The morphological adaptation of <Emphasis Type="Italic">Lithocodium aggregatum</Emphasis> Elliott (calcareous green alga) to cryptic microhabitats (Lower Aptian,Spain): an example of phenotypic plasticity

Lithocodium aggregatum Elliott is interpreted as a heterotrichale chlorophycean alga with a prostrate and erect system within a well-calcified tissue. Within Lower Aptian coral rubble of the western Maestrat Basin, Spain, it forms thick masses of juxtaposed crusts around the bioclasts. In achieving a rapid and complete encrustation of the available bioclasts, Lithocodium applied several strategies, e.g., filling voids of structured surfaces with a special fabric or forming erect extensions to bridge adjacent substrates. In these deposits, different types of poorly if ever illuminated cryptic microhabitats can be distinguished: (1) existing crypts such as empty shells or structural voids within bioclasts (2) crypts resulting from the complete encrustation of adjacent bioclasts by Lithocodium itself and (3) syndepositionally created crypts, e.g., boreholes produced by lithophagid bivalves. In these cases, Lithocodium developed a poorly calcified structure of large cells with thin microcrystalline walls indicating a high degree of variability (phenotypic plasticity). This cryptic growth stage is interpreted as an adaption to the poorly illuminated crypts (photoadaption) in order to maximize light capture for photosynthesis. The Lower Cretaceous Lithocodium is therefore not per se a cryptoendolithic microorganism, but may show adaptation to develop and survive also in these already existing or newly created niches.     

Ordovician Calcified Algae and Cyanobacteria, Northern Tarim Basin Subsurface, China

Boreholes between Kuqa and Korla, in the northern Tarim Basin, Xinjiang, penetrated Ordovician marine limestones at depths of 5–6.2km. From three boreholes 54 out of a total of 170 limestone samples contain calcified algae, cyanobacteria and associated Microproblematica. Calcified cyanobacteria (GirvanellaBotomaella, ?Subtifloria) account for 37 per cent of occurrences; green algae (Dasyporelleae and Vermiporella) 28 per cent; Microproblematica (NuiaBevocastriaRothpletzellaHalysis) 20 per cent; and ‘solenoporaceans’ 15 per cent GirvanellaNuia are common in the Early Ordovician deposits, and ‘solenoporaceans’ are abundant in the Mid‐Ordovician. Dasyporelleae and Vermiporella are most abundant in Mid–Late Ordovician samples. Calcified cyanobacteria are common throughout the limestone succession, but particularly in the Mid Ordovician part. MoniliporellaContextaPlexaTexturata, and Villosoporella, hitherto placed in the supposed red algal family Moniliporellaceae Gnilovskaya, are here regarded as dasycladalean green algae. Despite some omissions, this Tarim flora broadly resembles others from Kazakhstan, Baltica and North America, indicating the generally cosmopolitan nature of Ordovician calcified algae and cyanobacteria.     

Formation and diagenesis of modern marine calcified cyanobacteria

Calcified cyanobacterial microfossils are common in carbonate environments through most of the Phanerozoic, but are absent from the marine rock record over the past 65 Myr. There has been long-standing debate on the factors controlling the formation and temporal distribution of these fossils, fostered by the lack of a suitable modern analog. We describe calcified cyanobacteria filaments in a modern marine reef setting at Highborne Cay, Bahamas. Our observations and stable isotope data suggest that initial calcification occurs in living cyanobacteria and is photosynthetically induced. A single variety of cyanobacteria, Dichothrix sp., produces calcified filaments. Adjacent cyanobacterial mats form well-laminated stromatolites, rather than calcified filaments, indicating there can be a strong taxonomic control over the mechanism of microbial calcification. Petrographic analyses indicate that the calcified filaments are degraded during early diagenesis and are not present in well-lithified microbialites. The early diagenetic destruction of calcified filaments at Highborne Cay indicates that the absence of calcified cyanobacteria from periods of the Phanerozoic is likely to be caused by low preservation potential as well as inhibited formation.     

