Domain is a moving target for relational learning

The domain for relational learning was manipulated by varying the training set size for pigeons that had learned the same/different (S/D) concept. Six pigeons that had learned a S/D task with pairs of pictures with a set size of 1024 picture items had their training set size reduced to 8 items. Training on the reduced 8-item set was followed by transfer testing that was repeated four times. Transfer performance following reduction of the training set to 8 items was 9.2% less than it had been when the pigeons were trained with the 1024-item set, but 25.8% above chance. This partial abstract-concept learning remained constant over the four tests with novel stimuli. The results show that a broad domain established by a large expanding training set can once again become restricted by further training with a small training set.     

Superior abstract-concept learning by Clark's nutcrackers (Nucifraga columbiana)

The ability to learn abstract relational concepts is fundamental to higher level cognition. In contrast to item-specific concepts (e.g. pictures containing trees versus pictures containing cars), abstract relational concepts are not bound to particular stimulus features, but instead involve the relationship between stimuli and therefore may be extrapolated to novel stimuli. Previous research investigating the same/different abstract concept has suggested that primates might be specially adapted to extract relations among items and would require fewer exemplars of a rule to learn an abstract concept than non-primate species. We assessed abstract-concept learning in an avian species, Clark''s nutcracker (Nucifraga columbiana), using a small number of exemplars (eight pairs of the same rule, and 56 pairs of the different rule) identical to that previously used to compare rhesus monkeys, capuchin monkeys and pigeons. Nutcrackers as a group (N = 9) showed more novel stimulus transfer than any previous species tested with this small number of exemplars. Two nutcrackers showed full concept learning and four more showed transfer considerably above chance performance, indicating partial concept learning. These results show that the Clark''s nutcracker, a corvid species well known for its amazing feats of spatial memory, learns the same/different abstract concept better than any non-human species (including non-human primates) yet tested on this same task.     

Functional relationships for determining similarities and differences in comparative cognition

Pigeons, capuchin monkeys and rhesus monkeys were trained in nearly identical same/different tasks with an expanding 8-item training set and showed qualitatively similar functional relationships: increasing novel-stimulus transfer (i.e., concept learning) as a function of the training-set size and the level of transfer eventually becoming equivalent to baseline training performance. There were also some quantitative functional differences: pigeon transfer increases were more gradual and baseline-equivalent transfer occurred at a larger set size (256 items) than for monkeys (128 items). Other pigeon groups trained at 32 and 64-item initial set sizes showed improved transfer (relative to expanding the 8-item training set), equivalent to the monkey species’ transfer at these same training set sizes. This finding of equivalent concept learning over a portion of the functional range (8, 32, and 64 items or 64-4096 training pairs) is discussed in terms of species differences: carryover effects from smaller-set training, evolved neural systems, cognitive and cortical modules, and general distributed learning systems for “higher-order” cognitive abilities.     

Conceptual thresholds for same and different in old-(Macaca mulatta) and new-world (Cebus apella) monkeys

Learning of the relational same/different (S/D) concept has been demonstrated to be largely dependent upon stimulus sets containing more than two items for pigeons and old-world monkeys. Stimulus arrays containing several images for use in same/different discrimination procures (e.g. 16 identical images vs. 16 nonidentical images) have been shown to facilitate and even be necessary for learning of relational concepts ( [Flemming et al., 2007], [Wasserman et al., 2001] and [Young et al., 1997]). In the present study, we investigate the threshold at which a new world primate, the capuchin (Cebus apella) may be able to make such a discrimination. Utilizing a method of increasing entropy, rather than conventional procedures of decreasing entropy, we demonstrate unique evidence that capuchin monkeys are readily capable of making 2-item relational S/D conditional discriminations. In another experiment, we examine the supposed level of difficulty in making S/D discriminations by rhesus monkeys (Macaca mulatta). Whereas pigeons (Columba livia) and baboons (Papio papio) have shown marked difficulty simultaneously discriminating same from different arrays at all when composed of fewer than 8 items each, rhesus monkeys seem to understand that pairs of stimuli connote sameness and difference just the same (Flemming et al., 2007). With sustained accurate performance of 2-item S/D discriminations, both experienced and task-naïve rhesus monkeys appear quite certain in their conceptual knowledge of same and different. We conclude that learning of the same/different relational concept may be less dependent upon high levels of entropy contrast than originally hypothesized for nonhuman primates.     

Conceptualization of above and below relationships by an insect

Relational rules such as 'same' or 'different' are mastered by humans and non-human primates and are considered as abstract conceptual thinking as they require relational learning beyond perceptual generalization. Here, we investigated whether an insect, the honeybee (Apis mellifera), can form a conceptual representation of an above/below spatial relationship. In experiment 1, bees were trained with differential conditioning to choose a variable target located above or below a black bar that acted as constant referent throughout the experiment. In experiment 2, two visual stimuli were aligned vertically, one being the referent, which was kept constant throughout the experiment, and the other the target, which was variable. In both experiments, the distance between the target and the referent, and their location within the visual field was systematically varied. In both cases, bees succeeded in transferring the learned concept to novel stimuli, preserving the trained spatial relation, thus showing an ability to manipulate this relational concept independently of the physical nature of the stimuli. Absolute location of the referent into the visual field was not a low-level cue used by the bees to solve the task. The honeybee is thus capable of conceptual learning despite having a miniature brain, showing that such elaborated learning form is not a prerogative of vertebrates.     

