The effects of lorazepam on visual integration processes: How useful for neuroscientists?

The effects of psychotropic drugs can be seen as a reversible lesion of the central nervous system. We studied the effect of lorazepam, a benzodiazepine facilitating the fixation of GABA on the GABA_A receptor. Evidence will be reviewed that lorazepam affects integration processes, using static fragmented pictures or compound stimuli. It will be shown how these effects can be dissociated from other peripheral or central effects of the drug. As a whole, the results suggest that lorazepam improves the detection of discontinuities between collinear elements, but impairs the detection of discontinuities between two parallel line-ends. Recent results will be presented that support the hypothesis that lorazepam does not affect integration processes themselves but modulates the mechanisms involved in the processing of discontinuities, i.e., the processing of line-ends and/or the production of virtual lines orthogonal to the line-ends. An understanding of the effects of the drug might give insights into the way these signals are used and modulated by the visual system.     

Lorazepam,a Benzodiazepine,Induces Atypical Distractor Effects with Compound Stimuli: A Role for Line-ends in the Processing of Compound Letters

The present study was designed to dissociate the effect of a single dose of lorazepam (a benzodiazepine, GABA_A-agonist) on the processing of spatial frequency, visual integration, and attentional processes in healthy volunteers. The stimuli were composed of small letters spatially organized to form a large letter. The task was to decide which of two target letters was displayed, either as a small or the large letter. Effects of spatial frequency and attentional processing were explored by manipulating respectively the visual angle of the stimuli and the probability that the targetwould appear at the local or at the global level. The results showed no evidence of a lorazepam effect on spatial frequency or attentional processing, but there was an atypical distractor effect when the target was a large letter, related to the arrangement of line-ends. The results are discussed in relation to studies showing the involvement of GABA in the generation of neuronal responses to line-ends.     

Modulations of the processing of line discontinuities under selective attention conditions?

We examined whether the processing of discontinuities involved in figure-ground segmentation, like line ends, can be modulated under selective attention conditions. Subjects decided whether a gap in collinear or parallel lines was located to the right or left. Two stimuli were displayed in immediate succession. When the gaps were on the same side, reaction times (RTs) for the second stimulus increased when collinear lines followed parallel lines, or the reverse, but only when the two stimuli shared the same orientation and location. The effect did not depend on the global form of the stimuli or on the relative orientation of the gaps. A frame drawn around collinear elements affected the results, suggesting a crucial role of the "amodal" orthogonal lines produced when line ends are aligned. Including several gaps in the first stimulus also eliminated RT variations. By contrast, RT variations remained stable across several experimental blocks and were significant for interstimulus intervals from 50 to 600 msec between the two stimuli. These results are interpreted in terms of a modulation of the processing of line ends or the production of amodal lines, arising when attention is selectively drawn to a gap.     

Surface and Edge Information for Spatial Integration: A Saccadic-selection Search Task

Three experiments are described that investigate visual integration across space using a saccade selection paradigm. Subjects saccaded to a vertical target in the presence of a number of horizontal distractor items. Both horizontal and vertical items were composed of two identical elements, so, in order to locate the target, subjects had to integrate the pairs of elements together. We measured saccade accuracy, the proportion of saccades directed to the vertical target, together with saccade latency following display appearance. In Experiment 1 we found that saccade accuracy was improved by the items having either common surface properties or collinear edges. These effects were not a resultof increased display heterogeneity (Experiment 2), or a result of the introduction of a strong internal contour between the items (Experiment 3). These results show that for saccadic selection both surface and edge properties of items are processed. This in turn suggests thatearly visual processing encodes and exploits both types of information.     

