Seed Development

learning objectives INTRODUCTION

• Trace the origin of seeds.
Four hundred million years ago, plants moved out of the oceans to col-
• Follow the relationship onize land. Two major adaptations made this possible. The first was the
between flower parts and
evolution of the root. The root not only anchored the plant in soil but
seed parts.
also allowed the plant to obtain water and minerals no longer brought
• Explain the general parts
to the plant by ocean water. A second adaptation that increased a plant’s
of a seed.
success on land was the development of a vascular system. This allowed
• Describe the stages of seed
materials obtained by the root system to be efficiently transported to the
development.
leafy photosynthetic parts of the plant. However, the price of these
• Explain unusual types of seed
adaptations to land habitation was relative immobility. The first vascu-
formation.
lar plants (e.g., ferns) used spores to spread the result of sexual repro-
• Observe how plant hormones
duction. However, plants that used spores to reproduce required a wet
are important to seed
environment to allow male sperm to swim to fertilize the female egg.
development.
The development of the seed habit (dispersal of seeds rather than spores
• Describe ripening and dissem-
for reproduction) permitted plants to move away from perpetually wet
ination of fruits and seeds.
environments and colonize areas with drier climates. This initiated the
proliferation of the marvelous diversity found in seeds and their accom-
panying fruit structures. Seed-producing plants (especially angiosperms)
became incredibly successful, and it is estimated that there are currently
over 250,000 species of flowering plants, easily the most diverse group
found in the plant kingdom.
Propagation by seeds is the major method by which plants repro-
duce in nature, and one of the most efficient and widely used propaga-
tion methods for cultivated crops. Plants produced from seeds are
referred to as seedlings. Sowing seeds is the physical beginning of
seedling propagation. The seed itself, however, is the end product of a
process of growth and development within the parent plant, which is
described in this chapter.

REPRODUCTIVE LIFE CYCLES

OF VASCULAR PLANTS
Plant life cycles are characterized by alternate sporophytic and
gametophytic generations. The sporophyte is usually
plant-like in appearance with a diploid genetic composi-
tion. The sporophyte produces specialized reproductive
structures that facilitate gamete production through
meiosis. This initiates the gametophytic generation. Male
and female gametes have a haploid genetic composition,
and fusion of these gametes
(fertilization) results in a fertilization The
reproductive zygote (embryo) that sexual union of a male
restarts the sporophytic generation. and female gamete.

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Vascular plants are separated into those that dissemi- (usually wet conditions). The life cycle of a fern is
nate the next generation by spores or those who do so depicted in Figure 1. Spores are produced in sporangia
with seeds. within a sorus produced on the underside or edge of
the fern frond (sporophyte). The spore (1n) germi-
Seedless Vascular Plants nates and produces a small leafy structure called a pro-
Seedless vascular plants reproduce from spores, and thallus. On the mature prothallus, male (antheridia)
include horsetails (Equisetum), wiskferns (Psilotum), and female (archegonia) are formed. The male
lycopods (Lycopodium), Selaginella, and ferns. The antheridium releases the motile sperm (1n) that swims
spore is a protective structure that is tolerant of envi- into the archegonium uniting with a single egg cell
ronmental conditions, germinating when conditions (1n). Following fertilization, the zygote develops into a
are conducive for the gametophytic generation new fern.

(a)

(b)

(c)

(f )

(d)

(e)

Figure 1
A representative fern life cycle includes alternate sporophytic and gametophytic generations. (a) A mature fern sporophyte
produces fronds that typically produce (b) sori (spore producing structures) on the underside of the leaf-like frond. (c) Within the
sori are sporangia that contain the spores that initiate the gametophytic gerneration. (d) When the spore germinates it produces
a leaf-like gametophyte called the prothallus. On the prothallus, several female archegonia and many male antheridia are
formed. (e) Fertilization occurs when the male sperm unites with the female egg within the archegonium. (f) The resultant young
sporophyte becomes the long-lived fern. Adapted from Linda R. Berg. 1997. Introductory Botany. Saunders College Publishing.

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 seed development

Seed Plants located between the scales of the female cones.

The seed habit developed during the Devonian period Haploid male and female gametes fuse to form a
about 350 to 385 million years ago in an extinct group of diploid zygote that devel-
plants called the progymnosperms (47). ops into the embryo endosperm The major
Progymnosperms are only known from the fossil record within the seed. Storage storage tissue in seeds.
(Fig. 2) and produced seedlike structures enclosed in tissue (endosperm) in a It is derived from the
female tissue called cupules. They are considered the pro- gymnosperm seed is from haploid female
genitors to our current-day gymnosperms and the haploid female game- gametophyte in gym-
angiosperms. The seed habit is characterized by several tophyte. nosperms, while in
anatomical features that differentiate them from spore- Angiosperms are angiosperms it is the
producing plants: true flowering plants. result of gamete fusion
The term angiosperm that forms a triploid
1. Rather than producing a single spore type (homo- means “enclosed seeds” (3n) storage tissue.
spory), seed plants produce a separate female and refers to the female
megaspore and male microspore (heterospory). ovary tissue (carpels) that forms the fruit surrounding
2. The female gametophyte is retained on the mother angiosperm seeds. Angiosperms are the dominant plant
plant (sporophyte) and is enclosed within a protec- type on Earth with approximately 250,000 species,
tive maternal seed coat. compared with only about 8,000 living species of gym-
3. The ovule has an opening designed to receive nosperms. One reason for angiosperm success and
pollen that does not depend on water for male diversity is the mutualistic co-evolution of animals
gamete transfer. (especially insects) as pollinators and seed dispersers.
Seed plants are separated into gymnosperms and A representative angiosperm life cycle is presented in
angiosperms. Gymnosperms include the cycads, Figure 4. A key development in the angiospermic life
ginkgo, gnetophytes (Ephedra, Gnetum) and the cycle is the presentation of the female megagameto-
conifers (like pine, fir, and hemlock). The term gym- phyte as a multi-celled (usually 8) embryo sac within
nosperm means “naked seeds” and refers to the absence the ovule. Male gametes from the pollen enter the
of ovary tissue covering the seeds, which is a character- ovule. One gamete fuses with the egg cell to form a
istic of angiosperms (flowering plants). Pine is repre- zygote, and the second fuses with two polar nuclei to
sentative of a gymnosperm life cycle (Fig. 3). form the endosperm. This double fertilization is a char-
Conifers produce separate male and female reproduc- acteristic of angiosperms and leads to a triploid
tive cones (strobili) on the same plant. Male cones pro- endosperm rather than the haploid endosperm seen in
duce winged pollen that is dispersed by wind. Egg cells gymnosperms. Based on seedling morphology,
are produced within the female megagametophyte angiosperms can be separated into dicotyledonous
(seedlings with two
cotyledons) and dicots Produce seedlings
monocotledonous with two cotyledons.
(seedlings with one monocots Produce only a
cotyledon) plants. single modified cotyledon.

CHARACTERISTICS OF A SEED
A seed (20, 21) is a matured ovule containing an
embryo, storage reserve tissue, and a protective outer
covering (Figs. 5, 6). Seeds are the sexual reproductive
unit in a plant.

Figure 2 Embryo
Seed-producing plants evolved approximately 360 million
The embryo represents the new plant generation and
years ago, but most were not successful and became extinct.
Progymnosperms developed seeds enclosed within a cupule develops after the sexual union of the male and female
(arrow) and are thought to be the progenitors of the gametes during fertilization. Its basic structure is an
gymnosperms. embryo axis with growing points at each end—one for
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 seed development

(a)
(c)
(b)

(d) (f )

(e)

(g)

(h)

Figure 3
A representative gymnosperm life cycle. (a) A pine tree is a mature sporophyte. It produces separate male (b) and female
(c) reproductive structures. The male gametophytes are produced in a (d) staminate cone as winged pollen grains (e) spread
by the wind. The female gametophyte is produced within the female ovulate cone (f). The female egg cell (g) is fertilized by
the male sperm to produce a seed (h)—the next sporophytic generation.

the shoot and one for the root—and one or more dicotyledonous plants (such as bean or peach) have
cotyledons attached to the embryo axis. The basic two, and gymnosperms (such as pine or ginkgo) may
embryo types relative to the seed’s storage tissue is rep- have as many as fifteen. Embryo size in relation to the
resented in Figure 5. seed varies considerably (3, 48). In many seeds, the
The number of cotyledons in the embryo is used embryo occupies the entire inner seed (Figs. 5d, 6e),
to classify plants. Monocotyledonous plants (such as while others have small to miniature embryos (Figs. 5c,
coconut palm or grasses) have a single cotyledon, 6c).
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 seed development

(a)

(b)

(c)

(d)

Figure 4
A representative angiosperm life cycle. (a) Flowers are formed during the sporophytic generation. In the gametophytic
generation, (b) male gametophytes are produced within the anther as pollen grains and (c) the female gametophyte is produced
in the ovule within the ovary. (d) The seed is formed following male and female gamete fusion (fertilization), which reinitiates the
sporophytic generation.

