CHAPTER
Mineral Nutrition
10
L E A R N I NG
OB J E CTI V E S
䊉
1 List the four components of
soil, and give the ecological
significance of each.
䊉
2 Describe the factors involved
in soil formation.
䊉
3 Distinguish between macro-
nutrients and micronutri-
ents, and list the essential
elements.
䊉
4 Discuss tension– cohesion
and root pressure as mecha-
nisms to explain the rise of
water in xylem.
䊉
5 Discuss the pressure–flow
hypothesis of sugar trans-
location in phloem.
198
and Transport
in Plants
T
he coastal redwood (Sequoia sempervirens) is an immense evergreen
with deeply grooved, spongy, reddish brown bark that may be 30 cen-
timeters (12 inches) thick on a mature tree. It produces flat, bluish
gray needles and bears its seeds in small, oval cones that mature in
one season; needles and cones are about 2.5 centimeters (1 inch) in length.
Coastal redwoods require the high humidity and cool temperatures provided
by ocean fogs and do not grow farther inland than the fogs will penetrate. Conse-
quently, their growth is restricted to a narrow ribbon of land about 800 kilometers
(500 miles) long in the foothills along southwestern Oregon and the northern Cali-
fornia coast and no more than 30 kilometers (19 miles) from the Pacific Ocean.
Coastal redwoods are among the world’s tallest trees: the world’s tallest known
tree is a coastal redwood that is 113 meters tall (370 feet). Most mature redwood
trees are 61 to 83 meters (200 to 275 feet) tall, with trunks measuring 3 meters
(10 feet) in diameter. As a coastal redwood ages, the lower branches die and fall off,
leaving a tapering, columnar trunk that extends about 30 meters (100 feet) above
the ground. The trunk spreads out at its base to provide support.
In addition to being noted for their gigantic size, coastal redwoods are known
for their extreme longevity. They mature in 400 to 500 years, and individuals
1500 or more years old are not uncommon. The oldest known coastal redwood is
2200 years old.
Redwoods are named for their reddish heartwood, which is straight grained,
attractive, and resistant to decay. Redwood lumber is highly valued for such uses
as shingles, furniture, interior paneling, doors, windowsills, and fence posts. These
majestic trees have been overexploited for their valuable timber, and most of the
original forests have been cut down. As a result, only a few virgin groves of coastal
redwoods remain, and conservationists strongly support their protection.
The coastal redwood is an appropriate plant to feature in this chapter because
of its immense size. How do water and dissolved minerals (inorganic nutrients) move
from the roots to the needles at the top of the coastal redwood? How does sugar
produced in photosynthesis move from the needles to the living cells in the roots?
We now consider these questions and examine the soil from which plants obtain
water and minerals.
D. Falconer/PhotoLink/Getty Images

The majestic redwoods are some of the world’s tallest plants.

200 • CHAPTER 10 Mineral Nutrition and Transport in Plants

Soil Soil is composed of inorganic minerals,

organic matter, air, and water
Soil is the ground underfoot, a relatively thin layer of
Earth’s crust modified by the natural actions of weather, Four distinct components compose soil: inorganic min-
wind, water, and organisms. It is easy to take soil for eral particles (which make up about 45 percent of a typi-
granted. We walk on and over it throughout our lives, cal soil), organic matter (about 5 percent), water (about
but we rarely stop to think about how important it is 25 percent), and air (about 25 percent) (•Figure 10-2).
to our survival. Vast numbers and kinds of organisms The plants, animals, fungi, and microorganisms that in-
colonize soil and depend on it for shelter and food. Most habit soil interact with it, and minerals are continually
plants anchor themselves in soil, and from it they draw cycled from the soil to organisms, which use them in
water and essential minerals. Many elements essential their biological processes. When the organisms die, bac-
for plant growth are obtained directly from the soil. teria and other soil organisms decompose the remains,
Most plants cannot survive on their own without soil, so the minerals return to the soil.
and because we depend on plants for our food, neither The inorganic mineral particles, which come from
could humans exist without soil. weathered rock, provide anchorage and essential miner-
Soils are generally formed from rock (called parent als for plants, as well as pore space for water and air. Be-
material ) that is gradually broken down, or fragmented, cause different rocks consist of different minerals, soils
into smaller and smaller particles by biological, chemi- vary in mineral composition and chemical properties.
cal, and physical weathering processes (•Figure 10-1). Rocks rich in aluminum form acidic soils, for example,
Two important factors that work together in the weath- whereas rocks that contain silicates of magnesium and
ering of rock are climate and organisms. When plant iron form soils that may be deficient in calcium, nitro-
roots and other organisms living in the soil respire, they gen, and phosphorus. Also, soils formed from the same
produce carbon dioxide (CO2), which diffuses into the kind of parent material may not develop in the same
soil and reacts with soil water to form carbonic acid way, because other factors, such as weather, topography,
(H2CO3). Soil organisms such as lichens, which are and kinds of organisms, differ.
“dual organisms” composed of a fungus and an alga or
cyanobacterium, produce other kinds of acids. These Evaporation
acids etch tiny cracks, or fissures, in rock surfaces; wa- and transpiration
ter then seeps into these cracks. If the parent material
is located in a temperate climate, the alternate freez-
ing and thawing of the water during winter break off Precipitation
small pieces of rock, causing the cracks to enlarge. Small
plants then become established and send their roots into
the larger cracks, fracturing the rock further.
Topography, which is a region’s surface features,
Plant
such as the presence or absence of mountains and val- litter
leys, is also involved in soil formation. Steep slopes often
have little or no soil on them, because gravity continually Earthworms Runoff and
transports soil and rock down the erosion
slopes. Runoff from precipitation Minerial affected by
Organic additions topography
tends to amplify erosion on steep (underground) from soil nutrients
Soil
slopes. Moderate slopes, on the other organisms microorganisms
hand, may have less downhill trans- Fine particles and
port and therefore form deep soils. solubles are washed
The disintegration of solid rock downward Water
into finer and finer mineral particles Capillarity and table
and the accumulation of organic evaporation cause some
materials to rise Parent
material (discussed in the next sec- material
tion) in the soil take an extremely
long time, sometimes thousands of Loss of soluble materials in groundwater
years. Soil forms constantly as the
weathering of parent material be- FIGURE 10-1 The dynamics of soil formation.
neath already formed soil continues Weather, climate, topography, and organisms interact with Earth’s crust to form soil, the
to add new soil. material that supports life on land.
 Soil • 201

