Many of the recently discovered extrasolar giant planets move around their stars on highly eccentric orbits, and some with e [ges ] 0·7. Systems with planets within or near the habitable zone (HZ) will possibly harbour life on terrestrial-type moons if the seasonal temperature extremes resulting from the large orbital eccentricities of the planets are not too severe. Here we use a three-dimensional general-circulation climate model and a one-dimensional energy-balance model to examine the climates of either bound or isolated earths on extremely elliptical orbits near the HZ. While such worlds are susceptible to large variations in surface temperature, long-term climate stability depends primarily on the average stellar flux received over an entire orbit, not the length of the time spent within the HZ.

The search for life in the Universe is a fundamental problem of astrobiology and modern science. The current progress in the detection of terrestrial-type exoplanets has opened a new avenue in the characterization of exoplanetary atmospheres and in the search for biosignatures of life with the upcoming ground-based and space missions. To specify the conditions favourable for the origin, development and sustainment of life as we know it in other worlds, we need to understand the nature of global (astrospheric), and local (atmospheric and surface) environments of exoplanets in the habitable zones (HZs) around G-K-M dwarf stars including our young Sun. Global environment is formed by propagated disturbances from the planet-hosting stars in the form of stellar flares, coronal mass ejections, energetic particles and winds collectively known as astrospheric space weather. Its characterization will help in understanding how an exoplanetary ecosystem interacts with its host star, as well as in the specification of the physical, chemical and biochemical conditions that can create favourable and/or detrimental conditions for planetary climate and habitability along with evolution of planetary internal dynamics over geological timescales. A key linkage of (astro)physical, chemical and geological processes can only be understood in the framework of interdisciplinary studies with the incorporation of progress in heliophysics, astrophysics, planetary and Earth sciences. The assessment of the impacts of host stars on the climate and habitability of terrestrial (exo)planets will significantly expand the current definition of the HZ to the biogenic zone and provide new observational strategies for searching for signatures of life. The major goal of this paper is to describe and discuss the current status and recent progress in this interdisciplinary field in light of presentations and discussions during the NASA Nexus for Exoplanetary System Science funded workshop ‘Exoplanetary Space Weather, Climate and Habitability’ and to provide a new roadmap for the future development of the emerging field of exoplanetary science and astrobiology.

Gamma-ray bursts (GRBs) produce a flux of radiation detectable across the observable Universe. A GRB within our own galaxy could do considerable damage to the Earth's biosphere; rate estimates suggest that a dangerously near GRB should occur on average two or more times per billion years. At least five times in the history of life, the Earth has experienced mass extinctions that eliminated a large percentage of the biota. Many possible causes have been documented, and GRBs may also have contributed. The late Ordovician mass extinction approximately 440 million years ago may be at least partly the result of a GRB. A special feature of GRBs in terms of terrestrial effects is a nearly impulsive energy input of the order of 10 s. Due to expected severe depletion of the ozone layer, intense solar ultraviolet radiation would result from a nearby GRB, and some of the patterns of extinction and survivorship at this time may be attributable to elevated levels of UV radiation reaching the Earth. In addition, a GRB could trigger the global cooling which occurs at the end of the Ordovician period that follows an interval of relatively warm climate. Intense rapid cooling and glaciation at that time, previously identified as the probable cause of this mass extinction, may have resulted from a GRB.

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.

Despite the strong increase in observational data on extrasolar planets, the processes that led to the formation of these planets are still not well understood. However, thanks to the high number of extrasolar planets that have been discovered, it is now possible to look at the planets as a population that puts statistical constraints on theoretical formation models. A method that uses these constraints is planetary population synthesis where synthetic planetary populations are generated and compared to the actual population. The key element of the population synthesis method is a global model of planet formation and evolution. These models directly predict observable planetary properties based on properties of the natal protoplanetary disc, linking two important classes of astrophysical objects. To do so, global models build on the simplified results of many specialized models that address one specific physical mechanism. We thoroughly review the physics of the sub-models included in global formation models. The sub-models can be classified as models describing the protoplanetary disc (of gas and solids), those that describe one (proto)planet (its solid core, gaseous envelope and atmosphere), and finally those that describe the interactions (orbital migration and N-body interaction). We compare the approaches taken in different global models, discuss the links between specialized and global models, and identify physical processes that require improved descriptions in future work. We then shortly address important results of planetary population synthesis like the planetary mass function or the mass–radius relationship. With these statistical results, the global effects of physical mechanisms occurring during planet formation and evolution become apparent, and specialized models describing them can be put to the observational test. Owing to their nature as meta models, global models depend on the results of specialized models, and therefore on the development of the field of planet formation theory as a whole. Because there are important uncertainties in this theory, it is likely that the global models will in future undergo significant modifications. Despite these limitations, global models can already now yield many testable predictions. With future global models addressing the geophysical characteristics of the synthetic planets, it should eventually become possible to make predictions about the habitability of planets based on their formation and evolution.

