Growth Responses

CHAPTER

and Regulation
of Growth
LEARNING
O BJECTIVES
Discuss genetic and

G
rapes are one of the world's most important fruit crops. They are
environmental factors that
affect plant growth and eaten fresh as table grapes, squeezed for juices and wines, dried
development. to make raisins, and processed into jams, jellies, and canned fruit.
[2 Describe phototropism, The European grape (Vitis vinifera), which has been cultivated for
gravitropism, and thousands of years, probably originated in Europe between the Black and Caspian
thigmotropism. Seas, but humans have introduced it to temperate areas worldwide. More recently,
S List several ways in which grapes have spread into subtropical areas. Today, more than 5000 different grape
each of the following hor­
varieties exist, all descended from the European grape (see photograph). These dif­
mones affects plant growth
ferent varieties have black, purple, blue, red, pink, golden, pale green, or white fruits.
and development: auxin, gib-
berellin, cytokinin, ethylene, Although more than 90 percent of cultivated grapes are varieties of the Euro­
abscisic acid. pean grape, other important species of the genus Vitis originated in North America.
3 Relate which hormone or The North American species have a more robust odor and taste, their skins separate
hormones is/are involved in from the fruit pulp easily, and their roots are resistant to low temperatures and in­
each of the following biologi­ sect and fungal pests. Frequently, disease-susceptible European vines are grafted to
cal processes: leaf abscis­
disease-resistant American rootstock to produce high-quality European grapes.
sion, seed germination,
The grape plant is a climbing, deciduous, woody vine with coiled tendrils that
apical dominance.
attach the plant to its supports. Grape flowers are small, green, fragrant, and borne
[5 Explain how varying amounts
of light and darkness induce in clusters. After self-pollination and fertilization, clusters of fruits develop that vary
flowering, and describe the in size, form, and color depending on the variety. Botanically, the fruits are berries
role of phytochrome. and usually contain seeds within a soft, juicy pulp. Thompson seedless grapes, first
[6 Explain how temperature developed in the United States, and other seedless varieties are widely consumed
affects flower induction in because they are easier to eat.
certain plants.
Growers increase the size of grapes by spraying them with a natural plant hor­
[7 Define circadian rhythm, and
mone called gibberellin during their development. Almost all Thompson seedless
give an example.
grapes grown for direct human consumption are treated with gibberellin, which
S Give an example of a turgor
lengthens the branches that bear flowers and thereby gives the fruit clusters addi­
movement, and distinguish
between turgor movements tional space in which to grow. Another advantage of gibberellin treatment of grapes
and tropisms. is that the longer stems allow air to circulate around the individual berries, keeping
them dry and preventing destructive fungi from attacking them.
In this chapter we consider the role of chemicals called hormones in regulating
all aspects of plant growth (increase in size) and development (progressive changes
during an organism's life). These processes include not only seed germination and
the growth of seedlings into mature plants but also a plant's responses to changes in
various aspects of its environment, including temperature, light, gravity, and touch.
220
F. Schussler/PhotoLink/Getty Images

A variety of pinot grape (Vitis vinifera).

222 • ch ai ter 11 Growth Responses and Regulation o f Growth

Genetic, Hormonal, and

Environmental Controls
of Growth and Development
The ultimate control of plant growth and development
is genetic. If the genes required for development of a
particular trait—for example, the shape of the leaf, the
color of the flower, or the type of root system—are not
present, that characteristic does not develop. When a
particular gene is present, its expression—that is, how it
exhibits itself as an observable feature in an organism—
is influenced by several factors, including signals from
other genes and from the environment. The location of
a cell in the young plant body also has a profound effect
on gene expression during development. chemical sig­
nals from adjacent cells help a cell “perceive” its location
within the plant body. Each cell’s spatial environment
helps determine what that cell ultimately becomes.
Growth and development, including a plant’s re­
sponses to various changes in its environment, are regu­

© Ted Morrison/Jupiterimages
lated by plant hormones, organic compounds that act
as chemical signals between cells. Environmental cues,
such as changing day length and changing tempera­
tures, exert an important influence on gene expression
and hormone production, as they do on all aspects of
plant growth and development.
The initiation of sexual reproduction is often under
environmental control, particularly in temperate lati­ FIGURE 11-1 Black-eyed Susans.
tudes, and plants switch from vegetative growth to repro­ This plant (Rudbeckia hirta) produces flowers in response to the
shortening nights of spring and early summer.
ductive growth after receiving appropriate signals from
the environment. Many flowering plants are sensitive to
changes in the relative amounts of daylight and darkness
that accompany the changing seasons, and these plants respond to such an external stimulus by directional
flower in response to those changes (•Figure 11-1). growth—that is, the direction of growth depends on the
Other plants have temperature requirements that induce direction of the stimulus. Such a directional growth re­
sexual reproduction. Plants, then, continually perceive sponse, called a tropism, results in a change in the po­
information from the environment and use this informa­ sition of a plant part. Tropisms are irreversible and are
tion to help regulate normal growth and development. positive or negative, depending on whether the plant
grows toward the stimulus (a positive tropism) or away
from it (a negative tropism). Tropisms are under hor­
Growth Movements: monal control, which is discussed in the next section.
Most growing shoot tips exhibit positive photo-
Tropisms tropism by bending (growing) toward light (•Fig­
Plants exhibit movements in response to environmental ure 11-2). This growth response increases the likelihood
stimuli such as light, gravity, and touch. A plant may that stems and leaves receive adequate light for pho­
tosynthesis. Blue light (wavelengths between 400 and
PHOTOTROPISM The directional growth of a plant caused 500 nm) triggers the bending response of phototropism.
by light. (You may recall from Chapter 8 that blue light also in­
GRAVITROPISM Plant growth in response to the direction
duces stomata to open.)
of gravity. For a plant to have a biological response to light, it
must contain a light-sensitive substance called a photo­
THIGMOTROPISM Plant growth in response to contact with
receptor to absorb the light. The photoreceptor that ab­
a solid object.
sorbs blue light and triggers the phototropic response
 Growth Movements: Tropisms • 2 2 3

and other blue-light responses (such as stomatal open­

ing) is a family of yellow pigments called phototropins.
Most stem tips exhibit negative gravitropism by
growing away from the center of Earth, whereas most
root tips exhibit positive gravitropism (•Figure 11-3).
The root cap is the site of gravity perception in roots;
when the root cap is removed, the root continues to
grow, but it loses any ability to perceive gravity.
Special cells in the root cap have starch-containing
amyloplasts that collect toward the bottoms of the cells
in response to gravity, and several lines of evidence in­
dicate that these amyloplasts initiate the gravitropic re­
sponse. If the root is reoriented, as when a potted plant
is laid on its side, the amyloplasts tumble to a new po­
sition, always settling in the direction of gravity. The
gravitropic response (bending) occurs shortly thereafter
and involves the hormone auxin (discussed in the next
section).
Despite the movement of amyloplasts in response
to gravity, their role in gravitropism remains an open
question. Mutant Arabidopsis plants with few or no am-
yloplasts in their root cap cells exhibit some gravitropic
response when placed on their sides, suggesting that
Maryann Frazier/Photo Researchers, Inc.

roots do not necessarily need amyloplasts to perceive

gravity. (A . thaliana is a model research plant that plant
biologists use to study many aspects of plant growth
and development.)
Thigmotropism is growth in response to touching
an object. An example of thigmotropism is the twining
or curling growth of tendrils or stems, which helps at­
tach a climbing plant such as a vine to some type of sup­
FIGURE 11-2 Phototropism. port. Like phototropism and gravitropism, thigmotro-
A geranium (Pelargonium) was cut back, and the plant was pism involves auxin.
placed in a window. The new growth is in the direction of light,
and therefore the plant exhibits positive phototropism.

