The bioenergetic role of dioxygen and the terminal oxidase(s) in cyanobacteria

Owing to the release of 13 largely or totally sequenced cyanobacterial genomes (see and ), it is now possible to critically assess and compare the most neglected aspect of cyanobacterial physiology, i.e., cyanobacterial respiration, also on the grounds of pure molecular biology (gene sequences). While there is little doubt that cyanobacteria (blue-green algae) do form the largest, most diversified and in both evolutionary and ecological respects most significant group of (micro)organisms on our earth, and that what renders our blue planet earth to what it is, viz. the O(2)-containing atmosphere, dates back to the oxygenic photosynthetic activity of primordial cyanobacteria about 3.2x10(9) years ago, there is still an amazing lack of knowledge on the second half of bioenergetic oxygen metabolism in cyanobacteria, on (aerobic) respiration. Thus, the purpose of this review is threefold: (1) to point out the unprecedented role of the cyanobacteria for maintaining the delicate steady state of our terrestrial biosphere and atmosphere through a major contribution to the poising of oxygenic photosynthesis against aerobic respiration ("the global biological oxygen cycle"); (2) to briefly highlight the membrane-bound electron-transport assemblies of respiration and photosynthesis in the unique two-membrane system of cyanobacteria (comprising cytoplasmic membrane and intracytoplasmic or thylakoid membranes, without obvious anastomoses between them); and (3) to critically compare the (deduced) amino acid sequences of the multitude of hypothetical terminal oxidases in the nine fully sequenced cyanobacterial species plus four additional species where at least the terminal oxidases were sequenced. These will then be compared with sequences of other proton-pumping haem-copper oxidases, with special emphasis on possible mechanisms of electron and proton transfer.     

The role of the bidirectional hydrogenase in cyanobacteria

Cyanobacteria have tremendous potential to produce clean, renewable fuel in the form of hydrogen gas derived from solar energy and water. Of the two cyanobacterial enzymes capable of evolving hydrogen gas (nitrogenase and the bidirectional hydrogenase), the hox-encoded bidirectional Ni-Fe hydrogenase has a high theoretical potential. The physiological role of this hydrogenase is a highly debated topic and is poorly understood relative to that of the nitrogenase. Here the structure, assembly, and expression of this enzyme, as well as its probable roles in metabolism, are discussed and analyzed to gain perspective on its physiological role. It is concluded that the bidirectional hydrogenase in cyanobacteria primarily functions as a redox regulator for maintaining a proper oxidation/reduction state in the cell. Recommendations for future research to test this hypothesis are discussed.     

Estuarine diversity of tintinnids (planktonic ciliates)

In Chesapeake Bay, a large eutrophic and partially stratifiedestuary, we investigated diversity among tintinnids (Ciliophora,suborder Tintinninia) in September 1999. In contrast with thetypical estuarine pattern, tintinnid diversity was high andincreased with decreasing salinity from the mouth of the bayto the mid-bay region. Peak species numbers and diversity values[20–25 species, H' (ln) = 2.4–2.5] were found instations in the mesohaline (14–17     

Persistence and genetic diversity among strains of phycoerythrinrich cyanobacteria from the picoplankton of Lake Constance

Twelve strains of phycoerythrin (PE)-rich unicellular cyanobacteriaof the Synechococcus type were isolated from the pelagial ofLake Constance in different phases during the growth periodof the year 1994. By analysing the restriction fragment lengthpolymorphism (RFLP) of the DNA three new genotypes were distinguished.The persistence of one strain during the course of the yearwas demonstrated. The data set was compared with the RFLP observedin PE-rich Synechococcus strains isolated in former years fromthe same sampling site. The plurality in the picoplanktic isolatesand the possibility of cultivation of strains is discussed withrespect to the oligotrophication of Lake Constance.     

The role of plastidic cytochrome c in algal electron transport and photophosphorylation.

