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The micronutrients that are managed by growers and we will discuss include:







There are three additional micronutrients that have been classified as essential, but are generally not managed
by growers. These additional three nutrients, listed below, are rather managed under experimental conditions:




Forms and Functions of Micronutrients


Form: Iron is taken up by plants as either Fe2+ (ferrous cation) or Fe3+ (ferric cation).

Function: Iron is involved in photosynthesis, respiration, chlorophyll formation, and many enzymatic


Form: Boron is taken up by plants primarily as H3BO3 (boric acid) and H2BO3- (borate).

Function: Boron plays an important role in the movement and metabolism of sugars in the plant and
synthesis of plant hormones and nucleic acids. It also functions in lignin formation of cell walls.


Form: The primary form of manganese uptake is Mn2+ (manganous ion).

Function: Manganese is a component of enzymes and is also involved in photosynthesis and root
growth. Additionally, it is involved in nitrogen fixation.


Form: The Zn2+ cation is the predominate form taken up by plants.

Function: Zinc is a component of many organic complexes and DNA protein. It is also an important
enzyme for protein synthesis. Also, zinc is involved in growth hormone production and seed


Form: Molybdenum is primarily taken up as MoO42- (molybdate ion).

Function: It is involved in nitrogen fixation (conversion of N2 to NH4+) and nitrification (conversion of

NH4+ to NO3-).


Form: Copper is taken up as Cu2+ (cupric ion).

Function: Copper is also a component of enzymes, some of which are important to lignin formation in
cell walls. It is also involved in photosynthesis, respiration, and processes within the plant involving

The iron cycle includes both mineral and organic forms.
Mineral Iron

Iron may exist:

in the soil solution

includes soluble iron and organic matter complexes in the form of chelates

as primary minerals and/or precipitated minerals

cation exchange site on soil particles

Fe containing minerals may dissolve to replenish the soil solution as iron is removed by plants. Little iron is
retained by the cation exchange sites of soil particles as compared to base and acid cations.
Organic Iron

Organic cycling is an important process that ensures iron availability through the processes of mineralization and

Iron Chelation

Iron can also form strong complexes with organic matter known as chelates (a Greek word meaning claw).
Chelation occurs between soluble organic compounds and certain metals in the soil through processes involving
microorganisms. Chelates are very important in micronutrient management because chelation increases the
solubility and plant uptake of many metal micronutrients. We will encounter chelation again when discussing zinc,
copper and manganese.

The manganese cycle is very similar to the iron cycle. The manganese cycle, too, has four fractions:

manganese cations in soil solution

includes soluble manganese and organic matter complexes known as chelates

exchangeable manganese on soil particles (cation exchange sites)

primary and secondary manganese-containing minerals

soil organic matter

Like iron, little manganese is retained by the cation exchange sites of soil particles. Manganese may undergo
precipitation/dissolution, sorption/desorption on the CEC, mineralization/immobilization, and chelation.


Zinc cycling includes:

zinc cations in soil solution zinc

includes soluble zinc and organic matter complexes known as chelates

zinc retained by soil particles on the cation exchange sites

primary and secondary zinc-containing minerals

soil organic matter

Zinc bearing minerals can dissolve and supply zinc to the soil solution. Once in the soil solution, zinc can be
immobilized, taken up by plants, retained by soil particles, or chelated with soluble organic matter. Organic matter
containing zinc must undergo mineralization before it becomes available for plant uptake.

Like Zinc, the copper cycle includes:

Solution copper

Includes soluble copper and organic matter complexes known as chelates

Exchangeable copper on the cation exchange sites of soil particles

Primary and secondary copper minerals

Copper may be occluded, or buried, within the structures of various minerals, such as iron and
aluminum oxides

Organic copper

Copper is more tightly bound to organic matter than the other micronutrients

Copper deficiencies can occur in organic soils

Copper-containing minerals can dissolve and supply Zn to the soil solution. Like zinc, copper can be immobilized
by microorganisms, taken up by plants, or exchanged on soil particle surfaces. Copper may also form chelates
with soluble organic matter. Organic copper must be mineralized before it is available for plant uptake.

Unlike the previous metal micronutrients, molybdenum exists as an anion in the soil solution. Nonetheless, the
molybdenum cycle is similar to the others. The molybdenum cycle includes:

Soil solution

Exchangeable molybdenum on the anion exchange sites

Primary and secondary molybdenum minerals

Organic matter

Instead of being held onto the cation exchange capacity, molybdenum is held to soil particles with an anion
exchange capacity (including amorphous materials, iron oxides, acidic kaolin clays). Organic molybdenum
undergoes mineralization and immobilization.