Dennis Drenner

FIGURE 11-3 Gravitropism.

(a) A corn (Zea mays) seedling was turned on its side 3 days after germinating. In 1 hour,
the root and shoot tips had curved. (b) In 24 hours, the new root growth was downward (b)
(positive gravitropism), and the new shoot growth was upward (negative gravitropism). A
control seedling germinated at the same time is on the left.
r table 11-1 p Plant Hormones and Signaling Molecules
HORMONE SITE OF PRODUCTION PRINCIPAL ACTIONS
Auxins (e.g., IAA) Shoot apical meristem, young leaves, seeds Stem elongation, apical dominance, root initiation,
fruit development
Gibberellins (e.g., GA3) Young leaves and shoot apical meristems, Seed germination, stem elongation, flowering, fruit
embryo in seed development
Cytokinins (e.g., Zeatin) Roots Cell division, delay of leaf senescence, inhibition
of apical dominance, flower development, embryo
development, seed germination
Ethylene Stem nodes, ripening fruit, damaged or Fruit ripening, responses to environmental stress­
senescing tissue ors, seed germination, maintenance of apical hook
on seedlings, root initiation, senescence and ab­
scission in leaves and flowers, apical dominance
Abscisic acid Almost all cells that contain plastids Seed dormancy, responses to water stress
(leaves, stems, roots)
Brassinosteroids Shoots (leaves and flower buds), seeds, Cell division, cell elongation, light-mediated differ­
(e.g., Brassinolide) fruits entiation, seed germination, vascular development
Oligosaccharins Unknown May function in normal cell growth and develop­
ment, defense responses to disease organisms,
inhibition of flowering
Jasmonates Leaves? Probably many tissues Initiation of defenses against insect predators
(e.g., Jasmonic acid) or disease organisms, pollen development, root
growth, fruit ripening, senescence
Salicylic acid Wound (site of infection) Resistance to insect predators and disease
organisms
Systemin Wound (site of herbivore or pathogen Initiation of defenses against predators (herbivores)
attack) or disease organisms

Source: From Solomon, Berg, Martin Biology 8/e, Table 37-1, p. 791.

PROCESS OF SCIENCE

QUESTION: Which part of the grass coleoptile absorbs the directional light that triggers
phototropic growth?
HYPOTHESIS: The coleoptile tip perceives the directional light.
EXPERIMENT: Some canary grass coleoptiles were uncovered, some were covered only at the
tip, some had the tip removed, and some were covered everywhere but at the tip. The cov­
ers were impervious to light.

RESULTS AND CONCLUSION: After exposure to light coming from one direction, the uncovered
plants (a) and the plants with uncovered tips (d) grew toward the light. The plants with tips
covered (b) or removed (c) did not bend toward light. The Darwins concluded that some sub­
stance was produced in the tip and transmitted to the lower part that caused it to bend. ■

FIGURE 11-4 The Darwins' phototropism experiments.

 Plant Hormones • 2 2 5

Plant Hormones PROCESS OF SCIENCE

A plant hormone is an organic chemical compound that
acts as a chemical signal, eliciting a variety of responses QUESTION: Is there a chemical substance responsible for
elongation of coleoptiles, and can it be isolated?
that regulate growth and development. The study of
plant hormones and their effects is challenging because HYPOTHESIS: The factor responsible for coleoptile growth
hormones are effective in extremely small amounts. In is a diffusible chemical that can be isolated from coleoptile
tips.
addition, the effects of different plant hormones over­
lap, and it is difficult to determine which hormone, if EXPERIMENT: Coleoptile tips were placed on agar blocks
any, is the primary cause of a particular response. Plant for a period of time (a). The agar block was transferred to
a decapitated coleoptile. It was placed off center, and the
hormones may also stimulate a response at one con­
coleoptile was left in darkness (b).
centration and inhibit that same response at a different
concentration .
For many years, biologists studied five major classes
of plant hormones: auxin, gibberellin, cytokinin, ethyl­
ene, and abscisic acid. More recently, researchers have
uncovered compelling evidence for a variety of other
signaling molecules, such as brassinosteroids, oligo-
saccharins, jasmonates, salicylic acid, and systemin.
(•Table 11-1 summarizes plant hormones and hormone­
like signaling molecules discussed in this chapter.)

Auxins promote cell elongation

PROCESS OF SCIENCE Charles Darwin, the British naturalist
best known for developing the theory of natural selec­
tion to explain evolution, also provided the first evi­
dence for the existence of plant hormones. The experi­
RESULTS AND CONCLUSION: The coleoptile bent (see c),
ments that Darwin and his son Francis performed in the indicating that a chemical moved from the original coleop­
1870s involved phototropism in newly germinated ca­ tile tip to the agar block and from there to one side of the
nary grass seedlings. As in all grasses, the first part of a decapitated coleoptile. As a result, that side elongated
canary grass seedling to emerge from the soil is the cole- more than the side without the agar block. ■
optile, a protective sheath that encircles the stem. When V__________________________________________________ )
coleoptiles are exposed to light from only one direction, FIGURE 11-5 Isolating auxin from coleoptiles.
they bend toward the light. The bending occurs close to
the tip of the coleoptile.
The Darwins tried to influence this bending in optile, normal growth resumed. When he placed one of
several ways (•Figure 11-4). For example, they covered these agar blocks to one side of the tip of a decapitated
the tip of the coleoptile as soon as it emerged from the coleoptile in the dark, bending occurred (•Figure 11-5).
soil. When they covered that part of the coleoptile above This bending indicated that the substance had diffused
where the bend would normally occur, the plants did from the coleoptile tip into the agar and, later, from the
not bend. On other plants, they removed the coleoptile agar into the decapitated coleoptile. Went named this
tip and found that bending did not occur. When the bot­ substance auxin (from the Greek word for “enlarge” or
tom of the coleoptile where the curvature would occur “increase”). ■
was shielded from the light, the coleoptile bent toward
light. From these experiments, the Darwins concluded
that “some influence is transmitted from the upper to
HORMONE An organic chemical messenger that regulates
the lower part, causing it to bend.”
growth and development in plants and other multicellular
In the 1920s Frits Went, a young Dutch plant phys­ organisms.
iologist, isolated this “influence” from oat coleoptiles.
To do so, he removed the coleoptile tips and placed them AUXIN A plant hormone involved in growth and develop­
ment, including stem elongation, apical dominance, and root
on tiny blocks of agar for a period of time. When he put
formation on cuttings.
one of these agar blocks squarely on a decapitated cole-
226 • ch ai ter 11 Growth Responses and Regulation o f Growth

