Microbial geochemical calcium cycle

The participation of microorganisms in the geochemical calcium cycle is the most important factor maintaining neutral conditions on the Earth. This cycle has profound influence on the fate of inorganic carbon, and, thereby, on the removal of CO2 from the atmosphere. The major part of calcium deposits was formed in the Precambrian, when prokaryotic biosphere predominated. After that, calcium recycling based on biogenic deposition by skeletal organisms became the main process. Among prokaryotes, only a few representatives, e.g., cyanobacteria, exhibit a special calcium function. The geochemical calcium cycle is made possible by the universal features of bacteria involved in biologically mediated reactions and is determined by the activities of microbial communities. In the prokaryotic system, the calcium cycle begins with the leaching of igneous rock predominantly through the action of the community of organotrophic organisms. The release of carbon dioxide to the soil air by organotrophic aerobes leads to leaching with carbonic acid and soda salinization. Under anoxic conditions, of major importance is the organic acid production by primary anaerobes (fermentative microorganisms). Calcium carbonate is precipitated by secondary anaerobes (sulfate reducers) and to a smaller degree by methanogens. The role of the cyanobacterial community in carbonate deposition is exposed by stromatolites, which are the most common organo-sedimentary Precambrian structures. Deposition of carbonates in cyanobacterial mats as a consequence of photoassimilation of CO2 does not appear to be a significant process. It is argued that carbonates were deposited at the boundary between the "soda continent", which emerged as a result of subaerial leaching with carbonic acid, and the ocean containing Ca2+. Such ecotones provided favorable conditions for the development of the benthic cyanobacterial community, which was a precursor of stromatolites.     

Transformation of carbonate minerals in a cyano-bacterial mat in the course of laboratory modeling

A laboratory model of a cyano-bacterial mat with mineral layers of carbonates was used to examine the dynamics of the transformation of calcium-magnesium carbonate under the conditions of a soda lake. The activity of various organisms of the cyanobacterial community results in conditions under which the Ca-Mg carbonate precipitate undergoes changes. The crystal lattice of the initial carbonate is restructured; its mineralogical composition changes depending on the conditions of the mat. In magnesium calcites, which are formed under such low-temperature conditions, a rudimentary cation adjustment can occur with the formation of dolomite domains. These experiments confirm the hypothesis that the dolomite found in stromatolites is of a secondary origin and can be formed in the course of transformation of Ca-Mg carbonates under alkaline conditions in an alkaliphilic cyanobacterial community.     

Transformation of carbonate minerals in a cyano-bacterial mat in the course of laboratory modeling

A laboratory model of a cyano-bacterial mat with mineral layers of carbonates was used to examine the dynamics of the transformation of calcium-magnesium carbonate under the conditions of a soda lake. The activity of various organisms of the cyanobacterial community results in conditions under which the Ca-Mg carbonate precipitate undergoes changes. The crystal lattice of the initial carbonate is restructured; its mineralogical composition changes depending on the conditions of the mat. In magnesium calcites, which are formed under such low-temperature conditions, a rudimentary cation adjustment can occur with the formation of dolomite domains. These experiments confirm the hypothesis that the dolomite found in stromatolites is of a secondary origin and can be formed in the course of transformation of Ca-Mg carbonates under alkaline conditions in an alkaliphilic cyanobacterial community.     

Ecophysiology and polymorphism of the unicellular extremely natronophilic cyanobacterium <Emphasis Type="Italic">Euhalothece</Emphasis> sp. Z-M001 from Lake Magadi

Strain Z-M001 of a unicellular cyanobacterium, assigned by analysis of the 16S rRNA gene sequence to the phylogenetic group of the generic level Euhalothece, was isolated from soda Lake Magadi. It was shown that strain Z-M001, unlike all other known cultured and uncultured organisms of the Euhalothece group, is extremely natronophilic, and it was named accordingly “Euhalothece natronophila”. In its ecophysiological characteristics, it is comparable to extremely alkaliphilic organotrophic natronobacteria, which is essential for soda ecosystems, because cyanobacteria belong to primary producers. E. natronophila exhibits considerable morphological variability depending on the concentration of carbonates in the medium. The polymorphism of “ E. natronophila” is primarily connected to limitation by utilizable forms of carbon.     

Specific features of the system of carbonic anhydrases of alkaliphilic cyanobacteria

This review brings together our recent data on carbonic anhydrases of representatives of alkaliphilic cyanobacteria inhabiting soda lakes, which are considered as the relicts of the ancient terrestrial microbiota. The modern information about cyanobacterial carbonic anhydrases is based mainly on the study of model strains, such as Synechocystis and Synechococcus. Our results are compared with literature data. The role of carbonic anhydrases in the assimilation of inorganic carbon by cyanobacteria of soda lakes is discussed in terms of evolution of the CO₂-concentrating mechanism.     

