The evolution of siderophore production as a competitive trait

Microbes have the potential to be highly cooperative organisms. The archetype of microbial cooperation is often considered to be the secretion of siderophores, molecules scavenging iron, where cooperation is threatened by “cheater” genotypes that use siderophores without making them. Here, we show that this view neglects a key piece of biology: siderophores are imported by specific receptors that constrain their use by competing strains. We study the effect of this specificity in an ecoevolutionary model, in which we vary siderophore sharing among strains, and compare fully shared siderophores with private siderophores. We show that privatizing siderophores fundamentally alters their evolution. Rather than a canonical cooperative good, siderophores become a competitive trait used to pillage iron from other strains. We also study the physiological regulation of siderophores using in silico long‐term evolution. Although shared siderophores evolve to be downregulated in the presence of a competitor, as expected for a cooperative trait, privatized siderophores evolve to be upregulated. We evaluate these predictions using published experimental work, which suggests that some siderophores are upregulated in response to competition akin to competitive traits like antibiotics. Although siderophores can act as a cooperative good for single genotypes, we argue that their role in competition is fundamental to understanding their biology.     

微生物铁载体运输系统

铁是大多数生物生长所必需的元素,在生命活动中占有极其重要的地位.尽管自然界中铁元素含量丰富,但是大多敬好氧生物包括微生物都面临着铁缺乏的问题,铁载体运输系统是微生物获得铁元素的重要途径.本文就铁载体运输系统的组成、调控机制及当前微生物中的铁载体运输系统的进展作以综述.     

A Catecholic Siderophore Produced by the Thermoresistant Bacillus licheniformis VK21 Strain

Thermophilic and thermoresistant strains of bacilli were screened on a medium containing Chrome Azurol S for the producers of siderophores. It was found that the Bacillus licheniformis VK21 strain dramatically increases secretion of the metabolite, a chelator of Fe³⁺, in response to addition of manganese(II) salts. The growth of the producer on a minimal medium containing MnSO₄ under the conditions of iron deficiency is accompanied by the accumulation of a catecholic product, the content of which reaches maximum at the beginning of the stationary growth phase of culture. In the presence of FeCl₃, the amount of the catecholic product in the medium considerably decreases. The siderophore, called S_VK21, was isolated from the cultural medium and purified by reversed phase HPLC, and its siderophore function was confirmed by the test for the restoration of growth of producer cells in a medium containing EDTA. The UV spectrum of the siderophore has absorption maxima at 248 and 315 nm. According to the amino acid analysis and NMR spectrometry, the metabolite S_VK21 is 2,3-dihydroxybenzoyl-glycyl-threonine.     

A dicarboxylate with asymmetric coordination ability: Characterization of a two dimensional coordination polymer of copper(II) exhibiting spin canting

A copper(II) coordination polymer {[Cu₂(cpa)₂]}_n (1) (cpaH₂ = 4-carboxyphenoxyacetic acid) has been synthesized by hydrothermal technique. The X-ray structure of the compound shows that it is a 2D coordination polymer where Cu(II) ions are pentacoordinated in a square pyramidal fashion. The magnetic measurement of 1 in different magnetic fields suggest a weak ferromagnetic ordering taking place below 20 K, arising from spin canting phenomenon due to the non-coplanar basal planes of copper ions.     

Phenotypic plasticity of a cooperative behaviour in bacteria

There is strong evidence that natural selection can favour phenotypic plasticity as a mechanism to maximize fitness in animals. Here, we aim to investigate phenotypic plasticity of a cooperative trait in bacteria – the production of an iron‐scavenging molecule (pyoverdin) by Pseudomonas aeruginosa. Pyoverdin production is metabolically costly to the individual cell, but provides a benefit to the local group and can potentially be exploited by nonpyoverdin‐producing cheats. Here, we subject bacteria to changes in the social environment in media with different iron availabilities and test whether cells can adjust pyoverdin production in response to these changes. We found that pyoverdin production per cell significantly decreased at higher cell densities and increased in the presence of cheats. This phenotypic plasticity significantly influenced the costs and benefits of cooperation. Specifically, the investment of resources into pyoverdin production was reduced in iron‐rich environments and at high cell densities, but increased under iron limitation, and when pyoverdin was exploited by cheats. Our study demonstrates that phenotypic plasticity in a cooperative trait as a response to changes in the environment occurs in even the simplest of organisms, a bacterium.     

