Are Plants Computers?

Figure 35.1 Computer art?

he object in Figure 35.1 is not the creation of a computer genius with a flair
for the artistic. It is a head of romanesco, an edible relative of broccoli. Romanescos mesmerizing beauty is attributable to the fact that each of its smaller buds
resembles in miniature the entire vegetable (shown below). (Mathematicians refer
to such repetitive patterns as fractals.) If romanesco looks as if it were generated
by a computer, its because its growth pattern follows a repetitive sequence of
instructions. As in most plants, the growing shoot tips lay down a pattern of stem
. . . leaf . . . bud, over and over again. These repetitive developmental patterns are
genetically determined and subject to natural selection. For example, a mutation
that shortens the stem segments between leaves will generate a bushier plant. If
this altered architecture enhances the plants ability to access resources such as
light and, by doing so, to produce more offspring, then this trait will occur more
frequently in later generationsthe population will have evolved.
Romanesco is unusual in adhering so rigidly to its basic body organization. Most
plants show much greater diversity in their individual forms because the growth
of most plants, much more than in animals, is affected by local environmental
conditions. All adult lions, for example, have four legs and are of roughly the same
size, but oak trees vary in the number and arrangement of their branches. This is

because plants respond to challenges and opportunities in

their local environment by altering their growth. (In contrast,
animals typically respond by movement.) Illumination of a
plant from the side, for example, creates asymmetries in its
basic body plan. Branches grow more quickly from the illuminated side of a shoot than from the shaded side, an architectural change of obvious benefit for photosynthesis. The highly
adaptive development of plants is critical in facilitating their
acquisition of resources from their local environments.

All plants are able to

harvest diffuse resources
and concentrate them
in cells and tissues,
but their forms and
strategies are diverse.
These baobab trees in
Madagascar may live
to be hundreds of years
old despite drought
conditions, in part by
storing water in their
enormous trunks.

hotosynthetic plants carry out the most remarkable biochemistry of any terrestrial organisms.
Using the energy in sunlight and the simplest of starting materialscarbon dioxide, water,
and ions containing nitrogen, phosphorus, potassium, and other key atomsplants synthesize thousands of different carbohydrates, proteins, nucleic acids, and lipids. They use these compounds to build bodies that may live for thousands of years.
This feat is even more impressive when you consider that the simple starting materials that plants
need to grow are tiny and diffusecarbon dioxide molecules, water molecules, nitrate ions, and
other resources are usually found at low concentrations over a large area. To gather the raw materials required for their sophisticated biosynthetic machinery, a plants roots and shoots grow outward,
extending the individual into the soil and atmosphere.

In essence, a plants body harvests diffuse resources and concentrates them in cells and tissues. The structure of its body is dynamic, because most plants exhibit indeterminate growth; that is,
they grow throughout their lives. A 4750-year-old bristlecone pine
has roots and shoots that are still growing. In response to favorable
conditions, a plant sends shoots and roots in the most promising
directions, seeking light and the simple compounds it requires.
The contrast between the plant and animal way of life is striking. Most animals move around, eat concentrated sources of
food, and avoid stressful conditions. But plants stay in one place,
extend their roots and shoots to harvest diffuse resources, make
their own food, and cope with stress where they stand.

What Is the Basic Body Plan of Plants?

Plants live by harvesting energy from sunlight and by collecting water and mineral nutrients from the atmosphere
and the soil. Because these resources are sometimes limited,
plants must collect them from large areas, both above and
below ground. The plant is further challenged by its inability to move; a plant cannot, for example, relocate from a
dry, shady location to one that is wet and sunny.
The plant body plan allows plants to respond to these
challenges:

Stems, leaves, and roots enable a plant anchored to one

spot to capture scarce resources effectively, both above
and below the ground.

Plants can grow throughout their lifetimes, enabling

them to respond to environmental cues. A plant can
redirect its growth to exploit opportunities in its immediate environment; for example, it can extend its roots
toward a water supply.
Plant organs are organized into two systems :

The root system anchors the plant in place, absorbs water

and dissolved minerals, and stores the products of photosynthesis from the shoot system. The extreme branching
of plant roots and their high surface area-to-volume ratios
allow them to absorb water and mineral nutrients from the
soil efficiently.

