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F Bremmer

Publications and source records attributed to F Bremmer.

At least 19 recordsLinked to original sources
PubMed2002

Visual receptive field modulation in the lateral intraparietal area during attentive fixation and free gaze.

The receptive field (RF) of neurons recorded from the lateral intraparietal area (LIP) was quantified using a rapid, computer-driven mapping procedure. For each neuron, the RF was mapped: (1) during attentive fixation and (2) during free visual exploration. RF location, size and internal structure were modulated by the mapping context in over two-thirds of the recorded neurons. The major trend was a proportionally larger amount of neuronal visual resources allocated to central space during fixation, and an attenuated center-to-periphery gradient in the visual field representation during free gaze. A population approach shows that these spatial modulations are accompanied by changes in the signal-to-noise ratio of the information carried in the RF substructure. We related these neurophysiological observations to behavior, by comparing the characteristics of saccades elicited during fixation and free gaze. Together, the results suggest that the dynamics of LIP visual RFs may characterize both the state of engagement of attention and the power of resolution of visual analysis: during fixation, the neural population is locked in a filter state concentrating the processing resources at the fovea, while during free gaze, the population shifts to a detector state spreading the resources more evenly across the visual field.

Animals↗
PubMed2002

Directional asymmetry of neurons in cortical areas MT and MST projecting to the NOT-DTN in macaques.

The cortical projection to the subcortical pathway underlying the optokinetic reflex was studied using antidromic electrical stimulation in the midbrain structures nucleus of the optic tract and dorsal terminal nucleus of the accessory optic system (NOT-DTN) while simultaneously recording from cortical neurons in the superior temporal sulcus (STS) of macaque monkeys. Projection neurons were found in all subregions of the middle temporal area (MT) as well as in the medial superior temporal area (MST). Antidromic latencies ranged from 0.9 to 6 ms with a median of 1.8 ms. There was a strong bias in the population of cortical neurons projecting to the NOT-DTN for ipsiversive stimulus movement (towards the recording side), whereas in the population of cortical neurons not projecting to the NOT-DTN a more or less equal distribution of stimulus directions was evident. Our data indicate that there is no special area in the posterior STS coding for ipsiversive horizontal stimulus movement. Instead, a specific selection of cortical neurons from areas MT and MST forms the projection to the NOT-DTN and as a subpopulation has the same directional bias as their subcortical target neurons. These findings are discussed in relation to the functional grouping of cortical output as an organizational principle for specific motor responses.

Animals↗
PubMed2001

The perception of inferred action.

Our actions, and those of others, are often partly obscured from view. This complicates the sensory inputs that guide motor actions. In this issue of Neuron, Umilità and colleagues demonstrate that "mirror neurons" in ventral premotor cortex respond when monkeys observe hidden, but inferred, actions.

Animals↗
PubMed2001

Space coding in primate posterior parietal cortex.

Neuropsychological studies of patients with lesions of right frontal (premotor) or posterior parietal cortex often show severe impairments of attentive sensorimotor behavior. Such patients frequently manifest symptoms like hemispatial neglect or extinction. Interestingly, these behavioral deficits occur across different sensory modalities and are often organized in head- or body-centered coordinates. These neuropsychological data provide evidence for the existence of a network of polymodal areas in (primate) premotor and inferior parietal cortex representing visual spatial information in a nonretinocentric frame of reference. In the monkey, a highly modular structural and functional specialization has been demonstrated especially within posterior parietal cortex. One such functionally specialized area is the ventral intraparietal area (VIP). This area is located in the fundus of the intraparietal sulcus and contains many neurons that show polymodal directionally selective discharges, i.e., these neurons respond to moving visual, tactile, vestibular, or auditory stimuli. Many of these neurons also encode sensory information from different modalities in a common, probably head-centered, frame of reference. Functional imaging data on humans reveal a network of cortical areas that respond to polymodal stimuli conveying motion information. One of these regions of activation is located in the depth of human intraparietal sulcus. Accordingly, it is suggested that this area constitutes the human equivalent of monkey area VIP. The functional role of area VIP for polymodal spatial perception in normals as well as the functional implications of lesions of area VIP in parietal patients needs to be established in further experiments.

