Chapter 7

The Root System

THE FUNCTIONS OF ROOTS

TYPES OF ROOT SYSTEMS

Differences in the Design of Root systems

Help Plants Compete for Water and
Minerals
Plants Have Different Types of Roots

THE DEVELOPMENT OF ROOTS

The Root Tip Is Organized into Regions

and Is Protected by a Root Cap
The Root Apical Meristem Forms Three
Primary Meristems

THE STRUCTURE OF ROOTS

The Epidermis, Cortex, and Vascular

Cylinder are Composed of Specialized
Tissues
Lateral Roots Are Initiated in the Pericycle
The Vascular Cambium and the Cork
Cambium Partially Form From the Pericycle
Some Roots Have Special Functions

SUMMARY

PLANTS, PEOPLE, AND THE

ENVIRONMENT: The climbing fig and its
adhesive pads.

PLANTS, PEOPLE, AND THE

ENVIRONMENT: Myths and Popular Uses
of Roots.

IN DEPTH: How Do Roots Advertise their

Presence?

Copyright Terence M. Murphy, Thomas L. Rost, Michael G. Barbour 2015. All federal and state copyrights reserved.

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KEY CONCEPTS

1. The principal functions of roots are absorption of water and nutrients, conduction of
absorbed materials into the plant body, and anchorage of the plant in the soil. Many
roots have relationships with bacteria and fungi in the rhizosphere (soil zone near the
root).

2 . The root is initiated in the embryo as the radicle. It penetrates into the soil and
forms branches.

3 . The root tip is composed of the root cap, the root apical meristem, the region of
cell elongation, and the region of cell maturation.

4 . Roots are composed of the following tissues: epidermis, cortex, endodermis,

pericycle, xylem, and phloem.

5. The endodermis regulates ion movement into the xylem. The Casparian strip
embedded in the cell wall inhibits mineral movement through the wall. The pericycle
is the site of lateral root initiation and contributes to vascular cambium and cork
cambium formation.

7.1 TH E FUNCTIONS OF ROOTS

Although plant biologists have studied plants for hundreds of years, they have largely
ignored the root system and its functions. This is probably because roots are
underground, where they cannot be seen. Though they are hidden from view, roots
play a critical role in the everyday activities of plants. The main functions of roots are
anchorage of the plant body in the soil (or to a surface, in the case of some vines);
absorption of water and minerals from the soil; storage of foods; and conduction of
food and water from the soil and from storage reserves into the shoot. Other root
functions are also described in this chapter.
The root system becomes more complex as the plant grows from a single root
in a young seedling to a massive system of branched roots, often weighing tons in
large trees. During all stages in a plant's growth cycle, there is a balance between the
shoot system and the root system (Fig. 7.1). The root system must be able to supply
the shoot with sufficient water and mineral nutrients, and the shoot system must
manufacture enough food to maintain the root system.

The contact zone between the root surface and the soil is called the rhizosphere
(Fig. 7.2). This region, only a few millimeters thick, is a very interesting and unique
zone. Its complex chemistry includes organic material, gases, and nutrients from the
roots. Consequently, the bacteria and fungi near roots are often richer and more
diverse than in soil farther away. Soil fungi and bacteria form important symbiotic
(mutually beneficial) relationships with roots. These will also be discussed later in the
chapter.

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Every time you try to pull weeds you are reminded of the anchorage function of roots.
Plants with unusual roots provide anchorage in atypical ways. In ivy (Hedera helix)
and the climbing fig (Ficus pumila; see sidebar: The Climbing Fig And Its Adhesive Pads)
clusters of roots develop from the stems to form an unusual adhesive pad allowing

Figure 7.1. Root and shoot

systems of a tree at least 10 years
old. Note that the majority of the
roots lie within the upper 1 m of
soil and that the total volume of
roots is equal to or greater than
the shoot branches.

Figure 7.2. The soil area

immediately around a root is
called the rhizosphere. This area,
shown near the root tip, is rich in
soil microorganisms, bacteria
and fungi, and in nutrients from
the root body and from
sloughed-off root cap cells.

these vines to cling to vertical surfaces. Parasitic plants like dodder (Cuscuta sp.) sink
specialized roots into their host and then tap into its water and nutrient supply.
All plants need water. In herbaceous (non-woody) plants, water accounts for about 90% of
the plant's weight. Water is needed for all root processes and for every metabolic reaction.
Plants also need the dissolved salts and minerals, such as potassium, sulfur, phosphorus,
calcium, and magnesium, contained in the soil water. For these reasons, plants have
developed an elaborate system in roots for the absorption and conduction of water.
All roots, even slender ones with a primary function of absorption, may temporarily store
small amounts of food. For example, when sugar moves into roots more rapidly than can be
used by growing cells, it may be converted to starch and stored. During slow growth periods,
rather large quantities of starch are stored in woody roots of orchard trees. This food

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constitutes a reserve that is used when flowering and active growth are resumed in spring.
Carrots (Dacus carota), and beets (Beta vulgarus) are common root crops.

