North Korea experienced a food crisis after losing support from the Soviet Union, which had previously provided chemicals and petroleum needed for agriculture. Without these imports, North Korea could no longer produce sufficient fertilizer, which is essential for supplying nitrogen and other nutrients to crops. Nitrogen is the nutrient most often limiting for plant growth, and industrial fertilizer production requires significant energy from petroleum. As a result, North Korea's farming system failed due to the lack of fertilizer inputs, leading the country to experience widespread starvation.

1. 37 Plant Nutrition
Food is an essential commodity that separates prosperous nations from
struggling ones. For instance, North Korean agriculture met that en-
tire country’s food needs until about a decade ago. The country’s farm-
ers were highly efficient and productive. Its food crisis began with the
collapse of the Soviet Union, which had provided North Korea with chemicals and
petroleum. This loss of support was followed by three years of drought, hailstorms,
and floods. Today, North Korea is a starving country with a failed farming system.
Why should a desperate shortage of chemicals and petroleum affect a nation’s
agriculture? Crop production depends on several factors, but the one that is most
commonly limiting is a supply of nitrogen in a form usable by plants. All plants re-
quire the element nitrogen, which is an abundant component of proteins and nucleic
acids as well as chlorophyll and many other important biochemical compounds. If a
plant cannot get enough nitrogen, it cannot synthesize these compounds at a rate ad-
equate to keep itself healthy. To meet their crops’ need for nitrogen and other miner-
als, farmers in all parts of the world apply fertilizers of one kind or another. The in-
Nitrogen is Essential for Plant Growth
dustrial production of fertilizers is an energy-intensive process, and the energy In this experimental wheat field in
needed is most commonly obtained from petroleum. Without petroleum, North Bangladesh, nitrogen was withheld from
Korea cannot begin to provide the fertilizer needed to restore its crop production. the plot on the left. The resulting plants
were stunted and unhealthy.
In addition to nitrogen, plants need other
materials from their environment. In this
chapter, we will explore the differences be-
tween the basic strategies of plants and of an-
imals for obtaining nutrition. Then we will
look at what nutrients plants require and how
they acquire them. Because most nutrients
come from the soil, we will discuss the for-
mation of soils and the effects of plants on
soils. As any farmer can tell you, nitrogen is
the nutrient that most often limits plant
growth, so we will devote a section specifi-
cally to nitrogen metabolism in plants. The
chapter concludes with a look at carnivorous
and parasitic plants, which supplement their
nutrition in special ways.
The Acquisition of Nutrients
Every living thing must obtain raw materials
from its environment. These nutrients include

2. PLANT NUTRITION 717
the major ingredients of macromolecules: carbon, hydrogen, sources that are somehow brought to it. Most sessile animals
oxygen, and nitrogen. Carbon and oxygen enter the living depend primarily on the movement of water to bring them
world in the form of atmospheric carbon dioxide through the raw materials and energy in the form of food, but a plant’s
carbon-fixing reactions of photosynthesis. Hydrogen enters supply of energy arrives at the speed of light from the sun.
living systems through the light reactions of photosynthesis, However, with the exception of carbon and oxygen in CO2,
which split water. For carbon, oxygen, and hydrogen, pho- a plant’s supply of nutrients is strictly local, and the plant
tosynthesis is the gateway to the living world, and these may use up the water and mineral nutrients in its local envi-
elements are in plentiful supply. ronment as it develops. How does a plant cope with the
In the remainder of this chapter, we shall focus our atten- problem of scarce nutrient supplies?
tion on nitrogen, which is in relatively short supply for One way is to extend itself by growing in search of new
plants. The movement of nitrogen into organisms begins resources. Growth is a plant’s version of movement. Among
with processing by some highly specialized bacteria living in plant organs, the roots obtain most of the mineral nutrients
the soil. Some of these bacteria act on nitrogen gas, convert- needed for growth. By growing through the soil, they mine
ing it into a form usable by plants. The plants, in turn, pro- the soil for new sources of mineral nutrients and water. The
vide organic nitrogen (and carbon) to animals, fungi, and growth of leaves helps a plant secure light and carbon diox-
many microorganisms. ide. A plant may compete with other plants for light by out-
In addition to nitrogen, other mineral nutrients are es- growing and shading them.
sential to living organisms. The proteins of organisms con- As it grows, a plant—or even a single root—must deal with
tain sulfur (S), and their nucleic acids contain phosphorus (P). a variable environment. Animal droppings create high local
There is magnesium (Mg) in chlorophyll, and iron (Fe) in concentrations of nitrogen. A particle of calcium carbonate in
many important compounds, such as the cytochromes. the soil may make a tiny area alkaline, while dead organic mat-
Within the soil, these and other minerals dissolve in water, ter may make a nearby area acidic. Such microenvironments
forming a solution—called the soil solution—that contacts encourage or discourage the proliferation of a root system.
the roots of plants. Plants take up most of these mineral nu-
trients from the soil solution in ionic form.
Mineral Nutrients Essential to Plants
As roots grow through the soil, what important mineral nu-
Autotrophs make their own organic compounds trients do plants take up from their environment, and what
Plants, some protists, and some bacteria are autotrophs; that are the roles of those nutrients? Table 37.1 lists the mineral
is, they make their own organic (carbon-containing) com- nutrients that have been determined to be essential for plants.
pounds from simple inorganic nutrients—carbon dioxide, Except for nitrogen, they all derive from rock. All of them are
water, nitrogen-containing ions, and a few other soluble min- usually taken up from the soil solution.
eral nutrients. The plants provide carbon, oxygen, hydrogen, There are three criteria for calling something an essential
nitrogen, and sulfur to most of the rest of the living world. element:
Heterotrophs are organisms that require preformed organic
The element must be necessary for normal growth and
compounds as food. All heterotrophs depend directly or in-
reproduction.
directly on autotrophs as their source of nutrition.
The element cannot be replaceable by another element.
