Individual and sex differences in learning abilities of ravens

Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.     

Sex chromosome complement affects multiple aspects of reversal-learning task performance in mice

Determining the mechanisms by which the sex-chromosome complement (SCC) affects learning, attention, and impulsivity has implications for observed sex differences in prevalence, severity, and prognosis of psychiatric/neurodevelopmental disorders and syndromes associated with sex-chromosome aneuploidy. Here, Four Core Genotypes (FCG) mice were evaluated in order to assess the separable and/or interacting effects of gonads (testes vs. ovaries) and their secretions and/or SCC (XX vs. XY) acting via non-gonadal mechanisms on behavior. We tested FCG mice on a reversal-learning task that enables the quantification of aspects of learning, attention and impulsivity. Across testing phases (involving the initial acquisition of a spatial discrimination and subsequent reversal learning), overall error rate was larger in XY compared with XX mice. Although XX and XY groups did not differ in the total number of trials required in order to reach a preset performance criterion, analyses of reversal error types showed more perseverative errors in XY than XX mice, with no difference in regressive errors. Additionally, prepotent-response latencies during the reversal phase were shorter in XY males, as compared with both XX gonadal males and females of either SCC, and failures to sustain the observing response were more frequent in XY mice than XX mice during the acquisition phase. These results indicate that SCC affects the characteristic pattern of response selection during acquisition and reversal performance without affecting the overall learning rate. More broadly, these results show direct effects of the SCC on cognitive processes that are relevant to psychiatric/neurodevelopmental disorders and syndromes associated with sex-chromosome aneuploidies.     

Transient Decline in Hippocampal Theta Activity during the Acquisition Process of the Negative Patterning Task

Hippocampal function is important in the acquisition of negative patterning but not of simple discrimination. This study examined rat hippocampal theta activity during the acquisition stages (early, middle, and late) of the negative patterning task (A+, B+, AB-). The results showed that hippocampal theta activity began to decline transiently (for 500 ms after non-reinforced stimulus presentation) during the late stage of learning in the negative patterning task. In addition, this transient decline in hippocampal theta activity in the late stage was lower in the negative patterning task than in the simple discrimination task. This transient decline during the late stage of task acquisition may be related to a learning process distinctive of the negative patterning task but not the simple discrimination task. We propose that the transient decline of hippocampal theta activity reflects inhibitory learning and/or response inhibition after the presentation of a compound stimulus specific to the negative patterning task.     

Early learning in the common marmoset (Callithrix jacchus): Behavior in the family group is related to preadolescent cognitive performance

Early environment can have a major impact on development, with family life known to play an important role. Longitudinal studies can therefore help increase our understanding of variance in cognitive abilities in young animals, as well as over time. We followed 22 marmosets (11 male and 11 female) from infancy through to early adolescence. At 3 months old, the marmosets were trained to reliably touch a rewarded stimulus. At 5 months, behavior was observed within the natal group. At 9 months, the marmosets were given a visual discrimination task to assess learning ability. Mann–Whitney U tests found no sex or family size differences in number of errors at 3 or 9 months. While no significant relationships were found between behavior in the family and learning at 3 months, significant negative correlations were found between duration spent in locomotion and learning errors (p = .05), as well as between frequency of calm vocalizations and learning errors (p = .001) at 9 months. A U-shape curve was found between amount of social play and learning at 9 months. Positive family interactions, including moderate amounts of play, as well as calm individual behavior, may therefore be important in learning. This study sheds light on cognitive development in much younger marmosets than previously studied, and helps increase understanding of how individual differences in learning may arise.     

Characterisation of learning and memory deficits following NMDA receptor antagonism

Summary The effects of the non-competitive NMDA antagonist dizocilpine in tests of cognitive function have been compared with its effects on motor function in rats. Severe motor impairments were observed at doses above 0.1 mg/kg. Dizocilpine (0.075 mg/kg) had no effect on the acquisition of a spatial discrimination task in a Y-maze, but disrupted reversal learning. Both the acquisition and reversal of a visual discrimination task were impaired following dizocilpine (0.075 mg/kg). Dizocilpine (0.04 mg/kg) also disrupted performance of a fivechoice visual reaction time task. It is clear that dizocilpine can impair cognitive function at doses which do not induce pronounced motor dysfunction. The impairment induced by dizocilpine includes a disruption of spatial discrimination learning and a deficit in tasks with sustained attentional demands.     