7.2 TYPES OF ROOT SYSTEMS

There are two basic types of root systems: fibrous and tap. They are distinguished by the way
they develop and by their appearance. Many grasses and small garden plants, for instance,
when pulled up bring with them a massive clump of soil. This happens because the fibrous
root system of these plants consists of several main roots that branch to form a dense mass
(Fig. 7.3a, b). A typical annual grass such as corn ( Zea mays ) or rye (Elymus cereale) will build
an immense fibrous root system in one
growing season. A single rye plant 50 a b c
cm (20 in) tall, with 80 tillers (shoot
branches), may have a root system
surface area of about 210 m2 (1890 ft2),
compared to only about 5 m2 (45 ft2) for
its aboveground shoot system.
Plants with a large storage root,
like carrot (Daucus carota), have a tap
root system, consisting of one main
root from which lateral roots branch
(Fig. 7.3c). Some desert plants have a
rapidly growing tap root system that
enables them to penetrate the soil
quickly to reach deep sources of water.

Differences in the Design of Root

Systems Help Plants Compete for
Water and Minerals
Figure 7.3. Fibrous and tap root systems. (a)
Shallow, spreading, fibrous root system. (b)
Plants that grow in close proximity
Fibrous root system penetrating the soil evenly
compete for water, mineral nutrients,
up to 1 meter in this example. (c) Tap root
and light energy. As will be discussed
system, in which main primary root penetrates
in Chapter 26, plants reduce the
soil 2 meters or more.
effects of competition by utilizing
different parts of the environment,
including the soil. This is one reason
why root systems of different species also occupy different depths in the soil.
When plants growing in the prairie in the middle United States are carefully
excavated, three general categories of roots are evident, based on how deep the roots grow
(Fig. 7.3). Some grassland species, such as blue grama (Bouteloua gracilis), pos sess a very
shallow root system, most of the roots being within the top 15 cm (6 in) of soil. Other
species, such as buffalo grass (Buchloe dactyloides), have evenly distributed roots as deep
as 1.5 m (60 in). Still others, such as locoweed (Crotalaria sagittalis), have a tap root

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system, which lacks width but runs deep. By using different depths of the soil, these plants
reduce competition for moisture and dissolved minerals.

Plants Have Different Types of Roots

When seeds germinate, the embryonic root (the radicle--see Chapter 14) extends by the
division and elongation of cells to form the primary root (Fig. 7.4a). Tap root systems
develop from one primary root, which then forms lateral roots. Further branching results
in successive orders of roots. Fibrous root systems develop in a slightly different way. The
embryos of most grasses have a single radicle, but in addition several other embryonic
roots form just above the radicle; these are called seminal roots. The seminal roots emerge
soon after the radicle, and all of these roots branch, making a fibrous root system (Fig.
7.4b).
Roots called adventitious roots originate on leaves and stems. There are several
common examples of adventitious roots. In a young corn plant, soon after germination

a b
Figure 7.4. (a) Seedling of pea ( Pisum
sativum) with a tap root and several
lateral roots. (b) Seedling of wheat
(Triticum aestivum). Seminal roots
emerge from the hypocotyl (these are
adventitious roots since they do not
emerge from another root) and create the
fibrous root system.

prop roots develop on the stem just above the soil (Fig. 7.5a). Prop roots absorb water and
minerals, but they also support the plant in the soil.
Banyan (Ficus bengalensis) trees grow in the salty mud of tropical lagoons and tidal
marshes. Branches of these trees form adventitious roots--also called aerial roots, because
they are exposed to air. These extend down from branches into the soil, where they
enlarge and actually hold up the large branches (Fig. 7.5b). These roots absorb water and
nutrients, but their most important function is to prop up the stem. In India, merchants
once held open-air bazaars among the prop roots and expansive branches of the banyan.
Mangrove trees (Rhizophora mangle ) are native to low tidal shores and marshes in
tropical and subtropical regions. In the mangrove (Fig. 7.5c) small adventitious roots called

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 pneumatophores, stick up from the mud. These roots absorb oxygen and increase its
availability to the submerged roots. There are many other examples of adventitious roots.
Pieces of stem, such as a cane from a blackberry plant or a branch of willow, can be
induced to make roots from their cut ends simply by placing them in moist soil. Leaves
from Begonia and several other plants also can be rooted, simply by soaking them in water.
Many commercially important ornamentals are reproduced by root propagation from the
leaves or stems.

7.3 THE DEVELOPMENT OF ROOTS

The tips of functional roots are thin (Fig. 7.4a) and usually white. If you were to dig
up the roots of a big tree, you would see many very large roots, each of which could
be followed through its branches to a thin, white tip. These tiny root tips are
important parts of the root system because it is here, in just a few millimeters,
where many of the important functions of roots take place.

a b c

Figure 7.5. Types of adventitious roots. (a) Corn (Zea mays) stem showing prop roots, which
emerge from the stem just above the soil. These roots help support the shoot system. (b) Extensive
adventitious root system of mangrove (Rizophora mangle) growing in the tidal zone of Australia's
tropical coast. Note the many air roots (pneumatophores) sticking into the mud. (c) Banyan (Ficus
bengalensis) tree with an extensive aerial root system.

The Root Tip Is Organized into Regions and Protected by a Root Cap

In a longitudinal section of a root tip viewed through a microscope, it is apparent that

the cells are organized into three regions: the root apical meristem, the region of
elongation, and the region of maturation (Fig. 7.6a). The developmental events that
take place in the cells of each region are somewhat specific, but the regions do
overlap (Fig. 7.6b).
The root cap at the tip of the root apex protects the root apical meristem
(RAM), a group of small, regularly shaped cells, most of which are dividing. These

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 cells are organized into two different patterns (Figure 7.7a, b). A small, centrally
located part of the RAM is called the quiescent center (QC) (Fig. 7.8) because its cells
divide at an extremely slow rate. The function of the QC is not exactly known, but it
seems to be activated during times of acute stress. It may be a site for the synthesis of
plant hormones important for controlling root development.
Figure 7.6. Regions of the
root tip. (left) Radish
seedling ( Raphanus sativus)
showing the root apical
meristem, region of
elongation, and region of
maturation containing root
hairs. (right) Median
longitudinal section of root
tip. Procambium cells tend
to stop dividing before
ground meristem cells,
protoderm and pericycle
cells continue divided
farther back in the root. All
cells stop elongating at
approximately the same
point. Primary tissue cell
maturation also may occur
at different positions.

