Dissociation between the effects of damage to perirhinal cortex and area TE

Perirhinal cortex and area TE are immediately adjacent to each other in the temporal lobe and reciprocally interconnected. These areas are thought to lie at the interface between visual perception and visual memory, but it has been unclear what their separate contributions might be. In three experiments, monkeys with bilateral lesions of the perirhinal cortex exhibited a different pattern of impairment than monkeys with bilateral lesions of area TE. In experiment 1, lesions of the perirhinal cortex produced a multimodal deficit in recognition memory (delayed nonmatching to sample), whereas lesions of area TE impaired performance only in the visual modality. In experiment 2, on a test of visual recognition memory (the visual paired comparison task) lesions of the perirhinal cortex impaired performance at long delays but spared performance at a very short delay. In contrast, lesions of area TE impaired performance even at the short delay. In experiment 3, lesions of the perirhinal cortex and lesions of area TE produced an opposite pattern of impairment on two visual discrimination tasks, simple object discrimination learning (impaired only by perirhinal lesions), and concurrent discrimination learning (impaired only by TE lesions). Taken together, the findings suggest that the perirhinal cortex, like other medial temporal lobe structures, is important for the formation of memory, whereas area TE is important for visual perceptual processing.     

Intact visual perceptual discrimination in humans in the absence of perirhinal cortex

While the role of the perirhinal cortex in declarative memory has been well established, it has been unclear whether the perirhinal cortex might serve an additional nonmnemonic role in visual perception. Evidence that the perirhinal cortex might be important for visual perception comes from a recent report that monkeys with perirhinal cortical lesions are impaired on difficult (but not on simple) visual discrimination tasks. We administered these same tasks to nine amnesic patients, including three severely impaired patients with complete damage to perirhinal cortex bilaterally (E.P., G.P., and G.T.). The patients performed all tasks as well as controls. We suggest that the function of perirhinal cortex as well as antero-lateral temporal cortex may differ between humans and monkeys.     

Perirhinal cortex lesions impair feature-negative discrimination

The role of the perirhinal cortex in inhibitory eyeblink conditioning was examined. In Experiment 1, rats were given lesions of the perirhinal cortex or control surgery and subsequently trained with a feature-negative discrimination procedure followed by summation and retardation tests for conditioned inhibition. Perirhinal cortex lesions impaired, but did not prevent acquisition of feature-negative discrimination. Results from the summation test showed that rats with perirhinal cortex lesions could not generalize feature-negative discrimination to a new stimulus. There were no group differences during the retardation test. Experiment 2 showed that lesions of the perirhinal cortex did not impair simple excitatory conditioning. Experiment 3 showed that perirhinal cortex lesions had no effect on acquisition of a simple tone-light discrimination. The results suggest that the perirhinal cortex plays a role in eyeblink conditioning when using discrimination procedures involving overlapping stimuli.     

Monkey perirhinal cortex is critical for visual memory, but not for visual perception: Reexamination of the behavioural evidence from monkeys

Overdependence on discrimination learning paradigms to assess the function of perirhinal cortex has complicated understanding of the cognitive role of this structure. Impairments in discrimination learning can result from at least two distinct causes: (a) failure to accurately apprehend and represent the relevant stimuli, or (b) failure to form and remember associations between stimulus representations and reward. Thus, the results of discrimination learning experiments do not readily differentiate deficits in perception from deficits in learning and memory. Here I describe studies that do dissociate learning and memory from perception and show that perirhinal cortex damage impairs learning and/or memory, but not perception. Reanalysis and reconsideration of other published data call into further question the hypothesis that the monkey perirhinal cortex plays a critical role in visual perception.     

Monkey perirhinal cortex is critical for visual memory, but not for visual perception: reexamination of the behavioural evidence from monkeys.

