UNIT 6 SEED AND FRUIT

Page No.

6.1 Introduction
Objective
6.2 Seed
6.2.1 Parts of a Seed
6.2.2 How Seed Develops
6.3 Seed Appendages
6.3.1 Aril
6.3.2 Caruncle
6.3.3 Operculum
6.3.4 Wings and Hairs
6.4 Stored Metabolites
6.5 Development of Fruit
6.5.1 Follicle
6.5.2 Caryopsis
6.5.3 Legume
6.5.4 Berry
6.5.5 Drupe
6.6 Dispersal of Seeds
6.6.1 Autochory
6.6.2 Anemochory
6.6.3 Hydrochory
6.6.4 Zoochory
6.7 Parthenocarpy
6.8 Vivipary
6.9 Summary
6.10 Terminal Questions
6.11 Answers

6.1 INTRODUCTION

Sexual reproduction in flowering plants requires two processes-pollination and

fertilization, about which you have studied in Unit 3. Following double fertilization, the
zygote develops to form the embryo. The Primary endosperm nucleus divides and the
resulting nuclei further proliferate to eventually give rise to the nutritive tissue-
endosperm (Unit 4). Meanwhile the ovule undergoes a series of changes that transform
it into a protective container of the young sporophyte (Unit 5). In this unit you will
study various aspects of the development of seeds and fruits. A seed may be defined as
the unit of reproduction which has the embryo and commonly food-laden endosperm
enveloped by a seed coat derived from the integumentb) of the ovule. Beginning with
pollination, the ovary is activated to form the fruit, which encloses the developing seeds.
The fruit and the seed not only protect and nurture the young sporophyte, but also serve
the function of dispersal. Seeds of many plants remain viable for long periods in the
soil. They may even have a certain period of dormancy to ensure germination only
when conditions (temperature and moisture in particular) become suitable. Incidentally,
the food stored in the fruit wall and seed are also the main source of nutrition for man,
wild and domestic animals, bacteria and fungi.
Objectives
After studying this unit ,you should be able to:
explain the structure and development of the seed;
describe the various modifications in seed structure for effective dispersal;
know the nature of reserve materials in the seed for the nutrition of the young
sporophyte during seed germination and seedling establishment;
trace the changes that occur as the ovary develops into a fruit;
 classify the various types of fruits; Seed and Fruit

elucidate how some fruits such as banana develop without having seeds;
describe the relatively rare but interesting phenomenon of vivipaq, which involves
in situ germination of the seed while still enclosed and attached to the parent plant.

6t2 SEED
-
A seed is a mature ovule enclosing an embryonic plant, stored food material (in
endosperm, persistent nucellus or embryo itself) and a seed coat formed by one or two
integuments. In h broad sense the term seed is also applied to small one-seeded, dry
fruits (e.g., grains of wheat or barley which are in fact made up by fusion of fruit wall
and seed coat) or other disseminules (fruits with attached bracts, inflorescences or even
vegetative structures such as tubers and bulbils).
The size, shape, colour and surface of the seed show innumerable variations. Most
orchids have minute seeds like dust particles. Seeds of a majority of flowering plants are
a few millimetres in diameter (e.g., mustard, guava and poppy) or extend to a length of
about a centimetre (e.g., castor, cucumber and groundnut). Some tropical trees and lianas
have fruits with very large seeds. The double coconut, Lodoicea maldivica has bilobed
seeds as large as 10 cm weighing nearly 6 kg.
The seed surface may be smooth, wrinkled, striated, ribbed, furrowed or it may have a
variety of patterns on it. l'hk surface may be glossy (as in linseed and castor), fleshy or
pulpy (in Magnolia) or covered with hair (in cotton).
Parts of a Seed
Seed is attached to the fruit by a stalk, the funiculus (funicle). The prolongation of the
funiculus running along the seed and terminating at the chalaza is called raphe (Fig.
6.1). The funicular vascular supply is responsible for the flow of food reserves. When
the seed is separated from the funiculus a scar is left at the point of attachment which is
termed mum.
COTYLEDONS,

SEED
COAT

RADICLE

SHOOT

Fig. 6.1: Various parts commonly present in the seeds

How Seed Develops?

Corresponding with the development of embryo and endospenn the ovule, the
integument(s) and the nucellus also embark on certain changes which eventually result
in the formation of mature seed. The usual alterations are described with the help of a
few examples.
Nucellus: In a large majority of flowering plants the nucellus is gradually utilized by the
endosperm or embryo, In leguminous seeds, for example, the nucellus degenerates
completely. Sometimes, as observed in Euphorbia spp., nucellar cells near the micropyle
(termed epistase) and chalaza (hypostase) survive longer and may even persist in the
mature seed. In the black pepper fruit the bulk of the volume is occupied by the
persistent nucellus (Fig. 6.2), which is also the chief food storage tissue (endosperm is
relatively little). Such persisting nucellus in the seed is designated perispenn. In
Daphniphyllum himalayense the seed has copious endospenn surrounded by perisperm,
which is characterized by the presence of oil droplets and even protein crystals
(Fig. 6.3).
Plant Developmeml-I

. ,,
ENDOSPERM

Fig. 6.2: Black pepper-A fruiting branch of Peper nigrum. B-a pendulous dense
spike. C - whole and split fruits D -Fruits with seed cut in longitudinal
section.

PROEMBRYO

Fig. 6.3: Endosperm and seed coat of DaphniphyUum. A. L.S. of seed at globular
proembryo ,stage. B. Magnified to show scelerification of tegmen and presence
of crystalline protein reserves in perisperm

