Topographic Maps within Brodmann's Area 5 of macaque monkeys.

Brodmann's area 5 has traditionally included the rostral bank of the intraparietal sulcus (IPS) as well as posterior portions of the postcentral gyrus and medial wall. However, different portions of this large architectonic zone may serve different functions related to reaching and grasping behaviors. The current study used multiunit recording techniques in anesthetized macaque monkeys to survey a large extent of the rostral bank of the IPS so that hundreds of recording sites could be used to determine the functional subdivisions and topographic organization of cortical areas in this region. We identified a lateral area on the rostral IPS that we term area 5L. Area 5L contains neurons with receptive fields on mostly the shoulder, forelimb, and digits, with no apparent representation of other body parts. Thus, there is a large magnification of the forelimb. Receptive fields for neurons in this region often contain multiple joints of the forelimb or multiple digits, which results in imprecise topography or fractures in map organization. Our results provide the first overall topographic map of area 5L obtained in individual macaque monkeys and suggest that this region is distinct from more medial portions of the IPS.

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title: Neurofilament protein defines regional patterns of cortical organization in the macaque monkey visual system: A quantitative immunohistochemical analysis
abstract: Visual function in monkeys is subserved at the cortical level by a large number of areas defined by their specific physiological properties and connectivity patterns, For most of these cortical fields, a precise index of their degree of anatomical specialization has not yet been defined, although many regional patterns have been described using Nissl or myelin stains. In the present study, an attempt has been made to elucidate the regional characteristics, and to varying degrees boundaries, of several visual cortical areas in the macaque monkey using an antibody to neurofilament protein (SM132). This antibody labels a subset of pyramidal neurons with highly specific regional and laminar distribution patterns in the cerebral cortex. Based on the staining patterns and regional quantitative analysis, as many as 28 cortical fields were reliably identified. Each field had a homogeneous distribution of labeled neurons, except area V1, where increases in layer IVB cell and in Meynert cell counts paralleled the increase in the degree of eccentricity in the visual field representation. Within the occipitotemporal pathway, areas V3 and V4 and fields in the inferior temporal cortex were characterized by a distinct population of neurofilament‐rich neurons in layers II–IIIa, whereas areas located in the parietal cortex and part of the occipitoparietal pathway had a consistent population of large labeled neurons in layer Va. The mediotemporal areas MT and MST displayed a distinct population of densely labeled neurons in layer VI. Quantitative analysis of the laminar distribution of the labeled neurons demonstrated that the visual cortical areas could be grouped in four hierarchical levels based on the ratio of neuron counts between infragranular and supragranular layers, with the first (areas V1, V2, V3, and V3A) and third (temporal and parietal regions) levels characterized by low ratios and the second (areas MT, MST, and V4) and fourth (frontal regions) levels characterized by high to very high ratios. Such density trends may correspond to differential representation of corticocortically (and corticosubcortically) projecting neurons at several functional steps in the integration of the visual stimuli. In this context, it is possible that neurofilament protein is crucial for the unique capacity of certain subsets of neurons to perform the highly precise mapping functions of the monkey visual system.

title: Posterior parietal projections to the intraparietal sulcus of the rhesus monkey
abstract: SummaryA cyto- and myeloarchitectonic parcellation of the intraparietal sulcus in the rhesus monkey was correlated with the pattern of afferent connections from the parietal lobe as determined by autoradiographic techniques. Area PEa in the upper bank receives topographically-organized input from the ventral and caudal superior parietal lobule and the medial surface of the parietal lobe. Area POa in the lower bank is the recipient of a projection from the rostral inferior parietal lobule. Area IPd, situated in the depth of the intraparietal sulcus, receives converging input from the caudal superior parietal lobule, medial surface of the parietal lobe, and mid-and caudal inferior parietal lobule. Finally, intrinsic sequences of connections, directed from rostral to caudal and caudal to rostral, exist within both areas PEa and POa, each having a distinctive laminar pattern of origins and terminations.

title: Mapping of architectonic subdivisions in the macaque monkey, with emphasis on parieto‐occipital cortex
abstract: The intraparietal sulcus (IPS) of the macaque monkey contains numerous areas associated with different aspects of cortical function, including motor control as well as visual, somatosensory, vestibular, and possibly auditory processing. This study focuses largely on the architectonic organization of areas within and near the IPS, but also examines remaining portions of the hemisphere with which the IPS is interconnected. We charted the location of up to 72 architectonically distinct areas plus numerous architectonic zones in individuals over a region covering most of the cortical hemisphere. Identified cortical subdivisions (areas plus zones) were represented on computationally generated flat maps in relation to gyral and sulcal geography, thereby facilitating the analysis of consistent as well as variable aspects of the sizes and relative positions of subdivisions across animals. Using myelin and Nissl stains, plus immunohistochemical staining with the SMI‐32 antibody, 17 architectonic subdivisions were identified that are largely or entirely contained in the intraparietal and parieto‐occipital sulci. This includes four newly identified zones: a heavily myelinated lateral occipitoparietal zone, termed LOP; a strongly SMI‐32 immunoreactive zone termed 7t (near the tip of the IPS); plus medial and lateral subdivisions (VIPm and VIPl) of ventral intraparietal area (VIP), which was previously regarded as an anatomically homogeneous area. Within the superior temporal sulcus, we identified a densely myelinated zone termed the dorso‐posterior subdivision of the medial superior temporal area (MSTdp) that bordered middle temporal area (MT). We charted the extent of numerous other architectonically defined subdivisions throughout the cortical hemisphere by using criteria largely based on previous studies, but in some instances involving revised or expanded identification criteria.  2000.

title: Somatosensory properties of neurons in the superior parietal cortex (area 5) of the rhesus monkey.
abstract: The response of individual neurons within Brodmann's area 5 of the parietal lobe to physiological somesthetic stimuli was studied in unanesthetized rhesus monkeys. Most neurons were sensitive to light mechanical stimulation of the skin and deep tissue and/or joint rotation. These neurons were distinguishable from those of the primary somatosensory area by several characteristics: multiple joint interaction, ipsilateral receptive field, bilateral interaction, interaction between forelimb and hindlimb or trunk, and excitatory interaction of joint and skin stimuli. Some were highly selective, in that they responded only to certain critical patterns of stimuli involving both joint and skin. The results were viewed as supporting the hypothesis that area 5 is the site of higher order processing of somesthetic information received from the lemniscal system, and may give rise to the neural code of position and form of body and tactile objects in 3-dimensional space.

title: Posterior parietal association cortex of the monkey: command functions for operations within extrapersonal space.
abstract: Experiments were made on the posterior parietal association cortical areas 5 and in 17 hemispheres of 11 monkeys, 6 M. mulatta and 5 M. arctoides. The electrical signs of the activity of single cortical cells were recorded with microelectrodes in waking animals as they carried out certain behavioral acts in response to a series of sensory cues. The behavioral paradigms were one for detection alone, and a second for detection plus projection of the arm to contact a stationary or moving target placed at arm's length. Of the 125 microelectrode penetrations made, 1,451 neurons were identified in terms of the correlation of their activity with the behavioral acts and their sensitivity or lack of it to sensory stimuli delivered passively; 180 were studied quantitatively. The locations of cortical neurons were identified in serial sections; 94 penetrations and 1,058 neurons were located with certainty. About two-thirds of the neurons of area 5 were activated by passive rotation of the limbs at their joints; of these, 82% were related to single, contralateral joints, 10% to two or more contralateral joints, 6% to ipsilateral, and 2% to joints on both sides of the body. A few of the latter were active during complex bodily postures. A large proportion of area 5 neurons were relatively insensitive to passive joint rotations, as compared with similar neurons of the postcentral gyrus, but were driven to high rates of discharge when the same joint was rotated during an active movement of the animal...