Overdependence on discrimination learning paradigms to assess the function of perirhinal cortex has complicated understanding of the cognitive role of this structure. Impairments in discrimination learning can result from at least two distinct causes: (a) failure to accurately apprehend and represent the relevant stimuli, or (b) failure to form and remember associations between stimulus representations and reward. Thus, the results of discrimination learning experiments do not readily differentiate deficits in perception from deficits in learning and memory. Here I describe studies that do dissociate learning and memory from perception and show that perirhinal cortex damage impairs learning and/or memory, but not perception. Reanalysis and reconsideration of other published data call into further question the hypothesis that the monkey perirhinal cortex plays a critical role in visual perception.     

The perceptual-mnemonic/feature conjunction model of perirhinal cortex function.

The perirhinal cortex was once thought to be "silent cortex", virtually ignored by researchers interested in the neurobiology of learning and memory. Following studies of brain damage associated with cases of amnesia, perirhinal cortex is now widely regarded as part of a "medial temporal lobe (MTL) memory system". This system is thought to be more or less functionally homogeneous, having a special role in declarative memory, and making little or no contribution to other functions such as perception. In the present article, we summarize an alternative view. First, we propose that components of the putative MTL system such as the hippocampus and perirhinal cortex have distinct and dissociable functions. Second, we provide evidence that the perirhinal cortex has a role in visual discrimination. In addition, we propose a specific role for perirhinal cortex in visual discrimination: the contribution of complex conjunctive representations to the solution of visual discrimination problems with a high degree of "feature ambiguity". These proposals constitute a new view of perirhinal cortex function, one that does not assume strict modularity of function in the occipito-temporal visual stream, but replaces this idea with the notion of a hierarchical representational continuum.     

Selective lesions in the temporal-hippocampal region of the rat: effects on acquisition and retention of a visual discrimination task.

The first purpose of this study was to investigate whether lesions in the temporal region may affect acquisition or retention of a discrimination task. In Experiment 1, rats with lesions of the temporal cortex (TC), the lateral entorhinal cortex (LEC), or their interconnections were tested postoperatively in simultaneous brightness discrimination. The results show that neither TC lesions nor LEC lesions affected acquisition of the task, and only LEC lesions impaired retention. TC/LEC transections impaired both acquisition and retention. The second purpose was to investigate effects of hippocampal lesions and perforant path transections on the discrimination task (Experiment 2). Both hippocampal and perforant path lesions impaired acquisition of the task, whereas retention was unaffected. It is suggested that TC and LEC are primarily involved in information storing and that hippocampal function is primarily involved in information processing.     

The perceptual-mnemonic/feature conjunction model of perirhinal cortex function

The perirhinal cortex was once thought to be “silent cortex”, virtually ignored by researchers interested in the neurobiology of learning and memory. Following studies of brain damage associated with cases of amnesia, perirhinal cortex is now widely regarded as part of a “medial temporal lobe (MTL) memory system”. This system is thought to be more or less functionally homogeneous, having a special role in declarative memory, and making little or no contribution to other functions such as perception. In the present article, we summarize an alternative view. First, we propose that components of the putative MTL system such as the hippocampus and perirhinal cortex have distinct and dissociable functions. Second, we provide evidence that the perirhinal cortex has a role in visual discrimination. In addition, we propose a specific role for perirhinal cortex in visual discrimination: the contribution of complex conjunctive representations to the solution of visual discrimination problems with a high degree of “feature ambiguity”. These proposals constitute a new view of perirhinal cortex function, one that does not assume strict modularity of function in the occipito-temporal visual stream, but replaces this idea with the notion of a hierarchical representational continuum.     