By an improved isolation procedure chloroplasts could be obtained from the alga Bumilleriopsis filiformis (Xanthophyceae) which exhibited high electron transport rates tightly coupled to ATP formation. Uncouplers both stimulate electron transport and inhibit photophosphorylation. These chloroplasts retain almost all soluble cytochrome c-553 besides a membrane-bound cytochrome c-554.5 (=f-554.5). Sonification or iron deficiency removed the soluble cytochrome only with a concurrent decrease of electron transport from water to methyl viologen or to NADP and decreased non-cyclic and cyclic photophosphorylation. However, photosynthetic control and the P/2e ratios remain unaltered. In Bumilleriopsis, which apparently has no plastocyanin, the soluble cytochrome c-553 seemingly links electron transport between the bound cytochrome c and P-700.     

Preliminary studies of cyanobacteria, picoplankton, and virioplankton in the Salton Sea with special attention to phylogenetic diversity among eight strains of filamentous cyanobacteria

Cyanobacterial diversity in the Salton Sea, a high-salinity, eutrophic lake in Southern California, was investigated using a combination of molecular and morphological approaches. Representatives of a total of 10 described genera (Oscillatoria, Spirulina, Arthrospira, Geitlerinema, Lyngbya, Leptolyngbya, Calothrix, Rivularia, Synechococcus, Synechocystis) were identified in the samples; additionally, the morphology of two cultured strains do not conform to any genus recognized at present by the bacteriological system. Genetic analysis, based on partial 16S rRNA sequences suggested considerable cryptic genetic variability among filamentous strains of similar or identical morphology and showed members of the form-genus Geitlerinema to be distributed among three major phylogenetic clades of cyanobacteria. Cyanobacterial mats, previously described from the Sea were, in fact, composed of both filamentous cyanobacteria and a roughly equivalent biomass of the sulfur-oxidizing bacterium Beggiatoa, indicating their formation in sulfide rich regions of the lake. Flow cytometric analysis of the water samples showed three striking differences between samples from the Salton Sea and representative marine waters: (1) phycoerythrin-containing unicells, while abundant, were much less abundant in the Salton Sea than they were in typical continental shelf waters, (2) Prochlorococcus appears to be completely absent, and (3) small (3–5 m) eukaryotic algae were more abundant in the Salton Sea than in typical neritic waters by one-to-two orders-of-magnitude. Based on flow cytometric analysis, heterotrophic bacteria were more than an order of magnitude more abundant in the Salton Sea than in seawater collected from continental shelf environments. Virus particles were more abundant in the Salton Sea than in typical neritic waters, but did not show increases proportionate with the increase in bacteria, picocyanobacteria, or eukaryotic algae.     

The role of cyanobacteria in the assemblage of the Lakhanda Microbiota

This paper discusses prokaryotic and eukaryotic components of the Lakhanda Microbiota, which provides data on communities of organisms at the Meso-Neoproterozoic boundary (1010–1025 Ma). Communities of ancient organisms are fixed in mats and biofilms of various architecture. The significance of cyanobacteria in the development of highly differentiated communities of ancient organisms, found in the shallow-water terrigenous rocks of the Lakhanda Group at the Maya River (Uchuro-Maiskii District, southeastern Siberia), is discussed. The paper discusses the significance of clayey minerals for benthic organic life in the Late Precambrian and their effect on the preservation and taphonomy of fossil remains.     

The role of plastidic cytochrome c in algal electron transport and photophosphorylation

By an improved isolation procedure chloroplasts could be obtained from the alga Bumilleriopsis filiformis (Xanthophyceae) which exhibited high electron transport rates tightly coupled to ATP formation. Uncouplers both stimulate electron transport and inhibit photophosphorylation. These chloroplasts retain almost all soluble cytochrome c-553 besides a membrane-bound cytochrome c-554.5 (=f-554.5). Sonification or iron deficiency removed the soluble cytochrome only with a concurrent decrease of electron transport from water to methyl viologen or to NADP and decreased non-cyclic and cyclic photophosphorylation. However, photosynthetic control and the $\text{P}\text{2e}$ ratios remain unaltered.In Bumilleriopsis, which apparently has no plastocyanin, the soluble cytochrome c-553 seemingly links electron transport between the bound cytochrome c and P-700.     