Several naturally occurring auxins have been iden­ Shaded side

of coleoptile
tified, and some artificial auxins have been synthesized.
The most common naturally occurring auxin is indole-
acetic acid (IAA). The movement of auxin in the plant is
polar, or unidirectional. Auxin moves downward along
the shoot-root axis from its site of production, usually
the shoot apical meristem. Young leaves and seeds are
also sites of auxin production.
Auxin’s most characteristic action is promotion of
cell elongation in stems and coleoptiles. This effect, ap­
parently exerted by acidifying the cell walls, increases Light rays
their plasticity, enabling them to expand under the force
of the cell’s internal turgor pressure. (Turgor pressure is Illuminated
the pressure that develops against a cell wall when water side of
coleoptile
moves into the cell by osmosis.) Auxin’s effect on cell
elongation also provides an explanation for phototro-
pism. When a plant is exposed to a light from only one
direction, the auxin moves laterally to the shaded side of
the stem before moving down the stem by polar trans­
port. Because of the greater auxin concentration on the
shaded side of the stem, the cells there elongate more than
the cells on the lighted side, and the stem bends toward
the light (•Figure 11-6). As mentioned earlier, auxin is
also involved in gravitropism and thigmotropism. FIGURE 11-6 Unequal distribution of auxin causes phototropism.
Auxin exerts other effects on plants. For example, Auxin travels down the side of the stem or coleoptile away from
the light (black arrow), causing cells on the shaded side to elon­
some plants branch out very little when they grow.
gate. Therefore, the stem or coleoptile bends toward light.
Growth in these plants occurs almost exclusively from
the shoot apical meristem rather than from axillary
buds, which do not develop as long as the terminal bud
is present. Such plants are said to exhibit apical domi- used to stimulate root development on stem cuttings for
nance—the inhibition of axillary bud growth by the asexual propagation, particularly of woody plants with
shoot apical meristem (•Figure 11-7). In plants with horticultural importance (‘ Figure 11-8). The synthetic
strong apical dominance, auxin produced in the apical auxins 2,4-D and 2,4,5-T are used as selective herbicides
meristem inhibits axillary buds near the apical meristem (weed killers). These compounds kill plants with broad
from developing into actively growing shoots. When the leaves but, for reasons not completely understood, do
apical meristem is pinched off, the auxin source is re­ not kill grasses. Although both herbicides are similar
moved, and axillary buds grow to form branches. Api­ in structure to IAA, they disrupt the plants’ normal
cal dominance is often quickly reestablished, however, growth processes. Because many of the world’s most
as one branch begins to inhibit the growth of others. important crops are grasses (for example, wheat, corn,
Other hormones (ethylene and cytokinin, both dis­ and rice), both 2,4-D and 2,4,5-T can be used to kill
cussed later) are also involved in apical dominance. As broadleaf weeds that compete with these crops. The use
with many physiological processes, the interactions of of 2,4,5-T is no longer allowed in the United States be­
these hormones, rather than the action of one hormone cause of its association with dioxins, a group of mildly
alone, may be responsible for apical dominance. to very toxic compounds formed as by-products during
Auxin produced by developing seeds stimulates the the manufacture of 2,4,5-T.
development of the fruit. When auxin is applied to cer­
tain flowers in which fertilization has not occurred (and,
therefore, in which seeds are not developing), the ovary 1Not all seedless fruits are produced by treatment with auxin.
enlarges and develops into a seedless fruit. Seedless to­ As discussed in the chapter introduction, Thompson seedless
matoes are produced in this manner.1 Auxin is not the grapes result from treatment of the flowers and young fruits with
only hormone involved in fruit development, however. gibberellins, another group of hormones. In this case, fertiliza­
Some manufactured, or synthetic, auxins have tion occurs, but the embryos abort, and therefore the seeds fail
structures similar to that of IAA. A synthetic auxin is to develop.
 Plant Hormones • 2 2 7

(a) (b)
FIGURE 11-8 Honeysuckle cuttings treated with various concen­
trations of a synthetic auxin.
FIGURE 11-7 Auxin inhibits the development of axillary buds.
Many adventitious roots formed on honeysuckle (Lonicera
(a) When the tip of the plant (the source of auxin) is intact,
fragrantissima) cuttings placed in a higher auxin concentration
the axillary buds do not develop. (b) The tip of the plant was
(left), whereas fewer roots formed in a lower auxin concentration
removed. Because auxin is not moving down from the stem tip,
(middle). Cuttings placed in water (right) served as a control and
axillary buds develop into branches.
did not form roots in the same period.

One of the many controversial aspects of the Viet­ work, and the hormone was found to be one of a class
nam War was the defoliation program carried out by of compounds collectively called gibberellins. During
the United States in South Vietnam. From 1961 to 1971, the 1950s and 1960s, studies in the United States and
the United States sprayed Agent Orange, a mixture Great Britain showed that healthy plants also produce
of the herbicides 2,4-D and 2,4,5-T, over large areas of gibberellins. ■
South Vietnam to expose suspected hiding areas and Gibberellins are hormones involved in many nor­
destroy crops planted by the Vietcong and North Viet­ mal plant functions. The symptoms of the “foolish
namese troops. The negative impact of these herbicides seedling” disease are caused by an abnormally high gib-
on the environment is still being felt today. Many eco­ berellin concentration in the plant tissue (because both
logically important mangrove forests and commercially the plant and the fungus are producing gibberellin).
valuable hardwood forests were destroyed. In addition, Currently, dozens of naturally occurring gibberellins
the herbicide sprays caused health problems in the na­ are known, although many are probably inactive forms;
tive people and members of the U.S. military who were there are no synthetic gibberellins.
exposed to them in the Vietnamese jungles. The herbi­ Gibberellins promote stem elongation in many
cides contained traces of dioxin, which is now known plants. When gibberellin is applied to a plant, particu­
to cause birth defects in animals. Reportedly, the num­ larly certain dwarf varieties, this elongation may be
ber of birth defects and stillbirths in Vietnam increased spectacular. Some corn and pea plants that are dwarfs
during the herbicide spraying. Also, American veterans as a result of one or more mutations (changes in their
who were exposed to high levels of herbicides have more genetic material) grow to a normal height when treated
health problems than do other veterans, including cer­ with gibberellin (•Figure 11-9a). Short-stemmed, high-
tain types of cancer and skin diseases. yielding varieties of wheat have short stems because
they have a reduced response to gibberellin. Gibberel-
lins are also involved in bolting, the rapid elongation of
Gibberellins promote stem elongation a floral stalk that occurs naturally in many plants when
they initiate flowering (‘ Figure 11-9b).
PROCESSOFSCIENCE In the 1920s, a Japanese biologist was
studying a disease of rice in which the young rice seed­
lings grow extremely tall and spindly, fall over, and die.
The cause of the disease, dubbed the “foolish seedling”
disease, was a fungus that produces the chemical sub­ GIBBERELLIN A plant hormone involved in growth and
development, including stem elongation, flowering, and seed
stance gibberellin. Not until after World War II did
germination.
scientists in Europe and North America learn of this
228 • chap ter 11 Growth Responses and Regulation o f Growth

<C

1d w a rf - * I I dw a rf - 1 if. 1
1 *u

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(a) Effect of the continued application of (b) Bolting in Indian blanket (Gaillardia pulchella). Many
gibberellin on normal and dwarf corn (Zea biennials grow as a rosette (a circular cluster of leaves close
mays) plants. From left to right: dwarf, to the ground) during their first year (left) and then bolt when
untreated; dwarf, treated with gibberellin; they initiate flowering in their second year (right). Gibberellin
normal, treated with gibberellin; normal, triggers the rapid stem elongation.
untreated. This dwarf variety is a mutant
with a single recessive gene that impairs
gibberellin metabolism.

