Biomedical subjects
Publications and source records attributed to V Walsh.
Learning leads to neural changes often considered to be driven by 'smart' areas of the brain. A recent study of the cellular changes that underlie perceptual learning has found that plasticity in the primary visual cortex V1 is necessary for learning and the changes that correlate with learning are more complex than one might expect.
Much is known about the pathways from photoreceptors to higher visual areas in the brain. However, how we become aware of what we see or of having seen at all is a problem that has eluded neuroscience. Recordings from macaque V1 during deactivation of MT+/V5 and psychophysical studies of perceptual integration suggest that feedback from secondary visual areas to V1 is necessary for visual awareness. We used transcranial magnetic stimulation to probe the timing and function of feedback from human area MT+/V5 to V1 and found its action to be early and critical for awareness of visual motion.
Every neuroscientist knows that emotions are as much to do with the head as the heart, but as a number of new studies show, the heart - or rather the body - and the brain are by no means independent purveyors of feeling and emotion.
Music may be the food of love but it is also good fodder for cognitive scientists. Here we highlight a recent study of a neuropsychological patient who has lost her ability to read music, but not text, in the absence of any other musical deficit.
We aimed to use repetitive transcranial magnetic stimulation (rTMS) to disrupt speech with the specific objective of dissociating speech disruption according to whether or not it was associated with activation of the mentalis muscle. Repetitive transcranial magnetic stimulation (rTMS) was applied over two sites of the right and left hemisphere while subjects counted aloud and recited the days of the week, months of the year, and nursery rhymes. Analysis of EMG data and videotaped recordings showed that rTMS applied over a posterior site, lateral to the motor hand area of both the right and the left hemisphere resulted in speech disruption that was accompanied by activation of the mentalis muscle, while rTMS applied over an anterior site on the left but not the right hemisphere resulted in speech disruption that was dissociated from activation of the mentalis muscle. The findings provide a basis for the use of subthreshold stimulation over the extrarolandic speech disruption site in order to probe the functional properties of this area and to test psychological theories of linguistic function.
To investigate the hemispheric organization of a language-independent spatial representation of number magnitude in the human brain we applied focal repetitive transcranial magnetic stimulation (rTMS) to the right or left angular gyrus while subjects performed a number comparison task with numbers between 31 and 99. Repetitive TMS over the angular gyrus disrupted performance of a visuospatial search task, and rTMS at the same site disrupted organization of the putative "number line." In some cases the pattern of disruption caused by angular gyrus rTMS suggested that this area normally mediates a spatial representation of number. The effect of angular gyrus rTMS on the number line task was specific. rTMS had no disruptive effect when delivered over another parietal region, the supramarginal gyrus, in either the left or the right hemisphere.
Transcranial magnetic stimulation (TMS) can be conceptualized as a virtual lesion technique, capable of disrupting organized cortical activity, transiently and reversibly. The technique combines good spatial and temporal resolution and, moreover, because it represents an interference technique, can be said to have excellent functional resolution. The following is a review and discussion of the contribution which TMS has made to the study of vision, attention, development and plasticity and speech and language.
OBJECTIVE: To investigate the stability of visual phosphene thresholds and to assess whether they correlate with motor thresholds. BACKGROUND: Currently, motor threshold is used as an index of cortical sensitivity so that in transcranial magnetic stimulation (TMS) experiments, intensity can be set at a given percentage of this value. It is not known whether this is a reasonable index of cortical sensitivity in non-motor and hence whether it should be used in experiments where other cortical areas are targeted. Previous studies have indicated that phosphene threshold might be a suitable alternative in TMS studies of the visual system. METHOD: Using single pulse TMS visual phosphene and motor thresholds were measured in 15 subjects. Both thresholds were retested in seven of these subjects a week later. RESULT: Visual phosphene thresholds, though stable within subjects across the two sessions, showed greater variability than motor thresholds. There was no correlation between the two measures. CONCLUSION: TMS motor thresholds cannot be assumed to be a guide to visual cortex excitability and by extension are probably an inappropriate guide to the cortical excitability of other non-motor areas of the brain. Phosphene thresholds are proposed as a potential standard for inter-individual comparison in visual TMS experiments.
