hence  in  agriculture 
it  is  a  common  practice  to  grow  such  plants  and  to  plough 
in  the  crop  as  manure.*  In  a  soil  rich  in  nitrates,  leguminous 
plants  can  obtain  their  nitrogenous  food  without  the  aid 
of  these  bacteria,  and  in  such  cases  very  few,  if  any,  tubercles 
are  formed. 

180.      The     majority      of    the  Mycorhizae. 

higher  plants  cannot  directly  utilize  as  food  the  organic 
substances,  such  as  the  dead  remains  of  plants,  wood,  leaves, 
etc.,  which  compose  the  humus  of  the  Forester.  Hence 
such  plants  are  found  to  grow  normally  when  supplied 
solely  with  water  and  the  necessary  mineral  salts.  Many 
plants,  however,  such  as  pines,  oaks,  the  hazel  and  others, 
can  thrive  on  humus  soils,  and  in  the  majority  of  such 
plants  their  roots  are  found  to  be  living  in  symbiosis  with 
the  mycelia  of  fungi.  The  fungal  hyphse  are  sometimes 
found  chiefly  inside  the  cells  of  the  root  cortex,  a  few  fila- 

*lf  leguminous  plants  could  be  grown  continually  on  the  same  soil  the  supply 
of  nitrogen,  in  the  latter  might  clearly  be  increased  indefinitely,  but  unfortu- 
nately this  is  not  at  present  possible.  Experiments  in  Europe  have  shown  that 
clover  can  only  be  successfully  grown  on  the  same  soil  about  once  in  four  37ears 
and  sometimes  even  not  so  frequently.  The  reason  for  this  is  at  present  unknown. 


204 


Distant 

Symbiosis 

between 

Green  and 

Non-green 

Plants. 


Decay  of 
Wood. 


ments  here  and  there  extending  into  the  soil,  and  at  other 
times  they  cover  the  roots  externally  with  a  felt-like  mantle. 
Such  mycelial  growths  are  called  mycorliizas.  In  Europe 
it  has  been  found  that  if  trees  like  the  oak  and  pine  are  grown 
in  sterile  soil  and  are  thus  deprived  of  these  mycorhizas  their 
growth  is  retarded  and  sometimes  entirely  prevented.  The 
precise  nature  of  the  symbiotic  relation  in  this  case  is  not  yet 
known,  but  it  is  probable  that  the  host  supplies  the  fungus 
with  carbohydrates,  while  the  fungus  helps  the  host  to  obtain 
water,  mineral  salts  (probably  phosphates  and  potassium 
chiefly)  and  simple  nitrogenous  compounds. 

181.  The  above  are  all  instances 

of    close  symbiosis,   i.e.    in  which    two     organisms    live    in 
intimate  connection  with  each  other.     As  an  example  of  distant 
symbiosis  we  may  take  that  existing  between  green  and  non- 
green  plants. 

We  have  had  frequent  occasion  to  mention  above  the 
decomposing  action  of  many  bacteria  and  fungi  which  give 
rise  to  various  processes  popularly  called  rotting,  putrefaction, 
fermentation,  nitrification,  etc.,  such  organisms  being  able 
to  feed  on  complex  organic  materials  and  to  break  them  down 
into  simpler  substances  on  which  the  higher  green  plants 
feed.  By  the  activity  of  such  organisms,  for  instance,  the 
carbon  dioxide  required  by  green  plants  is  being  continually 
returned  to  the  air,  while  nitrogenous  organic  substances 
such  as  albuminoids,  amides,  and  others  are  also  broken  down 
and  nitrates  produced  which  are  the  chief  source  of  nitro- 
genous food  for  higher  plants.  Many  of  these  decomposing 
processes  are  accompanied  by  a  liberation  of  energy  in  the 
form  of  heat,  and  we  know  that  the  heat  may  be  so  great, 
in  a  stack  of  rotting  hay,  for  instance,  that  spontaneous  com- 
bustion is  produced. 

182.  Before    leaving   this   sub- 
ject of  decay   a    few    words   will    be   added    on    the    decay 
of    timber.       In    the    first    place    care    must    be   taken     to 
distinguish  between    the    terms   dead    and    decayed.      Plant 
tissues   become   dead  when   their   elements    lose   their  living 
protoplasmic   contents,    they   do   not   become   decayed   until 
they  have  undergone  a  further  process  in  which  they  become 
disintegrated  and  their  structure  is  destroyed.     A  piece   of 
sound,  Teak  heart- wood,  which  is  really  dead  tissue,  and  which 
is  known  to  be  the  best  and  most  durable  wood  for  construc- 
tion, compared  with  a  fragment  from  a  hollow  tree  which 


205 

can  be  crumbled  in  the  fingers  will  illustrate  the  difference. 
For  the  decay  and  destruction  of  structural  timber  fungi 
are  mainly  responsible,  their  hyphae  excreting  enzymes  which 
destroy  the  wood  tissue,  as  has  been  made  clear  in  the  examples 
of  Fomes  and  Trametes,  and  it  is  therefore  important  to 
remember  that,  as  a  general  rule,  the  conditions nrrost  favour- 
able for  the  development  of  such  fungi  are:— 

(1)  A  supply  of  organic  food  materials  such  as  carbohy- 

drates   and    proteid    substances    which    are    com- 
monly found  in  sapwood. 

(2)  Warmth. 

(3)  A   liberal   supply    of   moisture    and   absence   of   free 

ventilation. 

(4)  A  supply  of  oxygen. 

Thus  sapwood,  especially  if  kept  damp,  quickly  decays, 
and  in  the  case  of  posts  driven  into  the  ground  we  frequently 
see  that  decay  spreads  most  rapidly  in  that  portion  of 
the  post  situated  in  the  upper,  well-a3rated,  layers  of  soil. 

Seeing  also  how  fungal  hyphse  may  spread  from  a  decayed 
piece  of  wood  to  a  sound  piece  in  contact  with  it,  and  how 
easily  the  minute  spores  are  distributed,  it  is  clear  that  the 
greatest  care  must  be  taken  if  we  wish  to  preserve  sound 
wood  from  infection. 

SECTION  III.— INFLUENCES    OF    THE    SOIL   ON   PLANT   DEVE- 
LOPMENT. 

183.  For  the  healthy  develop-  Presence  of 

ment  of  plants  the  soil  must  contain  a  sufficient  supply  of  all  ^eoewaiy 
the    essential    mineral    salts.      These   may,    for   instance,   be  suitable  Con- 
so     diminished    by    the    competing    roots     of     neighbouring  dition  and  in 
plants     as    to    cause     disease    and     death,     the  absence     of  •^lltab|e. 
any  one  of  them  making  growth   impossible.     Further   these 

salts  must  exist  in  such  a  state  of  combination  as  to  enable 
the  plant  to  make  use  of  them  as  food  materials,  and  they 
must  not  be  present  in  excess.  The  best  nutrient  solution 
for  most  phanerogams,  for  example,  should  not  contain  more 
than  0*5  per  cent,  of  salts. 

184.  Any  substance  which  may  Presence  of 

cause    disease  or  death  in  plants  may  be  broadly  classed  as  a  Substances. 
Poison.     If  the  solution  of  salts  in  the  soil  becomes  concentrated 
beyond  a  certain  limit,  rarely  exceeding  five  per  cent,  owing  to 

the  osmotic  processes  being  interfered  with,  the  roots  of  plants 


206 

are  no  longer  able  to  obtain  their  necessary  supplies  of  water 
from  the  soil  and  are  thus  in  danger  of  desiccation.  At  the 
same  time  plants  vary  in  this  respect  and  there  are  many 
(usually  known  as  halophytes)  which  can  thrive  on  saline 
soil  and  are  able  to  recover  the  necessary  water  from  con- 
centrated solutions  of  salts.  There  are  also  many  substances 
which  have  a  more  directly  poisonous  effect  on  plants  and 
such  are  as  a  rule  the  salts  of  the  heavy  metals,  e.g.  sulphate 
of  zinc  and  others.  Free  acids  and  alkalies  are  also  very 
poisonous  even  when  dilute.  Carbon  dioxide  is  continually 
excreted  by  the  roots  of  plants  and,  in  water-logged  localities, 
the  accumulation  of  this  gas  may  have  a  poisonous  effect. 
The  escape  of  coal-gas  from  underground  pipes,  or  chemical 
solutions  from  factories,  such  as  dye-works,  may  also  kill  and 
injure  plants.  Again  many  essential  mineral  salts,  and  others 
which,  though  not  essential,  are  often  found  to  be  taken  up 
by  plants,  may  have  a  poisonous  effect  if  present  in  large 
quantities.  A  trace  of  iron  is  thus  essential  for  plants,  but 
a  very  dilute  solution  of  an  iron  salt  may  be  poisonous,  the 
quantity  of  the  salt  present  being  above  the  optimum.  Cal- 
cium appears  to  be  injurious  in  some  cases,  while  as  regards 
common  salt  it  is  not  clear  if  its  injurious  effect  is  merely 
due  to  its  osmotic  properties  or  to  a  more  direct  poisonous 
action.  In  the  majority  of  cases  the  exact  way  in  which 
the  poison  acts  is  not  known,  but  sometimes  at  all  events 
the  poisonous  substance  appears  to  enter  into  chemical 
combination  with  some  constituents  of  the  protoplasm,  much 
as  in  the  case  of  animals  carbon  monoxide  combines  with 
haemoglobin.  Humus  by  its  absorbing  action  has  a  most 
beneficial  effect  in  protecting  plants  from  poisons.  Poisonous 
metallic  salts  may  thus  be  strongly  held  by  the  humus  and  only 
allowed  to  pass  into  the  soil  in  very  dilute,  harmless  solutions. 
The  principal  product  which  is  excreted  in  large  quantities 
by  the  roots  of  higher  plants  is  carbon  dioxide,  and  this  being 
a  gas  rarely  accumulates  sufficiently  to  be  injurious. 

Many  plants  are  also  able  to  accumulate  considerable 
quantities  of  poisonous  substances  without  injury,  by 
depositing  them  in  dead  tissues,  or  in  living  ones  in  such  a  way 
that  they  cannot  penetrate  the  living  protoplasm. 

185'  Umess  the  root-hairs   are 
Water  and    able  to  obtain  a  sufficient  supply  of  water  and  oxygen  in  the 
Oxygen.        soil  they  cannot  perform  their  functions  properly,  and  a  good 
soil,  besides  being  sufficiently  moist,  must,  therefore,  be  open  and 
well  aerated.     The   amount  of  available   water  in   a    forest 


207 

soil  may  be  decreased  by  a  diminished  rainfall  and  by  the 
more  or  less  complete  failure  of  the  monsoon,  by  draining, 
by  the  removal  of  litter,  or  by  interrupting  the  leaf  canopy, 
and  this  may  cause  trees  to  become  stag-headed.  The  same 
result  may  be  caused  by  underplanting  trees  with  a  species 
which  absorbs  large  quantities  of  moisture  fronT  the  soil. 
In  an  ordinary  well-drained  soil  oxygen  is  always  present 
as  gas  in  the  interspaces  and  also  dissolved  in  the  water.  If 
this  oxygen  is  removed  the  root-hairs  are  killed  and  the  roots 
rot,  the  trees  becoming  stag-headed.  This  may  be  caused 
by  water  being  allowed  to  stagnate  around  the  roots  as  hap- 
pens in  swampy  ground,  and  it  commonly  occurs  also  in  dense 
soils  and  in  soils  with  an  impermeable  substratum.  The 
available  oxygen  in  such  cases  is  gradually  exhausted,  that 
utilized  by  the  roots  being  not  replaced  sufficiently  quickly 
by  the  admission  of  fresh  air.  The  same  result  may  be  caused 
by  other  factors  which  interfere  with  the  free  aeration  of  the 
soil,  such  as  heavy  grazing  which  hardens  and  increases  the 
density  of  the  soil.  The  roots  of  plants  grown  in  glazed 
pots  are  unable  to  obtain  a  sufficient  supply  of  oxygen  and 
often  rot  in  consequence.  Similarly  trees  growing  in 
towns  are  often  unhealthy,  owing  to  the  buildings  and 
close  pavements  preventing  sufficient  water  and  air  entering 
the  soil. 

The  free  aeration  of  the  soil  is  also  important  for  another 
reason.  We  have  seen  above  that  organic  substances,  such  as 
humus,  cannot  be  directly  used  by  the  higher  plants  as  food, 
although  such  substances  always  contain  a  certain  quantity 
of  the  elements,  such  as  phosphorus,  potassium,  nitrogen, 
etc.,  which  are  essential  for  the  existence  of  such  plants.  With- 
out the  aid  therefore  of  fungi  and  bacteria  which  break  down 
this  organic  material  and  liberate  these  elements  in  a  suitable 
condition,  they  could  not  be  utilized  by  the  higher  plants 
and  consequently  humus  would  accumulate  in  our  forests 
and  the  soil  would  gradually  become  poorer  in  the  essential 
food  materials,  until  eventually  no  trees  could  grow  in  it. 
Now  a  sufficient  supply  of  oxygen  is  absolutely  necessary 
for  the  existence  of  a  large  proportion  of  these  useful  fungi 
and  bacteria,  and  they  are  in  consequence  found  in  the  upper 
layers  of  the  soil,*  where,  moreover,  their  presence  is  most 

*  Experiments  have  shown  that  when  impure  sewage  water  containing  large 
numbers  of  bacteria  is  passed  through  sandy  soil  most  of  the  bacteria  are  retained 
in  the  superficial  well-aerated  layers  of  soil,  on  which  fact  the  system  of  filtering 
water  throug'i  sandy  soil  is  chiefly  based. 


208 


Accumula- 
tion of 
Starch  in 
the  Leaves 
indicates 
Root 
trouble. 


needed,  seeing  that  this  is  where  organic  debris  chiefly 
accumulates.  Thus  from  this  point  of  view  alone  we 
see  how  enormously  important  is  the  proper  aeration  of 
the  soil,  as  also  is,  of  course,  the  quantity  of  moisture  in  the 
soil,  and  its  temperature,  and  indeed  all  factors  which  may 
favour  or  retard  the  development  of  these  useful  living 
organisms.* 

186.  Now  if  for  any  of  the  above 

reasons  a  plant  is  unable  to  obtain  its  necessary  supplies  of  water 
and  mineral  salts  from  the  soil,  the  carbohydrates  manufactured 
in  the  leaves  cannot  be  converted  into  plastic  materials 
and  thus  be  conducted  away  to  the  growing  tissues  which 
require  them,  but,  owing  to  the  deficiency  of  water,  have  to  be 
put  aside  as  starch,  and  hence  an  accumulation  of  starch  grains 
in  the  leaves  is  often  an  indication  that  we  must  look  to  the 
roots  for  the  cause  of  a  disease. 


ture. 


SECTION  IV. — INFLUENCES   OF   THE  ATMOSPHERE    ON   PLANT 

DEVELOPMENT. 

187.  For    every    external    factor 

which  affects  the  growth  and  development  of  a  plant,  such 
as  temperature,  light,  etc.,  there  is,  as  has  already  been  noted 
in  Part  1 1 1,,  a,  certain  degree  of  intensity  called  the  optimum 
which  is  most  favourable  to  the  plant,  see  page  86.  Any 
Tempera8  *  considerable  departure  from  this  limit  may  cause  disease  and 
if  carried  far  enough  death.  Thus,  as  regards  temperature, 
the  death  of  plants  may  be  caused  either  by  excessive 
heat  or  cold.  The  power  of  a  plant  to  withstand  extremes 
of  temperature  depends  chiefly  on  the  species  of  plant  and  its 
stage  of  development.  Most  flowering  plants  are  killed  by 
a  long  exposure  to  a  temperature  of  45°  C.,  while  some  bacteria 
grow  well  at  a  temperature  of  70°  C.  Potato  plants  may  be 
killed  in  one  night  by  a  temperature  of — 4°  C,  while  some  bacteria 
can  withstand  a  temperature  of — 200°  C.  Khair  (Acacia  Catechu) 
and  Ber  (Zizyphus  Jujuba)  are  frost-resistant,  while  Aonla 
(Phyllanthus  Emblica)  is  frost-tender  in  the  climates  where 

*  In  connection  with  this  breaking  down  of  organic  material  by  fungi  and 
bacteria  it  is  interesting  to  note  that  this  process  may  not  only  result  in  the  pro- 
duction of  very  valuable  plant  food  materials,  but  may  also  convert  evil  smelling 
organic  substances,  such  as  are  contained  in  sewage,  into  simpler  substances  which 
are  absolutely  free  from  odour  and  inoffensive,  a  fact  which  is  utilized  in  the 
disposal  of  sewage  in  towns,  the  sewage  being  collected  in  tanks  where  it  is 
exposed  to  the  action  of  various  fungi  and  bacteria. 


209 

they  usually  grow.  Again,  as  regards  the  stage  of  de- 
velopment, we  know  that  many  trees  which  are  easily 
killed  by  extremes^  of  temperature  as  seedlings  are 
resistant  when  older,  and  that,  for  example,  flowers  and  young 
leaves  may  be  readily  injured  while  resting  buds  are  very 
resistant.  The  quantity  of  moisture  also  present"  in  a  plant 
materially  affects  its  power  of  withstanding  extremes  of  tem- 
perature. Many  mosses  and  lichens  are  easily  killed  when 
moist,  but  are  extremely  resistant  when  dry.  The  more  water 
contained  in  plant  tissues  the  more  likely  are  they  to  be 
injured  by  frost. 

188.  Excessive    heat  most    com- Effects  of 
monly  injures  plants  in  one  of  the   following  ways  : —  Excessive 

.ti.C£ii» 

(1)  By  causing  the  drying  up  and  death  of  leaves,  twigs, 

branches,  or  even  entire  plants  and  trees,  owing  to 
the  abstraction  of  water  from  the  plants  themselves 
and  from  the  soil  in  which  they  grow.  The  roots 
are  thus  unable  to  obtain  sufficient  moisture  from 
the  soil  to  make  good  the  losses  occasioned  by 
increased  transpiration.  This  form  of  injury  is 
most  destructive  in  India  in  years  when  the  more 
or  less  complete  failure  of  the  monsoon  has  lowered 
the  level  of  all  springs  and  considerably  reduced 
the  amount  of  moisture  ordinarily  available  in  the 
soil. 

(2)  By  splitting  the  stems  of  trees.     Intense  heat,  drying 

up  the  outer  tissues  of  a  stem  and  thus  causing  them 
to  shrink,  gives  rise  to  splits  in  the  wood  very 
similar  to  frost  cracks.  Thin-barked  species,  such 
as  Phyllanthus  Emblica,  may  thus  bo  found  split 
to  the  centre,  especially  in  years  of  drought. 

(3)  By  directly  scorching    and  killing  the  cambium  and 

other  living  tissues,  thus  causing  cancerous  patches 
on  the  stems,  or  the  death  of  entire  trees.  This 
is  often  caused  by  the  sun's  rays  impinging  on 
thin-barked  stems,  especially  those  which  have 
grown  up  in  shade  and  then  become  exposed  by 
the  felling  of  the  surrounding  growth,  as  happens  in 
the  case  of  standards  in  a  coppice. 

189.  Excessive  cold  usually  results  Effects  of 
in  one  of  the  following  modes  of  injury :— 

(1)  The  direct  freezing  and  killing  of  parts  of  plants,  such 
as  the  leaves,  twigs,  etc.,  and  even  of  entire  plants 


210 

and  trees.  The  injury  or  death  is  in  this  case 
caused  by  some  change  in  the  protoplasm  brought 
about  by  the  low  temperature  in  a  way  which  is  not 
yet  understood,  and  which  varies  for  different  species. 
On  freezing,  ice  usually  forms  in  the  intercellular 
spaces,  water  being  withdrawn  from  the  cells.  This 
withdrawal  of  water  and  drying  of  the  cell  contents 
appears  to  be  the  cause  of  death  in  some  cases.  As  a 
rule  the  rapidity  or  otherwise  of  the  freezing  or  thaw- 
ing has  no  specially  injurious  effect,  the  injury  being 
due  to  the  low  temperature,  although  we  can 
generally  only  see  if  a  plant  is  dead  or  alive  after 
thawing.  (This  action  must  be  distinguished  from 
that  of  indirect  desiccation,  (2)  below,  in  which  a 
sudden  rise  of  the  air  temperature  may  be  very 
injurious,  by  increasing  transpiration  from  the  leaves 
while  the  roots  are  still  inactive  in  frozen  soil. )  The 
temperature  of  plants  may  be  very  considerably 
reduced  by  radiation  which  may  result  in  the  tem- 
perature of  the  plant  being  as  much  as  8°  C.  below 
that  of  the  air.  Radiation  is  reduced  by  fogs, 
clouds,  dust,  or  smoke,  and  the  kindling  of  smoky 
fires  is  often  efficacious  in  preventing  frost  damage, 
as  also  are  artificial  coverings  of  various  sorts. 
The  temperature  of  the  leaves,  twigs,  and  more 
delicate  serial  portions  of  plants  chiefly  depends 
on  the  temperature  of  the  surrounding  air,  and 
hence  the  damage  done  by  frost  depends  to  a 
great  extent  on  the  locality.  The  absence  of  air 
currents  is  one  of  the  most  important  factors  which 
affect  the  air  temperature  and  hence  low-lying 
enclosed  situations  where  there  is  little  or  no  cir- 
culation of  air  are  particularly  liable  to  frost  damage. 
Such  are  the  grassy  blanks  so  frequently  seen  in 
Sal  (Shorea  robusta)  forests,  in  which  the  circulation 
of  air  is  prevented  by  the  dense  wall  of  high  Sal 
forest  surrounding  them  on  all  sides  and  where  the 
cold  air  collects  and  stagnates.  In  such  places 
all  Sal  shoots  are  killed  down  year  after  year  by 
frost,  whereas  small  patches  of  Sal  forest  may  often 
be  seen  in  the  same  locality  untouched  by  frost, 
owing  to  the  open  cultivated  ground  on  all  sides 
oi  it  promoting  the  free  circulation  of  air  currents. 
In  these  frosty  localities  the  cold  air  collects  like 


211 

water  in  a  lake,  and  the  height  to  which  it  rises  is 
usually  clearly  marked  by  a  definite  line  above 
which  growth  is  uninjured.  This  was  well  seen  in 
the  Sal  forests  of  the  Dehra  Dun  during  the 
severe  winter  of  1904-05,  a  definite  line  being  clearly 
visible,  running  along  the  flanks  of  the  Siwaliks, 
below  which  the  leaves  on  every  tree  were  brown 
and  dead,  those  above  the  line  being  green  and 
uninjured.  The  presence  of  large  quanti- 
ties of  water  in  the  soil  may  considerably  reduce 
the  air  temperature,  as  may  also  a  heavy  growth 
of  grass  which  reduces  the  temperature  by 
radiating  heat,  transpiring  moisture,  interfering 
with  the  free  circulation  of  air,  and  with  the  access 
of  heat  to  the  soil.  Observations  have  shown 
that  the  temperature  on  an  area  covered  with 
grass  may  be  16°  F.  lower  than  that  of  a  similar 
area  with  no  vegetation.  The  effect  of  frost 
depends  of  course  largely  on  the  condition  of  the 
plant  itself.  Consequently  two  trees  of  the  same 
species  may  be  very  differently  affected  in  different 
localities.  One  growing  in  a  damp  warm  valley 
may  be  killed,  while  another,  in  a  more  exposed 
situation  at  a  higher  elevation,  may  escape  injury ; 
the  former  owing  its  susceptibility  chiefly  to  the 
longer  period  of  its  vegetative  activity,  the  larger 
quantity  of  water  in  its  tissues  and  to  the  latter 
being  less  completely  lignified  or  otherwise  matured. 
(2)  The  indirect  drying  up  and  killing  of  plants  or  parts  of 
them,  the  injury  being  caused  in  precisely  the  same 
manner  as  in  the  case  of  drought.  When  the  soil 
temperature  falls  below  a  certain  point  the  root 
hairs  are  no  longer  able  to  absorb  the  necessary 
supplies  of  water.  While  the  roots  are  thus  inactive, 
the  leaves  may  be  actively  transpiring  moisture 
under  the  influence  of  bright  sun-light  and  dry  air- 
currents.  The  leaves  and  branches  may  thus  become 
completely  dried  up  and  killed.  Trees  which  retain 
their  leaves  in  winter  are  particularly  liable  to  this 
form  of  damage  and  mahua  (Bassia  latifolia)  and 
achar  (Buchanania  latifolia)  may  frequently  be  seen 
injured  in  this  way.  This  mode  of  injury  may  often 
be  recognised  by  the  leaves  commencing  to  die 
back  from  the  tip,  the  tissues  at  the  base  of  the 


212 

leaf-blade  and  especially  near  the  midrib  being 
often  uninjured  and  green,  while  the  rest  of  the 
leaf  is  shrivelled  and  brown,  i.e.  just  those  portions 
escape  injury  which  can  first  intercept  the  water- 
supply  on  its  entrance  into  the  leaf  and  which  only 
allow  that  quantity  of  water  to  pass  on  to  the 
more  distant  cells  which  is  in  excess  of  their  own 
needs.  See  Plate  XX.  The  damage  can  often  be  well 
seen  in  a  nursery  after  a  frosty  night,  no  injury  to 
the  leaves  being  usually  noticeable  until  the  plants 
have  been  exposed  to  the  bright  sunlight  for  some 
hours.  It  is  obviously  important  that  trees  grow- 
ing at  high  altitudes,  where  the  leaves  are  often 
exposed  to  the  bright  sunlight  when  the  roots  are 
in  soil  still  frozen,  should  be  able  to  prevent  trans- 
piration as  far  as  possible.  We  thus  find  that  the 
high-level  Rhododendron  campamdatum  and  Quercus 
semecarpifolia  have  the  undersurface  of  their  leaves 
covered  with  dense  tomentum.  With  regard  to 
this  form  of  injury  it  must  be  pointed  out  that 
roots  situated  in  the  superficial  layers  of  soil  are 
more  subject  to  the  effects  of  sudden  changes  in 
temperature,  while  those  in  the  deeper  layers  of  soil 
are  chiefly  affected  by  prolonged  cold  temperature. 
Thus  in  some  cases  we  find  seedlings  suffer  more 
than  older  plants,  while  in  other  cases  the  reverse 
happens.  In  the  latter  case  the  superficial  roots, 
under  the  influence  of  a  few  hours  hot  sunshine, 
having  been  sufficiently  warmed  to  become  active, 
whereas  the  deeper  roots  are  still  in  frozen  soil. 
An  artificial  covering-  of  straw,  dead  leaves,  etc., 
on  the  soil  over  the  roots  of  a  plant  may  often 
prevent  injury  by  frost,  by  moderating  the  tem- 
perature of  the  surface  soil,  and  it  is  probable 
that  the  beneficial  action  of  watering  plants  is 
often  due  to  the  temperature  of  the  roots  being 
raised  sufficiently  to  become  active  thereby. 

(3)  The  splitting  of  stems  of  trees.  When  the  temper- 
ature is  sufficiently  low  ice  is  always  formed 
in  plants,  usually  in  the  intercellular  spaces,  but 
in  wood,  where  intercellular  spaces  are  usually 
absent,  in  the  lumina  of  the  wood  elements. 
As  these  spaces  contain  considerable  quantities  of 
air  besides  water,  the  water  finds  ample  space  for 


213 

expansion  on  being  converted  into  ice.  During  the 
formation  of  ice,  water  is  abstracted  from  the  cell 
walls,  and  this  results  in  the  drying  and  shrinkage  of 
the  wood,  just  as  happens  when  cut  timber  is  dried 
in  the  air.  During  severe  cold,  especially  if  the 
temperature  falls  very  rapidly,  the  outer  wood  layers 
shrink  more  rapidly  than  the  inner  warmer  layers, 
and  longitudinal  cracks  and  fissures  are  caused 
in  the  stem.  Such  cracks  may  become  occluded  in 
the  usual  way,  but  subsequent  frost  frequently 
re-opens  the  wounds,  and  if  this  is  repeated  several 
times  prominent  ridges,  or  frost-ribs,  result. 

(4)  The  formation  of   cankers.     Frost  cankers  are  usually 

found  at  the  base  of  a  young  branch  which  has 
been  killed  by  frost  down  to  the  main  stem.  The 
callus  forming  at  the  base  of  the  dead  shoot  is 
again  killed  by  subsequent  frosts,  and  with  recur- 
ring frosts  the  cancerous  area  spreads.  The  growth 
of  the  callus  is  also  interfered  with  by  the  pressure 
of  the  dead  tissues  under  which  it  forms,  and 
such  diseased  areas  usually  heal  slowly,  if  at  all, 
and  aiford  a  favourable  point  of  attack  for  wound 
fungi.  Cankers  due  to  frost  may  be  distinguished 
from  those  due  to  fungi  as  they  only  increase  in  size 
in  frosty  years. 

(5)  The  uprooting   of   seedlings.     When  the  soil  freezes, 

the  expansion  which  the  water  in  its  interstices 
undergoes,  on  being  converted  into  ice,  raises  the 
surface  of  the  soil  and  pushes  up  the  roots  above 
their  original  position.  When  the  ice  melts  with 
the  thaw  the  particles  of  soil  fall  away  from  the  roots 
which  become  exposed  and  the  plant  falls  over  and 
dies. 

(6)  The  mechanical  bending   and   breaking  of   stems  or 

branches,  owing  to  the  accumulation  of  snow  or  ice 
upon  them.  On  hill  sides  the  pressure  of  snow 
against  the  upper  side  of  young  stems  often  causes 
the  latter  to  curve  outwards  at  the  base,  and  this 
curve,  being  retained  as  the  plant  grows  older,  is 
often  visible  in  mature  stems.  This  bending  over 
of  young  stems  often  also  ruptures  the  tissues  on  the 
uphill  side  and  causes  wounds  which  may  give 
access  to  injurious  fungi. 


214 

(7)  The  more  or  less  complete  destruction  of  flowers,  leaves, 
twigs,  and  young  stems,  by  hail.  The  damage  done 
depends  largely  on  the  state  of  development  of  the 
plant.  While  mature  leaves  may  only  be  penetrated 
by  the  hail-stones,  giving  rise  to  so-called  "  shot- 
holes,"  the  injury  being  more  or  less  local,  tender 
young  leaves  may  be  torn  into  shreds  and  the  tree 
completely  defoliated.  Such  complete  defoliation 
results  in  the  loss  of  valuable  food  materials  con- 
tained in  the  young  leaves,  while  the  tissues  have 
to  be  depleted  of  their  food  reserves  for  the  for- 
mation of  fresh  foliage,  this  reduction  in  the  avail- 
able food  supply  resulting  in  loss  of  increment  and 
diminished  production  of  flowers  and  seed. 

of  190.  All  plants  may  be  killed  by 

sufficiently  strong  light,  and  the  fact  that  disease-produc- 
ing bacteria  may  be  killed  by  a  sufficiently  long  exposure 
to  sunlight  is  obviously  of  great  practical  importance.  In 
green  plants  the  chlorophyll  corpuscles  are  usually  more 
subject  to  injury  than  the  rest  of  the  protoplasm.  If  the  light 
is  too  intense  these  corpuscles  lose  their  power  of  assimila- 
tion and  may  become  permanently  bleached.  The  necessity 
of  protecting  the  chlorophyll  corpuscles  from  intense  light  is 
indicated  by  the  presence  of  colouring  matters  such  as  antho- 
cyanin,  to  which  the  red  or  purple  colour  of  many  young 
leaves  is  due,  and  which  acts  as  a  protective  screen  to  the 
chlorophyll,  by  the  general  absence  of  chlorophyll  in  the 
epidermis,[and  by  the  fact  that  the  chlorophyll  corpuscles  when 
exposed  to  strong  light  arrange  themselves  in  such  a  way  in  the 
cells  as  to  be  least  exposed  to  its  action. 

