Disruptions to human speed perception induced by motion adaptation and transcranial magnetic stimulation

To investigate the underlying nature of the effects of transcranial magnetic stimulation (TMS) on speed perception, we applied repetitive TMS (rTMS) to human V5/MT+ following adaptation to either fast‐ (20 deg/s) or slow (4 deg/s)‐moving grating stimuli. The adapting stimuli induced changes in the perceived speed of a standard reference stimulus moving at 10 deg/s. In the absence of rTMS, adaptation to the slower stimulus led to an increase in perceived speed of the reference, whilst adaptation to the faster stimulus produced a reduction in perceived speed. These induced changes in speed perception can be modelled by a ratio‐taking operation of the outputs of two temporally tuned mechanisms that decay exponentially over time. When rTMS was applied to V5/MT+ following adaptation, the perceived speed of the reference stimulus was reduced, irrespective of whether adaptation had been to the faster‐ or slower‐moving stimulus. The fact that rTMS after adaptation always reduces perceived speed, independent of which temporal mechanism has undergone adaptation, suggests that rTMS does not selectively facilitate activity of adapted neurons but instead leads to suppression of neural function. The results highlight the fact that potentially different effects are generated by TMS on adapted neuronal populations depending upon whether or not they are responding to visual stimuli.     

The neural basis of the Enigma illusion: a transcranial magnetic stimulation study

The aim of this study was to test the role of the visual primary (V1) and the middle temporal area (V5/MT) in the illusory motion perception evoked by the Enigma figure. The Enigma figure induces a visual illusion that is characterized by apparent rotatory motion in the presence of a static figure. By means of repetitive transcranial magnetic stimulation (rTMS) we show that V5/MT is causally linked to the illusory perception of motion. When rTMS was applied bilaterally over V5/MT just prior to presentation of the Enigma figure, the perception of illusory motion was disrupted for approximately 400 ms resulting in a delayed illusion onset. In contrast, rTMS applied over V1 did not have any effect on the illusory perception of motion. These results show that V5/MT, a visual cortical area associated with real motion perception, is also important for the perception of illusory motion, while V1 appears not to be functionally involved in illusory motion perception.     

The contribution of color to motion processing in Macaque middle temporal area.

The chromatic properties of an image yield strong cues for object boundaries and thus hold the potential to facilitate the detection of object motion. The extent to which cortical motion detectors exploit chromatic information, however, remains a matter of debate. To address this further, we quantified the strength of chromatic input to directionally selective neurons in the middle temporal area (MT) of macaque cerebral cortex using an equivalent luminance contrast (EqLC) paradigm. This paradigm, in which two sinusoidal gratings, one heterochromatic and the other achromatic, are superimposed and moved in opposite directions, allows the sensitivity of motion detectors to heterochromatic stimuli to be quantified and expressed relative to the benchmark of sensitivity for a luminance-defined stimulus. The results of these experiments demonstrate that the chromatic contrast in a moving red-green heterochromatic grating strongly influences directional responses in MT when the luminance contrast in that same grating is relatively low; for such stimuli, EqLC is at least 5%. When luminance contrast is added to the heterochromatic grating, however, EqLC wanes sharply and becomes negative (-4%) when luminance contrast is sufficiently high (>17-23%). Thus, the chromatic properties of an object appear to confer little or no benefit to motion processing by MT neurons when sufficient luminance contrast concurrently exists. These data support a simple model in which chromatic motion processing in MT is almost exclusively determined by magnocellular input. Additionally, a comparison of neuronal and psychophysical data suggests that MT may not be the sole contributor to the perceptual experience elicited by motion of heterochromatic patterns, or that only a subset of MT neurons serve this function.     