Microbial crusts of the late jurassic: Composition,palaeoecological significance and importance in reef construction

Summary Upper Jurassic reefs contain variable amounts of calcareous microbial crusts. In examples from Portugal, Spain and southern Germany they occur within coral biostromes and bioherms, mixed coral-siliceous sponge reefs, siliceous sponge meadows and mudmounds, and build up thrombolities with or without additional reef metazoans. The crusts are of paramount importance for the establishment and development of positive buildups. Commonly, reef growth starts with crusts which develop from a narrow base and rapidly expand laterally by rising above the sea floor. Reef associations with little or no microbial crust normally did not develop distinct relief. The basic microbial crust type is characterised by a dense to peloidal, mostly clotted, hence thrombolitic fabric which developed due to calcification triggered by microbes. Morphological evidence for this organic nature are positive relief, bridge-structures, and the shape and arrangement of peloids. The basic thrombolitic crust type is a eurytopic feature, equally occurring in settings of different bathymetry, waterenergy, salinity and oxygen/nutrient concentrations. However, the crusts also comprise additional micro-encrusters of variable abundance and diversity. The concurrent occurrence of these encrusters and diversity trends allows discrimination between crusts of different environments, particularly of different water depths. Microbial crusts from non-reefal marine oncoids show both similarities and differences with reefal crusts. For some of the mostly enigmatic micro-encrusters new clues to their nature could be detected. For instance, bubble-like structures, formerly interpreted as sporangia inLithocodium could be identified as the foraminiferBullopora aff.laevis, possibly living as a parasite or symbiont in theLithocodium algal tissue.Lithocodium andBacinella are regarded as different organisms.‘Tubiphytes’ morronensis clearly represents a symbiotic intergrowth between a nubeculinellid foraminifer and a microbe of unknown nature. The main prerequisite for the occurrence of microbial crusts is a cessation of background sedimentation which commonly can be tied to rises in sea level. This results in the development of crust-rich reefs. Fluctuations in oxygen and nutrient levels are indicated by dysaerobic bivalves and richness in authigenic glauconite, and led to the microbes outcompeting reefal metazoans, and to the development of thrombolites. Such thrombolites occur at very different depths which is interpreted to be related to a rise of dysaerobic waters due to climatic buffering and lowering of oceanic circulation during sea level rises. Microbial crusts in modern reefs are largely restricted to shaded, cryptic settings which contrasts with the wide distribution of crusts in Upper Jurassic reefs. Microbial crusts were increasingly replaced by coralline red algae since the Late Mesozoic, but despite their restricted modern habitat seem to still play an important, commonly overlooked role in the stabilisation of reef framework.     

Significance of the biomineralizing microniche in a lyngbya (cyanobacterium) travertine

Observations of the spatial arrangement of Lyngbya (Phormidium) incrustatum tri‐chomes in a travertine crust allowed prediction of the frequency of four types of microniche favorable for photosynthetically induced calcification. The most frequent type of microniche occurred at “cross‐over”; points where two trichomes made contact at a single point. Such points occurred very frequently (～6000/mm² travertine). Microniches more favorable to photocalcification consist of groups of three or more trichomes lying side by side, enclosing narrow tubes of interstitial water. These configurations were far less frequent and their effects would be negligible in Lyngbya travertines. In the calcified cyanobacteria Rivularia and Schizothrix such configurations may be much more common. The existence of such microniches demonstrated that regions favorable for photocalcification occur in travertine‐encrusted cyanobac‐terial mats and may initiate carbonate precipitation. However, the bulk of the carbonate precipitated in such travertines is known to be formed through inorganic processes.     

Genetic diversity and classification of cyanobacteria in different Azolla species by the use of PCR fingerprinting

