Effects of collicular lesions in the hamster during visual discrimination. An analysis from computer-video actograms

Recent reports have suggested that lesions of the superior colliculus may impair the orienting or scanning movements of the head that are thought to serve a key function in visual discrimination. In the present investigation computer-video “actograms” were used to quantify the head movements of freely moving hamsters performing a simultaneous visual discrimination. Hamsters with collicular lesions did not differ from the controls in their head movement spectra, but there was a significant reduction in the incidence of the pauses in locomotion during which these movements are made. Intact animals showed slower and irregular progression interspersed with pauses during which they made scanning movements, whereas collicular hamsters made straighter runs. Although the rate of learning of lesioned hamsters was not impaired, our data strongly suggest that they used different orienting and learning strategies. When tested for the effects of novel stimuli, unrelated to the task, normal hamsters reacted strongly with active exploratory scanning movements, but collicular animals did not, although qualitative changes in their behaviour suggested that the novel stimuli were not being ignored.     

Visual sampling after lesions of the superior colliculus in rats

In two separate experiments, rats with bilateral lesions of the superior colliculus showed significantly poorer relearning of a horizontal/vertical stripe discrimination than control animals. In Experiment 1, all animals showed disruption of performance when a stimulus--response (S--R) separation was introduced by raising the stimuli above the site of responding. However, the colliculectomized rats were much more disturbed by the S--R separation than were animals in the control group. In Experiment 2, all animals showed lower performance levels when conflicting patterns were introduced into the upper portion of the stimulus doors, but this time the rats with collicular lesions were less disturbed than the control animals. It is suggested (a) that when the stimulus and response sites are discontiguous, animals must make an appropriate orienting response in order to effectively sample the visual stimuli and (b) that lesions of the superior colliculus alter performance by interfering with this orienting behavior. The impairment in relearning is tentatively attributed to the absence of preoperative overtraining on the discrimination task.     

Contrasting effects of lateral striate and superior colliculus lesions on visual discrimination performance in rhesus monkeys.

Rhesus monkeys with lesions of lateral striate cortex, monkeys with superior colliculus lesions, and unoperated monkeys were tested for retention of a preoperatively acquired pattern discrimination. The three groups of monkeys were then tested in two-choice, color, color discrimination tests, one involving varying degrees of stimulus-response (S-R) separation and the other, administered several months later, involving various directions of S-R separations. The monkeys were also tested in a series of two-choice pattern discriminations, following each of which they were tested for relearning when the patterns were masked with bars or circles. The monkeys with lateral striate lesions were moderately retarded in retention of the pattern discrimination, whereas those with superior colliculus lesions were not. The monkeys with colliculus lesions, but not those with lateral striate lesions, were impaired in both S-R separation tests, which demonstrates that their deficit was not transient or solely due to a difficulty in shifting the gaze in one direction. The lateral striate monkeys, unlike those with colliculus lesions, were dificient in relearning discriminations between masked patterns. These findings suggest that superior colliculus and striate cortex may be involved in two different aspects of attention: respectively, shifting attention (and orientation) from one spatial locus to another and maintaining attention on fixated stimuli. Alternative interpretations of the effects of the lesions, based on their retinotopic loci, are discussed.     

Stimulus control and auditory discrimination learning sets in the bottlenose dolphin

The learning efficiency of an Atlantic bottlenose dolphin was evaluated using auditory discrimination learning-set tasks. Efficiency, as measured by the probability of a correct response on Trial 2 of a new discrete-trial, two-choice auditory discrimination problem, reached levels comparable to those attained by advanced species of nonhuman primates. Runs of errorless problems in some cases rivaled those reported for individual rhesus monkeys in visual discrimination learning-set tasks. This level of stimulus control of responses to new auditory discriminanda was attained through (a) the development of a sequential within-trial method for presentation of a pair of auditory discriminanda; (b) the extensive use of fading methods to train initial discriminations, followed by the fadeout of the use of fading; (c) the development of listening behavior through control of the animal's responses during projection of the auditory discriminanda; and (d) the use of highly discriminable auditory stimuli, by applying results of a parametric evaluation of discriminability of selected acoustic variables. Learning efficiency was tested using a cueing method on Trial 1 of each new discrimination, to allow the animal to identify the positive stimulus before its response. Efficiency was also tested with the more common blind baiting method, in which the Trial 1 response was reinforced on only a random half of the problems. Efficiency was high for both methods. The overall results were generally in keeping with exceptations of learning capacity based on the large size and high degree of cortical complexity of the brain of the bottlenose dolphin.     

