Biomedical subjects
A Cowey
Publications and source records attributed to A Cowey.
The role of the 'face-cell' area in the discrimination and recognition of faces by monkeys.
Cortical neurons that are selectively sensitive to faces, parts of faces and particular facial expressions are concentrated in the banks and floor of the superior temporal sulcus in macaque monkeys. Their existence has prompted suggestions that it is damage to such a region in the human brain that leads to prosopagnosia: the inability to recognize faces or to discriminate between faces. This was tested by removing the face-cell area in a group of monkeys. The animals learned to discriminate between pictures of faces or inanimate objects, to select the odd face from a group, to inspect a face then select the matching face from a pair of faces after a variable delay, to discriminate between novel and familiar faces, and to identify specific faces. Removing the face-cell area produced no or little impairment which in the latter case was not specific for faces. In contrast, several prosopagnosic patients were impaired at several of these tasks. The animals were less able than before to discern the angle of regard in pictures of faces, suggesting that this area of the brain may be concerned with the perception of facial expression and bearing, which are important social signals in primates.
Defensive responses to looming visual stimuli in monkeys with unilateral striate cortex ablation.
A number of residual visual functions including detection, localization and discrimination of visual stimuli, have been demonstrated in the "blind" fields of monkeys and human patients following damage to striate cortex. We report here that avoidance movements of the head can also be elicited from monkeys with unilateral striate cortex ablations when a "looming" stimulus is presented within the hemianopic and presumably "blind" field. The possible role of the retinofugal projection to superior colliculus in the mediation of these defensive head movements is discussed.
Wavelength discrimination in blindsight.
In the circumscribed, long-standing, clinically absolute visual field defects of three patients with vascular lesions that involved the optic radiation and visual cortex, forced-choice discrimination between coloured stimuli was tested. Paired stimuli were matched for luminous efficiency on the basis of previous measurements of increment-threshold spectral sensitivity made in the same patients and at the same retinal positions. To different extents all patients could discriminate between narrowband wavelength stimuli. The results imply that despite the effects of retrograde degeneration on thalamic and retinal colour-processing channels, neurons which process wavelength information are still functional, although the information they transmit is not consciously perceived.
Patterns of inter- and intralaminar GABAergic connections distinguish striate (V1) and extrastriate (V2, V4) visual cortices and their functionally specialized subdivisions in the rhesus monkey.
Local GABAergic connections are undoubtedly important for the operation of cerebral cortex, including the tuning of receptive field properties of visual cortical neurons. In order to begin to correlate specific configurations of GABAergic networks with particular receptive field properties, we examined the arrangement of GABAergic neurons projecting to foci in compartments of known functional specialization in striate (area V1) and extrastriate (areas V2, V4) cortices of rhesus monkeys. GABAergic cells were detected autoradiographically following microinjections into supragranular, granular, or infragranular layers of 5, 10, or 50 nl of 3H-nipecotic acid, which selectively exploits the GABA reuptake mechanism. These injections produced complex inter- and intralaminar distributions of retrograde perikaryal labeling that was selective for GABA-immunopositive neurons and glia. The pattern of retrograde labeling depended on both the laminar and cytoarchitectonic location of injection sites. In all cases, a high density of labeled neurons was present in the immediate vicinity of injection sites, with the density of labeled neurons decreasing for the most part uniformly with horizontal distance. Injections in supragranular layers produced relatively widespread labeling (up to 1.5-1.7 mm from the center of injections) in upper layers, whereas in granular and infragranular layers, labeling was confined to a radius of 0.25-0.5 mm. Conversely, injections in infragranular layers produced labeling that was widest (up to 1 mm) in lower layers, but more laterally restricted in supragranular layers. Injections in granular layers, on the other hand, produced an even distribution of labeling, 0.6-1.0 mm in diameter, throughout all layers. Comparably placed injections in V1, V2, and V4 resulted in patterns of labeling that were distinguished by features including stepwise increases in the lateral extent of labeling from striate to extrastriate areas, and the circular versus markedly elongated intralaminar distribution of labeled neurons in V1 and V4 versus V2. Further, for superficial injections, labeling was present in all layers in V1 and V2, but did not extent below the top layer V in area V4. These findings offer clear examples of organizational differences in the intrinsic inhibitory connections of visual cortices. The results also demonstrate that the number of GABAergic neurons projecting to any spot in cortex decreases systematically with horizontal distance from the spot, and that radiolabeled cells do not coalesce to form slabs, columns, or clusters. This relatively even distribution of retrogradely labeled cells in the tangential plane is consistent with recent computer simulations (Worgotter and Koch, 1991) that suggest that inhibitory neurons broadly tuned as a population can produce the specific response properties of cortical neurons.(ABSTRACT TRUNCATED AT 400 WORDS)
Cortical area V4 and its role in the perception of color.
