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Microalgae, with cyanobacteria, are the major primary producers in aquatic, predominantly marine, ecosystems, contributing to the biogeochemical cycling of multiple elements despite their small instantaneous biomass. Their evolutionary history revealed by genomic analyses has shown a complex past that produces a polyphyletic group including organisms that have undergone primary, secondary and tertiary chloroplast endosymbiosis with genetic integration and also horizontal gene transfer. All but one genus of photosynthetic eukaryotes arose by endosymbiosis of a gloeomargarita-like β-cyanobacterium in a eukaryote with the retention of some genes in the plastid, the transfer of more genes to the eukaryote nucleus, and the loss of many others, to produce the Archaeplastida. A second, much later, endosymbiosis of an α-cyanobacterium in a euglyphid amoeba yielded Paulinella. The diversification of the Archaeplastida yielded Glaucophyta, Rhodophyta, Chlorophyta and Streptophyta. Secondary endosymbiosis of red algae in eukaryotes led to microalgae of the ‘red line’, that is, photosynthetic Ochrista (= stramenopiles), Haptophyta, Cryptophyta and Alveolata (dinoflagellates and chromerids). Secondary endosymbiosis of chlorophyte algae in eukaryotes yielded microalgae of the Chlorarachniophyta and Euglenophyta. The ‘red line’ of secondary endosymbionts are dominant marine phytoplankton, possibly related to the occurrence of chlorophyll c that has high absorbance of blue light that dominates in deep ocean waters.
The evolution of oxygenic photosynthesis had profound effects on the biogeochemistry of the planet. The increase in atmospheric oxygen levels brought about alterations to a range of biogeochemical processes involving changes in the availability of a host of elements, including nitrogen, sulfur and many metal ions such as iron and manganese, central to biological activities. Critically for photosynthetic organisms, the increase in oxygen levels in the atmosphere following the evolution of oxygenic photosynthesis and the Great Oxidation Event had consequences for the assimilation of inorganic carbon via the enzyme ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco). Although there are a number of alternative pathways leading to autotrophic CO2 assimilation, 99% of primary productivity on the planet is carried out by processes that involve Rubisco and the Benson–Calvin–Bassham cycle. The accumulation of O2 in the atmosphere also had major repercussions for increasing the energetic yield of the catabolism of photosynthate by allowing oxidative respiration, with a much greater ATP yield than from anaerobic fermentative processes. The interaction of O2 with UVC radiation led to the production of UVC- and UVB-absorbing O3. This also significantly influenced life on Earth and facilitated the colonisation of the upper ocean and terrestrial surface.
The Vestfold Hills are a 400 km2, isolated ice-free oasis in eastern Antarctica featuring large areas with translucent quartz rocks that provide habitat for hypolithic microbial communities underneath. We used high-throughput DNA sequencing of 16S and 18S ribosomal RNA amplicons to characterize bacterial and eukaryotic hypolithic communities across the Vestfold Hills. We found high-level, local heterogeneity in community structure consistent with limited dispersal between hypoliths. Hypolithic communities were dominated by heterotrophic Bacteroidetes (mean bacterial relative read abundance: 56%) as well as Cyanobacteria (35%), with the eukaryote component often dominated by Chlorophyta (43%). Small but significant proportions of the variation in microbial community composition and function were explained by soil salinity (5–7%) and water availability (8–11%), with distinct taxa associated with different salinities and water availabilities. Furthermore, many inferred bacterial metabolic pathways were enriched in hypolithic communities from either dry or high-salinity sites. Vestfold Hills hypolithic habitats are likely to be local refuges for bacterial and eukaryotic diversity. Gradients in soil salinity and water availability across the Vestfold Hills, in addition to the number and diversity of lake types and fjords as potential source populations, may contribute to the observed variation in the extremophile, hypolithic microbial community composition.
With the increasing need for architectural sustainability, biodesign offers a new approach to incorporating living organisms in building materials. Bacteria hold a range of biological activities that impact their environment, and which could enable the solidification of inorganic materials; this has already been seen with microbially-induced carbonate precipitation that strengthens bonds between sand particles. This paper describes the novel development of an additive co-fabrication manufacturing process, demonstrating an interdisciplinary approach of architecture and microbiology. Specifically, the activity of a biological deposition (i.e., cyanobacterial calcium carbonate precipitation) and its integration with that of a robotic deposition (i.e., a sand-based biomixture) within an architectural biofabrication workflow. Two bacterial strains were successfully grown in potential sand-based construction materials. Microbiological protocols, such as optical density and fluorescence measurements, were then applied to identify parameters, for harvesting light through photosynthesis and harnessing it to the sedimentation of calcium carbonate. Assessments of the proposed mechanical delivery system and printing properties enabled the outlining of a suitable robotic deposition system for sand-based mixtures. Through examinations of these microbiological and mechanical protocols, this paper outlines design strategies and tradeoffs for an integrated workflow, that corresponds with both the biological (micro) and architectural (macro) scales.