title: Posterior parietal cortex: functional properties of neurons in area 5 during an instructed-delay reaching task within different parts of space
abstract: The activity of single neurons was recorded in area 5 while monkeys made parallel arm movements within different parts of space in an instructed-delay reaching task. In this task: (1) extrinsic variables, such as the direction of movement, were dissociated from intrinsic ones, such as the joint configuration used to perform the movements; and (2) the early neural events related to the presentation of the visual stimulus concerning movement direction were dissociated in time from the later events linked to the execution of movement. Under these experimental conditions, cell activity in area 5 changed so that the population of preferred direction vectors of parietal neurons rotated in space in a way which predicted the rotation of the arm necessary to perform the task. This rotation occurred both during the “instructed-delay time,” when the monkey waited for the “go-signal,” and during the time interval surrounding the onset of movement. This suggests that reaching to visual targets in area 5 is coded by a mechanism combining somatic and visually derived information within a shoulder- or body-centered coordinate system and that instructed-delay time activity in area 5 reflects not only the composition of the direction signal for reaching but also the spatial configuration of the arm.

title: Parietal cortex area 5 and visuomotor behavior.
abstract: For years, area 5 of the parietal cortex was thought to be a somatic sensory structure. This view was challenged by recordings during active movements in alert behaving monkeys, which suggested that area 5 also contained populations of neurons that issued nonspecific, context-dependent "commands" about intended motor responses. Recent studies have revealed that area 5 generates a representation of the spatiotemporal form of arm movements and arm postures, although the parameter space of that representation is still controversial. They also showed that many area 5 cells are recipient to centrally generated signals about the motor relevance of external sensory signals. These new results support the hypothesis that area 5 contributes to the sensorimotor guidance of motor behavior, by contributing to the somatomotor and visuomotor transformations presumed to underlie visually guided behavior. However, area 5 appears to be less directly implicated than the premotor cortex in the process of selecting the nature of the response to instructional signals.

title: Representation of the midline trunk, bilateral arms, and shoulders in the monkey postcentral somatosensory cortex
abstract: A total of 1608 units were isolated from four hemispheres of two animals, and receptive fields (RFs) and submodalities were identified in 1162 units. Deep or skin submodality neurons were dominant in area 3a or area 3b, respectively. The deep/skin ratio increased as the recording site moved from area 3b to the more caudal areas. In areas 3a and 3b, neuronal RFs were almost exclusively on either the arm or trunk. In areas 2 and 5, neurons with RFs on the trunk decreased and those with RFs on the hand or covering more than one body part, etc. increased. We found a total of 107 neurons with bilateral RFs and 56 with ipsilateral RFs, while the rest (n=999) were with contralateral RFs. Bilateral or ipsilateral neurons of skin submodality (n=37) were found in areas 1, 2, and 5. Twenty six (70%) had RFs on the trunk and/or occiput, five on the forelimb, and the rest (n=6) on both the trunk and forelimb (the combined type). Among 33 skin bilateral neurons, 90% (n=30) had RFs across the midline. Bilateral or ipsilateral neurons responding to joint manipulation (n=104) were found in areas 2 and 5. Most of them were activated by manipulation of the shoulder and/or elbow (the proximal type, n=72, 69%). There were 25 neurons of the combined type (both the proximal and distal joints were effective, 24%). Bilateral or ipsilateral neurons of deep-others submodality (n=20) were found in areas 1, 2, and 5. The forelimb type (n=12, 60%) was dominant in this category. The combined-type neurons in both the skin- and joint-manipulation categories were found only or mostly in area 5. These results indicate the presence of hierarchical processing for bilateral as well as contralateral information within the arm/trunk region of the postcentral gyrus.

title: Bilateral receptive field neurons in the hindlimb region of the postcentral somatosensory cortex in awake macaque monkeys
abstract: Single-neuron activities were recorded in the hindlimb region of the primary somatosensory cortex and part of area 5 in awake Japanese monkeys. A total of 1050 units were isolated from five hemispheres of four animals. Receptive fields (RFs) and submodalities were identified for 90% of isolated neurons in areas 3a and 3b. The percentage decreased as the recoding site moved to the more caudal areas. Deep or skin submodality neurons were dominant in area 3a or area 3b, respectively. Deep submodality neurons increased in more caudal areas and were the majority in areas 2 and 5. These observations were consistent with those in the hand and/or digit or arm and/or trunk region. The identified neurons were classified by their RF positions into four types: the foot, leg, foot and leg, or hindlimb and other body parts type. Among 831 identified neurons, 33 neurons had bilateral RFs, 14 had ipsilateral RFs, and the rest (N=784) had contralateral RFs. The relative incidence of neurons with bilateral or ipsilateral RFs among identified neurons was less than 1% in areas 3a, 3b, and 1, and 16% or 25% in areas 2 or 5, respectively. Within areas 2 and 5, the percentage of neurons with bilateral or ipsilateral RFs was significantly smaller in the foot type (5%) than in other RF types (24–57%). RFs of the foot type were on the sole or single toe but never on multiple toes. These observations contrasted with the previous findings that neurons with bilateral RFs were more frequently seen in the hand and/or digit region and that RFs on multiple digit tips were dominant there. The present study thus demonstrated that neurons with bilateral RFs do exist in the hindlimb region. Similarly to the forelimb region, they were found mostly in areas 2 and 5, the caudalmost areas of the postcentral gyrus and hierarchically higher stages in information processing. The relative paucity of neurons with bilateral RFs on the foot, especially those with RFs on multiple toes, may reflect functional differences between the foot and the hand.

title: Neural representation of hand kinematics during prehension in posterior parietal cortex of the macaque monkey.
abstract: Studies of hand manipulation neurons in posterior parietal cortex of monkeys suggest that their spike trains represent objects by the hand postures needed for grasping or by the underlying patterns of muscle activation. To analyze the role of hand kinematics and object properties in a trained prehension task, we correlated the firing rates of neurons in anterior area 5 with hand behaviors as monkeys grasped and lifted knobs of different shapes and locations in the workspace. Trials were divided into four classes depending on the approach trajectory: forward, lateral, and local approaches, and regrasps. The task factors controlled by the animal-how and when he used the hand-appeared to play the principal roles in modulating firing rates of area 5 neurons. In all, 77% of neurons studied (58/75) showed significant effects of approach style on firing rates; 80% of the population responded at higher rates and for longer durations on forward or lateral approaches that included reaching, wrist rotation, and hand preshaping prior to contact, but only 13% distinguished the direction of reach. The higher firing rates in reach trials reflected not only the arm movements needed to direct the hand to the target before contact, but persisted through the contact, grasp, and lift stages. Moreover, the approach style exerted a stronger effect on firing rates than object features, such as shape and location, which were distinguished by half of the population. Forty-three percent of the neurons signaled both the object properties and the hand actions used to acquire them. However, the spread in firing rates evoked by each knob on reach and no-reach trials was greater than distinctions between different objects grasped with the same approach style. Our data provide clear evidence for synergies between reaching and grasping that may facilitate smooth, coordinated actions of the arm and hand.

