Habitability Is Binary, But It Is Used by Astrobiologists to Encompass Continuous Ecological Questions.

The term "habitability" is pervasive throughout the space sciences and astrobiology literature and is broadly used to describe an environment's ability to support life. Here, we argue that, while it is fundamentally a binary matter whether an organism can persist in an environment or not, these binary assessments lead to continuous ecological measurements that are often collected under the umbrella term "habitability" by astrobiologists. Although the use of habitability in this way has provided a framework for those studying the potential of environments to support life, including comparative analyses between terrestrial and extraterrestrial environments, it can also generate confusion and limit interdisciplinary understanding. Namely, differing ecological metrics used as proxies for habitability can yield differing conclusions depending upon the metrics chosen. Therefore, we suggest that in this continuous sense, the terms habitable and habitability lose meaning unless the specific scientific question and biological metric chosen to address it are defined. As a corollary, the search for universal single metrics to make habitability assessments is not to be encouraged, and as we argue, attempting to do so would oversimply analyses of the ability of environments to support life.

The Detection of Elemental Signatures of Microbes in Martian Mudstone Analogs Using High Spatial Resolution Laser Ablation Ionization Mass Spectrometry.

The detection and identification of biosignatures on planetary bodies such as Mars in situ is extremely challenging. Current knowledge from space exploration missions suggests that a suite of complementary instruments is required in situ for a successful identification of past or present life. For future exploration missions, new and innovative instrumentation capable of high spatial resolution chemical (elemental and isotope) analysis of solids with improved measurement capabilities is of considerable interest because a multitude of potential signatures of extinct or extant life have dimensions on the micrometer scale. The aim of this study is to extend the current measurement capabilities of a miniature laser ablation ionization mass spectrometer (LIMS) designed for space exploration missions to detect signatures of microbial life. In total, 14 martian mudstone analogue samples were investigated regarding their elemental composition. Half the samples were artificially inoculated with a low number density of microbes, and half were used as abiotic controls. The samples were treated in a number of ways. Some were cultured anaerobically and some aerobically; some abiotic samples were incubated with water, and some remained dry. Some of the samples were exposed to a large dose of γ radiation, and some were left un-irradiated. While no significant elemental differences were observed between the applied sample treatments, the instrument showed the capability to detect biogenic element signatures of the inoculated microbes by monitoring biologically relevant elements, such as hydrogen, carbon, sulfur, iron, and so on. When an enrichment in carbon was measured in the samples but no simultaneous increase in other biologically relevant elements was detected, it suggests, for example, a carbon-containing inclusion; when the enrichment was in carbon and in bio-relevant elements, it suggests the presences of microbes. This study presents first results on the detection of biogenic element patterns of microbial life using a miniature LIMS system designed for space exploration missions.

A Systematic Study of the Limits of Life in Mixed Ion Solutions: Physicochemical Parameters Do Not Predict Habitability.

This study investigated what defines the limits of life in mixed ion solutions. Better understanding these limits should allow us to better predict the habitability of extreme environments on the Earth and extraterrestrial environments. We systematically examined the response of Bacillus subtilis, a well characterized non-halophile model organism, to a range of solutions made from single and mixed salts up to their solubility limits and measured at what concentration growth was arrested, specifically exploring Na, Mg, and Ca cations and Cl, SO4, and ClO4 anions. We measured the physicochemical properties of the solutions to identify which properties correlated with the limits of growth. Individual salts imposed a growth limit specific to the combination of cation and anion, although we generally observe that chloride salts allow growth at lower water activity than sulfate salts, with perchlorate restricting growth even at the highest measured water activity. Growth was limited at a wide range of ionic strength, with no apparently correlation. Despite the theoretically counteracting disordering effects (chaotropic) of perchlorates and ordering effects (kosmotropic) effects of sulfates, when these salts were combined they instead additively narrowed the window for growth in both the Na and Mg cation systems, in the same manner as the combined effects of two chaotropic Ca salts. Our results imply that away from hard limits that might be imposed by physicochemical properties such as water activity, ionic strength or chaotropicity in highly concentrated brines, these properties do not set the limits of life. Instead these limits are highly specific to the salts and organisms in question. This specificity means that the habitability of extreme environments cannot be predicted, even with accurate measurements of the physicochemical conditions present.

Detectability of biosignatures in a low-biomass simulation of martian sediments.

