Accounting for herbaceous communities in process-based models will advance our understanding of "grassy" ecosystems.

Grassland and other herbaceous communities cover significant portions of Earth's terrestrial surface and provide many critical services, such as carbon sequestration, wildlife habitat, and food production. Forecasts of global change impacts on these services will require predictive tools, such as process-based dynamic vegetation models. Yet, model representation of herbaceous communities and ecosystems lags substantially behind that of tree communities and forests. The limited representation of herbaceous communities within models arises from two important knowledge gaps: first, our empirical understanding of the principles governing herbaceous vegetation dynamics is either incomplete or does not provide mechanistic information necessary to drive herbaceous community processes with models; second, current model structure and parameterization of grass and other herbaceous plant functional types limits the ability of models to predict outcomes of competition and growth for herbaceous vegetation. In this review, we provide direction for addressing these gaps by: (1) presenting a brief history of how vegetation dynamics have been developed and incorporated into earth system models, (2) reporting on a model simulation activity to evaluate current model capability to represent herbaceous vegetation dynamics and ecosystem function, and (3) detailing several ecological properties and phenomena that should be a focus for both empiricists and modelers to improve representation of herbaceous vegetation in models. Together, empiricists and modelers can improve representation of herbaceous ecosystem processes within models. In so doing, we will greatly enhance our ability to forecast future states of the earth system, which is of high importance given the rapid rate of environmental change on our planet.

The metabolic rate of the biosphere and its components.

We assessed the relationship between rates of biological energy utilization and the biomass sustained by that energy utilization, at both the organism and biosphere level. We compiled a dataset comprising >10,000 basal, field, and maximum metabolic rate measurements made on >2,900 individual species, and, in parallel, we quantified rates of energy utilization, on a biomass-normalized basis, by the global biosphere and by its major marine and terrestrial components. The organism-level data, which are dominated by animal species, have a geometric mean among basal metabolic rates of 0.012 W (g C)-1 and an overall range of more than six orders of magnitude. The biosphere as a whole uses energy at an average rate of 0.005 W (g C)-1 but exhibits a five order of magnitude range among its components, from 0.00002 W (g C)-1 for global marine subsurface sediments to 2.3 W (g C)-1 for global marine primary producers. While the average is set primarily by plants and microorganisms, and by the impact of humanity upon those populations, the extremes reflect systems populated almost exclusively by microbes. Mass-normalized energy utilization rates correlate strongly with rates of biomass carbon turnover. Based on our estimates of energy utilization rates in the biosphere, this correlation predicts global mean biomass carbon turnover rates of ~2.3 y-1 for terrestrial soil biota, ~8.5 y-1 for marine water column biota, and ~1.0 y-1 and ~0.01 y-1 for marine sediment biota in the 0 to 0.1 m and >0.1 m depth intervals, respectively.

Discovery of Chlorophyll d: Isolation and Characterization of a Far-Red Cyanobacterium from the Original Site of Manning and Strain (1943) at Moss Beach, California.

We have isolated a chlorophyll-d-containing cyanobacterium from the intertidal field site at Moss Beach, on the coast of Central California, USA, where Manning and Strain (1943) originally discovered this far-red chlorophyll. Here, we present the cyanobacterium's environmental description, culturing procedure, pigment composition, ultrastructure, and full genome sequence. Among cultures of far-red cyanobacteria obtained from red algae from the same site, this strain was an epiphyte on a brown macroalgae. Its Qyin vivo absorbance peak is centered at 704-705 nm, the shortest wavelength observed thus far among the various known Acaryochloris strains. Its Chl a/Chl d ratio was 0.01, with Chl d accounting for 99% of the total Chl d and Chl a mass. TEM imagery indicates the absence of phycobilisomes, corroborated by both pigment spectra and genome analysis. The Moss Beach strain codes for only a single set of genes for producing allophycocyanin. Genomic sequencing yielded a 7.25 Mbp circular chromosome and 10 circular plasmids ranging from 16 kbp to 394 kbp. We have determined that this strain shares high similarity with strain S15, an epiphyte of red algae, while its distinct gene complement and ecological niche suggest that this strain could be the closest known relative to the original Chl d source of Manning and Strain (1943). The Moss Beach strain is designated Acaryochloris sp. (marina) strain Moss Beach.

GISS-E2.1: Configurations and Climatology.

