The throwing shoulder in pitchers frequently exhibits a paradox of glenohumeral joint motion, in which excessive external rotation is present at the expense of decreased internal rotation. The object of this study was to determine the role of humeral head retroversion in relation to increased glenohumeral external rotation. Glenohumeral joint range of motion and laxity along with humeral head and glenoid version of the dominant versus nondominant shoulders were studied in 25 professional pitchers and 25 nonthrowing subjects. Each subject underwent a computed tomography scan to determine bilateral humeral head and glenoid version. The throwing group demonstrated a significant increase in the dominant shoulder versus the nondominant shoulder in humeral head retroversion, glenoid retroversion, external rotation at 90 degrees, and external rotation in the scapular plane. Internal rotation was decreased in the dominant shoulder. Total range of motion, anterior glenohumeral laxity, and posterior glenohumeral laxity were found to be equal bilaterally. The nonthrowing group demonstrated no significant difference in humeral head retroversion, glenoid retroversion, external rotation at 90 degrees or external rotation in the scapular plane between shoulders, and no difference in internal rotation at 90 degrees, total motion, or laxity. A comparison of the dominant shoulders of the two groups indicated that both external rotation at 90 degrees and humeral head retroversion were significantly greater in the throwing group.
As diurnal rodents with a well-developed visual system, squirrels provide a useful comparison of visual system organization with other highly visual mammals such as tree shrews and primates. Here, we describe the projection pattern of gray squirrel superior colliculus (SC) with the large and well-differentiated pulvinar complex. Our anatomical results support the conclusion that the pulvinar complex of squirrels consists of four distinct nuclei. The caudal (C) nucleus, distinct in cytochrome oxidase (CO), acetylcholinesterase (AChE), and vesicular glutamate transporter-2 (VGluT2) preparations, received widespread projections from the ipsilateral SC, although a crude retinotopic organization was suggested. The caudal nucleus also received weaker projections from the contralateral SC. The caudal nucleus also projects back to the ipsilateral SC. Lateral (RLl) and medial (RLm) parts of the previously defined rostral lateral pulvinar (RL) were architectonically distinct, and each nucleus received its own retinotopic pattern of focused ipsilateral SC projections. The SC did not project to the rostral medial (RM) nucleus of the pulvinar. SC injections also revealed ipsilateral connections with the dorsal and ventral lateral geniculate nuclei, nuclei of the pretectum, and nucleus of the brachium of the inferior colliculus and bilateral connections with the parabigeminal nuclei. Comparisons with other rodents suggest that a variously named caudal nucleus, which relays visual inputs from the SC to temporal visual cortex, is common to all rodents and possibly most mammals. RM and RL divisions of the pulvinar complex also appear to have homologues in other rodents.
The density of cells and neurons in the neocortex of many mammals varies across cortical areas and regions. This variability is, perhaps, most pronounced in primates. Nonuniformity in the composition of cortex suggests regions of the cortex have different specializations. Specifically, regions with densely packed neurons contain smaller neurons that are activated by relatively few inputs, thereby preserving information, whereas regions that are less densely packed have larger neurons that have more integrative functions. Here we present the numbers of cells and neurons for 742 discrete locations across the neocortex in a chimpanzee. Using isotropic fractionation and flow fractionation methods for cell and neuron counts, we estimate that neocortex of one hemisphere contains 9.5 billion cells and 3.7 billion neurons. Primary visual cortex occupies 35 cm 2 of surface, 10% of the total, and contains 737 million densely packed neurons, 20% of the total neurons contained within the hemisphere. Other areas of high neuron packing include secondary visual areas, somatosensory cortex, and prefrontal granular cortex. Areas of low levels of neuron packing density include motor and premotor cortex. These values reflect those obtained from more limited samples of cortex in humans and other primates.
Receptive fields of neurons in somatosensory area 3b of monkeys are typically described as restricted to part of a single digit or palm pad. However, such neurons are likely involved in integrating stimulus information from across the hand. To evaluate this possibility, we recorded from area 3b neurons in anesthetized owl monkeys with 100-electrode arrays, stimulating two hand locations with electromechanical probes simultaneously or asynchronously. Response magnitudes and latencies of single- and multiunits varied with stimulus conditions, and multiunit responses were similar to single-unit responses. The mean peak firing rate for single neurons stimulated within the preferred location was estimated to be ∼26 spike/s. Simultaneous stimulation with a second probe outside the preferred location slightly decreased peak firing rates to ∼22 spike/s. When the nonpreferred stimulus preceded the preferred stimulus by 10-500 ms, peak firing rates were suppressed with greatest suppression when the nonpreferred stimulus preceded by 30 ms (∼7 spike/s). The mean latency for single neurons stimulated within the preferred location was ∼23 ms, and latency was little affected by simultaneous paired stimulation. However, when the nonpreferred stimulus preceded the preferred stimulus by 10 ms, latencies shortened to ∼16 ms. Response suppression occurred even when stimuli were separated by long distances (nonadjacent digits) or long times (500 ms onset asynchrony). Facilitation, though rare, occurred most often when the stimulus onsets were within 0-30 ms of each other. These findings quantify spatiotemporal interactions and support the hypothesis that area 3b is involved in widespread stimulus integration.
