24.

1 Reproduction in Flowering Plants

Flowers are reproductive organs that are composed of four different kinds of specialized leaves:
sepals, petals, stamens, and carpels. The outermost circle (whorl) of floral parts contains the sepals. In
many plants, sepals are green and closely resemble ordinary leaves. Sepals enclose the bud before it
opens, and they protect the flower while it is developing. Petals, which are often brightly colored, are
found just inside the sepals. The color, number, and shapes of such petals attract insects and other
pollinators to the flower. Within the ring of petals are the structures that produce male and female
gametophytes. The stamens are the male part of the flower. Each stamen consists of a stalk called a
filament with an anther at its tip. Anthers are the structures in which the pollen grains—the male
gametophyte—are produced. In most angiosperm species, the flowers have several stamens. The
innermost floral parts are the carpels. Carpels produce and shelter the female gametophytes and, later,
seeds. Each carpel has a broad base forming an ovary, which contains one or more ovules where female
gametophytes are produced. The diameter of the carpel narrows into a stalk called the style. At the top
of the style is a feathery or sticky portion known as the stigma, which is specialized to capture pollen. A
pistil may consist of one carpel (with its ovary, style, and stigma), or it may comprise several carpels
joined together to form a single ovary; the whole unit is called a pistil. A typical flowering plant produces
both male and female gametophytes (monoecy). In some plants, however, male and female
gametophytes are produced on different plants (dioecy).

Ovary with two carpels?

The number of locules

(chamber within ovary) is
less than or equal the
number of carpels.

In vascular plants, which include angiosperms, the sporophyte is much larger than the gametophyte. In
angiosperms, male and female gametophytes live within the tissues of the sporophyte.

As a flower develops, sporogenous (spore-producing) cells form within the anther. Each sporogenous
cell undergoes meiosis producing four haploid microspores which are contained within a wall. After the
wall breaks down due to an enzyme, The unicellular microspores undergo mitosis forming a pollen grain
(microgametophyte) surrounded by a thick wall that contains the gametes (two sperm cells).
In the ovule, usually (it differs from flower species to flower species), a single diploid cell goes through
meiosis to produce four haploid cells, three of which disintegrate. The remaining cell undergoes mitosis
three times, producing eight nuclei. These eight nuclei and the surrounding membrane are called the
embryo sac. The embryo sac, contained within the ovule, makes up the female gametophyte of a
flowering plant. Next, cell walls form around six of the eight nuclei. One of the eight nuclei, near the
base of the gametophyte, is the nucleus of the egg—the female gamete. If fertilization takes place, this
egg cell will fuse with the male gamete to become the zygote that grows into a new sporophyte plant.

Pollination is the transfer of pollen to the female portions of the flower. Some angiosperms are
wind-pollinated, but most are pollinated by animals. These animals, mainly, insects, birds, and bats,
carry pollen from one flower to another. because wind pollination is less efficient than animal
pollination, wind-pollinated plants rely on favorable weather and sheer numbers of pollen grains to get
pollen from one plant to another. Animal pollinated plants have a variety of adaptations, such as bright
colors and sweet nectar, to attract animals. Animals, such as hummingbirds, have evolved body shapes
that enable them to reach nectar deep within certain flowers. In the case of the hummingbird, a long,
thin beak allows it to reach deep within flowers. Animal pollination forms a mutualistic relationship
between the pollinating animals and the animal-pollinated plant: The animals get food, pollen and
nectar, and the plant’s pollen is scattered to other plants. Insect pollination is more efficient than wind
pollination, which may be one of the main reasons angiosperms displaced gymnosperms as the
dominant land plants.
If a pollen grain lands on the stigma of a flower of the same species, it begins to grow a pollen tube. Of
the pollen grain’s two cells, one cell—the generative cell—divides and forms two sperm cells. The other
cell—the vegetative cell—becomes the pollen tube. The pollen tube contains a tube nucleus and the
two sperm cells. The pollen tube grows into the style, where it eventually reaches the ovary and enters
an ovule. Inside the embryo sac, two distinct fertilizations take place—a process called double
fertilization. First, one of the sperm nuclei fuses with the egg nucleus to produce a diploid zygote. The
zygote will grow into the new plant embryo. Second, the other sperm nucleus fuses with two polar
nuclei in the embryo sac to form a triploid (3N) cell. This cell will grow into a food rich tissue known as
the endosperm, which nourishes the seedling as it grows. The process of fertilization in angiosperms is
distinct from that found in other plants. Two fertilization events take place—one produces the zygote
and the other a tissue, called endosperm, within the seed. Double fertilization may be another reason
why the angiosperms have been so successful. By using endosperm to store food, the flowering plant
spends very little energy on producing seeds from ovules until double fertilization has actually taken
place (no energy is wasted on an unfertilized embryo sac).
Although angiosperms are best known for their patterns of sexual reproduction, many flowering plants
can also reproduce asexually. This process, known as vegetative reproduction, enables a single plant to
produce offspring genetically identical to itself. This process takes place naturally in many plants, and
horticulturists (gardeners) also use it as a technique to produce many copies of an individual plant.
Vegetative reproduction is the formation of new individuals by mitosis. It does not require gametes,
flowers, or fertilization.

