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Neuronal projections to the auditory cortex arise largely from the thalamus and other cortical areas muscle relaxant used for migraines proven skelaxin 400mg, although inputs from other structures have been demonstrated infantile spasms 9 month old order skelaxin 400mg. Many of these distinct areas have unclear functions and await further physiological investigation to determine their contributions to auditory processing spasms upper left abdomen order 400mg skelaxin. Organization of Auditory Cortex Thalamic Input Distinct cortical areas are generally identified by architectonic differences and confirmed by neuroanatomical tracing studies and neurophysiological properties muscle spasms 7 little words effective 400 mg skelaxin. These areas are typically subdivided into groups based upon their proximity and the types of subcortical projections they receive. This pathway appears to transmit strictly auditory information at short latencies using projection neurons having small receptive field sizes and arranged topographically in the thalamus. Rat A1 Auditory cortex Cat Central sulcus Lateral sulcus Human A1 Dorsal Auditory cortex Rostral Figure 1 Approximate locations of auditory cortex on the surface of rat, cat, and human cortex. Some areas in the human brain are visible only by cutting away overlying brain tissue. Auditory Cortex Structure and Circuitry 67 Of the distinct thalamocortical pathways, the ventral pathway projects to fewer cortical areas and connects to these areas with relatively patchy distributions predominantly in cortical laminae 3b and 4. The dorsal pathway contrasts with the ventral pathway in most if not all anatomical and physiological features. Compared with the ventral pathway, the dorsal pathway receives different, nontonotopic inputs from the inferior colliculus, projects largely to distinct areas of cortex, and sends less dense axonal collaterals more widely throughout most cortical laminae, although the largest projections are also to layers 3b and 4. The neurons composing this pathway exhibit less topographic organization, broader tuning, longer and more variable response latencies, and tendencies to habituate to ongoing stimuli. The projection patterns of the individual nuclei of the dorsal division appear to be more diverse than those of the ventral division. Its neurons typically respond at relatively short latencies (although with potential cell-to-cell variation) and some project with large axons, possibly indicating a fast connection from thalamus to cortex. These projections likely underlie associative, learning, and sensory integration functions. Area Parcellation Distinct areas of auditory cortex are typically discerned by cytoarchitectural differences with adjacent areas and the pattern of thalamic input. Additional criteria for distinctiveness typically include variation in functional responses, although most of the anatomically determined areas of auditory cortex have yet to be explored physiologically in any detail. In addition, certain developmental stages in some species yield differential staining properties of A1, with stains differentiating a variety of molecules, such as cytochrome oxidase, acetylcholinesterase, parvalbumin, and neurofilament protein. In primates, three groups of areas have been delineated based largely upon architecture, thalamocortical connections, interconnections with one another, and physiology. These areas appear to be tonotopically organized but are difficult to study with classical, simple sound stimuli. Connections of Auditory Cortex Local Structure of Primary Auditory Cortex A1 represents both the most studied auditory cortical area and is the focus of the most research. For many years even such a basic organizational structure as tonotopy the topographic representation of the cochlea of A1 was debated because of the great variety of functional responses encountered during physiological experiments. Many modern lines of evidence, however, demonstrate tonotopy in A1, but other potential topographic structures remain unclear. In tonotopic areas, each sound frequency is represented in a narrow strip along the surface of the cortex (see Figure 3), and many attempts have been made to identify other sound properties that may be mapped along each of these strips. Many sound properties are known to be mapped nonrandomly onto the surface of A1, and computational studies imply that this relatively large number (>5) makes it unlikely that any one of these parameters is mapped orthogonally to the tonotopic axis. Spatial segregation of processing appears to be true for frequency, as well, and evidence exists for substantial local frequency interactions in A1, currently of unknown function but potentially for the sensitive detection of frequency modulation sweeps and extraction of harmonicity underlying the perception of pitch both important components of species-specific vocalizations. The patchiness of other stimulus parameters, such as neuronal receptive field bandwidth, is predicted by computational topographic models. Local connections within cortical columns of A1 remain to be further elucidated, but, as with other primary sensory areas, it is likely that specificity of interlaminar connections in A1 will become more apparent with further experimentation. Layer 4 cells tend to receive most of their excitatory input from other layer 4 cells. Layer 2/3 pyramidal cells appear to receive most of their excitatory input from either other layer 2/3 cells or layer 4 cells, reminiscent of the lemniscal/paralemniscal projection dichotomy apparent in rodent barrel cortex. How these local intracolumnar connections interface with thalamocortical and corticocortical connections remains unknown, but potentially they represent multiple modes of information transfer through A1 to other cortical areas. Interconnections between Areas of Auditory Cortex Auditory cortical interconnections follow the parcellation created by thalamocortical projections.