Terrestrial analogue studies underpin almost all planetary missions and their use is essential in the exploration of our Solar system and in assessing the habitability of other worlds. Their value relies on the similarity of the analogue to its target, either in terms of their mineralogical or geochemical context, or current physical or chemical environmental conditions. Such analogue sites offer critical ground-truthing for astrobiological studies on the habitability of different environmental parameter sets, the biological mechanisms for survival in extreme environments and the preservation potential and detectability of biosignatures. The 33 analogue sites discussed in this review have been selected on the basis of their congruence to particular extraterrestrial locations. Terrestrial field sites that have been used most often in the literature, as well as some lesser known ones which require greater study, are incorporated to inform on the astrobiological potential of Venus, Mars, Europa, Enceladus and Titan. For example, the possibility of an aerial habitable zone on Venus has been hypothesized based on studies of life at high-altitudes in the terrestrial atmosphere. We also demonstrate why many different terrestrial analogue sites are required to satisfactorily assess the habitability of the changing environmental conditions throughout Martian history, and recommend particular sites for different epochs or potential niches. Finally, habitable zones within the aqueous environments of the icy moons of Europa and Enceladus and potentially in the hydrocarbon lakes of Titan are discussed and suitable analogue sites proposed. It is clear from this review that a number of terrestrial analogue sites can be applied to multiple planetary bodies, thereby increasing their value for astrobiological exploration. For each analogue site considered here, we summarize the pertinent physiochemical environmental features they offer and critically assess the fidelity with which they emulate their intended target locale. We also outline key issues associated with the existing documentation of analogue research and the constraints this has on the efficiency of discoveries in this field. This review thus highlights the need for a global open access database for planetary analogues.

Perchlorate (ClO4−) is widespread in Martian soils at concentrations between 0.5 and 1%. At such concentrations, perchlorate could be an important source of oxygen, but it could also become a critical chemical hazard to astronauts. In this paper, we review the dual implications of ClO4− on Mars, and propose a biochemical approach for removal of perchlorate from Martian soil that would be energetically cheap, environmentally friendly and could be used to obtain oxygen both for human consumption and to fuel surface operations.

Life on Earth has existed for at least 3.5 billion years. Yet, relatively little is known of its evolution during the first two billion years, due to the scarceness and generally poor preservation of fossilized biological material. Cyanobacteria, formerly known as blue green algae were among the first crown Eubacteria to evolve and for more than 2.5 billion years they have strongly influenced Earth's biosphere. Being the only organism where oxygenic photosynthesis has originated, they have oxygenated Earth's atmosphere and hydrosphere, triggered the evolution of plants --being ancestral to chloroplasts-- and enabled the evolution of complex life based on aerobic respiration. Having such a strong impact on early life, one might expect that the evolutionary success of this group may also have triggered further biosphere changes during early Earth history. However, very little is known about the early evolution of this phylum and ongoing debates about cyanobacterial fossils, biomarkers and molecular clock analyses highlight the difficulties in this field of research. Although phylogenomic analyses have provided promising glimpses into the early evolution of cyanobacteria, estimated divergence ages are often very uncertain, because of vague and insufficient tree-calibrations. Results of molecular clock analyses are intrinsically tied to these prior calibration points, hence improving calibrations will enable more precise divergence time estimations. Here we provide a review of previously described Precambrian microfossils, biomarkers and geochemical markers that inform upon the early evolution of cyanobacteria. Future research in micropalaeontology will require novel analyses and imaging techniques to improve taxonomic affiliation of many Precambrian microfossils. Consequently, a better understanding of early cyanobacterial evolution will not only allow for a more specific calibration of cyanobacterial and eubacterial phylogenies, but also provide new dates for the tree of life.

