Primary visual cortex in the brushtailed possum: receptive field properties and corticocortical connections

The corticocortical connections and receptive field properties of primary or striate visual cortex of the brushtailed possum, Trichosurus vulpecula, have been examined. In this Australian marsupial species primary visual cortex has connections with four other visual areas in the occipital lobe. In these adjacent visual areas fibers from striate cortex terminate mainly in layers 3 and 4 and in some cases also in layers 1 and 2. In all four areas return connections to striate cortex originate predominantly in layers 2 and 3, and to a much lesser extent in layers 5 and 6. Interhemispheric connections of striate cortex are limited to the boundary of striate and peristriate cortex. In addition to its cortical connections, striate cortex makes reciprocal connections with the claustrum. Most neurons in striate cortex are highly binocular. Of our sample of 113 visually responsive neurons, only 30% were orientation selective. On the basis of these observations we have compared striate cortex of the marsupial brushtailed possum with striate cortices of the American marsupial opossum and those of placental mammals.     

Recovery of vision after ischemic lesions: positron emission tomography

We used [18F]fluoro-2-deoxyglucose positron emission tomography (PET) to study serial changes in the local cerebral metabolic rate of glucose in 5 patients with ischemic lesions of the posterior afferent visual system causing homonymous visual field defects. All 5 patients had striking impairment of glucose metabolism in the striate cortex shortly after ictus. In 2 patients, visual field defects abated, and repeat PET scans showed reduced size of the metabolic lesion and improvement of striate metabolism. Three patients did not recover vision, and repeat PET scans did not show improvement. Patients who recovered vision had ischemic lesions outside the occipital lobe, while those who did not experience improvement had primary damage to the occipital lobe itself.     

Neural activity within area V1 reflects unconscious visual performance in a case of blindsight

Although lesions of the striate (V1) cortex disrupt conscious vision, patients can demonstrate surprising residual abilities within their affected visual field, a phenomenon termed blindsight. The relative contribution of spared "islands" of functioning striate cortex to residual vision, versus subcortical pathways to extrastriate areas, has implications for the role of early visual areas in visual awareness and performance. Here we describe the behavioral and neural features of residual cortical function in Patient M.C., who sustained a posterior cerebral artery stroke at the age of 15 years. Within her impaired visual field, we found preserved visual abilities characteristic of blindsight, including superior detection of motion, and above-chance discrimination of shape, color, and motion direction. Functional magnetic resonance imaging demonstrated a retinotopically organized representation of M.C.'s blind visual field within the lesioned occipital lobe, specifically within area V1. The incongruity of a well-organized cortex and M.C.'s markedly impaired vision was resolved by measurement of functional responses within her damaged occipital lobe. Attenuated neural contrast-response functions were found to correlate with M.C.'s impaired psychophysical performance. These results demonstrate that the behavioral features of blindsight may arise in the presence of residual striate responses that are spatially organized and sensitive to contrast variation.     

Cerebral areas mediating visual redirection of gaze: cooling deactivation of 15 loci in the cat

In humans, damage to posterior parietal or frontal cortices often induces a severe impairment of the ability to redirect gaze to visual targets introduced into the contralateral field. In cats, unilateral deactivation of the posterior middle suprasylvian (pMS) sulcus in the posterior inferior parietal region also results in an equally severe impairment of visually mediated redirection of gaze. In this study we tested the contributions of the pMS cortex and 14 other cortical regions in mediating redirection of gaze to visual targets in 31 adult cats. Unilateral cooling deactivation of three adjacent regions along the posterior bend of the suprasylvian sulcus (posterior middle suprasylvian sulcus, posterior suprasylvian sulcus, and dorsal posterior ectosylvian gyrus at the confluence of the occipital, parietal, and temporal cortices) eliminated visually mediated redirection of gaze towards stimuli introduced into the contralateral hemifield, while the redirection of gaze toward the ipsilateral hemifield remained highly proficient. Additional cortical loci critical for visually mediated redirection of gaze include the anterior suprasylvian gyrus (lateral area 5, anterior inferior parietal cortex) and medial area 6 in the frontal region. Cooling deactivation of: 1) dorsal or 2) ventral posterior suprasylvian gyrus; 3) ventral posterior ectosylvian gyrus, 4) middle ectosylvian gyrus; 5) anterior or 6) posterior middle suprasylvian gyrus (area 7); 7) anterior middle suprasylvian sulcus; 8) medial area 5; 9) the visual portion of the anterior ectosylvian sulcus (AES); 10) or lateral area 6 were all without impact on the ability to redirect gaze. In summary, we identified a prominent field of cortex at the junction of the temporo-occipito-parietal cortices (regions pMS, dPE, PS), an anterior inferior parietal field (region 5L), and a frontal field (region 6M) that all contribute critically to the ability to redirect gaze to novel stimuli introduced into the visual field during fixation. These loci have several features in common with cortical fields in monkey and human brains that contribute to the visually guided redirection of the head and eyes.     

