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PubMed1999

Design and models for estimating antagonist potency (pA2, Kd and IC50) following the detection of antagonism observed in the presence of intrinsic activity.

The antagonist activity of the metabotropic glutamate receptor ligand (2S,1'S,2'S)-2-methyl-2(carboxycyclopropyl)glycine (MCCG) was examined using the [35S]GTPgammaS binding and forskolin (FSK)-stimulated adenosine 3':5'-cyclic monophosphate (cAMP) assays with recombinant Chinese hamster ovary (CHO) cells expressing the G protein-coupled human subtype 2 metabotropic glutamate (hmGlu2) receptor. Whereas MCCG proved to be a partial agonist in the GTPgammaS binding assay, it not only antagonized the agonist effect of (IS,3R)-ACPD in the cAMP assay but further produced an anomalous increase of the cAMP level relative to baseline. The anomalous MCCG response was also observed following treatment of the cells with MCCG in the absence of added agonist. Determination of the glutamate concentration in the incubate at the start and end of the cAMP reaction revealed the existence of micromolar concentrations of cellularly released glutamate throughout the course of the assay, reaching levels which exceeded its reported affinity for the mGlu2 receptor. Considering MCCG's partial agonist effect in the GTPgammaS binding assay and its pseudo-inverse agonist effect in the cAMP assay, available methods of estimating its antagonist potency were inappropriate since the classical Schild method and the alternative model suggested by Waud both assume the antagonist to lack a concentration-response relationship. We derived an alternate design and models that permit estimation of the pA2 (pAx), Kd and IC50 for antagonists which produce a concentration related effect when applied by themselves. With their use, the data acquired in both assays support the designation of MCCG as a competitive antagonist of the hmGlu2 receptor and provide similar pA2 estimates between assays. In addition, the newly derived models and design permit the determination of antagonist potency for partial and inverse agonists so characterized in studies employing the Schild design.

Amino Acids, Dicarboxylic↗
PubMed1997

The phonological short-term store-rehearsal system: patterns of impairment and neural correlates.

Two left brain-damaged patients (L.A. and T.O.) with a selective impairment of auditory-verbal span are reported. Patient L.A. was unable to hold auditory-verbal material in the phonological store component of short-term memory. His performance was however normal on tasks requiring phonological judgements, which specifically involve the phonological output buffer component of the rehearsal process. He also showed some evidence that rehearsal contributed to the immediate retention of auditory-verbal material. Patient T.O. never made use of the rehearsal process in tasks assessing both immediate retention and the ability to make phonological judgements, but the memory capacity of the phonological short-term store was comparatively preserved. These contrasting patterns of impairment suggest that the phonological store component of verbal short-term memory was severely impaired in patient L.A., and spared, at least in part, in patient T.O. The rehearsal process was preserved in L.A., and primarily defective in T.O. The localisation of the lesions in the left hemisphere (L.A.: inferior parietal lobule, superior and middle temporal gyri; T.O.: sub-cortical premotor and rolandic regions, anterior insula) suggests that these two sub-components of phonological short-term memory have discrete anatomical correlates.

Adult↗
PubMed1999

Dissociation of attentional processes in patients with focal frontal and posterior lesions.

A location-based ('select-what, respond-where') priming task was used to examine three measures of selective attention (interference (INT), negative priming (NP), and inhibition of return (IOR)) as a function of focal brain pathology and the complexity of target selection. Control subjects showed different patterns of performance for the three attentional measures as a function of complexity, suggesting some independence among INT, NP, and IOR. Brain-damaged subjects showed significant response slowing, as well as a number of lesion-specific attentional abnormalities. Right frontal (including bifrontal) damage resulted in proportionally increased interference related to task complexity. Left posterior damage increased IOR in the most complex task, while left frontal damage reversed the control pattern of IOR as a function of complexity. Right hemisphere (right posterior and right frontal damage) pathology resulted in a virtual loss of negative priming at all levels of task complexity; left and bifrontal damage resulted in diminished NP only related to increases in the complexity of selection. INT, NP, and IOR are mediated by different brain regions and their expression can be modulated by the complexity of the selection task.

Analysis of Variance↗
PubMed2002

Modeling neural oscillations.

