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Provided for Non-Commercial Research and Educational Use. Not for Reproduction, Distribution Or Commercial Use Provided for non-commercial research and educational use. Not for reproduction, distribution or commercial use. This article was originally published in the Encyclopedia of Evolutionary Biology published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of the author's institution, for non- commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who you know, and providing a copy to your institution's administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution's website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's permissions site at: http://www.elsevier.com/locate/permissionusematerial Gontier, N. (2016) Symbiosis, History of. In: Kliman, R.M. (ed.), Encyclopedia of Evolutionary Biology. vol. 4, pp. 272–281. Oxford: Academic Press. © 2016 Elsevier Inc. All rights reserved. Author's personal copy Symbiosis, History of N Gontier, University of Lisbon, Lisbon, Portugal r 2016 Elsevier Inc. All rights reserved. Glossary Transformation and metamorphosis are precursors to Deism The idea that the universe follows natural, transmutation and evolution theory. mechanical laws even though it was created by a deity. The Symbiogenesis Evolutionary mechanism that results from deity is presumed not to intervene after its original creation hereditary symbiosis. act. Deism was introduced by Natural Philosophers and is Symbiont Any organism that engages in a symbiotic foundational for nineteenth century Natural History schools. relation with one or more organisms – often used to Ecology The study of the ‘household’ of organisms, designate the smaller partner in the symbiosis. broadly conceived as including biotic and abiotic Theism The idea that a supernatural being has created the environmental interactions, often depicted in hierarchy universe and intervenes in creation at will. This belief is theories ranging from populations and species to foundational for the Semitic religions as well as early ecosystems, biomes, and biospheres. Natural Theology schools. Metamorphosis Pre-evolutionary idea that living organisms can transform, or change in form. Defining Symbiosis called Social-Contract theoreticians because they founded sociopolitical, secular thought by debating the fair and just Symbiosis refers to the phenomenon whereby two or distribution of ‘public goods’ and ‘natural resources’ across more organisms with distinct genealogical, evolutionary nations, the ‘natural’‘division of labor,’ and the ‘common histories live in close association with one another (de Bary, goods’ (shared belief states) that socially bond individual 1878, 1879). Together, the host and its symbionts form a new citizens into a ‘common wealth’ (a nation) (Gontier, 2009). biological entity that is sometimes called a superorganism Their theorizing on the ‘naturalness’ by which humans ‘live (Spencer, 1876; Carrapiço, 2015), holobiont (Margulis, 1991; together,’ and what the most righteous sociopolitical societal Guerrero et al., 2013), or symbiome (Sapp, 2003: 33); and this structure is, would come to define liberal and social thought newly formed entity is considered to be a single unit, either of and lead to the rise of the major secular sociopolitical doctrines natural selection (through its hologenome, Rosenberg et al., of the nineteenth and twentieth centuries (libertarianism, lib- 2007; Zilber-Rosenberg and Rosenberg, 2008), or of other eralism, socialism, Marxism (communism), communitarian- evolutionary mechanisms. Symbionts can be acquired both ism, national socialism, and democracy). These sociopolitical vertically (during phylogenesis) and horizontally (during ideologies were formulated by drawing idealized analogies ontogenesis). When symbiosis becomes obligate and heredi- with the animal world (often in the form of fables), which tary, it can lead to evolution by symbiogenesis (Margulis and in turn founded preliminary researches on ‘animal societies’ Dolan, 2000: 157). But even without causing symbiogenesis, that developed into naturalistic, and eventually evolutionary symbiotic associations can affect an organism's adaptation, research on biological species. reproduction and fitness, species extinction or speciation, and It is well-known that Darwin (Burrow, 1972; Barrett, 1977; symbiosis can influence ecological systems (Brucker and Bowler, 1983; Smocovitis, 1996) was inspired by: Bordenstein, 2012; Margulis, 1991; Zook, 2015). 