Discovery of a remnant habitable environment by the Mars Science Laboratory in the sedimentary record of Gale Crater has reinvigorated the search for evidence of martian life. In this study, we used a simulated martian mudstone material, based on data from Gale Crater, that was inoculated and cultured over several months and then dried and pressed. The simulated mudstone was analysed with a range of techniques to investigate the detectability of biosignatures. Cell counting and DNA extraction showed a diverse but low biomass microbial community that was highly dispersed. Pellets were analysed with bulk Elemental Analysis - Isotope Ratio Mass Spectrometry (EA-IRMS), high-resolution Laser-ablation Ionisation Mass Spectrometry (LIMS), Raman spectroscopy and Fourier Transform InfraRed (FTIR) spectroscopy, which are all techniques of relevance to current and future space missions. Bulk analytical techniques were unable to differentiate between inoculated samples and abiotic controls, despite total levels of organic carbon comparable with that of the martian surface. Raman spectroscopy, FTIR spectroscopy and LIMS, which are high sensitivity techniques that provide chemical information at high spatial resolution, retrieved presumptive biosignatures but these remained ambiguous and the sedimentary matrix presented challenges for all techniques. This suggests challenges for detecting definitive evidence for life, both in the simulated lacustrine environment via standard microbiological techniques and in the simulated mudstone via analytical techniques with relevance to robotic missions. Our study suggests that multiple co-incident high-sensitivity techniques that can scan the same micrometre-scale spots are required to unambiguously detect biosignatures, but the spatial coverage of these techniques needs to be high enough not to miss individual cellular-scale structures in the matrix.

Using Science-Driven Analog Research to Investigate Extravehicular Activity Science Operations Concepts and Capabilities for Human Planetary Exploration.

Biologic Analog Science Associated with Lava Terrains (BASALT) is a science-driven exploration program seeking to determine the best tools, techniques, training requirements, and execution strategies for conducting Mars-relevant field science under spaceflight mission conditions. BASALT encompasses Science, Science Operations, and Technology objectives. This article outlines the BASALT Science Operations background, strategic research questions, study design, and a portion of the results from the second field test. BASALT field tests are used to iteratively develop, integrate, test, evaluate, and refine new concepts of operations (ConOps) and capabilities that enable efficient and productive science. This article highlights the ConOps investigated during BASALT in light of future planetary extravehicular activity (EVA), which will focus on scientific exploration and discovery, and serves as an introduction to integrating exploration flexibility with operational rigor, the value of tactical and strategic science planning and execution, and capabilities that enable and enhance future science EVA operations.

Strategic Planning Insights for Future Science-Driven Extravehicular Activity on Mars.

Short-term and long-term science plans were developed as part of the strategic planning process used by the Biologic Analog Science Associated with Lava Terrains (BASALT) science team to conduct two Mars-simulation missions investigating basalt habitability at terrestrial volcanic analog sites in 2016. A multidisciplinary team of scientists generated and codified a range of scientific hypotheses distilled into a Science Traceability Matrix (STM) that defined the set of objectives pursued in a series of extravehicular activity (EVA) campaigns performed across multiple field deployments. This STM was used to guide the pre-deployment selection of sampling stations within the selected Mars analog sites on the Earth based on precursor site information such as multispectral imagery. It also informed selection of hand-held instruments and observational data to collect during EVA to aid sample selection through latency-impacted interaction with an Earth-based Science Support Team. A significant portion of the pre-deployment strategic planning activities were devoted to station selection, ultimately the locations used for sample collection and EVA planning. During development of the EVAs, the BASALT science team identified lessons learned that could be used to inform future missions and analog activities, including the critical need for high-resolution precursor imagery that would enable the selection of stations that could meet the scientific objectives outlined in the STM.

A Low-Diversity Microbiota Inhabits Extreme Terrestrial Basaltic Terrains and Their Fumaroles: Implications for the Exploration of Mars.

