Chapter 37

PLANT NUTRITION

NUTRITIONAL REQUIREMENTS OF PLANTS   

The chemical composition of plants provides clues to their nutritional requirements   
Plants require nine macronutrients and at least eight micronutrients   
The symptoms of a mineral deficiency depend on the function and mobility of the
element

THE ROLE OF SOIL IN PLANT NUTRITION   

Soil characteristics are key environmental factors in terrestrial ecosystems   
Soil conservation is one step toward sustainable agriculture

THE SPECIAL CASE OF NITROGEN AS A PLANT NUTRIENT   

The metabolism of soil bacteria makes nitrogen available to plants   
Improving the protein yield of crops is a major goal of agricultural research

NUTRITIONAL ADAPTATIONS: SYMBIOSIS OF PLANTS AND SOIL MICROBES   

Symbiotic nitrogen fixation results from intricate interactions between roots and
bacteria 
Mycorrhizae are symbiotic associations of roots and fungi that enhance plant nutrition 
Mycorrhizae and root nodules may have an evolutionary relationship

NUTRITIONAL ADAPTATIONS: PARASITISM AND PREDATION BY PLANTS   

Parasitic plants extract nutrients from other plants   
Carnivorous plants supplement their mineral nutrition by digesting animals
 1. NUTRITIONAL REQUIREMENTS OF PLANTS

* The chemical composition of plants provides clues to their nutritional

requirements  
* Plants require 9 macronutrients and at least 8 micronutrients  
* The symptoms of a mineral deficiency depend on the function and
mobility of the element
 1.1. The chemical composition of plants
provides clues to their nutritional requirements

Watch a large plant grow from a tiny seed, and you cannot help wondering where all the
mass comes from
As it turns out, none of the early ideas about plant nutrition is entirely incorrect. Plants do
extract minerals from the soil.
Mineral nutrients are essential chemical elements absorbed from the soil in the form of
inorganic ions
By weight, the bulk of the organic material of a plant is derived not from water or soil
minerals, but from the CO2 assimilated from the atmosphere (fig 37.1).
Generally, > 90% of the water absorbed by plants is lost by transpiration, and most of the
water retained by the plant functions as a solvent, provides most of the mass for cell
elongation, and helps maintain the form of soft tissue by keeping cells turgid.
More than 50 chemical elements have been identified among the inorganic substances
present in plants, but it is unlikely that all these elements are essential.
Roots are able to absorb minerals somewhat selectively, enabling the plant to accumulate
essential elements that may be present in the soil in very minute quantities
 Fig 37-1. 
The uptake of nutrients
by a plant: an overview.
Roots absorb water and
minerals from the soil, with
mycorrhizae and root hairs
greatly increasing surface area
for absorption. Carbon dioxide,
the source of carbon for photo-
synthesis, diffuses into leaves
from the sur-rounding air
through stomata. (Plants also
need O2 for cellular
respiration, although the plant
is a net producer of O2.) From
these inorganic nutrients the
plant can produce all of its own
organic material.
 1.2. Plants require 9 macronutrients and at least 8 micronutrients

A particular chemical element is considered an essential nutrient if it is required for a plant

to grow from a seed and complete the life cycle, producing another generation of seeds.
Researchers can use a method known as hydroponic culture to determine which of the
mineral elements are actually essential nutrients (fig 37.2).
Such studies have helped identify 17 elements that are essential nutrients in all plants and
a few other elements that are essential to certain groups of plants
Elements required by plants in relatively large amounts are called macronutrients. There
are nine macronutrients in all, including the six major ingredients of organic compounds:
carbon (C), oxygen (O2), hydrogen (H2), nitrogen (N), sulfur (S), and phosphorus (P). The
other three macronutrients are potassium (K), calcium (Ca), and magnesium (Mg). (table
37.1)
Elements that plants need in very small amounts are called micronutrients. The eight
micronutrients are iron (Fe), chlorine (Cl), copper (Cu), manganese (Mn), zinc (Zn),
molybdenum (Mo), boron (Bo), and nickel (Ni). These elements function in plants mainly as
cofactors of enzymatic reactions
 Form vailable
Element Major Functions
to Plants