to 0.02 millimeter in diameter), and the smallest

particles are called clay (less than 0.002 millime-
ter in diameter). Sand particles are large enough to
easily see, and silt particles (about the size of flour
particles) are barely visible. Most individual clay
particles are too small to see with an ordinary light
microscope; they can be seen only under an elec-
tron microscope.
A soil’s texture affects many of that soil’s
Water film properties, in turn influencing plant growth. The
Root hair Air space clay component of a soil is particularly important
Nematode
in determining many of its characteristics, be-
cause clay particles have the greatest surface area
of all soil particles. If the surface areas of about 450
grams (1 pound) of clay particles were laid out side
by side, they would occupy 1 hectare (2.5 acres).
Each clay particle has predominantly nega-
tive electric charges on its outer surface that attract
and reversibly bind cations, which are positively
charged mineral ions, such as potassium (K⫹) and
magnesium (Mg2⫹). Because many cations are es-
sential for plant growth, cation absorption is an
important aspect of soil fertility. Roots secrete
protons (H⫹), which are exchanged for other posi-
tively charged mineral ions attracted to the surface
of soil particles, in a process known as cation ex-
change (•Figure 10-3). The plant’s roots absorb
these “freed” ions and the water that forms a film
around the soil particles.
In contrast, anions, which are negatively
Humus
charged mineral ions, are repelled by the negative
surface charges of clay particles and tend to remain
Clay in solution. Anions such as nitrate (NO3⫺) are of-
ten washed out of the root zone by water moving
Relative
Silt through the soil.
size
of soil Soil always contains a mixture of different-
Sand
particle sized particles, but the proportions vary from one
Gravel soil to another. A loam, which is an ideal agricul-
tural soil, has an optimal combination of differ-
FIGURE 10-2 Soil components. ent-sized soil particles: it contains approximately
Soil is composed of inorganic mineral particles (sand, silt, and clay), 40 percent each of sand and silt and about 20 per-
organic material (humus), soil air, and soil water. cent of clay. Generally, the larger particles provide
structural support, aeration, and permeability to
the soil, whereas the smaller particles bind together
The texture, or structural characteristic, of a soil into aggregates, or clumps, and hold minerals and water.
is determined by the percentages (by weight) of the dif- Soils with larger proportions of sand are not as desir-
ferent-sized inorganic mineral particles—sand, silt, and able for most plants, because they do not hold water and
clay—in it. The size assignments for sand, silt, and clay mineral ions well. Plants grown in such soils are more
give soil scientists a way to classify soil texture. Any susceptible to drought and mineral deficiencies. Soils
particles larger than 2 millimeters in diameter, called
gravel or stones, are not considered soil particles because
they do not have any direct value to plants. The largest SOIL The surface layer of Earth’s crust, consisting primarily
of fragmented and weathered grains of rocks; supports ter-
soil particles are called sand (0.02 to 2 millimeters in di-
restrial plants as well as many animals and microorganisms.
ameter), the medium-sized particles are called silt (0.002
202 • CHAPTER 10 Mineral Nutrition and Transport in Plants

2+
Ca Clay particle Ca
2+

K+
2+
2+ Mg
K+ Mg
Mg
2+
K+
2+ K+
Ca
K+ K+ 2+ Mg
2+

K+ 2+ Mg
Ca
2+ Ca
2+ 2+
2+ Mg Mg
2+ Mg
Mg 2+ 2+
Ca Mg 2+
K+ Ca
2+
Mg
+ K+ K+
H 2+
+ Mg
H 2+ 2+
Mg
+ 2+ K+ Mg
H Ca
2+
+ Ca
H

USDA/Soil Conservation Service

+
H

Root
K+ K+
hair
Mg
2+
K+
Mg
2+
K+
K+
2+
Ca 2+ 2+
Mg Ca 2+
2+ 2+
2+
Mg 2+ Ca Ca
Ca Mg Mg
2+
2+
Ca 2+
K+ K+
2+ 2+ Ca
K+
Ca Ca K+

FIGURE 10-4 Humus.

Soil that is rich in humus, partially decomposed organic material
FIGURE 10-3 Cation exchange. (primarily from plant and animal remains), has a loose, some-
Negatively charged clay particles bind to positively charged what spongy structure with several properties (such as increased
mineral cations in the soil, holding the cations in the soil. Roots water-holding capacity) that are beneficial for plants and other
pump out protons (H⫹), which are exchanged for the cations, organisms living in it.
facilitating cation absorption.

with larger proportions of clay are also not desirable for resistant to decay, but a succession of microorganisms
most plants, because they have poor drainage and often gradually reduces it to CO2, water, and mineral ions.
do not provide enough oxygen. Clay soils used in agri- Organic matter dominates some soils. Peat soils,
culture tend to get compacted, which reduces the num- which consist almost entirely of undecayed plant re-
ber of soil spaces filled by water and air. mains, may form where poor drainage limits decompo-
sition by soil-dwelling bacteria.
A soil’s organic matter consists of the
wastes and remains of soil organisms About 50 percent of soil volume
The organic matter in soil is composed of litter (dead is composed of pore spaces
leaves and branches on the soil’s surface); droppings Soil has numerous pore spaces of different sizes around
(animal dung); and the remains of dead plants, animals, and among the soil particles. Pore spaces occupy
and microorganisms in various stages of decomposi- roughly 50 percent of a soil’s volume and are filled with
tion. Microorganisms, particularly soil bacteria and varying proportions of soil air and soil water. Both air
fungi, decompose organic matter—that is, break it down and water are necessary to produce a moist but aerated
into simpler materials. During decomposition, essential soil that sustains plants and other soil-dwelling organ-
mineral ions are released into the soil, where they are isms (•Figure 10-5). Water is usually held in the smaller
bound by soil particles or absorbed by plant roots. Or- pores, whereas air is found in the larger pores. After a
ganic matter increases the soil’s water-holding capacity prolonged rain, almost all the pore spaces are filled with
by acting much like a sponge. For this reason, garden- water, but water drains rapidly from the larger pore
ers often add organic matter to soils, especially sandy spaces, drawing air from the atmosphere into those
soils, which are naturally low in organic matter (see spaces.
Plants and People: Managing Your Lawn and Garden Soil air contains the same gases as atmospheric air,
at Home). although they are usually present in different propor-
The partly decayed organic portion of the soil is tions. As a result of cellular respiration by roots and
referred to as humus (•Figure 10-4). Humus, which is other soil organisms, there are less oxygen and more
not a single chemical compound but a mixture of many carbon dioxide in soil air than in atmospheric air. (Re-
organic compounds, binds mineral ions and holds wa- call from Chapter 4 that aerobic respiration uses oxygen
ter. On average, humus persists in agricultural soil for and produces carbon dioxide.) Among the important
about 20 years, although certain components of humus gases in soil air are oxygen (O2), required by roots and
may persist for hundreds of years. Humus is somewhat soil organisms for aerobic respiration; nitrogen (N2),
 Soil • 203

PLANTS AND PEOPLE | Managing Your Lawn and Garden at Home

You can put to use some of what you
have learned in this chapter when man-
aging your lawn and garden at home.
Using compost and mulch can maintain
and improve your soil. Gardeners often
dispose of grass clippings, leaves, and
other plant refuse by bagging it for
garbage collection or burning it. Neither
action is desirable, however; treating
yard wastes as garbage contributes
more material to our already overbur-
dened landfills, and the burning of yard

USDA/Soil Conservation Service

wastes contributes to air pollution.
These materials are a valuable
resource for making compost, a natural
soil-and-humus mixture that improves
fertility and soil structure. Grass clip-
pings, leaves, weeds, sawdust, coffee
grounds, animal manure, ashes from
the fireplace or grill, shredded newspa-
pers, fruit peelings and other meat- and
cheese-free leftovers, and eggshells are
just some of the materials that can be
composted, that is, transformed into
compost by microbial action.
To make a compost heap, spread a
15- to 30-centimeter (6- to 12-inch)
layer of grass clippings, leaves, or other
plant material in a shady area, sprinkle
it with garden fertilizer or a thin layer of
animal manure, and cover it with sev-
eral centimeters of soil. Add layers as
you collect more organic debris. Water
the material thoroughly, and turn it over
with a pitchfork each month to aer-
ate it. Although it is possible to make
compost by just heaping it in layers, it
is more efficient to construct a compost
enclosure. An enclosed compost heap
is also less likely to attract animals.
When the compost is uniformly dark in

