Chapter 14

Seeds and Fruits

SEEDS

The Seed Is a Mature Ovule

Seed Structures Vary

GERMINATION

The Germination Process Differs Among

Plants Germination May Be Delayed by
Dormancy

FRUITS: RIPENED OVARIES

The Nature of the Ovary Determines the

Structure of the Fruit
Fruits May Be Simple or Compound
Simple Fruits Are from Single Ovaries
Compound Fruits Develop from Several
Ovaries
Not All Fruits Have Seeds

ADAPTATIONS FOR SEED DISPERSAL

Dispersal May Be by Wind, Water, and

Animals
Some Plants Have Evolved Anti-herbivore
Mechanisms
Distant Dispersal of Seeds Is Not a Universal
Aim

SUMMARY

PLANTS, PEOPLE, AND THE

ENVIRONMENT: Doctrine of Signatures

IN DEPTH: Key to Fruits

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KEY CONCEPTS

1. In flowering plants, seeds are the structures containing the embryo plant for the next
generation. Seeds are surrounded by a seed coat and contain the embryo axis and the
cotyledons. They contain either one cotyledon (monocotyledonous plants) or two
(dicotyledonous plants). Cotyledons contain stored food.

2. Germination of seeds involves the activation of processes in the embryo, such as

mobilization of food reserves and starting cell division and elongation. The embryo
radicle becomes the root system of the seedling plant, and the epicotyl becomes the shoot
system.

3. There are several different types of seeds. Seeds have differing mechanisms and
specialized structures for dispersal.

4. A fruit is a ripened ovary. There are several different types of fruits.

5. The function of the fruit is to aid in dispersal of the seeds. Several different vectors--
wind, water, and animals--are involved in fruit and seed dispersal.

14.1 SEEDS

Seeds and fruits are without doubt the most important source of food for people
and other animals, and they always have been. Seeds and fruits are filled with stored
foods intended to help the embryo germinate and grow--or to attract an animal to eat the
fruit and inadvertently carry the seeds away to spread them elsewhere. Rice (Oryza
sativa ), corn (Zea mays), and barley ( Hordeum vulgare ) grains are used for food by the
majority of the people in the world. Early people recognized the nutritional value of seeds
and fruits, and they harvested them from wild plants, and later they figured out how to
grow them for food.
Biologically, seeds are mature ovules that contain the embryonic plants of the next
generation. The tremendous production of seeds ensures the renewal of plant
populations. Each seed is constructed and packaged to ensure its dispersal to a favorable
site for successful germination and growth. The fruit is the packaging structure for the
seeds of flowering plants. In this chapter we will discuss the structure and development of
seeds and fruits and their adaptations for dispersal.

The Seed Is a Mature Ovule

The seed completes the process of reproduction initiated in the flower. Following
fertilization, the zygote develops into an embryo, the primary endosperm nucleus
develops the endosperm, and the integuments of the ovule develop into the seed coat.
For a short time after fertilization (Fig. 14.1b, d), the zygote nucleus divides
frequently while the primary endosperm nucleus divides rapidly to form the endosperm,
the nutrient-rich storage tissue that will feed the seed when it germinates. After the

2
 a c
b

g
e

Figure 14.1. Embryo and seed development in cotton (Gossypium hirsutum). (a) Ovule after double
fertilization. (b) Embryo sac after fertilization. (c) The zygote divides by mitosis, one of the two cells
is destined to become the embryo and the other the suspensor. (d) Early stage of embryo
(proembryo) and endosperm development. (e) Early embryo as a small globe of cells. (f) Heart-
shaped stage of the embryo with newly formed cotyledons. (g) The cotton seed is the mature
embryo, with highly folded cotyledons, surrounded by a seed coat. Note the seed coat fibers (really
epidermal hairs), for which cotton is harvested.

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endosperm has developed, the zygote nucleus divides to form a filament of several cells
(Fig. 14.1c). The cell farthest from the micropyle begins a series of divisions that produces
the early stage embryo or proembryo. At about the same time, the cell closest to the
micropyle elongates and divides, becoming the suspensor, which supports the embryo in
the endosperm (Fig. 14.1d, e). Further divisions result in a globular stage (Fig. 14.1e) and,
finally, a heart-shaped stage, after the two cotyledons have developed (Fig. 14.1f). In
addition, the embryo develops a radicle (the embryonic root) at one end and a shoot tip at
the other (Fig. 14.18).
The specific steps just described apply to cotton (Gossypium hirsutum ); there are
many variations in the details of embryo development for other plants. For example, the
major distinction between embryos of dicotyledonous and monocotyledonous plants is
the number of cotyledons (two or one, respectively).
While the embryo is developing, the nucellus, endosperm, and integuments are
also undergoing changes that are characteristic of the group of plants to which the seed
belongs. In the great majority of plants, the nucellus and endosperm are required only
for the initial stages of embryo development. This is particularly true of the nucellus,
which is generally used as a nutritive source in early embryo stages. It persists as a food
storage tissue, the perisperm, in seeds of sugar beet and many other species. The
endosperm persists as a food reserve in seeds of many monocot plants, such as onion
(Allium cepa) (Fig. 14.2); these include grasses of such major economic importance as rice
(Oryza sativa ) and serious weed pests such as yellow foxtail ( Setaria lutescens) (Fig. 14.3).
Endosperm persists as a food storage tissue in relatively few dicot seeds, castor bean
(Ricinus communis ) being an exception (Fig. 14.4).

Figure 14.2. (above) External view of onion Figure 14.3. Caryopses (grains) of yellow
seeds (Allium cepa). X11. (below) A foxtail grass (Setaria lutescens),. X17.
longitudinal section through an onion seed (above) External views. (below) Median
showing the embryo coiled within the section. 4
endosperm.
a b c

Figure 14.4. Castor bean (Ricinus communis) seed, a dicot with endosperm in the mature seed.
(left) External view. (center) Section showing edge view of embryo. (right) Section showing
flat view of embryo.

When food storage occurs within seed structure, the seeds of two dicot
the embryo, the normal vascular tissues plants--
of the embryo convey the solubilized food
to the meristems of the emerging plant, a
where it is required for growth. Food
stored in the endosperm, outside the
embryo, is absorbed through epidermal
cells of the embryo axis.
The integuments become the
seed coats in the mature seed. Scanning
electron microscopy shows the seed coats
to be variously and sometimes beautifully
sculptured (Fig. 14.5). The seed coat acts
as a protective shell around the embryo
and sometimes contains chemical
b
substances that inhibit the seed from
germinating until the temperature, light,
or moisture conditions are exactly right
for germination.

