Microbial Strategy Shift for Releasing Iron with Varying Mineral Substrates and their Accessibility: Implications for Iron Biomining

Summary Poor iron ( F e) availability in soil represents one of the most important limiting factors of agricultural production and is closely linked to physical, chemical and biological processes within the rhizosphere as a result of soil–microorganism–plant interactions. Iron shortage induces several mechanisms in soil organisms, resulting in an enhanced release of inorganic (such as protons) and organic (organic acids, carbohydrates, amino acids, phytosiderophores, siderophores, phenolics and enzymes) compounds to increase the solubility of poorly available F e pools. However, rhizospheric organic compounds (ROCs) have short half‐lives because of the large microbial activity at the soil–root interface, which might limit their effects on F e mobility and acquisition. In addition, ROCs also have a selective effect on the microbial community present in the rhizosphere. This review aims therefore to unravel these complex dynamics with the objective of providing an overview of the rhizosphere processes involved in F e acquisition by soil organisms (plants and microorganisms). In particular, the review provides information on (i) F e availability in soils, including mineral weathering and F e mobilization from soil minerals, ligand and element competition and plant‐microbe competition; (ii) microbe–plant interactions, focusing on beneficial microbial communities and their association with plants, which in turn influences plant mineral nutrition; (iii) plant–soil interactions involving the metabolic changes triggered by F e deficiency and the processes involved in exudate release from roots; and (iv) the influence of agrochemicals commonly used in agricultural production systems on rhizosphere processes related to F e availability and acquisition by crops.

The long-term stability of nuclear waste form borosilicate glasses can be evaluated by understanding the processes that effect the long-term alteration of glass and by comparing laboratory alteration of synthetic basalt and borosilicate glasses with the observed stability of naturally occurring basaltic glasses in diverse geologic environments. This paper presents detailed electron microprobe analyses of naturally altered basaltic glasses (with maximum ages of 10,000 to 20 million years) from low-temperature environments. These results are compared to laboratory data on the corrosion of a synthetic basaltic glass in MCC-1 tests (90°C, a SA/V of 0.1 cm−1 and time periods up to 182 days), MCC-2 tests (190°C, a SA/V of 0.1 cm−1 and time periods up to 210 days) and hydration tests in saturated water vapor (240°C, an estimated SA/V of ∼ 106 cm−1 and time periods up to 63 days). Additionally, laboratory induced hydration alteration of synthetic basalt and borosilicate glasses is compared. These preliminary experiments provide evidence that the alteration processes observed for natural basalt glasses are relevant to understanding the alteration of nuclear waste glass, as both appear to react via similar processes.

The long-term stability of nuclear waste form borosilicate glasses can be evaluated by understanding the processes that effect the long-term alteration of glass and by comparing laboratory alteration of synthetic basalt and borosilicate glasses with the observed stability of naturally occurring basaltic glasses in diverse geologic environments. This paper presents detailed electron microprobe analyses of naturally altered basaltic glasses (with maximum ages of 10,000 to 20 million years) from low-temperature environments. These results are compared to laboratory data on the corrosion of a synthetic basaltic glass in MCC-1 tests (90/sup 0/C, a SA/V of 0.1 cm/sup -1/ and time periods up to 182 days), MCC-2 tests (190/sup 0/C, a SA/V of 0.1 cm/sup -1/ and time periods up to 210 days) and hydration tests in saturated water vapor (240/sup 0/C, an estimated SA/V of approx. 10/sup 6/ cm/sup -1/ and time periods up to 63 days). Additionally, laboratory-induced hydration alteration of synthetic basalt and borosilicate glasses is compared. These preliminary experiments provide evidence that the alteration processes observed for natural basalt glasses are relevant to understanding the alteration of nuclear waste glass, as both appear to react via similar processes. 12 references, 6 figures, 1 table.

Iron acquisition by Mycobacterium tuberculosis residing within myeloid dendritic cells

Acid‐leached rinds and coatings occur in volcanic environments on Earth and have been identified using orbital spectroscopy on Mars, but their development is poorly understood. We simulated long‐term open‐system acidic weathering in a laboratory by repeatedly rinsing and submerging crystalline and glassy basalts in pH ~ 1 and pH ~ 3 acidic solutions for 213 days and compared their visible/near‐infrared (0.3–2.5 µm) and thermal infrared (5–50 µm) spectral characteristics to their microscopic physical and chemical properties from scanning electron microscopy (SEM). We find that while alteration at moderately low pH (~3) can produce mineral precipitates from solution, it has very little spectral or physical effect on the underlying parent material. In contrast, alteration at very low pH (~1) results in clear silica spectral signatures for all crystalline samples while glasses exhibit strong blue concave‐up near‐infrared slopes. SEM indicates that these spectral differences correspond to different modes of alteration. In glass, alteration occurs only at the surface and produces a silica‐enriched leached rind, while in more crystalline samples, alteration penetrates the interior to cause dissolution and replacement by silica. We confirm that glass is more stable than crystalline basalt under long‐term acidic leaching, suggesting that glass could be enriched and common in terrains on Mars that have been exposed to acidic weathering. Leached glasses are consistent with both OMEGA and Thermal Emission Spectrometer (TES) spectra of the Martian northern lowlands and may contribute to the high‐silica phases detected globally in TES Surface Type 2. Thus, both glass‐rich deposits and acidic weathering may have been widespread on Mars.

