A new view of the functional role of the left anterior cortex in language use is proposed. The experimental record indicates that most human linguistic abilities are not localized in this region. In particular, most of syntax (long thought to be there) is not located in Broca's area and its vicinity (operculum, insula, and subjacent white matter). This cerebral region, implicated in Broca's aphasia, does have a role in syntactic processing, but a highly specific one: It is the neural home to receptive mechanisms involved in the computation of the relation between transformationally moved phrasal constituents and their extraction sites (in line with the Trace-Deletion Hypothesis). It is also involved in the construction of higher parts of the syntactic tree in speech production. By contrast, basic combinatorial capacities necessary for language processing – for example, structure-building operations, lexical insertion – are not supported by the neural tissue of this cerebral region, nor is lexical or combinatorial semantics. The dense body of empirical evidence supporting this restrictive view comes mainly from several angles on lesion studies of syntax in agrammatic Broca's aphasia. Five empirical arguments are presented: experiments in sentence comprehension, cross-linguistic considerations (where aphasia findings from several language types are pooled and scrutinized comparatively), grammaticality and plausibility judgments, real-time processing of complex sentences, and rehabilitation. Also discussed are recent results from functional neuroimaging and from structured observations on speech production of Broca's aphasics. Syntactic abilities are nonetheless distinct from other cognitive skills and are represented entirely and exclusively in the left cerebral hemisphere. Although more widespread in the left hemisphere than previously thought, they are clearly distinct from other human combinatorial and intellectual abilities. The neurological record (based on functional imaging, split-brain and right-hemisphere-damaged patients, as well as patients suffering from a breakdown of mathematical skills) indicates that language is a distinct, modularly organized neurological entity. Combinatorial aspects of the language faculty reside in the human left cerebral hemisphere, but only the transformational component (or algorithms that implement it in use) is located in and around Broca's area.

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.

The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals.

Studies of animal culture have not normally included a consideration of cetaceans. However, with several long-term field studies now maturing, this situation should change. Animal culture is generally studied by either investigating transmission mechanisms experimentally, or observing patterns of behavioural variation in wild populations that cannot be explained by either genetic or environmental factors. Taking this second, ethnographic, approach, there is good evidence for cultural transmission in several cetacean species. However, only the bottlenose dolphin (Tursiops) has been shown experimentally to possess sophisticated social learning abilities, including vocal and motor imitation; other species have not been studied. There is observational evidence for imitation and teaching in killer whales. For cetaceans and other large, wide-ranging animals, excessive reliance on experimental data for evidence of culture is not productive; we favour the ethnographic approach. The complex and stable vocal and behavioural cultures of sympatric groups of killer whales (Orcinus orca) appear to have no parallel outside humans, and represent an independent evolution of cultural faculties. The wide movements of cetaceans, the greater variability of the marine environment over large temporal scales relative to that on land, and the stable matrilineal social groups of some species are potentially important factors in the evolution of cetacean culture. There have been suggestions of gene-culture coevolution in cetaceans, and culture may be implicated in some unusual behavioural and life-history traits of whales and dolphins. We hope to stimulate discussion and research on culture in these animals.

The tendency to choose a larger, more delayed reinforcer over a smaller, less delayed one has frequently been termed “selfcontrol.” Three very different research traditions – two models emphasizing the control of local contingencies of reinforcement (Mischel's social learning theory and Herrnstein's matching law) and molar maximization models (specifically optimal foraging theory) – have all investigated behavior within the self-control paradigm. A framework is proposed to integrate research from all three research areas. This framework consists of three parts: a procedural analysis, a causal analysis, and a theoretical analysis. The procedural analysis provides a common procedural terminology for all three areas. The causal analysis establishes that, in all three research traditions, self-control varies directly with the current physical values of the reinforcers; that is, choices increase with reinforcer amount and decrease with reinforcer delay. But self-control also varies according to past events to which a subject has been exposed, and according to current factors other than the reinforcers. Each of the three models has therefore incorporated these indirect effects on self-control by postulating unobservable mechanisms. In all three cases, these mechanisms represent a subject's behavior as a function of a perceived environment. The theoretical analysis demonstrates that evolutionary theory can encompass the research from all three areas by considering differences in the adaptiveness of self-control in different situations. This integration provides a better and more predictive description of self-control.