title: Understanding the parietal lobe syndrome from a neurophysiological and evolutionary perspective
abstract: In human and nonhuman primates parietal cortex is formed by a multiplicity of areas. For those of the superior parietal lobule (SPL) there exists a certain homology between man and macaques. As a consequence, optic ataxia, a disturbed visual control of hand reaching, has similar features in man and monkeys. Establishing such correspondence has proven difficult for the areas of the inferior parietal lobule (IPL). This difficulty depends on many factors. First, no physiological information is available in man on the dynamic properties of cells in the IPL. Second, the number of IPL areas identified in the monkey is paradoxically higher than that so far described in man, although this issue will probably be reconsidered in future years, thanks to comparative imaging studies. Third, the consequences of parietal lesions in monkeys do not always match those observed in humans. This is another paradox if one considers that, in certain cases, the functional properties of neurons in the monkey’s IPL would predict the presence of behavioral skills, such as construction capacity, that however do not seem to emerge in the wild. Therefore, constructional apraxia, which is well characterized in man, has never been described in monkeys and apes. Finally, only certain aspects, i.e. hand directional hypokinesia and gaze apraxia (Balint’s psychic paralysis of gaze), of the multifaceted syndrome hemispatial neglect have been described in monkeys. These similarities, differences and paradoxes, among many others, make the study of the evolution and function of parietal cortex a challenging case.

title: Functional dissociation of saccade and hand reaching control with bilateral lesions of the medial wall of the intraparietal sulcus: Implications for optic ataxia
abstract: The posterior parietal cortex (PPC) is essential for the integration of visuomotor information during visually guided reaching. Studies in macaque monkeys have demonstrated a functional specialisation around the intraparietal sulcus (IPS) with a more medial representation of hand movements ("parietal reach region") and a more lateral representation of saccadic eye movements (lateral intraparietal area, LIP). Here we present evidence for the validity of this concept with respect to the human parietal cortex. We recorded isolated and combined goal-directed eye-hand movements in normal control subjects and in a patient with bilateral parieto-occipital lesions and incomplete Balint's syndrome including severe optic ataxia (misreaching to visual targets). Brain lesions in the patient were caused by acute posterior leucoencephalopathy in association with aortic surgery because of Takayasu's arteritis. MRI scans showed bilateral line-shaped hemorrhagic lesions, restricted to the cortex at the medial banks of the intraparietal sulcus, but leaving its lateral banks largely intact. In the patient visually guided reaching was significantly dysmetric, whereas the metrics of visually guided saccades were within normal limits. Dysmetria was more pronounced for the right visual field, with a gross hypermetria. Variability of the movement improved when a delay of 5 or 10 s was introduced between target presentation and movement execution. Lesion data support the concept of a functional specialisation around the human IPS: The cortex medial to the IPS predominantly controls rapid goal-directed reaching movements, comparable to the parietal reach region in monkeys, whereas saccadic eye movements appear to be controlled rather by the cortex lateral to the IPS.

title: Thalamocortical connections of parietal somatosensory cortical fields in macaque monkeys are highly divergent and convergent.
abstract: We examined the organization and cortical projections of the somatosensory thalamus using multiunit microelectrode recording techniques in anesthetized monkeys combined with neuroanatomical tracings techniques and architectonic analysis. Different portions of the hand representation in area 3b were injected with different anatomical tracers in the same animal, or matched body part representations in parietal areas 3a, 3b, 1, 2, and areas 2 and 5 were injected with different anatomical tracers in the same animal to directly compare their thalamocortical connections. We found that the somatosensory thalamus is composed of several representations of cutaneous and deep receptors of the contralateral body. These nuclei include the ventral posterior nucleus, the ventral posterior superior nucleus, the ventral posterior inferior nucleus, and the ventral lateral nucleus. Each nucleus projects to several different cortical fields, and each cortical field receives projections from multiple thalamic nuclei. In contrast to other sensory systems, each of these somatosensory cortical fields is uniquely innervated by multiple thalamic nuclei. These data indicate that multiple inputs are processed simultaneously within and across several, "hierarchically connected" cortical fields.

title: Quantitative light and electron microscopic analysis of cytochrome oxidase‐rich zones in V II prestriate cortex of the squirrel monkey
abstract: Area 18 of V II of the prestriate cortex of the squirrel monkey was examined at both the light and electron microscopic (EM) levels for cytochrome oxidase (C.O.) activity. At the 17/18 border, the intense C.O. staining of lamina 4 abruptly ended and a new pattern continued for approximately 6 mm into the adjacent prestriate cortex. Here, periodic puffs of high C.O. activity appeared in laminae (lam.) 2 and 3, with the highest activity in lower 3 (3B) extending slightly into upper 4. There was a hint of a columnar pattern in that lam. 4 and especially 5 below the puffs were slightly more reactive than adjacent areas. A thin band of activity could also be seen in upper 5 (5A) and another one between 5 and 6. Tangential sections revealed that the puffs were arranged in alternating wide and narrow rows that radiated orthogonally from the 17/18 border. The puffs in the wider rows tended to be larger (700–1,100 μm in diameter) than those in the narrow rows (400–890 μm in diameter). The center‐to‐center spacing between the puffs was approximately 1,100 μm.

title: Silver staining of myelin by means of physical development.
abstract: For staining myelin with silver a physical development technique has been devised that can render visible the thinnest fibers in various animal species, including fishes and reptiles, even in the early phase of myelination and may be applied to both frozen and embedded materials. Its principle is as follows: Myelin can form and bind colloidal silver particles in a 0.1% ammoniacal silver nitrate solution of pH 7.5. The production of metallic silver by other tissue elements is suppressed by the sections pretreated with a 2:1 mixture of pyridine and acetic anhydride for 30 min. The colloidal silver particles bound in the myelin are enlarged to microscopic dimensions by a special physical developer.

title: Coding of intention in the posterior parietal cortex
abstract: To look at or reach for what we see, spatial information from the visual system must be transformed into a motor plan. The posterior parietal cortex (PPC) is well placed to perform this function, because it lies between visual areas, which encode spatial information1,2, and motor cortical areas. The PPC contains several subdivisions, which are generally conceived as high-order sensory areas3,4. Neurons in area 7a and the lateral intraparietal area fire before and during visually guided saccades. Other neurons in areas 7a and 5 are active before and during visually guided arm movements5–10. These areas are also active during memory tasks in which the animal remembers the location of a target for hundreds of milliseconds before making an eye or arm movement. Such activity could reflect either visual attention11–15 or the intention to make movements16–22. This question is difficult to resolve, because even if the animal maintains fixation while directing attention to a peripheral location, the observed neuronal activity could reflect movements that are planned but not executed22. To address this, we recorded from the PPC while monkeys planned either reaches or saccades to a single remembered location. We now report that, for most neurons, activity before the movement depended on the type of movement being planned. We conclude that PPC contains signals related to what the animal intends to do.