Seed Structures Vary

Seed structure varies widely between

species. This means that plants have
evolved many solutions to propagating
Figure 14.5. Sculptured seed coats. (a) Field
themselves successfully. We will briefly
bindweed (Convolvulus arvensis). X19. (b)
describe, as examples of variations in
California poppy (Eschscholtzia californica).
X118.

5
bean and castor bean--and two monocot a b
plants--a grass and onion.

COMMON BEAN The bean (Phaseolus

vulgaris) seed is kidney-shaped in outside
view. External structures on the seed
are the hilum, micropyle, and raphe (Fig.
14.6). The hilum is a large oval scar left
when the seed breaks away from its
placental connection, the funiculus. The
micropyle is a small opening in the seed
coat at one end of the hilum; it is the
opening through which the pollen tube
enters the ovule. The raphe is a ridge at c
the end of the hilum opposite the
micropyle and is the base of the
funiculus.
When the seed coat of a soaked
bean is removed, what remains is the
embryo; no endosperm is present. The
bean embryo consists of two fleshy
cotyledons and the embryo axis. The
embryo axis is composed of the
embryonic root or radicle at one end and
the embryonic shoot or epicotyl at the
other end. The hypocotyl is just below
Figure 14.6. Bean (Phaseolus vulgaris), a
the cotyledons.
typical dicot seed lacking an endosperm
when mature. (a) External side view. (b)
CASTOR BEAN The castor bean (Ricinus
External face view. (c) Opened to expose
communis ) seed has an external
embryo.
structure called the caruncle, which is a
spongy outgrowth of the outer seed coat.
The hilum and micropyle of the castor bean are covered by the caruncle, and the raphe
runs the full length of the seed (see Fig. 14.4). The caruncle functions in absorbing water,
which is needed during germination.
The castor bean embryo is embedded in a massive endosperm. The embryo
consists of two thin cotyledons, a very short hypocotyl, a small epicotyl, and a small
radicle. Castor bean oil was used by the Egyptians as a laxative; a paste made from the
seeds was also used as a treatment for toothache. These seeds should be handled with
care, however, because they contain a very toxic substance, called ricin. If ingested, ricin
causes nausea, muscle spasms, and convulsions; as few as eight seeds can cause death in
sensitive people.

GRASSES The so-called seed of grasses is really a fruit, the caryopsis or grain (see Fig.
14.3). It is a one-seeded, dry fruit in which the pericarp or ovary wall is firmly attached to

6
the seed coat and seed. The starchy endosperm constitutes the bulk of the caryopsis and
is surrounded by a layer of cells, the aleurone layer, that contain proteins and fats but
little or no starch.
The grass embryo has an axis with a shoot apex and a root apex. The shoot apex,
together with several rudimentary leaves, is ensheathed by a coleoptile. The radicle is
surrounded by the coleorhiza. A relatively large part of the grass embryo is a very
specialized, shield-shaped cotyledon called the scutellum. The outer cells of the scutellum
secrete enzymes that digest the adjacent stored foods in the endosperm upon
germination. These digested foods move from the endosperm through the scutellum to
the growing parts of the embryo.
The process of milling to make polished white rice removes the caryopsis fruit coat,
the entire protein-rich aleurone layer, and the outer layers of the endosperm. This
means that most of the nutritious protein of the grain is removed before the rice is
packaged for human consumption. The milled material, called bran, is now itself a popular
food.
Popcorn is also the fruit of a grass plant. Americans love popcorn and we eat
billions of gallons of it every year. Have you ever wondered why popcorn pops? Have you
ever wondered why so many kernels are left unpopped at the bottom of the kettle?
Scientists at the University of Illinois conducted experiments to answer these questions.
Popcorn pops when heated because steam builds up inside the grain. The resistance of
the fruit coat holds the steam back until the pressure becomes so high that the grain
bursts. All the starch inside becomes fluffy. The "duds" don't pop because their fruit coat
has cracks in it that allow the steam to escape.

ONION Like grasses, the onion (Allium cepa) is a monocot; but unlike grasses, its seed
coat encloses only a small amount of endosperm. The embryo is very simple, the radicle
and single cotyledon being quite prominent. The shoot apex is located close to the
midpoint of the axis and appears as a notch. The embryo is coiled, with the radicle end
usually pointing toward the micropyle (see Fig. 14.2).

14.2 GERMINATION

Germination, the first step in the growth of the embryo, begins with the uptake or
imbibition, of water. This is a critical step because seeds are quite dry, containing only 5
to 10% water. The cells of dry seeds are tightly packed with stored proteins, starch, and
lipids (Fig. 14.7a). This stored food is packaged into cytoplasmic organelles called protein
bodies, lipid bodies, and amyloplasts, which store starch (see Chapter 3). After imbibition,
enzymes are activated and rapidly released to digest the stored food into smaller
molecules which can then be transported and converted into energy needed for growth.
Consequently, the cells of imbibed embryos contain fewer storage organelles and more
mitochondria, ribosomes, and endoplasmic reticulum which are organelles involved in
metabolism (Fig. 14.7b).
The first indication that germination has begun is generally the swelling of the
radicle. It imbibes water rapidly and, bursting the seed coat and other coverings that may
be present, starts to grow downward into the soil.

7
a b

Figure 14.7. Cellular changes brought about by germination, demonstrated in transmission

electron micrographs of yellow foxtail grass (Setaria lutescens). (a) Scutellum cell in dry caryopsis
before germination; note the abundance of storage organelles such as lipid bodies and protein
bodies. X8,600. (b) Root cell after 65 hours of germination; the cell now has fewer storage organelles
and more organelles involved in metabolism such as mitochondria and leucoplasts. X6,500.