This chapter serves as an introduction to the book. It discusses the origin of Planet Earth and its Moon, their dependence on the Sun for energy, and the evolution of life on Earth. The evolution of the first living cell seems to have been a single event and all life on Earth is directly derived from this individual primary organism. The first life forms were anaerobic bacteria, but these later gave rise to photosynthesising cyanobacteria, which produced oxygen. The presence of oxygen eventually led to the emergence of aerobic animals and plants. The chapter then details the emergence of the oceans and supercontinents Pangea and Gondwanaland, the eventual break-up of the supercontinents and the development of the varied ecosystems which characterise Planet Earth at the present time.
Glacier phototroph blooms on the surfaces of ice sheets and glaciers cause albedo reduction, leading to increased melting rates. We observed seasonal changes in the abundance of phototrophs on the Qaanaaq Ice Cap in northwestern Greenland from June to August 2014, and reproduced these changes using numerical and empirical models. The phototroph community on the ice surface mainly consisted of the glacier alga Ancylonema nordenskioldii and the cyanobacterium Phormidesmis priestleyi. The glacier alga appeared on the ice surface in late June, after which its abundance increased exponentially throughout the melting period. A logistic growth model designed for snow algal growth reproduced the measured exponential increases, suggesting that growth could be explained using the model as a function of the ice melting duration. Cyanobacteria appeared and their abundance increased in late July but did not change exponentially thereafter. The abundance of cyanobacteria was explained with an empirical model expressed as a function of the amount of mineral dust on the bare ice surface. Our numerical and empirical models for reproducing glacier algae and cyanobacteria could be useful for quantifying the albedo reduction caused by their growth and the melt rates of the Greenland ice sheet and glaciers in the future.
In the only salt evaporation pond retaining its natural setting of the historic Salina di Cervia (Italy), the northernmost salterns of the Mediterranean area, a number of potentially preservable textures derive from the interactions between photosynthetic mat producers and the sedimentary substrate. These morphologies occur at the beginning of the taphonomic processes when repeated emerged-submerged conditions take place. In these conditions the cohesive nature of the diatom- and cyanobacterial-derived mucilage favours the stabilization of otherwise ephemeral structures. Surface micromorphologies for which diatoms and cyanobacteria have played some active role when still living in the soft microlayer and down to the sediment-water interface, such as during the gliding motility, can overcome the surface layer of most intense mixing (i.e., the taphonomically active zone) and keep traces of them in the fossil record either as body fossils or as texture contributors. Tiny microbial-derived remnants, such as filaments and biofilm strands of halotolerant microorganisms, while fragile upon their formation, can therefore stabilize as biosignatures when combined with salt precipitation. Halophilic and halotolerant ecosystems are models for life in extreme environments (analogue sites) with similarity to those strongly suspected to occur and/or have occurred on Mars and on other planetary bodies. The study of hypersaline systems such as Salina di Cervia which harbour diverse and abundant microbial life, can be relevant for astrobiology since it allows the investigation of potential biosignatures and their preservation, and of further understand the range of conditions and the planetary processes sustaining potentially habitable systems.
Lichens are a well-known symbiosis between a host mycobiont and eukaryote algal or cyanobacterial photobiont partner(s). Recent studies have indicated that terrestrial lichens can also contain other cryptic photobionts that increase the lichens’ ecological fitness in response to varying environmental conditions. Marine lichens live in distinct ecosystems compared with their terrestrial counterparts because of regular submersion in seawater and are much less studied. We performed bacteria 16S and eukaryote 18S rRNA gene metabarcoding surveys to assess total photobiont diversity within the marine lichen Lichina pygmaea (Lightf.) C. Agardh, which is widespread throughout the intertidal zone of Atlantic coastlines. We found that in addition to the established cyanobacterial photobiont Rivularia, L. pygmaea is also apparently host to a range of other marine and freshwater cyanobacteria, as well as marine eukaryote algae in the family Ulvophyceae (Chlorophyta). We propose that symbiosis with multiple freshwater and marine cyanobacteria and eukaryote photobionts may contribute to the ability of L. pygmaea to survive the harsh fluctuating environmental conditions of the intertidal zone.