Compartmentation of α- and β-Carbonic Anhydrases in Cells of Halo- and Alkalophilic Cyanobacteria Rhabdoderma lineare

The organization of carbonic anhydrase (CA) system in halo- and alkalophilic cyanobacteria Rhabdoderma lineare was studied by Western blot analysis and immunocytochemical electron microscopy. The presence of extracellular -CA (60 kD) in the glycocalyx, forming a tight sheath around the cell, and of two intracellular -CA is reported. One -CA (60 kD) is associated with polypeptides of photosystem II (PSII) and is a constitutive enzyme. Another -carbonic anhydrase (25 kD) was induced by low content of bicarbonate in the culture medium; this inducible CA was found in the fraction of total soluble proteins. The expressed synthesis of inducible -CA was accompanied by the increase in the intracellular pool of inorganic carbon, which suggests an important role of this enzyme in the functioning of CO₂-concentrating mechanism.     

Carbon isotope discrepancy between precambrian stromatolites and their modern analogs: Inferences from hypersaline microbial mats of the sinai coast

The isotopic composition of organic carbon from extant stromatolite-type microbial ecosystems is commonly slanted toward heavy ¹³ C values as compared to respective compositions of average organic matter (including that from Precambrian stromatolites). This seems the more enigmatic as the bulk of primary producers from benthic microbial communities are known to fix carbon via the C3 pathway normally entailing the sizable fractionations of the RuBP carboxylase reaction.There is reason to believe that the small fractionations displayed by aquatic microorganisms result from the limitations of a diffusion-controlled assimilatory pathway in which the isotope effect of the enzymatic reaction is largely suppressed. Apart from the diffusion-control exercised by the aqueous environment, transport of CO₂ to the photosynthetically active sites will be further impeded by the protective slime (polysaccharide) coatings commonly covering microbial mats in which gas diffusivities are extremely low. Ineffective discrimination against¹³C becomes, however, most pronounced in hypersaline environments where substantially reduced CO₂ solubilities tend to push carbon into the role of a limiting nutrient (brine habitats constitute preferential sanctuaries of mat-forming microbenthos since the emergence of Metazoan grazers 0.7 Ga ago). As the same microbial communities had been free to colonize normal marine environments during the Precambrian, the CO₂ concentration effect was irrelevant to the carbon-fixing pathway of these ancient forms. Therefore, it might not surprise that organic matter from Precambrian stromatolites displays the large fractionations commonly associated with C3 photosynthesis. Increased mixing ratios of CO₂ in the Precambrian atmosphere may have additionally contributed to the elimination of the diffusion barrier in the carbon-fixing pathways of ancient mat-forming microbiota.     

Carbonic anhydrase: Enzyme that has transformed the biosphere

The bases of modern type biosphere were laid down about two billion years ago during the predominance of prokaryotes on the Earth. Cyanobacteria changed radically the composition of the Proterozoic atmosphere by saturating it with photosynthetic oxygen. At the same time, large quantities of atmospheric CO₂ became sequestered in carbonates owing to mineralization of ancient cyano-bacterial communities; the latter have reached us in the form of laminated limestone deposits, termed stromatolites. The mechanism of carbonate depositing by cyanobacteria is still poorly understood. It is not yet clear whether physiological processes are involved in cell mineralization or if the outer membranes of cyanobacteria serve as a kind of crystallization center and arrange the structure for natural accumulation of sediments. We proposed that a key role in the mechanism of biomineralization belongs to the enzyme carbonic anhydrase (CA), which regulates the equilibrium between the inorganic carbon forms (C_i), including bicarbonate that participates in natural sedimentation of calcium. Since the deposition of calcium carbonate by prokaryotes occurs in the pericellular space and this deposition is controlled by pH, it seems likely that CA, localized on the periphery of cyanobacterial cells, is involved in stabilizing the external pH and in promoting cell mineralization. This review summarizes information concerning possible mechanisms of biogenic calcification (CaCO₃ deposition). The function of CA in the living cell and the role of this enzyme in biological processes are considered, and the data on localization of CA in cyano-bacterial cells are presented. Based on available evidence, a scheme is suggested to describe the role of extracellular CA in photosynthetic carbon assimilation and to relate this process with CaCO₃ deposition during mineralization of cyanobacteria.     