Motion standstill leads to activation of inferior parietal lobe

Previous studies on motion perception revealed motion-processing brain areas sensitive to changes in luminance and texture (low-level) and changes in salience (high-level). The present functional magnetic resonance imaging (fMRI) study focused on motion standstill. This phenomenon, occurring at fast presentation frequencies of visual moving objects that are perceived as static, has not been previously explored by neuroimaging techniques. Thirteen subjects were investigated while perceiving apparent motion at 4 Hz, at 30 Hz (motion standstill), isoluminant static and flickering stimuli, fixation cross, and blank screen, presented randomly and balanced for rapid event-related fMRI design. Blood oxygenation level-dependent (BOLD) signal in the occipito-temporal brain region MT/V5 increased during apparent motion perception. Here we could demonstrate that brain areas like the posterior part of the right inferior parietal lobule (IPL) demonstrated higher BOLD-signal during motion standstill. These findings suggest that the activation of higher-order motion areas is elicited by apparent motion at high presentation rates (motion standstill). We interpret this observation as a manifestation of an orienting reaction in IPL towards stimulus motion that might be detected but not resolved by other motion-processing areas (i.e., MT/V5).     

Repetitive transcranial magnetic stimulation of human area MT/V5 disrupts perception and storage of the motion aftereffect

Following adaptation to a moving stimulus, the introduction of a stationary pattern creates the illusion of motion. This phenomenon, known as the motion aftereffect (MAE), can be delayed by placing a blank storage interval between the adapting and test stimuli. Human motion selective area MT/V5 has been proposed as the likely neural origin of MAEs. To examine the role of MT/V5 in perceiving and storing MAEs, we applied repetitive transcranial magnetic stimulation (rTMS) to this area during a 10 s storage interval and while subjects perceived illusory motion. Our results show that rTMS disrupts perception of the MAE when it is delivered in the early parts of the storage period and when it is applied during the perceptual MAE itself. Stimulation of control regions corresponding to V1 or Cz did not affect the MAE. In addition, magnetic stimulation of dorsolateral prefrontal and posterior parietal cortices did not disrupt MAE perception. These data provide experimental support for the notion that MT/V5 subserves perception and storage of the motion aftereffect.     

Breaking camouflage: responses of neurons in the middle temporal area to stimuli defined by coherent motion

Camouflaged animals remain inconspicuous only insofar as they remain static. This demonstrates that motion is a powerful cue for figure-ground segregation, allowing detection of moving objects even when their luminance and texture characteristics are matched to the background. We investigated the neural processes underlying this phenomenon by testing the responses of neurons in the middle temporal area (MT) to 'camouflaged' bars, which were rendered visible by motion. These responses were compared with those elicited by 'solid' bars, which also differed from background in terms of their mean luminance. Most MT neurons responded strongly to camouflaged bars, and signaled their direction of motion with precision, with direction-tuning curves being only slightly wider than those measured with solid bars. However, the tuning of most MT cells to stimulus length and speed depended on the type of stimulus - in comparison with solid bars, responses to camouflaged bars typically showed more extensive length summation, weak end-inhibition, and stronger attenuation at high speeds. Moreover, the emergence of direction selectivity was delayed in trials involving camouflaged bars, relative to solid bars. Comparison with results obtained in the first (V1) and second (V2) visual areas, using similar stimuli, indicates that neural computations performed in MT result in significantly stronger and more accurate signals about camouflaged objects, particularly in situations in which these are relatively large and slow moving. These computations are likely to represent an important step in enabling cue-invariant perception of moving objects, particularly in biologically relevant situations.     