This paper describes the GISS-E2.1 contribution to the Coupled Model Intercomparison Project, Phase 6 (CMIP6). This model version differs from the predecessor model (GISS-E2) chiefly due to parameterization improvements to the atmospheric and ocean model components, while keeping atmospheric resolution the same. Model skill when compared to modern era climatologies is significantly higher than in previous versions. Additionally, updates in forcings have a material impact on the results. In particular, there have been specific improvements in representations of modes of variability (such as the Madden-Julian Oscillation and other modes in the Pacific) and significant improvements in the simulation of the climate of the Southern Oceans, including sea ice. The effective climate sensitivity to 2 × CO2 is slightly higher than previously at 2.7-3.1°C (depending on version) and is a result of lower CO2 radiative forcing and stronger positive feedbacks.

Albedos, Equilibrium Temperatures, and Surface Temperatures of Habitable Planets.

The potential habitability of known exoplanets is often categorized by a nominal equilibrium temperature assuming a Bond albedo of either ∼0.3, similar to Earth, or 0. As an indicator of habitability, this leaves much to be desired, because albedos of other planets can be very different, and because surface temperature exceeds equilibrium temperature due to the atmospheric greenhouse effect. We use an ensemble of general circulation model simulations to show that for a range of habitable planets, much of the variability of Bond albedo, equilibrium temperature and even surface temperature can be predicted with useful accuracy from incident stellar flux and stellar temperature, two known parameters for every confirmed exoplanet. Earth's Bond albedo is near the minimum possible for habitable planets orbiting G stars, because of increasing contributions from clouds and sea ice/snow at higher and lower instellations, respectively. For habitable M star planets, Bond albedo is usually lower than Earth's because of near-IR H2O absorption, except at high instellation where clouds are important. We apply relationships derived from this behavior to several known exoplanets to derive zeroth-order estimates of their potential habitability. More expansive multivariate statistical models that include currently non-observable parameters show that greenhouse gas variations produce significant variance in albedo and surface temperature, while increasing length of day and land fraction decrease surface temperature; insights for other parameters are limited by our sampling. We discuss how emerging information from global climate models might resolve some degeneracies and help focus scarce observing resources on the most promising planets.

The Sensitivity of Land-Atmosphere Coupling to Modern Agriculture in the Northern Midlatitudes.

Modern agricultural land cover and management are important as regional climate forcings. Previous work has shown that land cover change can significantly impact key climate variables, including turbulent fluxes, precipitation, and surface temperature. However, fewer studies have investigated how intensive crop management can impact background climate conditions, such as the strength of land-atmosphere coupling and evaporative regime. We conduct sensitivity experiments using a state-of-the-art climate model with modified vegetation characteristics to represent modern crop cover and management, using observed crop-specific leaf area indexes and calendars. We quantify changes in land-atmosphere interactions and climate over intensively cultivated regions situated at transitions between moisture- and energy-limited conditions. Results show that modern intensive agriculture has significant and geographically varying impacts on regional evaporative regimes and background climate conditions. Over the northern Great Plains, modern crop intensity increases the model simulated precipitation and soil moisture, weakening hydrologic coupling by increasing surface water availability and reducing moisture limits on evapotranspiration. In the U.S. Midwest, higher growing season evapotranspiration, coupled with winter and spring rainfall declines, reduces regional soil moisture, while crop albedo changes also reduce net surface radiation. This results overall in reduced dependency of regional surface temperature on latent heat fluxes. In central Asia, a combination of reduced net surface energy and enhanced pre-growing season precipitation amplify the energy-limited evaporative regime. These results highlight the need for improved representations of agriculture in global climate models to better account for regional climate impacts and interactions with other anthropogenic forcings.

Habitable Climate Scenarios for Proxima Centauri b with a Dynamic Ocean.

The nearby exoplanet Proxima Centauri b will be a prime future target for characterization, despite questions about its retention of water. Climate models with static oceans suggest that Proxima b could harbor a small dayside surface ocean despite its weak instellation. We present the first climate simulations of Proxima b with a dynamic ocean. We find that an ocean-covered Proxima b could have a much broader area of surface liquid water but at much colder temperatures than previously suggested, due to ocean heat transport and/or depression of the freezing point by salinity. Elevated greenhouse gas concentrations do not necessarily produce more open ocean because of dynamical regime transitions between a state with an equatorial Rossby-Kelvin wave pattern and a state with a day-night circulation. For an evolutionary path leading to a highly saline ocean, Proxima b could be an inhabited, mostly open ocean planet with halophilic life. A freshwater ocean produces a smaller liquid region than does an Earth salinity ocean. An ocean planet in 3:2 spin-orbit resonance has a permanent tropical waterbelt for moderate eccentricity. A larger versus smaller area of surface liquid water for similar equilibrium temperature may be distinguishable by using the amplitude of the thermal phase curve. Simulations of Proxima Centauri b may be a model for the habitability of weakly irradiated planets orbiting slightly cooler or warmer stars, for example, in the TRAPPIST-1, LHS 1140, GJ 273, and GJ 3293 systems.