Storage Reserves of haploid (1n) female perisperm Nucellus

storage and Storage tissue is designed gametophytic tissue (7). tissue that remains in
food reserves to sustain the germinating Perisperm is nucellar tis- the mature seed and
High-energy embryo until the seedling can sue from the female plant is used as storage
macromolecules produce its own resources and is diploid (2n). tissue.
like oils, through photosynthesis. For Seeds can be sepa-
carbohydrates, dicots, storage materials are rated into three basic storage reserve types that occur in
and protein that are contained in the endosperm, endospermic, non-endospermic, or unclassified
produced during cotyledons, and perisperm seeds (Table 1 and Figs. 5, 6).
seed development tissue. The endosperm is usu- In endospermic seeds, cotyledon growth is
and used for the ally the result of the fusion of arrested in dicots at different stages of development
early stages of seed two female and one male such that the embryo may be only one-third to one-
germination and nuclei during double fertil- half the size of the seed at the time it is ripe. The
seedling ization and is triploid (3n). remainder of the seed cavity contains large amounts of
emergence. However, in some plants, the endosperm or perisperm depending on the species.
endosperm ploidy level may Although the origin of the endosperm tissue is dif-
be higher (e.g., five-ploid in some members of the lily ferent, most monocot and gymnosperm seeds are
family and nine-ploid in peperomia). Storage tissue for endospermic.
monocots is the starchy endosperm (3n), and for gym- The pattern for reserve metabolism in non-
nosperms, the storage tissue is an endosperm consisting endospermic dicot seeds begins with an initial rapid

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Figure 5
The basic embryo types
found in seeds. Major forms
include: (a) Basal embryos
that have a high endosperm
to embryo ratio. This is
considered a more primitive
evolutionary condition;
(b) Peripheral embryos
surround and inner mass of
perisperm storage tissue;
(c) Axial embryos occupy the
center of the seed and
contain a significant amount
of endosperm; and (d) Foliate
embryos where the
cotyledons develop to
occupy most of the seed and
function as storage reserve
tissue. Color codes for these
images have the embryo in
green, endosperm in yellow,
perisperm in white, and seed
coverings are brown. Adapted
from Martin, A. C. 1946.

(a) (b) (c)

(d) (e)
Figure 6
Representative seed morphologies. (a) Gymnosperm (conifer) seeds have embryos with multiple cotyledons and use the female
gametophyte as reserve material. (b) Corn is an example of a monocot in the grass family. It has a peripheral embryo and a large
endosperm reserve. The outer protective layer is fruit tissue—pericarp. (c, d, and e) Each of the representative dicots has embryos
with two cotyledons. Magnolia has a small embryo and a large endosperm reserve. The fleshy outer covering is an aril derived
from the funiculus. Beet seeds have a curved embryo and utilize perisperm derived from nucellar tissue. In pear, the cotyledons
fill the seed and are used for storage reserve. The nutritive reserves in the endosperm have been transferred to the cotyledons,
so there is only a small remnant endosperm between the embryo and seed coat. The outer layer is fruit (pericarp) tissue.

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Table 1
C LASSIFICATION OF S EEDS
The following classification is based upon morphology of embryo and seed coverings. It includes, as examples, families
of herbaceous plants.
I. Seeds with dominant endosperm (or perisperm) as seed storage organs (endospermic).
A. Rudimentary embryo. Embryo is very small and undeveloped but undergoes further increase at germination (see
Fig. 5a, 6c Magnolia).
1. Ranunculaceae (Aquilegia, Delphinium), Papaveraceae (Eschscholtzia, Papaver), Fumariaceae (Dicentra),
Araliaceae (Fatsia), Magnoliaceae (Magnolia), Aquifoliaceae (Ilex).
B. Linear embryo. Embryo is more developed than those in (A) and enlarges further at germination (Fig. 5c).
1. Apiaceae (Daucus), Ericaceae (Calluna, Rhododendron), Primulaceae (Cyclamen, Primula), Gentianaceae
(Gentiana), Solanaceae (Datura, Solanum), Oleaceae (Fraxinus).
C. Miniature embryo. Embryo fills more than half the seed (Fig. 4c).
1. Crassulaceae (Sedum, Heuchera, Hypericum), Begoniaceae (Begonia), Solanaceae (Nicotiana, Petunia,
Salpiglossis), Scrophulariaceae (Antirrhinum, Linaria, Mimulus, Nemesia, Penstemon), Lobeliaceae (Lobelia).
D. Peripheral embryo. Embryo encloses endosperm or perisperm tissue (Fig. 4b).
1. Polygonaceae (Eriogonum), Chenopodiaceae (Kochia), Amaranthaceae (Amaranthus, Celosia, Gomphrena),
Nyctaginaceae (Abronia, Mirabilis).
II. Seeds with embryo dominant (nonendospermic); classified according to the type of seed covering (Fig. 4d).
A. Hard seed coats restricting water entry.
1. Fabaceae (Cercis, Gymnocladus, Gleditsia), Geraniaceae (Pelargonium), Anacardiaceae (Rhus), Rhamnaceae
(Ceanothus), Malvaceae (Abutilon, Altea), Convolvulaceae (Convolvulus).
B. Thin seed coats with mucilaginous layer.
1. Brassicaceae (Arabis, Iberis, Lobularia, Mathiola), Linaceae (Linum), Violaceae (Viola), Lamiaceae (Lavandula).
C. Woody outer seed coverings with inner semipermeable layer.
1. Rosaceae (Geum, Potentilla), Zygophyllaceae (Larrea), Balsaminaceae (Impatiens), Cistaceae (Cistus,
Helianthemum), Onagraceae (Clarkia, Oenothera), Plumbaginaceae (Armeria), Apocynaceae, Polemoniaceae
(Phlox), Hydrophyllaceae (Nemophila, Phacelia), Boraginaceae (Anchusa), Verbenaceae (Lantana, Verbena),
Labiateae (Coleus, Moluccela), Dipsacaceae (Dipsacus, Scabiosa).
D. Fibrous outer seed covering with more or less semipermeable membranous layer, including endosperm
remnant.
1. Asteraceae (many species).
III. Unclassified
A. Rudimentary embryo with no food storage.
1. Orchidaceae (orchids, in general).
B. Modified miniature embryo located on periphery of seed (Fig. 6b).
1. Poaceae (grasses).
C. Axillary miniature embryo surrounded by gametophyte tissue (Fig. 6a).
1. Gymnosperms (in particular, conifers).

Source: After Atwater (1).

nucellus Maternal growth of the embryo can still play an important role in controlling seed germi-
tissue in which the that digests the enclos- nation and dormancy.
megaspore mother ing nucellus. This is fol- The third storage reserve type occurs in
cell (also called lowed by expansion of unclassified seeds. These are seeds that have negligible
megasporocyte) the embryo through cell seed storage reserves like orchids. These tiny seeds rely
undergoes meiosis and division at the periphery on a fungal (mycorrhiza) symbiosis during germina-
forms the embryo sac. of the cotyledons that tion to provide the nutrition required for development
digests the developed and growth (63).
endosperm. In these seeds, the endosperm and/or the
nucellus is reduced to a remnant between the embryo Protective Seed Coverings
and the seed coat (integuments), and the cotyledons The protective seed covering layer surrounds the seed
function as the major storage tissue. Although the and provides physical protection; it may act to exclude
reduced endosperm may be only few cell layers thick, it water and gases. Seed coverings may consist of the seed
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coat, the remains of the nucellus and endosperm, and, Seed coverings provide mechanical protection for
sometimes, parts of the fruit. The seed coat, also the embryo, making it possible to handle seeds without
termed the testa, is derived from the integuments of the injury, and, thus, permitting transportation for long
ovule. During development, the seed coat becomes distances and storage for long periods of time. The seed
modified so that at maturity it presents an appearance coverings can also contribute to seed dormancy and
often characteristic of the plant family (18). Usually, control germination.
the outer layer of the seed coat becomes dry, somewhat Seeds may also contain additional surface struc-
hardened and thickened, and brownish in color. In par- tures that usually aid in seed dispersal. These include
ticular families, it becomes hard and impervious to arils, elaiosomes, caruncles, wings, and various plumes
water. On the other hand, the inner seed coat layers are of hairs (Fig. 8). Elaiosomes are particularly interesting
usually thin, transparent, and membranous. Remnants because they are nutrient-containing organs (especially
of the endosperm and nucellus are sometimes found oils) specifically designed to attract ants (4). The ants
within the inner seed coat, sometimes making a dis- use the elaiosome as a food source, and the plant bene-
tinct, continuous layer around the embryo. fits by ant dispersal of the seeds.
In some plants, parts of the fruit remain attached to
the seed so that the fruit and seed are commonly handled
together as the “seed.” In fruits such as achenes, caryopsis,
samaras, and schizocarps, the pericarp and seed coat layers REPRODUCTIVE PARTS
are contiguous (Fig. 7a). In others, such as the acorn, the OF THE FLOWER
pericarp and seed coverings separate, but the fruit covering Sexual reproduction (fusion of male and female
is indehiscent. In still others, such as the “pit” of stone gametes) occurs in the flower. The sexual cycle of plant
fruits (Fig. 7b) or the shell of walnuts, the covering is a reproduction starts with meiotic cell divisions that
hardened portion of the pericarp, but it is dehiscent (splits halve the number of chromosomes in male pollen cells
along an existing suture line) and usually can be removed and female cells in the embryo sac.
without much difficulty.
Pollen Development (Microsporogenesis)
Pericarp Seed coat
Male gametes are formed in the pollen grains
(microspores) that are produced within the stamen of
the flower (Fig. 9). Pollen or microspore mother cells
located within the stamen divide meiotically to form
tetrads (four haploid microspores). These are sur-
rounded by a nutritive cell layer called the tapetum.
The exine is the outer pollen layer that provides protec-
tion for the pollen grain. The exine tends to be smooth
in wind-pollinated plants and rough or spiked in
(a) insect-pollinated plants. A mature pollen grain typi-
Exocarp Endocarp
cally is two or three celled—one or two generative cells
Seed and a tube cell (Fig. 9). The tube cell functions during
pollen tube growth and the two generative cells are
involved in fertilization.

