The study of information-processing adaptations in the brain is controversial, in part because of disputes about the form such adaptations might take. Many psychologists assume that adaptations come in two kinds, specialized and general-purpose. Specialized mechanisms are typically thought of as innate, domainspecific, and isolated from other brain systems, whereas generalized mechanisms are developmentally plastic, domain-general, and interactive. However, if brain mechanisms evolve through processes of descent with modification, they are likely to be heterogeneous, rather than coming in just two kinds. They are likely to be hierarchically organized, with some design features widely shared across brain systems and others specific to particular processes. Also, they are likely to be largely developmentally plastic and interactive with other brain systems, rather than canalized and isolated. This article presents a hierarchical model of brain specialization, reviewing evidence for the model from evolutionary developmental biology, genetics, brain mapping, and comparative studies. Implications for the search for uniquely human traits are discussed, along with ways in which conventional views of modularity in psychology may need to be revised.hat is the nature of the brain mechanisms that give rise to human cognition, and how do these mechanisms evolve? Although it is clear that human cognition, like all organismal traits, must be accounted for by some combination of ancestral and derived brain processes, attempts to decompose human mental processes into functional components whose features have been shaped by the process of natural selection-i.e., adaptations-have been highly contested and controversial (1). The controversy centers on the difficulty of establishing whether a particular aspect of cognition or behavior is the result of an adaptation or adaptations, and in what way. Is a given cognitive ability in humans or any other species-for example, the ability to discriminate between different quantities of objects, to navigate spatially, or to learn to speak a language-the product of an adaptation specifically for that ability? Or is it just a specific instantiation of a more general ability, such as associative learning, or the general computational properties of neural networks? Or is it not the result of adaptations at all? Proposals about functional specialization have long been a source of debate in psychology and the brain sciences. In particular, there is little agreement over whether cognitive processes other than perceptual and motor processes-i.e., so-called higher-level processes-are specialized, and if so, how (2). At stake are both theoretical and empirical issues. Theoretically, although it is clear that the brain is the product of evolutionary processes, including natural selection, we cannot move past this simple truism if we are unable to answer the question of what adaptations it contains, or to distinguish the results of natural selection from the results of other processes. Empirically, a v...