Figure 7.7. The patterns of

RAM divisions. (a)
Longitudinal section of the
root tip of flax (Linum
grandiflorum). Notice that all
cell files connect directly to
specific tiers of cells just
above the root cap. X200. (b)
Longitudinal section of onion
(Allium cepa) root tip. In this
type of root apex the cell files
terminate at a group of cells
without any apparent
organization. X81.

a b

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 Figure 7.9. (above) Corn (Zea mays) root
cap with sloughed root cap cells in the
soil, X60.

Figure 7.8. (left) Three-dimensional

diagram of a root in longitudinal view to
show the relative positions of the primary
meristems and primary tissues. The
quiescent center is a small group of cells
that are metabolically quiescent.

Cells just apical to the QC divide and produce cells to form the root cap (Fig. 7.7, 7.8),
the thimble-shaped layer of cells that protect the RAM as the root elongates and pushes
through the soil. The root cap is also the site of gravity perception, which controls the
direction of root growth . Root cap cells are constantly being sloughed off at the very tip, but
new cells are added by the apical meristem. The sloughed-off cells can remain alive in the soil
for a time, where they provide nutrients for soil bacteria and fungi in the rhizosphere (Figs.
7.2, 7.9).
Just basal to the meristem region, toward the body of the root, is the region of
elongation. Careful examination of a longitudinal section of a root shows that the boundary
where cells stop dividing and start elongating is different for each tissue (see Fig. 7.6b). The
region of maturation is the site of root hair formation and the maturation of other cell

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types. The precise position of cell maturation in different cell files is variable; cells in some
files mature close to the tip, and others mature farther back (Fig. 7.6b).

The RAM Forms Three Primary Meristems

As mentioned in Chapter 4, cells change their structure according to their position, and in
roots the process of change begins in the root apical meristem. The RAM differentiates into
three primary meristems: the protoderm, ground meristem, and procambium (see Fig.
7.8). These then go on to become the primary tissues of the root, as described in the
following pages and as summarized in Table 7.1.

7.4 THE STRUCTURE OF ROOTS

The Epidermis, Cortex, and Vascular Cylinder Are Composed of Specialized Tissues

EPIDERMIS Protoderm cells differentiate into the epidermis, which in roots is composed
mostly of long epidermal cells. Some cells of the protoderm develop into root hairs (Fig.
7.10) by the extension of epidermal cell walls into the surrounding soil. Root hairs may be
quite long, but they are only one cell. The cell walls of root hairs are thin and composed
principally of cellulose and pectic substances. Root hairs tend to be sticky, so that soil
particles cling to them (Fig. 7.10b). In most plants, the life of any one root hair is short; it
functions for only a few days or weeks. New hairs are constantly forming at the apical
end of the roothair zone, while those at the basal end are dying. Thus, as the root
advances through the soil, fresh, actively growing root hairs are constantly coming into
contact with new soil particles. In the rye plant, root hairs develop at an average rate in
excess of 100 million per day.

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 Root hairs account for a major fraction of the surface area of the root tip, and
thus they generally play a major role in the absorption of water and nutrients from
the soil. Although nearly all ordinary land plants possess root hairs, a few plants, such
as some gymnosperm trees, like firs, apparently lack them. Also, many aquatic plants
have no root hairs. Moreover, land plants (corn, for example) that normally develop
root hairs when the root system grows in the soil may develop no root hairs when the
roots grow in water. In plants devoid of root hairs, absorption is accomplished entirely
through the epidermal cells.
The epidermis in roots is usually one cell layer thick; but in the aerial roots of certain
plants, such as orchids, a multilayered epidermis develops that stores and possibly absorbs
water from the moist air.

a b

Figure 7.10. The development of root hairs.

(a) Radish seedling ( Raphanus sativa). (b)
Stages in the development of root hairs.
Note that the external epidermal cell wall
protrudes and that the cell cytoplasm and
nucleus move into the root hair near the tip.
Root hairs are in close contact with soil particles
and increase the water absorptive surface of the root.

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Figure 7.11. (a) A dicot root cross section
from the area where maximum absorption
occurs. The epidermis contains root hairs.
The cortex has abundant air spaces
between parenchyma cells. The
endodermis is bounded by a Casparian
strip. The pericycle is one cell-layer thick.
The primary xylem is distributed into three
protoxylem points, with metaxylem in the
middle of the root. The primary phloem
alternates with primary xylem. Residual
procambium not shown) occurs between
the primary xylem and primary phloem.