Task integration as a factor in secondary-task effects on sequence learning

In several studies it has been found that implicit sequence learning is impaired by the concurrent performance of a secondary task. In most studies the task was to count high-pitched tones when high-pitched and low-pitched tones were presented in a random sequence. In this paper the hypothesis is examined that dual-task interference in the particular combination of tasks results from task integration, in particular from the learning of an integrated visual-auditory sequence in which every second (auditory) element is random. Instead of the tone-counting task a similar go/no-go task was used in which foot responses to high-pitched tones were performed. In Exp. 1 the sequence of tones was random in one condition, but in two other conditions repeated tone sequences of 5 and 6 elements were combined with visual sequences of 6 elements. Under dual-task test conditions, implicit learning of the visual as well as the auditory sequences was better with the auditory sequence of 6 elements than of 5 elements, while under single-task test conditions the nature of the tone sequence had no effect. In Exp. 2 the superior implicit learning with the 6-element sequence was replicated with different test procedures in which either the visual or auditory sequence was changed to random or in which the two sequences remained intact but were shifted by one element relative to each other. Randomization of the visual or auditory sequences not only impaired visual or auditory RT, respectively, but also impaired RT to stimuli in the other modality, and this cross-modal effect was almost as strong as the intra-modal effect of randomization. Finally, in Exp. 3 it was shown that integrated visual-auditory sequences are learned only when responses to both of them are required, but not when the tones can be neglected. These results are consistent with a conception of implicit learning as (at least partly) a basic and nonselective type of learning of all potentially behaviorally relevant relations between stimuli in the environment and one's own actions.     

Peak Shift on an Artificial Dimension

Blough (1975) proposed an elemental model of generalization and discrimination phenomena in which stimuli from physical dimensions, such as wavelength, are conceptualized as overlapping sets of elements. In order to test predictions generated by this model, we constructed an artificial 'dimension' of stimuli as a series of overlapping sets of arbitrary 'icons' (small, unrelated shapes). In Experiment 1, we trained pigeons to discriminate two neighbouring stimuli from this artificial dimension and then assessed their responses to other stimuli from the dimension. The results of these tests typified those obtained with stimuli from genuine physical dimensions-that is, we obtained positive and negative peak shifts. In Experiment 2, human subjects were given a similar discrimination task and produced an analogous set of results to those of the pigeons. These results support Blough's elemental theory and are not readily accommodated by Pearce's (1987) configural theory. In Experiment 3, both humans and pigeons were trained and tested with stimuli from a real physical dimension (luminance). The pigeons' results were again consistent with Blough's analysis, but those of the humans suggested the use of more sophisticated strategies that are unavailable to pigeons and that lie outside the scope of elemental models of discrimination learning.     

Expansion of Sidman's theory: The inclusion of prompt stimuli in equivalence classes

Stimulus equivalence is defined as the ability to relate stimuli in novel ways after training in which not all of the stimuli had been directly linked to one another. Sidman (2000) suggested all elements of conditional discrimination training contingencies that result in equivalence potentially become class members. Research has demonstrated the inclusion of samples, comparisons, responses, and reinforcers in equivalence classes. Given the evidence that all elements of a conditional discrimination become part of the class, the purpose of this study was to determine if class-specific prompts would also enter into their relevant equivalence classes. Experiment 1 investigated the inclusion of prompts in an equivalence class using abstract stimuli with neurotypical students enrolled in higher education courses. Experiment 2 systematically replicated Experiment 1 using meaningful stimuli and individuals diagnosed with autism spectrum disorder. The results of both experiments demonstrated that class-specific prompts became part of equivalence classes with the other positive elements of the contingency. The results are discussed in terms of class expansion and the potential impact on equivalence-based instruction.     

Holistic vs. attribute repetition effects in classifying stimuli

The effect of the just previous stimulus on the current response was determined in three speeded classification experiments. When the stimuli were perceptual patterns, and when they were words, there was marked facilitation if the previous stimulus was identical to the current stimulus. When successive perceptual stimuli were composed of identical elements, but in a changed configuration, there was no measureable sequential effect on “same” response times. These results are for integral stimuli. When stimuli are not integral, as with some studies in the literature, it is suggested that each two-dimensional nonintegral stimulus is treated by the observer as two objects. In those cases, responses can be facilitated by repetition of that object on which responses were based on the previous trial, and this can occur when a value of the other dimensions (other objects) has not repeated. Accordingly, the processing of integral and nonintegral stimuli must be considered separately when attempting to evaluate process models. Further, facilitation due to stimulus repetitions does not account for all of the differences in response times between various classification tasks. These additional differences may be attributed to a multidimensional range effect analogous to the range effects found in psychophysical tasks.     