Macronutrients
C CO2 Major component of plant’s organic compounds

O2 CO2 Major component of plant’s organic compounds

H2 H2O Major component of plant’s organic compounds

N NO3-, NH4+ Component of nucleic acids, proteins, hormones, and coenzymes

S SO42- Component of proteins, coenzymes

P H2PO42-, HPO42- Component of nucleic acids, phospholipids, ATP, several coenzymes

K K+ Cofactor that functions in protein synthesis; major solute functioning in water balance; operation of stomata
Ca Ca2+ Important in formation and stability of cell walls and in maintenance of membrane structure and permeability;
activates some enzymes; regulates many responses of cells to stimuli
Mg Mg2+ Component of chlorophyll; activates many enzymes

Micronutrients
Cl Cl- Required for water-splitting step of photosynthesis; functions in water balance
Fe Fe3+, Fe2+ Component of cytochromes; activates some enzymes
Bo H2BO3- Cofactor in chlorophyll synthesis; may be involved in carbohydrate transport and nucleic acid synthesis

Mn Mn2+ Active in formation of amino acids; activates some enzymes; required for water-splitting step of photosynthesis
Zn Zn2+ Active in formation of chlorophyll; activates some enzymes
Cu Cu+, Cu2+ Component of many redox and lignin-biosynthetic enzymes
Mo MoO42- Essential for nitrogen fixation; cofactor that functions in nitrate reduction

Ni Ni2+ Cofactor for an enzyme functioning in nitrogen metabolism

 Fig 37-2. 
Using hydroponic culture to
identify essential nutrients.
A researcher bathes the roots
of plants in solutions of
various minerals dissolved in
known concentrations.
Aerating the water provides
the roots with oxygen for
cellular respiration. A
particular mineral, such as
potassium, can be omitted
from the culture medium to
test whether it is essential to
the plants. If the element
deleted from the mineral
solution is an essential
nutrient, then the incomplete
medium will cause plants to
become abnormal in appear-
ance compared with controls
grown on a complete mineral
medium. The most common
symptoms of a mineral
deficiency are stunted growth
and discolored leaves
 1.3. The symptoms of a mineral deficiency
depend on the function and mobility of the element

The symptoms of a mineral deficiency depend partly on the function of that nutrient in
the plant.
For example, a deficiency of Mg, an ingredient of chlorophyll, causes yellowing of the
leaves, or chlorosis (fig 37.3).
Mineral deficiency symptoms depend not only on the role of the nutrient in the plant
but also on its mobility within the plant
The symptoms of a mineral deficiency are often distinctive enough for a plant
physiologist or farmer to diagnose its cause
One way to ensure optimal mineral nutrition is to grow plants hydroponically on
nutrient solutions that can be precisely regulated (fig 37.4)
Mineral deficiencies are not limited to terrestrial ecosystems, nor are they unique to
plants among photosynthetic organisms
 Fig 37-3. Magnesium deficiency in a tomato plant
Yellowing of the leaves (chlorosis) is the result of an inability to synthesize
chlorophyll, which contains Mg
 Fig 37-4. Hydroponic farming. In this apparatus,
a nutrient solution flows over the roots of lettuce growing on a slat
Perhaps astronauts living in a space station will one day grow their vegetables hydroponically, but
because of the expense, it is unlikely that this type of farming will relieve hunger here on Earth
 2. THE ROLE OF SOIL IN PLANT NUTRITION

* Soil characteristics are key environmental factors in

terrestrial ecosystems  
* Soil conservation is one step toward sustainable agriculture
 2.1. Soil characteristics
are key environmental factors in terrestrial ecosystems

