Biomedical subjects
S Shipp
Publications and source records attributed to S Shipp.
Functional demarcation of a border between areas V6 and V6A in the superior parietal gyrus of the macaque monkey.
We have compared physiological data recorded from three alert macaque monkeys with separate observations of local connectivity, to locate and characterize the functional border between two related but distinct visual areas on the caudal face of the superior parietal gyrus. We refer to these areas as V6 and V6A. The occupy almost the entire extent of the anterior bank of the parieto-occipital sulcus, V6A being the more dorsal. These two areas are strongly interconnected. Anatomically, we have defined the border as the point at which labelled axon terminals first adopt a recognizably 'descending' pattern in their laminar characteristics, after injections of wheatgerm agglutinin-horseradish peroxidase into the dorsal half of the gyrus (in presumptive V6A). A similar principle was used to recognize the same border by the pattern of input from area V5, except that in this case the relevant transition in laminar characteristics is that between an 'intermediate' pattern (in V6) and an 'ascending' pattern (in V6A). V6A was found to be distinct from V6 in a number of its physiological properties. Unlike V6, it contains visually unresponsive cells as well as units with craniotopic receptive fields ('real-position' cells), units tuned to very slow stimulus speeds, units with complex visual selectivities and units with activity related to attention. V6A was also found to have a larger mean receptive field size and scatter than V6. By contrast, response properties related to the basic orientation and direction of moving bar stimuli were indistinguishable between V6 and V6A, as was the influence of gaze direction on cell activity in the two areas. Two-dimensional maps of the recording sites allowed reconstruction of the V6/V6A border. For comparison, the anatomical results were rendered on two-dimensional maps of identical format to those used to summarize the physiological data. After normalizing for relative size, the physiological and connectional estimates of the border between V6 and V6A were found to coincide, at least within the range of individual variation between hemispheres. An architectonic map in the same format was also made from a hemisphere stained for myelin and Nissl substance. Area PO, defined by its general density of myelination was not distinct in this material, but several architectural features were traceable and one of these was also found to approximate the V6/V6A border. The particular criteria that distinguish V6 from V6A differ from a recent description of areas PO and POd in the Cebus monkey; we believe it most likely that PO and POd together may correspond to V6.
Visual processing. The odd couple.
Two distinct pathways in the visual cortex shadow each other from start to finish. Why?
Retinotopic maps in human prestriate visual cortex: the demarcation of areas V2 and V3.
We have used PET (positron emission tomography) to chart the mapping of the retina in human occipital visual cortex and hence to locate the secondary and tertiary visual areas, V2 and V3. A group of four non-selected male volunteers was presented with dynamic stimuli that were aligned with either the vertical or the right horizontal meridians (VM or HM) from 0 degree to 29 degrees eccentricity; the vertical stimuli were restricted to either the inferior or the superior hemifields. PET scans were performed using intravenous infusion of H215O and a Siemens-CTI 953B PET scanner with 3D data acquisition. Subjects received 18 scans, divided equally among the right HM, the superior VM, and the inferior VM. Data were analyzed with SPM software. The group average result confirmed our experimental hypothesis that human occipital visual cortex has retinotopic maps similar to those of the macaque monkey. Thus human areas V2 and V3 can be defined on the basis that the border between them is formed by the HM and that the outer border of V3 is demarcated by a second representation of the VM that runs approximately parallel to the primary representation of the VM at the V1/V2 border. Furthermore, as in many mammals, the extrastriate representation of the HM is "split", such that the superior contralateral quadrant is mapped in lower V2 and V3, occupying the ventral surface of human cortex, and the inferior contralateral quadrant is mapped in upper V2 and V3, which extend over the lateral and medial surfaces of each hemisphere. After stereotaxic normalization, the position of V3 defined by retinal topography was found to correspond to that surmised from our previous PET studies employing moving stimuli.
Segregation and convergence of specialised pathways in macaque monkey visual cortex.