The sway that the response-reinforcement framework (Spencer, Thorndike, Hull, Skinner) has held on the behavioral sciences for nearly a hundred years is finally ending. The strength of this framework lay in providing concepts and methods for studying the effects of hedonic (reinforcing) stimuli on the repetition of specified responses acquired in instrumental training situations of various kinds. Its weakness lay in the invalidity of its central assumptions, stimulus-response association and response-reinforcement, which could not deal with motor-equivalence and flexibility (or “intelligence”) in behavior. To the four decades of incisive criticism on particular theoretical and empirical grounds, a more comprehensive challenge to the response-reinforcement framework is now added by the newer ideas about the nature of cognitive, motivational, and response-production processes that have emerged from the work of ethologists, neuroscientists, and cognitive psychologists. An alternative framework, incorporating the newer ideas, is clearly needed.

The particular framework proposed here is based on the ideas of perceptual learning of stimulus-stimulus correlations and of a motivational (rather than reinforcing) role of hedonic (incentive) stimuli. According to it, an act is produced when its act-assembly is activated by a pexgo (perceptual representation) of a certain eliciting stimulus complex (ES). When certain eliciting stimuli are correlated with incentive stimuli, they acquire motivational properties that serve to strengthen the pexgos generated by those eliciting stimuli and thereby increase the probability of activation of the corresponding act-assemblies. Motivation thus influences response production, not by directly instigating “existing” responses, but by modulating the strength of pexgos of eliciting stimuli for the succession of acts that comprise a response. Therefore, a response is always constructed afresh on the basis of current perceptions; not even a stable and stereotyped response occurs as a mere activation of a preformed motor program. The topography of any response that emerges is determined by the nature of the motivational state and the momentary spatiotemporal distribution of eliciting stimuli of changing motivational valence.

By suggesting that the animal learns the overlapping and nested correlations between the stimulus events that commonly occur in a given situation, and by separating what is learned from the processes of response production, the proposed perceptual-motivational framework seems capable of dealing with the problems of motor equivalence and flexibility in adaptive behavior. Some implications of this approach for further behavioral and brain research on such problems as behavior modification, learning by observation of models, analysis of causality, and search for neural substrates of learning and response production, are outlined.

Elements of episodic memory (Tulving 1983b) consists of three parts. Part I argues for the distinction between episodic and semantic memory as functionally separate albeit closely interacting systems. It begins with a review of the 1972 essay on the topic (Tulving 1972) and its shortcomings, presents a somewhat more complete characterization of the two forms of memory than the one that was possible in 1972, and proceeds to discuss empirical and theoretical reasons for a tentative acceptance of the functional distinction between the two systems and its possible extensions. Part II describes a framework for the study of episodic memory, dubbed General Abstract Processing System (GAPS). The basic unit in such study is an act of remembering. It begins with the witnessing of an event and ends with recollective experience of the event, with related memory performance, or both. The framework specifies a number of components (elements) of the act of remembering and their interrelations, classified under two broad categories of encoding and retrieval. Part III discusses experimental research under the label of “synergistic ecphory.” Ecphory is one of the central elements of retrieval; “synergistic” refers to the joint influence that the stored episodic information and the cognitively present retrieval information exert on the construction of the product of ecphory, the so-called ecphoric information. The concept of encoding specificity and the phenomenon of recognition failure of recallable words figure prominently in Part III. The final chapter of the book describes a model, named the synergistic ecphory model of retrieval, that relates qualitative characteristics of recollective experience and quantitative measures of memory performance in recall and recognition to the conjunction of episodic-memory traces and semantic-memory retrieval cues.

The finding that women are attracted to men older than themselves whereas men are attracted to relatively younger women has been explained by social psychologists in terms of economic exchange rooted in traditional sex-role norms. An alternative evolutionary model suggests that males and females follow different reproductive strategies, and predicts a more complex relationship between gender and age preferences. In particular, males' preferences for relatively younger females should be minimal during early mating years, but should become more pronounced as the male gets older. Young females are expected to prefer somewhat older males during their early years and to change less as they age. We briefly review relevant theory and present results of six studies testing this prediction. Study 1 finds support for this gender-differentiated prediction in age preferences expressed in personal advertisements. Study 2 supports the prediction with marriage statistics from two U.S. cities. Study 3 examines the cross-generational robustness of the phenomenon, and finds the same pattern in marriage statistics from 1923. Study 4 replicates Study 1 using matrimonial advertisements from two European countries, and from India. Study 5 finds a consistent pattern in marriages recorded from 1913 through 1939 on a small island in the Philippines. Study 6 reveals the same pattern in singles advertisements placed by financially successful American women and men. We consider the limitations of previous normative and evolutionary explanations of age preferences and discuss the advantages of expanding previous models to include the life history perspective.