Identification and characterization of the main β-alanine uptake system in <Emphasis Type="Italic">Escherichia coli</Emphasis>

In Escherichia coli, -alanine is a direct precursor in the biosynthesis of pantothenic acid (vitamin B₅). Although a sufficient -alanine supply is crucial for biotechnological vitamin B₅ production, nothing was known about -alanine transport in E. coli until now. The aim of this work was the characterization of -alanine transport by E. coli and the identification and overexpression of the corresponding carrier-encoding gene for the rational improvement of pantothenic acid-producing strains. -Alanine uptake was found to be an active process catalyzed by the amino acid carrier CycA. The corresponding gene was cloned and overexpressed, resulting in an increase in the uptake rate, compared with the wild type. In all tested strains, this overexpression led to a strong sensitivity to -alanine, but not to the other CycA substrates, such as l-alanine, d-alanine, and glycine. This prevented a direct application for the improvement of pantothenic acid-producing strains by an enhanced precursor supply.     

The paleobiological record of photosynthesis

Fossil evidence of photosynthesis, documented in Precambrian sediments by microbially laminated stromatolites, cyanobacterial microscopic fossils, and carbon isotopic data consistent with the presence of Rubisco-mediated CO₂-fixation, extends from the present to ~3,500 million years ago. Such data, however, do not resolve time of origin of O₂-producing photoautotrophy from its anoxygenic, bacterial, evolutionary precursor. Though it is well established that Earth’s ecosystem has been based on autotrophy since its very early stages, the time of origin of oxygenic photosynthesis, more than 2,450 million years ago, has yet to be established.     

Laboratory Simulations of Cyanobacterial Mats of the Alkaline Geochemical Barrier

The goal of this work was to illustrate a possible interaction between the soda continent and the ocean. A laboratory simulation was undertaken of the development of alkaliphilic mat with calcium carbonate and calcium phosphate interlayers in the zone where ocean waters, containing calcium and manganese, come into contact with carbonate- and phosphate-rich alkaline waters. The macrostructure of the layered cyanobacterial mat turned out to be little dependent on the chemical conditions causing sediment formation. The chemical composition of freshly formed mineral interlayers of the mat was found to vary with the medium composition. The mineralogical composition of the sediment is determined by diagenesis conditions in its depth, which can cause mineral phase conversions.     

Polypeptides of photosystem II and their role in oxygen evolution

The linear, four-step oxidation of water to molecular oxygen by photosystem II requires cooperation between redox reactions driven by light and a set of redox reactions involving the S-states within the oxygen-evolving complex. The oxygenevolving complex is a highly ordered structure in which a number of polypeptides interact with one another to provide the appropriate environment for productive binding of cofactors such as manganese, chloride and calcium, as well as for productive electron transfer within the photoact. A number of recent advances in the knowledge of the polypeptide structure of photosystem II has revealed a correlation between primary photochemical events and a core complex of five hydrophobic polypeptides which provide binding sites for chlorophyll a, pheophytin a, the reaction center chlorophyll (P680), and its immediate donor, denoted Z. Although the core complex of photosystem II is photochemically active, it does not possess the capacity to evolve oxygen. A second set of polypeptides, which are water-soluble, have been discovered to be associated with photosystem II; these polypeptides are now proposed to be the structural elements of a special domain which promotes the activities of the loosely-bound cofactors (manganese, chloride, calcium) that participate in oxygen evolution activity. Two of these proteins (whose molecular weights are 23 and 17 kDa) can be released from photosystem II without concurrent loss of functional manganese; studies on these proteins and on the membranes from which they have been removed indicate that the 23 and 17 kDa species from part of the structure which promotes retention of chloride and calcium within the oxygen-evolving complex. A third water-soluble polypeptide of molecular weight 33 kDa is held to the photosystem II core complex by a series of forces which in some circumstances may include ligation to manganese. The 33 kDa protein has been studied in some detail and appears to promote the formation of the environment which is required for optimal participation by manganese in the oxygen evolving reaction. This minireview describes the polypeptides of photosystem II, places an emphasis on the current state of knowledge concerning these species, and discusses current areas of uncertainty concerning these important polypeptides.Abbreviations A 23187 ionophore that exchanges divalent cations with H⁺ - Chl chlorophyll - cyt cytochrome - DCPIP dichlorophenolindophenol - DPC diphenylcarbazide - EGTA ethyleneglycoltetraacetic acid - P680 the chlorophyll a reaction center of photosystem II - pheo pheophytin - PQ plastoquinone - PS photosystem - Q_A and Q_B primary and secondary plastoquinone electron acceptors of photosystem II - Sn (n=0, 1, 2, 3, 4) charge accumulating state of the oxygen evolving system - Signals II_vf, II_f and II_s epr-detectable free radicals associated with the oxidizing side of photosystem II - Z primary electron donor to the photosystem II reaction center The survey of literature for this review ended in September, 1984.     