A review of algal (including cyanobacterial) symbionts associated with lichen-forming fungi is presented. General aspects of their biology relevant to lichen symbioses are summarized. The genera of algae currently believed to include lichen symbionts are outlined; approximately 50 can be recognized at present. References reporting algal taxa in lichen symbiosis are tabulated, with emphasis on those published since the 1988 review by Tschermak-Woess, and particularly those providing molecular evidence for their identifications. This review is dedicated in honour of Austrian phycologist Elisabeth Tschermak-Woess (1917–2001), for her numerous and significant contributions to our knowledge of lichen algae (some published under the names Elisabeth Tschermak and Liesl Tschermak).
The presence of perchlorate in the Martian soil may limit in-situ resource utilization (ISRU) technologies to support human outposts. In order to exploit the desiccation, radiation-tolerant cyanobacterium Chroococcidopsis in Biological Life Support Systems based on ISRU, we investigated the perchlorate tolerance of Chroococcidopsis sp. CCMEE 029 and its derivative CCMEE 029 P-MRS. This strain was obtained from dried cells mixed with Martian regolith simulant and exposed to Mars-like conditions during the BIOMEX space experiment. After a 55-day exposure of up to 200 mM perchlorate ions, a tolerance threshold value of 100 mM perchlorate ions was identified for both Chroococcidopsis strains. After 40-day incubation, a Mars-relevant perchlorate concentration of 2.4 mM perchlorate ions, provided as a 60 and 40% mixture of Mg- and Ca-perchlorate, had no negative effect on the growth rate of the two strains. A proof-of-concept experiment was conducted using Chroococcidopsis lysate in ISRU technologies to feed a heterotrophic bacterium, i.e. an Escherichia coli strain capable of metabolizing sucrose. The sucrose content was fivefold increased in Chroococcidopsis cells through air-drying and the yielded lysate successfully supported the bacterial growth. This suggested that Chroococcidopsis is a suitable candidate for ISRU technologies to support heterotrophic BLSS components in a Mars-relevant perchlorate environment that would prove challenging to many other cyanobacteria, allowing a ‘live off the land’ approach on Mars.
Basidiolichens are generally poorly researched because of the very small number of species and their restriction to special niches. Lichenomphalia basidiolichens grow in considerable quantities in arctic and alpine habitats but they are inadequately studied in these habitats in Mediterranean areas. Based on morphological and phylogenetic analyses, we identified the different symbionts of L. meridionalis, collected in localities in Spain at altitudes ranging from 533 to 2200 m above sea level. The present study provides the first molecular data available for L. meridionalis. We found that a microindel of six bp within the nrITS2 could help to discriminate L. meridionalis from other species of the genus. Molecular analyses revealed the existence of two different green algal strains, both belonging to Coccomyxa subellipsoidea, a species shared with other Lichenomphalia lichens. Notably, the two chlorobiont strains associated with L. meridionalis were differentially distributed according to altitude, and samples having one of the two strains consistently also included cyanobacteria.
By investigating the survival and the biomarker detectability of a rock-inhabiting cyanobacterium, Chroococcidiopsis sp. CCMEE 029, the BIOMEX space experiment might contribute to a future exploitation of the Moon as a test-bed for key astrobiology tasks such as the testing of life-detection technologies and the study of life in space. Post-flight analyses demonstrated that the mixing of dried cells with sandstone and a lunar regolith simulant provided protection against space UV radiation. During the space exposure, dried cells not mixed with minerals were killed by 2.05 × 102 kJ m−2 of UV radiation, while cells mixed with sandstone or lunar regolith survived 1.59 × 102 and 1.79 × 102 kJ m−2, respectively. No differences in survival occurred among cells mixed and not mixed with minerals and exposed to space conditions in the dark; this finding suggests that space vacuum and 0.5 Gy of ionizing radiation did not impair the cells’ presence in space. The genomic DNA of dead cells was severely damaged but still detectable with PCR amplification of a short target, thus suggesting that short sequences should be targeted in a PCR-based approach when searching for traces of life. The enhanced stability of genomic DNA of dried cells mixed with minerals and exposed to space indicates that DNA might still be detectable after prolonged periods, possibly up to millions of years in microbes shielded by minerals. Overall, the BIOMEX results contribute to future experiments regarding the exposure of cells and their biomarkers to deep space conditions in order to further test the lithopanspermia hypothesis, the biomarker stability and the microbial endurance, with implications for planetary protection and to determine if the Moon has been contaminated during past human missions.
