Correlations between hand preference and cortical thickness in the secondary somatosensory (SII) cortex of the common marmoset, callithrix jacchus.

[Correction Notice: An erratum for this article was reported in Vol 123(2) of Behavioral Neuroscience (see record 2009-04037-027). In this article, there were errors in figure 4 on p.1347 (missing labels along the x-axis) and in figure 5 on p. 1348 (distortion in the reproduction of the panels). The corrected figures are included.] Cortical asymmetries are well established in humans for language and motor regions and correlate with handedness. Here the authors investigate structural differences in the hemispheres of left- and right-handed common marmosets using surface photography and histology. The hand preferences of 11 marmosets were assessed over their adult life span using a simple reaching task. A significant correlation was found between the length of the right lateral sulcus/brain weight and the % right-hand preference (r = .86, p = .001). Cortical thickness on the superior bank of the right lateral sulcus posteriorly was also positively correlated with % right-hand preference (r = .69, p = .025). Comparison of this site with previously published functional maps of the marmoset cortex show this area corresponds to SII, a region involved in tactile processing and somatosensory discriminations. It is suggested that the correlation between SII thickness and right-hand preference would be consistent with the fact that right-handed marmosets are more proactive than left-handers in exploring novel objects by touch. Enlargement of a cortical area involved tactile discriminations could be a precursor to the evolution of right-handedness as a population bias. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differences in social and vocal behavior between left- and right-handed common marmosets (Callithrix jacchus).

Common marmosets (Callithrix jacchus) show either a left- or right-hand preference for reaching to pick up food and they retain the same preference throughout adult life. We compared the behavior of 10 right-handed and 10 left-handed marmosets, matched for age and sex. They were presented with live crickets both when alone and when in their social group. The marmosets captured more crickets and the latency to capture the first cricket was shorter when they were in a group than when they were alone. This effect of social facilitation was significantly greater for right- than left-handed individuals. The number of vocalizations (tsik, crackle, very brief whistle, cough, and phee) produced by the left- and right-handed marmosets differed significantly: right-handed marmosets produced an increased number of all of these calls when the crickets were presented, whereas left-handed marmosets did not show a change from pretesting levels. The right-handed marmosets also produced more tsik (mobbing) calls than left-handed marmosets when they were presented with a fear-inducing stimulus and performed more head cocking and parallax movements than the left-handed marmosets. Hence, hand preference is associated with differences in exploratory and social behavior, the latter including vocal communication. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Responses of neurons of the first and second somatosensory zones of the cortex to peripheral, thalamic and cortical stimulation

Experiments were performed on cats immobilized with d-tubocurarine or myorelaxin. Neuronal responses were studied in the first somatosensory cortex (SI) to the second somatosensory cortex (SII), ventroposterior nucleus (VP) and contralateral forepaw stimulation. Besides, neuronal responses in SII to SI, VP and contralateral forepaw stimulation were also studied. It was shown that in SII the percentage of neurons excited by afferent volley with two or more synaptic change-overs in the cerebral cortex was larger than in SI. Neurons of SI and SII responded to cortical stimulation ortho- and antidromically, thus confirming the existence of bilateral cortico-cortical connections. Both in SI and SII, PSPs to cortical stimulation were similar in character to PSPs in the same neurons to VP stimulation. In 50.0% of SI neurons and 37.1 of SII neurons the difference in latencies of orthodromic spike potentials to VP and cortical stimulation was less than 1.0 ms. In 19.6% of SI neurons and 41.4% of SII neurons the latency of the response to cortical stimulation was 1.6-4.7 ms shorter than that of the response in the same neuron to VP stimulation. It is supposed that impulses from SI participate significantly in afferent activation of SII neurons.     

Sensorimotor integration in human primary and secondary somatosensory cortices

We measured somatosensory evoked fields (SEFs) to electric median nerve stimuli from eight healthy subjects with a whole-scalp 122-channel neuromagnetometer in two different conditions: (i) 'rest', with stimuli producing clear tactile sensation without any motor movement, and (ii) 'contraction' with exactly the same stimuli as in 'rest', but with the subjects maintaining sub-maximal isometric contraction in thenar muscles of the stimulated hand. The aim was to study the role of the primary (SI) and secondary somatosensory (SII) cortices in sensorimotor integration. The amplitude of the SI response N20m did not change with coincident isometric contraction, whereas P35m was significantly reduced. On the contrary, activation of contra- and ipsilateral SII cortices was significantly enhanced during the contraction. We suggest that isometric contraction facilitates activation of SII cortices to tactile stimuli, possibly by decreasing inhibition from the SI cortex. The enhanced SII activation may be related to tuning of SII neurons towards relevant tactile input arising from the region of the body where the muscle activation occurs.     

