The exaptive excellence of spandrels as a term and prototype
The exaptive excellence of spandrels as a term and prototype
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DOI：
10.1073/pnas.94.20.10750
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发表时间：
1997-09-30
影响因子：
11.1
通讯作者：
Gould, SJ
中科院分区：
文献类型：
--
作者：
Gould, SJ
1979年，列文丁和我借用了建筑学术语“spandrel”(以威尼斯的圣马可吊灯为例)，将出现的形式和空间类别指定为设计中另一个决定的必要副产品，而不是对自身直接用途的改编。这一提议产生了一部包含两个批评的大型文献：(I)术语上声称圣马可的斯堪的纳维亚山脉根本不是真正的斯堪的纳维亚山脉，(Ii)概念上声称它们是改编的，而不是副产品。圣马可吊坠的特征被明确定义为悬垂属性--它们必要的数量(4个)和形状(大致为三角形)--是不可避免的建筑副产品，无论吊坠本身的结构属性如何。术语Spandrel可能会从其对二维副产品的特定建筑用途扩展到“剩余空间”的一般性，这一定义适当地包括了圣马可悬垂。进化生物学需要这样一个明确的术语来描述作为副产品出现的特征，而不是适应，无论它们随后产生的作用是什么。生物悬崖的概念--包括这里给出的雌性鬣狗雄性生殖器的例子，蜗牛将脐带作为孵卵室的活生生的使用，巨大爱尔兰鹿的肩峰，以及人类心理的几个关键特征--支撑了对进化解释中过度依赖适应情景的批评。历史起源的原因必须始终与当前的效用分开；它们的融合严重阻碍了对生命历史中形式的进化分析。
In 1979, Lewontin and I borrowed the architectural term ''spandrel'' (using the pendentives of San Marco in Venice as an example) to designate the class of forms and spaces that arise as necessary byproducts of another decision in design, and not as adaptations for direct utility in themselves. This proposal has generated a large literature featuring two critiques: (i) the terminological claim that the spandrels of San Marco are not true spandrels at all and (ii) the conceptual claim that they are adaptations and not byproducts. The features of the San Marco pendentives that,ve explicitly defined as spandrel-properties-their necessary number (four) and shape (roughly triangular)-are inevitable architectural byproducts, whatever the structural attributes of the pendentives themselves. 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. Evolutionary biology needs such an explicit term for features arising as byproducts, rather than adaptations, whatever their subsequent exaptive utility. The concept of biological spandrels-including the examples here given of masculinized genitalia in female hyenas, exaptive use of an umbilicus as a brooding chamber by snails, the shoulder hump of the giant Irish deer, and several key features of human mentality-anchors the critique of overreliance upon adaptive scenarios in evolutionary explanation. 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.