Siderophores mediate reduced and increased uptake of cadmium by Streptomyces tendae F4 and sunflower (Helianthus annuus), respectively

Aims: As a toxic metal, cadmium (Cd) affects microbial and plant metabolic processes, thereby potentially reducing the efficiency of microbe or plant‐mediated remediation of Cd‐polluted soil. The role of siderophores produced by Streptomyces tendae F4 in the uptake of Cd by bacteria and plant was investigated to gain insight into the influence of siderophores on Cd availability to micro‐organisms and plants. Methods and Results: The bacterium was cultured under siderophore‐inducing conditions in the presence of Cd. The kinetics of siderophore production and identification of the siderophores and their metal‐bound forms were performed using electrospray ionization mass spectrometry. Inductively coupled plasma spectroscopy was used to measure iron (Fe) and Cd contents in the bacterium and in sunflower plant grown in Cd‐amended soil. Siderophores significantly reduced the Cd uptake by the bacterium, while supplying it with iron. Bacterial culture filtrates containing three hydroxamate siderophores secreted by S. tendae F4 significantly promoted plant growth and enhanced uptake of Cd and Fe by the plant, relative to the control. Furthermore, application of siderophores caused slightly more Cd, but similar Fe uptake, compared with EDTA. Bioinoculation with Streptomyces caused a dramatic increase in plant Fe content, but resulted only in slight increase in plant Cd content. Conclusion: It is concluded that siderophores can help reduce toxic metal uptake in bacteria, while simultaneously facilitating the uptake of such metals by plants. Also, EDTA is not superior to hydroxamate siderophores in terms of metal solubilization for plant uptake. Significance and Impact of the Study: The study showed that microbial processes could indirectly influence the availability and amount of toxic metals taken up from the rhizosphere of plants. Furthermore, although EDTA is used for chelator‐enhanced phytoremediation, microbial siderophores would be ideal for this purpose.     

Characterization of ‘Schizokinen’; a dihydroxamate-type siderophore produced by Rhizobium leguminosarum IARI 917

The Rhizobia comprise one of the most important groups of beneficial bacteria, which form nodules on the roots (rarely on the stems) of leguminous plants. They live within the nodules and reduce atmospheric nitrogen to ammonia, which is further assimilated by plants into required nitrogenous compounds. The Rhizobia in return obtain nutrition from the plant. Rhizobia are free-living soil bacteria and have to compete with other microorganisms for the limited available iron in the rhizosphere. In order to acquire iron Rhizobia have been shown to express siderophore-mediated iron transport systems. Rhizobium leguminosarum IARI 917 was investigated for its ability to produce siderophore. It was found to produce a dihydroxamate type siderophore under iron restricted conditions. The siderophore was purified and chemically characterized. The ESMS, MS/MS and NMR analysis indicate the dihydroxamate siderophore to be ‘schizokinen’, a siderophore reported to be produced by Bacillus megaterium that shares a similar structure to ‘rhizobactin 1021’ produced by Sinorhizobium meliloti 1021. This is the first report of production of schizokinen by a strain of R. leguminosarum, therefore it was carefully investigated to confirm that it is indeed ‘schizokinen’ and not a degradation product of ‘rhizobactin 1021’. Since ferric–siderophore complexes are transported across the outer membrane (OM) into the periplasm via an OM receptor protein, R. leguminosarum IARI 917 was investigated for the presence of an OM receptor for ‘ferric–schizokinen’. SDS PAGE analysis of whole cell pellet and extracted OM fractions indicate the presence of a possible iron-repressible OM receptor protein with the molecular weight (MW) of approximately 74 kDa.     