Ovule Development (Megasporogenesis)

The ovule begins development within the nucellus
of the female cones (gymnosperms) or flower
(b) (angiosperms) (Figs. 10 and 11). The nucellus is sur-
rounded by one or two integuments that grow to even-
Figure 7 tually cover the nucellus. A megaspore mother cell is
Fruit structures included as the “seed” unit. (a) Sunflower
initiated in the nucellus that divides and begins meio-
“seeds” actually include the entire fruit, called an achene.
(b) Plum is an example of a pome (stone fruit) where the sis. There are four linear nuclei formed at the
inner part of the fruit (endocarp) adheres to the seed and end of meiosis. Only one nucleus survives to divide
usually part of the seed unit.

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(a) (b) (c)

Figure 8
Specialized seed structures. (a) Red aril on a black seed in glory bower
(Clerodendrum). Arils are usually developed from outgrowth of the funiculus.
(b) Elaiosome on twinleaf (Jeffersonia). (c) Elaiosome in the euphorbia family is
called a caruncle (castor bean, Ricinus). Elaiosomes are nutrient rich and usually
derived from the outer layer of the seed coat. They are part of a strategy for seed
dissemination by ants called myrmecochory. (d) Apical hairs aid in wind dispersal
(d) of butterflyweed (Aesclepias).

Developing
tetrads of pollen Tube cell
grains (microspores)
Tapetum
Generative cell
Pollen sac

(a) (b)
Figure 9
Pollen development in a typical angiosperm. (a) Within the pollen sac, meiotic divisions give rise to the male gametes contained
within a pollen grain. The tapetum is a nutritive layer of cells enclosing the pollen grains. (b) Mature pollen grain containing a
tube and generative cell.

and form the archegonia in gymnosperms or the antipodal cells at the opposite end of the embryo sac,
contents of the embryo sac in angiosperms. In and the central cell with two polar nuclei.
angiosperms, the most common arrangement of cells in
the embryo sac is called the Polygonum type and occurs RELATIONSHIP BETWEEN
in about two-thirds of flowering plants (Fig. 12).
This type of embryo sac has seven cells (eight
FLOWER AND SEED PARTS
nuclei) that occupy specific locations that dictate The initiation of seed formation generally requires two
their function (72). These cells include the egg appara- processes—pollination and pollination The
tus consisting of a single egg and two synergid cells fertilization. Pollination is transfer of male pollen
located at the micropylar end of the embryo sac, three the transfer of pollen within to the female stigma.

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Figure 10
Development of the female
gametophyte in a representative
gymnosperm (pine). (a) The
megaspore mother cell (arrow)
develops in the female nucellar
tissue. (b) Two archegonia (red
arrow) form, each containing a
(a) (b) female egg cell (black arrow).

a single flower (self-pollination) or from separate flowers acts to guide the pollen micropyle An
(cross-pollination) to a receptive stigma. Pollen is trans- tube, while the genera- opening between the
ferred to the stigma by a variety of means including wind, tive nuclei will eventually integuments through
insects, and, in some cases, mammals. The basic parts of fuse with female egg which the pollen tube
an angiosperm flower are illustrated in Figure 13. The cells. The pollen tube enters the ovule.
pollen grain interacts with a receptive stigma and germi- enters the micropyle (a integuments Two
nates. A pollen tube grows down specialized cells in natural opening between layers of cells that
the style called transmitting cells toward the embryo the integuments) releas- develop between the
sac. The pollen tube con- ing the generative nuclei nucellus and embryo
transmitting cells tains three nuclei: one into the embryo sac. sac and become the
Specialized cells in the tube nucleus and two Fertilization is the seed coat.
style that conduct the generative nuclei (Fig. fusion of haploid (1n)
pollen tube to the 14). The tube nucleus male and female gametes inside the ovule. In gym-
ovule. nosperms, there is a single fertilization between the

Integuments

Figure 11
Development of the
embryo sac in a
representative angiosperm
Nucellus (lily). (a) The megaspore
mother cell develops in
(a) (b) (c) (d) the flower’s nucellar tissue.
(b) Meiosis results in one
viable and three
degenerative nuclei. (c and
e) Progenitor nucleus for
the embryo sac. (d, f, and
g) Embryo sac within the
ovule bounded by the
Embryo sac integuments and attached
Integuments to the ovary by the
funiculus. It is common for
Micropyle the ovule to turn during
development. The
Funiculus
orientation illustrated is
the most common form,
(e) (f ) (g) called anatropous.

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Meiosis I Meiosis II

2n 1n 1n

(a) Megasporogenesis (b) Megagametogenesis

Antipodals Embryo sac

Polar nuclei Figure 12

Development of the most common form of
Egg cell embryo sac (Polygonum type). (a) Initially a
mother spore or mother cell develops in the
nucellar tissue of the flower. Four haploid cells
Synergid Synergid
are formed during meiosis, but only one is
(c) retained. (b) It then divides to form the cell in
the embryo sac. Each cell has a distinct role.
(c, d) Three become antipodals, one is the
central cell with two polar nuclei, two become
(d) synergids, and one becomes the egg cell.

sperm and egg cells. In angiosperms, double fertiliza- and its two polar nuclei to form the 3n endosperm
tion occurs. (Fig. 15). The female synergid cells are closely associ-
Double fertilization (5) occurs when one gen- ated with the egg cell and function to attract and
erative nucleus fuses with the egg cell to form the guide male nuclei to the egg cell for fertilization (31).
zygote (2n embryo), Synergids produce a chemical that attracts the pollen
zygote The result of while the second gen- tube to the micropyle, arrests its growth, and ensures
sexual reproduction, erative nucleus fuses the proper release of the sperm cells into the ovule.
which forms the embryo. with the central cell Evidence suggests that the central cell signals the

Petal

Stigma

Anther

Style

Ovary

Receptacle

Pedicel

(a) (b)
Figure 13
In a typical angiosperm flower, floral organs are produced in separate whorls. The outermost whorl are the sepals (caylx), the
next are the petals (corolla), inside the petals are the male stamens, and innermost is the female pistil. Pollination occurs with
the transfer of pollen from the stamens to the stigma of the pistil. The pollen grain germinates and the pollen tube grows down
the style. Eventually, the pollen tube enters the ovule through the micropyle and deposits two male sperm cells. Fertilization
involves the fusion of the male and female cells in the embryo sac.
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 seed development

(a) (b) (c)

Figure 14
Pollen (male gametophyte). (a) Stamen pair opening along a suture line to shed
pollen. (b) Pollen on the stigma of hibiscus. (c) Close-up of pollen grain showing
(d) (e) the surface structure (exine). (d and e) A germinating pollen grain.

synergids to release the chemical attractant. The 4. Polar nuclei plus a generative nucleus become the
synergid cell degenerates soon after sperm cell release, endosperm.
permitting sperm cell access to the egg cell for fertil- 5. Egg cell fuses with one generative nucleus to form
ization and release of the second sperm cell to migrate embryo.
to the central cell. The exact function of antipodal 6. Integuments form the layers of the seed coat (also
cells is not completely understood, but they disinte- called testa).
grate soon after fertilization of the egg cell.
Fertilization in gymnosperms differs from
The relationship between flower tissue and subse-
angiosperms because they do not produce elaborate
quent parts of the fruit and seed for a typical
flower parts. There is no true stigma in gymnosperms.
angiosperm species is outlined as follows:
Rather, there is either a
1. Ovary grows into fruit tissue. stigmatic surface on the ovule Develops in the
2. Ovule becomes the mature seed. opening of the ovule or nucellus and is enclosed
3. Embryo sac is the inner part of the seed. a sugary pollination by the integuments.

