showed them to other monkeys with recording electrodes in their vlPFC. This experiment revealed that the majority of vlPFC neurons integrate auditory and visual information in a nonlinear manner. This finding is important because human speech perception also involves a considerable amount of audiovisual integration, as demonstrated by the McGurk effect (McGurk and MacDonald, 1976). Of course, audiovisual integration of vocalization-related stimuli is not identical to speech perception, which requires the integration of sounds and visual information with meanings. The latter type of integration still eludes the understanding of neurobiologists and is extremely difficult to study in monkeys. Nonetheless, the audiovisual integration that Romanski describes in monkeys is likely to have played a major role in the evolution of human language.

In Chapter 16, Jessica Cantlon compares the mathematical abilities of nonhuman primates and humans, especially human children. Although we often think that mathematics requires symbols (e.g., numbers and operators), simple math can be performed without symbols. For example, one can compare two images and estimate, even without counting, which image contains more items of a particular sort. This kind of analog numerical estimation can also be performed by human infants and nonhuman primates. Cantlon further reports that the analog math task activates homologous brain areas in the parietal cortex of both humans and monkeys. Collectively, the data strongly suggest that analog math abilities evolved long before the origin of Homo sapiens. This finding is fascinating, but how did symbolic math evolve? Was it built on top of the more ancient analog skill, using the ancient circuitry with only minor modifications? Or did symbolic math evolve out of symbolic communication (i.e., language)? At this point, the answer is unknown.

In the final Chapter 17, Clark Barrett dispels the notion—promulgated by some evolutionary psychologists—that adaptive specializations in the brain must be hard-wired modules. To grasp the argument, consider face-selective neurons in primate brains. Given the importance of conspecific faces in the lives of most primates, the distinct patches of face-selective neurons in monkey and human brains were likely shaped by natural selection. Nonetheless, the development of face-selective neurons probably depends on extensive experience with faces. Indeed, Barrett hypothesizes that selection generated not an innate face-processing module but a set of mechanisms that, given experience with faces, will generate a large number of neurons that selectively encode faces. Given other types of experience, the same mechanisms would (and do) generate patches of neurons selective for other kinds of behaviorally important stimuli. Stated succinctly, Barrett argues that natural selection generates developmental norms of reaction rather than experience-independent specialized modules. This idea extends evo-devo neurobiology into the realm of evolutionary psychology.

14

Human Brain Evolution: From Gene Discovery to Phenotype Discovery

TODD M. PREUSS

The rise of comparative genomics and related technologies has added important new dimensions to the study of human evolution. Our knowledge of the genes that underwent expression changes or were targets of positive selection in human evolution is rapidly increasing, as is our knowledge of gene duplications, translocations, and deletions. It is now clear that the genetic differences between humans and chimpanzees are far more extensive than previously thought; their genomes are not 98% or 99% identical. Despite the rapid growth in our understanding of the evolution of the human genome, our understanding of the relationship between genetic changes and phenotypic changes is tenuous. This is true even for the most intensively studied gene, FOXP2, which underwent positive selection in the human terminal lineage and is thought to have played an important role in the evolution of human speech and language. In part, the difficulty of connecting genes to phenotypes reflects our generally poor knowledge of human phenotypic specializations, as well as the difficulty of interpreting the consequences of genetic changes in species that are not amenable to invasive research. On the positive side, investigations of FOXP2, along with genomewide surveys of gene-expression changes and selection-driven sequence changes, offer the opportunity for “phenotype discovery,” providing clues to human phenotypic specializations that were previously unsuspected. What is more,

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Yerkes National Primate Research Center, Emory University, Atlanta, GA 30322. E-mail: tpreuss@emory.edu.