Chemiosmosis, Evolutionary Conflict, and Eukaryotic Symbiosis
Mutualistic symbiosis, in which individuals of different species cooperate and both benefit, has long been an evolutionary puzzle. Why should individuals of two different species cooperate? In this case, as in all others, cooperation is not automatic, but
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Chemiosmosis, Evolutionary Conflict, and Eukaryotic Symbiosis Neil W. Blackstone
Abstract Mutualistic symbiosis, in which individuals of different species cooperate and both benefit, has long been an evolutionary puzzle. Why should individuals of two different species cooperate? In this case, as in all others, cooperation is not automatic, but rather requires the mediation of evolutionary conflicts. In chemiosmosis, redox reactions produce a trans-membrane “proton-motive force” that powers energy-requiring reactions in most organisms. Chemiosmosis may also have a role in conflict mediation. Chemiosmosis rapidly produces considerable amounts of products, increasing the risk of end-product inhibition and the formation of dangerous by-products, such as reactive oxygen species. While several mechanisms can modulate chemiosmosis, potential negative effects can also be ameliorated by simply dispersing excess product into the environment. This “free lunch you are forced to make” can attract individuals of other species leading to groups, in which other organisms share the products that are released into the environment by the chemiosmotic cell or organism. Since the time of Darwin, evolutionary biology has recognized that groups are the key to the evolution of cooperation. With many small groups, chance associations of cooperators can arise, even if cooperation is selected against at the individual level. Groups of cooperators can then outcompete groups of defectors, which do not cooperate. Indeed, numerous symbioses may have arisen in this way, perhaps most notably the symbioses of host cells and chemiosmotic bacteria that gave rise to the eukaryotic cell. Other examples in which one partner relies on chemiosmotic products supplied by the other include lichens, corals or other metazoans and dinoflagellates, sap-feeding insects, and plant–rhizobia and plant–mycorrhiza interactions. More problematic are cases of gut microbiomes—for instance, those of termites, ruminants, and even human beings. Under some but not all circumstances, chemiosmosis can be co-opted into punishing defectors and enforcing cooperation, thus leading to mutualistic symbioses.
N. W. Blackstone (*) Department of Biological Sciences, Northern Illinois University, DeKalb, IL, USA e-mail: [email protected] © Springer Nature Switzerland AG 2020 M. Kloc (ed.), Symbiosis: Cellular, Molecular, Medical and Evolutionary Aspects, Results and Problems in Cell Differentiation 69, https://doi.org/10.1007/978-3-030-51849-3_9
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Introduction
Symbiosis, intimate relationships between different organisms, often at the cellular level, can range from parasitism to commensalism to mutualism. The last, in which both the host and the symbiont benefit, has attracted considerable attention. Why should individuals of two different species cooperate? Indeed, such examples seem to be contrary to Darwinian evolution and for some time were supposed to be just that (Blackstone 2016). Currently, however, explicit considerations of evolutionary conflict are re
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