CORTEX The root cortex is derived from ground meristem and is composed chiefly of
parenchyma cells. The innermost layer of the cortex, a single row of cells called the
endodermis (Figs. 7.11, 7.12a, b) plays a special role in controlling mineral accumulation by
the root. This is the role of the Casparian strip, a waxy material embedded in the upper,
lower (transverse), and side (radial) walls of endodermal cells (Fig. 7.12).
Water and dissolved minerals from the soil can move from cell to cell by two paths;
they can travel through the porous walls of the cortex and epidermis, or they can move
through the living cells (Fig. 7.12c, d). Movement through the cell wall is free movement
without any constraints. Movement into a living cell, however, is regulated because it
involves crossing the plasma membrane. Some substances can move across the membrane
by diffusion. Some minerals, for example potassium, can be moved across membranes
through special proteins ("channels") embedded in the membrane. Some proteins can
actually pump minerals into a living cell, even against a diffusion gradient.
The function of the endodermis is to guarantee that the minerals that finally reach
the vascular cylinder can do so only by first passing across at least one plasma membrane.
One reason this is important in roots is that it provides a mechanism to increase the
concentration of needed minerals through pumps in the endodermis cell membrane.
In roots of many plants, an exodermis containing Casparian strips occurs at the outer
layer of the cortex, just inside the epidermis. This layer is present in many grass roots and in
the aerial roots of orchids (Fig. 7.13). The exodermis apparently also functions to regulate ion
absorption and accumulation.

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 a b

Figure 7.12. Control of nutrient movement into the xylem is a function of the endodermis. (a, b)
The endodermis is a single layer of inner cortex cells which have a waxy strip (Casparian strip)
embedded into their transverse and radial cell walls. (c) The strip keeps water from moving
indiscriminately through the cell walls and into the vascular cylinder. (d) The Casparian strip makes
water move through the endodermal cells and in this way the plasma membrane can selectively
control the uptake of nutrients.

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 Figure 7.13. The Casparian strip in
exodermis cells of an aerial root of
Epidendrum orchid. The plasma
membrane has been pulled away
from the cell wall by soaking the roots
in salt solution. This SEM shows that
the membrane is attached at the
Casparian strips. X800.

VASCULAR CYLINDER The entire central cylinder of roots is composed of vascular tissue
that differentiates from the procambium cells (Fig. 7.14). In roots of dicot plants the primary
xylem usually consists of a central core of xylem elements organized into two or more
radiating points (Fig. 7.11a). In most monocot roots the very center of the root is composed of
parenchyma cells with the primary xylem and phloem forming in a ring (Fig. 7.11b). The first
xylem elements to mature, the protoxylem, develop at the outer points of the xylem (Figs.
7.11, 7.14b). Metaxylem, the last primary xylem to mature, differentiates in the center of the
vascular cylinder (Fig. 7.14b, c).
The protoxylem is capable of transporting water while the root is elongating, which
requires both the strength to withstand the forces that move water and still be flexible
enough to stretch as the root elongates. This dual ability comes from a secondary cell
wall in the shape of annular rings or spirals (see Fig. 4.9).
Metaxylem cells mature in regions of the root where elongation has been
completed. Because they are no longer required to elongate, they form thick
secondary cell walls with pits through which lateral exchange of water and minerals
may take place. Protoxylem cells often become crushed after the metaxylem
develops, but by then these cells are not needed.
In roots of monocots like asparagus ( Asparagus officinale ), a central region of
parenchyma cells forms (Fig. 7.15). This region is sometimes called a pith, which
refers to the location of ground tissue in the center of stems, which is formed from
ground meristems. However, in roots it is part of the vascular cylinder and originates
from procambium. Xylem (both protoxylem and metaxylem) of roots consists of
several other cell types, including vessel elements, tracheids, parenchyma, and
fibers.
Phloem cells form in the areas between the protoxylem arms (Figs. 7.11, 7.14).
The protophloem is actually the first part of the vascular system to become
functional. These cells form at the periphery of the phloem and function primarily
during root elongation. Metaphloem develops toward the inside and functions during
the plant's adult life. Phloem of roots may consist of parenchyma, fibers, sieve-tube
members, and companion cells.
The outer boundary of the vascular cylinder is the pericycle (Figs. 7.11, 7.14c).
This tissue is unique, in that it remains capable of dividing for a long time. It has
three functions in dicot roots: (1) It is the site where the development of lateral roots

13
 is initiated, (2) it contributes to the formation of vascular cambium, and (3) it
contributes to the formation of the cork cambium. In monocot roots no secondary
growth occurs, so pericycle is involved only in lateral root initiation. The pericycle is
usually one cell layer thick, but in some roots it has multiple layers.

a b

c d

Figure 7.14. Differentiation in the primary growth of buttercup (Ranunculus sp.) roots. (a) Immature
region, where no secondary walls have yet formed in the xylem. X170. (b) The cells of the
protoxylem have developed secondary walls (shown here stained red with safranin). This particular
root has four protoxylem points with phloem between. X190. (c) Fully mature root with all primary
tissues differentiated. Note that the endodermal cells adjacent to the protoxylem points are lacking
secondary walls. X170. (d) The same section as (c), but showing all tissues. X190.

Lateral Roots are Initiated in the Pericycle

The initiation of lateral roots at particular locations is controlled by chemical growth

regulators that cause pericycle cells to begin dividing at specific sites (Fig. 7.16a). The
lateral root primordia that result (Fig. 7.16d) continue to form new cells, which in turn
elongate. Endodermal cells outside the primordium also divide for a short time,
contributing cells to the tip of the new lateral root. As it expands (Fig. 7.16b), the lateral root
pushes its way through and destroys the cortical cells and the outer epidermis. The

14
 breakdown of these cortical cells is thought to be at least partly the result of digestive
enzymes released from the lateral root primordium. As the lateral root emerges, its cells
become organized into a root cap and root apical meristem (Fig. 7.16c, d). The wound
formed by lateral root emergence is quickly healed by the secretion of mucilage and waxy
substances by adjacent cortical cells. The vascular system of the main root axis and the
lateral roots is connected (Fig. 7.16e).