Most autotrophs are photosynthesizers—that is, they use
The requirement must be direct—that is, not the result of
light as their source of energy for synthesizing organic com-
an indirect effect, such as the need to relieve toxicity
pounds from inorganic raw materials. Some autotrophs,
caused by another substance.
however, are chemosynthesizers, deriving their energy not
from light, but from reduced inorganic substances, such as In this section, we’ll consider the symptoms of particular
hydrogen sulfide (H2S), in their environment. All chemosyn- mineral deficiencies, the roles of some of the mineral nutri-
thesizers are bacteria. As we’ll see below, some chemosyn- ents, and the technique by which the essential elements for
thetic bacteria in the soil contribute to the nutrition of plants plants were identified.
by increasing the availability of nitrogen and sulfur. There are two categories of essential elements: macronu-
trients and micronutrients (see Table 37.1).
How does a stationary organism find nutrients? Plants need macronutrients in concentrations of at least
Many heterotrophs can move from place to place to find the 1 gram per kilogram of their dry matter.*
nutrients they need. An organism that cannot move, termed *Dry matter, or dry weight, is what remains after all the water has been
a sessile organism, must obtain nutrients and energy from removed from a plant tissue sample.

3. 718 CHAPTER THIRT Y-SEVEN
37.1 Mineral Elements Required by Plants
ELEMENT ABSORBED FORM MAJOR FUNCTIONS
Macronutrients
Nitrogen (N) NO3– and NH4+ In proteins, nucleic acids, etc.
Phosphorus (P) H2PO4– and HPO4
2–
In nucleic acids, ATP, phospholipids, etc.
+
Potassium (K) K Enzyme activation; water balance; ion balance; stomatal opening
2–
Sulfur (S) SO 4 In proteins and coenzymes
2+
Calcium (Ca) Ca Affects the cytoskeleton, membranes, and many enzymes; second messenger
Magnesium (Mg) Mg2+ In chlorophyll; required by many enzymes; stabilizes ribosomes
Micronutrients
Iron (Fe) Fe2+ In active site of many redox enzymes and electron carriers; chlorophyll synthesis
Chlorine (Cl) Cl– Photosynthesis; ion balance
Manganese (Mn) Mn2+ Activation of many enzymes
Boron (B) B(OH)3 Possibly carbohydrate transport (poorly understood)
Zinc (Zn) Zn2+ Enzyme activation; auxin synthesis
Copper (Cu) Cu2+ In active site of many redox enzymes and electron carriers
Nickel (Ni) Ni2+ Activation of one enzyme
2–
Molybdenum (Mo) MoO4 Nitrate reduction
Nitrogen deficiency is not the only cause of chlorosis. In-
Plants need micronutrients in concentrations of less
adequate iron in the soil can also cause chlorosis because
than 100 milligrams per kilogram of their dry matter.
iron, although it is not contained in the chlorophyll molecule,
These two categories differ only with regard to the amounts is required for chlorophyll synthesis. However, iron defi-
required by plants. Both the macronutrients and the mi- ciency commonly causes chlorosis of the youngest leaves,
cronutrients are essential for the plant to complete its life cy- with their veins sometimes remaining green. The reason for
cle from seed to seed. How do we know if a plant is getting this difference is that nitrogen is readily translocated in the
enough of a particular nutrient? plant and can be redistributed from older tissues to younger
tissues to favor their growth. Iron, on the other hand, cannot
Deficiency symptoms reveal inadequate nutrition
Before a plant that is deficient in an essential element dies, it
usually displays characteristic deficiency symptoms, such as 37.2 Some Mineral Deficiencies in Plants
discoloration or deformation of its leaves. Table 37.2 describes DEFICIENCY SYMPTOMS
the symptoms of some common mineral deficiencies. Such
symptoms help horticulturists diagnose mineral nutrient de- Calcium Growing points die back; young leaves
are yellow and crinkly
ficiencies in plants. With proper diagnosis, appropriate treat-
Iron Young leaves are white or yellow with green
ment can be applied in the form of a fertilizer (an added veins
source of mineral nutrients).
Magnesium Older leaves have yellow in stripes between
Nitrogen deficiency is the most common mineral defi- veins
ciency in both natural and agricultural environments. Plants Manganese Younger leaves are pale with stripes of dead
in natural environments are almost always deficient in nitro- patches
gen, but they seldom display deficiency symptoms. Instead, Nitrogen Oldest leaves turn yellow and die pre-
their growth slows to match the available supply of nitrogen. maturely; plant is stunted
Crop plants, on the other hand, show deficiency symptoms if Phosphorus Plant is dark green with purple veins
a formerly abundant supply of nitrogen runs out. The visible and is stunted
symptoms of nitrogen deficiency include uniform yellowing, Potassium Older leaves have dead edges
or chlorosis, of older leaves. Chlorophyll, which is responsible Sulfur Young leaves are yellow to white with
yellow veins
for the green color of leaves, contains nitrogen. Without ni-
trogen there is no chlorophyll, and without chlorophyll, the Zinc Young leaves are abnormally small;
older leaves have many dead spots
yellow carotenoid pigments in the leaves become visible.

4. EXPERIMENT
Question: Is a particular ingredient of a growth medium an
essential plant nutrient?
be readily redistributed. Younger tissues that are actively
growing and synthesizing compounds needed for their METHOD Grow seedlings in a medium that lacks the element
in question (in this case, nitrogen)
growth show iron deficiency before older leaves, which have
already completed their growth.
Several essential elements fulfill multiple roles
Essential elements may play several different roles in plant Seedling grown Seedling grown
cells—some structural, others catalytic. Magnesium, as we in a complete in a medium
growth medium. lacking nitrogen.
have mentioned, is a constituent of the chlorophyll molecule
and hence is essential to photosynthesis. It is also required as
a cofactor by numerous enzymes involved in cellular respi-
ration and other metabolic pathways. RESULTS
Phosphorus, usually in phosphate groups, is found in
many organic compounds, particularly in nucleic acids and
in the intermediates of the energy-harvesting pathways of
photosynthesis and glycolysis. As we saw in Chapter 7, the
transfer of phosphate groups occurs in many energy-storing
and energy-releasing reactions, notably those that use or pro-
duce ATP. The addition or removal of phosphate groups is Growth is normal. Growth is abnormal.
also used to activate or inactivate enzymes.