Same/different concept learning by capuchin monkeys in matching-to-sample tasks

The ability to understand similarities and analogies is a fundamental aspect of human advanced cognition. Although subject of considerable research in comparative cognition, the extent to which nonhuman species are capable of analogical reasoning is still debated. This study examined the conditions under which tufted capuchin monkeys (Cebus apella) acquire a same/different concept in a matching-to-sample task on the basis of relational similarity among multi-item stimuli. We evaluated (i) the ability of five capuchin monkeys to learn the same/different concept on the basis of the number of items composing the stimuli and (ii) the ability to match novel stimuli after training with both several small stimulus sets and a large stimulus set. We found the first evidence of same/different relational matching-to-sample abilities in a New World monkey and demonstrated that the ability to match novel stimuli is within the capacity of this species. Therefore, analogical reasoning can emerge in monkeys under specific training conditions.     

Dynamic Sensorimotor Planning during Long-Term Sequence Learning: The Role of Variability,Response Chunking and Planning Errors

Many everyday skills are learned by binding otherwise independent actions into a unified sequence of responses across days or weeks of practice. Here we looked at how the dynamics of action planning and response binding change across such long timescales. Subjects (N = 23) were trained on a bimanual version of the serial reaction time task (32-item sequence) for two weeks (10 days total). Response times and accuracy both showed improvement with time, but appeared to be learned at different rates. Changes in response speed across training were associated with dynamic changes in response time variability, with faster learners expanding their variability during the early training days and then contracting response variability late in training. Using a novel measure of response chunking, we found that individual responses became temporally correlated across trials and asymptoted to set sizes of approximately 7 bound responses at the end of the first week of training. Finally, we used a state-space model of the response planning process to look at how predictive (i.e., response anticipation) and error-corrective (i.e., post-error slowing) processes correlated with learning rates for speed, accuracy and chunking. This analysis yielded non-monotonic association patterns between the state-space model parameters and learning rates, suggesting that different parts of the response planning process are relevant at different stages of long-term learning. These findings highlight the dynamic modulation of response speed, variability, accuracy and chunking as multiple movements become bound together into a larger set of responses during sequence learning.     

Auditory Same/Different Concept Learning and Generalization in Black-Capped Chickadees (Poecile atricapillus)

Abstract concept learning was thought to be uniquely human, but has since been observed in many other species. Discriminating same from different is one abstract relation that has been studied frequently. In the current experiment, using operant conditioning, we tested whether black-capped chickadees (Poecile atricapillus) could discriminate sets of auditory stimuli based on whether all the sounds within a sequence were the same or different from one another. The chickadees were successful at solving this same/different relational task, and transferred their learning to same/different sequences involving novel combinations of training notes and novel notes within the range of pitches experienced during training. The chickadees showed limited transfer to pitches that was not used in training, suggesting that the processing of absolute pitch may constrain their relational performance. Our results indicate, for the first time, that black-capped chickadees readily form relational auditory same and different categories, adding to the list of perceptual, behavioural, and cognitive abilities that make this species an important comparative model for human language and cognition.     

Absolute and relational control of a sequential auditory discrimination by pigeons (Columba livia)

Recent evidence indicates that pigeons can readily learn visual discriminations based on both absolute and relational stimulus factors. To examine how these two types of control function in their non-dominant auditory modality, we tested four pigeons in a go/no-go sequential auditory discrimination in which both absolute and relational cues were redundantly available. In this task, sequences of different sounds created from one set of pitches were reinforced, while different sequences created from another set of pitches and any same sequences made from either set of pitches were not. Across three experiments, we independently varied the relative discriminability of the absolute and relational components. The pigeons were consistently and primarily controlled by the absolute fundamental pitch of our notes in all of the experiments, although this was influenced by the range and arrangement of the pitches used in each set. A majority of the pigeons also demonstrated relational control when this component was made more salient. The more robust control exhibited by absolute factors is consistent with the comparative hypothesis that birds in general may have a well-developed aptitude for processing absolute pitch in many auditory settings. The relational control is consistent with our recent evidence of same/different auditory learning by pigeons.     

Behavioral and anatomical consequences of early versus late symbol training in macaques

Distinct brain regions, reproducible from one person to the next, are specialized for processing different kinds of human expertise, such as face recognition?and reading. Here, we explore the relationship between age of learning, learning ability, and specialized brain structures. Specifically, we ask whether the existence of reproducible cortical domains necessarily means that certain abilities are innate, or innately easily learned, or whether reproducible domains can be formed, or refined, by interactions between genetic programs and common early experience. Functional MRI showed that intensive early, but not late, experience caused the formation of category-selective regions in macaque temporal lobe for stimuli never naturally encountered by monkeys. And behaviorally, early training produced more fluent processing of these stimuli than the same training in adults. One explanation for these results is that in higher cortical areas, as in early sensory areas, experience drives functional clustering and functional clustering determines how that information is processed.     

