C H A P T E R

Roots and Soils

Overview
Learning Outcomes
How Roots Develop
Root Structure
The Root Cap
The Region of Cell Division
The Region of Elongation
The Region of Maturation
Specialized Roots
Food-Storage Roots
Water-Storage Roots
Propagative Roots
Pneumatophores
Aerial Roots
Contractile Roots
Buttress Roots
Parasitic Roots
ECOLOGY: Plants Need Roots
Mycorrhizae
Root Nodules
Human Relevance of Roots
Soils
Parent Material
Climate
Living Organisms and Organic
Composition
Topography
Soil Texture and Mineral Composition
Soil Structure
ECOLOGY: Metal-Munching Plants
Water in the Soil
Soil pH
SUMMARY
REVIEW QUESTIONS
DISCUSSION QUESTIONS
ADDITIONAL READING
LEARNING ONLINE

Aerial roots of a banyan tree (Ficus sp.) near the

door of Ta Prohm Temple in Cambodia.
© Glowimages/Punchstock RF
OVERVIEW
This chapter discusses roots, beginning with their functions and continuing with the development of roots from a seed. It covers the function
and structure of the root cap, region of cell division, region of elongation, and region of maturation (with its tissues). The endodermis and
pericycle are also discussed.
Specialized roots (food-storage roots, water-storage roots, propagative roots, pneumatophores, aerial roots, contractile roots, buttress
roots, parasitic roots) and mycorrhizae are discussed. This is followed by some observations on the economic importance of roots. After a
brief examination of soil horizons, the chapter concludes with a discussion of the development of soil texture, composition, and structure; and
water.
Learning Outcomes
1. State the primary functions and forms of roots.
2. Describe root regions, including the root cap, region of cell
division, region of cell elongation, and region of maturation
(including root hairs and all tissues) and list the function of
each.
3. Provide the specific functions of the endodermis and the
pericycle.
Y
ou have likely seen pictures of the destruction caused
by a tornado as it passed through a village or a city,
but have you seen what a twister can do to a forest?
Large trees may be snapped off above the ground or knocked
down, and branches may be stripped bare of leaves. Unless
the soil in the area happens to be thin, sandy, or loose, how-
ever, you will probably see relatively few trees completely
torn up by the roots and blown elsewhere. In the tropics, it
is indeed rare to find healthy palm trees uprooted even after
a hurricane.
Roots anchor trees firmly in the soil, usually through an
extensive branching network that constitutes about one-third
of the total dry weight of the plant. The roots of most plants
do not usually extend more than 3 to 5 meters (10 to 16 feet)
down into the earth; those of many herbaceous species are
confined to the upper 0.6 to 0.9 meter (2 to 3 feet). The roots
of a few plants, such as alfalfa, however, often grow more
than 6 meters (20 feet) into the earth. When the Suez Canal
was being built, workers encountered roots of tamarisk at
depths of nearly 30 meters (100 feet), and mesquite roots
have been seen 53.4 meters (175 feet) deep in a pit mine in
the southwestern United States. Some plants, such as cacti,
form very shallow root systems, but these systems still
effectively anchor the plants with a densely branching mass
of roots radiating out in all directions as far as 15 meters
(50 feet) from the stem.
Besides anchoring plants, roots absorb water and min-
erals in solution, mostly through “feeder” roots found in the
upper meter (3.3 feet) of soil. Some plants have roots that,
as well as anchoring and absorbing, store water or food, or
perform other specialized functions.
Some aquatic plants (e.g., duckweeds and water hyacinths)
normally produce roots in water, and many epiphytes (non-
parasitic plants that grow suspended without direct contact
66
4. State the differences among the various types of specialized
roots.
5. List at least 10 practical human uses of roots.
6. Describe how a good agricultural soil is developed from raw
materials.
7. Explain how soil constituents affect the availability of water to
plants and discuss the role of soil pH in nutrient availability.
with the ground—e.g., orchids) produce aerial roots. The great
majority of vascular plants, however, develop their root systems
in soils. The soils, which vary considerably in composition,
texture, and other characteristics, are discussed toward the end
of this chapter.
How Roots Develop
When a seed germinates, the tiny, rootlike radicle, a part
of the embryo (immature plantlet) within it, grows out
and develops into the first root. The radicle may develop
into a thick, tapered taproot, from which thinner branch
roots arise, or many adventitious roots may arise from
the stem, which is attached to the radicle and continuous
with it. Adventitious roots are those that do not develop
from another root but develop instead from a stem or leaf.
A fibrous root system, which may have large numbers of
fine roots of similar diameter, then develops from the adven-
titious roots (Fig. 5.1). Many mature plants have a combina-
tion of taproot and fibrous root systems.
The number of roots produced by a single plant may
be prodigious. For example, a single, mature ryegrass plant
may have as many as 15 million individual roots and branch
roots, with a combined length of 644 kilometers (400 miles)
and a total surface area larger than a volleyball court, all con-
tained within 0.57 cubic meter (20 cubic feet) of soil. Root
hairs (discussed in the section “The Region of Maturation”)
greatly increase the total surface area of the root.
Many plants, such as peas and carrots, whose seeds
have two “seed leaves”—commonly referred to as dicots—
have taproot systems with one, or occasionally more, pri-
mary roots from which secondary roots develop (see the
discussion of primary and secondary tissues in Chapter 4).

