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The first potential exoplanet biosignature detections are likely to be ambiguous due to the potential for false positives: abiotic planetary processes that produce observables similar to those anticipated from a global biosphere. Here we propose a class of methylated gases as corroborative `capstone' biosignatures. Capstone biosignatures are metabolic products that may be less immediately detectable, but have substantially lower false positive potential, and can thus serve as confirmation for a primary biosignature such as O$_2$. CH$_3$Cl has previously been established as a biosignature candidate, and other halomethane gases such as CH$_3$Br and CH$_3$I have similar potential. These gases absorb in the mid infrared at wavelengths that are likely to be captured while observing primary biosignatures such as O$_3$ or CH$_4$. We quantitatively explore CH$_3$Br as a new capstone biosignature through photochemical and spectral modeling of Earth-like planets orbiting FGKM stellar hosts. We also re-examine the biosignature potential of CH$_3$Cl over the same set of parameters using our updated model. We show that CH$_3$Cl and CH$_3$Br can build up to relatively high levels in M dwarf environments and analyze synthetic spectra of TRAPPIST-1e. Our results suggest that there is a co-additive spectral effect from multiple CH$_3$X gases in an atmosphere, leading to increased signal-to-noise and greater ability to detect a methylated gas feature. These capstone biosignatures are plausibly detectable in exoplanetary atmospheres, have low false positive potential, and would provide strong evidence for life in conjunction with other well established biosignature candidates., Comment: 18 pages, 11 figures, appendix. Published in ApJ

In the search for life on other planets, the presence of photosynthetic vegetation may be detectable from the colors of light it reflects. On the modern Earth, this spectral reflectance is characterized by an increase in reflectance between the red and near-infrared wavelengths, a "red edge". On planets orbiting different stellar types, red edge analogs may occur at other colors than red. Thus, knowing the wavelengths at which photosynthetic organisms preferentially absorb and reflect photons is necessary to detect red edge analogs on other planets. Using a numerical model that predicts the absorbance spectrum of extant photosynthetic pigments on Earth from Marosv\"olgyi and van Gorkom (2010), we calculate the absorbance spectrum for pigments on an Earth-like planet around F through late M type stars that are adapted for maximal energy production. In this model, cellular energy production is maximized when pigments are tuned to absorb at the wavelength that maximizes energy input from incident photons while minimizing thermal emission and costs to build the photosynthetic apparatus. We find that peak photon absorption for photosynthetic organisms around F type stars tends to be in the blue while for G, K, and early M type stars, red or just beyond is preferred. Around the coolest M type stars, these organisms may preferentially absorb in the near-infrared, possibly past one micron. These predictions are consistent with previous, qualitative estimates of pigment absorptance. Our predicted pigment absorbance spectra depend on both the stellar type and planetary atmospheric composition, especially atmospheric water vapor concentrations, which alter the availability of surface photons and thus the predicted pigment absorption. By constraining the absorbance spectra of alien, photosynthetic organisms, future observations may be better equipped to detect red edge analogs.

Here we review how environmental context can be used to interpret whether O2 is a biosignature in extrasolar planetary observations. This paper builds on the overview of current biosignature research discussed in Schwieterman et al. (2017), and provides an in-depth, interdisciplinary example of biosignature identification and observation that serves as a basis for the development of the general framework for biosignature assessment described in Catling et al., (2017). O2 is a potentially strong biosignature that was originally thought to be an unambiguous indicator for life at high-abundance. We describe the coevolution of life with the early Earth's environment, and how the interplay of sources and sinks in the planetary environment may have resulted in suppression of O2 release into the atmosphere for several billion years, a false negative for biologically generated O2. False positives may also be possible, with recent research showing potential mechanisms in exoplanet environments that may generate relatively high abundances of atmospheric O2 without a biosphere being present. These studies suggest that planetary characteristics that may enhance false negatives should be considered when selecting targets for biosignature searches. Similarly our ability to interpret O2 observed in an exoplanetary atmosphere is also crucially dependent on environmental context to rule out false positive mechanisms. We describe future photometric, spectroscopic and time-dependent observations of O2 and the planetary environment that could increase our confidence that any observed O2 is a biosignature, and help discriminate it from potential false positives. By observing and understanding O2 in its planetary context we can increase our confidence in the remote detection of life, and provide a model for biosignature development for other proposed biosignatures., Comment: 55 pages. The paper is the second in a series of 5 review manuscripts of the NExSS Exoplanet Biosignatures Workshop. Community commenting is solicited at https://nexss.info/groups/ebwww

