The distribution of fitness effects (DFE) for new mutations is fundamental for many aspects of population and quantitative genetics. In this study, we have inferred the DFE in the single-celled algaChlamydomonas reinhardtiiby estimating changes in the frequencies of 254 spontaneous mutations under experimental evolution and equating the frequency changes of linked mutations with their selection coefficients. We generated seven populations of recombinant haplotypes by crossing seven independently derived mutation accumulation lines carrying an average of 36 mutations in the haploid state to a mutation-free strain of the same genotype. We then allowed the populations to evolve under natural selection in the laboratory by serial transfer in liquid culture. We observed substantial and repeatable changes in the frequencies of many groups of linked mutations, and, surprisingly, as many mutations were observed to increase as decrease in frequency. Mutation frequencies were highly repeatable among replicates, suggesting that selection was the cause of the observed allele frequency changes. We developed a Bayesian Monte Carlo Markov Chain method to infer the DFE. This computes the likelihood of the observed distribution of changes of frequency, and obtains the posterior distribution of the selective effects of individual mutations, while assuming a two-sided gamma distribution of effects. We infer that the DFE is a highly leptokurtic distribution, and that approximately equal proportions of mutations have positive and negative effects on fitness. This result is consistent with what we have observed in previous work on a differentC.reinhardtiistrain, and suggests that a high fraction of new spontaneously arisen mutations are advantageous in a simple laboratory environment.

Abstract De novo mutations are central for evolution, since they provide the raw material for natural selection by regenerating genetic variation. However, studying de novo mutations is challenging and is generally restricted to model species, so we have a limited understanding of the evolution of the mutation rate and spectrum between closely related species. Here, we present a mutation accumulation (MA) experiment to study de novo mutation in the unicellular green alga Chlamydomonas incerta and perform comparative analyses with its closest known relative, Chlamydomonas reinhardtii. Using whole-genome sequencing data, we estimate that the median single nucleotide mutation (SNM) rate in C. incerta is μ = 7.6 × 10−10, and is highly variable between MA lines, ranging from μ = 0.35 × 10−10 to μ = 131.7 × 10−10. The SNM rate is strongly positively correlated with the mutation rate for insertions and deletions between lines (r > 0.97). We infer that the genomic factors associated with variation in the mutation rate are similar to those in C. reinhardtii, allowing for cross-prediction between species. Among these genomic factors, sequence context and complexity are more important than GC content. With the exception of a remarkably high C→T bias, the SNM spectrum differs markedly between the two Chlamydomonas species. Our results suggest that similar genomic and biological characteristics may result in a similar mutation rate in the two species, whereas the SNM spectrum has more freedom to diverge.

Extreme environmental fluctuations such as marine heatwaves (MHWs) can have devastating effects on ecosystem health and functioning through rapid population declines and destabilization of trophic interactions. However, recent studies have highlighted that population tolerance to MHWs is variable, with some populations even benefitting from MHWs. A number of factors can explain variation in responses between populations including their genetic variation, previous thermal experience and the cumulative heatwave intensity (°C d) of the heatwave itself. We disentangle the contributions of these factors on population mortality and post-heatwave growth rates by experimentally simulating heatwaves (7.5 or 9.2°C, for up to 9 days) for three genotypes of the Southern Ocean diatom Actinocyclus actinochilus. The effects of simulated heatwaves on mortality and population growth rates varied with genotype, thermal experience and the cumulative intensity of the heatwave itself. Firstly, hotter and longer heatwaves increased mortality and decreased post-heatwave growth rates relative to milder, shorter heatwaves. Secondly, growth above the thermal optimum before heatwaves exacerbated heatwave-associated negative effects, leading to increased mortality during heatwaves and slower growth after heatwaves. Thirdly, hotter and longer heatwaves resulted in more pronounced changes to thermal optima (Topt) immediately following heatwaves. Finally, there is substantial intraspecific variation in post-heatwave growth rates. Our findings shed light on the potential of Southern Ocean diatoms to tolerate MHWs, which will increase both in frequency and in intensity under future climate change.

The term "habitability" is pervasive throughout the space sciences and astrobiology literature and is broadly used to describe an environment's ability to support life. Here, we argue that, while it is fundamentally a binary matter whether an organism can persist in an environment or not, these binary assessments lead to continuous ecological measurements that are often collected under the umbrella term "habitability" by astrobiologists. Although the use of habitability in this way has provided a framework for those studying the potential of environments to support life, including comparative analyses between terrestrial and extraterrestrial environments, it can also generate confusion and limit interdisciplinary understanding. Namely, differing ecological metrics used as proxies for habitability can yield differing conclusions depending upon the metrics chosen. Therefore, we suggest that in this continuous sense, the terms habitable and habitability lose meaning unless the specific scientific question and biological metric chosen to address it are defined. As a corollary, the search for universal single metrics to make habitability assessments is not to be encouraged, and as we argue, attempting to do so would oversimply analyses of the ability of environments to support life.