At the level of cortical area V2, the various visual inputs to the cortex have reorganised to form 3 distinct channels. Anatomically these are embodied in the thick and thin dark stripes, and paler interstripes characteristic of cytochrome oxidase architecture. Do the outputs of these compartments remain segregated at higher levels of processing, or are they in turn combined and repackaged? To examine this question we have injected distinct orthograde tracers into the functionally distinct areas V4 and V5 of one hemisphere in 3 macaque monkeys (Macaca fascicularis). V4 is known to receive input from both thin stripes and interstripes of V2, but some parts of V4 receive only interstripe afferents, others receive a relatively greater contribution from the thin stripes. Thus V4 itself is thought to possess subcompartments of at least two distinct types, acting to extend the blob-thin stripe and interblob-interstripe pathways through V1 and V2. The experiments reported here reveal no further divergence between these channels: both types of V4 subcompartment make rather similar patterns of connection with further visual areas and subcortical structures. In contrast to V4, area V5 receives input from the thick stripes of V2. V4 and V5 are weakly interconnected, at best, and there is limited direct convergence in their two sets of ascending connections. For instance, both areas send output to area LIP; but V4 targets the dorsal half of the area, and V5 the ventral half, with some minor overlap. Projections to the superior temporal sulcus are also mainly separate, although we found instances of direct convergence in areas FST and possibly V4t. Segregation is also the rule for subcortical connections to the pulvinar from these two areas. In summary, the segregated outputs of V2 can remain largely distinct through at least two subsequent stages of cortical processing.
The brain activity related to residual motion vision in a patient with bilateral lesions of V5.
We have used the technique of PET to chart the cortical areas activated by visual motion in the brain of a patient with a severe impairment in the ability to recognize the motion of objects (akinetopsia), following bilateral lesions which have so far been presumed to include area V5. High resolution MRI of her brain showed that the zone occupied by area V5 had indeed been destroyed bilaterally. Positron emission tomography activation images, co-registered to the MRIs, showed three principal regions of the cortex activated by motion. These were located (i) bilaterally in the precuneus of superior parietal cortex (area 7 of Brodmann); (ii) bilaterally in the cuneus (a region considered to represent upper V3); (iii) in the left lingual and fusiform gyri (possibly lower V3 and adjacent areas). In contrast to normal subjects, there was no significant activation of area V1 or V2. The stimuli used for scanning were chosen by prior testing of the patient's visual capacities. The control stimulus was a static random distribution of light squares on a dark background. In the moving stimulus these squares moved coherently, the direction of motion changing periodically between the cardinal directions (left, right, up and down). It was ascertained that the patient could correctly identify these directions. We also found (i) that her occasional errors were always in the direction opposite to the motion presented, so that her identification of axis of motion (i.e. vertical or horizontal) was 100% correct; (ii) that when a few static squares were added to the moving the display her identification of direction fell to chance but her identification of the axis of motion remained 100%; (iii) that when a few squares moving opposite and orthogonal to the predominant direction of motion were incorporated, her performance on both direction and axis fell to chance; (iv) that she was unable to identify motion in oblique directions between the horizontal or vertical axes, always guessing one of the cardinal directions. In accounting for her residual vision in terms of cortex, which remains active, we hypothesize; (i) that the bilateral loss of V5 has affected direction sensitive mechanisms at other sites in the cortex which are interconnected with V5 and (ii) that in consequence her performance on our tests reflects the properties of dynamic orientation selective mechanisms that were also differentially activated by the stimuli used during scanning.
The cerebral activity related to the visual perception of forward motion in depth.
