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R A Andersen

Publications and source records attributed to R A Andersen.

At least 73 records · Page 4Linked to original sources

The analysis of complex motion patterns by form/cue invariant MSTd neurons.

Several groups have proposed that area MSTd of the macaque monkey has a role in processing optical flow information used in the analysis of self motion, based on its neurons' selectivity for large-field motion patterns such as expansion, contraction, and rotation. It has also been suggested that this cortical region may be important in analyzing the complex motions of objects. More generally, MSTd could be involved in the generic function of complex motion pattern representation, with its cells responsible for integrating local motion signals sent forward from area MT into a more unified representation. If MSTd is extracting generic motion pattern signals, it would be important that the preferred tuning of MSTd neurons not depend on the particular features and cues that allow these motions to be represented. To test this idea, we examined the diversity of stimulus features and cues over which MSTd cells can extract information about motion patterns such as expansion, contraction, rotation, and spirals. The different classes of stimuli included: coherently moving random dot patterns, solid squares, outlines of squares, a square aperture moving in front of an underlying stationary pattern of random dots, a square composed entirely of flicker, and a square of nonFourier motion. When a unit was tuned with respect to motion pattern producing the most vigorous response in a neuron was nearly the same for each class. Although preferred tuning was invariant, the magnitude and width of the tuning curves often varied between classes. Thus, MSTd is form/cue invariant for complex motions, making it an appropriate candidate for analysis of object motion as well as motion introduced by observer translation.

Animals↗

Electrical microstimulation suggests two different forms of representation of head-centered space in the intraparietal sulcus of rhesus monkeys.

We examined the effects of eye position on saccades evoked by electrical stimulation of the intraparietal sulcus (IPS) of rhesus monkeys. Microstimulation evoked saccades from sites on the posterior bank, floor, and the medial bank of the IPS. The size and direction of the eye movements varied as a function of initial eye position before microstimulation. At many stimulation sites, eye position affected primarily the amplitude and not the direction of the evoked saccades. These "modified vector saccades" were characteristic of most stimulation-sensitive zones in the IPS, with the exception of a narrow strip located mainly on the floor of the sulcus. Stimulation in this "intercalated zone" evoked saccades that moved the eyes into a particular region in head-centered space, independent of the starting position of the eyes. This latter response is compatible with the stimulation site representing a goal zone in head-centered coordinates. On the other hand, the modified vector saccades observed outside the intercalated zone are indicative of a more distributed representation of head-centered space. A convergent projection from many modified vector sites onto each intercalated site may be a basis for a transition from a distributed to a more explicit representation of space in head-centered coordinates.

Animals↗

The impact of alteration of polyunsaturated fatty acid levels on C6-aldehyde formation of Arabidopsis thaliana leaves.

C6-aldehydes are synthesized via lipoxygenase/hydroperoxide lyase action on polyunsaturated fatty acid (PUFA) substrates in plant leaves. The source pools and subcellular location of the processes are unknown. A close relationship is found between the composition of PUFA and the composition of C6-aldehydes. In the current study, this relationship was tested using the Arabidopsis PUFA mutant lines act1, fad2, fad3, fad5, fad6, and fad7. The results indicate that C6-aldehyde formation is influenced by the alteration of C18 PUFA levels. Mutants act1 and fad5, which are deficient in C16 unsaturated fatty acids, had wild-type levels of C6-aldehyde production. Mutants deficient in the chloroplast hexadecenoic acid/oleic acid desaturase (fad6) or hexadecadienoic acid/linoleic acid desaturase (fad7) had altered C6-aldehyde formation in a pattern similar to the changes in the PUFA. Mutations that impair phosphatidylcholine desaturase activity, such as fad2 and fad3, however, resulted in increased E-2-hexenal formation. The enzymes involved in C6-aldehyde production were partially characterized, including measurement of pH optima. The differences in C6-aldehyde formation among the fatty acid mutants of Arabidopsis appeared not to result from alteration of lipoxygenase/hydroperoxide lyase pathway enzymes. Investigation of the fatty acid composition in leaf phospholipids, glycolipids, and neutral lipids and analysis of the fatty acid composition of chloroplast and extrachloroplast lipids indicate that chloroplasts and glycolipids of chloroplasts may be the source or major source of C6-aldehyde formation in Arabidopsis leaves.

Aldehydes↗

Spatially tuned auditory responses in area LIP of macaques performing delayed memory saccades to acoustic targets.