Visual functions without the occipital lobe or after cerebral hemispherectomy in infancy

This paper investigates whether and to what extent vision with awareness is still possible in the whole visual field after loss of the occipital lobe of one or both cerebral hemispheres or after hemispherectomy in childhood. The visual functions of four children who suffered from unilateral or bilateral loss of the occipital lobe or who had been hemispherectomized were examined. The results show that even after unilateral loss of the striate and prestriate cortex the extent of the visual field may still be in the normal range. The residual visual functions may be mediated by intact extrastriate areas such as V5 and LO of the damaged cerebral hemisphere. It is also shown that even after complete hemispherectomy in early life the visual field may have a normal extent and that conscious visual perception in the whole visual field may be preserved. In hemispherectomized children, the remaining cerebral hemisphere or neural structures in the midbrain, including the superior colliculi and the praetectum, may be able to mediate these visual functions.     

Clinical features and pathophysiological mechanism of the hemianoptic complication after the occipital transtentorial approach

Objective

To obtain detailed insight into neuro-ophthalmological characteristics and pathophysiology of hemianoptic complications after occipital transtentorial surgery.

Methods

We reviewed the cases of 14 patients surgically treated by the occipital transtentorial approach. Treated lesions included 6 posterior third ventricle tumors, including pineal and tectal lesions, 3 falco-tentorial meningiomas, and 5 superior cerebellar lesions. The surgeries were performed by the unilateral occipital transtentorial approach with patients in the prone position.

Results

Visual functions were preoperatively normal in all patients. After surgery, 11 patients (79%) showed hemianoptic complications detected by a confrontation test in the immediate postoperative period. The condition began to improve in the early postoperative days. The visual field recovered completely in 6 patients within 10 days, 2 patients recovered within 3 months, and 3 patients complained of permanent visual field defects. Optometric neuro-ophthalmic evaluation in the early postoperative period failed to detect complete homonymous hemianopsia, but homonymous inferior quadrantanopia and scotomatous defects were observed in 6 patients. These visual field defects were permanent in 3 patients. Postoperative MRI showed no morphological abnormality except these three patients. Atrophic change of the occipital lobe with preservation of striate cortex was associated with persistent visual field defects in two patients. Cerebral blood flow evaluation by single photon emission computed tomography suggested that temporary local hyperperfusion of the retracted occipital region when visual field defect was present.

Conclusion

Hemianoptic visual field defects can recover via inferior quadrantanopia or scotomatous defect. All of these defects are attributable to injury to the optic radiation as well to the occipital lobe. Hyperperfusion of the retracted occipital region may underlie the pathophysiology of hemianoptic complications after the occipital transtentorial approach.     

“Blindsight”: Improvement of visually guided eye movements by systematic practice in patients with cerebral blindness

Patients with damage to the geniculo-striate visual pathways were trained to discriminate different positions of targets presented briefly in the blind region of their visual field. The results indicate that the ability to detect and localize stimuli improved markedly after the systematic training of visually evoked saccadic eye movements. It is concluded that even in the absence of the striate cortex detection and localization of stimuli remained still intact, and that this visual capacity is probably mediated by the extrastriate retionotecnal pathway.     

Bilateral altitudinal anopia caused by infarction of the calcarine cortex.