The shoot system of a plant consists of the stems, leaves,

and flowers. Broadly speaking, the leaves are the chief
organs of photosynthesis. The stems hold and display the
leaves to the sun and provide connections for the transport
of materials between roots and leaves.

Vascular plants, with few exceptions, rely on both systems

for survival. Roots are almost never photosynthetic; they
starve unless photo-synthates, the sugars and the other
carbohydrates produced during photosynthesis, are
imported from the shoot system. Conversely, the shoot
system depends on the water and minerals that roots absorb
from the soil.min
The iterative (repeating) unit of the vegetative shoot
consists of the internode, node, leaf, and axillary bud, but not
reproductive structures. An axillary bud is a lateral shoot
apex that allows the plant to branch or replace the main
shoot if it is eaten by an herbivore. A vegetative axillary bud
has the capacity to reiterate the development of the primary
shoot. When the plant has shifted to the reproductive phase
of development, these axillary buds may produce flowers or
floral shoots.

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Shoots and roots are composed of repeating modules called

phytomers. Each phytomer in the shoot consists of a node carrying one or more leaves; an internode, which is the interval of
stem between two nodes; and one or more axillary buds, each

of which forms in the angle (axil) where a leaf meets the stem.
A bud is an undeveloped shoot that can develop further to produce another leaf, a phytomer, a flower, or a flowering stem.
The axillary buds (also called lateral buds) are distinguished
from the bud at the end of a stem or branch, which is called a
terminal bud. If it becomes active, an axillary bud can develop
into a new branch, or an extension of the shoot system. The arrangement of leaves along the stem (called the phyllotaxy) is
characteristic of the plant species.
Plant roots also have a modular construction. In the roots,
each phytomer consists of a root segment between two
branches.

Most angiosperms are either monocots

or eudicots
As we saw in Section 29.3, most angiosperms belong to one of
two major clades. Monocots are generally narrow-leaved flowering plants such as grasses, lilies, orchids, and palms. Eudicots are broad-leaved flowering plants such as soybeans, roses,
sunflowers, and maples. These two clades, which account for
97 percent of flowering plant species, differ in several basic
characteristics:

Monocots have one cotyledon (leaf in the embryo),

whereas eudicots have two.

In eudicots, the vascular bundles in the stem are arranged

in concentric circles; in monocots they are scattered.

In monocots, the major leaf veins are usually parallel; in

eudicots they are reticulate, meaning they form a network.

Eudicots usually have taproot systems; monocots have fibrous root systems.

Monocot flowers have parts (petals and sepals) that occur

in threes; eudicots have floral parts that occur in fours or
fives.

Monocot pollen grains each have one furrow or pore; eudicot pollen grains have three.

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Figure 36.1 Diagram of a plant body. Branching root and

shoot systems create the plants architecture. Each root and shoot
has an apex that extends growth. Leaves are initiated at the nodes of
the shoot, which also contain axillary buds that can remain
dormant, grow to form lateral branches, or make flowers. A leaf
can be a simple blade or consist of multiple parts as shown here.
Roots, shoots, and leaves are all connected with vascular
(conducting) tissue.

Shoot apex

Flower

Stipule
Tendril
Axillary bud

Shoot

Vein
Leaflet Leaf

Internode

Blade
Petiole

Node
Vascular system

Primary root

Lateral root
Root

Root apex

34.1 Vegetative Organs and Systems The basic plant body

plan, with root and shoot systems, and the principal vegetative
organs are similar in eudicots and monocots, although there are
also some differences between the two clades.
Monocot

Eudicot
Terminal bud
Axillary bud
Phytomer
Node
The shoot system
consists of stems
and leaves, in
which photosynthesis
takes place.

Internode
Node
Internode
Petiole
Branch

Leaf
Blade
Stem

The root system

anchors the plant
and provides water
and nutrients for the
shoot system.

Roots

Roots and shoots are composed

of three types of tissues
Roots, shoots, and leaves all contain three basic types of tissues:
dermal, ground, and vascular tissue. Because each of these tissues extend through the root and shoot systems, they are called
tissue systems.
Plant cells contribute to three tissue systems. Dermal
tissue, primarily epidermis, is one cell layer thick in most
plants, and it forms an outer protective covering for the plant.
Ground tissue cells function in storage, photosynthesis, and
secretion, in addition to forming fibers that support and
protect plants. Vascular tissue conducts fluids and dissolved
substances throughout the plant body. Each of these tissues
and their many functions are described in more detail in
later sections.

key point
The tissue systems are continuous throughout the plant. For example, the vascular tissue system in a leaf is continuous
with the vascular tissue system in the stem to which it is attached.