Calcium plays many roles in plants. Its function in the pro- Conclusion: Nitrogen is an essential plant nutrient.
cessing of hormonal and environmental cues is a subject of
great biological interest, as we’ll see in the next chapter. Cal- 37.1 Identifying Essential Elements for Plants This dia-
cium also affects membranes and cytoskeletal activity, partic- gram shows the procedure for identifying nutrients essential
to plants, using nitrogen as an example.
ipates in spindle formation for mitosis and meiosis, and is a
constituent of the middle lamella of cell walls. Other elements,
such as iron and potassium, also play multiple roles in plants.
All of these elements are essential to the life of all plants. that they provided micronutrients that the investigators
How did biologists discover which elements are essential? thought they had excluded. Furthermore, because some mi-
cronutrients are required in such tiny amounts, there may be
enough in a seed to supply the embryo and the resultant sec-
Experiments were designed to identify ond-generation plant throughout its lifetime and leave
essential elements enough in the next seed to get the third generation well
An element is considered essential to plants if a plant fails to started. Indeed, simply touching a plant may give it a signif-
complete its life cycle, or grows abnormally, when that element icant supply of chlorine in the form of chloride ions from
is not available, or is not available in sufficient quantities. The sweat. Such difficulties make it necessary to perform nutri-
essential elements for plants were identified by growing plants tion experiments in tightly controlled laboratories with spe-
hydroponically—that is, with their roots suspended in nutrient cial air filters (to exclude microscopic salt particles in the air)
solutions without soil (Figure 37.1). In the first successful ex- and to use only chemicals that had been purified to the high-
periments of this type, performed a century and a half ago, est degree attainable by modern chemistry. Only rarely are
plants grew seemingly normally in solutions containing only new essential elements reported now. Either the list is nearly
calcium nitrate, magnesium sulfate, and potassium phosphate. complete, or perhaps, we will need more sophisticated tech-
Omission of any of these compounds made the solution inca- niques to add to it.
pable of supporting normal growth. Tests with other com- Where does the plant find its essential mineral nutrients?
pounds including these elements soon established the six How does it absorb them?
macronutrients— calcium, nitrogen, magnesium, sulfur, potas-
sium, and phosphorus—as essential elements.
Identifying essential elements by this experimental ap-
Soils and Plants
proach proved to be a more difficult task in the case of the Most terrestrial plants live their lives anchored to the soil. Of
micronutrients. In the nineteenth-century experiments on course, soils offer mechanical support for growing plants, but
plant nutrition, some of the chemicals used were so impure there are many other plant-soil interactions, some of which

5. 720 CHAPTER THIRT Y-SEVEN
are much more complex. Plants obtain their mineral nutri-
ents from the soil solution. Water for terrestrial plants also
A horizon
comes from the soil, as does the supply of oxygen for the
Topsoil
roots. Soil harbors bacteria, some of which are beneficial to
plant life. Soils may also contain organisms harmful to plants.
In this section, we will examine the composition, structure,
B horizon
and formation of soils. We will consider their role in plant nu- Subsoil
trition, their care and supplementation in agriculture, and
their modification by the plants that grow in them.
Soils are complex in structure
Soils are complex systems made up of living and nonliving C horizon
Weathering
components. The living components include plant roots as parent rock
well as populations of bacteria, fungi, protists, and animals (bedrock)
such as earthworms and insects (Figure 37.2). The nonliving
portion of the soil includes rock fragments ranging in size
from large rocks through sand and silt and finally to tiny par-
ticles called clay that are 2 µm or less in diameter. Soil also
contains water and dissolved mineral nutrients, air spaces, 37.3 A Soil Profile The A, B, and C horizons can sometimes be seen
and dead organic matter. The air spaces are crucial sources in road cuts such as this one in Australia. The dark upper layer (the A
of oxygen (in the form of O2) for plant roots. The character- horizon) is home to most of the living organisms in the soil.
istics of soils are not static. Soils change constantly through
natural phenomena such as rain, temperature extremes, and
the activities of plants and animals, as well as human activi- horizons, lying on top of one another. Mineral nutrients tend
ties—agriculture in particular. to be leached from the upper horizons—dissolved in rain or
The structure of many soils changes with depth, revealing irrigation water and carried to deeper horizons, where they
a soil profile. Although soils differ greatly, almost all soils con- are unavailable to plant roots.
sist of two or more recognizable horizontal layers, called Soil scientists recognize three major horizons (A, B, and C)
in the profile of a typical soil (Figure 37.3). Topsoil is the A
horizon, from which mineral nutrients may be depleted by
leaching. Most of the dead and decaying organic matter in
the soil is in the A horizon, as are most plant roots, earth-
Organic matter (from plants, worms, insects, nematodes, and microorganisms. Successful
animals, and fungi) agriculture depends on the presence of a suitable A horizon.
Topsoils are composed of different proportions of sand,
silt, and clay. In pure sand there are abundant air spaces be-
tween the relatively large particles, but sand is low in water
and mineral nutrients. Clay contains many mineral nutrients
and more water than sand does, but the tiny clay particles
Air pack tightly together, leaving little space to trap air. A little
Bacteria bit of clay goes a long way in affecting soil properties. A loam
is a soil that has significant amounts of sand, silt, and clay,
and thus has sufficient levels of air, water, and nutrients for
Quartz Air plants. Loams also contain organic matter. Most of the best
Air
H2O topsoils for agriculture are loams.
Air and water Below the A horizon is the B horizon, or subsoil, which is
the zone of infiltration and accumulation of materials leached
Aggregates of
from above. Farther down, the C horizon is the parent rock
clay particles 25 µm that is breaking down to form soil. Some deep-growing roots
37.2 The Complexity of Soil Even a tiny crumb of soil has both extend into the B horizon to obtain water and nutrients, but
organic and inorganic components. roots rarely enter the C horizon.

6. PLANT NUTRITION 721
A clay particle, which is negatively
Soils form through the weathering of rock charged, binds cations.
