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Biomedical subjects

P Stoerig

Publications and source records attributed to P Stoerig.

At least 19 recordsLinked to original sources

Visual detection in monkeys with blindsight.

Monkeys with unilateral striate cortical removal show residual visual abilities in their affected hemifield. To learn whether the monkeys, like patients with blindsight, lose the phenomenal representation of the visual stimuli they nevertheless respond to, we first studied their ability to localize a briefly presented target in either hemifield. By varying the luminance of the stimuli we determined their visual sensitivity, which was reduced by 0.3-1.5 log units in the impaired hemifield; suprathreshold stimuli yielded almost perfect performance. We then presented two tests designed to show whether the monkeys categorized visual stimuli in the impaired field in the same manner as they categorize them in the normal field. In the first test, they had to respond differently according to whether one or two lights were presented, with the relative position of the two stimuli in a pair being varied. Whenever one of the paired stimuli lay in the impaired hemifield, two of the three monkeys consistently ignored it, and responded as if it had been a single stimulus in the good field. In the second test, trials consisting of a single stimulus light were interleaved with blank trials. The monkey touched the position of the light or made a different response, indicating that no light had appeared. All three monkeys responded to a light of supra-threshold luminance presented in the impaired field as if it were a blank trial. These results suggest that monkeys with striate cortical destruction, like neurological patients with similar lesions, have blindsight rather than phenomenal vision when they have to detect brief static visual targets.

Animals

Blindsight in man and monkey.

In man and monkey, absolute cortical blindness is caused by destruction of the optic radiations and/or the primary visual cortex. It is characterized by an absence of any conscious vision, but stimuli presented inside its borders may nevertheless be processed. This unconscious vision includes neuroendocrine, reflexive, indirect and forced-choice responses which are mediated by the visual subsystems that escape the direct cerebral damage and the Ensuring degeneration. While extrastriate cortical areas participate in the mediation of the forced-choice responses, a concomitant striate cortical activation does not seem to be necessary for blindsight. Whether the loss of phenomenal vision is a necessary consequence of striate cortical destruction and whether this structure is indispensable for conscious sight are much debated questions which need to be tackled experimentally.

Animals

No blindsight following hemidecortication in human subjects?

Using a guessing paradigm we measured visual sensitivity in the blind and normal half-fields of four cerebrally hemidecorticated patients. In the blind field, sensitivity was reduced by approximately 3 long units. Stimuli which produced significant detection also evoked conscious sensations of light and colour. Control experiments showed that although sensitivity in the blind field depended in a normal fashion on background luminance it was independent of the luminance of a local platform, and showed no spatial summation. This residual vision can be explained by intraocular light diffusion and reflection.

Brain Diseases

Varieties of vision: from blind responses to conscious recognition.

Lesions in consecutive parts of the visual system cause visual deficits that spare increasingly complex residual functions. Patients with lesions up to and including primary visual cortex can show neuroendocrine, reflexive, implicit and forced-choice responses to visual stimulation but no conscious vision. In contrast, patients with lesions in higher visual cortical areas have conscious vision. Its lowest level is that of phenomenal vision, followed by object vision and recognition. These levels are dissociable. They require the integrity of different parts of the system.

Animals

Blindsight in monkeys.

Blindsight, the visually evoked voluntary responses of patients with striate cortical destruction that are demonstrated despite a phenomenal blindness, has attracted attention from neuroscientists and philosophers interested in problems of perceptual consciousness and its neuronal basis. It is assumed to be mediated by the numerous extra-geniculostriate cortical retinofugal pathways whose properties are studied primarily in monkeys. Like patients with blindsight, monkeys with lesions of the primary visual cortex can learn to detect, localize and distinguish between visual stimuli presented within their visual field defects. Although the patients deny seeing the stimuli they can nevertheless respond to (by forced-choice guessing) in their phenomenally blind fields, it is not known whether the monkeys experience the same absence of phenomenal vision. To determine whether they too have blindsight, or whether they actually see the stimuli in their field defects, monkeys who showed excellent detection in tasks where a visual stimulus was presented on every trial, albeit at different positions, were tested in a signal-detection task in which half the trials were blank trials, with no visual stimulus. They classified the visual stimuli presented in the field defect as blank trials, demonstrating, like patients, blindsight rather than degraded real vision.

Animals

Visual perception and phenomenal consciousness.

In the (re-)animated debate on consciousness we focus on three questions: Who has consciousness? What is its neuronal basis? What is its function? Regarding the first, we suggest that consciousness is exclusive to living organisms able to distinguish self from non-self. It may be restricted further to organisms who possess a repertoire of overt and covert behaviour which can be voluntarily modified and suppressed. This requires an intermediary neuronal net mediating between sensory input and behavioural output. What are the properties of this net which distinguish unalloyed information processing per se from conscious representation? To tackle this second question, we use the visual system and the functional losses that result from lesions at its different levels, and differentiate a reflexive, a phenomenal, and a consciously accessible stage of visual processing. We suggest that the latter two represent two distinct aspects of consciousness. Blindsight, a neurological example of visual processing in the absence of phenomenal vision, could help to elucidate the neuronal basis of phenomenality, and the special role of striate cortex. Like the patients, our monkeys with unilateral striate cortical removal show evidence not just of residual visual processing, but of the same absence of phenomenal vision, opening routes to further exploring the details of its neuronal implementation. The second aspect, conscious access to presently or previously processed information, is likely to require higher cortical structures, and may depend on the stage of phenomenal representations. In patients with blindsight, both aspects are lost, and it is conceivable that a loss of phenomenality generally causes a loss of conscious accessibility. One important function of phenomenal representations, our third question, would then be to allow conscious retrieval and manipulation of currently processed or formerly stored information, enabling organisms to consciously think and plan.

