In the 1970s, R.D. MacElroy coined the term ‘extremophile’ to describe microorganisms that thrive under extreme conditions (MacElroy 1974). This hybrid word transliterates to ‘love of extremes’ and has been studied as a straightforward concept for the past 40 years. In this paper, we discuss several ways the term has been understood in the scientific literature, each of which has different consequences for the distribution and importance of extremophiles. They are, briefly, Human-Centric, at the Edge of life’s habitation of (...) Morphospace, by appeal to Statistical Rarity, described by Objective Limits, and at the Limits of Impossibility for metabolic processes. Importantly, these concepts have co-existed, unacknowledged and conflated, for decades. Confusion and equivocation threaten to follow from the wildly varied inclusion or exclusion of organisms as extremophiles depending on the concept used. Under some conceptions, entire kinds of extremophiles become meaningless. Since our understanding of how life works is shaped by what we take to be its extremes, clarifying extremophily is key for many large-scale projects in biology, ecology, biotechnology and astrobiology. In what follows, we proceed as if a noncontroversial account of life is possible and that it is possible to find complex chemistry in the Universe that is similar enough to Life on Earth such that both may be considered instances of ‘life’ (but see Mariscal & Doolittle 2018). We raise, but do not address, the questions of whether the distribution of Life on Earth is representative of what we may find elsewhere in the Universe, whether the same kinds of extremophiles would exist given a replay of the tape of life. Additionally, each of these concepts assumes life based on some sort of biochemistry in this universe, effectively setting aside claims made by some artificial life proponents that their digital organisms are genuine examples of life (Langton 1989, Ray 1995). On the distinction between extremophilic and extremotolerant, we note that all accounts accept the latter as a broader category than the former, since ‘phily’ means love and tolerance is a prerequisite for love under any conception. Indeed, there will be many extremely impoverished environments where tolerance is the only option and thriving is precluded, e.g. Bacillus marismortui was extracted and grown from 250-million-year-old salt crystals in the Permian Salado Formation in an inactive yet persistent state (Vreeland et al., 2000). We also note that extremophily, as a functional category, is potentially applicable at many levels of the biological hierarchy. Extremophily at one level does not necessarily extend to higher and lower levels. For instance, a microorganism in isolation might be quite intolerant to certain environmental conditions yet flourish when subjected to the same conditions as a community or natural biofilm. Alternatively, a protein molecule might be quite stable or active under certain conditions even if the optimal environment for the organism containing it is far more mesophilic. There is an industry of artificially selecting organisms and proteins to adapt to extreme environments (see van den Burg & Eijsink 2002), providing some justification to consider ‘functioning at extremes’ as a worthwhile category of investigation. Finally, we also note certain physico-chemical ranges are rarely considered with respect to extremophily – timespan, size, nutrient availability (Hoehler & Jørgensen 2013), etc. – as well as some biological parameters – abundance, isolation, competition, etc. Perhaps scientific interest must also come into play as to the reason these criteria are not considered relevant. We return to this issue later. In the next section, we give five definitions of extremophily, show their benefits, drawbacks, and unintended yet unavoidable consequences. These arguments are summarized in Table 1 and represented visually in Figures 1 and 2. Given research on polyextremophiles, it seems Figure 2 is a more plausible representation of the state of current knowledge than Figure 1 (Harrison et al. 2013). Life is patchily distributed across various dimensions, which may reflect its contingent history, poor sampling, or (perhaps) fundamental limits. Figure 3 shows the conceptual flowchart for all of these views. In the following section, we take a step back to ask whether we should choose between these definitions and how such a judgment could be made. We argue for a limited pluralism, in which some, but not all, of the concepts are acceptable relative to certain practical and theoretical aims. (shrink)

The osmotic activity produced by internal, non‐permeable, anionic nucleic acids and metabolites causes a persistent and life‐threatening cell swelling, or cellular edema, produced by the Gibbs‐Donnan effect. This evolutionary‐critical osmotic challenge must have been resolved by LUCA or its ancestors, but we lack a cell‐physiology look into the biophysical constraints to the solutions. Like mycoplasma, early cells conceivably preserved their volume with Cl−, Na+, and K+‐channels, Na+/H+‐exchangers, and a light‐dependent bacteriorhodopsin‐like H+‐pump. Here, I simulated protocells having these ionic‐permeabilities and inhabiting (...) an oceanic pond before the Great‐Oxygenation‐Event. Protocells showed better volume control and stable resting potentials at lower external pH and higher temperatures, favoring a certain type of extremophile life. Prevention of Na+‐influx at night, with low bacteriorhodopsin activity, required deep shutdown of highly voltage‐sensitive Na+‐channels and extremely selective K+‐channels, two conserved features essential for modern neuronal encoding. The Gibbs‐Donnan effect universality implies that extraterrestrial cells, if they exist, may reveal similar volume‐controlling mechanisms. (shrink)

