Searching for signs of alien life-biosignatures-on distant planets requires general hypotheses on the evolution of biospheres, and how life interacts with and influences its planet. We will not be able to visit a distant planet to directly observe any alien life and thus measuring properties of a biosphere such as organism abundance or reproduction rate is impossible. This paper provides an overview of two life-environment coupled models and analyses how their results might aid in guiding our search for biosignatures. The first model is of a simple methanogen biosphere and demonstrates that the 'strength' (i.e. the concentration of a gas in the atmosphere) of the biosignature is only very minimally impacted by the population dynamics of the biosphere. This reduces the number of assumptions needed to predict biosignatures for a given metabolic pathway. The second model is of an evolving life-environment coupled system-the Tangled Nature model-and demonstrates that the most probable trajectory for an exo-biosphere is to diversify over time owing to a mechanism known as sequential selection with memory. These models can inform our search for alien life in the galaxy.This article is part of the discussion meeting issue 'Chance and purpose in the evolution of biospheres'.

Studying the origin of life and its prevalence in the universe offers a perspective that compels us to look after our irreplaceable home in the cosmos. An entrenched conception of humans as distinct from Nature prevents us from seeing and embracing our place in space and time, to our catastrophic detriment. We call on our colleagues to harness this unique perspective to connect their research with broader problems facing humanity, while leveraging the trust, credibility, and privilege of the scientific enterprise.

Life on Earth has experienced numerous upheavals over its approximately 4 billion year history. In previous work we have discussed how interruptions to stability lead, on average, to increases in habitability over time, a tendency we called Entropic Gaia. Here we continue this exploration, working with the Tangled Nature Model of co-evolution, to understand how the evolutionary history of life is shaped by periods of acute environmental stress. We find that while these periods of stress pose a risk of complete extinction, they also create opportunities for evolutionary exploration which would otherwise be impossible, leading to more populous and stable states among the survivors than in alternative histories without a stress period. We also study how the duration, repetition and number of refugia into which life escapes during the perturbation affects the final outcome. The model results are discussed in relation to both Earth history and the search for alien life.

ABSTRACT Life on Earth has experienced numerous upheavals over its approximately 4 billion year history. In previous work, we have discussed how interruptions to stability lead, on average, to increases in habitability over time, a tendency we called entropic Gaia. Here, we continue this exploration, working with the Tangled Nature Model of co-evolution, to understand how the evolutionary history of life is shaped by periods of acute environmental stress. We find that while these periods of stress pose a risk of complete extinction, they also create opportunities for evolutionary exploration which would otherwise be impossible, leading to more populous and stable states among the survivors than in alternative histories without a stress period. We also study how the duration, repetition and number of refugia into which life escapes during the perturbation affects the final outcome. The model results are discussed in relation to both Earth history and the search for alien life.

Methane is thought to have been an important greenhouse gas during the Archean, although its potential warming has been found to be limited at high concentrations due to its high shortwave absorption. We use the Met Office Unified Model, a general circulation model, to further explore the climatic effect of different Archean methane concentrations. Surface warming peaks at a pressure ratio CH$_4$:CO$_2$ of approximately 0.1, reaching a maximum of up to 7 K before significant cooling above this ratio. Equator-to-pole temperature differences also tend to increase up to pCH$_4$ $\leq$300 Pa, which is driven by a difference in radiative forcing at the equator and poles by methane and a reduction in the latitudinal extend of the Hadley circulation. 3D models are important to fully capture the cooling effect of methane, due to these impacts of the circulation.

In this article, we introduce some simple models, based on rolling dice, to explore mechanisms proposed to explain planetary habitability. The idea is to study these selection mechanisms in an analytically tractable setting, isolating their consequences from other details which can confound or obscure their effect in more realistic models. We find that the observable of interest, the face value shown on the die, "improves" over time in all models. For two of the more popular ideas, Selection by Survival and Sequential Selection, this is down to sampling effects. A modified version of Sequential Selection, Sequential Selection with Memory, implies a statistical tendency for systems to improve over time. We discuss the implications of this and its relationship to the ideas of the "Inhabitance Paradox" and the "Gaian bottleneck."

