Reproduction in Flowering Plants

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CHAPTER 2

SEXUAL REPRODUCTION IN - . .
FLOWERING PLANTS .fl \·
.x~~~r,.du;~'
f ~m, rWI
r] ~s>r 1(\ . rn

2.1 Flower-A Fascinating


Organ ofAngiosperms
2.2 Pre:fertilisation: Structures Are we· not lucky that plants reproduce sexually'? The
andEvents •myriads of flowers that we enjoy gazing at, the scents and
. the perfumes that we swoon over, the rich colours that
2.3 Double Fertilisation
attract us, are all there as an aid to sexual reproduction.
2.4 Post:fertilisation: Structures flowers do not exist only for us to be ·used for our own
andEvents
\
selfishne~ ! o~ enng plants show.8'Xllal repnilud:10_9.
A look at the diversity of structures of the inflorescences,
· 2.5 Apomixis and
flowers and floral parts, shows an ~azing range of
Polyembryony
adaptations to ensure formation pf the end products of
sexual reproduction, the fruits and seeds. In this chapter,
·ie'tus ui'.icterstaiid hie morphology"":stiucture and the
processes of s~xual reproduction in flowering plants
(angiosperms).

2.1 FtoWER - A FASCINATING ORGAN OF


ANGIOSPERMS
Human beings have had an intimate relationship with
flowers since time immemorial. Flowers. are objects of
aesthetic, ornamental, social, religious and cultural val.,JJe
-.:a
~ - - - -'71 ,-:t.~ ---.::s::-.,-
~.......

- they have always been~ s.ymboJS-for..cou\!.e..)cing


~ ~tant human f~: l~ gs such as lo~e. ~!f~cti~,
happiness, grief, mourning, etc. List at least.five.flowers
of omamentai valu e that are ·commonly cultivated at
<;c ieoq of- cu.k+t 11 a.¼,d' , ·
l:f'Ud ,~ , rrf\Oi\<.e.tiztn::
'(u · Figure 2 . 1 A diagrammatic representation of L:S. of a flower"
\JQl"'d on-a .
.i
O
lf)~
Lused
• . homes and in gardens. Find out the ~ s offive more }lowers that
"'
iri social and cultural celebrations in your family. Have you
II
ofjloriculture - what does it refer to?
.
(!o a biologist, flowers are qi~oIP.~ol~~cal and embryolo c evario
th
~1!!:-~ tes ~f ~ -al repr?ouction. In class XI, you have read a tyPiO
parts of a flower. Figure 2.1 will help you recall the parts of J!lO:
flower. Can you name the two parts in a flower in which th~
important units of sexual reproduction develop? Ard.~-~ .

,W 2.2 Pim-FERTILISATION' STRUCTURES AND ~ NTS .


,&, l-'\«-f &+e rn . • weJaJI
Much before the ac seen on a lant. the decision that -" ·
~w•
i ts going to flower has taken nlace. Several hormonal and struc ...-,eO
1
~U~sqnq are •
initiated which lead - - - - - - JoR
_to_the differentiattifn and
___,~er evefio~
v
u~ thefl~ __ _ _ _;,___ ---:.1btl
--•iiiiiiiiiiiiii: J \. 0
primordium. Inflorescences are formed which ~ tlcU'
~• and Qien the flowers. In the flower the male and female rep devel*
~Oftlt tmm «ld.i~ structures, the anclroecium and the gynoecium differentiate and 5~1°'.
dt.Jt..l,
'1,. • You would reco11ect that the androecium consists of awh orI of esr,,· .rtl:

,~"2.
rtorot "w:l 1 :~resenting the male reproductive organ and the gynoec~UJil rePr
"- e female reproductive oESan. -
_,
V Y V

EXUAL REPRQDU · . •~ . . \I/


·
1~
.
.2.1 Stamen Mi
CTION IN FLOWERING PLANTS

' crosporangtum and Pollen Gr


~~~:_=-_Pi ~f'~Qw [~~ ,JZ~ \~
gure 2.2a shows the two arts · ·
talk called the ftlam p of a typical - the long and slender
~ ed the antht;J. The ent, and th e ,U;nninal generally bilobed structure
attached to th th f-roxtmal end of the filament , • f-1\ ah, o c.ttte :
e _a)arn fth
e numbe o e Dower.
r and length of stamens are variabl:-fn .
owers of different species. If you were to collect a
stain . N'u.u th i,bt\l..frl 1
_en each from ten flowers (each from different ff) o (l ()+ht c.u-!,,, 1

species) and arrange them on a sllde, you would rflol) c,Sr ~' ~ •

. able to appreciate the large variation in s~e s~en Pollen grains

lil nature. Careful observation of each stamen


:der a dissecting microscope and making neat
. agrams would elucidate the range in shape and
attachment of anthers in different flowers.
AjYPical angiosperm .anther is bilobed with
each lobe ha~g1wo theca, i.e., they are dithecous
(Fi~e 2.2 6). Otten a longitudinal oove runs
lengthwise se aratin e . he . Let us
unclerstand the various types of tissues and their
organisation in the transvers~ section of an anther
(Figure 2.3 a). The bilobed nature of an anther is
~ ry_distinct in the tranw erse section_o[theanther.
The anther is a four-sided (tetragonal) structure
consisting of, fom;_ ~-5!,9,!porangia located at the (b)
comers, two in each lobe.
The microsporangia develop further and . (a)
becm:µe pollen sacs. They extendJbngitudinally . .
through the ~ength of an anther ana af[P@ed Figure 2-~!',typ~c~ s~en;
~ ollen grams. · -:Jert'.frree- dunens10nal cut section
Structure of microsporangium: In a transverse of an anlher •
section, a typical microsporangi.um appears near
s:trcular in o~tline7 t is gen'.erally surrouiiaedt>~_;V~ l~ s (Figure -nr if\ h6,d ror/ \6 .
2.3 b)- the epidermis, endothecium~middl«u&et§1and th~ ~-The
outer three wall la>,:ers perform the ,function of protecti?n and help in
t. 'vv .

dehiscence of anthe to release the pollen. The innermost walflayer is Q- cM®I.~


th~ tapet
(tapetum
=c.
nouns e e eve op~ -~~ e,_n !i:am_,s. e s
- - - ~
possess ense cytoplasm and generally have more than one
o
,
-.•~-••wwn-i;
--e J @a. . . -
,
lnucl,!;u~. Can you -· k of how tapetal cells could become 6i-nuclea.te1' 'f'(')O"' +t(T'.lt:½1
When the anther is oung, a group of compactly arranged homogenous
-- _..,_._. ..-:a- --
J ··
cells called the spor enous tissue occu ies the centre of each
.ii

"==
microsporangium. po~ ~o Id I 1'I' ~r-}f\ Cd\~ .;:-· -
Microsporogenesis : As the anttlier ijJvelops, the cells of the sporogenous
tissue undergo meiotic divisions to form micro~por tetrad What would -
be the ploidy of the cells of the tetra.d?
-4-i.-i.----,~+1---tConnective ----
/ ~ - -Epidermis
\+-',,-------Endothecium
~--1-4-~ - -sporogenotts
tissue

Tapetum

a
si
fr

-~ - s -1i
\\

l!'J-- -4iiii~~Pollen grains

'' /
"' · •
, , . ,'l
...OOii/)
-.....i!,n.,~
:,.i
,~""'
.