Overtraining, extinction and shift learning in a concurrent discrimination in rats

Two experiments examined the influence of overtraining on the reversal of a concurrent discrimination. After rats are trained to criterion on two different discriminations in the same apparatus, the reversal of one of these proceeds more rapidly than when both are reversed. If the reversal is conducted after overtraining on the original discriminations, then the opposite pattern of results is observed. That is, learning about the reversal of both discriminations is more rapid than when only a single discrimination is reversed. Experiment 1 replicated this effect and suggested that it is not caused by differences in the rate of extinction during reversal learning. In order to test a cue-association account for these findings, Experiment 2 examined the effect of exchanging the negative stimuli of a concurrent discrimination. This manipulation had a disruptive influence on performance, but only when subjects were not overtrained on the original discrimination.     

The effect of reduced spatial cues on the learning of successive visual reversals

The performance of rats over 12 brightness discrimination reversals was studied under two experimental conditions. Under one condition all visual cues external to the apparatus were eliminated so that only the relative positions of the discriminanda could serve as a visual cue to spatial position. Under the other condition all visual cues to position were eliminated. Under the former condition performance deteriorated with successive reversals but under the latter condition performance improved. Implications of these results for theories of successive reversal improvement were discussed and two possible explanations were suggested.     

Retrograde amnesia for visual memories after hippocampal damage in rats

It is generally believed that the hippocampus is not required for simple discrimination learning. However, a small number of studies have shown that hippocampus damage impairs retention of a previously learned visual discrimination task. We propose that, although simple discrimination learning may proceed in the absence of the hippocampus, it plays an important role in this type of learning when it is intact. In order to test the role of the hippocampus in simple discrimination learning, we performed a series of experiments utilizing a two-choice picture discrimination task. Our experiments confirm that rats readily learn simple two-choice picture discriminations after hippocampus damage. However, if such discriminations are first learned while the hippocampus is intact, subsequent hippocampus damage causes severe retrograde amnesia for the discriminations. Furthermore, retrograde amnesia for simple picture discriminations was equally severe when the interval between training and damage was 1 d or 60 d; remote picture memories are not spared. Similarly, the rule or schema underlying a recently or remotely acquired picture discrimination learning set was lost after hippocampus damage. The severity of retrograde amnesia for simple picture discriminations is negatively correlated with the volume of spared hippocampus tissue. Thus, the hippocampus plays an essential role in long-term memories supporting simple picture discriminations.     

Orientation to visual stimuli and the superior colliculus in the rat

Rats were trained in a six-choice jumping stand apparatus to enter the dark door, and avoid the five bright doors. Rats with bilateral superior collicular lesions were found to be severely impaired in this task, although further experiments showed that they were able to discriminate bright from dark stimuli and could perform correctly if allowed to approach each door in turn. It is suggested that the superior colliculus is important in orientation to visual cues, and there is some evidence that it is involved in orientation to brightness but not visual form cues.     

Same-different categorization in rats

Same-different categorization is a fundamental feat of human cognition. Although birds and nonhuman primates readily learn same-different discriminations and successfully transfer them to novel stimuli, no such demonstration exists for rats. Using a spatial discrimination learning task, we show that rats can both learn to discriminate arrays of visual stimuli containing all same from all different items and transfer this discrimination to arrays composed of novel visual items. These results are consistent with rats' engaging in same-different categorization. As such, they pave the way for investigations into the perceptual, cognitive, and neurobiological substrates of abstract categorization behavior.     

Studies in olfactory acuity. I: Measurement of olfactory thresholds in the rat

The capacity of the rat to respond to olfactory cues has been studied in three experimental situations: (i) an elevated Y-maze, (ii) a rectangular choice apparatus in which extraneous environmental cues were present, and (iii) a circular choice apparatus from which such cues were either excluded or randomized.

No rat showed any sign of learning on the Y-maze even after extensive training and the application of electric shock as deterrent for error.

Some rats succeeded in performing an olfactory discrimination in the rectangular choice apparatus but learning was retarded, and in some cases inhibited, by the counter-influence of non-olfactory factors.

All rats readily learned an identical discrimination when shielded from such cues in the circular choice apparatus which provides a satisfactory medium both for training and for the study of olfactory acuity.

Differences in the behaviour of rats on these three types of apparatus suggest first, that the formation of learned habits based on olfaction depends on the close contiguity of stimulus and reward, and second that the sense of smell possesses weak orientating properties by comparison with those exerted by other modalities. The findings thus help to explain differences in the ease or di+culty uith which olfactory discriminations have been established by earlier workers.     