A double dissociation between striate and extrastriate visual cortex for pattern motion perception revealed using rTMS

The neural mechanisms underlying the integration and segregation of motion signals are often studied using plaid stimuli. These stimuli consist of two spatially coincident dynamic gratings of differing orientations, which are either perceived to move in two unique directions or are integrated by the visual system to elicit the percept of a checkerboard moving in a single direction. Computations pertaining to the motion of the individual component gratings are thought to take place in striate cortex (V1) whereas motion integration is thought to involve neurons in dorsal stream extrastriate visual areas, particularly V5/MT. By combining a psychophysical task that employed plaid stimuli with 1 Hz offline repetitive transcranial magnetic stimulation (rTMS), we demonstrated a double dissociation between striate and extrastriate visual cortex in terms of their contributions to motion integration. rTMS over striate cortex increased coherent motion percepts whereas rTMS over extrastriate cortex had the opposite effect. These effects were robust directly after the stimulation administration and gradually returned to baseline within 15 minutes. This double dissociation is consistent with previous patient data and the recent hypothesis that both coherent and transparent motion percepts are supported by the visual system simultaneously and compete for perceptual dominance. Hum Brain Mapp 2009. © 2009 Wiley‐Liss, Inc.     

Visual detection of motion speed in humans: spatiotemporal analysis by fMRI and MEG

Humans take a long time to respond to the slow visual motion of an object. It is not known what neural mechanism causes this delay. We measured magnetoencephalographic neural responses to light spot motion onset within a wide speed range (0.4-500 degrees /sec) and compared these with human reaction times (RTs). The mean response latency was inversely related to the speed of motion up to 100 degrees /sec, whereas the amplitude increased with the speed. The response property at the speed of 500 degrees /sec was different from that at the other speeds. The speed-related latency change was observed when the motion duration was 10 msec or longer in the speed range between 5 and 500 degrees /sec, indicating that the response is directly related to the speed itself. The source of the response was estimated to be around the human MT+ and was validated by functional magnetic imaging study using the same stimuli. The results indicate that the speed of motion is encoded in the neural activity of MT+ and that it can be detected within 10 msec of motion observation. RT to the same motion onset was also inversely related to the speed of motion but the delay could not be explained by the magnetic response latency change. Instead, the reciprocal of RT was linearly related to the reciprocal of the magnetic response latency, suggesting that the visual process interacts with other neural processes for decision and motor preparation.     

Similarities and differences in motion processing between the human and macaque brain: evidence from fMRI

The present report reviews a series of functional magnetic resonance imaging (fMRI) activation studies conducted in parallel in awake monkeys and humans using the same motion stimuli in both species. These studies reveal that motion stimuli engage largely similar cortical regions in the two species. These common regions include MT/V5 and its satellites, of which FST contributes more to the human motion complex than is generally assumed in human imaging. These results also establish a direct link between selectivity of MT/V5 neurons for speed gradients and functional activation of human MT/V5 by three-dimensional (3D) structure from motion stimuli. On the other hand, striking functional differences also emerged: in humans V3A and several regions in the intraparietal sulcus (IPS) are much more motion sensitive than their simian counterparts.     

Response latencies of neurons in visual areas MT and MST of monkeys with striate cortex lesions

Cortical area, MT (middle temporal area) is specialized for the visual analysis of stimulus motion in the brain. It has been suggested [Brain 118 (1995) 1375] that motion signals reach area MT via two dissociable routes, namely a 'direct' route which bypasses primary visual cortex (area, striate cortex (V1)) and is specialized for processing 'fast' motion (defined as faster than 6 degrees/s) with a relatively short latency, and an 'indirect' route via area V1 for processing 'slow' motion (slower than 6 degrees/s) with a relatively long latency. We tested this proposal by measuring the effects of unilateral V1 lesions on the magnitudes and latencies of responses to fast- and slow-motion (depicted by random dot kinematograms (RDK) ) of single neurons in areas MT and medial superior temporal area (MST) of anaesthetized macaque monkeys. In the unlesioned hemisphere contralateral to a V1 lesion, response magnitudes and latencies of MT neurons were similar to those previously reported from MT neurons in normal monkeys, and there was no significant association between slow movement and long response latency (>100 ms), or between fast movement and short latency (< or =100 ms). V1 lesions led to diminished response magnitudes and increased latencies in area MT of the lesioned hemisphere, but did not selectively abolish MT responses to slow moving stimuli, or abolish long-latency responses to either slow- or fast-moving stimuli. Response magnitudes and latencies in area MST, which receives visual inputs directly from area MT and is also specialized for visual analysis of motion, were unaffected by V1 lesions (though we have shown elsewhere that directionally-selective responses in both areas were impaired by V1 lesions). Overall, the results are incompatible with the hypothesis that there are dissociable routes to MT specialized for processing separately fast and slow motion.     