Integuments The ovule has usually one or two integuments. After fertilization, there
may be initially a proliferation of cell layers in one or both the integuments. An
integument in which the cell layers increase is described as multiplicative. If the number
of cell layers in the integument remains the same as in the mature ovule then the
integument is regarded non-multiplicative. Alternatively, a process of disorganisation of
cells may begin at an early stage. In either case, as the seed matures, most of the cell
layers degenerate and get compressed. At the same time, some particular cell layers in
one or both the integuments persist and may become hard to form a protective sheath.
Cells of the protective layer often enldge in the anticlinal (perpendicular to the seed
surface) plane, and their walls become lignified and even cutinized. The characteristic
layer, if present, is described d sclerotic, mechanical, palisade or Malpighian layer.
Some seed biologists prefer to call the palisade-like cells as a layer of macrosclereids.
In a seed developing from a bitegrnic ovule, the persisting outer integument is termed
testa and the inner integument tegmen. In seeds originating from unitegmic ovules the
seed coat is loosely termed testa. Seeds with characteristic testa are called testal Seed
and those with prominent tegmen are described as tegmic. Seeds in which outer part
of the outer integument constitutes the mechanical layer are designated exotestal,
and those having hardened inner portion are endotestal. Similarly, seeds with outer
part of the inner integument modified as sclerotic zone are exotegmic and those with
inner layers fonning the protective sheath are endotegmic. In some plants which
have a stony or tough fruit wall, the seed coat may be thin and soft (coconut and
almond).
The histological changes that lead to the formation of the seed coat may be studied with
the help of examples of cotton, melon, mustard and bean.
In cotton (Gossypium spp.) the ovule has two integuments (Fig. 6.4) and both participate
in formation of the seed coat. At mature embrvo sac stage the outer integument is 4-6

.
Fig. 6.4: Seed coat development in Gossypium newaceurn A L.S. of ovule having
mature embryo sac. B. Portion of integuments from ovule after 2-3 days of
pollination.

cells thick and inner is 8-15 cells thick. The inner integument is multiplicative. Six days
after pollination the outer integument can be distinguished into three zones (Fig. 6.5):
(i) outer epidermis; (ii) outer pigmented zone of 2-5 layers having some tannin and
starch-filled cells; (iii) inner epidermis. In the inner integument, cells of the outer
epidermis start elongating radially. These epidermal cells enlarge many times their
original size and their walls become thick (Fig. 6.5.). This layer forms the sclerotic
layer of the mature seed coat. In the mature seed the inner integument has four zones:
outer palisade layer; a pigmented zone of 4 or 5 layers, inner colourless zone of 9 or 10
layers; inner epidermis. Thus, the mature seed coat has seven distinct zones made up by
both the integuments.

OUTER PIGMENTED

OUTER COLOURLESS

PALISADE LAYER

INNER PIGMENT

INNER COLOURLE

FRINGE LAYER

Fig. 65: Structure of integuments of Gossypium herbaceurn. A-Mature seed coat.

B-A lint hair C--A fuzz hair.
During development of seed coat in cotton, some of the outer epidermal cells of the
outer integument enlarge and then elongate outward to form hairs. These hairs, the
cotton of commerce, are single celled, thin-walled and attain a size of up to 40 mm.
Lint hairs are longer with characteristic twists, whereas fuzz hairs are short and without
twists.
In the gourd family, cucurbitaceae the ovules are bitegmic but the outer integument
alone forms the seed coat. In LuRa spp. at mature embryo sac stage the outer
integument is 10-15 layers thick and inner is 2 or 3 layered (Fig, 6.6). During seed
development the inner integument degenerates. The outer epidermal cells of the outer
integument elongate radially and develop rod-like thickenings on their radial walls (Fig.
6.6). A few layers of small pitted cells lie beneath the epidermis. The innermost layer of
this zone comprises radially elongated cells. A single layer of large, radially elongated
and somewhat bone-shaped cells occur next, constituting the palisade or mechanical
layer. Inside the palisade layer is a region of spongy parenchyma having thin walls and
intercellular air spaces.

Fig. 6.6: Seed-coat development in Luff;.(iii), inner integument; oi, outer integument;
I,, epidermis, l,, hypodermls; I,, sclerenchymatous layer; 1, ,aerenchyma). .
A. Portion of longisection of integuments before fertilization in L hermphr&.
B. Portion of longisection of mature seed-coat in L graveolens; chlorenchymatous
zone is not drawn.

In mustard, Brassica campestris the outer integument has 2-5 layers of cells and the
inner has up to 10 layers. Cells of the outer epidermis of outer integument become large
and filled with mucilage (Fig. 6.7). The subepidermal layer has tangentially elongated
cells which are gradually crushed. Inner epidermis of the outer integument forms the
sclerotic layer. The inner.integument gets obliterated, except for its inner epidermis
which forms a pigmented layer.

Fig. 6.7: T.S. of testa of Brassica sp. ewpidermis, sep-sub epidermis, Pal-palisade;
pil-pigment layer; al-4eurone layer of endosperm.

In leguminous seeds, such as those of Phaseolus lunarusi also the seed coat is derived
from the outer integument whereas the inner integument degenerates. The outer
epidermis of the testa forms the palisade layer Fig. 6.8). Its cells have a Sad and M t
characteristic light line running tangentially along the middle or close to the outer
walls. The light line is the result of intense refraction in the particular region of '
epidermal walls. Orientation of microfibrils that constitute the wall thickening is
responsible for the varying refraction. The subepidermal layer of cells differentiates
into hour-glass or funnel-shaped cells. Below this is the region of thin-walled
parenchymatous tissue with vascular bundles. The outer part of this tissue has well-
develo~edintercellular spaces.
The region of hilum in leguminous seed has an extraordinary organisation. The
attachment of funicle forms a disc-shaped structure which fits into the depression of
the hilum. The outer layer of cells of the head of the funicle also forms a palisade
layer (termed as counter-palisade), which is attached to the palisade layer of the testa
(Fig. 6.8). Both palisade and counter-palisade are interrupted in the centre by a
narrow groove which serves as an air passage in the ripening seed. The groove also
leads to a group of tracheids in the seed called the tracheid bar. On either side of the
bar is aerenchyma. Since the testa is impermeable to water, the tracheid bar also
serves as a 'hygroscopic valve' for absorbing moisture during seed ripening and
germination.

Fig. 6.8: Median L.S. of seed coat through hilum of Phaseolus aureus. A-photographic
representation of the region of hilum. B. Diagrammatic representation of funicle
region.

The seed coat of certain plants is fleshy or juicy. In the pomegranate, Punica
granatum, for example, the outer epidermal cells of the testa enlarge and become filled
with a sweet sap under considerable turgor pressure. This layer forms the juicy edible
part of the seed (Fig. 6.9). The inner part of the testa is hard and the tegmen is
membranous. In Magnolia spp. the inner integument forms the hard shell, whereas the
outer integument is fleshy and brightly-coloured. Pulpy and fleshy testa can be termed
sarcotesta.
Fig. 6.9: L.S. of seed (~ia*ammatic representation) of Punica gmnututn. Outer epidermis of
testa has radially elongated cells. Thesecells form fleshy part of seed.Solid black
portion is inner portion of outer integument and is made up of sclerenchyma.