Sociopaths are “outstanding” members of society in two senses: politically, they draw our attention because of the inordinate amount of crime they commit, and psychologically, they hold our fascination because most ofus cannot fathom the cold, detached way they repeatedly harm and manipulate others. Proximate explanations from behavior genetics, child development, personality theory, learning theory, and social psychology describe a complex interaction of genetic and physiological risk factors with demographic and micro environmental variables that predispose a portion of the population to chronic antisocial behavior. More recent, evolutionary and game theoretic models have tried to present an ultimate explanation of sociopathy as the expression of a frequency-dependent life strategy which is selected, in dynamic equilibrium, in response to certain varying environmental circumstances. This paper tries to integrate the proximate, developmental models with the ultimate, evolutionary ones, suggesting that two developmentally different etiologies of sociopathy emerge from two different evolutionary mechanisms. Social strategies for minimizing the incidence of sociopathic behavior in modern society should consider the two different etiologies and the factors that contribute to them.

Because little is known about the human trait of affiliation, we provide a novel neurobehavioral model of affiliative bonding. Discussion is organized around processes of reward and memory formation that occur during approach and consummatory phases of affiliation. Appetitive and consummatory reward processes are mediated independently by the activity of the ventral tegmental area (VTA) dopamine (DA)–nucleus accumbens shell (NAS) pathway and the central corticolimbic projections of the u-opiate system of the medial basal arcuate nucleus, respectively, although these two projection systems functionally interact across time. We next explicate the manner in which DA and glutamate interact in both the VTA and NAS to form incentive-encoded contextual memory ensembles that are predictive of reward derived from affiliative objects. Affiliative stimuli, in particular, are incorporated within contextual ensembles predictive of affiliative reward via: (a) the binding of affiliative stimuli in the rostral circuit of the medial extended amygdala and subsequent transmission to the NAS shell; (b) affiliative stimulus-induced opiate potentiation of DA processes in the VTA and NAS; and (c) permissive or facilitatory effects of gonadal steroids, oxytocin (in interaction with DA), and vasopressin on (i) sensory, perceptual, and attentional processing of affiliative stimuli and (ii) formation of social memories. Among these various processes, we propose that the capacity to experience affiliative reward via opiate functioning has a disproportionate weight in determining individual differences in affiliation. We delineate sources of these individual differences, and provide the first human data that support an association between opiate functioning and variation in trait affiliation.

How can we disentangle the neural basis of phenomenal consciousness from the neural machinery of the cognitive access that underlies reports of phenomenal consciousness? We see the problem in stark form if we ask how we can tell whether representations inside a Fodorian module are phenomenally conscious. The methodology would seem straightforward: Find the neural natural kinds that are the basis of phenomenal consciousness in clear cases – when subjects are completely confident and we have no reason to doubt their authority – and look to see whether those neural natural kinds exist within Fodorian modules. But a puzzle arises: Do we include the machinery underlying reportability within the neural natural kinds of the clear cases? If the answer is “Yes,” then there can be no phenomenally conscious representations in Fodorian modules. But how can we know if the answer is “Yes”? The suggested methodology requires an answer to the question it was supposed to answer! This target article argues for an abstract solution to the problem and exhibits a source of empirical data that is relevant, data that show that in a certain sense phenomenal consciousness overflows cognitive accessibility. I argue that we can find a neural realizer of this overflow if we assume that the neural basis of phenomenal consciousness does not include the neural basis of cognitive accessibility and that this assumption is justified (other things being equal) by the explanations it allows.

To describe phenomena that occur at different time scales, computational models of the brain must incorporate different levels of abstraction. At time scales of approximately 1/3 of a second, orienting movements of the body play a crucial role in cognition and form a useful computational level – more abstract than that used to capture natural phenomena but less abstract than what is traditionally used to study high-level cognitive processes such as reasoning. At this “embodiment level,” the constraints of the physical system determine the nature of cognitive operations. The key synergy is that at time scales of about 1/3 of a second, the natural sequentiality of body movements can be matched to the natural computational economies of sequential decision systems through a system of implicit reference called deictic in which pointing movements are used to bind objects in the world to cognitive programs. This target article focuses on how deictic bindings make it possible to perform natural tasks. Deictic computation provides a mechanism for representing the essential features that link external sensory data with internal cognitive programs and motor actions. One of the central features of cognition, working memory, can be related to moment-by-moment dispositions of body features such as eye movements and hand movements.