Patterns of lateral sensory cortical activation determined using functional magnetic resonance imaging

OBJECT: Functional magnetic resonance (fMR) imaging was performed in human volunteers to determine the lateral perisylvian cortical areas activated by innocuous cutaneous stimulation. METHODS: Eight volunteers who underwent 53 separate experiments form the basis of this report. Eight contiguous coronal slices were obtained using echoplanar fMR imaging techniques while participants were at rest and while somatosensory activation stimuli consisting of vibration or air puffs were delivered to various body areas. The data were analyzed using Student's t-test and cluster analysis to determine significant differences between the resting and activated states. The findings were as follows: the areas in the lateral cortex activated by the stimuli were the primary sensory cortex (SI), the second somatosensory area (SII), the insula, the superior parietal lobule, and the retroinsular parietal operculum (RIPO). Somatotopy was demonstrable in SI but not in the other areas identified. There was a surprisingly low correlation between the amount of cortex activated in the various areas, which could mean separate inputs and functions for the areas identified. The highest correlation was found between activity in SII and RIPO (0.69). CONCLUSIONS: The authors maintain that fMR imaging can be used to identify multiple lateral somatosensory areas in humans. Somatotopy is demonstrated in SI but not in the other lateral cortical sensory areas. The correlations between the amounts of cortex activated in the different lateral sensory areas are low. Recognition of the multiple lateral sensory areas is important both for understanding sensory cortical function and for safe interpretation of studies designed to identify the central sulcus by activating SI.     

Attention modulates both primary and second somatosensory cortical activities in humans: a magnetoencephalographic study

To clarify the role of primary and second somatosensory cortex (SI and SII) in somatosensory discrimination, we recorded somatosensory evoked magnetic fields during a stimulus strength discrimination task. The temporal pattern of cortical activation was analyzed by dipole source model coregistered with magnetic resonance image. Stimulus intensity was represented in SI as early as 20 ms after the stimulus presentation. The later components of SI response (latency 37.7 and 67.9 ms) were enhanced by rarely presented stimuli (stimulus deviancy) during passive and active attention. This supports an early haptic memory mechanism in human primary sensory cortex. Contra- and ipsilateral SII responses followed the SI responses (latency 124.6 and 138.3 ms, respectively) and were enhanced by attention more prominently than the SI responses. Active attention increased SII but not SI activity. These results are consistent with the concept of ventral somatosensory pathway that SI and SII are hierarchically organized for passive and active detection of discrete stimuli.     

Cortical connections of the frontoparietal opercular areas in the rhesus monkey

The connections of the frontoparietal opercular areas were studied in rhesus monkeys by using antero- and retrograde tracer techniques. The rostral opercular cortex including the gustatory and proisocortical motor (ProM) areas is connected with precentral areas 3, 1, and 2 as well as with the rostral portion of the opercular area which resembles the second somatosensory type of cortex (area SII) and the ventral portion of area 6. Its distant connections are with the ventral portion of prefrontal areas 46, 11, 12, and 13 as well as with the rostral insula and cingulate motor area (CMAr). The mid opercular region (areas 1 and 2) is connected with pre- and postcentral areas 3, 1, and 2 as well as SII. Additionally, it is connected with the ventral portion of area 6, area 44, area ProM, the gustatory area, and the rostral insula. Its distant connections are with area 4, the ventral portion of area 46, area 7b, and area POa in the intraparietal sulcus (IPS). The rostral parietal opercular region is connected with the postcentral portions of areas 3, 1, and 2; areas 5, 7, and SII; the gustatory area; and the vestibular area. Its other connections are with area 4, area 44, the ventral portion of area 46, area ProM, CMAr, and the supplementary motor area (SMA). The caudal opercular region is connected with the dorsal portion of area 3; areas 2, 5, and 7a; and areas PEa as well as IPd of IPS. It is also connected with area SII, insula, and the superior temporal sulcus. Its distant connections are with area 44; the dorsal portion of area 8 and the ventral portion of area 46; as well as CMAr, SMA, and the supplementary sensory area. This connectivity suggests that the ventral somatosensory areas are involved in sensorimotor activities mainly related to head, neck, and face structures as well as to taste. Additionally, these areas may have a role in frontal (working) and temporal (tactile) memory systems.     

