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In the hyper-arid Atacama Desert, microbial life thrives near its “dry limit” in scarcely distributed habitats. Fracture networks of salt-poor sand wedges outlining salt-cemented polygons on alluvial surfaces in the Yungay region (Chile) represent potential microbial habitats. The degree of soil habitability at the surface (0–5 cm depth) and subsurface (10–15 cm depth) of a polygon and adjacent sand wedge was assessed before and up to 42 days after a 20 mm simulated rain experiment through the abundance of phospholipid fatty acids (PLFAs). Mineralogical composition, salinity, pH, electrical and thermal conductivity, water content, and water activity were analyzed for their relevance to habitability. After wetting, the PLFA content exclusively increased steadily with time in the polygon subsurface indicating the growth of an indigenous bacterial community. This increased habitability is presumably related to the soil's ability to retain water for at least 6 weeks at this depth. The lack of a continuous growth signal at the surface is likely due to rapid desiccation. In the sand wedge subsurface, the increase in PLFA content is not continuous despite the water activity being >0.9. The reason for this remains unclear but indicates that not only water availability is relevant for habitability but also the here described soil heterogeneities might impact the detection of the microbial response. Yet, the increasing PLFA trend in the polygon subsurface emphasized its relevance as a saline microbial habitat in an otherwise hostile environment, which could have implications for the assessment of soil habitability on Mars

Polygonal grounds are landscape features commonly associated with periglacial environments originating from freeze-thawing cycles or frost-related processes. However, such a genesis is unlikely for polygonal grounds on alluvial surfaces in the warm and hyper-arid Atacama Desert due to the lack of enduring sub-zero temperatures and limited water availability, whereas a cracking mechanism based on thermal contraction and/or desiccation is more plausible. To differentiate between those mechanisms, we performed a quantitative morphometric terrain characterization in combination with a geochemical and sedimentological analysis on three polygonal networks located in the Yungay area of the Atacama Desert, Chile. Our data show that these sand wedge polygons differ from other polygonal features in the Yungay area such as salt polygons and mud crack polygons from playa environments in regard to composition, morphometry and topographical setting. The investigated polygonal soils are composed of siliciclastic sediment that is mainly cemented by sulfates (gypsum & anhydrite) in the shallow ground (~0–50 cm) and by nitrates and chlorides in the deeper ground (~50–100 cm) while being separated by about 1 m deep, salt-poor and V-shaped sand wedges. The low clay content (~2 wt%) makes an exclusive desiccation origin less relevant whereas the high salt content (63 wt%) and the high surface temperature variations make thermal contraction origin more likely. Morphometric data indicate a link between topography and polygon geometry, as the flat-centered polygons (mean size 3.9 to 4.7 m) are aligned either in slope direction or perpendicular to it, while being more elongated on steeper slopes, which is common to thermal contraction polygons. Although we cannot exclude that desiccation cracking plays a minor role for the formation of the here described polygons, we conclude that their genesis is dominated by thermal contraction.

The Atacama Desert is the oldest and driest non-polar desert on Earth. Millions of years of hyperaridity enabled salt accumulations through atmospheric deposition. These salts can serve as proxies to decipher the interaction between water and soil as well as to understand the habitability with changing environmental settings. Therefore, we investigated four soil profiles regarding their mineralogy, salt abundance, and sulfate stable isotopic composition. The profiles were located along an elevation transect in the hyperarid region southeast of Antofagasta, Chile. The two lower sites situated on the distal parts of inactive alluvial fan deposits were subject to occasional fog occurrences. The upper steeper-sloped sites experienced no fog but are subject to minimal erosion. In all soil profiles, sulfates are the dominant salts showing a downward transition from gypsum to anhydrite that is accompanied by an increase of highly soluble salts and a decrease of sulfate δ34S and δ18O values. These trends are consistent with downward directed water infiltration during rare rain events causing salt dissolution followed by precipitation within the deeper soil column. This conclusion is also supported by our Rayleigh fractionation model. We attribute the presence of anhydrite at > 40 cm depth to the cooccurrence of nitrate and chloride salts, which decreases water activity during sulfate precipitation and therefore drives anhydrite formation. Along the elevation transect, the total salt inventories of each profile show a trend for nitrates and chlorides concentration decreasing with elevation. This observation together with the sulfate stable isotopes indicates a fog-independent source and suggests remobilization of soluble salts through enhanced washout from hillslopes to alluvial fans. These findings are essential for assessing the long-term regional habitability of hyperarid environments and have also relevance for Mars.

