Auditory discrimination learning and acoustic cue weighing in female zebra finches with localized FoxP1 knockdowns.

Rare disruptions of the transcription factor FOXP1 are implicated in a human neurodevelopmental disorder characterized by autism and/or intellectual disability with prominent problems in speech and language abilities. Avian orthologues of this transcription factor are evolutionarily conserved and highly expressed in specific regions of songbird brains, including areas associated with vocal production learning and auditory perception. Here, we investigated possible contributions of FoxP1 to song discrimination and auditory perception in juvenile and adult female zebra finches. They received lentiviral knockdowns of FoxP1 in one of two brain areas involved in auditory stimulus processing, HVC (proper name) or CMM (caudomedial mesopallium). Ninety-six females, distributed over different experimental and control groups were trained to discriminate between two stimulus songs in an operant Go/Nogo paradigm and subsequently tested with an array of stimuli. This made it possible to assess how well they recognized and categorized altered versions of training stimuli and whether localized FoxP1 knockdowns affected the role of different features during discrimination and categorization of song. Although FoxP1 expression was significantly reduced by the knockdowns, neither discrimination of the stimulus songs nor categorization of songs modified in pitch, sequential order of syllables or by reversed playback were affected. Subsequently, we analyzed the full dataset to assess the impact of the different stimulus manipulations for cue weighing in song discrimination. Our findings show that zebra finches rely on multiple parameters for song discrimination, but with relatively more prominent roles for spectral parameters and syllable sequencing as cues for song discrimination.NEW & NOTEWORTHY In humans, mutations of the transcription factor FoxP1 are implicated in speech and language problems. In songbirds, FoxP1 has been linked to male song learning and female preference strength. We found that FoxP1 knockdowns in female HVC and caudomedial mesopallium (CMM) did not alter song discrimination or categorization based on spectral and temporal information. However, this large dataset allowed to validate different cue weights for spectral over temporal information for song recognition.

Color discrimination repetition distorts color representations.

Perceptual learning is the improvement of perceptual performance after repeated practice on a perceptual task. Studies on perceptual learning in color vision are limited. In this study, we measured the impact of color discrimination repetitions at a specific base color on color perception for entire hues. Participants performed five sessions of color discrimination training (200 or 300 trials per session) over five days, at colors on either the negative or positive direction of the L-M color axis, based on group assignment. We administered three color perception assessments (unique hues, color category boundaries, and color appearance) before and after the sessions to evaluate perceptual changes after training. The results showed declines in color discrimination thresholds after training, as expected. Additionally, the training influenced outcomes across all three assessment types. After the training, the perceived color appearance changed near the trained color along the stimulus hue, and some of the unique hues and the color category boundaries moved significantly toward the trained color. These findings indicate that short-term repetitions of color discrimination training can alter color representations in the visual system, distorting color perception around the trained color.

Reach-to-Grasp and tactile discrimination task: A new task for the study of sensory-motor learning.

Although active touch in rodents arises from the forepaws as well as whiskers, most research on active touch only focuses on whiskers. This results in a paucity of tasks designed to assess the process of active touch with a forepaw. We develop a new experimental task, the Reach-to-Grasp and Tactile Discrimination task (RGTD task), to examine active touch with a forepaw in rodents, particularly changes in processes of active touch during motor skill learning. In the RGTD task, animals are required to (1) extend their forelimb to an object, (2) grasp the object, and (3) manipulate the grasped object with the forelimb. The animals must determine the direction of the manipulation based on active touch sensations arising during the period of the grasping. In experiment 1 of the present study, we showed that rats can learn the RGTD task. In experiment 2, we confirmed that the rats are capable of reversal learning of the RGTD task. The RGTD task shared most of the reaching movements involved with conventional forelimb reaching tasks. From the standpoint of a discrimination task, the RGTD task enables rigorous experimental control, for example by removing bias in the stimulus-response correspondence, and makes it possible to utilize diverse experimental procedures that have been difficult in prior tasks.

Learning enhances representations of taste-guided decisions in the mouse gustatory insular cortex.