Loss of GluN2A‐containing NMDA receptors impairs extra‐dimensional set‐shifting

Glutamate neurotransmission via the N‐methyl‐d ‐aspartate receptor (NMDAR) is thought to mediate the synaptic plasticity underlying learning and memory formation. There is increasing evidence that deficits in NMDAR function are involved in the pathophysiology of cognitive dysfunction seen in neuropsychiatric disorders and addiction. NMDAR subunits confer different physiological properties to the receptor, interact with distinct intracellular postsynaptic scaffolding and signaling molecules, and are differentially expressed during development. Despite these known differences, the relative contribution of individual subunit composition to synaptic plasticity and learning is not fully elucidated. We have previously shown that constitutive deletion of GluN2A subunit in the mouse impairs discrimination and re‐learning phase of reversal when exemplars are complex picture stimuli, but spares acquisition and extinction of non‐discriminative visually cued instrumental response. To investigate the role of GluN2A containing NMDARs in executive control, we tested GluN2A knockout (GluN2A^KO), heterozygous (GluN2A^HET) and wild‐type (WT) littermates on an attentional set‐shifting task using species‐specific stimulus dimensions. To further explore the nature of deficits in this model, mice were tested on a visual discrimination reversal paradigm using simplified rotational stimuli. GluN2A^KO were not impaired on discrimination or reversal problems when tactile or olfactory stimuli were used, or when visual stimuli were sufficiently easy to discriminate. GluN2A^KO showed a specific and significant impairment in ventromedial prefrontal cortex‐mediated set‐shifting. Together these results support a role for GluN2A containing NMDAR in modulating executive control that can be masked by overlapping deficits in attentional processes during high task demands.     

Extensive Lesions in Rat Insular Cortex Significantly Disrupt Taste Sensitivity to NaCl and KCl and Slow Salt Discrimination Learning

While studies of the gustatory cortex (GC) mostly focus on its role in taste aversion learning and memory, the necessity of GC for other fundamental taste-guided behaviors remains largely untested. Here, rats with either excitotoxic lesions targeting GC (n = 26) or sham lesions (n = 14) were assessed for postsurgical retention of a presurgically LiCl-induced conditioned taste aversion (CTA) to 0.1M sucrose using a brief-access taste generalization test in a gustometer. The same animals were then trained in a two-response operant taste detection task and psychophysically tested for their salt (NaCl or KCl) sensitivity. Next, the rats were trained and tested in a NaCl vs. KCl taste discrimination task with concentrations varied. Rats meeting our histological inclusion criterion had large lesions (resulting in a group averaging 80% damage to GC and involving surrounding regions) and showed impaired postsurgical expression of the presurgical CTA (LiCl-injected, n = 9), demonstrated rightward shifts in the NaCl (0.54 log₁₀ shift) and KCl (0.35 log₁₀ shift) psychometric functions, and displayed retarded salt discrimination acquisition (n = 18), but eventually learned and performed the discrimination comparable to sham-operated animals. Interestingly, the degree of deficit between tasks correlated only modestly, if at all, suggesting that idiosyncratic differences in insular cortex lesion topography were the root of the individual differences in the behavioral effects demonstrated here. This latter finding hints at some degree of interanimal variation in the functional topography of insular cortex. Overall, GC appears to be necessary to maintain normal taste sensitivity to NaCl and KCl and for salt discrimination learning. However, higher salt concentrations can be detected and discriminated by rats with extensive damage to GC suggesting that the other resources of the gustatory system are sufficient to maintain partial competence in these tasks, supporting the view that such basic sensory-discriminative taste functions involve distributed processes among central gustatory structures.     

