Plant Physiology

and Reproduction
Chapter Concepts
10.1 Water and Mineral Transport
Evaporation pulls water and minerals from the
roots to the leaves in xylem. 170
Stomates must be open for evaporation to
occur. 172

10.2 Organic Nutrient Transport

Osmotic pressure pushes organic nutrients in
phloem, usually from the leaves to the
roots. 174

10.3 Plant Responses to Environmental Stimuli

Plants respond to outside stimuli by changing
their growth patterns. 176
Plant hormones regulate plant growth patterns.
176
Some plant responses are controlled by the
length of daylight (photoperiod). 178

10.4 Sexual Reproduction in Flowering Plants

In the plant life cycle, the pollen grain carries
sperm from ower to ower. 180
Seeds within fruits are the products of sexual
reproduction in owering plants. 181
Seeds must be dispersed and must germinate to
complete the plant life cycle. 184

10.5 Asexual Reproduction in Flowering Plants

Plants reproduce asexually. This ability can be
commercially utilized to mass produce identical
plants, sometimes after genetic engineering. 186

Flowering plants disperse their seeds in various ways. Milkweed

relies on the wind to carry seeds to new locations after they are
released from pods.

169

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Part 2

10-2

Plant Biology

Japanese farmer raises a bowl of steaming rice to his

lips as a Mexican mother serves freshly prepared tortillas and an American teenager reaches for another
slice of white bread. Rice, corn, and wheat are the staples of
diets throughout the world. Italians eat pasta, Irish and Germans consume potatoes, and Americans eat bread as a
source of carbohydrates, the nutrient that supplies most of
our energy needs. But plants also provide fats and minerals.
Then, too, plants have cell walls of cellulose that provide
much of the roughage in our diets. Eat your vegetables is
not the nagging expression it seemsit is a recognition that
plants provide us with all the essential nutrients that allow
our cells to continue metabolizing. The dietary importance of
plants gives ample reason for us to know something about
their anatomy and physiology.

10.1

Water and Mineral Transport M

Water and minerals enter a plant at the root, primarily

through the root hairs. From there water, along with minerals, moves across the tissues of a root until it enters xylem,
the vascular tissue that contains the hollow conducting cells
called tracheids and vessel elements (Fig. 10.1). The vessel elements are larger than the tracheids, and they are stacked one
on top of the other to form a pipeline that stretches from the
roots to the leaves. It is an open pipeline because the vessel
elements have no end walls separating one from the other.
The tracheids, which are elongated with tapered ends, form
a less obvious means of transport. Water can move across
the end and side walls of tracheids because of pits, or depressions, where the secondary wall does not form.
Water entering root cells creates a positive pressure called
root pressure that tends to push xylem sap upwards. It is estimated that root pressure, like atmospheric pressure, would be
able to raise water to 10.4 meters. However, since some trees
can be as tall as 120 meters, other factors must be involved.

Cohesion-Tension Theory of Xylem

Transport
The cohesion-tension theory of xylem transport explains
how water is transported to great heights against gravity
(Fig. 10.2). Because water molecules are polar, they adhere to
the walls of the vessel elements; and because of hydrogen
bonding, water molecules are cohesivethey cling together.
Cohesion of water molecules within the xylem pipeline is absolutely necessary for water transport in a plant. It causes
water to ll the pipeline completely, from the roots to the
leaves, and to resist any separation.
The other factor that causes water to rise in plants is
transpiration, the loss of water by evaporation. Much of the
water that is transported from the roots to the leaves evaporates and escapes from the leaf by way of the stomates,
openings where gas exchange occurs. A single corn plant
loses somewhere between 135 and 200 liters of water
through transpiration during a growing season.
The water molecules that evaporate are replaced by other
water molecules from the leaf veins. Therefore, transpiration
exerts a driving force by creating a negative pressurethat is,
a tensionwhich draws a column of water up the vessel elements from the roots to the leaves. As long as water molecules evaporate from the plant and the soil contains enough
moisture, the water column is pulled continuously upward.
The tension created by transpiration is effective only because of the cohesive property of water. Therefore, this explanation for xylem transport is called the cohesiontension theory.

Water is transported from the roots to the leaves

in xylem. The water column remains intact as
transpiration pulls water from the roots to the
leaves.

perforation
pits

a.

50 m

b.

31 m

pits

Figure 10.1 Conducting cells of xylem.

a. Tracheids are long, hollow cells with tapered ends. Water can move into and out of tracheids through pits only. b. Vessel elements are wider
and shorter than tracheids. Water can move from vessel element to vessel element through perforations (a large hole at each end). Vessel
elements can also exchange water with tracheids through pits.

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Chapter 10

Plant Physiology and Reproduction

171

The sun causes water to evaporate

the energy for xylem transport comes from the sun.
xylem in leaf vein

water molecules
outside air
stomate
mesophyll cells

Transpiration (evaporation) of water from leaves creates

tension that pulls the water column in xylem from the roots.

xylem

cohesion by
hydrogen bonding
between water
molecules
cell wall
adhesion due to
polarity of water
molecules

Water column is held together by cohesion;

adhesion keeps water column in place.

root hair
soil particles
xylem
water molecule

Water from soil enters xylem in root;

tension in water column extends from leaves to root.

Figure 10.2 Cohesion-tension theory of xylem transport.

Tension (a negative pressure) created by evaporation (transpiration) at the leaves pulls water from the root along the length of the plant.

Mineral Transport
Plants need only inorganic nutrients in order to produce all
the organic molecules that make up their bodies. Aside from
carbon, hydrogen, and oxygen obtained from carbon dioxide, the mineral elements listed in Table 10.1 are required by
plants. Minerals diffuse into root hairs, but eventually a
plant uses active transport to increase its uptake of minerals
which are carried along with water in xylem. A plant uses a
great deal of ATP for active transport of minerals into root
cells and xylem.
Human beings are fortunate that plants can concentrate minerals, for we often are dependent on them for our
basic supply of such minerals as calcium to build bones
and teeth and iron to help carry oxygen to our cells. Minerals like copper and zinc are cofactors for the functioning of
enzymes.

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Table 10.1 Inorganic Nutrients for Plants

Macronutrients
CaNO3

Ca, N (calcium, nitrogen)

NH4H2PO4

N, P (nitrogen, phosphorus)

KNO3

K, N (potassium, nitrogen)

MgSO4

Mg, S (magnesium, sulfur)

Micronutrients
Fe-EDTA

Fe (iron)

ZnSO4

Zn (zinc)

KCl

Cl (chlorine)

CuSO4

Cu (copper)

MnSO4

Mn (manganese)

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Part 2

10-4

Plant Biology

Open Stomate

Closed Stomate

50 m

50 m
H 2O

H 2O

guard
cell

H 2O

H 2O

K+
K+

stomate
K+ enters guard
cells and water
follows

K+ exits guard
cells and water
follows

Figure 10.3 Opening and closing of stomates.

Stomates open when water enters guard cells and turgor pressure increases. Stomates close when water exits guard cells and there is a loss of
turgor pressure.

Opening and Closing of Stomates

For a plant to photosynthesize, stomates must be open so
that CO2 will enter the leaves. But when stomates are open,
water is also exiting the leaves because transpiration is occurring. When a plant is water stressed, stomates close to
conserve water and then photosynthesis ceases.
The two guard cells on either side of a stomate regulate
whether the stomate is open or closed. When water enters
guard cells, their central vacuoles ll and the stomate opens.
When water exits guard cells, the stomate closes. Notice in
Figure 10.3 that the guard cells are attached to each other at
their ends and that the inner walls are thicker than the outer
walls. When water enters guard cells, cellulose microbrils
in the walls prevent radial expansion. Lengthwise expansion, which can occur, causes guard cells to buckle out from
the region of their attachment and the stomate opens.
When a plant is photosynthesizing, an ATP-driven
pump actively transports H out of the cell. Now potassium
ions (K) enter the guard cells and when water follows by
osmosis, the stomate opens. When the pump is not working,
K moves into surrounding cells, the guard cells lose water,
and the stomate closes.