A brief review of oscillatory activity in neurons and networks is given. Conditions required for neural oscillations are provided. Three mathematical methods for studying the coupling between neural oscillators are described: (i) weak coupling, (ii) firing time maps, and (iii) leaky integrate-and-fire methods. Several applications from macroscopic motor behavior to slice phenomena are provided.

Algorithms↗
PubMed2003

Social neuroscience, empathy, brain integration, and neurodevelopmental disorders.

Paul MacLean has investigated integrated brain functioning through selected brain lesions in animals that disturb circuits necessary for complex behaviors, such as social displays. MacLean is unique in his comparative neurobehavioral approach that emphasizes the evolutionary origins of parenting and social behaviors and the implications of brain changes in the evolution from reptiles (social displays) to mammals (nursing, audiovocal communication, play) to man (self-awareness, intentionality, social context) that link affect and cognition. Subjectively, how "looking with feeling toward others," the basic element in empathy, evolved has been a central concern of his. Neuroimaging studies of social cognition, mother-infant communication, moral behavior, forgiveness, and trust are consistent with particular brain systems being activated in cooperative social behaviors. The identification of mirror neurons is pertinent to MacLean's model of isopraxis and studies of thalamocortical resonances may be pertinent to his neurobehavioral models. Studies of behavioral phenotypes in human neurodevelopmental disorders are consistent with MacLean's model of brain circuits being linked to complex behaviors during development. In autistic disorder, the behavioral phenotype involves disrupted social communication, deviant imaginative play, and motor stereotypies. In Lesch-Nyhan syndrome (LNS), self-injury occurs in individuals with normal sensory systems intact who require and request physical restraint to prevent self-injury; they ask for assistance from others to prevent them from harming themselves. Autism involves the lack of subjective awareness of others intentions and LNS involves a failure in self-regulation and self-control of self-injurious behavior. MacLean's models laid the groundwork for studies focused on understanding brain functioning in these conditions.

Animals↗
PubMed2000

The spread of attention and learning in feature search: effects of target distribution and task difficulty.

We examined the roles of two determinants of spatial attention in governing the spread of perceptual learning, namely, stimulus location distribution and task difficulty. Subjects were trained on detection of a target element with an odd orientation imbedded in an array of light bars with otherwise uniform orientation. To assess the effects of target distribution on attention and learning, target positions were distributed so that attention was allocated not only to the target positions themselves, but also to intermediate positions where the target was not presented. Target detection performance substantially improved and improvement spread to match the induced window of spatial attention rather than only the actual target locations. To assess the effect of task difficulty on the spread of attention and learning, the target-distractor orientation difference and the time interval available for processing were manipulated. In addition, we compared performance of subjects with more versus with less detection difficulty. A consistent pattern emerged: When the task becomes more difficult, the window of attention shrinks, and learning becomes more localized. We conclude that task-specific spatial attention is both necessary and sufficient to induce learning. The spread of spatial attention, and thus of learning, is determined by the integrated effects of target distribution and task difficulty. We propose a theoretical framework whereby these factors combine to determine the cortical level of the focus of attention, which in turn enables learning modifications.

Adult↗
PubMed2001

Flanker effects in peripheral contrast discrimination--psychophysics and modeling.

We studied lateral interactions in the periphery by measuring how contrast discrimination of a peripheral Gabor patch is affected by flankers. In the psychophysical experiments, two Gabor targets appeared simultaneously to the left and right of fixation (4 degrees eccentricity). Observers reported which contrast was higher (spatial 2-alternative-forced-choice). In different conditions, Gabor flankers of different orientation, phase, and contrast were present above and below the two targets, at a distance of three times the spatial Gabor period. The data show that collinear flanks impair discrimination performance for low pedestal contrasts but have no effect for high pedestal contrasts. The transition between these two result patterns occurs typically at a pedestal contrast which is similar to the flanker contrast. For orthogonal flanks, we find facilitation at low pedestal contrasts, and suppression at intermediate contrasts. We account for this complex interaction pattern by a model that assumes that flankers can provide additive input to the target unit, and that they further contribute to the target's gain control, but only in a limited range of pedestal contrasts; once the target contrast exceeds a critical value, inhibition becomes subtractive rather than divisive. We further make specific propositions on how this model could be implemented at the neuronal level and show that a simple integrate and fire unit that receives time-modulated inhibition behaves in a fashion strikingly similar to the model inferred from the psychophysical data.