Galanthamine plus estradiol treatment enhances cognitive performance in aged ovariectomized rats

We hypothesize that beneficial effects of estradiol on cognitive performance diminish with age and time following menopause due to a progressive decline in basal forebrain cholinergic function. This study tested whether galanthamine, a cholinesterase inhibitor used to treat memory impairment associated with Alzheimer's disease, could enhance or restore estradiol effects on cognitive performance in aged rats that had been ovariectomized in middle-age. Rats were ovariectomized at 16–17 months of age. At 21–22 months of age rats began receiving daily injections of galanthamine (5 mg/day) or vehicle. After one week, half of each group also received 17ß-estradiol administered subcutaneously. Rats were then trained on a delayed matching to position (DMP) T-maze task, followed by an operant stimulus discrimination/reversal learning task. Treatment with galanthamine + estradiol significantly enhanced the rate of DMP acquisition and improved short-term delay-dependent spatial memory performance. Treatment with galanthamine or estradiol alone was without significant effect. Effects were task-specific in that galanthamine + estradiol treatment did not significantly improve performance on the stimulus discrimination/reversal learning task. In fact, estradiol was associated with a significant increase in incorrect responses on this task after reversal of the stimulus contingency. In addition, treatments did not significantly affect hippocampal choline acetyltransferase activity or acetylcholine release. This may be an effect of age, or possibly is related to compensatory changes associated with long-term cholinesterase inhibitor treatment. The data suggest that treating with a cholinesterase inhibitor can enhance the effects of estradiol on acquisition of a DMP task by old rats following a long period of hormone deprivation. This could be of particular benefit to older women who have not used hormone therapy for many years and are beginning to show signs of mild cognitive impairment. Potential mechanisms for these effects are discussed.     

Learning Differences between Feral Pigeons and Zenaida Doves: The Role of Neophobia and Human Proximity

Learning differences predicted from ecological variables can be confounded with differences in wariness of novel stimuli (neophobia). Previous work on feral pigeons ( Columba livia ), as well as on group-feeding and territorial zenaida doves ( Zenaida aurita ), reported individual and social learning differences predicted from social foraging mode. In the present study, we show that speed of learning a foraging task covaries with neophobia and latency to feed from a familiar dish in the three types of columbids. Pigeons were much faster than either territorial or group-feeding zenaida doves on all tests conducted in captivity, but showed unexpectedly strong neophobia in some urban flocks during field tests. Human proximity strongly affected performance in group-feeding doves both in the field and in captivity. They were slightly faster at learning than their territorial conspecifics in cage tests. In multiple regressions, species identity, but not social foraging mode, significantly predicted individual variation in learning, as did individual variation in neophobia. Wariness of novel stimuli and species differences associated with artificial selection appear to be more important than foraging mode and wariness of humans in accounting for learning differences between these columbids.     

Long Term Effects of Aversive Reinforcement on Colour Discrimination Learning in Free-Flying Bumblebees

The results of behavioural experiments provide important information about the structure and information-processing abilities of the visual system. Nevertheless, if we want to infer from behavioural data how the visual system operates, it is important to know how different learning protocols affect performance and to devise protocols that minimise noise in the response of experimental subjects. The purpose of this work was to investigate how reinforcement schedule and individual variability affect the learning process in a colour discrimination task. Free-flying bumblebees were trained to discriminate between two perceptually similar colours. The target colour was associated with sucrose solution, and the distractor could be associated with water or quinine solution throughout the experiment, or with one substance during the first half of the experiment and the other during the second half. Both acquisition and final performance of the discrimination task (measured as proportion of correct choices) were determined by the choice of reinforcer during the first half of the experiment: regardless of whether bees were trained with water or quinine during the second half of the experiment, bees trained with quinine during the first half learned the task faster and performed better during the whole experiment. Our results confirm that the choice of stimuli used during training affects the rate at which colour discrimination tasks are acquired and show that early contact with a strongly aversive stimulus can be sufficient to maintain high levels of attention during several hours. On the other hand, bees which took more time to decide on which flower to alight were more likely to make correct choices than bees which made fast decisions. This result supports the existence of a trade-off between foraging speed and accuracy, and highlights the importance of measuring choice latencies during behavioural experiments focusing on cognitive abilities.     