Background: The hyperarid core of the Atacama Desert is an extremely harsh environment thought to be colonized by only a few heterotrophic bacterial species. Current concepts for understanding this extreme ecosystem are mainly based on the diversity of these few species, yet a substantial area of the Atacama Desert hyperarid topsoil is covered by expansive boulder accumulations, whose underlying microbiomes have not been investigated so far. With the hypothesis that these sheltered soils harbor uniquely adapted microbiomes, we compared metagenomes and geochemistry between soils below and beside boulders across three distantly located boulder accumulations in the Atacama Desert hyperarid core. Results: Genome-resolved metagenomics of eleven samples revealed substantially different microbial communities in soils below and beside boulders, despite the presence of shared species. Archaea were found in significantly higher relative abundance below the boulders across all samples within distances of up to 205 km. These key taxa belong to a novel genus of ammonia-oxidizing Thaumarchaeota, Candidatus Nitrosodeserticola. We resolved eight mid-to-high quality genomes of this genus and used comparative genomics to analyze its pangenome and site-specific adaptations. Ca. Nitrosodeserticola genomes contain genes for ammonia oxidation, the 3-hydroxypropionate/4-hydroxybutyrate carbon fixation pathway, and acetate utilization indicating a chemolithoautotrophic and mixotrophic lifestyle. They also possess the capacity for tolerating extreme environmental conditions as highlighted by the presence of genes against oxidative stress and DNA damage. Site-specific adaptations of the genomes included the presence of additional genes for heavy metal transporters, multiple types of ATP synthases, and divergent genes for aquaporins. Conclusion: We provide the first genomic characterization of hyperarid soil microbiomes below the boulders in the Atacama Desert, and report abundant and highly adapted Thaumarchaeaota with ammonia oxidation and carbon fixation potential. Ca. Nitrosodeserticola genomes provide the first metabolic and physiological insight into a thaumarchaeal lineage found in globally distributed terrestrial habitats characterized by various environmental stresses. We consequently expand not only the known genetic repertoire of Thaumarchaeota but also the diversity and microbiome functioning in hyperarid ecosystems. Conclusion: We provide the first genomic characterization of hyperarid soil microbiomes below the boulders in the Atacama Desert, and report abundant and highly adapted Thaumarchaeaota with ammonia oxidation and carbon fixation potential. Ca. Nitrosodeserticola genomes provide the first metabolic and physiological insight into a thaumarchaeal lineage found in globally distributed terrestrial habitats characterized by various environmental stresses. We consequently expand not only the known genetic repertoire of Thaumarchaeota but also the diversity and microbiome functioning in hyperarid ecosystems.

The hyperarid core of the Atacama Desert is an extremely harsh environment previously thought to be colonized by only a few heterotrophic bacterial species. In addition, carbon and nitrogen cycling in these highly oligotrophic ecosystems are poorly understood. Here we genomically resolved a novel genus of Thaumarchaeota, Ca. Nitrosodesertus, found below boulders of the Atacama hyperarid core, and used comparative genomics to analyze their pangenome and site-specific adaptations. Their genomes contain genes for ammonia oxidation and the 3-hydroxypropionate/4-hydroxybutyrate carbon fixation pathway, indicating a chemolithoautotrophic lifestyle. Ca. Nitrosodesertus possesses the capacity for tolerating extensive environmental stress highlighted by the presence of genes against oxidative stress, DNA damage and genes for the formation of biofilms. These features are likely responsible for their dominance in samples with extremely low water content across three different boulder fields and eight different boulders. Genome-specific adaptations of the genomes included the presence of additional genes for UV resistance, heavy metal transporters, multiple types of ATP synthases, and divergent genes for aquaporins. Our results suggest that Thaumarchaeota mediate important carbon and nitrogen cycling in the hyperarid core of the Atacama and are part of its continuous and indigenous microbiome.