On  the  other  hand,  deficiency  of  light  may  be  no  less  injurious. 
In  the  absence  of  light,  green  chlorophyll  is  as  a  rule  not 
formed  and,  the  manufacture  of  food  materials  in  the  leaves 
being  impossible,  the  death  of  the  plant  by  starvation  ensues. 
We  know,  for  instance,  how  easily  so-called  ' '  light-demanding  ' : 
species  may  be  killed  by  shade.  Insufficient  light  causes  the 
diseased  condition  known  as  etiolation,  which  is  characterised 
by  the  development  of  abnormally  long  and  thin  internodes  and 
small,  or  very  thin,  yellowish  leaves,  with  unusually  watery 
tissues.  The  thickness  and  rigidity  of  the  cell  walls  is  also 
diminished,  and  the  laying  of  cereals  is  due  to  the  shading  of 
the  lower  portion  of  the  haulms  which  thus  become  too  weak 
to  support  the  weight  of  the  plants.  Unless  the  etiolated 


215 

condition  has  gone  too  far  the  plants  may  recover  with  the  access 
of  sufficient  light,  but,  owing  to  their  abnormally  thin  and  watery 
tissues,  they  are  particularly  liable  to  be  killed  or  damaged  by 
frost,  fungi,  and  other  injurious  influences. 

191.  Somewhat  similar  to  etiola-  E£Eect  ?f 
tion    is   the   condition    produced   by    an   excess   of   moisture 

in  a  plant.  This  is  chiefly  due  to  transpiration  not  being 
sufficiently  active  to  deal  with  the  quantity  of  water 
pumped  into  the  plant  by  the  roots.  The  tissues  become 
over-saturated  with  water,  food  materials  are  excessively 
diluted  and  can  only  be  transported  in  sufficient  quantity 
very  slowly  from  one  part  of  the  plant  to  another,  in  conse- 
quence of  which,  as  in  the  case  of  etiolation,  growth  in 
length  is  considerable,  but  the  watery  tissues  are  not  properly 
matured  and  are  very  liable  to  be  damaged  by  fungi,  insects, 
frost,  etc.  This  condition  is,  of  course,  distinguished  from 
etiolation  by  the  fact  that  green  chlorophyll  is  present  in  the 
leaves  and  that  the  disease  occurs  in  plants  which  have  access 
to  light.  Plants  with  an  abundance  of  water  available  for 
their  roots,  which  are  growing  in  a  saturated  atmosphere,  are 
liable  to  suffer  from  the  condition  here  described. 

192.  Winds,     besides     uprooting  Effect  of 

trees,   breaking    their  stems    and   branches   and   causing    the          ' 
development     of    stunted     and    misshapen    crowns     usually 
bending      away     from      the     prevailing       wind       direction, 
impoverish     the    soil   by  dispersing   dead   leaves   and  debris, 

thus  preventing  the  accumulation  of  humus  and  also  diminish 
the  available  supply  of  moisture  in  the  soil.  Winds  may  be 
also  exceedingly  destructive  to  plants  by  increasing  transpira- 
tion. Dry  winds  in  cold  frosty  weather  are  particularly 
injurious,  the  roots  in  the  cold  soil  being  then  unable  to  replace 
the  water  removed  by  transpiration,  the  plants  being  more  or 
less  injured,  if  not  killed,  by  desiccation. 

Observations  have  shown  that  the  velocity  of  a  wind  which, 
at  a  height  of  Ij  feet  above  the  ground,  was  only  22 '2  miles 
per  hour,  may  be  as  great  as  42*7  miles  per  hour,  at  a  height  of  51 
feet  above  the  ground.  Consequently  tall  plants  and  especially 
tree -growth  suffer  much  more  from  winds  than  shrubby  or 
herbaceous  growth.  It  should  be  noted  that  in  the  case  of  the 
Banana  the  tearing  of  the  large  leaves  by  the  wind  is  a  normal 
condition  leading  to  advantageous  aeration  of.  the  leaves. 
Winds  are  also  advantageous  in  the  way  of  distributing  pollen 
and  seeds  and  aiding  the  distribution  of  plants. 


216 


Presence  193.  The    presence   of  poisonous 

Substance  **  substances  in  the  air  may  cause  disaster  to  plants  just 
as  may  their  presence  in  the  soil.  More  than  4  per 
cent,  of  carbon  dioxide  in  the  air  is  injurious  to  many 
plants.  Sulphur  dioxide  and  fumes  of  hydrochloric  acid  are 
especially  harmful,  to  which  fact  is  to  be  attributed  the  injurious 
effect  of  an  excessive  amount  of  coal  smoke,  such  as  occurs  in 
many  towns,  and  of  the  fumes  from  iron-smelting  furnaces, 
alkali  and  other  chemical  works.  The  foliage  of  plants  thus 
injured  usually  turns  yellow,  and  the  plants  become  sickly  and 
may  eventually  die.  Trees  usually  suffer  more  than  herbaceous 
growth  and  forests  have  been  injured  at  a  distance  of  4|  miles 
from  the  place  of  origin  of  the  poisonous  fumes.  The  Plane 
(Platanus)  is  able  to  withstand  the  injurious  effect  of  coal  smoke 
and  grows  well  in  smoky  cities,  as  do  also  as  a  rule  maples, 
horse-chestnut  and  elms. 

Effect  of  194.  The      action      of     lightning 

Lightning.  in  causing  the  death  or  disease  of  plants  is  not  clearly  under- 
stood. In  some  cases  a  tree  struck  by  lightning  remains 
practically  uninjured  ;  a  narrow  strip  of  the  cortical 
tissues  is  separated  from  the  wood  and  the  wound  is  soon 
occluded.  In  other  cases,  where  the  external  signs  of  injury 
are  similar,  the  entire  tree  may  die.  Stems  of  trees  in  some 
cases  may  be  completely  barked  by  lightning,  in  other  cases  they 
may  be  shattered  into  fragments  and  occasionally  set  on  fire. 
The  injurious  effects  may  be  confined  to  a  single  tree,  or  the 
lightning,  passing  from  the  stem  originally  struck  to  others  near 
it,  may  damage  a  large  group  of  trees.  In  the  latter  case  the 
extensive  damage  appears  to  be  often  due  to  the  lightning 
passing  from  root  to  root  in  the  ground  owing  to  the  bad  con- 
ducting nature  of  the  soil.  In  such  a  case  the  injury  spreads 
from  a  certain  point  in  a  centrifugal  direction.  Trees  injured 
by  lightning  may  sometimes  remain  alive  for  4  or  5  years  and 
then  die. 

SECTION  V.  —  EFFECT  OF  FIRE  ON  PLANT-DEVELOPMENT. 

195.  The  injurious  action  of  fires 

consists  in  destroying  leaves,  flowers,  seeds,  twigs,  branches, 
seedlings  and  young  plants,  and  in  scorching,  and  more  or 
less  extensively  injuring,  the  roots,  cortex  and  cambium  of 
older  trees. 

Moreover,  the  soil  may  be  injuriously  affected  owing  to  the 
destruction  of  humus  and  to  a  decrease  in  the  amount  of  avail- 
able moisture.  Fires  may  thus  be  responsible  for  wounds  and 


Effect  of 


'217 

a  diminished  vitality,  which  render  trees  more  liable  to  injury 
by  fungi  and  insects,  for  loss  of  increment  and  for  interference 
with  reproduction.  It  has  been  noted  above  that  different 
species  vary  in  their  power  of  resisting  excessive  heat,  and 
while  some  plants  are  very  sensitive  to  damage  from  fires  others 
are  less  so.  The  effect  of  the  fire  on  any  particular  species  of 
plant  depends  mainly  on  the  intensity  of  heat  produced  by  the 
fire  and  on  the  state  of  development  of  the  plant  itself.  A  % 
fire  which  may  be  beneficial  shortly  after  the  fall  of  the  seed, 
by  hastening  germination,  would  be  very  destructive  after  ger- 
mination has  actually  commenced.  Seedlings  of  deciduous 
species  which  have  their  serial  parts  destroyed  by  fire 
in  the  resting  season  may  be  very  little  damaged,  while 
others  burnt  in  the  vegetative  season  may  be  seriously 
injured.  The  seeds  of  some  species  also  are  readily  destroyed 
by  fire,  while  those  of  others  are  protected  from  injury  by  hard 
or  corky  coverings,  etc.,  and  in  the  case  of  some  species  the  ger- 
mination of  the  seed  appears  to  be  aided  rather  than  hindered 
by  fire. 

196.  Finally,    it    must  never  be  Necessity  of 

forgotten  that  a  factor,    in  addition  to  any  direct  beneficial  j^jjjjf the 
or   injurious    action   which    it   has    on    the  development   of  Beneficial  or 
any  particular  plant,  may    also    exercise   a  by-no-means  less  injurious 
important      indirect     beneficial    or     injurious      action,      by  Eff?ct of 
affecting    the   development     of    correlated   organisms.     Thus68 
winds    may    be    directly     beneficial    in    aiding    fertilisation 
and  the  distribution  of  seeds  of  a  certain  species,  while  they  are 
indirectly  injurious  by  performing  the  same  services  for  an 
injurious  plant  competitor,  or  by  distributing  the  spores  of  an 
injurious    parasitic    fungus.     Again    fires    may  be    indirectly 
injurious  by  damaging  symbionts,  such  as  useful  soil  bacteria 
or  fungi,  and  indirectly  beneficial  by  destroying,  or  retarding 
the  development  of,  injurious  plant  competitors  or  parasites. 


218 


PART  VL-GEOGRAPHY. 

CHAPTER  L— FACTOKS  INFLUENCING  THE  DISTRIBU- 
TION OF   PLANTS. 

Factors  197.  The  principal  factors  affecting 

affecting       the  distribution  of  plants  will  first  be  shortly  considered  under 

Distribution.  ^  f0Howing  neads  ._ 

(1)  Water.  (5)  Air. 

(2)  Soil.  (6)  Existence  of  other  Plants. 

(3)  Temperature.  (7)  Existence  of  Animals. 

(4)  Light.  (8)  Fire. 

(9)   Action  of  Man. 

(l)  WATER.  198.  The  water  available  for  plants 

Available  depends  chiefly  on  the  rainfall.  In  deserts,  where  the  amount  of 
depends  moisture  available  for  plants  is  exceedingly  small,  owing  to  the 
chiefly  on  scanty  rainfall,  very  few  plants  are  able  to  survive  and  no  forests 
the  Rainfall.  exist.  In  India  "  really  thriving  forests  are  only  found  where  the 
rainfall  exceeds  40",  and  rich  luxuriant  vegetation  is  limited 
to  those  belts  which  have  a  much  higher  rainfall.  "  *  Owin<* 
to  the  low  temperature  prevailing  at  high  elevations,  mount 
ains  tend  to  condense  aqueous  vapour  and  thus  receive  more 
rain  than  the  adjoining  lowlands.  The  greatest  rainfall, 
however,  usually  occurs  at  a  comparatively  low  elevation  and 
at  higher  altitudes,  instead  of  periodic  heavy  falls  of  rain 
frequent  mists  and  light  rain  occur,  the  rarefied  air  having 
a  very  small  capacity  for  aqueous  vapour.  This  accounts  for 
the  luxuriant  growth  of  mosses  and  lichens  often  found  cover- 
ing the  stems  and  branches  of  trees  at  high  elevations.  The 
outer  ridges  of  a  range  of  hills,  also,  which  first  intercept  the 
currents  of  moisture -laden  air,  receive  more  rain  than  the  inner 
hills,  and  thus  Deoban,  for  instance,  on  the  outer  ridge  in  the 
North- West  Himalayas,  receives  a  greater  rainfall  than  either 
Mundali  or  Kathian,  while,  at  the  same  time,  Kalsi,  at  the  foot 
of  the  outer  hills  (elevation  1820'),  receives  a  greater  rainfall 

*  Cn  the  JJislribulion  of  Forests  in  India  by  Dietrich  Brandis,  p.  6. 


219 

than  Deobaii  at  the  top  of  the  outer  ridge    (elevation  9300'). 

The  amount  of   moisture   in  the   soil   may  be  largely  due  to  Also  on 

percolation  from  rivers,  canals,  or  lakes,  to  which  fact  is  largely  Percolation. 

due  the  characteristic  vegetation  of  river-banks  and  of  areas 

near  large  sheets  of  water. 

An  important  Indian  tree  which  requires  a  very  heavy 
rainfall  is  Ficus  elastica,  the  Caoutchouc  tree,  and  another 
tree  which  is  usually  confined  to  areas  near  rivers  or  to  swampy 
ground  is  Lagerstroemia  Flos-Reginae,  one  of  the  most  im- 
portant timber  trees  of  Assam  and  Burma.  On  the  other 
hand,  Prosopis  spicigera  is  a  tree  of  some  importance  which 
can  thrive  in  districts  with  a  very  small  rainfall. 

Under  this  head,  also,  must  be  considered    the  importance  Action  of 
of  water  as  an  agent  for   distributing  seeds.     The    sea  is   a  ^.a*e.r, m, . 

i  T        -i        •  ciii  rrn       Distributing 

great  obstacle  to  the  distribution  01  land  plants.  Ine  geed?. 
fruit  of  the  Cocoa  Nut  tree  is  provided  with  buoyant 
tissue  which  enables  it  to  float,  and  its  seeds  are  able  to 
germinate  after  prolonged  immersion  in  sea  water.  Many  other 
plants  have  similar  contrivances;  their  seeds  are  often 
distributed  great  distances  by  marine  currents,  and  they  are  thus 
able  to  establish  themselves  on  distant  shores. 

Water  is  largely  responsible  for  the  distribution  of  the 
spores  of  fungi,  while  rivers  and  streams  play  an  important 
part  in  distributing  the  seeds  of  many  plants.  The  seed  of 
Sissoo  (Dalbergia  Sissoo),  for  instance,  is  chiefly  distributed  by 
water,  and  that  of  Khair  (Acacia  Catechu)  frequently  so. 

Finally  must  be  noted  the  denuding  action  -of  water  which,  injurious 
by  washing  away  the  soil,  exposes  the  roots  of  plants,  thus  Denuding 
injuring  the  roots  more  or  less,  even  if  the  plants  are  not 
actually  up-rooted.  The  shrub  Rhabdia  lycioides,  common 
in  river-beds,  with  its  creeping,  rooting  branches,  is  well 
adapted  to  withstand  the  action  of  torrential  streams. 

199.   On    bare  rocks  an  insignifi-    (2)  SOIL. 

cant  vegetation,  consisting  chiefly  of   lichens,  alone  can  exist,  Depth  of 
and  on  very  shallow  soil,  capable  of  supporting  a  good  growth  Soil, 
of  grasses,  the  majority  of  forest  trees  cannot  thrive,  hence  the 
depth  of   the  soil  is  of  primary  importance. 

It  has  been  rioted  in  Part  V  above  that,  for  healthy  plant  Supply  of 
development,  the  essential  mineral  salts  must  be  present  in  the  Mineral 
soil,  and  these  must  exist  in  a  suitably  dilute  solution,  see  page  Salts> 
205.     As  the  solution  of  salts  in  the  soil  becomes  concentrated 
beyond  a  certain  limit,  plants  experience  increasing  difficulty  in 
obtaining  their  necessary  supply  of  water  from  the  soil.     Many 
substances,  also,  exercise  a  directly  poisonous   effect  on  plants 


220 


Supply  of 
Moisture. 


and  calcium  and  salts  of  iron  often  appear  to  be  injurious  in  this 
way.  Plants,  however,  vary  greatly  in  their  power  of  with- 
standing such  injurious  influences,  and  while  some  plants  can 
thrive  on  a  saline,  or  calcareous  soil,  others  cannot  exist  there, 
a  fact  which  is  largely  responsible  for  the  characteristic  vege- 
tation of  such  soils.  Among  important  Indian  trees,  Butea 
frondosa  is  remarkable  for  being  able  to  grow  on  soil  contain- 
ing large  quantities  of  salt. 

Supply  of  A  supply  of  °xygen  being  essential  for  the  healthy  action  of 

Oxygen.  the  roots  of  plants,  the  water  in  the  soil  must  be  well  aerated  and 
hence  not  stagnant.  Many  of  our  Indian  trees  are  exceedingly 
sensitive  in  this  respect  and  cannot  exist  unless  the  subsoil  is 
well  drained.  The  Deodar,  Pinus  longifolia,  Sal,  and  many 
others,  all  require  well- drained  soil.  On  the  other  hand,  many 
species  can  thrive  in  water-logged  localities,  e.g.  Butea  frondosa, 
Terminalia  Arjuna  and  others. 

The  physical  and  chemical  properties  of  the  soil  itself,  and 
of  the  subsoil,  influence  the  amount  of  available  moisture  to  a 
considerable  extent.  Finely  divided  soils,  such  as  clays,  have 
the  greatest  power  of  absorbing  water,  and  also,  being  very 
impermeable,  of  retaining  it.  Sand  on  the  other  hand,  being 
very  permeable,  can  retain  very  little  water.  Calcareous  soils 
again  have  very  little  power  of  absorbing  water,  while  those 
with  an  admixture  of  humus  are  capable  of  absorbing  and 
holding  large  quantities  of  water.  Among  Indian  trees  which 
thrive  best  on  clayey  soils  the  well-known  Sain,  or  Saj 
(Terminalia  tomeniosa),  may  be  mentioned.  Among  trees  often 
found  on  calcareous  soils  are  Khair  (Acacia  Catechu)  and  Satin 
Wood  (Chloroxylon  Swietenia).,  but  the  fact  that  both  these  trees 
are  commonly  also  found  on  non-calcareous  sand,  and  on  other 
soils  in  dry  forests,  indicates  that  the  important  factor  in  this 
case  is  the  small  amount  of  moisture  in  the  soil  and  not  the 
Characteris-  Presence  of  a  particular  chemical  constituent.  Two  very 
tics  of  characteristic  and  widely  distributed  types  of  Indian  soils  require 
Laterite  and  brief  mention  here,  viz.  (1)  Laterite  and  (2)  Regur,  generally 
known  as  Black,  or  Cotton,  Soil.  The  first  is  remarkable  for  its 
very  low  capacity  for  holding  water  and  for  containing  a  large 
percentage  of  iron.  The  Eng,  or  In  (Dipterocarpus  tuberculatus), 
a  valuable  timber  tree  of  Burma,  is  almost  exclusively  found 
on  laterite.  Many  other  species,  often  found  on  laterite, 
occur  just  as  frequently  also  on  other  dry  soils.  Regur  is 
generally  found  in  the  neighbourhood  of  trap  rocks  and  con- 
sists largely  of  clay  with  a  considerable  admixture  of  organic 


221 

substances.  It  is  characterized  by  its  great  adhesiveness 
when  wet  and  by  an  extraordinary  power  of  expansion  and 
contraction  under  the  influence  of  moisture  and  heat  respect- 
ively. In  the  dry  season  it  is  traversed  by  great  fissures, 
often  many  feet  in  depth.  As  a  rule  it  is  very  fertile,  especially 
for  herbaceous  plants  and  field  crops,  one  of  its  names  being 
due  to  its  suitability  for  the  cultivation  of  cotton.  The  Babul 
(Acacia  arabicd)  is  frequently  found  on  this  class  of  soil,  as 
also  is  Eutea  frondosa. 

Aspect,  slope  and  the  character  of  the  underlying  strata  Aspect, 
also  obviously  affect  the  loss  of  water  from  the  soil  through  Slope  and 
evaporation  and  drainage.  S^t?"18 

Finally,  there  is  the  important  factor  of  the  presence,  or  Effect  of 
absence,  of  vegetation  on  the  soil.  A  covering  of  forest  vegeta-  Vegetation, 
tion  protects  the  soil  from  insolation  and  the  effect  of  winds  and 
thus  diminishes  evaporation ;  it  leads  to  the  formation  of  humus 
which  absorbs  and  retains  water  well ;  the  force  of  rain  is  broken 
by  the  leaves,  twigs,  and  branches,  from  which  it  falls  gently, 
and  percolates  into  the  soil,  whereas  on  an  area  bare  of  vegeta- 
tion much  of  the  rain  water  runs  off  and  is  lost,  especially  on 
steep  slopes.  It  must  also  not  be  forgotten  that  the  rain  water 
contains  appreciable  quantities  of  useful  nitrogenous  compounds 
and  of  mineral  salts  which  are  essential  plant  food-materials,  so 
that,  by  intercepting  the  rain  water  alone,  humus  may  con- 
siderably increase  the  fertility  of  a  soil. 

The  character  of  the  soil  appears  to  be  the  principal  factor  Soil  the 
determining   the   distribution    of    the    Sal    tree,   this   species  Principal 
requiring     a     loose     well-drained     soil,     containing    a     con-  j-j^?*. 
siderable    proportion    of    humus.     It   cannot   thrive    on  the  mining  the 
heavy  soil  which  usually  overlies  trap    rocks  and,  in  Central  Distribution 
India,  the  extension  of  Sal  westwards    is  abruptly  checked of  Salt 
by  these  rocks,  Teak  forest  commencing  where  the  Sal  forest  ends. 

200.  The  effect  of  temperature  on  (3)  TEMPER- 

the  distribution  of  plants  has  been  recognised  from  very  early     ATURE- 
days,  and  it  was  found  that,  if  the   surface  of   the  globe  was  Vegetation 
divided  into  zones  of  the  same  mean  annual  temperature,  each  due  to  Tem- 
of    these    zones  was   characterized   by   possessing  particular  Perature- 
species  of  plants,  which  did  not  thrive  far  beyond  the  limits 
of  their  particular  zone.  These  zones  are  given  in  consecutive 
order  below,  that  with  the  highest  mean    temperature    first, 
and  that  with  the  lowest  last. 

I.   Tropical  Zone. — This  is  sometimes  sub- divided  into 
the  Equatorial,  Tropical  and  Sub-Tropical  Zones. 


222 

II.   Temperate  Zone, — This  is  also  usually  sub -divided 
into  the  Warm  Temperate    and    Cold    Temperate 
Zones. 
III.  Arctic  Zone. — This   is    sometimes  divided  into  the 

Sub -Arctic,  Arctic  and  Polar  Zones. 

The   Zones  in  the   northern   hemisphere  are  usually  termed 
Boreal  and  those  in  the  southern  hemisphere  Austral. 

The    Tropical  Zone  is  characterized  by  possessing  Palms, 
Bamboos,  Bananas,  Tree  Ferns,   Cycads  and  Screw 
.  j.  Pines  (Pandanus),  while  the  most    important   forest 

t  trees  in  this  Zone,  in  India,  belong  to  such  natural 

orders  as  Leguminosae,  Dipterocarpaceae,  Combre- 
taceae,  Urticaceae,  Meliaceae,  Verbenaceae,  Guttiferae 
and  others. 

In  the  Temperate  Zone  the  most  important  forest  trees 
•„  belong  to  the  Coniferae  and  Cupuliferae,  the  latter  in- 

cluding the  Oak,  Beech,  Chestnut,  Hazel,  Hornbeam, 
Birch,  and  Alder.  Other  characteristic  trees  are  the 
Poplar,  Willow,  Elm,  Walnut,  Maples,  and  Holly, 
while  among  shrubs,  Berberis,  Rhamnus,  and  Euony- 
mus,  are  common.  One  of  the  most  typical 
natural  orders  is  perhaps  that  of  the  Rosaceae. 
The  Arctic  Zone.  The  species  found  in  the  cold  tem- 
perate zone  also  extend  into  the  arctic  zone,  but,  in 
the  latter,  there  are  fewer  species,  and  the  vege- 
i  tation  is  especially  characterized  by  being  stunted. 

The  limit  of  tree  growth  is  usually  regarded  as  the 
natural  boundary  of  the  arctic  zone,  in  the  latter 
the  trees  becoming  shrubs,  while  shrubs  and  herbs 
are  much  reduced  in  height.  Plants  in  this  zone 
usually  have  small  leaves  and  well-developed  roots, 
while  large  and  brightly  coloured  flowers  are 
common. 

Although  the  above  suffices  to  illustrate  in  a  general  way  the 
of  Tempera-  importance  of  temperature  as  a  factor  influencing  the  distribu- 
ture.  t  ion  of  plants,  it  must  be  borne  in  mind  that  the  mean  annual 

temperature  is  very  much  less  important  than  the  extremes  of 
temperature  in  any  locality,  and  also  that  the  effect  of  a  certain 
temperature  on  a  particular  plant  depends  largely  on  the  season 
of  the  year  and  the  stage  of  development  of  the  plant.  Some 
plants  require  high  temperatures  throughout  the  year,  others 
uniformly  cold  temperatures,  while  yet  again  others  require  high 
temperatures  at  certain  seasons  and  low  temperatures  at  others. 
Thus  although  the  temperature  in  a  given  locality  may  be 


223 

sufficient  for  the  germination  of  the  seed  and  subsequent  growth 

of  a  particular  species,  it  may  not  be  sufficiently  high  to  enable 

the  plant  to  produce  fertile  seeds,  and  hence,  its  distribution  by 

seed  being  prevented,  its  chances  of  naturally,  establishing  itself 

are   small.     The  facts  that  luxuriant  vegetation  is  characteristic 

of  the  hottest  areas  in  the  tropics,  provided  there4s  sufficient 

moisture,  and  that  good  forests  exist  in  the  coldest  known 

spots  of  the  earth,  i.e.    in  parts   of  Siberia,    suffice  to  show 

that    other    factors    besides      temperature     are    responsible 

for  the  distribution  of  plants,  and  for  the  absence  of  trees  in 

parts  of  the  tropics  and    in  the  arctic    zone.     Since   a   similar  Low  Tem~ 

reduction  in  temperature  may  be  caused,  not  only  by  increas-  U^T0U!  to G 

ing  latitude,  but  also  by  increasing  elevation  above  sea-level,  increasing 

we  should  expect,   on   ascending  mountains,  to  find  charac-  Latitude  but 

teristic  zones,  or  regions,  of   vegetation  resembling  those  met  Elevation 

with  when  proceeding  from  the  equator  towards  the  poles,  and,  above  Sea 

to  a  great  extent,  this  is  the  case.     It  must,   however,  be  re-  l*vel. 

membered  that,   although  temperature  is  often  the  dominant 

factor  influencing  the  distribution  of  plants  in  both  cases,   on 

mountains  the  increasing  rarefaction  of  the  air  has  an  important 

effect  on    the  climate  and  therefore  on  the  vegetation,  which 

is  not  the  case  in  the  lowlands. 

Under  temperature,   also,   we  must   not    only  consider  the  importance 
temperature  of  the  atmosphere  but  also  that  of  the  soil,  seeing  peratureTof 
that,  when  the  soil  temperature  falls  below  a  certain  point,  the  the  Soil, 
roots  are  no  longer  able  to  absorb  the  necessary  supplies  of 
water  and  salts    from  the  soil.     If,  while  the   roots    are    thus 
inactive,  the  leaves  are  actively  transpiring  moisture  under  the 
action  of  bright  sunlight  and  dry  air,  the   plant   must  suffer 
from  desiccation,  as  has  already   been  pointed  out  in  Part  V 
above,   see   page  211.     Shallow-rooted,    low   plants,  such    as 
grasses,    on  the  whole  probably  suffer  less  in  this  respect  than 
do  trees. 

In  the  case  of  the  former,  as  the  temperature  rises  and 
transpiration  increases,  the  roots  situated  in  the  superficial  layers 
of  soil  are  soon  warmed  sufficiently  to  enable  them  to  make  good 
the  loss, while  the  tree  roots  situated  in  the  deeper  layers  of  cold 
soil  are  unable  to  do  so.  Moreover  low  plants,  with  their 
serial  portions  situated  in  the  calm,  damp,  lower  layers  of  the 
atmosphere,  do  not  transpire  so  actively  as  do  the  crowns  of 
trees  fully  exposed  to  the  action  of  drying  winds.  This  factor 
is  of  great  importance  in  high  mountains  where,  owing  to 
the  short  vegetative  season,  many  trees  find  it  necessary  to 
remain  in  leaf  throughout  the  year,  their  roots  being  in  winter 


224 

buried  in  frozen  soil  and  their  crowns  exposed  to  drying 
winds,  and  it  is  in  fact  mainly  responsible  for  the  absence  of 
trees  in  the  arctic  zone  and  at  high  elevations.  The  temper- 
ature of  the  soil  depends  partly  on  its  physical  properties, 
sand  possessing  a  far  greater  capacity  for  becoming  heated, 
for  instance,  than  clay ;  partly  on  its  depth,  the  deeper  layers 
only  being  affected  by  prolonged  winter  cold  or  summer  heat ; 
partly  on  the  aspect,  and  partly  also  on  the  presence  or  absence 
of  vegetation  and  the  character  of  such  vegetation. 

Temperature  is  the  principal  factor  determining  the  dis- 
tribution of  the  valuable  Hill  Forests  in  India  containing 
the  Deodar  and  other  important  Conifers  and  Oaks.  It 
also  appears  to  be  largely  responsible  for  the  distribution 
of  Teak,  no  good  natural  forests  of  this  species  occurring  north 
of  cold- weather  (i.e.,  November-February)  isotherm  65°.  An 
important  Indian  tree  which  appears  to  require  high  temper- 
atures throughout  the  year  is  the  Red  Sanders  (Pterocarpus 
santalinus). 

(4)  LIGHT.  201.  It  has    been  noted  in  Part  V 

above,  see  page  214,  that  light  may  injure  plants  by  being  (a) 
too  intense  or  (6)  too  feeble.  On  high  mountains  the  light  is 
more  intense  than  in  the  lowlands  and  is  of  importance  in 
regulating  the  distribution  of  plants  at  high  elevations-  On 
the  other  hand  the  too  feeble  light  makes  it  impossible  for 
many  plants,  e.  g.  grasses,  to  live  in  the  undergrowth  of  a 
dense  forest,  while  a  heavy  growth  of  grasses  may  in  turn,  by 
preventing  the  access  of  light,  be  responsible  for  the  death  of 
tree  seedlings.  Many  of  our  most  important  Indian  trees 
require  a  great  deal  of  light  and  cannot  grow  well  in  the  shade 
of  other  plants,  such  as  Teak,  Sissoo  (Dalbergia  Sissoo),  Khair 
(Acacia  Catechu),  Babul  (A.  arabica),  Blue  Pine  (Pinus  excelsa), 
and  the  Chir  (P.  longifolia),  while  others  can  stand  a  consider- 
able amount  of  shade,  e.  g.  Xylia  dolabriformis,  Sal  and  Sain. 

(5)  AIR.  202.  The  humidity  of  the  air  is  of 
Humidity  of  great  importance  for  plants,  inasmuch   as  (1)   the   supply  of 
the  Atmos-    wa^er  m  the  so{\  obtained  from  rain,  dew,  snow,  hail  and  from 

DilGrC' 

the  condensation  of  aqueous  vapour  depends  upon  it,  and  (2) 

it  regulates  the  degree  of  transpiration  from  the  aerial  parts  of 

plants  (transpiration  ceasing  in  saturated  air  and  increasing  in 

dry  air)  and  of  evaporation  from  the  soil.    Radiation  of  heat  is 

also  slower  in  moist  than  in  dry  air.    As  noted  in'  Part  V  above, 

Action  of      see  page  215,  winds  may  injure  plants   directly  by  uprooting, 

Winds.         breaking  and  rending  them,  and  also  indirectly  by  increasing 

transpiration,     The     second  mode  of    action  is  particularly 


225 

injurious  at  high  elevations  and  may  there  be  sufficient  to 
prevent  the  existence  of  trees.  Winds,  by  distributing  pollen, 
aid  in  the  fertilisation  of  flowers  and  production  of  seeds,  and 
also  help  in  the  distribution  of  seeds.  Grasses  are  thus  parti- 
cularly favoured  in  windy  localities  and  so  are  conifers  among 
trees.  The  fact  that  the  light,  large-winged  seed  of  the 
Blue  Pine  is  distributed  further  by  the  wind  than  is  the  seed  of 
the  Deodar  is  one  of  the  causes  which  handicap  the  latter 
in  its  struggle  for  existence  with  the  former.  Finally  the 
presence  of  poisonous  substances  in  the  air  may  make  it  pr=>sence  Of 
impossible  for  some  plants  to  survive  in  certain*  localities,  Poisonous 
e.g.  in  smoky  towns,  near  Iron  Works,  or  Chemical  Factories.  Substances. 