Embryo sac

Figure 15
Endosperm
Double fertilization in lily. One
sperm nucleus fuses with the egg
cell to form the zygote and the
Zygote other male nucleus fuses with the
polar nuclei to form the triploid
endosperm. (a) Shows the embryo
sac within the developing ovule.
(b) Is a close up of the embryo sac
showing the onset of cell division
(a) (b) following double fertilization.
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 seed development

BOX 1 GETTING MORE IN DEPTH ON THE SUBJECT

PLOIDY LEVELS IN PLANTS
With many angiosperms, the zygote is diploid (2n) and Crosses between species may fail to produce viable
divides to become the embryo; the endosperm is triploid seed because the species have different ploidy levels.
(3n) and develops into nutritive tissue for the developing Failure of the endosperm to develop properly can also
embryo. Terminology for ploidy levels in plants can be con- result in retardation or arrest of embryo development,
fusing. Ploidy indicates the number of sets of chromosomes and embryo abortion can result. This phenomenon is
in a plant. Not all plant species are diploid. Several impor- called somatoplastic sterility and commonly occurs when
tant crop plants, like potato, are tetraploid or even octa- two genetically different individuals are hybridized,
ploid like strawberry. However, the product of normal meio- either from different species (15, 16, 17) or from two
sis is still to produce gametes with half the original number individuals of different ploidy constitution. It can be a
of chromosomes. Therefore, current terminology for a barrier to hybridization in angiosperms but not in gym-
tetraploid species is 2n ⫽ 4x, where “x” is the number of nosperms (62), since the “endosperm” in these plants is
pairs of chromosomes. For example, in potato the ploidy haploid female gametophytic tissue. Embryos that show
level of the diploid would be expressed as 2n ⫽ 2x ⫽ 24, some growth from these types of crosses can be “res-
while the tetraploid would be 2n ⫽ 4x ⫽ 48. This means cued” by isolating these embryos and placing them in
that there are 24 chromosomes in the diploid and 48 in the tissue culture.
tetraploid.

drop exudes from the ovule to collect wind-borne cotyledon stage of development. There is rapid increase
pollen (62). In some species, like Ginkgo, the male in both fresh and dry weight. There are characteristic
gametes can be motile, but, in most cases, the pollina- stages of embryogenesis that occur during Stage I and
tion droplet pulls the pollen into the ovule and a pollen these are distinct for dicots, monocots, and gym-
tube is formed. Double fertilization does not occur in nosperms.
gymnosperms. However, the gymnosperm, Ephedra,
Embryo Differentiation in Dicots Although there
(in the Gnetophte group, which is possibly the progen-
are several variations on the types of angiosperm
itor line for the angiosperms) has a form of double fer-
tilization, but no endosperm results from the second
fertilization. Only angiosperms produce a true triploid Stages of Seed Development
Histo- Maturation
(3n) endosperm. In gymnosperms, haploid female differentiation Cell expansion
drying
gametophyte tissue surrounds the developing embryo
and performs the function of the endosperm. Fresh wt

STAGES OF SEED Water wt

Gram weight

DEVELOPMENT
Three physiological stages of development are recog-
nized in most seeds (Fig. 16). These include Dry wt
histodifferentiation, cell expansion (food reserve
deposits), and maturation drying. Figure 17 shows the
relative growth and development in lettuce seed (fruit),
showing the physiological stages of seed development Stage I Stage II Stage III
and days post-pollination. Days of Development

Figure 16
Stage I Histodifferentiation (Embryo The stages of seed development. The stages include
Differentiation) histodifferentiation (rapid increase in seed size due
predominantly to cell division), cell expansion (largest
Stage I is characterized by the differentiation of the increase in seed size for deposition of food reserves), and
embryo and endosperm mostly due to cell division. In maturation drying (dramatic loss in seed fresh weight due to
Stage I, the embryo reaches the beginning of the water loss). Redrawn from Bewley and Black, 1994.

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 seed development

Figure 17
Growth and development of the
fruit and seed in lettuce showing
the relative changes in seed size
during the three stages of seed
development. P, pericarp;
I, integuments; N, nucellus; EN,
endosperm; EM, embryo. Redrawn
from Jones 1927.

embryogenesis (8, 51, 59, 72), embryo formation in inner cell layers will develop into the procambium
shepherd’s purse (Capsella bursa-pastoris) has served as a and ground meristem.
good model for dicot embryogenesis and is very similar As the embryo enters the cotyledon stage, the
to Arabidopsis. Embryogenesis in dicots proceeds cotyledon primordia are evident in the heart-shaped
through the characteris- stage of embryogenesis (Fig. 18g–i). These primordia
proembryo The tic stages of develop- elongate to give a typical torpedo stage embryo (Fig.
earliest stages of ment. These include the 18g). In the torpedo stage, the embryo has organized to
embryo development proembryo, globular, form an apical meristem, radicle, cotyledons, and
before the embryo and heart, torpedo, and hypocotyl. The endosperm has been developing along
suspensor become cotyledon stages (Fig. with the embryo and providing nutrition for its
easily recognized. 18). growth. When the embryo reaches the mature stage
Following fertil- (Fig. 18j–l) in shepherd’s purse, the major storage tissue
ization of the egg and sperm nuclei, a proembryo is is the cotyledons, which now occupy most of the seed
initiated by a transverse cell division to form an cavity.
apical and basal cell (Figure 18a–c). The basal cell
forms the suspensor, while the apical cell forms the
embryo. The suspensor in dicots is usually a column Embryo Differentiation in Monocots Monocots have
of single or multiple cells. The suspensor functions to a more complex embryo structure in the mature seed
push the proembryo into the embryo sac cavity and compared with dicots, but early embryo development
to absorb and transmit nutrients to the proembryo. is similar (60). Embryogenesis in monocots includes
The embryo is supplied with nutrients for growth via the proembryo, globular, scutellar, and coleoptilar
the suspensor until later stages of embryo develop- stages (Fig. 19).
ment when the embryo is nourished by material from Following fertilization, an apical and basal cell is
the endosperm. There is also hormone signaling visible in corn (Zea mays) that initiates the proembryo
between the suspensor and embryo. In shepherd’s stage (Fig. 19a). The proembryo and globular stages are
purse, basal cell derivatives in the globular embryo similar to dicots, except that the suspensor is not a sin-
form the hypophysis that goes on to develop into the gle or double row of cells and is less differentiated (Fig.
radicle (Fig. 18d). Tissue differentiation becomes evi- 19b). In the late globular stage, the outer epidermal
dent in the sixteen-celled globular embryo layer is evident and a group of cells on one side of the
(Fig. 18d–f ). An outer layer of cells (protoderm) will proembryo divides more rapidly; these will give rise to
develop into epidermal cells of the embryo. The the embryo axis.
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 seed development

(a) Proembryo Stage (d) Globular Stage (g) Cotyledon Stage (j) Mature Stage

(b) (e) (h) Integuments (k)

Ovule Fruit wall
Fertilized Ovule Integuments
Ovule egg cell
Funiculus
Embryo Ovules
Endosperm Endosperm

Integuments
Funiculus
Embryo
Basal cell

Funiculus

Apical
(c) Fertilized (f ) Globular (i) Heart-shaped (l) meristem

Ovule egg cell Suspensor embryo embryo Seed

coat

Radicle

Basal Funiculus
cell
Cotyledons

Suspensor

Figure 18
Embryo development in a typical dicot (shepherd’s purse) showing the proembryo (a–c), globular (d–f), cotyledon (g–i), and
mature (j–l) stages. See text for detailed description of each stage.

The remnant of the cotyledon can be seen in the Finally, the embryo axis differentiates into
scutellar stage of development. Monocots have the plumule (shoot) and radicle in the coleoptilar
reduced the pair of cotyledons represented in dicot stage (Fig. 9d). In monocots, the embryo axis also has a
embryos to a single modified cotyledon termed the specialized tissue surrounding the shoot and root tissue
scutellum (Fig. 9c). The scutellum acts as conductive to aid in emergence during germination. These are the
tissue between the endosperm and embryo axis (Fig. coleoptile and coleorhiza, respectively (Fig. 9d–e).
9d–e).

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 seed development

Globular Stage

Proembryo Stage

) Endosperm

Coleoptile
(a) (b)

Scutellar Stage Coleoptilar Stage

Shoot axis

Radicle
Pericarp
Scutellum

Coleorhiza

(c) (d) (e)

Figure 19
Embryo development in a typical monocot (corn). See text for description of figure. (e) Cross section of a mature seed of corn
showing basic anatomical features.