I \
- i
I

\ ·. 'qjll66,i:r:P."'""" I 1
- / . (c} . J
#.ure 2 . ~Transverse section of a young anther;~ nlarged view of one microsporan~
showing wall layers~ mature dehisced anther
10
As each cell of the sporogenous tissue is capable of giving rise
1110th
, . microspore tetrad . E ach one i~ a g_otential _pollen or microspore
-9:ll. The process offormation ofnucrospo~ ~,;:'~~ m?th~
through meiosis is called microsporogenesis. e IlllCTOSporeS, as
· te
are foffiied , are arranged inacTusteroffottr cells-the mic~
(Figure-2 .3 a) . As the anthers mat~ ~'ctacl~ydrate, the ~~~
'
dissociate from each other and develop into pollen
giilns (Figure · u
, 22 .· Inside eac_fi:inlc"rosporangltifu~ver~!tlioUsand's- ofnrlet'O~ ~~
0
I •
,Bi;,a tns are formed that are released witli ffie defilscence
(Figure 2 .3 c). - -- --~- - - · ---- u
Polle0 ,&-...a.,; · bytes_: d6¢
&&&¥: The pollen grains represent the male g8;ffietop _
IfYo
1
touch the openedaii.'t hers ~fHibiscus or any other flower you wow '
°
depositlon 1"yellowish powdery - - --
p ollen grains on yotir fingers. SP 0dd
th ese gratns-011-a-dfop of water takefton a glass slide and observe u
SEXUAL REPRODU CTION IN FLOWERING PLANTS
'

( 1
"''
'~
... ·, -~

' ·\,
,--... .. l

,· -~ ' ....
r" , ,i i ..
C
' .. '"'4:

Figure 2.4 Scanning electron micrographs of a few pollen grains {a)

a microscope. You will really be amazed at the vruie"ty of ru·chitecture -


sizes. shapes, colours, designs - seen on tJ;ie..po~ains
from differen_t species (Figure 2.4).
Pollen grains are gen
,A. . ffT) ·
spherical measuring about
r ..
25-5 micro diameter. ~s a 1;rominent two-layereg
Vacuoles-1-=-hl:'ir-~F- i
' . e hard oufer layer called the exine IS made up of
--- ; !AL . • I
·spo~ogollenin which is 9ne of the most resistantc¥g~,; ~ ate~al
~ wn. Jt can withstand high temperatjires andS ongas1§s ;st Nudeus - ..,,__,,~.....,.
arui!!2!!: Noenzyme that degrades sporgpp))eOiP jssgzr8t
Jru2wn. ~~en ~ --~ e has prominent apertures calle~ e"! "'
p~~s where sporopollenin is abse:Rt. Pollen grains are well~- \I_ .
Ifreserved as fossils because of the spprgp.ofiernn:" presence of ·

The eE11e ~ b!ts 3; ~~i,!1~Ka_rr~~~gis an!~!!S.~ -..


do you think the exille should be hard? What is the \J-
junction of germ pore? The inner wall of the pollen grain is - Asymmelrtc°'.l-~~&!~
call
.

-su
_rr
ed th em

_o.,.
_~""--
l9$!
---
. tine. It 1s
. a.thin an d con tin uous 1ayer ma
. .
cellulose and 2.ectin. The cytoplasm of pollen grain is
. d e up of

~-d.]J;y~ -pJa;mzt WS:IJ,lQW~ Wli'en £fie'jJollen gYaTIHs


splindle

_,.., • Vegetative eel_ , ,._, .

e it contains_!WO~ells, _the~ v~~ ll an~ ! !


4

~
~Figure 2.5b). The vegetati! :Sell_is __b~e~, h~s ab~ t
food reserve and.,c!J~ g ~_lllar]y sg_aped !11.Jf l~J.¼S- The Generative cell---"~;,,;,i..._-:-,,
genei-ative cell is small and floats in the cytoplasm of the (bl
vegetative cell. It is spmdle shaped witll den~e cytoplasm and Figure 2.5 Enlarged view of f4'
a nucleus ~ In ,over 60iJe/'cent of angiospe~ :--pollengralns a pollen grain tetrad; stages
_ - ~---- of a microspore maturing into a
are shed at this 2-celled stage. in the remainiri.g spedes, the pollen grain
-----.-.----
generative cell divides mitotically t_o give rise to the two male
g ~ be~ llen grainssr.e sh~d @;ell~d stae ·. - ~stb~ af'eJ1\e t 'l'i
Pollen grains of many specie e. severe allergies and bro~1cbial te

affliction s in SO~,.P£._Opl e often leading to s.fu:.o~ic reseJratory


~ oide,rs - a~a. b~ nc:"11tis, etc. It may be mentioned that ~ niurr::,. > ~ar~+ht! , t~~ol7oc:liLlfl\
or c~o~ that came into India as a cont.aminanL \-Vith impor1 ed wheat, \
has become ubiquiwu:::, in occurrence and causes p~k~rgy.
- - - ----- _ .. _r- - '} .......
\ I/ •\Cl 1,01TI1' .... '!:,~~J.J_fl.~ w= ~
~l /: t.o.l~\,l'\ l , ~t) t --tc in Cl4) 'a
..;I'/ I-¥ · a1
("O. Pollen grams are rtcli. in nutrients. It has become a fashion !n
• )n f. o, c::tro~o-& o 11 ears to use pollen tablets a s food s ~1~p lements. In west~rn coun

~ r.uN
'ti~ ' m"'~ iarge n umber or pollen p roduct~ in the form of tablet! anct s
-~ C"\Of-
11'"1 .
~ lQ , ~
available in the markerl f2!!.en cor:is~pttoo hasbeen, c
tlle i,erfonnance of athletes and race horsS!!, (Figure 2.6).
e 0

~,f'"~ 6r..-oc:il,., . ,,.,.r 1 m 1'c '4'....-t ' _,,


30
- - ~ / l'OllenProducts . Bee Polleii°
~'!S'1m ... .__i, ~i;;:;:~ from England
f t-'IV. C • . ~ '"'

tr~: -.- p
· Natural Bee- collected Pollen
· in Tiny Golden Pods
A natural food from honey bees
NET WEIGHT .45 OZ.

I 1'418•• a. -.-
1!1,c~
• smdR..ut
~~:gr-c
t>~~ fOJ'(I
12.l'<ttte . o~\lct:60.a.
~.; ).ltn \J
~.-....
Figure
q r~
2.J Pollen prodi cts
. .
c;re.ti\ . .

I . ( !r~~+,'t) · .
'-°rfti\- a - JJ"Q 11 ( M~ Jonce they are shed, pollen grains have to larid on the sti. Fl
• ~U~n ff dN before they lose viability if they have to bring about fertilisation. HowI
'tj- 1 • do you think the pollen grains retain viability? The eriod for which po
---e:..l.M .
l
~.. ~ .t,ott 4 - '' · f--> . grains remain viable is higltly variable and to some extent aepends on
M ono ~ 2>1ai. prevaillng"teiiiperat~ and humidity. Tn-some cereals such as rice.
Cl
wheat, pollen grains lose viability within_30 minutes of their release,
ca
in some members of Rosaceae , Leguminoseae and Solanaceae, ,.
maintain viability for ~ - Yo~ may have heard Qf storing sem
sperms of many animals including humans for artificial insemination. Tl
is possible to store pollen grains of a large number of species for years s1
81
liquid nitrogen (-196°C) . Such stored pollen can be used as pollen
5

siinila'.ito seed banks, in crop breeding programmes. - -,


...
2 .2 .2 The Pistil, Megasporangium (ovule) an~ Embryo sa l
The gynoecium represents the female reproductive part of the flower. u
gynoecium may consist of a single pistil (monocatpellary) or may h3 t
more than one pistil (multicarpellary) . When there are inore than°
the pistils may be fused together (syncarpous) (Figure 2. 7b) or maY
' 27 . . .
free (apocarpous) (Figure 2 .7c) . Each pistil has three parts (Figure · . .1
the stigma, style and ovary. The stigma serves as a landing platfo
for pollen grains. The style is the elongated slender part beneath
stigma. The 6asal bulged part of the pistil is the OYUY.. Inside the 0

-
is the ovarian cavity (lo£_ule). The placenta is located inside . e O . ' 1
cavify. Recall the definition and types of placentatlon ffiat yoµsfi'.iID
'
SEXUAL REPROpUCTION IN FLOWERING PLANTS
QVUL£ 1'-rr~ c,,uM~ N )>

Stigma

I
Style
--,f - - - - - - Funicle
sugma ,......,--- Micropyle
Micropylar pole
Outer integument

Inner integument
-+--+--+-- Nucellus
Embryo sac
1--.J----1,---<1---

Ovary _ ___,~-Chalazal pole


._ ..--Thalamus

(c) (d)
(a) (b)
S.Q('\Ctlrro<A . .