1. The lateral intraparietal area (area LIP) of the macaque's posterior parietal cortex (PPC) lies in the dorsal stream of extrastriate visual areas. It receives extensive visual inputs and sends outputs to several eye movement centers. It contains neurons with visual and saccade-related responses suggesting a role of area LIP in programming saccadic eye movements to visual targets. Because primates can also orient to nonvisual stimuli, we investigated whether LIP neurons process stimuli of other modalities besides the visual one by comparing their activity in auditory and visual saccade tasks. 2. We recorded the activity of single neurons of Macaca mulatta monkeys while they performed memory saccades to acoustic and visual targets. We analyzed the activity during stimulus presentation (stimulus period, S) and during the delay (memory period, M) between stimulus presentation and the saccade to its remembered location. 3. Among 80 area LIP neurons tested, we found 44 that had S period and/or M period responses following presentation of the auditory stimulus. Most of these responses were spatially tuned, i.e., selective for the left or right stimulus location (27 of 29 S responses; 25 of 29 M responses). 4. The majority of neurons with responses in the auditory memory saccade task also responded in the visual version of the task. Eighty-nine percent (24/27) were clearly bimodal in the S period, and 88% (23/26) were bimodal in the M period. 5. Almost all the neurons with spatially tuned auditory responses that were bimodal were also spatially tuned in their visual responses (20/22 for S responses; 18/19 for M responses). The spatial tuning for the two modalities was the same in 85% (17/20) of the tested neurons for the S responses, and in 83% (15/18) of the tested neurons for the M responses. 6. Area LIP contains a population of neurons that respond to both visual and auditory stimuli. This result is consistent with our finding that the memory activity of many LIP cells encodes the next planned saccade. If cells are coding planned movements, they should be active independently of the sensory modality of the target for the movement, as was the case for most of the neurons described in the present study.

Acoustic Stimulation↗

Motor intention activity in the macaque's lateral intraparietal area. I. Dissociation of motor plan from sensory memory.

1. The lateral intraparietal area (area LIP) of the monkey's posterior parietal cortex (PPC) contains neurons that are active during saccadic eye movements. These neurons' activity includes visual and saccade-related components. These responses are spatially tuned and the location of a neuron's visual receptive field (RF) relative to the fovea generally overlaps its preferred saccade amplitude and direction (i.e., its motor field, MF). When a delay is imposed between the presentation of a visual stimulus and a saccade made to its location (memory saccade task), many LIP neurons maintain elevated activity during the delay (memory activity, M), which appears to encode the metrics of the next intended saccadic eye movements. Recent studies have alternatively suggested that LIP neurons encode the locations of visual stimuli regardless of where the animal intends to look. We examined whether the M activity of LIP neurons specifically encodes movement intention or the locations of recent visual stimuli, or a combination of both. In the accompanying study, we investigated whether the intended-movement activity reflects changes in motor plan. 2. We trained monkeys (Macaca mulatta) to memorize the locations of two visual stimuli and plan a sequence of two saccades, one to each remembered target, as we recorded the activity of single LIP neurons. Two targets were flashed briefly while the monkey maintained fixation; after a delay the fixation point was extinguished, and the monkey made two saccades in sequence to each target's remembered location, in the order in which the targets were presented. This "delayed double saccade" (DDS) paradigm allowed us to dissociate the location of visual stimulation from the direction of the planned saccade and thus distinguish neuronal activity related to the target's location from activity related to the saccade plan. By imposing a delay, we eliminated the confounding effect of any phasic responses coincident with the appearance of the stimulus and with the saccade. 3. We arranged the two visual stimuli so that in one set of conditions at least the first one was in the neuron's visual RF, and thus the first saccade was in the neuron's motor field (MF). M activity should be high in these conditions according to both the sensory memory and motor plan hypotheses. In another set of conditions, the second stimulus appeared in the RF but the first one was presented outside the RF, instructing the monkey to plan the first saccade away from the neuron's MF. If the M activity encodes the motor plan, it should be low in these conditions, reflecting the plan for the first saccade (away from the MF). If it is a sensory trace of the stimulus' location, it should be high, reflecting stimulation of the RF by the second target. 4. We tested 49 LIP neurons (in 3 hemispheres of 2 monkeys) with M activity on the DDS task. Of these, 38 (77%) had M activity related to the next intended saccade. They were active in the delay period, as expected, if the first saccade was in their preferred direction. They were less active or silent if the next saccade was not in their preferred direction, even when the second stimulus appeared in their RF. 5. The M activity of 8 (16%) of the remaining neurons specifically encoded the location of the most recent visual stimulus. Their firing rate during the delay reflected stimulation of the RF independently of the saccade being planned. The remaining 3 neurons had M activity that did not consistently encode either the next saccade or the stimulus' location. 6. We also recorded the activity of a subset of neurons (n = 38) in a condition in which no stimulus appeared in a neuron's RF, but the second saccade was in the neuron's MF. In this case the majority of neurons tested (23/38, 60%) became active in the period between the first and second saccade, even if neither stimulus had appeared in their RF. Moreover, this activity appeared only after the first saccade had started in all but two of