Animals↗
PubMed2001

Representation of the visual field in the lateral intraparietal area of macaque monkeys: a quantitative receptive field analysis.

The representation of the visual field in the primate lateral intraparietal area (LIP) was examined, using a rapid, computer-driven receptive field (RF) mapping procedure. RF characteristics of single LIP neurons could thus be measured repeatedly under different behavioral conditions. Here we report data obtained using a standard ocular fixation task during which the animals were required to monitor small changes in color of the fixated target. In a first step, statistical analyses were conducted in order to establish the experimental limits of the mapping procedure on 171 LIP neurons recorded from three hemispheres of two macaque monkeys. The characteristics of the receptive fields of LIP neurons were analyzed at the single cell and at the population level. Although for many neurons the assumption of a simple two-dimensional gaussian profile with a central area of maximal excitability at the center and progressively decreasing response strength at the periphery can represent relatively accurately the spatial structure of the RF, about 19% of the cells had a markedly asymmetrical shape. At the population level, we observed, in agreement with prior studies, a systematic relation between RF size and eccentricity. However, we also found a more accentuated overrepresentation of the central visual field than had been previously reported and no marked differences between the upper and lower visual representation of space. This observation correlates with an extension of the definition of LIP from the posterior third of the lateral intraparietal sulcus to most of the middle and posterior thirds. Detailed histological analyses of the recorded hemispheres suggest that there exists, in this newly defined unitary functional cortical area, a coarse but systematic topographical organization in area LIP that supports the distinction between its dorsal and ventral regions, LIPd and LIPv, respectively. Paralleling the physiological data, the central visual field is mostly represented in the middle dorsal region and the visual periphery more ventral and posterior. An anteroposterior gradient from the lower to the upper visual field representations can also be identified. In conclusion, this study provides the basis for a reliable mapping method in awake monkeys and a reference for the organization of the properties of the visual space representation in an area LIP extended with respect to the previously described LIP and showing a relative emphasis of central visual field.

Animals↗
PubMed2001

Polymodal motion processing in posterior parietal and premotor cortex: a human fMRI study strongly implies equivalencies between humans and monkeys.

In monkeys, posterior parietal and premotor cortex play an important integrative role in polymodal motion processing. In contrast, our understanding of the convergence of senses in humans is only at its beginning. To test for equivalencies between macaque and human polymodal motion processing, we used functional MRI in normals while presenting moving visual, tactile, or auditory stimuli. Increased neural activity evoked by all three stimulus modalities was found in the depth of the intraparietal sulcus (IPS), ventral premotor, and lateral inferior postcentral cortex. The observed activations strongly suggest that polymodal motion processing in humans and monkeys is supported by equivalent areas. The activations in the depth of IPS imply that this area constitutes the human equivalent of macaque area VIP.

Acoustic Stimulation↗
PubMed2000

Eye position effects in macaque area V4.

Recent studies revealed an influence of eye position on neuronal discharges in many dorsal stream areas of the macaque visual cortical system. This eye position information is thought to serve an implicit non-retinocentric representation of visual spatial information. The question arises of whether the two visual cortical pathways encode information in a common coordinate system, i.e. whether eye position effects exist also in ventral stream areas. We recorded 112 neurons from area V4 of two awake monkeys. Of these, 55 (49%) showed eye position effects. Like in dorsal stream areas, this modulatory influence was balanced out at the population level. Our data support the view of eye position effects as a general phenomenon in the macaque visual cortical system.

Analysis of Variance↗
PubMed2000

Responses to continuously changing optic flow in area MST.