Figure 7.15. Asparagus officinale root in

transverse section showing all tissues and
root regions. The vascular cylinder (stele)
has a pith of all parenchyma cells. X50.

a b d e

Figure 7.16. Lateral root formation. (a) Initiation of a lateral root of carrot (Daucus carota) by division
of cells in the pericycle. (b) Formation of the lateral root primordium. (c) The young root pushing
through the cortex. (d) Cleared pea (Pisum sativum) root showing root primordia. (e) Cleared pea
root showing vascular connections of the main axis of the root and the lateral roots.

15
The Vascular Cambium and the Cork Cambium Partially Form from the Pericycle

In long-lived dicot plants, the older regions of roots form secondary vascular tissues by
activating a secondary meristem, the vascular cambium. Secondary growth is initiated by
the division of pericycle cells and also some leftover or residual procambium cells located
between the arcs of xylem and phloem (Fig. 7.17a). Residual procambium cells are actually
procambial cells that did not develop into primary xylem or primary phloem. They are now
induced to divide, and they form secondary xylem to the inside and phloem to the outside
(Fig. 7.17b). After a time, the crescent-shaped region of dividing cells joins with the
pericycle, which also begins to divide, forming at least two layers of pericycle cells. The
inner layer joins to the residual procambium to form an intact ring of vascular cambium
(Fig. 7.17c). The outer layer remains as pericycle.
The secondary xylem or wood in roots with several years of secondary growth looks
very much like that of woody stems. The only difference is that in young roots primary
xylem occupies the middle of the root (Fig. 7.18a,b), whereas in young stems pith occupies
the middle.

Figure 7.17. Secondary

growth in a root. (a) At the
completion of primary
growth, an arc of residual
procambium cells remains
between the primary xylem
and primary phloem. The
pericycle is a complete
cylinder. (b) The residual
procambium starts to divide a b c
and joins with the pericycle
cells outside the xylem arms
to form a continuous
cylinder of vascular
cambium. The pericycle just
outside the protoxylem
points divides to form at
least two cell layers. The
inner layer joins with the
residual procambium to
form the vascular cambium; d
the outer layer stays part of
the pericycle cylinder
(c) The vascular cambium
forms secondary xylem internally and secondary phloem externally. The primary phloem is being
pushed outward. (d) A cylinder of vascular cambium produces secondary xylem to the inside and
secondary phloem to the outside. The primary xylem remains in the center of the root, the primary
phloem has been crushed, and the pericycle that remains will form the cork cambium. The cork
cambium forms the periderm after the epidermis and cortex die. The term bark refers to everything
outside the vascular cambium.

16
Continued growth expands the root and a
finally causes the splitting, sloughing off,
and destruction of the cortex and
epidermis (Fig. 7.17c). The pressure
created by this expansion apparently
stimulates the remaining ring of pericycle
cells to divide again. This last function of
the pericycle converts it into the cork
cambium, which forms the periderm (see
Fig. 4.20). The bark in woody roots (which
includes all cells from the vascular
cambium outward) appears similar to that
in stems; but it may be thinner and b
smoother on its outer surface (Fig. 7.17d).
Interestingly, there is only one
monocot plant, the dragon's blood tree
(Dracaena draco), that is known to have
secondary growth in roots. Even very
tall, tree-like monocots like palms lack
secondary growth in roots.

Figure 7.18. (a) Alfalfa (Medicago sativa) root

transverse section with some secondary
tissues and periderm on its periphery. X125.
(b) Woody root of oak (Quercus sp.). X77.

Some Roots Have Special Functions

Many plants have roots with special characteristics. This chapter has already
discussed examples of adventitious roots that arise from non-root origins and the
uniquely shaped roots of clinging vines. Other plants, especially those forming
partnerships with microorganisms, have specialized root structures. Some important
examples are haustorial roots, root nodules, mycorrhizae, and contractile roots.
Parasitic plants like dodder ( Cuscuta sp.) anchor themselves by sinking
haustorial roots into the vascular tissue of a host stem, thus tapping the host's water
and nutrient supply (Fig. 7.19).

17
 Although nitrogen is one of the most important elements needed by plants,
most plants cannot use atmospheric nitrogen (N 2 ) directly. Certain legumes, such as
peas (Pisum sativum ) and soybeans
(Glycine max ), are capable of fixing
nitrogen--that is, changing N 2 that
diffuses into the soil into NH 4 + a
(ammonium ion), which is usable
by the plant. Nitrogen fixation is
the result of an unusual
relationship between the
bacterium Rhizobium and the roots
of legumes (see Chapter 19; Fig.
19.15). Root cells are infected by
the passage of a thin infection
thread of the bacteria into root
hair cells and on through to the b
cortical cells. The bacteria then
divide and stimulate the cortical
cells to divide, thereby forming a
root nodule (Fig. 7.20). The
bacteria are the actual agents for
fixing the nitrogen.
Mycorrhizae are short, forked
roots common to as many as 90% of
Figure 7.19. (a) Haustorial roots of a plant
seed plants. These specialized root
parasite, dodder (Cuscuta sp.) infecting the
structures represent an association
stem of a host plant, X53. (b) External view of
of roots with a soil-borne fungus (see
dodder on tomato plants.
Chapter 20).
Two types of mycorrhizal
roots may occur, distinguished by
whether the fungus penetrates into
the root cells or not. Ectotrophic