Vegetative reproduction takes place in a number of ways. For example, new plants may grow from
roots, leaves, stems, or plantlets. Plants such as potatoes, strawberries, and many cacti can reproduce
vegetatively. Because vegetative reproduction does not involve pollination or seed formation, a single
plant can reproduce quickly. In addition, asexual reproduction allows a single plant to produce
genetically identical offspring. This enables well-adapted individuals to rapidly fill a favorable
environment. One of the obvious drawbacks of asexual reproduction is that it does not produce new
combinations of genetic traits, which may be valuable if conditions in the physical environment change.

Horticulturists often take advantage of vegetative reproduction. To propagate plants with desirable
characteristics, horticulturists use cuttings or grafting to make many identical copies of a plant or to
produce offspring from seedless plants. One of the simplest ways to reproduce plants vegetatively is by
cuttings. A grower cuts from the plant a length of stem that includes a number of buds containing
meristem tissue. That stem is then partially buried in soil or in a special mixture of nutrients that
encourages root formation. Grafting is a method of propagation used to reproduce seedless plants and
varieties of woody plants that cannot be propagated from cuttings. To graft, a piece of stem or a lateral
bud (the part where the stem and leaf meet) is cut from the parent plant and attached to another plant.
Grafting works only when the two plants are closely related, such as when a bud from a lemon tree is
grafted onto an orange tree. Grafting usually works best when plants are dormant, which allows the
wounds created by the cut to heal before new growth starts.
 24.2 Fruits and Seeds
The development of the seed, which protects and nourishes the plant embryo, contributed greatly to
the success of plants on land. The angiosperm seed, encased inside a fruit, was an even better
adaptation. Once fertilization of an angiosperm is complete, nutrients flow into the flower tissue and
support the development of the growing embryo within the seed. As angiosperm seeds mature, ovary
walls thicken to form a fruit that encloses the developing seeds. A fruit is a matured angiosperm ovary,
usually containing seeds. An exception is found in commercially grown fruits that are selectively bred to
be seedless. The term fruit applies to the sweet things we usually think of as fruits, such as apples,
grapes, and strawberries. However, foods such as peas, corn, beans, rice, cucumbers, and tomatoes,
which we commonly call vegetables, are also fruits. The ovary wall surrounding a simple fruit may be
fleshy, as it is in grapes and tomatoes, or tough and dry, like the shell that surrounds peanuts.

The seeds of many plants, especially those encased in sweet, fleshy fruits, are often eaten by animals.
The seeds are covered with tough coatings, allowing them to pass through an animal’s digestive system
unharmed. The seeds then sprout in the feces eliminated from the animal. These fruits provide nutrition
for the animal and also help the plant disperse its seeds—often to areas where there is less competition
with the parent plants. Seeds contained in fleshy, nutritious fruits are usually dispersed by animals.
Animals also disperse many dry fruits, but not necessarily by eating them. Dry fruits sometimes have
burs(hooks) or hooks that catch in an animal’s fur, enabling them to be carried many miles from the
parent plant. Seeds are also adapted for dispersal by wind and water. Seeds dispersed by wind or water
are typically contained in lightweight fruits that allow them to be carried in the air or in buoyant fruits
that allow them to float on the surface of the water. A dandelion seed, for example, is attached to a
dry fruit that has a parachutelike structure. This adaptation allows the seed to glide considerable
distances away from the parent plant. Some seeds, like the coconut, are dispersed by water. Coconut
fruits are buoyant enough to float in seawater for many weeks, enabling the seeds to reach and colonize
even remote islands.