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What turns the H pump on or off? It appears that the

blue-light component of sunlight is a signal for stomates to
open. There is evidence to suggest that a avin pigment absorbs blue light, and then this pigment sets in motion the cytoplasmic response that leads to activation of the H pump.
Similarly, there could be a receptor in the plasma membrane
of guard cells that brings about inactivation of the pump
when carbon dioxide (CO2) concentration rises, as might
happen when photosynthesis ceases. Abscisic acid (ABA),
which is produced by cells in wilting leaves, can also cause
stomates to close. Although photosynthesis cannot occur,
water is conserved.
If plants are kept in the dark, the stomates open and
close on a 24-hour basis just as if they were responding to
the presence of sunlight in the daytime and the absence of
sunlight at night. This means that there must be some sort of
internal biological clock that is keeping time. Circadian
rhythms (a behavior that occurs every 24 hours) and biological clocks are areas of intense investigation at this time.

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When stomates open, rst K and then water

enters guard cells. Stomates open and close in
response to environmental signalsthe exact
mechanism is being investigated.

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Biodiversity and Nitrogen Uptake by Plants

G. David Tilman of the University of Minnesota was familiar
with long-term experiments showing that amount of soil nitrogen usually limits biodiversity. The better competitors for soil
nitrogen are expected to be prevalent to the point that other
species do not have a chance to continue existing. Yet the grasslands he was studying often
had more than 100 plant
species coexisting in an area
the size of a few hectares
(10,000 square meters). This
made him hypothesize that
while competition for soil nitrogen might limit biodiversity,
some other factor must be promoting biodiversity. First he
studied the response of grassland plants to predation by
insect and mammalian herbivores, light availability, and
re. None of these seemed to
account for the biodiversity he
observed.
Finally, he found that
grassland plants differ in their
ability to disperse to new sites.
He discovered this by planting
over 50 different plant species
in several plots and recording
how well they could germinate, grow, and reproduce
there. The best competitors for
nitrogen (bunch grasses) allocated 85% of their biomass to
root growth and only 0.5% of
their biomass to making seeds.
Their massive root systems
make them able to spread out
where they are! Little bluestem
is an example of a bunchgrass

(Fig. 10A) The best dispersers allocated 40% of their biomass to

root and 30% of their biomass to seed. Bent grass is an example
of a poor competitor for soil nitrogen but a good disperser (Fig.
10B). The production of seeds allows these plants to occupy any
new sites available.
Tilman developed a mathematical model which showed
that such a tradeoff (root versus
seed allocation) could explain
the stable coexistence of a whole
range of plant species that differ
according to their ability to occupy present space or to disperse to new areas. Coexistence
occurs because better competitors for soil nitrogen are poorer
dispersers, and therefore they do
not occupy all sites. Better dispersers are better at nding and
occupying all possible sites. Because the grasses varied in their
ability to take up nitrogen and
disperse, a number of species
could coexist.
Tilman studied another issue
that relates to biodiversity. He
and colleagues were annually
sampling 207 permanent plots
when there was a drought in
1987-88. The 1988 drought was
the third worst in the past 150
years. They found that plots that
contained only one to four
species suffered a greater loss of
overall biomass than plots that contained 16 to 26 species. This
suggests to Tilman that high biodiversity does buffer an ecosystem against an environmental disturbance, and therefore it is
wise to conserve the biodiversity of ecosystems in all areas
whether in Minnesota, New Jersey, Oregon, or the tropics.

Figure 10A

Figure 10B

Little bluestem (Schizachyrium scoparium) is a North American

bunchgrass that prefers sandy or rocky habitats. It is an excellent
competitor for soil nitrogen because of its high allocation of roots.

Bent grass (Agrostis scabra) is a native North American prairie grass

that is a poor competitor for soil nitrogen. It disperses rapidly into
disturbed areas because of its high allocation of seed.

173

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Part 2

10.2

10-6

Plant Biology

Organic Nutrient Transport M

The leaves carry out photosynthesis and produce the sugar

sucrose, which is transported in the vascular tissue called
phloem to all parts of a plant. The movement of organic substances in phloem is termed translocation. Translocation
makes sugars available to those parts of a plant that are actively metabolizing and growing. The conducting cells in
phloem are sieve-tube elements, each of which typically has
a companion cell (Fig. 10.4). Sieve-tube elements contain cytoplasm but have no nucleus. Their end walls have pores and
resemble a sieve; therefore, the end walls are said to be sieve
plates. The sieve-tube elements are aligned vertically, and
strands of cytoplasm called plasmodesmata (sing., plasmodesma) extend from one cell to the other through the
sieve plates. Therefore, there is a continuous pathway for organic nutrient transport throughout the plant.
The smaller companion cell, which does have a nucleus,
is a more generalized cell than the sieve-tube element. It is
speculated that the companion cell nucleus controls and

sieve plate
with sieve pores

sieve-tube
element

companion cell

maintains the lives of both itself and the sieve-tube element,

and may help a sieve-tube element perform its translocating
function.

Pressure-Flow Theory of Phloem Transport

Chemical analysis of phloem sap shows that it is composed
chiey of sugar and that the concentration of organic nutrients is 1013% by volume. Samples for chemical analysis
most often are obtained by using aphids, small insects that
are phloem feeders. The aphid drives its stylet, a short
mouthpart functioning like a hypodermic needle, between
the epidermal cells and withdraws phloem sap from a sievetube element. The body of the aphid can be cut away carefully, leaving the stylet, which exudes phloem sap for
collection and analysis.
During the growing season the leaves are a source of
sugarthey are photosynthesizing and producing sugar.
This sugar is actively transported into sieve-tube elements,
and water follows passively by osmosis. Active transport is
possible because sieve-tube elements have a living plasma
membrane, and the necessary energy is provided by the
companion cells. The buildup of water within the sieve-tube
elements creates pressure, which starts a ow of phloem
sap. The roots (and other growth areas) are a sink for
sugarthe roots are removing sugar. It is actively transported out of the sieve-tube elements at this location. When
water follows passively by osmosis, phloem sap ows from
the leaves (source) to the roots (sink) (Fig. 10.5). This explanation for translocation in phloem is called the pressureow theory.
The pressure-ow model is supported by an experiment
in which two bulbs are connected by a glass tube. The rst
bulb contains solute at a higher concentration than the second bulb. Each bulb is bounded by a differentially permeable membrane, and the entire apparatus is submerged in
distilled water:

nucleus

flow of solution

cytoplasm

concentrated
sugar
solution

H2O
1

differentially
permeable
membranes
H2O

Figure 10.4 Sieve-tube elements of phloem.

Sieve-tube elements contain cytoplasm, and their at end walls have
a sieve plate with large sieve pores. Plasmodesmata connect the
cells one with the other. Each sieve-tube element has a companion
cell that has both cytoplasm and a nucleus.

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dilute
sugar
solution

H2O

Distilled water ows into the rst bulb because it has the
higher solute concentration. In this way a pressure difference is created that causes water to ow from the rst bulb
to the second and even to exit from the second bulb. As the
water ows, it carries solute with it from the rst to the second bulb.

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Chapter 10

Plant Physiology and Reproduction

175

Sucrose is actively
transported into phloem,
and water follows
by osmosis.

xylem
vessel

source
(mature
leaf
cells)

sieve-tube
element

phloem

companion
cell

This creates a positve

pressure that causes
sap to flow within phloem

water
molecule

sucrose
molecule

sieve
plate

sink
(root
and
other
growth
areas)

Sucrose is actively
transported out of
sieve-tube cells,
and water follows
by osmosis.