Bacterial siderophores promote plant growth: Screening of catechol and hydroxamate siderophores

The aim of the study was to determine the quality and quantity of siderophores produced by bacteria isolated from plants' roots. The second aim was to determine the effect of siderophores on plants growth (Festuca rubra L. and Brassica napus L.). The study was carried out using bacteria isolated from roots of: Arabidopsis thaliana L., F. rubra, and Agrostis capillaris L., growing on the heavy metals contaminated area. The chrome azurol sulfonate (CAS) test, Arnow's test for catechol siderophores, and Csaksy's test for hydroxamate siderophores were performed. Among the bacteria, 42 isolates (39%) had a positive result in the CAS. Endophytic bacteria were mostly producing the catechol siderophores. It was found that F. rubra is the plant which is linked with the highest number of siderophores producing bacteria. The highest concentration of siderophores was noted for ectorhizospheric bacteria associated with A. thaliana, hyperaccumulating plant. It was found that hydroxamate siderophores are mainly produced by ectorhizosphere and rhizoplane bacteria. The siderophores producing bacteria reduced the toxicity of metals and improved the phytoremediation. Siderophores treatment increased the growth of plants in the biological assay, growing on two different soils: one highly contaminated with heavy metals and the second strongly alkaline soil.     

Monocyclopentadienyl titanium complexes supported by functionalized carboxylate ligands

The reaction of [TiCp*Cl₃] with [Fe(η⁵-C₅H₅)(η⁵-C₅H₄COOH)] in the presence of NEt₃ yields [TiCp*{(OOC-C₅H₄)FeCp}₃] (1), (Cp = η⁵-C₅H₅). The alkyl complex [TiCp*Me₃] reacts with [FeCp(η⁵-C₅H₄-CH₂COOH)] or anthranilic acid rendering the tris-carboxylate titanium complexes [TiCp*{(OOCCH₂-C₅H₄)FeCp}₃] (2) and [TiCp*{(OOCC₆H₄NH₂)₃] (3), respectively. Complex 3 can be protonated with triflic acid to render [TiCp*{(OOCC₆H₄NH₂)₃].HOTf (4). The reaction of [TiCp*Me₃] with anthranilic acid in a 1:2 M ratio yields the alkyl carboxylate derivative [TiCp*Me{(OOCC₆H₄NH₂)₂] (5). Complex 5 reacts with ^tBuNC to render the iminoacyl complex [TiCp*(η²-MeCN^tBu){(OOCC₆H₄NH₂)₂] (6). The reaction of [TiCp*Cl₃] with the ferroceneacetic acid, gives [TiCp*Cl₂{(OOCCH₂-C₅H₄)FeCp}] (7). The [TiCp*Cl]₂(μ-O)[(ΟΟC-C₅H₄)₂Fe] (8) can be obtained by reaction of [TiCp*Cl₃] with [Fe(η⁵-C₅H₄-COOH)₂] in the presence of a base. The molecular structures of 1 and 8 have been established by X-ray diffraction methods.     

Hydrothermal syntheses, crystal structures and topological analyses of four new 3D coordination polymers with mixed ligands

Four new coordination polymers, namely [Ni_1.5(L)(fum)_0.5(mal)(H₂O)]·4H₂O (1), [Zn₂(L)(male)₂]·3H₂O (2), [Ni(L)(adi)_0.5(H₂O)]·0.5(adi)·2.5H₂O (3), and [Zn₂(L)(adi)₂]·5H₂O (4) (L, tetrakis(imidazol-1-ylmethyl)methane; fum, fumarate; mal, malate; male, maleate; adi, adipate) have been synthesized under hydrothermal condition and characterized by elemental analysis, IR, and single-crystal X-ray diffraction. In compound 1, the L, fum, and mal connect the Ni(II) atoms to form a three-dimensional framework. Topologically, the compound shows a trinodal (3,4)-connected (4·10²)(4²·8²·10²)(4²·8⁴) net. Interestingly, the fum and mal came from the configuration transformation and addition reaction of maleate during the synthesis, respectively. In compound 2, the L ligands link the Zn(II) centers to generate a 2D layer, which are further connected by the male ligands to furnish a 3D chrial 4-connected (3·7⁵) net. In compound 3, the adi ligands and the L ligands link the Ni(II) centers to form a 3D framework. Topologically, compound 3 displays a (4,5)-connected net with the Schläfli symbol of (4²·5²·6²)(4²·5³·6⁴·7). In compound 4, the L and adi ligands link the binuclear Zn(II) clusters to yield an intricate 3D (3,6)-connected net with the Schläfli symbol of (4·6²)(4³·6¹²). The fluorescence and thermal stability of complexes 1-4 have also been investigated.