Prosthetic Symbiosis

Abstract

Symbiosis refers to intimate relations between life forms that involve strong interdependence. Some well-known examples are the associations between flowers and bees, fungi and tree roots, the lichen symbiosis between fungi and algae, and even the mitochondria that make chemical energy for eukaryotic cells. In this list, the examples go from relatively loose associations to ones that are ever more tightly coupled. While the flower and the bee are interdependent products of co-evolution, they are still distinct and distinguishable organisms. For some biologists, including Lynn Margulis and Dorion Sagan (2002, 18), they do not count as examples of symbiosis because they do not live in close or “obligate” contact. The wasp can find other sources of food; the orchid can be pollinated by other species. But mycorrhizae, lichens, and mitochondria are much more integrated. Lichens, the example for which biologists first used the term symbiosis in the late 19th century (Sapp 1994, 6), look like a single organism even if their fungal and algal partners can still be observed separately under a microscope. Mitochondria are fully integrated into cells, living inside their membranes and utterly dependent on their hosts. While the word symbiosis commonly refers to mutual benefit, its more specialized meaning in biology is a continuum from mutualism to one-sided parasitism.In addition to proximity and benefit, symbiosis is about evolution. As Angela Douglas (2010) puts it, evolution as descent with modification is the dominant theory of how organisms vary and acquire new traits. The assumption of both Darwinism and Neo-Darwinism is that descent with modification is gradual. But symbiosis can be its own source of “modification” or evolutionary variation: There is unambiguous evidence that some traits of great evolutionary and ecological importance have been gained laterally from different, often phylogenetically distant, taxa. Some laterally acquired traits are novel for the recipient organism and they can be evolutionary innovations, i.e., “new acquired structures or properties which permit the assumption of a new function.” (Douglas 2010, 1) The term for evolution through symbiosis is symbiogenesis, which refers to a theory first outlined by the Russian biologist and lichenologist Konstantin Mereschkowski in papers from 1905 and 1910. The famous image of the tree of life, which fails to capture this reintegration of lineages noted by Douglas, gives way to a network. The tree does not only branch in one temporal direction, so that all branches lead back to common ancestors. The branches also fuse and reticulate like a mycelium. These two powerful images, tree and mycelium, illustrate two distinct processes of evolution (and there may be more, such as self-organization [Depew and Weber 1994], which no doubt plays a role in symbiosis as well). They are complementary, not mutually exclusive.Theoretical biology has seen many explanations of symbiosis accompanied by symptomatic figures of speech. Even though Margulis and Sagan (2002, 15–17) are quick to denounce the idea that Darwinian competition is “more natural” and less anthropomorphic than cooperation, they go on to criticize anthropomorphism in biology as such—and then to rely heavily on social figures of cooperation. And why not, when scholars have established so many times that metaphor is inseparable from scientific reasoning? But the stakes of anthropomorphism change if I am unsure where to draw the line between humans and the rest of life, whether I even should, and how widely the qualities that anthropomorphism ostensibly projects are already distributed among species. Jan Sapp (1994) details the wide range of social analogies that biologists have used to describe symbiosis, which include friendship, hospitality, neighborliness, oppression, slavery, imprisonment, love, games, cost–benefit economics, and more. 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