The feeling of fluent perception: a single experience from multiple asynchronous sources

Zeki and co-workers recently proposed that perception can best be described as locally distributed, asynchronous processes that each create a kind of microconsciousness, which condense into an experienced percept. The present article is aimed at extending this theory to metacognitive feelings. We present evidence that perceptual fluency-the subjective feeling of ease during perceptual processing-is based on speed of processing at different stages of the perceptual process. Specifically, detection of briefly presented stimuli was influenced by figure-ground contrast, but not by symmetry (Experiment 1) or the font (Experiment 2) of the stimuli. Conversely, discrimination of these stimuli was influenced by whether they were symmetric (Experiment 1) and by the font they were presented in (Experiment 2), but not by figure-ground contrast. Both tasks however were related with the subjective experience of fluency (Experiments 1 and 2). We conclude that subjective fluency is the conscious phenomenal correlate of different processing stages in visual perception.     

Differential activation patterns of occipital and prefrontal cortices during motion processing: Evidence from normal and schizophrenic brains

Visual motion perception is normally mediated by neural processing in the posterior cortex. Focal damage to the middle temporal area (MT), a posterior extrastriate region, induces motion perception impairment. It is unclear, however, how more broadly distributed cortical dysfunction affects this visual behavior and its neural substrates. Schizophrenia manifests itself in a variety of behavioral and perceptual abnormalities that have proved difficult to understand through a dysfunction of any single brain system. One of these perceptual abnormalities involves impaired motion perception. Motion processing provides an opportunity to clarify the roles of multiple cortical networks in both healthy and schizophrenic brains. Using fMRI, we measured cortical activation while participants performed two visual motion tasks (direction discrimination and speed discrimination) and one nonmotion task (contrast discrimination). Normal controls showed robust cortical activation (BOLD signal changes) in MT during the direction and speed discrimination tasks, documenting primary processing of sensory input in this posterior region. In patients with schizophrenia, cortical activation was significantly reduced in MT and significantly increased in the inferior convexity of the prefrontal cortex, an area that is normally involved in higher level cognitive processing. This shift in cortical responses from posterior to prefrontal regions suggests that motion perception in schizophrenia is associated with both deficient sensory processing and compensatory cognitive processing. Furthermore, this result provides evidence that in the context of broadly distributed cortical dysfunction, the usual functional specificity of the cortex becomes modified, even across the domains of sensory and cognitive processing.     

Is color “categorical perception” really perceptual?

Roberson and Davidoff (2000) found that color categorical perception (CP; better cross-category than within-category discrimination) was eliminated by verbal, but not by visual, interference presented during the interstimulus interval (ISI) of a discrimination task. On the basis of this finding, Roberson and Davidoff concluded that CP was mediated by verbal labels, and not by perceptual mechanisms, as is generally assumed. Experiment 1 replicated their results. However, it was found that if the interference type was uncertain on each trial (Experiment 2), CP then survived verbal interference. Moreover, it was found that the target color name could be retained across the ISI even with verbal interference (Experiment 3). We therefore conclude that color CP may indeed involve verbal labeling but that verbal interference does not necessarily prevent it.     

Separate but interacting recognition memory systems for different senses: the role of the rat perirhinal cortex

Two different models (convergent and parallel) potentially describe how recognition memory, the ability to detect the re-occurrence of a stimulus, is organized across different senses. To contrast these two models, rats with or without perirhinal cortex lesions were compared across various conditions that controlled available information from specific sensory modalities. Intact rats not only showed visual, tactile, and olfactory recognition, but also overcame changes in the types of sensory information available between object sampling and subsequent object recognition, e.g., between sampling in the light and recognition in the dark, or vice versa. Perirhinal lesions severely impaired object recognition whenever visual cues were available, but spared olfactory recognition and tactile-based object recognition when tested in the dark. The perirhinal lesions also blocked the ability to recognize an object sampled in the light and then tested for recognition in the dark, or vice versa. The findings reveal parallel recognition systems for different senses reliant on distinct brain areas, e.g., perirhinal cortex for vision, but also show that: (1) recognition memory for multisensory stimuli involves competition between sensory systems and (2) perirhinal cortex lesions produce a bias to rely on vision, despite the presence of intact recognition memory systems serving other senses.     