Mulches discourage the growth of weeds and help keep the soil damp. Organic mulches,
such as shredded bark (shown), have the added benefit of gradually decaying, thereby
increasing soil fertility.
color, is crumbly, and has a pleasant
“woodsy” odor, it is ready to use. The
time required for decomposition varies
from 1 to 6 months depending on what
the temperature is, what materials are
used, and how often the heap is turned
and watered.
Whereas compost is mixed into soil
to improve the soil’s fertility, mulch is
placed on the surface of soil, around
the bases of plants (see figure). Mulch
helps control weeds and increases the
amount of water in the upper levels of
the soil by reducing evaporation. It low-
ers the soil temperature in the summer
and extends the growing season slightly
by providing protection against cold in
the fall. It also decreases soil erosion
by lessening precipitation runoff.
Although mulches can consist of in-
organic materials such as plastic sheets
or gravel, natural mulches of grass
clippings, straw, chopped corncobs, or
shredded bark have the added benefit
of increasing the organic content of the
soil. Grass clippings are an effective
mulch when placed around the bases
of garden plants because they mat
together, making it difficult for weeds to
become established. You must replace
grass mulches often, however, because
they decay rapidly. Some gardeners
prefer mulches of more expensive ma-
terials such as shredded bark, because
they take longer to decompose and are
more attractive.

used by nitrogen-fixing bacteria; and carbon dioxide,
involved in soil weathering.
Soil water originates as precipitation, which drains
downward, or as groundwater (water stored in porous
underground rock), which rises upward from the water
table (the uppermost level of groundwater). Soil water
contains low concentrations of dissolved minerals that
enter the roots of plants when they absorb water. Wa-
ter not bound to soil particles or absorbed by roots per-
colates (drains down) through soil, carrying dissolved
minerals with it. The removal of dissolved materials
from soil by percolating water is called leaching. The
deposition of leached material in the lower layers of
soil is known as illuviation. Iron and aluminum com-
pounds, humus, and clay are some of the illuvial ma-
terials that gather in the subsurface portion of the soil.
 Soil particles
Soil water

Soil air
Soil air

Soil water
(with dissolved
minerals)

(a) In a wet soil, most of the pore space is (b) In a dry soil, a thin film of water is tightly
filled with water. bound to soil particles, and soil air occupies
most of the pore space.

FIGURE 10-5 Pore space.

Surface
litter

Topsoil

Root
nodules:
nitrogen-
fixing
bacteria Subsoil

Parent material
(bedrock)

Mite

Nematode Fungus

Root
Protozoa Bacteria

FIGURE 10-6 Diversity of life in fertile soil.

Soil-dwelling life-forms include plant roots, algae, fungi, earthworms, flatworms, roundworms,
insects, spiders and mites, bacteria, and burrowing animals such as moles and groundhogs.
 Soil • 205

Some substances are completely leached out of the soil

because they are so soluble that they migrate down to
the groundwater. It is also possible for water that is
moving upward through the soil to carry dissolved ma-
terials with it.

Soil organisms form a complex community

A single teaspoon of fertile agricultural soil may contain
millions of microorganisms such as bacteria, fungi, al-
gae, protozoa, and microscopic worms. Many other or-
ganisms also colonize soil, including plant roots, earth- Elements
worms, insects, moles, gophers, snakes, and groundhogs Elements in organic
(•Figure 10-6). Most numerous in soil are bacteria, incorporated matter
which number in the hundreds of millions per gram of into organic in animals
soil. Biologists have identified about 170,000 species of matter in
plants
soil organisms, but thousands remain to be identified.
Little is known about the roles of soil organisms, in part
because it is hard to study their activities under natural
conditions.
Bacteria and fungi are important, not only in de-
Organic
composing dead organic material in the soil but in nu- matter
trient cycling, the processes by which matter cycles in soil
from the living world to the nonliving physical envi-
ronment and back again. Nutrient cycles break down or Plant roots Decomposition
build up chemicals that plants need (•Figure 10-7). For absorb
minerals Minerals freed and
example, microorganisms are involved in most steps in available to plant
from soil
the nitrogen cycle, which supplies nitrogen, an essential
part of proteins and nucleic acids, to organisms (see Fig-
ure 26-19). FIGURE 10-7 Nutrient cycles.
Mineral nutrients cycle from the soil to organisms and then back
Worms are some of the most important organisms
to the soil.
living in soil. Earthworms, one of the most familiar soil
inhabitants, ingest soil and obtain energy and raw mate-
rials by digesting humus. Castings, bits of soil that have
most soils ranges from 4 to 8, but some soils are outside
passed through the gut of an earthworm, are deposited
this range. The soil of the Pygmy Forest in Mendocino
on or near the soil surface. In this way, minerals from
County, California, is extremely acidic (pH 2.8 to 3.9).
deeper layers are brought to upper layers. Earthworm
At the other extreme, certain saline soils in Death Val-
tunnels aerate the soil, and the worms’ waste products
ley, California, have a pH of 8.5.
and corpses add organic material to the soil.
Soil pH affects plants, partly because the solubility
Ants live in the soil in enormous numbers, con-
of certain minerals varies with differences in pH. Plants
structing tunnels and chambers that aerate it. Members
absorb soluble mineral elements but not insoluble forms.
of soil-dwelling ant colonies forage on the surface for
At a low pH, for example, the aluminum and manga-
bits of food, which they carry back to their nests. Not
nese in soil water are more soluble and are sometimes
all this food is eaten, however, and its eventual decom-
absorbed by roots in toxic amounts. (See Plants and the
position helps increase the organic matter in the soil.
Environment: Soil Salinization for an examination of
Many ants are also indispensable in plant reproduction
soils with too many minerals.) At a higher pH, certain
because they bury seeds in the soil (see Plants and the
mineral salts essential for plant growth, such as calcium
Environment: Seed Dispersal by Ants, in Chapter 9).
phosphate, become less soluble and thus less available to
plants.
Soil pH affects soil characteristics Soil pH greatly affects the leaching of minerals. An
and plant growth acidic soil has less ability than an alkaline soil to bind
As discussed in Chapter 2, soil acidity is measured with positively charged ions because the soil particles also
the pH scale, which ranges from 0 (extremely acidic) bind the abundant protons (•Figure 10-8). As a result,
through 7 (neutral) to 14 (extremely alkaline). The pH of certain minerals essential for plant growth, such as po-
206 • CHAPTER 10 Mineral Nutrition and Transport in Plants

THE ENVIRONMENT | Soil Salinization

PLANTS AND
Soils found in arid and semiarid areas
often contain high concentrations of
inorganic mineral salts. In these areas,
the amount of water that drains into
lower soil layers is minimal because
the little precipitation that falls quickly
evaporates. In contrast, humid climates
have enough precipitation to leach salts
out of the soils and into waterways and
groundwater. Irrigation of agricultural
fields in arid and semiarid areas often Root
hair Cell contents
results in their becoming increasingly cell
salty. Also, when irrigated soil becomes
waterlogged, salts may move upward
from groundwater to the soil surface, Water
where they are deposited as a crust. Water movement
Most plants cannot obtain all movement
the water they need from salty soil,
because such soil produces a water- Relative
balance problem. Under normal condi- salt
tions, the dissolved materials in the wa- concentration
tery cytoplasm of plant cells give them (a) Root hair (b)
a concentration of water lower than

