The anatomic pattern and left hemisphere size predominance of the planum temporale, a language area of the human brain, are also present in chimpanzees (Pan troglodytes). The left planum temporale was significantly larger in 94 percent (17 of 18) of chimpanzee brains examined. It is widely accepted that the planum temporale is a key component of Wernicke's receptive language area, which is also implicated in human communication-related disorders such as schizophrenia and in normal variations such as musical talent. However, anatomic hemispheric asymmetry of this cerebrocortical site is clearly not unique to humans, as is currently thought. The evolutionary origin of human language may have been founded on this basal anatomic substrate, which was already lateralized to the left hemisphere in the common ancestor of chimpanzees and humans 8 million years ago.

Symbolic gestures, such as pantomimes that signify actions (e.g., threading a needle) or emblems that facilitate social transactions (e.g., finger to lips indicating ''be quiet''), play an important role in human communication. They are autonomous, can fully take the place of words, and function as complete utterances in their own right. The relationship between these gestures and spoken language remains unclear. We used functional MRI to investigate whether these two forms of communication are processed by the same system in the human brain. Responses to symbolic gestures, to their spoken glosses (expressing the gestures' meaning in English), and to visually and acoustically matched control stimuli were compared in a randomized block design. General Linear Models (GLM) contrasts identified shared and unique activations and functional connectivity analyses delineated regional interactions associated with each condition. Results support a model in which bilateral modality-specific areas in superior and inferior temporal cortices extract salient features from vocalauditory and gestural-visual stimuli respectively. However, both classes of stimuli activate a common, left-lateralized network of inferior frontal and posterior temporal regions in which symbolic gestures and spoken words may be mapped onto common, corresponding conceptual representations. We suggest that these anterior and posterior perisylvian areas, identified since the mid-19th century as the core of the brain's language system, are not in fact committed to language processing, but may function as a modality-independent semiotic system that plays a broader role in human communication, linking meaning with symbols whether these are

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The cortical circuits subserving neural processing of human language are localized to the inferior frontal operculum and the posterior perisylvian region. Functional language dominance has been related to anatomical asymmetry of Broca's area and the planum temporale. The evolutionary history of these asymmetric patterns, however, remains obscure. Although testing of hypotheses about the evolution of language areas requires comparison to homologous regions in the brains of our closest living relatives, the great apes, to date little is known about normal interindividual variation of these regions in this group. Here we focus on Brodmann's area 44 in African great apes (Pan troglodytes and Gorilla gorilla). This area corresponds to the pars opercularis of the inferior frontal gyrus (IFG), and has been shown to exhibit both gross and cytoarchitectural asymmetries in humans. We calculated frequencies of sulcal variations and mapped the distribution of cytoarchitectural area 44 to determine whether its boundaries occurred at consistent macrostructural landmarks. A considerable amount of variation was found in the distribution of the inferior frontal sulci among great ape brains. The inferior precentral sulcus in particular was often bifurcated, which made it impossible to determine the posterior boundary of the pars opercularis. In addition, the distribution of Brodmann's area 44 showed very little correspondence to surface anatomy. We conclude that gross morphologic patterns do not offer substantive landmarks for the measurement of Brodmann's area 44 in great apes. Whether or not Broca's area homologue of great apes exhibits humanlike asymmetry can only be resolved through further analyses of microstructural components. Anat Rec Part A 271A: 276 -285, 2003. © 2003 Key words: Broca's area; brain evolution; language; great apes; chimpanzee; gorilla Broca's area, located in the inferior frontal gyrus (IFG) of humans, is a key component of the cortical circuitry that subserves language production. In approximately 95% of humans the left hemisphere is dominant for language (Branche et al., 1964), as demonstrated by functional imaging (Petersen et al., 1988) and cortical stimulation studies (Rasmussen and Milner, 1975;Ojemann, 1991). Whereas numerous studies of gross and microscopic structure have revealed anatomic asymmetries that may underlie this functional dominance in humans, an important unresolved question is whether this asymmetric pattern is evolutionarily novel to humans (autapomorphic) or is shared with our closest living relatives, the great apes (synapomorphic).At the microstructural level, Broca's area is comprised of Brodmann's areas 44 and 45 (Aboitiz and Garcia, 1997).