Inferior temporal lesions do not impair discrimination of rotated patterns in monkeys.

Ablation of inferior temporal cortex in the rhesus monkey produces a visual discrimination learning deficit. The severity of this deficit has often been found to be a function of task difficulty. This report concerns a type of visual discrimination problem that, although difficult, is not sensitive to inferior temporal lesions. Monkeys with anterior, posterior, and complete inferior temporal lesions were repeatedly unimpaired or only slightly impaired in learning to discriminate a pattern from the same pattern rotated 90 degrees or 180 degrees; yet they were very severely impaired in learning equally or more difficult discriminations of two different patterns. This demonstration that discrimination of orientation of patterns is relatively spared after inferior temporal lesions helps specify the pattern-recognition processes that require inferior temporal cortex.     

Detection of a deficit in visual discrimination of pattern following lesions of anterior cortex: role of type of motivation.

6 albino rats were trained under appetitive motivation to perform diametrically opposed visual discriminations of pattern via opposite eyes. Small lesions were subsequently placed unilaterally in anterior neocortex. Following a period of post-surgical recovery, subjects were re-trained on the diammetrically opposed discriminations in alternating 10-trial blocks. Every subject made more post-surgical errors on the discrimination mediated by the eye contralateral to the lesion. Since albino rats have virtually complete decussation of the visual system, this indicates selective impairment in mediation of performance of pattern discrimination in the part of the lesioned hemispheres. Thus, findings previously obtained in studies involving aversive motivation are general to appetitively motivated tasks.     

Elemental and Configural Processes in Patterning Discrimination Learning

Three experiments used appetitive conditioning with rats to examine the involvement of elemental and configural processes in positive and negative patterning discriminations. The first experiment demonstrated that negative and, to some extent, positive patterning discriminations were learned more rapidly when these discriminations consisted of stimulus elements that had previously been the reinforced as opposed to the non-reinforced elements of a simple discrimination. Experiment 2 revealed an excitatory summation effect during the early phase of negative patterning learning that depended upon discrimination pretraining. The final experiment demonstrated faster discrimination learning between the compound and the less salient, rather than the more salient, element of an instrumental patterning task. The present set of results were interpreted as reflecting the possibility, consistent with connectionist theory, that internal representations of the conditioned stimuli change over the course of a patterning discrimination.     

A computer touch-screen apparatus for training visual discriminations in rats.

We describe an operant conditioning apparatus that uses computerized touch-screen technology and is designed for the versatile and highly controlled testing of rats in a potentially wide variety of behavioral paradigms. Although computer-controlled touch-screen systems have been developed for use with pigeons, monkeys, and humans, analogous technologies and methods have not yet been developed for rats. The development of a touch-screen system for rats could enhance the efficiency of behavioral research with rats, and may offer a unique tool for studying animal learning. In the first test of the utility of the apparatus, 3 Sprague-Dawley rats learned to activate the touch screen only after the touch-screen panel was made slightly movable. These animals then learned to discriminate visual stimuli presented on the computer monitor, but only after the food magazine and pellet dispenser were moved to the rear of the chamber opposite the stimulus display and response window. In a test of the utility of the modified apparatus, 6 Long-Evans rats learned to activate the touch screen and learned one of three different simple discriminations using computer-generated, visually presented stimuli. A basic method for training rats to activate the computer touch screen and for visual discrimination training is described. Results show that rats learned to activate the touch screen and discriminate visual stimuli presented on a computer monitor. Potential applications and advantages of the touch-screen-equipped rat operant conditioning chamber are discussed.     

Overtraining, transfer to proprioceptive control and position reversal

If rats are overtrained on a visual discrimination they generally learn the reversal of the discrimination faster than non-overtrained rats. In position discriminations, however, this effect does not generally hold—indeed several investigators have found overtraining to retard position reversal. One of the important differences between the two types of problem seems to be the presence of irrelevant cues in visual discriminations, and their absence in position discriminations. It is suggested that a second important feature of position discriminations is that overtraining usually causes control of the maze running habit to be transferred from external to proprioceptive stimuli, and that successful reversal cannot normally occur until external control is re-established. In two experiments a study by Krechevsky and Honzik (1932) is repeated with certain modifications, with results that support this hypothesis; and a third experiment provides direct evidence of transfer to proprioceptive control in a T-maze. It is shown that this analysis will explain the apparently conflicting results of all recent position reversal experiments.     

