title: Multiple states of environmental regulation in well-mixed model biospheres.
abstract: 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.
authors.affiliation: Earth System Science, University of Exeter, UK. Electronic address: arwen.e.nicholson@gmail.com.
authors.forename: Arwen E
authors.initials: AE
authors.lastname: Nicholson
authors.affiliation: Natural Science and Psychology, Liverpool John Moores University, UK; School of Life Sciences, University of Lincoln, UK.
authors.forename: David M
authors.initials: DM
authors.lastname: Wilkinson
authors.affiliation: Earth System Science, University of Exeter, UK.
authors.forename: Hywel T P
authors.initials: HTP
authors.lastname: Williams
authors.affiliation: Earth System Science, University of Exeter, UK.
authors.forename: Timothy M
authors.initials: TM
authors.lastname: Lenton
mesh_terms: D017753:Ecosystem; D008954:Models, Biological
keywords: Agent-based model; Complexity; Feedback; Gaia; Self-regulation