title: Parietal area 5 neuronal activity encodes movement kinematics, not movement dynamics
abstract: SummaryA previous study reported that proximal-arm related area 5 neurons showed continuously-graded changes in activity during unloaded arm movements in different directions (Kalaska et al. 1983), which resembled the responses of primary motor cortex cells in several respects (Georgopoulos et al. 1982). We report here that loading the arm reveals an important difference between cell activity in the two areas. Loads were continuously applied to the arm in different directions. The loads produced large continuously-graded changes in muscle activity but did not alter the handpath or joint angle changes of the arm during the movements. The activity of most area 5 cells was only weakly affected by the loads, and the overall pattern of population activity was virtually unaltered under all load conditions. This indicates that area 5 activity encodes the invariant spatial parameters (kinematics) of the movements. In contrast, many motor cortex cells showed large changes in activity during loading, and so signal the changing forces, torques or muscle activity (movement dynamics; Kalaska et al. 1989).

title: The somatotopic organization of area 2 in macaque monkeys
abstract: Area 2 is a traditional architectonic subdivision of anterior parietal cortex in macaque monkeys, but its overall somatotopic organization and responsiveness to different types of somatic stimuli are poorly understood, and there are uncertainties concerning its rostral and caudal extent. The goals of the present study were to define the rostral and caudal borders of area 2 better, and to describe its overall organization and responsiveness. Somatic receptive fields were defined for hundreds of closely spaced microelectrode recording sites in postcentral parietal cortex of individual macaque monkeys anesthetized with ketamine. Electrophysiological and architectonic evidence suggested that a 3–4‐mm‐wide strip of cortex along the caudal border of area 1 includes all or most of area 2. The most lateral explored portion of area 2 adjoined the representation of the face in area 1. Much of this sector of area 2 was activated by cutaneous stimulation of the face, especially the chin, but more caudal parts of the head also were represented there. Medially, an adjacent sector of area 2 represented the hand. Rostrally, in the cortex within 1.5 mm from the area 1 border, the glabrous surfaces of digits 5 through 1 were represented in a mediolateral cortical sequence, and from tip to base in a rostrocaudal sequence, mirroring the organization in the adjacent portion of area 1. More caudally at this mediolateral level of area 2, digit tips and other phalanges were represented for a second time. The pads of the palm and the dorsal surfaces of the hand were represented laterally and medially within the portion of area 2 devoted to the hand. More medially, the wrist, forearm, and arm were represented in a lateromedial cortical sequence in area 2, roughly matching the mediolateral organization within the bordering area 1. However, immediately caudal to the representation of the occiput, neck, and shoulder in area 1, a rostrocaudal strip of cortex extending across area 2 represented the arm and forearm for a second time in area 2. This cutaneously activated strip of cortex extended into area 5, where the proximal portion of the hand was represented. More medially, next to the trunk representation in area 1, area 2 was devoted to the trunk and limbs. Next to the representations of the ankle, leg, and thigh in area 1, area 2 was activated from similar locations on the hindlimb. Most medially in area 2, the glabrous digits 1–5 of the foot were represented in a lateromedial sequence.

title: Organization of area 3a in macaque monkeys: Contributions to the cortical phenotype
abstract: The detailed organization of somatosensory area 3a was examined in macaque monkeys using multiunit electrophysiological recording techniques. By examining topographic relationships, changes in receptive field size, and the type of stimulus that neurons responded to, functional boundaries of area 3a were determined and related to architectonic boundaries. One striking observation was that the location of area 3a varied with respect to the central sulcus. In one‐half of the cases area 3a was on the rostral bank and fundus of the central sulcus and in the other half of the cases it was on the caudal bank and fundus of the central sulcus. In terms of topographic organization, we found that area 3a contains a complete representation of deep receptors and musculature of the contralateral body, and that the general organization of body part representations mirrors that of the primary somatosensory area, 3b. These results as well as results from studies of area 3a in ours and other laboratories indicate that area 3a is part of a network involved in proprioception, postural control, and the generation of coordinated movements. Further, comparative analysis of area 3a in a variety of species suggests that its construction is based, to a large extent, on the use of a particular body part rather than on innervation density.  2004.

title: Representations of the body surface in postcentral parietal cortex of Macaca fascicularis
abstract: The somatotopic organization of the postcentral parietal cortex of the Old World monkey, Macaca fascicularis, was determined with multi‐unit microelectrode recordings. The results lead to the following conclusions: (1) There are at least two complete and systematic representations of the contralateral body surface in the cortex of the postcentral gyrus. One representation is contained within Area 3b, the other within Area 1. (2) While there are important differences in the organization of the two representations, they are basically mirror‐images of each other. (3) Each representation maintains body‐surface adjacency by cortical adjacency in some mediolateral regions. In other regions, two types of discontinuities can be described: first, in which adjacent body surfaces are represented in separate cortical loci; second, in which adjacent cortical regions represent disparate body‐surface regions. The internal organization of each representation is better described as a composite of somatotopic regions (Merzenich et al., '78) than as a serial array of dermatomal bands, or as a “homunculus.” (4) While architectonic Area 2 responds to stimulation of deep body tissue, at least parts of Area 2 also respond to cutaneous stimulation. The organization of the cutaneous representation of the hand in Area 2 is basically a mirror‐image of the hand representation in Area 1. (5) Area 3a is activated by deep body tissue stimulation, suggesting the possibility of a fourth body representation within the traditional “S‐I” region of somatosensory cortex in macaques. In accord with a previous study in a New World monkey (Merzenich et al., '78), we suggest that the cutaneous representation in Area 3b be considered as SI proper, and that the cutaneous representation in Area 1 be termed the posterior cutaneous field. Furthermore, based on the orientation of the representations of the body surface, as well as other factors, we suggest that the representation in Area 3b is homologous to “SmI” (or “SI”) in non‐primates.