Categorical perception of familiar objects

We report three experiments where the categorical perception of familiar, three-dimensional objects was investigated. A continuum of shape change between 15 pairs of objects was created and the images along the continuum were used as stimuli. In Experiment 1 participants were first required to discriminate pairs of images of objects that lay along the shape continuum. Then participants were asked to classify each morph-image into one of two pre-specified classes. We found evidence for categorical perception in some but not all of our object pairs. In Experiment 2 we varied the viewpoint of the objects in the discrimination task and found that effects of categorical perception generalized across changes in view. In Experiment 3 similarity ratings for each object pair were collected. These similarity scores correlated with the degree of perceptual categorization found for the object pairs. Our findings suggest that some familiar objects are perceived categorically and that categorical perception is closely tied to inter-object perceptual similarity.     

Incidental encoding of numerosity in visual long-term memory

The visual system can readily extract numerosity information from brief experiences. This numerical perception is characterized by diminishing accuracy as numerosity increases, and impaired discrimination for similar quantities and large magnitudes. Here we assess whether these properties apply more broadly to numerosity in visual long-term memory. In surprise memory tests, we observed: Remarkable accuracy in estimating the number of repetitions of an exemplar image (Experiment 1a), that this accuracy decreased but remained high when estimating over categories (Experiments 1b and 1c), that numerical discrimination from memory exhibited psychophysical distance and size effects (Experiment 2), that these effects may derive from stored representations rather than post hoc approximation (Experiment 3a), and that they can reflect total elapsed experience in addition to discrete counts (Experiment 3b). Similar to how numerosity is readily extracted during visual perception, our results suggest that numerosity is encoded incidentally in visual long-term memory.     

Females excel at basic face perception

Females are generally better than males at recognizing facial emotions. However, it is not entirely clear whether and in what way females may also excel at non-affective face recognition. Here, we tested males and females on two perceptual face recognition tasks that involved only neutral expressions: detection and identity discrimination. On face detection (Experiment 1), females were significantly more accurate than males in detecting upright faces. This gender difference was reduced during inverted face detection, and not present during tree detection, suggesting that the magnitude of the gender difference for performance co-varies with the extent to which face processing mechanisms are involved. On facial identity discrimination (Experiment 2), females again outperformed males, particularly when face images were masked by visual noise, or the delay between comparison face images was extended from 0.5 to 3 s. These results reveal a female advantage in processing face-specific information and underscore the role of perceptual factors in socially relevant gender differences.     

Perception and recognition memory in monkeys following lesions of area TE and perirhinal cortex

Monkeys with lesions of perirhinal cortex (PR group) and monkeys with lesions of inferotemporal cortical area TE (TE group) were tested on a modified version of the delayed nonmatching to sample (DNMS) task that included very short delay intervals (0.5 sec) as well as longer delay intervals (1 min and 10 min). Lesions of the perirhinal cortex and lesions of area TE produced different patterns of impairment. The PR group learned the DNMS task as quickly as normal monkeys (N) when the delay between sample and choice was very short (0.5 sec). However, performance of the PR group, unlike that of the N group, fell to chance levels when the delay between sample and choice was lengthened to 10 min. In contrast to the PR group, the TE group was markedly impaired on the DNMS task even at the 0.5-sec delay, and three of four monkeys with TE lesions failed to acquire the task. The results provide support for the idea that perirhinal cortex is important not for perceptual processing, but for the formation and maintenance of long-term memory. Area TE is important for the perceptual processing of visual stimuli.     