In contrast to vertical gene transfer from parent to offspring, horizontal (or lateral) gene transfer moves genetic information between different species. Bacteria and archaea often adapt through horizontal gene transfer. Recent analyses indicate that eukaryotic genomes, too, have acquired numerous genes via horizontal transfer from prokaryotes and other lineages. Based on this we raise the hypothesis that horizontally acquired genes may have contributed more to adaptive evolution of eukaryotes than previously assumed. Current candidate sets of horizontally acquired eukaryotic genes may (...) just be the tip of an iceberg. We have recently shown that adaptation of the thermoacidophilic red alga Galdieria sulphuraria to its hot, acid, toxic‐metal laden, volcanic environment was facilitated by the acquisition of numerous genes from extremophile bacteria and archaea. Other recently published examples of horizontal acquisitions involved in adaptation include ice‐binding proteins in marine algae, enzymes for carotenoid biosynthesis in aphids, and genes involved in fungal metabolism.Editor's suggested further reading in BioEssays Jumping the fine LINE between species: Horizontal transfer of transposable elements in animals catalyses genome evolution Abstract. (shrink)

Astrobiologists are aware that extraterrestrial life might differ from known life, and considerable thought has been given to possible signatures associated with weird forms of life on other planets. So far, however, very little attention has been paid to the possibility that our own planet might also host communities of weird life. If life arises readily in Earth-like conditions, as many astrobiologists contend, then it may well have formed many times on Earth itself, which raises the question whether one or (...) more shadow biospheres have existed in the past or still exist today. In this paper, we discuss possible signatures of weird life and outline some simple strategies for seeking evidence of a shadow biosphere. Key Words: Weird life—Multiple origins of life—Biogenesis—Biomarkers—Extremophiles—Alternative biochemistry. Astrobiology 9, 241–249. (shrink)

Gaia, the grand idea -- Good citizens or selfish genes? -- Life at the edge : lessons from extremophiles -- Temperature paces life -- Icehouse earth -- Given enough time -- Evolutionary innovations and environmental change -- A stable or an unstable world? -- The puzzle of life's long persistence -- Conclusions.

A hard difficulty in Astrobiology is the precise definition of what life is. All living beings have a cellular structure, so it is not possible to have a broader concept of life hence the search for extraterrestrial life is restricted to extraterrestrial cells. Earth is an astronomical rarity because it is difficult for a planet to present liquid water on the surface. Two antagonistic bioethical principles arise: planetary protection and terraforming. Planetary protection is based on the fear of interplanetary cross-infection (...) and possible ecological damages caused by alien living beings. Terraforming is the intention of modifying the environmental conditions of the neighbouring planets in such a way that human colonisation would be possible. The synthesis of this antagonism is ecopoiesis, a concept related to the creation of new ecosystems in other planets. Since all the multicellular biodiversity requires oxygen to survive, only extremophile microorganisms could survive in other planets. So, it could be carried out a simulation of a meteorite by taking to other planets portions of the terrestrial permafrost, or ocean or soil, so that if a single species could grow, a new ecosystem would start, as well as a new Natural History. As a conclusion, ecopoiesis should be the bioethical principle to guide practices and research in Astrobiology. (shrink)

Is listening to music like looking through a kaleidoscope? Formalists contend that music is meaningless. Most music theorists concede that this austere thesis is surprisingly close to the truth. Nevertheless, they refute formalism with a little band of diffusely referential phenomena, such as musical quotation, onomatopoeia, exemplification, and leitmotifs. These curiosities ought to be pressed into a new campaign against assumptions that vagueness can only arise in the semantically lush setting of language. Just as the discovery of extremophilic bacteria led (...) biologists to revise their opinions about the scope and preconditions of life, the marginal forms of reference that survive in the semantic desert of absolute music should lead philosophers to revise their assumptions much about scope and preconditions of vagueness. (shrink)

In concert with the selective pressures affecting protein folding and function in the extreme environments in which they can exist, proteins in Archaea have evolved to present permanent molecular adaptations at the amino acid sequence level. Such adaptations may not, however, suffice when Archaea encounter transient changes in their surroundings. Post‐translational modifications offer a rapid and reversible layer of adaptation for proteins to cope with such situations. Here, it is proposed that Archaea further augment their ability to survive changing growth (...) conditions by modifying the extent, position, and, where relevant, the composition of different post‐translational modifications, as a function of the environment. Support for this hypothesis comes from recent reports describing how patterns of protein glycosylation, methylation, and other post‐translational modifications of archaeal proteins are altered in response to environmental change. Indeed, adjusting post‐translational modifications as a means to cope with environmental variability may also hold true beyond the Archaea. (shrink)

Two categories of life are currently recognized—chemosynthetic and photosynthetic—indicating their principal free energy resource as either chemicals or electromagnetic radiation. Building on recent developments in thermodynamics, we posit a third category of life—thermosynthetic life (TL)—which relies on environmental heat rather than traditional free energy sources. Since thermal energy is more abundant than chemicals or light in many settings, thermosynthesis offers compelling evolutionary possibilities for new life forms. Based on variants of standard cellular machinery, a physical model is proposed for the (...) conversion of thermal energy into biochemical work. Conditions favorable to thermosynthetic life and prospects for its discovery are assessed. Terrestrially, deep-subsurface unicellular anaerobic superthermophiles are deduced to be likely TL candidates. (shrink)