The effect of saline soil on water absorption by roots. (a) Normally there is a net move-
that of soil. As a result, water moves
ment of water (black arrows) into root cells from the soil. (b) When soil contains a high
by osmosis from the soil into plant amount of salt, water moves out of the roots into the soil, even when the soil is wet.
roots. When soil water contains a large
quantity of dissolved salts, however,
its concentration of water is lower than are genetically selected to tolerate high short supply. Also, many soils lack good
that in plant cells; consequently, water salt, are not productive in saline soil. drainage properties, so adding lots of
moves out of plant roots and into the In principle, the way to remove water simply causes them to become
salty soil (see figure). Not surprisingly, excess salt from saline soils is to add waterlogged. Even if the salt is flushed
most plants cannot survive under these enough water to leach it away. Al- out of the soil, it has to go somewhere.
conditions. Plant species that thrive in though this sounds straightforward, it The excess salt is usually carried to
saline soils have special adaptations is extremely difficult and in many cases groundwater or to rivers and streams,
that enable them to tolerate the high impossible. Recall that saline soils where it becomes a water pollutant.
amount of salt. Most crops, unless they usually occur in areas where water is in

tassium (K⫹), are leached more readily from acidic soil.
The optimum soil pH for most plant growth is 6.5 to
7.5, because most essential elements needed by plants
are available in that range.
Soil pH affects plants, and plants and other organ-
isms, in turn, influence soil pH. The decomposition of
humus and the cellular respiration of soil organisms and
roots decrease the pH of the soil.
Acid precipitation, a type of air pollution in which
sulfuric and nitric acids produced by human activities
fall to the ground as acid rain, sleet, snow, or fog, can
seriously decrease soil pH (see Plants and the Environ-
ment: Acid Rain, in Chapter 2). Acid precipitation is one
of several factors implicated in forest decline, the grad-
ual deterioration, and often death, of trees in many Eu-
ropean and North American forests (see Plants and the
Environment: The Effects of Air Pollution on Leaves,
in Chapter 8). Forest decline may be partially the result
of soil changes caused by acid precipitation. In central
European forests that have experienced forest decline, a
strong correlation exists between forest damage and soil
chemistry altered by acid precipitation.
 Soil • 207

Negatively charged ions of nitrogen salts—nitrate (NO3⫺) and ammonium

clay particles
2+
(NH4⫹). The nitrogen in the nitrogen salts ultimately
Mg
H
+ comes from atmospheric nitrogen (N2 ) and is converted
NH4+
to nitrate and ammonium by various microorganisms in
+ +
Na K
+ 2+
either the soil or the root nodules of certain plants (see
H Ca

K+
Chapter 26). The remaining 15 essential elements are
obtained from the soil as dissolved mineral ions. Their
2+
Mg
2+
NH4+ Ca
(a) In normal soil, K+
ultimate source is the parent material from which the
+
Na
positively charged mineral +Ca
2+
soil was formed.
Na
ions are attracted to the H
+ Mg
2+
Let us examine the main functions of essential el-
K+ NH +
negatively charged soil 4
ements. Carbon, hydrogen, and oxygen are found in
particles. the structures of all biologically important molecules,
including lipids, carbohydrates, nucleic acids, and pro-
+
H teins. Nitrogen is part of proteins, nucleic acids, and
Al 3+
+ chlorophyll.
H
H
+ Potassium, which plants use in fairly substantial
+ +
H H amounts, is not found in a specific organic compound
Al 3+ in plant cells. Instead, it remains as free K⫹ and plays a
+
(b) In acidified soil,
H
3+
key role in maintaining the turgidity of cells because it
protons (H⫹) displace the +
H
+ Al
is osmotically active. The presence of K⫹ in cytoplasm
H +
cations. Aluminum ions
+
Al 3+
H
causes water to pass through the plasma membrane into
released from inorganic H
the cell by osmosis. Potassium is also involved in stoma-
mineral particles when the tal opening (see Chapter 8).
soil becomes acidified also
adhere to soil particles.
Calcium plays a key structural role as a component
of the middle lamella (the cementing layer between the
cell walls of adjacent plant cells). Calcium ions (Ca 2⫹)
FIGURE 10-8 How acid alters soil chemistry.
are also important in a number of physiological roles in
plants, such as altering membrane permeability and cell
signaling (see Chapter 3).
Soil provides most of the Magnesium is critical for plants because it is part
minerals found in plants of the chlorophyll molecule. Phosphorus is a component
More than 90 naturally occurring elements exist on of nucleic acids, phospholipids (an essential part of cell
Earth, and more than 60 of these, including elements membranes), and energy transfer molecules such as ad-
as common as carbon and as rare as gold, are found in enosine triphosphate (ATP). Sulfur is essential because
plant tissues. Not all these elements are considered es- it is found in certain amino acids and vitamins. Many
sential for plant growth, however. plants accumulate silicon in their cell walls and inter-
Nineteen elements are essential for most, if not all, cellular spaces. Silicon enhances the growth and fertil-
plants (•Table 10-1). Ten of these are required in fairly ity of some species and may help reinforce cell walls.
large quantities (greater than 0.05 percent dry weight) Chlorine and sodium are micronutrients that help
and are therefore known as macronutrients. These in- maintain turgidity of cells. In addition to its osmotic
clude carbon, hydrogen, oxygen, nitrogen, potassium, role, chloride (Cl⫺) and sodium (Na⫹) ions are essential
calcium, magnesium, phosphorus, sulfur, and silicon. for photosynthesis.
The remaining nine micronutrients are needed in very
small, or trace, amounts for normal plant growth and
development. These include chlorine, iron, boron, man-
ganese, sodium, zinc, copper, nickel, and molybdenum. MACRONUTRIENT An essential element that is required in a
Four of the 19 elements—carbon, oxygen, hydro- fairly large amount for normal plant growth.
gen, and nitrogen—come directly or indirectly from
MICRONUTRIENT An essential element that is required in
soil water or from gases in the atmosphere. Carbon is very small amounts for normal plant growth.
obtained from carbon dioxide in the atmosphere during
photosynthesis. Oxygen is obtained from atmospheric OSMOSIS The net movement of water (the principal solvent
in biological systems) by diffusion through a selectively per-
oxygen and water. Water also supplies hydrogen to
meable membrane.
the plant. Plants absorb their nitrogen from the soil as
208 • CHAPTER 10 Mineral Nutrition and Transport in Plants

TABLE 10-1 Functions of Elements Required by Most Plants

ELEMENT SOURCE MAJOR FUNCTIONS
Macronutrients
Carbon Air (as CO2) In carbohydrate, lipid, protein, and nucleic acid molecules
Hydrogen Water In carbohydrate, lipid, protein, and nucleic acid molecules
Oxygen Water, air (as O2) In carbohydrate, lipid, protein, and nucleic acid molecules
Nitrogen Soil In proteins, nucleic acids, chlorophyll, certain coenzymes
Potassium Soil Osmotic and ionic balance; opening and closing of stomata; enzyme activator
(for 40⫹ enzymes)
Calcium Soil In cell walls; involved in membrane permeability; enzyme activator
Magnesium Soil In chlorophyll; enzyme activator
Phosphorus Soil In nucleic acids, phospholipids, ATP (energy transfer compound)
Sulfur Soil In certain amino acids and vitamins
Silicon Soil In cell walls