Common (and multiple) neural substrates for static and dynamic motion after-effects: A rTMS investigation

Introduction

Prolonged exposure to directional motion (adaptation) biases the perceived direction of subsequently presented test stimuli towards the opposite direction with respect to that of adaptation (i.e., motion after-effect; MAE). Different neural populations seem to be involved in the generation of the MAE, depending on the spatiotemporal characteristics of both adapting and test stimuli. Although the tuning mechanisms of the neural populations involved in the MAE have been psychophysically identified, the specific loci along the motion processing hierarchy where the different types of MAE take place is still debated.

Method

In this study, by using repetitive transcranial magnetic stimulation (rTMS) delivered during the inter-stimulus interval (ISI) between adapting and test patterns, we investigated the cortical locus of processing of static MAE (sMAE) and dynamic MAE (dMAE).

Results

Results showed that rTMS over V2/V3 or V5/MT decreased the perceived duration of both sMAE and dMAE, although rTMS over V2/V3 decreased mainly the perceived duration of sMAE.

Conclusions

sMAE and dMAE rely on the same cortical structures present at intermediate and low-levels of motion processing, although low-level visual areas (e.g., V2/V3) show a prevalence of neurons responsible for sMAE.     

Visual trajectory perception in humans: is it lateralized? Clues from online rTMS of the middle-temporal complex (MT/V5)

Inconsistent observations have been reported in the literature regarding the asymmetrical contribution of higher visual areas of the left and right hemispheres to visual motion processing. In the present experiment, we tested for hemispheric asymmetry of the middle-temporal complex (V5/MT), which is a key-component of the visual motion network, by using rTMS applied over left or right V5/MT during a visual trajectory perception task. The results showed that the effect of rTMS was to enhance individual hemispheric asymmetries present when the test was performed without rTMS. The more general meaning of these results is that there are robust individual hemispheric asymmetries in motion perception but no general pattern of hemispheric differences.     

The constructive nature of vision: direct evidence from functional magnetic resonance imaging studies of apparent motion and motion imagery

Echoplanar functional magnetic resonance imaging was used to monitor activation changes of brain areas while subjects viewed apparent motion stimuli and while they were engaged in motion imagery. Human cortical areas MT (V5) and MST were the first areas of the ‘dorsal’ processing stream which responded with a clear increase in signal intensity to apparent motion stimuli as compared with flickering control conditions. Apparent motion of figures defined by illusory contours evoked greater activation in V2 and MT/MST than appropriate control conditions. Several areas of the dorsal pathway (V3A, MT/MST, areas in the inferior and superior parietal lobule) as well as prefrontal areas including FEF and BA 9/46 responded strongly when subjects merely imagined moving stimuli which they had seen several seconds before. The activation during motion imagery increased with the synaptic distance of an area from V1 along the dorsal processing stream. Area MT/MST was selectively activated during motion imagery but not during a static imagery control condition. The comparison between the results obtained with objective motion, apparent motion and imagined motion provides further insights into a complex cortical network of motion-sensitive areas driven by bottom-up and top-down neural processes.     