Animals↗

Motor intention activity in the macaque's lateral intraparietal area. II. Changes of motor plan.

1. In the companion paper we reported that the predominant signal of the population of neurons in the lateral intraparietal area (area LIP) of the monkey's posterior parietal cortex (PPC) encode the next intended saccadic eye movement during the delay period of a memory-saccade task. This result predicts that, should be monkey change his intention of what the next saccade will be, LIP activity should change accordingly to reflect the new plan. We tested this prediction by training monkeys to change their saccadic plan on command and recording the activity of LIP neurons across plan changes. 2. We trained rhesus monkeys (Macaca mulatta) to maintain fixation on a light spot as long as this spot remained on. During this period we briefly presented one, two, or three peripheral visual stimuli in sequence, each followed by a delay (memory period, M). After the final delay the fixation spot was extinguished, and the monkey had to quickly make a saccade to the location of the last target to have appeared. The monkey could not predict which stimuli, nor how many, would appear on each trial. He thus had to plan a saccade to each stimulus as it appeared and change his saccade plan whenever a stimulus appeared at a different location. 3. We recorded the M period activity of 81 area LIP neurons (from 3 hemispheres of 2 monkeys) in this task. We predicted that, if a neuron's activity reflected the monkey's planned saccade, its activity should be high while the monkey planned a saccade in the neuron's motor field (MF), and low while the planned saccade was in the opposite direction. The activity of most of the neurons in our sample changed in accordance with our hypothesis as the monkey's planned saccade changed. 4. In one condition the monkey was instructed by visual stimuli to change his plan from a saccade in the neuron's preferred direction to a saccade planned in the opposite direction. In this condition activity decreased significantly (P < 0.05) in 65 (80%) of 81 neurons tested. These neurons' activity changed to reflect the new saccade plan even though the cue for this change was not presented in their RF. 5. As a control we randomly interleaved, among trials requiring a plan change, trials in which the monkey had to formulate two consecutive plans to make a saccade in the neuron's preferred direction. The activity remained unchanged (P < 0.05) in 22 of 31 neurons tested (79%), indicating that the neurons continued to encode the same saccade plan. 6. In a variant of the task, the cue to the location of the required saccade was either a light spot or a noise burst from a loudspeaker. Of 22 neurons tested in this task, 16 (73%) showed activity changes consistent with plan changes cued by visual or auditory stimuli. 7. Alterations in the monkey's intentions, even in the absence of overt behavior, are manifested in altered LIP activity. These activity changes could be induced whether visual or auditory cues were used to indicate the required plan changes. Most LIP neurons thus do not encode only the locations of visual stimuli, but also the intention to direct gaze to specific locations, independently of whether a gaze shift actually occurs.

Acoustic Stimulation↗

Eye-centered, head-centered, and intermediate coding of remembered sound locations in area LIP.

1. The lateral intraparietal area (LIP) of the posterior parietal cortex lies within the dorsal cortical stream for spatial vision and processes visual information to plan saccadic eye movements. We investigated how LIP neurons respond when a monkey makes saccades to the remembered location of sound sources in the absence of visual stimulation. 2. Forty-three (36%) of the 118 neurons sampled showed significant auditory triggered activity during the memory period. This figure is similar to the proportion of cells showing visually triggered memory activity. 3. Of the cells showing auditory memory activity, 44% discharged in an eye-centered manner, similar to the way in which LIP cells discharge for visually initiated saccades. Another 33% responded in head-centered coordinates, and the remaining 23% had responses intermediate between the two reference frames. 4. For a substantial number of cells in all three categories, the magnitude of the response was modulated by eye position. Similar orbital "gain fields" had been shown previously for visual saccades. 5. We propose that area LIP is either at the origin of, or participates in, the transformation of auditory signals for oculomotor purposes, and that orbital gains on the discharge are part of this process. 6. Finally, we suggest that, by the level of area LIP, cells are concerned with the abstract quality of where a stimulus is in space, independent of the exact nature of the stimulus.