We studied the temporal behavior and tuning properties of medial superior temporal (MST) neurons in response to constant flow-field stimulation and continuously changing flow-field stimulation (transitions), which were obtained by morphing one flow field into another. During transitions, the flow fields resembled the motion pattern seen by an observer during changing ego-motion. Our aim was to explore the behavior of MST cells in response to changes in the flow-field pattern and to establish whether the responses of MST cells are temporally independent or if they are affected by contextual information from preceding stimulation. We first tested whether the responses obtained during transitions were linear with respect to the two stimuli defining the transition. In over half of the transitions, the cell response was nonlinear: the response during the transition could not be predicted by the linear interpolation between the stimulus before and after the transition. Nonlinearities in the responses could arise from a dependence on temporal context or from nonlinearities in the tuning to flow-field patterns. To distinguish between these two hypotheses, we fit the responses during transitions and during continuous stimuli to the predictions of a temporally independent model (temporal-independence test) and we compared the responses during transitions to the responses elicited by inverse transitions (temporal-symmetry test). The effect of temporal context was significant in only 7.2% and 5.5% of cells in the temporal-independence test and in the temporal-symmetry test, respectively. Most of the nonlinearities in the cell responses could be accounted for by nonlinearities in the tuning to flow-field stimuli (i.e., the responses to a restricted set of flow fields did not predict the responses to other flow fields). Tuning nonlinearities indicate that a complete characterization of the tuning properties of MST neurons cannot be obtained by testing only a small number of flow fields. Because the cells' responses do not depend on temporal context, continuously changing stimulation can be used to characterize the receptive field properties of cells more efficiently than constant stimulation. Temporal independence in the responses to transitions indicates that MST cells do not code for second-order temporal properties of flow-field stimuli, i.e., for changes in the flow field through time that can be construed as paths through the environment. Information about ego-motion three-dimensional paths through the environment may either be processed at the population level in MST or in other cortical areas.

Action Potentials↗
PubMed1999

Linear vestibular self-motion signals in monkey medial superior temporal area.

The present study was aimed at investigating the sensitivity to linear vestibular stimulation of neurons in the medial superior temporal area (MST) of the macaque monkey. Two monkeys were moved on a parallel swing while single-unit activity was recorded. About one-half of the cells (28/51) responded in the dark either to forward motion (n = 10), or to backward motion (n = 11), or to both (n = 7). Twenty cells responding to vestibular stimulation in darkness were also tested for their responses to optic flow stimulation simulating forward and backward self-motion. Forty-five percent (9/20) of them preferred the same self-motion directions, that is, combined visual and vestibular signals in a synergistic manner. Thirty percent (6/20) of the cells were not responsive to visual stimulation alone. The remaining 25% (5/20) preferred directions that were antialigned. Our results provide strong evidence that neurons in the MST area are at least in part involved in the processing of self-motion.

Animals↗
PubMed1999

Eye position encoding in the macaque ventral intraparietal area (VIP).

Many neurons in area VIP encode the location of visual stimuli in a non-retinocentric frame of reference. In this context the question needed to be addressed whether the underlying coordinate transformation of the incoming visual signals could be generated within area VIP or whether this information would have to arrive from other areas. We tested 74 neurons in area VIP of two awake monkeys for an influence of eye position while animals performed a fixation task. More than half of the neurons (40/74) revealed an eye position effect. At the population level, however, this effect was balanced out. We suggest that local connections within area VIP could be used to generate an encoding of visual information in a non-retinocentric frame of reference.

Animals↗
PubMed1999

The use of optical velocities for distance discrimination and reproduction during visually simulated self motion.