203.  The  Forester  in  India  knows  (6)  Existence 

from  experience  that,  if  he  wishes  to  create  forests  oi  some    of  of  other 
his  valuable    species,   he   must  in    the  first  place  establish  on  Plant9- 
the    area    a    growth  ^of .  other     plants    under    the    shelter  of^any1"5 
of  which  his    valuable  species   can  be  successfully  introduced,  Species  on 
whereas  in  the  open  their  existence,  at  all  events  when  young,  °ther  P^t3 
would  be  impossible.     This  question    of  dependence  on  other  Existence 
plants  may  often  decide  which  species  shall  survive  in  a  given 
locality.     On  hot   dry   aspects,   for  instance,    shelter   during 
youth  is  essential  for  Deodar,  which  in  such  situations  usually 
establishes  itself  naturally  under  the  shade  of  species  of  Indi- 
gofera,  Desmodium,  and    others.     The  seedlings  of  the  Blue 
Pine  on  the  other  hand  grow  readily  in  the  open  and,  in  this 
respect,  this  species  is  favoured  in  its  struggle  for  existence 
with  the  Deodar.     The  Sal  also  usually  establishes  itself  best 
in  open  grassy  areas  if  it  is  preceded  by  a  growth    of  shrubs 
of  inferior  species,  such  as  Mallotus  philippinensis,  and  others, 
under  the  shade  of  which  the  young  Sal  are  able  to  exist,  a 
phenomenon  which   may  be  often  seen  on  the  edges  of  grassy 
blanks  where  the  Sal  forest  is   gradually   extending   and   en- 
croaching on  the  grassland.     A  very  interesting  case  on  record 
in    this    connection    is    that   of   the   Bhinga    Forest   of    the 
Bahraich  Division  of  Oudh.     In   1875  this     was  practically 
a  ruined  Sal  forest,  very  open  with  no  Sal  regeneration ;  closure 
to  grazing  and  the  encouragement  of  a  growth  of  Mallotus 
philippinensis  and  other  shrubs  has  gradually  resulted   in  the 
establishment    of  a    growth    of    young    Sal.     An  even  more 
remarkable    case  is    afforded    by   the  Spruce  and    Silver    Fir 
forests  of  the  North-Western    Himalayas.     Here    the    natural 
reproduction   of  these   species  is  usually  conspicuous   by  its 
absence,  and  if  a  clearing    is    made,   instead   of  good   fir  re- 
generation taking  place,    the  ground  is  at  once  occupied  by 


brambles,  willows,  poplars,  and  other  plants.  After  such  plants 
have  occupied  the  ground  for  some  time,  reproduction  fre- 
quently reappears  beneath  their  shade  and  again  establishes 
itself  on  the  area,  the  inferior  species  thus  appearing  to  have 
prepared  the  way  for  the  Spruce  and  Silver  Fir  and  to  have 
made  their  existence  once  more  possible. 

Although  in  all  these  cases  it  is  obvious  that  certain  plants 
are  in  some  way  helped  by,  and  are  in  a  measure  dependent 
for  their  existence  on,  others,  the  exact  way  in  which  this  help  is 
given  is  by  no  means  always  clear. 

In  some  cases  mere  shade  from  intense  light  may  be  the 
beneficial  factor.  In  other  cases  the  shade  may  be  beneficial  on 
account  of  its  effect  on  the  temperature  of  the  plant  and  soil, 
and  on  the  quantity  of  available  moisture  in  the  soil  and  air. 
In  other  cases  again  the  benefit  may  consist  solely  in  the  fact 
that  the  shade  has  been  sufficient  to  prevent  the  growth  and 
development  of  injurious  plant  competitors,  as  would  appear 
to  be  often  the  case  when  shrubs  prepare  the  way  for  trees  by 
killing  out  grasses. 

It  has  been  often  thought  that  plants  during  life  con- 
tinually excrete  poisonous  waste  products,  the  accumulation 
of  which  in  the  soil  might  render  impossible  the  continued 
existence  either  of  the  same  species  or  of  other  plants  and 
that  therefore  some  plants  were  able  to  poison  the  plants  they 
displaced  and  that  others  even  made  their  own  existence  im- 
possible after  a  certain  period.  Although  in  the  case  of  the 
higher  plants  this  has  not  yet  been  proved,  still  it  does  un- 
doubtedly hold  good  in  the  case  of  many  fungi  and  bacteria.  In 
the  case  of  a  particular  tree  therefore  occupying  a  given  area  for 
a  long  period,  it  is  possible  that  the  accumulation  of  substances 
excreted  by  fungi  and  bacteria  in  the  soil  may  render  impos- 
sible the  existence  of  those  symbiotic  fungi  and  bacteria  which 
are  employed  in  breaking  down  the  humus,  in  which  case 
the  soil  may  become  so  impoverished  that  it  can  no  longer 
support  a  crop  of  the  tree  in  question,  see  also  the  remarks  on 
page  95  in  connection  with  the  rotation  of  crops  and  also  those 
on  page  207.  Similarly  the  existence  of  the  symbiotic  fungi 
which  form  the  so-called  mycorhizas  may  be  prevented  for 
a  time,  and  the  natural  regeneration  of  certain  tree  species 
consequently  rendered  impossible.  Any  factor  injuriously 
affecting  these  symbiotic  organisms,  such  as  insufficiency  of 
oxygen  or  water,  excess  of  water,  too  low  a  temperature,  absence 
of  sufficient  organic  food  materials,  and  so  on,  would  obviously 
produce  the  same  result, 


227 

Enough,  however,   has   now   been   said  to   show   that   the  (6) ExIS' 
presence  of  an  injurious  plant  parasite,  or  competitor,   or  the  0/OTHER 
absence  of  a  useful  symbiont,  may  suffice  to  prevent  the  exist-  PLANT. 
ence  of  a  given  plant  in  a  particular  locality,  and  hence  it  will  be 
seen  how  necessary  it  is  to  avoid  generalisations  and  to  study  in 
detail  the  case  of  each  individual  species  in  different  localities. 
The  valuable  Sandal  tree,  we  have  seen  above,  depends  to    a, 
large  extent  on  other  plants  for  its  supply  of  water  and  mineral 
salts,  and  the  existence  of  such  plants  is  undoubtedly  an  im- 
portant factor  influencing  the  distribution  of  this  tree. 

204.  In  India  examples  of  forests  (7)Exis- 

which  have  been  practically  ruined  by  excessive  grazing  are  not  ^ANIMAI 
uncommon.     In  such  areas  species  which  are  readily  eaten  by  injurious 
cattle  are  rapidly  exterminated,  all  young  growth  being  destroy- and  Useful 

j  IT.        iu   •         •  4-i,    Action  of 

ed  and  there  being,  in  consequence,  no  young  stems  to  replace  the  ^^^ 
mature  trees  which  must  sooner  or  later  die.  Goats  are  parti- 
cularly destructive,  and  in  areas  continually  browsed  by  them, 
the  forest  is  frequently  reduced  to  a  scrub  of  scattered  thorny 
shrubs  and  small  trees,  which  alone  are  able  to  survive.  Some 
of  our  important  trees  are  not  readily  eaten  by  goats,  and  this 
fact  has  considerably  affected  the  distribution  of  such  trees  as 
Pinus  excelsa,  Terminalia  tomentosa,  and  Butea  frondosa.  Xylia 
dolabriformis  is  also  an  important  tree  which  is,  as  a  rule,  little 
eaten  by  cattle.  On  the  other  hand,  insects  and  birds  are 
largely  instrumental  in  helping  plants  to  exist  in  certain  localities 
by  pollinating  their  flowers,  while  birds  and  other  animals  are 
often  the  most  important  agents  for  scattering  seeds.  Many 
species  of  plants  may  thus  have  their  areas  of  distribution  con- 
fined to  that  of  definite  insects,  birds,  or  other  animals.  Red 
Clover  is  a  well-known  instance ;  this  plant,  being  exclusively 
pollinated  by  humble  bees,  is  only  able  to  form  fertile  seed 
where  humble  bees  exist.  In  India  the  flowers  of  Butea 
frondosa  are  largely  pollinated  by  the  rose-coloured  starling 
(Pastor  roseus) ;  the  seeds  of  species  of  Loranthus,  Ficus,  Morus, 
and  of  the  Sandal  tree  are  chiefly  distributed  by  birds,  while 
jackals  are  believed  to  be  largely  responsible  for  distributing  the 
seed  of  species  of  Zizyphus.  The  seeds  of  Babul  also  are  said  to 
germinate  best  after  they  have  passed  through  goats.  From 
such  instances  it  is  obvious  that  the  presence  of  an  injurious 
parasitic,  or  the  absence  of  a  useful  symbiotic,  animal  may  suffice 
to  explain  the  absence  of  a  particular  plant  in  a  certain  locality* 

205.  If  grasslands  are  burnt  many  (8)  Sire. 
of  the  most  valuable  fodder  grasses,  which  are  delicate  annuals, 

are  killed  out  and  only  the  coarser  species  remain.    Fires  also, 


228 


(9)  ACTION 
OF  MAN. 


Action  of 
MaD* 


Necessity 
of  avoiding 

Inclusions 
as  to  the 
Factors 


Distribution 
of  any  Plant 


by  destroying  seed,  seedlings  and  young  growth,,  may  in  time 
reduce  a  good  forest  to  open  grassland.  Here,  however,  as 
in  the  case  of  all  other  factors  influencing  the  distribution 
of  plants,  when  studying  their  effect  on  a  particular  species  of 
plant,  we  must  consider  not  only  their  direct  effect  on  that  plant 
alone  but  also  their  effect  on  all  those  organisms  which  influ- 
ence the  development  of  the  plant  concerned.  Thus  pro- 
tecting a  forest  from  fire  with  the  object  of  favouring  the 
development  of  a  particular  species  may  have  very  unexpected 
results  ;  for  the  growth  of  injurious  competitors  may  be  so 
much  favoured  by  the  fire  protection  as  to  enable  them  to 
oust  the  species  it  was  desired  to  protect.  This  is  reported 
to  have  occurred  in  certain  fire  -protected  forests  in  Burma, 
where  the  teak  is  in  danger  of  being  ousted  by  some  species 
of  bamboos. 

206.  On   the  one    hand,  man   is 

responsible  for  the  absence  of  forest  trees  over  enormous  areas 
which  have  been  cleared   for  cultivation,    or  more    gradually 
devastated   by     reckless   fellings,  fires    and  excessive  grazing  ; 

•/  o  o  ~ 

on  the  other  hand,  he  helps  many  plants  to  extend  their  range 
of  distribution  and  to  establish  themselves  in  areas  they  could 
not  have  reached  without  his  intervention.  Lantana  aculeata, 
for  instance,  a  native  of  America,  has  been  established  in 
Ceylon  and  India.  Anona  squamosa,  also,  introduced  from 
the  West  Indies,  is  now  wild  in  many  parts  of  India,  and 
several  other  instances  might  be  given. 

207.  There  are  thus  a  large  number 

o|  f  actOrs  which  influence  the  distribution  of  plants,  and  hence 
we  must  guard  against  hastily  ascribing  to  any  one   factor  a 
result  which  may  be  due  to  the   combined  action  of  several, 
an(j  agamst  concluding  that  the  factor,  which   appears  most 
obvious,  is  primarily  responsible  for  the  distribution  of  any 
particular  plant.     Thus  the  fact  that   Salai  (Boswellia  serrata) 

.  jg  usuai]y  found  in  barren  places  where  the  soil  is  very  poor 
and  shallow  must  not  lead  us  to  conclude  that  this  tree  requires 
such  soil  for  its  development  and  cannot  thrive  on  any  other. 
As  a  matter  of  fact  Salai  will  grow  well  in  good  deep  soil,  but 
in  nature  it  is  ousted  from  such  localities  by  stronger  competing 
trees  which  there  find  suitable  conditions  for  their  development, 
the  result  being  that  Salai  is  driven  into  the  barren  spots  where 
it  can  exist,  but  where  the  majority  of  other  trees  cannot.  Thus 
the  fact  that  in  nature  Salai  is,  as  a  rule,  only  found  on  very  poor 
rocky  soil  is  due  not,  as  might  be  supposed,  to  Salai  's  preference 
for  such  places?  but  to  the  presence  of  injurious  competitors  in 


229 

others.  Similarly  the  fact  that  the  Cocoanut  tree  (Cocos  nucifera) 
in  nature  usually  occurs  on  the  saline  soil  of  the  seashore 
formerly  led  to  the  belief  that  the  tree  required  a  large  quantity 
of  salt  in  the  soil.  The  fact  that  this  tree  grows  well  in  gardens 
on  ordinary  soil,  however,  disproves  this,  and  we  are  driven  to  the 
conclusion  that  the  tree  can  only  thrive  m  nature  on  saline  soils 
owing  to  the  fact  that,  in  such  localities,  the  majority  of  other 
plants  cannot  exist,  and  that  it  is  driven  out  of  more  favour- 
able localities  by  stronger  competitors. 

208.  Another     important     point  power  Of 

concerning  plant  distribution  is  the  fact  that  most  plants  possess  Adaptation 
a  considerable  power  of  adapting  themselves  to  a  new  environ-  *°  ^itff1"611* 
ment,  to  changed  conditions  of  existence.     We  very  often  hear, 
for  instance,  of  plants  which  have  become  acclimatised.     Some 
species  possess  this  power  in  a  high  degree,  others  are  less  adapt- 
able.    It  must  be  remembered  that  a  plant  which  thus  adapts 
itself  to  new  conditions  undergoes  a  more  or  less  fundamental 
change,  acquires  as  it  were  a  somewhat  different  constitution, 
and   this  may,  or  may  not,  be  manifested  by  an  obvious  change 
in  its  outward  form,  or  habit.  We  have  already  seen,  in  Part  V 
above,  that  certain  individuals   of  one  and  the  same   fungus 
nourished  by  different  hosts   may  differ  essentially  from  one 
another  although  they  cannot  be  distinguished  by  any  visible 
character,  see  page  199.    This  undoubtedly  also  occurs  in  higher 
plants,  but  in  many  cases  also  remarkable  changes  are  notice- 
able, some  of  which  have  been  already  mentioned  under  varia- 
tions in   Part  IV  above,  see   pages  152 — 155.     In  some  cases 
plants   show   the   change  they    have  undergone  by   flowering 
and  leafing  at   different  periods  of  the  year,  or  by  becoming 
more  or  less  deciduous,  or  evergreen. 

209.  A  plant  which  is  adapted  to  Xerophytes. 

thrive  in  a  locality  where  very  little  water  is  available  is  termed  ^^  and 
a  xerophyte.  Such  plants  are  usually  provided  with  a  large  and  T£-(  pophytes. 
well  developed  root-system  and  with  devices  for  preventing 
excessive  loss  of  moisture  by  transpiration.  Thus  they  fre- 
quently have  small  leaves  (the  transpiring  surface  being  thus 
reduced),  which  are  coriaceous,  or  fleshy,  with  a  thick-walled 
epidermis,  and  often  provided  with  a  protecting  covering  of 
hairs,  etc.  They  also  often  possess  water-storing  tissue.  They 
are  often  provided  with  thorns,  or  spines,  and  sometimes  with 
mobile  leaflets  which  close  up  in  strong  sunlight.  Hygrophytes, 
on  the  other  hand,  are  plants  adapted  to  thrive  in  a  locality 
where  an  abundance  of  water  is  always  available.  They  are 
usually  provided  with  contrivances  for  accelerating  trans- 


230 

piration,  oversaturation  of  the  tissues  with  water  being  the  chief 

danger  to  be  guarded  against  in  this  case.     Tropophytes  are 

plants  which  at  one  season  of  the  year  are  xerophytes  and  at 

another  hygrophytes  ;    such  are   many  trees   in  the  deciduous 

forests  of  India  which  are  hygrophilous  in  the  rains  and  xero- 

philous  at  other  seasons.     A  plant  growing  in  a  soil  containing 

abundance  of  water  need  not  necessarily  be  a  hygrophyte  ;  the 

water  may  for  instance  consist  of  a  concentrated  solution  of 

salts,  in  which  case  the  roots  can  only  obtain   the    necessary 

supply  of  water  and  salts  slowly  and  with  difficulty.     A  xero- 

phiious  structure  is  thus   necessitated.     Such  a  structure  may 

be   due   to  any  factor  which  either    favours  transpiration  or 

interferes  with  the  ready  absoiption  of  water  from  the  soil  by 

the  roots. 


231 


CHAPTER  II.— PRINCIPAL  TYPES  OF  VEGETATION, 

210.  The  vegetation  of  the  earth  Types  of 
may  be  broadly  divided  into  three  great  types  : —  Vegetation. 

(1)  Woodland  in  which  woody  plants  predominate. 

(2)  Grassland  'in  which  grasses  predominate,  usually  in 

company  with  other  herbaceous  plants. 

(3)  Desert  where  the  climatic  conditions  render  luxuriant 

vegetation  of  any  kind  impossible  and  only  a  few 
plants  are  able  to  survive. 

Grassland  containing  isolated  trees  is  usually  called  savannah. 
Of  each  of  these  great  types  a  multitude  of  varieties  exist 
which  however  do  not  concern  us  at  present.  The  distribution  Factors 


of  the  above  types  is  influenced  chiefly  by  three  factors,  viz.,'11  . 

(1)  Moisture  which  depends  principally  on  the  amount  and  Distribution. 

distribution  of  the  rainfall,  (2)  Soil  and  (3)  the  Action  of  Man. 

These  will  be  shortly  considered  below  : 

211.  Typical  deserts  are    charac- (i)  Moisture- 

teristic  of   areas  with  a  very   small  rainfall.      Grasses,   being  Different 
usually  shallow-rooted  plants,  depend  mainly  on  the  moisture  Mature 
in  the  surface  soil.  Typical  grassland,  therefore,  can  thrive  if  the  produce 
rainfall  is  sufficient  to  keep    the  surface   soil   moist   in    the  Desert, 
vegetative  season.     The  majority  of  trees  on  the  other  hand  a  'or 

depend  mainly  on  the  water  in  the  subsoil  and  therefore 
require  a  rainfall  sufficiently  great  to  keep  the  subsoil  per- 
manently moist.  No  more  remarkable  instance  of  the 
dependence  of  vegetation  on  moisture  can  perhaps  be  given 
than  that  quoted  below  : — 

'  The  station  of  Jacobabad  is  a  striking  example  of  the  effect 
of  water  supply  in  that  climate.  It  was  founded  in  1844  by 
General  Jacob,  in  the  midst  of  a  barren,  treeless  desert.  A 
canal  was  led  to  it  from  the  Indus,  and  now  the  plain  is  a  dense 
forest  of  babool  and  other  trees,  upwards  of  sixty  feet  high, 
sheltering  the  houses  and  gardens  of  the  inhabitants.  A  ride 
of  a  few  miles  takes  you  into  the  desert  which  skirts  the  hills 
-of  Beloochistan,  a  level  plain  of  splendid,  fertile,  alluvial  soil, 
but  hard,  naked,  and  barren,  like  a  threshing  floor,  without 
shrub,  herb,  or  grass,  except  in  the  vicinity  of  the  canals, 
where  vegetation  is  rich  and  luxuriant. "  * 

It  has  been  noted  above  that  in  mountains  the  greatest 
rainfall   usually   occurs  at  a  comparatively  low  elevation  and 

*  Sir  Dietrich  Brandis  op.  cit.,  p.  11. 


232 


(2)  SoiL 

Existence 
of  Desert, 
Grassland 

depend 
the  Soil. 


Pioneer 


becomes  less  as  the  elevation  increases.  Hence  in  high  moun- 
tains. also,  the  effect  of  moisture  on  the  vegetation  may  often 
be  clearly  seen.  On  the  lower  slopes  with  a  heavy  rainfall 
we  find  forests,  at  higher  altitudes  grassland  and  finally,  if  the 
line  of  perpetual  snow  does  not  extend  to  the  grassland,  desert. 
As  already  noted  also  the  quantity  of  water  available  for  deep- 
rooted  plants  at  high  elevations  depends  not  only  on  the 
amount  of  the  rainfall  but  to  a  great  extent  also  on  the  tempera- 
ture of  the  soil. 

212.  A    study    of    the    gradual 

changes  in  the  soil  and  vegetation  which  may  pften  be  seen 
occurring  in  an  area  covered  with  naked  rock,  before 
or  luxuriant  vegetation  finally  establishes  itself  there,  demonstrates 
ver^  clearty  the  dependence  of  the  type  of  vegetation  on 
the  soil.  The  rocks  are  first  weathered  and  broken  up, 
slowly  it  may  be  but  none  the  less  surely,  by  the  mechanical 
action  of  heat  and  cold  and  the  chemical  action  of  the 
atmosphere  and  water.  On  the  decomposing  rocks  and 
rocky  particles  small  algae  and  lichens  usually  soon  establish 
themselves.  Lichens  aid  in  the  work  of  decomposition  by 
keeping  the  rock  surface  moist  and  by  the  action  of  their 
absorbing  hyphae.  The  decaying  remains  of  such  plants, 
mixing  with  the  particles  of  rock,  produce  a  little  soil  on 
which  mosses,  grasses,  and  other  higher  plants  can  establish 
themselves.  In  cracks  and  clefts  in  the  rock  also  where 
more  soil  accumulates  than  elsewhere,  woody  plants  soon 
appear  which  by  the  action  of  their  roots  play  an  important 
part  in  the  disintegration  of  the  rock  and  in  the  formation 
of  new  soil.  Many  trees  and  shrubs  which  in  themselves  may 
not  be  valuable  on  account  of  their  timber  or  other  products 
are  for  this  reason  of  the  greatest  value  to  the  Forester,  they 
being  the  pioneers  which  prepare  the  way  for  more  valuable 
species  and  make  the  existence  of  the  latter  possible.  Among 
such  pioneer  plants  should  be  noted  the  very  common  and  wide- 
ly distributed  Salai  (Boswellia  serrata)  and  Kulu  (Sterculw 
urens).  As  this  preparatory  work  proceeds  and  the  accu- 
mulation of  soil  increases,  more  and  more  trees  succeed  in  estab- 
lishing themselves,  and  eventually  woodland  arises  where  at 
first  there  was  only  barren  rock.  Thus  here,  as  the  naked  rock 
becomes  converted  at  first  into  shallow  and  later  into  deep 
soil,  we  can  trace  corresponding  changes  in  the  vegetation,  the 
desert  at  first  passing  into  grassland,  while  the  latter  eventual- 
ly gives  way  to  woodland.  It  is  important  for  the  Forester  to 
realize  the  part  played  in  nature  by  the  pioneer  plants  noted 


233 

above  for,  on  the  one  hand,  much  money  has  been  uselessly 
spent  in  parts  of  India  on  sowings  and  other  operations  with 
the  object  of  forestalling  the  slow  operations  of  nature  by 
quickly  afforesting  areas  occupied  by  pioneer  plants  with  more 
valuable  forest  trees  and,  on  the  other  hand,  cases  have  occur- 
red where  the  premature  wholesale  removal  of  sueh  pioneers 
has  retarded  the  natural  evolution  of  valuable  forest  by  many 
decades. 

From  what  has  been  said  above  we  see  that  a  virgin  forest, 
in  a  sense,  represents  the  final  product  of  centuries  of  work  on  pjant 
the  part  of  nature,  the  final  result  of  countless  struggles  in  Generatkms- 
which  only  those  plants  survived  and  for  a  time  occupied  the  or  Cr°Ps- 
ground  which  were  able  to  exist  under  the  conditions  of  the 
environment  prevailing  at  the  time.  At  first,  the  absence 
of  a  suitable  nutritive  substratum  rendered  impossible  the 
existence  of  the  majority  of  plants,  and  minute  alga3  and 
lichens  were  left  in  undisturbed  possession.  Later,  the 
accumulation  of  humus  soil  which  could  retain  a  sufficient 
quantity  of  moisture  allowed  grasses  and  herbaceous  plants 
to  obtain  a  footing,  and  they  ousted  the  algae  and  lichens. 
Finally,  a  still  deeper  soil  with  moist  subsoil  enabled 
woodland  to  vanquish  the  grassland.  The  first  plant  settlers 
have  to  struggle  against  the  very  unfavourable  conditions 
of  the  non-living  environment,  while  later  arrivals  have  to 
face  a  keen  fight  for  existence  against  other  competing 
plants.  The  scientific  Forester,  it  is  true,  has  to  interfere  with 
the  operations  of  nature  to  a  certain  extent,  since  it  is  his  busi- 
ness to  see  that  the  most  valuable  plants,  from  his  point  of 
view,  are  favoured  as  much  as  possible  in  the  struggle  for  exist- 
ence. At  the  same  time  this  power  can  only  be  usefully  ex- 
ercised within  very  definite  limits  and  the  interference  to  be 
useful  must  be  intelligent  and  exercised  so  far  as  possible  in  the 
light  of  a  good  knowledge  of  the  principal  factors  at  work  in 
each  case.  Man  cannot,  for  instance,  establish  flourishing 
woodland  in  a  locality  which  can  only  support  grassland  and 
by  an  unintelligent  interference  with  the  vegetation  on  an  area 
he  may  reduce  woodland  to  a  desert. 

213.  A  phenomenon  analogous  in  Succession 

many  respects  to   the  succession  of  different  types  of  vegeta-  communities 
tion  noted  above  may  often  be    seen  in   the  Sub-Himalayan  on  alluvial 
tract  of  India.     Here  on  the   banks  of  boulders,  shingle  and  deposits 
sand  brought  down  by  the  rivers  the  first  arrival  is  frequent-  Simaia  aa 
ly  Dalbergia  Sissoo  which  in  such  places  forms  gregarious  forests.  Tract. 
This  species  with  its  deep  and  far-reaching  ro  Dt  .system  helps  to 


234 

fix  and  protect  these  "  flood-plains,"  as  they  are  sometimes 
called,  against  the  action  of  subsequent  floods,  while  the  soil 
is  gradually  enriched  by  the  accumulation  of  organic  debris. 
Sissoo,  being  a  light  demander,  the  forests  of  this  species  thin 
out  and  become  open  with  increasing  age,  thus  leaving  room 
for  various  miscellaneous  species  which  soon  establish  them- 
selves and  eventually,  ousting  the  light- demanding  Sissoo, 
give  rise  to  a  mixed  forest  of  inferior  species.  As  time  goes  on 
however  and  the  soil  is  still  further  improved  by  the  accumula- 
tion of  humus  and  by  the  continued  shelter  from  climatic  in- 
fluences, the  struggle  for  existence  between  the  competing  trees 
becomes  more  severe,  more  trees  finding  on  it  congenial  condi- 
tions for  their  development,  and  cases  may  often  be  seen  on  the 
older  alluvial  deposits  where  the  Sal,  partly  owing  to  its  capa- 
city of  withstanding  the  injurious  effects  of  a  considerable 
degree  of  shade,  has  been  able  not  only  to  obtain  a  footing  in 
the  mixed  forest  but  to  entirely  oust  other  miscellaneous  species 
and  to  form  pure  forests  of  its  own. 

Action  of  214.  From  these  two  instances  the 

Man  in        important    part  played  by    plants  themselves  in  forming  soil 

Ordinary   °  an<^  *n  a^ering  its  physical  and  chemical  properties  is  clearly 

Course  of      recognized,  and  we  see  that   some   plants    are   able   to   create 

Natural        ^ne  conditions    necessary    for    the   existence  of  others ;   this 

necessarily    Par^  °^  *ne   subject,   however,   has  been  more  fully  discussed 

Restricted,    in  Chapter  I  above  under  Existence  of  other  Plants,  see  pages  225, 

226.    Cases  such  as  these  also  show  us  that  in  nature  a  particular 

type  of  vegetation  is  not  always  able  to  maintain  possession  of 

an  area  for  an  indefinite  period,    and  that  an   area  carrying  a 

good  forest  of  a  particular  species  may  not  necessarily  be  able, 

in  the  natural  course  of  events,  after  the  removal  of  that  crop, 

to  at  once  produce  a  second  crop  of  the  same  species.     Further 

we  must  remember,  as   already    stated   above,   that  although 

man  can   to   a  certain    extent   interfere    with    and   alter  the 

process  of  natural   development  yet  his  power  in  this  respect 

is  limited. 

(3)  Action  of  215.  Several  cases    are  on  record 

Man.  in   India   in   which  man,   by  destroying  the  forest  growth  on 

orSesert!  hill-sides,  has  reduced  woodland  to  desert,  and  the  Forester  is 
Grassland,  often  now  employed  in  the  difficult  task  of  inducing  vege- 
and  Forest  tation  to  re-establish  itself  on  the  barren,  rocky  slopes  and 
depends  on  °^  transforming  the  desert  once  more  into  woodland.  In  many 
the  Action  of  cases,  however,  the  destructive  influence  of  man  has  not  pro- 
Ma11-  ceeded  so  far,  and  his  action  has  resulted  in  establishing 
grassland  where  flourishing  forests  once  existed.  Throughout 


235 

India  such  instances  are  extremely  common,  the  forest  having 
been  first  cleared  for  cultivation  and  grassland  now  occupying 
the  abandoned  fields.  It  has  been  well  said  that :  "  woodland 
and  grassland  stand  opposed  to  one  another  like  two  equally 
powerful  but  hostile  nations,  which  in  the  course  of  time  have  re- 
peatedly fought  against  one  another  for  the  dominion  over  the 
soil,"*  and  nowhere  does  the  Forester  perhaps  realise  the  truth 
of  this  more  strongly  than  in  India.  The  seedlings  of  many  of 
our  most  valuable  tree  species,  in  the  first  year  or  two  of  their 
existence,  may  only  attain  a  height  of  a  few  inches  above  the 
ground  and  have  no  chance  of  surviving  in  the  struggle  for 
existence  in  a  dense  mass  of  grasses  which  effectually  shut  out 
the  necessary  light. 

216.  In  many  parts  of  India  the  Conversion 

afforesting  of  these  grass  lands  is  one  of  the  most  difficult  pro-  of  Grassland 
blems  which  the  Forester  has  to  solve.  The  difficulty  of  the 1U 
task,  moreover,  is  often  increased  by  the  fact  that  these 
clearings  were  frequently  made  in  the  middle  of  dense  forests. 
Tall  tree-growth  therefore,  in  such  cases,  surrounds  the  clearings 
like  a  wall  which,  by  interfering  with  the  free  circulation  of 
air,  is  largely  responsible  for  the  grassy  area  becoming  what 
is  known  as  a  frost-hole,  where  only  the  most  frost-hardy 
species  can  exist.  In  many  cases  it  has  been  noticed  that 
if  such  grasslands  are  protected  from  fire  and  grazing  they 
become  in  time  naturally  reconverted  into  woodland.  At 
first  small  grasses  form  a  dense  mantle  completely  covering 
the  ground  and  hiding  it  from  view ;  the  grass-crop  then 
becomes  gradually  thinner,  and  the  small  species  are  replaced 
by  taller,  coarser  grasses,  under  the  shade  of  which  patches  of 
unoccupied  soil  are  visible.  Between  these  grasses  a  growth  of 
.shrubs  then  creeps  in,  the  shade  of  which  while  sufficing  to  kill 
off  the  grasses  still  allows  the  seedlings  of  many  tree  species  to 
establish  themselves.  Finally  the  thinning  out  of  the  shrub 
growth  allows  these  seedlings  to  develop,  and  the  area  becomes 
once  more  covered  with  forest.  That  this  process  has  occurred 
and  is  still  in  operation  in  many  of  our  fire-protected  forests 
there  can  be  no  doubt,  but  at  the  same  time  the  process  is  often 
exceedingly  slow.  Various  plans  have  been  resorted  to  in  differ- 
ent parts  of  India  with  the  object  of  hastening  the  naturally 
slow  progress  of  events.  In  some  cases  the  seeds  of  tree  species 
are  sown  with  low-growing  field  crops,  the  injurious  competi- 
tive action  of  which  is  less  injurious  than  that  of  the  wild 

*  Plant  Geography  by  Dr.  A.  F.  W.  Schimper,  Eng.  Edn.,  1903,  p.  162. 