Symbiotically associated cyanobacteria from 18 accessions within all known species in the genus Azolla were examined and classified by the use of polymerase chain reaction (PCR)-fingerprinting. A repetitive sequence specific for cyanobacteria, the short tandemly repeated repetitive (STRR) sequence, was used as a primer in the reaction. Cyanobacterial filaments isolated directly from the Azolla leaf cavity or contained within homogenised symbiotic Azolla tissue were used as templates. Based on the fingerprint pattern, distinct differences were demonstrated between cyanobacteria isolated from the Euazolla and Rhizosperma sections. In addition, individual fingerprints were obtained from all cyanobacteria isolated from the different Azolla species. The fingerprints were used to generate a phylogenetic tree. Three clusters were distinguished: one contained the four isolates from the section Euazolla, a second the isolate from Azolla filiculoides, and a third the three isolates from the section Rhizosperma. By the use of STRR-PCR fingerprinting, new data on the taxonomy of cyanobacteria in Azolla were obtained, which have been difficult to generate by other classification methods. PCR-fingerprinting may, therefore, be a valuable tool for diversity and classification studies of symbiotic cyanobateria from Azolla and, as co-evolution between the cyanobacteria and its corresponding host exists the method may also be useful for the taxonomy of Azolla. Received: 19 December 1998 / Accepted: 31 May 1999     

Effects of bacillamide and newly synthesized derivatives on the growth of cyanobacteria and microalgae cultures

The antialgal activity of newly synthesized bacillamides against several cyanobacteria and microalgae isolates was screened using a rapid 96-well microplate bioassay. Cultures were exposed to serial dilutions of each bacillamide derivative (0–160 μg mL⁻¹) in the microplate wells and daily optical measurements were used to estimate growth over a 216 h period. Inhibition values (%) were calculated from the estimated growth curves and inhibitory concentrations (IC₅₀-216 h) were obtained from the sigmoidal inhibition curves fitted by regression analysis. The effects of bacillamides on cell morphology and ultrastructure were also analysed by light and transmission electron microscopy. In general, the toxic cyanobacteria Microcystis aeruginosa, Aphanizomenon gracile, Anabaena circinalis and Anabaenopsis circularis were much more sensitive to bacillamides then the chlorophytes Ankistrodesmus falcatus and Scenedesmus obliquus. However, clear signs of morphological and ultrastructural changes induced by bacillamide were observed on both cyanobacteria and chlorophytes. Other cyanobacteria, namely the nostocalean Nodularia spumigena and the oscillatorialeans Leptolyngbya sp. and Planktothrix rubescens, exhibit higher tolerances to bacillamides, similar to that shown by different eukaryotic microalgae. Diatoms, on the other hand, proved to be quite as sensitive to most bacillamides as the most affected cyanobacteria. The properties of 5-iodo-Bacillamide (algicide or algistatic) were further investigated. This compound acted as an algistactic agent against eukaryotic algae and, depending on its concentration, acted as either an algicide or algistactic agent against most of the cyanobacteria tested. Although bacillamides cannot be considered as broad spectrum cyanobacterial algicides, different bacillamides might be of use in selectively controlling the growth of particular species of cyanobacteria.     

Utilization of hydrocarbons by cyanobacteria from microbial mats on oily coasts of the Gulf

Several pieces of evidence indicate that Microcoleu chthonoplastes and Phormidium corium, the predominant cyanobacteria in microbial mats on crude oil polluting the Arabian Gulf coasts, contribute to oil degradation by consuming individual n-alkanes. Both cyanobacteria grew phototrophically better in the presence of crude oil or individual n-alkanes than in their absence, indicating that hydrocarbons may have been utilized. This result was true when growth was measured in terms of dry biomass, as well as in terms of the content of biliprotein, the accessory pigment characteristic of cyanobacteria. The phototrophic biomass production by P. corium was directly proportional to the concentration of n-nonadecance (C₁₉) in the medium. The chlorophyll to carotene ratio of hydrocarbon-grown cyanobacteria did not decrease compared to the ratio in the absence of hydrocarbons, indicating that on hydrocarbons the organisms were not stressed. Comparing the fatty acid patterns of total lipids from hydrocarbon-grown cyanobacteria to those of the same organisms grown without hydrocarbons confirms that n-alkanes were taken up and oxidized to fatty acids by both cyanobacteria.     

