扩展适应

扩展适应（英语：exaptation）是指在演化过程中一种特征的功能发生了变化。例如，如果一个特征最初演化出来是为了实现某种特定的功能，但后来它又有了另一种其他用途，就可称之为“扩展适应”。鸟类羽毛的演化便是一个经典的例子：羽毛最初是为了保温而演化出来的，但在后来羽毛变得开始适应于飞翔。对扩展适应的兴趣与进化的过程和产物都息息相关：创造复杂特征（英语：Complex traits）的过程和可能不完善的产物（功能，解剖结构，生化试剂等等）^[1]^[2] 。扩展适应是由美国古生物学家斯蒂芬·古尔德（Stephen Jay Gould）和伊丽莎白·弗巴（英语：Elisabeth Vrba）（Elisabeth Vrba）提出的，以取代他们认为是目的论的术语“预适应”（英语：pre-adaptation）^[3]。

历史和定义

特征的功能在其进化历史中可能发生变化的想法起源于查尔斯·达尔文（Charles Darwin）（(Darwin 1859)）。多年以来，这种现象被标记为“预适应”，但由于“预适应”这一术语隐含了生物学中的目的论，与自然选择的基本原理相悖，故现已被“扩展适应”所取代^[4]。

1982年，斯蒂芬·古尔德和伊丽莎白·弗巴（英语：Elisabeth Vrba）引入了“扩展适应”一词，这是一些学者们^[a]探索的想法。

实例

扩展适应包括羽毛的共同选择，羽毛最初是为用于保温的和用于展示的功能而演化的，后来被用于鸟类的飞行。另一个例子则是早期鱼类的肺，它们后来演化为陆生脊椎动物的肺，但同时还通过扩展适应成为了一些鱼类体内用来调节浮力的器官——鳔。^[7]

注释

[a]
See Jacob (1977)^[5] and Mayr (1982)^[6] for references.

参考文献

[1]
Bock, W.J. Preadaptation and multiple evolutionary pathways. Evolution. 1959, 13 (2): 194–211. JSTOR 2405873. doi:10.2307/2405873.
[2]
Hayden, Eric J.; Ferrada, Evandro; Wagner, Andreas. Cryptic genetic variation promotes rapid evolutionary adaptation in an RNA enzyme (PDF). Nature. 2 June 2011, 474 (7349): 92–95 [2020-06-13]. PMID 21637259. doi:10.1038/nature10083. （原始内容存档 (PDF)于2020-08-06）.
[3]
Gould & Vrba 1982.
[4]
Gould, S. J.; Vrba, E. S. Exaptation - a missing term in the science of form. Paleobiology. 1982, 8 (1): 4–15. JSTOR 2400563.
[5]
Jacob, F. Evolution and tinkering. Science. 1977, 196 (4295): 1161–6. Bibcode:1977Sci...196.1161J. PMID 860134. doi:10.1126/science.860134.
[6]
Mayr, Ernst. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Harvard University Press. 1982 [2020-06-13]. ISBN 978-0-674-36445-5. （原始内容存档于2020-08-17）.
[7]
Colleen Farmer. Did Lungs and the Intracardiac Shunt Evolve to Oxygenate the Heart in Vertebrates?. Paleobiology (Paleontological Society). 1997, 23 (3): 358–372. JSTOR 2401109.

资料来源

（英文）Buss, David M., Martie G. Haselton, Todd K. Shackelford, et al. (1998) "Adaptations, Exaptations, and Spandrels," American Psychologist, 53 (May):533–548. http://www.sscnet.ucla.edu/comm/haselton/webdocs/spandrels.html （页面存档备份，存于互联网档案馆）
（英文）Darwin, Charles. On the origin and transitions of organic beings with peculiar habits and structure. On the Origin of Species 1st. London: John Murray. 1859: 179–186 [2020-06-13]. （原始内容存档于2016-03-04）.
（英文）Darwin, Charles. Difficulties of the theory. The Origin of Species 6th. London: John Murray. 1872: 138–143 [2020-06-13]. （原始内容存档于2020-08-06）.
（英文）Ehrlich, Paul, and Marcus Feldman (2003) "Genes and Culture: What Creates Our Behavioral Phenome?," Current Anthropology, 44 (February):87–107. Included are comments and a reply.
（英文）Gould, Stephen Jay; Vrba, Elisabeth S. Exaptation — a missing term in the science of form (PDF). Paleobiology. 1982, 8 (1): 4–15 [2020-10-11]. JSTOR 2400563. doi:10.1017/S0094837300004310. （原始内容 (PDF)存档于2018-05-08）.
（英文）Gould, Stephen Jay. Exaptation: A Crucial Tool for an Evolutionary Psychology. Journal of Social Issues. 1991, 47 (3): 43–65. doi:10.1111/j.1540-4560.1991.tb01822.x.
（英文）MacDonald, G.; Leary, M.R. Why does social exclusion hurt? The relationship between social and physical pain. Psychological Bulletin. 2005, 131 (2): 202–23. PMID 15740417. doi:10.1037/0033-2909.131.2.202.
（英文）"Preadaptation" Merriam-Webster Online Dictionary. 2009. Merriam-Webster Online. 22 January 2009 <http://www.merriam-webster.com/dictionary/preadaptation （页面存档备份，存于互联网档案馆）>

参阅

[7] [a]
See Jacob (1977)^[5] and Mayr (1982)^[6] for references.

[1] [1]
Bock, W.J. Preadaptation and multiple evolutionary pathways. Evolution. 1959, 13 (2): 194–211. JSTOR 2405873. doi:10.2307/2405873.

[2] [2]
Hayden, Eric J.; Ferrada, Evandro; Wagner, Andreas. Cryptic genetic variation promotes rapid evolutionary adaptation in an RNA enzyme (PDF). Nature. 2 June 2011, 474 (7349): 92–95 [2020-06-13]. PMID 21637259. doi:10.1038/nature10083. （原始内容存档 (PDF)于2020-08-06）.

[FOOTNOTEGouldVrba1982-3] [3]
Gould & Vrba 1982.

[4] [4]
Gould, S. J.; Vrba, E. S. Exaptation - a missing term in the science of form. Paleobiology. 1982, 8 (1): 4–15. JSTOR 2400563.

[Jacob1977-5] [5]
Jacob, F. Evolution and tinkering. Science. 1977, 196 (4295): 1161–6. Bibcode:1977Sci...196.1161J. PMID 860134. doi:10.1126/science.860134.

[Mayr1982-6] [6]
Mayr, Ernst. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Harvard University Press. 1982 [2020-06-13]. ISBN 978-0-674-36445-5. （原始内容存档于2020-08-17）.

[8] [7]
Colleen Farmer. Did Lungs and the Intracardiac Shunt Evolve to Oxygenate the Heart in Vertebrates?. Paleobiology (Paleontological Society). 1997, 23 (3): 358–372. JSTOR 2401109.

[1]

[2]

[3]

[4]

[a]

[7]

[5]

[6]