Microbes are active agents of environmental change. From the depths of the Earth's crust to the heights of the upper atmosphere, microorganisms alter the physicochemical conditions surrounding them. Their activity provides important ecosystem services to the wider biosphere, making essential elements such as carbon, sulfur, phosphorus, and iron available for higher organisms. One such environmental interaction is the weathering of minerals and rocks by microbial communities, a key process that underpins soil formation and global biogeochemical cycles. By facilitating mineral dissolution and rock degradation, microbes enhance the release of elements from their geological reservoirs and perform significant elemental transformations. Under what conditions microbes perform these activities and to what extent they impact their surrounding environment, are key topics in geobiology. In this chapter, various aspects of microbial mineral and rock weathering will be explored. Key concepts and terminology will be introduced, followed by an overview of the mechanisms used by microorganisms to perform weathering activity. Sections covering the methodological approaches used by researchers to study microbial rock weathering processes, including detecting the traces or “biosignatures” such microbial activity leaves behind on geological materials, will be provided. The effect of microbial rock weathering on the wider microbial ecosystem, including the endurance and functional capacity of microbial communities, is also explored. Finally, this chapter will discuss the emerging field of microbial biogeomorphology, the study of how biological activity at the micron scale can impact the environment at meter to kilometer scales, contributing to the processes that shape Earth's landscapes.

Environmental cues that regulate motility are poorly understood, but specific carbon and nitrogen sources, such as casamino acids (CAA), are known to stimulate motility in model organisms. However, natural environments are commonly more nutrient-limited than laboratory growth media, and the effect of energy-rich CAA on the motility of oligotrophic microorganisms is unknown. In this study, an extreme oligocarbotroph, Variovorax paradoxus YC1, was isolated from weathered shale rock within a disused mine level in North Yorkshire, UK. The addition of 0.1% CAA to minimal media significantly reduced the motility of YC1 after 72 h and inhibited swimming motility resulting in enhanced surface growth. We propose this response to CAA is a physiological adaptation to oligotrophy, facilitating the colonization of nutrient-rich environments.

Microbial rock weathering of shale forms an important part of global biogeochemical cycling and soil formation. Culture-independent analyses have revealed diverse microbial communities in weathered shale environments, yet few studies have attempted to discern the functional ecology of such communities in relation to their rock weathering capabilities. In this study, phenotypic plate assays were used to determine the abundance of microbes with different rock weathering phenotypic traits in weathered shale environments. A physicochemical parameter (pH) is shown to influence the abundance of aerobic rock weathering microbes in weathered shale. Iron and manganese oxidizers were restricted to acidic environments while siderophore producing and alkaline phosphatase producing microbes were largely confined to pH neutral environments. Furthermore, a clear separation in the spatial distribution of aerobic-iron oxidizing and siderophore-producing microbes, as defined by a pH gradient across the sites sampled, was demonstrated. Phylogenetic analysis of isolates revealed that siderophore-producing and alkaline phosphatase producing bacteria belonged to commonly identified rock weathering genera including Arthrobacter, Pseudomonas, and Streptomyces. These results enhance our understanding of how physicochemical parameters can define the composition and rock weathering potential of microbial communities.

Spontaneous mutations are the source of new genetic variation and are thus central to the evolutionary process. In molecular evolution and quantitative genetics, the nature of genetic variation depends critically on the distribution of effects of mutations on fitness and other quantitative traits. Spontaneous mutation accumulation (MA) experiments have been the principal approach for investigating the overall rate of occurrence and cumulative effect of mutations but have not allowed the phenotypic effects of individual mutations to be studied directly. Here, we crossed MA lines of the green alga Chlamydomonas reinhardtii with its unmutated ancestral strain to create haploid recombinant lines, each carrying an average of 50% of the accumulated mutations in a large number of combinations. With the aid of the genome sequences of the MA lines, we inferred the genotypes of the mutations, assayed their growth rate as a measure of fitness, and inferred the distribution of fitness effects (DFE) using a Bayesian mixture model. We infer that the DFE is highly leptokurtic (L-shaped). Of mutations with absolute fitness effects exceeding 1%, about one-sixth increase fitness in the laboratory environment. The inferred distribution of effects for deleterious mutations is consistent with a strong role for nearly neutral evolution. Specifically, such a distribution predicts that nucleotide variation and genetic variation for quantitative traits will be insensitive to change in the effective population size.