Distribution and ecology of tintinnids in the plankton of Lebanese coastal waters (eastern Mediterranean)

The tintinnids of Jounieh Bay in Lebanon were identified andtheir numerical abundance determined in horizontal and verticalnet samples from February 1979 to December 1980. Large fluctuationsin abundance and composition of the tintinnid fauna occurredduring the period of investigation. Two distinct peaks werenoted, a major one in May–June and a minor one in November–December.The first followed the main phytoplankton peak, suggesting apositive nutritional relationship between the two populations.The most abundant species were Tintinopsis beroidea and Eutintinnuslusus-undae. A few other species showed great abundance duringshorter periods. In all, 121 species were recorded during thisperiod, and of these 12 belong to the genus Tintinnopsis and11 to Eutintinnus.     

Phosphorus strategy in bloom-forming cyanobacteria (Dolichospermum and Microcystis) and its role in their succession

Dolichospermum (formerly Anabaena) and Microcystis cause harmful cyanobacterial blooms in freshwater ecosystems worldwide. Input reduction of both nitrogen (N) and phosphorus (P) are commonly recognized as basic ways of controlling blooms, but little is known about the roles of nutrients and their using strategy among cyanobacteria in triggering the succession of diazotrophic to non-diazotrophic cyanobacteria. In this study, we investigated in situ responses of cyanobactria to ambient P status during the transition from Dolichospermum flos-aquae to Microcystis spp. in Lake Taihu and Lake Chaohu. While dominant in phytoplankton community, D. flos-aquae experienced P deficiency as evidenced by qualitative detection of extracellular phosphatase via enzyme labeled fluorescence (ELF). The percentage of ELF-labelled D. flos-aquae cells was 33% when it dominated the phytoplankton community, and was 78% when it co-dominated with Microcystis spp., indicating an increase in P deficiency. Meanwhile, no ELF-labelled Microcystis cells were observed while polyphosphate body (PPB) were present, suggesting that Microcystis spp. were not P deficient. Additionally, the percentages of Microcystis cells containing PPB showed an inverted “U-shaped” relationship with concentrations on soluble reactive phosphorus (SRP). To validate the field observation, a laboratory study of the monocultures of the dominant cyanobacteria was conducted. Extracellular alkaline phosphatase activity (APA) and PPB accumulation were regulated by P availability in monocultures of D. flos-aquae. Interestingly, no cell bound extracellular phosphatase was found on Microcystis aeruginasa even in the culture without P supply. Consistently, the expressions of phosphatase encoding gene phoX showed no differences among the treatments. The way in which PPB accumulation occurred in Microcystis spp. in response to P availability in the cultures was similar to that observed in the field, demonstrating a strategy of energy conservation over P accumulation. The competitive advantage of Microcystis spp. was displayed at low P concentrations: where it could rapidly uptake and store inorganic P, which also increased the P deficiency of the coexisting phytoplankton species. Responses of P-transport gene pstS confirmed this hypothesis. The physiological and molecular mechanisms mentioned above enable Microcystis to survive and proliferate in environment with low available P supply more efficiently. In conclusion, different cyanobacterial species have distinct ways of responding to P availability, suggesting that the control of cyanobacterial blooms by targeted nutrient reduction is largely dependent upon the dominant species. P reduction is more effective in controlling diazotrophic cyanobacteria than non-diazotrophic cyanobacteria.     