The type of soil in a given area depends on the type of par-
Root hair
ent rock from which it formed, the climate, the landscape
features, the organisms living there, and the length of time
H+
that soil-forming processes have been acting (sometimes mil-
lions of years). Rocks are broken down into soil in part by K+
mechanical weathering, which is the physical breakdown—
without any accompanying chemical changes—of materials H+ – – – –
– Clay –
by wetting, drying, and freezing. The most important parts –– – – –
H+
of soil formation, however, include chemical weathering, the
chemical alteration of at least some of the materials in the
rocks. CO2 + H2O H2CO3 HCO3– + H+
Both the physical and chemical properties of soils depend
on the amounts and kinds of clay particles they contain.
These tiny particles, which bind mineral nutrients and ag- The cations are exchanged for hydrogen ions obtained
from carbonic acid (H2CO3 ) or from the plant itself.
gregate into larger particles, are extremely important to plant
growth. Clay is not produced merely by the mechanical 37.4 Ion Exchange Plants obtain mineral nutrients from the soil
grinding up of rocks. In addition to mechanical weathering, primarily in the form of positive ions; potassium is the example
shown here.
several types of chemical weathering are required:
Oxidation by atmospheric oxygen makes some essential
elements more available to plants.
Reaction with water (hydrolysis) releases some mineral
plant growth, called soil fertility, is determined in part by its
nutrients from the rock.
ability to provide nutrients in this manner.
Acids, carbonic acid in particular, free some essential ele-
Clay particles effectively hold and exchange cations, and
ments from their parent salts.
cations tend to be retained in the A horizon. However, there
These reactions leave the surface of clay particles with an is no comparable mechanism for exchanging anions, the
abundance of negatively charged chemical groups, to which negatively charged ions. As a result, important anions such
certain mineral nutrients bind. Let’s see how roots take up as nitrate (NO3–) and sulfate (SO42–)—the primary and direct
these mineral nutrients from clay particles. sources of nitrogen and sulfur, respectively—leach rapidly
from the A horizon. As a consequence of this leaching, the
primary soil reservoir of nitrogen is not in the form of nitrate
Soils are the source of plant nutrition ions. Most of the nitrogen in the A horizon is found in the
The availability of mineral nutrients to plant roots depends organic matter in the soil, which slowly decomposes to re-
on the presence of clay particles in the soil. The negatively lease nitrogen in a form that can be absorbed and used by
charged clay particles bind the cations of many minerals plants.
that are important for plant nutrition, such as potassium
(K+), magnesium (Mg2+), and calcium (Ca2+). To become
available to plants, these cations must be detached from the Fertilizers and lime are used in agriculture
clay particles. Agricultural soils often require fertilizers because irrigation
This task is accomplished by reactions with protons (hy- and rainwater leach mineral nutrients from the soil and be-
drogen ions, H+). These protons are released into the soil by cause the harvesting of crops removes the nutrients that the
roots, which also release CO2 through cellular respiration. plants took up from the soil during their growth. Crop yields
The CO2 dissolves in the soil water and reacts with it to form decrease if any essential element is depleted. Mineral nutri-
carbonic acid, which then ionizes to form bicarbonate and ents may be replaced by organic fertilizers, such as rotted
free protons (CO2 + H2O ~ H2CO3 ~ H+ + HCO3–). These manure, or by inorganic fertilizers of various types.
protons bind more strongly to the clay particles than do the
mineral cations, so they trade places with the cations in a ORGANIC AND INORGANIC FERTILIZERS. The three elements
process called ion exchange (Figure 37.4). Ion exchange puts most commonly added to agricultural soils are nitrogen
important cations back into the soil solution, from which they (N), phosphorus (P), and potassium (K). Commercial fertil-
are taken up by the roots. The capacity of a soil to support izers are characterized by their “N-P-K” percentages. A

7. 722 CHAPTER THIRT Y-SEVEN
5-10-10 fertilizer, for example, contains 5 percent nitrogen, Plants affect soil fertility and pH
10 percent phosphate (P2O5), and 10 percent potash (K2O) The soil that forms in a particular place depends on the types
by weight.* Sulfur, in the form of a sulfate, is also occasion- of plants growing there. Plant litter, such as dead fallen
ally added to soils. leaves, is the major source of the carbon-rich materials that
Either organic or inorganic fertilizers can provide the nec- break down to form humus—dark-colored organic material,
essary mineral nutrients for plants. Organic fertilizers release each particle of which is too small to be recognizable with the
nutrients slowly, which results in less leaching than occurs naked eye. Soil bacteria and fungi produce humus by break-
with a one-time application of an inorganic fertilizer. How- ing down plant litter, animal feces, dead organisms, and
ever, the nutrients from organic fertilizers are not immedi- other organic material. Humus is rich in mineral nutrients,
ately available to plants. Organic fertilizers also contain especially nitrogen that was excreted by animals. In combi-
residues of plant or animal materials that improve the struc- nation with clay, humus favors plant growth by trapping
ture of the soil, providing spaces for air movement, root supplies of water and oxygen for absorption by roots. Look-
growth, and drainage. Inorganic fertilizers, on the other ing at the big picture, we see that successful plant growth can
hand, provide a supply of soil nutrients that is almost im- create conditions that support further plant growth.
mediately available for absorption. Furthermore, inorganic Plants also affect the pH of the soil in which they grow.
fertilizers can be formulated to meet the requirements of a Roots maintain a balance of electric charges. If they absorb
particular soil and a particular crop. more cations than anions, they excrete H+ ions, thus lower-
ing the soil pH. If they absorb more anions than cations, they
pH EFFECTS ON NUTRIENTS. The availability of nutrient ions, excrete OH– ions or HCO3– ions, raising the soil pH.
whether they are naturally present in the soil or added as The mineral nutrient most commonly in short supply, in
fertilizer, is altered by changes in soil pH. The optimal soil both natural and agricultural situations, is nitrogen, despite
pH for most crops is about 6.5, but so-called acid-loving the fact that elemental nitrogen makes up almost four-fifths
crops such as blueberries prefer a pH closer to 4. Rainfall of Earth’s atmosphere. What is the reason for this scarcity?
and the decomposition of organic substances lower the pH Let’s consider how nitrogen is made available to plants.
