We propose that retinal-based phototrophy arose early in the evolution of life on Earth, profoundly impacting the development of photosynthesis and creating implications for the search for life beyond our planet. While the early evolutionary history of phototrophy is largely in the realm of the unknown, the onset of oxygenic photosynthesis in primitive cyanobacteria significantly altered the Earth's atmosphere by contributing to the rise of oxygen ~2.3 billion years ago. However, photosynthetic chlorophyll and bacteriochlorophyll pigments lack appreciable absorption at wavelengths about 500-600 nm, an energy-rich region of the solar spectrum. By contrast, simpler retinal-based light-harvesting systems such as the haloarchaeal purple membrane protein bacteriorhodopsin show a strong well-defined peak of absorbance centered at 568 nm, which is complementary to that of chlorophyll pigments. We propose a scenario where simple retinal-based light-harvesting systems like that of the purple chromoprotein bacteriorhodopsin, originally discovered in halophilic Archaea, may have dominated prior to the development of photosynthesis. We explore this hypothesis, termed the 'Purple Earth,' and discuss how retinal photopigments may serve as remote biosignatures for exoplanet research.

AbstractWe propose that retinal-based phototrophy arose early in the evolution of life on Earth, profoundly impacting the development of photosynthesis and creating implications for the search for life beyond our planet. While the early evolutionary history of phototrophy is largely in the realm of the unknown, the onset of oxygenic photosynthesis in primitive cyanobacteria significantly altered the Earth's atmosphere by contributing to the rise of oxygen ~2.3 billion years ago. However, photosynthetic chlorophyll and bacterio chlorophyll pigments lack appreciable absorption at wavelengths about 500–600 nm, an energy-rich region of the solar spectrum. By contrast, simpler retinal-based light-harvesting systems such as the haloarchaeal purple membrane protein bacteriorhodopsin show a strong well-defined peak of absorbance centred at 568 nm, which is complementary to that of chlorophyll pigments. We propose a scenario where simple retinal-based light-harvesting systems like that of the purple chromoprotein bacteriorhodopsin, originally discovered in halophilic Archaea, may have dominated prior to the development of photosynthesis. We explore this hypothesis, termed the ‘Purple Earth,’ and discuss how retinal photopigments may serve as remote biosignatures for exoplanet research.

Recent progress in extremophile biology, exploration of planetary bodies in the solar system, and the detection and characterization of extrasolar planets are leading to new insights in the field of astrobiology and possible distribution of life in the universe. Among the many extremophiles on Earth, the halophilic Archaea (Haloarchaea) are especially attractive models for astrobiology, being evolutionarily ancient and physiologically versatile, potentially surviving in a variety of planetary environments and with relevance for in situ life detection. Haloarchaea are polyextremophilic with tolerance of saturating salinity, anaerobic conditions, high levels of ultraviolet and ionizing radiation, subzero temperatures, desiccation, and toxic ions. Haloarchaea survive launches into Earth's stratosphere encountering conditions similar to those found on the surface of Mars. Studies of their unique proteins are revealing mechanisms permitting activity and function in high ionic strength, perchlorates, and subzero temperatures. Haloarchaea also produce spectacular blooms visible from space due to synthesis of red-orange isoprenoid carotenoids used for photoprotection and photorepair processes and purple retinal chromoproteins for phototrophy and phototaxis. Remote sensing using visible and infrared spectroscopy has shown that haloarchaeal pigments exhibit both a discernable peak of absorption and a reflective "green edge". Since the pigments produce remotely detectable features, they may influence the spectrum from an inhabited exoplanet imaged by a future large space-based telescope. In this review, we focus primarily on studies of two Haloarchaea, Halobacterium sp. NRC-1 and Halorubrum lacusprofundi.

We introduce a Bayesian method for guiding future directions for detection of life on exoplanets. We describe empirical and theoretical work necessary to place constraints on the relevant likelihoods, including those emerging from better understanding stellar environment, planetary climate and geophysics, geochemical cycling, the universalities of physics and chemistry, the contingencies of evolutionary history, the properties of life as an emergent complex system, and the mechanisms driving the emergence of life. We provide examples for how the Bayesian formalism could guide future search strategies, including determining observations to prioritize or deciding between targeted searches or larger lower resolution surveys to generate ensemble statistics and address how a Bayesian methodology could constrain the prior probability of life with or without a positive detection. Key Words: Exoplanets-Biosignatures-Life detection-Bayesian analysis. Astrobiology 18, 779-824.

We introduce a Bayesian method for guiding future directions for detection of life on exoplanets. We describe empirical and theoretical work necessary to place constraints on the relevant likelihoods, including those emerging from better understanding stellar environment, planetary climate and geophysics, geochemical cycling, the universalities of physics and chemistry, the contingencies of evolutionary history, the properties of life as an emergent complex system, and the mechanisms driving the emergence of life. We provide examples for how the Bayesian formalism could guide future search strategies, including determining observations to prioritize or deciding between targeted searches or larger lower resolution surveys to generate ensemble statistics and address how a Bayesian methodology could constrain the prior probability of life with or without a positive detection. Key Words: Exoplanets-Biosignatures-Life detection-Bayesian analysis. Astrobiology 18, 779-824.

Exoplanet science promises a continued rapid accumulation of new observations in the near future, energizing a drive to understand and interpret the forthcoming wealth of data to identify signs of life beyond our Solar System. The large statistics of exoplanet samples, combined with the ambiguity of our understanding of universal properties of life and its signatures, necessitate a quantitative framework for biosignature assessment Here, we introduce a Bayesian framework for guiding future directions in life detection, which permits the possibility of generalizing our search strategy beyond biosignatures of known life. The Bayesian methodology provides a language to define quantitatively the conditional probabilities and confidence levels of future life detection and, importantly, may constrain the prior probability of life with or without positive detection. We describe empirical and theoretical work necessary to place constraints on the relevant likelihoods, including those emerging from stellar and planetary context, the contingencies of evolutionary history and the universalities of physics and chemistry. We discuss how the Bayesian framework can guide our search strategies, including determining observational wavelengths or deciding between targeted searches or larger, lower resolution surveys. Our goal is to provide a quantitative framework not entrained to specific definitions of life or its signatures, which integrates the diverse disciplinary perspectives necessary to confidently detect alien life.

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 comprehensive 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 to maintain short-lived biogenic gases as atmospheric signatures. We focus particularly on advances made since the seminal review by Des Marais et al. The purpose of this work is not to propose new biosignature strategies, a goal left to companion articles in this series, but to review the current literature, draw meaningful connections between seemingly disparate areas, and clear the way for a path forward. Key Words: Exoplanets-Biosignatures-Habitability markers-Photosynthesis-Planetary surfaces-Atmospheres-Spectroscopy-Cryptic biospheres-False positives. Astrobiology 18, 663-708.

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.

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 comprehensive 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 to maintain short-lived biogenic gases as atmospheric signatures. We focus particularly on advances made since the seminal review by Des Marais et al. The purpose of this work is not to propose new biosignature strategies, a goal left to companion articles in this series, but to review the current literature, draw meaningful connections between seemingly disparate areas, and clear the way for a path forward. Key Words: Exoplanets-Biosignatures-Habitability markers-Photosynthesis-Planetary surfaces-Atmospheres-Spectroscopy-Cryptic biospheres-False positives. Astrobiology 18, 663-708.

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.