Abundance and biogeography of tintinnids (Ciliophora) and associated microzooplankton in the Southwestern Atlantic Ocean

Absolute abundances of foraminifers, polycystine and phaeodarian radiolarians, tintinnids, pteropods and early crustacean larvae and moults were assessed in a collection of 57 vertically stratified (0-100 m) net microplankton samples from 22 stations located between 34 and 58S (along 51-56°W), covered on 8-16 November 1994. Tintinnids were identified to species and measured in order to estimate their biomass from biovolume to carbon conversions. The distribution of the microzooplanktonic groups assessed was irregular and patchy, both geographically and vertically, and their abundances were characteristic of oceanic low to medium productivity environments. Tintinnid biomass was also generally low (0.05-0.40 g Cl^-1). With the exception of the tintinnids, associations between microzooplanktonic numbers and chlorophyll a were generally loose. Eighty-eight tintinnid taxa were recorded, yet only five accounted for 53% of the specimens identified. Multivariate (cluster) analysis of tintinnid specific distribution patterns clearly showed several distinct zones. From north to south, these are: Transition Zone (TZ), with three subzones, TZ north (34°S-38°S), TZ central (39°44S-44°S) and TZ south (46°S); Subantarctic Zone (SZ; 48-55°S); Polar Front Zone (55°30S); Antarctic Zone (AZ; 58-59°36S). Each of these was characterized by distinct tintinnid assemblages, abundance and biomass. With few exceptions, tintinnid cells were fairly evenly distributed throughout the upper 50 m. Taxonomic composition usually changed little with depth. Mean population depths were calculated for a subset of 35 tintinnids; 29 of these dwell preferably above 40 m. The spatial distribution of tintinnid species richness showed a more or less gradual decrease from north to south. Specific diversity and equitability generally increased with depth, and were higher in antarctic waters than the southern transitional and subantarctic ones; this trend is tentatively attributed to higher water column vertical stability south of the Polar Front.      

The role of cyanobacteria in crystallization of magnesium calcites

Laboratory experiments showed the effect of the cyanobacterium Microcoleus chthonoplastes on the formation of magnesium calcites, using model solutions (2.14M MgCl₂-0.05M CaCl₂-0.6M NaCl-0.18M NaHCO₃). The conditions of existence of cyanobacteria in such solutions in light or darkness significantly alter the structure of the sediment and the shape and size of the carbonate crystals. Cyanobacteria slow down crystallization due to the formation of exometabolites with a chelating effect, which leads to the precipitation of high-magnesium calcites. In the photosynthetic environment the presence of huntite (CaMg₃(CO₃)₄), possible forerunner of dolomite, is prominent.     

The species-rich assemblages of tintinnids (marine planktonic protists) are structured by mouth size

Many microbial taxa in the marine plankton appear super-saturated in species richness. Here, we provide a partial explanation by analyzing how species are organized, species packing, in terms of both taxonomy and morphology. We focused on a well-studied group, tintinnid ciliates of the microzooplankton, in which feeding ecology is closely linked to morphology. Populations in three distinct systems were examined: an Eastern Mediterranean Gyre, a Western Mediterranean Gyre and the California Current. We found that species abundance distributions exhibited the long-tailed, log distributions typical of most natural assemblages of microbial and other organisms. In contrast, grouping in oral size-classes, which corresponds with prey-size exploited, revealed a geometric distribution consistent with a dominant role of a single resource in structuring an assemblage. The number of species found in a particular oral size-class increases with the numerical importance of the size-class in the overall population. We suggest that high species diversity reflects the fact that accompanying each dominant species are many ecologically similar species, presumably able to replace the dominant species, at least with regard to the size of prey exploited. Such redundancy suggests that species diversity greatly exceeds ecological diversity in the plankton.     

The respiratory chain of blue-green algae (cyanobacteria)

Electron transport components on the way from reduced substrates to the terminal respiratory oxidase(s) are discussed in relation to analogous and/or homologous enzymes and electron carriers in the generally much better known bacteria, mitochondria and chloroplasts. The kinetic behaviour of the components, their localization within the cell and their evolutionary position are given special attention. Pertinent results from molecular genetics are also mentioned. The unprecedented role of cyanobacteria for our biosphere and our whole planet earth appears to deserve a more extended introductory chapter.     