Embryo Differentiation in Gymnosperms Compared within a single gymnosperm seed but rarely does more
with the more evolutionarily advanced angiosperms, than one of these embryos mature.
embryo formation in gymnosperms (62) differs in In pine (Pinus sp.), the fertilized egg cell divides
several important ways (Fig. 20). Most conspicuous is to form a free nuclear stage without cell walls
that seeds of gymnosperms are not contained within a between nuclei (Fig. 20b). Following cell wall forma-
carpel or ovary (fruit). The term gymnosperm means tion, cells organize to form an embryo tier of cells
“naked seeded.” Only a single fertilization occurs in and a suspensor tier (Fig. 20c). The suspensor differ-
gymnosperms (Fig. 20a). Therefore, there is also no entiates into a set of primary suspensor cells (rosette
true triploid endosperm in gymnosperms. Rather, the cells) and embryonal suspensor tubes. The suspensor
developing embryo is nourished by haploid female cells elongate and there are several cleavage events to
gametophyte tissue also referred to as an endosperm give multiple embryos (polyembryos) inside a single
(Fig. 20e). Pollination and fertilization may be seed (Fig. 20d). Usually, only one of these embryos
separated by months (up to 12 months in pine), and continues to develop. The proembryo differentiates
seed formation can take two seasons in some species. an epidermal layer (Fig. 20d) prior to the cotyledon
The pollen tube germinates soon after pollination but primordia becoming evident. The mature pine
must wait for the female gametophyte to complete embryo has multiple (usually eight) cotyledons com-
development before fertilization can proceed. After pared to two or one in the dicots and monocots (Fig.
fertilization, several embryos begin development 20d).

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 seed development

Fertilization Free nuclear stage Suspensor tier stage

Embryo
tier

(b)

Egg

Sperm Suspensor
tier

Rosette
(a) cells
(c)
Proembryo Stage Cotyledon Stage

(e)

Figure 20
(d) Embryo development in a typical gymnosperm (pine). See text for description of figure.

Stage II Cell Expansion substrates to ensure survival of the germinating

Stage II is a period of rapid cell enlargement—often seedling, but also provide essential food for humans
called seed filling—due to the accumulation of food and animals.
reserves (Fig. 16). This is an active period with large Food reserves are manufactured in the develop-
increases in DNA, RNA, and protein synthesis in the ing seed from photosynthate being “loaded” or
seed (7). The major food reserves include carbohydrates moved into the seed from the mother plant. The
(starch), storage proteins, and lipids (oils or fats). process of seed reserve accumulation requires the
Although different species may predominantly store a translocation of small molecular weight compounds,
particular food reserve (i.e., cereal grains store starch, such as sucrose, asparagine, glutamine, and minerals,
legumes store protein, and sunflower stores oil), most into the seed. In dicot seeds, there is a direct vascular
seeds contain all three types of food reserves (Table 2). connection (phloem, xylem) between the mother
Such substances not only provide essential energy plant and the seed through the funiculus (Fig. 21).

132
 seed development

Table 2
F OOD R ESERVES F OUND IN VARIOUS P LANT S PECIES
Average percent composition
Species Protein Oils Starch Major storage organ
Cereals 10–13% 2–8% 66–80% Endosperm
Oil palm 9% 49% 28% Endosperm
Legumes 23–37% 1–48% 12–56% Cotyledons
Rape seed 21% 48% 19% Cotyledons
Pine 35% 48% 6% Female gametophyte

Source: From (7, 18, 62).

A vascular strand usually runs through the funiculus embryo in this process, but may accumulate in the
and down one side of the integuments (seed coat), outer layers of the seed. There is no vascular connec-
allowing transfer of photosynthate and water into the tion between the mother plant and developing seed
developing seed (30). There is no direct vascular con- in monocots. Rather, there is a group of cells at the
nection from the seed coat to the nucellus, seed and mother plant interface called transfer cells
endosperm, or embryo, and assimilates must reach that facilitate the passage of photosynthate into the
the embryo by diffusion (75). Most viruses and large endosperm (61).
complex molecules are effectively screened from the

Endosperm
Embryo

Seed coat

Vascular
Funiculus Embryo
trace

(a)
Figure 21
(a) Longitudinal section
through a developing ovule
Seed of eastern redbud (Cercis
coat canadensis) about 57 days
Funiculus
post-anthesis (pollen
shedding) showing the
vascular connection between
the funiculus and the ovule.
Fruit pod
(b) Close-up of the vascular
trace. Note typical xylem
cells in the vascular trace.
Funiculus (c) Bean seed with funiculus
attached to the pod.
(b) (c) From Jones and Geneve (36).

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 seed development

BOX 2 GETTING MORE IN DEPTH ON THE SUBJECT

GENE EXPRESSION DURING SEED FILLING (69)
Specific mRNAs are required for the synthesis of storage Figure 22b shows the increase in mRNA that precedes the
compounds (7, 26, 70). The pattern of mRNA for storage accumulation of the storage protein, cruciferin, in rape
protein accumulation is similar for a number of proteins seed (Brassica napus) (24).
and mRNAs including phaseolin, legumin, and vicilin in Very specific genes are “turned on” during this stage
legumes; cruciferin in rape seed; and zein and hordein in of embryo growth (26, 67). These genes are only
cereals. A typical pattern for storage protein accumulation expressed during the embryogenesis stage of a plant’s life
is illustrated in Figure 22a for broad bean (Vicia faba). cycle. The mRNAs for storage proteins are no longer
This increase in storage protein is coincident with the translated after maturation drying and cannot be
increase in dry weight accumulation in Stage II embryos. detected in germinating seeds.

(a) (b)
Figure 22
Accumulation of storage proteins related to the stages of seed development. (a) Pattern of protein accumulation in broad
bean (Vicia faba) for vicilin and legumin, two major seed storage proteins in beans. (b) Accumulation of cruciferin protein
and its mRNA in rape seed (Brassica napus). Note how the mRNA for the protein is only expressed at high levels during
Stage II of seed development and is not detectable following maturation drying. Redrawn from Finkelstein and Crouch, 1987.

BOX 3 GETTING MORE IN DEPTH ON THE SUBJECT

BIOTECHNOLOGY OF SEED RESERVES (14)
The food reserves in seeds make up a major part of the transformation technology to insert new genes into crop
world’s diet both for human and livestock consumption. plants to produce storage proteins high in lysine or sulfur.
The nutritional quality of seeds can be improved by under- New germplasm is being developed that will increase the
standing the molecular genetics responsible for food nutritional yield of some of our major crop plants. For
reserve production. There are efforts through genetic example, genetic engineering of rice has resulted in grains
engineering to improve the amino acid content of storage containing beta-carotene that could serve as a major
proteins in seeds (44, 64). source of this essential nutrient for a large portion of the
Cereals and legumes are important to worldwide world’s population (27).
diets, and their yield and nutrition have been improved Plants that store oils in seeds are also the target of
significantly over years of conventional breeding. However, increased efforts to produce novel oils that can be used
most cereal proteins are nutritionally low in the essential for detergents, lubricants, and cooking oil that is healthier
lysine-containing amino acids, and legume seeds produce by producing lipids low in unsaturated fat (32). Canola
storage proteins low in essential sulfur amino acids. The (rape seed) and soybeans are major crops being bioengi-
amino acid profile of these seeds can be improved using neered to produce novel oils.

134
 seed development

Stage III Maturation Drying The low moisture level attained by dry seeds is a
Seeds at the end of Stage II of development have reached remarkable plant condition (9). Many plant tissues
physiological maturity cannot tolerate moisture levels much below ∼20 per-
perisperm Nucellus
(also called mass matu- cent on a fresh weight basis for a prolonged time. Dry
tissue that remains in
rity). Physiological matu- orthodox seeds can usually remain viable at 3 percent
the mature seed and is
rity is the time prior to to 5 percent moisture. Orthodox seeds prepare for
used as storage tissue.
maturation drying when maturation drying towards the end of Stage II prior to
the seed has reached maximum dry weight through physiological maturity. Abscisic acid (ABA) is the
reserve accumulation. Seeds at physiological maturity main signal for induction of desiccation tolerance.
can be removed from the fruit and show high germina- The physiological mechanisms for tolerating very dry
tion potential as measured by seed viability and vigor conditions are not totally understood, but they are
(52). Seeds that do not tolerate desiccation drying are
called recalcitrant seeds
recalcitrant seeds (see Box 4) and are usu-
Seeds that are unable ally shed from the plant
to withstand maturation at this stage without
drying. entering Stage III: mat-
uration drying.
Orthodox seeds tolerate maturation drying and
0.5
represent the condition of most crop seeds. Seeds in the
maturation drying stage are characterized by rapid water Seed Weight (mg)
loss (Fig. 23). There is no longer a vascular connection 0.4

with the mother plant through the funiculus. Water loss

occurs throughout the 0.3
funiculus The
attachment between
seed coat but may be
the ovary and the ovule.
more rapid where there 0.2
are natural openings at
hilum The scar left on the hilum (scar left 0.1
the seed coat after the on the seed coat after Time
funiculus abscises. funiculus detachment)
Figure 23
and micropyle. In species that develop impermeable seed Water loss in honeylocust (Gleditsia triacanthos) seeds during
coats as a form of dormancy, the final quantity of water development. Note the typical loss of chlorophyll during
leaves the seed at the hilum (34). maturation drying and the overall reduction in seed size.