Figure 2.7 4issected flower of Hibl cus showing pistil (other flJ)J'al parts have been removed);
%M~lticarpellaiy, sygc~s pistil of P2£,~ver ; ~A ~l;!tic_¥P.!i!,~ ae ~cru:ru>J.1S
· . gynoecium of Mwhel~ A diagrammatic view of a typical anatroe ous ovul~
.- O\J~ of") b O..~~ .\
~ t :;--
Class XI. Arising from the placenta are the megasporangia, commonly 0 o~t-
~ JLovui~.= rne numoer of ovules in--~ ovary may •be 29,e [email protected]_ p,'j~ ,' - t''t-l'H)
padw , mango) to many (papaya. water melon, orchids). .
The Megasporangium (Ovule) : Lef us familianseourselves with the@ ~a~oLU M o~T
structure of a typical angiosperm ovule (Figure 2.7d). The ovule is a small ,Corn~ on
,s \J Si_, ~cll.l~ ·
structure ~ched to the placenta by means of a stalk called funiE_le. '" o<rtia ~f
The body of the ovule fuses with fti~ de t!.1~-~ £ caj!_e_g 1lflYE}~Thus, (j) · rn,' a nJ
hiium represents the junction between ovule~fumcl~._Each ovule has -_ llOnuf\u.
one or two protective envelones calle~ _!_~~ents. In~en_!_s e~: ir,:l<@ n1 (l'"T'/Jt)
the nucellus except at the tip where a small' opening~called the micropyle · _ .
is organised. Opposite the micrcipylar en<Cis!he Chalaza, ri presenurig'f,l\ · i' \llllo
meoasaipart of the ovule. f
0f'fno/rn I.V l'\1f1)
Enclosed within the 'iiiteguments is a class of cells called the nucellus. 'f' mrrui
Cells o( the nucellus have abundant reseive food materials. Located1n the· 6
nucellus is the embryo sac or femaieganietophyte. An ovule generally has :.~:--:.....
~ ;;;::;;:.:_~:.:...:~.:.=:=~,,.;_~ - - -- - • , /
~..,~~.4-=t1±::::~;:;~~
.:,00
a single e131bryo sac formed from a megaspor~ 0 ,-,· a. {f-o c-'11~,. •
Meguporogenesls : The~ Q q of meiw.eores from. th~. ,.; flo'-«>11 .
~e asp~re m~thei' cell is called megasporo~ is~ ~ne~ally V • 1
.
differentiate a sm e megaspore mother cell (MMC) in the microp yl~ region
' .-"' - 47~ - ,. , .
e~ .i'vt-- - Nucellus ,..,_,..-- Nucellus

Megas11r, ,
,-- - Megaspore tetra(!
- Megaspore dyad
mother cell ·
CP+1
( C f'\O QJJ;~~
o"'~. .
(a) .
Micropylar end
~rr>
p-u (l &f

Chalazal end
. Antipod 1

Synergids~--14~H-t--

Egg ---t-w--...., ) .
Central
. cell
~ .------ffl--- Polar nuc
.polar - -- central c
nuclei

Antipodals ~ =---r-"'.'.:
~-- ~--Nil-- Egg

+-':=--t-li.f--.'?' Synergids

~~bu.r-q , fi ' (b)
(j U 11~~'11.fZ-= ==-- Filiform

~a,or~ J
·• '\ Micropylar end apparatus
PoL'100\l~ T'(ft sA<' . . (cl
. '. ['8bi- °i .
Figure 2.8 · (a) Parts of f
ij
ovule showing a large tnegaspore mother ce).l, a dyad and a tetrad
megaspores; 2, 4, and 8 -nucleate stages of embryo sac and a mature embryo sac: Id
diagrammatic representation of the mature embryo sac.

. t of the nucellus.· It is a large cell containing dense cytoplasm and


'it!-,, ' prominent nucle us . The MMC undergoes meiotic division. What iS
, importance of the MMC undergoing meiosis? Meiosis results i1l ,
production of four megaspores (Figure 2 .8a) .
i Female gametophyte : In ~ majority_ of flowering plant~
••-----• megaspores is functional while the other three degenerate.:. g,olYb

' i;,uul:).J?l f':


{: ~ 'c..
-~,·-~
t e.,t--e...lJLI!,
functional megaspore develops into the femal~
~). lhis method of emfiryo sac formation from a.s ingle m~gaspore IS:;,1
·monospo~ v0 L
0 pme!'t. What w ill be the ploidy ofthe cells 0Jt11e_
n
1

I . "1v1'ii1c. t&'functional megasp~re and female gametophyte?


~ro ntfu.u .

. ff~ -~ r," 'r~ -"--, ~


. ~l\i:i1',iJ, cM~ - ~ tn'"
i\t
• Of't -2_0
EXUAL " " oucnoN IN FLOWERING PLANT~
.
PIJ.unbo;=r ai blJ.Lrn \:.o ~
ea m
., - ~
/ !3, c. l ii"'
' ,-,., 0,
a
n J,,),n,·c.w.M - ·7,/~
- t "~ ~ .


" Let us study formation of the embryo sac In a little more detail. . p )II@\~.; h t.06 ~
Figure 2.8b). TI1e nucleus of the functional megaspore divides mltotlcally UTI· ~'r-f r. ~1tc
m~-. rntfdi·
cl'\
.
o form two nuclei which move to the_ oµp os il c poles. forming the 0
~,. -nucleate embryo sac. 1\vo more sequential mitotic nuclear divisions \ uc.Q.e, r:r~·fc.l CJ::t1 ,1--

l es ult in the formation of the 4-nucleate and later the 8-nucleate stages ,o em 0
"> c 'ofi~·_
· ·
f U1e embryo sac. It is of lnteres~ to note U1at ~hese mitotic divisions are G) P~DC\llf\ •
lricUy free nuclear. tl1at is. nuclear divisions are not followed IIiim.ediately ot.n erclll
y cell wall fonna tion. ~ ter_the 8-nucleate stage, cell walls are laid ~wn {!) !'.!s.fci6\c.
~ ~ f\Q.1..i'~ •
eading to the or amsat10n of the t ical female ·gametophyte_ ~ ~ ff' • of'\ ,
r_embry'!,!ac. O~serve the distribu~on of cells inside the em ryo sac ® ~-\f'O~rm·c-4 \J~o')la_,
Figur_e 2.8~, c) . SIX.of the eigh: ~ucle1 cP"e surrounded by cell walls and pQ.um'r:>~ , J)ru.\°' , hl~o.9~,
rgarused mto cells. the remannng two nuclei, called pol~r nuc!!'i are Pot f'lo~ 1 pu.,.mboJ~ c, ,
ituated below the egg apparatu s in the large central ce.11. f'!ro-orn ,·~·
There is a characteristic distribution of the cells within the embryo
ac. Tirree cells are grou~ed together at tl1e micr,opylar end an!,;_onsti~uJ e
e egg ap aratus. The egg apparatus. in tum, consists of two syner1~ds
d one egg cell. The synergids have speciaJ cellular thickenings at the
11irropylar ti called filiform a _aratus, which play an important role in
• ~uid~m'1 he pollen tubes into the s· ergid. Tirree cells are at thecfial'azal
-:id and are called the ,a ntipodals , The arge ~_!:al c~l1, as mentior~ed
_artier. has two polar n~ ~lei. Thus, a i an ios erm embryo sac,~a~
atun,2, though 8-nucleat 7-cellee:, ~Q be. 16 U..Q. Q..,\ 1 CJ:l)t,~ '!SctT1'"e
~ • .,,e_ L_M,·q·-op-,.;Q.ru N\.( d"\o.lO. ~ .. 1
2 .2 .3 Pollination · V · "o.. c..u.•~
. In the precedLTJ.g sections you have learnt that the maJe and femaJe gametes
Ort in 11owering plants are produq~d in the pollen grain and embryo sac,
respectively. As 170th types of gametes are non-m..?!!le, they have to be
brought together for fertilisati,an to occur. How is this achieved?