Magnesium-Dependent Ecto-ATP Diphosphohydrolase Activity in <Emphasis Type="Italic">Herpetomonas muscarum muscarum</Emphasis>

In the present work we characterized the ecto-ATP diphosphohydrolase activity of the trypanosomatid parasite Herpetomonas muscarum muscarum. This parasite hydrolyzed ATP at a rate of 15.52 nmol Pi/mg protein/min and this activity reached a maximum at pH 7.5. Classical inhibitors of acid phosphatases, such as sodium orthovanadate (N_aVO₃), sodium fluoride (NaF), and ammonium molybdate presented no effect on this activity. MgCl₂, ZnCl₂, and MnCl₂ stimulated the ATP hydrolysis by H. m. muscarum. The ecto-ATPase activity was insensitive to oligomycin and sodium azide, two inhibitors of mitochondrial Mg-ATPase, bafilomycin A₁, a V-ATPase inhibitor, ouabain, a Na⁺+K⁺-ATPase inhibitor and to levamizole, an inhibitor of alkaline phosphatase. An extracellular impermeant inhibitor 4,4-diisothiocyanostylbene 2,2-disulfonic acid (DIDS) and a inhibitor of some ecto-ATPases, suramin, which is also a competitive antagonist of P₂-purinergic receptors, promoted a great inhibition on the ATP hydrolysis. This enzyme is able to hydrolysis ATP, ADP, UTP, and UDP, but not GTP, GDP, CTP, or CDP. ADP inhibited the enzymatic activity in a concentration dependent manner, reaching 70% inhibition. Received: 17 September 2002 / Accepted: 19 November 2002     

Biotic and Abiotic Imprints on Mg-Rich Stromatolites: Lessons from Lake Salda,SW Turkey

Abstract

Modern hydrated Mg rich stromatolites are actively growing along the shallow shorelines of Lake Salda (SW Turkey). An integrated approach involving isotopic, mineralogical, microscopic, and organic/geochemical techniques along with culture-independent molecular methods were applied to various lake samples to assess the role of microbial processes on stromatolite formation. This study further explores the biosignature preservation potential of fossil stromatolites by comparing with textures, lipid profiles and isotopic composition of the modern stromatolites. Similar lipid profile and δ¹³C isotope values in active and fossil stromatolites argue that CO₂ cycling delicately balanced between photosynthetic and heterotrophic (aerobic) activity as in the active ones may have regulated stromatolite formation in the lake. A decrease in the exopolymeric substances (EPS) profile of the mat and concurrent hydromagnesite precipitation imply a critical role for EPS in the formation of stromatolite. Consistently, a discrete, discontinuous lamination and clotted micropeloidal textures with cyanobacterial remnants in the fossil stromatolites likely refer to partial degradation of EPS, creating local nucleation sites and allowing precipitation of hydrated Mg minerals and provide a link to the active microbial mat in the modern stromatolites. Our results for the first time provide strong evidence for close coupling of cyanobacterial photosynthesis and aerobic heterotrophic respiration on hydromagnesite textures involved in the stromatolite formation of Lake Salda. The creation of photosynthesis induced high-pH conditions combined with a change in the amount and properties of the EPS and the repetition of these processes over time seems to be a possible pathway for stromatolite growth in the lake. Understanding these microbial symbioses and their mineralized records may provide new insights on the formation mechanism of Mg-rich carbonates not only for terrestrial geological records but also for planetary bodies like Mars, where hydrated Mg-carbonate deposits have been identified in possible paleolake deposits at Jezero crater, the landing site of the NASA Mars 2020 rover.     

Fossilization of the cells of natronophilic endoevaporite cyanobacterium ‘<Emphasis Type="Italic">Euhalothece natronophila</Emphasis>’ in a modelling system

Laboratory simulation of fossilization of cyanobacterial cells in the high-carbonate medium in the presence of calcium was carried out for the haloalkaliphilic natronophilic cyanobacterium ‘Euhalothece natronophila’ Z-M001. This organism was isolated from the Magadi soda lake, where the bioherms consisting of mineralized coccoid cyanobacteria were found in the Quaternary sediments. The structural and chemical heterogeneity of the minerals produced during this process was established, with calcium carbonate and trona being the main products. The differences in the process of cyanobacterial cell carbonatization in soda lakes and marine or freshwater systems were determined. Initial precipitation of calcium carbonate was shown to occur due to a chemical reaction not involving cyanobacteria. At the subsequent stages, amorphous CaCO₃ is sorbed and crystallized on the surface of some of the cells within a cyanobacterial population, resulting in formation of a shell-like mineral layer. The cells embedded in trona in the same system were shown to undergo deformation and destruction. In both cases the mineralized cells were shown to lose their photosynthetic activity.     