1. Adam Smith’s liberal idea of free-market economy and his metaphor of the ‘invisible hand’; 2. Thomas Malthus’s notions of ‘scarcity of resources’ and Is Nature ‘Red in Tooth and Claw’ or ‘Social’–Origins ‘struggle for existence’; of Symbiosis Research in Natural History Research 3. Herbert Spencer and Thomas Huxley’s liberal ideas on and Its Relation to the Sociopolitical Sciences sociocultural ‘progress’; 4. Jean Baptiste Lamarck’s ideas of ‘adaptation’ and tele- Research on symbiosis originated in nineteenth century ological progress in biology; European Natural History Research. Natural History Research 5. Francis Bacon’s ‘induction theory’; marked the beginning of the modern sciences and en- 6. Charles Lyell’s ‘uniformitarianism’; and compassed the physical, biological, and cultural sciences 7. Wilhelm von Humboldt and Auguste Schleicher’s ‘natural (Gontier, 2011). Rather than adhering to ‘divine laws’ speci- genealogies’ of the Indo-European languages. fied in religious writings, utopian scholars such as Thomas More or Bernard Mandeville, and moral philosophers such as The early Darwinists did not distinguish the natural from Thomas Hobbes, Adam Smith, David Hume, and Jean-Jacques the sociocultural sciences, and their evolutionary theories Rousseau would search for ‘natural laws,’ and transition from were a natural extension of sociopolitical, liberal Hobbesian a theist to a deist worldview. These moral philosophers are thought – how the latter ideology defined the ‘natural’ 272 Encyclopedia of Evolutionary Biology, Volume 4 doi:10.1016/B978-0-12-800049-6.00015-9 Author's personal copy Symbiosis, History of 273 condition as well as the sociocultural and political life of which in turn specify the nature of the social lives of animals, humans and animals. For Hobbes (1651/2010), humans were in particular how they establish communal living and how like wolves, who in a ‘natural state’ found themselves at ‘war’ they share their food resources. In this context, he would with other humans because they wanted to defend their introduce the terms commensalism (commensaux), mutualism and individual freedom. Early Darwinists extended these ideas parasitism (Table 1). to the whole of nature and, in the words of Alfred Lord For van Beneden, Tennyson, they saw nature as ‘red in tooth and claw.’ A struggle for existence results from a scarcity of resources, When taking a closer look at the animal world, it does not take long leading to a natural selection of the fit, at the expense of the to find more than one analogy with human society. If I’m allowed to maladaptive (Huxley, 1888; Bouglé, 1909). say so, there is not one social position found in human society that is Etymological analyses of symbiosis jargon demonstrate not also found in the animal kingdom. Most of the animals live peacefully from the fruits of their labors, and exercise a profession that, prior to the introduction into biology, many of the that gives them life. But, at the sideline of these honest industrials, foundational concepts were also used to define sociocultural we also find miserable ones, who cannot do without the assistance of human relations and sociopolitical ideologies (Table 1). Early their neighbors, some of whom establish themselves as parasites in fi symbiologists, however, applied less-liberal or non-liberal, their organs, others as commensals (commensaux) that take pro t from the gains (labors) of the honest. (Van Beneden, 1875:2–3, my often socialist and communitarian language to characterize the translation) living arrangements present in the natural world – ideas that were inspired by Rousseau’s social-contract theory. Rousseau (1762/2001) understood humans as ‘good natured’ and in- Commensals or messmates are merely companions at the herently social beings that became corrupted by artificial so- table, they are allowed to dine with the host and feed on their cieties. The human family, one of the basic units of law, is neighbor’s catch. They can live inside or onside their host typified by its sharing of resources, and societal living is ideally (what we today call endo- or ectosymbiosis), and they can either characterized by reciprocal exchange and a fair (re)distribution live independently or forever remain fixed (today called fac- of natural resources. Reciprocal and altruistic social rule- ultative and obligate symbiosis). Parasites are those animals who following enables the formation of a ‘social-contract’ that in live at the expense of their neighbor, they take advantage of turn allows for the establishment of a ‘higher, common good’ their host and can endanger its life. Mutualists are animals that that founds the resurrection of the welfare state. As members live onto one another without being either parasites or mess- of the welfare state, persons become citizens that create a mates. They receive
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