SAQ 1
State whether the following statements are true or false. Write either T or F in the boxes
provided.
a) Seeds of flowering plants are always enclosed in a fruit. 1 I
b) Micropyle of the ovule is represented in the seed as hilum. [ 1
c) A true seed is formed from a ovule and it encloses embryo formed
with or without fertilization. I 1
d) In the majority of seeds the nucellus constitutes the nutritive tissue. 1
e) Malpighian layer of seed coat has radially elongated cells with thick
wall impregnated with lignin or cutin. [ 1
f) The seed of cotton is exotegmic. [ 1
g) Leguminous seed is endotestal. [ 1

6.3 SEED APPENDAGES

Seeds of certain plants have specialized outgrowths or envelopes. These structures

develop following fertilization from parts of the ovule or funicle. Here you will
study the origin and structure of some seed appendages. In a later part of this unit
you will learn more about how these seed appendages help in effective seed
dispersal.
Aril. It is an outgrowth that arises from the funicle or the testa near the raphe and
covers the seed partially or completely. It is often referred to as the third
integument. It is often fleshy or brightly coloured. The edible part of litchi fruit is
aril, which envelops the hard brown seed coat. In Myristica fragrans the hard seed
(nutmeg of commerce) is covered by a thin, irregular and bright-orange aril (that
gives us the precious spice mace). Seeds of Pithecellobium dulce, a leguminous tree,
have a fleshy red aril that partially surrounds the seed. In Crossosoma californicum
a fimbriate aril covers the seed on the sides (Fig. 6.10). Cells of the aril contain
oils, starch, sugars, pigments and aroma containing compounds. The appendage is
mostly an attraction for birds, which consume the aril and scatter the seeds. Seeds
of white water lily, Nymphaea alba have a spongy aril that provides buoyancy for
dispersal of seed by water.
 Seed and FI

-
Fig. 6.10: hature seed of Crossosoma calijornieum which is surrounded by imbriate aril.
Caruncle. This is a white, collar-like structure borne on the micropylar end of the
seed in many members of the Euphorbiaceae, such as castor, Ricinus communis. The
soft outgrowth capping the hard seed is formed by proliferation of the cells at the
tip of the outer integument. It is rich in starch and sugars. The violent opening of
the fruit (this is termed gun-shot mechanism) causes seeds to be thrown away a few
feet. It is believed that ants consume the caruncle and in the process carry the seed
further away. The caruncle is also considered to be hygroscopic and it may be
helpful in absorbing moisture for seed germination.
Operculum. The term operculum is applied to a plug-like structure formed in the .
micropylar portion of the seed by proliferation of cells at the tip of the inner
integument or the nucellus. An operculum has been observed in seeds of many
monocotyledonous families, such as the Commelinaceae, Musaceae, Lemnaceae and
Zingiberaceae, and a few dicotyledonous families like Bignoniaceae and
Nymphaeaceae. In L e m paucicostata, cells at the tip of the inner integument
undergo a remarkable expansion after fertilization and form a dome-shaped, stopper-
like operculum (Fig. 6.1 1). Cells of.the operculum are thick-walled and contain an
orange-red substance. During germination of the embryo the operculum becomes
detached from =st of the seed and facilitates emergence of the embryo.

Fig. 6.11: L.S. of mature seed of h n a paucicostakr. A prominent operculum can be seen
at the micropylar end. (After Maheshwari and Kapil, 1964).
4. W i gs and hairs. Seeds of certain plants have epidermal outgrowths or the
integuments themselves may form folds and projections that present a wing-like
appearance. In the tree Oroxylon sp. a thin, transparently white, circular or oval,
disc-like wing spreads all around the two-lobed seed (Fig. 6.12). The seed of
cucurbit Zanonia macrocarpa has wings measuring up to 10 cm. In the drumstick
plant Moringo oleifera the seed has three equidistant wings. Seed wings provide
large surface area, with optimum strength, combined with a minimum of
 material. A surprisingly large number of winged seeds can be arranged wmpactly
in a fruit. Wings help the seed to propel, sail or spin some distance away when

Fig. 6.12: W i g e d seed of Oroxylon.

Have you observed some tiny structures with lustrous white hairs floating around in the
air, specially in early summer? These are seeds which travel long distances with the
help of hairs. Seeds of milkweed, Calotropis procera have a tuft of hairs at one pole
(Fig. 6.13). Those of Adenium sp. of the Apocyanaceae have hairs at both ends. In
cotton and poplars the seeds have hair all over the seed coat surface. Hairs provide the
seed a large surface area without a corresponding increase in weight, thus helping in
dispersal by air. In some aquatic plants such as Nymphoides spp. the seed hairs are
filled with air and provide buoyancy to the seed to float and become distributed over a
large area.

Fig. 6.13: Mature and dehisced fruit of Calatropis, showing hairy seeds at one end.

SAQ 2
Given below are some inwmplete statements regarding seed appendages. Fill in the
blanks using appropriate words:

a) The sweet, edible part we relish in a litchi fruit is ...............................

b) Caruncle is involved in seed dispersal by ...............................
c) Aril is also known as the ...............................

d) In the drumstick fruit the seed has .............................. wings.

e) Operculum is produced by proliferation of cells at the tip of ...........................
or ...............................

f) ............................... and ..............................are two families in which the

members have seeds hairs.

g) In Oroxylon the seed has a .............................. for dispersal.

h) Several members of the family .... ........:................
! bear caiunculate seeds.
I 6.4 STORED METABOLITES
Seed and