The colonization of emergent continental landmass by microbial life was an evolutionary step of paramount importance in Earth history. Here we report direct fossil evidence for life on land 3,220 million years ago (Ma) in the form of terrestrial microbial mats draping fluvial conglomerates and gravelly sandstones of the Moodies Group, South Africa. Combined field, petrographic, carbon isotope and Raman spectroscopic analyses confirm the synsedimentary origin and biogenicity of these unique fossil mats as well as their fluvial habitat. The carbon isotope compositions of organic matter (δ13Corg) from these mats define a narrow range centred on −21‰, in contrast to fossil mats of marine origin from nearby tidal deposits that show δ13Corg values as low as −34‰. Bulk nitrogen isotope compositions (2

Extraterrestrial environments encompass physicochemical conditions and habitats that are unknown on Earth, such as perchlorate-rich brines that can be at least temporarily stable on the martian surface. To better understand the potential for life in these cold briny environments, we determined the maximum salt concentrations suitable for growth (MSCg) of six different chloride and perchlorate salts at 25°C and 4°C for the extremotolerant cold- and salt-adapted bacterial strain Planococcus halocryophilus. Growth was measured through colony-forming unit (CFU) counts, while cellular and colonial phenotypic stress responses were observed through visible light, fluorescence, and scanning electron microscopy. Our data show the following: (1) The tolerance to high salt concentrations can be increased through a stepwise inoculation toward higher concentrations. (2) Ion-specific factors are more relevant for the growth limitation of P. halocryophilus in saline solutions than single physicochemical parameters like ionic strength or water activity. (3) P. halocryophilus shows the highest microbial sodium perchlorate tolerance described so far. However, (4) MSCg values are higher for all chlorides compared to perchlorates. (5) The MSCg for calcium chloride was increased by lowering the temperature from 25°C to 4°C, while sodium- and magnesium-containing salts can be tolerated at 25°C to higher concentrations than at 4°C. (6) Depending on salt type and concentration, P. halocryophilus cells show distinct phenotypic stress responses such as novel types of colony morphology on agar plates and biofilm-like cell clustering, encrustation, and development of intercellular nanofilaments. This study, taken in context with previous work on the survival of extremophiles in Mars-like environments, suggests that high-concentrated perchlorate brines on Mars might not be habitable to any present organism on Earth, but extremophilic microorganisms might be able to evolve thriving in such environments.

Mobile genetic elements (MGEs) play an essential role in bacterial adaptation and evolution. These elements are enriched within bacterial communities from extreme environments. However, very little is known if specific genes co-occur with MGEs in extreme environments and, if so, what their function is. We used shotgun-sequencing to analyse the metagenomes of 12 soil samples and characterized the composition of MGEs and the genes co-occurring with them. The samples ranged from less arid coastal sites to the inland hyperarid core of the Atacama Desert, as well as from sediments below boulders, protected from UV-irradiation. MGEs were enriched at the hyperarid sites compared with sediments from below boulders and less arid sites. MGEs were mostly co-occurring with genes belonging to the Cluster Orthologous Group (COG) categories &ldquo, replication, recombination and repair,&rdquo, &ldquo, transcription&rdquo, and &ldquo, signal transduction mechanisms.&rdquo, In general, genes coding for transcriptional regulators and histidine kinases were the most abundant genes proximal to MGEs. Genes involved in energy production were significantly enriched close to MGEs at the hyperarid sites. For example, dehydrogenases, reductases, hydrolases and chlorite dismutase and other enzymes linked to nitrogen metabolism such as nitrite- and nitro-reductase. Stress response genes, including genes involved in antimicrobial and heavy metal resistance genes, were rarely found near MGEs. The present study suggests that MGEs could play an essential role in the adaptation of the soil microbiome in hyperarid desert soils by the modulation of housekeeping genes such as those involved in energy production.