When searching for inhabited exoplanets, understanding the boundaries of the habitable zone around the parent star is key. If life can strongly influence its global environment, then we would expect the boundaries of the habitable zone to be influenced by the presence of life. Here using a simple abstract model of `tangled-ecology' where life can influence a global parameter, labelled as temperature, we investigate the boundaries of the habitable zone of our model system. As with other models of life-climate interactions, the species act to regulate the temperature. However, the system can also experience `punctuations', where the system's state jumps between different equilibria. Despite this, an ensemble of systems still tends to sustain or even improve conditions for life on average, a feature we call Entropic Gaia. The mechanism behind this is sequential selection with memory which is discussed in detail. With this modelling framework we investigate questions about how Gaia can affect and ultimately extend the habitable zone to what we call the Gaian habitable zone. This generates concrete predictions for the size of the habitable zone around stars, suggests directions for future work on the simulation of exoplanets and provides insight into the Gaian bottleneck hypothesis and the habitability/inhabitance paradox.

The search for biosignatures necessitates developing our understanding of life under different conditions. If life can influence the climate evolution of its planet then understanding the behaviour of life-climate feedbacks under extreme conditions is key to determine the 'edges' of the habitable zone. Additionally understanding the behaviour of a temperature limited biosphere will help towards formulating biosignature predictions for alien life living under conditions very different to those on Earth. Towards this aim, we extend the 'ExoGaia Model' - an abstract model of microbial life living on a highly simplified 0-dimensional planet. Via their metabolisms, microbes influence the atmospheric composition and therefore the temperature of the planet and emergent feedback loops allow microbes to regulate their climate and maintain long term habitability. Here, we adapt the ExoGaia model to include temperature adaptation of the microbes by allowing different species to have different temperature 'preferences'. We find that rather than adapting towards the planet's abiotic conditions the biosphere tends to more strongly influence the climate of its planet, suggesting that the surface temperature of an inhabited planet might be significantly different from that predicted using abiotic models. We find that the success rate for microbial establishment on planets is improved when adaptation is allowed. However, planetary abiotic context is important for determining whether overall survival prospects for life will be improved or degraded. These results indicate the necessity to develop an understanding of life living under different limiting regimes to form predictions for the boundaries of the habitable zone.

With the characterisations of potentially habitable planetary atmospheres on the horizon, the search for biosignatures is set to become a major area of research in the coming decades. To understand the atmospheric characteristics that might indicate alien life we must understand the abiotic characteristics of a planet and how life interacts with its environment. In the field of biogeochemistry, sophisticated models of life-environment coupled systems demonstrate that many assumptions specific to Earth-based life, e.g. specific ATP maintenance costs, are unnecessary to accurately model a biosphere. We explore a simple model of a single-species microbial biosphere that produces CH4 as a byproduct of the microbes' energy extraction - known as a type I biosignature. We demonstrate that although significantly changing the biological parameters has a large impact on the biosphere's total population, such changes have only a minimal impact on the strength of the resulting biosignature, while the biosphere is limited by H2 availability. We extend the model to include more accurate microbial energy harvesting and show that adjusting microbe parameters can lead to a regime change where the biosphere becomes limited by energy availability and no longer fully exploits the available H2, impacting the strength of the resulting biosignature. We demonstrate that, for a nutrient limited biosphere, identifying the limiting nutrient, understanding the abiotic processes that control its abundance, and determining the biosphere's ability to exploit it, are more fundamental for making type I biosignature predictions than the details of the population dynamics of the biosphere.