The ipsilateral and contralateral connections of the fifth somatosensory area (SV) in the cat cerebral cortex

THE ipsilateral and contralateral corticocortical connections to the fifth somatosensory area (SV) in the feline cortex were determined from the location of retrogradely labelled cells following a single injection of HRP into SV. The injection was made into physiologically defined components of the body representation in SV. After injection of HRP into the face regions of SV, HRP-labelled cells were located ipsilaterally in areas 6 beta, 3b and 1-2 of the primary somatosensory (SI), in the second somatosensory (SII), third somatosensory (SIII), and fourth somatosensory (SIV) areas, along the ansate sulcus, and in areas 5a and 6a beta of the ipsilateral cortex, as well as in area 1-2 of SI and in SV of the contralateral cortex. On the other hand, after HRP had been injected into the trunk/hindlimb area, HRP-labelled cells were located in areas 3a, 1-2 of SI, in area 5, in SII, in SIII and in SIV of the ipsilateral cortex, as well as in area 1-2 of SI, and in SV of the contralateral cortex. The extent of these interconnections suggests that SV receives multiple sensory inputs and may function to integrate this information.     

Primary motor cortex asymmetry is correlated with handedness in capuchin monkeys (cebus apella).

Humans exhibit a population-wide tendency toward right-handedness, and structural asymmetries of the primary motor cortex are associated with hand preference. Reported are similar asymmetries correlated with hand preference in a New World monkey (Cebus apella) that does not display population-level handedness. Asymmetry of central sulcus depth is significantly different between left-handed and right-handed individuals as determined by a coordinated bimanual task. Left-handed individuals have a deeper central sulcus in the contralateral hemisphere; right-handed individuals have a more symmetrical central sulcus depth. Cerebral hemispheric specialization for hand preference is not uniquely human and may be more common among primates in general. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Comparative assessment of the role of the primary and secondary somatosensory regions of the cerebral cortex during conditioned reflex behavior in animals]

The first (SI) and second (SII) somatosensory cortical areas were ablated in one group of cats after preliminary learning of tactile differentiation of rough and smooth surfaces of the floor coating in a special chamber. Somatosensory areas were ablated in another group after learning an adequate choice of the reinforcement side in response to a bell and a metronome. Unilateral and bilateral ablation of SI affected but little the elaboration and achievement of the above acts. Unilateral and bilateral ablation of area SII resulted in a sharp impairment of the tactile differentiation of the surfaces (35% correct responses after unilateral and 26% after bilateral ablation) and a less pronounced disturbances in the choice of the side of reinforcement in response to the bell and the metronome (80% of correct choice after unilateral and 72% after a bilateral ablation.     

Hand preference predicts reactions to novel foods and predators in marmosets (Callithrix geoffroyi).

In chimpanzees (Pan troglodytes) and common marmosets (Callithrix jacchus), left-handed individuals are less likely than right-handed individuals to explore new objects and situations, suggesting a relationship between the hemispheric specialization of emotional states and motor function. To further explore this relationship and to test the hypothesis that fearfulness is related to hand preference, we assessed willingness to approach, sniff, and taste novel foods, and the duration of freeze reactions in response to hawk calls, in 18 Geoffroy's marmosets (Callithrix geoffroyi). In accordance with these hypotheses, left-handed marmosets were slower to explore novel foods and slower to emerge from a freeze response than right-handed marmosets. Hand preference and at least some features of temperament seem to be related in this and other species of primates. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Neural activity in SII modifies sensory evoked potentials in SI in awake rats

The function of the projection from the secondary somatosensory cortex (SII) to the primary somatosensory cortex (SI) in rats was investigated by recording sensory evoked potentials (SEP) in SI during glutamate activation and lidocaine blockade of SII. In anesthetized animals, glutamate stimulation of SII decreased SEP latency and increased SEP amplitude, whereas no changes were evident during lidocaine blockade of SII. In awake animals, a second, later component of the SEP appeared. This second component was almost completely eliminated during lidocaine blockade of SII. We conclude that the projection from SII to SI in rats slightly facilitates the SEP response in anesthetized animals and is responsible for a major portion of the late component of the SEP in awake animals.     