Figure 10.5 Pressure-ow theory of phloem transport.

Sugar and water enter sieve-tube elements at a source. This creates a positive pressure, which causes phloem contents to ow. Sieve-tube
elements form a continuous pipeline from a source to a sink, where sugar and water exit sieve-tube elements.

The pressure-ow theory of phloem transport can account for any direction of ow in sieve tubes if we consider that the direction of flow is always from source to
sink. In young seedlings, the cotyledons containing reserved food are a major source of sucrose, and roots are a
sink. Therefore, the flow is from the cotyledons to the
roots. In older plants, the most recently formed leaves can
be a sink and they will receive sucrose from other leaves
until they begin to maximally photosynthesize. When a
plant is forming fruit, phloem ow is monopolized by the
fruits, little goes to the rest of the plant, and vegetative
growth is slow.

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Because phloem sap ows from source to a sink, situations

arise in which there is a bidirectional ow within phloemnot
just at different times in the life cycle but even at the same
time! Bidirectional ow is occurring because different sieve
tubes can be conducting phloem sap in opposite directions.
Phloem transports organic nutrients in a plant.
Typically, sugar and then water enter sieve-tube
elements in the leaves. This creates pressure,
which causes water to ow to the roots, carrying
sugar with it.

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Part 2

10-8

Plant Biology

10.3

Auxin

Plant Responses to
Environmental Stimuli
Plants respond to environmental stimuli (e.g., light, day
length, gravity, and temperature) usually by changing their
growth patterns. Plant growth toward or away from a directional stimulus is called a tropism. Three well-known
tropisms, each named for the stimulus that causes the response, are

Phototropism: a movement in response to a light stimulus

Gravitropism: a movement in response to gravity
Thigmotropism: a movement in response to touch

Growth toward a stimulus is called a positive tropism and

growth away from a stimulus is called a negative tropism
(Fig. 10.6). Tropisms are due to differential growthone
side of an organ elongates faster than the other, and the result is a curving toward or away from the stimulus.
Among the principal internal factors that regulate such
responses are plant hormones. A hormone is a chemical
messenger produced in very low concentrations that has
physiological and/or developmental effects, usually in another part of the organism. Some plant hormones are made
in meristem and transported to other tissues; others move
directly from the tissue of origin to another tissue; and still
others are used where they are made.
Table 10.2 lists the major types of plant hormones and
their primary functions. Some of these are promoters of
growth and some are inhibitors of growth. These hormones
often interact to control physiological responses. Each naturally occurring hormone has a specic chemical structure.
Other chemicals, some of which differ only slightly from the
natural hormones, also affect the growth of plants. These
and the naturally occurring hormones are sometimes
grouped together and called plant growth regulators.

Auxin is a plant hormone whose effects have been studied

for a long time. The only known naturally occurring auxin is
indoleacetic acid (IAA). It is produced in shoot apical meristem and is found in young leaves and in owers and fruits.
Therefore, you would expect that auxin would affect many
aspects of plant growth and development. Apically produced auxin prevents the growth of axillary buds, a phenomenon called apical dominance. When a terminal bud is
removed deliberately or accidentally, the nearest axillary
buds begin to grow, and the plant branches. To achieve a
fuller look, one generally prunes the top (apical meristem) of
the plant. This removes apical dominance and causes more
branching of the main body of the plant.

Figure 10.6 Positive phototropism.

The stem of a plant bends toward the light as it grows.

Table 10.2 Plant Hormones

Type

Primary Example

Notable Function

Auxin

Indoleacetic acid (IAA)

Promotes cell elongation in stems; phototropism, gravitropism, apical dominance;

formation of roots, development of fruit

Gibberellins

Gibberellic acid (GA)

Promote stem elongation; release some buds and seeds from dormancy

Cytokinins

Zeatin

Promote cell division and embryo development; prevent leaf senescence and promote
bud activation

Abscisic acid

Abscisic acid (ABA)

Resistance to stress conditions; causes stomatal closure; maintains dormancy

Ethylene

Ethylene

Promotes fruit ripening; promotes abscission and fruit drop; inhibits growth

Growth Promoters

Growth Inhibitors

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Chapter 10

Plant Physiology and Reproduction

177

After tips are placed on agar,

agar is cut into blocks.

Coleoptile tip
is intact.

Coleoptile tip
is removed.

Block is placed
to one side of
coleoptile.

Curvature occurs
beneath block.

Figure 10.7 Demonstrating phototropism.

Oat seedlings are protected by a hollow sheath called a coleoptile. After a tip is removed and placed on agar, an agar block placed on one side of
the coleoptile can cause it to curve.

The application of a weak solution of auxin to a

woody cutting causes roots to develop. Auxin production
by seeds also promotes the growth of fruit. As long as
auxin is concentrated in leaves or fruits rather than in the
stem, leaves and fruits do not fall off. Therefore, trees can
be sprayed with auxin to keep mature fruit from falling to
the ground.
Auxin is involved in gravitropism as well as phototropism. After gravity has been perceived, auxin moves to the
lower surface of roots and stems. Thereafter, roots curve
downward and stems curve upward.

cell wall
H+
ATP

ATP

H+

ATP

H+

receptor

Phototropism and Auxin

The role of auxin in the positive phototropism of stems
has been studied for quite some time. The experimental
material of choice has been oat seedlings with coleoptile
intact. (A coleoptile is a protective sheath for the young
leaves of the seedling.) For example, if the tips of the
coleoptiles are cut off and placed on agar (a gelatinlike
material), and then an agar block is placed to one side of a
tipless coleoptile, the shoot will curve away from that
side. The bending occurs even though the seedlings are
not exposed to light (Fig. 10.7). Apparently, the agar
blocks contain a chemical that was produced by the coleoptile tips.
Because blue light in particular causes phototropism
to occur, it is believed that a yellow pigment related to the
vitamin riboflavin acts as a photoreceptor for light. Following reception, auxin migrates from the bright side to
the shady side of a stem. The cells on that side elongate
faster than those on the bright side, causing the stem to
curve toward the light. It is now known that when a plant
is exposed to unidirectional light, auxin moves to the
shady side, where it binds to receptors and activates an
ATP-driven H pump (Fig. 10.8). As hydrogen ions (H)

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auxin

ATP

ATP
H+

H+

Figure 10.8 Auxin mode of action.

After auxin binds to a receptor, the combination stimulates an H
pump so that hydrogen ions (H) are transported out of the
cytoplasm. The resulting acidity causes the cell wall to weaken, and
the electrochemical gradient causes solutes to enter the cell. Water
follows by osmosis and the cell elongates.

are being pumped out of the cell, the cell wall becomes
acidic, breaking hydrogen bonds within cellulose. Cellulose fibrils are weakened, and activated enzymes further
degrade the cell wall. The electrochemical gradient established by the H pump causes solutes to enter the cell,
and water follows by osmosis. The turgid cell presses
against the cell wall, stretching it so that elongation
occurs.