The accumulation of thousands of boulder-sized clasts into boulder fields in the Atacama Desert has been linked to seismic-driven downslope transport, a rare sedimentary process corroborated by this study. We surveyed boulder arrangements occurring in the Atacama Desert and identified three accumulation types for further investigation: a small circular boulder cluster (BC), a long channelized boulder stream (BS), and a wide convex-shaped boulder field (BF). Drone-based photogrammetric techniques and field observations were used to generate high-quality digital elevation models and orthophotos to determine boulder count, size, coverage, orientation, lithology and local topography. Our data shows that the arrangement of boulder accumulations corresponds with the shape of the accommodation space and the boulder input, where BCs occur at the center of completely confined topographic depressions, BSs occur along laterally confined and incised hill slopes with boulders stacked above each other, and BFs occur on largely unconfined shallow and low-relief slopes with a distinct boulder front. A general downslope increase of average boulder size and coverage was measured in all boulder accumulations and a long-axis orientation of boulders parallel to the transport direction was observed for the BS. Based on these results and the lack of fluvial transport indicators, we conclude that transport and arrangement of boulder accumulations are largely controlled by the interplay of topography and seismic-driven boulder transport, resulting in unique landscape features present in the hyper-arid Atacama Desert. Such sedimentary transport processes are rare on Earth but potentially play a greater role on other arid planetary surfaces that are covered by boulders and subject to sufficient seismic activity.

It is well known that dissolved salts can significantly lower the freezing point of water and thus extend habitability to subzero conditions. However, most investigations thus far have focused on sodium chloride as a solute. In this study, we report on the survivability of the bacterial strain Planococcus halocryophilus in sodium, magnesium, and calcium chloride or perchlorate solutions at temperatures ranging from +25°C to −30°C. In addition, we determined the survival rates of P. halocryophilus when subjected to multiple freeze/thaw cycles. We found that cells suspended in chloride-containing samples have markedly increased survival rates compared with those in perchlorate-containing samples. In both cases, the survival rates increase with lower temperatures; however, this effect is more pronounced in chloride-containing samples. Furthermore, we found that higher salt concentrations increase survival rates when cells are subjected to freeze/thaw cycles. Our findings have important implications not only for the habitability of cold environments on Earth but also for extraterrestrial environments such as that of Mars, where cold brines might exist in the subsurface and perhaps even appear temporarily at the surface such as at recurring slope lineae.It is well known that dissolved salts can significantly lower the freezing point of water and thus extend habitability to subzero conditions. However, most investigations thus far have focused on sodium chloride as a solute. In this study, we report on the survivability of the bacterial strain Planococcus halocryophilus in sodium, magnesium, and calcium chloride or perchlorate solutions at temperatures ranging from +25°C to −30°C. In addition, we determined the survival rates of P. halocryophilus when subjected to multiple freeze/thaw cycles. We found that cells suspended in chloride-containing samples have markedly increased survival rates compared with those in perchlorate-containing samples. In both cases, the survival rates increase with lower temperatures; however, this effect is more pronounced in chloride-containing samples. Furthermore, we found that higher salt concentrations increase survival rates when cells are subjected to freeze/thaw cycles. Our findings have important implications not only for the habitability of cold environments on Earth but also for extraterrestrial environments such as that of Mars, where cold brines might exist in the subsurface and perhaps even appear temporarily at the surface such as at recurring slope lineae.