title: Thalamic projections to areas 3a, 3b, and 4 in the sensorimotor cortex of the mature and infant macaque monkey
abstract: Area 3a in the macaque monkey, located in the fundus of the central sulcus, separates motor and somatosensory cortical areas 4 and 3b. The known connections of areas 4 and 3b differ substantially, as does the information which they receive, process, and transfer to other parts of the central nervous system. In this analysis the thalamic projections to each of these three cortical fields were examined and compared by using retrogradely transported fluorescent dyes (Fast Blue, Diamidino Yellow, Rhodamine and Green latex microspheres) as neuron labels. Coincident labeling of projections to 2–3 cortical sites in each monkey allowed the direct comparison of the soma distributions within the thalamic space of the different neuron populations projecting to areas 3a, 3b, and 4, as well as to boundary zones between these cortical fields. The soma distribution ofthalamic neurons projecting to a small circumscribed zone (diameter = 0.5–1.0 mm) strictly within cortical area 3a (in region of hand representation) filled out a “territory” traversing the dorsal half of the cytoarchitectonically defined thalamic nucleus, VPLc (abbreviations as in Olszewski [1952] The Thalamus of the Macaca mulatta. Basel: Karger). This elongate, rather cylindrical, territory extended caudally into the anterior pulvinar nucleus, but not forward into VPLo. The rostrocaudal extent of the thalamic territory defining the soma distribution of neurons projecting to small zones of cortical area 3b was similar, but typically extended into the ventral part of VPLc, filling out a medially concavo‐convex laminar space. Two such territories projecting to adjacent zones of areas 3a and 3b, respectively, overlapped and shared thalamic space, but not thalamic neurons. Contrasting with the 3a and 3b thalamic territories, the soma distribution of thalamic neurons projecting to a circumscribed zone in the nearby motor cortex (area 4) did not penetrate into VPLc, but instead filled out a mediolaterally flattened territory extending from rostral VLo, VLm, VPLo to caudal and dorsal VLc, LP, and Pulo. These territories skirted around VPLc. All three cortical areas (4, 3a, and 3b) also received input from distinctive clusters of cells in the intralaminar Cn.Md. It is inferred that, in combination, the thalamic territories in areas 3a, 3b, and 4 (and also area 1 and 2), which would be coactive during the execution of a manual task, constituted a lamellar space extending from VLo rostrally to Pul.o caudally. How Pul.o neuron populations relate to the more rostral populations within the same thalamic territory projecting to a localized cortical zone remains uncertain. Within the medially located territories the distribution of the neuron population in Pul.o was spatially continuous with the more rostral thalamic cells projecting to the same localized cortex, but in lateral thalamic territories these 2 populations were usually spatially discontinous. In the newborn macaque an orderly change in the territorial projections to localized zones in area 4, 3a, and 3b was also demonstable. However, thalamic nuclear projections were more expansive than in the mature animal. As well as the VPLc input, a third of the thalamic input to area 3a was now from VLo, VPLo, and VLm. Area 4 also had a significant input from VPLc, an input not observed in the mature macaque.

title: A sequential representation of the occiput, arm, forearm and hand across the rostrocaudal dimension of areas 1, 2 and 5 in macaque monkeys
abstract: Receptive fields were determined for recording sites in cortical areas 1, 2 and 5 in macaque monkeys. While a mixture of cutaneous and deep receptors were represented in most regions of area 2, a specific mediolateral level of area 2 was highly responsive to cutaneous input. Anteroposterior rows of recording sites across areas 1, 2 and 5 at this mediolateral location revealed a continuous sequence of cutaneous receptive fields proceeding from the occiput and face in area 1 to the forearm and hand in area 5.

title: Lesions in Posterior Parietal Area 5 in Monkeys Result in Rapid Behavioral and Cortical Plasticity
abstract: We examined the effects of focal lesions of posterior parietal area 5 in macaque monkeys on bimanual behavior performed with and without visual guidance. The animals were trained on two reaching tasks and one tactile texture discrimination task. Task 1 simply involved reaching toward and grasping a reward from one of five well positions. Task 2 required the monkey to use both hands simultaneously to obtain a reward. The tactile texture discrimination task required the monkey to signal the roughness of a passively delivered texture using its jaw. After lesions to area 5, the monkeys showed a decrease in hand use for tasks 1 and 2 and an inability to perform task 2 in specific locations in visual space. These deficits recovered within several days. No deficits were observed in the tactile texture discrimination task or in an analgesic control monkey. Electrophysiological recordings made just before the lesion, immediately after the lesion, and 2 months after the lesion demonstrated that cortical areas just rostral to the lesioned area 5, and areas 1 and 2, were topographically reorganized and that receptive fields for neurons in these fields changed location on the body surface. These cortical map changes are correlative and may, in part, contribute to the rapid behavioral recovery observed. The mechanism for such rapid changes may be the unmasking of existing divergent and convergent thalamocortical connections that are part of the normal cortical circuitry.

title: The relationship of corpus callosum connections to electrical stimulation maps of motor, supplementary motor, and the frontal eye fields in owl monkeys
abstract: Microstimulation and anatomical techniques were combined to reveal the organization and interhemispheric connections of motor cortex in owl monkeys. Movements of body parts were elicited with low levels of electrical stimulation delivered with microelectrodes over a large region of precentral cortex. Movements were produced from three physiologically defined cortical regions. The largest region, the primary motor field, M‐I, occupied a 4‐6mm strip of cortex immediately rostral to area 3a. M‐I represented body movements from tail to mouth in a grossly somatotopic mediolateral cortical sequence. Specific movements were usually represented at more than one location, and often at as many as six or seven separate locations within M‐I. Although movements related to adjoining joints typically were elicited from adjacent cortical sites, movements of nonadjacent joints also were produced by stimulation of adjacent sites. Thus, both sites producing wrist movements and sites producing shoulder movements were found next to sites producing digit movements. Movements of digits of the forepaw were evoked at several locations including a location rostral to or within cortex representing the face. Overall, the somatotopic organization did not completely correspond to previous concepts of M‐I in that it was neither a single topographic representation, nor two serial or mirror symmetric representations, nor a “nesting about joints” representation. Instead, M‐I is more adequately described as a mosaic of regions, each representing movements of a restricted part of the body, with multiple representations of movements that tend to be somatotopically related.

title: Architectionis, somatotopic organization, and ipsilateral cortical connections of the primary motor area (M1) of owl monkeys
abstract: The ipsilateral cortical connections of primary motor cortex (M1) of owl monkeys were revealed by injecting WGA‐HRP and fluorescent tracers into M1 sites identified by intracortical microstimulation. In some of the same animals, the extent and somatotopic organization of M1 was determined by making detailed microstimulation movement maps and relating the results to cortical architectonics. Thus, delineation of M1 was based on a combination of physiological and anatomical characteristics. M1 comprised most, but not all, of the cortex rostral to area 3a where movements were evoked at low levels of current (40 μA or less). Analysis of somatotopic patterns and architectonics placed some of the low‐threshold sites in a ventral premotor field (PMV) and the dorsomedially situated supplementary motor area (SMA). Movements were also reliably elicited from a dorsal premotor area (PMD) at higher currents. M1 was characterized by a somatotopic global organization, representing hindlimb, trunk, forelimb, and face movements in a mediolateral sequence, and a mosaic local organization, with a given movement typically represented at several different sites. Architectionically, M1 was characterized by the absence of a granular layer IV and the presence of very large layer V pyramidal cells. However, M1 was not uniform in structure: pyramidal cells were larger caudally than rostrally, a feature we used to distinguish caudal (M1c) and rostral (M1r) subdivisions of the area. M1 resembles Brodmann's area 4, although the rostral subdivision has probably been considered as part of area 6 by some workers. Tracer injections of M1 revealed somatotopically distributed connections with motor areas PMD, PMV, and SMA, as well as in somatosensory areas 3a, 1, 2, and S2. Weaker connections were with area 3b, posterior parietal cortex, the parietal ventral area (PV), and cingulate cortex. M1r and M1c differed connectionally as well as architectonically, M1c being connected primarily with somatosensory areas, while M1r was strongly connected with both non‐primary motor cortex and somatosensory cortex. These results indicate that M1 interacts directly with at least three non‐primary motor areas and at least six somatosensory areas.