The Germination Process Differs Among Plants

Although the succeeding steps of germination are essentially similar in all plants, there
are variations. In the germination of beans, peas, castor beans, and onions, a structure
with a sharp hook is first forced upward through the soil. The structure forming the hook
is different in each case. In bean (Phaseolus vulgaris ), the hypocotyl elongates (Fig. 14.8).
In pea (Pisum sativum ), the epicotyl elongates (Fig. 14.9). In both cases, cotyledons and
shoot apex remain below ground at first. The hook straightens once it is above ground
and exposed to light. In the case of bean, the straightening of the hypocotyl raises the
cotyledons and shoot apex toward the light. This is called epigeal germination. When the
pea epicotyl straightens, the cotyledons remain below ground, and only the apex and first
leaf are raised upward. This is called hypogeal germination (Fig. 14.9).
Castor bean (Ricinus communis ) has epigeal germination (Fig. 14.10). It is different
from common beans in that its cotyledons first function as absorbing organs, facilitating
the transfer of food from the endosperm to the rest of the seedling. When the reserve
food supply in the endosperm is exhausted, the cotyledons of the castor bean embryo
emerge from the seed coat. They enlarge, become green, carry out photosynthesis for a
time, and then eventually wither and die.
In onion (Allium cepa), a sharply bent cotyledon breaks the soil surface and slowly
straightens out. The cotyledon of the onion is tubular, and its base encloses the shoot
apex (Fig. 14.11). The first leaf finally emerges through a small opening at the base of the
cotyledon.

8
 In grasses such as corn (Zea mays ), the situation is more complex. The shoot and
root are enveloped by tubular sheaths of the coleoptile (Fig. 14.12) and coleorhiza. The
primary root rapidly pushes through the coleorhiza. Adventitious roots then arise from
the lower nodes of the stem. The coleoptile elongates and emerges above ground,
becoming 2 to 4 cm (about 1 to 2 in) long. At this time, the uppermost leaf pushes its way
through the coleoptile and, growing rapidly, becomes part of the photosynthesizing shoot.

Figure 14.8. Stages in the epigeal germination of a bean (Phaseolus vulgaris) seed.

9
Figure 14.9. Stages in the hypogeal germination of a pea (Pisum sativum) seed.

Figure 14.10. Stages in the germination of a castor bean (Ricinus communis) seed.

10
Figure 14.11. Stages in the germination of an onion (Allium cepa) seed.

Figure 14.12. Stages in the germination of corn (Zea mays). After the primary root emerges, it
branches to form the root system. Adventitious roots emerge from the lower stem, and prop roots
form to hold the stem upright. The emerging young leaves are protected by a sheath-like coleoptile.

11
Germination May Be Delayed by Dormancy

Seeds can remain viable for remarkably long periods. In one study, jars containing seeds
from several different plant species were buried. At 5- and 10-year intervals, the jars were
opened, and the seeds were tested for germination. Most species remained viable for at
least 10 years, and one species, the moth mullein (Verbascum blattaria ), germinated after
more than 100 years. This is not a record for seed longevity, however. Viable seeds from
the Oriental lotus (Nelumbo nucifera ) have been removed from archaeological sites
known to be more than 1000 years old.
Many viable seeds will not germinate even when supplied with water, oxygen, and
a favorable temperature, because they are in a state of dormancy (inability to germinate
because of reduced physiological activity). Various factors can break dormancy. For
instance, light is necessary for the germination of some lettuce ( Lactuca) species.
Scarring or breaking through the seed coat is required before the seeds of some plants
(including many kinds of legumes) will germinate; their hard, dense seed coats restrict the
movement of water and gases. In orchids, seeds are dispersed while the embryos are
immature, and they must develop further before germinating. Seeds of some cool-
temperate zone plants, (for instance, gooseberry, Ribes speciosum ), will not germinate
unless they are first subjected for a time to temperatures close to freezing, while moist.
At the other extreme, the seeds of some pines ( Pinus ) will not germinate unless they have
been subjected to the rather high heat of a fire. Another type of dormancy is produced by
natural chemical inhibitors, which occur in many fruits or in seed coats--for example, in
yellow foxtail grass ( Setaria lutescens).

14.3 FRUITS: RIPENED OVARIES

A fruit, the ripened ovary of a flower, is an important auxiliary structure in the sexual life
cycle of angiosperms. Fruits protect seeds, aid in their dispersal, and may be a factor in
timing their germination. Because fruits are highly constant in structure, even when
grown in different environments, they play an important role in the classification of
angiosperms. The kay at the end of the chapter, "Key to Fruits," will help you identify the
different types of fruits described in this section.
In everyday usage, the term fruit usually refers to a juicy and edible structure such
as an apple (Malus sp.), plum (Prunus sp.), peach (Prunus persica), or grape (Vitis vinifera ).
Structures that are commonly called vegetables, such as string beans (Phaseolus vulgaris ),
eggplant ( Solanum melongena), okra ( Hibiscus esculentus ), squash ( Cucurbita sp.), tomato
( Lycopersicon esculentum ) and cucumber (Cucumis sativus ), are all fruits in a botanical
sense, as are grains of corn (Zea mays ), and oats (Avena sativa), and other cereals.

The Nature of the Ovary Determines the Structure of the Fruit

In spite of considerable variation along family lines, fruits share basic developmental and
anatomical characteristics. For instance, all fruit development is initiated by fertilization
which stimulates the ovary wall to undergo development and differentiation into three
layers. The fruit wall (which develops from the ovary wall) is called the pericarp; its three
more or less distinct layers (in order, beginning with the outermost) are exocarp,

12
mesocarp, and endocarp (Fig. 14.13).
When the fruit is mature, floral
structures--such as pedicel, calyx,
withered stamens, style and stigma,
and even remnants of the corolla--may
also be present.
Tissues other than the ovary
wall that form part of a fruit, are
referred to as accessory. Much of the
fruit of pineapple (Ammas sp.), apple
(Malus sp.), and strawberry (Fragaria
sp.) can be called accessory. In the
Figure 14.13. Mature almond (Prunus amygdalus)
strawberry, the edible portion is a
fruit showing split outer portion of ovary walls
thickened, pulpy central receptacle in
(pericarp)--endocarp, mesocarp and ectocarp.
which achenes (dry, one-seeded fruits)
are embedded.