Successful navigation through an environment requires precise monitoring of direction and distance traveled ("path integration" or "dead reckoning"). Previous studies in blindfolded human subjects showed that velocity information arising from vestibular and somatosensory signals can be used to reproduce passive linear displacements. In these studies, visual information was excluded as sensory cue. Yet, in our everyday life, visual information is very important and usually dominates vestibular and somatosensory cues. In the present study, we investigated whether visual signals can be used to discriminate and reproduce simulated linear displacements. In a first set of experiments, subjects viewed two sequences of linear motion and were asked in a 2AFC task to judge whether the travel distance in the second sequence was larger or shorter than in the first. Displacements in either movement sequence could be forward (f) or backward (b). Subjects were very accurate in discriminating travel distances. Average error was less than 3% and did not depend on displacements being into the same (ff, bb) or opposite direction (fb, bf). In a second set of experiments, subjects had to reproduce a previously seen forward motion (passive condition), either in light or in darkness, i.e., with or without visual feedback. Passive displacements had different velocity profiles (constant, sinusoidal, complex) and speeds and were performed across a textured ground plane, a 2-D plane of dots or through a 3-D cloud of dots. With visual feedback, subjects reproduced distances accurately. Accuracy did not depend on the kind of velocity profile in the passive condition. Subjects tended to reproduce distance by replicating the velocity profile of the passive displacement. Finally, in the condition without visual feedback, subjects reproduced the shape of the velocity profile, but used much higher speeds, resulting in a substantial overshoot of travel distance. Our results show that visual, vestibular, and somatosensory signals are used for path integration, following a common strategy: the use of the velocity profile during self-motion.

Adult↗
PubMed1999

Gaze effects in the cerebral cortex: reference frames for space coding and action.

Visual information is mapped with respect to the retina within the early stages of the visual cortex. On the other hand, the brain has to achieve a representation of object location in a coordinate system that matches the reference frame used by the motor cortex to code reaching movement in space. The mechanism of the necessary coordinate transformation between the different frames of reference from the visual to the motor system as well as its localization within the cerebral cortex is still unclear. Coordinate transformation is traditionally described as a series of elementary computations along the visuomotor cortical pathways, and the motor system is thought to receive target information in a body-centered reference frame. However, neurons along these pathways have a number of similar properties and receive common input signals, suggesting that a non-retinocentric representation of object location in space might be available for sensory and motor purposes throughout the visuomotor pathway. This paper reviews recent findings showing that elementary input signals, such as retinal and eye position signals, reach the dorsal premotor cortex. We will also compare eye position effects in the premotor cortex with those described in the posterior parietal cortex. Our main thesis is that appropriate sensory input signals are distributed across the visuomotor continuum, and could potentially allow, in parallel, the emergence of multiple and task-dependent reference frames.

Animals↗
PubMed1999

Perception of self-motion from visual flow.

Accurate and efficient control of self-motion is an important requirement for our daily behavior. Visual feedback about self-motion is provided by optic flow. Optic flow can be used to estimate the direction of self-motion ('heading') rapidly and efficiently. Analysis of oculomotor behavior reveals that eye movements usually accompany self-motion. Such eye movements introduce additional retinal image motion so that the flow pattern on the retina usually consists of a combination of self-movement and eye movement components. The question of whether this 'retinal flow' alone allows the brain to estimate heading, or whether an additional 'extraretinal' eye movement signal is needed, has been controversial. This article reviews recent studies that suggest that heading can be estimated visually but extraretinal signals are used to disambiguate problematic situations. The dorsal stream of primate cortex contains motion processing areas that are selective for optic flow and self-motion. Models that link the properties of neurons in these areas to the properties of heading perception suggest possible underlying mechanisms of the visual perception of self-motion.

Journal Article↗
PubMed1998

Eye position encoding in the macaque posterior parietal cortex.

In two previous studies, we had demonstrated the influence of eye position on neuronal discharges in the middle temporal area, medial superior temporal area, lateral intraparietal area and area 7A of the awake monkey (Bremmer et al., 1997a,b). Eye position effects also have been found in visual cortical areas V3A and V6 and even in the premotor cortex and the supplementary eye field. These effects are generally discussed in light of a coordinate transformation of visual signals into a non-retinocentric frame of reference. Neural network studies dealing with the eye position effect succeeded in constructing such non-retinocentric representations by using model neurones whose response characteristics resembled those of 'real' neurones. However, to our knowledge, response properties of real neurones never acted as input into these neural networks. In the present study, we thus investigated whether, theoretically, eye position could be estimated from the population discharge of the (previously) recorded neurones and, if so, we intended to develop an encoding algorithm for the position of the eyes in the orbit. The optimal linear estimator proved the capability of the ensemble activity for determining correctly eye position. We then developed the so-called subpopulation encoding of eye position. This algorithm is based on the partition of the ensemble of neurones into two pairs of subpopulations. Eye position is represented by the differences of activity levels within each pair of subpopulations. Considering this result, encoding of the location of an object relative to the head could easily be accomplished by combining eye position information with the intrinsic knowledge about the retinal location of a visual stimulus. Taken together, these results show that throughout the monkey's visual cortical system information is available which can be used in a fairly simple manner in order to generate a non-retinocentric representation of visual information.