Contrast Sensitivity↗
PubMed2001

Darkness filling-in: a neural model of darkness induction.

A model of darkness induction based on a neural filling-in mechanism is proposed. The model borrows principles from both Land's Retinex theory and BCS/FCS filling-in model of Grossberg and colleagues. The main novel assumption of the induction model is that darkness filling-in signals, which originate at luminance borders, are partially blocked when they try to cross other borders. The percentage of the filling-in signal that is blocked is proportional to the log luminance ratio across the border that does the blocking. The model is used to give a quantitative account of the data from a brightness matching experiment in which a decremental test disk was surrounded by two concentric rings. The luminances of the rings were independently varied to modulate the brightness of the test. Observers adjusted the luminance of a comparison disk surrounded by a single ring of higher luminance to match the test disk in brightness.

Adult↗
PubMed2002

Computing relief structure from motion with a distributed velocity and disparity representation.

Recent psychophysical experiments suggest that humans can recover only relief structure from motion (SFM); i.e., an object's 3D shape can only be determined up to a stretching transformation along the line of sight. Here we propose a physiologically plausible model for the computation of relief SFM, which is also applicable to the related problem of motion parallax. We assume that the perception of depth from motion is related to the firing of a subset of MT neurons tuned to both velocity and disparity. The model MT neurons are connected to each other laterally to form modulatory interactions. The overall connectivity is such that when a zero-disparity velocity pattern is fed into the system, the most responsive neurons are not those tuned to zero disparity, but instead are those having preferred disparities consistent with the relief structure of the velocity pattern. The model computes the correct relief structure under a wide range of parameters and can also reproduce the SFM illusions involving coaxial cylinders. It is consistent with the psychophysical observation that subjects with stereo impairment are also deficient in perceiving motion parallax, and with the physiological data that the responses of direction- and disparity-tuned MT cells covary with the perceived surface order of bistable SFM stimuli.

Cues↗
PubMed2002

A model of speed tuning in MT neurons.

We have shown previously that neurons in the middle temporal (MT) area of primate cortex have inseparable spatiotemporal receptive fields-their response profiles exhibit a ridge that is oriented in the spatiotemporal frequency domain, and this orientation predicts the neurons' preferred speed. When measured in spatiotemporal frequency space, such MT spectral receptive field (SRF) properties are closely matched to the spectrum generated by a moving edge. In contrast, V1 neurons have SRF properties that are poorly matched to moving edge spectra, indicating that V1 neurons are not tuned to a particular image speed but rather to specific spatial and temporal frequencies. Here we describe a neural mechanism based directly on the properties of V1 neurons that is able to explain the SRF change that occurs between V1 and MT. We outline the theory behind this transformation and posit an explanation for how the visual system extracts true speed (independent of spatial frequency) from retinal image motion. We tested this speed model against our MT neuron data and found that it provides an excellent account of speed tuning in MT.

Animals↗
PubMed2002

Symmetry detection and visual attention: a "binary-map" hypothesis.

Recent research suggests that human symmetry-detection mechanisms cannot simultaneously compare different colors across the axis of symmetry (Nature 399 (1999) 115). In the present study, observers were required to judge symmetry in arrays composed of elements varying not only in color, but also in size, spatial frequency and orientation. In every case, response times increased with the number of different levels of a given feature. It is proposed that this increase reflects a sequential strategy whereby coarse "binary maps" are created by attentional filtering, and the symmetry of each map is then checked. Experiment 2 required observers to detect "pseudo-symmetry" (symmetry in feature values defined relative to an arbitrary featural boundary); the ease with which this task was accomplished supported the binary map hypothesis. The results suggest that (1) symmetry detection is spatially imprecise, and (2) attentional gating can operate prior to symmetry detection in the visual pathway.

Adult↗
PubMed2002

Action-perception dissociation in response to target acceleration.