Contribution of color signals to ocular following responses

Ocular following responses (OFRs) are elicited at ultra‐short latencies (< 60 ms) by sudden movements of the visual scene. In this study, we investigated the roles of color signals in OFRs in monkeys. To make physiologically isoluminant sinusoidal color gratings, we estimated the physiologically isoluminant points using OFRs and found that the physiologically isoluminant points were nearly independent of the spatiotemporal frequency of the gratings. We recorded OFRs induced by the motion of physiologically isoluminant color gratings and found that OFRs elicited by the motion of color gratings had different spatiotemporal frequency tuning from those elicited by the motion of luminance gratings. Additionally, OFRs to isoluminant color gratings had smaller peak responses, suggesting that color signals weakly contribute to OFRs compared with luminance signals. OFRs to the motion of stimuli composed of luminance and color signals were also examined. We found that color signals largely contributed to OFRs under low luminance signals regardless of whether color signals moved in the same or opposite direction to luminance signals. These results provide evidence of the multichannel visual computations underlying motor responses. We conclude that, in everyday situations, color information contributes cooperatively with luminance information to the generation of ocular tracking behaviors.     

Convergence of parvocellular and magnocellular information channels in the primary visual cortex of the macaque

We investigated whether responses of single cells in the striate cortex of anaesthetized macaque monkeys exhibit signatures of both parvocellular (P) and magnocellular (M) inputs from the dorsal lateral geniculate nucleus (dLGN). We used a palette of 128 isoluminant hues at four different saturation levels to test responses to chromatic stimuli against a white background. Spectral selectivity with these isoluminant stimuli was taken as an indication of P inputs. The presence of magnocellular inputs to a given cortical cell was deduced from its responses to a battery of tests, including assessment of achromatic contrast sensitivity, relative strengths of chromatic and luminance borders in driving the cell at different velocities and conduction velocity of their retino-geniculo-cortical afferents. At least a quarter of the cells in our cortical sample appear to receive convergent P and M inputs. We cannot however, exclude the possibility that some of these cells could be receiving a convergent input from the third parallel channel from the dLGN, namely the koniocellular (K) rather than the P channel. The neurons with convergent P and M inputs were recorded not only from supragranular and infragranular layers but also from the principal geniculate input recipient layer 4. Thus, our results challenge classical ideas of strict parallelism between different information streams at the level of the primate striate cortex.     

Detailed spatiotemporal brain mapping of chromatic vision combining high‐resolution VEP with fMRI and retinotopy

Neuroimaging studies have identified so far, several color‐sensitive visual areas in the human brain, and the temporal dynamics of these activities have been separately investigated using the visual‐evoked potentials (VEPs). In the present study, we combined electrophysiological and neuroimaging methods to determine a detailed spatiotemporal profile of chromatic VEP and to localize its neural generators. The accuracy of the present co‐registration study was obtained by combining standard fMRI data with retinotopic and motion mapping data at the individual level. We found a sequence of occipito activities more complex than that typically reported for chromatic VEPs, including feed‐forward and reentrant feedback. Results showed that chromatic human perception arises by the combined activity of at the least five parieto‐occipital areas including V1, LOC, V8/VO, and the motion‐sensitive dorsal region MT+. However, the contribution of V1 and V8/VO seems dominant because the re‐entrant activity in these areas was present more than once (twice in V8/VO and thrice in V1). This feedforward and feedback chromatic processing appears delayed compared with the luminance processing. Associating VEPs and neuroimaging measures, we showed for the first time a complex spatiotemporal pattern of activity, confirming that chromatic stimuli produce intricate interactions of many different brain dorsal and ventral areas.     