Exoplanet Biosignatures: Future Directions.

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 Biosignatures: At the Dawn of a New Era of Planetary Observations.

The rapid rate of discoveries of exoplanets has expanded the scope of the science possible for the remote detection of life beyond Earth. The Exoplanet Biosignatures Workshop Without Walls (EBWWW) held in 2016 engaged the international scientific community across diverse scientific disciplines, to assess the state of the science and technology in the search for life on exoplanets, and to identify paths for progress. The workshop activities resulted in five major review papers, which provide (1) an encyclopedic review of known and proposed biosignatures and models used to ascertain them (Schwieterman et al., 2018 in this issue); (2) an in-depth review of O2 as a biosignature, rigorously examining the nuances of false positives and false negatives for evidence of life (Meadows et al., 2018 in this issue); (3) a Bayesian framework to comprehensively organize current understanding to quantify confidence in biosignature assessments (Catling et al., 2018 in this issue); (4) an extension of that Bayesian framework in anticipation of increasing planetary data and novel concepts of biosignatures (Walker et al., 2018 in this issue); and (5) a review of the upcoming telescope capabilities to characterize exoplanets and their environment (Fujii et al., 2018 in this issue). Because of the immense content of these review papers, this summary provides a guide to their complementary scope and highlights salient features. Strong themes that emerged from the workshop were that biosignatures must be interpreted in the context of their environment, and that frameworks must be developed to link diverse forms of scientific understanding of that context to quantify the likelihood that a biosignature has been observed. Models are needed to explore the parameter space where measurements will be widespread but sparse in detail. Given the technological prospects for large ground-based telescopes and space-based observatories, the detection of atmospheric signatures of a few potentially habitable planets may come before 2030. Key Words: Exoplanets-Biosignatures-Remote observation-Spectral imaging-Bayesian analysis. Astrobiology 18, 619-626.

Exoplanet Biosignatures: A Review of Remotely Detectable Signs of 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.

Exoplanet Biosignatures: A Framework for Their Assessment.

Finding life on exoplanets from telescopic observations is an ultimate goal of exoplanet science. Life produces gases and other substances, such as pigments, which can have distinct spectral or photometric signatures. Whether or not life is found with future data must be expressed with probabilities, requiring a framework of biosignature assessment. We present a framework in which we advocate using biogeochemical "Exo-Earth System" models to simulate potential biosignatures in spectra or photometry. Given actual observations, simulations are used to find the Bayesian likelihoods of those data occurring for scenarios with and without life. The latter includes "false positives" wherein abiotic sources mimic biosignatures. Prior knowledge of factors influencing planetary inhabitation, including previous observations, is combined with the likelihoods to give the Bayesian posterior probability of life existing on a given exoplanet. Four components of observation and analysis are necessary. (1) Characterization of stellar (e.g., age and spectrum) and exoplanetary system properties, including "external" exoplanet parameters (e.g., mass and radius), to determine an exoplanet's suitability for life. (2) Characterization of "internal" exoplanet parameters (e.g., climate) to evaluate habitability. (3) Assessment of potential biosignatures within the environmental context (components 1-2), including corroborating evidence. (4) Exclusion of false positives. We propose that resulting posterior Bayesian probabilities of life's existence map to five confidence levels, ranging from "very likely" (90-100%) to "very unlikely" (<10%) inhabited. Key Words: Bayesian statistics-Biosignatures-Drake equation-Exoplanets-Habitability-Planetary science. Astrobiology 18, 709-738.

Climates of Warm Earth-like Planets I: 3-D Model Simulations.