Despite the lack of ipsilateral receptive fields (RFs) for neurons in the hand representation of area 3b of primary somatosensory cortex, interhemispheric interactions have been reported to varying degrees. We investigated spatiotemporal properties of these interactions to determine the following: response types, timing between stimuli to evoke the strongest bimanual interactions, topographical distribution of effects, and their dependence on similarity of stimulus locations on the two hands. We analyzed response magnitudes and latencies of single neurons and multineuron clusters recorded from 100-electrode arrays implanted in one hemisphere of each of two anesthetized owl monkeys. Skin indentations were delivered to the two hands simultaneously and asynchronously at mirror locations (matched sites on each hand) and nonmirror locations. Since multiple neurons were recorded simultaneously, stimuli on the contralateral hand could be within or outside of the classical RFs of any given neuron. For most neurons, stimulation on the ipsilateral hand suppressed responses to stimuli on the contralateral hand. Maximum suppression occurred when the ipsilateral stimulus was presented 100 ms before the contralateral stimulus onset ( p Ͻ 0.0005). The longest stimulus onset delay tested (500 ms) allowed contralateral responses to recover to control levels ( p ϭ 0.428). Stimulation on mirror digits did not differ from stimulation on nonmirror locations ( p ϭ 1.000). These results indicate that interhemispheric interactions are common in area 3b, somewhat topographically diffuse, and maximal when the suppressing ipsilateral stimulus precedes the contralateral stimulus. Our findings point to a neurophysiological basis for "interference" effects found in human psychophysical studies of bimanual stimulation.
Tree shrews are small squirrel-like mammals that are the closest living relative to primates available for detailed neurobiological study. In a recent study (Remple et al. [2006] J. Comp. Neurol. 497:133-154), we provided anatomical and electrophysiological evidence that the frontoparietal cortex of tree shrews has two motor fields (M1 and M2) and five somatosensory fields (3a, 3b, S2, somatosensory caudal area [SC], and parietal ventral area [PV]). In the present study, we injected anatomical tracers into M1, M2, 3a, 3b, SC, and posterior parietal cortex to establish the ipsilateral cortical connections of these areas. The results provide evidence for a number of new cortical areas including medial motor and somatosensory areas (MMA and MSA), three posterior parietal areas (PPd, PPv, and PPc), and an area ventral to temporal inferior cortex (TIV). Ml receives topographic projections from M2, MMA, 3a, and PPv, and nontopographic connections from the temporal anterior and dorsal areas (TA and TD), PPc, TIV, and MSA. The connections of M2 are similar to those of M1, except that M2 receives denser projections from TIV, PPc, and dorsal frontal cortex and sparser input from M1. Areas 3a, 3b, and SC receive dense topographic projections from each other, S2, and PV and sparser connections from PPd and PPv. Area 3a receives additional input from posterior parietal and temporal regions and from M1 and MMA. Overall, the frontoparietal connections of tree shrew cortex are most similar to those of prosimian primates and quite different from those of more distant relatives such as rats.
Tactile discrimination depends on integration of information from the discrete receptive fields (RFs) of peripheral sensory afferents. Because this information is processed over a hierarchy of subcortical nuclei and cortical areas, the integration likely occurs at multiple levels. The current study presents results indicating that neurons across most of the extent of the hand representation in monkey primary somatosensory cortex (area 3b) interact, even when these neurons have separate RFs. We obtained simultaneous recordings by using a 100-electrode array implanted in the hand representation of primary somatosensory cortex of two anesthetized owl monkeys. During a series of 0.5-s skin indentations with single or dual probes, the distance between electrodes from which neurons with synchronized spike times were recorded exceeded 2 mm. The results provide evidence that stimuli on different parts of the hand influence the degree of synchronous firing among a large population of neurons. Because spike synchrony potentiates the activation of commonly targeted neurons, synchronous neural activity in primary somatosensory cortex can contribute to discrimination of complex tactile stimuli.classical receptive field ͉ neuronal synchrony ͉ primate ͉ two-point stimulation ͉ Utah array H umans and other primates use their hands to make tactile discriminations that guide choices and actions. The vast majority of these choices are based on stimuli that are presented to sites across the hand or hands. Transformation of these scattered information sources from receptors in distinct patches of skin into one percept presumes integration within the central nervous system. This integration could occur at several levels, but here we consider the primary somatosensory cortex (S1 or area 3b). Though area 3b contains a detailed somatotopic representation of the hand and neurons with small receptive fields (RFs), considerable integration across hand locations may occur at this level via horizontal connections within area 3b.One way of examining neuronal interactions is through spike timing synchrony. When the spikes of two neurons occur together more often than expected by chance, we can infer that those neurons are part of the same local network. The two neurons may receive a common input that drives the synchronous firing, or the neurons may be synaptically connected. Our focus was on quantifying integration in anesthetized owl monkeys (Aotus trivirgatus) in the form of spike synchrony of neuronal activity in layers 2/3 of S1. Analysis of firing rate will be presented elsewhere.Using a 100-electrode array (Cyberkinetics) covering 4 mm ϫ 4 mm of cortical area ( Fig. 1 A and B), we examined sensory input integration in the area 3b hand representation by focusing on correlations in spike timing in pairs of neurons responding to the same or different stimulus probes (1-mm-diameter contact surface). We examined interactions across, rather than within, digit and palm pads to determine the extent of spatial integration in the area 3b hand rep...
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