During the first two years of human life a common neural substrate (roughly Broca's area) underlies the hierarchical organization of elements in the development of speech as well as the capacity to combine objects manually, including tool use. Subsequent cortical differentiation, beginning at age two, creates distinct, relatively modularized capacities for linguistic grammar and more complex combination of objects. An evolutionary homologue of the neural substrate for language production and manual action is hypothesized to have provided a foundation for the evolution of language before the divergence of the hominids and the great apes. Support comes from the discovery of a Broca's area homologue and related neural circuits in contemporary primates. In addition, chimpanzees have an identical constraint on hierarchical complexity in both tool use and symbol combination. Their performance matches that of the two-year-old child who has not yet developed the neural circuits for complex grammar and complex manual combination of objects.

Considerable evidence from preclinical and clinical investigations implicates disturbances of brain dopamine (DA) function in the pathophysiology of several psychiatric and neurologic disorders. We describe a neural model that may help organize theseindependent experimental observations. Cortical regions classically associated with the limbic system interact with infracortical structures, including the nucleus accumbens, ventral pallidum, and dorsomedial nucleus of the thalamus. In our model, overactivity in forebrain DA systems results in the loss of lateral inhibitory interactions in the nucleus accumbens, causing disinhibition of pallidothalamic efferents; this in turn causes rapid changes and a loss of focused corticothalamic activity in cortical regions controlling cognitive and emotional processes. These effects might be manifested clinically by some symptoms of psychoses. Underactivity of forebrain DA results in excess lateral inhibition in the nucleus accumbens, causing tonic inhibition of pallidothalamic efferents; this perpetuates tonic corticothalamic activity and prevents the initiation of new activity in other critical cortical regions. These effects might be manifested clinically by some symptoms of depression. This model parallels existing explanations for the etiology of several movement disorders, and may lead to testable inferences regarding the neural substrates of specific psychopathologies.

It is hypothesized that creole languages are largely invented by children and show fundamental similarities, which derive from a biological program for language. The structures of Hawaiian Pidgin and Hawaiian Creole are contrasted, and evidence is provided to show that the latter derived from the former in a single generation. A realistic model of the processes of Creole formation shows how several specific historical and demographic factors interacted to restrict, in varying degrees, the access of pidgin speakers to the dominant language, and hence the nature of input to the children of those speakers. It is shown that the resulting similarities of Creole languages derive from a single grammar with a restricted list of categories and operations. However, grammars of individual Creoles will differ from this grammar to a varying extent: The degree of difference will correlate very closely with the quantity of dominant-language input, which in turn is controlled by extralinguistic factors. Alternative explanations of the above phenomena are surveyed, in particular, substratum theory and monogenesis: Both are found inadequate to account for the facts. Primary acquisition is examined in light of the general hypothesis, and it is suggested that the bioprogram provides a skeletal model of language which the child can then readily convert into the target language. Cases of systematic error and precocious learning provide indirect support for the hypothesis. Some conjectures are made concerning the evolutionary origins of the bioprogram and what study of Creoles and related topics might reveal about language origins.

Tactical deception occurs when an individual is able to use an “honest” act from his normal repertoire in a different context to mislead familiar individuals. Although primates have a reputation for social skill, most primate groups are so intimate that any deception is likely to be subtle and infrequent. Published records are sparse and often anecdotal. We have solicited new records from many primatologists and searched for repeating patterns. This has revealed several different forms of deceptive tactic, which we classify in terms of the function they perform. For each class, we sketch the features of another individual's state of mind that an individual acting with deceptive intent must be able to represent, thus acting as a “natural psychologist.” Our analysis will sharpen attention to apparent taxonomic differences. Before these findings can be generalized, however, behavioral scientists must agree on some fundamental methodological and theoretical questions in the study of the evolution of social cognition.