Acoustic Stimulation↗

Head position signals used by parietal neurons to encode locations of visual stimuli.

The mechanism for object location in the environment, and the perception of the external world as stable when eyes, head and body are moved, have long been thought to be centred on the posterior parietal cortex. However, head position signals, and their integration with visual and eye position signals to form a representation of space referenced to the body, have never been examined in any area of the cortex. Here we show that the visual and saccadic activities of parietal neurons are strongly affected by head position. The eye and head position effects are equivalent for individual neurons, indicating that the modulation is a function of gaze direction, regardless of whether the eyes or head are used to direct gaze. These data are consistent with the idea that the posterior parietal cortex contains a distributed representation of space in body-centred coordinates.

Animals↗

Accumulation of metallothionein and its multiple forms by zinc, cadmium and dexamethasone in human peripheral T and B lymphocytes and monocytes.

Metallothionein (MT) is produced at high rates in isolated monocytes, and T and B lymphocytes during induction in vitro. At optimal concentrations, 125 microM for Zn and 10 microM for Cd and dexamethasone (dex), MT was demonstrated after only 2 h in some cases, and in all cell types substantial levels were measured after 1 day of exposure to all three inductors. With Cd, lower amounts of MT were found, but maximum levels were reached faster than with Zn. The same result was found for dex compared to Zn. Zn and dex in combination showed the same accumulation rate as Zn alone. If the inductors were used in lower concentrations than optimal, reduced accumulation rates occurred, particularly during the first part of the exposure period. No MT was found for concentrations below 5 microM Zn, 1 microM Cd or 0.5 microM dex. The constitutive levels of MT (mean +/- S.E.M.) were 0.11 +/- 0.05, 0.54 +/- 0.3, 0.06 +/- 0.05 and 0.15 +/- 0.08 nmol Cd bound/5 x 10(6) unseparated mononuclear cells (MNC), monocytes, T lymphocytes and B lymphocytes, respectively. Monocytes accumulated 19 times and B lymphocytes 6 times more MT than T lymphocytes after 2 days of exposure to 125 microM Zn. Despite these differences in accumulated amounts of MT, the fold accumulation values were rather similar between the cell types, reflecting corresponding variations in background MT levels. After exposure of unseparated MNC to 125 microM Zn for 2 days, removal of the metal caused constitutive MT levels to be reestablished in 5 days. Five different MT forms, all capable of Cd complexation, were demonstrated in these cells. These forms had kinetically different behaviour during Zn exposure among the cell types, and the response to Cd was different from the Zn response. The results indicate metals to be closely controlled in MNC and emphasize a role for multiple MT forms in the process of regulation.

B-Lymphocytes↗

Integration of motion and stereopsis in middle temporal cortical area of macaques.

The primate visual system incorporates a highly specialized subsystem for the analysis of motion in the visual field. A key element of this subsystem is the middle temporal (MT) cortical area, which contains a majority of direction-selective neurons. MT neurons are also selective for binocular disparity (depth), which is perplexing given that they are not sensitive to motion through depth. What is the role of disparity in MT? Our data suggest an important link between disparity and transparent motion detection. Motion signals in different directions tend to inhibit each other within a given MT receptive field. This inhibition has an averaging effect which minimizes MT responses to random motion signals created by light intensity changes and other non-motion stimuli (motion noise). But, in the absence of disparity cues, inhibition may also occur between surfaces moving in different directions through the same part of the visual field (transparent motion), thus impairing the detection of either surface. Here we show that inhibition in MT occurs mainly between motion signals with similar disparities. Transparent surface movements at different depths are thus represented independently in MT (that is, without inhibiting each other) whereas spurious motion signals from a given surface tend to cancel out. To our knowledge, these results provide the first evidence for a functional integration of motion and disparity in MT.

Animals↗

Cladistic analyses of combined traditional and molecular data sets reveal an algal lineage.