Action of 1,25-dihydroxyvitamin d3 on calcium metabolism in the crustacean Orchestia cavimana during the molt cycle

• 1.1. The possible involvement of vitamin D₃ in the calcium metabolism of the terrestrial crustacean Orchestia during the molt cycle was further investigated by measuring the effects of administration of exogenous 1,25 (OH)₂D₃ on three parameters of the calcium balance in two different compartments of the body.
• 2.2. At the hemolymph level, a strong hypocalcémie effect was observed in intermolt and early premolt. Within the posterior caeca of the midgut, stimulation of calcium storage and calcium release were noticed during a short period surrounding the time of exuviation, with concomitant variations of the epithelial carbonic anhydrase activity.
• 3.3. These results, together with other data, are discussed to determine the possible functions of vitamin D₃, or related molecules, in the calcium turnover within the different compartments of the body, according to the successive stages of the molt cycle.

     

Carbohydrate Structural Units in Glycosphingolipids as Receptors for Gal and GalNAc Reactive Lectins

Glycosphingolipids (GSLs) contain many carbohydrate epitopes or crypto-glycotopes for Gal and GalNAc reactive lectins. Many of them are in the nervous system and function as important receptors in various life processes. During the past two decades, 11 mammalian structural units have been used to express the binding domain of applied lectins. They are: F, GalNAc1 3GalNAc; A, GalNAc1 3Gal; T, Gal1 3GalNAc; I, Gal1 3GlcNAc; II, Gal1 4GlcNAc; B, Gal1 3Gal; E, Gal1 4Gal; L, Gal1 4Glc; P, GalNAc1 3Gal; S, GalNAc1 4Gal, and Tn, GalNAc1 Ser(Thr). Although 10 of them occur in GSLs, only 3 (L , S , and T ) are found in human brain, and 2 (L and II ) are present in the inner structures of human blood group active GSLs. In the families of gangliosides, L and II represent 55% of the total structural units, while the other three units (T , P , and S ) constitute the rest. To facilitate the selection of lectins that could serve as structural probes, the carbohydrate binding specificities of Gal/GalNAc reactive lectins have been classified according to their highest affinity for the structural units and their binding properties expressed by decreasing order of reactivity. Hence, the binding relation between GSLs and Gal/GalNAc specific lectins can be established.     

Age and growth,reproduction and diet of a sublittoral population of the rock goby Gobius paganellus (Teleostei,Gobiidae)

Colonisation of extremely acidic waters (pH 3) by aquatic angiosperms occurs widely, but is poorly documented. Unlike acid rain affected and other naturally acidic aquatic ecosystems, waters with pH 3 usually have a high conductivity, with high concentrations of SO₄ ^2- and often high concentrations of Fe³⁺, other heavy metal ions and Al³⁺. Where Fe³⁺ concentration is high, as in many mine waters, it provides a strong buffering system. In such waters, the biogeochemical Fe cycle exerts over water chemistry and the availability of nutrients and carbon for organisms. Biological activity is limited by low concentrations of phosphorus and inorganic carbon (DIC), which in this pH range is essentially all in the form of dissolved CO₂. A number of angiosperms grow in such waters including Phragmites australis, Typha spp. and Juncus bulbosus, though the last is the only one reported to grow totally submerged in waters with pH 3 . J. bulbosus occurs in many lignite mining lakes in Lusatia (north eastern Germany) with pH 3. The characteristics and possible survival strategies for this and other species are discussed.     

UDP-galactose:lactosylceramide α-galactosyltransferase activity in human placenta

The activity of UDP-Gal: LacCer galactosyltransferase in human placenta was studied by using crude homogenate and Triton CF-54 extract as the source of enzyme. Transfer of galactose to lactosylceramide was optimal in the presence of 0.1% Triton CF-54 and Mn²⁺ at pH 6.3, and the reaction product was susceptible to -galactosidase.Abbreviations LacCer lactosylceramide (Gal1-4Glc1-1Cer) - Gb₃ globotriaosylceramide (Gal1-4Gal1-4Glc1-1Cer) - Gb₄ globoside (GalNAc1-3Gal1-4Gal1-4Glc1-1Cer) - TLC thin-layer chromatography - GC/MS gas chromatography/mass spectrometry - NMR nuclear magnetic resonance - EDTA ethylenediamine tetraacetic acid - CHAPS 3-[(3-cholamidopropyl)dimethylammonio]-1-propanesulfonate