title: Converging evidence from microstimulation, architecture, and connections for multiple motor areas in the frontal and cingulate cortex of prosimian primates
abstract: In the present study, somatotopic organization, architectonic features, and patterns of connections were used to define motor areas in the frontal and cingulate cortex of the prosimian primate Galago garnetti. Sites throughout portions of the motor cortex were electrically stimulated with microelectrodes at the approximate depth of layer V. In some of the same animals, injections in primary motor cortex (M1), and in the spinal cord, revealed patterns of connections with physiologically identified motor areas. Results were related to cortical architecture in brain sections processed for Nissl, myelin, cytochrome oxidase, acetylcholinesterase, or neurofilaments. Evidence was obtained for a number of fields previously identified in simian primates, including M1, dorsal premotor field with caudal (PMDc) and rostral (PMDr) divisions, ventral premotor area (PMV), supplementary motor area (SMA), presupplementary motor area (pre‐SMA), frontal eye field (FEF), and cingulate motor areas, CMAr and CMAc located rostrally and caudally, respectively. In addition, we distinguished area 6Ds of Preuss and Goldman‐Rakic ( 1991a ) between PMV and PMDc, and a more posterior cingulate sensorimotor area (CSMA) with motor connections that may correspond to the supplementary sensory area of monkeys. Areas M1, SMA, PMDc, PMV, CMAr, CMAc, and CSMA projected to the spinal cord, while all of these areas and 6Ds projected to M1. Although area M1 had the lowest stimulation thresholds for evoked movements, movements were also evoked from the other motor areas, as well as from somatosensory areas 3a and 3b. These results indicate that prosimian galagos have a complex of motor areas that closely resembles that in monkeys and suggest that at least 10 motor fields emerged early in primate evolution.  2000.

title: Organization of frontoparietal cortex in the tree shrew (Tupaia belangeri). I. Architecture, microelectrode maps, and corticospinal connections
abstract: Despite extensive investigation of the motor cortex of primates, little is known about the organization of motor cortex in tree shrews, one of their closest living relatives. We investigated the organization of frontoparietal cortex in Belanger's tree shrews (Tupaia belangeri) by using intracortical microstimulation (ICMS), corticospinal tracing, and detailed histological analysis. The results provide evidence for the subdivision of tree shrew frontoparietal cortex into seven distinct areas (five are newly identified), including two motor fields (M1 and M2) and five somatosensory fields (3a, 3b, S2, PV, and SC). The types of movements evoked in M1 and M2 were similar, but M2 required higher currents to elicit movements and had few connections to the cervical spinal cord and distinctive cyto‐ and immunoarchitecture. The borders between M1 and the anterior somatosensory regions (3a and 3b) were identified primarily from histological analysis, because thresholds were similar between these regions, and differences in corticospinal neuron distribution were subtle. The caudal (SC) and lateral (S2 and PV) somatosensory fields were identified based on differences in architecture and distribution of corticospinal neurons. Myelin‐dense modules were identified in lateral cortex, in the expected location of the oral, forelimb, and hindlimb representations of S2, and possibly PV. Evidence for a complex primate‐like array of motor fields is lacking in tree shrews, but their motor cortex shares a number of basic features with that of primates, which are not found in more distantly related species, such as rats.  2006.

title: Distinct nature of directional signals among parietal cortical areas during visual guidance.
abstract: We examined neuronal signals in the monkey medial superior temporal area (MST), the medial intraparietal area (MIP), and the lateral intraparietal area (LIP) during visually guided hand movements. Two animals were trained to use a joystick to guide a spot to a target. Many neurons responded in a direction-selective manner in this guidance task. We tested whether the direction selectivity depended on the direction of the stimulus spot or the direction of the hand movement. First, in some trials, the moving spot disappeared transiently. Second, the mapping between the hand direction and the spot direction was reversed on alternate blocks of trials. Third, we recorded the spot's movement while the animals moved the joystick and then played back that movement while the animals fixated without moving the joystick. Neurons in the three parietal areas conveyed distinct directional information. MST neurons were active and directional only on visible trials in both joystick-movement mode and playback mode and were not affected by the direction of hand movement. MIP neurons were mainly directional with respect to the hand movement, although some MIP neurons were also selective for stimulus direction. MIP neurons were much less active in playback mode. LIP neurons were active and directional in both joystick-movement mode and playback mode. Directional signals in LIP were unrelated to planning saccades. The selectivity of LIP neurons also became evident hundreds of milliseconds before the start of movement. Since the direction of movement was consistent throughout a block of trials, these signals could provide a prediction of the upcoming direction of motion. We tested this by alternating blocks of trials in which the direction was consistent or randomized. The direction selectivity developed earlier on trials in which the upcoming direction could be predicted. These results suggest that LIP neurons combine "bottom-up" visual motion signals with extraretinal, predictive signals about stimulus motion.

title: Discrimination between active and passive head movements by macaque ventral and medial intraparietal cortex neurons
abstract: An important prerequisite for effective motor action is the discrimination between active and passive body movements. Passive movements often require immediate reflexes, whereas active movements may demand suppression of the latter. The vestibular system maintains correct body and head posture in space through reflexes. Since vestibular inputs have been reported to be largely suppressed in the vestibular nuclei during active head movements, we investigated whether head movement‐related signals in the primate parietal cortex, a brain region involved in self‐motion perception, could support both reflex functions and self‐movement behaviour. We employed a paradigm that made available direct comparison of neuronal discharge under active and passive movement conditions. In this study, we demonstrate that a population of intraparietal (VIP (ventral) and MIP (medial)) cortex neurons change their preferred directions during horizontal head rotations depending on whether animals have performed active movements, or if they were moved passively. In other neurons no such change occurred. A combination of these signals would provide differential information about the active or passive nature of an ongoing movement. Moreover, some neurons' responses clearly anticipated the upcoming active head movement, providing a possible basis for vestibular‐related reflex suppression. Intraparietal vestibular neurons thus distinguish between active and passive head movements, and their responses differ substantially from those reported in brainstem vestibular neurons, regarding strength, timing, and direction selectivity. We suggest that the contextual firing characteristics of these neurons have far‐reaching implications for the suppression of reflex movements during active movement, and for the representation of space during self‐movement.

title: Anterior Regions of Monkey Parietal Cortex Process Visual 3D Shape
abstract: The intraparietal cortex is involved in the control of visually guided actions, like reach-to-grasp movements, which require extracting the 3D shape and position of objects from 2D retinal images. Using fMRI in behaving monkeys, we investigated the role of the intraparietal cortex in processing stereoscopic information for recovering the depth structure and the position in depth of objects. We found that while several areas (CIP, LIP, and AIP on the lateral bank; PIP and MIP on the medial bank) are activated by stereoscopic stimuli, AIP and an adjoining portion of LIP are sensitive only to depth structure. Furthermore, only these two regions are sensitive to both the depth structure and the 2D shape of small objects. These results indicate that extracting 3D spatial information from stereo involves several intraparietal areas, among which AIP and anterior LIP are more specifically engaged in extracting the 3D shape of objects.