Competing responses and the processing of irrelevant information

A number of studies have shown that performance of identification and discrimination tasks is detrimentally affected by irrelevant information, yet other studies have failed to find such decrements. It is suggested that these contradictory findings depend on whether S must make difficult discriminations among the relevant stimuli, the irrelevant stimuli, or between the relevant and irrelevant stimuli. The role of irrelevant information in these tasks is to enhance or amplify the competihg responses engendered by the difficult discriminations. Irrelevant information enhances competing responses by increasing the information processing requirements of a task. The results of various studies of irrelevant information seem to be in good agreement with the assertions.     

Evidence for an association between the discriminative stimulus and the response-outcome association in instrumental learning

In 4 experiments, rats received 1 of several outcomes for engaging in various instrumental responses in the presence of discriminative stimuli. Discriminative stimuli shared some response-outcome relations but not others. When a response was subsequently extinguished in the presence of 1 discriminative stimulus, that produced relatively more decrement in responding in other stimuli that shared the same response-outcome relation. Other discriminative stimuli, in the presence of which that response had been reinforced by other outcomes and in which the original outcome had reinforced another response, were less affected. Moreover, postextinction devaluation of that outcome suggested that the particular response-outcome relation extinguished had undergone decrement. These results suggest that discriminative stimuli have relatively specific associations with the response-outcome relations that obtain in their presence.     

Attenuated audiovisual integration in middle-aged adults in a discrimination task

Numerous studies have focused on the diversity of audiovisual integration between younger and older adults. However, consecutive trends in audiovisual integration throughout life are still unclear. In the present study, to clarify audiovisual integration characteristics in middle-aged adults, we instructed younger and middle-aged adults to conduct an auditory/visual stimuli discrimination experiment. Randomized streams of unimodal auditory (A), unimodal visual (V) or audiovisual stimuli were presented on the left or right hemispace of the central fixation point, and subjects were instructed to respond to the target stimuli rapidly and accurately. Our results demonstrated that the responses of middle-aged adults to all unimodal and bimodal stimuli were significantly slower than those of younger adults (p < 0.05). Audiovisual integration was markedly delayed (onset time 360 ms) and weaker (peak 3.97%) in middle-aged adults than in younger adults (onset time 260 ms, peak 11.86%). The results suggested that audiovisual integration was attenuated in middle-aged adults and further confirmed age-related decline in information processing.     

Visual form integration and discontinuity detection

Two experiments were conducted to determine if discontinuity detection limits the visual integration of two stimuli presented successively. If this is the case, then presenting two dim stimuli should permit better integration than presenting two bright stimuli. In Experiment 1, five observers named the position of the missing dot in a 5 x 5 dot matrix. Twelve randomly selected dots were presented in each of two stimulus presentations. The stimuli varied in intensity in a ratio of 15:1 and stimulus onset asynchrony (SOA) varied between 50 msec and 80 msec. At the longer SOAs, presenting two bright stimuli led to a higher percentage of correct responses than presenting two dim stimuli. There was no difference between those conditions at the shorter SOAs. In Experiment 2, 12 naive observers rated the amount of integration of the same stimuli on a 7-point scale, under the same presentation conditions. Rated integration was higher with two dim stimuli than with two bright stimuli. These apparently contradictory results are explained in terms of Hawkins and Shulman's distinction between two kinds of visual persistence. It is concluded that subjects can respond to integration and discontinuity detection separately. No evidence was found that discontinuity detection limits the integration.     

Influence of physical and semantic information in a categorisation task of fragmented forms.