The texture and chemical composition of soil are major factors determining what kinds
of plants can grow well in a particular location, be it a natural ecosystem or an agricultural
region
2.1.1. Texture and composition of soils
Soil has its origin in the weathering of solid rock
The eventual result of all this activity is topsoil, a mixture of particles derived from rock,
living organisms, and humus, a residue of partially decayed organic material.
The topsoil and other distinct soil layers, or horizons, are often visible in vertical profile
where there is a roadcut or deep hole (fig 37.5)
The texture of topsoil depends on the size of its particles, which are classified in a range
from coarse sand to microscopic clay particles.
The most fertile soils are usually loams, made up of roughly equal amounts of sand, silt
(particles of intermediate size), and clay
Topsoil is home to an astonishing number and variety of organisms. A teaspoon of soil
has about 5 billion bacteria that cohabit with various fungi, algae and other protists, insects,
earthworms, nematodes, and the roots of plants. The activities of all these organisms affect
the physical and chemical properties of the soil
Humus, an important component of topsoil, is the decomposing organic material formed by
the action of bacteria and fungi on dead organisms, feces, fallen leaves, and other organic
refuse.
Humus prevents clay from packing together and builds a crumbly soil that retains water
but is still porous enough for the adequate aeration of roots
Humus is also a reservoir of mineral nutrients that are returned gradually to the soil as
microorganisms decompose the organic matter

Fig 37-5. 
Soil horizons
This researcher is
photographing a vertical
profile of three soil layers,
or horizons, in a
Tennessee cotton field.
2.1.2. The availability of soil water and minerals

After a heavy rainfall, water drains away from the larger spaces of the soil, but smaller
spaces retain water because of its attraction for the soil particles
Some of this water adheres so tightly to the hydrophilic soil particles that it cannot be
extracted by plants.
The film of water bound less tightly to the particles is the water generally available to
plants (fig 37.6a).
It is not pure water, but a soil solution containing dissolved minerals. Roots absorb this
soil solution
The presence of clay in a soil helps prevent the leaching (draining away) of mineral
nutrients during heavy rain or irrigation because the finely divided particles provide so
much surface area for binding minerals
Minerals that are negatively charged, such as nitrate (NO3-), phosphate (H2PO4), and
sulfate (SO42-), are usually not bound tightly to soil particles and thus tend to leach away
more quickly.
Positively charged minerals are made available to the plant when hydrogen ions in the
soil displace the mineral ions from the clay particles.
This process, called cation exchange, is stimulated by the roots themselves, which
secrete H+ and compounds that form acids in the soil solution (fig 37.6b).
Fig 37-6. The availability of soil water and minerals
 2.2. Soil conservation is one step toward sustainable agriculture

Both of these land uses left the soil exposed to the danger of erosion by the winds
that constantly sweep over the area (fig 37.7)
To understand soil conservation, we must begin with the premise that agriculture is
unnatural
In forests, grasslands, and other natural ecosystems, mineral nutrients are usually
recycled by the decomposition of dead organic material in the soil
Each year, the fertility of the soil diminishes unless fertilizers are applied to replace
the lost minerals
Prudent fertilization, thoughtful irrigation, and the prevention of erosion are 3 of
the most important goals of soil conservation
 Fig 37-7. Poor soil conservation has contributed to ecological disasters such as the Dust Bowl.
The widespread planting of wheat and the raising of cattle by homesteaders in the southwestern plains
states in the early part of the 20th century, in combination with recurrent droughts, left the land
susceptible to wind erosion. The organic matter, clay, and silt in the soil were carried great distances by
the winds, in some cases darkening the sky as far as the Atlantic coast. Sand and heavier materials
drifted against houses, fences, and barns. In the 1930s, hundreds of thousands of people abandoned
their homesteads. Many became destitute migrant laborers in California. Their plight was immortalized
as a symbol of the Great Depression in John Steinbeck’s The Grapes of Wrath
2.2.1. Fertilizers