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Part 2

10-10

Plant Biology

Other Plant Growth Regulators

Now that the formulas for many plant hormones are known,
it is possible to synthesize them, as well as related chemicals,
in the laboratory. Collectively, these substances are known as
plant growth regulators. Many scientists hope plant growth
regulators will bring about an increase in crop yield, just as
fertilizers, irrigation, and pesticides have done in the past.
In high concentrations, some auxins are used widely in
agriculture as herbicides to prevent the growth of broadleaved plants. In addition to their use in weed control, the
synthetic auxins known as 2,4D and 2,4,5T were used as defoliants during the Vietnam War. Even though 2,4D has been
used for over 35 years, we do not yet know how it works.
Apparently, it is structurally different enough from natural
auxin that plants cannot store it in an inactive form and a
plants own enzymes cannot break it down. Its concentration rises until metabolism is disrupted, cellular order is lost,
and the cells die.
Other plant hormones also have agricultural and commercial uses. When gibberellins are sprayed on grapefruit
(Fig. 10.9), the peel stays youthfulgreen, rm, and unappealing to various types of fruit ies. Yet the treated fruit is
fully ripe and sweetperfect for juicing or eating. Gibberellins are used to stimulate seed germination and
seedling growth of some grains, beans, and fruits. They also
increase the size of some mature plants. Treatment of sugarcane with as little as 2 oz per acre increases the cane yield by
more than 5 metric tons. The application of either auxins or
gibberellins can cause an ovary and accessory ower parts
to develop into fruit, even though pollination and fertilization have not taken place. In this way, it is sometimes possible to produce seedless fruits and vegetables, or bigger,
more uniform bunches with larger fruit.
Because cytokinins retard the aging of leaves and other
organs, they are sprayed on vegetables to keep them fresh
during shipping and storage. Such treatment of holly, for example, allows it to be harvested many weeks prior to a holiday.
Plant growth regulators are used in tissue culture when
seedlings are grown from a few cells in laboratory glassware. New varieties of food crops with particular characteristicssuch as tolerance to herbicides and insectsare
being developed by genetically engineering the original
cells. One day it may be possible to endow most crops with
the ability to utilize atmospheric nitrogen and/or make a
wider range of proteins.
Several synthetic inhibitors are used to oppose the action of auxins, gibberellins, and cytokinins normally present
in plants. Some of these can cause leaves and fruit to drop at
a time convenient to the farmer. Removing leaves from cotton plants aids in their harvest, and thinning the fruit of
young fruit trees results in larger fruit as the trees mature.
Retarding the growth of other plants sometimes increases
their hardiness. For example, an inhibitor has been used to
reduce stem length in wheat plants, so that the plants do not
fall over due to heavy winds and rain.

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Figure 10.9 Grapefruit being sprayed with gibberellins.

When sprayed with gibberellins, the peel stays green and rm so that
fruit ies look for another place to lay their eggs. The tougher skin
improves shipping and can be turned yellow by an application of
ethylene.

The commercial uses of ethylene were greatly increased with the development of ethylene-releasing compounds. Ethylene gas is injected into airtight storage
rooms to ripen bananas, honeydew melons, and tomatoes. It will also degreen oranges, lemons, and grapefruit
when the rind would otherwise remain green because of
a high chlorophyll level. When sprayed on certain fruit
and nut crops, ethylene increases the chances that the
fruit will detach when the trees are shaken at harvest
time.
Today, elds and orchards are often sprayed with synthetic growth regulators.

Photoperiodism
A response based on the proportion of light to darkness in a
24-hour cycle is called photoperiodism. Photoperiodic
responses in plants are particularly obvious in the temperate zone. In the spring, plants respond to increasing day
length by initiating growth; in the fall, they respond to
decreasing day length by ceasing growth processes. Day

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Chapter 10

critical length

Plant Physiology and Reproduction

critical length

cocklebur

179

clover

flower

flash

a. Short-Day (Long-Night) Plant

flash

b. Long-Day (Short-Night) Plant

Figure 10.10 Effect of day length on two types of plants.

Some plants ower only during a particular season. The cocklebur owers when days are short (a), and clover owers when days are long
(b). The length of the night is the determining factor, proven by interrupting a longer-than-critical-length night with a ash of light. Dependency
on the photoperiod is exploited by commercial greenhouses, which use it to grow plants out of season.

length controls owering in some plants; for example, violets and tulips ower in the spring, asters and goldenrods
ower in the fall.
Long-day plants initiate owering when the days get
longer than a certain minimum value, or critical length.
Short-day plants initiate owering when the days get
shorter than a critical length. Day-length-neutral plants are
insensitive to the length of the day. The cocklebur is a shortday plant; if a long night is interrupted by a ash of light, it
will not ower (Fig. 10.10). Clover, on the other hand, is a
long-day plant; if a long night is interrupted by a ash of
light, it will still ower. Interrupting the day with darkness
has no effect. This shows that the length of continuous darkness, not the day length, actually controls owering.
Florists use information about photoperiodism in order
to provide us with owers at a particular time of year or out
of season. They only need to know how long it takes for a
certain type of plant to ower, and then they start manipulating the photoperiod in order to have owers ready on
time. If chrysanthemums, which ower in the fall, are desired in the spring, they use blackout shades to articially
produce long nights. On the other hand, orists switch on
incandescent lights so that irises, which are spring plants,
are available in the winter. These same techniques make it
possible to always have poinsettias at Christmas and lilies at
Easter.
If owering is dependent on night and day length,
plants must have a photoreceptor to detect these periods.
The photoreceptor for photoperiodism is phytochrome, a

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blue-green leaf pigment that alternately exists in these two

forms:

(daytime)
red
light
Pr
(inactive form)

far-red
light
(shade and evening)

Pfr
(active form)

It could be that owering is initiated in some plants according to the time of day phytochrome changes its structure.
The pigment could be part of a biological clock system that
controls owering. Phytochrome is also believed to be involved in controlling a plants growth pattern in a low-light
versus a high-light environment.

Hormones often coordinate the response of plants

to various stimuli. The proportion of stimulatory
and inhibitory hormones often determines the
particular response. Phytochrome is a pigment
involved in response of plants to the photoperiod
which affects owering in some plants.

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Plant Biology

10.4

pistil

Sexual Reproduction in
Flowering Plants
Sexual reproduction is the rule in owering plants. This may
come as a surprise to those who never thought of plants in
terms of male and female. However, sexual reproduction is
dened properly as reproduction requiring gametes, often
an egg and a sperm. In a owering plant, the structures that
produce the egg and sperm are located within the ower.

stigma anther
style
filament
ovary

stamen

180

petal
ovule
sepal

zygote
sporophyte

Structure of Flowers
Figure 10.11 shows the parts of a typical ower. The sepals,
most often green, form a whorl about the petals, the color of
which accounts for the attractiveness of many owers. The
size, shape, and color of a ower attracts a specic pollinator.
Flowers pollinated by the wind often have no petals at all.
In the center of the ower is a small vaselike structure,
the pistil, which usually has three parts: the stigma, an enlarged sticky knob; the style, a slender stalk; and the ovary,
an enlarged base. The ovary contains one or more ovules,
which play a signicant role in reproduction. Grouped about
the pistil are the stamens, each of which has two parts: the
anther, a saclike container, and the lament, a slender stalk.
Not all owers have sepals, petals, stamens, and a pistil.
Those that do are said to be complete and those that do not
are said to be incomplete. Flowers with only stamens are
called staminate owers and those with only pistils are
called pistillate owers. If staminate owers and pistillate
owers are on one plant, as in corn, the plant is monoecious.
If staminate and pistillate owers are on separate plants, the
plant is dioecious. Holly trees are dioecious and, if red
berries are a priority, it is necessary to acquire a plant with
staminate owers and another with pistillate owers.

diploid (2n)

fertilization

meiosis

haploid (n)
megaspore
egg

sperm

megagametophyte
(embryo sac)

microspore

microgametophyte
(pollen grain)

Figure 10.11 Alternation of generations in a owering

plant.
Flowering plants are heterosporous; they produce microspores and
megaspores. A microspore becomes a pollen grain, which is a
microgametophyte that produces sperm. A megaspore becomes an
embryo sac, or megagametophyte, that produces an egg. When a
sperm joins with an egg, the resulting zygote develops into an
embryo, retained within a seed.