In the coming years and decades, advanced space- and ground-based observatories will allow an unprecedented opportunity to probe the atmospheres and surfaces of potentially habitable exoplanets for signatures of life. Life on Earth, through its gaseous products and reflectance and scattering properties, has left its fingerprint on the spectrum of our planet. Aided by the universality of the laws of physics and chemistry, we turn to Earth's biosphere, both in the present and through geologic time, for analog signatures that will aid in the search for life elsewhere. Considering the insights gained from modern and ancient Earth, and the broader array of hypothetical exoplanet possibilities, we have compiled a state-of-the-art overview of our current understanding of potential exoplanet biosignatures including gaseous, surface, and temporal biosignatures. We additionally survey biogenic spectral features that are well-known in the specialist literature but have not yet been robustly vetted in the context of exoplanet biosignatures. We briefly review advances in assessing biosignature plausibility, including novel methods for determining chemical disequilibrium from remotely obtainable data and assessment tools for determining the minimum biomass required for a given atmospheric signature. We focus particularly on advances made since the seminal review by Des Marais et al. (2002). The purpose of this work is not to propose new biosignatures strategies, a goal left to companion papers in this series, but to review the current literature, draw meaningful connections between seemingly disparate areas, and clear the way for a path forward., Comment: Open Access Article. 46 pages, 13 figures

A focus of many astrobiology questions resides on the interplay between biochemistry and geochemistry. This biotic-abiotic interface is intimately connected to habitability. Computational systems biology provides a paradigm to characterize this interface. Here we describe the application of constraint-based metabolic modeling to characterize photosynthetic microorganisms inhabiting a modern analog to the Earth’s primordial ocean. Biochemistry is captured in a metabolic network that is converted to a mathematical model. Geochemistry results in constraints that limit the capabilities of the metabolic network. The combination of the two within a single framework can elucidate the co-evolution of life and the environment.

Our understanding of long-term organic matter preservation comes mostly from studies in aquatic systems. In contrast, taphonomic processes in extremely dry environments are relatively understudied and are poorly understood. We investigated the accumulation and preservation of lipid biomarkers in hyperarid soils in the Yungay region of the Atacama Desert. Lipids from seven soil horizons in a 2.5 m vertical profile were extracted and analyzed using GC-MS and LC-MS. Diagnostic functionalized lipids and geolipids were detected and increased in abundance and diversity with depth. Deeper clay units contain fossil organic matter (radiocarbon dead) that has been protected from rainwater since the onset of hyperaridity. We show that these clay units contain lipids in an excellent state of structural preservation with functional groups and unsaturated bonds in carbon chains. This indicates that minimal degradation of lipids has occurred in these soils since the time of their deposition between greater than 40,000 and 2 million years ago. The exceptional structural preservation of biomarkers is likely due to the long-term hyperaridity that has minimized microbial and enzymatic activity, a taphonomic process we term xeropreservation (i.e., preservation by drying). The degree of biomarker preservation allowed us to reconstruct major changes in ecology in the Yungay region that reflect a shift in hydrological regime from wet to dry since the early Quaternary. Our results suggest that hyperarid environments, which comprise 7.5 percent of the continental landmass, could represent a rich and relatively unexplored source of paleobiological informationon Earth.