Prehistoric farmers may have started fertilizing their fields after noticing that grass grew
faster and greener where animals had defecated
In developed nations today, most farmers use commercially produced fertilizers
containing minerals that are either mined or prepared by industrial processes
A fertilizer marked "10-12-8," for instance, is 10% nitrogen (as ammonium or nitrate),
12% phosphorus (as phosphoric acid), and 8% potassium (as the mineral potash).
Manure, fishmeal, and compost are referred to as "organic" fertilizers because they are
of biological origin and contain organic material that is in the process of decomposing.
However, before the elements in compost can be of any use to plants, the organic
material must be decomposed to the inorganic nutrients that roots can absorb
To fertilize judiciously, a farmer must pay close attention to the pH of the soil. Soil pH not
only affects cation exchange but also influences the chemical form of all minerals
A major problem with acid soils, particularly in tropical areas, is that aluminum (Al)
dissolves in the soil at low pH and becomes toxic to roots
2.2.2. Irrigation

Even more than mineral deficiencies, the unavailability of water most often limits the
growth of plants
As the world population continues to grow, more and more acres of arid land will have
to be cultivated.
Another problem is that irrigation in an arid region can gradually make the soil so salty
that it becomes completely infertile. Salts dissolved in the irrigation water accumulate in
the soil as the water evaporates
New methods of irrigation may reduce the risks of running out of water or losing
farmland to salinization (salt accumulation)
For instance, drip irrigation is now used as an alternative to flooding fields for many of
the crops and orchards in Israel and the western United States.
In another approach to solving some of the problems of dryland farming, plant
breeders are working to develop varieties of plants that require less water
 3. THE SPECIAL CASE OF NITROGEN AS A PLANT NUTRIENT

* The metabolism of soil bacteria makes nitrogen available to plants 

* Improving the protein yield of crops is a major goal of agricultural research

Of all mineral elements, nitrogen is the one that most often limits the growth of
plants and the yields of crops. Plants require nitrogen as an ingredient of
proteins, nucleic acids, and other important organic molecules .
3.1. The metabolism of soil bacteria makes nitrogen available to plants

The atmosphere is nearly 80% nitrogen that is gaseous N2, and plants cannot use
nitrogen in that form. For plants to absorb nitrogen, N2 must first be converted 
ammonium (NH4+) or nitrate (NO3-).
Over the short term, the main source of nitrogenous minerals is the decomposition of
humus by microbes, including ammonifying bacteria (fig 37.9): called nitrogen fiaxation
N2 is lost from this local cycle when soil microbes called denitrifying bacteria convert
NO3-  N2, which diffuses from the soil to the atmosphere
All life on earth depends on nitrogen fixation. Soil is populated by several species of free-
living bacteria that are among the nitrogen-fixing prokaryotes.
The conversion of atmospheric N2  ammonia (NH3) is a complicated, multi-step
process, but we can simplify nitrogen fixation by just indicating the reactants and products
by nitrogenase
Nitrogen-fixing bacteria are most abundant in soils rich in organic material, which
provides fuel for cellular respiration

N2 (organic matter)  NH4+  nitrifying bacteria  NO3-  plant  NO3- / organic

compounds
 Fig 37-9. The role of soil bacteria in the nitrogen nutrition of plants
Ammonium is made available to plants by two types of soil bacteria: those that fix atmospheric N2
(nitrogen-fixing bacteria) and those that decompose organic material (ammonifying bacteria).
Although plants absorb some ammonium from the soil, they absorb mainly nitrate, which is
produced from ammonium by nitrifying bacteria. Plants reduce nitrate back to ammonium before
incorporating the nitrogen into organic compounds. Xylem transports nitrogen from roots to shoots
in the form of nitrate, amino acids, and various other organic compounds, depending on the
species
 3.2. Improving the protein yield of crops
is a major goal of agricultural research

Improving the quality and quantity of proteins in crops is a major goal of agricultural
research
Plant breeding has resulted in new varieties of corn, wheat, and rice that are enriched
in protein
However, many of these "super" varieties have an extraordinary demand for nitrogen,
which is usually supplied in the form of commercial fertilizer
The industrial production of ammonia and nitrate from atmospheric nitrogen is, like
biological nitrogen fixation, very expensive in energy costs. A chemical factory making
fertilizer consumes large quantities of fossil fuels
The use of new catalysts based on the mechanism by which nitrogenase fixes nitrogen
may make commercial fertilizer production less costly in the future
Another strategy that could potentially increase protein yields of crops is to improve the
productivity of symbiotic nitrogen fixation, a process we examine in the next section.
 4. NUTRITIONAL ADAPTATIONS:
SYMBIOSIS OF PLANTS AND SOIL MICROBES