title: REVIEW: The functional organization of the intraparietal sulcus in humans and monkeys
abstract: In macaque monkeys, the posterior parietal cortex (PPC) is concerned with the integration of multimodal information for constructing a spatial representation of the external world (in relation to the macaque's body or parts thereof), and planning and executing object‐centred movements. The areas within the intraparietal sulcus (IPS), in particular, serve as interfaces between the perceptive and motor systems for controlling arm and eye movements in space. We review here the latest evidence for the existence of the IPS areas AIP (anterior intraparietal area), VIP (ventral intraparietal area), MIP (medial intraparietal area), LIP (lateral intraparietal area) and CIP (caudal intraparietal area) in macaques, and discuss putative human equivalents as assessed with functional magnetic resonance imaging. The data suggest that anterior parts of the IPS comprising areas AIP and VIP are relatively well preserved across species. By contrast, posterior areas such as area LIP and CIP have been found more medially in humans, possibly reflecting differences in the evolution of the dorsal visual stream and the inferior parietal lobule. Despite interspecies differences in the precise functional anatomy of the IPS areas, the functional relevance of this sulcus for visuomotor tasks comprising target selections for arm and eye movements, object manipulation and visuospatial attention is similar in humans and macaques, as is also suggested by studies of neurological deficits (apraxia, neglect, Bálint's syndrome) resulting from lesions to this region.

title: The genitals and gluteal skin are represented lateral to the foot in anterior parietal somatosensory cortex of macaques
abstract: Detailed electrophysiological maps of the representations of trunk and adjacent body parts in area 3b and area 1 of somatosensory cortex were obtained in three macaque monkeys ( Macaca mulatta and Macaca radiata ) of either sex. A total of 211 microelectrode penetrations 250-300 w m apart resulted in 1,190 recording sites. During penetrations deep into the posterior bank of the central sulcus, recordings were made every 300 w m to depths of 6-7 mm until sites unresponsive to somatic stimuli were reached. Cortex was later cut parasagittally and sections were stained for cytochrome oxidase (CO) or Nissl substance. Contrary to expectations from earlier reports, the genitalia were represented lateral to the representations of the foot in cortex along the area 3b/1 border. The gluteal skin including the gluteal pads and the base of the tail were also represented in this section of cortex. Only a small region of cortex was devoted to the genitalia, and neurons in this cortex had receptive fields that were large and typically included skin of the inner thigh and belly. The lower, middle and upper trunk were represented more laterally, followed by the neck, upper head and arm. The receptive fields on the trunk were roughly the same size as those for the middle and lower trunk and slightly smaller on the upper trunk.

title: The Segregation of Function in the Nervous System
abstract: Publisher Summary It is clear, from the advent of more powerful electrophysiological and anatomical methods revealing sensory representations and patterns of connections in the brain, that advanced mammals—such as cats and monkeys—have many sensory representations within separate areas in cortex and nuclei in the thalamus. This chapter discusses the need of the various subdivisions of the sensory system. The basic subdivision of sensory systems is the area or nucleus. Areas and nuclei—as equivalent structures—contain systematic sensory representations, are architectonically distinct, interconnect with a number of other structures in a unique pattern, are distinguished by the response characteristics of the population of neurons, and uniquely contribute to the sensory-perceptual processing of the system. Species differ in number of areas and nuclei, and increases in numbers have occurred independently in many lines of evolution. Advanced mammals have many sensory representations within each major sensory system. These representations are complexly interconnected, and most must participate in the processing of a sensory event within the system. The multitude of subcortical efferent connections of each area suggests that most cortical representations influence behavior, memory, and emotion in a rather direct manner.

title: In Search of a Unifying Theory of Complex Brain Evolution
abstract: The neocortex is the part of the brain that is involved in perception, cognition, and volitional motor control. In mammals it is a highly dynamic structure that has been dramatically altered in different lineages, and these alterations account for the remarkable variations in behavior that species exhibit. When we consider how this structure changes and becomes more complex in some mammals such as humans, we must also consider how the alterations that occur at macro levels of organization, such as the level of the individual and social system, as well as micro levels of organization, such as the level of neurons, synapses and molecules, impact the neocortex. It is also important to consider the constraints imposed on the evolution of the neocortex. Observations of highly conserved features of cortical organization that all mammals share, as well as the convergent evolution of similar features of organization, indicate that the constraints imposed on the neocortex are pervasive and restrict the avenues along which evolution can proceed. Although both genes and the laws of physics place formidable constraints on the evolution of all animals, humans have evolved a number of mechanisms that allow them to loosen these constraints and often alter the course of their own evolution. While this cortical plasticity is a defining feature of mammalian neocortex, it appears to be exaggerated in humans and could be considered a unique derivation of our species.

title: Mechanisms of self-motion perception.
abstract: Guiding effective movement through the environment is one of the visual system's most important functions. The pattern of motion that we see allows us to estimate our heading accurately in a variety of environments, despite the added difficulty imposed by our own eye and head movements. The cortical substrates for heading perception include the medial superior temporal area (MST) and the ventral intraparietal area (VIP). This review discusses recent work on these two areas in the context of behavioral observations that establish the important problems the visual system must solve. Signals relevant to self motion are both more widespread than heretofore recognized and also more complex because they are multiplexed with other sensory signals, such as vestibular, auditory, and tactile information. The review presents recent work as a background to highlight important problems that remain unsolved.

title: Attention, intention, and priority in the parietal lobe.
abstract: For many years there has been a debate about the role of the parietal lobe in the generation of behavior. Does it generate movement plans (intention) or choose objects in the environment for further processing? To answer this, we focus on the lateral intraparietal area (LIP), an area that has been shown to play independent roles in target selection for saccades and the generation of visual attention. Based on results from a variety of tasks, we propose that LIP acts as a priority map in which objects are represented by activity proportional to their behavioral priority. We present evidence to show that the priority map combines bottom-up inputs like a rapid visual response with an array of top-down signals like a saccade plan. The spatial location representing the peak of the map is used by the oculomotor system to target saccades and by the visual system to guide visual attention.

title: Receptive field integration and submodality convergence in the hand area of the post‐central gyrus of the alert monkey.
abstract: 1. An exploration of the occurrence of different functional cell types was made in the three cytoarchitectural subdivisions (areas 3, 1 and 2) of the hand area of the post‐central gyrus of the monkey. The functional properties of 632 cells were studied using the transdural micro‐electrode recording method. 2. Over half of the neurones studied (57%) belonged to the class of simple skin neurones that were related either to rapidly adapting (272 neurones) or slowly adapting (seventeen neurones) cutaneous receptors or to both (seventy‐one neurones). The simple skin neurones were particularly common in the anterior part of S I where they constituted 60% of the cells. More complicated cutaneous neurones made up 10% of the total sample. They were more common in the posterior part of the gyrus. 3. Altogether ninety‐two neurones (15%) were related to subcutaneous or deeper receptors. Another seventy‐one neurones (11%) exhibited convergence of skin input and input from deep receptors. A smaller group of forty‐seven undamaged neurones (7%) were unrelated to stimuli of the types described above. 4. In tangential electrode penetrations made along the anterior and posterior banks of the gyrus, functional columns were found to be 500 micrometers wide on the average; this width is comparable with that of ocular dominance columns and visual orientation hypercolumns. 5. Correlation of the functional types of cells with cytoarchitecture showed that the complexity of the functional properties of the neurones increased posteriorly. The receptive field size also increased toward posterior. The changes that take place in the functional properties of cells when moving across different cytoarchitectural areas suggests intracortical information processing which leads to handling of larger body regions and more complex combinations of information in the cellular elements of the posterior part of the post‐central gyrus.