Animals↗
PubMed1998

Eye position effects on the neuronal activity of dorsal premotor cortex in the macaque monkey.

Visual inputs to the brain are mapped in a retinocentric reference frame, but the motor system plans movements in a body-centered frame. This basic observation implies that the brain must transform target coordinates from one reference frame to another. Physiological studies revealed that the posterior parietal cortex may contribute a large part of such a transformation, but the question remains as to whether the premotor areas receive visual information, from the parietal cortex, readily coded in body-centered coordinates. To answer this question, we studied dorsal premotor cortex (PMd) neurons in two monkeys while they performed a conditional visuomotor task and maintained fixation at different gaze angles. Visual stimuli were presented on a video monitor, and the monkeys made limb movements on a panel of three touch pads located at the bottom of the monitor. A trial begins when the monkey puts its hand on the central pad. Then, later in the trial, a colored cue instructed a limb movement to the left touch pad if red or to the right one if green. The cues lasted for a variable delay, the instructed delay period, and their offset served as the go signal. The fixation spot was presented at the center of the screen or at one of four peripheral locations. Because the monkey's head was restrained, peripheral fixations caused a deviation of the eyes within the orbit, but for each fixation angle, the instructional cue was presented at nine locations with constant retinocentric coordinates. After the presentation of the instructional cue, 133 PMd cells displayed a phasic discharge (signal-related activity), 157 were tonically active during the instructed delay period (set-related or preparatory activity), and 104 were active after the go signal in relation to movement (movement-related activity). A large proportion of cells showed variations of the discharge rate in relation to limb movement direction, but only modest proportions were sensitive to the cue's location (signal, 43%; set, 34%; movement, 29%). More importantly, the activity of most neurons (signal, 74%; set, 79%; movement, 79%) varied significantly (analysis of variance, P < 0.05) with orbital eye position. A regression analysis showed that the neuronal activity varied linearly with eye position along the horizontal and vertical axes and can be approximated by a two-dimensional regression plane. These data provide evidence that eye position signals modulate the neuronal activity beyond sensory areas, including those involved in visually guided reaching limb movements. Further, they show that neuronal activity related to movement preparation and execution combines at least two directional parameters: arm movement direction and gaze direction in space. It is suggested that a substantial population of PMd cells codes limb movement direction in a head-centered reference frame.

Action Potentials↗
PubMed1997

Spatial invariance of visual receptive fields in parietal cortex neurons.

Spatial information is conveyed to the primary visual cortex in retinal coordinates. Movement trajectory programming, however, requires a transformation from this sensory frame of reference into a frame appropriate for the selected part of the body, such as the eye, head or arms. To achieve this transformation, visual information must be combined with information from other sources: for instance, the location of an object of interest can be defined with respect to the observer's head if the position of the eyes in the orbit is known and is added to the object's retinal coordinates. Here we show that in a subdivision of the monkey parietal lobe, the ventral intraparietal area (VIP), the activity of visual neurons is modulated by eye-position signals, as in many other areas of the cortical visual system. We find that individual receptive fields of a population of VIP neurons are organized along a continuum, from eye to head coordinates. In the latter case, neurons encode the azimuth and/or elevation of a visual stimulus, independently of the direction in which the eyes are looking, thus representing spatial locations explicitly in at least a head-centred frame of reference.

Animals↗