(a)
Figure 5.1   Root systems. (a) A fibrous root system of a grass. (b) A taproot system of a California poppy.
Monocotyledonous plants (e.g., corn and rice, whose seeds
have one “seed leaf”—commonly referred to as monocots),
on the other hand, have fibrous root systems. Adventitious
and other types of roots may develop in both dicots and
monocots. In English and other ivies, lateral adventitious
roots that aid in climbing appear along the aerial stems,
and in certain plants with specialized stems (e.g., rhizomes,
corms, and bulbs; see Fig. 6.14), adventitious roots are the
only kind produced.
Root Structure
Close examination of developing young roots usually reveals
four regions, or zones. Three of the regions are not sharply
defined at their boundaries. The cells of each region grad-
ually develop the form of those of the next region, and the
extent of each region varies considerably, depending on the
species involved. These regions are called (1) the root cap,
(2) the region of cell division, (3) the region of elongation,
and (4) the region of maturation (Fig. 5.2).
The Root Cap
The root cap is composed of a thimble-shaped mass of
parenchyma cells covering the tip of each root. It is quite
large and obvious in some plants, while in others, it is
nearly invisible. One of its functions is to protect from
damage the delicate tissues behind it as the young root
tip pushes through abrasive soil particles. The root cap
has no equivalent in stems. The dictyosomes of the root
cap’s outer cells secrete and release a slimy substance that
lodges in the walls and eventually passes to the outside.
Roots and Soils 67
(b)
The cells, which are replaced from the inside, constantly
slough off, forming a slimy lubricant that facilitates the
root tip’s movement through the soil. This mucilaginous
lubricant also provides a medium favorable to the growth
of beneficial bacteria that add to the nitrogen supplies
available to the plant (see the discussion of the nitrogen
cycle in Chapter 25).
The root cap, whose cells have an average life of less
than a week, can be removed from a living root, and when
this is done, a new root cap is produced. Until the root cap
has been renewed, however, the root seems to grow ran-
domly instead of more or less downward, suggesting that
the root cap also functions in the perception of gravity (see
gravitropism in Chapter 11). It is known that amyloplasts
(plastids containing starch grains) act as gravity sensors,
collecting on the sides of root-cap cells facing the direction
of gravitational force. When a root that has been growing
vertically is artificially tipped horizontally, the amyloplasts
tumble or float down to the “bottom” of the cells in which
they occur. The root begins growing downward again within
30 minutes to a few hours. The exact nature of this gravita-
tional response is not known, but there is some evidence that
calcium ions in the amyloplasts influence the distribution of
growth hormones in the cells.
The Region of Cell Division
Cells in the region of cell division, which is composed of
an apical meristem (a tissue of actively dividing cells) in
the center of the root tip, produce the surrounding root cap.
Most of the cell divisions take place next to the root cap at
the edges of this inverted cup-shaped zone, located a short
distance behind the actual base of the meristem. Here the
68 Chapter 5

region of
maturation

protoderm
region of ground meristem
elongation
procambium

apical meristem
(region of apical meristem
root cap cell division) (region of
cell division)

root cap

(a) (b)

Figure 5.2   A longitudinal section through a dicot root tip. (a) Regions of the root. (b) Locations of the primary meristems of the root.

cells divide every 12 to 36 hours, while at the base of the
meristem, they may divide only once in every 200 to 500 hours.
The divisions are often rhythmic, reaching a peak once or
twice each day, usually toward noon and midnight, with rela-
tively quiescent intermediate periods. Cells in this region are
mostly cubical, with relatively large, more or less centrally
located nuclei and a few very small vacuoles.
In both roots and stems, the apical meristem soon sub-
divides into three meristematic areas: (1) The protoderm
gives rise to an outer layer of cells, the epidermis; (2) The
ground meristem, to the inside of the protoderm, produces
parenchyma cells of the cortex; (3) The procambium,
which appears as a solid cylinder in the center of the root,
produces primary xylem and primary phloem (Fig. 5.3).
Pith (parenchyma) tissue, which originates from the ground
meristem, is generally present in stems but is absent in most
dicot roots. Grass roots and those of most other monocots,
however, do have pith tissue.

epidermis
cortex

endodermis
epidermis pericycle
cortex
passage
pith cell
primary
xylem
primary
phloem

pith

(a) (b)

Figure 5.3   A cross section of a monocot (greenbrier—Smilax) root. (a) Complete view. ×40. (b) Enlargement showing partial section of the
root interior. ×100. (a-b) © James E. Bidlack
 Roots and Soils 69

The Region of Elongation
The region of elongation, which merges with the apical
meristem, usually extends about 1 centimeter (0.4 inch) or
less from the tip of the root. Here the cells become several
times their original length and somewhat wider. At the same
time, the tiny vacuoles merge and grow until one or two large
vacuoles, occupying up to 90% or more of the volume of each
cell, have been formed. Only the root cap and apical meri-
stem are actually pushing through the soil, because no further
increase in cell size takes place above the region of elonga-
tion. The usually extensive remainder of each root remains
stationary for the life of the plant. If a cambium is present,
however, there normally is a gradual increase in girth through
the addition of secondary tissues produced by the cambium.
The Region of Maturation
Most of the cells mature, or differentiate, into the various
distinctive cell types of the primary tissues in this region,
which is sometimes called the region of differentiation,
or root-hair zone. The large numbers of delicate hairs that
develop from many of the epidermal cells give the root-hair
zone its name. The root hairs, which absorb water and min-
erals, adhere tightly to soil particles (Fig. 5.4) with the aid
of microscopic fibers they produce and greatly increase the
total absorptive surface of the root. Differentiation is dis-
cussed further in Chapter 11.
The root hairs are not separate cells; rather, they are
tubular extensions of specialized epidermal cells. In fact,
the nucleus of the epidermal cell to which each is attached
often moves out into the root hair. They are so numerous that
they appear as a fluffy mass to the naked eye, typically num-
bering more than 38,000 per square centimeter (250,000
per square inch) of surface area in roots of plants such as
corn; they seldom exceed 1 centimeter (0.4 inch) in length.
A single ryegrass plant occupying less than 0.6 cubic meter
(20 cubic feet) of soil was found to have more than 14 billion
root hairs, with a total surface area almost the size of a foot-
ball field.
When a seedling or plant is moved, many of the deli-
cate root hairs are torn off or die within seconds if exposed
to the sun, greatly reducing the plant’s capacity to absorb
water and minerals in solution. This is why plants should be
watered, shaded, and pruned after transplanting until new
root hairs have formed. In any growing root, the extent of the
root-hair zone remains fairly constant, with new root hairs
being formed toward the root cap and older root hairs dying
back in the more mature regions. The life of the average root
hair is usually not more than a few days, although a few live
for a maximum of perhaps 3 weeks.
The cuticle (see Fig. 4.11), which may be relatively
thick on the epidermal cells of stems and leaves, is thin
enough on the root hairs and epidermal cells of roots in
the region of maturation (Fig. 5.5) to allow water to be
absorbed but still sufficient to protect against invasion by
bacteria and fungi.
The cells of the cortex, a tissue composed of paren-
chyma cells lying between the epidermis and inner tissues,
mostly store food. This tissue, which may be many cells
thick, is similar to the cortex of stems except for the presence
of an endodermis at its inner boundary (Fig. 5.6). The endo-
dermis consists of a single-layered cylinder of compactly
arranged cells whose walls are impregnated with lignin and
suberin. These substances form bands, called Casparian
strips, which can be found on the radial and transverse cell
walls. The plasma membranes of the endodermal cells are
fused to the Casparian strips, which are perpendicular to
the root’s surface; they prevent water from passing through
the otherwise permeable (porous) cell walls. The Casparian

epidermalcell
epidermal cell

airspace
air space

roothair
root hair

developing
developing
root
roothair
hair

cell
cellofofcortex
cortex
(a)
(a) (b)
(b)