BOX 4 GETTING MORE IN DEPTH ON THE SUBJECT

RECALCITRANT SEEDS
After developing seeds reach physiological maturity, they The biological basis for this inability in recalcitrant
proceed to desiccate (orthodox seeds), germinate on the seeds to tolerate drying is not well understood (6).
plant (vivipary), or bypass complete desiccation (recalci- Arabidopsis is an orthodox seeded species and its
trant seeds). By definition, a recalcitrant seed loses viabil- mutants have been a very useful tool for physiologists try-
ity after drying, while orthodox seeds tolerate drying (7). ing to study a variety of processes in plants. Arabidopsis
Germination in recalcitrant seeds must proceed soon after mutants have been found with reduced levels of ABA, LEA
maturity or the seeds must be stored under conditions proteins, and carbohydrates, and these mutants are
that prevent drying. Examples of storage life for some impaired for tolerance to drying. These substances are
recalcitrant seeds stored at high humidity include coffee thought to be critical for survival in orthodox seeds during
(Coffee arabica) for 10 months, coconut (Cocus nucifera) for desiccation drying. It would seem logical that recalcitrant
16 months, and oak (Quercus) for 20 months, compared to seeds have reduced ABA levels or that they are impaired
decades or years for many orthodox seeds. Recalcitrant for the production of LEA proteins or some carbohy-
seeds present challenges for propagators and limit germ drates. However, most recalcitrant species produce these
plasm conservation because of their inability to store. substances at almost normal levels. The true nature of
orthodox seeds Seeds that tolerate maturation drying recalcitrance drying remains to be found for this interest-
and survive at less than 10 percent moisture. ing group of seeds (23).

135
 seed development

correlated to an increase in sugars (especially di- and

oligosaccharides) and LEA (late embryogenesis abun-
dant) proteins (see Box 5) (33). These are thought to
preserve proteins and membranes by replacing
the water function as cells become dry and enter a
highly viscous state termed glassy (water replacement
theory).
As indicated earlier, seeds also acquire the ability
to germinate in Stage II prior to maturation drying.
Usually, this potential to germinate is not expressed in
orthodox seeds unless the fruit is removed from the
plant and the seeds are gradually dried (38, 39).
Germination of seeds prematurely on the plant without
Figure 24
desiccation drying is termed precocious germination or Precocious or viviparous germination occurs when the seed
vivipary (Fig. 24 and Box 6). It is usually the result of a prematurely germinates in the fruit. This is the result of the
mutation in the ability to produce or perceive ABA. developing seed not completing the third stage of
During normal seed development, the seed does not development—maturation drying. The cause of precocious
germinate prior to maturation drying because of high germination is usually the inability of the embryo to produce
or perceive abscisic acid (ABA). ABA is a potent germination
ABA content in the seed and, for some seeds, the low
inhibitor and one of its roles during seed development is to
water potential in the fruit coverings caused by high prevent precocious germination. The tomato illustrated here
salt and sugar content. is most likely an ABA production mutant.
Following maturation drying, the seed can be
considered in a quiescent or dormant condition.
Quiescent seeds fail to germinate because they are UNUSUAL TYPES OF SEED
dry. Exposing quiescent seeds to a favorable environ-
ment will induce them to germinate. Dormant seeds
DEVELOPMENT
fail to germinate even under favorable environmental Apomixis and polyembryony represent variations from
conditions. There are several ecological advantages to the normal pattern of zygote formation and embryoge-
seed dormancy and it is a common feature of many nesis. Although related, they are not necessarily the
seeds. Over years of selection, dormancy has been same phenomenon. Apomixis
bred out of most economically important crop is the asexual development of apomixis
species. seeds that represent clonal Asexual seed
duplicates of the mother plant. production.

BOX 5 GETTING MORE IN DEPTH ON THE SUBJECT

GENE EXPRESSION DURING MATURATION DRYING
Maturation drying can be considered a “switch,” ending drying below 15 percent moisture (55). LEA proteins
the seed’s developmental program and preparing the seed appear to help the seed adjust to a dry condition. In addi-
for germination (22, 37, 40). Synthesis of developmental tion, the seed is also protected during desiccation by an
proteins stops prior to drying, and a new set of proteins is increase in certain sugars and oligosaccharides that also
synthesized (38, 39); a major set of these proteins is called provide stabilization to proteins and membranes (2, 10).
LEA (late embryogenesis abundant) proteins (33). LEA pro- Also during maturation drying, mRNAs for early ger-
teins are synthesized in response to water loss in the seed. mination are produced (19, 67). These are called con-
LEA proteins are very stable and hydrophilic (attracts served mRNA because they are stored in the dry seed and
water), and possibly function as desiccation protectants by expressed early in germination. Although conserved
stabilizing membranes and proteins as the seed dries. mRNAs are lost in the first few hours of germination, they
There are many ecological advantages to the production allow the seed to produce proteins essential for germina-
of a dry seed for seed dissemination and seed survival. tion before the embryo regains the capacity to synthesize
However, there are few living organisms that can survive new mRNAs.

136
 seed development

BOX 6 GETTING MORE IN DEPTH ON THE SUBJECT

PRECOCIOUS GERMINATION OR VIVIPARY
Precocious germination or vivipary is the phenomenon in many species including cereal grains (wheat and corn),
which seeds precociously germinate without maturation fleshy fruits (citrus and tomato), and nuts (pecan).
drying. These seeds germinate in the fruit while still Precocious germination is considered a genetic mutation,
attached to the plant (Fig. 24). Precocious germination but occurrence of precocious germination can be modi-
occurs naturally in some species like mangrove (Rhizophora fied by the environment (71). Expect increased precocious
mangle). In mangrove, precocious germination is an adap- germination in susceptible species during periods of wet
tation to growing in a wet (swampy) environment. Embryos weather (7).
germinate directly on the tree to produce seedlings with a The genetics of viviparous mutants in corn has been
long, javelin-shaped root (Fig. 25). The seedling eventually most extensively studied (50). Up to nine genes have been
falls and becomes embedded in the mud below (65). associated with precocious germination in corn. The com-
vivipary Germination of a seed while it is still attached mon feature in viviparous mutants is reduced production,
to the mother plant. or insensitivity to abscisic acid (ABA). This supports the
For most plant species, however, precocious germina- role for ABA in maintaining the embryo in the develop-
tion is undesirable. Premature seed sprouting occurs in mental mode through maturation drying.

(a)

Figure 25
Precocious (viviparous) germination
in mangrove (Rhizophora mangle).
(a and b) Note the protrusion of the
radicle from the fruit while it is still
attached to the plant. (c) After
sufficient radicle growth the fruit
will fall from the plant and embed
in the soft marshy soil around the
(b) (c) mother plant.

polyembryony Two types of apomixis will be the same as the seed parent. Seed production
The development of are known: gameto- via apomixis is asexual. Such clonal seedling plants
multiple embryos within phytic and sporophytic are known as apomicts. Some species or individuals
the same seed. apomixis (41, 68). produce only apomictic embryos and are known as
Polyembryony means obligate apomicts; however, the majority of apomic-
that more than one embryo develops within a single tic species produce both apomictic and sexual embryos
seed, sometimes many (Fig. 26). on the same plant and are known as facultative
apomicts (46).
Apomixis Apomixis can be further divided into gametophytic
Apomixis (53, 54, 59) is the production of an embryo versus sporophytic apomixis (see Box 7). From a horti-
that bypasses the usual process of meiosis and fertiliza- cultural production standpoint, sporophytic apomixis
tion. The genotype of the embryo and resulting plant is the most significant because it is the type of seed
137
 seed development

Nonrecurrent Apomixis In nonrecurrent apomixis,

meiosis does occur and an embryo arises directly from
the egg nucleus without fertilization. Since the egg is
haploid, the resulting embryo will also be haploid.
This case is rare and primarily of genetic interest. It
does not consistently occur in any particular kind of
plant, as do recurrent apomixis and adventitious
embryony.

Polyembryony (46)
In 1719, Leeuwenhoek reported the first account of poly-
embryony in plants when he observed the production of
twin embryos in Citrus. Polyembryony is the production of
additional embryos within a seed other than the normal sex-
ual embryo (43). The multiple embryos could be all sexual
or a mixture of sexual and asexual (apomictic) embryos.
Four types of polyembryony are recognized in angiosperms:
Figure 26 1. After the normal sexual embryo begins to form,
Polyembryony in trifoliate orange (Poncirus trifoliata) seeds as additional embryos can “bud-off ” from the proem-
shown by the several seedlings arising from each seed. One
seedling, usually the weakest, may be sexual; the others arise
bryo (found in Asparagus, Tulipa gesneriana, and
apomictically from cells in the nucellus and are diploid copies Hamamelis) or suspensor cells (found in Acanthus).
of the mother plant. The result is a sexual embryo and multiple copies
of that sexual embryo.
2. Adventive embryony results in additional embryos
production that predominates in Citrus, mango formed from cells in nucellar (found in Citrus,
(Mangifera), and mangosteen (Garcinia) and allows for Mangifera, and Garcinia) or integuments (found in
clonal understock production from seeds for grafting or Spiranthes cernua). The result is one sexual and
budding (11). Gametophytic apomixis results in multiple multiple asexual embryos.
clonal embryos developing from nucellar (rarely, integu- 3. Multiple embryo sacs may be formed within a sin-
ment tissue) surrounding a normally developing sexual gle ovule (seed). This has been observed in species
embryo sac. The seed usually contains one sexual embryo of cotton (Gossypium). The result can be multiple
and multiple asexual embryos (Fig. 27). Often the sexual embryos from separate fertilizations or mul-
seedling developing from the sexual embryo is easily tiple asexual embryos from aposporic apomixis
identified as the weakest seedling in the group. This type (Box 7).
of apomixis is a form of polyembryony and is termed 4. Additional embryos may result from a synergid cell
adventitious embryony (also nucellar embryony and functioning as an egg cell. This can result from
nucellar budding). fertilization of the synergid by a male sperm cell or

Figure 27
Development of nucellar
embryos in Citrus. Left: Stage
of development just after
fertilization showing zygote and
remains of pollen tube. Note
individual active cells (shaded)
of the nucellus, which are in the
initial stages of nucellar
embryony. Right: A later stage
showing developing nucellar
embryos. The large one may be
the sexual embryo. Redrawn from
Gustafsson, 1946.