Micronutrients
Chlorine Soil Ionic balance; involved in photosynthesis
Iron Soil Involved in photosynthesis and respiration
Boron Soil In cell walls; involved in nucleic acid metabolism and in cell growth
Manganese Soil In enzymes involved in respiration and nitrogen metabolism; required for photosynthesis
Sodium Soil Involved in photosynthesis; substitutes for potassium in osmotic and ionic balance
Zinc Soil In enzymes involved in respiration and nitrogen metabolism
Copper Soil In enzymes involved in photosynthesis
Nickel Soil In enzymes (urease) involved in nitrogen metabolism
Molybdenum Soil In enzymes involved in nitrogen metabolism

Six of the micronutrients (iron, manganese, zinc,
copper, nickel, and molybdenum) are associated with
various plant enzymes, often as enzyme activators, and
are involved in certain enzymatic reactions. Boron,
present in cell walls, is also involved in nucleic acid me-
tabolism and cell growth.
PROCESS How do botanists determine
OF SCIENCE whether an element is essential?
It is impossible to conduct mineral nutrition experi-
ments by growing plants in soil, because soil contains
too many elements. Therefore, nutritional studies re-
quire different methods. One of the most useful meth-
ods of testing whether or not an element is essential
is hydroponics, which is the cultivation of plants in
aerated water to which mineral salts are added (•Fig-
ure 10-9). Hydroponics has commercial applications
in addition to its scientific use (see Plants and People:
Commercial Hydroponics).
If botanists suspect that a particular element is es-
sential for plant growth, they grow plants in a nutri-
ent solution that contains all known essential elements
except the one in question. If the plants grown in the
absence of that element do not develop normally or
complete their life cycle, the element may be essential.
Additional criteria are used to confirm whether an ele-
ment is essential. For example, botanists must demon-
strate that the element has a direct effect on the plant’s
metabolism and that the element is essential for a wide
variety of plants.

Fertilizers replace essential
elements missing from the soil
Soil is a valuable natural resource on which humans de-
pend for food. Many human activities may generate or
aggravate soil problems. For example, mineral depletion
may occur in soils that are farmed.
In a natural ecosystem, essential minerals removed
from the soil by plants are returned when the plants or
the animals that have eaten the plants die and are de-
composed. An agricultural system disrupts this pattern
of nutrient cycling when the crops are harvested. Plant
material containing minerals is removed from the nutri-
ent cycle, and the harvested crops fail to decay and re-
lease their nutrients back to the soil. Over time, soil that
is farmed inevitably loses its fertility, that is, its ability
to produce abundant crops (•Figure 10-10). Homeown-
ers often mow their lawns and remove the clippings,
similarly preventing decomposition and cycling of min-
erals that were in the grass blades (see Plants and People:
 Soil • 209

PROCESS OF SCIENCE Managing Your Lawn and Garden at

Home on page 203).
The essential material (water,
QUESTION: How does nitrogen deficiency affect a crop plant such as a tomato?
sunlight, or some essential element)
HYPOTHESIS: Tomatoes grown in the absence of nitrogen exhibit recognizable that is in shortest supply usually lim-
symptoms.
its plant growth. This phenomenon is
EXPERIMENT: The plants are grown in a liquid solution of mineral nutrients through sometimes called the concept of limit-
which air is bubbled to allow the roots to respire. The liquid solution for the ex-
ing resources. The three elements that
perimental plants contains all known nutrient minerals except nitrogen. The liquid
solution for control plants contains all known nutrient minerals, including nitrogen.
are most often limiting resources for
plants are nitrogen, phosphorus, and
potassium. To sustain the productivity
of agricultural soils, farmers periodi-
cally add fertilizers to depleted soils
to replace the minerals that limit plant
growth.
The two main types of fertilizers
are organic and commercial inorganic.
Organic fertilizers include such natu-
ral materials as cow manure, crop resi-
dues, bone meal, and compost. Green
manure, a special type of organic fer-
Lawrence Migdale/Photo Researchers, Inc.

tilizer, is a crop such as alfalfa or sweet

clover that is planted and then plowed
into the soil to decompose instead of
being harvested. Frequently, plants
grown as green manure have nitrogen-
fixing bacteria living in root nodules,
thereby increasing the amount of ni-
trogen in the soil. Organic fertilizers
RESULTS AND CONCLUSION: The control plant is on the left, and the experimental are complex, and their exact compo-
plant is on the right. Compared to the control, tomatoes grown in the absence sitions vary. The mineral nutrients in
of nitrogen exhibit retarded growth and have small leaves that are light green to organic fertilizers become available
pale yellow in color. The plants eventually die (not shown).
to plants only as the organic material
decomposes. For that reason, organic
fertilizers are slow acting and long
lasting. Another advantage of organic
fertilizers is that, in ways that are not
completely understood, they change
the types of organisms living in the
soil, sometimes suppressing micro-
organisms that cause certain plant
diseases.
Commercial inorganic fertilizers
are manufactured from chemical com-
pounds, and their exact compositions
are known. Because they are soluble,
they are immediately available to
plants. Commercial inorganic fertil-
izers are available in the soil for only
Edgar E. Webber

a short period (relative to organic fer-

tilizers), because they quickly leach
away, often into groundwater or sur-
face runoff, and pollute the water.
FIGURE 10-9 A hydroponics experiment. Most commercial inorganic fertilizers
210 • CHAPTER 10 Mineral Nutrition and Transport in Plants

PLANTS AND PEOPLE | Commercial Hydroponics

Botanists have long used hydropon-
ics, the practice of growing plants in
an aerated solution of mineral salts, to
determine which elements are essential
for plant growth. Initially, entrepreneurs
hailed hydroponics as the scientific way
to grow plants in places where soil was
poor or unavailable. The expenses in-
volved in commercial cultivation of pro-

Joseph H. Bailey/National Geographic/Getty Images

duce for human consumption, however,
prevented hydroponics from becoming
more than a curiosity. However, techni-
cal improvements have revived interest
in commercial hydroponics (see figure).
Hydroponics has great potential in
several places. It is being tried ex-
perimentally in desert countries in the
Middle East, where the soil is too arid
to support cultivation and water is un-
available for irrigation. When plants are
A greenhouse with lettuce growing hydroponically.
grown hydroponically in greenhouses,
little water is used in comparison with
traditional irrigation methods. Hydro- (Most foods consumed by Americans trenches through which a thin layer of
ponics is also being tried in temperate travel an average of 2100 kilometers, or water and nutrients trickle. Because
climates, particularly to produce crops 1300 miles.) most of the roots are exposed to the air,
in winter. The main disadvantage of hydropon- they get adequate aeration. The nutri-
Hydroponics has other advantages. ics is the expense. Plants grown hydro- ent solution is saved and reused, which
Because it is possible to grow crops ponically must be given a nutrient solu- cuts down on water and mineral costs.
hydroponically under conditions in tion that is continually monitored and Although hydroponics will probably
which disease-causing microorganisms adjusted. Heating and lighting costs are never replace traditional agriculture, it
and insect pests are completely absent, high. Aeration of the roots was a major is a viable alternative in certain situ-
the crops are not exposed to chemical expense until developments such as ations. As new techniques are devel-
pesticides. Hydroponics is also used the nutrient-film technique helped cut oped, hydroponics may become even
to grow crops near their area of use, costs. In the nutrient-film technique, more common.
thus reducing transportation costs. plants are grown in gently sloping

contain the three elements—nitrogen, phosphorus, and
potassium—that are usually the limiting resources in
plant growth. The numbers on fertilizer bags (for exam-
ple, 10-20-20) tell the relative concentrations of each of
the three elements (N, P, and K).
One advantage of inorganic fertilizers over organic
fertilizers is that one knows precisely how much of a
particular element is being applied to the soil. Varying
the relative concentrations of nitrogen, phosphorus, and
potassium causes different growth responses in plants.
When growing a lettuce crop, for example, it is best to
use a fertilizer with a high nitrogen content, because ni-
trogen stimulates vigorous vegetative growth (growth
of leaves, stems, and roots) rather than reproduction
(growth of flowers, fruits, and seeds). The application of
a fertilizer with a high nitrogen content around tomato
plants, however, causes a low production of tomatoes.
Although the roots, stems, and leaves of the tomato
plants grow vigorously with a high-nitrogen fertilizer,
reproduction is not stimulated; the lush plants form few
flowers and therefore few fruits.
It is environmentally sound to avoid or limit the
use of manufactured fertilizers, for several reasons.
First, because of their high solubility, inorganic fertiliz-