In a large majority of seeds food is stored in the cells of the endosperm. In coconut,
wheat and castor bean for example, it is tlie endosperm which stores the bulk of the
food reserves. Food stored in endosperm is utilized by the embryo during development
and seed germination. In Unit 4 you have already learnt that endosperm is usually a
triploid tissue derived from the fusion product of a male gamete (brought by pollen
tube) and two polar nuclei in the central cell of the embryo sac. The endosperm
surrounds the embryo all around and is ideal for nurturing the embryo till the seedling
begins to photosynthesize and becomes antotrophic.
A second sear/& of nutritive tissue in some seeds is the perisperm, which represents
the persisting nucellus. Nucellus is observed in some monocot families, such as the
Zingiberaceae (to which turmeric and ginger belong), and a few dicotyledonous families,
including the Piperaceae (e.g., black pepper) and the Nymphaeaceae (e.g., lotus). In
Canna the chalazal cell of the ovule divides repeatedly to form a starch-containing tissue
called chalazosperm. Mature seeds containing persistent endosperm or perisperm are
I called albuminous and those lacking them exalbuminous..
Mature leguminous seeds lack endosperm. In pea, gram (chick pea) groundnut and a
whole range of pulses the large cotyledons of the embryo take up the function of
storage of food. In cotton seed also the large, lobed and folded cotyledons (Fig. 6.14)
are the main repository of nutrients.

HYPOCOTYL

COTYLEDON

RESIN DUCT

ENDOSPERM

Fig. 6.14: L.S. of cotton seed kernel showing embryo witb intricate folds of the
cotyledons dark dots are glands.
The embryo in a germinating seed has critical requirements to mobilize food reserves.
It needs a source of carbon skeletal precursors, and a source of energy to assemble the
precursors for building complex compounds. Energy is not only provided by
carbohydrates but also by the stored lipids. Their oil content varies$om 30% of dry
weight in sunflower to 50% in castor and groundnut and even%i&er in coconut and oil
palm.
Carbohydrates may be stored in the seed by way of thickened walls of the endosperm
(as in the date palm and coffee beans) or cotyledons (in balsam and garden nasturtium).
More often, cells of endosperm and cotyledons contain starch grains. Cereal grains such
as wheat and rice contain 70430% starch, mainly in the endospenn. In beans,
cotyledonary cells may have as much as 50% starch.
Seeds are often described as starch type (cereals) or oil type (castor, linseed). However,
nearly all seeds have in addition protein reserves for supplying nitrogenous compounds
to the young seedling till it becomes capable of absorbing nitrogen from the soil witb
the help of roots. Protein reserves are also needed for rapid synthesis of enzymes
Plant Development-I required for digestion o f starch at the time of germination. Storage proteins occur in the
form of discrete protein bodies, often termed aleurone grains. These grains also contain
some minerals. Protein bodies occur in the endosperm and embryo in most plants.
Soybean seeds are particularly rich in protein content. However, in some others such as
the cereal grains these are concentrated in the specialized outer layer of the endosperm
called the aleurone layer. The aleurone protein bodies become active during seed
germination, triggered by the gibberellins released by the embryo. Tbis results in the
production of the enzyme a-amylase which digests starch present in rest of the
-'endosperm. Grain legumes are rich in reserve proteins. For example, groundnut contains
about 25% and soybean nearly 40% protein.

SAQ 3
State whether the following statements are m e or false by indicating T or F in tbe box
provided.
a) In a majority of seeds the endosperm is the main reservoir of nutrients. [ 1
b) Leguminous seeds have well-developed perisperm. [ 1
c) Castor seeds are exalbuminous. [ 1
d) Food reserves in the seed are meant for tiding over the period of
dormancy. 1 1
e) Proteins occur in only those seeds which lack starch and lipids. [ 1
f) In wheat barley grains proteins occur mainly m the cells of the
outermost layer of the endosperm called the aleurone layer. 1 1
g) During seed gemination, enzymes responsible for digestion of starch
are produced in the seed itself. 1 1

6.5 DEVELOPMENT OF FRUIT

Concment with the development of the seed(s), the ovary is transformed into a fruit.
The fruit protects the seeds and allows their release or germination. In primitive families
such as the Magnoliaceae the fruit opens while still on the plant and the seed itself is
the unit of dispersal. However, in most of the flowering plants the function of dispersal
is at least partly transferred to the fruit.

SEPAL
BUNDLE
PETAL
BUNDLE

THE DORSAL AND

VENTRAL BUNDLES

Fig. 6.15: Cross-section of the fruit of Pyrus malus (diagrammatic representation).

A true fruit develops from the carpel, specifically from its ovary. However, in many
so-called fruits, organs or tissues in addition to those of the ovary participate in
protection and dispersal of seed. Examples of accessory tissues or organs contributing to
fruit formation are many. In strawberry, Fragaria spp. the floral receptacle extends to
form the fleshy edible part of the fruit. In apple, Pyrus malus the floral tube formed by
the floral organs and the receptacle around the inferior ovary, together constitute the
bulk of the fruit (Fig.6.15). In both these in$inces the edible fruit is product of
carpellary and accessory tissues. On the other hand, in jackfruit, Artocarpus integrifolia
I
the perianth and in pineapple, Ananas comosus the bracts surrounding the flowers in an Seed and
inflorescence proliferate to contribute to formation of the fruit. Where organs other than
gynoecium participate in forming a fruit, the fruit is termed a false fruit or pseudocarp.
The wall of a true fruit is termed pericarp. The mature pericarp is often made up of
three distinct regions. In mango, for instance, the outer skin or peel represents the
exocarp or epicarp. The fleshy and juicy middle portion is the mesocarp. The inner shell
or stone is formed by the endocarp.
Fruits of different plants display a rich diversity in size, shape, structure, and hardness.
Chemical constituents and dispetsal mechanisms. From the morphological standpoint
they are classified into a few types based on two criteria. The main criterion is the
degree of hardness of the fruit wall or pericarp whether it is dry and hard or soft and
fleshy. The second criterion is the ability of the fruit to dehisce or remain intact after
ripening. Based on these criteria some important types of fruits are classified in
Table 6.1.
Table 6.1: Types of Fruits

Dry fruits

Indebiscent fruits

Developing from a single carpel

1. Achene. Contains only one seed which is loose inside the fruit (buttercup).
2. Caryopsis. It is like an achene but the pericarp and the testa of the single seed
become fused (wheat, corn).
3. Samara. A winged one-seeded fruit (maple).
Developing from a compound gynoecium with several carpels
4. Nut. A single-seeded fruit that develops from an ovary that originally contains
several carpels of which all but one degenerate. Mature nut has one carpel and one
seed (walnut).
Dehiscent Fruits