FIGURE 11-9 Effects of gibberellin.

Gibberellins cause stem elongation by stimulating tion, researchers showed that the release of gibberellin
cells to divide as well as elongate. The actual mechanism from the embryo triggers the synthesis of an enzyme
of cell elongation differs from that caused by auxin, that digests starch in the endosperm. As a result, glu­
however. Recall that IAA-induced cell elongation in­ cose becomes available for absorption by the embryo as
volves acidifying the cell wall. In gibberellin-induced it resumes growth. In addition to mobilizing food re­
cell elongation, cell-wall acidification does not occur; serves in certain newly germinated seeds, application
instead, gibberellin increases the cell wall’s ability to of gibberellins substitutes for low-temperature or light
expand. requirements for germination in seeds such as lettuce,
Gibberellins affect several reproductive processes in oats, and tobacco.
plants. They stimulate flowering, particularly in long-
day plants (discussed later in the chapter). In addition,
gibberellins can substitute for the low temperature that Cytokinins promote cell division
biennials require before they begin flowering. If gibber- PROCESSOFSCIENCE During the 1940s and 1950s, research­
ellins are applied to biennials during their first year of ers were trying to find substances that might induce
growth, flowering occurs without exposure to a period plant cells to divide in tissue culture, a technique in
of low temperature. Gibberellins, like auxin, affect fruit which cells are isolated from plants and grown in a nu­
development. As mentioned in the chapter introduction, trient medium. They discovered that cells divided when
agriculturists apply gibberellins to several varieties of placed in a culture containing coconut milk. Because
grapes to produce larger fruits. coconut milk has a complex chemical composition, in­
Gibberellins are also involved in the seed germi­ vestigators did not chemically identify the division-
nation of certain plants (cereals and other grasses). In inducing substance for some time. Finally, researchers
a classic experiment involving barley seed germina­ isolated the active substance from a different source.
 Plant Hormones • 2 2 9

They called it cytokinin because it induces cell division, Senescence is accelerated in cells of plant parts that are
or cytokinesis. In 1963, researchers identified the first cut, such as flower stems. Botanists think plants have to
plant cytokinin (in corn), and since then similar mol­ have a continual supply of cytokinins from the roots.
ecules have been identified in other plants. Biologists Cut stems, of course, lose their source of cytokinins and
have also synthesized several cytokinins. ■ therefore age rapidly.
Cytokinins promote cell division and differen­ Despite their involvement in many aspects of plant
tiation of young, relatively unspecialized cells into ma­ growth and development, cytokinins have not been ap­
ture, more specialized cells in intact plants. They are a proved for many commercial applications other than
required ingredient in any plant tissue culture medium plant tissue culture. However, in the mid-1990s, mo­
because they must be present in all dividing plant cells. lecular biologists used genetic engineering to develop
In tissue culture, cytokinins interact with auxin during tobacco plants that produce more cytokinin and there­
the formation of plant organs such as roots and stems. fore live longer (‘ Figure 11-11). This discovery has the
For example, in tobacco tissue culture, a high ratio of potential to increase the longevity and productivity of
cytokinin to auxin induces shoots to form, whereas a certain crops.
low ratio of cytokinin to auxin induces roots to form
(•Figure 11-10).
Cytokinins and auxin also interact in the control
of apical dominance, in which the shoot apical meristem
suppresses the growth of axillary buds. Here their re­
lationship is antagonistic: auxin inhibits the growth of
axillary buds, and cytokinin promotes their growth.
One effect of cytokinins on plant cells is to delay
the aging process. Plant cells, like all living cells, go
through a natural aging process known as senescence.

Cell division Cell division

without differentiation with differentiation

(a) Initial (b) Callus (c) Roots (d) Shoots

explant

FIGURE 11-10 Hormones and organ formation when propagating

tobacco by tissue culture.
(a) A fragment of tissue explant from the center of a tobacco
stem is placed in a culture medium. A complete plant can form
from the tissue fragment because each cell of the fragment
contains all the genetic information for the entire organism. FIGURE 11-11 Cytokinin synthesis and delay of senescence.
Varying amounts of auxin and cytokinin in the culture media The tobacco (Nicotiana tabacum) on the left was genetically
produce different growth responses. (b) Nutrient agar containing engineered to produce additional cytokinin as it aged, whereas
a moderate amount of both auxin and cytokinin caused cells to the unaltered tobacco plant of the same age on the right served
divide and form a callus (a clump of disorganized, undifferenti­ as a control. Note the extensive senescence and death of older
ated cells). (c) When callus is transplanted to a medium with a leaves on the control plant.
higher relative amount of auxin, roots differentiate. (d) Shoot
growth is stimulated by a medium with a higher relative amount CYTOKININ A plant hormone involved in growth and devel­
of cytokinin. Plants grown using tissue culture techniques can opment, including cell division and delay of senescence.
be transferred to soil and grown normally.
230 • chap ter 11 Growth Responses and Regulation o f Growth

Ethylene promotes abscission

and fruit ripening
During the early 20th century, scientists observed
that the gas ethylene has several effects on plant
growth, but not until 1934 did they demonstrate
that plants produce ethylene. This natural hor­
mone influences many diverse plant processes.
Ethylene inhibits cell elongation, promotes seed
germination, promotes apical dominance, and is
involved in plant responses to wounding or inva­
sion by disease-causing microorganisms.