Pollination is the mechanism to achieve this objective. Transfer
of pollen grains (shed frof!l the anther) to the stigma of a pistil i -
termed pollination. Flowering plants have evolved an amazing ar r j
of adaptations to achieve pollination. They make use of extr 1al
agents to achieve pollination. Can you list the possible rxternal
agents?
1
Kinds of Pollination : Depen~ ~tpn,..gie source of po len~ pollinati on
can divided into three t ~-
Autogamy : In this type, po~nation ~s achi_~ d wtt~~1 the S;J n::
0ower. Transfer of pollen grains from th~ stp~r !'2.~ ~ g1lia ~•_f the
same Hower (Figure 2.9a) . In a norinal flower '.::~e~s ~ d
exposes tfie 1!fl~rs~anoJP.t §..~)grpa ,_oompiu atltogamy is rarher
rare. Autogamy in such (lowers requires,synch rpny ii, ·., }lien Telea~
°¥.<l.5Ligma~ ptrvip and aJso, the.~n q~~~!!!!2 t~~~hou]cl
CCC L('"{i, I

lie close to each other so that self-po


can occur. Some plants such as .
(common Eansy), Oxalis, and Corn
produce tw~ x.p..;;e__s___~ f:a-.....,.Ul\~
chasmogamous flowers which are8
flowers of other speci~ th e~osect
and stigma, and cleistogamous flow
do not open at all (Figure 2.9c). In such ti ;,,i'

/
the anthers ~d stigma lie close to each
(a) When anthers dehisce in the ·flower b'
_,_, pollen grains come in contact With the s I
to effect pollination. Thus, ~ ist~g

r
• ff,:_•_~· ••. . /u'/ flowers are invcl;iably- autogamous as
,.,,,. no chance . of cross-pollen land ing 0 ~
.
stigma. C~~togamous flowers rodu
. .·
assured seed-set even in the absence
.. , .._-
,
r
·, t_r;.
.
_) ' ~ .
~-
pollin~tors. o,'.i"°y ou tiiiiiRlffiifclels~
. , ..,,_,
., :
f'.- ·•-· . 1
advanlag&us or disadvantageous to gener
plant? Why? -, i!ils.l
(b)
Geitonogamy-Transfer of pollen gr · ro!Xle
the anther to t4,,e sj;!~ of =--- the 11
t]Je s~ t _P l,ant.f Aiffioug bl)'llP
- ----:~:;.._.!!l'g
~---=_;:. . ,-
· unctionall cro,ss-po ·
nt, gene
Jie;I
-Chasmogarnous ' :t.:a .c w ~ ~r- ...
flower ~ ~Y: ~..e.tlie.p rA ma
. _/ tlie s ~~g!ant.
.__Jffi) Xenogamy - Transfer of pollen grains grow i
anther to the stigma of.a cli!Igrent plant ~ 1aq\i
2.9b). This is tht_,o_Ely ..!Y.J>e of p,2,l lina~ ji:ha
duri£-g ~!!-Jnation_~~gs geneticall~
types of pollen!~ ~ ~ ~s!,!~a. .
Agents of Pollin.ation : Plants use. ~ o! bioti<J!.. ,,
- Cleistogarnous and.water) and c~ ~-~ti_: (animals) agents to achl
flowers
pclffifation. ~ ~!j!Y of pl~ ts 1;1,s e biotic ~¾ot
pollination. OnJy a small proportion of plants .
(c) - -agents.
a b iotic - Pollen grains coming in contact " t
1
ltii~I
the stigma is achance factor in both wind ~ d wa
· - . . - · tieS SU
pollinatio n . To compensate for this uncert:aJ.Il dll 11.~
ij,.._
Figure 2.9 (;rf Self-pollinat/:!d flowers; - - . f0 l'l lt"''
associated loss of pollen grains, the flowers to tJII ~ i
, ::f;l{l,,Cross pollinated flowers;
. Cleistogamous flowers eno~ous amount of pollen when compare n0~ ~
number of ovules available for pollination.
CTION IN FLOWERING PLANTS
SEXUAL REPRODU

b wffid ls more common


( 'PoUln&tlon y_
..:.--~u pornnattons. Wind pollitlaffim
aiiiongs~ abl~ ~ at the 0~ n grains are..Qs!i!
also require k 5 ! that they can be
n -stlc_!)'
and no .. .-1 ... d currents. They often
-c= rted In w 1u
transpo ed stamens (so that the
posses~ w~cli:j)ersed Into wind currents,
pollens are d large often-feathery stigma
2
Figure ·!.;p:-boriie ollen a1nJ. Wind -
to easily owers often ~ave a single ovule m
po · .. and numerous flowers packed Into
~ch ~ c e · a familiar example ts the ~
an infl .,. ' thing b t th
cob - the tassels you see are no u e
- - and style whlch wave
stigma - In the wind· to
trap pollen grams. Wind-pollination is quite
common in gra.sse~- -----,..... .
Pollination by water is qu ite rare in
flowering plants and is limited to about 30
genera, mostly monocotyledpns. As against
ffiis." you would recall that water is a regular
mode of transport for the male gametes among
the lower plant groups such as algae.
bryophytes and pteridophytes.lt i~ believeo;.
particularly for some pi:ytjphytes anGl Fi~2.10 A Wind-Rollinated plant showing
pteridophytes, that their distribution is limitt ,d CQ,mpast jnftorecence and well-
Because of the need for water for the transpo1t exposed stamens
of male gametes and fertilisation. Solne r--:f \Oft \ t\ll-2

f'~'~
(!r~t.j
e'xamples of water pollinated plants _are ~ lliJne~ a:hct drillh whic ~ lb~
grow in fresh water and se! eral marin! ~'!;~ s such as ster; Not ~ac -
all aquatic plants use water for pollination . 11.l._~1:lajo~ty of aquatic nlants c,btNe I
rt~
po~o.e.

arw
-
'SUchaswater hyacinth watf[ lily. the fllJFSF~.eru~rg~,~ ve the level '
of waiefaiicl are pollinated by insects or 'ivind as in most of the l"-.'-:-d
piants. In Vcillisne[ia, the female flower reach the ~~fa~~~aftr Sytti'e '
.. . .

long stalk and the male ~owers £LE.2!!~!lJtr~~~~~~:,~ed on to the


. surface of water. They are s_,an:te~ e a ~si~~ly by water currents (Figure
~- l la); some"of them eventually reach the fem.ale flowers and the stigma.
In another group of water pollinated plants. such as seagraslies, female
fl~~ remain submer&ed In water and the pollen f;:~s are rel~
inside the water. Pollen grains in iii.any such spe'"cies a~e long, bbon like
and they are carried passively Inside the water; some of them re ch the
sti~ and acffl'"~ ofiina.Mo}i:-ifi~t ~t~'"water1~a,¥as
grains are protected from wetting by a muci!ag!:aous cove g. !' h~ ·. _L.~
Botii'Wfii'd and ; at; pollinated fi~wers ai=~ ~ ot ~ I)· coi~ ,_
;rtuf cl do
~29~ nec4tr. What would be the reason for lhis? . -- - -
&.~ ' P'~"
~ I \ i'r, lu~a.
(~ODJJ'll")

z
~s 01.NO
· f"~ -
!'lc
7l'
fo/Z_Q •nett\ '"
p(!a.nt ':) . ~e\J~ /
Zdx5l5
sec::b
'M<! cf\CWJ.S.. w. .
i\J.~

~ a>+t)~a . (Fr,;_,:,..r&
· . . . nn I V
rtt
l n.JC!..d---
81
. c_~ ~ t OG'I O CZ,V O VU,..)1-1! • M . .ty [fl
__ --= ::::: ,.. - aJon o oweruig
+ I 'Female .,. .,. .. )~ ' ) a range of ani.tna]s as ; 1clt\i.