Developing from a single carpel

5. Follicle. A pod-like fruit that splits open on the ventral side (larkspur).
6. Legume. The pod splits open on both ventral and dorsal sides (pea, beans).
Developing from a syncarpous ovary with two or more carpels.
I 7. Capsule. The dehiscence is along fusion lines of carpels (Hypericum), along dorsal
bundles or lines'of dehiscence in each carpel (Iris), by splitting transversely into top
and bottom portions (Primula) or by small pores which develop in the pericarp
(below the persistent but dried stigma poppy).
8. Siliqua. A pod-like fruit of two carpels with a false septum dividing the locule.
When the fruit ripens the two valves separate and the seeds remain attached to the
septum (mustard).
Fleshy Fruits

9. Berry. Exocarp, mesocarp and endocarp are distinguishable but soft or fleshy. The
fleshy pericarp encloses one or many seeds (grapes, tomato).
10. Drupe. Like a berry except that endocarp is thick and hard (peach, mango).

11. Pepo. Like a berry, but the exocarp is hard forming a rind (pumpkin, squash).
Schizocarpic Fruits

There are fruits that develop from multilocular ovaries that separate when ripe into
individual achenes, each representing a carpel (Malva, Abutilon).
Plant Development-I Development of fruit can be studied by selecting a few representative examples of
follicle, caryopsis, legume, capsule, berry and drupe.

1. Follicle. In larkspur, Delphinium sp. the exocarp develops from the outer epidermis
and sometime the hypodermis of the ovary wall. It consists of thick-walled cells
(Fig. 6.16). Mesocarp is parenchymatous. The endocarp, formed from inner
epidermis, consists of thick-walled cells. At maturity the pericarp dries up and the
follicle opens along the line of fusion of the carpels.

OUTER
EPlDERMlS nf-I-<

Fig. 6.16: T.S. of the follicle wall of Delphinium.

2. Caryopsis. In cereals each carpel has one ovule and therefore the mature fruit has,
just one seed. During maturation, very little or no cell divisions are necessary in the
ovary wall. The pericarp and the remains of the integuments of the seed get
completely fused. In wheat caryopsis three main regions can be distinguished: (i)
the caryopsis wall which includes the pericarp, seed coat and remains of nucellus;
(ii) endosperm; (iii) embryo (Fig. 6.17). The pericarp can be distinguished into five
layers (Fig. 6.18): (i) epidermis; (ii) hypodermis; (iii) zone of thin-walled cells; (iv)
cross cells; (v) tube cells. The outer epidermis and hypodermis together form the
exocarp, having thick-walled compressed cells. Inside the exocarp are one or a few
layers of thin-walled parenchymatous cells. These are followed by the cross cells
that have thick walls with characteristic pits elongated transversely to the cell. The
tube cells constitute the inner epidermis of the pericarp. These cells have thinner
walls than cross cells but these too are pitted. In the mature caryopsis testa is
destroyed but tegmen is discernible along with one or two layers of nucellus.

Fig. 6.17: A L.S. of Grain of Triticum B. L.S. of embryo of Trificum.

3. Legume. Outer epidermis of the ovary usually forms the exocarp of the leguminous
pod. Next few cell layers constitute the mesocarp with thick-walled parenchyma.
The endocarp consists of sclerenchyma on the inside of which is epidermis or a few
layers of parenchyma and epidermis. In Astragalus macrocarpus, for example, inside
the sclerenchymatous part of endocarp there is a thin-walled hypodermis and an
 epidermis (Fig.6.19). Vascular bundles are located in the mesocarp accompanied by
some sclerenchymatous cells.
The two valves of the dried legume usually twist, resulting in their opening and
scaaering of seeds. The twisting and opening of valves is brought about by
shrinkage of the sclerenchymatous cell layers which are obliquely oriented.

Flg. 6.18: Portion of CS. through the caryopsis of W u m at mature stage of development

EPIDERMIS

R'

Fig. 6.19: AstrugcJIu macrocarpus (oblique d n ) .

4. Berry. In a berry much of the @carp is fleshy or juicy. In tomato.Lycopersicon

esculentwn even the placenta on which the seeds are borne is fleshy (Fig. 6.20).
The exocarp consists of an epidermis and 3 or 4 layers of collenchymatouc cells.
The epidermis is covered by a cuticle. Mesocarp consists of large, thin-wailed cells
1 with abundant intercellular spaces. There is a large increase in cell volume in this
zone of @carp. With the development of ovules, a M pollination, the
I parenchymatous tissue of the placenta grows around the funiculi. This
r parenchymatous tissue continues to pmlifemw furha and envelops the seeds .
cOrnPle@ly.
 FLESHY,
PARENCHYMATOUS
TISSUE

Fig. 6.20: Diagrammatic, ~epresentationof T.S.of Lycospersion esculcnhmi~after fertilization.

5. Drupe. In the drupe of peach, Prunus persica most of the cell divisions occur in the
ovary wall before fertilization or soon after it. Further growth of the fruit is
accomplished mainly by cell enlargement. To begin with, cell enlargement occurs in
all directions, but later expansion is mainly in radial direction. This is more so in
the inner portion of the mesocarp. At maturity the outer epidermis has thick cuticle
and unicellular hairs. Mesocarp has loose parenchyma. Endocarp has sclereids and
forms the stone of the fruit.
SAQ 4
State whether the following statements are m e or false by indicating T or F in the box
against each.
a) A m e seed develops within a carpel. [ 1
b) Apple is called a false fruit because it has no fertile seeds. [ 1
c) In mango the entire pericarp is fleshy and the seed coat is hard. [ 1
d) In the caryopsis of wheat the pericarp and seed coat are fused together. [ 1
e) A follicle opens along the line of marginal fusion of the carpel . [ 1
f) In a tomato fruit only the endocarp is edible. [ 1