Dermal tissue system

Vascular tissue system
Ground tissue system

(a) Leaf
Dermal tissue system
Vascular tissue system
Ground tissue system

(b) Stem
Dermal tissue system
Vascular tissue system
Ground tissue system

(c) Root

Figure 33-2 Animation The three tissue systems in the plant body
This figure shows the distribution of the ground tissue system, vascular tissue system, and dermal tissue
system in a herbaceous eudicot such as Arabidopsis.
Cengage Learning

predict What do you think would

happen if the vascular tissue system
were to become discontinuous in the
stem but the dermal and ground tissue systems were to remain intact?

Meristems elaborate the body plan throughout the plants life

When a seed sprouts, only a tiny portion of the adult plant exists. Although embryo cells can undergo division and differentiation to form many cell types, the fate of most adult cells is
more restricted. Further development of the plant body depends on the activities of meristems, specialized cells found in
shoot and root apices, as well as other parts of the plant.

Overview of meristems
Meristems are clumps of small cells with dense cytoplasm and
proportionately large nuclei that act as stem cells do in animals. That is, one cell divides to give rise to two cells, of which
one remains meristematic, while the other undergoes differentiation and contributes to the plant body (figure 36.3). In this
way, the population of meristem cells is continually renewed.
Molecular genetic evidence supports the hypothesis that animal stem cells and plant meristem cells may also share some
common pathways of gene expression. Extension of both root
and shoot takes place as a result of repeated cell divisions and
subsequent elongation of the cells produced by the apical
meristems. In some vascular plants, including shrubs and
most trees, lateral meristems produce an increase in root and
shoot diameter.

cell division. Plant meristems

consist of cells that divide to
give rise to a differentiating
daughter cell and a cell that
persists as a meristem cell.

Meristem cell
Cell division

Many herbaceous plants (that is, plants with fleshy, not woody
stems) exhibit only primary growth, but others also exhibit
secondary growth, which may result in a substantial increase
of diameter. Secondary growth is accomplished by the lateral
meristemsperipheral cylinders of meristematic tissue within
the stems and roots that increase the girth (diameter) of gymnosperms and most angiosperms. Lateral meristems form from
ground tissue that is derived from apical meristems. Monocots
are the major exception (figure 36.5).
Although secondary growth increases girth in many
nonwoody plants, its effects are most dramatic in woody

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Meristem cell

Differentiated cell

Cell division

Meristem cell

Differentiated cell

Cell division

Differentiated cell

Apical meristems
Apical meristems are located at the tips of stems and roots
(figure 36.4). During periods of growth, the cells of apical meristems divide and continually add more cells at the tips. Tissues
derived from apical meristems are called primary tissues, and
the extension of the root and stem forms what is known as the
primary plant body. The primary plant body comprises the
young, soft shoots and roots of a tree or shrub, or the entire
plant body in some plants.

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moves through the soil. In contrast, leaf primordia shelter the

growing shoot apical meristem, which is particularly susceptible to desiccation because of its exposure to air and sun.
The apical meristem gives rise to the three tissue systems
by first initiating primary meristems. The three primary meristems are the protoderm, which forms the epidermis; the
procambium, which produces primary vascular tissues (primary xylem for water transport and primary phloem for nutrient transport); and the ground meristem, which differentiates
further into ground tissue. In some plants, such as horsetails
and corn, intercalary meristems arise in stem internodes
(spaces between leaf attachments), adding to the internode
lengths. If you walk through a cornfield on a quiet summer
night when the corn is about knee high, you may hear a soft
popping sound. This sound is caused by the rapid growth of the
intercalary meristems. The amount of stem elongation that occurs in a very short time is quite surprising.