Fruits May Be Simple or Compound

There are three main categories of fruits. Simple fruits are derived from a single ovary.
They may be dry or fleshy; the ovary may be composed of one or more carpels, and the
fruit may be dehiscent (splits open when mature) or indehiscent (does not split open).
Compound fruits are composed of more than one fruit. There are two types: aggregate
fruits and multiple fruits. Aggregate fruits are derived from many separate ovaries of a
single flower, all attached to a single receptacle (for example, strawberry, Fig. 14.14).
Multiple fruits, such as pineapple (Fig. 14.15), are the enlarged ovaries of several flowers
grown more or less together into a single mass. The receptacle of some multiple fruits,
such as fig (Ficus sp.), enlarges and is actually the edible part (Fig. 14.16).
In classifying the different kinds of fruits, the following criteria are taken into
account:

1. the structure of the flower from which the fruit develops

2. the number of ovaries involved in fruit formation
3. the number of carpels in each ovary
4. the nature of the mature pericarp (whether the fruit wall is dry or fleshy)
5. whether or not the pericarp splits (dehisces) at maturity
6. if the pericarp dehisces, the manner of its splitting
7. the role, if any, that accessory tissues may play in formation of the mature fruit

Simple Fruits Are from Single Ovaries

Simple fruits come in several forms. Their pericarp (fruit wall) may be dry or fleshy. If
dry, the pericarp may or may not dehisce.

13
 Figure 14.14 (upper left). Strawberry (Fragaria),
an aggregate fruit. The tiny individual fruits,
called achenes, are simple, dry fruits. They are
embedded in an enlarged fleshy receptacle.

Figure 14.15 (upper right). Pineapple (Ananas

comosus), a multiple fruit. The individual fruits,
called berries, are embedded in the swollen edible
inflorescence axis. Jagged-edged bracts extend
out over the fruits.

Figure 14.16 (left). The fruit and flowers of fig

(Ficus carica). (a) Fleshy receptacle. Internal
views showing (b) staminate and (c) pistillate
flowers.

PERICARP DRY AND DEHISCENT The legume or pod is the type of fruit found in nearly
all members of the pea family (Fabaceae). A pod arises from a single carpel, which at
maturity generally dehisces along two sides (Fig. 14.17). In the pea (Pisum sativa ) pod, the
shell is the pericarp, and the pea is the seed. Pods may be spirally twisted or curved, as in
Scotch broom (Cytissus scoparius ). However, a number of legumes such as alfalfa
(Medicago sativa ) have pods that do not dehisce.
An example of a follicle fruit is the magnolia (Magnolia grandiflora ) (Fig. 14.18).
The follicle develops from a single carpel and opens along only one side.
Capsules are simple fruits derived from compound ovaries (an ovary composed of
more than one carpel). Each carpel produces a few to many seeds. Capsules dehisce in
various ways along the top surface. Poppy (Papaver sp.) is an example (Fig. 14.19).
The silique is the characteristic fruit of members of the mustard family
(Brassicaceae). The silique (Fig. 14.20) is a dry fruit derived from a superior ovary
consisting of two locules. At maturity, the dry pericarp separates into three portions; the
seeds are attached to the central, persistent portion.

14
Figure 14.17. Opened pea pod (Pisum Figure 14.18. Dehiscing follicle of a Magnolia
sativum) showing developing seeds attached sp.
to carpel margins.

a b

Figure 14.19. Poppy (Papaver sp.) capsule. (a) Side view Figure 14.20. Silique of Mathiola sp.
before dehiscence. (b) Mature poppy capsule dehiscing
by pores at the top. (c) Cross section of poppy capsule
showing the position of seeds.
15
PERICARP DRY AND INDEHISCENT The achene is a dry, one-seeded fruit. Sunflower
(Helianthus annuus ) achenes are usually called seeds, but carefully breaking open the
pericarp reveals that the seed is inside. The pericarp is easily separated from the seed
coat, which is a thin, filmy layer surrounding the sunflower embryo (Fig. 14.21).
The caryopsis or grain is the fruit of the grass family (Poaceae), which includes rice
(Oryza sativa ) and wheat ( Triticum aestivum ). The grain is a dry, one-seeded, indehiscent
fruit (see Fig. 14.3). It differs from the achene in that pericarp and seed coat are firmly
united all the way around the embryo.
The samara may be a one-seeded simple fruit, as in elm (Ulmus sp.), or a two-
seeded one, as in maple (Acer sp.) (Fig. 14.22). These fruits are typified by an outgrowth of
the ovary wall, which forms a winglike structure that aids in seed dispersal.
The schizocarp is a fruit characteristic of the carrot family (Apiaceae), which
includes celery (Apium graveolens). The schizocarp consists of two carpels that split, when
mature, along the midline into two one-seeded, indehiscent halves (Fig. 14.23).
The term nut is popularly applied to a number of hard-shelled fruits and seeds.
Botanically speaking, a typical nut is a one-seeded, indehiscent dry fruit with a hard or
stony pericarp (shell). Examples are chestnut ( Castanea sp.) and walnut (Juglans sp.). An
acorn, the fruit of the oak (Quercus sp.) (Fig. 14.24), is partially enclosed by a hardened
cup. The outer husk of the walnut, which is removed during processing, is composed of
bracts, perianth, and the outer layer of the pericarp. The hard shell is the remainder of
the pericarp. Note that unshelled almonds (Prunus sp.), are really not nuts but fleshy
fruits known as drupes, from which the hulls--exocarp and mesocarp--have been removed.
Brazil nuts (Bertholleda excelsa) are seeds, not nuts, and the unshelled peanut (Arachis
hypogaea ) is really a pod.

Figure 14.21 (left). Achene of sunflower (Helianthus annuus), unopened and opened to show
attachment of seed.

Figure 14.22 (center). Two-seeded samara of maple (Acer).

Figure 14.23 (right). Schizocarp of cow-parsnip (Heracleum hirsutum); left is face view, and right is
side view.