Significance It has remained an unresolved question whether microorganisms recovered from the most arid environments on Earth are thriving under such extreme conditions or are just dead or dying vestiges of viable cells fortuitously deposited by atmospheric processes. Based on multiple lines of evidence, we show that indigenous microbial communities are present and temporally active even in the hyperarid soils of the Atacama Desert (Chile). Following extremely rare precipitation events in the driest parts of this desert, where rainfall often occurs only once per decade, we were able to detect episodic incidences of biological activity. Our findings expand the range of hyperarid environments temporarily habitable for terrestrial life, which by extension also applies to other planetary bodies like Mars., Traces of life are nearly ubiquitous on Earth. However, a central unresolved question is whether these traces always indicate an active microbial community or whether, in extreme environments, such as hyperarid deserts, they instead reflect just dormant or dead cells. Although microbial biomass and diversity decrease with increasing aridity in the Atacama Desert, we provide multiple lines of evidence for the presence of an at times metabolically active, microbial community in one of the driest places on Earth. We base this observation on four major lines of evidence: (i) a physico-chemical characterization of the soil habitability after an exceptional rain event, (ii) identified biomolecules indicative of potentially active cells [e.g., presence of ATP, phospholipid fatty acids (PLFAs), metabolites, and enzymatic activity], (iii) measurements of in situ replication rates of genomes of uncultivated bacteria reconstructed from selected samples, and (iv) microbial community patterns specific to soil parameters and depths. We infer that the microbial populations have undergone selection and adaptation in response to their specific soil microenvironment and in particular to the degree of aridity. Collectively, our results highlight that even the hyperarid Atacama Desert can provide a habitable environment for microorganisms that allows them to become metabolically active following an episodic increase in moisture and that once it decreases, so does the activity of the microbiota. These results have implications for the prospect of life on other planets such as Mars, which has transitioned from an earlier wetter environment to today’s extreme hyperaridity.

Cavities are considered plausible and favorable habitats for life on early Earth. In such microenvironments, organisms may have found an adequate protection against the intense ultraviolet radiation that characterized the Archean ozone-free atmosphere. However, while there is clear evidence that benthic life existed in the Paleoarchean, the oldest traces of cavity-dwelling microbes (coelobionts) have been found in Neoarchean rocks. Here we present the results of a detailed investigation of early silicified cavities occurring in the oldest well-preserved siliciclastic tidal deposits, the 3.22 Ga Moodies Group of the Barberton Greenstone Belt (South Africa). Downward-growing microstromatolitic columns composed of kerogenous laminae are commonly present in planar, bedding-parallel, now silica-filled cavities that formed in sediments of the peritidal zone. In-situ δ13CPDB (PDB—Peedee belemnite) measurements of the kerogen range from –32.3‰ to –21.3‰ and are consistent with a biogenic origin. Scanning electron microscopy analysis of the silicified cavities shows well-preserved chains of cell-sized molds that are interpreted as fossil filamentous microorganisms. The geological context, the morphology of the microstromatolites, the δ13C composition of the kerogen, and the presence of microfossils all suggest that a microbial community inhabited the cavities. These results extend the geological record of coelobionts by ∼500 m.y., supporting the view that cavities were among the first ecological niches to have been occupied by early microorganisms.

Microbial life was well established and widespread by the Paleoarchean; however, the degree of evolutionary advancement such as microbial motility, intra- and inter-species interactions, phototropism, or oxygenic photosynthesis by that time remains highly debated. The 3.22 Ga Moodies Group in the Barberton Greenstone Belt (BGB, South Africa) are Earth's oldest well-preserved siliciclastic tidal deposits. They exhibit a unique assemblage of microbial mats, providing an excellent opportunity to decipher the morphological adaptations of microbial communities to different paleoenvironmental settings. The fossil mats are preserved as kerogenous laminations (0.5–1 mm thick) that can be traced laterally for ∼15 km in a ∼1000 m-thick succession of fine- to coarse-grained tidal sandstones and conglomerates. We here present a detailed stratigraphic and depositional facies analysis, documenting the association of the three principal mat morphotypes with specific environmental settings: (1) planar-type in coastal floodplain, (2) wavy-type in intertidal, and (3) tufted-type in upper inter- to supratidal facies. All mat types indicate a flourishing phototrophic biota; moreover, the tufted morphology suggests an intricate level of coordinated growth commonly known from cyanobacterial mats in modern environments.

