Wilkins & Wakefield suggest that changes in the hominid brain made it uniquely “preadaptive” for language, yet no precursor functions served as adaptive substrates to the emergence of language. We present contrary evidence that the ability to discriminate and process rapid and complex auditory information is a cross-species function subserving communication processes including, but not limited to, human speech perception. We suggest that auditory temporal processing served as an evolutionary precursor to speech processing and consequent language development in humans.

Three issues are addressed in this commentary. (1) Wilkins & Wakefield are commended for placing the complex behavior they discuss within an evolutionary matrix. (2) They err on a number of points in regard to their treatment of this complex behavior. These involve (a) their emphasis on the evolution of conceptual structure rather than language, (b) their equation of meaning with reference, (c) their minimalist view of learning theory, and (d) their separation of the evolution of speech from that of (...) language. (3) They adopt a framework for neural preconditions that has been clearly discredited on both behavioral and neuroanatomical grounds. Finally, recent research involving modern neuroimaging techniques casts further doubt on the authors' postulated functions for the neural areas they discuss. (shrink)

If POT (parieto-occipital-temporal junction) reorganization came earlier in australopithecines than in Homo, it is likely that the selective pressures were different, and not necessarily directed toward language. The brain endocast evidence for the POT in A. afarensis is actually better than it is for early Homo.

This target article presents a plausible evolutionary scenario for the emergence of the neural preconditions for language in the hominid lineage. In pleistocene primate lineages there was a paired evolutionary expansion of frontal and parietal neocortex (through certain well-documented adaptive changes associated with manipulative behaviors) resulting, in ancestral hominids, in an incipient Broca's region and in a configurationally unique junction of the parietal, occipital, and temporal lobes of the brain (the POT). On our view, the development of the POT in (...) our ancestors resulted in the neuroanatomical substrate consistent with the ability for representations in modality-neutral association cortex and, as a result of structure-imposing interaction with Broca's area, the hierarchically structured Evidence from paleoneurology and comparative primate neuroanatomy is used to argue that Homo habilis (2.5–2 million years ago) was the first hominid to have the appropriate gross neuroanatomical configuration to support conceptual structure. We thus suggest that the neural preconditions for language are met in H. habilis. Finally, we advocate a theory of language acquisition that uses conceptual structure as input to the learning procedures, thus bridging the gap between it and language. (shrink)

Toolmaking requires motor skills that in turn require handedness, so that there is no competition between the two sides of the brain. Thus, handedness is not necessarily linked to vocalization but to the origin of causal beliefs required for making complex tools. Language may have evolved from these processes.

Recent evidence in natural and semi-natural settings has revealed a variety of left-right perceptual asymmetries among vertebrates. These include preferential use of the left or right visual hemifield during activities such as searching for food, agonistic responses, or escape from predators in animals as different as fish, amphibians, reptiles, birds, and mammals. There are obvious disadvantages in showing such directional asymmetries because relevant stimuli may be located to the animal's left or right at random; there is no a priori association (...) between the meaning of a stimulus (e.g., its being a predator or a food item) and its being located to the animal's left or right. Moreover, other organisms (e.g., predators) could exploit the predictability of behavior that arises from population-level lateral biases. It might be argued that lateralization of function enhances cognitive capacity and efficiency of the brain, thus counteracting the ecological disadvantages of lateral biases in behavior. However, such an increase in brain efficiency could be obtained by each individual being lateralized without any need to align the direction of the asymmetry in the majority of the individuals of the population. Here we argue that the alignment of the direction of behavioral asymmetries at the population level arises as an “evolutionarily stable strategy” under “social” pressures occurring when individually asymmetrical organisms must coordinate their behavior with the behavior of other asymmetrical organisms of the same or different species. Key Words: asymmetry; brain evolution; brain lateralization; development; hemispheric specialization; laterality; lateralization of behavior; social behavior; theory of games. (shrink)

The species-specific organizational property of speech is a continual mouth open-close alternation, the two phases of which are subject to continual articulatory modulation. The cycle constitutes the syllable, and the open and closed phases are segments framescontent displays that are prominent in many nonhuman primates. The new role of Broca's area and its surround in human vocal communication may have derived from its evolutionary history as the main cortical center for the control of ingestive processes. The frame and content components (...) of speech may have subsequently evolved separate realizations within two general purpose primate motor control systems: (1) a motivation-related medial system, including anterior cingulate cortex and the supplementary motor area, for self-generated behavior, formerly responsible for ancestral vocalization control and now also responsible for frames, and (2) a lateral system, including Broca's area and surround, and Wernicke's area, specialized for response to external input (and therefore the emergent vocal learning capacity) and more responsible for content. (shrink)

Many people have argued that the evolution of the human language faculty cannot be explained by Darwinian natural selection. Chomsky and Gould have suggested that language may have evolved as the by-product of selection for other abilities or as a consequence of as-yet unknown laws of growth and form. Others have argued that a biological specialization for grammar is incompatible with every tenet of Darwinian theory – that it shows no genetic variation, could not exist in any intermediate forms, confers (...) no selective advantage, and would require more evolutionary time and genomic space than is available. We examine these arguments and show that they depend on inaccurate assumptions about biology or language or both. Evolutionary theory offers clear criteria for when a trait should be attributed to natural selection: complex design for some function, and the absence of alternative processes capable of explaining such complexity. Human language meets these criteria: Grammar is a complex mechanism tailored to the transmission of propositional structures through a serial interface. Autonomous and arbitrary grammatical phenomena have been offered as counterexamples to the position that language is an adaptation, but this reasoning is unsound: Communication protocols depend on arbitrary conventions that are adaptive as long as they are shared. Consequently, language acquisition in the child should systematically differ from language evolution in the species, and attempts to analogize them are misleading. Reviewing other arguments and data, we conclude that there is every reason to believe that a specialization for grammar evolved by a conventional neo-Darwinian process. (shrink)

During the first two years of human life a common neural substrate 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. (shrink)

Wilkins & Wakefield (1995) provide a thoughtful contribution to our understanding of language origins. In this commentary I attempt to define the relationship between object-manipulation and primate brain function further by reviewing research on aimed throwing and the production and use of stone tools by tufted capuchin monkeys (Cebtis apella). I propose that examining the relation between brain function and object-manipulation inCebuswill provide insight into the preconditions of language in our hominid ancestors.