Figure 5.4   (a) A radish (Raphanus) seedling shortly after germination, showing the root hair zone. (b) A diagram of an enlargement of a
longitudinal section of a small portion of a root-hair zone, showing root hairs in contact with soil particles. Note that in most soils there is normally
considerably less air space between soil particles than is depicted in this image. (a) © Kingsley Stern
70 Chapter 5
epidermis
cortex
(a)
Figure 5.5   A cross section of a dicot (buttercup—Ranunculus) root. (a) Complete view. ×40. (b) Enlargement of the root center (vascular
cylinder). ×400. (a-b) © James E. Bidlack
cortex
endodermis
pericycle
passage cell
Figure 5.6   A portion of the endodermis of a buttercup
(Ranunculus) root. ×1,000. © James E. Bidlack
strip barrier forces water and dissolved substances entering
and leaving the central core of tissues to pass through the
plasma membranes of the endodermal cells or their plasmo-
desmata. This regulates the types of minerals absorbed and
transported by the root to the stems and leaves. The plasma
membranes tend to exclude harmful minerals while gener-
ally retaining useful ones.
An endodermis is rare in stems but so universal in
roots that only three species of plants are known to lack a
root endodermis. In some roots, the epidermis, cortex, and
endodermis are sloughed off as their girth increases, but in
those roots where the endodermis is retained, the inner walls
of the endodermal cells eventually become thickened by the
addition of alternating layers of suberin and wax. Later, cel-
lulose and sometimes lignin are also deposited. Some endo-
dermal cells, called passage cells, may remain thin-walled
and retain their Casparian strips for a while, but they, too,
eventually tend to become suberized.
A core of tissues, referred to collectively as the
vascular cylinder, lies to the inside of the endodermis. Most
of the cells of the vascular cylinder conduct water or food
endodermis
primary xylem
primary phloem
pericycle
passage cell
(b)
in solution, but lying directly against the inner boundary of
the endodermis is an important layer of parenchyma tissue
known as the pericycle. This tissue, which is usually one
cell wide, may in some plants be a little wider. The cells of
the pericycle may continue to divide even after they have
matured. Lateral (branch) roots and part of the vascular
cambium of dicots arise within the pericycle (Fig. 5.7).
In most dicot and conifer roots, the primary xylem con-
sists of a solid central core of water-conducting cells (e.g.,
tracheids; vessels). In cross section, this first root xylem
usually loosely resembles the rear view of a rocket with
fins. The fins, generally referred to as arms, tend to taper
toward their tips and terminate just inside of the thin, cylin-
drical pericycle layer. There are usually four of these arms,
with some plants having two, three, or several. Lateral roots
arise toward the outside of these xylem arms, from within
the pericycle. Dividing cells cause the pericycle to bulge out
into the cortex and the endodermis usually ruptures before
the lateral root emerges. As the new root emerges, it pushes
aside or crushes the cortex cells and eventually breaks
through the epidermis. Parenchyma cells between the vas-
cular tissues of the originating and lateral roots differentiate
into vascular elements and establish connections between
the old and new tissues.
The primary xylem surrounds pith parenchyma cells
in monocot roots and those of a few dicots; in such plants,
the arms may not be well defined. Primary phloem, which
conducts food, forms in discrete patches between the xylem
arms of both dicot and monocot roots.
A vascular cambium develops from parts of the pericy-
cle and other parenchyma cells between the xylem arms and
phloem patches in most dicots and conifers. This cambium
at first follows the starlike outline of the primary xylem as
it starts producing secondary phloem to the outside and
secondary xylem to the inside. Eventually, however, the
position of the cambium gradually shifts so that instead of
appearing as patches and arms, the secondary conducting

