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An evolutionary constraint is a bias or limitation in genotypic or phenotypic variation that a biological system produces. Striking phenotypic examples include the absence of photosynthesis in higher animals, and the general lack of teeth in the lower jaw of frogs. Constraints can influence the spectrum of evolutionary adaptations and innovations that are accessible to living things. Based on the cause of constrained phenotypic variation, one can distinguish physicochemical, selective, genetic, and developmental constraints. The latter class of constraints emerges from the processes that produce phenotypes from genotypes. This chapter examines these four causes for molecules, regulatory circuits, and metabolic networks in the genotype space framework. This framework shows that processes of phenotype formation cause the three other classes of constraints. It can help us appreciate why causes of constrained variation are often entangled and not clearly separable. The chapter also shows that the kind of evolutionary stasis that occurs during punctuated and episodic evolution is a consequence of genetic constraints, whose origins the genotype space framework can readily explain.

This chapter deals with the theoretical analysis of evolutionary constraint in producing convergent evolution. It investigates the implications of theoretically possible, but nevertheless nonexistent, ecological roles on Earth for the concept of convergent evolution, and also analyzes the phenomenon of convergent evolution with respect to the spectrum of existent, nonexistent, and impossible biological forms. The chapter shows that the evolution of development on Earth may have been a two-stage process, and also suggests that possible, but nonexistent, biological forms can be visualized using the techniques of theoretical morphology.

Adaptations are the products of natural selection—traits that evolved because they proved useful at some level. Often adaptations evolved because they enhanced the survival or reproductive output of individuals, but selection can also operate at other levels—genes, individuals, populations, and species. Sometimes a genetic change has positive effects on all levels. A mutation that increases the survival of its carrier would increase in frequency; populations that become fixed for that allele may be less prone to extinction, which in turn may increase the longevity of that species. Other times there can be conflicting fitness effects at the different levels. This chapter explores the power of natural selection in shaping the living world by investigating the complexities of multilevel selection, biological solutions to heterogeneity and unpredictability in the environment, and how interactions between species can shape evolution. However, the chapter starts by investigating factors that may constrain adaptive evolution.