Hydration traits determine much of a lichen’s distribution pattern along a climatic gradient but the mechanisms involved are still incompletely known. A higher abundance of large external cephalodia in wet oceanic than in drier climates has previously been reported in Lobaria amplissima. This study aims to quantify how much more rain L. amplissima thalli with external cephalodia would need to fill their internal water holding capacity (WHCinternal) than thalli without. The mean WHCinternal was 1·8 times higher in thalli with external cephalodia than in those without. The WHCinternal when converted to mm rain needed to saturate an average specimen was 1·37 mm (min–max: 0·55–3·8 mm) for a cephalodiate thallus, whereas an average thallus without external cephalodia needed just 0·76 mm (min–max: 0·36–1·3 mm). Known dewfall rates and rates of water uptake from humid air are far below what is needed to saturate even the cephalodiate thallus with the lowest WHCinternal, implying a stronger dependency on rain for thalli with external cephalodia. Thus, the observed trends in this study are consistent with earlier reports of decreasing frequency of external cephalodia from wet to drier climates.
Cyanobacteria (blue-green algae) are undesirable in ponds used to raise fish for human food. Management of cyanobacterial communities in aquaculture ponds has been hindered by the small number of herbicides approved for use in food-fish ponds and by the lack of selectivity toward cyanobacteria for those herbicides that are approved for that use. To facilitate development of additional management options, a rapid bioassay utilizing 96-well cell culture plates was developed for screening herbicides and other phytotoxins for selective toxicity toward cyanobacteria. Oscillatoria cf. chalybea and Selenastrum capricornutum were chosen as representatives of cyanobacteria (Cyanophyta) and green algae (Chlorophyta), respectively. In the bioassay, wells of the cell culture plates were inoculated with cyanobacterial or unialgal culture. One of five herbicides (atrazine, diuron, endothall, fluridone, or paraquat) was then added to the wells at various concentrations, and absorbance (650 nm) was measured at 24-h intervals. Growth of treated cultures relative to untreated cultures was used to determine relative toxicity of the herbicide to the two organisms. Paraquat was the most selective of the herbicides tested and was over 10-fold more toxic to O. cf. chalybea than to S. capricornutum. This method was demonstrated to be a rapid, effective, and highly reproducible bioassay to identify selective algicides.
Small-island coral reef ecosystems are usually closely coupled to the activities of human inhabitants. Ahus Island (Papua New Guinea) is an isolated Pacific island with a rapidly growing population, heavy reliance on marine resources and limited infrastructure. We hypothesized that untreated sewage was driving distinct benthic assemblages around Ahus and neighbouring uninhabited Onetah. At sites with varying proximities to beach toilets, fore-reef herbivorous fish biomass and benthic composition were measured alongside reef-flat sedimentary oxygen consumption (SOC); a high SOC rate reflects high organic input into coastal waters, thus serving as a potential indicator of sewage input. Fish biomass was low (17.1–20.1 g m–2), but consistent between sites. However, cyanobacteria dominated the fore-reef closest to toilets (62 ± 3%) with highest reef-flat SOC, whereas hard corals dominated furthest away (63 ± 1%), where SOC was lowest. To our knowledge, this is the first study that used SOC to detect local differences in sewage pollution. The results indicate that whilst corals can maintain their dominance on overfished reefs, additional sewage stress may drive pronounced benthic shifts, highlighting the urgency to improve small-island waste management.
Allelopathy has been suggested as a mechanism of interference in several weed species. Allelochemicals released from certain weed species influence the growth and yield of crop species. Several laboratory studies present circumstantial evidence of the occurrence of allelopathy as a causative agent in weed–crop agroecosystems. Field evidence, however, is still lacking. In this paper, the significance of field studies is argued in terms of a multifaceted approach to allelopathy, and mugwort is used as an example. Previous research demonstrated the allelopathic potential of mugwort; however, experiments were not carried out in a natural environment. Inderjit and Foy (1999) have demonstrated that chemical characteristics (pH, inorganic ions, and phenolics) of soil amended with mugwort leaf leachate were altered when compared to unamended soil. We have analyzed the mugwort-infested field soil and compared its chemical characteristics with those of amended soils. No definite trend, in terms of influence of mugwort on soil chemistry, was observed. Results indicate the importance of field studies in order to obtain ecologically relevant data from laboratory studies. Field situations are often complex in terms of the presence of interfering flora. Cyanobacteria, for example, play an important role in weed–crop interactions in rice paddy soils. Allelochemicals released from weed species present in the paddy field may influence nitrogen-fixing cyanobacteria in addition to their phytotoxic effects to the paddy crop. Significance of phytotoxic effects of weed species on crop growth, and N2-fixing potential of cyanobacteria in paddy soils is discussed.