Figure 5.7   A cross section through a dicot (willow—Salix) root showing the origin of a lateral (branch) root. Photomicrograph by G.S. Ellmore
tissues appear as concentric cylinders. The primary phloem,
in particular, may be sloughed off and lost as secondary tis-
sues are added.
In woody plants, a second cambium, the cork cambium,
normally arises in the pericycle outside of the vascular cam-
bium and gives rise to cork tissue (periderm). Cork cells,
which are dead at maturity, are impregnated with suberin
and are impervious to moisture; similar tissues are produced
in stems. Although there are exceptions, monocot roots gen-
erally have no secondary meristems and therefore no sec-
ondary growth.
In both roots and stems (see Chapter 6), growth may
be determinate or indeterminate. Determinate growth is
growth that stops after an organ such as a flower or a leaf
is fully expanded or after a plant has reached a certain size.
Indeterminate growth occurs in trees and other perenni-
als where new tissues are added indefinitely, season after
season.
Natural grafting can take place between roots of dif-
ferent trees of the same species, especially in the tropics.
The roots unite through secondary growth when they come
in contact with one another, but the details of the uniting
process are not yet known. One unfortunate aspect of this
grafting is that if one tree becomes diseased, the disease can
be transmitted through the grafts to all the other trees con-
nected to it.
Roots and Soils 71
epidermis
pericycle
lateral root
xylem
phloem
cortex
Specialized Roots
As mentioned earlier, most plants produce a fibrous root
system, a taproot system, or, more commonly, combina-
tions of the two types. Some plants, however, have roots
with modifications that adapt them for performing specific
functions as well as the absorption of water and minerals in
solution.
Food-Storage Roots
Most roots and stems store some food, but in certain plants,
the roots are enlarged and store large quantities of starch and
other carbohydrates (Fig. 5.8), which may later be used for
extensive growth. In sweet potatoes and yams, for example,
extra cambial cells develop in parts of the xylem of branch
roots and produce large numbers of parenchyma cells. As a
result, the organs swell and provide storage areas for large
amounts of starch and other carbohydrates. Similar food-
storage roots are found in the deadly poisonous water hem-
locks, in dandelions, and in salsify. In carrots, beets, turnips,
and radishes, the food-storage tissues are actually a combi-
nation of root and stem. Although the external differences
are not obvious, approximately 2 centimeters (0.8 inch) at
the top of an average carrot is derived from stem tissue that
merges with the root tissue below.
72 Chapter 5
Figure 5.8   Sweet potato (Ipomoea) plants. Note the food-
storage roots. © Martin Harvey/Getty Images RF
Water-Storage Roots
Some members of the Pumpkin Family (Cucurbitaceae) pro-
duce huge water-storage roots. This is particularly charac-
teristic of those that grow in arid regions or in those areas
where there may be no precipitation for several months of
the year. In certain manroots (Marah), for example, roots
weighing 30 kilograms (66 pounds) or more are frequently
produced (Fig. 5.9), and a root of one calabazilla plant
(Cucurbita perennis) was found to weigh 72.12 kilograms
(159 pounds). The water in the roots is used by the plants
when the supply in the soil is inadequate.
Propagative Roots
Many plants produce adventitious buds (buds appearing in
places other than stems) along the roots that grow near the
surface of the ground. The buds develop into aerial stems
called suckers, which have additional rootlets at their bases.
The rooted suckers can be separated from the original root
and grown individually. Cherries, apples, pears, and other
fruit trees often produce suckers. The adventitious roots of
rice-paper plants (Tetrapanax papyrifera) and tree-of-heaven
(Ailanthus altissima) can become a nuisance in gardens,
often producing propagative roots 10 meters (33 feet) or
more from the parent plant. Horseradish (Rorippa armora-
cia), Canada thistle (Cirsium arvense), and some other weeds
have a remarkable ability to reproduce in this fashion as well
as by means of seeds. In the past, this capacity has made
them difficult to control, but some biological controls being
Figure 5.9   A manroot (Marah) water-storage root that weighs
over 25.3 kilograms (60 pounds). Courtesy of Robert A. Schlising
investigated (see Appendix 2) may be an answer to the prob-
lem in the future.
Pneumatophores
Water, even after air has been bubbled through it, contains
less than one-thirtieth the amount of free oxygen found in
the air. Accordingly, plants growing with their roots in water
may not have enough oxygen available for normal respira-
tion in their root cells. Some swamp plants, such as the black
mangrove (Avicennia nitida) and the yellow water weed
(Ludwigia repens), develop special spongy roots, called
pneumatophores, which extend above the water’s surface
and enhance gas exchange between the atmosphere and the
subsurface roots to which they are connected (Fig. 5.10).
The woody “knees” of the bald cypress (Taxodium disti-
chum), which occurs in southern swamps (see Fig. 22.19),
were in the past believed to be pneumatophores, but there is
no conclusive evidence for this theory.
Aerial Roots
Velamen roots of orchids (Fig. 5.11), prop roots of corn and
banyan trees, adventitious roots of ivies, and photosynthetic
roots of certain orchids are among various kinds of aerial
roots produced by plants. It was formerly assumed that the
epidermis of velamen roots, which is several cells thick,
aided in the absorption of rain water. It appears, however,
it may function more in preventing loss of moisture from
the root. Corn prop roots, produced toward the base of the
stems, support the plants in a high wind. Some tropical
plants, including the screw pines and various mangroves,
produce sizable prop roots extending for several feet above
the surface of the ground or water. Debris collects between
them and helps create additional soil.

(a)
(b)
Figure 5.10   (a) Pneumatophores (foreground) of tropical
mangroves rising above the sand at low tide. The pneumatophores
are spongy outgrowths from the roots beneath the surface.
Pneumatophores facilitate the exchange of oxygen and carbon
dioxide for the roots, which grow in areas where little oxygen
is otherwise available to them. (b) Close-up photograph of
pneumatophores. (a-b) Majority World/Getty Images
Roots and Soils 73
Figure 5.11   The aerial (velamen) roots of orchids have a thick
epidermis that reduces water loss from internal tissues. © Kingsley
Stern
Many of the tropical figs or banyan trees produce roots
that grow down from the branches until they contact the soil.
Once they are established, they continue secondary growth
and look just like additional trunks (Fig. 5.12). Banyan trees
may live for hundreds of years and can become very large.
In India and southeast Asia, there are several banyan trees
that have almost 1,000 root-trunks and have circumferences
approaching 450 meters (1,476 feet). The oldest is estimated
to be about 2,000 years old.
The vanilla orchid, from which we obtain vanilla fla-
voring, produces chlorophyll in its aerial roots and, through
photosynthesis, can manufacture food with them. The
adventitious roots of English ivy, Boston ivy, and Virginia
creeper appear along the stem and aid the plants in climbing.
Contractile Roots
Some herbaceous dicots and monocots have contractile
roots that pull the plant deeper into the soil. Many lily bulbs
are pulled a little deeper into the soil each year as new sets
of contractile roots are developed (Fig. 5.13). The bulbs con-
tinue to be pulled down until an area of relatively stable tem-
perature is reached. Plants such as dandelions always seem
to have the leaves coming out of the ground as the top of the
stem is pulled down a small amount each year when the root
contracts. The contractile part of the root may lose as much
as two-thirds of its length within a few weeks as stored food
is used and the cortex collapses.
74 Chapter 5

Figure 5.12   A banyan (Ficus) tree with many large prop roots that have developed from the branches. © Kingsley Stern

(a) (b) (c) (d)

Figure 5.13   A lily bulb over three seasons is pulled deeper into the soil by the action of contractile roots. (a) A small bulb produced
during the first growing season. (b) Contractile roots pull the newly formed bulb down several millimeters during the first season. (c) The bulb is
pulled down farther the second season. (d) The bulb is pulled down even farther the third season. The bulb will continue to be pulled down in
succeeding seasons until it reaches an area of relatively stable soil temperatures.

Buttress Roots Parasitic Roots

Some tropical trees growing in shallow soils produce huge, Some plants, including dodders, broomrapes, and pinedrops,
buttresslike roots toward the base of the trunk, giving them have no chlorophyll (which is necessary for photosynthesis)
great stability (Fig. 5.14). Except for their angular appear- and have become dependent on chlorophyll-bearing plants for
ance, these roots look like a part of the trunk. their nutrition. They parasitize their host plants via peglike
 Roots and Soils 75

the air and rain. Indian pipe is referred to as an epiparasite

KEY THEME : ecology because it is a parasite of fungi in the soil; some scientists
also use the term myco-heterotroph to describe Indian pipe
because this plant obtains some or all of its carbon from the
fungus rather than from photosynthesis.
Plants Need Roots

projections called haustoria (singular: haustorium), which (a)

develop along the stem in contact with the host. The haustoria
penetrate the outer tissues and establish connections with the
xylem and phloem (Fig. 5.15). Some green plants, including
Indian warrior and the mistletoes, also form haustoria. These
haustoria, however, apparently aid primarily in obtaining
water and dissolved minerals from the host plants, because
the partially parasitic plants are capable of manufacturing
at least some of their own food through photosynthesis. The
term saprophyte, which refers to organisms that lack chloro-
phyll and obtain nutrients from organic matter, is most appro-
priately used to describe the way many fungi obtain nutrition.
It has been used in the description of some plants, but this is
not correct, and more accurate terminology can be applied,
depending on where plants obtain their nutrients. Orchids,
for instance, are often considered as epiphytes because they
grow on other plants and obtain moisture and nutrients from