The evolutionary adaptability of life to extreme environments is astounding given that all life on Earth is based on the same fundamental biochemistry. The range of some physicochemical parameters on Earth exceeds the ability of life to adapt, but stays within the limits of life for other parameters. Certain environmental conditions such as low water availability in hyperarid deserts on Earth seem to be close to the limit of biological activity. A much wider range of environmental parameters is observed on planetary bodies within our Solar System such as Mars or Titan, and presumably even larger outside of our Solar System. Here we review the adaptability of life as we know it, especially regarding temperature, pressure, and water activity. We use then this knowledge to outline the range of possible habitable environments for alien planets and moons and distinguish between a variety of Planetary Environment Types (PETs). Some of these types are present in our Solar System, others are hypothetical. Our schematic categorization of alien habitats is limited to life as we know it, particularly regarding to the use of solvent (water) and energy source (light and chemical compounds).

The origin and evolution of life has undoubtedly had a major impact on the evolution of Earth's oceans and atmosphere. Recent studies have suggested that bioactivity may have had an even deeper impact and may have caused a change in the redox-state of the mantle and provided a path for the formation of continents. We here present a numerical model that assumes that bioactivity increases the continental weathering rate and that relates the sedimentation rate to the growth of continents and to the hydration of the mantle using elements of plate tectonics and mantle convection. The link between these factors is provided by assuming that an increase of the thickness of the sedimentary layer of low permeability on top of a subducting oceanic slab will reduce its dewatering upon subduction. This in turn leads to a greater availability of water in the source region of andesitic partial melt, resulting in an enhanced rate of continental crust production, and to an increased regassing rate of the mantle. The mantle in turn responds by reducing the mantle viscosity while increasing the convective circulation rate, degassing rate and plate speed. We use parameters that are observed for the present Earth and gauge uncertain parameters such that the present day continental surface area and mantle water concentration can be obtained. Our steady state results show two stable fixed points in a phase plane defined by the fractional continental surface area and the water concentration in the mantle, one of them pertaining to a wet mantle and the continental surface area of the present day Earth, and the other to a dry mantle and a small continental surface area. When the sedimentation rate is reduced, both fixed points move and the area of attraction of the latter fixed point increases systematically. We conclude that if the presence of life has increased the continental weathering rate, as is widely believed, and led to the observables of a wet mantle and a continental surface coverage of roughly 40%, an abiotic Earth would likely have evolved toward a dry mantle with a small continental surface area instead.

article Two Patagonian lakes studied here, Lago Sarmiento and Laguna Amarga, are located within the orographic rain shadow formed to the east of the Southern Patagonian Ice Field in the Andes Range. Major thrombolite colonies are present in Lago Sarmiento, whereas widespread stromatolites occur in Laguna Amarga. Based on the characterization of the hydrologic system of these two lakes, together with an estimation of the isotopic balance and an analysis of the equilibrium conditions between the water and biologically induced carbonates, it is concluded that the microbialites of Lago Sarmiento are better suited as paleotemperature indicators than those of Laguna Amarga. Lago Sarmiento thrombolites contain unique carbonate mineral species in which carbonate precipitation occurs close to isotopic equilibrium and where the variation in water temperature controls fractionation of the stable oxygen isotope. The results indicate that at 1215 cal yr Bp the level of the lake was at 85 m a.s.l with a temperature close to 9.3 °C, was at 82 m a.s.l. at 600 cal yr Bp with a temperature close to 8.5 °C. This coincides with the timing of the Northern Hemisphere Medieval Warming Period. At 183 cal yr Bp the level of the lake was at 80 m a.s.l with a cooler temperature close to 7.7 °C, representing a colder period coinciding with the timing of the Little Ice Age (LIA). An interesting outcome of this study is that it reinforces the idea that the δ 13 C signal in carbonate deposits can be an effective tool in distinguishing between inorganic and biologically induced precipitation.