The patient reported here had a bilateral inferior altitudinal hemianopia from lesions of the calcarine (striate) cortex of the occipital lobes. The only significant pathologic findings were bilateral calcarine artery occlusive disease, with infarcts of the striate cortex on both sides. The ages of the infarcts appeared compatible with the clinical development of the respective visual field defects. The rest of the visual system was anatomically intact.     

Saccade induced cortical activation in patients with post-stroke visual field defects

Substantial disability in patients with hemianopia results from reduced visual perception. Several studies have shown that these patients have impaired saccades but may improve search strategies with appropriate training of saccades. We used fMRI to study the representation of saccades in patients with post-stroke hemianopia to the left. Brain activation during visually guided saccades was measured in 10 patients with a pure occipital cortical lesion causing homonymous hemianopia and in 10 healthy control subjects. Differences in activation between rest and saccades and between controls and patients were assessed with statistical parametric mapping (SPM'99). In normal subjects, significant activation was found in the frontal and parietal eye fields bilaterally and in the supplementary eye field. These areas were also activated in patients, however, to a lesser degree. In contrast, an area of increased activation in patients was found in the posterior parietal cortex of the (non-affected) left hemisphere. Visual field defects after striate lesions are associated with changes in the frontoparietal network underlying the cortical control of saccades.     

Developmental lesions of visual cortex influence control of reaching

We examined visually guided reaching movements in a young adult (EW) who had extensive bilateral lesions in the visual cortex since birth. EW lacked a right occipital lobe and ventral portions of the left and had poor visual acuity (3/400), yet could point to visual targets as quickly as 9 controls with visual cortex lesions acquired in adulthood and 4 adults without neurological disease. However, EW's endpoint variability and hand movement path curvature were much greater, especially for left hand movements, in concert with large sensorimotor transformation errors. Experimental reduction of acuity (to 3/240 or worse) in the normal controls produced symmetric increases in endpoint variability but did not change hand path curvature, indicating that EW's impaired movements were not due to poor vision alone. Results suggest that visual cortex in early life supports the development of lifelong neural mechanisms for the planning and control of reaching movements.     

Effects of straite and occipital cortical lesions of visual discrimination in the rabbit

Learning of brightness and pattern discrimination in normal rabbits and in rabbits with striate and occipital lesions was compared. Those with striate lesions did not differ from normal rabbits in learning brightness discriminations but took significantly longer to learn patterns consisting of striations differing by 90° and 45°. No rabbit with striate lesions learned to discriminate either striations differing by 22° or a cross/ circle discrimination. Those with occipital lesions did not differ from normal rabbits in learning brightness or pattern discriminations. The relationship between receptive field characteristics of neurons in striate cortex and the effects of striate lesions on visual discrimination is discussed.     

Study of the human visual cortex: direct cortical evoked potentials and stimulation.

The authors studied the visual cortex of 15 patients undergoing studies for medically intractable epilepsy. Although the subdural and strip electrode placement varied in each of these patients, there were enough electrodes over the visual cortex to complete studies involving evoked potentials and direct cortical stimulation. Visual evoked potentials were elicited using two check sizes (50 and 16 min) for pattern reversal studies, 50 min checks for on-off stimulation, 50 min checks for horizontal and vertical hemifields and simple flash for the VEP. These studies demonstrated that the pattern reversal and on-off stimuli caused very complex, multipotential waveforms in striate and vision associational cortex that do not resemble the response obtained at the scalp. Different volumes of visual cortex are activated by stimulation with 16 min checks, 50 min checks and simple flash. Flash activates the largest volume of visual cortex and it is likely that this finding is what makes this test of so little value clinically. Direct cortical stimulation shows that colored responses are generated primarily in the posterior striate cortex and inferior occipital lobe, while movement is primarily generated by the visual association cortex. No complex visual images were obtained by stimulation of either the striate cortex or visual association cortex. The brain mechanisms that lead to formed visual images remain to be identified.     