Instrumental discrimination: Reversal learning in the monotreme Tachyglossus aculeatus setosus

Tasmanian echidnas, Tachyglossus aculeatus setosus, were tested in serial discrimination-reversal tasks, comparing their operant behaviour in reversal-shift and non-reversal-shift conditions, using positional and visual/tactile cues. Learning-set formation of the pattern characteristic of eutherian mammals was observed in all tests for all subjects, but performances were superior in tests of positional discrimination and in reversal-shift conditions. The results were consistent with those obtainable for eutherian mammals in comparable tests and suggest that echidnas can be used as subjects in experimental studies of operant learning.     

Effects of atropine on operant test battery performance in rhesus monkeys.

The acute behavioral effects of atropine sulfate were assessed using a battery of complex food-reinforced operant tasks that included: temporal response differentiation (TRD, n = 7); delayed matching-to-sample (DMTS, n = 6), progressive ratio (PR, n = 8), incremental repeated acquisition (IRA, n = 8), and conditioned position responding (CPR, n = 8). Performance in these tasks is thought to depend primarily upon specific brain functions such as time perception, short-term memory and attention, motivation, learning, and color and position discrimination, respectively. Atropine sulfate (0.01-0.56 mg/kg iv), given 15-min pretesting, produced significant dose-dependent decreases in the number of reinforcers obtained in all tasks. Response rates decreased significantly at greater than or equal to 0.03 mg/kg for the learning and discrimination tasks, at greater than or equal to 0.10 mg/kg for the motivation and short-term memory and attention tasks, and at greater than or equal to 0.30 mg/kg for the time perception task. Response accuracies were significantly decreased at doses greater than or equal to 0.10 mg/kg for the learning, discrimination, and short-term memory and attention tasks, and at greater than or equal to 0.30 mg/kg for the time perception task. Thus, the order of task sensitivity to any disruption by atropine is learning = color and position discrimination greater than time perception = short-term memory and attention = motivation (IRA = CPR greater than TRD = DMTS = PR). Thus in monkeys, the rates of responding in operant tasks designed to model learning and color and position discrimination were the most sensitive measures to atropine's behavioral effects. Accuracy in these same task was also disrupted but at higher doses. These data support the hypothesis that cholinergic systems play a greater role in the speed (but not accuracy) of performance of our learning and discrimination tasks compared to all other tasks. Accuracy of responding in these and the short-term memory task, all of which involve the use of lights as visual stimuli, was more sensitive to disruption by atropine than those tasks which did not utilize such strong visual stimuli.     

Implementing long‐term baselines into primate tool‐use studies

Studies on primate tool‐use often involve the use of baseline conditions, as they allow for the examination of any differences in the subjects' behavior before and after the introduction of a tool‐use task. While these baseline conditions can be powerful for identifying the relative contributions of individual and social learning for the acquisition of tool‐use behaviors in naïve (usually captive) subjects, many have criticized them for being too short, and not allowing enough time for the behavior to develop spontaneously. Furthermore, some wild tool‐use behaviors such as chimpanzee nut‐cracking require animals to manipulate and familiarize themselves with the materials of the behavior within a “sensitive learning period” before it develops later on in life. One solution to this problem is to implement long‐term baselines, in which, with collaboration with zoological institutions, the materials of the behavior are left in the enclosure for an extended period. The keepers would then be asked not to demonstrate or train the animals in the target behavior, but to report back to the researchers if they observe the behavior emerge during this extended period. Alongside keeper reports, video cameras could be installed in the enclosure to minimize the chance of false negatives and to allow for coding and inter‐rater reliability to be carried out on the videos. These long‐term baselines therefore provide extended enrichment opportunities for the animals, alongside allowing the zoological institution to publicize their involvement with the study and guests to observe animals interacting with different testing apparatuses and tools. Finally, long‐term baselines can provide invaluable insight on the individual and social learning abilities of primates as well as the potential development stages and sensitive learning periods required for specific behaviors.     