Illustration Services
Ethylene also has a major role in many aspects
of development, including fruit ripening. As a fruit
ripens, it produces ethylene, which triggers an ac­
celeration of the ripening process. This induces the
fruit to produce more ethylene, which further ac­
celerates ripening. The expression “one rotten ap­ FIGURE 11-12 Ethylene and fruit ripening.
ple spoils the lot” is true. A rotten apple is one that Both boxes of tomatoes were picked at the same time, while green.
The tomatoes in the box on the right were exposed to an atmosphere
is overripe and produces large amounts of ethylene,
containing ethylene for 3 days, whereas the tomatoes on the left
which diffuses and triggers the ripening process in were not.
nearby apples. Humans use ethylene commercially
to uniformly ripen bananas and tomatoes. These fruits engineer crops and horticultural plants that are more re­
are picked while green and shipped to their destination, sistant to drought.
where they are exposed to ethylene before they are de­ The low temperatures of winter are also a type of
livered to grocery stores (•Figure 11-12). stress on plants. A winter adaptation that involves ab­
Ethylene, along with auxin, is involved in leaf se­ scisic acid is dormancy in seeds. Many seeds have high
nescence and abscission. As a leaf ages (as autumn ap­ levels of abscisic acid in their tissues and do not ger­
proaches, for deciduous trees in temperate climates), the minate until spring, after winter snows and rain have
level of auxin in the leaf decreases. concurrently, cells washed out the abscisic acid. In a corn mutant unable to
in the abscission layer at the base of the petiole (where synthesize abscisic acid, the seeds germinate as soon as
the leaf will break away from the stem) begin producing the embryos are mature, even while attached to the ear
ethylene, which in turn stimulates weakening of the cell (•Figure 11-13).
walls in this location. Abscisic acid is not the only hormone involved in
seed dormancy. For example, addition of gibberellin
Abscisic acid promotes seed dormancy reverses the effects of dormancy. In seeds, the level of
abscisic acid decreases during the winter, and the level
Abscisic acid was discovered simultaneously in 1963
of gibberellin increases. Cytokinins are also implicated
by two independent research teams. Despite its name,
in breaking dormancy. Once again you see that a single
abscisic acid does not induce abscission in most plants.
activity such as seed dormancy may be controlled by
Instead, abscisic acid is involved in a plant’s response
the interaction of several hormones. The plant’s actual
to stress and in seed dormancy, a temporary state
response may result from changing ratios of hormones
of arrested physiological activity when growth does
rather than the effect of each individual hormone.
not occur even when environmental conditions are
favorable.
As an environmental stress hormone, abscisic acid
Additional signaling molecules
particularly promotes changes in plant tissues that affect growth and development
are water stressed. The level of abscisic acid increases Biologists continue to discover new plant hormones and
dramatically in the leaves of plants exposed to severe hormone-like signaling molecules. Many of these sig­
drought conditions. The high level of abscisic acid in the naling molecules are involved in defensive responses of
leaves triggers the closing of stomata, which saves the plants to disease organisms and insects. Here we briefly
water that the plant would normally lose by transpira­ consider five groups—brassinosteroids, oligosaccharins,
tion, thereby increasing the plant’s likelihood of sur­ jasmonates, salicylic acid, and systemin.
vival. As knowledge of cell signals relating to abscisic Although steroid hormones have crucial roles in
acid increases, botanists hope to use this information to animals, we are just beginning to understand their roles
 Plant Hormones • 2 3 1

in plants. The brassinosteroids (BRs) are a group of ste­ tical value in controlling certain insect pests without the
roids that function as plant hormones. BRs are involved use of chemical pesticides.
in several aspects of growth and development. Arabidop- For centuries people chewed willow (Salix) bark to
sis mutants that cannot synthesize BRs are dwarf plants treat headaches and other types of pain. Salicylic acid
with reduced fertility. Researchers reverse this defect by was first extracted in the 19th century from willow bark
applying BR. Studies of these mutants suggest that the and is chemically related to aspirin (acetylsalicylic acid).
brassinosteroids are involved in multiple processes such Recently, biologists have shown that salicylic acid is a
as cell division, cell elongation, light-induced differen­ signaling molecule that helps plants defend against in­
tiation, seed germination, and vascular development. sect pests and disease-causing agents such as viruses.
Oligosaccharins are carbohydrate fragments that Although many animal hormones are polypep­
consist of short, branched chains of sugar molecules. tides, the first plant polypeptide with hormonal prop­
They are present in extremely small quantities in cells erties, called systemin, was not isolated until 1991. In
and active at much lower concentrations (100 to 1000 response to wounding by insects, systemin may trigger
times as low) than hormones such as auxin. Some oligo- the plant to produce molecules that disrupt insect diges­
saccharins trigger the production of phytoalexins (from tion, thereby curbing leaf damage done by caterpillars
the Greek phyto, “plant,” and alexi, “to ward off”), an­ and other herbivorous (plant-eating) insects. The dis­
timicrobial compounds that limit the spread of plant covery of systemin in tomato leaves prompted a flurry
disease organisms such as fungi. Other oligosaccharins of research that resulted in the discovery of additional
inhibit flowering and induce vegetative growth (that is, polypeptides in other plants.
the growth of leaves, stems, and roots).
Jasmonates are lipid-derived hormones that affect
several plant processes, such as pollen development, root PROCESS Progress is beingmade
growth, fruit ripening, and senescence. They are also inidentifyingthe elusive
OFSCIENCE

produced in response to the presence of insect pests and flower-promotingsignal

disease-causing organisms. Jasmonates may be of prac- Experiments in which different tobacco species are
grafted together indicate that additional flower-
promoting and flower-inhibiting substances may exist.
N icotiana silvestris is a long-day tobacco plant (it flow­
ers when exposed to short nights; discussed later in the
chapter). A variety of N . tabacum is a day-neutral to­
bacco plant (day length does not regulate its flowering;
also discussed later in the chapter). When a long-day to­
bacco is grafted to a day-neutral tobacco and exposed
to short nights, both plants flower (•Figure 11-14). The
day-neutral tobacco plant flowers sooner than it nor­
mally would.
For many years biologists have found evidence that
a flower-promoting substance, florigen, may be induced
in the long-day plant and transported to the day-neutral
plant through the graft union, causing the day-neutral
tobacco plant to flower sooner than expected. An intact
plant may produce florigen in the leaves and transport it
in the phloem to the shoot apical meristem. There, it in­
duces a transition from vegetative to reproductive devel-
opment—that is, to a meristem that produces flowers.
When a botanist grafts a long-day tobacco to a day-
neutral tobacco and exposes them to long nights, nei­
ther plant flowers. As long as these conditions continue,