16
 PERICARP FLESHY The fruits in
this category are popular for food.
They feature a fleshy fruit wall
(pericarp). The fleshy part is
usually attractive to animals, who
eat the fruit and in turn carry
away the seeds. The seeds found
in these fruits tend to have a hard
seed coat that is not broken down
as the seed passes through the
animal and is deposited in its
feces.
a b Cherry, almond, peach,
and apricot (all are Prunus sp.), in
the rose family (Rosaceae), are
Figure 14.24. Nut. (a) An acorn of oak (Quercus) is
examples of drupes. The olive
actually a nut, the cup are fused bracts. (b) Internal
(Oleo sp.) (family Oleaceae) fruit
view of acorn, showing the embryo.
is also a drupe. Derived from a
single carpel, the drupe is usually one-seeded. It has a hard endocarpconsisting of thick-
walled sclereids (see Chapter 4), and a thin exocarp forms the skin. The mesocarp is the
edible fleshy portion. The pit of a cherry is a seed, with a thin seed coat, plus the stony
inner layer (endocarp) of the ovary wall. I n almond fruit (see Fig. 14.13), the mesocarp is
fleshy like a typical drupe when the fruit is young. As it develops, however, the mesocarp
becomes hard and dry and forms the hull. The shell of the almond is endocarp. This is an
instance in which the seed, not the outer part, is the edible part of a drupe.
A berry is a fleshy type of fruit that is derived from a compound ovary. Usually,
many seeds are embedded in the flesh (Fig. 14.25), which is pericarp, although the line of
demarcation may be difficult to see. It comes as a surprise to many that tomatoes, lemons
(Citrus limon ), and cucumbers ( Cucurbita sp.) are berries, but strawberries (Fragaria sp.)
and blackberries (Rubus sp.), in spite of their common names, are not. These are good
examples of botanical names and common names that don't match.
Lemons, oranges, limes, and grapefruits (all Citrus sps.) are a type of berry called a
hesperidium. The thick, leathery rind (peel) with numerous oil cavities is exocarp and
mesocarp; the thick, juicy pulp segments (endocarp) are composed of several wedge-
shaped locules (Fig. 14.26). The juice forms in juice sacs or vesicles that are outgrowths
from the endocarp walls. Each mature juice sac is composed of many living cells filled
with juice. The fruits of watermelon (Citrullus vulgaris ), cucumber, and squash--all
members of the cucumber family (Cucurbitaceae)--are a kind of berry called a pepo (Fig.
14.27). The outer wall (rind) of the fruit consists of receptacle tissue that surrounds and is
fused with the exocarp. The flesh of the fruit is principally mesocarp and endocarp.
Apples (Malus sp.) and pears (Pyrus sp.), both in the Rosaceae family, are examples
of pomes. This fruit is derived from a flower with an inferior ovary (Fig. 14.28). The flesh
is enlarged hypanthium (a fleshy floral tube), and the core is from the ovary.

17
 sepal

a
pericarp

placenta

locule

b b

Figure 14.25 (above left). Berry of tomato (Lycopersicon esculentum). (a) External view. (b) Cross
section.

Figure 14.26 (above right) A citrus fruit is a hesperidium. (a) Flower of organge (Citrus sinensis),
showing a lengthwise section of maturing ovary. (b) Cross section and external view of mature
fruit.

Figure 14.27. Cucumber (Cucumis sativus) fruit

is a berry.

18
 Figure 14.28. Pome fruit of apple (Malus
sylvestris). (a) Median longitudinal
section of flower showing maturing fruit.
(b) Median section of fruit. (c) Cross
section of fruit.

b c

Compound Fruits Develop from Several Ovaries

An aggregate fruit is formed from numerous carpels of one individual flower. These fruits
are made up of many simple fruits attached to a fleshy receptacle. The strawberry flower
has numerous separate carpels on a single receptacle. Each carpel develops into an
achene (see Fig. 14.14). Flowers of raspberry, blackberry, and other species of Rubus have
essentially the same structure as strawberry, except that the attached fruits are small
drupes (Fig. 14.29).
A multiple fruit is formed from individual ovaries of several flowers, all clumped
together. The fig (Ficus sp.) (see Fig. 14.16) and the pineapple (Ananas comosus ) (see Fig.
14.15) are examples of multiple fruits; the individual fruits composing them are drupes in
fig and berries in pineapple. The fig fruit we eat is an enlarged, fleshy receptacle. Its
flowers are small and attached to the inner wall of the receptacle.

Not All Fruits Have Seeds

In some plants, normal fruit may develop without seeds being enclosed. Fruits that
develop without fertilization are called parthenocarpic; consequently, such

19
 fruits are seedless. Thompson seedless
grapes (Vitis sp.) were thought to be
parthenocarpic until it was shown that
fertilization does take place but that the
ovules fail to mature into seeds. Such
situations have led to a broader use of
the word parthenocarpy to mean simply,
seedless fruits.
Parthenocarpic (seedless) fruits are quite
regularly produced in such cultivated
plants as eggplant, navel orange,
banana, pineapple, and some varieties of
apple and pear. In certain plants,
seedless fruits may be induced by pollen
that is incapable of fertilizing the ovules.
For example, in some orchids, placing
dead pollen or a water extract of pollen
upon the stigma may start fruit
development. Parthenocarpy is
commercially induced in some plants by
spraying the blossoms with dilute
Figure 14.29. Aggregate fruit of the blackberry aqueous solutions of growth substances
(Rubus ursinus). The individual fruits are like auxin.
drupes.

14.4 ADAPTATIONS FOR SEED DISPERSAL

The role of ripe fruit is threefold: to aid in the dispersal of the seeds inside; to deter
inappropriate seed-dispersing animals from taking the fruit or seed; and to protect the
seeds from herbivores who merely consume seeds but do not disperse them. It is
important to realize that there is no nutritional relationship between the fruit and the
seeds within it. That is, the stored food in the fruit cannot be utilized by dormant seeds or
by germinating seedlings. The only stored food available to seedlings is in the endosperm
and cotyledons within the seed itself.
Both seeds and fruits are rich in a variety of chemical resources: sugar, starch, protein,
lipid, amino acids, and a variety of secondary compounds. The average caloric value of
this material is about 5100 kilocalories (kcal) per gram dry weight, a value approaching
that of healthy animal tissue (about 6000 kcal per gram). In contrast, leaf, root, stem, and
other vegetative tissues average only 4000 kcal per gram. This means that it costs the
plant more to make seeds and fruits than it does to make vegetative organs. This
expense is necessary to ensure that the materials present inside seeds and fruits will
guarantee the successful dissemination of the seeds. Several different mechanisms have
c), float in seawater, attract the eye and the stomach of a bird (Fig. 14.30g), entice an ant
to carry a seed, or permit the fruit to hook onto the hairs of a passing mammal (Fig.
14.30d-f). Just as in pollination, the animal vectors used are sometimes rewarded,

20
sometimes exploited; in other words, the relationship is sometimes mutualistic, sometimes
parasitic.