A major objective in the exploration of Mars is to test the hypothesis that the planet hosted life. Even in the absence of life, the mapping of habitable and uninhabitable environments is an essential task in developing a complete understanding of the geological and aqueous history of Mars and, as a consequence, understanding what factors caused Earth to take a different trajectory of biological potential. We carried out the aseptic collection of samples and comparison of the bacterial and archaeal communities associated with basaltic fumaroles and rocks of varying weathering states in Hawai'i to test four hypotheses concerning the diversity of life in these environments. Using high-throughput sequencing, we found that all these materials are inhabited by a low-diversity biota. Multivariate analyses of bacterial community data showed a clear separation between sites that have active fumaroles and other sites that comprised relict fumaroles, unaltered, and syn-emplacement basalts. Contrary to our hypothesis that high water flow environments, such as fumaroles with active mineral leaching, would be sites of high biological diversity, alpha diversity was lower in active fumaroles compared to relict or nonfumarolic sites, potentially due to high-temperature constraints on microbial diversity in fumarolic sites. A comparison of these data with communities inhabiting unaltered and weathered basaltic rocks in Idaho suggests that bacterial taxon composition of basaltic materials varies between sites, although the archaeal communities were similar in Hawai'i and Idaho. The taxa present in both sites suggest that most of them obtain organic carbon compounds from the atmosphere and from phototrophs and that some of them, including archaeal taxa, cycle fixed nitrogen. The low diversity shows that, on Earth, extreme basaltic terrains are environments on the edge of sustaining life with implications for the biological potential of similar environments on Mars and their exploration by robots and humans.

Opportunities and Challenges of Promoting Scientific Dialog throughout Execution of Future Science-Driven Extravehicular Activity.

Science-driven, human spaceflight missions of the future will rely on regular and interactive communication between Earth- and space-based teams during activity in which astronauts work directly on Mars or other planetary surfaces (extravehicular activity, EVA). The Biologic Analog Science Associated with Lava Terrains (BASALT) project conducted simulated human missions to Mars, complete with realistic one-way light time (OWLT) communication latency. We discuss the modes of communication used by the Mars- and Earth-based teams, including text, audio, video, and still imagery. Real-time communication between astronauts in the field (extravehicular, EV) and astronauts in a communication relay station (intravehicular, IV) was broadcast over OWLT, providing important contextual information to the Science Backroom Team (SBT) in Mission Control. Collaborative communication between the Earth- and Mars-based teams, however, requires active communication across latency via the Mission Log. We provide descriptive statistics of text communication between IV and SBT in a high-fidelity, scientifically driven analog for human space exploration. Over an EVA, the SBT sent an average of ∼23 text messages containing recommendations, requests, and answers to questions, while the science-focused IV crew member (IV2) sent an average of ∼38 text messages. Though patterns varied, communication between the IV and SBT teams tended to be highest during ∼50-150 min into the EVA, corresponding to the candidate sample search and presampling instrument survey phases, and then decreased dramatically after minute ∼200 during the sample collection phase. Generally, the IV2 and SBT used ∼4.6 min to craft a reply to a direct question or comment, regardless of message length or OWLT, offering a valuable glimpse into actual time-to-reply. We discuss IV2-SBT communication within the context of case examples from an EVA during which communication failures affected operations in the field. Finally, we offer recommendations for communication practices for use in future analogs and, perhaps, science-driven human spaceflight.

The BASALT Research Program: Designing and Developing Mission Elements in Support of Human Scientific Exploration of Mars.

The articles associated with this Special Collection focus on the NASA BASALT (Biologic Analog Science Associated with Lava Terrains) Research Program, which aims at answering the question, "How do we support and enable scientific exploration during human Mars missions?" To answer this the BASALT team conducted scientific field studies under simulated Mars mission conditions to both broaden our understanding of the habitability potential of basalt-rich terrains on Mars and examine the effects of science on current Mars mission concepts of operations. This article provides an overview of the BASALT research project, from the science, to the operational concepts that were tested and developed, to the technical capabilities that supported all elements of the team's research. Further, this article introduces the 12 articles that are included in this Special Collection.

Basaltic Terrains in Idaho and Hawai'i as Planetary Analogs for Mars Geology and Astrobiology.