New view of the organization of the pulvinar nucleus in Tupaia as revealed by tectopulvinar and pulvinar-cortical projections

The projections of the superficial layers of the superior colliculus to the pulvinar nucleus in Tupaia were reexamined by injecting WGA-HRP into the tectum. The main result was finding two different patterns of terminations in the pulvinar nucleus: a zone remote from the lateral geniculate nucleus, which occupies the dorsomedial and caudal poles of the pulvinar nucleus, was almost entirely filled with terminals in every case irrespective of the location of the injection site; and a second division of the pulvinar nucleus, adjacent to the lateral geniculate nucleus, contained irregular patches--much more densely populated--and the distribution of patches varied from case to case. We call the first projection "diffuse" and the patchy projection "specific." Next we injected several divisions of the extrastriate visual cortex to find the cortical target of each pathway. The diffuse path terminates in the ventral temporal area (Tv). The specific path terminates in the dorsal temporal area (Td) and area 18. We speculated about the significance of the two pathways: the specific path may be responsible for the preservation of vision after removal of the striate cortex; the diffuse path may have an important place in the evolution of the visual areas of the temporal and occipital lobe. We argued that the target of the diffuse path is in a position to relate limbic and visual impulses and relay the product of such integration to the other visual areas, striate as well as extrastriate cortex.     

Magnetoencephalographic fields from patients with spontaneous and induced migraine aura.

We investigate and characterize the magnetoencephalographic waveforms from patients during spontaneous and visually induced migraine aura. Direct current neuromagnetic fields were measured during spontaneous onset of migraine auras in 4 migraine patients, and compared with recordings from 8 migraine-with-aura patients and 6 normal controls during visual stimulation of the occipital cortex. Complex direct current magnetoencephalographic shifts, similar in waveform, were observed in spontaneous and visually induced migraine patients, but not in controls. Two-dimensional inverse imaging showed multiple cortical areas activated in spontaneous and visually induced migraine aura patients. In normal subjects, activation was only observed in the primary visual cortex. Results support a spreading, depression-like neuroelectric event occurring during migraine aura that can arise spontaneously or be visually triggered in widespread regions of hyperexcitable occipital cortex.     

Afferent basis of visual response properties in area MT of the macaque. II. Effects of superior colliculus removal

In a previous study (Rodman et al., 1989), we found that many neurons in the middle temporal area (MT) of the macaque monkey remain visually responsive and directionally selective after striate cortex lesions or cooling. In the present study, we examined the effects of superior colliculus (SC) lesions and combined lesions of striate cortex and the SC on the visual properties of MT neurons. Removal of the SC alone had no effect on the proportion of visually responsive cells, strength of direction selectivity and direction tuning, orientation tuning, receptive field size, or binocularity in MT. There was, however, a slight increase in response strength to both stationary and moving slit stimuli. In contrast to the minor effects of SC lesions alone, addition of an SC lesion to striate cortex damage abolished all visual responsiveness in area MT. The results indicate that pathways damaged by the SC lesion are not necessary for most of the properties of MT neurons found in the intact animal, although these pathways are capable of sustaining considerable visual responsiveness and direction selectivity when striate input is removed.     

Topographic organization of human visual areas in the absence of input from primary cortex.

Recently, there has been evidence for considerable plasticity in primary sensory areas of adult cortex. In this study, we asked to what extent topographical maps in human extrastriate areas reorganize after damage to a portion of primary visual (striate) cortex, V1. Functional magnetic resonance imaging signals were measured in a subject (G.Y.) with a large calcarine lesion that includes most of primary visual cortex but spares the foveal representation. When foveal stimulation was present, intact cortex in the lesioned occipital lobe exhibited conventional retinotopic organization. Several visual areas could be identified (V1, V2, V3, V3 accessory, and V4 ventral). However, when stimuli were restricted to the blind portion of the visual field, responses were found primarily in dorsal extrastriate areas. Furthermore, cortex that had formerly shown normal topography now represented only the visual field around the lower vertical meridian. Several possible sources for this reorganized activity are considered, including transcallosal connections, direct subcortical projections to extrastriate cortex, and residual inputs from V1 near the margin of the lesion. A scheme is described to explain how the reorganized signals could occur based on changes in the local neural connections.     

Cerebral color blindness: an acquired defect in hue discrimination.