The role of heterocytes in the physiology and ecology of bloom-forming harmful cyanobacteria

Dolichospermum flos-aquae and Cylindrospermopsis raciborskii are two cyanobacteria species which cause harmful blooms around the world. Both these species share the capacity to fix atmospheric nitrogen in heterocytes (cell where fixation occurs). While Dolichospermum can express heterocytes at rather regular intervals across the filament, Cylindrospermopsis can only express heterocytes at the end of the filament. The aim of this study was to experimentally assess the role of heterocyte position in the eco-physiological responses of these bloom forming cyanobacteria. Replicated monocultures of each species were grown at different eutrophication scenarios (limiting and sufficient nitrogen and phosphorus concentrations, in factorial design). Dolichospermum reached high biomass regardless of the nitrogen (and phosphorus) provided, suggesting that this species could bloom in situations with and without nitrogen limitation. In contrast, Cylindrospermopsis reached high biomass only when nitrogen supply was high; its biomass was 15-20 times lower when relying on nitrogen fixation. Hence, despite its ability to fix nitrogen, blooms of Cylindrospermopsis would be expected only under high total nitrogen availability. In Dolichospermum heterocytes occurred only in the scenarios without supplied nitrogen while in Cylindrospermopsis heterocytes occurred regardless of nitrogen availability. Yet, in both species nitrogen fixation occurred (heterocytes were functional) only when nitrogen was limiting, and nitrogen fixation increased significantly at higher phosphorus concentration. Finally, in the absence of supplied nitrogen, filament length in Dolichospermum was the longest, while filaments in Cylindrospermopsis were the shortest (up to 13 times shorter than at nitrogen sufficiency). Therefore, heterocyte expression in Dolichospermum, and filament length in Cylindrospermopsis seem good proxies of nitrogen fixation. The eco-physiological responses recorded here help understand the distribution of these species along nutrient gradients in nature.     

Dynamics of prokaryotic picoplankton community in the central and southern Adriatic Sea (Croatia)

This paper addresses the dynamics of the prokaryotic picoplankton community in the coastal and open sea areas of the central Adriatic and in the coastal area of the southern Adriatic. This involved the study, from January to December 2005, of bacteria (total number of non-pigmented bacteria; high nucleic acid content (HNA) bacteria; low nucleic acid content (LNA) bacteria), cyanobacteria (Synechococcus and Prochlorococcus) and heterotrophic nanoflagellates. During the warmer seasons, in the mainly oligotrophic area under investigation into the Adriatic Sea, bacterial densities and bacterial production have shown an increase in values and domination of the LNA group of the bacterial population. In contrast, in those areas influenced by karstic rivers, the domination of HNA bacteria in total abundance of non-pigmented bacteria and high values of bacterial production was estimated throughout the investigated period. Our results show the importance of both HNA and LNA bacterial groups in the total bacterial activity throughout the investigated area. The biomass of bacteria was mostly predominant in the prokaryotic community, while within the autotrophic community Synechococcus biomass mostly predominated. During the warmer seasons, an increase in autotrophic biomass was observed in relation to non-pigmented biomass. The importance of predation in controlling bacteria by heterotrophic nanoflagellates was pronounced during the warmer period and in the coastal areas.     

The plastidic pentose phosphate translocator represents a link between the cytosolic and the plastidic pentose phosphate pathways in plants

