Chapter 38 Angiosperm Reproduction and Biotechnology
Concept 38.1 Pollination enables gametes to come together within a flower
The life cycles of angiosperms and other plants are characterized by an alternation of
generations, in which haploid (n) and diploid (2n) generations take turns producing each
other.
The diploid plant, the sporophyte, produces haploid spores by meiosis.
These spores divide by mitosis, giving rise to multicellular male and female haploid plants
—the gametophytes.
The gametophytes produce gametes—sperm and eggs.
Fertilization results in diploid zygotes, which divide by mitosis to form new sporophytes.
In angiosperms, the sporophyte is the dominant generation, the conspicuous plant we see.
Over the course of seed plant evolution, gametophytes became reduced in size and
dependent on their sporophyte parents.
Angiosperm gametophytes are the most reduced of all plants, consisting of only a few
cells.
In angiosperms, the sporophyte produces a unique reproductive structure, the flower.
Male and female gametophytes develop within the anthers and ovules, respectively, of a
sporophyte flower.
Pollination by wind, water, or animals brings a male gametophyte (pollen grain) to a
female gametophyte contained in an ovule embedded in the ovary of a flower.
Union of gametes (fertilization) takes place within the ovary.
Ovules develop into seeds, while the ovary itself develops into the fruit around the seed.
Flower Structure
Flowers, the reproductive shoots of the angiosperm sporophyte, are typically composed of
four whorls of highly modified leaves called floral organs, which are separated by very short
internodes.
Unlike the indeterminate growth of vegetative shoots, flowers are determinate shoots in
that they cease growing once the flower and fruit are formed.
The four kinds of floral organs are the sepals, petals, stamens, and carpels.
Their site of attachment to the stem is the receptacle.
Sepals and petals are sterile.
Sepals, which enclose and protect the floral bud before it opens, are usually green and
more leaflike in appearance than the other floral organs.
In many angiosperms, the petals are brightly colored and advertise the flower to insects
and other pollinators.
Stamens and carpels are the male and female reproductive organs, respectively.
A stamen consists of a stalk (the filament) and a terminal anther containing chambers
called pollen sacs.
The pollen sacs produce pollen.
A carpel has an ovary at the base and a slender neck, the style.
At the top of the style is a sticky structure called the stigma that serves as a landing
platform for pollen.
Within the ovary are one or more ovules.
Some flowers have a single carpel.
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In others, several carpels are fused into a single structure, producing an ovary with two
or more chambers, each containing one or more ovules.
The term pistil is sometimes used to refer to a single carpel or to a groups of fused
carpels.
Depending on the species, flowers can vary in the presence or absence of sepals,
petals, stamens, or carpels.
• Complete flowers have all four basic floral organs.
• Incomplete flowers lack one or more of these organs.
• For example, most grass flowers lack petals. Some incomplete flowers are sterile,
lacking functional stamens and carpels.
Flowers also vary in size, shape, color, odor, and arrangement of floral organs.
Much of this diversity represents adaptation to specific groups of pollinators
Symmetry
Flowers can differ in symmetry. In bilateral symmetry, the flower can be divided into two
equal parts by a single imaginary line.
In radial symmetry, the sepals, petals, stamens, and carpels radiate out from a center.
Any imaginary line through the central axis divides the flower into two equal parts.
Floral organs can also be either fused or separate. For instance, daffodil petals are fused into
a funnel.
Ovary Location
The location of the ovary may vary in relation to the stamens, petals, and sepals.
An ovary is called superior if those parts are attached below it, semi-inferior if they are
attached alongside it, and inferior if they are attached above it
Floral Distribution
Floral distribution can also differ. Some species have individual flowers, while others have
clusters called inflorescences.
Reproductive Variations
Most species have a single type of flower that has functional stamens and carpels.
All complete flowers and some incomplete flowers have functional stamens and carpels.
In most incomplete flowers, stamens or carpels are either absent or nonfunctional.
Incomplete flowers that have only functional stamens are called staminate, and those with
only functional carpels are called carpellate.
If staminate and carpellate flowers are on the same plant, the species is said to be
monoecious (“one house”).