Many seeds will not grow when they first mature. Instead, these seeds enter a period of dormancy,
during which the embryo is alive but not growing. The length of dormancy varies in different species.
Germination is the resumption of growth of the plant embryo. Environmental factors such as
temperature and moisture can cause a seed to end dormancy and germinate. Before germinating,
seeds absorb water. The absorbed water causes food-storing tissues to swell, cracking open the seed
coat. Through the cracked seed coat, the young root emerges and begins to grow. The shoot—the part
of the plant that will grow above ground—emerges next.

Cotyledons are a flowering plant’s first leaves. Their job is to store nutrients and then transfer them to
the growing embryo as the seed germinates. Monocots have a single cotyledon, which usually remains
underground while it passes nutrients to the young plant. The growing monocot shoot emerges from
the soil protected by a sheath. In dicots, which have two cotyledons, there is no sheath to protect the tip
of the young plant. Instead, the upper end of the shoot bends to form a hook that forces its way through
the soil. This protects the delicate tip of the plant, which straightens as it emerges into the sunlight. In
some species, the cotyledons appear above ground as the plant emerges, while in others, such as the
garden pea, the cotyledons remain underground.
Seed dormancy can be adaptive in several ways. For one, it can allow for long-distance dispersal. And it
also allows seeds to germinate under ideal growth conditions. The seeds of most temperate (the region
between the tropics and polar regions where the temperature is mild) plants, for example, germinate in
the spring, when conditions are best for growth. For some species, a period of cold temperatures during
which the seeds are dormant is required before growth can begin. Seeds can easily survive winter cold,
but many young green plants cannot. The period of cold that is required is long enough that seeds will
not germinate until the dangerous winter season has passed allowing the plant to start germinating for
the extended period of time following winter when favorable conditions are present. Sometimes, only
extreme environmental conditions can end seed dormancy. Some pine trees, for example, produce
seeds in cones that remain sealed until the high temperatures generated by forest fires cause the cones
to open. The high temperature both activates and releases the seeds, allowing the plants to reclaim the
forest quickly after a fire.
 24.3 Plant Hormones
Hormones are chemical signals produced by living organisms that affect the growth, activity, and
development of cells and tissues. In plants, hormones may act on the same cells in which they are made,
or they may travel to different cells and tissues like animal hormones typically do. Plant hormones serve
as signals that control the development of cells, tissues, and organs. They also coordinate responses
to the environment. The two functions fit together well because plants respond to the environment
mainly by changing their development.

Cells in an organism affected by a particular hormone are called target cells. To respond to a hormone, a
cell must contain hormone receptors—usually proteins—to which hormone molecules bind. The
response that results will depend on what kinds of receptors are present in the target cell. One kind of
receptor might alter metabolism; a second might speed growth; a third might inhibit cell division. Thus,
depending on the receptors present, a given hormone may affect roots differently from stems or
flowers—and the effects may change as the developing organs add or remove receptors. Cells that do
not contain receptors are generally unaffected by hormones.

In 1880, Charles Darwin and his son Francis published the results of a series of experiments exploring
the mechanism behind a grass seedling’s tendency to bend toward light as it grows. The results of their
experiments suggested that the tip of the seedling somehow senses light. The Darwins hypothesized
that the tip produces a substance that regulates cell growth. More than forty years later, the regulatory
substances produced by the tips of growing plants were identified and named auxins. Auxins stimulate
cell elongation and the growth of new roots, among other roles that they play. They are produced in the
shoot apical meristem and transported to the rest of the plant.

One of the effects of auxins is to stimulate cell elongation. In the Darwins’

experiment, when light hits one side of the shoot, auxins collect in the shaded
part of the shoot. This change in concentration stimulates cells on the dark
side to lengthen. As a result, the shoot bends away from the shaded side and
toward the light.
Auxins also regulate cell division in meristems. As a stem grows in length, it produces lateral buds. As
you may have observed, the buds near the apex grow more slowly than those near the base of a plant.
The reason for this delay is that growth at the lateral buds is inhibited by auxins. Because auxins move
out from the apical meristem, the closer a bud is to the stem’s tip, the more it is inhibited. This
phenomenon is called apical dominance. Apical dominance occurs when the shoot (main stem or
branch) apex inhibits or controls the growth of lateral (side; auxiliary) buds. If you snip off the tip of a
plant, these lateral buds begin to grow more quickly. The plant becomes bushier. This is because the
apical meristem—the source of the growth-inhibiting auxins—has been eliminated.