Isoflurane anesthesia blunts cerebral responses to noxious and innocuous stimuli: a fMRI study

We used functional magnetic resonance imaging to determine how isoflurane affected cerebral neuronal activation resulting from noxious and innocuous stimuli. Five male volunteers were subjected to mild electrical shock and tactile stimuli applied to the hand. During low (0.7%) and moderate (1.3%) isoflurane anesthesia the stimuli were repeated and a supramaximal electrical shock was also applied. Tactile stimulation activated bilateral SI and SII, but resulted in no significant activation at low or moderate anesthesia. Electrical shock activated contralateral SI and bilateral SII; low anesthesia completely abolished this response. The supramaximal stimulus activated the caudate nucleus and bilateral thalamus at low anesthesia; these responses were diminished at moderate anesthesia. Isoflurane anesthesia blunts cerebral responses to somatosensory stimuli, and the absence of cortical activation during supramaximal stimulation suggests that noxious-induced movement is generated in lower CNS structures.     

Representations of graphomotor trajectories in the human parietal cortex: evidence for controlled processing and automatic performance

The aim of this study was to identify the cerebral areas activated during kinematic processing of movement trajectories. We measured regional cerebral blood flow (rCBF) during learning, performance and imagery of right-hand writing in eight right-handed volunteers. Compared with viewing the writing space, increases in rCBF were observed in the left motor, premotor and frontomesial cortex, and in the right anterior cerebellum in all movement conditions, and the increases were related to mean tangential writing velocity. No rCBF increases occurred in these areas during imagery. Early learning of new ideomotor trajectories and deliberately exact writing of letters both induced rCBF increases in the cortex lining the right intraparietal sulcus. In contrast, during fast writing of overlearned trajectories and in the later phase of learning new ideograms the rCBF increased bilaterally in the posterior parietal cortex. Imagery of ideograms that had not been practised previously activated the anterior and posterior parietal areas simultaneously. Our results provide evidence suggesting that the kinematic representations of graphomotor trajectories are multiply represented in the human parietal cortex. It is concluded that different parietal subsystems may subserve attentive sensory movement control and whole-field visuospatial processing during automatic performance.     

Lipoma arborescens of the knee: MR characteristics in 13 joints

We describe a technique for mapping out human somatosensory cortex using functional magnetic resonance imaging (fMRI). To produce cortical activation, a pneumatic apparatus presented subjects with a periodic series of air puffs in which a sliding window of five locations moved along the ventral surface of the left arm in a proximal-to-distal or distal-to-proximal direction. This approach, in which the phase-delay of the stimulus can be used to produce somatotopic maps of somatosensory cortex, is based on a method used to generate retinotopic maps of visual cortex. Functional images were acquired using an echoplanar 1.5T scanner and a T2*-weighted spiral acquisition pulse sequence. The periodic series of air puffs created phase-related activation in two cortical regions of the contralateral parietal lobe, the posterior bank of the central sulcus and a more posterior and lateral region.     

Hand preference of the common marmoset (Callithrix jacchus): Problem solving and responses in a novel setting.

Common marmosets (Callithrix jacchus) with a right-hand preference displayed shorter latencies to enter a novel room containing novel structures and objects, touched more objects, and performed more touches and more parallax movements than marmosets with a left-hand preference. These results are consistent with specialization of the right hemisphere (left hand) for fear and negative emotional states and specialization of the left hemisphere (right hand) for approach and positive emotional states. There were no effects of age or sex on any of these behaviors. This relationship between exploration and hand preference did not occur when the marmosets were tested in the home cage with a novel problem, although significant effects of both age and dominance were found in solving the problem. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Cortical activation during fast repetitive finger movements in humans: dipole sources of steady-state movement-related cortical potentials