Alternation of Generations
Plants have a life cycle called alternation of generations because two generations are involved: the sporophyte and the
gametophyte. The sporophyte is a diploid (2n) generation
that produces haploid (n) spores by meiosis. A ower produces two types of spores: microspores and megaspores.
Microspores are produced in the anthers of stamens, and
megaspores are produced within ovules (Fig. 10.11). A microspore becomes a pollen grain, which upon maturity is a
sperm-containing microgametophyte, also called the male gametophyte. A megaspore becomes an egg-containing embryo sac, which is a megagametophyte, also called the female
gametophyte. Following fertilization, the zygote develops
into an embryo located within a seed. When the seed germinates, the new sporophyte plant begins to grow.
Figure 10.12 shows these same steps in greater detail.
Within an ovule, a megaspore (mega means large) parent cell
undergoes meiosis to produce four haploid megaspores.
Three of these megaspores disintegrate, leaving one func-

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tional megaspore, which divides mitotically. The result is the

megagametophyte, or embryo sac, which typically consists
of eight haploid nuclei embedded in a mass of cytoplasm.
The cytoplasm differentiates into cells, one of which is an
egg and another of which is the central cell with two nuclei
(called the polar nuclei).
The anther has pollen sacs, which contain numerous microspore (micro means small) parent cells. Each parent cell
undergoes meiosis to produce four haploid cells called microspores. The microspores usually separate, and each one
becomes a pollen grain. At this point, the young pollen grain
contains two cells, the generative cells and the tube cell.
Pollination occurs when pollen is windblown or carried by insects, birds, or bats to the stigma of the same
type of plant. Only then does a pollen grain germinate
and develop a long pollen tube. This pollen tube grows
within the style until it reaches an ovule in the ovary. At
this time, too, the generative cell divides mitotically, pro-

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Plant Physiology and Reproduction

Chapter 10

In pollen sacs, microspore

parent cell produces
microspores.

181

microspore
parent
cell
stigma
meiosis

pistil
anther

stamen

style

In ovule, megaspore
parent cell produces
megaspores.

filament

Microspores
develop into
microgametophytes
(pollen grains).

ovule
ovary
4 microspores

megaspore parent
cell undergoes
meiosis

mitosis
tube cell

sepal
3 megaspores
disintegrate

pollen grain
(microgametophyte)
generative cell

mitosis
sperm

One megaspore becomes

embryo sac
(megagametophyte).

pollen tube

polar
nuclei

embryo
Ovule develops into a seed
which contains the
embryonic sporophyte and
endosperm.

endosperm

embryo sac
(megagametophyte)
egg
cell

seed
coat
seed

fertilization
One sperm from microgametophyte
fertilizes egg; another sperm joins
with polar nuclei to produce endosperm.

View 1

View 2

Figure 10.12 Life cycle of a owering plant.

The owering plant life cycle involves production of eggs and sperm by gametophyte generations and development of an embryo-containing seed.

ducing two sperm, which have no flagella. On reaching

the ovule, the pollen tube discharges the sperm. These
two fusions are known as double fertilization. One of the
two sperm migrates to and fertilizes the egg, forming a
zygote, and the other sperm migrates to and unites with
the two polar nuclei, producing a 3n (triploid) endosperm
nucleus.
The endosperm nucleus divides to form endosperm,
food for the developing plant. The zygote develops into a
multicellular embryo and the ovule wall hardens and becomes the seed coat. A seed is a structure formed by matu-

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ration of an ovule; it consists of a sporophyte embryo,

stored food, and a seed coat. The ovary, and sometimes
other oral parts, develops into a fruit.
Flowering plants produce an egg within each ovule
of an ovary, and sperm within pollen grains. They
have double fertilizationone fertilization
produces the zygote; the other produces
endosperm. The ovule now becomes a seed and
the ovary becomes a fruit.

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Plant Biology

bring about primary growth in a plant; the shoot apical

meristem is responsible for aboveground growth, and the
root apical meristem is responsible for underground growth.
Monocots, unlike dicots, have only one cotyledon. Another important difference between monocots and dicots is
the manner in which nutrient molecules are stored in the
seed. In a monocot, the cotyledon rarely stores food; rather,
it absorbs food molecules from the endosperm and passes
them to the embryo. During the development of a dicot embryo, the cotyledons usually store the nutrient molecules
that the embryo uses. Therefore, in Figure 10.13 we can see
that the endosperm seemingly disappears. Actually, it has
been taken up by the two cotyledons. In a plant embryo, the
epicotyl is above the cotyledon and contributes to shoot development; the hypocotyl is that portion below the cotyledon that contributes to stem development; and the radicle
contributes to root development. The embryo plus stored
food is now contained within a seed.

endosperm. The ovule now becomes a seed and

the ovary becomes a fruit.

Development of the Plant Embryo

Stages in the development of a dicot embryo are shown in
Figure 10.13. After double fertilization has taken place, the
single-celled zygote lies beneath the endosperm nucleus. The
endosperm nucleus divides to produce a mass of endosperm
tissue surrounding the embryo. The zygote also divides,
forming two parts: the upper part is the embryo, and the
lower part is the suspensor, which anchors the embryo and
transfers nutrients to it from the sporophyte plant. Soon the
cotyledons, or seed leaves, can be seen. At this point, the dicot embryo is heart shaped. Later, when it becomes torpedo
shaped, it is possible to distinguish the shoot apex and the
root apex. These contain apical meristems, the tissues that

endosperm
nucleus
zygote
a.
endosperm
b.

embryo

suspensor
epicotyl
c.
cotyledons
appearing
shoot
apex

seed
coat
f.

d.
hypocotyl
cotyledons

bending
cotyledons

root
apex

radicle

e.

Figure 10.13 Development of a dicot embryo.

a. The unicellular zygote lies beneath the endosperm nucleus. b, c. The endosperm is a mass of tissue surrounding the embryo. The embryo is
located above the suspensor. d. The embryo becomes heart shaped as the cotyledons begin to appear. e. There is progressively less endosperm
as the embryo differentiates and enlarges. As the cotyledons bend, the embryo takes on a torpedo shape. f. The embryo consists of the epicotyl
(represented here by the shoot apex), the hypocotyl, and the radicle which includes the root apex.

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Chapter 10

Plant Physiology and Reproduction

183

b.
one portion
of ovary

a.

Figure 10.14 Fruit diversity.

a. The almond fruit is eshy, with a single seed enclosed by a hard
covering. b. The tomato is derived from a compound ovary. c. Each
blackberry is from a ower with many ovaries.

Fruits and Seeds

In owering plants, seeds are enclosed within a fruit, which
develops from the ovary and at times, from other accessory
parts. Although peas, beans, tomatoes, and cucumbers are
commonly called vegetables, botanists categorize them as
fruits. A fruit is a mature ovary that usually contains seeds.
As a fruit develops from an ovary, the ovary wall thickens to become the pericarp. In eshy fruits, the pericarp is at
least somewhat eshy; peaches and plums are good examples of eshy fruits. In almonds, the eshy part of the pericarp is a husk removed before marketing. We crack the
remaining portion of the pericarp to obtain the seed (Fig.
10.14a). An apple develops from a compound ovary, but
much of the esh comes from the receptacle, which grows
around the ovary. Its more obvious that a tomato comes
from a compound ovary, because in cross section, you can
see several seed-lled cavities (Fig. 10.14b).
Dry fruits have dry pericarps. Legumes, such as peas
and beans, produce fruits that split along two sides, or
seams. Not all dry fruits split at maturity. The pericarp of a
grain is tightly fused to the seed and cannot be separated
from it. A corn kernel is a grain, as are the fruits of wheat,
rice, and barley plants.