Nature-based solutions to climate change are growing policy priorities yet remain hard to quantify. Here we use remote sensing to quantify direct and indirect benefits from community-led agroforestry by The International Small group and Tree planting program (TIST) in Kenya. Since 2005, TIST-Kenya has incentivised smallholder farmers to plant trees for agricultural benefit and to sequester CO 2 . We use Landsat-7 satellite imagery to examine the effect on the historically deforested landscape around Mount Kenya. We identify positive greening trends in TIST groves during 2000–2019 relative to the wider landscape. These groves cover 27,198 ha, and a further 27,750 ha of neighbouring agricultural land is also positively influenced by TIST. This positive ‘spill-over’ impact of TIST activity occurs at up to 360 m distance. TIST also benefits local forests, e.g. through reducing fuelwood and fodder extraction. Our results show that community-led initiatives can lead to successful landscape-scale regreening on decadal timescales.

The Gaia hypothesis considers the life-environment coupled system as a single entity that acts to regulate and maintain habitable conditions on Earth. In this paper we discuss three mechanisms which could potentially lead to Gaia: Selection by Survival, Sequential Selection and Entropic Hierarchy. We use the Tangled Nature Model of co-evolution as a common framework for investigating all three, using an extended version of the standard model to elaborate on Gaia as an example of an entropic hierarchy. This idea, which combines sequential selection together with a reservoir of diversity that acts as a 'memory', implies a tendency towards growth and increasing resilience of the Gaian system over time. We then discuss how Gaian memory could be realised in practice via the microbial seed bank, climate refugia and lateral gene transfer and conclude by discussing testable implications of an entropic hierarchy for the study of Earth history and the search for life in the universe. This paper adds to the existing taxonomy of Gaia hypotheses to suggest an "Entropic Gaia" where we argue that increasing biomass, complexity and enhanced habitability over time is a statistically likely feature of a co-evolving system.

The search for habitable exoplanets inspires the question - how do habitable planets form? Planet habitability models traditionally focus on abiotic processes and neglect a biotic response to changing conditions on an inhabited planet. The Gaia hypothesis postulates that life influences the Earth's feedback mechanisms to form a self-regulating system, and hence that life can maintain habitable conditions on its host planet. If life has a strong influence, it will have a role in determining a planet's habitability over time. We present the ExoGaia model - a model of simple 'planets' host to evolving microbial biospheres. Microbes interact with their host planet via consumption and excretion of atmospheric chemicals. Model planets orbit a 'star' which provides incoming radiation, and atmospheric chemicals have either an albedo, or a heat-trapping property. Planetary temperatures can therefore be altered by microbes via their metabolisms. We seed multiple model planets with life while their atmospheres are still forming and find that the microbial biospheres are, under suitable conditions, generally able to prevent the host planets from reaching inhospitable temperatures, as would happen on a lifeless planet. We find that the underlying geochemistry plays a strong role in determining long-term habitability prospects of a planet. We find five distinct classes of model planets, including clear examples of 'Gaian bottlenecks' - a phenomenon whereby life either rapidly goes extinct leaving an inhospitable planet, or survives indefinitely maintaining planetary habitability. These results suggest that life might play a crucial role in determining the long-term habitability of planets.

Recently postulated mechanisms and models can help explain the enduring 'Gaia' puzzle of environmental regulation mediated by life. Natural selection can produce nutrient recycling at local scales and regulation of heterogeneous environmental variables at ecosystem scales. However, global-scale environmental regulation involves a temporal and spatial decoupling of effects from actors that makes conventional evolutionary explanations problematic. Instead, global regulation can emerge by a process of 'sequential selection' in which systems that destabilize their environment are short-lived and result in extinctions and reorganizations until a stable attractor is found. Such persistence-enhancing properties can in turn increase the likelihood of acquiring further persistence-enhancing properties through 'selection by survival alone'. Thus, Earth system feedbacks provide a filter for persistent combinations of macroevolutionary innovations.