ETHYLENE A gaseous plant hormone involved in growth and

FIGURE 11-13 Abscisic acid and seed germination. development, including leaf abscission and fruit ripening.
In a corn (Zea mays) mutant that does not produce abscisic ABSCISIC ACID A plant hormone involved in growth and
acid, some of the kernels have germinated while still on the ear, development, including dormancy and responses to stress.
producing roots (arrows).
232 • ch ai ter 11 Growth Responses and Regulation o f Growth

the day-neutral plants do not flower even

when they would normally do so. In this case,
the long-day tobacco may produce a flower-
inhibiting substance that is transported to the
day-neutral tobacco through the graft union.
This substance prevents the day-neutral to­
bacco from flowering.
Although biologists have not yet isolated
and chemically characterized all flower pro­
moters and flower inhibitors, plant biologists
working with Arabidopsis reported the identi­
fication of the elusive florigen signal in 2005.
A gene is activated (turned on) in Arabidopsis
leaves, and a messenger RNA (mRNA) mole­
cule is transcribed from this gene. This mRNA
molecule, thought to be florigen, travels in the
phloem from the leaf to the shoot apical meri-
stem. There, it is translated into a protein that
alters gene expression (turning genes on or
off) in cells at the meristem, thereby initiating FIGURE 11-14 Evidence of a flower-promoting substance.
When a long-day tobacco (Nicotiana silvestris) is grafted to a day-neutral
flower development. ■
tobacco (N. tabacum) and both plants are exposed to a long-day, short-
night regimen, they both flower. The day-neutral plant flowers sooner than it
Plant hormones act normally would, presumably because a flower-promoting substance passes
from the long-day plant to the day-neutral one through the graft. In 2005,
by signal transduction after years of searching, biologists identified florigen as a messenger RNA
Researchers have used Arabidopsis mutants to molecule.
better understand the biology of plant hor­
mones. For example, some mutants have defects in sig­ a plant to the relative lengths of daylight and darkness.
nal transduction. In signal transduction, a receptor in Initiation of flowering at the shoot apical meristem is
a cell’s plasma membrane converts the hormone, which one of several processes that are photoperiodic in many
is outside the cell, into a signal that causes some change plants. ilants are classified into four main groups on
inside the cell; that change ultimately leads to a physi­ the basis of how photoperiodism affects their transition
ological response. Many plant hormones bind to recep­ from vegetative growth to flowering: short-day, long-
tors located in the plasma membrane, triggering enzy­ day, intermediate-day, and day-neutral plants.
matic reactions of some sort. Short-day plants (also called long-night plants)
Let us consider a specific example, involving the flower when the night length is equal to or greater than
plant hormone auxin (•Figure 11-15). As shown in step some critical period (•Figure 11-16). The initiation of
o , the receptor for auxin is located in the plasma mem­ flowering in short-day plants is due to the long, uninter­
brane and has a shape that binds to the auxin molecule. rupted period of darkness rather than the short period
The binding of auxin to its receptor catalyzes the attach­ of daylight (•Figure 11-17 F ). The minimum critical
ment of a molecule, called ubiquitin, to proteins known night length varies considerably from one plant species
to inhibit certain genes (see step U). Whenever ubiquitin to another but falls between 12 and 14 hours for many.
is attached to protein molecules, the cell destroys those Examples of short-day plants are florist’s chrysanthe­
molecules (5 ). As a result, the genes that were inhibited mum, cocklebur, and poinsettia, all of which typically
by those proteins are now turned on O ), resulting in flower in late summer or fall. ioinsettias, for example,
changes in cell growth and development. initiate flower buds in early October in the Northern
Hemisphere and flower about 8 to 10 weeks later, hence
their traditional association with Christmas. Short-day
plants detect the lengthening nights of late summer or
Photoperiodism fall, and they flower at that time.
In the early 20th century, Wightman Garner and Henry Long-day plants (also called short-night plants)
Allard, two American plant physiologists working at flower when the night length is equal to or less than
an agricultural research center in Maryland, discovered some critical length (•Figure 11-17 y ). Plants such as
that day length (or, more correctly, night length) was in­ spinach, black-eyed Susan (see Figure 11-1), and Ara-
volved in flowering. Photoperiodism is any response of bidopsis flower in late spring or summer and are long-
 Photoperiodism • 2 3 3

Many plant hormones activate genes through signal transduction.

V___________________________________________________________________________________________________________ J
FIGURE 11-15 How auxin works.
The numbered steps are explained in the text.

day plants. These plants detect the shortening nights of when they are exposed to days and nights of intermedi­
spring and early summer, and they flower at that time. ate length.
Intermediate-day plants do not flower when day Some plants, called day-neutral plants, do not ini­
length is either too long or too short. Sugarcane and tiate flowering in response to seasonal changes in the
coleus are intermediate-day plants. These plants flower period of daylight and darkness but instead respond to
some other type of stimulus, external or internal. To­
mato, dandelion, string bean, and pansy are examples of
day-neutral plants. Many of these plants originated in
the tropics, where day length does not vary appreciably
during the year. (In contrast, short-day, long-day, and
intermediate-day plants are temperate species.)

Phytochrome detects day length

The main photoreceptor for photoperiodism and many
other light-initiated plant responses is phytochrome, a
family of about five blue-green pigment proteins. A mix­
Dennis Drenner

ture of phytochrome proteins is present in cells of all

vascular plants examined so far.
Much current knowledge of phytochrome is based
on various mutant Arabidopsis plants that do not pro-
FIGURE 11-16 Short-day plants.
The chrysanthemum (Chrysanthemum) is a short-day plant and
therefore flowers only when exposed to long nights. Two cuttings PHOTOPERIODISM The physiological response (such as
from the same plant were planted in separate pots. The plant flowering) of plants to variations in the length of daylight and
on the left, which flowered, received short days (8 hours of day­ darkness.
light) and long nights (16 hours of darkness) for several weeks. PHYTOCHROME A blue-green proteinaceous pigment
The plant on the right, which remained vegetative, received long involved in many plant responses to light, independent
days (16 hours of daylight) and short nights (8 hours of dark­ of photosynthesis.
ness) during the same period.
234 • chap ter 11 Growth Responses and Regulation o f Growth

PROCESS OFSCIENCE length (660 nm). In the process, the

shape of the molecule changes to the
second form of phytochrome, Pfr,
QUESTION: Is day length or night length the critical factor affecting flowering?
so designated because it absorbs far-
HYPOTHESIS: The plant measures day length, not night length: Short-day plants red light, which is red light with a
flower when the day length is short, and long-day plants flower when the day length
relatively long wavelength (730 nm)
is long.
(•Figure 11-18).
EXPERIMENT: Short-day plants (chrysanthemum) and long-day plants (black-eyed
far-red light, the Pfr reverts to the
Susan) were exposed to different light-dark regimes.
original form, Pr. Pfr, the active
Day Night form of phytochrome, triggers or in­
hibits responses such as flowering.
What does a pigment that ab­
sorbs red light and far-red light
(a) Short days (b) Long days (c) Short days (d) Long days have to do with daylight and dark­
and long nights and short nights and long nights (interrupted with ness? Sunlight consists of various
(interrupted with a brief period of
amounts of the entire spectrum of
a brief period of dark) and short
visible light, in addition to ultravio­
light) nights
let and infrared radiation. Because
sunlight contains more red light
than far-red light, however, when
a plant is exposed to sunlight, the
level of Pfr increases.
Why is phytochrome important
to plants? Timing of day length and
Chrysanthemum (short-day/long-night plant)
darkness is the most reliable way
for plants to measure the change
from one season to the next. This
measurement, which synchronizes
the stages of plant development, is
crucial for survival, particularly in
environments where the climate has
an annual pattern of favorable and
Black-eyed Susan (long-day/short-night plant) unfavorable seasons.