236 

grasses,  while  the  cultivation  prevents  the  development  of  such 
grasses.  In  other  cases  grazing  has  been  allowed  with  the 
object  of  keeping  down  the  grass-growth  until  the  tree-seedlings 
have  become  established,  and  elsewhere,  quick-growing,  accom- 
modating tree-species  have  been  introduced  artificially  into  the 
grassland  in  the  hope  that  they  will  oust  the  grasses.  Providing 
the  soil  is  suitable,  the  greater  the  rainfall,  the  more  quickly 
as  a  rule  does  woodland  reassert  itself  in  India  on  grasslands 
from  which  the  forest  has  been  cleared,  and  vice  versa. 
.  217.  Root-suckers  which  are  capa- 

Roo  "suckers  ^e  of  attaining  a  height  of  several '  feet  in  one  year  are 
Favoured  obviously  more  able  to  hold  their  own  in  such  grasslands  than 
in  the  are  minute  seedlings,  and  species  which  reproduce  them- 

with^  6        selves  readily  by  root-suckers  are  thus  often  able  to  successfully 
Grasses.        conquer  the  grasses  in  such  areas.     Evidence   of  this  is  often 
seen   in    India    in   the   practically   pure   forests  of   Ougeinia 
dalbergioides  and  Diospyros  tomentosa  which  now  occupy  the 
sites  of  old  abandoned  fields. 


237 

CHAPTER    III.— PRINCIPAL   TYPES  OF  INDIAN 
FORESTS. 

218.  For   the  purposes  of  a  brief 

and  general   description  the  forests   of  India  may  be  divided 
broadly  into  the  following  types  :— 

(1)  Arid-country  Forests. 

(2)  Deciduous  Forests. 

(3)  Evergreen  Forests. 

(4)  Hill  Forests. 

(5)  Tidal  or  Littoral  Forests. 

It  must  be  remembered  that  as  each  type  of  forest  often  passes 
very  gradually  into  a  different  type,  the  boundary  line  between 
the  various  types  cannot  always  be  clearly  denned;  moreover 
differences  in  the  soil,  local  conditions  affecting  the  amount  of 
moisture  available,  and    other  factors  may  entirely  alter  the 
type  of  forest  over  small  areas,  which  cannot  be  taken  into  con- 
sideration in  a  broad  and  general  account.     The  approximate 
distribution,  however,  of  these  types  is  shown  in  Plate  XXII 
areas  bearing  no  forest  not  being  separately  shown.     In  each  of 
these  main  types  also  numerous  sub -types  of  course  exist  which 
it  is  impossible  to  note  in  detail  here.     The  principal  factor  Factors 
responsible  for  the  distribution  of  types  1  to  3  is  the  amount  of  responsible 
available  moisture  as  measured  approximately  by  the  annual  Distribution, 
rainfall,  which  is  clearly  shown  by  a  comparison  of  Plates  XXI 
and  XXII ;  type  4  owes  its  existence  mainly  to  low  temperature 
caused  by  the   elevation   above  sea-level,   while   type  5  is  due 
mainly  to  the  large  quantities  of  salts  dissolved  in  the  sea-water 
and  to  the  action  of  the  tides. 

219.  The  principal  tract  of  country  (i)  Arid- 

which,  with  reference  to  its   rainfall,  may  be  called  arid,  the  country 
annual  rainfall  being  less  than  20  inches,  occurs  in  the  North-  Forests- 
Western  corner  of  India,  including  Sindh,  the  southern  portion 

of  the  Panjab,  and  a  large  part  of  Rajputana.  A  large 
portion  of  this  area  is  occupied  by  desert,  the  principal 
woodlands  being  the  so-called  rakhs  of  the  Panjab  in  the  north 
and  the  belts  of  the  riverside  forest  along  the  Indus  and  its  main 
tributaries.  The  rakhs  occupy  the  high  ground  between  the 
rivers  and  are  poor,  scanty  woods  of  small  trees  and  shrubs,  the 
most  characteristic  species  being  Prosopis  spicigera  and  species 
of  Salvadora  and  Capparis.  Owing  to  the  great  depth  of  the 
subterranean  water  the  species  occurriDg  in  these  rakhs  are 
often  characterised  by  possessing  roots  of  immense  length  and  a 
taproot  of  Prosopis  measuring  86  feet  in  length  has  been  exhibit- 


238 

ed.     The  riverside  forests  occupy  the  alluvial    soil  watered  by 
the  rivers,  either  by  percolation,  or  by  the  annual  floods,   and: 
they  owe  their  existence  to  the  supply  of  moisture  thus   obtained. 
The   principal   species   are     Babul    (Acacia   arabica),   Populus 
euphratica  and  species    of  Tamarix.     Both    the  babul  and  the 
poplar    occur    more    or    less    pure     over    considerable    areas 
and  at  other  times  as  standards  over  an  underwood  of  Tamarix. 
These  forests  are  often  washed  away  as    the  river   changes  its 
course,  and  the  fresh  alluvial  deposits  thrown  up  by   the  river 
are  quickly  covered  with  a  dense  growth  of    seedlings.     This 
arid  region  occupies  the  extreme  eastern  end  of  the  great  belt 
of  desert  which  stretches  from  Northern  Africa,  through  Arabia, 
South   Persia   and   Baluchistan,  to    beyond   the    Indus.     The 
comparatively   small  number  of  species   occurring  in  it  is  a 
noticeable  feature,  and  it  is  also  remarkable  that  some  of  them 
occur  both  in    the    African  and   Indian   area,    e.g.    Salvadcra 
persica  and  Capparis  spinosa.     The  forests  of  this  arid   Indian 
region  become  richer  in   species   towards  the  north  and   east, 
passing  then  gradually  into  deciduous  forests.    Towards  the  east 
Ancgeissus  pendula  is  a  common  and  characteristic  species,  it 
being  the  chief  forest  tree  in  Meywar  and  Merwara. 

220.  A  reference  to  the  map  Plate 
XXII  will   show  the  enormous    area   which  is  included    in 

Ous  j?  orcst-s.  .  r»      i          i      •  i  t*  i"i  •  i 

the  region  01  the  deciduous  forests  which,  with  reference 
to  its  rainfall,  may  be  called  the  moist  Indian  Eegion,  the 
annual  rainfall  ranging  from  20  to  70  inches.  These  are 
commercially  the  most  valuable  forests  of  India,  including  as 
they  do  the  greater  part  of  the  Teak  and  Sal  forests,  besides 
other  valuable  species  such  as  the  Sandal,  Ked  Sanders,  Sissoo 
and  many  others.  In  the  drier  regions  this  type  of  forest  merges 
into  arid  forest  and  in  moist  localities  into  evergreen  forest,  but 
on  the  whole  the  growth,  while  better  than  that  in  the  arid 
forests,  is  not  so  luxuriant,  and  the  trees  do  not  attain  such  large 
dimensions  as  in  the  evergreen  forests.  In  the  deciduous 
forests  the  great  majority  of  the  trees  are  deciduous  in  the  dry 
season.  While  a  large  number  of  species  are  found  almost 
universally  distributed  in  this  type  of  forest,  others  are  confined 
more  or  less  to  local  areas.  Among  the  most  widely  distributed 
species  are  the  following  : — 

Terminalia  iomentosa  (The  Sain  or  Saj,  usually  on  clayey 
soil). 

Terminalia  belerica. 

Terminalia  Chebula. 

Lagerstrcemia  parviflora. 


239  .  .   / 

Butea  frondusa  (Palas). 

Bombax  malabaricum. 

Buchanania  latifolia. 

Adina  cordifolia. 

Stephegyne  parvifolia. 

Mqle  Marmelos. 

Cassia  Fistula. 

Acacia  Catechu  (The  Khair,  often  gregarious  on  '  flood- 
plains'  and  scattered  in  dry  forest). 

Albizzia  odoratissima. 

Bauhinia  racemosa. 

Bauhinia  variegata. 

Erythrina  suberosa  (in  dry  forests). 

Eugenia  Jambolana  (in  moist  places  and  along  river 
banks). 

Ficus  glomerata  (in  moist  places  and  along  river 
banks). 

Ficus  hispida  (common  in  shady  moist  places). 

Ficus  infectoria. 

Flacourtia  Ramontchi  (on  dry  hills). 

Gardenia  turgida  (in  dry,  rocky  places). 

Grewia  asiatica. 

Holarrhena  antidysenterica. 

Hymenodictyon  excelsum  (in  dry  forests). 

Kydia  calycina. 

Phyllanthus  Emblica. 

Schleichera  trijuga. 

Schrebera  swietenioides. 

Stereospermum  suaveolens. 

Garuga  pinnata. 

Zizyphus  Jujuba. 

Zizyphus  CEnoplia. 

Zizyphus  rugosa. 

Boswellia  serrata  (not  in  Burma)  )  i      T.-II 

StenuKa  urens.  '  )  on  dry  rocky  bulls. 

Odina  Wodier  (  in  dry  forests). 
Careya  arborea. 
Bassia  latifolia  (the  mahua). 
Dendrocalamus  strictus. 
Phoenix  humilis. 

Anogeissus  latifolia  (not  in  Burma), 
imong  climbers  the  commonest  are — 
Butea  superba. 
Bauhinia  Vahlii. 


240 

Acacia  pennata. 
Acacia  caesia. 
Cryptolepis  Buchanani. 
Vitis  sp. 

Parasitic  Loranthaceae  are  also  very  common  in  these 
forests.  The  limits  of  the  distribution  of  some  of  the  most  im- 
portant species  of  these  forests,  i.e.  of  Sal,  Teak,  Sandal  and 
Red  Sanders,  are  shown  in  Plate  XXIII. 

The  Sal  is  strongly  gregarious,  and  is  by  far  the  most 
numerous  tree  in  the  forests  in  which  it  occurs.  In  the  Sub- 
Himalayan  tract  the  Sal  forests  usually  occupy  the  higher 
tracts  of  old  alluvium  lying  between  the  rivers,  while  the  more 
recent  alluvium,  consisting  of  beds  of  shingle  and  sand,  in  and 
near  the  river  beds,  is  often  covered  with  gregarious  forests  of 
Sissoo  and  Khair. 

In  damp  places  in  the  Sal  forests,  the  Indian  Red  Cedar 
(Cedrela  Toona),  Trewia  nudiflora,  and  Pterospermum  aceri- 
folium  are  common. 

The  Teak,  unlike  the  Sal,  usually  occurs  in  mixed  forests,) 
where  it  frequently  forms  only  a  small  proportion  of  the  stock. 
Although  the  Sal  and  Teak  areas  meet  in  Central  India  and  to  a 
certain  extent  overlap,  these  trees  are  not  found  together.  An 
isolated  patch  of  Sal  for  instance  occurs  inside  the  Teak  area,  at 
Pachmarhi,  in  the  Central  Provinces.  Teak  is  occasionally 
found  practically  pure  on  alluvial  ground,  but  its  best  growth 
is  obtained  on  well  drained  slopes.  A  large  number  of  species 
found  in  the  Indian  Teak  area  are  also  found  in  the  Teak  forests 
of  Burma  and,  besides  those  already  mentioned,  the  Ironwood 
of  Pegu  and  Arracan  (Xylia  dolabriformis)  should  be  noted.  On 
the  other  hand  several  species  found  in  the  Indian  area  do  not 
extend  to  Burma  and  vice  versa.  In  the  Indian  area  the  follow- 
ing are  important  trees  : — 

Pterocarpus  Marsupium  (Bijasal). 
Pterocarpus  santalinus  (Red  Sanders). 
Hardwickia  binata  (Anjan). 

Dalbergia  latifolia  (Shisham,  or  Bombay  Black  Wood). 
Chloroxylon  Swietenia  (Satin  Wood). 
Ougeinia  dalbergioides  (Sandan,  or  Tinas). 
Santalum  album  (Sandal). 

The  Burman  Teak  forests  on  the  other  hand  contain  such 
species  as — 

Dillenia  parviflora. 
Homaiium  tomentosum. 


241 

Cassia  siamect. 
Pterocarpus  macfocarpus. 

Peculiar  species  of  Sterculia. 

And  above  all  are  characterised  by  possessing  many 
different  bamboos  such  as — 

Bambusa  Tulda. 

Bambusa  polymorpha. 

Cephalostachyum  pergracile. 

Dendrocalamus  longispathus. 

Oxytenanthera  albociliata. 

In  the  Burman  deciduous  forest  area  also  are  found  the  so- 
called  Eng,  or  In  forests,  usually  on  formations  of  latente.  Dip- 
terocarpus  tuberculatus  is  here  the  most  characteristic  tree  with 
which  occur  the  following  : — 

Pentacme  suavis, 
Shorea  obtusa, 
Dillenia  pulcherrima, 
Dalbergia  cultrata, 

and  others.  The  well-known  Andaman  Padauk  tree,  Ptero- 
carpus dalbergioides,  is  found  in  the  Andaman  Islands,  usually 
in  deciduous  forests. 

221.  These  are  found,  as  will  be  (3)  Evei" 
seen  from  the  maps,  in  the  wet  region  which  has  an  annual 
rainfall  exceeding  70  inches.  They  are  in  their  characteristics 
the  most  essentially  ' '  tropical ' '  of  our  Indian  forests.  They 
are  especially  remarkable  on  account  of  the  large  number  of 
evergreen  species  which  they  contain,  their  luxuriant  vegeta- 
tion, and  the  enormous  dimensions  attained  by  some  of 
their  trees.  A  rich  growth  of  giant  climbers  (lianes)  and  of 
epiphytes  usually  covers  the  stems  of  the  trees,  and  the  under- 
growth is  frequently  so  dense  as  to  be  practically  impenetrable. 
The  number  of  species  is  very  much  greater  than  in  the 
deciduous  forests.  The  general  characteristics  of  this  class  of 
forest  are  excellently  described  in  the  following  passage  : — 

"  In  a  tropical  evergreen  forest,  growing  under  favourable 
conditions,  we  find  four  storeys  of  vegetation.  Immediately 
covering  the  soil  are  seedlings  mixed  with  shrubs  and  herbaceous 
species  and  in  the  next  zone,  or  storey,  small,  or  medium-sized 
evergreen  trees  50 — 75  feet  high.  The  top  canopy  of  great  ever- 
greens, often  150  feet  above  the  ground,  is  crowned  by  giant, 
sometimes  deciduous,  trees,  of  which  Tetrameles  nudiftora  is  one 
of  the  most  common  and  remarkable.  The  tree  stems  are  in 
many  instances  covered  with  epiphytic  Utricularias,  Orchids, 


242 

Aroids,  and  Ferns.  In  North  Kanara  where  the  evergreen 
tropical  Kans  (  a  local  name  for  evergreen  forests)  are  contiguous 
to,  or  surrounded  by,  mixed  deciduous  forest  the  divergence 
between  the  classes  of  vegetation  is  very  striking.  There  is 
considerable  physical  relief  in  passing  abruptly  from  the  strong 
glaring  sunlight  of  the  open  deciduous  jungle  in  the  hot  season 
to  the  cooler  atmosphere  and  deep,  somewhat  gloomy,  shade 
of  the  lofty  evergreens.  The  bewildering  diversity,  height  and 
size  of  the  trees,  the  universal  green  and  general  absence  of 
colour,  the  great  climbers  with  fantastic  shaped  stems,  the 
epiphytic  orchids,  aroids,  and  ferns,  the  general  stillness  and 
apparent  absence  of  animal  life,  appeal  to  the  naturalist,  who 
is  satisfied  that  here  at  least  the  action  of  man  has  not  affected 
and  changed  the  original  flora  of  these  truly  primeval  forests. 
The  principal  forces  of  nature  are  in  constant  action,  and  there 
is  no  annual  period  of  rest,  corresponding  to  the  winter  in  tem- 
perate and  arctic  regions  or  the  hot  seasons  in  the  dry  tropics. 
On  the  shady,  moist,  well-covered  soil,  the  growth  is  continuous, 
and  the  struggle  for  existence  amongst  the  many  species  in  the 
zones  of  vegetation  is  very  great.  The  principal  causes  prevent- 
ing the  predominance  of  any  one  genus  or  species  over  more  than 
a  limited  area  are  to  be  found  in  the  very  favourable  conditions 
in  which  this  strongly  differentiated  and  extremely  rich  flora 
is  placed.  * 

It  is  very  difficult  to  correctly  identify  on  the  spot  many 
of  the  high  trees  growing  in  the  Kans.  The  rapid  and  conti- 
nuous growth  produces  generally  a  thin,  smooth,  greyish 
bark  with  scarcely  any  rhytidome.  The  great  height  of  many 
of  the  stems  prevents  examination  of  their  foliage,  the 
flowers  are  also  often  inconspicuous  and  appear  at  different 
seasons  of  the  year.  These,  together  with  the  diversity 
of  the  species,  make  a  satisfactory  interpretation  of  the  flora 
almost  impossible.  In  the  smaller,  less  varied  and  more  open 
deciduous  forests  such  examination  is  not  attended  with  similar 
difficulties.  It  is  of  common  occurrence  to  see  a  tall  tree  in  full 
bloom  in  the  evergreens  and  to  be  unable  to  procure  specimens 
of  the  flowers,  except  by  felling  or  sending  up  a  native  climber, 
both  usually  very  tedious  operations.  Much  of  the  evergreen 
region  of  North  Kanara  is  somewhat  difficult  of  access,  as  the 
dense  undergrowth  often  bars  the  way  of  the  observer.  The 
forest  pathways  are  also  usually  bounded  by  monotonous  walls 
of  verdure,  without  the  relieving  colour  of  conspicuous  flowers. ' '  * 

*  The  Distribution  of  the  Forest  Flora  of  the  Bombay  Presidency  and  Sind  by 
W.  A.  Talbot.  Indian  Forester,  Vol.  XXXII,  pp.  56—58. 


243 

Bamboos,  being  as  a  rule  light-demanders,  are  usually 
absent  in  the  evergreen  forest  although  they  are  found  along 
open  water-courses,  in  clearings  and  on  the  borders  of  the  forest. 
It  is  only  possible  here  to  mention  a  very  few  of  the  most 
characteristic  and  useful  trees  and  genera  found  in  this  type  of 
forest  as  follows  : —  >. 

Tetrameles  nudi flora. 

Alstonia  scholar  is. 

Antiaris  toxicaria. 

Chukrasia  tabularis. 

Dipterocarpus. 

Diospyros. 

Dysoxylum. 

Canarium.  •..   „     . 

Swintonia. 

Sterculia. 

Lagerstrcemia  Flos  Reginae. 

Mesua  ferrea. 

Vitex. 

Cedrela  Toona. 

Pterospermum. 

Mangifera  indica. 

Calophyllum. 

Myristica. 

Artocarpus. 

Ficus. 

Cinnamomum, 

Aporosa. 

Memecylon. 

Garcima. 

It  will  be  noticed  that  a  few  species  found  in  the  deciduous 
forests  also  extend  into  the  evergreens,  such  as  Cedrela  Toona. 

In  many  places  an  intermediate  type  of  forest  can  be  more 
or  less  clearly  distinguished  in  which  many  species  of  the  deci- 
duous forests  are  found,  but  where  the  luxuriance  of  the  vege- 
tation rather  resembles  that  of  the  evergreen  forests.  In  such 
areas  trees  like  Teak,  Xylia  dolabriformis,  Dalbergia  latifolia  and 
Terminalia  tomentosa  are  found  and  there  attain  large 
dimensions. 

Among  climbers  of  the  evergreen  forest  many  species  of 
Calamus  are  common,  while  screw  pines  and  palms  are  often 
met  with,  among  the  latter  Arenga,  Caryota,  Livistona  and 
Licuala  being  noticeable. 

O  .•. 


244 

222.  The     most    important   area 

(4)  Hill        of   hill  forest  in  British  territory  is  situated  in  the  North -West 

inn?*    ts  Himalaya,  as  will  be  seen  from  the  map,  Plate  XXII.     A  brief 

in  North-      description  of  the  zones  of  forest  traversed  on  ascending  the 

We?  em        Himalayas,  starting,  say,  from   Kalsi  in  the  Dehra  Dun   and 

Himalaya,     proceeding  via  Chakrata,   will  give   a   fair  idea   of  the  chief 

characteristics  of  this  type   of   forest.     The  different  zones  of 

vegetation  may  be  conveniently  numbered  serially  as  below, 

beginning  with  the  lowest,  it  being  of  course  remembered  that 

the  transition  from  any  one  zone  to  the  next  is  often  gradual 

and  not  sharply  denned. 

ZONE  I.— Elevation  1500'— 3000 . 

SAL  predominates,  mixed  with  several  species  of  the  deci- 
duous forests,  such  as  Terminalia  tomentosa,  Anogeis- 
sus      latifolia,       Ougeinia      dalbergioides,      Adina 
cordifolia,  Odina  Wodier  and  others. 
The    Ghir    Pine    (Pinus   longifolid)    also   occurs   as 
scattered  trees,  or  in   small    patches,    and   the 
Sal  here  usually  attains  only  small  dimensions. 
Common  and  characteristic  species  also  in  these  outer 
hills  are — 

Boehmeria  rugulosa,  Bauhinia  retusa  and  Engelkardtia 
spicata. 

ZONE  II.— Elevation  3000'— 6000'. 

CHIK  PINE  predominates,  mixed  with  small  trees  of  Zone 
I  at  lower  elevations  and  higher  up  with — 
The    Ban    Oak     (QuercAis    incana)     Rhododendron 

arboreum,  Pieris  ovalifolia. 

In  valleys  Quercus  glauca,  Meliosma  pungens, 
Albizzia  Julibrissin,  Rhus  Cotinus,  Ccltis 
australis  and  Olea  glandulifera  are  common. 

ZONE  III.— Elevation  6000'— 8000'. 

DEODAR  predominates,  mixed  with  Rhododendron 
arboreum,  Ban  Oak,  Moru  Oak  (Quercus  dilatata), 
Blue  Pine  (Pinus  excelsa),  Spruce  (Picea  Morinda] 
and  many  broad-leaved  trees  such  as  Betula  alnoides, 
Populus  ciliata,  Horse-chestnut,  Walnut,  Elm,  Hazel, 
Hornbeam,  Maples,  and  Holly.  Among  shrubs 
occurring  in  this  zone  are — Berberis,  Euonymus, 
Rhamnus,  Abelia,  Viburnum.,  Lonicera,  Deutzia, 
Indigofera,  Desmodium,  Spiraea,  Rubus,  Rosa, 
Cotoneasier,  and  Salix.  A.mong  climbers  are 


245 

Clematis,  Holbcellia  latifolia,  Hydrangea  altissima, 
Vitis  semicordata  (Himalayan  "  Virginia  Creeper  ") 
and  the  Ivy. 

Above   7000  feet  Quercus  dilatata  usually  replaces 

the  Ban  Oak. 
In  this  zone  also  occur— 

The  Cypress  (Cupressus  torulosa)  often  on  lime- 
stone precipices. 

The  Box  (Buxus  sempervirens)  usually  on  calcareous 
soil,  but  also  on  other  kinds  of  soil  in  moist 
sheltered  valleys. 

The  Yew  ( Taxus  baccata)  in  shady  places,  but  perhaps 
more  often  met  with  in  the  next  higher  zone. 

ZONE  IV.— Elevation  8000'— 1 1000'. 

KARSHU  OAK  (Q.  semecarpifolia),  which  is  often  gregarious, 
predominates.  In  its  zone  are  found  the  Spruce, 
Silver  Fir  (Abies  Pindrow)  and  some  broad-leaved 
species.  Rhododendron  campanulatum  is  also  found 
which  extends  into  the  next  zone. 

ZONE  V. — Elevation  above  11000'. 

WHITE  BIRCH  (Betuta  utilis)  predominates,  often  with  a 
tangled  undergrowth  of  the  shrubby,  gregarious 
Rhododendron  Anthopogon.  In  this  zone  also 
occurs  another  shrubby  Rhododendron,  R. 
lepidotum,  and  species  of  Juniper,  e.g.  Juniperus 
recurva  on  the  northern  slopes  of  Chansil. 

Beyond  the  region  of  shrubs,  if  the  snow-line  does  not  inter- 
vene, we  find  grasses  with  Gentians,  Edel  Weiss  (Leontopodium 
alpinum]  and  other  herbs  and  then  perpetual  snow. 

223.  A  detailed  description  of  the  Hill  Forest 

hill  forests  of  the  Eastern  Himalaya,  Assam,  and  Burma  can-  of  Eastern 
not  be  given   here  although  botanically  they    are  more  inter-  As^mt'an'd 
estmg  than  the  hill  forests  of  the  Western  Himalaya.     While  Burma, 
some    species,  as  might  be   expected,    are  widely  distributed 
throughout  the  hill  forests,  others  are  more  or   less  confined  to 
local  areas. 

Rhododendron  arboreum  and  Taxus  baccata  for  instance  are 
found  throughout  the  Himalaya,  in  Assam,  and  in  Burma. 
On  the  other  hand  each  region  possesses  certain  species  of  its 


246 

own,  and  those  mentioned  below  are  more   or  less  confined  to 
the  regions  shown  : — 

Kegion.  Conifers.  Oaks. 

Western  Himalaya.  (1)  Deodar.  (1)  Quercus  dilatata. 

(2)  Cupressus  torulosa.    (2)  Q.  incana. 

(3)  Pinus  Gerardiana     (3)  Q.  Ilex. 

(inner  dry  hills  of  the 
Himalayas  and  in  Af- 
ghanistan). 

(4)  Abies  Pindrow. 

(5)  Juniperus        com- 
munis. 

(6)  J.  macropoda. 

Eastern  Himalaya.     (1)  Tsuga  Brunoniana  (1)  Q.  lanuginosa. 

(Indian     Hemlock 
Spruce). 

(2)  Larix  Griffithii. 
Assam.  Cephalotaxus  Mannii.  (1)  Q.  Olla. 

(2)  Q.  xylocarpa. 
Burma.  Pinus  MerJcusii  (I)  Q.  calathiformis. 

(2)  Q.  Brandisiana. 

(3)  Q.  Lindleyana. 

(4)  Q.  eumorpha. 

The  following  conifers  are  found  both  in  the  West  and 
East  Himalaya : — 

Pinus  excelsa.,  Picea  Morinda,  Juniperus  Wallichiana, 
Pinus  longifolia,  Juniperus  renurva. 

Pinus  Khasya  is  found  in  Assam  and  Burma,  often  form- 
ing pure  forests  with  an  undergrowth  of  grass. 

Quercus  semecarpifolia  and  Q.  glauca  are  found  throughout 
the  Himalaya  and  also  in  Assam.  Several  oaks  occur  both  in 
the  Eastern  Himalaya  and  in  Assam,  e.  g.  Q.  lamellosa,  Q.  pachy- 
phylla,  Q.  spicata  and  Q.  dealbata. 

Several  also  occur  both  in  Assam  and  Burma,  e.g.  Q.  semiser- 
rata,  Q.  mespilifolia,  Q.  polystacliya,  Q.  truncata  and  Q.  Hel- 
feriana. 

Common  to  the  Eastern  Himalava,  Assam,  and  Burma  are — 

Q.  serrata,  Q.  Griffithii,  Q.  fenestrata,  Q.  lineata,  species  of 
Castanopsis,  Bucklandia,  Machilus,  PTicebe  and  Nyssa.  Notice- 
able features  of  some  of  the  hill  forests  of  the  Eastern  Himalaya 
are  the  occurrence  of  a  large  number  of  Rhododendrons,  many 
of  which  are  gregarious  shrubs  forming  dense  thickets,  and 


247 

the  presence  of  several  magnoliaceous  trees  with  beautiful 
flowers,  e.g.  Magnolia  Campbellii. 

224.  These  forests    are  found  on  (5)  Tidal  or 

the  mud-banks  bordering  the  sea  and  tidal  rivers.     The  trees  Littoral 
are  here  never  large,   and  the  commonest  species  are  those  I 
usually  known  collectively  as  mangroves,  whence  fhisTtype  of 
forest  is  often  called  mangrove-forest.    Owing  to  their  peculiar 
environment,  the  plants  in  this  class  of  forest  have  to  contend 
chiefly  with — 

(1)  The  action  of  the  wind,  waves,  and  tides  tending  to 

uproot  the  trees  growing  in  the  soft  mud. 

(2)  The  excess  of  salts  in  the  water  around  the  roots. 

(3)  The  difficulty  of  obtaining  sufficient  oxygen  for  their 

roots. 

(4)  The  danger  of  having  their  seedlings  submerged  and 

killed  by  the  rising  tide. 

We  should  therefore  naturally  expect  that  comparatively 
few  species  would  *"be  able  to  exist  under  these  unfavourable 
conditions  and  that  those  which  are  able  to  survive  would 
possess  certain  definite  and  well-marked  characteristics. 

In  the  littoral  forests  of  India  and  Burma  "  the  species  of  tree 
which  forms  the  advanced  line  along  the  sea  and  which,  by  its 
slow  forward  march,  causes  a  gradual  elevation  of  the  coast, 
is  Rhizophora  mucronata.  No  mangrove-tree  is  better  equip- 
ped for  resisting  the  movements  of  the  tide  on  the  soft  mud, 
for  propagating  itself  under  these  difficult  conditions,  and 
for  recovering  from  the  frequently  quite  undilute  salt  sea- 
water,  the  water  lost  in  transpiration.  The  scaffolding  of  bow- 
shaped  stilt-roots  supporting  the  stem  represents  a  complete 
system  of  anchors,  which  is  strengthened  by  new  roots  growing 
down  from  the  branches  to  balance  the  growth  of  the  crown." 
(In  other  species  of  mangrove  growing  further  from  the  sea 
these  anchoring  roots  are  less  strongly  developed  or  are  alto- 
gether absent.)  :  The  leaves  possess  a  marked  xerophilous 
structure  with  a  thick  cuticle.  protected 

stomata,  and  especially  a  large-celled  thin- walled  aqueous  tissue, 
the  dimensions  of  which  increase  with  the  age  of  the  leaf  and 
with  the  corresponding  rise  in  the  amount  of  salt  contained.  Old 
leaves  serve  essentially  as  water- reservoirs  for  the  younger 
leaves."* 

*  Schimper  op.  cit.,  p.  396. 


248 

In  order  to  insure  a  sufficient  supply  of  oxygen  for  the  rootd, 
many  trees  in  these  forests  are  supplied  with  so-called  pneumato- 
phores,  or  aerating  roots.  In  some  species  these  grow  up  from 
the  ground  and  look  like  thick  shoots  of  asparagus,  in  others 
the  roots  bend  up  out  of  the  ground  and  form  knee-like  struc- 
tures, while  in  some  -cases  the  upper  surface  of  the  roots 
alone  projects  above  the  ground.  These  serial  roots,  or  portions 
of  roots,  are  usually  covered  with  thin  cork  and  possess  abundant 
lenticels.  "  The  mode  of  propagation  is  most  remarkable  in 
Rhizophora  mucronata,  which  in  this  respect  agrees  in  the  main 
with  the  other  Rhizophoraceae  living  in  the  mangroves.  The  fruit 
leathery  and  indehiscent  and  about  the  size  of  a  hazel-nut, 
soon  after  the  completion  of  its  growth  is  pierced  at  its  summit 
by  the  green  hypocotyl,  as  the  embryo  does  not  undergo  any 
period  of  rest,  but  continues  to  develop  without  interruption. 
The  hypocotyl  in  R.  mucronata  is  club-shaped  and  attains  a 
length  of  60  centimetres,  sometimes  even  more,  before  it  falls 
down,  leaving  behind  it  the  fused  cotyledons  which  served  as 
absorbing  organs.  As  its  lower  end  is  thicker,  the  seedling  falls 
vertically,  with  its  root-tip  downwards  into  the  mud,  and  within 
a  few  hours  develops  roots  that  fix  it  firmly."* 

The  principal  genera  in  these  forests  are — 

Rhizophora.  Carapa. 