~ , f() t'cNrbtm . flower
\ agen ts . Bees , bu tte fl .0 11Jii
. Cd"'t-d.lJJt o r::-::- - -- r tes
,t b eetles. wasps. ants"7i'- ·
sugma - . _ _ . 101:hs'
,, 1 i..+cr, (s~·cts and liunirnin~ir '
'h:-.e. ahl(1 i?>Q) • b ats are the common"' ds]
- . --~ Po
• ~ •~n::,t_oct\1'8
,L ~mri~ .dagef!tS. (F1gy re 2 . 1 1b).
fan imals: nn sec&:;, parfi~
t~ oQ,Q / · rarelhe d ominant '6ic5Bc po
. agents. Even larger ?-Uilnais
~ ~-. Male fl ower Female t )is some primates (leitiurs)
flower Q (tree-dwelling) r odeii'is · ·
,as:w ~n l a --.:---i----~~-- - , 0 r
pl..a11. t-- r epWes (geC'.ko ) iz~ d and g
t,o:x .. ~ izard) h ave also b een reporte(!
t· poffinators in s ome species.
(a )
O ft e n Gowers .9f anilll
pollinated p lants are specifi
adap ted for a particular spectes
animal .
·Majority of insect-p?lltna
flowers are l~e. colourful,
and rich in nectar.
----- ------
When
are sma l!_,~umber of flow
the fl

make them conspicuous.


~ tt~~~t~d- -t ~ ers by co
and/ or fragrance he
p flie
secre e o o our s to a

--
t animals. To sustain animal
the flowers have to provide re
to the animals . N<:>clar and
grains are the !cllil 1a0}9!.a l r~
(b ) Forti~est.ing the .reward(s)
th e flower the animal visitm;
Figure 2. l ~ ollination by water in Vallisr1eHa ; in contact with the ant.hers ci\Jld
Insect pollination . . stigma. The body of the anim~

..t< r • •30 a coating of pollen grains. wh:~,p


genera By sti,b,y in animal pollinated flowers . When the anim_al cariiii
"', ~• r ' - ,.
pollen on its body comes in contact wilh the stigma . it bnngs 8
pollination . et
In some species floraJ rewards are in providing safe places to lar,;
an example is that of th e: lall f' st llowrr ol Amorplwphallus (the ~
itself is abou t 6 ff'et in IieiFWi. A similar relationship e5sts bet\~
spec ies QI ~ h cllld the_pla n:_:} ir ,a wh ere both species - moth a~

~ubo. "'-otn.
-------
SEXUAL REPR Oo
ucnO N IN FLOWERING PLANTS

lete Uieir !if w· Uiout each other. The moth


t cnnnot comp
P1an - ' s c ,Qs in the Jocule of Ole ovary a11d Lhe n~wcr, in urn . geLs
depQ§lts 1 TI 1001
Tue larvae. of U1c 1110LPfcm nc ou I ol the eggs as
pollinated by ie n .'
,d.:zrSl1\rt de,relopm[_ .
tl:e S N · . observe some flowers oj the following planls (or any
Why don t yot 1 . .
vailable to you): Cucumber. Mango. Peepal, Conander. Papaya,
oUl~rs a ia.. Cotton. Tobacco, Rose. Lemon. Eucalyptus , Banana? Try to
Onwn.~ ·ch animals visit them and whether they could be
f ind out L.V 111 fl ·
v: u'II Jmve to patiently observe the owers over ajew days
pollinators. 10 ·
;FF. ,,,ta.mes of the day . You could also lry to see whether there
and at dw ere,'
. orrelati.on in the chw:acteristics of a flower to the animal that
t~ ~n~t cearefaUy observe if any of the visitors come in contact with the

: : ~ and the s~gma as only such visitor~ can bring about pollination. Q..i..ltOJ) al-· ~D(l) ~
ec 5 consume ollen or the 'nectar wit1wut 1:iringing about } frtt1" ,~ J • Q~01-t
JJ<!llt,wtfDn. suc1lfli5rid visitors are nfferred to as pollen!'.!;;_~ robberS . ",LI.I'~• t <f'UJ>c-!I 't
You may or may not be able to ident{Ty the pollinators. but you will surely 9...+-lae \j N_ct\e.rf .
e11_joy your efforts! . ·
~w.::u ~ ·CJJ...)
Outbreedin,g D~vices~ f f i o w eringplants produce herma fuoclite ~ f ' I · ~ ~~ \ , , .
11owers and pollen grains are · e to come in contact with the stigma of 1' · tto'l'
. e same owq. Continued se~-E.oJ?1~on result in in1a_rees;)ing.de.RL~~$.ion. rtend.~ Oto
~lowering plants have ?evelopea man~ devices to discourage self- · ~/
pollLnation and to en~ourage cross-pollination. In some species, ollen l>lc.H ~G,~
release and sti ma rece tivi · . Either the pollen is ·
released before the stigma becomes receptive or stigma becomes receptive
much before the .release of pollen. II,l.some other species, th! ~ th~r !11d ~
- .
51
stigma are placed at different ositions so that the oUen canno!.._:?~:..u · -, I ..... rrw.lo\I \J
contact wi the stigma of the same flower. Both these devices prevent
a_utog'imy. 'Il}e third device to ~revent inbre~1in,.S,i,~ s![-i? SOT patib~l! ...., . .SSI
This is a genetic mechanism and prevents self-pollen (from the same flower ITT . f?" u·
or other flowers of the s~e plant) from fertili;ing theovu~j~~~~~~
po.llen ~TI.$11_mlo n or po~~ t h e . . Q i s . t i l. _A nother device to
l ~"'T~
·
eiir
prevent self-pollination is the production of unisexual flowers. If both male Q:Qof\a.~
and female flowers arepresent on the same plant suffi~ castor and maize t t't J'a~
lmonoecious), it pr~ents autogamy but not geitono amy'."insevera"ispecie~ Br:s_~~~ . rooc..w.e,
such !ls papaya, ma e an em e owers arc present on different plants.

,re, ,
that is eacnplant is eith~r male or female (clioPcy);This condition prev~nts .
both au~ gamy and geitonogamy. . frepott p ,1 = pe.,_r . ~,
7tif'' S lfl'lA,..)
L
•--1<1>

Pollen-p1stil Interaction : Pollina tion does n ~ guarantee the transfer \j


of the righ t type of pollen (compatible pollen of the same s pe cies as the
stigma). Often, pol~en of the wrong type. either from other species or from
th e ~mne plant (if it is self-incompatible), also lan d on the s tigma. The
piSlil has the ability to recognise the pollen , \-vhether it is of the right lype
1 (com~a~ble) or of the wrong type (incompatible). If it is of the right type ,
th e pistil accepts the pollen and promotes post-pollina tion events that

'•
If
Egg~

SYner&ld,

Longitudinal section of a flower


showing growth of pollen tube
(a)
(b) (c)
Polar nuclei
Central cell
Egg,-tt---,,..,~ )
Plasm a •
nucleus ,,.__,.. c:!l~---tt:7 Male gametes
,=--rHr-Synergtd

IJ.lllfOl".tt--.F iliform
·apparatus
Male gametes
Vegetative ·
nucleus
(d) (e)

Figure 2.12 -{a) Pollen grains germinating on the stigm~ Pollen . tube~._growing througi/ t
style~ L.S. of pistil showing path of pollen tube growt.b,;,id) e~arged view of al
egg apparatus .showing entry of pollen tube into a synergid; , C e( Discharge of malt
gametes into a synergid and the movements of the sperms . one into the egg an!
the other into the central cell

leads to fertilisation. If the pollen is of the wrong type. the pistil rejects-th
pollen by preventing pollen gennination on the stigma or the pollen tub
growth in the style. The ability of the pistil to re<;:ogp_tse the pollenf~
py its acce ta:nce- or~f e ectlon is the result of a continuous dialog¥
between pollen grain and the pistil. This dialogue is db c e
compolllints of the p ollen inte~ cting with those of the pistil. It is 0 n1Y 1
recent' years that botanists have been able to identify som e of thepolle
an0 pistil components and the interactions leading to the rec 0 e- ...... 1t1or
followed by acceptance or rejection.