(b)

Figure 5.15   (a) Pale stems of a parasitic plant (dodder—Cuscuta)

twining about other vegetation. (b) A close-up view of dodder, showing
the peglike haustoria that penetrate the tissues of the host plant. ×5.
Figure 5.14   Buttress roots of a tropical fig tree. © Kingsley Stern (a-b) © Kingsley Stern
76 Chapter 5

Mycorrhizae upon the association for survival. (Mutualism is a form of

More than 95% of land plants have fungi associated with
their roots and these associations are termed mycorrhizae
(Fig. 5.16). The association is mutualistic; that is, both
the fungus and the root benefit from it and are dependent
sym­biosis; see page 299.) For some mycorrhizae the fun-
gal hyphae penetrate the root cortex but the hyphae do not
enter the cells. On the root surface, the hyphae form a thin,
encrusting layer called the mantle. This type of mycorrhiza is
termed ectotrophic and is found in trees like pines and oaks.

normal mycorrhizal fungal hyphae

roots roots
mycorrhizal
sheath
longitudinal section
through part of the epidermis
root
cortex of root

endodermis
pericycle
phloem
xylem

(b)

(a)

root root
endodermis phloem xylem cortex ectomycorrhizae

endomycorrhizae

(c) (d)

Figure 5.16   Mycorrhizae. (a) A longitudinal drawing of a root with ectomycorrhizae (visible on the right outside of the root). (b) A diagram
of a cross section of a root with ectomycorrhizae. See Figs. 5.5b and 5.6 for definitive photomicrographs of most of the tissues in this diagram.
(c) Photomicrograph of a cross section of a root around which ectomycorrhizae have formed a mantle ×40. The fungal cells have not penetrated
deeper than the outermost layers of root cells. (d) A cross section of a few root parenchyma cells with endomycorrhizae ×100. The endomycorrhizae
develop and flourish within the parenchyma cells. (c-d) © Kingsley Stern
 Roots and Soils 77

In another type of mycorrhiza, the hyphae invade the cells of

the cortex and a mantle is not present. This type of mycor-
rhiza is called endotrophic and is found in the majority of
woody and herbaceous land plants.
Mycorrhizae are essential to the normal growth and
development of forest trees, as well as most herbaceous
plants, and they facilitate the uptake of water and inorganic
ions. The fungus is able to absorb and concentrate phospho-
rus much better than it can be absorbed by the root hairs.
In fact, if mycorrhizal fungi have been killed by fumigation
or are otherwise absent, many plants appear to have consid-
erable difficulty absorbing phosphorus, even when the ele-
ment is abundant in the soil. Hence, making phosphorus
readily available for absorption is a very important function
of these fungi. In exchange for the phosphorus made avail-
able to the plant by fungi, sugars and amino acids are pro-
vided by plants, which are essential for the fungi to survive.
Some plants do not seem to need mycorrhizae unless there
are barely enough essential elements for healthy growth pres-
ent in the soil. Plants with mycorrhizae develop few root hairs
compared with those growing without an associated fungus.
Mycorrhizae have been shown to be particularly susceptible
to acid rain (discussed in Chapter 25); this may signal major
problems for our coniferous forests in the future if the problem
of acid rain is not solved. Methyl bromide, used in the past to
sterilize seed beds, kills all soil organisms, including mycor-
rhizae; its continued use in the United States has been banned.

Root Nodules
Although almost 80% of our atmosphere consists of nitrogen
gas, plants cannot convert the nitrogen gas to usable forms.
A few species of bacteria, however, produce enzymes with
which they can convert nitrogen into nitrates and other nitrog-
enous substances readily absorbed by roots. Members of the
Legume Family (Fabaceae), which includes peas, beans,
alfalfa, and a few other plants such as alders, form associa-
tions with certain soil bacteria that result in the production of
numerous small swellings called root nodules that are clearly
visible when such plants are uprooted (Fig. 5.17). The nod-
ules contain large numbers of nitrogen-fixing bacteria.
A substance exuded into the soil by plant roots stimulates
Rhizobium bacteria, which, in turn, respond with another
substance that prompts root hairs to bend sharply. A bacte-
rium may attach to the concave side of a bend and then invade
the cell with a tubular infection thread that does not actually
break the host cell wall and plasma membrane. The infection
thread grows through to the cortex, which is stimulated to
produce new cells that become a part of the root nodule; here
the bacteria multiply and engage in nitrogen conversion. (See
also the discussion of the nitrogen cycle in Chapter 25.)
Root nodules should not be confused with root knots,
which are also swellings that may be seen in the roots of toma-
toes and many other plants. Root knots develop in response
to the invasion of tissue by small, parasitic roundworms
(nematodes). Unlike bacterial nodules, root knots are not
Figure 5.17   Root nodules on the roots of bur clover (Medicago
polymorpha). The somewhat popcorn-like nodules contain bacteria that
convert nitrogen from the air into forms that can be used by the plant. ×15.
© Kingsley Stern
beneficial, and the activities of the parasites within them can
eventually lead to the premature death of the plant.
Human Relevance of Roots
Roots have been important sources of food for humans since
prehistoric times, and some, such as the carrot, have been
in cultivation in Europe for at least 2,000 years. A number
of cultivated root crops involve biennials (i.e., plants that
complete their life cycles from seed to flowering and back
to seed in two seasons). Such plants store food in a swollen
taproot during the first year of growth, and then the stored
food is used in the production of flowers in the second sea-
son. Among the best-known biennial root crops are sugar
beets, beets, turnips, rutabagas, parsnips, horseradishes, and
carrots. Other important root crops include sweet potatoes,
yams, and cassava. Cassava (Fig. 5.18), from which tapioca
is made, forms a major part of the basic diet for millions of
inhabitants of the tropics. With a minimum of human labor,
it yields more starch per hectare (about 45 metric tons, the
equivalent of 20 tons per acre) than any other cultivated
78 Chapter 5
Figure 5.18   Cassava (Manihot esculenta) plants. Note the food-storage roots on the plants that
have been dug up. © Shelley H. Jansky
crop. Minor root crops, including relatives of wild mustards,
nasturtiums, and sorrel, are cultivated in South America and
other parts of the world.
Several well-known spices, including sassafras, sar-
saparilla, licorice, and angelica, are obtained from roots.
Sweet potatoes are used in the production of alcohol in
Japan. Some important red to brownish dyes are obtained
from roots of members of the Madder Family (Rubiaceae),
to which coffee plants belong. Drugs obtained from roots
include aconite, ipecac, gentian, and reserpine, a tranquil-
izer. A valuable insecticide, rotenone, is obtained from the
barbasco plant, which has been cultivated for centuries as
a fish poison by primitive South American tribes. When
thrown into a dammed stream, the roots containing rotenone
cause the fish to float but in no way poison them for human
consumption. In tobacco plants, nicotine produced in the
roots is transported to the leaves. Other uses of roots are dis-
cussed in Chapter 24.
Soils
The soil is a dynamic, complex, constantly changing part of
the earth’s crust, which extends from a few centimeters deep
in some places to hundreds of meters deep in others. It is
essential not only to our existence but also to the existence
of most living organisms. It has a pronounced effect on the
plants that grow in it, and they have an effect on it.
If you dig up a shovelful of soil from your yard and
examine it, you will probably find a mixture of ingredients,
including several grades of sand; rocks and pebbles; powdery
silt; clay; humus; dead leaves and twigs; clods consisting of
soil particles held together by clay and organic matter; plant
roots; and small animals, such as ants, pill bugs, millipedes,
and earthworms. Also present, but not visible, would be mil-
lions of microorganisms, particularly bacteria, fungi, and, of
course, air and water.
The soil became what it is today through the inter-
action of a number of factors: climate, parent material,
topography of the area, vegetation, living organisms, and
time. Because there are thousands of ways in which these
factors may interact, there are many thousands of differ-
ent soils. The solid portion of a soil consists of mineral
matter and organic matter. Pore spaces, shared by variable
amounts of water and air, occur between the solid parti-
cles. The smaller pores often contain water, and the larger
ones usually contain air. The sizes of the pores and the
connections between them largely determine how well the
soil is aerated.
If you were to dig down 1 or 2 meters (3 to 6 feet) in an
undisturbed area, a soil profile of three intergrading regions
called horizons would probably be exposed (Fig. 5.19). The
horizons show the soil in different stages of development,
and the composition varies accordingly. The upper layer,
usually extending down 10 to 20 centimeters (4 to 8 inches),
is called the topsoil. It is usually subdivided into a darker
upper portion called the A horizon and can have a lighter
lower portion below it called the E horizon. The E horizon is
not always present and usually occurs in older, well-developed
soils. The A portion contains more organic matter than the
layers below. It can be divided into different parts such as
the A1 and A2 horizons.