Plastids are the site of the reductive and the oxidative pentose phosphate pathways, which both generate pentose phosphates as intermediates. A plastidic transporter from Arabidopsis has been identified that is able to transport, in exchange with inorganic phosphate or triose phosphates, xylulose 5-phosphate (Xul-5-P) and, to a lesser extent, also ribulose 5-phosphate, but does not accept ribose 5-phosphate or hexose phosphates as substrates. Under physiological conditions, Xul-5-P would be the preferred substrate. Therefore, the translocator was named Xul-5-P/phosphate translocator (XPT). The XPT shares only approximately 35% to 40% sequence identity with members of both the triose phosphate translocator and the phosphoenolpyruvate/phosphate translocator classes, but a higher identity of approximately 50% to glucose 6-phosphate/phosphate translocators. Therefore, it represents a fourth group of plastidic phosphate translocators. Database analysis revealed that plant cells contain, in addition to enzymes of the oxidative branch of the oxidative pentose phosphate pathway, ribose 5-phosphate isomerase and ribulose 5-phosphate epimerase in both the cytosol and the plastids, whereas the transketolase and transaldolase converting the produced pentose phosphates to triose phosphates and hexose phosphates are probably solely confined to plastids. It is assumed that the XPT function is to provide the plastidic pentose phosphate pathways with cytosolic carbon skeletons in the form of Xul-5-P, especially under conditions of a high demand for intermediates of the cycles.     

Toxic cyanobacteria Nodularia spumigena in the diet of Baltic mysids: Evidence from molecular diet analysis

A PCR-based method was used to detect toxic cyanobacteria Nodularia spumigena in the diet of Baltic mysids, Mysis mixta and Mysis relicta. The decay in detectability of Nodularia DNA in mysid stomachs and feces following the cyanobacterium consumption was examined in laboratory with special references to (1) marker size (780 bp vs. 200 bp), (2) mysid developmental stage (juveniles vs. subadults), and (3) feeding regime after consuming the cyanobacteria (continuous vs. interrupted feeding). The Nodularia DNA could be reliably detected in mysid stomachs and feces by PCR technique. In the mysid with interrupted feeding, the calculated half-lives of N. spumigena DNA in the mysid stomachs were 1.2 and 6.1 h for 780 and 200 bp fragments, respectively. Continuous feeding, however, facilitated decay in the detectability, most likely due to increased gut evacuation rate. In stomachs of the field-collected mysids, the Nodularia DNA was detected with high frequencies, 60% in M. mixta and 51% in M. relicta. Moreover, it was higher in immature mysids than in adults and correlated with stomach fullness in age-specific manner: in juveniles and subadults, stomachs containing Nodularia were significantly fuller, while in adults, the presence of the cyanobacteria was associated with empty stomachs. This suggests greater habitat overlap for juvenile mysids and N. spumigena and thus higher encounter and consumption rates. These findings contribute to a growing body of evidence that cyanobacteria in the Baltic Sea are important food for grazers.     

莱州湾大型砂壳纤毛虫群落季节变化

为揭示莱州湾砂壳纤毛虫群落季节变化规律, 在莱州湾设置8个站位, 于2011年5-11月及2012年3-4月进行了9个航次的调查, 用浅海III型浮游生物网由底至表垂直拖网采集砂壳纤毛虫。结果表明, 莱州湾3-11月砂壳纤毛虫物种丰富度的变化范围为5-19, 周年变化呈现一峰两谷的趋势。丰度的范围为0-318 ind./L, 丰度较大(> 50 ind./L)的种类有运动类铃虫(Codonellopsis mobilis)和清兰拟铃虫(Tintinnopsis chinglanensis)。各月平均丰度随时间的变化趋势为双峰型, 最大值出现在7月(63 ind./L), 次峰值出现在5月(48 ind./L), 最小值出现在3月(2 ind./L)。黏着壳种类在3-11月均有出现, 透明壳种类仅在温度较高(> 15°C)的6-9月出现。各月的优势种数目为1种(5月)到8种(8月), 其中运动类铃虫在所有月中都是优势种, 对砂壳纤毛虫丰度周年的变化规律产生较大影响。使用各月所有种类的平均丰度对各月砂壳纤毛虫群落进行聚类分析, 得到两个群落(相似度30%): 群落I(7-9月)和群落II(3-6月、10-11月), 说明砂壳纤毛虫群落发生了明显的季节变化。砂壳纤毛虫的物种丰富度、丰度与环境因子(温度、盐度)均没有明显的相关性。     

The role of phytoplankton in the diet of the bladderwort Utricularia australis R.Br. (Lentibulariaceae)

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