Transcranial magnetic stimulation (TMS) is a means of stimulating the brain from outside the skull with little, and occasionally no discomfort for the subject. A single TMS pulse, lasting less than 1 ms, can briefly disrupt the normal activity of a targeted region of the brain for tens of milliseconds, allowing the effects of disruption on specific perceptual and cognitive tasks to be measured behaviorally. Rapid, repeated pulses can disrupt activity for correspondingly longer periods. The reversibility of the effects make it possible to create 'virtual patients' who can be tested in the same way as actual patients with real brain damage in order to explore regional functional specialization. Although several aspects of TMS continue to be evaluated, such as its safety, the extent and localization of the effective region of induced electrical current, the importance of the waveform of the pulse, the configuration and positioning of the coil, its productivity has been firmly established in little more than 10 years of systematic use. Examples of the latter are given from investigations of the nature of visual phosphenes produced by TMS applied to different regions of the visual cortex in normal subjects and subjects with occipital or ocular damage in an attempt to reveal the role of visual cortex in visual awareness.
It is widely agreed that the right posterior parietal cortex has a preeminent role in visuospatial and orienting attention. A number of lines of evidence suggest that although orienting and the preparation of oculomotor responses are dissociable from each other, the two are intimately related. If this is true, then it should be possible to identify other attentional mechanisms tied to other response modalities. We used repetitive transcranial magnetic stimulation (rTMS) to demonstrate the existence of a distinct anterior parietal mechanism of motor attention. The critical area for motor attention is anterior to the one concerned with orienting, and it is lateralized to the left hemisphere in humans.
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Direct stimulation of visual cortex can produce illusory flashes of light, called phosphenes. Here we describe the spatial and motion properties of phosphenes produced by transcranial magnetic stimulation in normal subjects and in two subjects with peripheral or cortical blindness. The totally retinally blind subject experienced normal phosphenes, apart from their concentration in the centre of the visual field, whereas the hemianopic subject, lacking area V1, did not experience phosphenes when his surviving extrastriate visual areas were stimulated. In the absence of V1, magnetically induced activity was unable to generate a conscious visual percept in the field defect.
The two cerebral hemispheres are specialised for different cognitive functions, and which hemisphere's strategy is superior depends on the nature of the task. A new study of split-brain patients has provided another unexpected insight: the two hemispheres use different strategies when performing a guessing task.
Primate visual areas V4 and TEO are important for many aspects of visual perception and ablation of these areas leads to a wide range of deficits in visual discrimination, attention to less salient items, recognition of visually transformed objects, visual grouping and in visual memory. All these studies demonstrate that monkeys with V4 or TEO lesions have higher perceptual thresholds or are slower or less accurate than normal monkeys on a particular visual task. Here we show that when monkeys with V4 or TEO lesions perform a simple discrimination task on which they are unimpaired, they perform the task differently from normal monkeys. We examined visual priming in a feature detection task and discovered that it is diminished by lesions of TEO and abolished by lesions of V4. The results support the hypothesis, based on a recent demonstration of visuotopic priming in humans, that areas V4 and TEO are indispensable for normal visual form priming.
We applied single-pulse transcranial magnetic stimulation (TMS) to the occipital pole of healthy subjects while they performed a forced-choice visual letter-identification task. We found three separate periods when TMS suppressed performance; the first period is best explained by TMS-induced blinking whereas the last two periods are best explained by TMS-induced disruption of letter-processing in the early visual cortex. Unexpectedly, we also found that TMS-induced suppression progressively disappeared during three weeks of repeated TMS experiments. However, it was only suppression during the last two periods that disappeared; suppression during the first period remained undiminished. When subjects were then presented with dimmer letters, suppression reappeared. The most likely explanation is a practice-induced increase in neuronal activity in the early visual cortex.
Some visual attributes, such as colour, are purely visual, but others, such as orientation and movement, can be perceived by touch or audition. A magnetic stimulation study has now shown that the perception of tactile orientation may be influenced by visual Information.
Fifteen years after its introduction by Anthony Barker, transcranial magnetic stimulation (TMS) appears to be 'coming of age' in cognitive neuroscience and promises to reshape the way we investigate brain-behavior relations. Among the many methods now available for imaging the activity of the human brain, magnetic stimulation is the only technique that allows us to interfere actively with brain function. As illustrated by several experiments over the past couple of years, this property of TMS allows us to investigate the relationship between focal cortical activity and behavior, to trace the timing at which activity in a particular cortical region contributes to a given task, and to map the functional connectivity between brain regions.
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