In the version of this Article originally published, author Magali Ader was wrongly linked to affiliation 3; it should have been affiliation 2. This error has now been corrected in the online versions.

Nitrates are salts of the nitric acid (HNO3). It has been estimated that one quarter of the initial Martian atmospheric molecular nitrogen today is fixed in the form of nitrates in the soil. A possible formation process of these nitrates is the dissociation of the atmospheric molecular nitrogen through shock heating caused by impacts or through photochemical reactions (Smith et al. 2014). Nitrates are highly soluble in water, hence nitrates entrained in theMartian▶ cryosphere may contribute to the generation of brines near the surface. Because nitrates are a fundamental source of nitrogen for life on Earth, the detection of nitrates on Mars is an important parameter for the assessment of potential habitable environments. Recent analyses of “Rocknest” sand samples at Gale crater by the NASA rover “Curiosity” tentatively indicate the presence of nitrates.

On the basis of acquired thermotolerance and cryotolerance, the optimal heat shock and cold shock temperatures have been determined for Deinococcus radiodurans. A heat shock at 42 degrees C maximized survival at the lethal temperature of 52 degrees C and a cold shock at 20 degrees C maximized survival after repeated freeze-thawing. Enhanced survival from heat shock was found to be strongly dependent on growth stage, with its greatest effect shortly after phase. Increased synthesis of a total of 67 proteins during heat shock and 42 proteins during cold shock were observed by two-dimensional polyacrylamide gel electrophoresis (2D PAGE) and autoradiography. Eight of the most highly induced heat shock proteins shown by 2D PAGE were identified by MALDI-MS as Hsp20, GroEL, DnaK, SodA, Csp, Protease I, and two proteins of unknown function.

Some molecules, particularly aromatics, have high molar extinction coefficients at wavelengths in the damaging ultraviolet radiation region of the spectrum between 200 and 400 nm. Thus, under a UV radiation flux in which these wavelengths are represented, it could be argued that a selection pressure would exist for a UV transparent biochemistry in which they were not represented. This hypothesis is explored using data made available from proteomics, focusing particularly on tryptophan, against which a selection pressure could exist on present-day Earth as a result of its absorbance shoulder at wavelengths greater than 290 nm. The abundance of tryptophan in whole proteomes is lower than expected from the degeneracy of the genetic code. A lower usage of tryptophan is found in the cytochrome c oxidase polypeptide I of UV-exposed organisms compared to nocturnal and subterranean organisms, but not in ATP synthase chain A. Examination of the amino acid composition of photolyase, an enzyme that requires exposure to light to function, shows that the tryptophan abundances exceed those of the total proteome of most organisms and the abundances expected from the degeneracy of the genetic code. This is also true for cytochrome c oxidase, another enzyme that makes extensive use of the electron transfer properties of tryptophan. We suggest that the selection pressure for the use of tryptophan caused, among other factors, by the uses of delocalised pi-electrons that this aromatic provides in active sites and binding motifs outweighs the selection pressure for UV transparency. This trade-off explains the lack of conclusive evidence for a UV transparent selection pressure. We suggest that this trade-off applies to the stacked pi-electrons of DNA. It offers a solution to the long-standing paradox of why the macromolecule responsible for the faithful replication of information has high absorbance in the damaging UV radiation region of the spectrum.