6.6 DISPERSAL OF SEEDS

A plant usually bears many fruits and innumerable seeds. If all the seeds produced by a
plant were to germinate in the immediate vicinity, this will have several disadvantages.
The resulting seedlings would compete intensely among themselves for space, light,
water and minerals. They will be more vulnerable to atrack by pests and pathogens.
Moreover, there will be greater chances for backcrossing, in the resulting progeny,
rendering them genetically inferior.
To overcome these problems most plants have evolved one or another mechanism for
dispersal of seeds over a wide area. Fruits of some plants have built-in mechanism for
dispersing their seeds to considerable distances (autochory). Other plants depend on
external agencies such as air (anemochory), water (hydrochory) and animals (&hory)
to disseminate their fruits or seeds.
1. Autochory. This mechanism of self-dispersal is based on forceful expulsion of the
seed from the fruit because of desiccation or turgidity of the cells of the pericarp.
In balsam, Impatiens spp., for example, the fruit is a cylindrical capsule fonned by
the, fusion of five carpels (Fig. 6.21 A). The fruit wall comprises three regions of
which the middle is made up of radially elongated cells with high turgor pressure
(Fig. 6.21 B). This zone is termed expansion zone. In the dry, ripe capsule the cells
of the expansion zone are in a state of high tension. However, the inner portion of
the pericarp consists of 2 or 3 layers of collenchyma which offer resistance. At this
stage even a mild touch or jerk results in separation of the carpels at the base (Fig.
6.21 C). The five carpels instantly curl inward throwing the seeds at a distance of
about 2 metres.
In Arcenthobium sp, the fruit is a pseudoberry enclosing a single seed. The mesocarp is
formed by viscin cells. When the fruit is detached from its stalk the cells of the viscid
 Seed and Fruit
layer generate a high pressure and the pericarp contracts to hurl the seed out with great
force (Fig. 6.22).

\1B OUTER
EPIDERMIS

RESISTANCE EXPANSION TISSUE

TISSUE

Fig. 6.21: A L.S. of closed fruit of Impatiens. L.S. through pericarp showing elongated ceUs
of middle region with high turgor pressure. A fruit the valves of which have curled
inwards and thus have ejected the seeds.

Fig. 6.22: Diagrammatic representation of explosive seed discharge in Arcenthobium.

2. Anemochory. Seeds that are dispersed by air currents are usually light or are
provided with special structures to help them remain air-borne for long periods.
Since a large proponion of seeds are wasted, anemochorus plants usually produce a
profuse quantity of seeds per plant. Some of the orchids, for example, produce as
much as seven hundred million seeds per plant and the seeds are so tiny that these
are blown away like dust particles. The orchid seeds have an undifferentiated
embryo and lack endospenn (Fig. 6.23).

You have studied earlier in this unit about seeds and fruits of several species with hairs
or wings that propel them even in mild wind.
Winged fruits and seeds are a characteristic feature of some tall trees. In maple the
wings represent expansions of the pericarp.

Fig. 6.23: A seed of Q p r i p e b . Embryci q n be seen without endoepcrm through

transparent seed-coat.
,
3: Hydrochory. Plants that grow in or along the bank of water bodies often utilise
water as an agency for dissemination of fruits and seeds. Coconut, Cocos nucifcra
is an excellent example of a fruit which has spread to different continents because
of its ability to float over hundreds or even thousands of kilometres. The fruit has a
smooth, waterproof exocarp, followed by a fibrous and air-filled mesocarp and a
hard endocarp. The seed with a thin seed coat retains its viability for more than
three months. During this period the fruit can travel as long as 4300 km.Seeds of
lotus and some cypresses have air-filled cortical tissues.
4. Zoochory. Some fruits are eaten by animals and the seeds are passed out with the
excreta (endozoochory). Plums, Lantana, grapes, figs and guava are examples of
some of the fruits eaten by birds. Fleshy part of the fruit is digested in the gut of
the bird and the seeds are expelled along with the droppings. Using the excreta as
manure, many of such seeds germinate. It is believed that the seeds of Ficus species
germinate only after they have passed through the gut and have been subjected to
the scarification in the gullet by small pebbles and digestive fluids'in the alimentary
canal.
In plants like neem, maulsari etc., the seeds are too large. The birds eat the pulp and
discard the seeds below their perch. Fruits and seeds of many other plants are carried by
animals externally (exozoochory), sticking to their body or mouth parts. Seeds of
mistletoe, (Viscum album) are dispersed widely because they adhere to the beaks of
birds feeding on its fruits. Fruits of many members of the Asteraceae have spines (e.g.,
Xanthium) or hooks (BidCm)with the help of which they stick to the bodies of animals
and get distributed over a large area.
Mammals like squirrels, monkeys and even goats (these are used by fanners to
encourage germination of seed species of Acacia nitotica (feeding pods) play an
important role in dispersal. Humans serve as active agents of deliberate distribution of
seeds to raise plants useful to them.

6.7 PARTHENOCARPY
-
It is generally observed that the fruit develops after fertilization and it has fertile seeds
inside it. However, this is not always so. Fruits of certain varieties of plants, such as
edible banana (Musa sp.), tomato (Lycopersicon esculentum), orange (Citrus sp.) and
 grapes (Vifis vinifera), develop without seeds. In the seed-bearing bananas the three Seed' a d Fndt
l d e s of the beny are occupied by large seeds, whereas in the parthenocarpic varities,
the ovules degenerate and the cells of the pericarp and septae proliferate to form the
,
I

I
pulp regions (Fig. 6.24). The phenomenon of formation of fruit without fertilization is
known as parthenocarpy. Parthenocarpic development may require pollination
(stimulative parthenocarpy), or it may occur even without pollination (vegetative
p-enocarpy ).
n

Fig. 6.24: T.S.Fruits of developing parthenocarpic and seed beating varieties of banana.
(a) ~akbenocarpicfruit at the time of emergence of the inflorescence.
(b) Parthenocarpic fruit 8 weeks after emergence, showing the pulp (p) invading
the ovarian cavity (oc) or locule. (c) Seeded M t at the time of emergence
(d) Seed,ed fruit 8 weeks after emergence. Very little pulp is present around the
lo&, which are occupied by the enlarging seeds. 0, ovuk; s. seed, vb, vascular
bundle.