of the soil. Such acidification can be reversed by liming—
the application of compounds commonly known as lime,
such as calcium carbonate, calcium hydroxide, or magne-
Nitrogen Fixation
sium carbonate. The addition of these compounds leads to The Earth’s atmosphere is a vast reservoir of nitrogen in the
the removal of H+ ions from the soil. Liming also increases form of nitrogen gas (N2). However, plants cannot use N2 di-
the availability of calcium to plants. rectly as a nutrient. It is a highly unreactive substance—the
Sometimes, on the other hand, a soil is not acidic enough. triple bond linking the two nitrogen atoms is extremely sta-
In this case, sulfur can be added in the form of elemental sul- ble, and a great deal of energy is required to break it. How,
fur, which soil bacteria convert to sulfuric acid. Iron and some then, is nitrogen made available for the synthesis of proteins
other elements are more available to plants at a slightly acidic and nucleic acids?
pH. Soil pH testing is useful for home gardens and lawns as
well as for agriculture. The test results indicate what amend-
ments should be made to the soil.
SPRAY APPLICATION OF NUTRIENTS. Spraying leaves with a
nutrient solution is another effective way to deliver some
essential elements to growing plants. Plants take up more
copper, iron, and manganese when these elements are
applied as foliar (leaf) sprays than when they are added to
the soil. Such foliar application of mineral nutrients is
increasingly used in wheat production, but fertilizers are
still delivered most commonly by way of the soil.
The relationship between plants and soils is not a one-way
affair—soils affect plants, but plants also affect soils.
*The analysis is by weight of the nutrient-containing compound and not
as weights of the elements N, P, and K. A 5-10-10 fertilizer actually does
contain 5 percent nitrogen, but only 4.3 percent phosphorus and 8.3 per- 37.5 Root Nodules Large, round nodules are visible in the root
cent potassium on an elemental basis. system of a pea plant. These nodules house nitrogen-fixing bacteria.

8. PLANT NUTRITION 723
A few species of bacteria have an enzyme that enables Bacteria of the genus Rhizobium fix nitrogen only in close,
them to convert N2 into a more reactive and biologically use- mutualistic association with the roots of plants in the legume
ful form by a process called nitrogen fixation. These prokary- family. The legumes include peas, soybeans, clover, alfalfa,
otic organisms—nitrogen fixers—convert N2 to ammonia and many tropical shrubs and trees. The bacteria infect the
(NH3). There are relatively few species of nitrogen fixers, and plant’s roots, and the roots develop nodules in response to
their biomass is small relative to the mass of other organisms their presence. The various species of Rhizobium show a high
that depend on them for survival on Earth. This talented specificity for the species of legume they infect. Farmers and
group of prokaryotes is just as essential to the biosphere as gardeners coat legume seeds with Rhizobium to make sure the
are the photosynthetic autotrophs. bacteria are present. Some farmers alternate their crops,
planting clover or alfalfa occasionally to increase the avail-
able nitrogen content of the soil.
Nitrogen fixers make all other life possible The legume–Rhizobium association is not the only bacterial
By far the greatest share of total world nitrogen fixation is association that fixes nitrogen. Some cyanobacteria fix nitro-
performed biologically by nitrogen-fixing prokaryotes, which gen in association with fungi in lichens or with ferns, cycads,
fix approximately 170 million Mg (megagrams or metric or nontracheophytes. Rice farmers can increase crop yields
tons) of nitrogen per year. About 80 million Mg is fixed in- by growing the water fern Azolla, with its symbiotic nitrogen-
dustrially by humans. A smaller amount of nitrogen is fixed fixing cyanobacterium, in the flooded fields where rice is
in the atmosphere by nonbiological means such as lightning, grown. Another group of bacteria, the filamentous actino-
volcanic eruptions, and forest fires. Rain brings these atmos- mycetes, fix nitrogen in association with root nodules on
pherically formed products to the ground. woody species such as alder and mountain lilacs.
Several groups of bacteria fix nitrogen. In the oceans, vari- How does biological nitrogen fixation work? In the four
ous photosynthetic bacteria, including cyanobacteria, fix nitro- sections that follow, we’ll consider the role of the enzyme ni-
gen. In fresh water, cyanobacteria are the principal nitrogen fix- trogenase, the mutualistic collaboration of plant and bacter-
ers. On land, free-living soil bacteria make some contribution ial cells in root nodules, the need to supplement biological
to nitrogen fixation, but they fix only what they need for their nitrogen fixation in agriculture, and the contributions of
own use and release the fixed nitrogen only when they die. plants and bacteria to the global nitrogen cycle.
Other nitrogen-fixing bacteria live in close association with
plant roots (Figure 37.5). They release up to 90 percent of the
nitrogen they fix to the plant and excrete some amino acids Nitrogenase catalyzes nitrogen fixation
into the soil, making nitrogen immediately available to other Nitrogen fixation is the reduction of nitrogen gas. It proceeds
organisms. The plant obtains fixed nitrogen from the bac- by the stepwise addition of three pairs of hydrogen atoms to
terium, and the bacterium obtains the products of photosyn- N2 (Figure 37.6). In addition to N2, these reactions require
thesis from the plant. Such associations are excellent exam- three things:
ples of mutualism, an interaction between two species in
which both species benefit. They are also examples of sym-
biosis, in which two different species live in physical contact 37.6 Nitrogenase Fixes Nitrogen Throughout the chemical reac-
tions of nitrogen fixation, the reactants are bound to the enzyme
for a significant portion of their life cycles. nitrogenase. A reducing agent transfers hydrogen atoms to nitrogen,
and eventually the final product—ammonia—is released.
2 A reducing agent transfers 3 The final products—two molecules
three successive pairs of of ammonia—are released, freeing
1 The enzyme nitrogenase binds the nitrogenase to bind another N2
hydrogen atoms to N2.
a molecule of nitrogen gas. molecule.