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 seed development

from autonomous cell divisions in reduced syn- PLANT HORMONES AND SEED
ergids resulting in a haploid apomictic embryo.
DEVELOPMENT
These types are found in Pennisetum, Tamarix, and
Solanum. In general, concentrations of plant hormones are high
in seeds compared with other parts of the plant (7).
Vegetative Apomixis Seeds were the first tissue where several of the plant
hormones were discovered and studied in detail. All of
The term apomixis has been used in the past for any form
the major hormones have been associated with seed
of vegetative propagation. Today, its usage has been
development (57, 58). Plant hormones are involved in
restricted to asexual production of an embryo within the
seed development in several ways:
ovule of flowering plants. However, some references still
include the term vegetative apomixis to describe the pro- 1. growth and differentiation of the embryo
duction of other structures besides an embryo. In some 2. accumulation of food reserves
cases, vegetative buds or bulbils are produced in the 3. storage for use during germination and early
inflorescence in place of flowers. This occurs in Poa bul- seedling growth, and
bosa and some Allium, Agave, and grass species. 4. growth and development of fruit tissue

BOX 7 GETTING MORE IN DEPTH ON THE SUBJECT

APOMIXIS (42)
Most seed plants produce embryos after the fusion of hap- rather than the normal 1n cells found in the embryo sac.
loid female and male gametes. However, in a small propor- Otherwise, the embryo sac has a normal appearance. This
tion of plants, embryos form spontaneously from cells that type of apomixis is common in the Asteraceae and in some
bypass meiosis. These embryos have the same genotype grass species. In apospory, the megaspore mother cell
as the mother plant, and this process is called apomixis. undergoes normal meiosis, but the resulting cells usually
The term apomixis is from the Greek apo, meaning degrade before they are fertilized. Additional cells in the
from or away from, and mixis, to mingle or mix. Apomixis nucellus become densely cytoplasmic and take on the role of
literally means “away from mixing” and refers to the pro- apomictic initials bypassing meiosis to form cells in their own
duction of new plants without mixing of gametes. Apomixis embryo sacs within the same ovule. Aposporous apomicts
occurs in over 300 species from at least 35 plant families are found in the Rosaceae, Asteraceae, and in some grasses.
(12). Most display facultative apomixis. It is most common In some apomictic species, such as Crepis, dandelion
in the daisy (Asteraceae), grass (Graminaceae), rose (Taraxacum), bluegrass (Poa), and onion (Allium), there is
(Rosaceae), and citrus (Rutaceae) families. no need for the stimulus of pollination; in others [e.g.,
There are two general categories for apomixis. These species of guayule (Parthenium), raspberry (Rubus), apple
are gametophytic and sporophytic apomixis. (Malus), some grasses (Poa species), and coneflower
(Rudbeckia)], pollination appears to be necessary, either
Gametophytic Apomixis to stimulate embryo development or to produce a viable
During normal embryogenesis, female egg cells are derived endosperm.
from the megaspore mother cell found in the nucellus tissue
of the flower. The megaspore mother cell undergoes reduc- Sporophytic Apomixis
tion division (meiosis) to produce haploid nuclei that take up Sporophytic apomixis is also known as adventitious embry-
typical locations in the embryo sac. In gametophytic ony. It has been extensively studied because it is the type of
apomixis, the megaspore mother cell degenerates or apomixis that occurs in Citrus and mango (Mangifera) (Figs.
bypasses meiosis. Unreduced cell(s) divide to produce 26 and 27). In adventitious embryony, the megaspore
reproductive cells that take their normal location in the mother cell undergoes normal meiosis and forms a normal
embryo sac. The unreduced nucleus corresponding to the sexual embryo sac. These cells are fertilized by male sperm
egg cell undergoes spontaneous or parthenogenic divisions cells as in normal embryogenesis. However, at about the
to form an embryo without fertilization from a male gamete. time the first divisions begin in the sexual embryo, cells in
In some cases, polar nuclei can autonomously form the nucellus begin abnormal cell divisions leading to multi-
endosperm, but, in many cases, an unreduced polar nuclei ple embryos forming in the micropylar region of the ovule.
will fuse with a male sperm cell to form the endosperm. This Since these are derived from mother plant cells, the result-
process is known as pseudogamy. ing embryos are asexual. These asexual embryos do not
There are two types of gametophytic apomixis called produce their own embryo sac (thus the term sporophytic
diplospory and apospory. These differ because of the apomixis). Rather, they grow into the embryo sac of the sex-
location in the nucellus where apomictic cells arise. In ual embryo and share its sexually derived endosperm. The
diplospory, the megaspore mother cell does not undergo result is a single seed that can contain a single sexual and
or complete meiosis. Rather, it divides to produce 2n multiple asexual embryos.

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 seed development

BOX 8 GETTING MORE IN DEPTH ON THE SUBJECT

SIGNIFICANCE OF APOMIXIS
Apomixis is significant in agriculture and horticulture (Poa pratensis), ‘King Ranch’ bluestem (Andropogun),
because the seedling plants have the same genotype as the ‘Argentine’ Bahia grass (Paspalum notatum), and ‘Tucson’
mother plant (28). This asexual process eliminates variability side oats grama (Bouteloua curtipendula).
and “fixes” the characteristics of any cultivar immediately. There has been a recent resurgence in research con-
However, the apomictic life cycle has the same juvenile cerning apomixis. It has been known for some time that
period found in sexually derived seeds. apomixis is an inherited trait and that the gene maps to a
Only a few economically important food crops exhibit single chromosome (68). This indicates that there is a sin-
apomixis. These include Citrus, mango (Mangifera), and gle apomixis gene and that if that gene is isolated it could
mangosteen (Garcinia). All three have adventitious embry- be used to genetically engineer apomixis into important
ony. They have mainly been exploited as clonal seedling crop plants. The major benefit would be that apomixis
understocks for grafting and budding because they are would fix hybrid vigor (heterosis) in crops that now require
virus-free, show seedling vigor, and are uniform. However, costly crossing between inbred parents. It would also be a
sexual embryos can also be produced and can exhibit simple way to eliminate virus in traditionally vegetatively
unwanted variability. Use of DNA fingerprinting is being propagated crops like potato. The seed produced would
used to separate sexual and asexual embryos for under- have the same genetics of the parents, but because the
stock production. embryo is derived from a single cell, it should be virus-
Several grass species and cultivars are facultative free.
gametophytic apomicts. These include Kentucky bluegrass

Auxin Gibberellins were originally thought to play only

Free and conjugated forms of indoleacetic acid (IAA) a minor role in seed development. Gibberellin-
are abundant in developing seeds. Free IAA is high dur- deficient mutants in tomato and Arabidopsis gener-
ing cell division stages of development (Stages I and II) ally show normal seed development only affecting
and is essential for normal embryo and endosperm final seed size. However in pea, gibberellins are
development. An auxin gradient is required to establish required for embryo growth (66). In gibberellin-
appropriate bipolar symmetry during embryo develop- deficient mutants that show reduced gibberellin
ment. Mutations that cause seeds to have low auxin biosynthesis, gibberellin is required to sustain
production or reduced auxin transport generally result
in malformed embryos with fused cotyledons and poor
endosperm development (74).
Conjugated forms of IAA are abundant in mature
seeds and during germination. Free IAA is released
from the conjugated forms for utilization during early
seedling growth. There is evidence that auxin from the
developing seed signals the fruit to continue to develop
(Fig. 28). Fruits usually abscise if seeds abort or are
unfertilized. Auxin applied to tomato or strawberry can
induce parthenocarpic fruit development (see Box 9).