A model for the organization of language in the adult humans brain is derived from electrical stimulation mapping of several language-related functions: naming, reading, short-term verbal memory, mimicry of orofacial movements, and phoneme identification during neurosurgical operations under local anesthesia. A common peri-Sylvian cortex for motor and language functions is identified in the language dominant hemisphere, including sites common to sequencing of movements and identification of phonemes that may represent an anatomic substrate for the “motor theory of speech perception.” This is surrounded by sites related to short-term verbal memory, with sites specialized for such language functions as naming or syntax at the interface between these motor and memory areas. Language functions are discretely and differentially localized in association cortex, including some differential localization of the same function, naming, in multiple languages. There is substantial individual variability in the exact location of sites related to a particular function, a variability which can be partly related to the patient's sex and overall language ability and which may depend on prior brain injury and, perhaps subtly, on prior experience. A common “specific alerting response” mechanism for motor and language functions is identified in the lateral thalamus of the language–dominant hemisphere, a mechanism that may select the cortical areas appropriate for a particular language function.

What determines what we see? In contrast to the traditional “modular” understanding of perception, according to which visual processing is encapsulated from higher-level cognition, a tidal wave of recent research alleges that states such as beliefs, desires, emotions, motivations, intentions, and linguistic representations exert direct, top-down influences on what we see. There is a growing consensus that such effects are ubiquitous, and that the distinction between perception and cognition may itself be unsustainable. We argue otherwise: None of these hundreds of studies – either individually or collectively – provides compelling evidence for true top-down effects on perception, or “cognitive penetrability.” In particular, and despite their variety, we suggest that these studies all fall prey to only a handful of pitfalls. And whereas abstract theoretical challenges have failed to resolve this debate in the past, our presentation of these pitfalls is empirically anchored: In each case, we show not only how certain studies could be susceptible to the pitfall (in principle), but also how several alleged top-down effects actually are explained by the pitfall (in practice). Moreover, these pitfalls are perfectly general, with each applying to dozens of other top-down effects. We conclude by extracting the lessons provided by these pitfalls into a checklist that future work could use to convincingly demonstrate top-down effects on visual perception. The discovery of substantive top-down effects of cognition on perception would revolutionize our understanding of how the mind is organized; but without addressing these pitfalls, no such empirical report will license such exciting conclusions.

Maximization theory, which is borrowed from economics, provides techniques for predicing the behavior of animals - including humans. A theoretical behavioral space is constructed in which each point represents a given combination of various behavioral alternatives. With two alternatives - behavior A and behavior B - each point within the space represents a certain amount of time spent performing behavior A and a certain amount of time spent performing behavior B. A particular environmental situation can be described as a constraint on available points (a circumscribed area) within the space. Maximization theory assumes that animals always choose the available point with the highest numerical value. The task of maximization theory is to assign to points in the behavioral space values that remain constant across various environmental situations; as those situations change, the point actually chosen is always the one with the highest assigned value.

Maximization theory is an alternative to reinforcement theory as a description of steady-state behavior. Situations to which reinforcement theory has been directly applied (such as reinforcement of rats pressing levers and pigeons pecking keys in Skinner boxes) and situations to which reinforcement theory has occasionally been extended (such as human economic behavior and human self-control) can be described by maximization theory. This approach views behavior as a quantitative outcome of the interaction of the putative instrumental response, the reinforcer, and the other activities available in the situation. It provides new insight into these situations and, because it takes context into account, has greater predictive power than reinforcement theory.

Females' tendency to place a high value on protecting their own lives enhanced their reproductive success in the environment of evolutionary adaptation because infant survival depended more upon maternal than on paternal care and defence. The evolved mechanism by which the costs of aggression (and other forms of risk taking) are weighted more heavily for females may be a lower threshold for fear in situations which pose a direct threat of bodily injury. Females' concern with personal survival also has implications for sex differences in dominance hierarchies because the risks associated with hierarchy formation in nonbonded exogamous females are not offset by increased reproductive success. Hence among females, disputes do not carry implications for status with them as they do among males, but are chiefly connected with the acquisition and defence of scarce resources. Consequently, female competition is more likely to take the form of indirect aggression or low-level direct combat than among males. Under patriarchy, men have held the power to propagate images and attributions which are favourable to the continuance of their control. Women's aggression has been viewed as a gender-incongruent aberration or dismissed as evidence of irrationality. These cultural interpretations have “enhanced” evolutionarily based sex differences by a process of imposition which stigmatises the expression of aggression by females and causes women to offer exculpatory (rather than justificatory) accounts of their own aggression.