Lateral meristems

Figure 36.3 Meristem

Meristem cell

Both root and shoot apical meristems are composed of

delicate cells that need protection (see figure 36.4). The root
apical meristem is protected by the root cap, the anatomy of
which is described later on. Root cap cells are produced by the
root meristem and are sloughed off and replaced as the root

plants, which have two lateral meristems. Within the

bark of a woody stem is the cork cambiuma lateral meristem that contributes to the outer bark
of the tree. Just beneath the bark is the vascular
cambiuma lateral meristem that produces
secondary vascular tissue. The vascular cambium
forms between the xylem and phloem in vascular
bundles, adding secondary vascular tissue to both
of its sides.
Secondary xylem is the main component of wood. Secondary phloem is very close to the outer surface of a woody
stem. Removing the bark of a tree damages the phloem and
may eventually kill the tree. Tissues formed from lateral meristems, which comprise most of the trunk, branches, and older
roots of trees and shrubs, are known as secondary tissues and
are collectively called the secondary plant body.

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Young leaf
primordium
Shoot apical
meristem
Older leaf
primordium
Lateral bud
primordium
100 m
dermal tissue
ground tissue
vascular tissue

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Root apical
meristem
Root cap
400 m

Figure 36.4 Apical

meristems. Shoot and root apical
meristems extend the plant body above
and below ground. Leaf primordia protect
the fragile shoot meristem, while the root
meristem produces a protective root cap in addition
to new root tissue.

Learning Outcomes Review 36.1

The root system anchors plants and absorbs water and nutrients, whereas
the shoot system, consisting of stems, leaves, and owers carries out
photosynthesis and sexual reproduction. The three general types of tissue
in both roots and shoots are dermal, ground, and vascular tissue. Primary
growth is produced by apical meristems at the tips of roots and shoots;
secondary growth is produced by lateral meristems that are peripheral and
increase girth.

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Why are both primary and secondary growth necessary

in a woody plant?

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The Importance of Surface Area/Volume

Relationships
Before exploring the nature of root and shoot systems in more
detail, its important to recognize a key structural relationship
that is critical to their function.
Root and shoot systems both function in absorptionof water and key ions, or of light, respectively. Absorption takes place
across a surface. But the cells that use the absorbed molecules
and light occupy a volume. Thus, a plant body is more efficient as
an absorption-and-synthesis machine when it has a large surface
area relative to its volume.
FIGURE 37.2 illustrates this point. In this example, the cells in a
plant are represented by cubes; the side of each cell is 50 m long.
Thus, each face of a cell has a surface area of 50 * 50 = 2500 m2;
each cell has a volume of 50 * 50 * 50 = 125,000 m3. Follow the
calculations in the figure and note that:

If 64 cells are arranged in a cube, the surface area/volume relationship is 0.0300/m.

If 64 cells are arranged in a long tube, the surface area/volume

relationship is 0.0425/m.

If 64 cells are arranged in a flat sheet, the surface area/volume

relationship is 0.0525/m.

This simple exercise has an important punch line: Tubes and

sheets have much more surface area relative to their volume than
cubes. Its no surprise, then, that the absorptive regions of a root
system are tubelike, and the absorptive regions of a shoot system are the flattened structures called leaves. Storage tissues such
as tubers and seeds have a low surface-area-to-volume ratio because they are not involved in absorption.

Thick structure (64 cells)

50 m

Tubelike structure
(64 cells)

Flattened structure
(64 cells)

Surface area = 240,000 m2

(96 cell surfaces x 2500 m2/cell surface)

Surface area = 340,000 m2

(136 cell surfaces x 2500 m2/cell surface)

Surface area = 420,000 m2

(168 cell surfaces x 2500 m2/cell surface)

Volume = 8,000,000 m3
(64 cells x 125,000 m3/cell)

Volume = 8,000,000 m3
(64 cells x 125,000 m3/cell)

Volume = 8,000,000 m3
(64 cells x 125,000 m3/cell)

Surface area/volume = 0.0300/m

Surface area/volume = 0.0425/m

Surface area/volume = 0.0525/m

FIGURE 37.2 The Morphology of Roots and Leaves Gives Them a High Surface-Area-to-Volume Ratio. In
this example, the thick structure represents a tree trunk or potato-like storage organ; the tubelike structure
represents a root; the flattened structure represents a leaf. Note that each schematic structure has the same
number of cells and the same total volumebut a very different surface area.