The present study investigates the photobiont diversity of the boreal felt lichen, Erioderma pedicellatum. Previously sampled genetic data from Newfoundland were reanalyzed and new sequence data (16S rDNA, rbcLX) of the boreal felt lichen from Alaska (USA), Kamchatka (Russia), and North Trøndelag (Norway) were generated. The highest genetic diversity of the photobiont is found in Alaska and Kamchatka, indicating that these may be the primary sources of the species in the Northern Hemisphere. In Newfoundland, the photobiont of E. pedicellatum was screened on leaves of the symbiotic liverwort Frullania asagrayana and it was found to occur on trees where no other lichens were present, demonstrating that the geographical distribution, and possibly also the ecological requirement of the photobiont of E. pedicellatum, is wider than that of the lichen phenotype. Finally, a postulated association between the occurrence of the vegetatively reproducing Coccocarpia palmicola and the occurrence of the compatible photobiont of E. pedicellatum on the same tree could not be established.
The biology and ecology of calcarean sponges are not as well understood as they are for demosponges. Here, in order to gain new insights, particularly about symbiotic relationships, the calcarean sponge Leucetta prolifera was sampled from south-western Australia and examined for its assumed photosymbionts. Pulse amplitude modulated fluorometry and extraction of photopigments established that the sponge was photosynthetic. Molecular analysis of the bacterial symbionts via sequencing of the V1–V3 region of the 16S rDNA gene confirmed that between 5 and 22% of all sequences belonged to the phylum Cyanobacteria, depending on the individual sample, with the most dominant strain aligning with Hormoscilla spongeliae, a widely distributed sponge symbiont. Analysis of fatty acids suggested that the sponge obtains nutrition through photosynthates from its symbionts. The relationship is assumed to be mutualistic, with the sponge receiving dietary support and the cyanobacteria sheltering in the sponge tissues. We list all Calcarea presently known to harbour photosymbionts.
Even though technological advances could allow humans to reach Mars in the coming decades, launch costs prohibit the establishment of permanent manned outposts for which most consumables would be sent from Earth. This issue can be addressed by in situ resource utilization: producing part or all of these consumables on Mars, from local resources. Biological components are needed, among other reasons because various resources could be efficiently produced only by the use of biological systems. But most plants and microorganisms are unable to exploit Martian resources, and sending substrates from Earth to support their metabolism would strongly limit the cost-effectiveness and sustainability of their cultivation. However, resources needed to grow specific cyanobacteria are available on Mars due to their photosynthetic abilities, nitrogen-fixing activities and lithotrophic lifestyles. They could be used directly for various applications, including the production of food, fuel and oxygen, but also indirectly: products from their culture could support the growth of other organisms, opening the way to a wide range of life-support biological processes based on Martian resources. Here we give insights into how and why cyanobacteria could play a role in the development of self-sustainable manned outposts on Mars.
Microbial mats are common in polar streams and often dominate benthic biomass. Climate change may be enhancing the variability of stream flows in the Antarctic, but so far studies investigating mat responses to disturbance have been limited in this region. Mat regrowth was evaluated following disturbance by experimentally scouring rocks from an ephemeral McMurdo Dry Valley stream over two summers (2001–02 and 2012–13). Mats were sampled at the beginning and resampled at the end of the flow season. In 2012–13, mats were additionally resampled mid-season along with previously undisturbed controls. In 2001–02 rocks regained 47% of chlorophyll a and 40% of ash-free dry mass by the end of the summer, while in 2012–13 rocks regrew 18% and 27%, respectively. Mat stoichiometry differed between summers, and reflected differences in biomass and discharge. Oscillatoria spp. were greatest on scoured rocks and Phormidium spp. on undisturbed rocks. Small diatoms Humidophila and Fistulifera spp. increased throughout the summer in all mats, with the latter more abundant in scoured communities. Collectively, these data suggest that mats are variable intra-annually, responsive to hydrology and require multiple summers to regrow initial biomass once lost. These results will aid the interpretation of long-term data, as well as inform Antarctic Specially Managed Area protocols.