Cytokinins are plant hormones that are produced in growing roots and in developing fruits and seeds.
Cytokinins stimulate cell division, they interact with auxins to help balance root and shoot growth and
stimulate the regeneration of tissues damaged by injury. Cytokinins also delay the aging of leaves and
play important roles in the early stages of plant growth. Cytokinins often produce effects opposite to
those of auxins. For example, root tips make cytokinins and send them to shoots; shoot tips make auxins
and send them to roots. This exchange of signals can restore lost organs and keep root and shoot
growth in balance. Auxins stimulate the initiation of new roots, and they inhibit the initiation and
growth of new shoot tips. Cytokinins do just the opposite. So if a tree is cut down, the stump will often
make new shoots because auxins have been removed and cytokinins accumulate near the cut.

For years, farmers in Japan knew of a disease that weakened rice plants by causing them to grow
unusually tall. The plants would flop over and fail to produce a high yield of rice grain. Farmers called the
disease the “foolish seedling’’ disease. In 1926, Japanese biologist Eiichi Kurosawa discovered that a
fungus, Gibberella fujikuroi, caused this extraordinary growth. His experiments showed that the fungus
produced a growth-promoting substance. In fact, the chemical produced by the fungus mimicked
hormones produced naturally by plants. These hormones, called gibberellins, stimulate growth and may
cause dramatic increases in size, particularly in stems and fruits.

Gibberellins also interact with another hormone, abscisic acid, to control seed dormancy. Abscisic acid
inhibits cell division, thereby halting growth. Recall that seed dormancy allows the embryo to rest until
conditions are good for growth. When seed development is complete, abscisic acid stops the seed’s
growth and shifts the embryo into a dormant state. The embryo rests until environmental events shift
the balance of hormones. Such events may include a strong spring rain that washes abscisic acid away.
(Gibberellins do not wash away as easily.) Without the opposing effect of abscisic acid, the gibberellins
can signal germination. Abscisic acid and gibberellins have opposite effects, much like auxins and
cytokinins. The opposing effects of plant hormones contribute to the balance necessary for
homeostasis.

One of the most interesting plant hormones, ethylene, is a gas. Fruit tissues release small amounts of
the hormone ethylene, stimulating fruits to ripen. Ethylene also plays a role in causing plants to seal off
and drop organs that are no longer needed. For example, petals drop after flowers have been
pollinated, leaves drop in autumn, and fruits drop after they ripen. In each case, ethylene signals cells at
the base of the structure to seal off from the rest of the plant by depositing waterproof materials in
their walls.
Plant sensors that detect environmental stimuli signal elongating organs to reorient their growth. These
growth responses are called tropisms. Plants respond to environmental stimuli such as light, gravity,
and touch. A tropism is positive if the affected plant part grows toward the stimulus. The response is
negative if the plant part grows away from the stimulus.

• The tendency of a plant to grow toward a light source is called phototropism. This response can
be so quick that young seedlings reorient themselves in a matter of hours. Recall that changes in
auxin concentration are responsible for phototropism. Experiments have shown that auxins
migrate toward shaded tissue, possibly due to changes in membrane permeability in response
to light.
• Auxins also affect gravitropism, the response of a plant to gravity. For reasons still not
understood, auxins migrate to the lower sides of horizontal roots and stems. In horizontal
stems, the migration causes the stem to bend upright. In horizontal roots, however, the
migration causes roots to bend downward (as they inhibit the growth of roots).
• Some plants even respond to touch, a process called thigmotropism. Vines and climbing plants
exhibit thigmotropism when they encounter an object and wrap around it. Other plants, such as
grape vines, have extra growths called tendrils (slender threadlike appendages, often growing in
a spiral form, that stretch out and twine around any suitable support.) that emerge near the
base of the leaf and wrap tightly around any object they encounter.
Some plant responses are so rapid that it would be a mistake to call them tropisms. Plants like Mimosa
pudica and the Venus Flytrap exhibit this rapid response. When an insect lands on a flytrap’s leaf, it
triggers sensory cells on the inside of the leaf, sending electrical signals from cell to cell. A combination
of changes in osmotic pressure and cell wall expansion causes the leaf to snap shut, trapping the insect
inside.

In the early 1920s, scientists discovered that tobacco plants flower according to their photoperiod, the
number of hours of light and darkness they receive. Additional research showed that many other plants
also respond to changing photoperiods, a response called photoperiodism. Photoperiodism is a major
factor in the timing of seasonal activities such as flowering and growth. It was later discovered that a
plant pigment called phytochrome is responsible for plant responses to photoperiod. Phytochrome
absorbs red light and activates a number of signaling pathways within plant cells. By mechanisms that
are still not understood completely, plants respond to regular changes in these pathways. These
changes determine the patterns of a variety of plant responses.