As mentioned earlier, following compatible pollination. t h ~
germinate,!i..Dn.ihe stigma~to-pn >auee-&- pollen..t.ub, oullh one~
. ~ w
germ pores (Figure 2.12a) . The contents o e po en grain move ill
.a,.,~'(ft:~'." 1l t~" [ " ,, "" "' t& -it -\ti
EXUAL REt RooucnoN 1N FLOWERING PLANTS -Gtrtlo) n ?ol..Q_Qu == d)Q.rn4c. \'-
'i' a,u\4Q.. R-eo. <;:~ •l\o~t\o" \fo'!' ~ :,q C\v - CIJ

ollen tube. Pollen tube grows th'ilou~~~ :nr;ues dfth~stigma and E'"""?'\
le and reaches the ovary (Figure 2.12b, c). You would recall Uiat In --,.uk:R \JI~ O
ome plants, pollen grains are shed at two-celled condition (a vegetative
ell and a generative cell). In such plants, th'egenerative cell divides and ~ chtb3" lf'I~~
rms the two male gametes d the owth of ollen tube in th sti J.. (M
n plants w ch shed pollen in the three-celled condition, pollen tub<;_s ~uonl\,O,
11. e be · . Pollen tube, after reaching (l'l')<M- C......... , Au...
·'II (o(l)ITtll, Ul'L-U;IJ Q<l"I
e ovary, enters the ovule through the micropyle and then enters one of - -r'C'.ii~:J:i"'lj':""":'~ fl.a
es e ·ds throu h the filiform a aratus (Figure 2.12d, e). Many r ~£ iR,t,, !J. ~ -
tudies have shown that flliform apparatus present at the micropylar part r.'°'U 0 ~ ,
fthe s er "ds · the en of ollen tube. All these events-from ~rrFHjw-ldtt.. e.~
ollen aeposition on the stigma until pollen tubes enter th vul are ~ ~ sae ~ ,
ogether referred to as ollen- istil interaction. As porn e er,r'lfl°I' . eJ'd ,
ollen-p1stil interaction i~ a dynamic process_!Qvolvin_g_pollen recogni,tion
ollowed b romotion or inhibition of the olle~ e knowledge gained
this area would help the plant breeder in manipulating pollen-pistil·GlQ:,·~f\O:t-\~: t~
teraction, .even in incompatible pollinations, to get desired hybriqs.
You can easily ~tudy pollen germination by dusting some pollen from Mutl•d ·
ow~:5 such as E:_a, chic){pea, Crotalari.a, balsam and ~ a on a glas~ ,p,~ ~8~ , " , es;_ ct. '°"fl~
ontaining a dro of su ar solution (about: 10 er c~t). After abouj( J _ . !~ -
. utes, observe the slide under th lo
_.. ns of the microscope. You
likely to see poll~n tubes coming out of.the pollen grains.
As you shall learn in the chapter on plant breeding (Chapter 9) , a

-
reeder is interested in crossing different species and often genera to ,,
combine desirable characters to pro9-uce commercially 'superior' varieties.
ificial hybridisation is one of the !!!,aj_gf _app_Jo_a_£:ge~ ~
1
. @provefleiit programme: In such cros~ing experiments it is important
cl to make sure that only the desired pollen grains are used for pollination
and the stigma is protected from~ ,!;,.~ a~ ~nJfy~~':!n~~~2!k_n).
This is achieved by emasculation and bagging techniques. ·
.,.·-x."'.!!t~
If the female parent bears bisexual flo~~!:._~_,Are1;1_oval of ant!,i~rs_f!:~ ..
- -~ - - ~ \ l t l S - " ~ <-~ •

the fl6wer efore the anther dehisces . using a pair of forceps is r . J


-----. -..~----- - .
neces~. This step is referred to as emasculation. Emasculated flowers
pave iab~ covered with a bag of.slli_ta6jf.~~e, gen'erally made up of 2_~ r
paper, to prevent contaminai,io.n .ofit&.stigma..wtthunwanttd.P.Ollen. This
'- .
process is called bagging. When the stigma of bagged flower attains
receptivity, mature pollen grains collected from anthers of the male parent
are dusted on the stigma, and the flowers are rebaggeo. and the fruits
allowed to develop.
If the female parent produces unisexual flowers, there is qo need for
emasculation. The female flower bud~ are bagg~d before the flowers og_en.
Id! When the stigma becomes receptive, pollination is carried out using the
ie de~ired pollen and the flower rebagged.
--n'tfi ~ ~\A ~ m.
. 'l~ ~ ~ I C \
2.3 DoUBLE PEaTJLISATION · '\J ,0cJ.,'OJLlffi
<I" ~ •

• 'o)
After ente1i n one of U1e s er idt:, . the ollen tube releases the tw
gametes into the c:,·tnpbsm of U1c · rgid . One of the male g~
moves towards U1e egg cell and fuses w 1, Ii 1 t '- nucleus thus completui_ 1
Z,i<coue<ui "':I r syngainy. This results in the formation oi" diploid cell, the zygot i ~
o-«>s:c °'"','!'!-' , u· other male gamete moves 1owards the two polar nucleilocated in the ' •

1
\
:!j .m!f/',J.r
... HI,
cell and fuses with tl1em to produce a biploid nrimlln'. endosperm n"'ru,, .
\PEN) (Flgun, 2. 13a). As thls involves the fusion of three haploid n~ :•
1s"termed triple fusion. Since two types of fusions, . syngamy and.!'1!1
- -l!ll!~
I Pl.. '.1ill.J fusion take place in an embryo sac the phenomenon is termed d tl'j
*I<'%...,~..., . fertilisation. an event uni ue to flowering plants. 'J!!e ceritral ce~

~\l
RiJb«'V~ 4· ) ' " ~tp etl:

I
t>ot9w-. ion becomes the primary endosperm cell (PEC) and~
-\QI, I • .· m O 1e endosperm while the zygote develops into an em~

1 ®ti'- Degenerating - ·
synergids ,~ cJJ
oL~ . . r_ r-o-u.d&f\.l.lf() / \l tt.i ~u.tl1
, Zygote (2n) -fl...;_lj• M_~( C) r,M- {'-- \ (I •••J
QJU..e.ll'

-'
I \ r~ -1:)
Primary endosperm
cell (PEC) , Suspensor
Primru.y endosperm
nucleus (3n)
(PEN)

Cotyledon
eart-ShapedI
Embryo
-~ -
Degenerating - ~-
antipodal cells (~ Jr•tu t,) Globular m2!2'.?
r Plumu~
rvlature
(a) .,- 'r -- .-- ,-, ... (bl
rnbi>f'P~ ..u)l 4 <m'
fc
L Figure 2 . 1~

';:j
Fertilised
. embryo sac showing zygot e an d p nmarv
) S tag~~I'}\ embryo development in a dicot [ h
~J,:i'I'. , / ,t
: \j"""Endosp 'ii- \ o11:~ l~ ( 'IEN)'
.
<"o/19'"'
-
p-:.
erm uc eus P :

ip,·1,~~,. . .t-
11
;'.-~p'j'.~iu~ed size as ompruced to(~
u d~ 1.9..i o,...\.
2 4 'L
P \½f " ~ 8 M<\ ~ ,. ' (- ~Q.,rt J. ~ b,Q ,
• ost!.FE~ll,ISATl9N ; &J:JltJCT~ AND E\TENTS
Following
d 1 double fertilisation ' events 0 f en d osperm and embryo.-•

.
} 34 eve opment, maturation of ovule(s) into seed(s) d . ..
( 't,
I
, .·'. collectively termed post-fertilisat1·0nevents. . an ovary mto fni1t, are

orcb'l,'d eb ~ .

!
l
-flt- W\

~ e..,o,,~
t • fod.o~-tttf\o c.eor- 2 .4 .1 Endosperm
• -u-1c:1 cPo.e .
_ oer,ath ta .
'En dosperm deyelopment precedes emb.ryo develo
P ry en osperm c~lL!;iJ.'tld.es ~ a e an

I f
1
/ <Y\~ o-5.'f ~b'ffi

·e.~>tW"D
I 8eW M+
roQJrn .
·RiU, WN.a.t- "' ~rcl\.j
.
-~
""°\3
t .,, PnAf l'Nt+~
, Coc.Qf\u:t ::: o,''ti
- - - -::..:.:..:.:-

) •
· s-1• ·
i ~ £ 1~ I
.2:, !::>Oll..m tW..m,'c~o~.