Alleviation of forgetting of discrimination learning

Reactivation treatments intended to alleviate forgetting in rats were tested in two discrimination tasks—instrumental learning with brightness or with spatial location as the discriminanda. For memories acquired 28 days earlier, forgetting was alleviated with both tasks when testing was preceded by certain reactivation treatments. Therefore, the generality of the effectiveness of reactivation treatments for alleviating forgetting is extended to include discrimination learning and response measures other than go, no-go behavior.     

A primacy effect in monkeys when list position is relevant

In Experiment 1 (1a and 1b), Rhesus monkeys (Macaca mulatta) learned lists of two-choice visual discriminations in which list position was relevant to discrimination performance. For example, Stimulus A was the rewarded stimulus if it was presented at List Position 1, but was not rewarded if it was presented at any other position in the list; similarly, Stimulus B was rewarded only at List Position 2, and so on. In learning these lists, all animals showed a marked primacy effect. In Experiment 2 (2a and 2b), Rhesus monkeys and Cynomolgus monkeys (M. fascicularis) learned lists of visual discriminations in which each visual stimulus occupied a fixed position in a list, but list position was not relevant to discrimination performance. For example, Stimulus E was always rewarded, and was always presented at List Position 1. To increase the salience of list beginning as a distinctive event, successive presentations of the list were separated by 24-hr intervals. In Experiment 2 there was no primacy effect, however. These results show for the first time that a primacy effect can be obtained in visual discrimination learning by monkeys. Furthermore, they suggest that it is obtained only when list position is relevant to the discrimination learning task.     

Acquisition of discrimination learning of patterns identical in configuration in macaques (Macaca mulatta and M. fuscata)

The acquisition of discrimination of five pairs of pattern cues in a Wisconsin General Testing Apparatus by 223 naive macaque monkeys was compared. The pairs of discriminanda were identical in configuration but varied slightly in either the size of the cue or the size of the background plaque; thus, the degree of separation of the cue from the fringe of the plaque, the response site, was slightly different for each pair of discriminanda. These small differences in cue-response separation had marked effects on the rate of acquisition of the discriminations. Even an increment of separation as small as 0.5 cm resulted in a remarkable retardation of the acquisition. This retardation was due entirely to prolonged performance at the chance level, and not to a slow rate of improvement from the chance to a criterion level. The finding indicates that the difficulty in the acquisition learning on pattern tasks depends largely on the difficulty of attending to the pattern cues at small cue-response separations.     

Discrimination of shape reflections and shape orientations by Columba livia

By using a free-operant instrumental discrimination procedure, it was demonstrated that pigeons find two-dimensional mirror-image visual forms more difficult to distinguish than otherwise similar forms. Variations in orientation of the discriminanda exacerbated the relative confusability of mirror images. No significant difference was found in the pigeons' performance whether the birds were discriminating vertically or horizontally reflected mirror-image pairs. Mirror images of shapes were also shown to be less discriminable than upside-down versions of shapes. The similarity of mirror-image patterns is discussed in relation to the generalized recognition of bilaterally symmetrical forms by pigeons. Pigeons found an orientation discrimination task involving a 45 degree tilt comparatively hard. A second experiment with a discrete-trial conditional paradigm confirmed that discriminations of shape orientations can be difficult for these birds. The addition of shape cues improved the performance on the orientation discrimination task, more so when arbitrary shapes were employed than when mirror images were used, which indicates again that the latter were more difficult to discriminate than the former. The relative insensitivity to shape orientations is ascribed to normal ecological demands on pigeons.     

Recognition memory for complex visual discriminations is influenced by stimulus interference in rodents with perirhinal cortex damage

Rats with perirhinal cortex (PRC), hippocampal, or sham lesions were trained on a successive discrimination go/no-go task to examine recognition memory for an array of visual objects with varying interference among the objects in the array. Rats were trained to recognize a target array consisting of four particular objects that could be presented in any one of four possible configurations to cover baited food wells. If the four target objects were presented, the rat should displace each object to receive food. However, if a novel object replaced any one or more of the target objects, then the rat should withhold its response. The number of novel objects presented on nonrewarded trials varied from one to four. The fewer the number of novel objects in the array, the more interference the array shared with the target array, therefore increasing task difficulty. An increased number of novel objects should result in less interference with the target array and decreased task difficulty. Although accuracy was slightly lower in rats with hippocampal lesions compared with controls, the learning of the groups was not statistically different. In contrast, rats with PRC lesions were significantly impaired in learning compared with both control and hippocampal-lesioned rats. The results suggest that recognition memory for complex visual discriminations is affected by stimulus interference in rodents with PRC damage.