mycorrhizae are found in roots of such trees as pines (Pinus), birches (Betula),
willows (Salix), and oaks (Quercus). This type causes a drastic change in the root
shape (Fig. 7.21), but the fungus does not penetrate the root cells. Instead, the
fungus penetrates between the cell walls of the cortex, and it forms a covering
sheath (or mantle) of fungal hyphae around the entire root. These mycorrhizal
roots are about 0.5 cm (0.2 in) long; they lack a root cap and have a simple vascular
cylinder. Endotrophic mycorrhizae do not form a mantle over the root, and the
fungus actually enters the cortex cells (Fig. 7.22). Mycorrhizae make roots more
efficient in mineral absorption, but they are apparently not absolutely essential for
the growth of the usual host plants (see Chapter 11). This is known because plants
that are artificially fed adequate nutrients can grow without mycorrhizae.
Mycorrhizae may also be beneficial to their host plants by secreting hormones or
antibiotic agents that reduce the potential of plant disease.

18
 Roots of the dandelion (Taraxacum officinale), water hyacinth (Hyacintha
orientalis), and some other plants are capable of contracting, which keeps above
ground parts near the soil surface. This contraction is caused by the radial
expansion (or, in some instances, the collapse) of cells in the root cortex (Fig. 7.23).

a a

Figure 7.20. Bacterial nodules in legume

roots. (a) Legume plants form bacterial
nodules on roots. (b) Bacteria enter the plant
by passing through a tiny infection thread
which penetrates root hairs. Once inside the
host, the bacteria penetrate to the cortex of
the root forming a swollen mass of cells filled Figure 7.21. Ectotrophic mycorrhizae. (a) The
with bacteria. The bacteria, called Rhizobium typical Y-branched form in pine roots ( Pinus
sp., fix nitrogen which then passes up the pinaster). (line scale = 1mm). (Used with
plant body in a usable form. permission from J. Parlade, IRTA-Departament
de Proteccio Vegetal, Barcelona, Spain) (b)
Transverse section through an infected
mycorrhizal root showing the mantle of fungal
hyphae, and the hyphae growing between the
cortical cell walls, X240.

19
Figure 7.22. Photograph of Coralloriza sp.
rhizome with endotrophic mycorrhizae,
showing fungal hyphae actually inside the
host cells. (Note: A rhizome is actually a stem
which grows underground. This example is
being used, however, because it shows the
nature of the infection process very well, and
it looks the same in root cells. X735.

Figure 7.23. Contractile roots of water

hyacinth (Hyacinthus orientalis). Notice the
wrinkled surface at the base of these roots
where the contraction occurs.

KEY TERMS

adventitious roots procambium

Casparian strip prop roots
ectotrophic mycorrhizae protoderm
endodermis radicle
endotrophic mycorrhizae region of elongation
exodermis region of maturation
fibrous root system residual procambium
ground meristems root apical meristems
haustorial roots root hairs
lateral root primordial root nodule
pericycle seminal roots
pneumatophores tap root system

20
SUMMARY

1. The principal functions of roots are absorption of water and nutrients, conduction
of absorbed materials and food, and anchorage of the plant in the soil.

2. The primary root develops from the radicle in the embryo. It generally
penetrates the soil to some depth; if it dominates, a tap root system results, with a
main root axis and branches.

3. Fibrous root systems are formed by seminal roots arising in the embryo in
addition to the radicle. Grasses are good examples of plants with fibrous roots.

4. Adventitious roots may arise on stems and leaves.

7. The root tip is composed of (a) the root cap, which protects (b) the meristematic
region; (c) a region of elongation; and (d) a region of maturation, characterized
externally by root hairs and internally by the formation of primary vascular tissues.

6. The epidermis forms as the outer tissue of the root. Water is absorbed through
the epidermal cells and the root hairs. The next layer is the cortex. Its cells mainly
store nutrients. The endodermis is the innermost cell layer of the cortex. The
Casparian strip is a waxy substance found in the radial and transverse walls. Water
cannot move across the Casparian strip. Therefore, all water with dissolved
nutrients must pass through the protoplasts of endodermal cells. An exodermis is
present just inside the epidermis in many roots; it may also have Casparian strips
and function in ion absorption and regulation.

7. In cross section, the primary xylem in dicot roots is star-shaped, with protoxylem
at the points. Primary phloem arises between the arms of primary xylem.

8. Roots of certain grasses usually have central parenchyma and many protoxylem
points.

9. The pericycle, a row of cells internal to the endodermis, represents the outermost
row of cells of the vascular cylinder. Cells of pericycle may eventually initiate the
differentiation of the vascular cambium and the cork cambium in dicot roots. The
pericycle initiates lateral roots in both dicots and monocots.

10. A vascular cambium originates in dicot roots from procambium cells between
primary xylem and phloem and from pericycle cells exterior to the radiating points
of primary xylem. The vascular cambium forms secondary xylem internally and
secondary phloem externally. The resulting increase in diameter stretches and
tears the endodermis, cortex, and epidermis. A cork cambium develops from the
pericycle and forms the periderm.

21
11. Haustorial roots from parasitic plants penetrate into the host. Bacterial nodules
occur in roots of nitrogen-fixing legumes. Mycorrhizae are roots infected by
beneficial fungi. Contractile roots pull the shoot tight to the soil surface.