RESULTS AND CONCLUSION: A short-day plant flowers when it is grown under long-
night conditions (a), but it does not flower when exposed to either (c) long nights Phytochrome is involved
interrupted with a brief period of light or (d) long days interrupted with a brief pe­ in many other responses
riod of darkness. A long-day plant flowers when it is grown under short-night condi­
to light
tions (b) but does not flower when grown under long-night conditions (a) unless the
Phytochrome is involved in the light
long night is interrupted with a brief period of light (c). These data indicate that the
plant is actually measuring the dark period, not day length. The original hypothesis isrequirement that some seeds have
disproved. ■ for germination. Seeds with a light
requirement must be exposed to
FIGURE 11-17 Photoperiodic responses of short-day and long-day plants. light containing red wavelengths.
Exposure to red light converts Pr to
Pfr, and germination occurs. Many
duce a particular type of phytochrome. Biologists study­ temperate species with small seeds require light for ger­
ing the responses of plants that do not produce an indi­ mination. (Larger seeds generally do not have a light re­
vidual phytochrome have concluded that the individual quirement.) This adaptation enables the seeds to germi­
forms of phytochrome have both unique and overlap­ nate at the optimal time. During early spring, sunlight,
ping functions. including red light, penetrates the bare branches of over­
Each member of the phytochrome family exists in lying deciduous trees and reaches the soil between the
two forms and readily converts from one form to the other trees. As spring temperatures warm, the seeds on the
after absorption of light of specific wavelengths. One soil absorb red light and germinate. During their early
form, designated Pr (for red-absorbing phytochrome), growth, the newly germinated seedlings do not have to
strongly absorbs red light with a relatively short wave­ compete with the taller trees for sunlight.
 Temperature and Reproduction • 2 3 5

i
jii____ -< P LA N TS^EN VIR O N M EN T I P h y t o c h r o m e a n d S h a d e A v o id a n c e

Can plants sense the presence of Since the 1970s, botanists have Pfr forms. This signal triggers a series of
nearby plants? The answer is yes: They recognized the environmental factor responses that cause the shade-avoiding
not only detect the presence of nearby that triggers shade avoidance: Plants plant, which is adapted to full-light envi­
plants, which are potential competi­ perceive changes in the ratio of red ronments, to grow taller or flower earlier.
tors, but they also change the way they to far-red light that results from the When a plant is using many of its re­
grow and develop in response. Many presence of nearby plants. The leaves sources for stem elongation, it has fewer
plants, from small herbs to large trees, of neighboring plants absorb much resources to allocate for new leaves and
compete for light, a response known as more red light than far-red light. (Recall branches, storage tissues, or repro­
shade avoidance, in which plants tend from Chapter 4 that the green pigment ductive tissues. However, for a shade­
to grow taller when closely surrounded chlorophyll strongly absorbs red light avoiding plant that is shaded by its neigh­
by other plants. If successful, the during photosynthesis.) In a densely bors, a rapid increase in stem length is
shade-avoiding plant projects its new plant-populated area, the ratio of red advantageous, because once this plant
growth into direct sunlight, increasing light to far-red light (r/fr) decreases, af­ is taller than its neighbors, it obtains
its chances of survival. fecting the equilibrium between Pr and a larger share of unfiltered sunlight.

Other processes under the influence of phyto­

chrome include sleep movements in leaves (discussed
Temperature
later); shoot dormancy; leaf abscission; and pigment for­ and Reproduction
mation in flowers, fruits, and leaves. (Also see Plants and You have seen that light is an important mechanism af­
the Environment: Phytochrome and Shade A voidance) fecting seasonal responses, such as flowering. Tempera­
ture also affects seasonal responses: certain plants have a
temperature requirement that must be met if they are to
flower. The promotion of flowering by exposure to low
temperature is known as vernalization. The part of the
plant that must be exposed to low temperature varies.
For some plants, recently germinated seedlings have a
“cold” requirement. For other plants, the moist seeds of
some plants must be exposed to low temperature for a
period of several weeks for flowering to occur after the
seeds germinate and grow.
In some plant species, the requirement of a low-
temperature period is absolute, meaning that they will
not flower unless they have been vernalized. Other spe­
cies flower sooner if exposed to low temperatures but
will still flower at a later date if they are not exposed to
low temperatures.
Examples of plants with a low-temperature re­
quirement include annuals such as winter wheat, which
grow, reproduce, and die in 1 year, and biennials such as
carrots, which take 2 years to complete their life cycles.
Winter wheat is planted in the fall and germinates at
that time. The young seedlings are exposed to cold dur­
ing the winter and subsequently flower after resuming
growth the following spring.

FIGURE 11-18 Phytochrome.

This pigment occurs in two forms, designated Pr and Pfr, and VERNALIZATION The low-temperature requirement for flow­
readily converts from one form to the other. Red light converts ering in some plant species.
Pr to Pfr, and far-red light converts Pfr to Pr.
236 • ch ai ter 11 Growth Responses and Regulation o f Growth

Carrots and other bien­

nials grow vegetatively the
first year and store surplus
food in their roots ( Fig­
ure 11-19). If the roots are
not harvested, the plants
flower and reproduce sexu­
ally during the second year,
after exposure to the low
temperatures of winter.
Carrots growing in a warm
environment and not ex­
posed to low temperatures
continue vegetative growth

© Derrick Ditchburn/Visuals Unlimited

indefinitely and do not initi­
ate sexual reproduction.
An external stimulus to
which a plant responds, such
as low temperature, may be
influenced by internal con­
ditions, such as hormone
levels in the plant. It is pos­ (a) The carrot (Daucus carota), a biennial (b) After the plant remains dormant through
sible, for example, to elimi­ plant, grows vegetatively the first year, storing the winter, the energy stored in the root is
nate the low-temperature surplus food in its storage root. used during the plant’s second year to flower
requirement for flowering and reproduce.
in biennials by treating the
plants with the hormone FIGURE 11-19 Temperature requirements for flowering.
gibberellin.

at the time of day that pollinating insects are foraging

Circadian Rhythms for food, reproduction will be unsuccessful.
Almost all organisms, including plants, animals, and Circadian rhythms in plants affect such biological
microorganisms, appear to have an internal timer, or processes as gene expression, the rate of photosynthesis,
biological clock, that approximates a 24-hour cycle, the and the opening and closing of stomata. Sleep move­
time it takes for Earth to rotate around its own axis. ments observed in the common bean and other plants are
Such internal cycles are known as circadian rhythms another example of a circadian rhythm (•Figure 11-20).
(from the Latin circum, “around,” and diurn, “daily”). During the day, bean leaves are horizontal, possibly for
Circadian rhythms help an organism respond to the optimal light absorption, but at night the leaves fold
time of day, whereas photoperiodism enables an organ­ down or up, a movement that orients them perpendicu­
ism to detect the time of year. lar to their daytime position. The biological significance
Why do plants and other organisms have circa­ of sleep movements is unknown at this time.
dian rhythms? iredictable environmental changes, such When constant environmental conditions are main­
as the sunrise and sunset, occur during the course of tained, circadian rhythms repeat every 20 to 30 hours,
each 24-hour period. These predictable changes may at least for several days. In nature, the rising and setting
be important to an individual organism by causing it to of the sun reset the biological clock so that the cycle re­
change its behavior (in the case of animals) or its physi­ peats every 24 hours. What happens if a plant’s circadian
ological processes. It is thought that circadian rhythms clock is mutated so that the day-night cycle is not resyn­
help an organism synchronize repeatable daily activities chronized? In 2005, biologists identified mutants of the
so that they occur at the appropriate time each day. If, model plant Arabidopsis in which the circadian clock is
for example, an insect-pollinated flower does not open not synchronized to match the external day-night cycle.
The mutant plants contained less chlorophyll, fixed less
carbon by photosynthesis, grew more slowly, and had
CIRCADIAN RHYTHM A biological activity with an internal
less of a competitive advantage than Arabidopsis plants
rhythm that approximates the 24-hour day.
with a normal circadian clock.
 Turgor Movements • 2 3 7