Learning to discriminate overlapping gustatory stimuli that predict distinct outcomes-a feat known as discrimination learning-can mean the difference between ingesting a poison or a nutritive meal. Despite the obvious importance of this process, very little is known about the neural basis of taste discrimination learning. In other sensory modalities, this form of learning can be mediated by either the sharpening of sensory representations or the enhanced ability of "decision-making" circuits to interpret sensory information. Given the dual role of the gustatory insular cortex (GC) in encoding both sensory and decision-related variables, this region represents an ideal site for investigating how neural activity changes as animals learn a novel taste discrimination. Here, we present results from experiments relying on two-photon calcium imaging of GC neural activity in mice performing a taste-guided mixture discrimination task. The task allows for the recording of neural activity before and after learning induced by training mice to discriminate increasingly similar pairs of taste mixtures. Single-neuron and population analyses show a time-varying pattern of activity, with early sensory responses emerging after taste delivery and binary, choice-encoding responses emerging later in the delay before a decision is made. Our results demonstrate that, while both sensory and decision-related information is encoded by GC in the context of a taste mixture discrimination task, learning and improved performance are associated with a specific enhancement of decision-related responses.

Investigating individual learning behaviour of dogs during a yes/no detection task.

Detection dogs are frequently tested for their ability to detect a variety of targets. It is crucial to comprehend the processes for odour learning and the consequences of training on an expanding set of target scents on performance. To properly evaluate their ability to identify the target, the only true measure is the dogs' initial response to novel sources, since this excludes learning effects. In this study, we evaluated the individual learning processes of three detection dogs that were pre-trained to differentially respond to a faecal sample of a mare in oestrus (S+) and a faecal sample of the same mare in di-oestrus (S-). After reaching criterion during a test with known training samples, the dogs were tested for generalization to a novel source. Average responses to S+ and S- were calculated as a function of presentation sequence, and Signal Detection Theory was used to further analyse characteristic differences in learning. The results of this study suggest that the ability of individual scent detection dogs to learn within an olfactory discrimination test varies considerably. The information obtained in this study could be helpful for mitigation training. We show that through careful monitoring of individual learning processes, the strategy each dog followed becomes apparent: especially the observations on the dogs' responses to first encounters with novel sample sources. This provides us with more detailed information than the more traditional sensitivity and specificity measures and allows us to better predict the dog's capabilities.

Shift in excitation-inhibition balance underlies perceptual learning of temporal discrimination.

Temporal perceptual learning (TPL) constitutes a unique and profound demonstration of neural plasticity within the brain. Our understanding for the neurometabolic changes associated with TPL on the other hand has been limited in part by the use of traditional fMRI approaches. Since plasticity in the visual cortex has been shown to underlie perceptual learning of visual information, we tested the hypothesis that TPL of an auditory interval involves a similar change in plasticity of the auditory pathway and if so, whether these changes take place in a lower-order sensory-specific brain area such as the primary auditory cortex (A1), or a higher-order modality-independent brain area such as the inferior parietal cortex (IPC). This distinction will inform us of the mechanisms underlying perceptual learning as well as the locus of change as it relates to TPL. In the present study, we took advantage of a new technique: proton magnetic resonance spectroscopy (MRS) in combination with psychophysical measures to provide the first evidence of changes in neurometabolic processing following 5 days of temporal discrimination training. We measured the (E)xcitation-to-(I)nhibition ratio as an index of learning in the right IPC and left A1 while participants learned an auditory two-tone discrimination task. During the first day of training, we found a significant task-related increase in functional E/I ratio within the IPC. While the A1 exhibited the opposite pattern of neurochemical activity, this relationship did not reach statistical significance. After timing performance has reached a plateau, there were no further changes to functional E/I. These findings support the hypothesis that improvements in temporal discrimination relies on neuroplastic changes in the IPC, but it is possible that both areas work synergistically to acquire a temporal interval.

Dynamic regulation of corticostriatal glutamatergic synaptic expression during reversal learning in male mice.

