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62 T. Sachs<br />

the development of varied biological patterns. The mature stomata of Sansevieria<br />

are more orderly or predictably spaced than the early, reversible<br />

stages of stomata initiation (Kagan and Sachs 1991). “Neural Darwinism”<br />

is a mechanism by which appropriate nerve connections are selected from<br />

excess alternatives (Edelman 1987). Within cells, microtubules are maintained<br />

according to functional roles that are established after they have<br />

been formed (Kirschner and Mitchison 1986).<br />

It may be useful to consider the adaptive significance of the generation<br />

of tree form by developmental selection rather than strict programs. Competition<br />

between branches requires the wasteful development of structures<br />

that are not maintained. There is no doubt that dead branches below trees<br />

represent considerable organic material. Yet these branches did carry out<br />

photosynthesisforaslongastheywereproductiveandresourcessuchas<br />

bound nitrogen can be withdrawn from dying plant organs (Habib et al.<br />

1993). Losses of substrates could be balanced by the advantages of gradual<br />

selection, in which information about the value of an existing branch<br />

feeds back to its survival and further development. This testing of varied<br />

possibilities could provide for a robust outcome.<br />

A major advantage of developmental section may be in its relation to<br />

plasticity. It is possible that in a predictable, ideal world strict programs<br />

could result in superior biological forms. The real world, however, is far<br />

from predictable. In the case of trees, the presence of competing neighbors,<br />

damage due to herbivores and parasites and possible local failures<br />

in the development of the plant itself require a pronounced developmental<br />

plasticity. Developmental selection uses the very same processes and<br />

genetic information to generate form and to insure plastic responses to unpredictable<br />

conditions (Sachs 2002). This parsimony might have adaptive<br />

significance both because simpler developmental systems are required and<br />

because the various component are all optimally integrated.<br />

<strong>References</strong><br />

Barlow PW (1989) Meristems, metamers and modules and the development of shoot and<br />

root systems. Bot J Linn Soc 100:255–279<br />

Berleth T, Sachs T (2001) Plant morphogenesis: long-distance coordination and local patterning.<br />

Curr Opin Plant Biol 4:57–62<br />

Cline MG (1994) The role of hormones in apical dominance. New approaches to an old<br />

problem in plant development. Physiol Plant 90:230–237<br />

Edelman GM (1987) Neural Darwinism – the theory of neuronal group selection. Basic<br />

Books, New York<br />

Frank SA (1996) The design of natural and artificial adaptive systems. In: Rose MR, Lauder<br />

GV (eds) Adaptation. Academic, San Diego, pp 451–505<br />

Frank SA (1997) Developmental selection and self-organization. BioSystems 40:237–243

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