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c.

many ovaries

Some fruits develop from several individual ovaries. A

blackberry is an aggregate fruit in which each portion is derived from a separate ovary of a single ower (Fig. 10.14c).
The strawberry is also an aggregate fruit, but each ovary becomes a one-seeded fruit called an achene. The esh of a
strawberry is from the receptacle. In contrast, a pineapple
comes from the fruit of many individual owers attached to
the same eshy stalk. As the ovaries mature, they fuse to
form a large, multiple fruit.
In owering plants, the seed develops from the
ovule and the fruit develops from the ovary.

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Plant Biology

Dispersal and Germination of Seeds

For plants to be widely distributed, their seeds have to be
dispersedthat is, distributed preferably long distances
from the parent plant. Following dispersal, the seeds germinate: they begin to grow so that a seedling appears.

Dispersal of Seeds
Plants have various means to ensure that dispersal takes
place. The hooks and spines of clover, bur, and cocklebur attach to the fur of animals and the clothing of humans. Birds
and mammals sometimes eat fruits, including the seeds,
which are then defecated (passed out of the digestive tract
with the feces) some distance from the parent plant. Squirrels and other animals gather seeds and fruits, which they
bury some distance away.
The fruit of the coconut palm, which can be dispersed
by ocean currents, may land many hundreds of kilometers
away from the parent plant. Some plants have fruits with
trapped air or seeds with inated sacs that help them oat in
water. Many seeds are dispersed by wind. Woolly hairs,
plumes, and wings are all adaptations for this type of

dispersal. The seeds of an orchid are so small and light that

they need no special adaptation to carry them far away. The
somewhat heavier dandelion fruit uses a tiny parachute
for dispersal. The winged fruit of a maple tree, which contains two seeds, has been known to travel up to 10 kilometers from its parent. A touch-me-not plant has seed pods
that swell as they mature. When the pods nally burst, the
ripe seeds are hurled out.
Animals, water, and wind help plants disperse their
seeds.

Germination of Seeds
Some seeds do not germinate until they have been dormant
for a period of time. For seeds, dormancy is the time during
which no growth occurs, even though conditions may be favorable for growth. In the temperate zone, seeds often have
to be exposed to a period of cold weather before dormancy
is broken. In deserts, germination does not occur until there
is adequate moisture. This requirement helps ensure that
seeds do not germinate until the most favorable growing

View 1

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seed coat
plumule
hypocotyl-radicle axis

;;;;;;;;;;
;;;;;;;;;;
;;;;;;;;;;
;;;;;;;;;;
cotyledon

epicotyl

a.

first true
leaves

cotyledon

withered
cotyledons

hypocotyl

seed coat

seed coat

hypocotyl

hypocotyl

primary root

secondary root

primary
root

b.

Figure 10.15 Common garden bean, a dicot.

a. Seed structure. b. Germination and development of the seedling. Notice that there are two cotyledons, and the leaves are net veined.

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Chapter 10

season has arrived. Germinationan event that takes

place if there is sufcient water, warmth, and oxygen to sustain growthrequires regulation, and both inhibitors and
stimulators are known to exist. It is known that eshy fruits
(e.g., apples, pears, oranges, and tomatoes) contain inhibitors so that germination does not occur until the seeds
are removed and washed. In contrast, stimulators are present in the seeds of some temperate zone woody plants. Mechanical action may also be required. Water, bacterial action,
and even re can act on the seed coat, allowing it to become
permeable to water. The uptake of water causes the seed
coat to burst.

Dicots Versus Monocots

As mentioned, the embryo of a dicot has two seed leaves,
called cotyledons. The cotyledons, which have absorbed the
endosperm, supply nutrients to the embryo and seedling,
and eventually shrivel and disappear. If the two cotyledons
of a bean seed are parted, you can see a rudimentary plant
(Fig. 10.15). The epicotyl bears young leaves and is called a
plumule. As the dicot seedling emerges from the soil, the

View 1

Plant Physiology and Reproduction

185

shoot is hook shaped to protect the delicate plumule. The

hypocotyl becomes part of the stem and the radicle develops
into the roots. When a seed germinates in darkness, it etiolatesthe stem is elongated, the roots and leaves are small,
and the plant lacks color and appears spindly. Phytochrome,
a pigment that is sensitive to red and far-red light (page 179),
regulates this response and induces normal growth once
proper lighting is available.
A corn plant is a monocot that contains a single cotyledon. Actually, the endosperm is the food-storage tissue in
monocots and the cotyledon does not have a storage role.
Corn kernels are actually fruits, and therefore the outer
covering is the pericarp (Fig. 10.16). The plumule and radicle are enclosed in protective sheaths called the coleoptile
and the coleorhiza, respectively. The plumule and the radicle burst through these coverings when germination
occurs.
Germination is a complex event regulated by many
factors. The embryo breaks out of the seed coat and
becomes a seedling with leaves, stem, and roots.

View 2

pericarp
endosperm
cotyledon

;;;;;;;;;;
;;;;;;;;;;
;;;;;;;;;;
;;;;;;;;;;
coleoptile
plumule
radicle

coleorhiza

a.

first leaf

coleoptile

coleoptile

prop roots
(adventitious roots)

coleorhiza

radicle

primary root

primary root

b.

Figure 10.16 Corn, a monocot.

a. Grain structure. b. Germination and development of the seedling. Notice that there is one cotyledon and the leaves are parallel veined.

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Plant Biology

10.5

Asexual Reproduction in
Flowering Plants
Because plants contain nondifferentiated meristem tissue, or
cells which can revert to meristematic activity (totipotency),
they routinely reproduce asexually by vegetative propagation.
In asexual reproduction there is only one parent, instead of
two as in sexual reproduction. Violets will grow from the
nodes of rhizomes (underground horizontal stems), and
complete strawberry plants will grow from the nodes of
stolons (aboveground horizontal stems). White potatoes are
actually portions of underground stems, and each eye is a
bud that will produce a new potato plant if it is planted with
a portion of the swollen tuber. Sweet potatoes are modied
roots of sweet potato plants that can be propagated by planting sections of the root. You may have noticed that the roots
of some fruit trees, such as cherry and apple trees, produce
suckers, small plants that can be used to grow new trees.

b.

Propagation of Plants in Tissue Culture

Hybridization, the crossing of different varieties of plants
or even species, is routinely done to produce plants with
desirable traits. Hybridization, followed by vegetative
propagation of the mature plants, will generate a large
number of identical plants with these traits. But tissue culture, growth of a tissue in an artificial liquid culture
medium, has led to micropropagation, a commercial method
of producing thousands, even millions, of identical
seedlings in a limited amount of space (Fig. 10.17). One favorite method to accomplish micropropagation is by meristem culture. If the correct proportions of auxin and
cytokinin are added to a liquid medium, many new shoots
will develop from a single callus, a group of nondifferentiated cells. When these are removed, more shoots form.
Since the shoots are genetically identical, the adult plants
that develop from them, called clonal plants, all have the
same traits.

c.

d.

a.

Figure 10.17 Micropropagation.

a. Sections of carrot root are cored, and thin slices are placed in a liquid nutrient medium. b. After a few days, the cells form a nondifferentiated
callus. c. After several weeks, the callus begins sprouting cloned carrot plants. d. Eventually the carrot plants can be moved from the culture
medium and potted.

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Chapter 10

The culturing of leaf, stem, or root tissues has led to a

technique called cell suspension culture. Rapidly growing calluses are cut into small pieces and shaken in a liquid nutrient
medium so that single cells or small clumps of cells break off
and form a suspension. These cells will produce the same
chemicals as the entire plant. For example, cell suspension
cultures of Cinchona ledgeriana produce quinine, and those of
Digitalis lanata produce digitoxin. Scientists envision that it
will also be possible to maintain cell suspension cultures in
bioreactors for the purpose of producing chemicals used in
the production of drugs, cosmetics, and agricultural chemicals. If so, it will no longer be necessary to farm plants simply
for the purpose of acquiring the chemicals they produce.