Living organisms on Earth are characterized by three necessary features: a set of internal instructions encoded in DNA (software), a suite of proteins and associated macromolecules providing a boundary and internal structure (hardware), and a flux of energy. In addition, they replicate themselves through reproduction, a process that renders evolutionary change inevitable in a resource-limited world. Temperature has a profound effect on all of these features, and yet life is sufficiently adaptable to be found almost everywhere water is liquid. The thermal limits to survival are well documented for many types of organisms, but the thermal limits to completion of the life cycle are much more difficult to establish, especially for organisms that inhabit thermally variable environments. Current data suggest that the thermal limits to completion of the life cycle differ between the three major domains of life, bacteria, archaea and eukaryotes. At the very highest temperatures only archaea are found with the current high-temperature limit for growth being 122 °C. Bacteria can grow up to 100 °C, but no eukaryote appears to be able to complete its life cycle above ∼60 °C and most not above 40 °C. The lower thermal limit for growth in bacteria, archaea, unicellular eukaryotes where ice is present appears to be set by vitrification of the cell interior, and lies at ∼−20 °C. Lichens appear to be able to grow down to ∼−10 °C. Higher plants and invertebrates living at high latitudes can survive down to ∼−70 °C, but the lower limit for completion of the life cycle in multicellular organisms appears to be ∼−2 °C.

We quantify the scientific potential for exoplanet imaging with the mid-infrared E-ELT Imager and Spectrograph (METIS) foreseen as one of the instruments of the European Extremely Large Telescope (E-ELT). We focus on two main science cases: (1) the direct detection of known gas giant planets found by radial velocity (RV) searches; and (2) the direct detection of small (1–4 R⊕) planets around the nearest stars. Under the assumptions made in our modelling, in particular on the achievable inner working angle and sensitivity, our analyses reveal that within a reasonable amount of observing time METIS is able to image >20 already known, RV-detected planets in at least one filter. Many more suitable planets with dynamically determined masses are expected to be found in the coming years with the continuation of RV-surveys and the results from the GAIA astrometry mission. In addition, by extrapolating the statistics for close-in planets found by Kepler, we expect METIS might detect ≈10 small planets with equilibrium temperatures between 200 and 500 K around the nearest stars. This means that (1) METIS will help constrain atmospheric models for gas giant planets by determining for a sizable sample their luminosity, temperature and orbital inclination; and (2) METIS might be the first instrument to image a nearby (super-) Earth-sized planet with an equilibrium temperature near that expected to enable liquid water on a planet surface.

The Biological Universe (Dick 1996) analysed the history of the extraterrestrial life debate, documenting how scientists have assessed the chances of life beyond Earth during the 20th century. Here I propose another option – that we may in fact live in a postbiological universe, one that has evolved beyond flesh and blood intelligence to artificial intelligence that is a product of cultural rather than biological evolution. MacGowan & Ordway (1966), Davies (1995) and Shostak (1998), among others, have broached the subject, but the argument has not been given the attention it is due, nor has it been carried to its logical conclusion. This paper argues for the necessity of long-term thinking when contemplating the problem of intelligence in the universe. It provides arguments for a postbiological universe, based on the likely age and lifetimes of technological civilizations and the overriding importance of cultural evolution as an element of cosmic evolution. And it describes the general nature of a postbiological universe and its implications for the search for extraterrestrial intelligence.

We have used 16S ribosomal genes to estimate the bacterial contents of Lake Vostok accretion ice samples at 3551 m and 3607 m, both containing sediment inclusions and formed 20000–15000 yr ago. Decontamination proved to be a critical issue, and we used stringent ice chemistry-based procedures and comprehensive biological controls in order to restrain contamination. As a result, up to now we have only recognized one 16S rDNA bacterial phylotype with confident relevance to the lake environment. It was found in one sample at 3607 m depth and represents the extant thermophilic facultative chemolithoautotroph Hydrogenophilus thermoluteolus of beta-Proteobacteria, and until now had only been found in hot springs. No confident findings were detected in the sample at 3551 m, and all other phylotypes revealed (a total of 16 phylotypes, 336 clones including controls) are presumed to be contaminants. It seems that the Lake Vostok accretion ice is actually microbe-free, indicating that the water body should also be hosting a highly sparse life. The message of thermophilic bacteria suggests that a geothermal system exists beneath the cold water body of Lake Vostok, what is supported by the geological setting, the long-term seismotectonic evidence from 4He degassing and the ‘18O shift’ of the Vostok accretion ice. The seismotectonic activity that seems to operate in deep faults beneath the lake could sustain thermophilic chemolithoautotrophic microbial communities. Such a primary production scenario for Lake Vostok may have relevance for icy planets and the approaches used for estimating microbial contents in accretion ice are clearly relevant for searching for extraterrestrial life.