We present a large ensemble of simulations of an Earth-like world with increasing insolation and rotation rate. Unlike previous work utilizing idealized aquaplanet configurations we focus our simulations on modern Earth-like topography. The orbital period is the same as modern Earth, but with zero obliquity and eccentricity. The atmosphere is 1 bar N2-dominated with CO2=400 ppmv and CH4=1 ppmv. The simulations include two types of oceans; one without ocean heat transport (OHT) between grid cells as has been commonly used in the exoplanet literature, while the other is a fully coupled dynamic bathtub type ocean. The dynamical regime transitions that occur as day length increases induce climate feedbacks producing cooler temperatures, first via the reduction of water vapor with increasing rotation period despite decreasing shortwave cooling by clouds, and then via decreasing water vapor and increasing shortwave cloud cooling, except at the highest insolations. Simulations without OHT are more sensitive to insolation changes for fast rotations while slower rotations are relatively insensitive to ocean choice. OHT runs with faster rotations tend to be similar with gyres transporting heat poleward making them warmer than those without OHT. For slower rotations OHT is directed equator-ward and no high latitude gyres are apparent. Uncertainties in cloud parameterization preclude a precise determination of habitability but do not affect robust aspects of exoplanet climate sensitivity. This is the first paper in a series that will investigate aspects of habitability in the simulations presented herein. The datasets from this study are opensource and publicly available.

Was Venus the First Habitable World of our Solar System?

Present-day Venus is an inhospitable place with surface temperatures approaching 750K and an atmosphere 90 times as thick as Earth's. Billions of years ago the picture may have been very different. We have created a suite of 3-D climate simulations using topographic data from the Magellan mission, solar spectral irradiance estimates for 2.9 and 0.715 Gya, present-day Venus orbital parameters, an ocean volume consistent with current theory, and an atmospheric composition estimated for early Venus. Using these parameters we find that such a world could have had moderate temperatures if Venus had a rotation period slower than ~16 Earth days, despite an incident solar flux 46-70% higher than Earth receives. At its current rotation period, Venus's climate could have remained habitable until at least 715 million years ago. These results demonstrate the role rotation and topography play in understanding the climatic history of Venus-like exoplanets discovered in the present epoch.

Photosystem trap energies and spectrally-dependent energy-storage efficiencies in the Chl d-utilizing cyanobacterium, Acaryochloris marina.

Acaryochloris marina is the only species known to utilize chlorophyll (Chl) d as a principal photopigment. The peak absorption wavelength of Chl d is redshifted ≈40nm in vivo relative to Chl a, enabling this cyanobacterium to perform oxygenic phototrophy in niche environments enhanced in far-red light. We present measurements of the in vivo energy-storage (E-S) efficiency of photosynthesis in A. marina, obtained using pulsed photoacoustics (PA) over a 90-nm range of excitation wavelengths in the red and far-red. Together with modeling results, these measurements provide the first direct observation of the trap energies of PSI and PSII, and also the photosystem-specific contributions to the total E-S efficiency. We find the maximum observed efficiency in A. marina (40±1% at 735nm) is higher than in the Chl a cyanobacterium Synechococcus leopoliensis (35±1% at 690nm). The efficiency at peak absorption wavelength is also higher in A. marina (36±1% at 710nm vs. 31±1% at 670nm). In both species, the trap efficiencies are ≈40% (PSI) and ≈30% (PSII). The PSI trap in A. marina is found to lie at 740±5nm, in agreement with the value inferred from spectroscopic methods. The best fit of the model to the PA data identifies the PSII trap at 723±3nm, supporting the view that the primary electron-donor is Chl d, probably at the accessory (Chl(D1)) site. A decrease in efficiency beyond the trap wavelength, consistent with uphill energy transfer, is clearly observed and fit by the model. These results demonstrate that the E-S efficiency in A. marina is not thermodynamically limited, suggesting that oxygenic photosynthesis is viable in even redder light environments.

Spectral signatures of photosynthesis. I. Review of Earth organisms.