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Two types of neural constraints have been invoked: modalityspecificity and domain-specificity spasms right flank proven 400 mg skelaxin. The second class of theories muscle relaxant elderly trusted skelaxin 400mg, based on the correlated structure principle spasms during sleep buy 400mg skelaxin, assumes that the organization of conceptual knowledge in the brain is a reflection of the statistical co-occurrence of object properties in the world spasms 1983 download generic skelaxin 400mg. Neuropsychological evidence, and more recently findings from functional neuroimaging, have figured centrally in attempts to evaluate extant theories of the organization of conceptual knowledge. Here we outline the main theoretical perspectives as well as the empirical phenomena that have been used to inform these perspectives. Modality-Specific Hypotheses the first class of theories based on the neural structure principle assumes that the principal determinant of the organization of conceptual knowledge is the sensorymotor modality. For instance, the knowledge that hammers are shaped like a T would be stored in a semantic subsystem dedicated to representing the visual structure of objects, while the information that hammers are used to pound nails would be represented in a semantic subsystem dedicated to functional knowledge of objects. There have been many proposals based on the modality-specific assumption (Beauvois, 1982; Warrington and McCarthy, 1983, 1987; Warrington and Shallice, 1984; Allport, 1985; Martin et al. One way to distinguish between these proposals concerns whether, and to what extent, conceptual knowledge is assumed to be repre- sented independently of sensory-motor processes. Central to such proposals is the notion of simulation, or the automatic reactivation of sensory-motor information in the course of conceptual processing. Toward the other end of the continuum are modality-based hypotheses of the organization of conceptual knowledge that assume that sensory-motor systems may be damaged without compromising the integrity of conceptual knowledge (Martin et al. Domain-Specific Hypotheses A second class of proposals based on the neural structure principle assumes that the principal determinant of the organization of conceptual knowledge is semantic category. For instance, in this view, it may be argued that conceptual knowledge of conspecifics and conceptual knowledge of animals are represented and processed by functionally dissociable processes/systems. Crucially, in this view, the first order principle of organization of conceptual processing is semantic category and not the modality through which that information is typically processed. One proposal along these lines, the Domain-Specific Hypothesis (Caramazza and Shelton, 1998), argues that conceptual knowledge is organized by specialized (and functionally dissociable) neural circuits innately determined to the conceptual processing of different categories of objects. However, not all Domain-Specific theories assume that the organization of the adult semantic system is driven by innate parameters. Feature-Based Hypotheses the class of hypotheses based on the correlated structure principle has focused on articulating the structure of 105 106 Category-Specific Knowledge semantic memory at the level of semantic features. There are many and sometimes diverging proposals along these lines; common to all of them is the assumption that the relative susceptibility to impairment (under conditions of neurological damage) of different concepts is a function of statistical properties of the semantic features that comprise those concepts. For instance, on some models, the degree to which features are shared by a number of concepts is contrasted with their relative distinctiveness (Devlin et al. Another dimension that is introduced by some theorists concerns dynamical properties of damage in the system; for instance, Tyler and Moss assume that features that are more correlated with other features will be more resistant to damage, due to greater reciprocal activation (or support) from those features with which they are correlated (but see Caramazza et al. Distinctive features, on the other hand, will not receive as much reciprocal support, and will thus be more susceptible to damage. More recently, theorists have expanded on the original proposal of Tyler and colleagues, adding dimensions such as familiarity, typicality, and relevance. Feature-based models of semantic memory have in general emphasized an empirical, bottom up, approach to modeling the organization of semantic memory, usually drawing on feature generation tasks. For this reason, featurebased models have been useful in generating hypotheses about the types of parameters that may contribute to the organization of conceptual knowledge. Clues from Cognitive Neuropsychology Neuropsychological studies of patients with semantic impairments have figured centrally in developing and evaluating the hypotheses outlined above. Of particular importance has been a clinical profile described as category-specific semantic deficit. Patients with categoryspecific semantic deficits present with disproportionate or even selective difficulty for conceptual knowledge of stimuli from one semantic category compared to other semantic categories. For instance, the reports of categoryspecific impairment by Warrington and her collaborators. Since those seminal reports, the phenomenon of category-specific semantic deficit has been documented by a number of investigators (for recent reviews of the clinical evidence, see Humphreys and Forde, 2001; Tyler and Moss, 2001; Capitani et al.
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