Boron exists in the soil as:

soil solution boron

exchangeable boron on the anion exchange capacity sites

primary and secondary boron minerals

Boron and organic matter complexes

Boron is the only nonmetal micronutrient described in this section. H3BO3 is most common form of boron in soils
that have a pH between 5 and 9. The exchangeable boron buffers changes in the boron levels of the soil solution.
Organic matter supplies plant available boron. Boron should be carefully managed when applied to the soil since
the range between boron sufficiency and toxicity levels is very narrow.

Factors affecting micronutrient availability


Soil pH: The availability of iron may be limited in soils with high pH, especially in arid, calcareous soils.

Excessive liming can induce iron deficiencies.

Soil Moisture and Aeration: Poorly aerated soils with excessive moisture in calcareous soil can
promote iron deficiencies.

However, flooding of non-calcareous soils can improve iron availability.

Organic Matter: Organic matter improves iron availability due to chelation, which increases iron
solubility. Additions of manure can increase chelation.

Interactions with other nutrients: Excessive amounts of other micronutrients, particularly copper,
manganese, zinc and molybdenum, can decrease iron availability


Soil pH: Soils with high pH have limited manganese availability since manganese precipitates at high

Overliming soils can cause Mn deficiencies.

Soil Moisture and Aeration: High soil moisture and poor aeration increases the availability of
manganese due to an increase in solubility.

Organic Matter: Manganese availability increases with the addition of natural organic matter (i.e.
compost) due to favorable chelation which increases the level of exchangeable and solution

Climate: Wet conditions and warm temperatures increase manganese availability.

Interactions with other nutrients: High amounts of copper, iron, and zinc may induce manganese


Soil pH: Zinc availability decreases as pH increases.

Overliming decreases Zn solubility.

Zn adsorption: Though the relative amount of zinc on the cation exchange capacity is low, zinc is
attracted and held tightly to magnesite, dolomite and CaCO3 minerals. As a result, soils containing
these minerals can develop zinc deficiencies.

Organic Matter: Soluble zinc chelates increase zinc availability.

Climate: Cool, wet weather generally has a negative effect on zinc availability.

Flooding: Flooding generally decreases zinc availability.

Increasing soil temperatures increases zinc availability.

However, lowering the pH of flooded soils may increase zinc availability.

Interactions with other nutrients: Copper, iron, manganese, and phosphorus can interfere with zinc


Soil texture: Copper availability is lower in highly leached, coarse textured soils.

Soil pH: Copper availability decreases as pH increases, primarily due to decreased solubility of copper

Organic matter: Copper forms very tight bonds with organic matter (more so than any other
micronutrient), which may reduce its availability in organic (peat and muck) soils.

Buried Cu: Copper may be occluded, or buried, within the structure of clay minerals and oxides.
Occluded Cu is not available to plants.

Interactions with other nutrients: Copper availability to plants may be reduced when zinc, iron, and/or
phosphorus contents are high in the soil solution.


Soil pH: Unlike the other micronutrients, the availability of molybdenum increases with increasing pH.

As a result, liming acidic soils increases molybdenum availability.

Fe/Al oxides: Molybdenum is strongly held onto the surfaces of aluminum and iron oxides, which
reduces its availability.

Interactions with other nutrients: Copper and manganese can reduce the uptake of molybdenum by
plants. Phosphate enhances molybdenum uptake.

Soil moisture: Low levels of soil moisture reduce molybdenum availability.


Soil pH: Boron availability decreases as pH increases.

Liming can temporarily induce boron deficiencies, or lessen boron toxicities.

Soil organic matter: Organic matter increases boron availability.

Soil texture: Highly leached, coarse textured soils tend to have low boron availability.

Plant factors: The range between boron sufficiency and boron toxicity is very narrow. Crop sensitivity to
boron varies, and it is important to become familiar with the boron sensitivity of your crop.

Interactions with other nutrients: Crops are less sensitive to boron when there is ample amount of
calcium. This is because calcium acts to reduce boron availability. Boron may become deficient when
the Ca:B range is greater than 1,200:1.

Soil Moisture: Dry environments reduce the availability of boron.