Crop plant
Dead
organic
material
Absorption of
minerals by roots
FIGURE 10-10 Why a soil that is farmed needs fertilizer.
As plant and animal remains decompose in natural environments, nutrients are cycled back
to the soil for reuse. In agriculture, much of the plant material is harvested. Because the
mineral nutrients in the harvested portions are unavailable to the soil, the nutrient cycle is
broken. For this reason, fertilizer is added to the soil periodically.
ers cause water pollution. Second, manufactured fertil-
izers do not improve the water-holding capacity of the
soil as organic fertilizers do. Third, the manufacture of
inorganic fertilizers requires a great deal of fossil fuels
and is therefore not energy conserving.
Transport in the
Plant Body
Now that we have discussed soil and its importance to
plants, we examine internal transport in the vascular
system of the plant (•Figure 10-11). Roots obtain water
and dissolved minerals from the soil. Once inside roots,
these materials are transported upward to stems, leaves,
flowers, fruits, and seeds. Furthermore, sugar mole-
cules manufactured in the leaves by photosynthesis are
transported in solution (dissolved in water) throughout
Transport in the Plant Body • 211
Minerals removed with
harvested plant material
Minerals added as
inorganic fertilizer
Minerals in soil
Mineral loss
from leaching
the plant, including the subterranean roots. Water and
dissolved minerals are transported from roots to other
parts of the plant in xylem, whereas dissolved sugar is
translocated in phloem.
Xylem transport and phloem translocation do not
resemble the movement of materials in animals because
in plants nothing circulates in a system of vessels. Water
and minerals transported in xylem travel in one direc-
tion only (upward), whereas translocation of dissolved
sugar may occur upward or downward in separate
phloem cells. In addition, xylem transport and phloem
translocation differ from internal transport in animals
because movement in both xylem and phloem is driven
largely by natural physical processes rather than by a
pumping organ, or heart.
How, exactly, do materials travel in the continuous
system of the plant’s vascular tissues? First we examine
water and its movement through the plant, and later we
discuss the translocation of dissolved sugar.
212 • CHAPTER 10 Mineral Nutrition and Transport in Plants

According to the tension–

Sugar molecules
manufactured in leaves by
photosynthesis are
transported in phloem
throughout plant,
including into roots.
FIGURE 10-11 An overview of transport in vascular plants.
Xylem transport is explained on the right side of the figure (start at the bottom and
work upward), whereas phloem transport is on the left side.
cohesion model, transpiration
in the leaves pulls water
Most water that plant
absorbs is transpired into up the stem
atmosphere.
Currently, most botanists consider the
tension– cohesion model, first proposed
in 1896 by Irish botanist Henry H.
Dixon, to be the dominant mechanism of
xylem transport in most plants. Accord-
ing to the tension– cohesion model, also
known as the transpiration– cohesion
model, water is pulled up the plant as a
result of a tension produced at the top
of the plant (•Figure 10-13). The evapo-
Once inside roots, water and minerals rative pull of transpiration causes this
are transported upward in xylem to
stems, leaves, flowers, fruits, and seeds. tension, which resembles sucking a liq-
uid up a straw. Most water loss from
transpiration takes place through the
Roots obtain numerous stomata present on leaf and
water and stem surfaces. The tension extends from
dissolved the leaves, where most transpiration oc-
minerals from
soil. curs, down the stems and into the roots.
It draws water up the stem xylem to leaf
cells that have lost water as a result of
transpiration and pulls water from root
xylem into stem xylem. As water is pulled
upward, additional water from the soil is
drawn into the roots. Thus, the pathway
of water movement is as follows:

Soil ¡ root tissues (epidermis, cortex, and so forth) ¡ root

Transport in Xylem
Water initially moves horizontally into the roots from
the soil, passing through several tissues until it reaches
the xylem (see Chapter 6). Once the water is in the tra-
cheids and vessel elements (see Figure 5-6) of the root
xylem, it travels upward through a continuous network
of xylem cells from root to stem to leaf (•Figure 10-12).
The dissolved minerals are carried along passively in the
water, although their initial uptake from the soil may
be active (discussed shortly). The plant does not expend
any energy of its own to transport water, which moves
as a result of natural physical processes. The movement
of water in the xylem is the most rapid of any movement
of materials in plants (•Table 10-2).
How does water move to the tops of plants? It is
either pushed up from the bottom of the plant or pulled
up to the top of the plant. Although plants use both
mechanisms, current evidence indicates that most water
is transported through the xylem by being pulled to the
top of the plant.
xylem ¡ stem xylem ¡ leaf mesophyll ¡ atmosphere
This upward pulling of water is possible only as
long as there is an unbroken column of water in the
xylem throughout the plant. Water forms an unbroken
column in a plant because of the cohesiveness of water
molecules. Recall from Chapter 2 that water molecules
exhibit cohesion, that is, they are strongly attracted to
one another because of hydrogen bonding. In addition,
the adhesion of water to the walls of the xylem cells,
also the result of hydrogen bonding, is an important fac-
tor in maintaining an unbroken column of water. Thus,
the cohesive and adhesive properties of water enable it
to form a continuous column that is pulled up through
the xylem.
Is the tension– cohesion model powerful enough
to explain the rise of water in the tallest plants? Plant
biologists have calculated that the tension produced by
transpiration is strong enough to pull water upward
150 meters (500 feet) in tubes the diameter of xylem
 Transport in Xylem • 213

Main leaf vein

Small veins

Leaf mesophyll cells

James Mauseth, University of Texas

FIGURE 10-12 Network of veins in leaf tissue.