Substrate: + 2H + 2H + 2H H H
H
H
Nitrogen gas (N2) N
H N
H
N N
H
H H H
H
H H
H
Product:
N N N N H N N H
H
N N H Ammonia (NH3)
Reduction Reduction Reduction
Nitrogenase
Nitrogenase Binding of
substrate

9. Root hairs Cortical cells Root hair
Rhizobia
1 Root hairs release chemical
signals that attract Rhizobium.
Infection
thread 2 Rhizobium proliferates and
causes an infection thread
to form.
Root tip
3 The infection thread grows
into the cortex of the root.
4 The infection thread releases bacterial
cells, which become bacteroids in the
root cells. Nod factors from bacteria
cause cortical cells to divide.
Nodule
Bacteroids in Uninfected cell
infected cell
Nodule
37.7 A Nodule Forms Rhizobium develops the ability to fix
nitrogen only after entering a legume root. The diagrams show the
sequence of events in nodule formation. The photograph shows
bacteroids of Rhizobium japonicum in vesicles within a soybean
root cell. A portion of an uninfected root cell is seen on the right. Bacteroids 5 The nodule forms from rapidly
dividing, infected cortical cells.
a strong reducing agent to transfer hydrogen atoms to
protein leghemoglobin in the cytoplasm of the nodule cells.
N2 and to the intermediate products of the reaction
Leghemoglobin is a close relative of hemoglobin, the oxy-
a great deal of energy, which is supplied by ATP
gen-carrying pigment of animals. Some plant nodules con-
the enzyme nitrogenase, which catalyzes the reaction
tain enough of it to be bright pink when viewed in cross sec-
(Depending on the species of nitrogen fixer, either respiration tion. Leghemoglobin, with its iron-containing heme groups,
or photosynthesis may provide both the necessary reducing transports enough oxygen to the bacteroids to support their
agent and ATP.) respiration.
Nitrogenase is so strongly inhibited by oxygen that its
presence in biochemical extracts was obscured and its dis-
covery delayed because investigators had not thought to seek Some plants and bacteria work together to fix nitrogen
it under anaerobic conditions. It is therefore not surprising Neither free-living Rhizobium species nor uninfected legumes
that many nitrogen fixers are anaerobes and live in environ- can fix nitrogen. Only when the two are closely associated in
ments with little or no O2. Because this crucial enzyme is so root nodules does the reaction take place. The establishment
inhibited by O2, it was at first surprising that legumes respire of this symbiosis between Rhizobium and a legume requires
aerobically, as do Rhizobium. Investigation of the root nodules a complex series of steps, with active contributions by both
where nitrogenase is found revealed how the enzyme could the bacteria and the plant root (Figure 37.7). First the root re-
operate there. leases flavonoids and other chemical signals that attract soil-
Within a root nodule, O2 is maintained at a low level suf- living Rhizobium to the vicinity of the root. Flavonoids trig-
ficient to support respiration, but not so high as to inactivate ger the transcription of bacterial nod genes, which encode
nitrogenase. The plant makes this possible by producing the Nod (nodulation) factors. These factors, secreted by the bac-

10. PLANT NUTRITION 725
teria, cause cells in the root cortex to divide, leading to the Plants and bacteria participate in the global
formation of a primary nodule meristem. The meristem gives nitrogen cycle
rise to the plant tissue that constitutes the nodule. The nitrogen released into the soil by nitrogen fixers is pri-
Among the products of the meristem is a layer of cells that marily in the form of ammonia (NH3) and ammonium ions
excludes O2 from the interior of the nodule. The function of (NH4+). Although ammonia is toxic to plants, ammonium
leghemoglobin is to carry O2 across this barrier. Within a nod- ions can be taken up safely at low concentrations. Soil bacte-
ule, the bacteria take the form of bacteroids within membra- ria called nitrifiers, which we described in Chapter 27, oxidize
nous vesicles. Bacteroids are swollen, deformed bacteria that ammonia to nitrate ions (NO3–)—another form that plants
can fix nitrogen—in effect, nitrogen-fixing organelles. can take up—by the process of nitrification (Figure 37.8). Soil
The partnership between bacterium and plant in nitrogen- pH affects the uptake of nitrogen: Nitrate ions are taken up
fixing nodules is not the only case in which plants depend on preferentially under more acidic conditions, ammonium ions
other organisms for assistance with their nutrition. Another under more basic ones.
example is that of mycorrhizae, root–fungus associations in The steps that we have followed so far are carried out by
which the fungus greatly increases the absorption of water bacteria: N2 is reduced to ammonia in nitrogen fixation and
and minerals (especially phosphorus) by the plant (see Fig- ammonia is oxidized to nitrate in nitrification. The next steps
ure 31.16). A growing body of evidence suggests that nodule are carried out by plants, which reduce the nitrate they have
formation depends on some of the same genes and mecha- taken up all the way back to ammonia. All the reactions of
nisms that allow mycorrhizae to develop. nitrate reduction are carried out by the plant’s own en-
zymes. The later steps, from nitrite (NO2–) to ammonia, take
place in the chloroplasts, but this conversion is not part of
Biological nitrogen fixation does not always meet photosynthesis. The plant uses the ammonia thus formed to
agricultural needs
Bacterial nitrogen fixation is not sufficient to support the
needs of agriculture. Traditional farmers used to plant dead 37.8 The Nitrogen Cycle Nitrogen fixation, nitrification,
fish along with corn so that the decaying fish would release nitrate reduction, and denitrification are the components of an
essential chemical cycle that converts atmospheric nitrogen gas
fixed nitrogen that the developing corn could use. Industrial into ammonium ions and nitrate ions—forms of nitrogen that
nitrogen fixation is becoming ever more can be taken up by plants—and returns N2 to the atmosphere.
important to world agriculture because
of the degradation of soils and the need Some denitrifying bacteria
can oxidize ammonia back
to feed a rapidly expanding population. N2
to nitrogen gas, which
Most industrial nitrogen fixation is returns to the atmosphere.
done by a chemical process called the
Haber process, which requires a great
DENITRIFICATION
deal of energy. An alternative is urgently
needed because of the rising cost of en-
Plants reduce
ergy. At present, in the United States, the nitrate ions back to
manufacture of nitrogen-containing fer- ammonia, the form
in which nitrogen NH4+
tilizer takes more energy than does any
is incorporated Re
other aspect of crop production. In bio- into proteins. cy Nitrogen-fixing NITROGEN
c li
logical systems, nitrogen fixation re- NITRATE ng bacteria FIXATION
to s
REDUCTION o il
quires a great deal of ATP.