The color and lightness vision of three monkeys with bilateral removal of cortical area V4 and three unoperated controls were tested by measuring their ability to discriminate between two rows of colored or gray stimuli. In one row, the stimuli were ordered in terms of either chromaticity or luminance, whereas in the other row they were disordered. Their ability to select the odd-one-out in an array of colors or grays and to select the colored patch from an array of achromatic grays was also assessed. Unlike an achromatopsic patient tested previously in an identical fashion, monkeys with V4 lesions performed indistinguishably from controls in the oddity test. The animals lacking V4 were slightly impaired at discriminating between ordered and disordered arrays of colors or grays, but the color impairment was no more severe than the impairment with grays. These deficits were readily accounted for in terms of the conspicuous deficits in pattern discrimination apparent in a nine-choice pattern oddity task. The results do not support the view that cortical area V4 in the monkey is the homolog of the cortical "color center" in humans, located in the lingual and fusiform gyri and damage to which leads to the clinical syndrome of cerebral achromatopsia, unless it is the additional damage to underlying white matter that leads to the severe color disorder in patients.
Visual perception. Grasping the essentials.
Explore the source record for details and available documents.
Direct and indirect retinal input into degenerated dorsal lateral geniculate nucleus after striate cortical removal in monkey: implications for residual vision.
We removed the striate cortex of one cerebral hemisphere in a macaque monkey, causing almost total retrograde degeneration of the corresponding dorsal lateral geniculate nucleus (dLGN) and extensive transneuronal degeneration of ganglion cells in the corresponding hemi-retina of each eye. The rare surviving geniculate projection neurons were retrogradely labelled by horseradish peroxidase (HRP) from extra-striate cortex and retinogeniculate terminals were labelled by an intraocular injection of HRP. Retinal terminals in the degenerated dLGN made synaptic contact exclusively with the dendrites of interneurons immunopositive for gamma-aminobutyric acid (GABA) in both parvocellular and magnocellular regions of dLGN. As well as being post-synaptic to retinal terminals these vesicle-containing dendrites were pre- and postsynaptic to other similar dendrites, and presynaptic to relay cells. Surviving labelled projection neurons received retinal input indirectly, via both the GABA-immunopositive interneurons and GABA-immunonegative terminals characteristic of those from the superior colliculus. In the degenerated, as opposed to the normal dLGN, about 20% of retinal terminals were GABA-immunopositive and GABA-immunoreactivity was prominently elevated in the ganglion and amacrine cell layers of the degenerated half of the retina. The optic nerve also contained numerous GABA-immunopositive axons but very few such axons were found in a normal optic nerve processed in identical manner. The surviving pathways from the retina must underlie the visual abilities that survive striate cortical removal in monkeys and human patients and may involve the degenerated dLGN as well as the mid-brain.
The role of the corpus callosum and extra striate visual areas in stereoacuity in macaque monkeys.
Stereoacuity was measured in six normal male rhesus monkeys by requiring them to indicate the closer of two vertical line targets. Their stereoacuity ranged from 13-23 arc sec and was as good as that of human observers tested in the same way. Sectioning the splenium of the corpus callosum, either before or after removing the foveal representation in visual area 2, had no effect on stereoacuity thresholds, indicating that the splenium is not essential for the detection of small retinal disparities at the visual mid-line. In contrast, removal of the foveal representation in V2 permanently and markedly elevated stereoacuity thresholds, almost to the level observed in two monkeys after removal of the representation in striate cortex of the central 5 degrees of the retina. Posterior infero-temporal ablation also impaired stereoacuity, though less strikingly. In these two monkeys, the effect of subsequent damage to the rostral superior colliculi was examined. One animal was unimpaired. In the other, in which the pre-tectum was damaged, the stereoacuity threshold was substantially raised, most plausibly as a consequence of imperfect binocular fixation.
The neurobiology of blindsight.
Some patients can respond to visual stimuli presented within their clinically absolute visual field defects that have been caused by partial destruction of striate cortex. This puzzling phenomenon of looking, pointing, detecting and discriminating without seeing has been called blindsight, and has fascinated philosophers and neuroscientists alike as a spotlight on the nature of unconscious or covert awareness, and the means it provides of studying the visual information carried by pathways other than the major route through the striate cortex.