Copepod grazing in a summer cyanobacteria bloom in the Gulf of Finland

Blooms of the cyanobacteria Nodularia spumigena and Aphanizomenon flos-aquae dominated the phytoplankton assemblages of the western Gulf of Finland and the eastern side of the northern Baltic Sea in late July–August, 1992. The bloom overlapped the peak seasonal contributions of the dominant mesozooplankton herbivores in the region, the copepods Acartia bifilosa and Eurytemora affinis and the cladoceran Bosmina longispina maritima. Using radio-labelling techniques; the copepods were offered one of the cyanobacteria, Nodularia, as well as the 10–54 µm fraction of the natural phytoplankton assemblage. In general, incorporation rates of the labelled phytoplankton into the copepods declined with increasing contributions of the cyanobacteria. For both copepods, incorporation was inversely related to total phytoplankton biomass, whether measured as chlorophyll, total cells or cyanobacteria biomass. The very low rates for Acartia (< 0.8 µl [copepod h]^–1) indicated that this copepod was likely starving in the cyanobacteria bloom, consistent with the generally poor condition of the animal observed in the laboratory. The other major mesozooplanktor, B. longispina maritima, ingested substantially more cyanobacterial biomass than the two copepods, based on HPLC-identified cyanobacteria-specific pigment echinenone in the gut. Bloom carbon provided < 1% and < 4% of the daily rations for Acartia and Eurytemora, respectively. Total copepod demand in the cyanobacteria blooms was trivial, < 1% of bloom biomass consumed daily. These results suggest that copepod herbivory is relatively unimportant in dissipating summer cyanobacteria blooms in the Gulf of Finland.     

The Absorption and Fluorescence Spectra of the Cyanobacterial Phycobionts of Cryptoendolithic Lichens in the High-Polar Regions of Antarctica

The algologically pure cultures of the green–brown cyanobacterium Chroococcidiopsissp. and three cyanobacteria of the genus Gloeocapsa, the blue–green Gloeocapsa sp.₁, the brown Gloeocapsa sp.₂, and the red–orange Gloeocapsa sp.₃, were isolated from sandstones and rock fissures in the high-polar regions of Antarctica. These cyanobacteria are the most widespread phycobionts of cryptoendolithic lichens in these regions. The comparative analysis of the absorption and the second-derivative absorption spectra of the cyanobacteria revealed considerable differences in the content of chlorophyll a and in the content and composition of carotenoids and phycobiliproteins. In addition to phycocyanin, allophycocyanin, and allophycocyanin B, which were present in all of the cyanobacteria studied, Gloeocapsa sp.₂ also contained phycoerythrocyanin and Gloeocapsa sp.₃ phycoerythrocyanin and C-phycoerythrin (the latter pigment is typical of nitrogen-fixing cyanobacteria). The fluorescence spectra of Gloeocapsa sp.₂ and Gloeocapsa sp.₃ considerably differed from the fluorescence spectra of the other cyanobacteria as well. The data obtained suggest that various zones of the lichens may be dominated either by photoheterotrophic or photoautotrophic cyanobacterial phycobionts, which differ in the content and composition of photosynthetic pigments.     

Charophyte-rich microfacies in the Barremian of the Eastern Iberian Chain (Spain)

Non-marine carbonate microfacies based on charophytes are a useful palaeoenvironmental tool that has been poorly developed to date. In the Barremian of the Maestrat Basin (Eastern Iberian Chain), five such microfacies are described and interpreted in terms of sedimentology and palaeoecology: (1) Microfacies of clavatoracean remains are mudstones and wackestones rich in utricles and well-preserved charophyte thalli of the genus Clavatoraxis, representing in situ deposition of clavatoracean meadows. (2) Microfacies of filamentous algae and clavatoraceans are mudstones with a high proportion of calcified filaments attributed to cyanobacteria and green algae. This facies may correspond to the open lake benthos. (3) Microfacies of Intraclasts and Charophyte remains are packstones formed on the shores of freshwater lakes or brackish swamps that underwent subaerial desiccation and wave reworking. (4) Microfacies of Porocharacean remains are wackestones and packstones with abundant gyrogonite fragments of genus Porochara. They are related to the reworking of a porocharacean meadow in shallow brackish marshes. (5) Microfacies of Munieria grambasti Bystricky 1976 fragments are grainstones attributed to the reworking of Munieria-dominated meadows. In consequence to these results, a palaeoecological model based on charophyte remains is proposed as a useful tool in carbonate facies analysis.     