In contrast to the traditional view that striate visual cortex (area 17) is surrounded by two homogeneous cortical areas (areas 18 and 19), recent studies have shown that mammalian extrastriate visual cortex contains several anatomically and functionally distinct subregions. One such region, the V-4 complex of the rhesus monkey, is highly specialized for the analysis of color information, suggesting that a lesion in a homologous region might produce a defect in color vision while sparing other visual functions. We have studied a patient whose clinical syndrome supports this suggestion: a 44-year-old man with normal color vision suffered two cerebral infarctions that produced first a right and then a left superior homonymous quadrantanopia and also caused prosopagnosia, topographical disorientation, and severely impaired color vision. Computed tomography demonstrated extensive lesions in both inferior occipital lobes in the territories of the lateral branches of the posterior cerebral arteries, involving the lingual and medial occipitotemporal gyri bilaterally; these gyri contain the inferior portion of striate cortex and segments of extrastriate visual cortex. The patient had no difficulty in giving the correct color names associated with common objects presented either verbally or in outline drawings. Standardized testing with the Farnsworth-Munsell 100-hue test, the Nagel anomaloscope, and a method that tests for just-noticeable differences between monochromatic stimuli all showed that the patient's ability to distinguish one color from another was markedly imparied but not totally absent. In contrast, visual acuity, reading, visually guided eye movements, and stereopsis were normal. Cells in the V-4 complex of monkey extrastriate cortex are highly specialized for distinguishing one color from another; the hue discrimination deficit that was demonstrated in this patient with cerebral color blindness indicates that a region or regions with similar function has been damaged.     

Epileptic activity of the occipital lobe. Clinico-electroencephalographic contribution

Our analysis of the course of illness in 14 patients, whose common electroencephalographic characteristic was epileptogenic activity in the occipital area, showed very different clinical symptoms. The first group comprised patients who presented bilateral amaurosis. In four of these cases, the occipital hypersynchronous EEG activity was merely a secondary symptom of either ischaemic hypoxia or of a degenerative process in the occipital visual cortex and was not responsible for the genesis of the actual blindness. In two further cases of monosymptomatic temporary loss of vision, it was difficult to make a differential diagnosis between ictal blindness, respectively status epilepticus amauroticus occurring in a occipital lobe epilepsy and a migraine attack involving the basilar territory. The second group comprised five patients with paroxysmal visual hallucinations respectively illusions. Three of them suffered from hallucinations of the elementary type, respectively flickering fits in the hemianopic field, symptoms which are based on discharges in the visual cortex of the occipital lobe. In a case of one patient with complex visual hallucinations as well as in a further case with visual illusions, it was not possible to find out with certainty their place of origin. A study of these cases shows that the cortical or sub-cortical functional disturbance within the visual system causing the various optical deformations and visual hallucinations, form an inhomogeneous group with different etiology. In the only patient belonging to the third group, whose seizures were i.a. characterized through motor phenomena in the field of the ocular organs and the tonic lateral turning movement of the bulbi of the eyes and of the head, an occipital epileptic crisis with spread of discharges from the occipital pole to the frontomesial surface should be assumed. The occurrence of complex partial seizures, respectively generalized tonic-clonic attacks in two patients of the fourth group who have definite epileptogenic EEG-activity in the occipital area, can be explained by a propagation of paroxysmal activity to the temporal lobe or to the motor cortex. Because of the marked tendency to propagation of the hypersynchronous activity originating in the occipital lobe, many combinations of sensory and/or motor symptoms can occur within the frame-work of occipital epileptic seizures. On the basis of one scalp EEG finding, no final localizing conclusions may be drawn here.     

Location of the human posterior eye field with functional magnetic resonance imaging.

The frontal eye field and parietal eye field are known to be involved during visually guided saccades. As the location of the human parietal eye field is not yet well known, functional MRI was used during such a saccade task to better localise this field. Besides activity in visual areas of the occipital cortex, bilateral activity was seen in the precentral sulcus, corresponding to the frontal eye field, and in the deep region of the intraparietal sulcus. It is suggested that this intraparietal area, bordering areas 39 and 40 of Brodmann, corresponds to the human parietal eye field.