Figure 14.30. Fruits

and seeds showing
various devices
aiding in dispersal.
a b c
Wind: (a) Clematis.
X4. (b) Dandelion
(Taraxacum vulgare).
X5. (c) Seed of
Coulter's big-cone
pine (Pinus coulteri).
X1. Attachment: (d)
Cranesbill
(Geranium). X3. (e)
f
foxtail (Hordeum
hispida). X3. (f) Bur
clover (Medicago
denticulata). X4. (g)
Fleshy edible fruit of
Cotoneaster . X2.
d e g Violent dehiscence of
pericarp. (h) Vetch
(Vicia sativa). X2. (i)
California poppy
(Eschscholtzia
californica). X2.

h i

Dispersal May Be by Wind, Water, and Animals

Common abiotic vectors for fruit and seed dispersal are wind and water. Winged and
plumed fruits (Fig. 14.30a-c) are common adaptations for wind dispersal. In some cases,
the seeds are ballistically exploded by a violent dehiscence of the pericarp (Fig. 14.30h, i).

21
 Some sedges (Carex sp.) have a fruit with a membranous envelope containing air, and
these are spread by floating on water. The coconut (Cocos sp.) is a tropical group of plants
famous for growing on midoceanic islands thousands of miles from other land. The
coconut fruit is capable of floating for many days and then germinating when washed
onto a sandy beach and leached of salt by rainwater (Fig. 14.31a). Many other tropical
beach plants have similar (although much smaller) floating fruits. A number of weed
species of irrigated farmland are dispersed by water along irrigation canals. In deserts
the smoke tree (Dalea) and desert willow (Chilopsis ) growing along arroyos (dry stream
beds) have hard seeds that are carried away from the parent by flash floods (Fig. 14.31b).
The rushing water also pushes the seeds against rocks, scraping the seed coats and
scarifying them. Without that scarification, the seeds would remain dormant. Water in
this case is more than a dispersal agent; it pretreats the seed and makes it receptive to
germination cues.

a b

Figure 14.31. Plants with water-dispersed fruits

or seeds. (a) Coconut germinating on a tropical
beach. The coconut fruit, or husk, is fibrous, and
the seed within is large and buoyant, capable of
floating hundreds or even thousands of miles in
seawater. (b) The seeds of these desert shrubs
are carried away from the parents by flash flood
waters in the arroyo.

Common animal (biotic) vectors include ants, birds, bats, rodents, fish, ruminants,
and primates. They are attracted to fruit by its color, position, seasonal availability, odor,
and taste. Sometimes, the vector eats only the fruit and discards the seeds; this is true of
some primates. In other cases, the vector swallows the seeds unchewed; after passing

22
unharmed through the gut, the seeds are excreted some distance from where they were
consumed. This is often the case with birds. Birds are attracted to fleshy, colored berries
and, after dining on them, may fly long distances before regurgitating or excreting the
hard seeds. Cattle eat legume pods of mesquite (Prosopis fuliflora) and Acacia in
southwestern grasslands and later pass many undamaged seeds out in their excrement.
These germinate, and the seedlings grow well in their fertilized microenvironment. In
yet other cases, the animals eat many seeds but cache others. Squirrels and jays, for
example, may carry walnuts, hickory nuts, acorns, and pine seeds from parent trees to
distant hiding places. Apparently, they forget some of the caches and never revisit them;
in effect, they have planted these seeds, and clusters of seedlings will emerge later.
Ants are responsible for dispersing many seeds of herbs in temperate-zone forests
and grasslands. They harvest the hard, small seeds and deposit them in granaries below
ground. Some seeds escape consumption and germinate. Other plant species have
elaiosomes, or food bodies, at one end of their seeds (Fig. 14.32). Ants harvest the seeds
only for that reward and then toss them out. Discarded outside the ant nest (several
meters from the parent plant), the unharmed seed may then germinate.
The relationships above are mutualistic because there is some reward for the
animal. Plants can also use animals in a more parasitic fashion; in such cases, there is no
reward. For instance, seeds of some aquatic and marsh plants stick to the feet of birds in
mud and are carried for long distances. Mistletoe (Phoradendron sp.), a parasite of other
plants, has naturally sticky seeds, and birds can carry them on their feet to new host
trees. Seeds with beards, spines, hooks, or barbs catch a ride to a new site by adhering to
animal hair and human clothing (see Fig. 14.30e, f).

Figure 14.32. Elaisomes. (left) Flannel bush (Fremontodendron californicum), a chaparral shrub of
California. (right) Each seed of the flannel bush has an elaiosome at one end of each seed. The
seeds are harvested by ants, taken to their nests, and stripped of their elaisomes, and discarded,
whereupon the seeds are free to germinate without competing with the parent plant.

Some Plants Have Evolved Anti-herbivore Mechanisms

At the same time that the fruit attracts dispersal vectors, it must repel herbivores.
Mechanisms to discourage herbivores include reducing the time of fruit availability,
making the fruit or seed coat physically hard, and making the fruit or endosperm
chemically repellent.

23
 Many perennial plant species do not reproduce every year, or at least the
magnitude of their reproduction varies from year to year. Such species produce fruit and
seed abundantly only during what are called mast years. Because food supply is a limiting
factor in population size, the relatively low amount of seed produced in off years keeps the
number of seed eaters in check. As a result, seed-eating mammal, bird, and insect
populations are not large enough to consume all the seeds available during a mast year, so
some seeds escape consumption and germinate.
Those species of plants that do reproduce every year often limit the time when
ripe fruit is available. Large fruits that require a long time to develop remain green, hard,
and relatively small until just before the final maturation stages. Then color, texture, size,
and sweetness change suddenly, and the fruit is available to herbivores for only a brief
time. This limits the number of fruits (and seeds) they can consume before dispersal.
Our common notion of fruit is a juicy, soft organ, but many plants produce fruits
that are partly or completely dry and hard. Examples include drupes, which have a hard
endocarp, and such completely hard fruits as nuts. Biting or boring insects are prevented
from invading the fruit or seed by the sclerenchyma tissue (usually sclereids), which also
prevents the seed from being damaged by the grinding action in the crops of birds or the
mouths of chewing mammals. Legume seed coats are notoriously hard and often pass
through animal guts unharmed.
Chemical protection mechanisms are widespread and diverse. Many fruits are rich
in secondary compounds, chemicals produced by a plant partly or entirely for the effect
they have on other organisms. In the case of herbivore defense, the effect is negative
and often toxic. Secondary compounds in fruits and seeds include lectins (which cause red
blood cells to clump), enzyme inhibitors, cyanogens (which release cyanide, a potent
nerve toxin), saponins (a detergent), alkaloids (like opium), and unusual amino acids.
When present in the endosperm, these secondary compounds may later have a primary
function as well; that is, they may be metabolized by the seedling into valuable nontoxic
resources.
During the course of evolution, it appears that the metabolic quirks of at least one
species of animal have successfully defused each chemical defense originated by seeds
and fruits. This is referred to as coevolution. Thus, there are specialized insects capable
of eating plant tissue toxic to nearly every other animal. In some cases--as in the larvae
of the monarch butterfly, which is able to feed on milkweeds rich in metabolic by-products
called cardiac glycosides--the animal not only eats the plant but uses the toxin for its own
benefit, to deter a predator.
An excellent example of plant-herbivore coevolution is the series of plant defenses
and herbivore responses summarized in Table 14.1 for a group of closely related tropical
legumes. The plants exhibit a range of herbivore defenses, involving chemistry, texture,
size, and timing. For each defense, however, some species of weevil has evolved a
solution.