The chromophyte algae are a large and biologically diverse assemblage of brown seaweeds, diatoms, and other golden algae classified in 13 taxonomic classes. One subgroup (diatoms, pedinellids, pelagophytes, silicoflagellates, and certain enigmatic genera) is characterized by a highly reduced flagellar apparatus. The flagellar apparatus lacks microtubular and fibrous roots, and the flagellum basal body is attached directly to the nucleus. We hypothesize that the flagellar reduction is the result of a single evolutionary series of events. Cladistic analysis of ultrastructural and biochemical data reveals a monophyletic group that unites all taxa with a reduced flagellar apparatus, supporting our hypothesis. Phylogenetic analyses of 18S rRNA gene sequence data provide strong resolution within most of the major groups of chromophytes but only weakly resolve relationships among those groups. Some of the molecularly based most parsimonious trees, however, also unite the taxa with a reduced flagellar apparatus, although the diatoms are not included in this lineage. This grouping is further supported by a posteriori character weighting of the molecular data, suggesting that flagellar reduction occurred at least twice in parallel evolutionary series of events. To further test our hypothesis of a single evolutionary reduction in the flagellar apparatus, we combine the two data sets and subject the hybrid data matrix to parsimony analysis. The resulting trees unite the diatoms with the other reduced flagellar apparatus algae in a monophyletic group. This result supports our hypothesis of a single evolutionary reduction and indicates the existence of a previously unrecognized lineage of algae characterized by a highly reduced flagellar apparatus. Further, this study suggests that the traditional classification of the diatoms with the chrysophytes and xanthophytes in the division (= phylum) Chrysophyta, as presented in most textbooks, is unsatisfactory and that a significantly different classification should be employed.

Base Sequence↗

V1 responses to transparent and nontransparent motions.

It is well known that a stimulus composed of two independent sets of random dots moving in opposite directions produces a percept of two overlapping transparent surfaces moving across each other, while a counterphase grating composed of two identical sine wave gratings drifting in opposite directions does not. We recorded from the directionally selective V1 cells of behaving macaque monkeys using these two types of stimuli in order to investigate the physiological basis of transparent motion perception. Previous single-unit recording experiments from our laboratory indicated that many V1 cells respond well to transparent random dot patterns, while MT cells' responses to the same patterns are strongly suppressed in comparison with their preferred direction responses. This observation alone would seem to suggest that V1 activity could better explain transparent motion perception than MT activity. However, one could argue to the contrary based on the psychophysical observation that there is a motion threshold elevation under the transparency condition. We decided to determine the correlation between V1 activity and the transparent motion perception directly by recording from V1 cells using both transparent random dot patterns and nontransparent counterphase gratings. It is found that V1 cells on the average could not reliably tell the two types of patterns apart. Our results further the idea that additional processing beyond V1 is involved in transparent motion analysis.

Animals↗

Recovering three-dimensional structure from motion with surface reconstruction.

This paper addresses the computational role that the construction of a complete surface representation may play in the recovery of 3-D structure from motion. We first discuss the need to integrate surface reconstruction with the structure-from-motion process, both on computational and perceptual grounds. We then present a model that combines a feature-based structure-from-motion algorithm with a smooth surface interpolation mechanism. This model allows multiple surfaces to be represented in a given viewing direction, incorporates constraints on surface structure from object boundaries, and segregates image features onto multiple surfaces on the basis of their 2-D image motion. We present the results of computer simulations that relate the qualitative behavior of this model to psychophysical observations. In a companion paper, we discuss further perceptual observations regarding the possible role of surface reconstruction in the human recovery of 3-D structure from motion.

Algorithms↗

Structure-from-motion: perceptual evidence for surface interpolation.

Dynamic random-dot displays representing a rotating cylinder were used to investigate surface interpolation in the perception of structure-from-motion (SFM) in humans. Surface interpolation refers to a process in which a complete surface in depth is reconstructed from the object depth values extracted at the stimulus features. Surface interpolation will assign depth values even in parts of the object that contain no features. Such a "fill-in" process should make the detection of featureless stimulus areas ("holes") difficult. Indeed, we demonstrate that such holes in our rotating cylinder can be as wide as one-quarter of the stimulus before subjects can reliably detect their presence. Subjects were presented with a variation on the rotating cylinder in which all dots were oscillating either in synchrony or asynchronously. Subjects perceive a rigidly rotating cylinder even when such a percept is not in agreement with the physical stimulus. To reconcile this discrepancy between actual and perceived stimulus we propose that individual points contribute to a surface based object representation and that in this process the visual system looses access to the identity of the individual features that make up the surface. Finally we are able to explain a variety of previously documented perceptual peculiarities in the perception of structure-from-motion by arguing that the perceptual interpretation of the object's boundaries influences the surface interpolation process. These findings offer strong perceptual evidence for a process of surface interpolation and are also physiologically plausible given results from recordings in awake behaving monkey cortical areas V1 and MT. The companion paper demonstrates how such a surface interpolation process can be incorporated into a structure-from-motion algorithm and how object boundaries can influence the perception of structure-from-motion as has been demonstrated before and in this paper.