A dioecious (“two houses”) species has staminate flowers and carpellate flowers on
separate plants.
Gametophyte Development and Pollination
Anthers and ovules bear sporangia, structures where spores are produced by meiosis and
gametophytes develop.
Pollen grains, each consisting of a mature male gametophyte surrounded by a spore wall, are
formed within pollen sacs (microsporangia) of anthers.
An egg-producing female gametophyte, or embryo sac, forms within each ovule.
In angiosperms, pollination is the transfer of pollen from an anther to a stigma.
o If pollination is successful, a pollen grain produces a structure called a pollen tube, which
grows and digests its way down into the ovary via the style and discharges sperm in the
vicinity of the embryo sac, resulting in fertilization of the egg.
The zygote gives rise to an embryo, and as the embryo grows, the ovule that contains it
develops into a seed.
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The entire ovary, develops into a fruit containing one or more seeds, depending on the
species.
Fruits, which disperse by dropping to the ground or being carried by wind or animals, help
spread seeds some distance from their source plants.
When light, soil, and temperature conditions are suitable, seeds germinate and the embryo
carried in the seed grows and develops into a seedling.
Within the microsporangia (pollen sacs) of an anther are many diploid cells called
microsporocytes, also known as microspore mother cells.
Each microsporocyte undergoes meiosis, forming four haploid microspores, each of which
can eventually give rise to a haploid male gametophyte.
A microspore undergoes mitosis and cytokinesis, producing two separate cells called the
generative cell and tube cell.
Together, these two cells and the spore wall constitute a pollen grain, which at this stage of
its development is an immature male gametophyte.
The spore wall usually exhibits an elaborate pattern unique to the particular plant species.
During maturation of the male gametophyte, the generative cell passes into the tube cell. The
tube cell now has a completely free-standing cell inside it (the generative cell).
o The tube cell produces the pollen tube, a structure essential for sperm delivery to the
egg.
During elongation of the pollen tube, the generative cell usually divides and produces two
sperm cells, which remain inside the tube cell. The pollen tube grows through the long style
of the carpel and into the ovary, where it then releases the sperm cells in the vicinity of an
embryo sac.
One or more ovules, each containing a megasporangium, form within the chambers of the
ovary.
o One cell in the megasporangium of each ovule, the megasporocyte (or megaspore mother
cell), grows and then goes through meiosis, producing four haploid megaspores.
The details of the next steps vary extensively, depending on the species.
In most angiosperm species, only one megaspore survives.
This megaspore continues to grow, and its nucleus divides by mitosis three times without
cytokinesis, resulting in one large cell with eight haploid nuclei.
Membranes then partition this mass into a multicellular female gametophyte the embryo sac.
At one end of the embryo sac are three cells: the egg cell and two cells called synergids.
The synergids flank the egg cell and function in the attraction and guidance of the pollen tube
to the embryo sac.
At the opposite end of the embryo sac are three antipodal cells of unknown function.
The remaining two nuclei, called polar nuclei, are not partitioned into separate cells but
instead share the cytoplasm of the large central cell of the embryo sac.
The ovule, which will eventually become a seed, now consists of the embryo sac and two
surrounding integuments (layers of protective sporophytic tissue that eventually develop into
the seed coat).
Pollination, the transfer of pollen from anther to stigma, is the first step in a chain of events
that can lead to fertilization.
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o This step is accomplished in various ways. In some angiosperms, including grasses and
many trees, wind is a pollinating agent.
o In such plants, the release of enormous quantities of pollen compensates for the
randomness of this dispersal mechanism.
At certain times of the year, the air is loaded with pollen grains, as anyone
plagued with pollen allergies can attest.
o Some aquatic plants rely on water to disperse pollen. Most angiosperms, however,
depend on insects, birds, or other animals to transfer pollen directly to other flowers.
Mechanisms that prevent self-fertilization.
Some flowers self-fertilize or “self,” but most angiosperms have mechanisms that make this
difficult or impossible.
The various barriers that prevent self-fertilization contribute to genetic variety by ensuring
that sperm and eggs come from different parents.
Dioecious plants cannot self-fertilize because they are unisexual.