A three-dimensional general-circulation climate model is used to simulate climates of Earth-like planets with extreme axial tilts (i.e. ‘obliquities’). While no terrestrial-planet analogue exists in the solar system, planets with steeply inclined spin axes may be common around nearby stars. Here we report the results of 12 numerical experiments with Earth-like planets having different obliquities (from 0° to 85°), continental geographies, and levels of the important greenhouse gas, CO2. Our simulations show intense seasonality in surface temperatures for obliquities [ges ]54°, with temperatures reaching 80–100 °C over the largest middle- and high-latitude continents around the summer solstice. Net annual warming at high latitudes is countered by reduced insolation and colder temperatures in the tropics, which maintains the global annual mean temperature of our planets to within a few degrees of 14 °C. Under reduced insolation, seasonal snow covers some land areas near the equator; however no significant net annual accumulation of snow or ice occurs in any of our runs with obliquity exceeding the present value, in contrast to some previous studies. None of our simulated planets were warm enough to develop a runaway greenhouse or cold enough to freeze over completely; therefore, most real Earth-like planets should be hospitable to life at high obliquity.

Solar radiation is the primary energy source for surface planetary life, so that pigments are fundamental components of any surface-dwelling organism. They may therefore have evolved in some form on Mars as they did on Earth. Photosynthetic microbes are major primary producers on Earth, but are concurrently vulnerable to ultraviolet (UV) damage. Using non-intrusive laser Raman spectroscopy to recognize the component parts of biomolecules, we have shown not only the abundance of microbial photosynthetic and photoprotective pigments in situ, but also their spatial distribution within their microhabitat. This essential aspect of their screening or avoidance survival strategies is lost on extraction with solvents. This precise approach is eminently suited to analysis of epilithic (surface) and endolithic (within rocks) communities in Antarctic desert habitats, which are putative analogues of early Mars. Raman spectra for key biomolecules (e.g. the UV screen parietin and the antioxidant β-carotene in epilithic lichens) enable not only the detection of organics in light-stratified habitats, but also the characterization of unknown pigments. Typical biomarkers of astrobiological relevance in our Raman spectral database include scytonemin (a UV screen), chlorophyll (primary photosynthetic pigment), phycocyanin (accessory pigment for shade adaptation) and a hopanoid extracted from 2·5 Gya microbial stromatolite from Australia. This compound dates from the same time period when a wetter Mars could have had a potentially flourishing surface microbial community of its own. Analyses with a laboratory Raman instrument have been extended to a novel miniature Raman spectrometer, operating at the same optimal excitation wavelength (1064 nm) via an In-Ga-As detector. After evaluation in Antarctica, this instrument will be space-qualified for a proposed Mars rover mission to detect biomolecules in the near-surface sediment profile of palaeolakes, using experience with Antarctic biomarkers to interpret alien spectra of fundamental components, without the need for prior knowledge of the identity of the target compounds.

We report the first in situ observation of hollow organic globules in any extraterrestrial material using the Tagish Lake carbonaceous chondrite. Data from analytical transmission electron microscopy, Raman and micro-Fourier-transform infrared (FTIR) spectroscopy indicate that the globules consist of aliphatic and oxygenated functions. The hollow spherical morphologies are strikingly similar to the material produced by the laboratory simulation of ultraviolet photolysis of interstellar ice analogues and subsequent aqueous processing, suggesting that the organic globules in the Tagish Lake meteorite may be extremely primitive organic material that formed before or during the formation of the solar system. The FTIR organic signatures also show strong similarities to the membrane-like products formed from hydrothermal reaction of an OH-bearing amino acid in the presence of hydrous minerals. The survival of the structures in the Tagish Lake sample indicates that primitive meteorites must have delivered these structures to the early Earth as a possible precursor to life.

Various Mars missions have detected Cl atoms, chlorides and perchlorates in martian surface materials. The global soils, in particular, always contain significant levels of observable Cl. Direct evidence points to this Cl being in the form of both chlorides and perchlorates, and possibly also chlorates and other oxychlorines. The most widespread measurements have been of Cl atoms, and cannot discern the chemical form. However, from separate evidence of perchlorate obtained at high latitudes (Phoenix lander) and low latitudes (Curiosity rover), it is likely that perchlorates are widespread, albeit in varying proportions relative to the total amount of ubiquitous Cl.