We have used the technique of PET to chart the areas of human cerebral cortex specifically responsive to an optical flow stimulus simulating forward motion in depth over a flat horizontal surface. The optical flow display contained about 2000 dots accelerating in radial directions away from the focus of expansion, which subjects fixated at the centre of the display monitor. Dots remained of constant size, but their density decreased from the horizon, lying across the middle of the screen, to the foreground at the lower screen margin; the top half of the display was void. For the control stimulus the dot motions were randomized, removing any sensation of motion in depth and diminishing the impression of a flat terrain. Comparison of the regional cerebral blood flow (rCBF) elicited by the optical flow and control stimuli was thus intended to reveal any area selectively responsive to the radial velocity field that is characteristic of optical flow in its simplest natural form. Six subjects were scanned, and analysed as a group. Four subjects were analysed as individuals, their PET data being co-registered with MRIs of the cerebrum to localize rCBF changes to individual gyri and sulci. There were three main areas of activation associated with optical flow: the dorsal cuneus (area V3) and the latero-posterior precuneus (or superior parietal lobe) in the right hemisphere, and the occipito-temporal ventral surface, in the region of the fusiform gyrus, in both hemispheres. There was no significant activation of V1/V2, nor of V5. These results show that higher stages of motion take place in both the 'dorsal' and 'ventral' visual pathways, as these are commonly conceived, and that both may be fed by area V3. The information potentially derivable from optical flow concerns the direction of heading, and the layout of the visual environment, a form of three-dimensional structure-from-motion. The perceptual division of labour between the various activated areas cannot be directly inferred, though it is a reasonable supposition that the parietal activation reflects the utility of optic flow for guiding self-motion.
Visuotopic organization of the lateral suprasylvian area and of an adjacent area of the ectosylvian gyrus of cat cortex: a physiological and connectional study.
We have explored the visuotopic organization of the territory surrounding the middle suprasylvian sulcus (MSS) of cat cerebral cortex by electrophysiological mapping, and by tracing the topography of its cortical and subcortical connections using wheatgerm-agglutinin horseradish peroxidase (WGA-HRP). Observations from the two approaches were concordant, and confirmed the presence of two separate visual areas in the MSS that approximate, but do not exactly correspond, to the location and internal organization of the posterior medial and posterior lateral lateral suprasylvian (PMLS, PLLS) areas of Palmer et al. (1978). We define as part of the lateral suprasylvian (LS) area the territory on the medial bank and caudal end of the lateral bank of the MSS that receives a topographically organized projection from the region of area 17 representing the lower visual quadrant. This territory is connected with other structures that are themselves striate-recipient (cortical areas 18 and 19, and the lateral division of the lateral posterior (LPl) nucleus), and with a variety of nuclei that receive direct retinal input, such as the C-laminae of the LGd, the medial interlaminar nucleus (MIN), and the superficial layers of the superior colliculus (SC). Its connections with the LPl, LGd, MIN, and SC correspond topographically with the input from area 17. Revised maps of area LS were produced from the physiological and connectional data: its rostral border is formed by a representation of lower visual elevations with the horizontal meridian represented caudally, and its lateral border is formed by the vertical meridian; area LS shares a representation of the center of gaze with the visual area of the lateral bank at its caudal end. The adjacent lateral bank area has larger receptive fields than area LS, and very different connectivity. It receives no input from area 17 and little input from striate-recipient structures, including area LS, but instead is connected to more remote extrastriate visual areas, such as the anterior ectosylvian visual (AEV) area in insular cortex, and to zones of the thalamus in receipt of tectal input (LPm and the lateromedial-suprageniculate nuclear complex). According to both mapping approaches, the lateral bank area contains representations of both the upper and lower visual quadrants but a rather limited degree of visuotopic order. We refer to it as the posterior ectosylvian visual (PEV) area, because it appears to be functionally and connectionally dissociated from area LS, but is possibly a functional antecedent of area AEV.
Organization of reciprocal connections between area 17 and the lateral suprasylvian area of cat visual cortex.