24
Distant Dispersal of Seeds Is Not a Universal Aim

The benefit of fruit and seed dispersal is the spread of a species far from its parent.
There is a cost as well, however. Many fruits and seeds are wasted because they are
eaten and deposited in places inappropriate for successful germination and seedling
establishment. In some stressful habitats, only a very few safe sites exist, and these are
scattered within a large, hostile area. Because parent plants ordinarily already reside
within one of the safe sites, it is advantageous to prevent or limit dispersal away from the
parents.
One method of limiting dispersal is self-planting. The morphology of the fruit
lends itself to lodging near the parent and drilling into the ground. Many grasses have
long, bent awns (slender bristles) that twist as air humidity fluctuates. The awns function
as levers that drive the grain into the soil. Stiff hairs at the base of the grain prevent it
from pulling backwards, out of the soil. A few nongrass herbs, such as cranesbill (see Fig.
14.30d), employ a similar technique.
The peanut (Arachis hypogaea) inclines its fertilized flowers down to the ground,
and the fruits become buried as they mature. Seeds never leave the immediate proximity
of the parent.
Sea rocket, a common annual beach plant of temperate-zone shores, has a
bipartite fruit. The top half is easily dislodged when mature, and its corky texture allows
the enclosed seed to float and be carried to distant beaches via ocean currents. The
bottom half is firmly attached to the parent, and its seed is buried with the dead parent by
shifting sand at the end of the growing season (Fig. 14.33). The following year, hundreds
of seedlings mark the place where the parent plant grew the year before. This two-

25
pronged dispersal strategy serves both to spread the species and to maintain it in safe
sites year after year, even though it is an annual plant.
Plant species of isolated islands, when compared to close relatives that grow on
distant continents, often exhibit a loss of dispersal mechanisms. It is possible that they
evolved on continents first and then by rare chance were spread to islands, where the
process of evolution modified their seeds and fruits so that dispersal became very limited,
in keeping with the limited size of the islands.

Figure 14.33. Two-pronged

a b seed-dispersal strategy. (a)
Sea rocket (Cakile
maritima) is a common,
succulent annual plant
along many coastlines.
Note the immature, green
fruits along some stems.
(b) Diagram of the fruit.
Each part has one seed.
The top half is carried away
by ocean currents, and the
bottom half stays with the
parent.

KEY TERMS

abiotic endosperm multiple fruits

achene epicotyl nucellus
aggregate fruits epigeal nut
aleurone layer exocarp parthenocarpic
berry follicle pepo
biotic fruit pericarp
capsules grain perisperm
caruncle hesperidium pod
caryopsis (grain) hilum pomes
coleoptile hypocotyl radicle
coleorhiza hypogeal raphe
cotyledons imbibition samara
dehiscent integuments schizocarp
dicotyledonous legume (pod) scutellum
drupes mesocarp seed coats
elaiosomes micropyle silique
endocarp monocotyledonous suspensor

26
SUMMARY

1. A seed consists of a plant embryo surrounded by a seed coat. Seeds may store food
within or outside the embryo. In most dicotyledonous plants such as bean, food is
stored in the two cotyledons. Cotyledons may also serve as absorbing and, later, as
photosynthesizing organs. Food may be stored in an endosperm rather than in the
cotyledons.

2. In seeds of monocotyledonous plants, food is usually stored in an endosperm. In grasses,

such as corn, the single cotyledon-like structure (scutellum) is a specialized organ that
absorbs the nutrients from the endosperm. In seeds such as those of onion, the cotyledon
emerges from the seed coat and becomes green, but its tip continues to absorb food from
the endosperm.

3. The first step in germination is the imbibition of water. This water facilitates the
activation of enzymes involved in digesting stored food, which is converted to energy for
growth.

4. In germination, the cotyledons may be elevated above the ground (epigeal), sometimes
becoming photosynthetically active, or they may remain below the ground (hypogeal).

5. Mature seeds may be dormant and, depending on the species and the immediate
environment, may remain viable and dormant from a few months to many years.

6. Dormancy is usually broken by providing the seed with moisture, oxygen, and a favorable
temperature. Other factors, such as light, the removal of chemical inhibitors, or the
destruction of the seed coat, may be required in some instances.

7. A fruit is a ripened ovary plus other closely associated floral parts.

8. There are three different kinds of fruits, classified on the basis of the number of
ovaries and flowers involved in their formation:

a. simple fruits, derived from a single ovary.

b. aggregate fruits, derived from a number of ovaries belonging to a single
flower and on a single receptacle.
c. multiple fruits, derived from a number of ovaries of several flowers more
or less grown together into one mass.

9. Simple fruits may have either a dry or a fleshy pericarp. If the pericarp is dry, it
may be either dehiscent (splitting at maturity to allow seeds to escape) or
indehiscent (not splitting).
10. The role of the fruit is to aid in the dispersal of the seeds within and to deter
herbivores from eating the seeds without dispersing them. There is no nutritional
link between the fruit and the seed of the germinating seedling.

27
11. Vectors of fruit and seed dispersal include wind, water, and animals. Common
animal vectors include ants, birds, bats, rodents, ruminants, and primates. The
animals are sometimes rewarded with food for their dispersal activities, but at
other times they simply carry seeds and fruits to other sites.