Ceriops.  Avicennia. 

Kandelia.  Sonneratia. 

Bruguiera.  Lumnitzera. 

Among  shrubs  Acanthus  ilicifolius  (with  leaves  like  holly 
and  blue  flowers)  and  Mqiceras  are  common. 

The  two  small  palms  Nipa  trulicans  and  Phoenix  paludosa 
form  gregarious  thickets  in  the  Sundarbans  and  littoral  forests 
of  Burma.  Common  climbers  are  species  of  Denis  ;  epiphytes 
are  scarce  and  there  are  usually  no  mosses.  Ferns  and  grasses 
often  form  the  undergrowth  in  the  drier  spots. 

Further  inland  above  high-tide  mark,  on  ground  which  is 
only  occasionally,  if  ever,  flooded,  the  following  are 
characteristic :  — 

Hentif.ra  Fomes.  The  Sundri,  which  is  the  most  important 
tree  in  the  Sundarbans. 

Thespesia  populnea. 
Hibiscus  iiliaceus. 

*  Schimpor  op.  cit.,  pp.  396—398. 


249 

Excce:aria  Agnllocha, 

Cerbera  Odottam. 

Scaevola. 

Clerodendrcn  inerme. 

Erythrina  indica, 

Pongamia  glabra. 

Casuarina  equisetifolia. 

Pandanus  tectorius. 

225.  The    principal    types      of  Riparian 

Indian  forests  have  now  been  considered  in  detail  above,  and  it  F01^3*8- 
only  remains  to  briefly  mention  the  sub-type,  often  called 
riparian  forest,  which  is  found  along  the  banks  and  in  the  beds 
of  rivers  and  streams  and  in  swampy  places.  Forests  in  such 
localities  usually  have  an  abundant  supply  of  fresh  water 
available  in  the  soil,  and  consequently  the  belt  of  land  imme- 
diately bordering  perennial  streams  in  the  arid  region  may  be 
able  to  support  forest  of  the  ordinary  deciduous  type,  while 
similar  streams  in  the  zone  of  deciduous  forests  may  be  fringed 
with  more  or  less  evergreen  forest.  At  the  same  time  the 
conditions  under  v  hich  the  plants  of  these  forests  exist  are 
often  peculiar  in  many  respects. 

The  ground  on  which  they  grow  is,  for  instance,  often  liable 
to  more  of  less  prolonged  submersion  in  water  at  certain  seasons 
which,  by  causing  stagnation  of  water  around  the  roots,  may 
eiiectually  prevent  the  latter  from  performing  their  functions 
on  account  of  the  want  of  sufficient  oxygen.  A  very  common 
species  in  riparian  tracts  is  Ficus  cjomerata  which  sheds  its 
leaves  during  the  rainy  season,  possibly  owing  to  the  roots 
being  then  unable  to  perform  their  normal  functions. 

In  addition  to  this  also  many  of  our  Indian  rivers  and  streams 
are  entirely  dry  for  several  months  in  the  year,  when  the  sandy 
shingly  soil  in  and  near  their  beds  becomes  dried  and  excessively 
heated  to  a  considerable  depth.     Other    factors   to  be  consi- 
dered also  are  the  action  of   floods  in   washing  away  the  soil 
and  exposing  the  roots  and  the  utility  of  water  as  a  seed  distri- 
butor, those  species  possessing  devices  favouring  this  mpde  of 
seed-dispersion  having  an  advantage  in  the  struggle  for  exist- 
ence in  riparian  tracts,  as  has  been   noted  above  in  the  case 
of  Sissoo  and  Khair.    It  is  therefore  not  surprising  that  many 
species  which  are  characteristic  of  riparian  tracts,  are  not  often 
iound  elsewhere.     Among  such  may  be  mentioned— 
Anogeissus  acuminata. 
Barrincjtonia  acutangula. 
Eugenia  sp. 


250 

Ixora  sp. 
Tamarix  sp, 
Pongamia  glabra.  ' 
Trewia  nudiflora. 
Terminalia  Arjuna. 
Ficus  hispida. 
F.  glomerata. 
Homonoia  sp. 
Salix  tetrasperma. 

As  indicating  that  the  plants  in  these  forests  often  find  it 
difficult  to  obtain  their  necessary  supply  of  moisture,  it  is 
noticeable  that  Anogeissus  acuminata  and  Eugenia  Heyneana, 
which  are  common  riparian  species,  are  characterized  by  being 
small-leaved  species  of  their  respective  genera,  small  leaves 
being  as  a  rule  very  characteristic  of  xerophytes,  while  several 
plants  which  occur  in  riparian  tracts  are  also  often  found  in 
very  dry  localities,  e.g.— 

Anogeissus  acuminata  (occasionally), 

Vitex  Ne^undo, 

Streblus  asper, 

Acacia  Catechu, 

Balanites  Roxburghii, 

Capparis  aphylla, 
and  others. 

An  interesting  and  perhaps  one  of  the  most  widely  distri- 
buted plant  characteristic  of  dry  river-beds  is  Rhabdia  lycioides, 
with  its  minute  leaves  and  creeping,  rooting  branches  which 
enable  it  to  withstand  successfully  the  action  of  violent 
floods. 

Distribution  226.  The  distribution  of  all   im- 

of  important  portant  Indian  trees,  so  far  as  it  is  at  present  known,  is  given 

Species         ^n  detail  in  Brandis'  Indian  Trees  and   in    Gamble's   Manual 

of  Indian  Timbers  and  will     not     be  repeated     here.       The 

approximate 'limits  of  the  distribution  of  Teak,    Sal,  Deodar, 

Sandal,  Red  Sanders  and  Caoutchouc    have,    however,    been 

indicated  in  the  map,  Plate  XXIII. 

The  principal  factors  responsible  for  the  distribution  of  the 
chief  types  of  forest  have  been  given  above,  and  in  a  few  cases 
the  factors  which  appear  to  have  a  considerable  influence 
on  the  distribution  of  individual  species  have  been  incidentally 
mentioned.  In  the  present  state  of  our  knowledge  it  is  impos- 
sible to  indicate,  with  any  degree  of  certainty,  the  factors  which 
determine  the  distribution  of  many  of  our  important  species. 


INDEX. 


A 

PAGE. 

Abdia        244 
Abies          191 

PiOB 

^Estivation         .         .         .         .             50 
Afforesting  grasslands           .         .           236 
Agaricus  melleus          .         .         .           133 
(see  also  Armillaria  mellea). 

„     Pindrow    .         .         .    184,  245,  246 
Abnormal  growth        .         .         .18,  79 
Abruptly-pinnate         ...             25 
Abrus  precatorius         .         .         .             41 
Absorption  by  plants  83,  86-88,  91,  93,  94, 
96,  206,  207,  211 
„   soil      ...            94 
Acacia       65 
.       arabica    .         .       134,221,224,238 
caesia       .         .         .         .41,  240 
Catechu    .  16,  25,  208,  219,  220,  224, 
239,  250 
Farnesiana       .         .         .41 
leucophlcea        ...             14 
pennata  ...        29,  67,  240 
Acanthus  ilicifolius     .         .         .           248 
Acaulescent  plants     ...             10 
Accessory  buds            ...              34 

Age  of  stems       .         .         .                17,33 
Aggregate  fruit  ....             59 
Agricultural  crops,  yield  of  .         .             95 
Ailanthus  excelsa         ...             65 
Air,  its  effect  on  plant-distribution           224 
Alae  (wings)         ....             56 
Alangium  Lamarckii  .         .         .           153 
Albizzia  JuUbrissin     .          .         .           244 
„       odoratissima  .         .         .41,  239 
„       procera            ...             15 
„       stipulata         .         .         .31,  65 
Albumen    58 
„       crystals      [.         .         .            70 
Albuminoids       ....           204 
Albuminous  seed          .         .          .58,  143 
„          substances        .         .            70 
Alburnum            .         .         .          .             17 
Alcohol       129,  132 

Acclimatised      ....           229 
Accrescent          ....             63 
Accumbent         ....             59 
Acer  caesium      ...           21,  29,  33 
Achar         211 
Achene       .....             61 
Achlamydeous    ....             43 
Acicular     .         .         .                   .             23 
Acorn        .....             61 
„      cup           ....             60 

Alcoholic  fermentation          .          .    131-133 
Alder         222 
Aleurone-grains           ...             70 
Algae        .            86,  127,  137,  146,  232,  233 
Alstonia  scholaris         .         .         .           243 
Alternate  leaves.         .         .         .        •       9 
Alternation   of  gener- 
ations    .         .         .       139,  142,  146,  1*7 
Amides      204 
Ammonia  .          .          .          .         .     68,  129 

Acropetal  development        .         .           5,  9 
Actinomorphic   ....             46 
Acuminate          ....             24 
Acute         24 
Adaptation,  power  of  .              154  199  229 

Ammonium        ....             94 
Ample  inflorescence     ...             37 
Amplexicaul       ....             25 
Anabolism          ....             90 

Adhesion    .....             48 
Adina  cordifolia           .         .         .  239,  244 
Adnate       48,52 

Anatomy,  Vegetable"  .         .         .   2,68-84 
Anatropous         ....             55 
Andaman  Padauk       .         .         .           241 

„      stipules            ...             31 
Adventitious,  branches         .         .             13 
„           buds      ...             33 
„           roots     .         .         .           5,  6 
„           structures       .         .           170 
jEcidia        136,197 
^Ecidiospores       ....  136,  197 
JEcidium  Berberidis    .          .     197,  198,  200 
„        montanum    .         .         .           200 
„        Thomsoni     .         .         .           135 
JBgiceras  .....           248 
Mgle  Marmelos.         .         .          .13,  239 

Andrcecium         .         .         .         .             41 
Anemophilous     .         .         .         .           Ill 
Angiosperrns       .         .57,59,143,145-147 
„          classification  of        .           145 
Animal  Kingdom         ...               1 
Animals     .          .         .1,2,  116,    180,  182 
„       their  effect    on  plant-dis- 
tribution        ....           227 
Anjan        .         .         .         .           6,  65,  240 
Annual      .         .         .         .66,  67,  80,  176 
„      rings      .         .         .         .       17,  78 
Annular  vessels  ....       73,  7  6 

Crating  root      ....           248 
Oration  of  soil    .         .         .     206-208,  220 
Mrinl,  roots        ...        8,  107,  185 
Msculus  indica  .         .         .         c      33,  34 

Anogeissus  acuminata           .      65,  249,  250 
„          latifolia      .          15,  29,  239,  244 
„          pendula      .         .         ,          238 
Anona  squamosa          .         .         ,           228 

11 


INDEX. 


PAGE. 

Antagonistic  symbiosis  .  .  180 

Anterior  side  of  flower  .  .  46 
Anther  ....  41, 51,  52 

Antheridia  ....  138,  140 

Anthocyanin  .  .  .  69, 214 

Anliaris  toxicaria  .  .  .  243 

Anti-clockwise  twiner  .  .  10 

Antidesma  diandrum  ...  29 

Antipetalous  ....  52 

Ants 183 

Aonla 208 

Apetalous  ....  43 

Aphides  ,  ...  133 

Apocarpous  ,  '  .  .  .  53 

Aporosa  .....  243 

Apothecium  .  .  .  .  131 

Apple 63 

Aquatic  plants  .  .  .  .  185 

„  roots  ....  8 

Arched  nerves  .  .  .  .  21 

Archegonia  ....  138,  141 

Arctic  Zone  ....  222 

Arcuate  nerves  .  .  .  .  21 

Arenga  .  .  .  .  .  243 

Arhar 67 

Arid-country  Forests  .  .  .  237-238 

Aril 58 

Arisaema  Wallichianum  .  .  41 
Armillaria  mellea  .  .  .  133,  193 

Aroids  .  .  .  .  242 

Arrangement  of  leaves  .  .  9,  32,  33 

„  ,,  parts  of  flower  .  44,  47 
Artificial  classification  .  118-119,124 

Artocarpus  ....  243 

Arundinaria  falcata  ...  30 

,,  spnthiflora  .  .  21 

Ascending  ovule  ...  54 

„  stems  ...  10 
Ascent  of  water  in  plants  .  96-98,  168-169 

Ascocarps  ....  131 
Ascomycetes  ....  131,  137 

Ascus  .....  131 

Aseptate  hyphse  .  .  .  130 

Asexual  generation  .  .  .  139 
Asexual  reproduction  .  108,  156,  158 
Assimilation  .  .  .90,  101-104 

Asymmetric  ....  46 

Atavism  .  .  .  .  .  157 
Atmosphere,  its  influence  on 

plant-development  .  208-216 
„  its  influence  on 

plant-distribution  224 

Auricled  .  .  .  .  24 

Austral  Zones  ....  222 

AutoBcious  ....  198 

Autumn  tints  ....  29 

Availability  ....  94 

Available  water  .  .  .  206, 218 

Avicennia  ....  248 

Axial  embryo  ....  59 

Axil  .'....  9 

Axile  placentation  ...  54 

Axillary  huds  ....  9 
flower  .  .  .37 


PAGE. 
Babul         .         134,  221,  224,  227,  231,  238 

Bacilli 127 

Bacillus  radicicola        .         .         .  129, 203 
„      tetani      ....  129 

„      typhi      .   '      .          .         .  129 

Bacillus  vulgar  is          .         .          .  129 

Bacteria  .  127-129,  146,  154,  176,  179, 
187,203,204,207,208,214, 
226. 

,,        nitrifying      .         .         .  129 

Bacterium  aceli  ....   127,  129 
„         acidilactici  .          .  129 

Balanites  Roxburghii  .          .          .  250 

Balsams    .....  75 

Bamboos  .  10,  11,  14,  19,  21,  30,  56,  59, 
67, 145, 176, 182, 183,  222, 

228,  243 

Bambusa  polymorpha  .         ,         .  183,  241 

Tulda  ...  241 

Banana      .          .         .21, 22,  66,  215,  222 

Ban  Oak    ....        7,244,245 

Banyan      .         .          .          .    5, 8,  185,  186 

Bardayella  deformans  .         .  135 

Bark  .      .         .          14,15,16,18,80 

„    internal     .          .          .          .18,80 

Barley 95 

Barrinatonia  acutangula       .  249 

Basal  nerves  .  .  .  .21,22 
Base,  leaf—  .  .  .  .  24,  30 
Basidia  .....  133 

Basidiomycetes  .  .  .  .  131,  133 
Basifixed  .....  52 

Bassia  latifolia  .         .  7,  65,  132,  211,  239 

Bast 75,78 

„     fibres         ....  78 

„     internal     ....        18, 80 

Bastards 150 

Bauhinia  malabarica  ...  29 

„       racemosa      .         ,         .  239 

„       retusa  .         .         .  244 

Vahlii         11, 17,  18,  79,  184,  239 

,,        variegata       .         .         .  239 

Bean 56,203 

Beech 191,222 

Bees  .          .        113,  114,  177,  178,  227 

Beet 7 

Bel 13,36 

Bell-shaped        ....  50 

Ber 208 

Berberis     .         .         .36,  196,  222,  244 

aristata  .          .         .     15,200 

„       coriaria.         .         .          .  200 

Lycium        30,  31,  45, 112-115, 197, 

200 
,,  „       fertilisation   of 

flowers  of 

Berberis  nepalensis       .         .         .     30,  137 
Berchemia  ftoribunda  . 
Berry         .....  61 

Be  tula  alnoides   ....  244 

Betula  utilis        .         .         .         .15,  245 
Bi-collateral  vascular  bundle 
Biennial     ,  .         .       66,67 


INDEX. 


Ill 


Bifarious  leaves 
Bifoliolate 
Bijasal 
Bilabiate   . 
Bilocular    . 
Biological  sciences 
Biology 
Bipinnate 
Birch 

white 


PAGE. 

9 

26 

240 

50 

51 

1 

1 

25 

15-17,  63,  80,  222 
245 


Birds         .  42,  111,  116,  182,  201,  227 

Biserrate 22 

Bisexual    .....  45 

Black  Soil  ....  220 

Blackwood  240 


Bleeding,  of  stumps     . 
Bloom 
Blue  Pine  . 
Bcehmeria  rugulosa 
Bombax  malabaricum  . 
Bombay  Blackwood    . 
Borassus  flabellifer 
Bordeaux  mixture 
Bordered-pit 
Boreal  Zones 


97 

64 

183,  224,  225,  244 
244 

.  8,  14,  65,  239 
240 

18 
189 

71 
222 


Boswellia  serrata       15,   171,228,  232,  239 
Botany      .....  1 

„      subdivisions  of  2 

Bougainvillea      ....  47 

Box 245 

Bracken  Fern     .         .         .          .     11,  140 
Bracteate  .         .  *      .          .          .  37 

Bracteoles.         ...          37, 44, 45 
Bracts        ...         37, 40, 44-47,  60 

Brambles 226 

Branches  .          .          .          .10, 12-14, 65 

„       development  of     .          .  82 

Branching.          .        5,  12,  13,  37,  52,  64,  65 

Branchlets          .         .         .          .       14, 65 

Brinjal 153 

Browsing,  by  cattle     ...  67 

Bruguiera  ....  248 

Bruises 167 

Bryophyta  .         .         .     126,  137,  147 

Buchanania  latifolia    .  15,16,211,239 

Bucklandia         ....  246 

Budding 175 

„      of  Yeast        .         .          .  130,  132 
Buds          .         .          .9,  10,  33,  34,  46,  48 

„     accessory  ....  34 

„     position  of          ...  9 

Bud-scales          .          .         .         .       33, 34 

Bulb 11,  109 

Bulbils 11,  109 

Bundles,  vascular        .  18-20,74-79,84 

Butea  frondosa    .  56,220,221,227,239 

„     superba     ....  239 

Buttresses.         .          .          .         .         8,  14 
Buxus  sempervirens     .         .         .  245 


Caducous  . 
Csespitose  stems 


Cajanus  indicus 
Calamus    . 

tenuis    . 
Calcareous  soil    . 
Calcium     . 
Callus 

'alophyllum 
Calyptra 
ialyx 

„      limb 

„      throat 

„      tube 
Cambium  . 

cork  . 
Campanulate 
Campylotropous 
Canala,  resin 
Canarium 
Canescent 
Cane-sugar 
Cankers 
Caoutchouc 

Tree 
Cap, root — 
Capitulum 


PAGE. 

67 

243 

10 

.  220,245 

71,93, 206,  220 

165,  169, 170 

-       .  243 

139 

43, 50, 60 
43 
43 

.       43, 49 

76,  77,  79 

80 

50 

54 

.       18,74 

243 

64 

103 

213 

.       71, 74 

.    219,250 

82 

38 


Capparis 47,  237 

aphylla          ...  250 

spinosa          .         .         .31,  238 

Capsule 60 

Carapa 248 

Carbohydrate,  manufacture  of  a  .         3,  89 

101,  102 

Carbon       .          .          .68,  70,  91,  93,  94 

Carbonaceous  food       .          .          .    178-180 

Carbon  dioxide  .     2,  85,  89,  94,  101,  102, 

103,  132,    179,  180, 

204,  206,  21 C 

Carbon  dioxide,  excreted  by  root- 
hairs       .....  94 
Careya  arborea    ....           239 
,,        herbacea  ....  67 
Carina        .          .          .          .        _             56 
Carissa  spinarum         .          .         .       15, 67 

Carpel 53,  143 

Carpinus  viminea         .          .          .14,  34 
Carpophore         ....  54 

Carrot        .....  7 

Cartilaginous      ....  63 

Caruncle   .....  58 

Caryopsis  .          .          .          .  61 

Caryota 243 

Cassia  Fistula    .         .         .         29,41,239 
•  .  241 

Castanopsis         ....  246 

Castor-Oil  Plant  ...  52 

Casuarina  cquisetifolia         .         .  249 

Catkin 38 

Caudate 24 

Caudex      .         .          .         .         .  11 

Cauliflower          ....  156 

Cedar,  Red          ....  240 

Cedars 145 

Cedrela  Tooni     .         .         .6.  60,  240,  243 
Cedrus  Libani  par  Deoiara  .121,  191 


INDEX. 


PAGE. 

Cell  .         .         .         .         .         .  69 

contents      ....  69 

dead  ...  72 

division       .         .         .         .  71 

guard 84,100 

—sap  ....  69 

shape  of      .  72 

wall 69 

thickening  of  71 

Cellular  plants  .  .  .  .126,146 
Cellulose  ....  69,71,90 
Celtis  australis  ....  244 

Centrifugal  inflorescence      .         .  39 

Centripetal  „  39 

Cepkalostachyum  perrjracile  .          .  241 

Ce-phalotaxus  Mannii  -          .         .  246 

Cerbera  Odollam  ...  249 

Cenops 248 

Chalaza 54 

Channelled          ....  14 

Characters  .         .       117,  119,  124,  125 

important  .          .  120 

latent         .          .          .  157 

of  ovules    .          .         .       54,  55 
„  perianth          .          .  43 

„  petiole  ...  29 

„  pistils    .          .         .53, 54 
„  stamens  .         .       51, 52 

Characters,  specific      .         .         .  120 

Chartaceous        ....  63 

Chestnut 222 

Chir  ....    136,  224,  244 

Chlorophyll    1,  2,  70,  75,  94,  102,  214,  215 

,,      corpuscles  .     70,  214 

„      light  absorbed  by  .          102 

Chloroxylon  Swietenia  .  220,  240 

Chrysomyxa  Himalense      .    . 

Chukrasia  tabvlaris  243 

Cilia  ....     127,  138,  188 

Ciliate        ...  22 

Cinnamomum     ....  243 

„  Camphora     .         .  21 

Tamala         .     •    .  29 

Circinnate  .         .         .         .35,  139 

Cladode 12 

Class 122 

Classification      .         .          .    2,  3,  117-164 

artifici.il         .     118-119,  124 

natural  .        118-119,  123-125 

„          necessity  for     .         .         117 

„          of  Angiosperms        .  145 

„  fungi  .         .  131 

„  „  Gymnosperms      .  144 

„          unit  of   .         .         .  119 

Clavate 62 

Claw 49 

Clay 220 

Cleft  ....          22,  23,  44 

Cleistogamic  flowers    .         .         .  116 

Cleistothecium   .         .         .         .  131 

Clematis 55, 245 

,,      montana        ...  31 

Cterodendron  inerme    .         .         .  249 

,.  terratum      .        .  67 


PAGE. 

Climbing  plants  .      10,  33,  65,  66,184 

Climbing  plants,    are   sometimes 

erect 67,  153 

Clockwise-twiner          ...  10 

Closed  vascular  bundle          .          .  76 

Close  symbiosis  ....  180 

Clover  .  .  .  177,  178,  203,  227 
Club-mosses  .  .  .  141,142,  147 
Cluster-cups  .  .  .  .  197 

Cobra  Plant        ....  41 

Cocci 127 

Cocculus  laurijolius  .  .  18,  28,  79 
Coccus  .....  60 

Cocoanut  Tree  .         .         .  219,  229 

Cocos  nucifera     ....  229 

Cohesion    .....  48 

Cohort 122 

Cold,  excessive  ....  209-214 
Collateral  host  ....  198 

„        vascular  bundle    .          .  75 

Collection  of  specimens         .         .  162 

Collenchyma  .  .  .  .  73, 75 
Colocasta  antique/rum  .  .  .  100 

Colour  of  bark  ....  15 

„  „  leaves  .  .  .28,  29 
„  „  perianth  .  .  42, 43,  111 
„  „  twigs  ...  80 

Combretaceae       ....  222 

Combretum  nanum       ...  67 

„          ovalifolium  .         .  65 

Communities,  succession  of  plant  .  233 

Compact,  inflorescence          .          .  37 

Competition  .  .  6, 109, 148,  184 
Competitors  .  .  180,183,226,228 
Compound  corymb  ...  38 

cyme          ...  39 

leaf  ....  23 

pistil          ...  53 

raceme       ...  38 

umbel        ...  38 

Compressed         .          .          .         .  61 

Concavo-convex  ...  62 

Concentric  vascular  bundle  .          .  76 

Conditions  of  plant  life         .         .  85 

„        „  existence  in  a  highly 

organised  plant        .  91 

Conduplicate       ....  35 

Cone  .          .         .          .56,  144,  145 

Conical 7,61 

Conidiophore  .  .  .  .  131,  187 
Conidium  ....  131,  187,  189 
Coniferae  .  .  .  144, 145, 147,  222 
Conifers  .  18,  74,  78,  101,  168,  191,  194, 
224,  225,  246 

Connate  .  ...  25,  43,  48 
Connective  .  .  .  ,  01,  53 
Contorted  ....  51 

Contraction,  of  root    ...  83 

Contrivances,  for  facilitating  cross- 
fertilisation     .         .         .          .  Ill 
Conversion,     of  grassland     into 

forest     .          .          .         .  i  235 

Convolute  ....  35 

Coppice,  leaves  of       .  .    28,  153 


INDEX. 


Coppice-shoots   . 

PAGE. 
.  169,  171 

PAGE. 

Cyme,  helicoid  .                                          40 
„     racemiform                 .         .             40 
„     scorpioid                     .         .             40 
„     trichotomous              .         .             39 
„     umbelliform               .         .             40 
Cymose  branching       -—      ,  __      12,  37,  38 
„         inflorescence            .         .        38-40 
Cypress      ....     136,145,245 
„       Swamp           ;                               67 

Cordate 
Coriaceous 
Coriaria  nepalensis 
Cork           .          .         72, 
„    cambium  . 
Corm 

24 
63 
.       14,  32,  107 
75,  80,  81,  82,  165 
80 
.     11,  109 

Corneous   . 
Cornus  macrophylla 
Corolla 

63 
.       21,  41 
.      43,  50 

„      limb 
„      papilionaceous 
„      throat    . 
„      tube 
Corona 
Corpuscles 
„            chlorophyll 
Cortex 
Corylus  Colurna 
Corymb 
Corymbiform 
Cotoneaster 
Cotton  Plant 

43 

56 
43 

43 
50 
70 
.     70,  214 
14,  81 
.       17,  65 
38 
40 
244 
44.  58 

D 

Dahlia       .                   ...               7 

Dalbergia  ......             56 
„        cultrata        .         .         .           241 
„        latifolia        .         .         .  240,  243 
„        paniculate,  .         .         .18,  29 
Sissoo          .          6,219,224.233 
Date  Palm          ....             63 
Dead  cells           ....             72 
„     tissue         ....           204 
Death,  of  plants.          .          .          .            176 
Decay        .          .          .          .  •       .  129,  179 
,,       of  wood    ....           204 
Deciduous  63  67 

Cotton  Soil         ....           220 
Cotyledons          .         .          .         .       58,  59 
Counter-clockwise  twiner     .         .            .10 
Crateriform         ....             50 
Creeping  stems  ....             10 
Crenate      .....             22 
Crops,  mixed      ....             95 
„     rotation  of          .         .         .95,  226 
.,      succession  of      .         .                     233 
Cross-fertilisation        .          .          .    110-115 
„             „             contrivances 
for    facilitat- 
ing      .          .           Ill 
„     illegitimate         .         .          .112,  123 
Crown,  shape  of          ...             65 
Crumpled  ...                                KI 

„           Forests     .         .      230,237-241 
plants       .          .     100,  101,  229 
Decumbent         .                   .                 10  65 

Decurrent           ....             25 
Decussate  leaves          ...               9 
Deeringia  celosioides    .         .         .             li 
Definite  inflorescence  ...             39 
Dehiscence,  of  anther            .         .       41,51 
Dehiscent  fruit  ....             60 
Deltoid      ....                       24 

Dendrocifamus  longispathus           .           241 
„                sntwuo            .           .              239 
Dense  inflorescence               .         .             37 
Dentate     .          .                                          22 

Crustaceous 
Cryptogams 
Cryptole.pis  Buchanani 
Crystals     . 

63 
.     126,  146,  183 
240 
71 

Denuding  action  ot  water     .         .219,  249 
Deodar      .   13,  18,  65,  183,    220,   224.  225, 
244,  '246,  250 
„       Forests  .          .     183,  224,  225,  244 
„       root-disease              .      134,  191-193 
Depressed           ...                       62 
Derris        ....                     248 

Culms 
Culm-sheath 
Cuneate     . 
Cup,  of  acorn 
Cupressus  torulosa 
Cup-shaped 
Cupuliferae 
Ciiscuta  reflexa    . 
Cuspidate 
Cuticle 
Cuticularised 
Cutin 
Cuttings    . 
Cycadaceae 
Cycads 

11 
30 
24 
60 
15,  136,  245,  246 
50 
222 
201 
24 
75,  101,  154,  247 
75 
75 
.    170,171 
.    144,  147 
.  147,  222 

Descriptive  terms,  for  petals  and 
sepals  .         .             49 
,,              ,,      for  shape      of 
fruit  and  seed              61 
Desert        .....    231-234 
„       plants      .          .          .          .          loo 
Desmodium         ....  225,  244 
„           piilchellum         .         .            41 
„           tiliaefolium         .         .       26,  31 
Determinate  inflorescence    .          .             39 
Determination,  of  natural  groups           123 
Deiitzia       .....           244 
Development,  abnormal       .         .             79 
„          of  plant-members    .            77 
„           of  root  .         .         «   6,  7,  82 
„          of  secondary  mem- 
bers  .                               82 

Cycas 
„    pectinata     . 
„    Rumphii 
Cylindrical 
Cyme,  corymbiform    . 
„     dichotomous    . 

.     79,  144 
144 
144 
61 
40 
39 

VI 


INDEX. 


PAGE. 

Development,  of  vascular  bundles.  76 

Dextrorse  twiners        ...  10 

Diadelphous       ....  52 

Diageotropic       ....  106 

Diagrams,  floral           ...  57 

Diaheliotropic    ....  107 

Diandrous.          ....  45 

Diastase    .....  103 

Dichasium           ....  13 

Dichlamydeous  ....  43 

Dichogamy         .          .         .          .  Ill 

Dichotomous  branching        .         .  12,  13 

„          cyme      ...  39 

Dicotyledons        59,77,78,101,145-147,  168 

„          root  of             .         .  79, 81 

„          stem  of  .         .          .  77 

„          wood  of  78 

Didymous           ....  62 

Didynamous       ....  52 

Digitately-veined         ...  21 

Dillenia  parviflora       .         .         .  240 

,,       pulcherrima    .         .         .  241 

Dimerous            .          .          .          .  45 

Dimorphic  flowers       .          .          .  112,  123 

„          species       .         .         .  154 

Dioecious  .....  45,  111 

Dioscorea  .         .         .         .  11,  145 

Diosmosis           ....  86 

Diospyros  .....  243 

„        lomentosa     .         .  7,  15,  236 

Diplostomonous           ...  52 

Dipterocarpaceae           .         .         .  222 

Dipterocarpus     ....  243 

,,           tuberculatus    .         .  220,  241 

Disassimilation  ....  90 

Disc  .          .          .          .          .49,  113 

Discoid 61 

Disease,  definition  of  .          .          .  176 

„       rarely  due  to  one  factor  .  181 

„        spike—.         '-.          .          .  202 

Diseases    ....       129,  176-217 

„        investigation  of      .          .  182,  202 

„        symptoms  of           .         .  181 

Dissemination  of  seeds          .          .  115 

Dissepiments      ....  53 

Distant  symbiosis         .          .          .  180,  204 

Distichous           .          .          .  9,  32 

Distribution,  of  important  species  250 

Distribution,  of  plants  depends  on 

action  of  man  .          .          .         228 

Distribution,  of  plants  depends  on 

action  of  air    ....  224 
Distribution,  of  plants  depends  on 

action  of  animals      .          .          .  227 
Distribution,  of  plants  depends  on 

action  of  fire   ....  227 
Distribution,  of  plants  depends  on 

action  of  light           .          .          .  224 
Distribution,  of  plants  depends  on 

action  of  other  plants        .          .  225 
Distribution,  of  plants  depends  on 

action  of  soil     .          .          .          .219 
Distribution,  of  plants  depends  on 

action  ot  temperature       .          .  221 


PAGE. 