Increment-threshold spectral sensitivity in blindsight. Evidence for colour opponency.
In the circumscribed visual field defects of 3 patients, increment-threshold spectral sensitivity was measured with a guessing paradigm. Nine 116', 200 ms narrowband stimuli with maximum transmission between 450 and 660 nm were presented on a white background of photopic or scotopic luminance. Sensitivity measured in the blind field was compared with that at matched positions in the patients' normal hemifield, and with that at corresponding positions in 2 control subjects. Results show that spectral sensitivity in the blind field, albeit reduced by up to 1 log unit, shows normal dependence on adaptation level, reflecting rod activity under scotopic, and cone activity under photopic conditions. Characteristic discontinuities in the spectral sensitivity curve seen under light adaptation are evidence for colour-opponent processes, presumably involving primate beta retinal ganglion cells.
Effect of exposure to high pressure on subsequent spatial learning and memory in rats.
The effects of high helium pressure on the subsequent acquisition of spatial memory were studied in male rats. Thirty-two rats were exposed to 65 ATA helium-oxygen pressure for 4.2 days, decompressed (total time in chamber 5 days), and then tested in an eight-arm radial maze. Thirty-two control rats were exposed in the chamber to 1 ATA air. Each rat had 20 sessions in the maze (2 sessions/day for 10 days), and the number of correct (visiting an arm not previously visited to obtain the reward pellet) and incorrect choices (visiting a previously visited arm) were recorded. Statistical analysis showed that the rats exposed to 65 ATA performed significantly better than 1-ATA controls during the first 8 of 20 sessions. This effect was most pronounced in sessions 5-8. Results for sessions 9-20 showed that the pressure-treated rats still made more correct choices but to an extent that did not always reach statistical significance. Possible explanations include the pressure-treated rats performing better because of hunger after a lower food consumption at pressure. Alternatively, pressure itself may enhance proposed mechanisms of spatial memory such as long-term potentiation.
Fibre organization of the monkey's optic tract: I. Segregation of functionally distinct optic axons.
The fibre organization of the monkey's optic tract was examined by implanting pellets of horseradish peroxidase into different locations within the tract, or into the superior colliculus and pretectum. Retinae were examined for the distribution, size, and morphological types of retrogradely labelled ganglion cells; optic tracts were examined for the distribution of anterogradely and retrogradely labelled axonal profiles; and lateral geniculate nuclei were examined for the distribution of anterogradely labelled processes within distinct geniculate laminae. Localized implants in the optic tract produced retrograde labelling of ganglion cells across wide regions of the retinal surface. The maximum density of labelled cells was always substantially less than the total ganglion cell density known to be present at those retinal loci. Distinct retinal ganglion cell types were labelled from differing regions within the optic tract: implants into the deep (dorsal) portion of the tract, far removed from the outer, pial, surface, retrogradely labelled predominantly P beta retinal ganglion cells, whereas implants into the superficial (ventral), subpial, part of the tract retrogradely labelled primarily the other retinal ganglion cell types, i.e., the P alpha, P gamma, and P epsilon cells. Within any given class of axon, there is a mapping of the centroperipheral retinal axis across the deep-to-superficial dimension of the tract, but this retinotopy is extremely coarse. Anterograde labelling of axonal terminations within the lateral geniculate nucleus showed a corresponding specificity for distinct geniculate laminae, the deep implants labelling the parvocellular laminae, superficial implants labelling the magnocellular laminae. Implants into the visual centres of the midbrain produced retrograde axonal labelling rostral to the lateral geniculate nucleus only in the superficial part of the optic tract. These results demonstrate that the monkey's optic tract is not a simple topographic mapping of retinal eccentricity. Rather, the primary organizational principle is that of a segregation of functionally distinct optic axon classes. As fibre order in the mammalian optic tract is also a chronological index of axonal arrival during development, the present results provide specific predictions about the temporal order of ganglion call genesis and axonal addition within the visual pathway. They also provide an anatomical basis for the functionally selective visual impairments that may arise following local damage to the optic tract in humans.
Fibre organization of the monkey's optic tract: II. Noncongruent representation of the two half-retinae.