title: Processing of tactile and kinesthetic signals from bilateral sides of the body in the postcentral gyrus of awake monkeys
abstract: Earlier studies pointed out that in the primary somatosensory cortex (SI) the receptive fields (RF) of bilateral neurons were related exclusively to the body midline. We recently found a substantial number of neurons with bilateral RFs on hand digits, shoulders/arms or legs/feet in the caudalmost part (areas 2 and 5) of the postcentral gyrus in awake macaque monkeys. The RFs of these neurons were generally of the most complex types found in this region of the cortex, and thus they were considered to be at the highest level along the hierarchical chain of information processing. We conclude that there are two types of bilateral RFs in the postcentral gyrus, one representing the midline structures such as the intraoral cavity, chin or trunk and the other related to limb structures such as fingers, hands, arms, shoulders, legs and girdles. Functional significance of the bilateral activity could be understood in behavioral context as it is seen more extensively in the body parts where bilateral coordination is essential.

title: Parallel Evolution of Cortical Areas Involved in Skilled Hand Use
abstract: Dexterous hands, used to manipulate food, tools, and other objects, are one of the hallmarks of primate evolution. However, the neural substrate of fine manual control necessary for these behaviors remains unclear. Here, we describe the functional organization of parietal cortical areas 2 and 5 in the cebus monkey. Whereas other New World monkeys can be quite dexterous, and possess a poorly developed area 5, cebus monkeys are the only New World primate known to use a precision grip, and thus have an extended repertoire of manual behaviors. Unlike other New World Monkeys, but much like the macaque monkey, cebus monkeys possess a proprioceptive cortical area 2 and a well developed area 5, which is associated with motor planning and the generation of internal body coordinates necessary for visually guided reaching, grasping, and manipulation. The similarity of these fields in cebus monkeys and distantly related macaque monkeys with similar manual abilities indicates that the range of cortical organizations that can emerge in primates is constrained, and those that emerge are the result of highly conserved developmental mechanisms that shape the boundaries and topographic organizations of cortical areas.

title: Somatosensory System: Organizational Hierarchy from Single Units in Monkey Area 5
abstract: The receptive fields of single cells in area 5 of monkey parietal cortex were studied by extracellular recording. Cells were driven primarily by gentle manipulation of multiple joints residing on one or more limbs. Both excitatory and inhibitory convergence were demonstrated. It is postulated that the multijoint receptive fields of area 5 are the result of convergence from single-joint cells of the primary receiving area. An analogy is drawn between the modification of information in the visual and somatosensory systems.

title: Change in motor plan, without a change in the spatial locus of attention, modulates activity in posterior parietal cortex.
abstract: The lateral intraparietal area (LIP) of macaque monkey, and a parietal reach region (PRR) medial and posterior to LIP, code the intention to make visually guided eye and arm movements, respectively. We studied the effect of changing the motor plan, without changing the locus of attention, on single neurons in these two areas. A central target was fixated while one or two sequential flashes occurred in the periphery. The first appeared either within the response field of the neuron being recorded or else on the opposite side of the fixation point. Animals planned a saccade (red flash) or reach (green flash) to the flash location. In some trials, a second flash 750 ms later could change the motor plan but never shifted attention: second flashes always occurred at the same location as the preceding first flash. Responses in LIP were larger when a saccade was instructed (n = 20 cells), whereas responses in PRR were larger when a reach was instructed (n = 17). This motor preference was observed for both first flashes and second flashes. In addition, the response to a second flash depended on whether it affirmed or countermanded the first flash; second flash responses were diminished only in the former case. Control experiments indicated that this differential effect was not due to stimulus novelty. These findings support a role for posterior parietal cortex in coding specific motor intention and are consistent with a possible role in the nonspatial shifting of motor intention.

title: Saccade-related activity in the parietal reach region.
abstract: In previous experiments, we showed that cells in the parietal reach region (PRR) in monkey posterior parietal cortex code intended reaching movements in an eye-centered frame of reference. These cells are more active when an arm compared with an eye movement is being planned. Despite this clear preference for arm movements, we now report that PRR neurons also fire around the time of a saccade. Of 206 cells tested, 29% had perisaccadic activity in a delayed-saccade task. Two findings indicate that saccade-related activity does not reflect saccade planning or execution. First, activity is often peri- or postsaccadic but seldom presaccadic. Second, cells with saccade-related activity were no more likely to show strong saccadic delay period activity than cells without saccade-related activity. These findings indicate that PRR cells do not take part in saccade planning. Instead, the saccade-related activity in PRR may reflect cross-coupling between reach and saccade pathways that may be used to facilitate eye-hand coordination. Alternatively, saccade-related activity may reflect eye position information that could be used to maintain an eye-centered representation of intended reach targets across eye movements.

title: A common reference frame for movement plans in the posterior parietal cortex
abstract: Orchestrating a movement towards a sensory target requires many computational processes, including a transformation between reference frames. This transformation is important because the reference frames in which sensory stimuli are encoded often differ from those of motor effectors. The posterior parietal cortex has an important role in these transformations. Recent work indicates that a significant proportion of parietal neurons in two cortical areas transforms the sensory signals that are used to guide movements into a common reference frame. This common reference frame is an eye-centred representation that is modulated by eye-, head-, body- or limb-position signals. A common reference frame might facilitate communication between different areas that are involved in coordinating the movements of different effectors. It might also be an efficient way to represent the locations of different sensory targets in the world.

title: The organization and connections of anterior and posterior parietal cortex in titi monkeys: do New World monkeys have an area 2?
abstract: We used multiunit electrophysiological recording techniques to examine the topographic organization of somatosensory area 3b and cortex posterior to area 3b, including area 1 and the presumptive area 5, in the New World titi monkey, Callicebus moloch. We also examined the ipsilateral and contralateral connections of these fields, as well as those in a region of cortex that appeared to be similar to both area 7b and the anterior intraparietal area (7b/AIP) described in macaque monkeys. All data were combined with architectonic analysis to generate comprehensive reconstructions. These studies led to several observations. First, area 1 in titi monkeys is not as precisely organized in terms of topographic order and receptive field size as is area 1 in macaque monkeys and a few New World monkeys. Second, cortex caudal to area 1 in titi monkeys is dominated by the representation of the hand and forelimb, and contains neurons that are often responsive to visual stimulation as well as somatic stimulation. This organization is more like area 5 described in macaque monkeys than like area 2. Third, ipsilateral and contralateral cortical connections become more broadly distributed away from area 3b towards the posterior parietal cortex. Specifically, area 3b has a relatively restricted pattern of connectivity with adjacent somatosensory fields 3a, 1, S2 and PV; area 1 has more broadly distributed connections than area 3b; and the presumptive areas 5 and 7b/AIP have highly diverse connections, including connections with motor and premotor cortex, extrastriate visual areas, auditory areas and somatosensory areas of the lateral sulcus. Fourth, the hand representation of the presumptive area 5 has dense callosal connections. Our results, together with previous studies in other primates, suggest that anterior parietal cortex has expanded in some primate lineages, perhaps in relation to manual abilities, and that the region of cortex we term area 5 is involved in integrating somatic inputs with the motor system and across hemispheres. Such connections could form the substrate for intentional reaching, grasping and intermanual transfer of information necessary for bilateral coordination of the hands.