Behavioral flexibility, one of the core executive functions of the brain, has been shown to be an essential skill for survival across species. Corticostriatal circuits play a critical role in mediating behavioral flexibility. The molecular mechanisms underlying these processes are still unclear. Here, we measured how synaptic glutamatergic α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor (AMPAR) and N-methyl-D-aspartic acid receptor (NMDAR) expression dynamically changed during specific stages of learning and reversal. Following training to well-established stages of discrimination and reversal learning on a touchscreen visual task, lateral orbitofrontal cortex (OFC), dorsal striatum (dS) as well as medial prefrontal cortex (mPFC), basolateral amygdala (BLA) and piriform cortex (Pir) were micro dissected from male mouse brain and the expression of glutamatergic receptor subunits in the synaptic fraction were measured via immunoblotting. We found that the GluN2B subunit of NMDAR in the OFC remained stable during initial discrimination learning but significantly increased in the synaptic fraction during mid-reversal stages, the period during which the OFC has been shown to play a critical role in updating outcome expectancies. In contrast, both GluA1 and GluA2 subunits of the AMPAR significantly increased in the dS synaptic fraction as new associations were learned late in reversal. Expression of NMDAR and AMPAR subunits did not significantly differ across learning stages in any other brain region. Together, these findings further support the involvement of OFC-dS circuits in moderating well-learned associations and flexible behavior and suggest that dynamic synaptic expression of NMDAR and AMPAR in these circuits may play a role in mediating efficient learning during discrimination and the ability to update previously learned associations as environmental contingencies change.

Conditional discrimination learning by pigeons: Stimulus-response chains or occasion setters?

In conditional discrimination, the conditional stimulus or sample indicates which of two choice or comparison stimuli is associated with a reinforcer. Two hypotheses have been proposed concerning the role of the sample stimulus. According to Hull (1952), the sample and the response to the correct comparison form a stimulus-response chain. According to Skinner (1938), however, the sample serves as an occasion setter, setting the occasion for the choice of the correct comparison stimulus. In a conditional discrimination, if the sample stimulus forms part of a stimulus-response chain, then presenting the sample in the absence of the comparison stimuli should weaken the association. If the sample serves as an occasion setter, however, presenting the sample alone should not weaken its occasion-setting ability. In two experiments we tested these predictions. In Experiment 1, following conditional discrimination training with vertical and horizontal line samples and red and green comparison stimuli, we found that the presentation of the samples without the comparison stimuli (followed sometimes by a reinforcer) had little effect on conditional discrimination accuracy. In Experiment 2, two different houselights served as samples. When we presented the samples without comparison stimuli and without the reinforcers we found similar results. The results support the hypothesis that in conditional discrimination, the samples serve as occasion setters. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Interference of same/different learning by a spatial discrimination.

Same/different learning by pigeons has long been of interest to experimental psychologists. In one of these procedures, matching-to-sample, responses to a sample stimulus result in the presentation of two comparison stimuli, one of which matches the sample, the other of which does not, and choice of the matching stimulus is reinforced. Evidence of a matching concept has been found when transfer has been found to new stimuli. Given the transfer results, it is surprising that acquisition of two matching tasks (or two mismatching tasks), has not been found to be any faster than one matching and one mismatching task (i.e., two compatible tasks do not appear to facilitate each other). In the present experiment, we asked if matching acquisition involving three colors would be retarded if the correct response to a fourth color was not matching but was spatial (e.g., if the sample is red choose the red comparison, if the sample is green choose the green comparison, if the sample is yellow choose the yellow comparison, but if the sample is blue choose the left comparison). We found that acquisition of this task was slower than acquisition of a four color matching task (i.e., when the sample was blue, the blue comparison was correct). The results suggest that there is an interaction among matching associations, such that common rules facilitate learning compared with having to learn an inconsistent (spatial) rule. This result provides further evidence of the development of a matching concept by pigeons.

Intermixed rapid exposure to similar stimuli reduces the effective salience of their distinctive features.

Intermixed exposure to two similar stimuli, for example, AX and BX, improves subsequent discrimination between them compared to blocked exposure (the intermixed/blocked effect). Salience modulation models, developed mainly from research with nonhuman animals and exposure to widely spaced similar stimuli, explain this effect in terms of increased salience of the unique elements, A and B. Conversely, the results from experiments initially conducted with humans and exposure to close spaced similar stimuli have led to the suggestion that it is the development of well-unitized representations of unique elements that leads to better discrimination, leaving the unique elements with less effective salience. The experiments carried out here aim to replicate the intermixed/blocked effect in rats using an exposure procedure with rapid succession between stimuli and to assess the effective salience of unique elements. In Experiment 1, an aversion to a new flavor, Y, was conditioned and then an external inhibition test with AY was given. In Experiment 2, an aversion to A was conditioned and its extinction was measured on unreinforced trials. In Experiment 3, an aversion to AY was conditioned and the associated aversion to Y was measured. We found after rapid intermixed preexposure a reduction in generalization from the aversive Y element to the compound AY (Experiment 1) as well as a reduction in A's salience (Experiments 2 and 3) compared to the effects of blocked preexposure. The results are discussed in terms of the various mechanisms underlying perceptual learning, which appear to depend on the details of the task. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Social odor discrimination and its enhancement by associative learning in the hippocampal CA2 region.