In plants with bisexual flowers, a variety of mechanisms may prevent self-fertilization.
For example, in some species stamens and carpels mature at different times.
Alternatively, they may be arranged in such a way that it is mechanically unlikely that an
animal pollinator could transfer pollen from the anthers to the stigma of the same flower.
The most common anti-selfing mechanism is self-incompatibility, the ability of a plant
to reject its own pollen and that of closely related individuals.
If a pollen grain from an anther happens to land on a stigma of a flower on the same plant,
a biochemical block prevents the pollen from completing its development and fertilizing an
egg.
The self-incompatibility systems in plant are analogous to the immune response of animals.
Both are based on the ability of organisms to distinguish “self” from “nonself.”
The key difference is that the animal immune system rejects nonself, but self-
incompatibility in plants is a rejection of self.
Recognition of “self” pollen is based on genes for self-incompatibility, called S-genes, with
dozens of different alleles in a population.
If a pollen grain and the carpel’s stigma have matching alleles at the S-locus, then the
pollen grain fails to initiate or complete the formation of a pollen tube.
Because the pollen grain is haploid, it will be recognized as “self” if its one S-allele
matches either of the two S-alleles of the diploid stigma.
Although self-incompatibility genes are all referred to as S-loci, such genes have evolved
independently in various plant families.
As a consequence, self-recognition blocks pollen tube growth by different molecular
mechanisms.
In some cases, the block occurs in the pollen grain itself, called gametophytic self-
incompatibility.
In some species, self-recognition leads to enzymatic destruction of RNA within the
rudimentary pollen tube.
RNases are present in the style of the carpel, and they can enter the pollen tube and
attack its RNA only if the pollen is of a “self” type.
In other cases, the block is a response by the cells of the carpel’s stigma, called sporophytic
self-incompatibility.
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In some species, self-recognition activates a signal transduction pathway in epidermal
cells that prevents germination of the pollen grain.
Germination may be prevented when cells of the stigma take up additional water,
preventing the stigma from hydrating the relatively dry pollen.
Basic research on self-incompatibility may lead to agricultural applications.
Many agricultural plants are self-compatible.
Plant breeders sometimes hybridize different varieties of a crop plant to combine the best
traits of the varieties and counter the loss of vigor that can result from excessive
inbreeding.
To maximize hybrid seed production, breeders currently prevent self-fertilization by
laboriously removing anthers from the parent plants that provide the seeds or by
developing male sterile plants.
Eventually, it may be possible to impose self-incompatibility on species that are normally
self-compatible.
Concept 38.2 After fertilization, ovules develop into seeds and ovaries into
fruits
Double fertilization
After landing on a receptive stigma, the pollen grain absorbs moisture and germinates,
producing a pollen tube that extends down the style toward the ovary.
The nucleus of the generative cell divides by mitosis to produce two sperm, the male
gametes.
The germinated pollen grain contains the mature male gametophyte.
Directed by a chemical attractant, possibly calcium, the tip of the pollen tube enters the
ovary, probes through the micropyle (a gap in the integuments of the ovule), and
discharges two sperm within the embryo sac.
Both sperm fuse with nuclei in the embryo sac.
One sperm fertilizes the egg to form the zygote.
The other sperm combines with the two polar nuclei to form a triploid nucleus in the
central cell.
This large cell will give rise to the endosperm, a food-storing tissue of the seed.
The union of two sperm cells with different nuclei of the embryo sac is termed double
fertilization.
Double fertilization ensures that the endosperm will develop only in ovules where the egg
has been fertilized.
This prevents angiosperms from squandering nutrients.
Normally nonreproductive tissues surrounding the embryo have prevented researchers from
visualizing fertilization in plants, but recently, scientists have been able to isolate sperm cells
and eggs and observe fertilization in vitro.
The first cellular event after gamete fusion is an increase in cytoplasmic Ca 2+ levels, which
also occurs during animal gamete fusion.
In another similarity to animals, plants establish a block to polyspermy, the fertilization of
an egg by more than one sperm cell.
In plants, this may be through deposition of cell wall material that mechanically
impedes sperm.