A1 horizon (crushed fine sandy loam,
dark brown)
A2 horizon (crushed fine sandy
loam, brown)
Bt 1 horizon (crushed sandy
loam, brown)
2Bt 2 horizon (broken face sandy
clay loam, reddish brown)
2Bt 3 horizon (broken face sandy
clay loam, yellowish red)
2C horizon (crushed stratified
coarse sand, strong brown)
Figure 5.19   A soil profile showing horizons that depict different
stages of development. Steven McGowen, USDA-NRCS
The next 0.3 to 0.9 meter (1 to 3 feet) section is called
the B horizon, or subsoil. It usually contains more clay and
is lighter in color than the topsoil. As with other soil hori-
zons, lowercase letters after the uppercase letter may be
used to describe characteristics within the horizon, such as
Bw (weakly colored or structured) and Bt (translocation of
silicate clay), as well as numbers after these labels to des-
ignate layers within the horizon. A number preceding the
horizon letter may also be used if the soil consists of differ-
ent parent materials. The C horizon at the bottom may vary
from about 10 centimeters (4 inches) to several meters (6 to
10 feet or more) in depth; it may even be absent. It is com-
monly referred to as the soil parent material and extends
down to bedrock.
Parent Material
The first step in the development of soil is the formation
of parent material from existing rocks that have not yet
been broken down into smaller fragments. Parent mate-
rial accumulates through the weathering of three types of
rock, which originate from various sources. These sources
and types include volcanic activity (igneous rocks);
depositing by glaciers, water, or wind (sedimentary
Roots and Soils 79
rocks); and changes in igneous or sedimentary rocks
brought about by great pressures, heat, or both (meta-
morphic rocks).
Climate
Climate varies greatly throughout the globe, and its role
in the weathering of rocks varies correspondingly. In des-
ert areas, for example, there is little weathering by rain,
and soils are poorly developed. In areas of moderate rain-
fall, however, well-developed soils are common. In some
areas of high rainfall, the excessive flow of water through
the soil may leach out important minerals. Similar leach-
ing out of important minerals may occur when gar-
den sprinklers are left on all night. Many gardeners and
house-plant enthusiasts have stunted or killed the very
plants they were trying to foster by “drowning” them with
too much water or too frequent watering. As a general
rule, plants should not be watered unless the soil surface
feels dry.
In areas where there are great temperature ranges, rocks
may split or crack as their outer surfaces expand or con-
tract at different rates from the material beneath the surface.
When water in rock crevices freezes, it expands and causes
further cracks and splits. The breaking up of rocks contrib-
utes to the development of soil.
Living Organisms
and Organic Composition
There are many kinds of organisms in the soil, as well as
roots and other parts of plants. In the upper 30 centimeters
(1 foot) of a good agricultural soil, living organisms con-
stitute about one-thousandth of the total weight of the soil.
This may not sound significant, but it amounts to approxi-
mately 6.73 metric tons per hectare (3 tons per acre).
Bacteria and fungi present in the soil decompose organic
matter, which accumulates when leaves fall and plants and
animals die. (This process is further discussed under the
section on composting in Chapter 17.) Roots and all other
living organisms produce carbon dioxide, which combines
with water in the soil and forms an acid, thereby increasing
the rate at which minerals dissolve. Ants and other insects,
earthworms, burrowing animals, and birds all alter the soil
through their activities and add to its organic content, either
through wastes that they deposit or through the decomposi-
tion of their bodies when they die. Humus, which consists
of partially decomposed organic matter, gives some soils a
dark color.
The total organic composition of a soil varies greatly.
An average topsoil might consist of about 25% air, 25%
water, 48% minerals, and 2% organic matter. Soils in low,
wet areas, where a lack of oxygen keeps microorganisms
from their normal activities, may contain as much as 90%
organic matter. Except in legumes and a few other plants,
80 Chapter 5
almost all of the nitrogen utilized by growing plants, as well
as much of the phosphorus and sulfur, comes from decom-
posing organic matter. In addition, as organic matter breaks
down, it produces acids, which, in turn, decompose miner-
als. Other roles of organic matter in the soil are discussed in
the “Soil Structure” section.
Topography
If the land is steep, soil may wash away or erode through
the action of wind, water, and ice as soon as it is weath-
ered from the parent material. It has been estimated that
more than 20 metric tons of topsoil per hectare (8.2 tons
per acre) are washed away annually from some prime
croplands in the central United States.
If an area is flat and poorly drained, pools and ponds
may appear in slight depressions when it rains. If these bod-
ies of water cannot drain quickly, the activities of organisms
in the soil are interrupted, and the development of the soil is
arrested. The ideal topography for the development of soil is
one that permits drainage without erosion.
Soil Texture and Mineral
Composition
Soil texture refers to the relative proportions of sand, silt,
and clay in a given soil (Table 5.1).
Sands are usually composed of many small particles
bound together chemically or by a cementing matrix material.
Table 5.1   Soil Mineral Components as Classified by the U.S. Department of Agriculture
Silt consists of particles that are mostly too small to be seen
without a lens or a microscope.
Clay particles are so tiny that they can’t be seen with
even a powerful light microscope, although they can be
seen with an electron microscope. Individual clay parti-
cles are called micelles. Micelles are somewhat sheetlike,
negatively charged, and held together by chemical bonds.
The negative charges attract, exchange, or retain posi-
tively charged ions. Many of the positively charged ions,
such as magnesium (Mg++) and potassium (K+), which
are needed for normal plant growth, are absorbed with
water by the roots. Clay is plastic in nature because the
water that adheres tightly to the surface of the particles
acts both as a binding agent and a lubricant. Physically,
clay is a colloid—that is, a suspension of particles that
are larger than molecules but that do not settle out of a
fluid medium.
The best agricultural soils are usually loams, which are
a mixture of sand, clay, and organic matter. The better loams
have about 40% silt, 40% sand, and 20% clay. Light soils
have a high sand and low clay content. Heavy soils have
high clay content. Coarse soils, which have larger particles,
are porous and don’t retain much water, while clay soils have
high water content and allow little water to pass through.
Over half the composition by weight of mineral matter
is oxygen. Other elements commonly present are hydrogen,
silicon, aluminum, iron, potassium, calcium, magnesium,
and sodium. However, soil obtains hundreds of different
mineral combinations from its parent material.