Although the hippocampus is crucial for social memory, how social sensory information is combined with contextual information to form episodic social memories remains unknown. Here, we investigated the mechanisms for social sensory information processing using two-photon calcium imaging from hippocampal CA2 pyramidal neurons (PNs)-which are crucial for social memory-in awake head-fixed mice exposed to social and non-social odors. We found that CA2 PNs represent social odors of individual conspecifics and that these representations are refined during associative social odor-reward learning to enhance the discrimination of rewarded compared with unrewarded odors. Moreover, the structure of the CA2 PN population activity enables CA2 to generalize along categories of rewarded versus unrewarded and social versus non-social odor stimuli. Finally, we found that CA2 is important for learning social but not non-social odor-reward associations. These properties of CA2 odor representations provide a likely substrate for the encoding of episodic social memory.

Prior experience modifies acquisition trajectories via response-strategy sampling.

Few studies have considered how signal detection parameters evolve during acquisition periods. We addressed this gap by training mice with differential prior experience in a conditional discrimination, auditory signal detection task. Naïve mice, mice given separate experience with each of the later correct choice options (Correct Choice Response Transfer, CCRT), and mice experienced in conditional discriminations (Conditional Discrimination Transfer, CDT) were trained to detect the presence or absence of a tone in white noise. We analyzed data assuming a two-period model of acquisition: a pre-solution and solution period (Heinemann EG (1983) in The Presolution period and the detection of statistical associations. In: Quantitative analyses of behavior: discrimination processes, vol. 4, pp. 21-36). Ballinger. http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.536.1978andrep=rep1andtype=pdf ). The pre-solution period was characterized by a selective sampling of biased response strategies until adoption of a conditional responding strategy in the solution period. Correspondingly, discriminability remained low until the solution period; criterion took excursions reflecting response-strategy sampling. Prior experience affected the length and composition of the pre-solution period. Whereas CCRT and CDT mice had shorter pre-solution periods than naïve mice, CDT and Naïve mice developed substantial criterion biases and acquired asymptotic discriminability faster than CCRT mice. To explain these data, we propose a learning model in which mice selectively sample and test different response-strategies and corresponding task structures until they exit the pre-solution period. Upon exit, mice adopt the conditional responding strategy and task structure, with action values updated via inference and generalization from the other task structures. Simulations of representative mouse data illustrate the viability of this model.

Do you see what I see? Testing horses' ability to recognise real-life objects from 2D computer projections.

The use of 2-dimensional representations (e.g. photographs or digital images) of real-life physical objects has been an important tool in studies of animal cognition. Horses are reported to recognise objects and individuals (conspecifics and humans) from printed photographs, but it is unclear whether image recognition is also true for digital images, e.g. computer projections. We expected that horses trained to discriminate between two real-life objects would show the same learnt response to digital images of these objects indicating that the images were perceived as objects, or representations of such. Riding-school horses (N = 27) learnt to touch one of two objects (target object counterbalanced between horses) to instantly receive a food reward. After discrimination learning (three consecutive sessions of 8/10 correct trials), horses were immediately tested with on-screen images of the objects over 10 image trials interspersed with five real object trials. At first image presentation, all but two horses spontaneously responded to the images with the learnt behaviour by contacting one of the two images, but the number of horses touching the correct image was not different from chance (14/27 horses, p > 0.05). Only one horse touched the correct image above chance level across 10 image trials (9/10 correct responses, p = 0.021). Our findings thus question whether horses recognise real-life objects from digital images. We discuss how methodological factors and individual differences (i.e. age, welfare state) might have influenced animals' response to the images, and the importance of validating the suitability of stimuli of this kind for cognitive studies in horses.

Selective enhancement of neural coding in V1 underlies fine-discrimination learning in tree shrew.

