Plants and Inorganic Nutrients

Chapter 4
 Learning objectives
Understand the methods used to study
plant mineral nutrition

Understand the distinction between

essential and beneficial nutrients and
between macronutrients and
micronutrients
 Learning objectives
Understand the biological roles for the 14
essential mineral elements

Understand the critical and deficient

concentrations for essential elements

Understand the symptoms associated with

nutrient deficiency and toxicity
 Plant and inorganic nutrients
Unlike heterotrophs, plants are autotrophic,
which makes them capable of surviving in an
inorganic environment.

Plant nutrition is treated in two aspects.

Organic nutrition (e.g., photosynthesis)
Inorganic nutrition

The study of the inorganic nutritional

requirements of plants is called mineral
nutrition.
 Methods in mineral nutrition
The impetus to study plant mineral nutrition was
a need to increase crop productivity to feed
growing populations.

Some of the early researchers were:

N. T. de Saussure
C. S. Sprengel
J-B Boussingault
J. Sachs
J. B. Lawes and J. H. Gilbert
 Methods in mineral nutrition
By the middle of the nineteenth century,
many pieces of the nutritional puzzle were
beginning to fall into place.

In 1860, Julius Sachs, a prominent

German botanist, demonstrated for the
first time that plants could be grown to
maturity in defined nutrient solutions in the
complete absence of soil.
 Methods in mineral nutrition
An important milestone was the
development of superphosphate as a
fertilizer.

This development, and others, expanded

the use of fertilizers to increase crop
productivity.
 Methods in mineral nutrition
A significant development in the study and
understanding of mineral nutrition was the
development of hydroponic solutions to
simulate the nutrient environment.

The first nutrient solution was developed

by Sachs and included many, but not all,
of the necessary plant nutrients.
 Methods in mineral nutrition
Table 4.1
 Methods in mineral nutrition
More balanced nutrient solutions were
developed by Hoaglund and included nearly all
of the necessary nutrients.

The base solution has been modified by many

researchers and is thus referred to as a
modified Hoaglunds solution.

These nutrients solutions provided nutrients in

higher concentrations than typical in the soil.
 Methods in mineral nutrition
Table 4.2
 Methods in mineral nutrition
Hydroponic solutions must be aerated to
prevent anoxia.

Light needs to be excluded from the

solution to prevent the growth of algae.

The plant must be supported in the

solution so it will grow.
 Methods in mineral nutrition
Figure 4.1
 Methods in mineral nutrition
There are some drawbacks to the use of
hydroponic solutions.
Depletions zones can develop unless the
solution is mixed.
In order to maintain nutrient levels, the
hydroponic solution has to be replenished via:
Slop culture.
Drip culture.
Subirrigation.
A nutrient film technique.
 Methods in mineral nutrition
Figure 4.2
 Essential nutrients
Essential nutrients are those that:
Must be provided to plants in order for it to
complete its life cycle.
Are part of some essential plant constituent or
metabolite.
The nutrient must be involved in an essential
biological process.

There are 17 mineral nutrients that meets

these requirements.
 Essential nutrients
Essential nutrients are placed into two
categories based upon the concentration
in plant tissues.
Macronutrients are found in concentrations
in >10 mmole kg-1 DW.
Micronutrients, or trace elements, are found
in concentrations in <10 mmole kg-1 DW.
 Essential nutrients
The essential macronutrients are:
Hydrogen
Carbon
Oxygen
Nitrogen
Potassium
Calcium
Magnesium
Phosphorus
Sulfur
 Essential nutrients
The essential micronutrients are:
Chlorine
Boron
Iron
Manganese
Zinc
Copper
Nickel
Molybdenum
 Essential nutrients
Determining essentiality of a mineral
nutrients is a difficult process.

Studies must use high purity reagents and

extremely clean materials to prevent small
amounts of nutrients from contaminating
the experiment.
 Beneficial nutrients
Beneficial nutrients may not be
universally required by plants, but may
promote plant growth and development if
present.

Beneficial elements include:

Sodium
Silicon
Cobalt
Selenium
 Beneficial nutrients
Sodium for example is required by C4 plants but
less so by C3 plants.

Silicon increases the rigidity and elasticity of cell

walls and the resistance to lodging.

Cobalt is required by legumes.

Some plants accumulate high concentrations of

selenium.
 Nutrient functions and deficiency
symptoms
A plant species requirement for a
particular nutrient is defined as the critical
concentration.

The critical concentration is the

concentration of a nutrient in the plant
tissue just below that which provides for
maximum growth.
 Nutrient functions and deficiency
symptoms
At tissue concentrations above the critical
concentration, the nutrient is present in
adequate amounts.

At concentrations in excess of the critical

concentration, nutrients can be toxic.

At tissue concentrations below the critical

concentration, the nutrient is deficient.
 Nutrient functions and deficiency
symptoms
Figure 4.3
 Nutrient functions and deficiency
symptoms
Plants that are deficient in a nutrient may
display visual symptoms such as:
Chlorosis, the loss of chlorophyll pigment.
Necrosis, the death of cells and tissues.

The location of deficiency symptoms in a

plant depend upon the mobility of the
nutrient in the phloem.
 Nitrogen
Plants generally acquire nitrogen as
ammonium or nitrate.

Nitrogen is required for the synthesis of

macromolecules such as amino acids,
proteins, and nucleic acids.

Nitrogen stimulates shoot growth more so

than root growth.
 Nitrogen
Symptoms of nitrogen deficiency include:
Chlorosis of the older leaves first, and then
the younger leaves.
An accumulation of anthocyanin pigments.
 Phosphorus
Phosphorus is acquired as phosphate and
the plant may have to convert organic
forms to inorganic forms to get phosphate.