Depth Perception↗

Cytotoxicity and accumulation of Hg, Ag, Cd, Cu, Pb and Zn in human peripheral T and B lymphocytes and monocytes in vitro.

1. The cytotoxic effects of various heavy metals were assayed by trypan blue exclusion in vitro in human peripheral immune cells separated to high purity. T and B lymphocytes and monocytes were equally sensitive to metals. The individual metals could be ranked in the following decreasing order of cytotoxic potency, Hg approximately Ag > Cd approximately Cu > Pb approximately Zn, based on exposure time and concentration needed to give a particular percentage of dead cells. 2. The cytotoxic effects became irreversible after about 13 hr of metal exposure. 3. Examination by scanning electron microscopy showed that the heavy metals caused serious destruction of the cell membranes. 4. Solubility and uptake of metals into the cells were studied and discussed in relation to the cytotoxic effects. It was concluded that metal binding to cell surfaces or precipitate formation could inhibit ordinary uptake, thereby affecting cytotoxicity. For Pb in monocytes this appeared to lead to uptake of non-toxic complexes, probably by phagocytosis.

Adult↗

Transparent motion perception as detection of unbalanced motion signals. I. Psychophysics.

Our visual system can solve the difficult problem of representing multiple motions in the same part of the visual space, the motion transparency problem. We investigated the conditions under which transparent motion perception occurs through psychophysical observations, using a series of visual displays composed of two simple patterns moving in opposite directions. We found that whenever a display has finely balanced opposing motion signals in all local regions, it is perceptually nontransparent. The displays that appeared transparent always contain locally unbalanced motion signals, with some local regions having net motion signals in one direction and some other regions in the opposite direction. These interdigitating net motion signals in both directions appear to be integrated separately to form two overlapping transparent surfaces. Displays that were spatially balanced could be made perceptually transparent if the two components moving in opposite directions were at different stereo depth planes or had different spatial frequency contents. Our results can be explained by proposing a disparity- and spatial frequency-specific suppression stage in the motion pathway, at which motion signals of different directions, but of the same disparity and spatial frequency contents, locally inhibit each other. Such a mechanism would suppress noise input to the motion system, which generally activates several direction channels simultaneously, and would still not eliminate activity evoked by transparent surfaces that are at different depths or have different textures.

Cues↗

Transparent motion perception as detection of unbalanced motion signals. II. Physiology.

We investigated how the primate visual system solves the difficult problem of representing multiple motion vectors in the same part of the visual space--the problem of motion transparency. In the preceding companion article we reported that displays with locally well-balanced motion signals in opposite directions are perceptually nontransparent (i.e., one does not see two coherent moving surfaces) and that transparent displays always contain locally unbalanced motion signals. This is exemplified by our paired and unpaired dot patterns. Although both types of stimuli contain two sets of dots moving in opposite directions, the former is locally well balanced and appears like flicker while the latter gives a perception of two transparent surfaces. In this article we report our physiological recordings from areas V1 and MT of behaving monkeys, comparing single-cell responses to the paired and the unpaired dot patterns. Although a small proportion of directionally selective V1 cells responded differently to the two types of patterns, the average V1 responses could not reliably distinguish between the paired and the unpaired stimuli. A large fraction of MT cells, on the other hand, responded significantly better to the unpaired dot patterns than to the paired ones. Furthermore, the average response of all MT cells to the unpaired dot patterns was significantly higher than that to the paired dot patterns. These results demonstrate a neural correlate of the perceptual transparency at the level of MT. On the other hand, V1 cells do not generally discriminate between the transparent and nontransparent stimuli, indicating that V1 activity is not well correlated with the perception of motion transparency. Our results are consistent with a two-stage model for motion processing: the first stage measures local motion and the second stage introduces suppression if different directions of motion are present at a local region of the visual field. The first stage is located primarily in V1 and the second stage primarily in MT. Finally, we found a strong and negative correlation between the degree of the opponent-direction suppression of MT cells and their responses to flicker noise stimuli. This result suggests that one of the fundamental roles of the opponent-direction suppression in MT is noise reduction.

Animals↗