In maize, this barrier is established within 45 seconds after the initial sperm fusion with
the egg.
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From Ovule to Seed
After double fertilization, the ovule develops into a seed, and the ovary develops into a fruit
enclosing the seed(s).
As the embryo develops, the seed stockpiles proteins, oils, and starch.
Initially, these nutrients are stored in the endosperm.
Later in seed development in many species, the storage function is taken over by the
swelling storage leaves (cotyledons) of the embryo itself.
Endosperm Development
Endosperm development usually precedes embryo development.
After double fertilization, the triploid nucleus of the ovule’s central cell divides, forming a
multinucleate “supercell” having a milky consistency.
It becomes multicellular when cytokinesis partitions the cytoplasm between nuclei.
Cell walls form, and the endosperm becomes solid.
Coconut “milk” is an example of liquid endosperm and coconut “meat” is an example
of solid endosperm.
The endosperm is rich in nutrients, which it provides to the developing embryo.
In most monocots and some dicots, the endosperm also stores nutrients that can be used
by the seedling after germination.
In many dicots, the food reserves of the endosperm are completely exported to the
cotyledons before the seed completes its development, and consequently the mature seed
lacks endosperm.
Embryo Development
The first mitotic division of the zygote is transverse, splitting the fertilized egg into a basal
cell and a terminal cell.
The terminal cell gives rise to most of the embryo.
The basal cell continues to divide transversely, producing a thread of cells, the suspensor,
which anchors the embryo to its parent.
The suspensor functions in the transfer of nutrients to the embryo from the parent.
The terminal cell divides several times and forms a spherical proembryo attached to the
suspensor.
Cotyledons begin to form as bumps on the proembryo.
A eudicot, with its two cotyledons, is heart-shaped at this stage.
Only one cotyledon develops in monocots.
After the cotyledons appear, the embryo elongates.
Cradled between cotyledons is the embryonic shoot apex with the apical meristem of the
embryonic shoot.
At the opposite end of the embryo axis is the apex of the embryonic root, also with a
meristem.
After the seed germinates, the apical meristems at the tips of the shoot and root sustain
primary growth as long as the plant lives.
Structure of the Mature Seed
During the last stages of maturation, a seed dehydrates until its water content is only about
5–15% of its weight.
The embryo stops growing and becomes dormant until the seed germinates.
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The embryo and its food supply are enclosed by a protective seed coat formed by the
integuments of the ovule.
In the seed of a common bean, the embryo consists of an elongate structure, the embryonic
axis, attached to fleshy cotyledons.
Below the point at which the fleshy cotyledons are attached, the embryonic axis is called
the hypocotyl; above it is the epicotyl.
At the tip of the epicotyl is the plumule, consisting of the shoot tip with a pair of
miniature leaves.
The hypocotyl terminates in the radicle, or embryonic root.
While the cotyledons of the common bean supply food to the developing embryo, the seeds
of some dicots, such as castor beans, retain their food supply in the endosperm and have
cotyledons that are very thin.
The cotyledons will absorb nutrients from the endosperm and transfer them to the embryo
when the seed germinates.
The embryo of a monocot has a single cotyledon.
Members of the grass family, including maize and wheat, have a specialized cotyledon
called a scutellum.
The scutellum is very thin, with a large surface area pressed against the endosperm, from
which the scutellum absorbs nutrients during germination.
The embryo of a grass seed is enclosed by two sheathes, a coleorhiza, which covers the
young root, and a coleoptile, which covers the young shoot.
From Ovary to Fruit
As the seeds are developing from ovules, the ovary of the flower is developing into a fruit,
which protects the enclosed seeds and aids in their dispersal by wind or animals.
Fertilization triggers hormonal changes that cause the ovary to begin its transformation
into a fruit.
If a flower has not been pollinated, fruit usually does not develop, and the entire flower
withers and falls away.
The wall of the ovary becomes the pericarp, the thickened wall of the fruit, while other parts
of the flower wither and are shed.
In some angiosperms, other floral parts contribute to the fruit.
In apples, the fleshy part of the fruit is derived mainly from the swollen receptacle, while
the core of the apple fruit develops from the ovary.