Research on biological nitrogen fixa- NH3
NH4+
tion is being vigorously pursued, with NO3–
commercial applications very much in Denitrifying Bacteria fix N2 from
bacteria the atmosphere
mind. One line of investigation centers
Oxidation producing ammonia
on recombinant DNA technology as a and ammonium ions.
NO2–
means of engineering new plant–bac- Nitrobacter
Nitrosomonas,
terium associations that produce their NITRIFICATION Nitrosococcus
own nitrogenase. Currently there are at-
tempts to transfer genes from Rhizobium
into bacteria that already live in the Nitrifying bacteria oxidize
ammonia to nitrate ions.
roots of cereal plants.

11. 726 CHAPTER THIRT Y-SEVEN
manufacture amino acids, from which the plant’s proteins The Venus flytraps have specialized leaves with two
and all its other nitrogen-containing compounds are formed. halves that fold together. When an insect trips trigger hairs
Animals cannot reduce nitrogen, and they depend on plants on a leaf, its two halves come together, their spiny margins
to supply them with reduced nitrogenous compounds. interlocking and imprisoning the insect. The leaf then se-
Bacteria called denitrifiers return nitrogen from animal cretes enzymes that digest its prey. The leaf absorbs the
wastes and dead organisms to the atmosphere as N2. This products of digestion, especially amino acids, and uses them
process is called denitrification (see Chapter 27). In combi- as a nutritional supplement.
nation with leaching and the removal of crops, denitrifica- Pitcher plants produce pitcher-shaped leaves that collect
tion keeps the level of available nitrogen in soils low. small amounts of rainwater. Insects are attracted into the
This global nitrogen cycle is complex. It is also essential for pitchers either by bright colors or by scent and are prevented
life on Earth: Nitrogen-containing compounds constitute 5 to from getting out again by stiff, downward-pointing hairs.
30 percent of a plant’s total dry weight. The nitrogen content The insects eventually die and are digested by a combination
of animals is even higher, and all the nitrogen in the animal of enzymes and bacteria in the water. Even rats have been
world arrives there by way of the plant kingdom. found in large pitcher plants.
Sundews have leaves covered with hairs that secrete a
clear, sticky, sugary liquid. An insect touching one of these
Carnivorous and Heterotrophic Plants hairs becomes stuck, and more hairs curve over the insect
Some plants that are found primarily in nitrogen-deficient and stick to it as well. The plant secretes enzymes to digest
soils augment their nitrogen and phosphorus supply by cap- the insect and eventually absorbs the carbon- and nitrogen-
turing and digesting flies and other insects. There are about containing products of digestion.
450 of these carnivorous species, the best-known of which are None of the carnivorous plants must feed on insects to
Venus flytraps (genus Dionaea; Figure 37.9a), sundews (genus survive. They can grow adequately without insects, but in
Drosera; Figure 37.9b), and pitcher plants (genus Sarracenia). their natural habitats they grow faster and are a darker green
Carnivorous plants are normally found in boggy regions when they succeed in capturing insects. They use the addi-
where the soil is acidic. Most decomposing organisms require tional nitrogen from the insects to make more proteins,
a less acidic pH to break down the bodies of dead organisms, chlorophyll, and other nitrogen-containing compounds.
so relatively little nitrogen is recycled into these acidic soils. Thus far in this chapter we have considered the mineral
Accordingly, the carnivorous plants have adaptations that al- nutrition of plants. As you already know, another crucial as-
low them to augment their supply of nitrogen by capturing pect of plant nutrition is photosynthesis—the principal
animals and digesting their proteins. source of energy and carbon for plants them-
selves and for the biosphere as a whole. Not
(b)
all plants, however, are photosynthetic au-
(a) totrophs. A few, in the course of their evolu-
tion, have lost the ability to sustain them-
selves by photosynthesis. How do these
plants get their energy and carbon?
A few plants are heterotrophic parasites
that obtain their nutrients directly from the
living bodies of other plants. Perhaps the
most familiar parasitic plants are the mistle-
toes and dodders (Figure 37.10). Mistletoes
are green and carry on some photosynthesis,
but they parasitize other plants for water and
mineral nutrients and may derive photosyn-
thetic products from them as well. Mistletoes
and dodders extract nutrients from the vas-
cular tissues of their hosts by forming ab-
Dionaea muscipula Drosera rotundifolia
sorptive organs called haustoria, which invade
the host plant’s tissues. Another parasitic
37.9 Carnivorous Plants Some plants have adapted to nitrogen-poor environments plant, the Indian pipe, once was thought to
by becoming carnivorous. (a) The Venus flytrap obtains nitrogen and phosphorus from
the bodies of insects trapped inside the plant when its hinges snap shut. (b) Sundews obtain its nutrients from dead organic matter.
trap insects on sticky hairs. Secreted enzymes will digest the carcass externally. It is now known to get its nutrients, with the

12. PLANT NUTRITION 727
Deficiency symptoms suggest what essential element a plant
lacks. Review Table 37.2
Tendrils of Biologists discovered the requirement for each essential ele-
dodder ment by growing plants on hydroponic solutions lacking that
element. Review Figure 37.1. See Web/CD Tutorial 37.1
Soils and Plants
Soils are complex systems with living and nonliving compo-
Dodder flowers
nents. They contain water, air, and inorganic and organic sub-
stances. They typically consist of two or three horizontal zones
called horizons. Review Figures 37.2, 37.3
Soils form by mechanical and chemical weathering of rock.
Plants obtain some mineral nutrients through ion exchange
between the soil solution and the surface of clay particles.
Review Figure 37.4
Farmers use fertilizers to make up for deficiencies in soil min-
Host stem eral nutrient content, and they apply lime to raise low soil pH.
Plants affect soils in various ways, such as by adding organic
material, removing nutrients (especially in agriculture), and
changing pH.