Transport of materials throughout the plant body involves xylem and phloem, which are found
in leaves as veins that branch extensively. No leaf cell is far from a vein.

vessels. Because the tallest trees are about 117 meters
(375 feet) high, the tension– cohesion model easily ac-
counts for the transport of water.
Root pressure pushes water
in a root up through the stem
In the less important mechanism for water transport,
known as root pressure, water that moves into a plant’s
roots from the soil is pushed up through the xylem to-
ward the top of the plant. Water moves from soil into
root cells by osmosis. The accumulation of water in
root tissues produces a pressure that forces the water up
through the xylem.
Guttation, the forceful release of liquid water
through special openings (hydathodes) in leaves (see
Chapter 8), results from root pressure. Root pressure
exerts an influence in smaller plants, particularly in the
spring when the soil is wet, but it clearly does not cause
water to rise 100 meters (330 feet) or more in the tall-

TENSION– COHESION MODEL The mechanism by which

TABLE 10-2 Xylem and Phloem Transport Rates water and dissolved minerals may be transported in xylem:
in Selected Plants water is pulled upward under tension due to transpiration,
MAXIMUM RATE MAXIMUM RATE while maintaining an unbroken column in xylem because of
IN XYLEM IN PHLOEM cohesion.
PLANT (CM/HR) (CM/HR) TRANSPIRATION Loss of water vapor from a plant’s aerial
Conifer 44,120 48 parts.
Woody eudicot 44,400 120 COHESION The tendency of like molecules to adhere or stick
Herbaceous eudicot/monocot 46,000 168–660 together.
Herbaceous vine 15,000 72
ADHESION The tendency of unlike molecules to adhere to
Source: Adapted from J. D. Mauseth, Botany: An Introduction to Plant Biology,
one another.
2nd ed., Philadelphia, Saunders College Publishing, 1995. ROOT PRESSURE The pressure in xylem sap that occurs as
Note: Xylem and phloem rates are from different plants within each general group a result of water moving into roots from the soil.
and should be used for comparative purposes only.
214 • CHAPTER 10 Mineral Nutrition and Transport in Plants

FIGURE IN FOCUS
Transpiration is the driving force of the tension– cohesion model.

•
•••••••
••••

•••
••••
••
••
••
•••
••
Stoma

•
••
••
••
11Water vapor transpires

•••
•••
Water from the surfaces of leaf

•••••••••••
molecules mesophyll cells to the
drier atmosphere through
stomata. This produces a

•••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••
tension that pulls water
out of leaf xylem toward
mesophyll cells.
••
••
••
•••
•••

22 Cohesion of water molecules,

caused by hydrogen bonding,

allows unbroken columns of water
to be pulled up narrow vessels
and tracheids of stem xylem.

3 This, in turn, pulls water up root

xylem, forming a continuous
column of water from root xylem to
stem xylem to leaf xylem. Move-
ment of water upward in root
•••••••••••••

produces a pull that causes soil

water to diffuse into the root.
••

••••
•••••
•••
••••
••
•••
•••
•••
•
••
•••
••••••••

Root
hair

FIGURE 10-13 The tension– cohesion model.

To follow the figure, start at the top and work downward.

est plants. Furthermore, root pressure does not occur to Roots selectively absorb minerals
any appreciable extent in summer (when water is often Minerals are available to plants as ions dissolved in soil
not plentiful in the soil), yet water transport is greatest water. Because the concentrations of various mineral
during hot summer days. ions in xylem sap (fluid in the xylem) are different from

those in soil water, plants appear to selectively accumu-
late the mineral ions they require. Most minerals prob-
ably enter the root by passing from one cell’s cytoplasm
to the next (the symplast) or along the interconnected
cell walls (the apoplast) until they reach the Casparian
strip of the endodermis (see Chapter 6). At this point
minerals enter the endodermal cells by passing through
their plasma membranes.
Dissolved mineral ions pass through plasma mem-
branes by active transport. In active transport, the
mineral ions move against the concentration gradient
(that is, from an area of low concentration to an area
of high concentration of that mineral) through special
channels in the membrane. One of many reasons root
cells require sugar and oxygen for cellular respiration
is that active transport of mineral ions across biological
membranes requires the expenditure of energy, usually
in the form of ATP. From the endodermis, mineral ions
enter the xylem of the root and are conducted to the rest
of the plant.
Translocation in Phloem
The sugar produced during photosynthesis is converted
into sucrose before being loaded into the phloem and
translocated to the rest of the plant. Sucrose is the pre-
dominant photosynthetic product carried in phloem.
Phloem sap also contains much smaller amounts of
other materials, such as amino acids, organic acids, pro-
teins, hormones, certain minerals, and sometimes dis-
ease-causing plant viruses. Translocation in phloem is
not as rapid as xylem transport (see Table 10-2).
Fluid within phloem tissue moves both upward and
downward. Sucrose is translocated in individual sieve
tubes from a source, an area of excess sugar supply (usu-
ally a leaf), to a sink, an area of storage (as starch) or
sugar use such as a root, an apical meristem, a fruit, or
a seed.
The pressure–flow hypothesis
explains translocation in phloem
Current experimental evidence supports the transloca-
tion of dissolved sugar in phloem by the pressure–flow
hypothesis, first proposed in 1930 by the German bota-
nist Ernst Münch. The pressure–flow hypothesis states
that dissolved sugar moves in phloem because of a pres-
sure gradient—that is, a difference in pressure. The pres-
sure gradient exists between the source, where the sugar
is loaded into the phloem, and the sink, where the sugar
is removed from the phloem.
Translocation in Phloem • 215
At the source, the dissolved sucrose moves from a
leaf’s mesophyll cells, where it was manufactured, into
the companion cells, which load it into the sieve-tube el-
ements of phloem. This loading occurs by active trans-
port, a process that requires ATP (•Figure 10-14). The
ATP supplies energy to pump protons out of the sieve-
tube elements, producing a proton gradient that drives
the uptake of sugar into the sieve-tube elements. The
sugar therefore accumulates in the sieve-tube elements
at the source. As a result of the increase in dissolved
sugars in the sieve-tube elements, water moves by osmo-
sis from the xylem cells into the sieve tubes, increasing
the turgor pressure (hydrostatic pressure) inside them.
Thus, phloem loading at the source is as follows:
Proton pump moves H⫹ out of sieve-tube element ¡ sugar is
actively transported into sieve-tube element ¡ water diffuses
from xylem into sieve-tube element ¡ turgor pressure increases
within sieve tube
At its destination (the sink), sugar is unloaded
by various methods, both active and passive, from the
sieve-tube elements. With the loss in sugar, water moves
out of the sieve tubes by osmosis and into surrounding
cells. Most of this water diffuses back to the xylem and
is transported upward again. This water movement de-
creases the turgor pressure inside the sieve tubes at the
sink. Thus, phloem unloading at the sink proceeds as
follows:
Sugar is transported out of sieve-tube element ¡ water diffuses
out of sieve-tube element and into xylem ¡ turgor pressure
decreases within sieve tube
The pressure–flow hypothesis explains the move-
ment of dissolved sugar in phloem because of a pressure
gradient. The difference in sugar concentrations between
the source and the sink causes translocation in phloem,
as water and dissolved sugar flow along the pressure gra-
dient. This pressure gradient pushes the sugar solution
en masse through the phloem much as water is forced
ACTIVE TRANSPORT The energy-requiring movement of a
substance across a membrane from a region of lower concen-
tration to a region of higher concentration.
PRESSURE– FLOW HYPOTHESIS The mechanism by which
dissolved sugar may be transported in phloem; caused by a
pressure gradient between the source (where sugar is loaded
into the phloem) and the sink (where sugar is removed from
the phloem).
TURGOR PRESSURE The internal pressure of water against
the cell wall.
216 • CHAPTER 10 Mineral Nutrition and Transport in Plants

FIGURE IN FOCUS
In phloem, sucrose moves from sources to sinks.

4 Water exits by
osmosis (blue
arrows) and
enters the xylem,
where transpira-
tion pulls it to the
leaves again.

2 The increase in
pressure forces the
fluid through the sieve
tube toward the root.

1 At the source
(leaf), companion
cells actively load
sugars (red arrows)
into sieve-tube
3 At the sink (root), sugar is
elements. Water
flows osmotically actively and/or passively
(blue arrows). loaded (red arrows) into root
cells such as parenchyma
cells in the root cortex.

