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90<br />

Ramesh Chander Kuhad et al.<br />

in solubility and transport of iron, hydroxamate siderophores are also<br />

involved in iron storage.<br />

Based on the chemical nature of their coordination sites, microbial<br />

siderophores are classified as hydroxamates, catecholates, carboxylates and<br />

mixed type. Hydroxamates are produced both by bacteria and fungi. In most<br />

fungi, a mixture of siderophores is produced which varies depending on cultivation<br />

conditions. Aspergilli produce ferricrocin accompanied by fusarinines,<br />

while certain penicillia produce ferrichrome accompanied by coprogen.<br />

Similar observations have been made with Neurospora, Gliocladium,<br />

Trichoderma and Agaricus bisporus (Neilands and Leong 1986). Fusarinines<br />

(fusigens) produced by species of Fusarium and Penicillium are linear and<br />

cyclic hydroxamic acids joined by ester bonds. Ericoid mycorrhizal fungi produce<br />

ferrichrome and fusarinines.<br />

Varieties of bacterial hydroxamates are known. Ferrioxamine, produced by<br />

actinomycetes, Nocardia and Pseudomonas stutzeri, is a cyclic trihydroxamate.<br />

Citrate hydroxamates are characterized by the presence of two hydroxamates<br />

and one citrate group as ligand, it is a linear citratehydroxamic acid<br />

obtained from Klebsiella pneumonia and several enteric bacteria. Catecholate<br />

siderophores are generally less diverse than the hydroxamates, and are conjugated<br />

to amino acids or polyamine backbones. Species of Bacillus, Aeromaonas<br />

and Erwinia are known to produce catecholate siderophores. Carboxylate<br />

(complexone) siderophores are produced by Rhizopus microsporus,<br />

Rhizobium meliloti and Staphylococcus hycius. Pyoverdines, the mixed types<br />

form a wide class of mixed siderophores showing a great variety of structures.<br />

Some strains of fluorescent pseudomonads produce hydroxamate siderophores<br />

(ferribactin) in addition to pyoverdine siderophores.<br />

The <strong>plant</strong> growth-promoting rhizobacteria (PGPR) owe their <strong>plant</strong> growth<br />

promoting activity to their stronger siderophores with higher stability constants<br />

that outgrow the other bacterial population in competition for iron and<br />

finally displace them from the root <strong>surface</strong>. The siderophore-producing PGPR<br />

have become important in the biological control of <strong>plant</strong> pathogens (Glick<br />

and Bashan 1997).<br />

5 Conclusions<br />

Microorganisms play an essential role in the functioning and sustainability of<br />

soil ecosystems including biogeochemical cycling of nutrients and biodegradation.<br />

Recent advances in soil community analysis using molecular and biochemical<br />

approaches have helped us understand the enormous microbial<br />

diversity and their functional significance in nutrient recycling in soil and<br />

<strong>plant</strong> development. Soil diversity exceeds that of aquatic environments and<br />

provides a great resource for the biological exploitation of novel organisms,<br />

processes and products. Microbes isolated from soil and developed as biofer-

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