Metal-Munching Plants
Soil Structure
Soil structure refers to the arrangement of the soil particles
into groups called aggregates. Aggregates in sands and
gravels show little cohesion, but most agricultural soils
have aggregates that stick together. Structure develops
when colloidal particles clump together, mostly as a result
of the activities of soil organisms, freezing, and thawing. If
KEY THEME : ecology
the individual granules do not become coated with organic
matter, they may continue to clump until they become
clods.
Productive agricultural soils are granular soils with
pore spaces that occupy between 40% and 60% of the total
volume of the soil. The pores contain air and water, and
their sizes are more important than their total volume. Clay
soils, for example, have more pore space than sandy soils,
81
82 Chapter 5
but the pores are so small that water and air are restricted
in their movement through the soil. When the pores are full
of water, air is kept out, and there is not enough oxygen for
root growth. Sandy soils have large pores, which drain by
gravity soon after they are filled. The water is replaced by
air, but too much air speeds up nitrogen release by micro-
organisms. Plants can’t use the nitrogen that quickly, and
much of it is lost.
Water itself, as we have noted, can be harmful. Under
anaerobic conditions (marked by the absence of oxygen),
too much water leaches mineral nutrients and slows the
mineralization process. Too much water also slows the
release of nitrogen, interferes with plant growth, and
accelerates the breakdown of nitrates to the extent that
virtually all the nitrates may be lost in as little as half
an hour.
Water in the Soil
Water in the soil occurs in three forms. Hygroscopic water
is physically bound to the soil particles and is unavailable
to plants. Gravitational water drains out of the pore spaces
after a rain. If drainage is poor, it is this water that interferes
with normal plant growth. Plants are mainly dependent on
the third type, capillary water, which is water held against
the force of gravity, in pores of the soil. The structure and
organic matter of the soil—which enable the soil to hold
water against the force of gravity—the density and type of
vegetational cover, and the location of underground water
tables largely determine the amount of capillary water avail-
able to the plant.
The ancient Incas of Peru knew that water would rise
just so far in some areas. Where the water table was close
to the surface, they removed the upper 0.6 meter (2 feet)
of soil and planted their crops down in the hollowed-out
areas so that the roots would be able to reach the capil-
lary water. They knew that in some areas having sandy
soils and low annual precipitation, soils could be com-
pacted by a heavy roller to create finer capillaries to raise
water from below. This technique is effective only if the
available water is within 1.5 to 3.0 meters (5 to 10 feet)
of the surface and if the soils do not contain too much silt
or clay.
After rain or irrigation, water in the soil drains away by
gravity. The water remaining after such draining is referred
to as the field capacity of the soil. Field capacity is mainly
governed by the texture of the soil, but the structure and
organic content also influence it to a certain extent. Plants
readily absorb water from the soil when it is at, or near,
field capacity. As the soil dries, the film of water around
each soil particle becomes thinner and more tightly bound
to the soil particle and less likely to enter the root. If water
is not added to the soil, eventually a point is reached at
which the rate of absorption of water by the plant is insuf-
ficient for its needs, and the plant wilts permanently. The
soil is then said to be at the permanent wilting point. In
clay soils, the permanent wilting point is reached when the
water content drops below 15%, while in sandy soils, the
permanent wilting point may be as low as 4%. Available
water is soil water between field capacity and the perma-
nent wilting point.
Soil pH
The pH (acidity or alkalinity) of a soil affects both
the soil and the plants growing in it in various ways.
Cranberries, for example, thrive under acidic conditions,
but a soil that is unusually acid or alkaline may be toxic to
the roots of other plants, and mycorrhizae do not survive
in soils having pH extremes. These conditions, however,
do not normally directly affect plants nearly as much as
they affect nutrient availability. For example, alkalinity
causes minerals, such as copper, iron, and manganese,
to become less available to plants, while acidity, if high
enough, inhibits the growth of nitrogen-fixing bacteria.
Acid soils tend to be common in areas of high precipita-
tion where significant amounts of bases are leached from
the topsoil.
It is a common agricultural practice to counteract soil
acidity by adding compounds of calcium or magnesium in a
process known as liming. Alkaline soils can be made more
acidic by the addition of sulfur, which is converted by bacte-
ria to sulfuric acid. The addition of some nitrogenous fertil-
izers may have the same effect.
SUMMARY
1. Roots anchor plants and absorb water and minerals
in solution. A germinating seed radicle becomes the
first root. Taproots with branch roots, or adventitious
roots that become a fibrous root system, develop from
the radicle or the stem just above it. Many plants have
combinations of both systems.
2. Four zones, or regions, of young roots are recognized:
(1) A protective root cap that also aids in the perception
of gravity. (2) A region of cell division. Its apical
meristem subdivides into a protoderm, which produces
the epidermis; a ground meristem, which produces the
cortex; and a procambium, which produces primary
xylem and primary phloem. (3) A region of elongation
in which the cells produced by the apical meristem
become considerably longer and slightly wider. (4) A
region of maturation in which the cells mature into the
distinctive cell types of primary tissues.
3. Some of the epidermal cells in the region of
maturation develop root hairs; the root hairs greatly
increase the absorptive surface of the root. The
tissues that mature in this region are similar to those
of stem tips, but pith is absent in most dicot roots and
originates from the procambium in monocot roots.