The purpose of this study was to investigate whether information about the acceleration characteristics of a moving target can be used for both action and perception. Also of interest was whether prior movement experience altered perceptual judgements. Participants manually intercepted targets moving with various acceleration, velocity and movement time characteristics. They also made perceptual judgements about the acceleration characteristics of these targets either with or without prior manual interception experience. Results showed that while aiming kinematics were sensitive to the acceleration characteristics of the target, participants were only able to perceptually discriminate the velocity characteristics of target motion, even after performing interceptive actions to the same targets. These results are discussed in terms of a two channel (action-perception) model of visuomotor control.

Acceleration↗
PubMed2002

Failure of direction identification for briefly presented second-order motion stimuli: evidence for weak direction selectivity of the mechanisms encoding motion.

We sought to investigate why the direction of second-order motion, unlike first-order motion, cannot be identified when the stimulus exposure duration is brief (<200 ms). In a series of experiments observers identified both the orientation (vertical or horizontal) and the direction (left, right, down or up) of a drifting sinusoidal modulation (0.93 c/ degrees ) in either the luminance (first order) or the contrast (second order) of a two-dimensional noise carrier. All motion stimuli were equated for visibility, and the duration was varied using the method of constant stimuli. Performance was measured for second-order motion over a range of drift temporal frequencies (0.63-5.04 Hz) and for first-order motion stimuli composed of two, opposite drifting modulations in luminance of unequal modulation depth. Orientation-identification performance was nearly 100% correct for both first-order and second-order motion stimuli, even at the briefest stimulus duration tested (26.49 ms). Direction identification for first-order motion was also typically good with brief presentations, but was poor for second-order motion when the exposure duration was < approximately 200 ms. Importantly increasing either the drift temporal frequency of second-order motion or the bidirectional nature of the first-order motion patterns produced comparable levels of performance for the two varieties of motion (i.e. the minimum duration required for reliable direction identification could be equated). As orientation-identification performance for the first-order and second-order motion stimuli was comparably good and minimally affected by duration, the marked differences on the direction-identification task must be specific to mechanisms that encode drift direction, rather than spatial structure. We propose that second-order motion detectors are much less selective for stimulus direction than first-order motion sensors, and thus are more susceptible to the deleterious effects of limiting stimulus duration (which introduces spurious motion in the opposite direction, particularly at low drift rates). Alternative explanations based on the delayed propagation of second-order motion signals or the temporal characteristics of the underlying motion mechanisms are not supported by our findings.

Humans↗
PubMed2002

Color opponency is an efficient representation of spectral properties in natural scenes.

The human visual system encodes the chromatic signals conveyed by the three types of retinal cone photoreceptors in an opponent fashion. This opponency is thought to reduce redundant information by decorrelating the photoreceptor signals. Correlations in the receptor signals are caused by the substantial overlap of the spectral sensitivities of the receptors, but it is not clear to what extent the properties of natural spectra contribute to the correlations. To investigate the influences of natural spectra and photoreceptor spectral sensitivities, we attempted to find linear codes with minimal redundancy for trichromatic images assuming human cone spectral sensitivities, or hypothetical non-overlapping cone sensitivities, respectively. The resulting properties of basis functions are similar in both cases. They are non-orthogonal, show strong opponency along an achromatic direction (luminance edges) and along chromatic directions, and they achieve a highly efficient encoding of natural chromatic signals. Thus, color opponency arises for the encoding of human cone signals, i.e. with strongly overlapping spectral sensitivities, but also under the assumption of non-overlapping spectral sensitivities. Our results suggest that color opponency may in part be a result of the properties of natural spectra and not solely a consequence of the cone spectral sensitivities.

Color Perception↗
PubMed2002

The perceived speed of drifting chromatic gratings is mechanism-dependent.

The perceived speed of chromatic motion was investigated for gratings that stimulate each chromatic mechanism [L-M and S-(L+M)] in isolation and for gratings that stimulate both chromatic systems. The observers' task consisted of adjusting the speed of a drifting achromatic grating to match the perceived speed of an isoluminant chromatic grating, drifting at 8 deg/s (temporal frequency of 4 Hz). Every observer reported a substantial decrease in perceived speed for chromatic gratings modulated along the S-(L+M) (blue-yellow) cardinal axis compared to other directions in color space. One observer even reported motion standstill for gratings modulated along this axis. Further testing demonstrates that the perceived speed of an isoluminant chromatic grating depends solely on the extent to which it stimulates the L-M (red-green) mechanism. Thus, under the conditions that were tested, the S-(L+M) postreceptoral mechanism does not appear to contribute significantly to determining the perceived speed of chromatic motion.

