Three theories currently compete to explain the conceptual deficits that result from brain damage: sensory-functional theory, domain-specific theory, and conceptual structure theory. We argue that all three theories capture important aspects of conceptual deficits, and offer different insights into their origins. Conceptual topography theory (CTT) integrates these insights, beginning with A. R. Damasio’s (1989) convergence zone theory and elaborating it with the similarity-in-topography (SIT) principle. According to CTT, feature maps in sensory-motor systems represent the features of a category’s exemplars. A hierarchical system of convergence zones then conjoins these features to form both property and category representations. According to the SIT principle, the proximity of two conjunctive neurons in a convergence zone increases with the similarity of the features they conjoin. As a result, conjunctive neurons become topographically organised into local regions that represent properties and categories. Depending on the level and location of a lesion in this system, a wide variety of deficits is possible. Consistent with the literature, these deficits range from the loss of a single category to the loss of multiple categories that share sensory-motor properties.

A key issue in cognitive neuroscience concerns the neural representation of conceptual knowledge. Currently, debate focuses around the issue of whether there are neural regions specialised for the processing of specific semantic attributes or categories, or whether concepts are represented in an undifferentiated neural system. Neuropsychological studies of patients with selective semantic deficits and previous neuroimaging studies do not unequivocally support either account. We carried out a PET study to determine whether there is any regional specialisation for the processing of concepts from different semantic categories using picture stimuli and a semantic categorisation task. We found robust activation of a large semantic network extending from left inferior frontal cortex into the inferior temporal lobe and including occipital cortex and the fusiform gyrus. The only category effect that we found was additional activation for animals in the right occipital cortex, which we interpret as being due to the extra visual processing demands required in order to differentiate one animal from another. We also carried out analyses in specific cortical regions that have been claimed to be preferentially activated for various categories, but found no evidence of any differential activation as a function of category. We interpret these data within the framework of cognitive accounts in which conceptual knowledge is represented within a nondifferentiated distributed system.

& How objects are represented and processed in the brain is a central topic in cognitive neuroscience. Previous studies have shown that knowledge of objects is represented in a featurebased distributed neural system primarily involving occipital and temporal cortical regions. Research with nonhuman primates suggest that these features are structured in a hierarchical system with posterior neurons in the inferior temporal cortex representing simple features and anterior neurons in the perirhinal cortex representing complex conjunctions of features (Bussey & Saksida, 2002; Murray & Bussey, 1999). On this account, the perirhinal cortex plays a crucial role in object identification by integrating information from different sensory systems into more complex polymodal feature conjunctions. We tested the implications of these claims for human object processing in an event-related fMRI study in which we presented colored pictures of common objects for 19 subjects to name at two levels of specificity—basic and domain. We reasoned that domain-level naming requires access to a coarsergrained representation of objects, thus involving only posterior regions of the inferior temporal cortex. In contrast, basic-level naming requires finer-grained discrimination to differentiate between similar objects, and thus should involve anterior temporal regions, including the perirhinal cortex. We found that object processing always activated the fusiform gyrus bilaterally, irrespective of the task, whereas the perirhinal cortex was only activated when the task required finer-grained discriminations. These results suggest that the same kind of hierarchical structure, which has been proposed for object processing in the monkey temporal cortex, functions in the human. &

The way in which meaning is represented and processed in the brain is a key issue in cognitive neuroscience, which can be usefully addressed by functional imaging techniques. In contrast to previous imaging studies of semantic knowledge, which have primarily used blocked designs, in this study we use an eventrelated fMRI (erfMRI) design, which has the advantage of enabling events to be presented pseudorandomly, thus reducing strategic processes and enabling more direct comparison with psychological behavioral studies. We used a semantic categorization task in which events were words representing either artifact or natural kinds concepts. Significant areas of activation for semantic processing included inferior frontal lobe bilaterally (BA 47) and left temporal regions, both inferior (BA 36 and 20) and middle (BA 21). These are areas that have been identified in previous neuroimaging studies of semantic knowledge. However, there were no significant differences between artifact and natural kinds concepts. These results are consistent with our previous imaging studies using blocked designs and suggest that conceptual knowledge is represented in a unitary, distributed neural system undifferentiated by domain of knowledge. These findings demonstrate that event-related designs can generate activations that are similar to those seen in blocked designs investigating semantics and, moreover, offer a greater capacity for interpretation free from the confounds of block effects. © 2002 Elsevier Science (USA)

The work described in this chapter is motivated by the conviction that a cognitive theory of semantic memory is best-suited to investigate the functional and neural bases of the semantic memory system. The advantage of this approach is that detailed hypotheses about the structure and function of the semantic system can be formulated and then tested in behavioral experiments with healthy individuals and neurologically impaired patients. The challenge is then to identify the neural correlates of these experimentally validated cognitive structures and processes, i.e., their neural substrates and mechanisms. The cognitive model provides a detailed framework for this investigation which, when combined with the appropriate functional-neuroanatomical technique, provides the potential to meet this challenge.

How objects are represented and processed in the brain remains a key issue in cognitive neuroscience. We have developed a conceptual structure account in which category-specific semantic deficits emerge due to differences in the structure and content of concepts rather than from explicit divisions of conceptual knowledge in separate stores. The primary claim is that concepts associated with particular categories (e.g., animals, tools) differ in the number and type of properties and the extent to which these properties are correlated with each other. In this review, we describe recent neuropsychological and neuroimaging studies in which we have extended our theoretical account by incorporating recent claims about the neuroanatomical basis of feature integration and differentiation that arise from research into hierarchical object processing streams in nonhuman primates and humans. A clear picture has emerged in which the human perirhinal cortex and neighbouring anteromedial temporal structures appear to provide the neural infrastructure for making fine-grained discriminations among objects, suggesting that damage within the perirhinal cortex may underlie the emergence of category-specific semantic deficits in brain-damaged patients. Understanding the nature and organization of conceptual knowledge in the brain requires

■ Research on the spatio-temporal dynamics of visual object recognition suggests a recurrent, interactive model whereby an initial feedforward sweep through the ventral stream to prefrontal cortex is followed by recurrent interactions. However, critical questions remain regarding the factors that mediate the degree of recurrent interactions necessary for meaningful object recognition. The novel prediction we test here is that recurrent interactivity is driven by increasing semantic integration demands as defined by the complexity of semantic information required by the task and driven by the stimuli. To test this prediction, we recorded magnetoencephalography data while participants named living and nonliving objects during two naming tasks. We found that the spatio-temporal dynamics of neural activity were modulated by the level of semantic integration required. Specifically, source reconstructed time courses and phase synchronization measures showed increased recurrent interactions as a function of semantic integration demands. These findings demonstrate that the cortical dynamics of object processing are modulated by the complexity of semantic information required from the visual input. ■

fMRI activation to conceptual stimuli to evaluate methods for

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