Distribution,  of  plants  depends  on 

action  of  water 
Distribution,  of 

„  seed  58,  109,  115, 

„  ,,  sexual  organs 

„  „  types  of  forests    . 

Divaricate  .... 

Divided  leaf       .... 

Division     . 

„      cell—     . 

Dodder      . 

Dolichandrone  falcata  . 

Dormant  buds    .... 

Dorsal  raphe       .... 
,,      side          . 

Dorsal  suture      .... 

Dorsifixed  .... 

Drupe        ... 

Dry-rot      .... 

Duramen  ..... 

Duration,  of  plant  life 

,,          „     ,,     members 

Dwarf  shoots      .... 

Dysoxylum          .... 


218 


182 


219, 

227 
45 

237 

65 

22,23 

122 
71 

201 
41 
34 
55 
52 
53 
52 
61 

134 
17 

176 
03 
13 

243 


Ebracteate  . 
Eccentric  embryo 
Echinate  .. 
Economic  Botany 
Edelweiss  .. 
Egg-plant  . 


E 


216, 


42,  55,  58.  59, 
.      73, 


58 
3,  89,  90, 


Elements,  of  tissues 

Ellipsoid    . 

Elliptical   . 

Elm 

Elongated  shoots 

Emarginate 

Embryo     . 

Embryonic  tissue 

Embryo-sac 

Endocarp 

Endophytes 

Endosmose 

Endosperm 

Energy,  in  plants 

Eng 

Enrjelhardtia  spicafa    . 

Entire  margin     .... 

Entomophilous  .... 

Envelopes,  floral — 

Enzymes  103,  132,  188,  192, 

Ephzdra  Gerardiana 

Epicalyx   ..... 

Epicarp     ..... 

Epicotyl    ..... 

Epidermis     75,  80,  82,   84:  96,  99 

Epigynous          .... 

Epipetalous         .... 

Epiphytes  .          .       185,241, 

Equatorial  Zone 

Erect  inflorescence 


37 

59 

64 

o 

245 
139 
65 
73 
61 
24 

222, 244 
13 
24 

138,  143 
170,  176 
55,143 
60,61 
185 
86 

,  143-145 

102,  104 

220,  241 

244 

22 

111 

41 

201,  205 
145 
44 

60,61 
59 

•101,  214 
49 
52 

242,  248 

221 

37 


INDEX. 


vii 


Erect  ovule 
„     stem 
Erythrina  indica 

„        suberosa 
Essential  food-materials 


PAGE. 
54 

.       10, 65 
249 

15,  17,  171,  239, 
•  93 

„  „  power  of 

plants  to  obtain  .  .  .94,  95 
Essential  mineral  salts  .  .  205,219 
Ethereal  oils  .  .  .  29,71,74 
Etiolation  ....  214 

Eugenia     .....  249 

,,        Heyneana       .         .         .  250 

Jambolana     .         .          7,22,239 
„      operculata       ...  41 

Euonymus          ....  222,  244 
„         Hamiltonianus    .         .  28 

Euphorbia  ....       65, 74 

Evergreen  ....  67 

„         Forests       .         .         .    241-243 
plants          .        07,  100, 101,  229 
Evolution,  theory  of    .          .          .  159 

Exalbuminous  seed     .         .         .     58,  143 
Excess  of  salts   .          .          .          .205,247 
Excessive  cold    ....    209-214 
heat   ....  209 

,,         moisture      .          .          .  215 

Exccecaria  Agallocha  .         .         .  249 

Excretion  .         .         .       90,206,226 

Exfoliation,  of  bark     .         .         .16,  81 
Existence,  conditions  of,  in  a  high- 
ly organised  plant      .  91 
Struggle  for  96, 148-149, 159, 177- 
178 

Exosmose  ....  86 

Exserted  stamens        ...  52 

Exstipulate         ....  31 

External  embryo         ...  59 

Extra-axillary    ....  34 

Extremes,  of  temperature    .         .  208,  222 

Extrorse 52 

Eyes,  of  Potato  .         ...  11 


Factors  causing  disease         .     177,  181,  217 
„       influencing  plant  distribu- 
tion   ....  218-230 
„      influencing   relations     be- 
tween organisms  .         .  178 

Falcate 24 

False  axis            .         .                   .  12  ,40 

dichotomous  branching       .  12 
dichotomy           .          .          . 

dissepiments       ...  53 

rings           ....  17 

trichotomous  branching       .  12,  13 

whorls       .         .         .  9, 65 

Family 122 

Fascicle     .....  40 

Fascicled  leaves            .          .          .  £,13 

Fats           .....  71 

Female  llower  45 


PAGE. 

Ferment  Organisms     .          .         .  132 

Fermentation     .          .       129, 131,   132, 204 

Ferns         .  22,  139-141,  147,  242,  248 

„   life-history  of      .         .         .  ]4i 

„   Tree  .          .         .  11, 19, 140, 222 

Fertilisation        .         T          4i',  55, 143, 145 

,,  cross        .         .          .     110-115 

„  ,,    contrivances 

for  facilitating  . 
,,  of  Bryophyta    . 

„  of  flowers  of  Berberis 

Lycium 
„  of  flowers  of  Salvia 

lanata  . 

„  of  Pteridophyta 

self—       .    '     . 
Fertility,  of  species 
Fibres        .... 

,,      bast — 
Fibrous  root 
Ficus 

bengalensis 
Cunia 
elaslica 
glomerata  . 
hispida 
infectoria  . 
pumila 
religiosa    . 
scandens    . 
Figs 

Filament   . 
Filices 

Filiform 

Filtering  water  .... 
Fimbriate  .... 

Fir  Forests  of  N.-W.  Himalaya 
Fire,  effect  on  plant-distribution 

of 227-228 

,,     its  effect  on   plant-develop- 
ment       .... 
Fires  .... 

Firs Ill,  145 

Fission  Fungi     .         .         .         .  128 

Fissures,  in  bark 
Flacourtia  Cataphracta 
„      Ramontchi 
Flagella     . 
Flemingia  congesta 

„        stricta 
Fleshy 
Floccose     . 
Flood-plains        .          .• 
Flora 
Floral-axis 

diagrams    . 
envelopes  . 
formulae     . 
parts 
receptacle 
shorthand 


111 

141 

112-115 

112, 113 
141 

110-115 
123 

.       73, 91 

78 

7 

31,34,186,227,243 

5,22,   185 

-2  37 

.  173,219 

.     239,  249,  250 

.  239,250 

239 

27 

185 

8 

74 

41,  51,  52 
139 
29 
207 
22 
225 


216 
67 


Flower 


bud 
parts  ot 


15 
13 
239 
127 
14 
14 
63 
64 

.  234,239 
117,119,121,124 
.       48, 49 
57 
41 
57 

41,44,46 
.42,  49,  60 
57 

3,4,37,41-57,143 

46 

41,  44,  46 


vm 


INDEX. 


PAGE. 

Flower,  parts  of,  are  leaves  .         .  46 

j,  „     „  position    and  num- 

ber of  44 

„      symmetry  of    ...  46 

Flowering  plants          .         .         .  126,  142 
„        shoot  ....  37 

Flowerless  plants         .         .         .  126 

Flowers     .         .         .         .         .  126 

„      green—.          ...  47 

„      types  of  which  may  cause 

difficulty     ...  55 

Fluctuating  variability        152-155, 157, 160 
Fluted  stems  ...  14 

Follicle 60 

Fomes  annosus   .         .         .         .  134 

(See  also  Trameles  radiciperda.) 
„       life  history  of    .          .          .    191-193 
„       Pappiamis        .          .          .  134 

Food-materials,  plant  3,  20,  89,  91,  93,  103, 

104,  178-180 

„          „  storage  of   .         .  7 

Foot  of  fern       ....  141 

Forests,  Arid-country  .         .         .  237,  238 
„       Deciduous      .         .         .    238-241 
„       Evergreen      .         .         .    241-243 
„       HiH        .         .         .      224,  244-247 
„       Littoral          .         .         .    247-249 
„       Riparian         .         .         .    249-250 
„       Tidal     ....    247-249 
„       types  of  and  factors  res- 
ponsible for  their  distri- 
bution        ...  237 
Formulae,  floral  —      ...  57 

Free 48 

Free-central        ....  54 

.Fronds       .         .         .         .         .  139 

Frost 67 

„       cankers   ....  213 

„       holes       ....  235 

„       injury  by          .         .      209-213,215 
,,       resistant  species        .         .  208 

»       ribs          ....  213 

„       tender  species  .         .         .  208 

Fruit 3,59 

„     aggregate  ....  59 

„     descriptive  terms  for  shape 

of  ....  61 

„     multiple    ....  59 

„     simple        ....  59 

„ — sugar         ....  103 

^  „     types  of     .          .          .          .       60, 61 

Fugacious  ....  63 

Functions  of  plants     .         .        2,3,85,86 

„          of  higher  plants  .  93 

Fundamental  tissue     .         .          74,  75,  84 

Fungi   2,  127,  129-137,  146,  156,  179,  183, 

186,  204-205,  207,  208,  213,  219,  226 

Funicle 54 

Funnel-shaped  ....  50 

Furcate  venation .       .         .         .22,  139 

Fusiform 7,  61 

Fusion,  root—    ....    170,175 


Galls 

Gambleola  cornuta 

Gamete 

Gamo-petalous  . 
„      phyllous 
,,      sepalous  . 

Garcinia    . 

Gardenia  turgida 

Garuga  pinnata  . 

Gases,  in  the  plant 


PAGE. 
156 
137 
126 

43,48 
43 

43,48 
243 

28,  239 
239 
82, 84, 87,  89 


Generations,  alternation  of  .     139, 142, 146 
„  plant,  succession  of  .  233 

Generic  name     ....  121 

Gentians    .....  245 

Genus        ...  .    121-123 

Geographical  Botany  .         .      2,  3,  218-250 
Geotropism         .          .         .         .  105 

Germinating  seeds       ...  103 

Germination"      ....  59 

Germs        .         .         .         .         .  127 

Gibbous 50 

Gills 133 

Girdling    ....       168-170,  185 
Glabrescent        ....  64 

Glabrous 64 

Gland-dotted      .         .         ...  29 

Glands       .         .  29,49,63,66,74,114 

Glandular  hairs  .          .          .  65 

Glaucous    .....  64 

Globose  ....  61 

Gloriosa  siipefba  .         .         .  31 

Glucose 97,  103 

Glume,  flowering         ..  56 

„       outer       ....  56 

Gmelina  arborea  ...  16 

Gnarled  stems  • .         :         .  14 

Gnetaceae    ....     144,  145,  147 

Gnetum 9,  145 

Goats 227 

Gonophore  ....  47 

Gossypium  ....  44 

Gouania  leptosiachya    .         .         .  13 

Grafting    .....  174 

Grain,  silver        .         .         .         .  17 

Grains,  aleurone          ...  70 

„        starch    .         .         .  70,  97 

Grape-sugar        .         .          .         .     97,  132 

Grasses      .6,11,21,30,42,50,56,    59,100, 

111,  134,  184,  219,  223,  224, 

225-227,  231-235,     245,  248 

Grassland  .         ...    231-236 

„  afforestation  of  .  .  235-236 
Gravity,  its  effect  on  plants.  .  105-107 
Grazing  .  .  96,156,207,227,236 
Green-flowers  ....  47 

„      plants       .         .          .     178-180,204 
Gregarious          ....     67,  111 

Grewia 47, 61 

„     asiatica      ....  239 

,,     pilosa        ....  63 

Groups,  natural  .         .         .  122,  123 

Growing-apex     .         .         .  69, 76 


INDEX. 

. 
IX 

PAGE. 

PAGE. 

Growing  points  .         . 

.      72,  76 

Hilum        .... 

.    -        55 

Growth      .         .         .    68,85,89,104,105 

Hirsute      .... 

64 

i            i 

1  8    7Q 

64 

,,        abnormal 
„        retarded  by  light 
Guard-cells 

.           .        lo,  rj 
.  104,  105 
.     84,  100 

Hoary        .... 

Holarrhena  antidysenterica    , 

64 
.  116,239 

Gum 

78,  82,  168 

Holbodlia  latifolia 

.     26,  245 

Guti 

.  172,  173 

Holly         .         .         . 

.  222,244 

Gutta-percha 
Guitiferae  . 

74 
222 

Holoptelea  integrifolia 
Homalium  tomentosum 

31 
240 

Gymnosperms    .    57,  77, 
„  _         bast  of 

78,  143-145,  147 

78 

Homologous 
Homonoia 

36 
250 

i            „            classification  of      .           144 

Hooks        .... 

.     10,  116 

„           .root  of 

79 

Horizontal  ovule 

54 

,,            stem  of 

77 

Hornbeam 

14,  222,  244 

Gymnosporangium  Cunningham- 
ianum    .....           136 

Horse  Chestnut 
Horsetails 

33,  216,  244 
141,  142,  147 

Gynobasic 

54 

Host          .... 

180 

Gynoacium 

.       41,  53 

„    collateral  . 

198 

Gynophore 

47 

„    intermediate 

198 

Humble-bees 

.  177,227 

Humus      .    94,203,206,207, 

220,  221,  226 

Hyblaea  puera    . 

178 

H 

Hybrids     .... 

150-152,  160 

Hydrangea  altissima    .         . 

245 

Habit 

65 

Hydrogen            .          . 

68,  70,  93,  94 

Hail,  damage  by 

214 

Hydrotropic 

106 

Hairs                   .       33,  35,  58,  63-65,  75,  99 

Hydrotropism 

106 

„    on  seed 

.     58,  116 

Hygrophyte 

229 

,,    root  — 

83 

Hymenial  layer  . 

133 

Half  -inferior 

49 

Hymenium 

133 

Halophytes 

206 

Hymenodittyon  excdsum 

.     16,  239 

Hamiltonia  suavcolcHs 

RK 

Hvphse 

.  130,  188 

Hardness,  of  wood 

•               •                     UO 

17 

Hypocotyl 

59 

Hardwickia  binata 

6,  65,  240 

Hypocrateriform 

50 

Harra 

7 

Hypogynous 

48 

Hastate 

24 

,,      leaf 

25 

Haustoria 

130,  188,  200-202 

I 

Hazel 

.     203,  222,  244 

Head 

38 

Illegitimate-cross 
Imbibition 
Imbricate 

.   112,  123 
87 
fil 

Healing  of  Wounds      . 
Healthy  development 

165 
177 

Heart-wood        .          .     17,  78,  98,  168,  204 
Heat,  excessive  ....           209 
Helicoid  cyme    ....             40 

Impari-pinnate  . 
Impatiens 
In     

•                          t-J-L 

25 

116 
.  220,  241 

Heliotropism 
Hemlock  Spruce,  Indian 

107 

246 

Inarching  .... 
Included  stamens 

174 

52 

Hepaticae  . 
Heptapleurum  venulosum 
Herbaceous 
Herbarium 

137 
41 

10,  66 
.  163,  164 

Incumbent 
Indefinite  inflorescence 
Indehiscent  fruit 
Indeterminate  inflorescence 

59 
39 

60 
39 

Herbs 

66  67 

Indian  Forests,  distribution  of     .           237 
„         „         types  of      .         .           237 
Indigofera           ....  225,  244 
Induplicate,  valvate  —         .          .             51 
Indusium  .....           140 
Inferior     43,49 
,,    radicle      ...                      if 

Heredity    . 
Heritiera  Fomes 
Hermaphrodite  . 
Hetercecious 
Heterophyllous  . 
Heterophylly 

156 
248 
45 
198 
27 
.       28,  33 

Heterostyled 

112 

,,    side  of  flower     .         . 

'.             46 

Heterostyly        . 

112 

Inflorescence 

37-41 

Hexamerous 

45 

mixed 

4.0 

Hibiscus  Bosa-sinensis 
„        tiliaceus 

44 
248 

„           types  of           . 
Infructescence    . 

• 

.       37-40 
69 

Hill-Forests 

.     224,244-247       Infundibuliform.         .        '. 

50 

INDEX. 


PAGE,     i 

PAGE. 

Innate         .... 

52    {    Lamella,  middle  — 

.     72,  192 

Insect-pollination 

.     42,  111        Lamina      .... 

20  49 

Insects         .   42,  111,  112,  115, 

156,  182,227       Lanceolate     ... 

23 

Integuments 

54    ,    Lantana  aciileala 

.   178,228 

Intercellular-spaces       .        73, 

74,84,91,99    |    Larches      . 

145 

Intermediate  forms     .          4, 

123,125,155    !    Larix  Grifflthii    . 

246 

,,               host 

198    j    Latent  character 

157 

Internal  bark 

18,  80    j    Lateral  branching 

12 

In  tern  odes 

9    j          „      nerves    . 

21 

„         in  the  flower 

44/47,48             „       plane      . 

46 

Inter  petiolar  stipule     . 

.      "31,  34             „      roots 

.     5,6,82 

Interruptedly-  pinnate 

25 

,,      section    . 

46 

Intramarginal  vein 

22 

Laterite  (soil) 

.  220,  241 

Introrse      .... 

52 

Latex         ... 

74 

Investigation,  of  diseases 

.   182,  202 

La  ticiferous-  tubes 

74 

Involucel  .... 

38 

Lax  inflorescence 

37 

Involucre  .... 

38,  39,  44 

Lavers 

172 

Involute    .... 

35       Leaf-apex 

24 

,,     ,  valvate  — 

51 

,,    base  .... 

24  30 

Iron            .          .     18,93,94, 

102,  206,  220 

„    blade 

20 

Ironwood  of  Pegu 

240 

^compound 

23 

Irregular  flower 

46 

,    margin 

.       22,  23 

Irritability 

68,  85,  105 

,    mosaic 

27 

Isostemonous 

52 

,    scars 

16,  20,  35 

Ivy  .         .         .         .         . 

.8,27,245 

,    season  for  coming  into  . 

7 

Ixora         .... 

250 

,    segments    . 

22 

,    sheath 

30 

,    shedding     . 

82 

,    simple 

23 

J 

,    stalk 

20 

Leaflet       .... 

23 

Jackals      .... 

.   116,227 

„     distinguished  from  leaf 

26 

Jamun       .... 

7 

Leaves       .         .     3,  4,  9,  13,  20-36,  46,  153 

Junipers    .... 

145 

„      arrangement  of 

9,32 

Juniperus] 

191 

,,J    [colour  of 

.       28,  29 

„         communis    . 

246 

„      development  of 

82 

„        macropoda  . 

246 

„      metamorphosed 

13,  31,  34 

„         recur  va        .         . 

.  245,  246    j          ,      movements  of 

107 

,          WallicJiiana 

246 

,      of  young  plants 

28 

,       parts  of  flower  are     . 

46 

,       polymorphic    . 

27 

K 

,       position  of 

9 

,       protection  of  young 

.       35,  36 

Kanddia   .... 

248 

,       shape  of 

.       23,  32 

Kans          .... 

242 

,      smell  of  . 

29 

Karshu  Oak 

245 

,      structure  of 

84 

Katabolism 

90 

,       taste  of  . 

29 

Keel  

56 

Leea  aspera 

67 

Kernel,  of  seed   . 

58 

Legitimate  cross 

112 

Khair         .           16,  25,  208,  219,  220,  224, 

Legume      .... 

60 

239,  240,  219 

Legummosae 

222 

Kinds  of  stems   . 

10 

„          and  bacteria 

203 

„      „  tissues 

73 

Leguminous  plants,  absorption 

of 

Kingdom,  Animal 

1 

nitrogen  by     . 

203 

„        Vegetable    . 

i 

Lenticels     .          .          .           16,81,186,248 

„            „           subdivisions  of  126-147 

Leontopodium  alpinum 

245 

Knot?,  in  wood  . 

168 

Lepidote    .... 

65 

Kulu          .... 

232 

Leptadenia  reliculala  . 

41 

Kydia  calycina   . 

239 

Liane         .... 

.     11,  241 

Lichens      .      137,  186,  203,  209 

,  218,  219, 

232,  233 

L 

Lichi          .... 

58 

Licuala      .... 

243 

Lagtrstrcemia  Flos-Eegines    . 

.  219,243 

Life,  duration  of  plant  — 

17<i 

„           jtarviftora        . 

.     29,  238 

„    plant  —  ,  conditions  of  . 

85 

INDEX. 


Light  absorbed  by  chlorophyll 
„     demanders          .          .       85 
„     growth  retarded  by     . 
,,     injury  by  . 

„      its  effect  on  plant-distribu- 
tion 

„     its    effect    on    plant-move- 
ments . 

„     required  for  assimilation 
„     variations  caused  by  . 

Lightning,  injury  by    . 

Lignified     .... 

Lignin         .... 

Ligulate      .          . 

Ligule         .... 

Liguliform 

Lily,  Water 

Limb,  calyx 
„       corolla 
„       perianth  . 

Lime  .... 

Linear         .          .          .      .    . 

Linnaeus,  sexual  system  of    . 

Linseed       .... 

Litsaea  angustifolia 

Littoral  Forests  . 

Liverworts 

Livistona   .... 

Lobed         .... 

Lobes,  leaf — 

Loculicidal 

Lodicule    .... 

Lomentum 

Longitudinal  section    . 

Lonicera    .... 

Loose  inflorescence 

Loranthaceae 

Loranthus 

,,  longiflorus  . 

„  pulvsmlenius 

Lower  side  of  flower 

Lumnitzera 

Lycopersicum  esculentum 

Lyrately-pinnate 


M 

Machilus   .... 

Macrosporangia 

Macrospores 

Magnesium 

Magnolia  Campbellii   . 

Mahua       .  .    7,  06,  132 

Maize          .... 

„     smut  of     . 
Male  flower 
Malformations    . 
Mallotus  philippinensis 
Maltose       .... 
Man,  action  of,  effect  on  plant-dis- 
tribution 

„    action  of,  effect  on  type  of 
Vegetation       . 


PAGE. 

PAGE. 

102 

Man,  action  of,  is  limited      .         .           234 

>,  214,  224 

Mangifera  indica         .         .         .           243 

.  104,  105 

Mango        .          17,  29,  61,    63,  133,  174,  185 

.  214,  215 

Mangrove            ....       8,  247 

Mangrove-forests          .          .          .            247 

224,  226 

Manufacture,    of     organic    -food- 

. 

materials          .          .                    .         3,  89 

107 

Manures,  plant  ....              95 

101,  102 

Maples       ....     21  \5,  222,  244 

153 

Marcescent          ....              63 

216 

Margin,  leaf  —     ....              22 

72 

Marginal  vein     ....             22 

72 

Maximum  intensity     ...             86 

50 

Mealy        .....             64 

.       30,50 

Median  plane      ....             46 

50 

,,     section  ....             46 

47 

Medullary  rays  .          .          .17,  77,  98,  166 

43 

Meliaceae  222 

43 

Meliola      133,  186 

43 

Meliosma  pungens       ,         .         .           244 

74 

Members,  plant  ....                2 

23 

„            ,,     development    and 

118 

structure  of       .              77 

63 

,,           „     duration  of            .             63 

133 

„           ,,     texture  of     .          .             63 

.    247-249 

„        secondary,  development 

.  137,  147 

of            ...             82 

243 

Membranous       ....             63 

22-23,  44 

Memecylon                    .         .         .           243 

.       23,  24 

Meristematic  tissue      .          .          .73,  76 

60 

Merulius  lacrymans     .         .         .           134 

56 

Mesocarp    .....       60,  61 

60 

Mesua  jerrea       .         .          .          .           243 

46 

Metabolism          .          .          .          .       89,  90 

244 

Metamorphosed  leaves           .           13,  31,  34 

37 

„              stems           .          .             13 

240 

Metamorphosis  ....             36 

.   116,227 

Microbes    .          .          .          .          .           127 

201 

Micropyle            ....              54 

64 

Microsporangia  ....   142,  143 

46 

Microspores        ....   142,  143 

248 

Middle-lamella    .          .         .          .     72,  192 

187 

Midrib        21,22 

25 

Millettia  auriculata      ...             30 

Mimosa  pudica  .          .         .68,  106,  108 

Mineral  crystals           ...             71 

Mineral  salts,  essential          .    205,  219,  221 

Minimum  intensity      ...             86 

24$ 

Mixed  crops        ....             95 

142,  143 

,,     inflorescence    ...             40 

142,  143 

Mobile  leaflets    .          .  ,       .          .           229 

93 

Moisture,  effect  of,  on  type  of  vege- 

247 

tation         .          .          .   231,  233 

2   211,  239 

,,        excessive      .          .          .           215 

8,118 

Monadelphous    ....             52 

134 

Monandrous        ....              45 

45 

Moniliform          ....              62 

156 

Monochlamydeous     ...              43 

225 

Monocotyledons  .          .    59,  78,  79,  145,  147 

103 

„             root  of         .         .       79,  81 

s- 

„            stem  of                             78 

228 

Monoecious           .         .         .         .  45,  111 

->f 

Monopodial  branching          .          12,  37,  38 

234 

„          inflorescence     .         .            38 

Xll                                                            IN] 
PAGE. 

Monopodium       .         .         .         .            12 
Monstrosities      ....           156 
Moringa  pterygosperma        .         .             26 
Morphology,  Vegetable         .         .           2,  5 
Moru  oak  .....           244 
Morus        227 
Mosaic,  leaf  —     ....             27 
Mosses,      .  137-139,  147,  209,  218,  232,  248 
„    ,club—    ....           141 
„      life  history  of  .         .         .           139 
Moulds        129 
Movement,  of  protoplasm     .           68,  69,  85 
Movements,  plant       .         .         .   105-108 
sleep         ...           107 
Mucronate           ....             24 
Multiple  fruit      ....             59 
Musci        137-139 
Mushrooms         ....  129,  133 
Mussaenda          ....            47 
Mutants     .....           155 
Mutations           .         .         .         .  155,  160 

DEX. 

PAGE. 
Obcordate.          ...                       <>5 

Obdiplostemonous       ...            52 
Oblanceolate       ....             25 
Oblique-leaf        .                             .22 
„        plane     .....             46 
„       section  ....            46 
Oblong       24 

Obovate    25 

Obtuse       ....                       24 

Occlusion,  of  wounds  .         .         .    165-167 
Ocroa         31 

Odd-pinnate       ....             25 
OdinaWodier     .         .           16,29,239,244 
Oenothcra  Lamarckiana       .         .            155 
Oil-glands           .          .         .         .       29  74 

Oil?  29  71,  74 

Olca  slan<?  ulifera          .          .          .           244 
Oniou         .....             11 
Oophvte    .          ,          .        139,  141-145,  147 
Ousphere  .          .          .       126,  138,  141,  143 
Oospore     ....     138,  141,  143 
Open  vascular  bundle           .         .             76 
Operculum          .                             .           139 
Opposite  leave?  ....               9 
Optimum  intensity      .          .          .86,  208 
Opuniia  Dillenii.         .         .         .             12 
Orange       ....  29,  30,  74,  133 
Orbicular  ....                       24 

Mutilation           .         .         .         .           156 
Mycelium  .         .         .         .         .           130 
Mycorhizas         .         .         .     203-204,226 
Myrisiica  ....                    243 

N 

Naked  budi,        ....            33 
Napiforru  root    ....              7 
Natural  classification  .      118,119,122-125 
„       grafting          .         .         .         .175 
„       groups  ....  122,  123 
„       order     ....           122 
„       selection         .                   .           157 
Nectar       .         .         .         42,49,111,114 
Nectary     49,  114 
Negatively  geotropic  .         .         .           106 
„         heliotropic         .         .           107 
Nelumbium  speciosum          .         .            25 
Nerves       .....             20 
Nipa  iruticans    ....           248 
Nitrates     ...        94,  129,  203,  204 
Nitrifying  bacteria       .          .          .           129 
Nitrogen    .  68,  70,  93,  94,  95,  129,  203,  207 
Nodes        9 
Nomenclature     .         .         .         .           121 
Non-essential  substances      .         .       95,  96 
Normal  roots      ....              5 
Nucellus    .....            54 
Nucleus     .         .         .         .          69,  70,  71 
Number,  of  parts  of  flower  .         .            44 
Nut  .                                                           61 
Nutlet          60 
Nutrition.         .         .    68,85,89,104,153 
Nymphaea          ....            47 
Nyssa        246 

0 

Oak           .    6,  17,  58-60,  118,  203,  222,  224 
Oats,  smut  of                       .                    134 

Orchids      ....                     241 

„       roots  of           ...               8 
Order,  Natural  .          .          .          .           122 
Organic  material         .         .          .     178-179 
,,          „       manufacture  of    .        3,  89 
Organised  ferments      .          .         .           132 
„         plant,  highly        .          .         2,  91 
„         substance    ...             87 
Organisms,f  actors  influencing  rela- 
tions between       .          .           178 
Organs,  plant      .          .          .         .           2,  3 
,,           „      functions  of  .          .               3 
»,           „       reproductive           .               4 
»,           ,,      sexual  ...             45 
„           „      vegetative     .         .               4 
Origin,  of  species          .          .          .    148-161 
Ornithophilous   .          .          .          .           HI 
Orobanche  indic.a          .         .         .     11,200 
Oroxylum  indicum        .         .28,58,59,116 
Orthotropous      ....             54 
Osmosis     ....                 86-87 

Osseous     .....             63 
Ougeinia  dalbergioides    14,  37,  236,  240,  244 
Oval           ....                       24 

Ovary        ....        41,54,145 
Ovate         24 

Overlap 

Ovoid        ....                       61 

Ovule        41,  143 
„      characters  of     .         .          .        54,55 
Oxalate,  calcium          .         .         .             71 
Oxalic  acid          ....             93 
Oxygen     .   68,  70,  85,  93,  94,  101,  103,  205- 
207,  220,  247,  249 
Oxytenanthera  albociliata      .         .          241 

INDEX.                                          xiii 

p 

PAGE. 

PAGE. 

241 

Perfoliate  25 

Palas          ... 

239 

Perianth    .         .         .         .41,  43,  44,  143 

Palea 

66 

„      limb     ....            43 

Paleaceous 

63 

„      throat  ....             43 

Palisade  tissue    . 

84 

»      tube              ...            43 

Palmate  lf»af 

OK 

Perioarp                            ,           _          <               59 

Palmately-cleft  .         . 
fejp-             divided 

.                .                      *-'J 

23 
23 

Periderm  .         .         .         .         ,      80  81 

Peridermium  brevius  .         .         .           136 

lobed 

23 

„            complanatum           .           136 

,                   parted 

23 

Peridium             .          .         .         .           197 

trifoliolate 

26 

Perigynous         .          .         .         .48,49 

veined 

21 

Perisperm           .          .         .          .58,  143 

Palmatifid 

23 

Peristome           .         .          .         .           139 

Palmatisect        .         . 

23 

Perithecium        ....  131    133 

Pal  minerved 

21 

Permanent  tissue        ...             73 

Palms        .     11,  13.  14, 

18,  38,  59,  65,  78, 

Persistent                              «.                       63 

145,  176,  222,  243 

Petaloid  perianth       .                    .             43 

„     Sago 

.  144,  147 

Petals        .         .         .                   43,44,47 

Pandanus 

.       S,  222 

„     descriptive  terms  for            .             49 

„        lecforius 

249 

Petiole       .         .     20,29-31,33,34,35,37 

Panicle 

40 

Petiolule             .          .          .          .      23  37 

,,       corymbiform 

40 

Phanerogams     .         126,  142-147,  183,  200 

„      cymose  . 