Three types of parthenocarpy are generally recognised: (i) genetical (ii) environmental,
(iii) chemically-induced.
Genetical parthenocarpy is obsmed in many species cultivated for their fruit It arises
due to mutations or hybridization. Generally parthenocarpic varieties have sterile pollen
so that the pollination stimulus is available, but fertilization Qes not take place. As a
result, a fruit is produced which has no seeds. The famous navel orange has arisen h m
a normal seeded Citrus variety through mutation in an axillary bud, which formed a
branch bearing seedless fruits. Several cultivated varieties of grapes and cucumber have
also resulted h m bud mutations.
Environmental parthenocarpy is the result of some environmental conditions such as
h s t or low temperature which interferes with the normal reproductive process. In
.
tomato, for instance, cultivation under low t e m p t u r e and high light intensity can .
induce parthenocarpy. Under these conditions pollination is so poor that seeds are not
produced but ovary is activated to form the fruit
Induced parthenocarpy involves lreatment of the flowers with certain plant growth
regulators. Auxins and gibberellins at low concentrations (about lC7- 104 M) have
been successfully utilised for induction of parthenocarpy in a large numbex of plants
which otherwise bear seeded fruits. Seedless guava, tomato, and straw- fruits have
been obtained by this method.
Paabenocarpy is important in horticulm because seedless fruits are more convenient to
consume and are particularly suitable for the industry manufacturing jams, jellies and
fruit juices. Gibberellins also cause fruit enlargement (in grapes which are considered
commercially beneficial for packaging and marketing, and also cause looseness of
b19ches).
6.8 VIVIPARY

In flowering plants the seed or fruit generally is dispersed and germination occurs when
the conditions are congenial for growth. However, in some plants growing along the sea
shores or in mangrove areas the soil is too saline or*salty for seed germination.
Moreover, in such places the seed and the young seedlings are likely to get washed
away by the tide. Mangrove plants (e.g., Rhizophora spp.) have, therefore, evolved a
unique adaptation for ensuring seed germination and nurturing of the young sporophyte.
The young seedling grows out of the intact fruit and hangs with its pointed radicular
end facing downward. Once the seedling attains a large size it drops down and
penetrates the soil. Roots are produced rapidly by the radicle and the seedling is fixed
firmly in the soil. By this time the seedling would have developed an extensive
photosynthetic tissue to ensure its establishment and has been compared to the similar
state observed in mammals. Certain bamboos and forest trees also practice vivipary.
This mechanism of rearing the young one by the parent is termed vivipary. It is a
specialized characteristic evolved by mangrove plants as a strategy for survival.
SAQ 5
Fill in the blanks in the statements given below wlth appropriate words.

a) Dispersal of seeds by means of wind is termed ............................

b) ........................... involves built-in mechanism in the fruit for expulsion of seed.
c) Seeds of the banyan tree are mostly dispersed by ............................
d) Edible banana is a ...........................fruit.
e) ........................... and ........................... are plant growth regulators
employed for induction of parthenocarpy.

f) Vivipary is observed in ........................... plants.

6.9 SUMMARY

In this unit we have studied the development of fruits and seeds. You have seen that the
fruit and seed svucture display immense diversity. We have examined the nature of
reserve materials and the sites of their storage in the seed. The interesting aspects of the
range of seed dispersed agencies have been discussed. The formation of seedless fruits
(parthenocarpy)and their importance in horticulture have been explained. The rather rare
phenomenon of vivipary in plants has been highlighted as an adaplive feature by the
mangroves.
What we have learnt can be summarized as follows:
A true seed is a fertilized ovule. It contains an embryo and generally the
endosperm. These are enclosed by the seed coat derived from the integuments.
Seed protects the young sporophyte and serves as an efficient propagule.
One or both the integuments form the seed coat. A layer called palisade or sclerotic
layer is usually differentiated which fonns the protective shell.
In some plants Perisperm (persistant nucellus) forms an additional nutritive tissue. In
many seeds, principally leguminous seeds, the cotyledons perform the function of
storing reserve food.
Seeds may have appendages such as aril, caruncle, operculum or wings and hairs
which help in their dispersal.
 carbohydrates (in the form of starch and cell wall materials), lipids and proteins Seed and Fruit
are the chief sources of nutrition for the germinating embryo and young seedling.
A true frbit develops from the gynoecium as a result of stimulus provided by
pollination and/or fertilization.
Fruits are classified on the basis of their water content (fleshy or dry), number of
carpels and locules, number of seeds and importantly on the criterion of their ability
to dehisce or not.
Fruits and seeds are variously adapted for self-dispersal (autochory). or dispersal
by agencies such as wind (anemochory), water (hydrochory) and animals
(zoochory).
Parthenocarpy involves formation of fruits without fertilization so that no fertile
seeds are produced. Varieties of horticultural plants can evolve by mutationsnr
hybridization to bear seedless fruits. Parthenocarpy can also be induced by
environmental factors and application of growth hormones regulators.
\
Vivipary is a unique phenomenon observed chietly in mangrove plants. It involves
nurturing and germination of the.young sporophyte while the fruit is still attached to
the parent plant.

The study of fruits and seeds becomes fascinating by a visit to a garden. crop field,
open land or a forest. You will need observant eyes, probably a knife and of necessity a
magnifying glass. The types of fruits and seeds you have studied in this Unit can be
found in your immediate surroundings, some even in a vegetable market or kitchen
garden! The spectacle of seed dispersal can be observed in nature at all places and at all
time.

6.10 TERMINAL QUESTIONS

1) What are the chief advantages of the seed habit?

2) Which are the food storage tissues in the seed? In what form is the food stored?
3) Name some seed appendages and describe how they help in dispersal or germination
of seed.
4) What is a true fruit? Why are apple and Jack fruit considered false fruits?
5) What are the chief criteria for the classification' of fruits?
6) What parts of the seed or fruit is edibleluseful in the following plants:
i) banana
ii) tomato
iii) coconut
iv) groundnut

vi) litchi
vii) cotton
viii) castor
ix) pineapple
x) strawberry
xi) pomegranate
xii) mustard

7) How are the fruitslseeds of the following plants dispersed? What type of adaptations
have been developed by these plants?
Plant Development-I i) coconut
ii) mistletoe
iii) banyan
iv) balsam
v) milkweed

8) What is the commercial importance of parthenocarpic fruits? How can seedless

fruits be induced artificially?

9) What is vivipary? How does ,it help the mangrove species to survive in their saline/
estuarine habitat?