A freshwater analog for the production of Epiphyton‐like microfossils

Calcified microbial microfossils—often interpreted as cyanobacteria—were important components of Precambrian and Paleozoic limestones, but their paucity in modern marine environments complicates our ability to make conclusive interpretations about their taxonomic affinity and geologic significance. Freshwater spring‐associated limestones (e.g., travertine and tufa) serve as terrestrial analogs to investigate mineralization in and around aquatic biofilms on observable timescales. We document the diagenesis of calcite fabrics associated with the freshwater algae Oocardium stratum, an epiphytic colonial green algae (desmid) known for producing stalks of extracellular polymeric substances (EPS) and passively producing a bifurcating tubular calcite monocrystal. Bifurcating EPS stalks produced by Oocardium colonies can become calcified and preserved in ancient carbonate deposits. Calcified micritic EPS stalks have a filamentous morphology, show evidence of branching, and maintain uniformity in diameter thickness throughout the mm‐scale colony, much like the enigmatic calcimicrobe Epiphyton. We provide a mechanism by which calcification associated with a colonial semispherical micro‐organism produces microfossils that deceptively resemble filamentous forms. These findings have implications for the use of morphological traits when assigning taxonomic affinities to extinct microfossil groups and highlight the utility of calcifying freshwater modern environments to investigate microbial taphonomy.     

Lithic cyanobacterial communities in the polyextreme Sahara Desert: implications for the search for the limits of life

The hyperarid Sahara Desert presents extreme and persistent dry conditions with a limited number of hours during which the moisture availability, temperature and light allow phototrophic growth. Some cyanobacteria can live in these hostile conditions by seeking refuge under (hypolithic) or inside (endolithic) rocks, by colonizing porous spaces (cryptoendoliths) or fissures in stones (chasmoendoliths). Chroococcidiopsis spp. have been reported as the dominant or even the only phototrophs in these hot desert lithic communities. However, the results of this study reveal the high diversity of and variability in cyanobacteria among the sampled habitats in the Sahara Desert. The chasmoendolithic samples presented high coccoid cyanobacteria abundances, although the dominant cyanobacteria were distinct among different locations. A high predominance of a newly described cyanobacterium, Pseudoacaryochloris sahariense, was found in hard, compact, and more opaque stones with cryptoendolithic colonization. On the other hand, the hypolithic samples were dominated by filamentous, non-heterocystous cyanobacteria. Thermophysiological bioassays confirmed desiccation and extreme temperature tolerance as drivers in the cyanobacterial community composition of these lithic niches. The results of the present study provide key factors for understanding life strategies under polyextreme environmental conditions. The isolated strains, especially the newly described cyanobacterium P. sahariense, might represent suitable microorganisms in astrobiology studies aimed at investigating the limits of life.     

Comparative Study of Cyanobacteria as Biocatalysts for the Asymmetric Synthesis of Chiral Building Blocks

The three representative cyanobacteria, Synechococcus PCC7942, Anabaena variabilis, and Nostoc muscorum, were studied for their ability to asymmetrically reduce the prochiral ketones 2′‐3′‐4′‐5′‐6′‐pentafluoroacetophenone, ethyl 4‐chloroacetate, 4‐chloroacetophenone, and ethylbenzoylacetate to the corresponding chiral alcohols. Photosynthesis as well as respiration was applied for intracellular regeneration of the NAD(P)H cofactor. It was shown for the first time that all cyanobacteria were able to reduce the prochiral ketones asymmetrically without light for cofactor regeneration. By comparison of the cell specific product formation capacities of cyanobacteria with typical heterotrophic whole cell biocatalysts in batch processes, it is shown that comparable or, in some cases, better performances at high enantiomeric excess (ee > 99.8 %) are obtained. As a consequence of a generally strong product inhibition, in situ product removal must be applied in order to restore process efficiency when using cyanobacteria as biocatalysts.     