Phosphates are found in the sugar-

phosphates involved in metabolism.

Phosphates are also a structural

component of nucleic acids.
 Phosphorus
Phosphate is also found as a component
of ATP.

Phosphorus deficiency symptoms include:

Over-greening of the leaves.
Accumulation of anthocyanins.
Malformation of leaves and necrosis.
Symptoms in the older leaves initially.
 Potassium
Potassium is the primary osmolyte in plants
used for water relations.

Potassium activates or is a cofactor for a wide

range of enzymes, including those in
photosynthesis and respiration.

Potassium deficiency symptoms include:

Chlorosis and mottling in older leaves
Necrosis, especially in grasses.
 Sulfur
Sulfate is the only form of sulfur plants
take up.

Sulfur deficiencies are rare.

Sulfur deficiency symptoms include:

Generalized chlorosis.
Symptoms primarily in the young leaves.
 Sulfur
Sulfur has biological roles in:
Disulfide bonds in proteins.
Vitamins and coenzymes.
Iron-sulfur proteins in electron transport.
Secondary compounds, such as the
thiocyanates and isothiocyanates.
 Calcium
Calcium is taken up as Ca2+.

Calcium is important:
As a second messenger.
For mitosis and the spindle fibers.
The structure of the cell wall.
The integrity of the cell membrane.
 Calcium
Calcium deficiency symptoms appear in
the young leaves.
The meristematic region is affected because
of the importance of Ca2+ to mitosis.
Young leaves are deformed and necrotic.

In hydroponic solution, Ca2+ deficiency

affects the roots.
 Magnesium
Magnesium is taken up as Mg2+.

Biological roles for magnesium include:

Serving as the cofactor for chlorophyll.
Stabilizing ribosome structure.
Activation of several enzymes, including those
in photosynthesis.
Serving as a cofactor in ATP-mediated
reactions.
 Magnesium
Magnesium deficiency symptoms include:
Chlorosis, particularly in the leaf veins.
Symptoms in the older leaves.
 Iron
Iron can be taken up as Fe2+ or Fe3+,
depending upon the plant species.

Iron is required for:

Chlorophyll synthesis.
Iron-sulfur proteins in electron transport.
Nitrogen fixation in legumes.
Redox enzymes.
 Iron
One reason that plants experience iron
deficiency is because iron can have a
limited solubility in soil.

The deficiency can be minimized as iron is

provided in a chelated form, such as with
ethylenediaminetetraacetic acid
(EDTA).
 Iron
Because of the problems with iron
solubility, plants have specific
mechanisms to acquire this nutrient.
Protons are released to acidify the soil.
Ligands like caffeic acid are released.
In non-grasses, a ferric reductase is used to
reduce solubilized Fe3+ to Fe2+ for uptake.
Grasses secrete phytosiderophores to
solubilize and take up Fe3+.
 Iron
Figure 4.4
 Iron

Figure 4.5
 Iron
Figure 4.6
 Boron
Boron can be present primarily as the
neutral chemical species H3BO3.

Roles for boron are unclear but include:

A contribution to cell wall structure.
Potential roles in carbohydrate metabolism.
Potential roles in cell division and elongation.
 Boron
Deficiency symptoms include:
Decreased root elongation.
Decreased cell division.
Distorted roots, including a stubby, bushy
appearance or cork-screw roots.
Shortened internodes.
 Copper
Copper is present in soils as Cu2+ and Cu+,
depending upon the redox conditions.

Copper is primarily a cofactor for various

oxidative enzymes, such as superoxide
dismutase.

Copper is also a component of electron

carriers in the chloroplast and
mitochondrion.
 Copper
Symptoms of copper deficiency are more
prominent in soils with high organic matter
because Cu2+ binds strongly to organic
matter.

Symptoms of copper deficiency include:

Stunted growth.
Distortion of young leaves.
 Zinc
Roots take up Zn2+.

Zinc is an activator of enzymes, including:

Alcohol dehydrogenase.
Carbonic anhydrase.

Zinc is also involved in auxin metabolism,

perhaps with respect to tryptophan
synthesis.
 Zinc
Symptoms of zinc deficiency include:
Chlorosis.
Shortened internodes.
Smaller leaves.
 Manganese
Manganese occurs in several chemical species
in soil, but is taken up as Mn2+.

Manganese is a cofactor for several enzymes,

including decarboxylase and dehydrogenase
enzymes involved in the respiratory carbon
cycle.

Manganese can substitute for magnesium in

ATP-mediated reactions.
 Manganese
Manganese is critical for the
magnoprotein involved in oxygen
evolution during photosynthesis.

Manganese deficiency symptoms include:

Gray speck in cereal grains.
Chlorosis.
Discoloration and deformities in legume
seeds.
 Molybdenum
Molybdenum is taken up primarily as
molybdate (MoO42-).

Molybdenum is required for dinitrogenase

and nitrate reductase.

Deficiency symptoms include:

Chlorosis and necrosis.
Whiptail, the deformation of young leaves.
 Chlorine
Chlorine is rarely deficient in plants and
then only in laboratory experiments.

Chloride is a counterion for water relations

and charge balance.

Chloride is also required for cell division

and photosynthesis.
 Nickel
Although considered an essential nutrient,
the essentiality of nickel has been difficult to
establish.

Nickel is required for the function of

enzymes in nitrogen metabolism, particularly
in legumes.
Urease, which is involved in the metabolism of
the ureides formed by legumes after nitrogen
fixation.
Hydrogenase
 Micronutrient toxicity
There is a critical toxicity level for each
micronutrient, above which the essential
nutrients can become toxic.

Toxicity symptoms are often similar to

those of nutrient deficiency.
Chlorosis
Inhibition of root growth