Two experiments were carried out to investigate the influence of structural and semantic information in the categorisation of visual forms. In experiment 1 the stimuli were pairs of 'well structured' or 'poorly structured' fragmented forms differing in structural properties including convexity, collinearity, and closure of the elements, and in their 'nameability'. The influence of structural and semantic information was tested by means of a task involving the presentation to both visual hemifields separately of stimuli having within-category physical similarity. For well structured forms, subjects were asked to judge if two sequentially presented forms belonged to the same semantic category. For poorly structured forms, subjects were required to decide if two stimuli belonged to the same previously learnt category of forms. For the two types of stimuli, one category was composed of physically similar items, and the other was composed of physically dissimilar items. The results show a marked advantage for the category composed of physically similar forms when the stimuli are well structured but no differences between the two categories when the stimuli are poorly structured. This suggests a facilitation in the computation of global shape information for forms having collinear and closed elements. The only effect of semantic information was a tendency towards a right visual field advantage for same-category pairs of well structured forms. The pattern of results for well structured fragmented forms (experiment 1) and forms in which fragments had been joined in order to complete the contour (experiment 2) were similar, suggesting that structured incomplete forms can be processed in the same way as forms having a continuous contour. Hypotheses about the organisational processes of forms are proposed in the discussion.     

Interactions between perceptual organization based on Gestalt laws and those based on hierarchical processing

Our recent research with compound stimuli (Han, Humphreys, & Chen, 1999a) suggests that grouping between local elements facilitates the perception of global structure, whereas encoding closure in local elements enhances their segmentation. The present study presents further evidence supporting this assertion. Experiment 1 first developed a new paradigm in which grouping between local elements was manipulated. Subjects responded to the orientations of perceptual groups consisting of local arrows or triangles embedded in background crosses. Responses to the orientations of the groups were slowed as a function of the increased contrast of the crosses, indicating that the strength of grouping between local arrows or triangles was gradually weakened by increasing the contrast of the crosses. Using a similar paradigm, Experiments 2 and 3 investigated the role of Gestalt factors in hierarchical analysis. Global arrows or triangles made up of local arrows, or triangles were embedded in background crosses. Subjects responded to global or local stimuli in terms of orientation or closure. Increasing the contrast of the background crosses produced stronger effects on global responses than on local responses and resulted in elimination of the global precedence effect and emerging of a local precedence effect, which was stronger for closure discrimination than for orientation discrimination. These results provide new evidence supporting our previous claim about the role of Gestalt factors in hierarchical analysis.     

Detection and discrimination of subjective contours defined by offset gratings

Three experiments were conducted to refine our understanding of the mechanisms that encode subjective contours. In Experiment 1, discrimination thresholds (stimulus onset asynchronies [SOAs] yielding 81% correct) were measured in a backward masking paradigm for subjective contours defined by offset gratings. For large apertures, thresholds increased as carrier frequency increased. For the smallest aperture, thresholds were a U-shaped function of carrier frequency. Experiment 2 showed that these threshold results were generally consistent with the rated strength of the subjective contours. Experiment 3 showed that detection thresholds (SOAs yielding 81% correct) again increased with carrier spatial frequency, increased for obliquely oriented carriers, and, for a particular frequency and orientation of the carrier, were lower when the subjective contour was orthogonal to the carrier. All of these results are well explained by a two-stage process in which a second-layer filter integrates the responses of end-stopped mechanisms to the terminators defining the subjective contour. In the model, the end-stopped mechanisms have low-pass sensitivity to carrier spatial frequency, and the sizes of the second-layer filters are proportional to the scale of the end-stopped mechanisms from which they draw their input.     

A model of the neural mechanisms underlying multisensory integration in the superior colliculus

Much of the information about multisensory integration is derived from studies of the cat superior colliculus (SC), a midbrain structure involved in orientation behaviors. This integration is apparent in the enhanced responses of SC neurons to cross-modal stimuli, responses that exceed those to any of the modality-specific component stimuli. The simplest model of multisensory integration is one in which the SC neuron simply sums its various sensory inputs. However, a number of empirical findings reveal the inadequacy of such a model; for example, the finding that deactivation of cortico-collicular inputs eliminates the enhanced response to a cross-modal stimulus without eliminating responses to the modality-specific component stimuli. These and other empirical findings inform a computational model that accounts for all of the most fundamental aspects of SC multisensory integration. The model is presented in two forms: an algebraic form that conveys the essential insights, and a compartmental form that represents the neuronal computations in a more biologically realistic way.     