Color Perception↗
PubMed2002

Acquisition of visual shape primitives.

Shape primitives have long been proposed as components for object models in the visual system, and account for a considerable body of behavioral findings. While a large amount of effort has been devoted to the study of detection of these parts in the scenes, no research has been undertaken simulating the acquisition of these representations. We present a model which suggests how the shape primitives may be learned by experience in a self-organized fashion. This model offers the first successful unsupervised learning of shape primitives which are as complex as object parts and can serve as intermediate representations for various objects. The algorithm uses synthetic gray-level objects, each composed of several parts (primitives or else), and shape primitives emerge as a result of partial matches between several objects. Our algorithm does not use any a priori knowledge about any attributes of the patterns to be learned; and the recurrence of these visual patterns in various objects is the only basis for their emergence as new features.

Algorithms↗
PubMed2002

The extent of crowding in peripheral vision does not scale with target size.

Identifying a target is more difficult when distracters are present within a zone of interaction around the target. We investigated whether the spatial extent of the zone of interaction scales with the size of the target. Our target was a letter T in one-of-four orientations. Our distracters were four squared-thetas in one-of-two orientations, presented one in each of the four cardinal directions, equidistant from the target. Target-distracter separation was varied and the proportion of correct responses at each separation was determined. From these the extent of interaction was estimated. This procedure was repeated for different target sizes spread over a 5-fold range. In each case, the contrast of the target was adjusted so that its visibility was constant across target sizes. The experiment was performed in the luminance domain (grey targets on grey background) and in the chromatic domain (green target on equiluminant grey background). In the luminance domain, target size had only a small effect on the extent of interaction; these interactions did not scale with target size. The extents of interaction for chromatic stimuli were similar to those for luminance stimuli. For a fixed target visibility, decreasing the duration of the stimulus resulted in an increase in the extent of interaction. The relevance of our findings is discussed with regard to a variety of proposed explanations for crowding. Our results are consistent with an attention-based explanation for crowding.

Attention↗
PubMed2002

Global speed processing: evidence for local averaging within, but not across two speed ranges.

A primary task of the visual system is to extract the direction and speed of animate objects from the retinal image. We examined global speed processing by determining how local speeds are integrated and whether integration occurs across all speeds or within fixed speed ranges. The first experiment addressed how local motion signals are combined to determine the speed of an object in motion. Observers judged the speed of a moving cloud of dots that took a random walk in direction while the dots inside the cloud moved somewhat independently of the cloud itself. The apparent speed of the cloud of dots is found to change in proportion with the dot speed and is well predicted by calculating the average speed resulting from nearest neighbour matches across stimulus frames. The second experiment addressed whether local speeds are combined across all speeds or within fixed speed ranges for the detection of global motion. Global dot motion (GDM) stimuli that moved in a radial or rotational directions moving at a low speed of 1.2 degrees /s or a high speed of 9.6 degrees /s were used to measure the thresholds for detecting structured motion as a function of the speed of noise dots (0 degrees /s-10.8 degrees /s) added to the stimulus. With low-speed targets, only additional noise dots moving at low speeds interfered with signal detection. High-speed targets were only interfered with by dots moving at high speeds. This finding established the existence of at least two independent speed tuned systems in the range of speeds tested. Experiment 3 investigated how speed signals are combined within a system to determine the global speed. Using sectored radial GDM stimuli the perceived speed of the fastest dots was measured as a function of whether the speed of the dots in alternate sectors either activated the high or low-speed systems. Averaging only occurred when dots were all within the sensitivity range of the high-speed system, however, if alternate sectors activated separate speed systems, averaging did not occur. Thus local speeds are averaged, independent of direction, to derive a global speed estimate, but averaging only occurs within, and not across, speed tuned mechanisms.

Adult↗