40 

Phellem     80 

„      racemiform 

40 

Phelloderm         ....             80 

„      umbelliform    . 

40 

Phellogen  80  81 

Papilionaceous   .         . 

56 

Phloem       ....                76-79 

Pappose    . 

50 

Phlogacanthus  thyrsi  fiorus     »         .             41 

Pappus 

50 

Phoebe        .         .         .         .         .            246 

Parallel  -venation 

20 

Phcenix      16,  28 

Parasites   . 

180,185,  186,227 

Phcfnix  acaulis  ....               10 

„        wound  — 

194 

,,       humilis            .         .         .           239 

Parasitic  Phanerogams 

.    200-202 

„       paiudosa         .         .         .            248 

Parasitism  of  Santalum  album       .           202 

Phosphates         .         .         .         .  '     94,  204 

Parenchyma 

,       73,  75 

Phosphorus        .          68,  70,  93,  95,  183,  207 

„            spongy  . 

84 

Phycomycetes      ....    131,  187 

Parietal     . 

53 

Phyllanthus  Emblica  .   15,  26,  67,  208,  209, 

Pari-pinnate       .         . 

25 

239 

Parted  leaf 

.       22,  23 

Phyllotaxis         ....             32 

Partite    „           . 

92 

Phyllotaxy          ....              32 
Physiology,  Vegetable          .         .2,86-116 
Phytophthora  infestans          .         .           131 
„                  „        life  history 

Parts,  of  flower  ....            41 
„       „     „      are  leaves     .         .            46 
»»  '    »    »»      position  and  nuni- 

"L.             £                                                                                                     *  * 

ber  of 

44 

of            .    187-190 

Pastor  roseus 

227 

,,            omnivora         .         .    190-191 

Pathology,  Vegetable  . 

2,3 

Picea         191 

Pea  . 

.     31,  203 

„     Morinda              33,  135,  184,  244,  246 

„    flowers  of  . 
Pear. 
Pedate  leaf 

56 
136 
26 

Picrasma  quassioides   ...             25 
Pieris  ovalifolia  ....           244 
Pileus        .....           133 

Pedately-com  pound    . 

26 

Pilose         .....             64 

„       nerved          . 
Pedicel      .         .         . 

26 
37 

Pine  (Pinus)       .   12,  13,  33,  56,  58,  67,  111, 
145   191   203 

Pedicellate 
Peduncle   . 
Pedunculate 

37 
37 
37 

„    Blue  .         .         .       183,  224,  225,  244 
„    flowers  of    ....             58 

Pinnae         25 

Pendulous  branches    . 
„         inflorescence 

'.         \            65 
37 

K.A 

Pinnate  leaf        .          .         ...             25 
Pinnately-cleft  ....             23 
—  divided    ...             23 

,,         ovule          . 
Penninerved       .         . 
Pcniacme  suavis 
Pentamerous 
Percolation         . 

•            *                u4 
21 
241 
45 
219 

—lobed       ...             23 
—  parted     ...             23 
—trifoliolate        .         .             26 
—  yeined      ...             21 

Perennial  .         .         . 

66,67,80,  176 

Pinnatifid            ....            23 

XIV 


INDEX. 


Pinnatiaect 
Pinnules  . 
Pinus  excelsa 

„  Gerardiana 
„  Khasya  . 
„  longi folia 


PAGE. 

23 
26 

134,  136,  194,  224,  2^7, 

244,  246 

246 

246 

15,  18,  136,  220,  224,  244, 
246 
246 

.  232,  233 
185 
63 

41-43,  47  53,  54 
.       53, 54 


„      Merkusii 
Pioneer  plants    . 
Pipal 
Pips 
Pistil 

Pistil,  characters  of 

Pistillate  flower  ...  45 

Pith  ....  16,  17,  77 

Pits  ......  71 

Placenta    .....  53 

Placentation       ....  53 

Plaited       .....  51 

Plane  (Platanus)          .          .          .     34,  216 
,,     lateral       ....  46 

„     median      ....  46 

„     oblique      ....  46 

Plano-convex      ....  62 

Plant-communities      .          .          .  233 

,,     crops,  succession  of    .          .  233 

„ —  distribution,  factors  affect- 

218 

3,20,89,91,93,  103, 
104,  178-180 


mg 
food-materials 


generations,  succession  of  . 
highly  organised 
life,  conditions  of 

,,    duration  of 
members,  development  and 

structure  of  . 
members,  duration  of 
members,  texture  of  . 
movements 


233 

2,91 

85 

176 


77 
63 
63 
105-108 

Plants 1,2 

deciduous        .        67, 100,  101, 229 
„      dependent  on  other  plants  225,  226 
„      effect  on  plant-distribu- 
tion ...  225 
„      evergreen        .         67,  100,  101, 229 
„      functions  of    .          .2,  3, 85,  86 
„      green      .                   .      178-180,204 
„      gregarious       ...  67 
„      higher,  functions  of          .  93 
„      pioneer  .                             .    232,  233 
„      sporadic           ...  67 
„      their  influence  on  plant- 
development         .          .            183 
Platanus     .          .          .          .        ..34,216 

Plicate 35,51 

Plumule 59 

Pneumatophores  .          .         .  248 

Pogostemon  plectranthoides   .         .  29 

Poisonous  substances  93,205,206,  216,225 
„  „          excretion  of 

by  plants  226 

Polar  Zone  ...          222 


Pollard- shoots    . 
Pollen 

,,     grains 
„     sacs 
„     tube 

Pollination         .  42,  111 

Pollinia 
Poly-adelphous  . 

,,    androus 
Polygamous 
Polygonum    > 
Polymorphic  leaves    . 
„  plants    . 

Poly-petalous 
,,    phyllous     . 
,,    sepalous 
Pongamia  glabra 
Poplar       .... 
Poppy        .  . 

Populus  ciliata  . 

,,        euphratica     . 
Pores          .... 

,,     water 
Position,  of  leaves  and  buds. 

,,        of  parts  of  flower 
Positively  geotropic     . 
,,         heliotropic  . 
,,         hydrotropic 
Posterior  side  of  flower 
Potassium.          .         .     93-95, 
Potato        .... 

„      disease    . 
Premna  latifolia 
Prepotent  pollen 
Preservation,  of  specimens  . 
Pressure,  root 

Prickles      .... 
Prickly  Pear 
Primary  nerve    . 

„       root 
Primula 

Prinsepia  utilis  . 
Procumbent  stems 
Products,  waste 
Promycelium 
Prosopis  spicigera 
Prostrate  stems 
Protandrous 

Protection,  of  young  leaves 
Proteid  grains 
Prothallium 
Protogynous 
Protonema 
Protophloem 
Protoplasm          .         .  1, 

„         continuity  of 
Protoxylem 
Pruinose    . 
Pruning     . 
Psoralea  corylifolia 
Pteridophyta         .          .        126, 
Pleris  aquilitta     ,         ,         t 


PAGE. 


171 

41,51,53,111 

.      53,  143 

.      51,  143 

.      55, 143 

,115,182,22? 

53 

52 

45 

.  46, 111 
31 
27 

.  128,  135 
43 
43 
43 

17,  249,  250 

.  222,226 

45 

244 

238 

17,  18,  74 

100 

9 

44 
105 
107 
108 
46 

183, 204,  207 

11,  109,  208 

131,  187-190 

29 

.   115,  151 

103 

96,  97 

63 

12 

21 

5 

112 

16,  34 

10 

.       69,  90 
196 

.  219,237 

10,  65 

.   Ill,  113 

.       35, 36 

.     70, 205 

140, 142,  143 

111 

138 

76 

68-70,  85-90 
72 

.      76, 79 
64 

.'    167,  168 
66 

139-142,  147 
11,140 


INDEX.                                                         XV 

PAGE. 

PAGE. 

Pterocarpus  dalbergioides 

241 

Racemose  inflorescence         .          .       38,  39 

„           macrocarpus 

241 

Radial  vascular  bundles       .          .             79 

„          Marsupium 

240 

Radicle      .....             58 

,,          santalinus 

.  224,240 

,,       inferior  .     <     .          .          .             60 

Pterospermum 

243 

,,       superior           ...             60 

„            acerifolium 

240 

Radish       .....               7 

Puberulous 

64 

Rainfall     .          .          .       218^237,  238,  241 

Pubescent 
Puccinia  graminis 

64 
135 

,,        its  effect^on  type  of  vege- 
tation "                 .          .           231 

„       life  history  of 
Pulvinule  . 

.    195-199 
.     30,  108 

„       its  effect  on  type  of  forest          237 
Rain-water          .          .          .          .            221 

Pulvinus    . 

.     30,  108 

Rakhs        ...                    .           237 

Pure  forests 
Putamen    . 

178 
61 

Rambling  plants          .         .          .             11 
Ramenta  .....           140 

Putrefaction 

.     129,  179,  204 

Randia  dumetorum       .         .       '  .             13 

Pyrausta  machceralis    . 

>          178 

Raphe        .....             55 

Pyrene 

61 

Raw  food-materials     ...               3 

Pyriform    . 
Pyrus  lanala 

62 
64 

Rays,  medullary  —     .          .17,77,98,  166 
,,      of  umbel  ....             38 

„     Pashia 
Pythium    . 

.  13,  16,  28,  136 
131 

Receptacle,  of  flower  .          .          42,  49,  60 
„         of  inflorescence            .             39 

Reciprocal  symbiosis,  .          .          .           180 

Reclining  stems            ...              10 

Red  Sanders       .          .      224,  238,  240,  250 

Quadrangular  stems    . 
Quadri-foliolate 

14 

26 
51 

Regular  flower    ...          .          .             46 
Regulation  of  transpiration            .      99-101 
Regursoil           ....           220 

,,       locular    .          . 
Quercus 
,,        Brandisiana   . 
„       calathiformis  . 
,,        dealbata. 
,,       dilatata 
„       eumorpha 

.   118,  121 

246 
246 
246 
184,  244  245,  246 
246 

Reinwardtia  trigyna     .          .          .     63,  1  12 
Relations,  between  organisms        .           178 
Reniform  .....             24 
Repand      22 
Repent  stems      ....             10 
Reproduction      .    68,  85,  108-110,  156,  158, 

,,       jenestrata 
,,       glauca    . 
„        Griffithii 
,,        Helferiana 

246 
.  244,  246 
246 
246 

Reserve  materials        .         .        97,  169,  171 
Resin          ....        71,  168,  194 
Resin-canals    ") 
ducts      I             •   .     •         18,74,145 

llfr 

246 

,  .           U.UL/LS          j 

,  ,               L  (OX                •                   • 

,,        incana   . 

7,  21,  29,  244,  246 
246 

Respiration         .     68,  85,  89,  103,  104,  179 
Response  to  a  stimulus          .         .             68 

„       lanuginosa 
,      Lindleyana 
,       lineata   . 

246 
246 
246 

Resting  spores    .          .       126,  128,  190,  196 
Reticulate  venation     .          .          .             20 
„         vessels        ...            73 

,        mespilifolia     . 
Olla 

246 
246 

Retuse       .....              24 
Revolute   .....             35 

,        pachyphylla    . 
,       polystachya     . 
,       semecarpifoha 

246 
246 
17,  28,  184,  212, 

Rhabdia  lycioides          .          .          .219,  250 
Rhachis     25,  37 
Rhamnus  .                                          222,  244 

•             .                                                                                            I  c> 

245,  246 

,,         virgatus 

„        semiserrata 

246 

Rhizoids    138 

,,       serrata  .         . 

246 

Rhizome    11,66 

„       spicata  . 

246 

Rhizomorphs      .          .       130,  133,  134,  193 

,,       truncata 

246 

Rhizophora         .                            .       8,248 

,,       xylocarpa 

246 

,,         mucronata           .         .  247,  248 

Quisqualis  indica 

11 

Rhododendron  Anthopogon    .         .           245 
arboreum        .      136,244-245 

,,             campanulatum    136,  212,  245 

B 

„             lepidotum       .         .           245 

Rhus  Cotinus      ....           244 

.  121,  123 

,,    vernicifera           .         .         .           122 

,,    importance  of     . 

149 
38 

„     Wattichii  ....           122 
Rhytidome          .          .          .          .81,  242 

Raceme      ... 
Racemiform 

40 

Ribs           20 

XVI 


INDEX. 


PAGE. 

Ringing 168-170 

Rings,  annual     .          .         .         .       17, 78 

Riparian  Forests          .         .  .  249-250 

Root          .         3-8,11,79,81,107,109,185 

„     aerating      ....  248 

„    cap 82 

„    contraction  of.         .         .  83 

„  development  of  .  .  .  6, 82 
„  fusions  ....  170,  175 
„  hairs  ....  83 

„         „     absorption  by    .     86, 93, 94, 96 
206,  207,  211 

„  pressure  .  .  .  .  96, 97 
„  shortening  of  .  .  83 

„    stock          ....  66 

„    structure  of         ...  79 

„    suckers        .         .          7,109,171,236 
Root  system       ....  5 

„  „  development  of  .  6, 7, 82 
„  „  types  of  .  .  .  5, 6 
„  tubercles  ....  129,203 
„  wood  of  ....  79 

Ro&a 31,244 

„    moschala    ....  43 

Bosaceae  .....  222 
Rosdlinia  bunodes  .  .  .  133 

Roses         ....          47,64,65 

Rotate 50 

Rotation  of  crops  .  .  .95, 226 
Rotting  .  .  .  .  129,  179,  204 
Roughness  of  bark  .  .  .  15 

Rubus 64,244 

„     lasiocarjMs        8,  11,  15,  64,  83,  109 
Rudiments         ....  56 

Rugose 64 

Ruminate  albumen 

Runcinate.         .         .         .         .  22 

Runner      .         .         .         ...     11,  109 

Rust- fungi          .         .         .         .  135 

Ruats         ....     129,  135,  195 


S 


Saccate      .         . 
Sacchar^mycetes 
Sagittate   .         . 
„        leaf     . 

. 

50 
131 

24 
25 
144,  147 

Sam           .         . 
Sai    . 

.     117,  220, 

224,  238 
220,  238 

Sal    .  6,7,15,60,63,67,116.117,133,210, 
220-221,  224,  225,  234,  238,   240, 
244,  250 
„    Forests       210,  211,  221,  22o,  234,  240 

Salai 228,232 

Saline  soil  .         .         .    206,220,229 

Salix         .         .  244 

„    bahylonif,a  ...  65 

„    daphnoidea          .         .         .  15 

„    ietrasperma         .         .         .          250 

Salt 206 

Salts,  excess  of   .         .  '      .         .205,  247 

Salvadora       .  ...          237 

pertica        ...          238 


PAGE. 

Salver-shaped     ....  50 

Salvia  lanata      ....    112-113 

„     fertilisation  of  flowers  of     .  112 

Samara 61 

Sand 220 

Sandal       .  18,202,227,238,240,250 

Sandan 240 

Sandy  soils          ....  94 

Santalum  album  .         .         .          240 

„         „      parasitism  of        .          202 

Sap,  cell — 69 

„     rising  of  .         .  96 

Sapium  sebiferum        ...  29 

Saprophytes       .         .         .     179,  191,  193 

Sap-wood 17,  205 

Sarcocarp  ....  61 

Sarmentose         .         .         .         .  11 

Satin  Wood        ....  220,240 
Savannah  ....  231 

Saxifraga  ligulata         .          .    •  63 

Scabrous 64 

Scaevola 249 

Scalariform  vessels      ...  74 

Scale-insects       ....   133.  181 

„    like  leaves.         .         .         .       11,  12 

Scales 63,65 

„      bud—     .         .         .         .       33,34 

„      of  bark  ...  16 

Scaly  buds          ....  33 

Scandent  stems  .         .         .         .  10 

Scape         .....  11 

Scaiious     .....  63 

Scars,  leaf  .          .          .  16,20,35 

„      of  bud  scales         ...  33 

.,      of  stipules          .         .         .  :?1 

Scattered  leaves.         ...  9 

Scent,  of  flowers  .         .         .43,111 

,,     of  wood    ....  18 

Schizocarp          ....  60 

Schizomycelf.8      .         .          .         .  128 

SrMeichera  trijuga       .         .         .  133, 239 

Schrebera  su-ietenioides          .         .          239 

Scion 174 

Sclereuchyma     ...         .        73, 75,  101 
Scorpioid  cyme  ....  40 

Screw  Pines        .         .         .         8,222,243 
Scrophulariace.ae,          .         .         .  187 

Seaweeds 127 

Secondary  growth        .          .          .77, 79 
„        members,  development 

of  ....  82 

21 
5 

123 
46 
46 
46 
46 
46 
63 

59,  109 
28 
190-191 


„         nervros 
„         roots  . 

Section,     . 

„        lateral  . 
„       longitudinal   . 
„       median 
„        oblique  . 

„       transverse 
Sedum  rosulalum         . 
Seedlings 

leaves  of 
damping  off  of 


Seeds         .     3,  42,  54,  55,  58,  59,  70,  109, 

126, 143 


INDEX. 


XVH 


PAGE. 

Seeds,  descriptive  terms  tor  shape  of         61 
„     distribution  of           .     58,  109,  115, 
182,  219,  227 
„     germinating      .         .         .           103 
Segments  leaf              -         •          •      22,  23 

PAGE, 

Sinuate      22 
Sissoo           6,  7,  25,  116,  219,  224,  234,  238, 
240,  249 

„     Forests      .                   ,         .   233,  240 
Skimmia  Laureola                                       29 
Sleep-movements                   -      —  -»—         107 
Smell,  of  leaves                      .                        29 
Smilax       .                             .                      145 
„       parvifolia                   .         .             21 
Smut-fungi          .                    .                     134 
Snow,  damage  by                  •.          .           213 
Soil,  absorption  by                                       94 
„    calcareous  .                    .          .  220,  24  5 
„    cotton         .                   .         .           220 
„    depth  of      .                                       219 
„    effect  of  vegetation  on          .221,  234 
„    formation  of         ...           232 
„    its  effect  on  plant  develop- 
ment       ....    205-208 
„    its  effect  on  plant  distribu- 
tion                                         219-221 

Selection'            ....    157-160 
„        natural        .         .         .           157 
Selective  power  of  protoplasm       .             87 
Self-fertilised      .         .         -         .           HO 
Kemecarfiui  Anacardium       . 
Semi-amplexicaul         .          » 
Sepaloid  perianth        ...             43 
Sepals       ....      43,45,47,49 
„     descriptive  terms  for 
Septate  hyphse   .         .         .         .           130 
Septiddal            ....              60 
Sept.fragat          . 

Serrate       ..... 

Sesbania  aculeata         ...            66 
Sessile                           20,  37,  42,  51,  51,  59 
Seta            139 

„    its  effect  on  the  type  of  vege- 
tation     ....            232 
„    laterite        ....    220,241 
„    moisture  in            206-208,220-221,233 
„    oxygen  in    .          ,       206-208,220,249 
„    regur           ....           220 
„    saline          .          .          .     206,  220,  229 
„    temperature  of    .       86,  100,  208,  211, 
212,  223,  232 

„    variations  caused  by     .         .           153 
,,    water-logged        .          .          .           220 
„    well-drained         .         .         .  207,220 
Solanaeeae           ....           187 
Solanum  Melongena     .         .         .           153 
„      tuberosum       .         .         .           187 
„      verbascifolium          .         .             29 
Solitary  flowers  .                             .37 
Sonneratia           .... 
Sorus           140 
Spaces,  intercellular     .       73,  74,  84,  91,  9S 
Spadix                           • 
Spathe        

Sewage,  disposal  of                .        ,.           208 
Sexes,  separation  of     . 
Sexual  generation        .          .          .           139 
Sexual  organs,  distribution  of        .             45 
„      reproduction    .          .     108,  156,  158 
„      system  of  classification       .           118 
Shade-bearers               .         .          .     85,224 
Shape,  of  cells     .... 
fruit    .... 
„           leaves           .         .         .       23,  32 
„           root     .... 
„           seeds  .                                        61 
„           stems  .... 
Sheath,  leaf        ....             30 
Shedding  of  leave?       . 
Shield  (in  budding)      .         .         .           175 
Shining  leaves    .... 
„       stems     ....             15 
Shiaham    240 
Shoot                   .         .         .         .  3,  4,  6,  7 

„     development  of 
„      flowering 
„     pollard     .... 
„     stool         ....           171 
STiorea  obtusa      ....           241 
„       robu^.a     .         .         .          .117,210 
Shortening,  of  roots     . 
Shorthand,  floral          ...              57 
Shot-holes. 
Shrubby  plants 
Shrubs'      .                   ...      66,67 
oieve-nlates                   ... 
„     tubes                   .          .         74,91,169 
Silica 
Silky          .                   .                                64 
Silver  Fir  .                   .17,  65,  168,  225,  245 
„      grain                   .         • 
Simple,  fruit 
„      leaf                  ...             23 
„       pistil                 ...              53 
ginistrorse  twiners       ...              10 

Species        ....      117,119-125 
„      constancy  of   . 
„      distribution  of  important.            250 
hybrids             ...           150 
origin  of           .         .         .    148-161 
.I      sub          ....   120,123 
„         „   importance  of    . 
Specific  characters        .         •         •           120 
,,      name      . 
Specimens,  collection  of 
preservation  of    . 
Spectrum,  of  chlorophyll       .                     102 
Spermatia            •                   •         '100 
Spermatozoid      .                   •      »*y,  100,1  A 
Spherical    .                              • 
Spheroidal 

Spike          .                             '         '           909 
„  'disease       .                  •                    •'"'* 

Spikelet      .                             • 

XV111 


INDEX. 


PAGE. 

PAOE, 

Spines             10,  12,  13,  32,  34,  36,  64,  116, 

Straggling  plants         .         .                      11 

153,  229 

Strands,  leaf                 .         .         .             20 

Spiraea      .         . 

244 

Strawberry                   .         .         11,  60,  109 

Spiral  vessels 

73,76 

Streblus  asper              .         .         .           250 

Spirillum 

127 

Striate       .                  ...            14 

Spirits,  manufacture  of                            132 

Strigose     .                  ...            64 

Spirocheete 
Splitting-Fungi 

128 
128 

Striking,  of  cuttings    .         .          .           171 
Strobilanthes       .          .   50,67,176,182,184 

Spongy  parenchyma     . 

84 

„           Wattichii          .         .67,  184 

Spontaneous  variations                              155 

Strophiole           ....             58 

Sporadic     . 

67 

Structure,  of  leaves     ...             84 

Sporangia 

131,  140 

„          „  plant-members         .            77 

Spores 

126 

,,root        .         .         .             79 

Sporidia     . 

134,136,196 

„          „  stem       .         .          14,  16,  77 

Sporogonium 

138 

„          „  vascular  bundles      .            75 

Sporophore 

.   133,  192,  194 

Struggle  for  existence         96,  148,  149,  159, 

Sporophyll  . 

140-143,  147 

177,  178 

„        female. 

.  142,  143 

Style          42,54 

>,        male 

.  142,  143 

Sub-  Arctic  Zone          ...           222 

Sporophyte 

139 

class  122 

Sports 

Iffl 

coriaceous                                              63 

Spruce    .  33,  65,  135,  145,  168,  225,  244,  245 

divisions  of  Botany       .         .              2 

Spurred      ... 

*n 

Vf*orpi"fl  Vil  A   TCiti  or. 

Stamen      .         .          41,43,47,51-53,143 

jj                 jj      V  vgC  U£v  UIU     -IVIIJ.^- 

dom            .         .    126-147 

„      characters  of    . 

.       51,  52 

Kingdom     ....           122 

Staminate  flower 

45 

opposite  leaves     ...              9 

Staminode 

5) 

order            ....           122 

Staminodium 

5 

species         .         .         .         .    120,  123 

Standard 

56 

„      importance  of    .         .           149 

Staphylea  Emodi 

63 

terranean  roots     ...               8 

Starch       .    3,70,89-92,97,101.103,208, 

Tropical  Zone       ...           221 

„      grains 

.      70,  97 

Suberised  .....             72 

Stellate  hairs 

65 

Subtending  bract         ...             46 

Stem 

.      3,4,9-19,65 

leaf  ....               9 

kinds  of     . 

10 

Subulate    23 

of  Dicotyledons 

.      77,  78 

Succession    of    crops,    or    plant 

„  Gymnosperms 

.      77,  78 

generations     ....           233 

„  Monocotyledons      .         .      78,  79 

Succulent  .....             63 

„  Tree  Ferns 

140 

^     „         fruit  ....             61 

shape  of     . 

14 

Suckers,  see  Haustoria! 

structure  of        . 

.  14,  16,  77 

Suckers,  root      .         .           7,109,171,236 

Stephegyne  parvifolia 

15,31,34,239 

Sucking  force,  in  leaves        .         .             98 

Sterculia    . 

.       65,  241,  243 

Suffrutescent      ....             66 

„      urens    . 

.       15,  232,  239 

Suffruticose        ....             66 

Stere.ospe.rmum  suaveolens     .         .           239 

Sugar,  cane         .         .         .         .           103 

Sterigma   . 

196 

„     fruit         ....           103 

Sterile  flower 

46 

Sugarcane           .         .         .         .  5,  6,  131 

Sterility     . 

.     112,  123,  150 

,,           adventitious  roots  of.          5,  6 

Sterilize,  to 

128 

Sulcate      14 

Stigma 

42,  54,  145 

Sulphur     ....           68,  70,  93 

Stigmatic  surface 

42 

Sundri       248 

Stilt  roots 

8,  247 

Superficial  placentation        .                      64 

Stimulus    . 

68,  85,  106 

,,         root-system         .         .              6 

Stipe 

133 

Superior    48,49 

Stipel 

31 

„      radicle          ...             60 

Stipellate  . 

31 

,,      side  of  flower          .         .             46 

Stipitate    . 

59 

Suppression  of  floral  parts    .         .             55 

Stipulate   . 

31 

Suspended  ovule         '  .         .         .             54 

Stipules     . 

,       30,  34 

Survival  of  the  Fittest          .         .            157 

Stock 

.      66,  174 

Sutures,  of  carpel        .         .         . 

Stolons 

11,  109 

Swamp  Cypress            ...             67 

Stomata,     .         . 

.  48,99-101,247 

Swelling,  of  wood 

Stone,  of  fruit     . 

61 

Swintonia           ....           243 

Stool-shoots 

71 

Symhionts          -         .       180,202-204,226 

IN 

PAGE. 

Symbiosis           .         .        180,  182,  202-204 
„         between  Green  and 
Non-green  Plants                204 
„          distant       .         .             180,204 
Symmetry  of  flower    .         .                     46 
Symplocos           ...                       29 
Sympodial  branching  .         .                      12 
Sympodium        .         .         .                 12,  40 
Symptoms  of  disease  .         .             181,  182 
Syncarpous         ...                       53 
Syngenesious      ...                       52 
Synonym  ....                     122 
Synopsis  of  sub-divisions  of  Vege 
table  Kingdom         .         .                     146 
Systematic  botany      .         .                        2 
Systems,  of  classification      .                    118 
„       tissue  ...                       74 

T 
Tamarix    238,  250 

DEX.                                                      XIX 

PAGE. 

Tetramdes  midiflora    .         ,         .241,  243 
Tetramerous       ....             4,1 
Texture,  of  bark          ...             15 
„      of  plant  members  .         .            63 
Thallophytes       .         .          .             126,  146 
Thallus                ....           127 

Theca,  of  moss    .          .     —  ;  —     .           138 
Thecse                  ....            61 
Thespesia  populnea     .         .         .           248 
Thickening,  of  cell-wall         .•         .             71 
Thickness,  of  bark        .          .         .14,  15 
Thorns       .          .          .          .31,  153,  229 
Throat,  calyx     .... 
„        corolla  ....             43 
„        perianth         ...             43 
Thuja                 ....           191 

Thymus  Serpyttum      ...             29 
Thyrsus     40 
Tidal  Forests      ....    247-249 
Tides                            .         .         .  237,247 

Tinas         240 
Tissues      73 

Tissue-system     ....            74 
Toadstools          ....   129,  133 
Tobacco  plant    . 
Toddalia  aculeata                   .         •           153 
Tomato      187 
Tomentose 
Tongue,  of  layers         .         •         •           173 
Toon                            ...        6,  18 

Tannins     71,  78 

Tap-root    .         .         .         .         .           5,  6 
Taste  of  leaves   ....             29 
Taxodium  dislichum    ...             67 
Taxus  bacf.ata     ....          245 
Teak          .         6,  7,  14-18,  20,  21,  33,  37, 
168,  178,  183,  204,  221, 
224,  228,  238,  240,  243,    250 
„    Forests   178,  183,  221,  224,    240,   243 
Tectona  grandis  ....               6 
Tegmen     58 
Tegumentary  tissue    .         .         .74,  75 
Teleuto-spores    .          .          .     135,  136,  196 
Temperate  Zone,  cold           .         .           222 
„            „     warm         .         .           222 
Temperature      .         .         .         .85,  102 
„            effect  on  plant-move- 
ments                  .           107 
„             extremes  of    .          .  208,  222 
„             influence  on  growth            104 
„             its   effect  on   plant 
distribution      221-224,  237 
of  soil    .       86,100,208,211, 
212,  223,  232 
Tendrils    ...            10,  13,  31,  107 
Tendu       ... 
Terete       ...                  .14 
Terminal  buds    . 
„       flower           .                  .             37 
Terminalia  Arjuna      .                15,  220,  250 
„        belerica      .                  .     41,238 
Chebvla      .                  .      7,238 
tomentosa  .     117,  220,  227,  238, 
243,  244 
Terme,  descriptive,  for  petals  and 
sepals           .            49 
„             „           for    shape  of 
fruit  and  eeed              61 
Terrestrial  plants        .         .         -           185 
Tertiary  nerves  . 
Testa         
Tetradycamous           « 

Toothed                        .          .         .22,  44 

Torulose    62 
Torus         ....          42,49,71 
Torus,  of  pits      .... 
Tracheae    .                                                    73 
Tracheids  .         .         .        73,74,78,91,98 
Trametes  Pini     .... 
„     life-history  of         .  194,195 
„       radiciperda       .      .         •           191 
(see  also  Fames  annosus.) 
Transpiration     .  88,  98,  212,  215,  223,  229 
current           .         .     98,169 
„           regulation  of  .          .      99-101 
Transport,  of  substances  in  plant  91,  97,  98 
103 
Transverse  section 
Tree  Ferns          .          .           11,19,140,222 
Trees                   .         .         .66,67,225,231 
„     and  Grasses        .         .    184,224,226 
„       „    Strobilanthea 
Trewia  nudiflora          .         .      41,240,250 
Triandrous          . 
Triangular           ....             24 
Tribe          123 
Trichotomous  branching       .         •      12,  !•* 
cyme    . 
Tri-foliolate        ....            Jj 
Trimerous          .         •         •         F-ftriS 
Trimorphio  flowers      .         •         •  lw»  "*j! 
Tri-pinnate         . 
Triquetrous        .                                         ** 
Tristichous 
Tropical  Zone     . 

XX 


INDEX. 


PAGE. 

Tropophyte        ....  229,230 

Trumpet-shaped          ...  50 

Truncate  .          ....  24 

Tsuga  Brunoniana      „         .         -  246 

Tube,  calyx        .          .          .         .       43, 49 

„     corolla      ....  43 

„     perianth  ....  43 

Tuber 11,109 

Tuberculate        ....  64 

Tuberous  root    ...          7, 11,  109 

Tubular 50 

Tufts,  of  leaves  ....  13 

Turbinate  ....  62 

Turgescence        .         .         .87, 88,  96,  108 

Turgid 88 

Turnip        .         .         .         .  7,95,156 

Twigs         ....  14,65,80 

Twining  stems    .         .         .  10 

Twisted  aestivation      .  51 

Types,  of  flowers  which  may  cause 

difficulty       .         .         .  55 

of  forests          ...          237 
„     „         factors    respon- 
sible for  their  distribution .  237 
of  fruit    .         .         .         .60,61 
„  inflorescence          .         .        37-41 
,,  vegetation     .         .         .  '        231 
„            „           factors    in- 
fluencing their    distribu- 
tion             ...  231 
selection  of,  for  description        4,  27 


TT 


Umbel        .....  38 

Umbelliform       ....  40 

Umbellule            ....  38 

Undershrub        .         .         .         .  66,  «7 
Undulate  .         .                   . 