I I· I I) ,/.}
gJ
SEXUAL REPRODUCTION IN FLOWERING PLA NTS
\ .
-.:
en do ~perm tlssue..t h e cells of thi s l issu e a re, lilkd wi th ~__:,,,-------- Plumule
rt'...; 1-rv1 food m a terials and arc 11 -rr1 fur til e. nul rition of
t he deve lopj n!' cmha:0 1 In th <t_ n , ....,, cummrJ11 1 w
end osperm devel opment. th f', PE N 11n ergoes successi ve ~....._,++-- --t;?Cotyledons
n u c lea r divl~lons to give rl~ <' to fre e n u clei. This s tage of
en dos perm development i s called fr, t 111wl•"1r ,,,, <Jo• 1wnn. J-Iypocotyl
Sub se qu ent ly cel l wa ll for m.;'uon oc c urs 3nct Cfit:
1

en doseerm bec9 m(''-> r1 llufar. Th <.:. Ill\JlM,ler of free nud i;:i


for mrct b rfor<' rellt1bri""lliot1 vari es greo1Uy.'¥be m onu'r
l#nte-r from tender coconut that ou are familiar with, l s Radicle
:(fothtng b ut fr e e -nuclear endosperm ma e up o f i..~~-Root cap
'
thousan s o an the surroun white e :-is
tli: cellular endosperm . Pt:h.tf"'o.
(a)
Endosperm may either be completely consumed by the _
developing emb,!Yo (e.g .. pea, groundnut. beans) before seej.
maturation or it may gers~ t in f!ie mature ;ee~ (e.g. casto~ --1--- Scu tell '-!m
and c;oconut) and be used up during seed germination. Sp,it
open some seeds o} castor, peas, '6eans, grounanilf.'fruit of
coconut and look for the endospenn in each case. Fitl.d out

and maize. IV
whether the endospenn is persistent in cereals - wheat, rice
(!> <IJrt i&..bUJ1111'<u>u..> .

2 .4.2 Embryo
lfmb~~ lops at the micropylru: encLoLthe embryo sac wh~re,_.
the ~ ote is situated. Most zygotes divide only after certa.il!.,
amount of endosperm is formed. This is an adag_tation to
provide assured nutrition to the develo in emb o. Though-
the seeds differ greatly, the early stages of embryo development
(embryogeny) are sjmj)ar ip botlJ. WOPQGOt~ n ~ : a,,
dicotyledons. Figure 2.13_depicts the stages of embryogeny in
a dicofyledonous embryo . . T!!; ~_:>t!:, Si':e~ :!_~~.:, Coleorhiza
_proembqo and subsequently to thS, globular, heU1-!iha~d·
and mature embryo. (b)
typical aicotyledonous embryo (Figure 2.14a). consists
of an embryonal axis and two cotyledo,pj, The portion of Figure 2.14 (a) A typical dicot
embryo: (b) L.S·. of an
1

e,Elbryonal axis above the level of cotyledons is the epi<:_otrl, embryo of grass
which terminates with the plumule or ~ m ti)?• The cylindrical
portiorQ ~Q§£Jbe level a( cotyieaons is hypoco!J;l-_truif .. ..;!"'

terminates at its lower end in the radicle or root tip. The root
. 35
tip is covered with a root cap. - ·
~as of monocotyledons (Figure 2. 14 b) possess only
one cotyledon. In the gra~s family !,t1e cotyl~op i_s _sall ~cl
, scutellum that is situated towards ..Qll.Ll!<;!L_Ua ~~-rs)l pf.-the
·Z?1bryona! axis. At.its lower end , the e,!llbrYonal~ ~ ~ t he
' <: ------ '
\'f
~)
'X'eo,·Q ~at--
· radical and rooJ, cap enclosed lo an u
Of'\
ffn,~i l1flrlto..u,
(~~) t f\Cb . c = _ _ _ a. The P..: :.O.:.: '
rt:;:;io::.:n~-..:o;.;.f-==--=:.:=.:,==~;_.;;.,;;;..;;:::::.::.;:.::..._:
o/1~ ""°-''S- c
t)
attachment of scutellum is as a shoo

1
t
few leaf primordia enclosed cture, the
'"1.°'i~1'Q.-;) cttt{
, (t sw, ) t~~ b"Soak a few seeds in w er (say of w a , maize, pe
ground nut) overnight. Tilen split the seeds and observe
r =-- - c----;n~ parts of the embryo and the seed.
~ d f - n« e&.\
2 .4 .3 Seed
· I 1.-n
-:.~an
~ gi;'o::;s::!p:.:e;rm
. :.:::s;..·th
.::.::ei.;s~e~e~d~is~ ~
t he~fin
= al ;.;.r::od:;:.u:;:.c~t~o~f.:;s~exu
~ al
~ r~et!;r~o::;d~ul9
ction, lt
Rum,nott Chf).L) eq ~Q . often described as a fe sea ovule.7 Seeds are formed inside fruits
R.o\..i.fth . (J_I seed typically consists of seed coat(s), cotyledon(s) and an embryo ·l,

~;f
~ t i ~ flT'


~ vQ.~s,~ Qo.-a_' \J . The cotyledons (Figure 2 .15a) of the embryo are simple ~
struct,~l(f~

\I enerallvthick and swollen due to stora e of food resezye,~[~ ~gu~


Moscu~
I'\ :r~sy~ -t
II"{)'\
Mature see s may be non-albuminous or ex-albumino~
albuminous seeds have no residual endosperm as it is compJet
fo\-<lU...i ~-._,, clJ
i
consumed during embryo development (e .g ., ea, groundnu~
c_eil_t1 Jo.:t---c.ho .I ~buminous seeds retain a part of endosperm as it is not comp
~1.,QL\.u I ~tt-e · . u during embryo development (e.g.,wwheat, maize, barley, caste
\,~sttf'f'(T) Occasionally. in some see s such as black w~cx: auci fieeftemnants
'-n r_ nucellu&We also pel'ststent. This residual, persistent nucellus is
·1 ~ - -..
l . l'tf\~on . S:\1.h ~"'1~ ~cf'6 Integuments of ovules harden as tough protective seed coau
ci ll) ~~u m · (Figure 2.15a). The micropvl<;_ remains as ·a small pore in the seed co~
;i,... CJ This facilitates entry of oxygen and water ~t~!h_e~ d durin · enninati<Xi
I I \}
Cl.\\-lS\JJT) ·
- -=--------..,;J,J -- " - - \. . . _____........._____
As t1ie seed matures, its water content is reduced and seed,s beco
I.
r~lativeJy dJX (10-15,ps.r _cent moisture by ma§~). The general metabo·
I activity of tl!.e i , ~ ~ ~ ~own,._ , The embryo may ~
j inactivity called dormancy, or if favourable conditions are availab
(adequate-~ oisture, oxygen and suitabieterri.perature), they germinate
t As ovules mature into seeds, the ovary develops into frin£, te~ a
transformation of ovules into seeds~wl. ~y~ry into ff!Iit procee
simultaneously. The wall of the ovary develops into the wall of fruit c

.ts~c-
I I _ _:_~_ _:_-'!i~=~:::;;:I.;=:~ pericarp. The fruit~ mar be fies~ as ill g§ va, or~ge, ~ go, e~
' fi .,jl "'- ":~~ 1 may b e 2!)', as in groundnu~ and ~ . etc. Many fruits have evolv
I j mechanisms for dispersal of seeds. Recall the classification of fruits
\. ~ their dispersal mechanisms that you have studied in an earlier class. Is
there any relationship between number of ovules in an ovary and IN
number of seeds present in a fruit?