Spatial discrimination performance of Drosophila melanogaster: Test-retest assessments and a reinterpretation

Two experiments found significant correlations between the performance of experimental flies, which were retrained on a spatial discrimination task, and yoked controls which were trained for the first time after having been allowed to explore the maze. The proportion of variance accounted for by the correlations equalled or exceeded that found for other variables in previous experiments. The data suggest that the performance of individual experimental flies and that of their control counterparts are not independent, seriously jeopardizing an instrumental learning interpretation of Drosophila performance.     

Acquisition versus Consolidation of Auditory Perceptual Learning Using Mixed-Training Regimens

Learning is considered to consist of two distinct phases–acquisition and consolidation. Acquisition can be disrupted when short periods of training on more than one task are interleaved, whereas consolidation can be disrupted when a second task is trained after the first has been initiated. Here we investigated the conditions governing the disruption to acquisition and consolidation during mixed-training regimens in which primary and secondary amplitude modulation tasks were either interleaved or presented consecutively. The secondary task differed from the primary task in either task-irrelevant (carrier frequency) or task-relevant (modulation rate) stimulus features while requiring the same perceptual judgment (amplitude modulation depth discrimination), or shared both irrelevant and relevant features but required a different judgment (amplitude modulation rate discrimination). Based on previous literature we predicted that acquisition would be disrupted by varying the task-relevant stimulus feature during training (stimulus interference), and that consolidation would be disrupted by varying the perceptual judgment required (task interference). We found that varying the task-relevant or -irrelevant stimulus features failed to disrupt acquisition but did disrupt consolidation, whereas mixing two tasks requiring a different perceptual judgment but sharing the same stimulus features disrupted both acquisition and consolidation. Thus, a distinction between acquisition and consolidation phases of perceptual learning cannot simply be attributed to (task-relevant) stimulus versus task interference. We propose instead that disruption occurs during acquisition when mixing two tasks requiring a perceptual judgment based on different cues, whereas consolidation is always disrupted regardless of whether different stimulus features or tasks are mixed. The current study not only provides a novel insight into the underlying mechanisms of perceptual learning, but also has practical implications for the optimal design and delivery of training programs that aim to remediate perceptual difficulties.     

Auditory Discrimination Learning: Role of Working Memory

Perceptual training is generally assumed to improve perception by modifying the encoding or decoding of sensory information. However, this assumption is incompatible with recent demonstrations that transfer of learning can be enhanced by across-trial variation of training stimuli or task. Here we present three lines of evidence from healthy adults in support of the idea that the enhanced transfer of auditory discrimination learning is mediated by working memory (WM). First, the ability to discriminate small differences in tone frequency or duration was correlated with WM measured with a tone n-back task. Second, training frequency discrimination around a variable frequency transferred to and from WM learning, but training around a fixed frequency did not. The transfer of learning in both directions was correlated with a reduction of the influence of stimulus variation in the discrimination task, linking WM and its improvement to across-trial stimulus interaction in auditory discrimination. Third, while WM training transferred broadly to other WM and auditory discrimination tasks, variable-frequency training on duration discrimination did not improve WM, indicating that stimulus variation challenges and trains WM only if the task demands stimulus updating in the varied dimension. The results provide empirical evidence as well as a theoretic framework for interactions between cognitive and sensory plasticity during perceptual experience.     

Second-Order Conditioning and Conditioned Inhibition: Influences of Speed versus Accuracy on Human Causal Learning

In human causal learning, excitatory and inhibitory learning effects can sometimes be found in the same paradigm by altering the learning conditions. This study aims to explore whether learning in the feature negative paradigm can be dissociated by emphasising speed over accuracy. In two causal learning experiments, participants were given a feature negative discrimination in which the outcome caused by one cue was prevented by the addition of another. Participants completed training trials either in a self-paced fashion with instructions emphasising accuracy, or under strict time constraints with instructions emphasising speed. Using summation tests in which the preventative cue was paired with another causal cue, participants in the accuracy groups correctly rated the preventative cue as if it reduced the probability of the outcome. However, participants in the speed groups rated the preventative cue as if it increased the probability of the outcome. In Experiment 1, both speed and accuracy groups later judged the same cue to be preventative in a reasoned inference task. Experiment 2 failed to find evidence of similar dissociations in retrospective revaluation (release from overshadowing vs. mediated extinction) or learning about a redundant cue (blocking vs. augmentation). However in the same experiment, the tendency for the accuracy group to show conditioned inhibition and the speed group to show second-order conditioning was consistent even across sub-sets of the speed and accuracy groups with equivalent accuracy in training, suggesting that second-order conditioning is not merely a consequence of poorer acquisition. This dissociation mirrors the trade-off between second-order conditioning and conditioned inhibition observed in animal conditioning when training is extended.     