Field research target regions within two basaltic geologic provinces are described as Earth analogs to Mars. Regions within the eastern Snake River Plain of Idaho and the Big Island of Hawai'i, the United States, provinces that represent analogs of present-day and early Mars, respectively, were evaluated on the basis of geologic settings, rock lithology and geochemistry, rock alteration, and climate. Each of these factors provides rationale for the selection of specific targets for field research in five analog target regions: (1) Big Craters and (2) Highway lava flows at Craters of the Moon National Monument and Preserve, Idaho, and (3) Mauna Ulu low shield, (4) Kilauea Iki lava lake, and (5) Kilauea caldera in the Kilauea Volcano summit region and the East Rift Zone of Hawai'i. Our evaluation of compositional and textural attributes, as well as the effects of syn- and posteruptive rock alteration, shows that basaltic terrains in Idaho and Hawai'i provide a way to characterize the geology and major geologic substrates that host biological activity of relevance to Mars exploration. This work provides the foundation to better understand the scientific questions related to the habitability of basaltic terrains, the rationale behind selecting analog field targets, and their applicability as analogs to Mars.

Growth, Viability, and Death of Planktonic and Biofilm Sphingomonas desiccabilis in Simulated Martian Brines.

Aqueous solutions on Mars are theorized to contain very different ion compositions than those on Earth. To determine the effect of such solutions on typical environmental micro-organisms, which could be released from robotic spacecraft or human exploration activity, we investigated the resistance of Sphingomonas desiccabilis to brines that simulate the composition of martian aqueous environments. S. desiccabilis is a desiccation-resistant, biofilm-forming microbe found in desert crusts. The viability of cells in both planktonic and biofilm forms was measured after exposure to simulated martian brines. Planktonic cells showed a loss of viability over the course of several hours in almost all of the seven brines tested. Biofilms conferred greater resistance to all the brines, including those with low water activity and pH, but even cells in biofilms showed a complete loss of viability in <6 h in the harsher brines and in <2 days in the less harsh brines. One brine, however, allowed the microbes to maintain viability over several days, despite having a water activity and pH lower and ionic strength higher than brines that reduced viability over the same timescales, suggesting important ion-specific effects. These data show that biofilm-forming cells have a greater capacity to resist martian aqueous extremes, but that evaporative or deliquescent brines are likely to be destructive to many organisms over relatively short timescales, with implications for the habitability of Mars and for micro-organisms dispersed by robotic or human explorers.

Correlations Between Life-Detection Techniques and Implications for Sampling Site Selection in Planetary Analog Missions.

We conducted an analog sampling expedition under simulated mission constraints to areas dominated by basaltic tephra of the Eldfell and Fimmvörðuháls lava fields (Iceland). Sites were selected to be "homogeneous" at a coarse remote sensing resolution (10-100 m) in apparent color, morphology, moisture, and grain size, with best-effort realism in numbers of locations and replicates. Three different biomarker assays (counting of nucleic-acid-stained cells via fluorescent microscopy, a luciferin/luciferase assay for adenosine triphosphate, and quantitative polymerase chain reaction (qPCR) to detect DNA associated with bacteria, archaea, and fungi) were characterized at four nested spatial scales (1 m, 10 m, 100 m, and >1 km) by using five common metrics for sample site representativeness (sample mean variance, group F tests, pairwise t tests, and the distribution-free rank sum H and u tests). Correlations between all assays were characterized with Spearman's rank test. The bioluminescence assay showed the most variance across the sites, followed by qPCR for bacterial and archaeal DNA; these results could not be considered representative at the finest resolution tested (1 m). Cell concentration and fungal DNA also had significant local variation, but they were homogeneous over scales of >1 km. These results show that the selection of life detection assays and the number, distribution, and location of sampling sites in a low biomass environment with limited a priori characterization can yield both contrasting and complementary results, and that their interdependence must be given due consideration to maximize science return in future biomarker sampling expeditions. Key Words: Astrobiology-Biodiversity-Microbiology-Iceland-Planetary exploration-Mars mission simulation-Biomarker. Astrobiology 17, 1009-1021.

NOMAD spectrometer on the ExoMars trace gas orbiter mission: part 2-design, manufacturing, and testing of the ultraviolet and visible channel.

NOMAD is a spectrometer suite on board the ESA/Roscosmos ExoMars Trace Gas Orbiter, which launched in March 2016. NOMAD consists of two infrared channels and one ultraviolet and visible channel, allowing the instrument to perform observations quasi-constantly, by taking nadir measurements at the day- and night-side, and during solar occultations. Here, in part 2 of a linked study, we describe the design, manufacturing, and testing of the ultraviolet and visible spectrometer channel called UVIS. We focus upon the optical design and working principle where two telescopes are coupled to a single grating spectrometer using a selector mechanism.