Unifoliolate  compound  leaf .         .  30 

Unil  ocular           .         .         .         .  51 

Unisexual           ....  45 

Unit  of  classification   .         .         .  119 

Upper  side  of  flower    ...  46 

Urceolate  .....  50 
Uredinaceae        .         .         .      131,135,195 

Uredospores       .         .         .         .  195 

Urn-shaped        .  50 

Urticacece 222 

Ustilaginaceae     ....  131,  134 

Ustilago  Maydis.         .         .         .  135 

Utricularia         .         .         .         .  241 


Vaeuoles    . 
Valeriana  Wallichii     . 
Valvate 

,,        induplicate   . 

,.        involute 
Variability,  fluctuating 
Variation  , 


69 
50 
51 
51 
51 

152-1S5,  157,  160 
110,  152-156,  229 


PAGE. 

Variation  due  to  fungi          .         „  156 

„              „     insects       .         .  156 

„              „     mutilation           .  156 

„        spontaneous         .         .  „   155 

Varietal  name    ....  121 
Variety       ....     121-123,  149 

hybrids           .         .         .  150 
Vascular-bundle           .        .   18-20,74-79,84 

„       development  of  76 

„        structure  of  .          .         .  75, 76 

„        tissue  .         .        ..         „  74, 75 

„        plants            ...  126 

Vegetable  Kingdom     ...  1 
„                „      subdivisions 

of      .         .  126-147 
„                „        subdivisions 
of,  synopsis 

of    '           .  146 

Vegetation,  types  of    .         .         .  231 
„             ,,       „  succession  of 

different   .  232-234 
„             „      „  their   distri- 
bution  de- 
pends on    ac- 
tion of  man..  234 
,,             y,      „     their   distri- 
bution depends 
on  moisture     .  231 
„             „      „    their    distri- 
bution depends  on 
soil         ...  232 
„          zones  of    .         .         .  221 
Vegetative  reproduction    .  .         .  108 
Veinlets     „         .          .                   ,  20 
Veins         .         .          ,  20 
Velameti                       „  8 
Venation  .                  ...  20-22 
„        in  Ferns      .         .         .139 
„         ,,  Mosses   ,         .         .  138 
Ventral  raphe     .          .         .         „  55 
„       side        ...»  52 
„       suture    ,  53 
Ventricose           ....  50 
Verbascum  Thapsus     ...  25 
Verbenaceac         ....  222 
Vernation            .         .         „         .35, 139 
Verrucose            ....  64 

Versatile 52,  111 

Verticils    .....  9 
Vessels      ...        73,  74,  76,  91,  98 

Vexillum 56 

Vibrio 127 

„      cholerae  .         .         .         .  129 

Viburnum           ....  244 

„         cotinifolium          .  30, 35, 36 

„        fattens         .         .         .  "        29 

Villous 64 

Violets 115 

Virginian  Creeper,  Indian     .         .  245 

Viscid        .....  65 

Viscnm 201 

Vitcx 243 

„    Negundo    ....  250 

Vitis 24o 

„    semicordata         .         .  245 


INDEX. 

XXI 

PAGE. 

PAGE. 

W 

Wood,  knots  in  . 

168 

„      of  root     . 

79 

Walnut      ....       16,222,244 

„      sap  — 

17,  205 

Waste-products            .         .         .69,  90 

,,      wound  —  . 

166 

Water,  absorption  of,  by  plants     83,  86,  88, 

Wocdfordia  floribunda 

112 

91,93,96,211 

Woodland 

231-236 

„     ascent  of  in  plants    .    96-98,  168-169 

Woody  roots      .            — 

8 

„      available          .         .         .    206,218 

„      stems     . 

10,  66 

„      culture    .         .         .         .93,95 

Woclly      . 

64 

„      denuding  action  of    .         .219,  249 

Wound-parasite 

194 

„      distribution  of  seeds  by  116,  219,  249 

Wounds     . 

165-175 

„       imbibition  of    .         .          .  •           87 

„        healing  of 

165 

„      in  plants           ...             93 

Wound-wood 

166 

„      its  effect  on  plant-distribu- 

tion     ....  218,219 

,,       its  effect  on  the  type  of 

X 

vegetation    .         .         .    231,232 

Xerophyte          .... 

229,  250 

„      Lily          ....             47 

75  77 

„      pores       .         .                   .            100 

Xylia  dolabriformis     18.  224,  227, 

240,  243 

„      required  for  growth  .         .           104 

Wax           .         .         .         .33,    35,  64,  75 

( 

Weeds        148 

y 

Weight,  of  wood           .         .         .             17 

Wendlaiidia  exserla      ...             41 

131  132 

Wheat,  flower  of          .         .         .             56 

„    Plant         .... 

130 

„         rust  of  ....           195 

Yew            

245 

„         smut  of.          .         .         .           134 

Yield,  of  agricultural  crops  . 

95 

Wheel-shaped     ....             50 

Young  plants,  leaves  of 

28 

Whorls      .         .         .         .      9,  33.  34,  65 

Willow      222,  226 

Wind,  distribution  of  seeds  by    58  116,  225 

z 

„     injury  by           .         .            215,  224 

„     pollination         .         .              42,  111 

Zanthoxylum  alctium    . 

29 

„     utility  of.         .         .            215,  225 

8  118 

Wine,  manufacture  of           .                     132 

llfi  227 

Winged  fruit       .         .         .               61,116 

„        Jujuha 

208,  239 

„       petiole  ...                       30 

„        Oenoplia 

239 

„       seed       ...               58,  116 

„        rugosa 

239 

Wings  (alas)        ...                      56 

Zones  of  vegetation 

221 

Witches'  brooms          .         .             156,  200 

1    182 

Wood        .         .      14,  17,  18,  72,  75,  78,79 

Zoospore   . 

188 

„      decay  of  .         .         .         .           204 
„      heart       .             17,  78,  98,  168,  204 

Zygomorphic 

46 

CALCUTTA 

STJMMNTENDENT    GOVERNMENT   FEINTING,    INDIA 
8,  HASTINGS  STBEET 


EXPLANATION    OF    THE    FIGURES. 

PLATE   I. 


Fig.  i.  Germinating  acorn  of  Quercus  incana.  («),  the  short  shoot  of  the  young 
seedling  bearing  small  scale-like  leaves  (i)  (V).  (c),  the  primary  root 
which  here  becomes  a  vigorous  tap-root,  (e)  (#),  the  secondary  lateral 
roots.  (d\  the  petioles  of  the  cotyledons.  The  latter  in  this  case  are 
thick  and  fleshy  and  remain  inside  the  acorn,  below  the  ground,  in 
germination. 

„  2.  Seedling  of  Quercus  incana  with  the  cotyledons  (e)  removed  from  the 
acorn,  otherwise  lettering  as  before 

„  3.  Young  wheat  plant,  (#).  wheat-grain  from  which  the  plant  has  sprung 
and  which  contains  the  single  cotyledon  ;  (d),  stem  bearing  green  leaves 
(')  >  (^)  and  (c)  roots.  In  this  case  there  is  no  vigorous  primary  root 
forming  a  tap-root,  and  a  number  of  roots  of  approximately  equal  vigour 
(&)  (3).  are  found  springing  from  the  base  of  the  stem  (rf>.  Vigorous 
young  adventitious  roots  (c)  (c)  have  also  subsequently  developed 
higher  on  the  stem  (d),  just  below  the  green  leaves.  The  active  roots 
and  their  branches  are  seen  to  have  particles  of  soil  clinging  to  them 
which  are  adhering  firmly  to  the  living  roots-hairs,  but  the  elongating 
tips  of  the  roots  (/)  (/)  have  no  root-hairs,  and  are  free  of  foil 
particles,  as  are  also  most  of  the  older  roots  (<5)  (<5)  and  their  branches, 
their  root-hairs  having  died  off,  and  they  themselves  having  ceased  to 
grow. 

„  4.  Base  of  a  stem  of  Sugarcane  showing  adventitious  roots  (a)  (a)  springing 
from  the  stem  ;  (J),  a  bud. 

All  the  figures      X  |. 


Fig.  3 


Fig.  2. 


PLATE   I. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   II. 


Fig.  I.  Diagramatic  plans  showing  the  course  followed  by  twining  stems,  (a), 
dextrorse  stem  moving  in  a  counter  •  or  anti-clockwise  direction  ;  (&), 
sinistrorse  stem  moving  in  a  clockwise  direction,  z'.<?.,  in  the  direction 
followed  by  the  hands  of  a  clock  or  watch. 

„      2.     Diagrams  showing  (a)  monopodial  branching  ;  (Z>),  sympodial  branching. 

In  (c),  the  sympodium,  or  false  axis,  has  become  straightened  and  has  a 

superficial   resemblance   to  a  monopodium.     (d)  (<f),  lateral    branches 

springing  from  the  leaf-axils. 

,,     3.     Twigs   of  Pyrus    Pashia   showing   the   relative    development    of    shoots 

produced  in  one  year.       X  f  - 

Some  of  the  buds  (a)  (a)  have  remained  dormant,  others  have  produced 
dwarf  spineless  shoots  (b)  (&),  others  have  produced  dwarf  spiny  shoots 
00  (c)>  and  others  have  produced  long  branches  (d~)  (d~). 


Fig.  3 


Fig.  1.  « 


Fig    SJ. 


PLATE  II 


EXPLANATION    OF    THE    FIGURES. 

PLATE  III. 


Fig.l.  (a).  Spine  of  Aegle  Marmelos.  The  spine  arises  in  the  axil  of  the  trifolio- 
late  leaf  (V)  and  at  its  apex  bears  a  rudimentary  leaf  (c),  thus 
indicating  that  it  is  a  branch.  X  \. 

„     2.     (a).     Compound  spine   of  Flacourtia  Cataphracta  bearing   three  normal 

green  leaves  (5)  (i)  and  thus  indicating  that  it  is  a  branch.      X-J- 

„     3.     Stem  of  Gouania  leptostachya.     The  tendrils  (a)  (a)  arise  in  the  axils  of 

the  leaves  (J>)  (J),  and  they  themselves  frequently  give  rise  to  normal 

leaves  as  at  (c),  thus  indicating  that  they  are  branches.      X  £. 


PLATE  III. 


EXPLANATION    OF    THE    FIGURES: 

PLATE   IV. 


Fig.  I.     Pennincrvcd  leaf  of  Qutrcus  incana.     (a),  midrib  ;  (J),  straight  secondary 

nerves  ;  (<:),  reticulate  tertiary  nerves.      X  f  • 
„      a.     Palminerved  leaf  of  Acer  caesium  with  straight  primary  nerves.     X  §. 

3.     Penninerved  leaf  of  Cornus  macrophylla,  with  arcuate  secondary  nerves. 

X  f. 
„     4.     Leaf  of  Smilax  parvifolia  with  curved  basal  nerves.     X  \> 

(a),  broad  leaf  base;  (&),  tendrils. 
5.     Penninerved  leaf  of  Cinnamomum  Camphora.     (a),  strongly  developed  pair 

of  lateral  nerves.      X  \- 
„     6.     Portion  of  leaf-blade  of  a  Fern,     (a),  midrib  from  which  spring  furcate 

lateral  nerves.      X  \. 
„      7.     Pedately-parted  leaf  of  Delphinium.      X  |. 


.4? 


Fig.  5. 


PLATE  ir. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  V. 


Fig.  I.  Pinnately  trifoliolate  leaf  of  Desmodium  tiluefolium.  (a)  (a),  petiolulee  of 
leaflets  ;  (£)  (£),  stipels  ;  (c),  rhachis  ;  (d),  petiole.  X  r 

„  2.  Palmately  5-foliolate  leaf  of  Holboellia  latifolia ;  (a)  (a\  petiolules  ;  (3), 
petiole.  X  f  • 

»  3-  Upper  portion  of  pinnate  leaf  of  Berberis  nepalensis ;  (a),  rhachis.  Note 
the  distinct  joint  (ft)  at  the  base  of  the  terminal  leaflet  and  at  the 
insertion  of  the  lateral  leaflets.  X  J. 

„  4.  Leaf  of  Berberis  Lycium.  Note  the  distinct  joint  (a)  between  the  leaf- 
blade  and  leaf-base.  X  -J- 


PLATE  V. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  VI. 


Fig.  i.    Creeping  stem  of  Ivy  (Hedera  Helix)  with  aerial  roots  (a).     Note    great 

variety  in  the  shape  of  the  leaves.     X  f . 
„     2.     Erect  fruiting  branch  of  Ivy.    Note  how  the   leaves  differ  from     those   in 

Fig.  i.     X  \. 
„      3.     Stem  of  Berber  is  Lycium.     X  \. 

(a),  normal  leaf  in  the  axil  of  which  a  shoot  bearing  two  small  leaves    is 

developing.     Note  the  minute  stipules  at  (d~)  ;  ( J),  a  three-pronged  spine 

which  is  really  a  leaf  as  indicated    by  the  fact  that  a  leafy    branche 

is  developing  in  its  axil.     Note  the  minute  stipules  still  visible  at  (<?). 

(0,  a  three-pronged  green  leafy  structure  intermediate  between  the  normal 

leaf  (a)  and  the  spine  (J).    The  leaf-nature  is  revealed  by  the  presence 

of  the  leafy  branch  in  its  axil.     Note  also  its  minute  stipule  at  (/) . 


Fig. 


PLATE  VI. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  VII. 


Fig.  I.  Twig  of  Acer  casium  from  the  terminal  bud  of  which  four  leaves  have 
developed.  At  (a)  (a)  the  bases  of  the  petioles  of  3  leaves  which  have 
been  cut  off  are  shown.  (6),  scars  marking  the  position  of  the  bud- 
scales  which  have  just  been  shed,  (c)  (c),  scars  marking  the  position 
of  the  bud-scales  which  have  similarly  been  shed  at  the  beginning  of  the 
annual  growth  in  previous  years.  The  twig  thus  shows  the  completed 
growth  of  3  years  and  is  just  commencing  its  4th  year's  growth. 
„  2.  Twig  of  Carpinus  viminea  which  has  just  commenced  its  annual  growth. 
(a)  to  (£),  part  of  the  winter  twig  ;  (^)  to  (c),  part  of  the  spring  shoot 
which  has  just  developed  from  the  terminal  bud  of  the  winter  twig.  At 
(3),  the  lowest  brown  bud-scales  have  fallen  off,  but  the  pair  of  scaly 
stipules  at  (</),  which  flank  the  petiole  of  the  lowest  leaf,  were  actually 
present  as  bud-scales  in  the  winter-bud,  and  at  their  tips  the  dark  brown 
portions  which  were  exposed  on  the  outside  of  the  winter-bud  are  still 
visible  At  (e)  (*),  the  stipules  are  larger,  but  in  them  also  the  dark 
portions  which  were  exposed  in  the  winter-bud  are  still  visible.  At  (c), 
the  stipules  are  normal  and  no  dark  portions  are  visible.  In  this  case 
the  bud-scales  are  obviously  modified  stipules. 


.3rd  y* 


fig    i 


PLATE  VII. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  VIII. 


Figs.  I — 8.  Showing  the  transition  from  an  obvious  bud-scale  (i),  taken  from  the 
exterior  of  the  bud,  to  a  normal  leaf  (8),  taken  from  the  interior  of 
the  bud,  observed  in  an  opening  bud  of  the  Horse  Chestnut  (Aesculus 
indic(i),  indicating  that  the  winter  bud-scales  are  really  leaves  which 
have  been  checked  in  their  development  at  an  early  stage  and 
converted  into  protective  coverings  for  the  bud.  Minute  points 
representing  the  rudimentary  leaflets  are  recognisable  at  the  tips  of 
the  scales  (i)  to  (4).  X  -j- 

„  9.     Young  shoot  of  Stephegyne  parvifolia.  («),  base  of  petiole  of  leaf  which 

has  been  cut  off  to  show  the  large  interpetiolar  stipules  (?>)  (/>). 
The  terminal  bud  is  concealed  and  protected  by  the  stipules,  one  of 
which  (0)  is  facing  the  observer.  X  -f . 

„  10.  Stem  of  Prinsepia  ut'ilis ;  (a),  leaf  in  the  axil  of  which  is  situated  a 
bud  (3)  and  also  a  stout  spine  (tf),  the  latter  protecting  the  bud 
(<5).  (d)  (rf),  other  axillary  buds,  the  subtending  leaves  of  which 
have  fallen  off.  At  (<?),  the  axillary  bud  has  produced  a  leafy 
shoot.  One  of  the  spines  bears  a  leaf  (/)  showing  the  spines  to  be 
branches.  X  4- 


Fig.  10 


PLATE  VIII. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   IX. 


Fig.  i.    Diagram  of  involute  vernation. 
„     2.  „          revolute          „ 

„      3.  „          conduplicate  vernation. 

,,4.  „          convolute  „ 

,.      5.  „          circinnate  „ 

„     6.  „          plicate  „ 

(a)  (a)  throughout  is  the  midrib ;  (<$),  the  upper  surface,  and  (<:),  the 

lower  surface,  or  back  of  the  leaf. 

Figs.  7 — 9.  Stages  in  the  development  of  the  leaves  of  Viburnum  colinifolium. 
The  opposite  leaves  (a)  (a),  when  very  young,  are  erect,  and  with 
their  upper  surfaces  in  close  contact,  only  the  lower  surface  being 
exposed  to  the  light  and  air.  The  lamina  is  plicately  folded,  and  the 
delicate  green  tissue  is  protected  by  the  close-set  framework  of  nerves 
which  alone  are  exposed  on  the  lower  surface.  As  development 
proceeds  the  folds  of  green  tissue  are  flattened  out  and  the  leaf  is 
thrown  outward  and  downward,  the  upper  surface  being  finally 
exposed  to  the  full  rays  of  the  sun  as  in  Fig.  9. 
The  petioles  of  the  young  leaves  are  grooved  on  their  upper  surface, 
so  that  space  is  provided  for  the  terminal  bud  between  them.  In 
Fig.  8,  (b)  represents  the  terminal  bud  in  cross  section,  and  (c)  (c) 
the  closely  adpressed  petioles  also  in  section. 
Figs.  7  and  8  slightly  enlarged,  Fig.  9.  X  J. 


Fig.  4 


Fig.  5. 


PLATE    IX. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  X. 


Fig.  I.     Diagram  of  a  raceme. 
,,2.  „  „     corymb. 

„      3-          „  „     spike. 

„     4.          „          „     umbel ;  (c~),  the  involucre. 
„      5.          „  „    head  or  capitulum  ;  (<:))  the  involucre. 

„     6.          „          „     compound    umbel ;    (c),    involucre ;    (a1)   (d),    umbellules ; 

(')  (*)i  involucels  ;  (/),  rays. 
„     7-          „          „     panicle. 
„     8.          „          „    dichotomous  cyme. 
„     9.          „          „     helicoid  „ 

„    10.          „          „    scorpioid  „ 

Throughout  (a)  (a)  are  flowers;  (<5)  (<5),  the  bracts. 
„    II.    Diagram  of  a  hypogynous  flower;   (a),  sepal;   (6),  petal;  (c~),  stamen; 

(d),  ovary. 

„  12.  and  13.     Diagrams  of  a  perigynous  flower — letters  as  before. 
„  14.     Diagram  of  an  epigynous  flower — letters  as  before. 
„  15.  „          valvate —  aestivation. 

„  16.  „          valvate — induplicate          „ 

„  17.  „  „          involute  „ 

„  18.  „          of  an  imbricate  „ 

„  19.  „          contorted  „ 

Overlap  to  the  right. 


Fig    13 


Fig    12 


Fig.  13. 


Fig.  14. 


Fig.  15. 


Fig.  16. 


Fig.  17. 


Fig.  18. 


Fig.   19. 


PLUTE  X. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  XI. 


Fig.  i.     Winter  twig  of  Odina  Wodier.     (a),  leaf-scara — note  their  shape  and  alter- 
nate arrangement. 

„      2.     Winter  twig  of  Hymenodictvon  excelsum.     (0),  leaf-scars — note  their  shape 
and  opposite  arrangement. 

„      3.     Photograph  of  Bombax  malabariciim.     Note  the  branches  in  whorls. 

„      4.     Photograph  of  Bombax  malaharicum  showing  the  woody  buttresses  at  the 
base  of  the  stem. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   XII. 


Fig.  I.     Seed  of  Oroxylum  indicum.         X  $. 

(a),   membranous  wing   developed   from    the   testa ;    (fi),   the  embryo 

enveloped  in  the  testa. 
„      2.     Embryo  of  Oroxylum  indicum  extracted  from  the  seed.         X  \. 

(a),   radicle  ;    ($),   lower  surface   of   one   cotyledon,   its  upper   surface 

being  placed  flat  against  that  of  the  opposite  cotyledon,  which  is  not 

visible  in  the  drawing.     The  plumule  also  is  not  visible. 
„      3.     Young  seedling  of  Oroxylum  indicum.     (a),  (5),  the  two  cotyledons  with 

their  upper  surfaces  separating  from  each  other ;  (c),  the  hypocotyl  ; 

(<Z),  primary  root  ;  (c),  secondary  lateral  roots.         X  \- 
„     4.     The  same  at  a  later  stage  ;    (/),  the  minute  terminal  bud  of  the  stem, 

otherwise  letters  as  before.         X  T* 
„      5.     The    same  at  a    still  later   stage;  (^),    the  first  pair  of  foliage  leaves;  A, 

the  epicotyl,  otherwise  letters  as  before.         X  T. 
The  root  not  shown. 


PLATE   XII. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   XIII. 


Fig.  I.     Erect  branch  of  Coriaria  nepalensis  seen  from  the  side.   X -j-- 

Note  the  decussate  leaves  arranged  in  four  ranks  on  the  4-angled  stem 
with  their  upper  surfaces  at  right  angles  to  the  rays  of  light  coming 
from  above.    The  stem  is  not  twisted. 
„     2.     Horizontal  branch  of  Coriaria  nepalensis,  seen  from  above.     X  \. 

Note  the  distichous  leaves  with  their  upper  surfaces  practically  horizontal 
and  at  right  angles  to  the  rays  of  light  coming  from  above.     Also  the 
twisting  of  the  4-angled  stem  caused  by  the  efforts  of  the  leaves  to 
place  themselves  at  right  angles  to  the  incident  rays  of  light. 
In  both  figures  one  side  of  the  4-angled  stem  is  shown  shaded  throughout. 


PLATE    XIII. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   XIV. 


Fig.  I.     Lever-apparatus  dissected  from  flower  of  Salvia  lanata.     X  4     (a),  (a), 

filaments  ;  (<5),  pouch  formed  by  the  metamorphosed  lower  anther-lobes  ; 

(c~),  pollen-bearing  anthers ;  (d),  upper  arm  of  connective.     If  the  pouch 

(&)  is  pushed  back  the  anthers  (c)  descend  as  indicated  by  the  arrows. 
„     2.     Flower  of  Salvia  lanata.     X  3  with  half  the  calyx  and  corolla  removed 

showing  the  lever-apparatus  in   position,     (a),  hypogynous   disc  which 

excretes  nectar  ;  (i),  ovary;  (<?),  style  ;   (<?),  stigma.     The  arrow  shows 

the  direction  in  which  pressure  is  applied  by  a  bee. 
„     3.     The  same  with  the   pouch-like  trap  door  pushed  back  and  the  anthers 

consequently   brought    down  close   to    the   landing  stage.     Letters  as 

before. 
„     4.     Complete  flower  ot  Salvia  lanata.     x  3.     (a),  position  of  the  stigma  in  a 

young  flower  ;   (i),  its  position  in  an  older  flower. 
„     5.     A  stamen  of  Berberis  Lycium. 

(a),  from  the  back,  the  anther  valves  are  closed. 

(b~),  also  from  the  back,  the  right-hand  valve  is  opening  and  beginning  to 

move  upwards  carrying  the  pollen  with  it. 
The   left-hand   valve   after   completing  the   upward   motion   has   turned 

inwards  so  that  the  mass  of  pollen  faces  the  centre  of  the  flower. 
(c),  from  the  front  showing  both  the  pollen-covered  valves  facing  the  centre 

of  the  flower. 
„     6.     Expanded   flower   ol    Berberis    Lycium.     X  5.     (a),   inner  large   yellow 

sepals  ;  (i),  outer  petals  ;  (c),  inner  petals  ;  (d~),  filaments  ;  (#),  nectary. 
„     7.     Single  petal  of  Berberis  Lycium  showing  the  position  of  the  two  large 

orange  nectaries  at  the  base. 
„     8      Showing  position  of  a  stamen   which  has  sprung  up  on  being  irritated, 

(a),  stimatic  surface  ;  (J),  ovary. 


Fig.  3. 


Fig.  4 


Fig.  6 


Fig.  5. 


Fig.  7. 


Fig.  8. 


PLATE    XIV. 


EXPLANATION    OF    THE    FIGURES. 

PLATE  XV. 


Fig,  I.    (a).    Barclayella  deformans  on  Picea  Morinda.      x  {•    Note  the  orange- 
red  curved  needles  on  the  attacked  shoot  and  compare  them  with 
the  straight  healthy  needle  shown  below. 
(3).     Cone  scale  of  Picea  Morinda  attacked  by  the  Barclayella,  showing 

the  beds  of  teleuto-spores.         x  £. 
„     2.    Peridermium  brevius  on  Pinus  excelsa.    Note  the  reddish-yellow  blisters. 

The  needles  have  been  cut  short  for  convenience.         X  \. 
,,     3.     Teleuto-form  of  Gymnosporangium  Cunninghamianutn  on  Cupressus  torulosa. 

Note  the  gelatinous  spore-masses.         x  \. 
„     4.     Aecidium  form  of  Gymnosporangium  Cunninghamianutn  on  Pyrus  Pashia. 

Note  the  tubular  aecidia.         x  -f. 

„     5.     Gambleola  cornuta  on  Berberis  nepalensis.     Note  the  black  hairs  of  teleuto- 
spores.         X  \, 

(All  after  Butler). 


PLATE  XV. 


EXPLANATION    OF    THE    FIGURES. 

PLATE   XVI. 


Phytophthora  infestans. 

Fig.  i.     Back  of  potato  leaflet  attacked  by  the  fungus  showing  the  characteristic 

dark  spots  with  pale  mouldy  margins. 
.,    2.     Conidiophores  bearing  conidia  emerging  from  a  stoma  on  the  underside 

of  a  leaflet.     Highly  magnified. 
„    3.     Conidium  with  protoplasm  dividend  into  blocks,  the  latter  being  liberated 

as  zoospores  in  Fig.  (4). 

„    5.     Zoospore  coming  to  rest  and  losing  its  cilia. 
„    6.    A  zoospore  which  has  germinated. 

„    7.    A  conidium  which  has  germinated  directly  and  sent  out  a  germ-tube. 
Figs-  (3),  (4),  (5),  (6)  and  (7)  very  highly  magnified. 


PLATE   XVI. 

PHYTOPHTHORA  INFESTANS. 


EXPLANATION    OF    THE    FIGURES. 

PLATE    XVII. 


Fig.  I.  Sporophore  of  Fames  annosus  on  deodar  showing  the  brown  tuberculate 
incrustation,  and  bracket-like  active  portion  with  white  hymenium 
below ;  a,  a,  brown,  mostly  sterile  portion  ;  b,  b,  recent  spore-forming 
portion. 

„    ^.    White  mycelial  sheets  under  the  bark  of  the  collar. 

,,  3.  Drawing  of  a  portion  of  the  surface  of  a  block  of  wood  cut  from  a  diseased 
tree,  showing  the  white  areas  in  the  wood. 

„  4.  Tissue  elements  modified  by  the  action  of  the  parasite,  a,  a,  cells  of 
the  wood  parenchyma-bearing  starch  ;  t>,  bore-holes  of  the  hyphae  with 
a  filament  extending  across  between  two  ;  0,  spiral  cracking  of  the  wall ; 
d,  pitted  vessel  with  a  hypha  penetrating  it ;  *,  network  from  the 
crossing  of  two  spirals,  the  trachea  is  seen  with  the  upper  wall  removed 
so  that  the  network  is  made  up  of  the  spiral  of  the  lower  wall  of  this 
trachea  and  that  of  the  upper  wall  of  the  subjacent  one  ;  f,  dissolution 
of  the  middle  lamella  which  results  in  the  isolation  of  the  elements. 

„    5.     A  rhizomorph. 

(All  after  Butler), 


PLATE    X*ll. 
Fig.  1. 

FOMES  ANNOSUS.  Fs. 


PLATE    XVII. 
Figs,  fi-5 

POMES  ANNOSUS.  F». 


(a) 


(b) 


PLATE    XVIII. 

TRAMETES  PINI,  FRIES. 

(«).    Sporophore. 

(b).    Wood  of  Plnus  excelta  rotted  by  the  fungus. 
(After  Butler). 


EXPLANATION    OF    THE    FIGURES. 

PLATE   XIX. 


Puccinia  graminis.     Pers. 

Fig.  I.     Wheat  stem  and  leaf  with  teleuto-spores .     x  f 
„      2.     Ureao-spores.      The  one   on   the   right   has  developed  two   germ-tubes. 

Highly  magnified. 
,.      3-     Ideuto-spores.     The   one   on   the   right  has  germinated,  the   upper  cell 

producing  a  pro-mycelium  (a)  from  which  spring  sterigmata,  bearing 

sporidia  (J).     Highly  magnified. 
„     4-     Under-surface  of  leaf  of  Serberis  Lycium  showing  the  ceci&ia  on  a  dark 

brown  patch,     x  \. 

„      5.    An  open  aecidium  from  above.     Magnified. 
„     6.     Transverse  section  (of  Berleris  leaf,     (a),  is  lower  surface  of  leaf.     Two 

secidia  are   shown,  one   of  which  has  not  yet  burst  through  the  leaf 

epidermis.     Highly  magnified. 


PLATE   XII, 
PUGCiNIA  GRAMINIS  PERS. 


PLATE    XX. 

LEAF  OF  A  YOUNG  MAHUA  PLANT  (BASS1A  LATIFOLIA)  INJURED 
INDIRECTLY  BY  FROST,    x*. 


Pboto.-Mechl.  and  Litho.  Dept..  TbontMon  Oollege,  Hoorkee. 


THE     AVERAGE     ANNUAL     DISTRIBUTION 

REFERENCES. 

8ELOW     20     INCHES  C_ I 

20    TO    70     INCHES  d 
ABOVE     70     INCHES  f~!  ZU 


PLATE    XXI. 


Pboto.-Ziuoo..  January,  1908.— No. 29 .so  -uoo. 


IE     RAINFALL. 


DISTRIBUTION     OF     THE     PRINCIPAL     TYI 


REFERENCES 


ARID-COUNTRY     FORESTS 

DECIDUOUS     FORESTS 

EVER-QREEN     FORESTS 

HILL     FORESTS  C 

TIDAL     OR     LITTORAL     FORESTS  L. 


L_ 
L 


PLATE  XXII 


!    V.OO 


OF     FORESTS 


LIMITS     OF     THE     DISTRIBUTION     OF     THE     PRir 

REFERENCES. 

LIMKT    OF    TEAK     (TECTONA     GRANDI8) 

„  SAL    (8HOREA     ROBU8TA) 

DEODAR    (CEDRU3     LIRANI) 

CAOUTCHOUC  (FICU8  ELASTICAX___ 
RED  8ANDER8  (PTEROCARPU8  SANT 
SANDAL  (SANTALUM  ALBUM).  _ 


PLATE    XXIII. 


AL     KINDS     OF     TREES. 


Ao.  2350-2. 1100 


S) 


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