Visual discrimination improves with training, a phenomenon that is thought to reflect plastic changes in the responses of neurons in primary visual cortex (V1). However, the identity of the neurons that undergo change, the nature of the changes, and the consequences of these changes for other visual behaviors remain unclear. We used chronic in vivo 2-photon calcium imaging to monitor the responses of neurons in the V1 of tree shrews learning a Go/No-Go fine orientation discrimination task. We observed increases in neural population measures of discriminability for task-relevant stimuli that correlate with performance and depend on a select subset of neurons with preferred orientations that include the rewarded stimulus and nearby orientations biased away from the non-rewarded stimulus. Learning is accompanied by selective enhancement in the response of these neurons to the rewarded stimulus that further increases their ability to discriminate the task stimuli. These changes persist outside of the trained task and predict observed enhancement and impairment in performance of other discriminations, providing evidence for selective and persistent learning-induced plasticity in the V1, with significant consequences for perception.

Replication of a skills assessment for auditory-visual conditional discrimination training.

Auditory-visual conditional discrimination training (e.g., receptive identification training, listener responses; AVCD) is ubiquitous in early intervention and special education programs. Nevertheless, some learners with Autism Spectrum Disorder (ASD) do not appear to benefit from this training despite use of empirically validated treatments. To prevent exposure to extended training that does not lead to learning, a skills assessment that measures skills related to AVCD training will be useful for educators and practitioners. The current study replicated the skills assessment developed and evaluated by Kodak et al. (2015) with 8 participants with ASD who received behavior analytic intervention that included at least 1 goal related to AVCD training. Two of the 8 participants mastered all skills included in the assessment except scanning. In addition, 5 participants' responding failed to reach mastery during subsequent exposure to AVCD training, which further demonstrated the predictive utility of the skills assessment.

The impact of hyperlinks, skim reading and perceived importance when reading on the Web.

It has previously been shown that readers spend a great deal of time skim reading on the Web and that this type of reading can affect comprehension of text. Across two experiments, we examine how hyperlinks influence perceived importance of sentences and how perceived importance in turn affects reading behaviour. In Experiment 1, participants rated the importance of sentences across passages of Wikipedia text. In Experiment 2, a different set of participants read these passages while their eye movements were tracked, with the task being either reading for comprehension or skim reading. Reading times of sentences were analysed in relation to the type of task and the importance ratings from Experiment 1. Results from Experiment 1 show readers rated sentences without hyperlinks as being of less importance than sentences that did feature hyperlinks, and this effect is larger when sentences are lower on the page. It was also found that short sentences with more links were rated as more important, but only when they were presented at the top of the page. Long sentences with more links were rated as more important regardless of their position on the page. In Experiment 2, higher importance scores resulted in longer sentence reading times, measured as fixation durations. When skim reading, however, importance ratings had a lesser impact on online reading behaviour than when reading for comprehension. We suggest readers are less able to establish the importance of a sentence when skim reading, even though importance could have been assessed by information that would be fairly easy to extract (i.e. presence of hyperlinks, length of sentences, and position on the screen).

Selective Interruption of Auditory Interhemispheric Cross Talk Impairs Discrimination Learning of Frequency-Modulated Tone Direction But Not Gap Detection and Discrimination.

Functional hemispheric lateralization is a basic principle of brain organization. In the auditory domain, the right auditory cortex (AC) determines the pitch direction of continuous auditory stimuli whereas the left AC discriminates gaps in these stimuli. The involved functional interactions between the two sides, mediated by commissural connections, are poorly understood. Here, we selectively disrupted the interhemispheric cross talk from the left to the right primary AC and vice versa using chromophore-targeted laser-induced apoptosis of the respective projection neurons, which make up 6-17% of all AC neurons in Layers III, V, and VI. Following photolysis, male gerbils were trained in a first experimental set to discriminate between rising and falling frequency-modulated (FM) tone sweeps. The acquisition of the task was significantly delayed in lesioned animals of either lesion direction. However, the final discrimination performance and hit rate was lowest for animals with left-side lesioned commissural neurons, demonstrating that also information from the left AC is relevant for FM direction learning. Photolysis after successful learning did not affect the retrieval of the learned task, indicating that the disruption during learning was not because of a general functional impairment. In a second experimental set, the gerbil's ability to detect and discriminate small silent gaps of varying length within FM sweeps was tested. This ability was also preserved after interhemispheric disruption. Taken together, interhemispheric communication between the left and right AC is important for the acquisition of FM tone direction learning but not for its retrieval and for gap detection and gap duration discrimination.SIGNIFICANCE STATEMENT Hemispheric lateralization of neuronal functions such as speech and music processing in humans are common throughout the brain; however, the involved interhemispheric interactions are ill-defined. Here, we show that the selective photolytic disruption of auditory cortical commissural connections in rodents impairs the acquisition but not retrieval of a frequency-modulated tone direction discrimination task. The final discrimination performance and hit rate was lowest for animals with lesioned left-to-right-side projections; thus, although right auditory cortex is dominant, left auditory cortex is also relevant for learning this task. The detection and discrimination of small gaps within the tone sweeps remain intact, suggesting a pathway for the processing of these temporal structures, which could be independent from the lesioned interhemispheric cross talk.