The lateral suprasylvian (LS) area (or Clare-Bishop area) is a region of visual cortex in the cat which has been defined as an isolated projection zone of area 17 (V1 or striate cortex) within the suprasylvian sulcus. We have studied the overall topography and detailed pattern of connection between these two visual areas following injections of WGA-HRP into one or the other. The projection from area 17 to LS is formed largely (approximately 90%) from supragranular layer neurons that are distributed, in the coronal plane, in multiple regularly spaced patches. These patches are especially prominent in regions of area 17 representing central vision along and around the horizontal meridian. In reconstructions of serial coronal sections, and in flatmounts of the same region, the patches are seen to align so that in the plane tangential to the cortical surface they appear as a system of parallel bands whose main axis of elongation is rostro-ventral to caudo-dorsal, or near parallel to the area 17/18 border. The mean periodicity of the bands is about 1.0 mm. The projection from area 17 terminates mainly in layers 4, 3, and 2 of area LS, and also appears patchy in the coronal plane. Reconstruction of the cortical surface view again reveals a system of rostrocaudal bands, but with a mean periodicity of 2 mm. The back projection is less periodically organized, arising predominantly (approximately 80%) from a continuous sheet of infragranular neurons in area LS and terminating mainly in layer 1 of area 17, across the underlying patch and interpatch zones of the supragranular projection cells. However, neurons in layers 2 and upper 3 of area LS, which form the minority origin of the back projection, are mostly located in columnar registration with the patches of area 17 terminals. The bands of supragranular layer neurons projecting to area LS are aligned obliquely to the iso-orientation domains of area 17, indicating a further component to its organization. It is suggested that this may correspond to a segregation of the X and Y channels in area 17, with outputs to area LS selectively arising from the Y pathway, in accordance with previous reports.
The functional logic of cortical connections.
Patterns of anatomical connections in the visual cortex form the structural basis for segregating features of the visual image into separate cortical areas and for communication between these areas at all levels to produce a coherent percept. Such multi-stage integration may be a common strategy throughout the cortex for producing complex behaviour.
Location of chromophoric residues in ribonuclease T1 by solvent perturbation difference spectroscopy.
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Segregation of pathways leading from area V2 to areas V4 and V5 of macaque monkey visual cortex.
V5 and V4 are areas of macaque monkey prestriate visual cortex that are specialized for involvement in different aspects of visual perception, namely motion for V5 (refs 1-4) and colour vision, with other possible functions, for V4 (refs 2, 5-9). Thus, it is unlikely that they should be fed the same information for further processing, yet both receive a strong input from patches of the upper layers of V2 (refs 10, 11), the area immediately adjoining the primary visual cortex, V1. V2, however, seems to comprise functionally distinct subregions, which can be revealed by staining the tissue for the mitochondrial enzyme cytochrome oxidase. Here we report that V4 and V5 are connected with separate cytochrome oxidase-defined subregions of V2, suggesting that cortical pathways dealing with motion and colour perception are segregated in their passage through V2, and reinforcing evidence for functional specialization in the visual cortex.
Area V5 of the human brain: evidence from a combined study using positron emission tomography and magnetic resonance imaging.
In pursuing our work on the organization of human visual cortex, we wanted to specify more accurately the position of the visual motion area (area V5) in relation to the sulcal and gyral pattern of the cerebral cortex. We also wanted to determine the intersubject variation of area V5 in terms of position and extent of blood flow change in it, in response to the same task. We therefore used positron emission tomography (PET) to determine the foci of relative cerebral blood flow increases produced when subjects viewed a moving checkerboard pattern, compared to viewing the same pattern when it was stationary. We coregistered the PET images from each subject with images of the same brain obtained by magnetic resonance imaging, thus relating the position of V5 in all 24 hemispheres examined to the individual gyral configuration of the same brains. This approach also enabled us to examine the extent to which results obtained by pooling the PET data from a small group of individuals (e.g., six), chosen at random, would be representative of a much larger sample in determining the mean location of V5 after transformation into Talairach coordinates. After stereotaxic transformation of each individual brain, we found that the position of area V5 can vary by as much as 27 mm in the left hemisphere and 18 mm in the right for the pixel with the highest significance for blood flow change. There is also an intersubject variability in blood flow change within it in response to the same visual task. V5 nevertheless bears a consistent relationship, within each brain, to the sulcal pattern of the occipital lobe. It is situated ventrolaterally, just posterior to the meeting point of the ascending limb of the inferior temporal sulcus and the lateral occipital sulcus. In position it corresponds almost precisely with Flechsig's Feld 16, one of the areas that he found to be myelinated at birth.