Fruits are classified into several types, depending on their developmental origin.
A typical fruit is derived from a single carpel or several fused carpels and is called a
simple fruit.
Some simpler fruits are fleshy, like a peach, while others are dry, like a pea pod.
An aggregate fruit results from a single flower that has more than one carpel, each
forming a small fruit.
The fruitlets are clustered together on a single receptacle, like a raspberry.
A multiple fruit develops from an inflorescence, a group of flowers tightly clustered
together.
When the walls of the ovaries thicken, they fuse together and form one fruit, as in a
pineapple.
The fruit usually ripens about the same time as its seeds are completing their development.
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For a dry fruit such as a soybean pod, ripening is a little more than senescence of the fruit
tissues, which allows the fruit to open and release the seeds.
The ripening of fleshy fruits is more elaborate, its steps controlled by the complex
interactions of hormones.
Ripening results in an edible fruit that serves as an enticement to the animals that help
spread the seeds.
The “pulp” of the fruit becomes softer as a result of enzymes digesting components of
the cell walls.
Color changes from green to red, orange, or yellow.
The fruit becomes sweeter as organic acids or starch molecules are converted to sugar.
Seed Germination
As a seed matures, it dehydrates and enters a dormancy phase, a condition of extremely low
metabolic rate and a suspension of growth and development.
Conditions required to break dormancy and resume growth and development vary between
species.
Some seeds germinate as soon as they are in a suitable environment.
Others remain dormant until some specific environmental cue causes them to break
dormancy.
Seed Dormancy: Adaptation for Tough Times
Seed dormancy increases the chances that germination will occur at a time and place most
advantageous to the seedling.
For example, seeds of many desert plants germinate only after a substantial rainfall,
ensuring enough water to complete development.
Where natural fires are common, many seeds require intense heat to break dormancy,
allowing them to take advantage of new opportunities and open space.
Where winters are harsh, seeds may require extended exposure to cold.
Small seeds such as lettuce require light for germination and break dormancy only if they
are buried near the surface.
Other seeds require a chemical attack or physical abrasion as they pass through an
animal’s digestive tract before they can germinate.
The length of time that a dormant seed remains viable and capable of germinating varies
from a few days to decades or longer.
It depends on the species and on environmental conditions.
Most seeds are durable enough to last for a year or two until conditions are favorable for
germination.
Thus, the soil has a pool of nongerminated seeds that may have accumulated for several
years.
This is one reason vegetation reappears so rapidly after a fire, drought, flood, or some
other environmental disruption.
From Seed to Seedling
Germination of seeds depends on imbibition, the uptake of water due to the low water
potential of the dry seed.
This causes the expanding seed to rupture its seed coat and triggers metabolic changes in
the embryo that enable it to resume growth.
Enzymes begin digesting the storage materials of endosperm or cotyledons, and the
nutrients are transferred to the growing regions of the embryo.
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The first organ to emerge from the germinating seed is the radicle, the embryonic root.
Next, the shoot tip must break through the soil surface.
In garden beans and many other dicots, a hook forms in the hypocotyl, and growth pushes
it aboveground.
Stimulated by light, the hypocotyl straightens, raising the cotyledons and epicotyl.
As it rises into the air, the epicotyl spreads its first foliage leaves (true leaves).
These foliage leaves expand, become green, and begin making food by photosynthesis.
After the cotyledons have transferred all their nutrients to the developing plant, they
shrivel and fall off the seedling.
Corn and other grasses, which are monocots, use a different method for breaking ground
when they germinate.
The coleoptile, the sheath enclosing and protecting the embryonic shoot, pushes upward
through the soil and into the air.
The shoot tip then grows straight up through the tunnel provided by the tubular coleoptile.
The tough seed gives rise to a fragile seedling that will be exposed to predators, parasites,
wind, and other hazards.
Because only a small fraction of seedlings endure long enough to become parents, plants
must produce enormous numbers of seeds to compensate for low individual survival.
This provides ample genetic variation for natural selection to screen.
However, flowering and fruiting in sexual reproduction is an expensive way of plant
propagation in terms of the resources consumed.
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