Microbial pyrite oxidation is an important driver of biological weathering within shale, making a significant contribution toward biogeochemical cycling, bedrock expansion, and soil formation. These processes are of global importance, both within natural systems and in anthropogenic environments. Despite its significance, there is a lack of research that directly investigates microbe- pyrite interactions within shale. In this study, we use both field and laboratory approaches to inspect microbial pyrite oxidation in weathered shale environments within North Yorkshire, UK. Incubation of polished pyrite samples within iron-oxidizing enrichment cultures in vitro resulted in extensive colonization and surface pitting, demonstrating the weathering potential of shale microbial communities. Mineral samples were buried for 1 year within the floor of a shale rock mine, to explore pyrite bioweathering in situ. Image analysis revealed the formation of dissolution channels by microbial filaments, a novel mechanism of pyrite oxidation that broadens the taxonomic range of known microbe-pyrite interactions in weathered shale.

The UK Centre for Astrobiology (UKCA) was set up in 2011 as a virtual center to contribute to astrobiology research, education, and outreach. After 5 years, we describe this center and its work in each of these areas. Its research has focused on studying life in extreme environments, the limits of life on Earth, and implications for habitability elsewhere. Among its research infrastructure projects, UKCA has assembled an underground astrobiology laboratory that has hosted a deep subsurface planetary analog program, and it has developed new flow-through systems to study extraterrestrial aqueous environments. UKCA has used this research backdrop to develop education programs in astrobiology, including a massive open online course in astrobiology that has attracted over 120,000 students, a teacher training program, and an initiative to take astrobiology into prisons. In this paper, we review these activities and others with a particular focus on providing lessons to others who may consider setting up an astrobiology center, institute, or science facility. We discuss experience in integrating astrobiology research into teaching and education activities. Key Words: Astrobiology-Centre-Education-Subsurface-Analog research. Astrobiology 18, 224-243.

Habitability is a widely used word in the geoscience, planetary science, and astrobiology literature, but what does it mean? In this review on habitability, we define it as the ability of an environment to support the activity of at least one known organism. We adopt a binary definition of "habitability" and a "habitable environment." An environment either can or cannot sustain a given organism. However, environments such as entire planets might be capable of supporting more or less species diversity or biomass compared with that of Earth. A clarity in understanding habitability can be obtained by defining instantaneous habitability as the conditions at any given time in a given environment required to sustain the activity of at least one known organism, and continuous planetary habitability as the capacity of a planetary body to sustain habitable conditions on some areas of its surface or within its interior over geological timescales. We also distinguish between surface liquid water worlds (such as Earth) that can sustain liquid water on their surfaces and interior liquid water worlds, such as icy moons and terrestrial-type rocky planets with liquid water only in their interiors. This distinction is important since, while the former can potentially sustain habitable conditions for oxygenic photosynthesis that leads to the rise of atmospheric oxygen and potentially complex multicellularity and intelligence over geological timescales, the latter are unlikely to. Habitable environments do not need to contain life. Although the decoupling of habitability and the presence of life may be rare on Earth, it may be important for understanding the habitability of other planetary bodies.

coveR Fred Hoyle (shown here in 1965) was a giant of astrophysics whose ideas influenced a generation of scientists.

Toby Samuels and Natasha Nicholson report on a debate over the pros and cons of turning humans into martians.

What is astrobiology? Which fields does it comprise and what makes an astrobiologist? Ask five scientists and you may end up with six different definitions. This issue was raised at the first symposium of the European network of Astrobiology Graduates (AbGradE), held last year in Edinburgh, when discussing whether the attendees' fields of study were represented in the astrobiology community.

Toby Samuels, Lena Noack, Cyprien Verseux and Paloma Serrano look back at the first symposium of the new network Astrobiology Graduates in Europe, held in Edinburgh in October 2014.

Manipulation of the soil microbiota associated with crop plants has huge promise for the control of crop pathogens. However, to fully realize this potential we need a better understanding of the relationship between the soil environment and the genes and phenotypes that enable microbes to colonize plants and contribute to biocontrol. A recent 2 years of investigation into the effect of wheat variety on second year crop yield in the context of take-all fungal infection presented the opportunity to examine soil microbiomes under closely defined field conditions. Amplicon sequencing of second year soil samples showed that Pseudomonas spp. were particularly affected by the wheat cultivar grown in year one. Consequently, 318 rhizosphere-associated Pseudomonas fluorescens strains were isolated and characterized across a variety of genetic and phenotypic traits. Again, the wheat variety grown in the first year of the study was shown to exert considerable selective pressure on both the extent and nature of Pseudomonas genomic diversity. Furthermore, multiple significant correlations were identified within the phenotypic/genetic structure of the Pseudomonas population, and between individual genotypes and the external wheat field environment. The approach outlined here has considerable future potential for our understanding of plant-microbe interactions, and for the broader analysis of complex microbial communities.