Gibberellins
Various forms of gibberellins are abundant during
seed development (Stages I and II). Most of the Figure 28
biochemistry known about gibberellins was first Strawberry “fruit” (receptacle) enlargement requires auxin
investigated in developing seeds. Active forms decline from the developing seed (actually the fruit-achene). Notice
how the only swelling in the receptacle tissue is around the
at seed maturity and are replaced by conjugated
developing achenes (red arrow). The black arrow shows a
forms of gibberellins. Like auxin, these conjugated non-fertilized seed where you can still see the style and
forms of gibberellins are utilized during germination. stigma attached. There is no swelling in this area because
there is no developing seed to provide the auxin.

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 seed development

BOX 9 GETTING MORE IN DEPTH ON THE SUBJECT

PARTHENOCARPY
For many plant species, pollination is the stimulus for the A number of species have been bred to naturally
beginning of fruit development. Continued fruit growth form parthenocarpic fruit. For example, parthenocarpy
depends on seed formation. The number of seeds within a is essential for greenhouse cucumber fruit production
fruit strongly affects fruit size in species like apple and straw- because there are no reliable insect pollinators in the
berry. Fruit that develop without seed formation (seedless) greenhouse. Other species (tomato, grape, some tree
are called parthenocarpic fruit. Two types of parthenocarpy fruits) will form parthenocarpic fruit if sprayed with
are recognized in plants (73). Vegetative parthenocarpy auxin or gibberellin. Interestingly, some species will
takes place in species like pear or fig, where the fruit devel- only form parthenocarpic fruit if treated with auxin
ops even without pollination. Stimulative parthenocarpy (tomato), while others require gibberellin (grape). The
takes place only after pollination but does not require fertil- developing seed is the normal source for auxin and gib-
ization or seed set for continued fruit growth. Grapes can berellin for fruit growth. Both gibberellin and auxin are
form seedless fruit by stimulative parthenocarpy. factors critical to fruit growth and interact during nor-
parthenocarpy The formation of fruit without seeds. mal fruit development.

embryo growth in the first few days following polli- transcription factors thought to be master regulators
nation. It appears that the suspensor is the source for of seed maturation (29). However, there is cross talk
this gibberellin and that gibberellin from the suspen- among auxin, ABA, and gibberellin via these four
sor is required for further development until the regulator genes.
embryo grows sufficiently to receive hormones and
nutrition from the endosperm.
Like auxin, gibberellins produced from the seed
Ethylene
may also signal fruit development (56). Pea pods con- Significant amounts of ethylene are produced through-
taining aborted seeds can continue development fol- out seed development as seen in Brassica species (35, 49).
lowing application of gibberellic acid. Gibberellins can Although the role of ethylene during seed development
also induce parthenocarpic fruit development in crops has not been extensively studied, it is interesting that eth-
like grapes (see Box 9). ylene production is high in developing Brassica embryos
when embryos begin to “degreen” during maturation
drying. In most seeds, embryos contain chlorophyll and
Cytokinins
are green during Stages I and II of development. There is
Several free and conjugated forms of cytokinins are a dramatic loss in chlorophyll during maturation drying
high in developing seeds. The highest concentration of while embryos “degreen” and appear yellow. Ethylene
cytokinins is found during the cell division stages of has a documented role in leaf senescence and could sup-
embryogenesis (Stage I and early Stage II). Cytokinins port embryo “de-greening.” Ethylene probably plays
appear to be supplied by the suspensor during histodif- only a minor role during seed development. Ethylene
ferentiation. The cytokinins-to-auxin ratio plays a key mutants of several species produce apparently normal
role in controlling shoot apical meristem formation, seeds.
and this association appears to be important in the dif- There is an interesting interaction between ethyl-
ferentiation phase of Stage I embryos. ene and ABA in controlling programmed cell death in
corn endosperm (76). In corn, endosperm cells die prior
Abscisic Acid (ABA) to maturation drying. There must be a mechanism in
ABA levels are high in the maturation phase of devel- place that programs these cells to die, while adjacent
oping seeds (Stage II). ABA has been shown to have a aleurone and embryo cells continue the maturation
major role in all the major features of seed matura- process. It appears that ethylene is differentially pro-
tion. ABA mutants typically show reduced storage duced in the endosperm cells, and the response to that
reserve synthesis, reduced tolerance to drying, and ethylene induces a senescence response in endosperm
premature germination prior to maturation drying. cells but not the other cell types. This differential
ABA has a major influence on all four major response is partly due to the ability of the cells to
genes (ABI3, FUS3, LEC1, and LEC2) that code respond to ABA.
141
 seed development

RIPENING AND DISSEMINATION a. species whose fruits ripen early in summer, drop to
the ground, and contain seeds that germinate
Specific physical and chemical changes that take immediately (some maples, poplar, elm)
place during maturation and ripening of the fruit b. species whose seeds mature in autumn and remain
lead to fruit senescence and dissemination of the in moist soil over winter (oak); and
seed. One of the most obvious changes is the drying c. species from warm, humid tropics (citrus). These
of the pericarp tissues. In certain species, this leads are called recalcitrant seeds (see Box 6), which
to dehiscence and the discharge of the seeds from the produce special problems in handling.
fruit. Changes may take place in the color of the
fruit and the seed coats, and softening of the fruit Seeds of species with fleshy fruits may become
may occur. dry but are enclosed with soft flesh that can decay and
Seeds of most species dehydrate at ripening and cause injury. In most species, this fleshy tissue should
prior to dissemination. Moisture content drops to be removed to prevent damage from spontaneous heat-
30 percent or less on the plant. The seed dries further ing or an inhibiting substance. In some species, how-
during harvest, usually to about 4 percent to 6 percent ever (e.g., Mahonia and Berberis), the fruits and seeds
for storage. Germination cannot take place at this level may be dried together (45).
of dryness, so it is an important basis for maintaining Many agents accomplish seed dispersal. Fish,
viability and controlling germination. birds, rodents, and bats consume and carry seeds in
In certain other species, seeds must not dry below their digestive tracts (25). This is often a function of
about 30 percent to 50 percent or they will lose their the type of fruit produced by that species (Table 3).
ability to germinate (13). These plants include

Table 3
D IFFERENT T YPES OF F RUITS
Type of fruit Description Example
Dry Fruits Indehiscent Fruits
1. Caryopsis Pericarp and seed coat are fused forming a Most often in monocots like corn
single seed. and wheat
2. Samara A one-seeded fruit with a specialized wing for Maple, ash, and elm
wind dissemination.
3. Achene A one-seeded fruit. Strawberry, sunflower, and clematis
4. Nut Fruit develops from an ovary with multiple Walnut and hazelnut
carpels, but only one survives.
5. Utricle Single-seeded fruit with inflated pericarp. Chenopodium
Dehiscent Fruits
1. Follicle Pod-like fruit from a single carpel that splits on Delphinium and columbine
one side.
2. Legume Pod that opens on both sides. Bean, locust, and pea
3. Capsule There are numerous types of dry capsules that Poppy, iris, and lily
open along different suture lines near top
of fruit.
4. Silique Develops from two carpels and opens along Cabbage and arabidopsis
two suture lines.
Fleshy Fruits 1. Berry A fleshy fruit with many seeds with an endocarp, Tomato and grape
mesocarp, and exocarp that are soft.
2. Drupe Has a hard endocarp. Peach, cherry, and fringe tree
3. Pome Has a papery endocarp. Apple and pear
4. Pepo Outer endocarp forming hard rind. Squash and pumpkin
5. Hesperidium Similar to a pepo but endocarp is not hard. Orange and lemon
6. Multiple fruits Several fruits aggregated into a single Blackberry (multiple drupes),
structure. pineapple, and mulberry
Schizocarpic Schizocarp Fruits develop so that locules in an ovary Sycamore, carrot, and parsley
Fruits separate into separate single-seeded units.

142
 seed development

Fruits with spines or hooks become attached to the fur be taken great distances and often become a source of
of animals and are often moved considerable distances. weeds in cultivated fields. Some plants (e.g., Impatiens
Wind dispersal of seed is facilitated in many plant and Oxalis) have mechanisms for short-distance dis-
groups by “wings” on dry fruits; tumbleweeds can persal, such as explosive liberation of seeds. Human
move long distances by rolling in the wind. Seeds car- activities in purposeful shipment of seed lots all over
ried by moving water, streams, or irrigation canals can the world are, of course, effective in seed dispersal.

DISCUSSION ITEMS
Knowledge of seed development is most important for 2. How does the seed storage tissue differ among a
understanding various aspects of seed quality. The envi- monocot, dicot, and gymnosperm?
ronment during seed development and the conditions 3. Compare zygotic and apomictic seed development.
during seed harvest are critical to producing quality 4. How are the stages of embryogenesis similar and
seeds. To evaluate problems related to seed quality, a different in shepherd’s purse vs. corn?
fundamental understanding of seed development, espe- 5. What might be the ecological advantages of vivip-
cially seed filling (deposition of food reserves) and seed ary as demonstrated by mangrove plants?
desiccation (maturation drying), are most important. 6. How is the scutellum of a monocot similar to
1. What are the three differences between pollination and/or different from the cotyledons in a dicot?
and fertilization?

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