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to a particular auditory tone results in increased cortical magnification for that frequency in A1 (Zhang et al., 2001). These changes in the internal organization of a sensory field and neuron response properties are similar to the types of differences observed across species and can be induced early in development by altering the sensory environment in which the animal develops.

Thus, a high degree of phenotypic variability can be induced without invoking genetic mechanisms that control brain development. The cortex has evolved to match the sensory environment in which it develops and produce highly adaptive behavior for that context. Although we have focused this review on how sensory systems and cortical areas are modified, if one considers both social and cultural influences on the brain as complex patterns of sensory stimuli that groups of brains generate, then the same rules of construction and modification apply. However, as with genes, the environmental factors that generate phenotypic variability also serve to constrain the types of changes that can be made to the brain. For example, although photons can be differentially distributed in an aquatic, cave, or terrestrial environment, they have the same intrinsic properties, are uniformly defined as a discrete quantum of electromagnetic energy, are always in motion, and in a vacuum travel at the speed of light. These immutable characteristics of a stimulus that the nervous system must detect, transduce, and ultimately translate, constrain the evolution and construction of the effector organ that initially captures some portion of the spectrum of this energy, and also impacts how higher-level structures transmit specific information about its presence, magnitude, and dispersal within an environment.

CONCLUSIONS

We have discussed phenotypic variability across and within species and conclude that the ways in which animals and brains change are limited and predictable. Further, we show that a specific characteristic, such as the size of a cortical field, can be generated by different genetic mechanisms and/or activity-dependent mechanisms. Thus, similar features of organization that have independently arisen in different lineages may not have similar underpinnings. Examination of variability at multiple levels of organization indicates that although genes are not directly related to a specific behavior, they covary with aspects of body and brain organization, which in turn covary with the targets of selection (Fig. 6.7). For example, the wing of a bat is constructed in development through complex interactions between genes and morphogens. Slight variations in the amount, location, and timing of these factors can generate phenotypic diversity within a population. The presence of the highly derived wing with its



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