Animals

Retinal ganglion cells labelled from the pulvinar nucleus in macaque monkeys.

In order to study the distribution and morphological classes of retinal ganglion cells that can be retrogradely labelled from the pulvinar nucleus, we made two iontophoretic injections of horseradish peroxidase into the pulvinar in each hemisphere of five macaque monkeys. The retrogradely labelled ganglion cells projecting to or through the pulvinar nucleus were examined in retinal whole-mounts. They comprise all three major ganglion cell classes. Primate gamma cells formed the great majority of classifiable cells and, like the primate alpha cells that were found in much smaller numbers, they were already known to send axons to the superior colliculus and to the pretectal complex. In contrast, the primate beta cells were hitherto thought to project solely to the dorsal lateral geniculate nucleus. This primate beta cell projection to an extrageniculate target could account in part for the substantial number of primate beta cells that escape transneuronal retrograde retinal degeneration following striate cortical ablation, and might contribute to the residual visual sensitivity that survives destruction of striate cortex and the degeneration of the lateral geniculate nucleus.

Animals

The visual system and levels of perception: properties of neuromental organization.

To see whether the mental and the neural have common attributes that could resolve some of the traditional dichotomies, we review neuroscientific data on the visual system. The results show that neuronal and perceptual function share a parallel and hierarchical architecture which is manifest not only in the anatomy and physiology of the visual system, but also in normal perception and in the deficits caused by lesions in different parts of the system. Based on the description of parallel hierarchical levels of active information processing in the visual brain, we suggest a concept of dissociable levels of perception, advocating that the phenomenal perception and recognition is realized in the functional integrity of a network of reciprocal cortico-cortical connections. The properties shared by neuronal and perceptional functions provide a basis for a neuromental monism in which both functions are attributed a causal role.

Cerebral Cortex

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.

Adult

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.

Animals

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.

Animals

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.

Adult

Projection patterns of surviving neurons in the dorsal lateral geniculate nucleus following discrete lesions of striate cortex: implications for residual vision.

In four monkeys with long-standing partial ablation of the striate cortex pellets of horseradish peroxidase were placed in either the striate cortex immediately adjacent to the ablation, or in the extrastriate cortex of the ventral prelunate gyrus, i.e. in visual area V4. We examined the dorsal lateral geniculate nucleus to see whether surviving neurons, within the region that shows retrograde degeneration as a result of the cortical lesion, project to remaining striate cortex and/or to extrastriate cortex. Neurons labelled from extrastriate cortex were found throughout the degenerated region, whereas neurons labelled from striate cortex were confined to the border between the normal and degenerated region of the nucleus. This shows that isolated neurons found within the degenerated region survive striate cortex damage because they project to an extrastriate visual area, and not because their terminals depart from the otherwise strict topographic representation of the lateral geniculate nucleus on to striate cortex.

Animals

Transneuronal retrograde degeneration of retinal ganglion cells after damage to striate cortex in macaque monkeys: selective loss of P beta cells.

We examined the retinae of two monkeys whose left striate cortex had been removed eight years previously and compared the transneuronally degenerated hemiretina of each eye with the normal hemiretina, and with the retinae of normal monkeys. All retinae were prepared as whole mounts. One from each pair was stained with Cresyl Violet; the other was reacted for horseradish peroxidase two days after placing pellets of the enzyme in the optic nerve. Measurements of ganglion cell density in the Nissl-stained retina of the contralateral right eye showed that approximately 80% of retinal ganglion cells were missing in the central 30 degrees of the degenerated hemiretinae. More peripherally the percentage loss was less extensive. Measurements of cell soma size and dendritic field size of peroxidase-labelled classified surviving cells in the degenerated temporal hemiretina of the ipsilateral eye showed them to be morphologically normal. In comparison with the normal hemiretina, however, the mean soma size at three selected eccentricities was larger than normal, suggesting selective loss of smaller ganglion cells. Classification of peroxidase-labelled ganglion cells in the normal and degenerated hemiretinae revealed that the population of P beta cells was reduced by as much as 85% in the degenerated region. There was comparable change in the density of P alpha or P gamma cells. The degeneration of the great majority of P beta cells, which are believed to be the morphological substrate of ganglion cells with small and colour-opponent receptive fields, must set limits on the visual sensitivity and discrimination that survive damage to striate cortex.

Animals

Residual target detection as a function of stimulus size.

In the visual field defects of 5 patients with damage in the postgeniculate part of the primary visual system, residual detectability of an achromatic intensity target was tested as a function of target diameter, which was 10, 27, 44, 69 and 116 min of arc of visual angle. The set of 5 targets was presented at 3 different intensity ranges on a background of low photopic luminance. Within this size range residual detectability was a V-shaped function of target size with a dip at 44 min of arc. This result is discussed in relation to the centre/surround organization of receptive and perceptive fields.

Adult

Plasticity and rigidity in the representation of the human visual field.

Neuronal plasticity in the mammalian visual system has been studied with a variety of experimental methods like induction of artificial squint and eye rotation. To investigate neuronal plasticity in the human visual system, we examined a patient with a congenital convergent squint of his left eye, who later suffered a vascular lesion in his left occipital lobe that led to an incomplete hemianopia in his right visual field. The examination revealed that the visual field representation in the striate cortex is rigidly prewired with reference to the anatomical fovea. In contrast, plasticity in the oculomotor system enables the patient to use a functional visual axis that does not correspond to the anatomical fovea. Local alterations of sensitivity within the visual field that indicate interactions among non-corresponding retinal points provide additional evidence of functional plasticity.

Esotropia