12. Fruits protect seeds from herbivores by the timing of their ripening, their
hardness, and their chemical composition. Fruits may contain secondary
compounds that deter feeding by all but the most metabolically specialized
herbivores.

13. Some fruits prevent or limit dispersal, thus ensuring that the site occupied
by the parent plant will be occupied by its offspring well into future growing
seasons.

Questions

1. A seed is actually a mature ovule. Define each of the following terms:

integuments
seed coat
suspensor
embryo
cotyledon
hilum
raphe
micropyle
radicle
epicotyl

2. Describe the processes that occur during germination.

3. What is seed dormancy? Why is it an important process?

4. A fruit is a ripened ovary. What are the functions of fruits?

5. What are the differences between simple, aggregate, and multiple fruits?

6. Buy several different fruits at the grocery store and use the "Key to Fruits" in
this chapter to identify their fruit types.

7. One role of fruits is to aid in seed dispersal. Describe two different ways that
fruits aid in dispersal by abiotic factors. Describe two different ways that fruits aid
dispersal by biotic factors.

28
PLANTS, PEOPLE, AND THE ENVIRONMENT: Doctrine of Signatures

Because seeds and fruits are so

important, stories, myths, and legends
abound. Even the Bible describes
"forbidden fruit" in the Garden of Eden.
In the 1500s the Doctrine of Signatures
was widely believed. The idea of the
Doctrine was that the appearance of a
plant or plant part would reveal its inner
secrets and possible uses to people.
Others expanded this idea to include (Echium)
shape, color, and smell as indicators of
use.
The scales of a pinecone, for example,
look like teeth, so medieval people made
a concoction of pinecones mixed with
vinegar to gargle for teeth and gum
problems. (Actually, the vinegar
probably did the trick by itself.) Seeds of
viper's bugloss ( Echium ) resemble a
snake's head, so the belief arose that a
mixture made from these seeds could be
used as a remedy for snakebite. Seeds of
snapdragon (Antirrhinum majus ) worn in
a linen bag around the neck were
supposed to prevent one from being
bewitched. Some of these stories may
have held a little truth simply by
happenstance; but their real importance
(Antirrhinum majus) lies in the fact that they arose in the first
place because people found plants, and
their seeds and fruits, important to their
very survival.

29
IN DEPTH: Key to Fruits

A key is a tool to help students identify

things when there are several possible
choices. This key can be used to identify
the different types of fruits. It is a
dichotomous key, in which there are two
choices at each level. For example, the
first level asks if the fruit is formed from
a single ovary or from several. Progress
through each level in the key to
determine the fruit type.

I. Fruit formed from a single ovary of one flower: Simple fruits

A. Pericarp fleshy
1. The ovary wall fleshy and containing one or more carpels and seeds:
Berry (tomato, Lycopersicon sp.)
a. Ovary wall with a hard rind: Pepo (watermelon, Cucumis melo)
b. Ovary wall with a leathery rind: Hesperidium (orange, Citrus sp.)
2. Only a portion of the pericarp fleshy
a. Exocarp thin; mesocarp fleshy; endocarp stony; single seed
and carpel: Drupe (cherry, Prunus sp.)
b. Outer portion of pericarp fleshy, inner portion papery,
floral tube fleshy; several seeds and carpels: Pome (apple,
Malus sp.)
B. Pericarp dry
1. Dehiscent fruits
a. Composed of one carpel
i. Splitting along two margins: Legume or Pod. (pea,
Pisum sp.)
ii. Splitting along one margin: Follicle. (individual
fruits in Magnolia multiple fruit)
b. Composed of two or more carpels
i. Dehiscing in one of four different ways:
Capsule (poppy)
ii. Separating at maturity, leaving a persistent
partition wall: Silique (mustard)
2. Indehiscent fruits
a. Pericarp bearing a winglike growth: Samara (maple)
b. Pericarp not bearing a winglike growth
i. Two or more carpels, united when immature,
splitting apart at maturity: Schizocarp (carrot)
ii. One carpel; if more, not splitting apart at
maturity; one-seeded fruits
a). Seed united to the pericarp all around:
Caryopsis or Grain (rice)

30
 b). Seed not united to the pericarp all
around
i). Fruit large, with thick, stony wall:
Nut (walnut)
ii). Fruit small, with thin wall:
Achene. (sunflower)

II. Fruits formed from several ovaries

A. Fruits developing from one flower: Aggregate fruit (classify the
individual fruits in key for simple fruits)(strawberry)
B. Fruits formed from several flowers: Multiple fruit (classify individual
fruits in key for simple fruits)(pineapple)

31
Photo Credits

Figure 14.1 – 1st ed

Figure 14.2a – Thomas L. Rost; b – 1st ed
Figure 14.3a – Thomas L. Rost; b – 1st ed
Figure 14.4 – 1st ed, originally from Weier
Figure 14.5 –1st ed
Figure 14.5 – 1st ed
Figure 14.7 a&b – Thomas L. Rost
Figure 14.8 – 1st ed
Figure 14.9- 1st ed
Figure 14.10 – 1st ed
Figure 14.11 – 1st ed
Figure 14.12 – Starr and Taggart
Figure 14.13 – 1st ed
Figure 14.14 – Mike Barbour
Figure 14.15 – [Link]
Figure 14.16 – 1st ed
Figure 14.17 – Thomas L. Rost
Figure 14.18 – Mike Barbour
Figure 14.19 – 1st ed
Figure 14.20 – Thomas L. Rost
Figure 14.21 – Thomas L. Rost
Figure 14.22 – 1st ed
Figure 14.23 – 1st ed
Figure 14.24a – Thomas L. Rost; b 1st ed
Figure 14.25 – (a) [Link]
(b) Adityamadhav83/tomato (cc)
Figure 14.26a – 1st ed; b Prof. Judy Jernstedt
Figure 14.27 – Thomas L. Rost
Figure 14.28 – 1st ed
Figure 14.29 – [Link]
brambles-blackberry-morning/blackberry-fruit-photos-9219/
Figure 14.30 – 1st ed
Figure 14.31a – Inga Spence © 1995/Tom Stack & Assoc; b.
[Link]
Figure 14.32 – Robert S. Boyud, Auburn University
Figure 14.33 – 1st ed

Doctrine of Signatures: [Link]

singular-remedy-against-poyson/
[Link]

Key to fruits: [Link]

32