Response inhibition in Extinction

In three experiments, rats were given a nonreinforced choice between two instrumental responses, in the presence of stimuli that had separately signalled the various outcomes those responses had earned. Whether those stimuli had been trained in a Pavlovian or instrumental procedure, they selectively enhanced the transfer response with which they shared an outcome. However, a subsequent test showed that this greater responding to a stimulus during extinction produced a reduced ability of that stimulus to augment its transfer response. These results are consistent with the development of an inhibitory S-R association during extinction.     

The influence of stimulus properties on multisensory processing in the awake primate superior colliculus.

Multisensory integration is a process whereby information converges from different sensory modalities to produce a response that is different from that elicited by the individual modalities presented alone. A neural basis for multisensory integration has been identified within a variety of brain regions, but the most thoroughly examined model has been that of the superior colliculus (SC). Multisensory processing in the SC of anaesthetized animals has been shown to be dependent on the physical parameters of the individual stimuli presented (e.g., intensity, direction, velocity) as well as their spatial relationship. However, it is unknown whether these stimulus features are important, or evident, in the awake behaving animal. To address this question, we evaluated the influence of physical properties of sensory stimuli (visual intensity, direction, and velocity; auditory intensity and location) on sensory activity and multisensory integration of SC neurons in awake, behaving primates. Monkeys were trained to fixate a central visual fixation point while visual and/or auditory stimuli were presented in the periphery. Visual stimuli were always presented within the contralateral receptive field of the neuron whereas auditory stimuli were presented at either ipsi- or contralateral locations. Many of the SC neurons responsive to these sensory stimuli (n = 66/84; 76%) had stronger responses when the visual and auditory stimuli were combined at contralateral locations than when the auditory stimulus was located on the ipsilateral side. This trend was significant across the population of auditory-responsive neurons. In addition, some SC neurons (n = 31) were presented a battery of tests in which the quality of one stimulus of a pair was systematically manipulated. A small proportion of these neurons (n = 8/31; 26%) showed preferential responses to stimuli with specific physical properties, and these preferences were not significantly altered when multisensory stimulus combinations were presented. These data demonstrate that multisensory processing in the awake behaving primate is influenced by the spatial congruency of the stimuli as well as their individual physical properties.     

Perceptual organization of line configurations: Is visual awareness necessary?

We examined whether configuring, which determines the appearance of grouped elements as a global shape, requires visual awareness, using a priming paradigm and two invisibility-inducing methods, CFS and sandwich masking. The primes were organized into configurations based on closure, collinearity, and symmetry (collinear primes), or on closure and symmetry (noncollinear primes). The prime-target congruency could be in configuration or in elements. During CFS, no significant response-priming was observed for invisible primes. When masking induced invisibility, a significant configuration response-priming was found for collinear and noncollinear primes, visible and invisible, with larger magnitude for the former. An element response-priming of equal magnitude was evident for visible and invisible noncollinear primes. Our results suggest that configuring can be accomplished in the absence of visual awareness when stimuli are rendered invisible by sandwich masking, but it benefits from visual awareness. Our results also suggest sensitivity to the available grouping cues in unconscious processing.     

Observing responses and discrimination learning

An operant response in the pigeon, whose performance results in exposure to the discriminative stimuli, is described and suggested as an experimental analogue for “observing.” Such an operant response is then used to explore the relationship between observing responses and discrimination learning in a variety of discrimination situations, of progressively increasing complexity. In general, the results support the contention that the development and maintainence of observing responses is closely related to the degree of differential behaviour manifested toward the discriminative stimuli. Certain modifications are suggested in the theoretical formulation underlying the concept of “observing responses.”