Alteration of basalt hyaloclastite at the off-axis Sea Cliff hydrothermal field, Gorda Ridge

Characterization of LuxI/LuxR and their regulation involved in biofilm formation and stress resistance in fish spoilers Pseudomonas fluorescens

Alteration textures were examined in subglacial (hyaloclastite) deposits at Valafell, Southern Iceland. Pitted and 'elongate' alteration features are observed in the glass similar to granular and tubular features reported previously in deep-ocean basaltic glasses, but elongate features generally did not have a length to width ratio greater than five. Elongate features were found in only 7% of surfaces. Crystalline basalt clasts, which are incorporated into the hyaloclastite, did not display elongate structures. Pitted alteration features were poorly defined in crystalline basalt, comprising only 4% of the surface compared to 47% in the case of basaltic glass. Examination of silica-rich glass (obsidian) and rhyolite similarly showed poorly defined pitted textures that comprised less than 15% of the surface and no elongate features were observed. These data highlight the differences in alteration textures between terrestrial basaltic glass and previously studied deep-ocean and subsurface basaltic glass, and the important role of mineralogy in controlling the type and abundance of alteration features. The hyaloclastite contains a diverse and abundant bacterial population, as determined by 16S rDNA analysis, which could be involved in weathering the glass. Despite the presence of phototrophs, we show that they were not involved in the production of most alteration textures in the basaltic glass materials we examined.

Temporal dynamics of litter quality, soil properties and microbial strategies as main drivers of the priming effect

Organisms have developed different strategies to cope with environmental conditions of low Fe availability based on the exudation of reducing, ligating, and acidifying compounds. In the context of Fe acquisition from soil, the effects of these reactive compounds have generally been considered independent and additive. However, highly efficient Fe acquisition strategies may rely on synergistic effects between reactive exudates. In the present study, we demonstrate that synergistic effects between biogenic ligands and a reductant (ascorbate) can occur in Fe mobilization from soil. Synergistic Fe mobilization was found for all ligands examined (desferrioxamine B (DFOB), 2'-deoxymugineic acid (DMA), esculetin, and citrate). The size and duration of the synergistic effect on Fe mobilization varied with ligand: larger effects were observed for the sideorphores compared to esculetin and citrate. For DFOB, the synergistic effect lasted for the 168 h duration of the experiment; for DMA, an initial synergistic effect turned into an antagonistic effect after 4 h because of enhanced mobilization of competing metals; and for esculetin and citrate, the synergistic effect was temporary (less than 24 h). Our results demonstrate that synergistic effects greatly enhance the reactivity of mixtures of compounds known to be exuded in response to Fe limitation. These synergistic effects could be decisive for the survival of plants and microorganisms under conditions of low Fe availability.

Heavy metal and sulfur transport during subcritical and supercritical hydrothermal alteration of basalt: Influence of fluid pressure and basalt composition and crystallinity

The interaction of a single bacterial species (Burkholderia fungorum) with basaltic rocks from the ICDP HSDP2 drill core and synthetic basaltic glasses was investigated in batch laboratory experiments to better understand the role of microbial activity on rock alteration and Fe mobilization. Incubation experiments were performed with drill core basaltic rock samples to investigate differences in the solution chemistry during biotic and abiotic alteration. Additionally, colonization experiments with synthetic basaltic glasses of different Fe redox states and residual stresses were performed to evaluate their influence on microbial activity and surface attachment of cells. In biotic incubation experiments bacterial growth was observed and the release of Fe and other major elements from drill core basaltic rocks to solution exceeded that of abiotic controls only when the rock sample assay was nutrient depleted. The concentration of dissolved major elements in solution in biotic colonization experiments with synthetic basaltic glasses increased with increasing residual stress and Fe(II) content. Furthermore, the concentration of dissolved Fe and Al increased similarly in biotic colonization experiments indicating that their dissolution might be triggered by microbial activity. Surface morphology imaging by SEM revealed that cells on basaltic rocks in incubation experiments were most abundant on the glass and surfaces with high roughness and almost absent on minerals. In colonization experiments, basaltic glasses with residual stress and high Fe(II) content were intensely covered with a cellular biofilm. In contrast, glasses with high Fe(III) content and no residual stress were sparsely colonized. We therefore conclude that structurally bound Fe is most probably used by B. fungorum as a nutrient. Furthermore, we assume that microbial activity overall increased rock dissolution as soon as the environment becomes nutrient depleted. Our results show that besides compositional effects, other factors such as redox state and residual stress can control microbial alteration of basaltic glasses.