The histological structure of the calcified lung of the fossil coelacanth Axelrodichthys araripensis (Actinistia: Mawsoniidae)

Abstract: The palaeohistological study of the calcified internal organ of Axelrodichthys araripensis Maisey, 1986, a coelacanthiform from the Lower Cretaceous of Brazil (Crato (Aptian) and Santana (Albian) formations of the Araripe Basin), shows that the walls of this organ consist of osseous blades of variable thickness separated from each other by the matrix. This indicates that, in the living individuals, the walls were reinforced by ossified plates, probably separated by conjunctive tissue. This calcified sheath present in Axelrodichthys, as well as in other fossil coelacanths, lies in ventral position relative to the gut and its single anterior opening is located under the opercle, suggesting a direct connection with the pharynx or the oesophagus. The calcified organ of Axelrodichthys, like that of other fossil coelacanths, is here regarded as an ‘ossified lung’ and compared with the ‘fatty lung’ of the extant coelacanth Latimeria. The reinforcement of the pulmonary walls by the overlying osseous blades could be interpreted as a means of adapting volumetric changes in the manner of bellows, a necessary function for ventilation in pulmonary respiration. Other functional hypotheses such as hydrostatic and/or acoustic functions are also discussed.     

Associative Cyanobacteria Isolated from the Roots of Epiphytic Orchids

Associative cyanobacteria were isolated from the rhizoplane and velamen of the aerial roots of the epiphytic orchids Acampe papillosa, Phalaenopsis amabilis, and Dendrobium moschatum and from the substrate roots of A. papillosa and D. moschatum. Cyanobacteria were isolated on complete and nitrogen-free variants of BG-11 medium. On all media and in all samples, cyanobacteria of the genus Nostoc predominated. Nostoc, Anabaena, and Calothrixwere isolated from the surface of the A. papillosa aerial roots, whereas the isolates from the substrate roots were Nostoc, Oscillatoria,and representatives of the LPP group (Lyngbia, Phormidium, and Plectonema, incapable of nitrogen fixation). On the D. moschatum substrate roots, Nostoc and LPP group representatives were also found, as well as Fischerella. On the aerial roots of P. amabilis and D. phalaenopsis grown in a greenhouse simulating the climate of moist tropical forest, cyanobacteria were represented by Nostoc, LPP group, and Scytonema in D. phalaenopsis and by Nostoc, Scytonema, Calothrix, Spirulina, Oscillatoria, and the LPP group in P. amabilis. For D. moschatum, the spectra of cyanobacteria populating the substrate root rhizoplane and the substrate (pine bark) were compared. In the parenchyma of the aerial roots of P. amabilis, fungal hyphae and/or their half-degraded remains were detected, which testifies to the presence of mycorrhizal fungi in this plant. This phenomenon is attributed to the presence of a sheath formed by cyanobacteria and serving as a substrate for fungi.     

Underestimated biodiversity as a major explanation for the perceived rich secondary metabolite capacity of the cyanobacterial genus Lyngbya

Marine cyanobacteria are prolific producers of bioactive secondary metabolites responsible for harmful algal blooms as well as rich sources of promising biomedical lead compounds. The current study focused on obtaining a clearer understanding of the remarkable chemical richness of the cyanobacterial genus Lyngbya. Specimens of Lyngbya from various environmental habitats around Curaçao were analysed for their capacity to produce secondary metabolites by genetic screening of their biosynthetic pathways. The presence of biosynthetic pathways was compared with the production of corresponding metabolites by LC‐ESI‐MS² and MALDI‐TOF‐MS. The comparison of biosynthetic capacity and actual metabolite production revealed no evidence of genetic silencing in response to environmental conditions. On a cellular level, the metabolic origin of the detected metabolites was pinpointed to the cyanobacteria, rather than the sheath‐associated heterotrophic bacteria, by MALDI‐TOF‐MS and multiple displacement amplification of single cells. Finally, the traditional morphology‐based taxonomic identifications of these Lyngbya populations were combined with their phylogenetic relationships. As a result, polyphyly of morphologically similar cyanobacteria was identified as the major explanation for the perceived chemical richness of the genus Lyngbya, a result which further underscores the need to revise the taxonomy of this group of biomedically important cyanobacteria.