Spandrel (biology)

The term "spandrel" originates from architecture, where it refers to the roughly triangular spaces between the top of an arch and the ceiling.

Etymology

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 widely 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 had no real relative advantage to survival, they were termed spandrels. In the biological sense, a "spandrel" might result from a requirement inherent in the body plan of an organism, or as a byproduct of some other constraint on adaptive evolution.

In response to the position that spandrels are just small, unimportant byproducts, Gould and Lewontin argue that "we must not recognize that small means unimportant. Spandrels can be as prominent as primary adaptations". A main example used by Gould and Lewontin is the human brain. Many secondary processes and actions come in addition to the main functions of the human brain. These secondary processes and thoughts can eventually turn into an adaptation or provide a fitness advantage to humans. Just because something is a secondary trait or byproduct of an adaptation does not mean it has no use.

In 1982, Gould and Vrba introduced the term "exaptation" for characteristics that enhance fitness in their present role but were not built for that role by natural selection. Exaptations may be divided into two subcategories: pre-adaptations and spandrels. Spandrels are characteristics that did not originate by the direct action of natural selection and that were later co-opted for a current use. Gould saw the term to be optimally suited for evolutionary biology for "the concept of a nonadaptive architectural by-product of definite and necessary form – a structure of particular size and shape that then becomes available for later and secondary utility".

Criticism of the term

Gould and Lewontin's proposal generated a large literature of critique, which Gould characterised as being grounded in two ways. First, a terminological claim was offered that the "spandrels" of Basilica di San Marco were not spandrels at all, but rather were pendentives. Gould responded, "The term spandrel may be extended from its particular architectural use for two-dimensional byproducts to the generality of 'spaces left over', a definition that properly includes the San Marco pendentives."

Other critics, such as Daniel Dennett, further claimed (in Darwin's Dangerous Idea and elsewhere) that these pendentives are not merely architectural by-products as Gould and Lewontin supposed. Dennett argues that alternatives to pendentives, such as corbels or squinches, would have served equally well from an architectural standpoint, but pendentives were deliberately selected due to their aesthetic value. Critics such as H. Allen Orr argued that Lewontin and Gould's oversight in this regard illustrates their underestimation of the pervasiveness of adaptations found in nature.

Response to criticism

Gould responded that critics ignore that later selective value is a separate issue from origination as necessary consequences of structure; he summarised his use of the term 'spandrel' in 1997: "Evolutionary biology needs such an explicit term for features arising as byproducts, rather than adaptations, whatever their subsequent exaptive utility ... Causes of historical origin must always be separated from current utilities; their conflation has seriously hampered the evolutionary analysis of form in the history of life." Gould cites the masculinized genitalia of female hyenas and the brooding chamber of some snails as examples of evolutionary spandrels.

Gould (1991) outlines some considerations for grounds for assigning or denying a structure the status of spandrel, pointing first to the fact that a structure originating as a spandrel through primary exaptation may have been further crafted for its current utility by a suite of secondary adaptations, thus the grounds of how well crafted a structure is for a function cannot be used as grounds for assigning or denying spandrel status. The nature of the current utility of a structure also does not provide a basis for assigning or denying spandrel status, nor does he see the origin of a structure as having any relationship to the extent or vitality of a later co-opted role, but places importance on the later evolutionary meaning of a structure. This seems to imply that the design and secondary utilization of spandrels may feed back into the evolutionary process and thus determine major features of the entire structure. The grounds Gould does accept to have validity in assigning or denying a structure the status of spandrel are historical order and comparative anatomy. Historical order involves the use of historical evidence to determine which feature arose as a primary adaptation and which one appeared subsequently as a co-opted by-product. In the absence of historical evidence, inferences are drawn about the evolution of a structure through comparative anatomy. Evidence is obtained by comparing current examples of the structure in a cladistic context and by subsequently trying to determine a historical order from the distribution yielded by tabulation.