* Symbiotic nitrogen fixation results from intricate interactions

between roots and bacteria 
* Mycorrhizae are symbiotic associations of roots and fungi that
enhance plant nutrition 
* Mycorrhizae and root nodules may have an evolutionary
relationship

The roots of plants belong to subterranean communities that include a diversity of

other organisms. Among those organisms are certain species of bacteria and fungi
that have coevolved with specific plants, forming symbiotic relationships with roots
that enhance the nutrition of both partners. The two most important examples are
symbiotic nitrogen fixation (roots and bacteria) and the formation of mycorrhizae
(roots and fungi).
 4.1. Symbiotic nitrogen fixation results
from intricate interactions between roots and bacteria

Many plant families include species that form symbiotic relationships with nitrogen-fixing
bacteria that give roots a built-in source of fixed nitrogen for assimilation into organic
compounds.
Most of the research on symbiotic nitrogen fixation has focused on agriculturally
important members of the legume family, including peas, beans, soybeans, peanuts,
alfalfa, and clover.
A legume’s roots have swellings called nodules composed of plant cells that contain
nitrogen-fixing bacteria of the genus Rhizobium ("root living").
Inside the nodule, Rhizobium bacteria assume a form called bacteroids, which are
contained within vesicles formed by the root cell (fig 37.10). Each legume is associated
with a particular species of Rhizobium.
fig 37.11 describes the steps in the development of root nodules after bacteria enter
through what is called an infection thread.
The symbiotic relationship between a legume and nitrogen-fixing bacteria is
mutualistic, with both partners benefiting
Some root nodules have a reddish color owing to a molecule called leghemoglobin
 Fig 37-10. Root nodules on legumes
(a) Pea plant root. The bumps on this pea plant root are nodules containing symbiotic bacteria.
The bacteria fix nitrogen and obtain photosynthetic products supplied by the plant. (b) Bacteroids
in a soybean root nodule. In this TEM, a cell from a root nodule of soybean is filled with
bacteroids in vesicles. The cell on the left is uninfected
 Fig 37-11. 
Development of a
soybean root nodule
4.1.1. Symbiotic nitrogen fixation and agriculture

One year a nonlegume such as corn is planted, and the following year alfalfa or some
other legume is planted to restore the concentration of fixed nitrogen in the soil.
Instead of being harvested, the legume crop is often plowed under so that it will
decompose as "green manure" (fig 37.12).
To ensure that the legume encounters its specific Rhizobium, the seeds are soaked in
a culture of the bacteria or dusted with bacterial spores before sowing.
Many plant families besides legumes include species that benefit from symbiotic
nitrogen fixation.
For example, Rice, a crop of great commercial importance, benefits indirectly from
symbiotic nitrogen fixation. Rice farmers culture a water fern called Azolla in their
paddies. The fern has symbiotic cyanobacteria that fix nitrogen and increase the fertility
of the rice paddy

4.1.2. The molecular biology of root nodule formation in legumes

Rhizobium  signal  root (plant) gene-regulating protein  nodulation genes 

Nod factor  infection thread (fig 37.11)
Rhizobium bacteria  Gene fix N2 (gene Nif)  transfer to plant  plant modified to fix
N2
 Fig 37-12. Crop rotation and "green manure”
" The "green manure" being mulched into the soil of this Washington State farm is sweet
clover, a legume with root nodules containing nitrogen-fixing bacteria. Every third year, clover
is planted and the crop is plowed under. This improves the physical structure and nitrogen
content of the soil for growing wheat and other crops during the other two years of the crop
rotation cycle. Crop rotation, especially when the legume is mulched instead of harvested,
reduces the need for manufactured fertilizers
Fig 37-13. Molecular biology of root nodule formation
 4.2. Mycorrhizae are symbiotic associations of
roots and fungi that enhance plant nutrition