FIGURE 10-14 The pressure– flow hypothesis.

Sugar is actively loaded into the sieve-tube element at the source. As a result, water moves
into the sieve-tube element. At the sink, the sugar is actively or passively unloaded, and wa-
ter moves from the sieve-tube element to the xylem. The pressure gradient within the sieve
tube, from source to sink, causes translocation from the area of higher turgor pressure (the
source) to the area of lower turgor pressure (the sink).

through a hose. Such pressure-driven flow of a solution
is known as bulk fl ow, or mass fl ow.
The actual translocation of dissolved sugar in the
phloem does not require metabolic energy. However,
the loading of sugar at the source and the active unload-
ing of sugar at the sink require energy derived from
ATP to move the sugar across cell membranes by active
transport.
PROCESS OF SCIENCE Although the pressure–flow hypoth-
esis adequately explains current data on phloem trans-
location, much remains to be learned about this process.
Phloem translocation is difficult to study in plants. Be-
cause phloem cells are under pressure, cutting into the
phloem to observe it relieves the pressure and causes the
contents of the sieve-tube elements (the phloem sap) to
exude and mix with the contents of other severed cells
that are also unavoidably cut.
In the 1950s botanists began to use a unique re-
search tool to avoid contaminating the phloem sap:
aphids, which are small wingless insects that insert their
mouthparts into phloem sieve tubes for feeding (•Fig-
ure 10-15). The pressure in the punctured phloem drives
the sugar solution through the aphid’s mouthpart and
into its digestive system. When the aphid’s mouthpart is
 Study Outline • 217

PROCESS OF SCIENCE

QUESTION: How can phloem sap be studied without cutting non-phloem cells that would
contaminate the sap?
HYPOTHESIS: An aphid mouthpart can be used to penetrate a single sieve-tube element.
EXPERIMENT: After allowing aphids to insert their mouthparts into phloem of a stem,
researchers anesthetized the feeding aphids with CO2 and used a laser to cut their bodies
from their mouthparts. The mouthparts remained in the phloem and functioned like minia-
ture pipes.

© 2007 by the American Association for the Advancement of Science

© Dr. Dennis Kunkel/Visuals Unlimited

Mouthpart Sieve-tube
element

Mouthpart

(a) Aphid feeding on a bean leaf; the aphid’s mouthpart (red) (b) LM of phloem cells, showing an aphid mouthpart
is penetrating a vein. penetrating a sieve-tube element.

RESULTS AND CONCLUSION: About 1 mm3 of phloem sap exuded from each severed mouth-
part per hour for several days, so researchers were able to collect and analyze the composi-
tion of the sap.

FIGURE 10-15 Collecting and analyzing phloem sap.

severed from its body by a laser beam, the sugar solution The identity and proportions of translocated sub-
continues to flow through the mouthpart at a rate pro- stances are also determined using severed aphid mouth-
portional to the pressure in the phloem. This rate can be parts. This technique has verified that in most plant
measured, and the effects on phloem transport of differ- species, the sugar sucrose is the primary carbohydrate
ent environmental conditions—varying light intensities, transported in phloem; however, some species transport
darkness, and mineral deficiencies, for example—can be other sugars, such as raffinose, or sugar alcohols, such
determined. as sorbitol.

ST U DY OU T LI N E
䊉
1 List the four components of soil, and give the ecologi- mineral particles (sand, silt, and clay) come from weath-
cal significance of each. ered parent material and constitute most of what we call
Four distinct components compose soil: inorganic min- soil. Organic matter consists of litter, droppings, and the
eral particles, organic matter, water, and air. Inorganic remains of dead plants, animals, and microorganisms
218 • CHAPTER 10 Mineral Nutrition and Transport in Plants
that are in various stages of decomposition. Pore spaces
of different sizes around and among the soil particles are
filled with varying proportions of soil air and soil water;
plants and other soil-dwelling organisms need soil miner-
als, soil air, and soil water.
䊉
2 Describe the factors involved in soil formation.
Soils form from rock, called parent material, that is grad-
ually fragmented into smaller and smaller particles by
biological, chemical, and physical weathering processes.
Two factors that work together in the weathering of
rock are climate and organisms. Topography, a region’s
surface features, is also involved in soil formation. Steep
slopes often have little or no soil on them because grav-
ity continually transports soil and rock down the slopes;
runoff from precipitation tends to amplify erosion on
steep slopes. The disintegration of solid rock into small
mineral particles and the accumulation of organic mate-
rial take an extremely long time, sometimes thousands
of years.
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3 Distinguish between macronutrients and micronutri-
ents, and list the essential elements.
Ten essential elements are required in fairly large quan-
tities and are known as macronutrients: carbon, hydro-
gen, oxygen, nitrogen, potassium, calcium, magnesium,
phosphorus, sulfur, and silicon. Nine micronutrients are
needed in very small amounts for normal plant growth
and development: chlorine, iron, boron, manganese,
sodium, zinc, copper, nickel, and molybdenum.
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4 Discuss tension– cohesion and root pressure as mecha-
nisms to explain the rise of water in xylem.
In the tension– cohesion model, water is pulled up the
plant as a result of a tension produced at the top of the
plant by the evaporative pull of transpiration, the loss of
water vapor from the aerial parts of plants. The tension
draws water up the stem xylem to leaf cells that have
lost water as a result of transpiration and pulls water
REVIEW QU E STIONS
1. List the four components of soil, and tell how each
is important to plants.
2. Explain how weathering processes convert rock
to soil.
3. What criteria have biologists used to determine
which elements are essential for plant growth?
4. Contrast macronutrients and micronutrients.
from root xylem into stem xylem. As water is pulled
upward, soil water is drawn into the roots. This upward
pulling of water is possible only as long as there is an
unbroken column of water in the xylem throughout the
plant. Water forms an unbroken column because of the
cohesion among water molecules, which are strongly
attracted to one another by hydrogen bonding. The
adhesion of water to the walls of xylem cells also helps
maintain an unbroken column of water. In the less
important mechanism for water transport, known as
root pressure, water that moves into a plant’s roots from
the soil is pushed up through the xylem toward the top
of the plant; the accumulation of water in root tissues
produces a pressure that forces the water up through the
xylem.
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5 Discuss the pressure– flow hypothesis of sugar trans-
location in phloem.
The pressure–flow hypothesis states that dissolved sugar
moves in phloem because of a pressure gradient—that
is, a difference in pressure. The pressure gradient exists
between the source, where the sugar is loaded into the
phloem, and the sink, where the sugar is removed from
the phloem. At the source, the dissolved sucrose moves
from a leaf’s mesophyll cells, where it was manufac-
tured, into the companion cells, which load it into the
sieve-tube elements of phloem by active transport,
a process that requires ATP. As a result of the increase
in dissolved sugars in the sieve-tube elements, water
moves by osmosis from the xylem cells into the sieve
tubes, increasing the turgor pressure (hydrostatic pres-
sure) inside them. At the sink, sugar is unloaded from
the sieve-tube elements, and water moves out of the
sieve tubes by osmosis and into surrounding cells. Most
of this water diffuses back to the xylem. This water
movement decreases the turgor pressure inside the sieve
tubes at the sink.
5. Give a role for each of the following essential ele-
ments: nitrogen, magnesium, phosphorus, potas-
sium, and calcium.
6. Compare organic and commercial inorganic
fertilizers.
7. Briefly describe how the tension– cohesion model
explains the rise of water in the tallest trees.