Nutrition and Growth of Bacteria (page 1)

(This chapter has 6 pages)
Kenneth Todar, PhD
Nutritional Requirements of Cells
Every organism must find in its environment all of the substances required for energy generation and cellular
biosynthesis. The chemicals and elements of this environment that are utilized for bacterial growth are referred to
as nutrients or nutritional requirements. Many bacteria can be grown the laboratory in culture media which are
designed to provide all the essential nutrients in solution for bacterial growth. Bacteria that are symbionts or obligate
intracellular parasites of other cells, usually eucaryotic cells, are (not unexpectedly) difficult to grow outside of their
natural host cells. Whether the microbe is a mutualist or parasite, the host cell must ultimately provide the nutritional
requirements of its resident.
Many bacteria can be identified in the environment by inspection or using genetic techniques, but attempts to isolate
and grow them in artificial culture has been unsuccessful. This, in part, is the basis of the estimate that we may know
less than one percent of all procaryotes that exist.
The Major Elements
At an elementary level, the nutritional requirements of a bacterium such as E. coli are revealed by the cell's elemental
composition, which consists of C, H, O, N, S. P, K, Mg, Fe, Ca, Mn, and traces of Zn, Co, Cu, and Mo. These elements
are found in the form of water, inorganic ions, small molecules, and macromolecules which serve either a structural or
functional role in the cells. The general physiological functions of the elements are outlined in Table 1.

Table 1. Major elements, their sources and functions in bacterial cells.

% of dry
organic compounds
Main constituent of cellular material
or CO2
H2O, organic
Constituent of cell material and cell
compounds, CO2, water; O2 is electron acceptor in
and O2
aerobic respiration
NH3, NO3, organic Constituent of amino acids, nucleic
compounds, N2
acids nucleotides, and coenzymes
H2O, organic
Main constituent of organic
Hydrogen 8
compounds, H2
compounds and cell water
Constituent of nucleic acids,
Phosphorus 3
nucleotides, phospholipids, LPS,
phosphates (PO4)
teichoic acids
SO4, H2S, S ,
Constituent of cysteine, methionine,
organic sulfur
glutathione, several coenzymes
Main cellular inorganic cation and
Potassium 1
Potassium salts
cofactor for certain enzymes
Inorganic cellular cation, cofactor for
Magnesium 0.5
Magnesium salts
certain enzymatic reactions
Inorganic cellular cation, cofactor for
Calcium salts
certain enzymes and a component of
Component of cytochromes and
Iron salts
certain nonheme iron-proteins and a
cofactor for some enzymatic reactions

Trace Elements
Table 1 ignores the occurrence of trace elements in bacterial nutrition. Trace elements are metal ions required by
certain cells in such small amounts that it is difficult to detect (measure) them, and it is not necessary to add them to
culture media as nutrients. Trace elements are required in such small amounts that they are present as "contaminants"
of the water or other media components. As metal ions, the trace elements usually act as cofactors for essential
enzymatic reactions in the cell. One organism's trace element may be another's required element and vice-versa, but
the usual cations that qualify as trace elements in bacterial nutrition are Mn, Co, Zn, Cu, and Mo.
Carbon and Energy Sources for Bacterial Growth
In order to grow in nature or in the laboratory, a bacterium must have an energy source, a source of carbon and other
required nutrients, and a permissive range of physical conditions such as O 2 concentration, temperature, and pH.
Sometimes bacteria are referred to as individuals or groups based on their patterns of growth under various chemical
(nutritional) or physical conditions. For example, phototrophs are organisms that use light as an energy source;
anaerobes are organisms that grow without oxygen; thermophiles are organisms that grow at high temperatures.
All living organisms require a source of energy. Organisms that use radiant energy (light) are called phototrophs.
Organisms that use (oxidize) an organic form of carbon are calledheterotrophs or (chemo)heterotrophs. Organisms
that oxidize inorganic compounds are called lithotrophs.

The carbon requirements of organisms must be met by organic carbon (a chemical compound with a carbon-hydrogen
bond) or by CO2. Organisms that use organic carbon areheterotrophs and organisms that use CO2 as a sole source of
carbon for growth are calledautotrophs.
Thus, on the basis of carbon and energy sources for growth four major nutritional types of procaryotes may be defined
(Table 2).

Table 2. Major nutritional types of procaryotes

Nutritional Type

Energy Source








Chemoautotrophs or
Chemoheterotrophs or

compounds, e.g. CO2
H2, NH3, NO2, H2S

Cyanobacteria, some
Purple and Green
Some Purple and
Green Bacteria
A few Bacteria and
many Archaea
Most Bacteria, some

Almost all eucaryotes are either photoautotrophic (e.g. plants and algae) or heterotrophic (e.g. animals, protozoa, fungi).
Lithotrophy is unique to procaryotes and photoheterotrophy, common in the Purple and Green Bacteria, occurs only in a
very few eucaryotic algae. Phototrophy has not been found in the Archaea, except for nonphotosynthetic light-driven
ATP synthesis in the extreme halophiles. journal fix mikronutrient