Selective facilitative effect of post-trial reticular stimulation in discriminative learning in the rat

Ninety-eight Sprague-Dawley rats, implanted with electrodes in the mesencephalic tegmentum (reticular activating system, RAS) served as subjects in two experiments. In the first experiment (n = 42) we investigated the effects of a RAS stimulation (5 μ A, 300 Hz, 90 sec in duration) on the acquisition of a positively reinforced light–dark discrimination in a T-maze. In the second experiment (n = 56) the reinforcement and the treatment were dissociated by comparing the effects of the RAS stimulation administered after correct or incorrect choices, during the same discrimination task.In the two experiments, despite large differences in learning conditions, the results show a considerable learning facilitation by administering the RAS stimulation immediately after each trial. This facilitation does not seem to be due to an interaction between reinforcement and stimulation, since the results of experiment 2 show the maximum facilitation in animals stimulated after each (non-reinforced) error, compared to subjects stimulated after each (reinforced) correct choice. These results are discussed both in terms of consolidation processes and in terms of comparison of the cue values of S⁺ and S^- in a discriminative learning situation.     

Behavioral compensations in a positional learning and memory task by aged monkeys

The present experiment assessed learning and memory of a positional task by evaluating behavioral strategies as well as accuracy of a task in four young and four aged monkeys. They were tested in a delayed response (DR) task that has been widely used to study animal models of aging. The task consisted of two phases; an acquisition of the task and a positional memory test with five delay times (1-30 s). There was no clear difference between age groups in the number of trials needed for acquisition of the task. However, an analysis of behavior revealed differences in behavioral characteristics displayed during testing. The young monkeys showed various irrelevant behaviors during the execution of the task. In contrast, the aged monkeys consistently concentrated on the task exhibiting no behaviors irrelevant to the task. These results showed than the aged monkeys' performance was supported by a different behavioral strategy from the young monkeys. The results of the memory test were similar to those of the acquisition on the accuracy and the behavior. The aged monkeys depended on behavioral cues to preserve their positional memory, especially during the task. The present study suggests that cognitive impairments in aged monkeys can be compensated for by employing behavioral strategies.     

Sometimes slower is better: slow-exploring birds are more sensitive to changes in a vocal discrimination task

Animal personality, defined as consistent individual differences across context and time, has attracted much recent research interest in the study of animal behaviour. More recently, this field has begun to examine how such variation arose and is maintained within populations. The habitat-dependent selection hypothesis, which posits that animals with differing personality types may fare better (i.e. have a fitness advantage) in different habitats, suggests one possible mechanism. In the current experiment, we tested whether slow- and fast-exploring black-capped chickadees (Poecile atricapillus), determined by performance in a novel environment exploration task, perform differentially when the demands of an acoustic operant discrimination (cognitive) task were altered following successful task acquisition. We found that slow-exploring birds learn to reverse previously learned natural category rules more quickly than faster exploring conspecifics. In accordance with the habitat-dependent selection hypothesis, and previous work with great tits (Parus major), a close relative of the black-capped chickadee, our results suggest that fast-exploring birds may perform better in stable, predictable environments where forming a routine is advantageous, while slow-exploring birds are favoured in unstable, unpredictable environments, where task demands often change. Our results also support a hypothesis derived from previous work with great tits; slow-exploring birds may be generally more flexible (i.e. able to modify their behaviour in accordance with changes in environmental stimuli) in some learning tasks.