· In most plants, by the time the fruit develops from the ovary, otbeI
floral parts degenerate and fall off, However, in a few species such as a~ ·
strawbeny, cashew, etc., the thalamus also contributes to fruit foilil!tiO!l -
Such frui~ e- called fals~ fruits (Figure 2.15b). Most_f!:uits
develop only from the ovary and arecalled true fru!ts. Although.ID JllO~ ciel
of the species, fruits are the results of fertilisation, there are a.few spe
iEXUAL REPRODUCTION IN FLOWERING PLANTS

Seed coat
.r 1cropyle]
· Endosperm

Cotyledon hoot apical


meristem
Hypocotyl
Cotyledo::.:c
ns; ....__ root axis
r~~1 fCf\-s--rue._]
-
Enqospe

oleoptile
Hypocotyl Scutellum1----1J.....a:1--
root axis
_.,_-Radicle
Root tip
Endosperm
V.-- - ~ '-1--1---Coleorhiza
-~='=\=-t__N_<-_t~
_ l_9N IG~

Thalamus

\
: j
\ j

''-
\

Mesocarp
(b)
- .

Figure 2.15 /structure of some seeds.jtr) .....--,-,.,,....,,...,_


False fruits of apple and strawberry

in which fruits develop without fertilisation . Such fruits are called


parthenocarpic '" ts. Banana is one such example. P~~~S,arp~ can
e induced throu e ap · lication of growth hormones an~ ch fruits ..
are seedless. US~ ,Rcr coC.O n.\rl-/ poflr}'f ,
Seeds offer several advantaMes to angiosperms. (6irstly. since
reproductive processes stFch as pollination and fertilisation ar~
independent of water, seed formation is more' dependable. Also seeds have
better adaptive
. strategies for dispersal to new habitats and help the species . .
e,o,
to coloni!',e m . a·s, · As _thev
. ot 11er rue · have s111Ticir11t
l f l lond 10 1;:i~r v e
t 8, v
seed lings are nourished until they are ca~ab e o P 10 osyn lhesis on 0
O\.\rn . The hard seed c;;oat prni4<l~s protcclio_n l~ the y~u~1~ e~ . t
products of S C'}.." l~ll reproduction . U1ey eneiatc n ew Cnct1r conibin~i
leading to vruiations. .
~l·t·d i:-- tlic basis ol ~ur ag1i<'td Lure. Och dr~llon and dorm.~
1 miJ( u rc seeds are crucial for stora e of seed_s wl11ch can be usect as
1
throu<1'hou t the yeru· and also to raise crop m t 1c n ex sea . a
imagine agricu lture in 1e a sence o·t. see d s , 01 ~ m . e pr ce of ti
r which genninate straight away soon after fo1?1iation and cannot be SloJ:
How long do the seeds remain alive alter they are dispersed?
I

period again varies greatly. In a fews ecies the seeds lose ~ ~


oxo.Q} !l .<- a few2;;onths. Seeds of-; 121"ge numl:5er of species live or severa} y~
. Some seeds can remain alive for hundreds o_f years. There_a r ~
records of very old yet viable seeds. The oldest 1s that ?f a lt;!pip e-.l_~
arcticus excavated froiu Arctic Tundra. The seed ermmafecranct fl~erl;
a.fter"'an estimated record of 10,000 years of dorrpaocy. A recent recor ·
2000 years old~ able seed is of the date palm, P h o e ~ ~
discovered during the archeological excavation at King Herodj Pil a'~
near the Dead Sea . , ·
· After completing a brief account of sexual reproduction ofJ1owe~
plants it would be worth attempting to comprehend the enorma~
reproductive capacity of some flowering plants by asking the Jollowifc
questions: How many eggs are present in an embryo sac? How mM
embryo sacs are present in an ovule? How many ovules are present fu
an ovary? How many ovaries are present in a typical.flower? How ~
flowers are present on_a tree? And so on.. .
Can you think of some plants in which fruits contain very lar!Ji
number of seeds. Orchid _fruits are one s_us:fl. p];eg2_ry_ and e ~
contain thousands o[ tiau seesl§. Similar is the case in fruits of son'l
_earasiti.c species _such as Q!o,!:a;2che and..§t111Ja. Have you s-een a ani
seed of Ficus? How l:3![J!_i!j t/fe_tree of Fic;JE,_ d_~veloped [9TT]Jha!
seed. How many billions of seeds does each 'Ficus tree produce? Can
ydiL imagine any other example in which such a tiny structure can
~ Adve~ produce such a large biomass over.the years?
·a,,,.. ·
, (~~-t~ \ ~ .5 JAPoMIXIs AND POLYEMBRYONY _

'f... 1
~ gh seeds, in general are the products offertilisation, a few flowerW
38 ~ec..u.~ plants such as some s ec1es of J\sleraceae and ,Irasscs. h ave evo ve 8
~ Ji speci mechanism, to produce s('ed,s without fer · is;~J~';', called a iiJ]j:is.
•------.• lJ · a ts ruit prod uction w ithout er i isation called ? Thus , apomLxiSis•

I ·
(Jorffi:'of asexual reproduction that inimics sexual renro<lucLion . Ther;_!~
several ways of clcvelopmen t ol. apomictic
- - ·· ., seeds. In some s pecies. . . thl
diploid eg~ cell is fo m1ed without redu cUo n divis ion ·a1id dcvt'l uµ51ii1r
1§1
t,:e"embiyil wflh;:)Ui Ii sa~on as in nffii'iy Cirn ,_;:" • '. , ·.1 )
;2
SEXUAL REPRODUCTION IN FLOWERiflG ~TS

s rrounding the embryo sac start


dividing. protru e in o e em ryo s c an eve op into the em ryos. n
1!1:0 of more
such species each ovule contains many em b ryos . Occurrence -
than one embryo in a seed is referred to as polyembryohy. TaJce out
some seeds o.[o~ and squeeze them Observe th't:many em15ryos of
stzesaruI's
hapes @iij"r;agi seedi,Count the number of embryos
in each seed. What0 wou'f!- be the genetic nature of apomicttc embryos? <,,-n)
Can they be called clones? ·
Hybnd varieties of sever ~ f our food and vegetable crops are being
extensively cultivated. Cultivation of hy~rids has tremendously increased
productivi~ ofthe problems of hybrids is that hybrid seeds nave
lto1ie roduced every year. if the seeds collected from hyqrids are sow1\¥'
the Iants in the ro en will se e ate and do not maintain y
(cliarac ers. Production of hybrid seeds is costly and ~ nee the cost 0
hybn1i'seeds become too e ensive for the fanners. If these hybrids are
"irufcreiiiro apomicts, there is no segregation o c aracters in the hybrid
progeny. Then the fanners can keep on using the hybrid seeds to raise
new crop year after year_ and he does not have to buy hybrid seeds every
year. Because of the importance of apomixis in hybrid seed industry,
active research is going .o n in many laboratories· aroun e wo
· unde;stantl .ffie enetics o - a omoos an . t'ffimt~ nes-

SUMMARY \:4;;:
J ~r

Flowers are the seat of sexual reproduction in angiosperms. In the flower.'


androecium consisting of stamens represents the male reproductive l
organs and gynoecium consisting of pistils represents the female J
reproductive organs.
A 'typical anther is bilobed, dithecous and tetrasporangiate. Pollen
grains develop inside the microsporangia. Four wall layers, the
epidermis, endothecium, middle layers and the tapetum surround the·
microsporangium. Cells of the sporogenous tissue lying in the centre of
the microsporangium, undergo meiosis (microsporogenesis) to form
.tetrads_of microspores. Individual microspores mature into pollen grains.
Pollen grains represents the male gametophytic generation. The
pollen grains have a ~o-layered wall, the outer exine and inner intine.
The exme is made up of sporopollenin and has germ pores. Pollen grains
ma:y have two cells {a vegetative cell and generative cell) or three cells (a .

I
vegetative cell and two male gametes) at the time of shedding.
The pistil has three parts - the stigma, style and the ovary. Ovules
are preseqt in the ovary. The ovules have a s_talk called funicle, pro~ective
. integument(s), and an opening called micropyle~ central tlspue I~
©!e nuceiius In whlcH tlie -arches ortum differentiates. A f ell of tlie , ·
archesportum the me as ore mo er ce div!_des meiotic ~d one of
t e megaspores forms the embryo sac e m ale ame o yte). Tlie
ma1ifre embryo sac is T-celled and 8-nucleate. At the micropylar end is

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