6.11 ANSWERS

Self Assessment Questions

True
False
True
False ,
True
True
:False
aril
b) ants
. C) third integument
d) three
e) inner integument or nucellus
f) Apocyanaeae and Asclepiadaceae
,

g) wing
h) Euphorbiaceae
3. a) True
b) False
c) False
d) False
e) False
f) True
g) True
4. a) True
b) False
C) False
d) True
e) True
f) False
5. a) anemochory
b) autochory
c) birds
d) parthenocarpic
e) auxins and gibberellins
 Terminal Questions
1. The seed not only protects and nurtures the young sporophyte, but also serves as a
propagule for wider distribution. It may remain viable for long periods, or stay dormant
till conditions are suitable for germination and growth of the seedling.

2. Endosperm, cotyledons, perisperm and rarely the chalazosperm are the food storage
tissues of the seed. Carbohydrates (in the form of starch and wall materials), lipids and
proteins are the chief stored food reserves.

3. Aril is consumed by birds and the animals and the discarded seed gets dispersed
widely. The caruncle is sought after by ants, which carry the seeds far from where they
have fallen after dehiscence of the fruit. In some aquatic plants the spongy aril provides
buoyancy to the seed to stay afloat and travel long distances. During seed germination,
the operculum (a lid-like structure at the micropylar part of seed) gets detached and
facilitates the emergence of embryo to form the seedling. Wings and hairs also help in
dispersal of seed by wind.

4. A true fruit develops from a carpel. Apple is regarded a false fruit because the floral

t
! tube and the receptacle around the inferior ovary also contribute to the fruit wall. In the
fruit of the jack tree the perianth of the flowers in the inflorescence also proliferate and
contribute to the edible portion.

5. The chief criteria for classification of fruit are: degree of hardness, number of
carpels and locules; number of seeds per locule; ability of the fruit to dehisce or stay
intact.

6. i) Mesocarp, endocarp and placentae

ii) The whole fruit (sometimes including seed) minus the persistant calyx.
iii) The whole seed.

iv) Whole seed (sometimes without seed coat, and sometimes only cotyledons).

v) Hypanthium (basal portions of perianth and stamens) and the fleshy receptacle
vi) Aril

vii) Lint hairs yield textile fibre fuzz hair which are used for high class cellulose
acetatelnitrate and the cotyledons give oil. The oil cake is used as manure or a
medium for fungi~bacteriain industry or as source of gossypol.

viii) Endosperm

ix) Bracts

x) Receptacle of the fruit (flower) plus individual fruits developed fiom all the
carpels (including seeds)

xi) Testa
xii) The whole seed is crushed for oil.

7. i) by sea water; impervious exocarp, buoyant mesocarp and hard endocarp

ii) by birds; seeds are sticky

iii) mostly by birds; the figs (Syconia) consumed by birds, and the tiny fruits
bearing hard seed (one each) emerge ready for germination

iv) autochory; high turgor pressure in the mesocarp

v) by wind; seed hairs

8. Seedless fruits are more convenient to consume. These are particularly suitable for
fruit preservation industries.
 Seedless fruits can be evolved by mutations or hybridization and environmental
conditions such as low temperature. Application of growth regulators can induce
parthenocarpy in some plants.
9) Vivipary involves germination of seed and nurturing of the young sporophyte in the
fruit while it is still attached to the parent plant. It is an adaptation to tbe pP;culiar
environmental conditions that prevail in mangroves ecosystems. If a seed of
mangrove is liberated it is likely to be carried away by the tidal waters. Moreover,
germination of seed and growth of young seedling may be seriously impaired by the
high salt content. As an adaptation the seedling in mangrove plants separate from
the parent only when it is sufficiently large and capable of establishment in saline
watea as an autotroph.
GLOSSARY
androecim : male reproductive organs of a plant; stamens taken collectively
archesporium :.a cell or mass of cells, dividing to form microspore mother cells.
anticlinal : line of division of cells at right angles to the surface of apex of a growing
point.
apomixis : a reproductive process without fertilization in plants, akin to parthenogenesis
but including development from cells other than ovules, as apogarny and apospory.
cleistogamy : the condition of having flowers which never open, and are self pollinated.
dehiience : the spontaneous opening .of an organ or structure along certain lines or in a
definite direction.

embryogeny : the process by which the embryo is formed.

embryo sac : the megaspore in angiosperms, containing the female gemetophyte.
endosperm : the nutritive tissue of most seeds.
endothecium : one of the wall layers of anther that helps in the dehiscence of the
anther.

endothelium : synonymous-integumentary tapetum; specialized cells having nutritive

role, these develop from the inner-most layer of the integument and closely surround the
nucellus.
generative cell : the smaller of the two cells of the pollen grain, divides and forms the
sperms.
haustorim : an outgrowth of embryo sac which extends to the nutritive tissue to draw
nutrition. - -

indehiscent :'fruits which do not open to release seeds, but the whole fruit is shed h m .
the plant.
massula : a group of pollen grains that occur in a mass.
megaspore : it develops to form the female garnetophyte, or the embryo sac.
microsporangium : pollen sac; a sporangium containing a large number of microspores.
microspore : the cell from which pollen grain - the male gametophyte, develops.
parthenogenesis : reproduction without fertilization by male gamete.
periclinal : division parallel to the surface of the cell, or the apex of the growing point.
pollen tube : a tubular structure developed from pollen grain after pollination, it grows
towards the ovule, carrying male gametes to the embryo sac.
polyembryony : formation of several embryos in one ovule.
proembryo : an embryonic structure preceding true embryo.
suspensor : a chain of cells, developed from hypobasal segment of angiosperm zygote; it
attaches embryo to the embryo sac.

tapetum : nutritive layer investing the sporogenous tissue in a sporangium; it supplies

nutrition to the developing microspores.
tetrad : 4 spores formed after the 1st and 2nd meiotic division of the microspore mother
cell.
FURTHER READING

1. Bhojwani, S.S. & B h a t n a g ~S.P. 1993. The Embryology of Angiosperms. Vikas

Publishing House Pvt. Ltd., New Delhi.
2. Maheshwari, P. 1950. An Introduction to the Embryology of Angiospem. Tata-
McGraw Hill Publishing Company Ltd., New Delhi.
3. Shivanna, K.R. & Johri, B.M. 1985. The Angiosperm Pollen: Structure and
Fwrczion. Wiley, New Delhi.,