Effects of preexposure on stimulus discrimination: an investigation of the mechanisms responsible for human perceptual learning

The effect of preexposure on human perceptual learning was investigated in four experiments. In Experiments 1a and 1b, participants were preexposed to one pair of visual stimuli on an intermixed schedule (AX/BX) and one on a blocked schedule (CX_DX). The ability to discriminate between AX and BX and between CX and DX was then assessed by examining the extent to which key presses assigned to each member of a stimulus pair generalized to the other member (Experiment 1a) and by looking at the accuracy of same-different responses (Experiment 1b). Stimuli were more easily discriminated following intermixed than following blocked preexposure on both the generalization and same-different tasks. This suggests that two stimuli are more perceptually distinct after intermixed preexposure. Experiments 2a and 2b investigated the mechanisms responsible for perceptual learning using same-different tasks. The results support the suggestion that the enhanced discrimination observed after intermixed preexposure is due to increases in the salience of the unique elements.     

A comparison of the effects of spatial separation on apparent motion in the auditory and visual modalities

In the present investigation, the effects of spatial separation on the interstimulus onset intervals (ISOIs) that produce auditory and visual apparent motion were compared. In Experiment 1, subjects were tested on auditory apparent motion. They listened to 50-msec broadband noise pulses that were presented through two speakers separated by one of six different values between 0 degrees and 160 degrees. On each trial, the sounds were temporally separated by 1 of 12 ISOIs from 0 to 500 msec. The subjects were instructed to categorize their perception of the sounds as "single," "simultaneous," "continuous motion," "broken motion," or "succession." They also indicated the proper temporal sequence of each sound pair. In Experiments 2 and 3, subjects were tested on visual apparent motion. Experiment 2 included a range of spatial separations from 6 degrees to 80 degrees; Experiment 3 included separations from .5 degrees to 10 degrees. The same ISOIs were used as in Experiment 1. When the separations were equal, the ISOIs at which auditory apparent motion was perceived were smaller than the values that produced the same experience in vision. Spatial separation affected only visual apparent motion. For separations less than 2 degrees, the ISOIs that produced visual continuous motion were nearly equal to those which produced auditory continuous motion. For larger separations, the ISOIs that produced visual apparent motion increased.     

The Bright and Dark Sides of Performance-Dependent Monetary Rewards: Evidence From Visual Perception Tasks

Studies have shown that performance-dependent monetary rewards facilitate visual perception. However, no study has examined whether such a positive effect is limited to the rewarded task or may be generalized to other tasks. In the current study, two groups of people were asked to perform two visual perception tasks, one being a reward-relevant task and the other being a reward-irrelevant task. For the reward-relevant task, the experimental group received performance-dependent monetary rewards, whereas the control group did not. For the reward-irrelevant task, both groups were not rewarded. The two tasks were randomly intermixed trial by trial (Experiment 1) or presented block by block (Experiment 2) or session by session (Experiments 3a, 3b, and 3c). Results showed that performance-dependent monetary rewards improved participants' performance on the relevant task in all experiments and impaired their performance on the irrelevant task in Experiments 2, 3a, 3b, and 3c. These results suggested that monetary rewards might incur a cost on reward-irrelevant tasks. Finally, the benefit of monetary rewards disappeared when they were no longer provided during the final session. This is the first study that reveals both the bright and dark sides of the performance-dependent monetary rewards in visual perception.     

Effects of preexposure on stimulus discrimination: An investigation of the mechanisms responsible for human perceptual learning

The effect of preexposure on human perceptual learning was investigated in four experiments. In Experiments 1a and 1b, participants were preexposed to one pair of visual stimuli on an intermixed schedule (AX/BX) and one on a blocked schedule (CX_DX). The ability to discriminate between AX and BX and between CX and DX was then assessed by examining the extent to which key presses assigned to each member of a stimulus pair generalized to the other member (Experiment 1a) and by looking at the accuracy of same–different responses (Experiment 1b). Stimuli were more easily discriminated following intermixed than following blocked preexposure on both the generalization and same–different tasks. This suggests that two stimuli are more perceptually distinct after intermixed preexposure. Experiments 2a and 2b investigated the mechanisms responsible for perceptual learning using same–different tasks. The results support the suggestion that the enhanced discrimination observed after intermixed preexposure is due to increases in the salience of the unique elements.