Nitrogen Fixation
A few species of soil bacteria are responsible for almost all
nitrogen fixation. Some nitrogen-fixing bacteria live free in the
37.10 A Parasitic Plant Tendrils of dodder wrap around other soil; others live symbiotically as bacteroids within the roots of
plants. This parasitic plant (genus Cuscuta) obtains water, sugars, and plants.
other nutrients from its host through tiny, rootlike protuberances that
penetrate the surface of the host. In nitrogen fixation, nitrogen gas (N2) is reduced to ammonia
(NH3) or ammonium ions (NH4+) in a reaction catalyzed by
nitrogenase. Review Figure 37.6
Nitrogenase requires anaerobic conditions, but the bacteroids
in root nodules require oxygen for their respiration. Leghemo-
help of fungi, from nearby actively photosynthesizing plants. globin helps maintain the oxygen supply to the bacteroids at the
Hence it, too, is a parasite. proper level.
The formation of a nodule requires an interaction between
Dwarf mistletoe is a serious parasite in forests of the west- the root system of a legume and Rhizobium bacteria. Review
ern United States, destroying more than 3 billion board feet Figure 37.7
of lumber per year. However, parasitic plants are a much Nitrogen-fixing bacteria reduce atmospheric N2 to ammonia,
more urgent problem in developing countries. Striga (witch- but most plants take up both ammonium ions and nitrate ions.
Nitrifying bacteria oxidize ammonia to nitrate. Plants take up
weed) imperils more than 300 million sub-Saharan Africans nitrate and reduce it back to ammonia, a feat of which animals
by attacking their cereal and legume crops. In the Middle are incapable. Review Figure 37.8. See Web/CD Activity 37.1
East and North Africa, Orobanche (broomrape) ravages many Denitrifying bacteria return N2 to the atmosphere, completing
crops, especially vegetables and sunflowers. the global nitrogen cycle. Review Figure 37.8
Carnivorous and Heterotrophic Plants
Chapter Summary Carnivorous plant species are autotrophs that supplement
their nitrogen supply by feeding on insects.
The Acquisition of Nutrients A few heterotrophic plants are parasitic on other plants. Some
Plants are photosynthetic autotrophs that can produce all the parasitic plants have major effects on crops, especially in devel-
organic compounds they need from carbon dioxide, water, and oping countries.
minerals, including a nitrogen source. They obtain energy from
sunlight, carbon dioxide from the atmosphere, and nitrogen-
containing ions and mineral nutrients from the soil. Self-Quiz
Plants explore their surroundings by growing rather than by
movement. 1. Macronutrients
a. are so called because they are more essential than
Mineral Nutrients Essential to Plants micronutrients.
b. include manganese, boron, and zinc, among others.
Plants require 14 essential mineral elements, all of which
c. function as catalysts.
come from the soil solution. Several of these essential elements
d. are required in concentrations of at least 1 gram per
fulfill multiple roles. Review Table 37.1
kilogram of plant dry matter.
The six mineral nutrients required in substantial amounts are e. are obtained by the process of photosynthesis.
called macronutrients; the eight required in much smaller
amounts are called micronutrients. Review Table 37.1

13. 728 CHAPTER THIRT Y-SEVEN
2. Which of the following is not an essential mineral element 8. Nitrate reduction
for plants? a. is performed by plants.
a. Potassium b. takes place in mitochondria.
b. Magnesium c. is catalyzed by the enzyme nitrogenase.
c. Calcium d. includes the reduction of nitrite ions to nitrate ions.
d. Lead e. is known as the Haber process.
e. Phosphorus 9. Which of the following is a parasite?
3. Fertilizers a. Venus flytrap
a. are often characterized by their N-P-O percentages. b. Pitcher plant
b. are not required if crops are removed frequently enough. c. Sundew
c. restore needed mineral nutrients to the soil. d. Dodder
d. are needed to provide carbon, hydrogen, and oxygen to e. Tobacco
plants. 10. All carnivorous plants
e. are needed to destroy soil pests. a. are parasites.
4. In a typical soil, b. depend on animals as a source of carbon.
a. the topsoil tends to lose mineral nutrients by leaching. c. are incapable of photosynthesis.
b. there are four or more horizons. d. depend on animals as their sole source of phosphorus.
c. the C horizon consists primarily of loam. e. obtain supplemental nitrogen from animals.
d. the dead and decaying organic matter gathers in the B
horizon.
e. more clay means more air space and thus more oxygen For Discussion
for roots.
1. Methods for determining whether a particular element is
5. Which of the following is not an important step in soil essential have been known for more than a century. Since
formation? these methods are so well established, why was the essen-
a. Removal of bacteria tiality of some elements discovered only recently?
b. Mechanical weathering
c. Chemical weathering 2. If a Venus flytrap were deprived of soil sulfates and hence
d. Clay formation made unable to synthesize the amino acids cysteine and
e. Hydrolysis of soil minerals methionine, would it die from lack of protein? Explain.
6. Nitrogen fixation is 3. Soils are dynamic systems. What changes might result when
a. performed only by plants. land is subjected to heavy irrigation for agriculture after
b. the oxidation of nitrogen gas. being relatively dry for many years? What changes in the
c. catalyzed by the enzyme nitrogenase. soil might result when a virgin deciduous forest is cut down
d. a single-step chemical reaction. and replaced by crops that are harvested each year?
e. possible because N2 is a highly reactive substance. 4. We mentioned that important positively charged ions are
7. Nitrification is held in the soil by clay particles, but other, equally impor-
a. performed only by plants. tant, negatively charged ions are leached deeper into the
b. the reduction of ammonium ions to nitrate ions. soil’s B horizon. Why doesn’t leaching cause an electrical
c. the reduction of nitrate ions to nitrogen gas. imbalance in the soil? (Hint: Think of the ionization of
d. catalyzed by the enzyme nitrogenase. water.)
e. performed by certain bacteria in the soil. 5. The biosphere of Earth as we know it depends on the exis-
tence of a few species of nitrogen-fixing prokaryotes. What
do you think might happen if one of these species were to
become extinct? If all of them were to disappear?