Why do plants reflect in the green and have a "red edge" in the red, and should extrasolar photosynthesis be the same? We provide (1) a brief review of how photosynthesis works, (2) an overview of the diversity of photosynthetic organisms, their light harvesting systems, and environmental ranges, (3) a synthesis of photosynthetic surface spectral signatures, and (4) evolutionary rationales for photosynthetic surface reflectance spectra with regard to utilization of photon energy and the planetary light environment. We found the "near-infrared (NIR) end" of the red edge to trend from blue-shifted to reddest for (in order) snow algae, temperate algae, lichens, mosses, aquatic plants, and finally terrestrial vascular plants. The red edge is weak or sloping in lichens. Purple bacteria exhibit possibly a sloping edge in the NIR. More studies are needed on pigment-protein complexes, membrane composition, and measurements of bacteria before firm conclusions can be drawn about the role of the NIR reflectance. Pigment absorbance features are strongly correlated with features of atmospheric spectral transmittance: P680 in Photosystem II with the peak surface incident photon flux density at approximately 685 nm, just before an oxygen band at 687.5 nm; the NIR end of the red edge with water absorbance bands and the oxygen A-band at 761 nm; and bacteriochlorophyll reaction center wavelengths with local maxima in atmospheric and water transmittance spectra. Given the surface incident photon flux density spectrum and resonance transfer in light harvesting, we propose some rules with regard to where photosynthetic pigments will peak in absorbance: (1) the wavelength of peak incident photon flux; (2) the longest available wavelength for core antenna or reaction center pigments; and (3) the shortest wavelengths within an atmospheric window for accessory pigments. That plants absorb less green light may not be an inefficient legacy of evolutionary history, but may actually satisfy the above criteria.

Spectral signatures of photosynthesis. II. Coevolution with other stars and the atmosphere on extrasolar worlds.

As photosynthesis on Earth produces the primary signatures of life that can be detected astronomically at the global scale, a strong focus of the search for extrasolar life will be photosynthesis, particularly photosynthesis that has evolved with a different parent star. We take previously simulated planetary atmospheric compositions for Earth-like planets around observed F2V and K2V, modeled M1V and M5V stars, and around the active M4.5V star AD Leo; our scenarios use Earth's atmospheric composition as well as very low O2 content in case anoxygenic photosynthesis dominates. With a line-by-line radiative transfer model, we calculate the incident spectral photon flux densities at the surface of the planet and under water. We identify bands of available photosynthetically relevant radiation and find that photosynthetic pigments on planets around F2V stars may peak in absorbance in the blue, K2V in the red-orange, and M stars in the near-infrared, in bands at 0.93-1.1 microm, 1.1-1.4 microm, 1.5-1.8 microm, and 1.8-2.5 microm. However, underwater organisms will be restricted to wavelengths shorter than 1.4 microm and more likely below 1.1 microm. M star planets without oxygenic photosynthesis will have photon fluxes above 1.6 microm curtailed by methane. Longer-wavelength, multi-photo-system series would reduce the quantum yield but could allow for oxygenic photosystems at longer wavelengths. A wavelength of 1.1 microm is a possible upper cutoff for electronic transitions versus only vibrational energy; however, this cutoff is not strict, since such energetics depend on molecular configuration. M star planets could be a half to a tenth as productive as Earth in the visible, but exceed Earth if useful photons extend to 1.1 microm for anoxygenic photosynthesis. Under water, organisms would still be able to survive ultraviolet flares from young M stars and acquire adequate light for growth.

Detectability of planetary characteristics in disk-averaged spectra II: synthetic spectra and light-curves of earth.

Spatially and spectrally resolved models were used to explore the observational sensitivity to changes in atmospheric and surface properties and the detectability of surface biosignatures in the globally averaged spectra and light-curves of the Earth. Compared with previous efforts to characterize the Earth using disk-averaged models, a more comprehensive and realistic treatment of the surface and atmosphere was taken into account here. Our results are presented as a function of viewing geometry and phases at both visible/near-infrared (0.5-1.7 microm) and mid-infrared (5-25 microm) wavelength ranges, applicable to the proposed NASA-Terrestrial Planet Finder visible coronagraph and mid-infrared interferometer and to the ESADarwin mission architectures. Clouds can change the thermal emission by as much as 50% compared with the cloud-free case and increase the visible albedo by up to 500% for completely overcast cases at the dichotomy phase. Depending on the observed phase and their distribution and type, clouds can also significantly alter the spectral shape. Moreover, clouds impact the detectability of surface biosignatures in the visible wavelength range. Modeling the disk-averaged sensitivity to the "red-edge," a distinctive spectral signature of vegetation, showed that Earth's land vegetation could be seen in disk-averaged spectra, even with cloud cover, when the signal was averaged over the daily time scale. We found that vegetation is more readily discriminated from clouds at dichotomy (50% illumination) rather than at full phase. The detectability of phytoplankton was also explored, but was found to be more difficult to detect in the disk-average than land vegetation.