Poorly crystalline Fe oxides are sources of Fe to plants. The adsorption of humic substances (HS) on these oxides alters its reactivity and stability in soils, and thus may affect Fe mobilization and uptake by plants from these compounds. This work aimed at studying how the adsorption of HS on Fe oxides affects its use as Fe source by two plant species with different Fe acquisition strategies, white lupin (Strategy I) and wheat (Strategy II). To this end, two completely randomized experiments, one with each plant, were carried out using a calcareous growing media and involving increasing amounts of HS adsorbed on ferrihydrite (0, 16, 60, and 97 mg C g–1) which was used as Fe source. The highest HS rate was the only treatment that significantly increased Fe uptake in wheat relative to control without HS. This was related to a decreased concentration of Fe in poorly crystalline oxides in the growing media. On the contrary, HS did not affect significantly Fe uptake by lupin. However, in this crop, the highest HS rate decreased the concentration of Fe in oxides relative to the lowest HS rate, without significant differences with other treatments. Thus, the effect of adsorbed HS on Fe uptake differed in two plants with different Fe acquisition strategies. The increased Fe uptake in wheat at the highest HS rate can be explained at least in part by an increased Fe mobilization from oxides by plant roots. These findings provide new insights on the role of soil organic matter on plant Fe nutrition.

Many bacteria employ chemical communication to coordinate group behaviors, a process that has been termed quorum sensing (QS). Although many different classes of bacteria use QS, the mechanism that has emerged as common in gram-negative proteobacteria is based on acyl-homoserine lactone (acyl-HSL) signal molecules. QS is an important field of study because of its important regulatory roles in pathogenic and environmental bacteria. Acyl-homoserine lactone (Acyl-HSL) QS controls virulence in many plant and animal pathogens such as Pseudomonas aeruginosa, Burkholderia cepacia, and Agrobacterium tumefaciens. This chapter focuses on the integration of acyl-HSL QS and such stress responses. In P. aeruginosa, QS controls the expression of numerous virulence factors such as extracellular enzymes (LasB elastase, LasA protease, alkaline protease), secondary metabolites (pyocyanin, hydrogen cyanide, pyoverdin), and toxins (exotoxin A). Microarray studies revealed that the las and rhl systems together control the expression of more than 300 genes. The majority of the associated genes are predicted to encode secreted factors and secretion machinery, confirming the notion that the core function of las QS is to control the expression of extracellular factors. Indeed, a recent population analysis of concurrently isolated P. aeruginosa from individual Cystic fibrosis (CF) lung infections revealed great heterogeneity of QS phenotypes and genotypes within as well as among patients. This finding suggests that a single selective mechanism, whether of social or nonsocial nature, is unlikely to explain the emergence of QS variants during CF infection.

Arbuscular mycorrhizal (AM) fungi can improve iron (Fe) acquisition of their host plants. Here, we report a characterization of two components of the high-affinity reductive Fe uptake system of Rhizophagus irregularis, the ferric reductase (RiFRE1) and the high affinity Fe permeases (RiFTR1-2). In the extraradical mycelia (ERM), Fe deficiency induced activation of a plasma membrane-localized ferric reductase, an enzyme that reduces Fe(III) sources to the more soluble Fe(II). Yeast mutant complementation assays showed that RiFRE1 encodes a functional ferric reductase and RiFTR1 an iron permease. In the heterologous system, RiFTR1 was expressed in the plasma membrane while RiFTR2 was expressed in the endomembranes. In the ERM, the highest expression levels of RiFTR1 were found in mycelia grown in media with 0.045mM Fe, while RiFTR2 was upregulated under Fe-deficient conditions. RiFTR2 expression also increased in the intraradical mycelia (IRM) of maize plants grown without Fe. These data indicate that the Fe permease RiFTR1 plays a key role in Fe acquisition and that RiFTR2 is involved in Fe homeostasis under Fe-limiting conditions. RiFTR1 was highly expressed in the (IRM), which suggests that the maintenance of Fe homeostasis in the IRM might be essential for a successful symbiosis.