Indexación: Web of Science; Scielo RESUMEN. Una serie de morrenas glaciares se distribuyen desde el margen oriental de la cuenca de drenaje Torres del Paine a cerca del actual margen de los Campos de Hielo Patagónico, junto con una serie de terrazas lacustres regional continuos relacionados con las fluctuaciones glaciales. La geomorfología, apoyada por la evidencia de sedimentos del lago, indica la existencia de un único paleolago proglacial en esta zona, aquí denominado el Gran Tehuelche paleolago. Este concepto ayuda a aclarar la cronología de eventos glaciales y conduce a una mejor comprensión de la evolución del sistema hidrológico de la zona de Torres del Paine. Avances glaciales referidos anteriormente como ocurrieron A, B y C durante el Último Máximo Glacial y alimentados con la Gran Tehuelche paleolago con agua de deshielo, que le permite alcanzar su máxima extensión. El descubrimiento de trombolitos en Laguna Amarga sugiere que el drenaje de la paleolago hacia el fiordo de Última Esperanza tuvo lugar en 7113 Cal. años AP, después de la fusión de una barrera de hielo que existía durante el avance glacial antes. Esto dio lugar al desarrollo de un sistema hidrológico fluvio-lacustre complejo que persiste hasta nuestros días. Palabras clave: Patagonia, último máximo glacial, Younger Dryas, trombolitos. RESUMEN. Un grupo de morrenas glaciales estan Distribuidas desde el margen Este de la cuenca de drenaje de Torres del Paine Hacia el margen real de los Campos de Hielo Patagónicos. Las morrenas se observan en conjunto con las Naciones Unidas Grupo de terrazas Lacustres Regionales, las Cuales estan vinculadas y una las fluctuaciones glaciales. La geomorfología y Evidencias de sedimentos Lacustres indicano la existence De unico ONU lago proglacial, referido En Este Estudio de Como Gran Paleolago Tehuelche. Este Concepto ayuda a clarificar la cronología de los eventos glaciales y permite Una mejor Comprensión de la Evolución del Sistema hidrológico del sector de Torres del Paine. Los eventos glaciales, previamente Referidos Como Avance A, B y C, ocurrieron Durante el Último Máximo Glacial y alimentaron con Aguas de fusión al Gran Paleolago Tehuelche, permitiéndole Alcanzar su alcalde extensión. El Descubrimiento de trombolitos en Laguna Amarga sugiere Que El desagüe del paleolago Ocurrió Hace 7.113 Cal. Por años AP el Seno de Última Esperanza, producto de la fusión De Una barrera glaciar existente Durante los Avances glaciales Anteriores. LUEGO del drenaje sí Desarrollo en Un complejo Sistema hidrológico Que Persiste Hasta el presente. Palabras clave: Patagonia, Último Máximo Glacial, Dryas Temprano, Trombolitos. http://ref.scielo.org/wgc2qs

Some molecules, particularly aromatics, have high molar extinction coefficients at wavelengths in the damaging ultraviolet radiation region of the spectrum between 200 and 400 nm. Thus, under a UV radiation flux in which these wavelengths are represented, it could be argued that a selection pressure would exist for a UV transparent biochemistry in which they were not represented. This hypothesis is explored using data made available from proteomics, focusing particularly on tryptophan, against which a selection pressure could exist on present-day Earth as a result of its absorbance shoulder at wavelengths greater than 290 nm. The abundance of tryptophan in whole proteomes is lower than expected from the degeneracy of the genetic code. A lower usage of tryptophan is found in the cytochrome c oxidase polypeptide I of UV-exposed organisms compared to nocturnal and subterranean organisms, but not in ATP synthase chain A. Examination of the amino acid composition of photolyase, an enzyme that requires exposure to light to function, shows that the tryptophan abundances exceed those of the total proteome of most organisms and the abundances expected from the degeneracy of the genetic code. This is also true for cytochrome c oxidase, another enzyme that makes extensive use of the electron transfer properties of tryptophan. We suggest that the selection pressure for the use of tryptophan caused, among other factors, by the uses of delocalised pi-electrons that this aromatic provides in active sites and binding motifs outweighs the selection pressure for UV transparency. This trade-off explains the lack of conclusive evidence for a UV transparent selection pressure. We suggest that this trade-off applies to the stacked pi-electrons of DNA. It offers a solution to the long-standing paradox of why the macromolecule responsible for the faithful replication of information has high absorbance in the damaging UV radiation region of the spectrum.