For many plants, two photoreceptors—the red

light-absorbing phytochrome and the blue/ultraviolet-
A light-absorbing cryptochrome—are implicated in
resetting the biological clock. Certain parts of the phy­
tochrome molecule are like clock proteins in fruit flies,
fungi, mammals, and bacteria; this molecular evidence
strongly supports the circadian clock role of phyto­
chrome. The evidence for cryptochrome as a clock pro­
tein is also convincing. First discovered in plants, cryp­
tochrome counterparts are found in the fruit fly and
mouse biological clock proteins. Possibly both photo­
receptors are involved in resetting the biological clock
in plants: Researchers have evidence that phytochrome
Dennis Drenner

and cryptochrome sometimes interact to regulate simi­

lar responses.
(a) Leaf position at noon. (b) Leaf position at midnight.
Turgor Movements
FIGURE 11-20 Sleep movements in the common bean
The sensitive plant (Mimosa pudica) dramatically folds
(Phaseolus vulgaris).
It is not known why some plants exhibit sleep movements, but
its leaves and droops in response to touch (or to an elec­
they occur on an approximate 24-hour cycle, regardless of the trical, chemical, or thermal stimulus) (•Figure 11-21).
amount of light or darkness to which the plants are exposed. The response, which typically occurs in a few seconds,
spreads throughout the plant even if only one leaflet is
initially touched. When a sensitive plant leaf is touched,
an electrical impulse moves down the leaf to special
cells housed in an organ at the base of the petiole, called

Leaflet
Dennis Drenner

(a) The sensitive plant before being touched.

Cross-sectional views
(c) How the folding and drooping occurs. Pulvini occur in three areas: the
Dennis Drenner

base of each leaflet, the base of each cluster of leaflets, and the base of each
leaf. Only changes in the pulvini at the bases of leaflets are shown. (Top right)
A section through two leaflets, showing their pulvini when the leaf is undis­
(b) The sensitive plant several seconds after being turbed. (Bottom right) A section through the two leaflets, showing how a loss
touched. of turgor produces the folding of the leaves.

FIGURE 11-21 Turgor movements in the sensitive plant (Mimosa pudica).

238 • chap ter 11 Growth Responses and Regulation o f Growth

the pulvinus (pl., pulvini). The pulvinus is a somewhat causes the leaf movement. Such turgor movements are
swollen joint that acts as a hinge. When the electrical temporary and reversible. The movement of potassium
signal reaches cells in the pulvinus, the electrical sig­ ions and water back into the pulvinus cells causes the
nal induces a chemical signal that increases membrane plant part to return to its original position, although re­
permeability to certain ions. A loss of turgor (rigidity covery takes several to many minutes longer than the
or distension caused by absorption of water) occurs in original movement.
certain pulvinus cells as potassium ions exit through The mechanism by which the Venus flytrap leaf
the now-permeable plasma membrane, causing water to closes is similar to the mechanism of the sensitive plant.
leave the cells by osmosis. The sudden change in turgor An electrical signal, which moves more rapidly than in
the sensitive plant, induces a chemical signal that causes
a movement of potassium ions out of certain cells, fol­
TURGOR MOVEMENT A temporary plant movement that re­
lowed by the exit of water. The loss of turgor causes the
sults from changes in internal water pressure in a plant part.
leaf to snap shut.

STUD Y O U T L I N E
0) Discuss genetic and environmental factors that affect 4 Relate which hormone or hormones is/are involved in
plant growth and development. each of the following biological processes: leaf abscis­
Plant growth and development are controlled by both sion, seed germination, apical dominance.
internal genetic factors and external environmental Many plant processes may be the result of interactions
factors. The location of a cell in the young plant body of several hormones rather than the effect of a single
affects gene expression during development, causing hormone. Ethylene, along with auxin, is involved in leaf
some genes in that cell to be turned off and others to abscission; as a leaf ages, the level of auxin in the leaf
be turned on. Many factors in the physical environment decreases, and the level of ethylene increases. Gibberel-
(such as changing day length, variation in precipitation, lins are involved in the seed germination of certain plants
and temperature) determine gene expression and affect (cereals and other grasses); application of gibberellins
plant growth and development. substitutes for low-temperature or light requirements for
germination in seeds such as lettuce, oats, and tobacco.
42 Describe phototropism, gravitropism, and Ethylene also promotes seed germination, whereas ab­
thigmotropism. scisic acid inhibits seed germination. Apical dominance
Tropisms are directional growth responses and are is the inhibition of axillary bud growth by the apical meri-
permanent. Phototropism is the directional growth of stem. In plants with strong apical dominance, auxin pro­
a plant caused by light. Gravitropism is plant growth in duced in the shoot apical meristem inhibits axillary buds
response to the direction of gravity. Thigmotropism is near the apical meristem from developing into actively
plant growth in response to contact with a solid object. growing shoots. Cytokinins and ethylene also affect api­
cal dominance: cytokinins promote the growth of axillary
43 List several ways in which each of the following hor­
buds, and ethylene inhibits axillary bud development.
mones affects plant growth and development: auxin,
gibberellin, cytokinin, ethylene, abscisic acid. 5 Explain how varying amounts of light and darkness in­
Hormones are organic chemical messengers in plants duce flowering, and describe the role of phytochrome.
and other multicellular organisms. Biologists have Photoperiodism is any physiological response (such as
identified five major classes of plant hormones (auxin, flowering) to variations in the length of daylight and
gibberellin, cytokinin, ethylene, and abscisic acid) as darkness. Some plants are short-day plants, some are
well as a variety of signaling molecules. Auxin is involved long-day plants, and others are intermediate-day plants.
in growth and development, including stem elongation, In these cases, the plant is actually measuring the
apical dominance, and root formation on cuttings. Gib­ length of the dark period. In day-neutral plants, photo­
berellin is involved in stem elongation, flowering, and period does not affect flowering. Phytochrome is a blue-
seed germination. Cytokinin is involved in cell division green proteinaceous pigment involved in many plant
and delay of senescence. Ethylene is a gaseous hormone responses to light, independent of photosynthesis. There
involved in leaf abscission and fruit ripening. Abscisic are about five different phytochrome proteins; each
acid is involved in dormancy and responses to stress. exists in two forms and readily converts from one form