Digital embryos: a novel technical approach to investigate perceptual categorization in pigeons (Columba livia) using machine learning.

Pigeons are classic model animals to study perceptual category learning. To achieve a deeper understanding of the cognitive mechanisms of categorization, a careful consideration of the employed stimulus material and a thorough analysis of the choice behavior is mandatory. In the present study, we combined the use of "virtual phylogenesis", an evolutionary algorithm to generate artificial yet naturalistic stimuli termed digital embryos and a machine learning approach on the pigeons' pecking responses to gain insight into the underlying categorization strategies of the animals. In a forced-choice procedure, pigeons learned to categorize these stimuli and transferred their knowledge successfully to novel exemplars. We used peck tracking to identify where on the stimulus the animals pecked and further investigated whether this behavior was indicative of the pigeon's choice. Going beyond the classical analysis of the binary choice, we were able to predict the presented stimulus class based on pecking location using a k-nearest neighbor classifier, indicating that pecks are related to features of interest. By analyzing error trials with this approach, we further identified potential strategies of the pigeons to discriminate between stimulus classes. These strategies remained stable during category transfer, but differed between individuals indicating that categorization learning is not limited to a single learning strategy.

Mice with exonic RELN deletion identified from a patient with schizophrenia have impaired visual discrimination learning and reversal learning in touchscreen operant tasks.

The Reelin gene (RELN) encodes a large extracellular protein, which has multiple roles in brain development and adult brain function. It activates a series of neuronal signal transduction pathways in the adult brain that function in synaptic plasticity, dendritic morphology, and cognitive function. To further investigate the roles of Reln in brain function, we generated a mouse line using the C57BL/6 J strain with the specific Reln deletion identified from a Japanese patient with schizophrenia (Reln-del mice). These mice exhibited abnormal sociality, but the pathophysiological significance of the Reln deletion for higher brain functions, such as learning and behavioral flexibility remains unclear. In this study, cognitive function in Reln-del mice was assessed using touchscreen-based visual discrimination (VD) and reversal learning (RL) tasks. Reln-del mice showed normal learning in the simple VD task, but the learning was delayed in the complex VD task as compared to their wild-type (WT) littermates. In the RL task, sessions were divided into early perseverative phase (sessions with <50% correct) and later learning phase (sessions with ≥50% correct). Reln-del mice showed normal perseveration but impaired relearning ability in both simple RL and complex RL task as compared to WT mice. These results suggest that Reln-del mice have impaired learning ability, but the behavioral flexibility is unaffected. Overall, the observed behavioral abnormalities in Reln-del mice suggest that this mouse model is a useful preclinical tool for investigating the neurobiological mechanism underlying cognitive impairments in schizophrenia and a therapeutic strategy.

How to turn the corner: Discrimination of path shapes in rats.

The internal representation of a path shape is an element that constructs an internal representation of an entire route or environment. In the present study, we examined the ability of rats to discriminate path shapes. The rats learned to discriminate between an oval-shaped runway and a square-shaped one and to respond to one of two response boxes on the two sides of the runways. After the learning sessions, we tested which of the inner and outer walls the rats used as cues for discrimination using different wall shapes. The results suggest that the rats used the shape of the inner walls for the discrimination. Subsequently, the learning sessions, in which different shapes of the inner and outer walls were used, continued. There was a tendency for the rats to show better performance when the shape of the inner walls was congruent with the rule in the original learning, suggesting again that the rats used the shape of the inner wall for the discrimination. In addition, similar results were obtained when the task was conducted in the dark, suggesting that rats can discriminate path shapes using non-visual information.