Spandrel (biology)

In evolutionary biology, a spandrel is a phenotypic trait that is a byproduct of the evolution of some other characteristic, rather than a direct product of adaptive selection. Stephen Jay Gould and Richard Lewontin brought the term into biology in their 1979 paper "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme". Adaptationism is a point of view that sees most organismal traits as adaptive products of natural selection. Gould and Lewontin sought to temper what they saw as adaptationist bias by promoting a more structuralist view of evolution. The term "spandrel" originates from architecture, where it refers to the roughly triangular spaces between the top of an arch and the ceiling. The term was coined by paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their paper "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme" (1979). Evolutionary biologist Günter P. Wagner called the paper "the most influential structuralist manifesto". In their paper, Gould and Lewontin employed the analogy of spandrels in Renaissance architecture, such as the curved areas of masonry between arches supporting a dome that arise as a consequence of decisions about the shape of the arches and the base of the dome, rather than being designed for the artistic purposes for which they were often employed. The authors singled out properties like the necessary number of four spandrels and their specific three-dimensional shape. At the time, it was thought in the scientific community that everything an animal has developed that has a positive effect on that animal's fitness was due to natural selection or some adaptation. Gould and Lewontin proposed an alternative hypothesis: that due to adaptation and natural selection, byproducts are also formed. Because these byproducts of adaptations that had no real relative advantage to survival, they were termed spandrels.