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Deaf signers have extensive experience using their hands to communicate. Using fMRI, we examined the neural systems engaged during the perception of manual communication in 14 deaf signers and 14 hearing non-signers. Participants passively viewed blocked video clips of pantomimes (e.g., peeling an imaginary banana) and action verbs in American Sign Language (ASL) that were rated as meaningless by non-signers (e.g., TO-DANCE). In contrast to visual fixation, pantomimes strongly activated fronto-parietal regions (the mirror neuron system, MNS) in hearing non-signers, but only bilateral middle temporal regions in deaf signers. When contrasted with ASL verbs, pantomimes selectively engaged inferior and superior parietal regions in hearing non-signers, but right superior temporal cortex in deaf signers. The perception of ASL verbs recruited similar regions as pantomimes for deaf signers, with some evidence of greater involvement of left inferior frontal gyrus for ASL verbs. Functional connectivity analyses with left hemisphere seed voxels (ventral premotor, inferior parietal lobule, fusiform gyrus) revealed robust connectivity with the MNS for the hearing non-signers. Deaf signers exhibited functional connectivity with the right hemisphere that was not observed for the hearing group for the fusiform gyrus seed voxel. We suggest that life-long experience with manual communication, and/or auditory deprivation, may alter regional connectivity and brain activation when viewing pantomimes. We conclude that the lack of activation within the MNS for deaf signers does not support an account of human communication that depends upon automatic sensorimotor resonance between perception and action.

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The subarcuate fossa of the petrosal bone houses the petrosal lobule of the cerebellar paraflocculus. Although the subarcuate fossa can be extensive, little is known about its relative size and distribution in primates. Studies indicate parafloccular involvement with cerebellar areas coordinating vestibular, visual, auditory, and locomotor systems. Hypotheses have proposed a role for the paraflocculus in vestibular-oculomotor integration, caudal muscle control, autonomic function, and visual-manual predation. This study examines the morphology and relative extent of the subarcuate fossa/petrosal lobule in a range of living primates. Methods include study of postmortem specimens representing nine mammalian orders, and qualification of the volume of the subarcuate fossa and endocranial cavity in 155 dry primate crania of 36 genera. Results show that, in mammals, the size and morphology of the petrosal lobule is directly related to that of the subarcuate fossa. Craniometric analysis shows that the ratio of subarcuate fossa volume to endocranial volume is largest in lemuriforms. The largest ratio is in Microcebus and Hapalemur. Lorisids show a significant reduction in the size of the subarcuate fossa to almost 50% below the lemuriform mean. Tarsius is near the lemuriform mean. Among platyrrhines, the ratio is high, but significantly reduced compared to lemuiforms. The highest platyrrhine ratio is seen in Ateles, the lowest in Saimiri and Alouatta. Atelids are significantly elevated compared to cebids. In cercopithecids, the fossa is significantly reduced compared to platyrrhines. The trend toward reduction of the cercopithecid fossa is most pronounced in Theropithecus and least evident in Presbytis. In hominoids, the fossa is present only in Hylobates. In great apes and humans, other than Gorilla, the petromastoid canal occupies a similar location to the subarcuate fossa of other primates, but is not homologous to it. Neither the subarcuate fossa nor the petromastoid canal are present in Gorilla. A graded reduction of the subarcuate fossa/petrosal lobule is evident among primates which evolved later in time. The relative size of this cerebellar lobule within primates may reflect size-related factors and/or degree of neocortical evolution as these relate to usage of a specific sensory-mediated locomotor behavior. The subarcuate fossa may serve as an indicator to the differentiation of the petrosal lobule of the paraflocculus in fossil forms.