Fast repetitive finger movements are associated with characteristic EEG patterns described in humans as steady-state movement-related cortical potentials (ssMRCPs). The objective of the present study was to determine the electrical generators of ssMRCPs (movement rate, 2 Hz) by dipole modelling. The generators for the initial ssMRCP phase (peak approximately 60 msec before EMG onset) were located in the central region bilaterally, with largely radial orientation, consistent with activation of the crown of the precentral gyrus. The generator of the next phase (peak approximately 10 msec after EMG onset) was located in the contralateral central region with tangential posterior orientation, consistent with activation of the anterior wall of the central sulcus. The postmovement phase (peak approximately 95 msec after EMG onset) was explained by another source in the contralateral central region with tangential anterior orientation, consistent with activation of the posterior wall of the central sulcus. This pattern probably corresponds to a sequence of activation of the bilateral dorsal premotor cortex, contralateral primary motor, and primary somatosensory cortex that takes place within approximately 200 msec around EMG onset. Steady-state movement-related cortical potentials in combination with dipole modelling provide a novel, noninvasive approach to assessing changes of human cortical premotor, motor, and somatosensory activation in the millisecond range.     

Neuronal activity of somatosensory cortex in a cross-modal (visuo-haptic) memory task

Studies have shown that in the monkey's associative cerebral cortex, cells undergo sustained activation of discharge while the animal retains information for a subsequent action. Recent work has revealed the presence of such "memory cells" in the anterior parietal cortex (Brodmann's areas 3a, 3b, 1, and 2)--the early stage of the cortical somatosensory system. Here we inferred that, in a cross-modal visuo-haptic short-term memory task, somatosensory cells would react to visual stimuli associated with tactile features. Single-unit discharge was recorded from the anterior parietal cortex--including areas of hand representation--of monkeys performing a visuo-haptic delayed matching-to-sample task. Units changed firing frequency during the presentation of a visual cue that the animal had to remember for making a correct tactile choice between two objects at the end of a delay (retention period). Some units showed sustained activation during the delay. In some of them that activation differed depending on the cue. These findings suggest that units in somatosensory cortex react to visual stimuli behaviorally associated with tactile information. Further, the results suggest that some of these neurons are involved in short-term active memory and may, therefore, be part of cross-modal memory networks.     

Magnetic source imaging combined with image-guided frameless stereotaxy: a new method in surgery around the motor strip

OBJECTIVE: In this study, information about the localization of the central sulcus obtained by magnetic source imaging (MSI) was intraoperatively translated to the brain, using frameless image-guided stereotaxy. In the past, the MSI results could be translated to the surgical space only by indirect methods (e.g., the comparison of the MSI results, displayed in surface renderings, with bony landmarks or blood vessels on the exposed brain surface). METHODS: Somatosensory evoked fields were recorded with a MAGNES II biomagnetometer (Biomagnetic Technologies Inc., San Diego, CA). Using the single equivalent current dipole model, the localization of the somatosensory cortex was superimposed on magnetic resonance imaging with a self-developed contour fit program. The magnetic resonance image set containing the magnetoencephalographic dipole was then transferred to a frameless image-guided stereotactic system. Intraoperatively, the gyrus containing the dipole was identified as the postcentral gyrus, using neuronavigation, and the next anterior sulcus was regarded as the central sulcus. With intraoperative cortical recording of somatosensory evoked potentials, this assumption was verified in each case. RESULTS: In all cases, the preoperatively assumed localization of the central sulcus and motor cortex with MSI agreed with the intraoperative identification of the central sulcus using the phase reversal technique. CONCLUSION: The combined use of MSI and a frameless stereotactic system allows a fast orientation of eloquent brain areas during surgery. This may contribute to a safer and more radical surgery in lesions adjacent to the motor cortex.     

Mnemonic neuronal activity in somatosensory cortex

Single-unit activity was recorded from the hand areas of the somatosensory cortex of monkeys trained to perform a haptic delayed matching to sample task with objects of identical dimensions but different surface features. During the memory retention period of the task (delay), many units showed sustained firing frequency change, either excitation or inhibition. In some cases, firing during that period was significantly higher after one sample object than after another. These observations indicate the participation of somatosensory neurons not only in the perception but in the short-term memory of tactile stimuli. Neurons most directly implicated in tactile memory are (i) those with object-selective delay activity, (ii) those with nondifferential delay activity but without activity related to preparation for movement, and (iii) those with delay activity in the haptic-haptic delayed matching task but no such activity in a control visuo-haptic delayed matching task. The results indicate that cells in early stages of cortical somatosensory processing participate in haptic short-term memory.