The representations of the two half-retinae were examined in the monkey's optic tract. Intravitreal injections of tritiated amino acids were made to reveal the distributions of the crossed and uncrossed populations of optic axons, while localized implants of horseradish peroxidase (HRP) were made into different regions of the optic tract in order to examine the distributions and morphological types of retrogradely labelled cells at corresponding loci in the two half-retinae. Crossed and uncrossed optic axons are intermingled throughout most of the optic tract, but uncrossed axons are very sparse or absent along both the deep and superficial extremes of the tract. Implants of HRP into the deeper regions of the tract demonstrate that the crossed and uncrossed optic axons of the P beta retinal ganglion cells are slightly out of binocular registration, with the uncrossed map being shifted to a slightly superficial location relative to the crossed map. The optic axons for the remaining cell classes, revealed by implants of HRP into the superficial portion of the tract, are much more conspicuously out of binocular registration (in particular, the P alpha optic axons); but in their cases, the uncrossed optic axons are shifted to deeper locations relative to the crossed optic axons. Further evidence that these optic axon classes are markedly out of binocular registration comes from the two optic tracts of a bilaterally destriated monkey, in which most of the P beta optic axons have undergone a transneuronal retrograde degeneration. Following a uni-ocular injection of tritiated amino acids, the distributions of the remaining crossed and uncrossed axonal labelling occupied different positions within the tract rather than being intermingled, with the uncrossed optic axons situated deep to the majority of crossed optic axons. These results demonstrate that the optic chiasm does not combine binocularly corresponding optic axons of similar type. They also demonstrate that noncongruent field defects should be a common consequence of damage to the optic tract in humans. If the fibre order in the mammalian optic tract arises as a consequence of the sequence of axonal addition during development, then differences in the relative times of genesis for nasal and temporal members of any cell class, and/or differences in the relative pathlengths between the eye and two optic tracts, may produce the fibre ordering described herein.
Pathways mediating resolution in the primate retina.
In recent years there has been a dramatic increase in knowledge of the anatomy of the primate retina that relates to the pathways involved in visual resolution. The density of cones at the fovea has been shown to have a surprising degree of individual variability and the cone distribution is asymmetric about the fovea with a greater density of cones in nasal than temporal retina. Information about the fine detail of our visual world is carried to the dorsal lateral geniculate nucleus of the thalamus in two parallel pathways. These pathways originate from two morphologically distinct types of ganglion cell; the M-ganglion cells project to the magnocellular layers, P-ganglion cells to the parvocellular layers. Different roles for the two types of ganglion cell in mediating spatial visual resolution have been proposed. These cannot be determined using available physiological and anatomical data.
Sensitivity to eye gaze in prosopagnosic patients and monkeys with superior temporal sulcus ablation.
Accuracy at perceiving frontal eye gaze was studied in monkeys and human subjects using a forced-choice detection task on paired photographs of a single human face. Monkeys learned the task readily, but after bilateral removal of the banks and floor of the superior temporal sulcus (STS) they failed to perform the task efficiently. This result is consistent with the conclusion, based on recordings from single cells in awake, behaving monkeys [Perret et al., Physiological Aspects of Clinical Neuro-ophthalmology, Chapman & Hall, London, 1988] that this region of the temporal lobe is important for coding information about eye-gaze of a confronting animal. Human subjects were given identical stimuli in a task where they were asked to detect "the face that is looking straight at you". Human performance is sensitive to the degree of angular deviation from the frontal gaze position, being poorest at small angular deviations from 0 degrees. This was also true of monkeys viewing these stimuli, pre- and post-operatively. Compared with normal controls, two humans prosopagnosics were impaired at this task. However the extent of impairment was different in the two patients. These findings are related to earlier reports (including those for patients with right-hemisphere damage without prosopagnosia), to normal performance with upright and inverted face photographs, and to notions of independent subsystems in face processing.
Development and retraction of a crossed retinal projection to the inferior colliculus in neonatal pigmented rats.
A transient aberrant projection from the retina to the contralateral inferior colliculus was demonstrated in pigmented rats in both whole-brains and sections following intra-ocular injection of horseradish peroxidase. The projection was prominent on the day of birth but reached its maximum density and extent after injection on day 1, when it covered at least a third of the inferior colliculus. It was absent or nearly absent by day 5. Its consistency, size, orderliness and systematic retraction suggest that it is not merely a developmental accident.
Retinal topography of the neonatal crossed aberrant exuberant projection to the inferior colliculus in the pigmented rat.
The retinal topography of the neonatal transient projection to the inferior colliculus was investigated in pigmented rats by injecting the retrograde fluorescent tracer Fast Blue into the colliculi. The results show that the projection arises from a small population of ganglion cells scattered across the entire contralateral retina, and that the transient projection is therefore not merely an overshoot of axons from the peripheral nasal retina whose appropriate target is the caudal pole of the superior colliculus.