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Evolution__3rd_Edition

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588 PART 5 / Macroevolution<br />

Summary<br />

1 Morphological change in evolution usually occurs<br />

by changes in developmental processes. The identification<br />

of genes that influence development is a major<br />

area of modern biology, and its methods can be applied<br />

to study the relations of development and evolution, a<br />

field known as “evo-devo.”<br />

2 Heterochrony refers to evolutionary changes in the<br />

relative timing and rate of different developmental<br />

processes. For instance, the time of reproduction may<br />

shift relative to somatic development. Also, shape<br />

changes can result from changes in growth gradients,<br />

and D’Arcy Thompson’s transformational diagrams<br />

can be interpreted in terms of heterochrony.<br />

3 Regulatory genes influence the expression of other<br />

genes, and evolutionary change can result from changed<br />

regulatory relations among genes as well as changes in<br />

the sequence of genes.<br />

4 Structures, such as the eyes of insects and vertebrates,<br />

that had been thought to be non-homologous,<br />

Further reading<br />

have been found to be developmentally controlled by<br />

the same gene. Insect and vertebrate eyes may share an<br />

element of homology, but it is uncertain what the level<br />

of the homology is.<br />

5 The number of Hox genes increased from perhaps<br />

two to seven near the origin of the triploblastic<br />

Bilateria, and quadrupled from 13 to 52 near the origin<br />

of vertebrates. Hox genes control spatial differentiation<br />

within the body during development, and increases in<br />

the number of Hox genes may be associated with<br />

increases in developmental complexity.<br />

6 Changes in the expression of developmental genes<br />

are likely achieved by gains, losses, and changes in the<br />

regulatory elements (particularly enhancers) of those<br />

genes.<br />

7 Some forms of life may be more evolvable than<br />

others: that is, be more likely to undergo innovative<br />

evolutionary change. The origin of genetic switches<br />

may have made life more evolvable.<br />

General developmental biology texts, such as Gilbert (2000) and Wolpert (2002)<br />

contain chapters on evolution, as well as developmental biology background. Wilkins<br />

(2001), Carroll et al. (2001), and Hall (1998) are texts more specifically on evo-devo.<br />

Proceedings of the National Academy of Sciences (2000), vol. 97 (9), pp. 4424–540 contains<br />

the proceedings from a conference on evo-devo. Gerhart & Kirschner (1997) is a<br />

stimulating book, more about the evolution of cells, but containing much relevant<br />

material for this chapter. Meyerowitz (2002) gives an evo-devo comparison of plants<br />

and animals.<br />

Gould’s (1977a) book discusses the history of recapitulatory ideas and modern<br />

work on heterochrony. Gould (2002b) contains further material. Raff (1996) is a more<br />

recent general book, and Levinton’s (2001) even broader book also covers the topic.<br />

Both Gould and Raff are good on heterochrony, but see also the review article by<br />

Klingenberg (1998), the web-page on heterochrony (and on D-Arcy Thompson’s<br />

transformations) by Horder in www.els.org, and the think-piece by Smith (2001).<br />

Britton & Davidson (1971) is an early work discussing gene regulation and evolution.<br />

See also the introductory article by A.C. Wilson (1985), the recent book by Davidson<br />

(2001), as well as the general references and some further references below.<br />

..

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