A stellar evolution computer model has been used to determine changes in the luminosity L and effective temperature Te of single stars during their time on the main sequence. The range of stellar masses investigated was from 0.5 to 1.5 times that of the Sun, each with a mass fraction of metals (metallicity, Z) from 0.008 to 0.05. The extent of each star's habitable zone (HZ) has been determined from its values of L and Te. These stars form a reference framework for other main sequence stars. All of the 104 main sequence stars known to have one or more giant planets have been matched to their nearest stellar counterpart in the framework, in terms of mass and metallicity, hence closely approximating their HZ limits. The limits of HZ, for each of these stars, have been compared to their giant planet(s)'s range of strong gravitational influence. This allows a quick assessment as to whether Earth-mass planets could exist in stable orbits within the HZ of such systems, both presently and at any time during the star's main sequence lifetime. A determination can also be made as to the possible existence of life-bearing satellites of giant planets, which orbit within HZs. Results show that about half of the 104 known extrasolar planetary systems could possibly have been housing an Earth-mass planet in HZs during at least the past billion years, and about three-quarters of the 104 could do so for at least a billion years at some time during their main sequence lives. Whether such Earth-mass planets could have formed is an urgent question now being investigated by others, with encouraging results.

Gale crater formed from an impact on Mars ∼3.6 billion years ago. It hosts a central mound nearly 100 km wide and ∼5 km high, consisting of layered rocks with a variety of textures and spectral properties. The oldest exposed layers contain variably hydrated sulphates and smectite clay minerals, implying an aqueous origin, whereas the younger layers higher on the mound are covered by a mantle of dust. Fluvial channels carved into the crater walls and the lower mound indicate that surface liquids were present during and after deposition of the mound material. Numerous hypotheses have been advocated for the origin of some or all minerals and layers in the mound, ranging from deep lakes to playas to mostly dry dune fields to airfall dust or ash subjected to only minor alteration driven by snowmelt. The complexity of the mound suggests that multiple depositional and diagenetic processes are represented in the materials exposed today. Beginning in August 2012, the Mars Science Laboratory rover Curiosity will explore Gale crater by ascending the mound's northwestern flank, providing unprecedented new detail on the evolution of environmental conditions and habitability over many millions of years during which the mound strata accumulated.

Cold environments are common throughout the Galaxy. We are conducting a series of experiments designed to probe the low-temperature limits for growth in selected methanogenic and halophilic Archaea. This paper presents initial results for two mesophiles, a methanogen, Methanosarcina acetivorans, and a halophile, Halobacterium sp. NRC-1, and for two Antarctic cold-adapted Archaea, a methanogen, Methanococcoides burtonii, and a halophile, Halorubrum lacusprofundi. Neither mesophile is active at temperatures below 5 °C, but both cold-adapted microorganisms show significant growth at sub-zero temperatures (−2 °C and −1 °C, respectively), extending previous low-temperature limits for both species by 4–5 °C. At low temperatures, both H. lacusprofundi and M. burtonii form multicellular aggregates, which appear to be embedded in extracellular polymeric substances. This is the first detection of this phenomenon in Antarctic species of Archaea at cold temperatures. The low-temperature limits for both psychrophilic species fall within the temperature range experienced on present-day Mars and could permit survival and growth, particularly in sub-surface environments. We also discuss the results of our experiments in the context of known exoplanet systems, several of which include planets that intersect the Habitable Zone. In most cases, those planets follow orbits with significant eccentricity, leading to substantial temperature excursions. However, a handful of the known gas giant exoplanets could potentially harbour habitable terrestrial moons.

While early eukaryotic life must have been unicellular, multicellular lifeforms evolved multiple times from protistan ancestors in diverse eukaryotic lineages. The origins of multicellularity are of special interest because they require evolutionary transitions towards increased levels of complexity. We have generated new sequence data from the nuclear large subunit ribosomal DNA gene (LSU rDNA) and the SSU rDNA gene of several unicellular opisthokont protists – a nucleariid amoeba (Nuclearia simplex) and four choanoflagellates (Codosiga gracilis, Choanoeca perplexa, Proterospongia choanojuncta and Stephanoeca diplocostata) to provide the basis for re-examining relationships among several unicellular lineages and their multicellular relatives (animals and fungi). Our data indicate that: (1) choanoflagellates are a monophyletic rather than a paraphyletic assemblage that independently gave rise to animals and fungi as suggested by some authors and (2) the nucleariid filose amoebae are the likely sister group to Fungi. We also review published information regarding the origin of multicellularity in the opisthokonts.