This commentary criticizes Wilkins & Wakefield's thesis that the neurological precursors of language provide a cognitive Rubicon to linguistically divide human from nonhuman primates. A causal model of their theory is presented, followed by a discussion of the relationship between brain expansion and tool use, Broca's area and the parietaloccipital-temporal junction (POT).

Wilkins & Wakefield assign importance to motor systems but skip from anatomy to cognitive structure with little attention to behavior. Organisms, no matter how sophisticated, that do not behave in accord with what they know will fall by the evolutionary wayside. Facts about behavior can supplement the authors' theory, whose hierarchical structures can accommodate an evolutionary scenario in which a million years or more of functionally varied utterances mainly limited to single words is followed by an explosion of linguistic diversity (...) with the development in the last 50,000 years or so of syntactically organized multiple word utterances. (shrink)

The precise manner in which language serves its communicative function suggests that natural selection, rather than exaptation or reappropriation, played the major role in its evolution. Natural selection is more readily invoked, I suggest, if it is assumed that language originated as a system of manual gestures, and later switched to an oral mode.

The problem of how certain structure–function composites of high complexity could have evolved gradually and by natural selection has been with us at least since Charles Darwin admitted how difficult it was to explain, “his” theory, the origins of “organs of extreme perfection and complication” – such as the eyes of higher animals. Human language capacity is another evolutionary achievement of extraordinary perfection and complexity. Like other skilled human activities, it involves both central (neural) and peripheral (vocal and respiratory) complexes. (...) The reduction of these to simpler building stones to which evolutionary principles may be applied is staggeringly difficult. (shrink)

This response to continuing commentary addresses brain-hand relationships in Cebus apella (as introduced in West-ergaard's commentary), the evolutionary and acquisition parallels between music and language (suggested by Lynch), and the potential behavioral linguistic consequences of the evolutionary neurobiology in Australopithecus africanus and Homo habilis (discussed by Tobias). Finally, we reiterate the importance of well informed, multidisciplinary approaches to the study of the emergence of human species-specific cognition, especially linguistic capacity.

This response clarifies the nature of reappropriation and the definition of language. It explicates the relationship between neural systems and language and between homology and evolutionary gradualism. Through a review of ape capacities in the realms of language and tool use, it distinguishes human language acquisition from nonhuman learning. Finally, it suggests the appropriate sorts of evidence on which to base further evolutionary arguments relevant to the origins of language.

Contra Wilkins & Wakefield, we argue that an evolutionarily inspired approach to language must consider different facets of language (i.e., more than syntax and semantics), and must explore the possibility of nonhuman precursors. Several examples are discussed, illustrating the power of the comparative approach in illuminating our understanding of language evolution.

Despite some sound basic assumptions, Wilkins & Wakefield portray a Homo habilis too linguistically sophisticated to fit in with the subsequent fossil record and thereby lose a reasoned explanation for human innovativeness. They err, too, in accepting a single-level model of conceptual structure and in deriving initial linguistic units from calls, a process far more dubious than the derivation of home-sign from naive gesture.

The strong predominance of right-handedness appears to be a uniquely human characteristic, whereas the left-cerebral dominance for vocalization occurs in many species, including frogs, birds, and mammals. Right-handedness may have arisen because of an association between manual gestures and vocalization in the evolution of language. I argue that language evolved from manual gestures, gradually incorporating vocal elements. The transition may be traced through changes in the function of Broca's area. Its homologue in monkeys has nothing to do with vocal control, (...) but contains the so-called “mirror neurons,” the code for both the production of manual reaching movements and the perception of the same movements performed by others. This system is bilateral in monkeys, but predominantly left-hemispheric in humans, and in humans is involved with vocalization as well as manual actions. There is evidence that Broca's area is enlarged on the left side in Homo habilis, suggesting that a link between gesture and vocalization may go back at least two million years, although other evidence suggests that speech may not have become fully autonomous until Homo sapiens appeared some 170,000 years ago, or perhaps even later. The removal of manual gesture as a necessary component of language may explain the rapid advance of technology, allowing late migrations of Homo sapiens from Africa to replace all other hominids in other parts of the world, including the Neanderthals in Europe and Homo erectus in Asia. Nevertheless, the long association of vocalization with manual gesture left us a legacy of right-handedness. Key Words: cerebral dominance; gestures; handedness; hominids; language evolution; primates; speech; vocalization. (shrink)

ProbablyHomo habilisis two species not one; similarly for Pan troglodytes. Although amenable to training, in naturePan paniscusmay be a “specialized insular dwarf.” Language is uniquely human, but symbolic behavior and intelligence are widespread among animals with little respect for phylogenetic closeness toHomo sapiens.

Wilkins & Wakefield are clearly right to separate linguistic capacity from communicative ability, if only because other animal species have one without the other. But I question the abruptness of the demarcation they make between a period when hominids evolved enriched conceptual representation for other reasons entirely, and a subsequent later stage when language use became an adaptation.