Plants such as chrysanthemums and poinsettias flower when days are short and are therefore called
short-day plants. Plants such as spinach and irises flower when days are long and are therefore known
as long-day plants.
Phytochrome also regulates the changes in activity that prepare many plants (deciduous ones: those
that shed their leaves part of the year, opposite of evergreen) for dormancy as winter approaches.
Recall that dormancy is the period during which an organism’s growth and activity decrease or stop. As
cold weather approaches, deciduous plants turn off photosynthetic pathways, transport materials
from leaves to roots, and seal off leaves from the rest of the plant.

In temperate regions, many flowering plants lose their leaves during the colder months. At summer’s
end, the phytochrome in leaves absorbs less light as days shorten and nights become longer. Auxin
production drops, but the production of ethylene increases. The change in the relative amounts of these
two hormones starts a series of events that gradually shut down the leaf. As chlorophyll breaks down,
other pigments that have been present all along—including yellow and orange carotenoids—become
visible for the first time. The brilliant reds come from freshly made anthocyanin pigments. Hormones
also produce important changes in apical meristems. Instead of continuing to produce leaves, meristems
produce thick, waxy scales that form a protective layer around new leaf buds. Enclosed in its coat of
scales, a terminal bud can survive the coldest winter days. At the onset of winter, xylem and phloem
tissues pump themselves full of ions and organic compounds. The resulting solution acts like antifreeze
in a car, preventing the tree’s sap (fluids transported in phloem and xylem) from freezing.
 24.4 Plants and Humans
Many scholars now trace the beginnings of human civilization to the cultivation of crop plants. Evidence
suggests that agriculture developed separately in many parts of the world about 10,000 to 12,000 years
ago. Once people discovered how to grow plants for food, the planting and harvesting of crops tended
to keep them in one place for much of the year, leading directly to the establishment of social
institutions. Even today, agriculture is the principal occupation of more human beings than any other
activity. Thousands of different plants—nearly all of which are angiosperms—are raised for food in
various parts of the world. Yet, despite this diversity, much of human society depends upon just a few of
these plants. Worldwide, most people depend on a few crop plants, such as rice, wheat, soybeans, and
corn, for the bulk of their food supply. The same crops are also used to feed livestock. For nutrition,
most of humanity worldwide depends on the endosperm of only a few carefully cultivated species of
grass, such as wheat, corn, and hay.

The discovery and introduction of new crop plants have frequently changed human history. The
efficiency of agriculture has been improved through the selective breeding of crop plants and
improvements in farming techniques. Recall that selective breeding is a method for improving a species
by allowing only organisms with certain traits to produce the next generation. The corn grown by Native
Americans, for example, was developed more than 8000 years ago from teosinte, a wild grass found in
Mexico. Further selective breeding has produced modern-day corn. In more recent times, other familiar
crops have been the product of selective breeding. Sugar beets, the source of most refined sugar from
the United States, were produced from the ordinary garden beet using selective breeding. Plants as
different as cabbage, broccoli, and Brussels sprouts have been developed from a single species of wild
mustard.

Between 1950 and 1970, a worldwide effort to combat hunger and malnutrition led to dramatic
improvements in farming techniques and crop yields. This effort came to be called the green revolution
because it greatly increased the world’s food supply. Green revolution technologies enabled many
countries to end chronic food shortages and, in some cases, become exporters of surplus food. At the
heart of the green revolution was the use of high-yield varieties of seed and fertilizer. For thousands of
years, farmers have added essential nutrients in the form of natural fertilizers such as animal manure.
Animal manure, however, can contaminate crops if it contains pathogens such as Salmonella,
Campylobacter, or E. coli. Many farmers use artificial fertilizers. Fertilizers are labeled with three
numbers that reflect the percentage by weight of three elements: nitrogen (N), phosphorus (P), and
potassium (K). A bag of garden fertilizer labeled “20-10-5’’ is 20 percent nitrogen, 10 percent
phosphorus, and 5 percent potassium by weight. Fertilizers and pesticides must be used with great care.
Overfertilizing can kill crop plants by putting too high a concentration of salts into the soil. The intensive
use of fertilizers can also affect the groundwater. When large amounts of nitrogen- and phosphate-
containing fertilizers are used near wetlands and streams, runoff from the fields may contaminate the
water. Pesticides can also pose a health risk. Chemical pesticides are poisons, and they have the
potential to harm wildlife and leave dangerous chemical residues in food.

Plants produce the raw materials for our homes and clothes, and some of our most powerful and
effective medicines.