Genetic Engineering of Plants

Some plants will grow from protoplasts, naked cells that
will take up foreign DNA in tissue culture medium. This
form of genetic engineering has been most successful in dicots because monocots, such as corn and wheat, tend not to

e are in the midst of a pollination crisis due to a decline in the population

of honeybees and many other insects,
birds, and small mammals that transfer
pollen from stamen to stigma. Pollinator
populations have been decimated by pollution, pesticide use, and destruction or
fragmentation of natural areas. Belatedly,
we have come to realize that various types
of bees are responsible for pollinating such
cash crops as blueberries, cranberries, and
squash, and are partly responsible for pollinating apple, almond, and cherry trees.
Why are we so short-sighted when it
comes to protecting the environment and
living creatures like pollinators? Because

Plant Physiology and Reproduction

187

grow from protoplasts. But new methods have been developed to introduce DNA into plant cells that have a cell wall.
For example, a device called a particle gun can bombard
plant cells with DNA-coated microscopic metal particles.
Many types of plant cells in tissue culture, including corn
and wheat, have been genetically engineered using a particle gun. Later, adult plants are generated.
Various crops have been engineered to be resistant to viral infections, insect predation, and herbicides that are
judged to be environmentally safe. If crops are resistant to a
broad-spectrum herbicide and weeds are not, then the herbicide can be used to kill the weeds. The hope is that in the
future it will be possible to produce plants that have a higher
protein content and that require less water and fertilizer.
The ability of plants to reproduce asexually has led
to the generation of plants in tissue culture. This, in
turn, has promoted the genetic engineering of
plants.

pollinators are a resource held in common.

The term commons originally meant a
piece of land where all members of a village were allowed to graze their cattle. The
farmer who thought only of himself and
grazed more cattle than his neighbor was
better off. The difculty is, of course, that
eventually the resource is depleted and
everyone loses.
So, when a farmer or property owner
uses pesticides he is only thinking of his
eld or his lawn, and not the good of the
whole. The commons can only be protected if citizens have the foresight to enact
rules and regulations by which all abide.
DDT was outlawed in this country in part

because it led to the decline of birds of

prey. Similarly, we need legislation to protect pollinators from those factors that kill
them off. As a society, we need legislation
to protect pollinators because it helps protect the food supply for all of us.

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1.

Are you willing to stop using pesticides

that kill pollinators if it means your lawn
will suffer? Why or why not?
2. Are you willing to pressure your
representatives for legislation to protect
pollinators? Why or why not?
3. Are you willing to turn your lawn and
garden into a haven for pollinators? Why
or why not?

transported into phloem at a source, and water follows by osmosis. The

resulting increase in pressure creates a ow, which moves water and
sucrose to a sink.

Summarizing the Concepts

10.1 Water and Mineral Transport
Water transport in plants occurs within xylem. The cohesiontension model of xylem transport states that transpiration (evaporation
of water at stomates) creates tension, which pulls water upward in
xylem. This method works only because water molecules are cohesive.
Stomates open when guard cells take up potassium (K) ions and
water follows by osmosis. Stomates open because the entrance of water
causes the guard cells to buckle out.

10.2 Organic Nutrient Transport

Transport of organic nutrients in plants occurs within phloem. The
pressure-ow theory of phloem transport states that sugar is actively

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10.3 Plant Responses to Environmental Stimuli

Plant hormones control plant responses to environmental stimuli. Tropisms are growth responses toward or away from unidirectional stimuli. When a plant is exposed to light, auxin moves laterally from the
bright to the shady side of a stem. Thereafter, the cells on the shady side
elongate and the stem moves toward the light.
Plant hormones most likely control photoperiodism. Short-day
plants ower when the days are shorter (nights are longer) than a critical length, and long-day plants ower when the days are longer
(nights are shorter) than a critical length. Some plants are day-length
neutral. Phytochrome, a plant pigment that responds to daylight, is

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Plant Biology

believed to be a part of a biological clock system that in some unknown

way brings about owering.

10.4 Sexual Reproduction in Flowering Plants

Flowering plants have an alternation of generations life cycle, which
includes separate microgametophytes and megagametophytes. The
pollen grain, the microgametophyte, is produced within the stamens of
a ower. The megagametophyte is produced within the ovule of a
ower. Following pollination and fertilization, the ovule matures to become the seed and the ovary becomes the fruit. The enclosed seeds contain the embryo (hypocotyl, epicotyl, plumule, radicle) and stored food
(endosperm and/or cotyledons). When a seed germinates, the root appears below and the shoot appears above.

10.5 Asexual Reproduction in Flowering Plants

Many owering plants reproduce asexually, as when the nodes of
stems (either aboveground or underground) give rise to entire plants,
or when an isolated root produces new shoots. Micropropagation, the
production of clonal plants utilizing tissue culture, is now a commercial venture. The particle-gun technique allows foreign genes to be introduced into plant cells, which then develop into adult plants with
particular traits.

Studying the Concepts

1. Explain the cohesion-tension theory of water transport. 170
2. What mechanism controls the opening and closing of
stomates by guard cells? 172
3. Explain the pressure-ow theory of phloem
transport. 174
4. Name ve plant hormones and state their functions. 176
5. How is auxin believed to bring about elongation of cells so
that positive phototropism occurs. 176
6. With regard to photoperiodism, how was it shown that the
length of darkness, not the day length, actually controls
owering? 179
7. What is phytochrome, and what are two possible functions of
phytochrome in plants? 179
8. Describe how the megagametophyte (female gametophyte)
forms in owering plants. 180
9. Describe how the microgametophyte (male gametophyte)
forms in owering plants. 180
10. Contrast the monocot seed and its germination with the dicot
seed and its germination. 185
11. What is one favorite method of micropropagating
plants? 186

Testing Yourself
Choose the best answer for each question.
1. Stomates are usually open
a. at night, when the plant requires a supply of oxygen.
b. during the day, when the plant requires a supply of carbon
dioxide.
c. whenever there is excess water in the soil.
d. All of these are correct.

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2. What is the role of transpiration in water and mineral transport? Transpiration

a. causes water molecules to be cohesive and cling together.
b. occurs in the vessel elements and not the tracheids which
do have end walls.
c. is the force that causes sugar to be transported in xylem
sap.
d. creates the tension that draws a column of water up the
vessel elements.
e. Both a and d are correct.
3. The pressure-ow model of phloem transport states that
a. phloem sap always ows from the leaves to the root.
b. phloem sap always ows from the root to the leaves.
c. water ow brings sucrose from a source to a sink.
d. Both a and c are correct.
4. Root hairs do not play a role in
a. oxygen uptake.
b. mineral uptake.
c. water uptake.
d. carbon dioxide uptake.
5. After an agar block is placed on one side of an oat seedling, it
bends only if
a. unidirectional light is present.
b. the agar block contains auxin.
c. unidirectional light is present and the agar block contains
auxin.
d. the agar block is acidic.
e. the agar block is acidic and unidirectional light is present
and the agar block contains auxin.
6. Which of these is a correct statement?
a. Both stems and roots show positive gravitropism.
b. Both stems and roots show negative gravitropism.
c. Only stems show positive gravitropism.
d. Only roots show positive gravitropism.
7. Short-day plants
a. are the same as long-day plants.
b. are apt to ower in the fall.
c. do not have a critical photoperiod.
d. will not ower if a short-day is interrupted by bright light.
e. All of these are correct.
8. A plant requiring a dark period of at least 14 hours will
a. ower if a 14-hour night is interrupted by a ash of light.
b. not ower if a 14-hour night is interrupted by a ash of light.
c. not ower if the days are 14 hours long.
d. Both b and c are correct.
9. How is the megaspore in the plant life cycle similar to the
microspore?
Both
a. have the diploid number of chromosomes.
b. become an embryo sac.
c. become a gametophyte that produces a gamete.
d. are necessary to seed production.
e. Both c and d are correct.
10. Which of these is mismatched?
a. polar nucleiplumule
b. egg and spermzygote
c. ovuleseed
d. ovaryfruit

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10-21

Chapter 10

11. Label the arrows and dots in this diagram. The icon
represented by the dots plays what role in the opening of
stomates?