Neural basis of photo/chromatic sensitivity in adolescence

Purpose: To determine a psychophysiological basis for age visual sensitivity to chromatic and achromatic stimuli. Methods: We investigated the effects of achromatic and four isoluminant color combinations (blue/red, blue/green, green/red, and blue/yellow), luminance ratio changes in color combinations (blue/red; 1:1, 3:4, 4:3) and contrast changes (3 to 100%) on steady‐state electroretinograms (ERGs) and visual evoked potentials (VEPs) in 32 healthy teenagers and 30 young adults. Results: We found that (1) dual peaks at 9 and 18 Hz with a dip at 12 Hz were observed in VEPs with all isoluminant color combinations, (2) VEP responses were significantly enhanced and the 12‐Hz dip became unclear with luminance ratio changes between two colors with a nonantagonistic relationship (blue/red), and (3) VEP amplitudes were significantly increased when the contrast was increased. These characteristics were more evident in teenagers than young adults; however, ERGs were qualitatively similar between the two groups. Discussion: The visual cortex is differently modulated by different color‐luminance combinations, and higher sensitivity to color‐luminance combinations in the visual cortex in teenagers is responsible for the high prevalence of photo/chromatic sensitivity in adolescence.     

Effects of repetitive transcranial magnetic stimulation on voice and speech in Parkinson's disease

OBJECTIVE: To investigate the effects of repetitive transcranial magnetic stimulation (rTMS) on vocal function in Parkinson's disease (PD). MATERIAL AND METHODS: Two different sets of rTMS parameters were investigated on 30 patients with PD: active or sham 15 Hz rTMS of the left dorsolateral prefrontal cortex (LDLPFC) (110% of motor threshold (MT), 3000 pulses per session) and active 5 Hz rTMS of the primary motor cortex (M1)-mouth area (90% MT, 2250 pulses per session). A blind rater evaluated speech characteristics (acoustic and perceptual analysis of voice) and voice-related quality of life (V-RQOL). RESULTS: rTMS of LDLPFC resulted in mood amelioration and subjective improvement of the V-RQOL only (71.9% improvement, P < 0.001), but not in objective measures such as fundamental frequency (P = 0.86) and voice intensity (P = 0.99). On the other hand, rTMS of M1-mouth induced a significant improvement of the fundamental frequency (12.9% for men and 7.6% for women, P < 0.0001) and voice intensity (20.6%, P < 0.0001). CONCLUSIONS: Our findings provide initial evidence that rTMS of the primary motor cortex might yield a beneficial effect on vocal function in PD.     

A lesion of cortical area V2 selectively impairs the perception of the direction of first-order visual motion

Lesions of area MT/V5 in monkeys and its presumed homologue, the motion area, in humans impair motion perception, including the discrimination of the direction of global motion in random dot kinematograms. Here we report the results of similar tests on patient TF, who has a discrete and very small, unilateral infarct in the medial superior part of the right occipital cortex. Structural MRI, co-registered in software with a standardized human brain atlas, reveals that the lesion involves area V2. The patient was impaired in his retinotopically corresponding left lower quadrant on several motion tasks including discrimination in random dot kinematograms of direction, speed and motion-defined discontinuity. He was also impaired on tasks selectively involving first-order motion based on luminance contrast but not on second-order motion based on texture contrast. The results show that even though area MT/V5 is intact, motion perception is abnormal and, in particular, his perception of first-order motion is impaired.     

经颅重频磁刺激对大鼠脑缺血再灌流损伤早期运动皮层兴奋性和神经功能的影响

目的 观察经颅重频磁刺激（rTMS）对大鼠脑缺血再灌流损伤早期运动皮层兴奋性和神经功能的影响。方法测定Wistar大鼠右后肢运动阚值（MT）．制作左侧大脑中动脉栓塞（MCAO）再灌流模型．给予rTMS（1次／d）。再灌流72h处死取脑。比较大鼠MCAO再灌流损伤不同时间MT、神经功能评分，脑梗死体积的变化及rTMS的影响。结果MCAO再灌流损伤使大鼠出现局灶性梗死灶，MT升高；神经功能障碍且其程度随损伤时间延长而愈加明显．表现为功能评分升高；rTMS可改善大鼠MCAO再灌流损伤72h的MT和功能障碍程度。减小梗死体积。尤其改善神经功能障碍的程度与对照组比较差异有显著性（P=0．004）。结论rTMS可能对早期缺血脑组织有保护作用．对缺血性脑卒中有进一步研究、应用前景。