Questions

1. Identify in a diagram or photograph the following root structures and tissues:

root cap cortex

root apical meristem (RAM) endodermis
region of elongation Casparian strip
region of maturation pericycle
root hair vascular cylinder (xylem and phloem)
epidermis

2. State how each item listed in question 1 contributes to the function of the root.

3. Discuss the tissues and cells involved in mineral uptake and transport in a root.

4. Describe the differences between primary and secondary tissues in a root. Where are
these located? Make a labeled diagram to show both primary and secondary xylem and
phloem.

5. Make a diagram to show the position of cork cambium and vascular cambium in a dicot
root.

6. Describe the structure and function of the root cap.

7. Describe two symbiotic associations involving roots (root nodules and mycorrhizae).
What are the microorganisms involved, and how does the association alter the structure
of the root? How do both partners benefit from the association?

8. Describe two modified roots.

22
PLANTS, PEOPLE, AND THE ENVIRONMENT: The climbing fig and its adhesive pads.

Everybody has seen vines growing on

the sides of buildings and fences. Have
you ever wondered how they manage to
climb so high and stick to different
surfaces?
Climbing vines have developed three
different growth strategies– twining,
winding and clinging. Twining vines like
Wisteria ( Wisteria sinensis ) encircle
poles and branches by twisting their
stems just like winding a rope around a
pole. Winding vines like clematis
(Clematis hybrida) support themselves by
tendrils. Tendrils are modified leaves Figure 1. Climbing fig plant ( Ficus pumila)
that can twist around poles and other growing on a privacy wall in Davis, CA.
things and attach to them. A third type
are called clinging vines and English ivy
(Hedera helix) is probably the best-
known example.
Another common plant that uses this strategy is the climbing fig (Ficus pumila).
Clinging vines climb surfaces by means of clusters of adventitious roots that emerge from
internodes and form into an interesting unique and sticky structure called an adhesive
pad.
Ficus pumila, the climbing fig, is commonly used as an ornamental to cover walls and
fences (Fig. 1). It is a very aggressive plant and it sprouts shoots that rapidly cover almost
any surface, including glass windows. Perhaps the first report of the clinging vine
mechanism in climbing fig was by Charles Darwin (1875) in his book, The Movements and
Habits of Climbing Plants. In his report he notes that roots on the stems secreted a
“viscid fluid” possibly “India-rubber” that adheres to any surface.

Adhesive Pads Hold the Climbing Fig to Any Surface

The adhesive pad in F. pumila develops from a cluster of adventitious roots that are
initiated just basal to a node near the tips of shoots (Figs. 2, 3). Adventitious roots are
initiated in pairs on either side of a vascular bundle at the second to third internodes of
young stems. After emerging through the cortex and epidermis of the stem, the
adventitious root clusters elongate until they are a few millimeters long (Fig. 2). The roots
then stop elongating, root hairs form on them, and they secrete a very sticky cement
substance. The clusters of adventitious roots and their root hairs all stick together
forming the adhesive pad (Fig. 3). If the adventitious roots do not touch a substrate they
usually dry up; if they touch moist soil they tend to branch and change to a terrestrial
form. If you ever see a climbing fig try pulling one away from the wall; you will see how
strongly they stick.

23
Fig. 2 Fig. 3

Figure 2. Tip of a climbing fig branch showing the position of an adhesive pad (red circle).
Figure 3. Close up view of an adhesive pad taken through a glass plate. Note that some of the
adventitious roots have merged and some are separate. Hairs on the surfaces of these roots secrete
a sticky substance that glues the pad to any surface.

24
PLANTS, PEOPLE, AND THE ENVIRONMENT: Myths and Popular Uses of Roots

Over the millennia, through trial

and error, ancient peoples developed
traditional uses--medical and otherwise--
for berries, seeds, leaves, stems, and
roots. Much of this plant lore has little
validity in medical fact, but some
traditional plants have proven to be
effective remedies.
During the Middle Ages in Europe,
most people believed that the way a plant
looked (its shape or color), how it
smelled, or some other characteristic
provided clues about its uses for people.
This was called the Doctrine of
Signatures. For example, the roots of
the mandrake ( Mandragora officinarum, Figure 1. These are colored drawings of
Fig. 1) are thick and fleshy and tend to be mandrake plants originally from an old
irregularly branched; with a little German Herbal. Herbals were among the very
imagination the shape of a person could first books ever made. They contained
be seen in a root carefully extracted from drawings of plants and sometimes discussed
the soil. In the Middle Ages, people how they could be used. (From Lewis, W.H.
assigned considerable importance to the and M.P.F. Elvin-Lewis. 1977. Medical Botany:
manlike appearance of these roots; they Plants Affecting Man's Health. Wiley-
believed that such roots could bring good Interscience. N.Y. Used with permission of
fortune. In some cases the roots were Wiley-Interscience.)
even ground up and eaten as a love
potion. Mandrake root doesn't seem to
have much real medicinal value. It
doesn't really give anyone good luck, and
it certainly doesn't work as a love potion;
but in some cases an infusion
made from mandrake root may help control a cough.
Another root that looks vaguely like a person is the ginseng ( Panax
quinquefolium). In Asia, ginseng root has been considered a cure-all for many
ailments. Ginseng also grows in the United States, but it has only recently become
popular. Nowadays you can find it in most health food stores. Most likely, the subtle,
supposedly restorative power associated with ginseng is psychological: it helps
because you think it will. However, some recent evidence suggests that chemicals in
ginseng roots do act to calm some people.
A root with more widely accepted utility comes from the cassava ( Manihot
esculenta), a shrubby South American plant. The whitish latex that exudes from a
cut plant contains hydrocyanic acid, a deadly poison used by Brazilian Indians to
make poisoned arrows. The interesting thing, however, is that when the latex is
removed, the rich, starchy pulp is a good food source. Native people ground up

