Phytosiderophores, organic

Reabsorption of the ligand plus its metal partner is a necessary requirement of processes like Fe acquisition by phytosiderophores (32). However, whether or not reabsorption of diffusates, which undoubtedly occurs in solution cultures (45), has a significant role to play is uncertain, largely because in soil most diffusates (sugars, amino acids, and other organic acids) are readily utilized by microorganisms or adsorbed by soil colloids. 

Nutrient availability also plays a major role in exudation, with deficiencies in N, P, or K often increasing the rate of exudation (218). It is believed that nutrient deficiency may trigger the release of substances such as organic acids or nonproteinogenic amino acids (phytosiderophores), which may enhance the acquisition of the limiting nutrient (219,220). An example here might be the release of phenolic acids such as caffeic acid in response to iron deficiency, which results in an increase in uptake of the cation (221). 

In the first case the mechanisms are based on an increased reducing capacity of Fe(lll)-chelates, a necessary step in the uptake process, with a concurrent increase in acidification and release of organic acids into the rhizosphere in the latter case molecules having high affinity for Fe (phytosiderophores) are synthesized and released into the rhizosphere when Fe is lacking. 

In the rhizosphere, microorganisms utilize either organic acids or phytosiderophores to transport iron or produce their own low-molecular-weight metal chelators, called siderophores. There are a wide variety of siderophores in nature and some of them have now been identified and chemically purified (54). Pre.sently, three general mechanisms are recognized for utilization of these compounds by microorganisms. These include a shuttle mechanism in which chelators deliver iron to a reductase on the cell surface, direct uptake of metallated siderophores with destructive hydrolysis of the chelator inside the cell, and direct uptake followed by reductive removal of iron and resecretion of the chelator (for reviews, see Refs. 29 and 54). 

These data strongly suggest that siderophore production by root-colonizing microorganisms is induced only at a level neeessary to supplement that which is not provided by phytosiderophores and organic acids released during the plant iron stress response. Thus, the plant iron stress response may control iron availability to microorganisms in the rhizosphere. 

Dissolved organic carbon and plant- or microbe- produced phytosiderophores increase the solubility of most trace elements in arid and semi-arid soils. This is especially important in arid regions. High pH increases the solubility of organic matter as dissolved organic carbon in arid soils. Copper, lead and nickel have a strong tendency to form complexes, while Cd complexes are weaker. Zinc, cobalt and manganese are... 

Dissolved organic molecules have many acidic functions (hydroxol and carbonic groups) to complex trace elements and their compounds to form soluble chelates. This is one of the reasons why solubility and bioavailability of trace elements in the rhizosphere are higher than bulk soils. At the same time, many organic acids also directly dissolve trace elements and their compounds in soils. Plant-produced phytosiderophores facilitate elements, such as Fe and Zn, uptake by plants (Zhang et al., 1991 Romheld, 1991 Hopkins et al., 1998). However, Shenker et al. (2001) did not find significant uptake of the Cd-phytosiderophores complex by plant roots. 

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