Recently postulated mechanisms and models can help explain the enduring 'Gaia' puzzle of environmental regulation mediated by life. Natural selection can produce nutrient recycling at local scales and regulation of heterogeneous environmental variables at ecosystem scales. However, global-scale environmental regulation involves a temporal and spatial decoupling of effects from actors that makes conventional evolutionary explanations problematic. Instead, global regulation can emerge by a process of 'sequential selection' in which systems that destabilize their environment are short-lived and result in extinctions and reorganizations until a stable attractor is found. Such persistence-enhancing properties can in turn increase the likelihood of acquiring further persistence-enhancing properties through 'selection by survival alone'. Thus, Earth system feedbacks provide a filter for persistent combinations of macroevolutionary innovations.

A long-standing objection to the Gaia hypothesis has been a perceived lack of plausible mechanisms by which life on Earth could come to regulate its abiotic environment. A null hypothesis is survival by pure chance, by which any appearance of regulation on Earth is illusory and the persistence of life simply reflects the weak anthropic principle - it must have occurred for intelligent observers to ask the question. Recent work has proposed that persistence alone increases the chance that a biosphere will acquire further persistence-enhancing properties. Here we use a simple quantitative model to show that such 'selection by survival alone' can indeed increase the probability that a biosphere will persist in the future, relative to a baseline of pure chance. Adding environmental feedback to this model shows either an increased or decreased survival probability depending on the initial conditions. Feedback can hinder early life becoming established if initial conditions are poor, but feedback can also prevent systems from diverging too far from optimum environmental conditions and thus increase survival rates. The outstanding question remains the relative importance of each mechanism for the historical and continued persistence of life on Earth.

We modify the Tangled Nature Model of Christensen et. al. so that the agents affect the carrying capacity. This leads to a model of species-environment co-evolution where the system tends to have a larger carrying capacity with life than without. We discuss the model as an example of an entropic hierarchy and some implications for Gaia theory.

The Tangled Nature Model—a biologically inspired model of evolutionary ecology—is described, simulated, and analyzed to show its applicability in organization science and organizational ecology. It serves as a conceptual framework for understanding the dynamics in populations of organizations. A salient dynamical feature of this model is the spontaneous generation of a symbiotic group of core organizations. This core, consisting of several dominating species, introduces a mesoscopic level between that of the individual and the whole system. Despite prolonged periods of stability, this core is disrupted at random by parasitic interactions causing sudden core rearrangements. The size distribution of the core organizations is log-normal as predicted by theory and supported by empirical findings. As a simple application of the model, we study the adaptation of organizations to changes in resource availability in terms of population size, population diversity, and ecological efficiency. We find evidence that a temporary reduction in resources forces a consolidation resulting in a sustained increase in overall efficiency, suggesting that such reductions can be applied strategically to drive incremental improvements.

The Gaia hypothesis postulates that life influences Earth's feedback mechanisms to form a self regulating system. This provokes the question: how can global self-regulation evolve? Most models demonstrating environmental regulation involving life have relied on alignment between local selection and global regulation. In these models environment-improving individuals or communities spread to outcompete environment degrading individuals/communities, leading to global regulation, but this depends on local differences in environmental conditions. In contrast, well-mixed components of the Earth system, such as the atmosphere, lack local environmental differentiation. These previous models do not explain how global regulation can emerge in a system with no well defined local environment, or where the local environment is overwhelmed by global effects. We present a model of self-regulation by 'microbes' in an environment with no spatial structure. These microbes affect an abiotic 'temperature' as a byproduct of metabolism. We demonstrate that global self-regulation can arise in the absence of spatial structure in a diverse ecosystem without localised environmental effects. We find that systems can exhibit nutrient limitation and two temperature limitation regimes where the temperature is maintained at a near constant value. During temperature regulation, the total temperature change caused by the microbes is kept near constant by the total population expanding or contracting to absorb the impacts of new mutants on the average affect on the temperature per microbe. Dramatic shifts between low temperature regulation and high temperature regulation can occur when a mutant arises that causes the sign of the temperature effect to change. This result implies that self-regulating feedback loops can arise without the need for spatial structure, weakening criticisms of the Gaia hypothesis that state that with just one Earth, global regulation has no mechanism for developing because natural selection requires selection between multiple entities.