25
cassava root, placed it in a sack, and hung it from a tree; when all the juice dripped
out, the meal that remained could be made into cakes and eaten. It was called
farina. Now cassava is grown commercially. You are probably familiar with the
starch-rich pellets manufactured from cassava root, called tapioca.
Native Americans also have a rich plant lore that includes roots. The roots of
several plants were used as a source of soap. One of these--soap weed ( Yucca sp.)--
is now used as a component of popular shampoos. Yucca root juice was also used as
glue to fasten feathers to arrows. One type of soap root was used to induce vomiting
as a step in ritual purification ceremonies. Another type was thrown in streams to
stupefy fish so that they could be harvested without using fishing gear. Roots are
obviously important to the plant, but many also have uses for people

26
IN DEPTH: How Do Roots Advertise their Presence?

Certain connections between plant roots and fungi are extremely important
for both members. In a mycorrhizal association, fungi get a rich source of
carbohydrates, which provide carbon and energy for growth. Plants get a source of
inorganic nutrients. Phosphate is particularly important, since phosphates have
very limited solubility in soil solutions. But how does the association form? How do
the two members find each other? Is it just random chance?
Recent research has identified
a class of compounds, strigolactones
(right), synthesized in the roots of all
plants. These compounds serve to
control growth in plants--they tend to
inhibit branching in shoots--but they
are also exuded from roots and can
serve as a signal to attract mycorrhizal
fungi [particularly ascomycete fungi
(see Chapter 20) that form arbuscular
mycorrhizae]. Strigolactones tend to break down in the soil, so that there is a
gradient of concentration, dropping steeply away from the root (or from the fungus's
standpoint, rising steeply as it grows toward the root).
The name "strigolactone" is
derived from the plant genus Striga
(witchweed), reflecting an unfortunate
side-effect of the exudation of these
compounds. Striga is the genus of
several species of parasitic plants.
Striga seeds germinate and form a
root-to-root connection with a nearby
plant, for instance maize or sugar
cane, removing carbohydrate and
other nutrients from the host plant
and eventually killing it.
Strigolactones stimulate the
germination of Striga seeds. And just
as mycorrhizal fungi zero in on roots
by detecting the gradient of
strigolactones, so may Striga roots.

Right: Striga (witchweed) plant (photo

by Randy Westbrooks, Invasive Plant
Control, Inc., Bugwood.org)

For more information: http://www.biomedcentral.com/17417007/12/19

Photo Credits

27
Figure 7.1 = Rost 1st
Figure 7.2 = New art Wadsworth
Figure 7.3 = Rost 1st
Figure 7.4 = Rost 1st
Figure 7.5 = (a) www.botgard.html (UCLA) (b) www.ecolibrary.org (c) Wikimedia Commons
(cc) Hans Hillewaert
Figure 7.6 = a & b Rost 1st
Figure 7. 7 a = longitudinal section of the root tip of flax – Thomas L. Rost
Figure 7.7 b = longitudinal section of onion – Thomas L. Rost
Figure 7.8 = Rost 1st
Figure 7.9 = Rost 1st – photo from Margaret McCully used with her permission.
Figure 7.10 = (a) http://www.instructables.com/id/MicroGreen-Machine-hydroponic-micro-
greens-at-vir/step4/Watch-and-wait; (b) Rost 1st
Figure 7.11 = Rost 1st
Figure 7.12 = all in Rost 1st; a – from Starr and Taggart; b – Thomas L. Rost; c & d – Starr and
Taggart
Figure 7.13 = Thomas L. Rost
Figure 7.14 a = differentiation in the primary growth of buttercup- immature region – Thomas
L. Rost
Figure 7.14 b = differentiation in the primary growth of buttercup- secondary walls – Thomas
L. Rost
Figure 7.14 c = differentiation in the primary growth of buttercup- fully mature – Thomas L.
Rost
Figure 7.14 d = differentiation in the primary growth of buttercup- showing all tissues –
Thomas L. Rost
Figure 7.15 = photo of asparagus officinale root in transverse section – Thomas L. Rost
Figure 7.16 a-c = Rost 1st, originally from Weier 6th
Figure 7.16 d & e = photo of root cleared to show cont. vascular connection – Maude Hinchee
Figure 7.17 = Rost 1st; redrawn from Weier 6th
Figure 7.18 = a & b – Thomas L. Rost (a appeared in Rost 1st, b is new)
Figure 7.19a = Rost 1st
Figure 7.19b = Rost anatomy teaching collection
Figure 7.20a&b = Starr and Taggart (I think)
Figure 7.21 a = ectotrophic mycorrhizae- typical Y-branched from in pine roots - Used with
permission from J. Parlade, IRTA-Departament de Proteccio Vegetal, Barcelona, Spain)
Figure 7.21 b = ectotrophic mycorrhizae- transverse section through an infected mycorrhizal
root – originally in Weier 6th ed also in Rost 1st ed.
Figure 7.22 = photograph of Coralloriza – Thomas L. Rost
Figure 7.23 = Judy Jernstedt, UC, Davis – Rost 1st
Sidebar – Ficus pumila – Figures 1-3 – Thomas L. Rost

28