12. Label the following diagram of alternation of generations in

owering plants.
a.
(2n)

j.

i.
h.

diploid (2n)
b.

haploid (n)
c.

g.

e.

d.

f.
(n)

Thinking Scientically
1. A twig with leaves is placed in the top of an open tube that
contains water above mercury. 171
a. Atmospheric pressure alone is sufcient to raise mercury
only 76 cm (76 cm mercury = 10.5 m water). What is
atmospheric pressure, and of what signicance is this
nding for a tree that is 120 m high?
b. Why does the mercury rise higher than 76 cm when a twig
with leaves is placed in the top of the tube?
c. What does the experiment suggest about the ability of
transpiration to raise water to the top of tall trees?
2. Scientists wish to produce a strain of corn that is resistant to a
particular pesticide. 186187
a. Why would they choose to genetically engineer a diploid
corn cell instead of an egg or sperm?
b. How would they acquire diploid corn cells?
c. How would they acquire adult plants from the genetically
engineered corn cells?
d. If they want to sell genetically altered corn seeds from
these plants, how would they acquire seeds?

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Plant Physiology and Reproduction

189

Understanding the Terms

alternation of generations
180
anther 180
auxin 176
cohesion-tension theory 170
cotyledon 182
cytokinin 178
double fertilization 181
egg 181
embryo 181
embryo sac 180
endosperm 181
epicotyl 182
ethylene 178
lament 180
ower 180
fruit 183
gametophyte 180
generative cell 180
germination 185
gibberellin 178
guard cell 172
hormone 176
hypocotyl 182
megaspore 180
microspore 180
mineral 171
ovary 180
ovule 180
petal 180

phloem 174
photoperiodism 178
phototropism 177
phytochrome 179
pistil 180
plasmodesma 174
plumule 185
polar nuclei 180
pollen grain 180
pollen sac 180
pollination 180
pressure-ow theory 174
radicle 182
seed 181
seed coat 181
sepal 180
sink 174
source 174
spore 180
sporophyte 180
stamen 180
stigma 180
stomate 172
style 180
tissue culture 186
transpiration 170
tropism 176
tube cell 180
xylem 170
zygote 181

Match the terms to these denitions:

a.
In plants, a growth response toward or away from
a directional stimulus.
b.
In the life cycle of a plant, the haploid generation
that produces gametes.
c.
Explanation for phloem transport; osmotic pressure following active transport of sugar into phloem brings
about a ow of sap from a source to a sink.
d.
In seed plants, a small spore that develops into
the sperm-producing microgametophyte; pollen grain.
e.
Plant pigment that induces a photoperiodic
response in plants.

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Part 2

10-22

Plant Biology

Using Technology
Your study of plant physiology and reproduction is supported by
these available technologies:

Essential Study Partner CD-ROM

Plants Translocation, Plant Responses,
Reproduction
Visit the Mader web site for related ESP activities.

Exploring the Internet

The Mader Home Page provides resources and tools as
you study this chapter.

http://www.mhhe.com/biosci/genbio/mader

Further Readings for Part 2

Arakawa, T., and Langridge W. R. H. May 1998. Plants are not just
passive creatures! Nature Medicine 4(5):550. Plants are being
used as bioreactors to produce foreign proteins for human
immunity.
Balick, M. J., and Cox, P. A. 1996. Plants, people, and culture: The
science of ethnobotany. New York: Scientic American Library. This
interesting, well-illustrated book discusses the medicinal and
cultural uses of plants, and the importance of rain forest
conservation.
Busch, R. October/November 1998. The value of autumn leaves.
National Wildlife 36(6):32. Article examines the physiology
behind autumn leaf colors.
Canny, M. J. March/April 1998. Transporting water in plants.
American Scientist 86(2):152. The cohesion-tension theory of
water transport is discussed in detail.
Chrispeels, M., and Sadava, D. 1994. Plants, genes and agriculture.
Boston: Jones and Bartlett Publishers. Teaches plant biology in
an agricultural context.
Gibson, A. C. November 1998. Photosynthetic organs of desert
plants. Bioscience 48(11):911. The specialized leaf and stem
adaptations in desert plants may be to maximize photosynthetic
rates and energy production, rather than to conserve water.
Glenn, E. P., et al. August 1998. Irrigating crops with seawater.
Scientic American 279(2):76. Certain useful salt-tolerant plants
can ourish on seawater irrigation.
Lee, D. W. January/February 1997. Iridescent blue plants.
American Scientist 85(1):56. Some tropical plants produce a blue
color, as do some insects, by using layered lters.

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Levetin, E., and McMahon, K. 1996. Plants and society. Dubuque,

Iowa: Wm. C. Brown Publishers. Basic botany and the impact of
plants on society are topics covered in this introductory text.
Luoma, J. R. March 1997. The magic of paper. National Geographic
191(3):88. The paper-making process is discussed in this article.
Martinez del Rio, C. February 1996. Murder by mistletoe. Natural
History 105(2):85. The quintral is a mistletoe that parasitizes
cacti.
Mauseth, J. 1995. Botany: An introduction to plant biology. 2d ed.
Philadelphia: Saunders College Publishing. Emphasizes
evolution and diversity in botany and general principles of
plant physiology and anatomy.
Milot, C. 17 July 1998. Plant biology in the genoma era. Science
281(5375):331. Genomic data can be used to make plants that
produce more vitamins and minerals, and can help explain
physiological processes, such as owering.
Moore, R., and Clark, W. D., et al. 1998. Botany. 2d ed. Dubuque,
Iowa: WCB/McGraw-Hill. This introductory botany text
stresses the process of science while presenting the evolution,
anatomy, and physiology of plants.
Niklas, K. J. February 1996. How to build a tree. Natural History
105(2):48. Article discusses the properties of wood.
Northington, D., and Goodin, J. R. 1996. The botanical world. 2d ed.
St. Louis: Times-Mirror/Mosby College Publishing. This is an
account of plant interactions and basic physiology.
Pitelka, L. F., et al. September/October 1997. Plant migration and
climate change. American Scientist 85(5):464. A relationship
between plant migration and climate change, as evidenced by
the fossil record and computer models, is discussed.
Redington, C. 1994. Plants in wetlands. Dubuque, Iowa:
Kendall/Hunt Publishing Company. Explains how specic
plants interact within the wetlands ecosystem.
Schmiedeskamp, M. December 1997. Pollution-purging poplars.
Scientic American 277(6):46. Hybrid poplar trees may be used to
break down toxic organic compounds in the soil.
Seymour, R. S. March 1997. Plants that warm themselves. Scientic
American 276(3):104. Some plants generate heat to keep
blossoms at a constant temperature.
Stern, K. 1997. Introductory plant biology. 7th ed. Dubuque, Iowa:
Wm. C. Brown Publishers. Presents basic botany in a clear,
informative manner.
Stryer, L. 1995. Biochemistry. 5th ed. New York: W. H. Freeman and
Company. Chapter 22 of this text presents an advanced but
understandable treatment of photosynthesis.
Sze, P. 1998. A biology of the algae. 3d ed. Dubuque, Iowa:
WCB/McGraw-Hill. A text that introduces algae morphology,
evolution, and ecology to the botany major.
Walsh, R. May 1997. The chocolate bug. Natural History 106(4):54.
The cacao trees survival depends upon a tiny insect, which
dictates where cacao will grow.

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