Mycorrhizae ("fungus roots") are modified roots consisting of symbiotic associations of

fungi and roots (see fig 31.18 and 36.8). The symbiosis is mutualistic
the fungus increases the surface area for water uptake and selectively absorbs
phosphate and other minerals from the soil and supplies them to the plant.
The fungi of mycorrhizae also secrete growth factors that stimulate roots to grow and
branch.
And the fungi produce antibiotics that may help protect the plant from pathogenic
bacteria and pathogenic fungi in the soil.
Mycorrhizae are not oddities; they are formed by almost all plant species
4.2.1. The two main types of Mycorrhizae

The modified roots formed from the symbiosis of fungi and plants take two major
forms: ectomycorrhizae and endomy corrhizae.
In ectomycorrhizae, the mycelium (mass of branching hyphae; see Chapter 31)
forms a dense sheath, or mantle, over the surface of the root (fig 37.14a).
Hyphae extend from the mantle into the soil, greatly increasing the surface area for
water and mineral absorption
Ectomycorrhizae are especially common in woody plants, including trees of the pine,
spruce, oak, walnut, birch, willow, and eucalyptus families
In contrast to ectomycorrhizae, endomycorrhizae do not have a dense mantle
ensheathing the root (fig 37.14b).
It takes a microscope to see the fine fungal hyphae that extend from the root into the
soil
Endomycorrhizae, much more common than ectomycorrhizae, are found in over 90%
of plant species, including important crop plants such as corn, wheat, and legumes.
4.2.2. Agricultural importance of Mycorrhizae

Roots can be transformed into mycorrhizae only if they are exposed to the
appropriate species of fungus
For example, inoculating pine seeds with spores of mycorrhizal fungi promotes
the formation of mycorrhizae by the seedlings. Pine seedlings so infected grow
more vigorously than trees without the fungal association.
 Fig 37-14. 
Mycorrhizae
 4.3. Mycorrhizae and root nodules
may have an evolutionary relationship

Why does the mycorrhizae and root nodules may have evolutionary relationship ?
 5. NUTRITIONAL ADAPTATIONS:
PARASITISM AND PREDATION BY PLANTS

* Parasitic plants extract nutrients from other plants 

* Carnivorous plants supplement their mineral nutrition by digesting animals

Symbiotic nitrogen fixation and mycorrhizae underscore the relationship between

plants and their environment, which includes the other organisms that interact with
plants. We conclude this chapter by exploring predation and parasitism as two other
types of plant adaptations that enhance nutrition through interactions with other
organisms
 5.1. Parasitic plants extract nutrients from other plants

The mistletoe we find tacked above doorways during the holiday season lives in
nature as a parasite on oaks and other trees.
Mistletoe is photosynthetic, but it supplements its nutrition by using projections
called haustoria to siphon xylem sap from the vascular tissue of the host tree
Indian pipe obtains its nutrients from trees indirectly via its association with fungal
hyphae of the host tree’s mycorrhizae (fig 37.15b)
Plants called epiphytes (from the Greek epi , upon, and phyton , plant) are
sometimes mistaken for parasites.
An epiphyte is an autotrophic plant that nourishes itself but grows on the surface of
another plant, usually on the branches or trunks of trees
 Fig 37-15. 
Parasitic plants
 5.2. Carnivorous plants
supplement their mineral nutrition by digesting animals
Living in acid bogs and other habitats where soil conditions are poor (especially in nitrogen) are plants
that fortify themselves by occasionally feeding on animals

Fig 37-16. Carnivorous plants

(a) The Venus flytrap is a mod ified leaf with two lobes that close together rapidly enough to capture an
insect. Prey that enters the trap touches sensory hairs, initiating an electrical impulse that triggers closure
of the trap. Glands in the trap then secrete digestive enzymes, and nutrients are later absorbed by the
modified leaf. In spite of its name, the flytrap catches more ants and grasshoppers than it does flies. (b)
Pitcher plants use a pitfall to capture insects. Insects slip into a long water-filled funnel. After the insect
drowns, it is digested by enzymes secreted into the water.