4. At its inner boundary, the cortex has an endodermis
with Casparian strips made of lignin and suberin.
5. Next to the endodermis toward the center of the root
are parenchyma cells constituting the pericycle.
Branch roots and the vascular cambium arise in the
pericycle.
6. In dicot roots, the primary xylem usually first forms a
solid core with two to several arms in the center of the
root; a pith may be present in monocot roots.
7. Primary phloem is first produced in discrete patches
between the primary xylem arms, but the tissues
eventually appear as concentric cylinders. In woody
plants, a cork cambium usually arises in the pericycle
and produces cork tissues similar to those of stems.
Roots may graft together naturally. There are no leaves
in roots.
8. Specialized roots include those for food or water
storage; pneumatophores; aerial roots (velamen roots,
prop roots, photosynthetic roots, and adventitious
roots); contractile roots; and buttress roots. Haustoria
are peglike, parasitic roots. Mycorrhizae are
mutualistic associations between roots and fungi.
Some plants have nitrogen-fixing bacteria in nodules
on their roots.
9. Root crops include sugar beets, beets, turnips,
rutabagas, parsnips, carrots, sweet potatoes, yams,
and cassava. Several spices are obtained from roots.
Other uses of roots include the production of alcohol
and the extraction of dyes, drugs, insecticides, and
poisons.
10. Soils contain a mixture of ingredients, including
sands, rocks, silt, clay, humus, dead organic
matter, plant roots, small animals, microorganisms,
plus air and water, within pore spaces of various
sizes.
11. A vertical column of soil exhibits horizons; the
topsoil is divided into an upper A horizon and a lower
E horizon. The B horizon (subsoil) usually contains
more clay and is lighter in color than the topsoil. The
C horizon (bottom portion) constitutes the weathered
soil parent material.
12. Living organisms in the soil decompose organic matter,
the source of most important plant nutrients. Animals
also cultivate the soil. Soil erosion is affected by
topography.
13. Soil texture pertains to the relative proportions of sand,
silt, and clay. More than half of the composition by
weight of mineral matter is oxygen.
14. Soil structure is the arrangement of the soil particles
into aggregates. Good soils are highly granular and
have pore spaces that constitute about half the total
volume.
15. Water in the soil occurs as hygroscopic water,
gravitational water, and capillary water.
Roots and Soils 83
16. The field capacity of the soil is the amount
of water that remains after the rest of the water
has drained away by gravity. Soil reaches the
permanent wilting point when plants wilt
permanently because they can no longer extract
enough water from the soil for their needs.
Available water is soil water between field capacity
and the permanent wilting point.
REVIEW QUESTIONS
1. Distinguish between a tiny root and a root hair. What is
the function of a root hair?
2. What is the difference between parasitic roots and
mycorrhizae?
3. If you were shown cross sections of a young root and a
young stem from the same dicot plant, how could you
tell them apart?
4. What is the function of the root cap, and from which
meristem does it originate?
5. How do endodermal cells differ from other types of
cells?
6. Where do branch roots originate?
7. List some spices and drugs obtained from roots.
8. What is soil parent material? Where does it come from,
and how does it become soil?
9. What is the difference between soil texture and soil
structure?
10. What types of soil water are recognized? What is
available soil water?
DISCUSSION QUESTIONS
1. Japanese gardeners regularly trim away parts of the
root system to assist in dwarfing a Bonsai plant. A
plant’s food is obtained through photosynthesis in the
leaves; can you suggest why trimming the roots can
cause dwarfing?
2. It was suggested that roots perceive gravity
through the root cap. Would it really matter if roots
grew randomly in the soil instead of responding to
gravity?
3. From the viewpoint of the plant, can you suggest a
practical reason for branch roots originating internally
instead of at the surface?
4. When you eat a yam or a sweet potato, what kinds of
compounds and cells are you consuming?
5. Persons associated with commercial nurseries and
greenhouses often sterilize their soil by heating it to get
rid of pests, but then they have to wait for a short time
after it has cooled to use it. Why?
 Rev.confirming Pages

84 Chapter 5

Miller, R., and D. T. Gardiner. 2000. Soils in our environment,

ADDITIONAL READING
9th ed. Paramus, NJ: Prentice-Hall.
Ridge, R. W. W., and A. M. Emons (Eds.). 2000. Root hairs: Cell
Altman, A., and Y. Waisel (Eds.). 1998. Biology of root formation
and molecular biology. New York: Springer-Verlag.
and development. Hingham, MA: Kluwer Academic.
Varma, A. (Ed.). 2008. Mycorrhiza: State of the art genetics and
Brady, N. C., and R. R. Weil. 2007. The nature and properties of molecular biology, eco-function, biotechnology,
soil, 14th ed. Paramus, NJ: Prentice-Hall. eco-physiology, structure and systematics. New York:
Charman, P., and B. Murphy (Eds.). 2007. Soils: Their properties Springer-Verlag.
and management, 3d ed. New York: Oxford University
Press. LEARNING ONLINE
Davis, T. D., and B. E. Haissig (Eds.). 1994. Biology of
adventitious root formation. New York: Plenum Press.
Evert, R. F. 2006. Esau’s plant anatomy: Meristems, cells, and
tissues of the plant body: Their structure, function, and Visit our website at http://connect.mheducation.com for additional
development, 3d ed. Hoboken, NJ: Wiley Interscience. information and learning tools.

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