Intermodal transfer from a visual to an auditory discrimination using an errorless learning procedure

Errorless learning is a technique developed by Terrace [Terrace, H.S., 1963a. Discrimination training with and without “errors”. J. Exp. Anal. Behav. 6, 1-27] to train stimulus discriminations with few or no errors. In the first replication of the original findings, errorless learning was also shown to transfer successfully between two visual discriminations without errors [Terrace, H.S., 1963b. Errorless transfer of a discrimination across two continua. J. Exp. Anal. Behav. 6, 223-232]. In the present experiment, we extended the errorless learning procedure to an intermodal transfer, from a discrimination between red and green colors to a discrimination between high and low tones. The pigeons were divided into two groups: an Experimental Group, which learned both discriminations through errorless learning, and a Control Group, which learned them through trial-and-error. Results showed that pigeons from the Experimental Group learned the red-green discrimination with significantly fewer errors than the Control Group and that errorless learning is effective in transferring from a visual to an auditory discrimination.     

Orientation discrimination and categorization of photographs of natural objects by pigeons

In Experiment 1, pigeons trained to discriminate rightside-up and upside-down orientations of slides of natural scenes with humans successfully transferred to new slides of the same kind. Experiment 2 revealed that both the orientations of the human figures and of the background scenes controlled the discrimination. When they were oppositely oriented, the background orientation cue was dominant. In Experiment 3 slides showing objects on a white background were presented either rightside up or upside down, with each slide presented in one orientation only. One group of pigeons learned to classify the slides according to their orientations. The other group learned to classify the slides according to arbitrary groupings. When the slides were shown rotated by 180 degrees, the latter group continued to discriminate the individual slides (i.e., the pigeons showed orientation invariance). The former group classified the rotated slides according to their orientations (i.e., orientation discrimination). In Experiment 4, pigeons learned the orientation discrimination with separate sets of human and bird figures. Partial reversal training in one object class transferred to the rest of stimuli in this object class but did not to the other object class. These results suggest that pigeons can learn to discriminate photographs on the basis of orientation but that orientation-based equivalence relationship is not formed between object classes.     

Fast-TIPL occurs for salient images without a memorization requirement in men but not in women

Recent research of task-irrelevant perceptual learning (TIPL) demonstrates that stimuli that are consistently presented at relevant point in times (e.g. with task-targets or rewards) are learned, even in the absence of attention to these stimuli. However, different research paradigms have observed different results for how salient stimuli are learned; with some studies showing no learning, some studies showing positive learning and others showing negative learning effects. In this paper we focused on how the level of processing of stimuli impacts fast-TIPL. We conducted three different experiments in which the level of processing of the information paired with a target was manipulated. Our results indicated that fast-TIPL occurs when participants have to memorize the information presented with the target, but also when they just have to process this information for a secondary task without an explicit memorization of those stimuli. However, fast-TIPL does not occur when participants have to ignore the target-paired information. This observation is consistent with recent models of TIPL that suggest that attentional signals can either enhance or suppress learning depending on whether those stimuli are distracting or not to the subjects' objectives. Our results also revealed a robust gender effect in fast-TIPL, where male subjects consistently show fast-TIPL, whereas the observation of fast-TIPL is inconsistent in female subjects.     

Transfer of the Nonmatch-to-Goal rule in Monkeys across Cognitive Domains

To solve novel problems, it is advantageous to abstract relevant information from past experience to transfer on related problems. To study whether macaque monkeys were able to transfer an abstract rule across cognitive domains, we trained two monkeys on a nonmatch-to-goal (NMTG) task. In the object version of the task (O-NMTG), the monkeys were required to choose between two object-like stimuli, which differed either only in shape or in shape and color. For each choice, they were required to switch from their previously chosen object-goal to a different one. After they reached a performance level of over 90% correct on the O-NMTG task, the monkeys were tested for rule transfer on a spatial version of the task (S-NMTG). To receive a reward, the monkeys had to switch from their previously chosen location to a different one. In both the O-NMTG and S-NMTG tasks, there were four potential choices, presented in pairs from trial-to-trial. We found that both monkeys transferred successfully the NMTG rule within the first testing session, showing effective transfer of the learned rule between two cognitive domains.     

Natural conceptual behavior in squirrel monkeys (saimiri sciureus): An experimental investigation

Natural conceptual discriminations have been tested in many different species, including pigeons and a variety of non-human primates. The ability of four male squirrel monkeys (Saimiri sciureus) to learn and use the natural concept ‘squirrel monkey’ was investigated in this study. After a training phase, subjects were presented with novel stimuli in transfer and test trials. All subjects performed at a rate significantly above chance on the first test trial (p<.001), indicating that squirrel monkeys can utilize natural concepts in the laboratory.     

Prefrontal cortex activity related to abstract response strategies

Many monkeys adopt abstract response strategies as they learn to map visual symbols to responses by trial and error. According to the repeat-stay strategy, if a symbol repeats from a previous, successful trial, the monkeys should stay with their most recent response choice. According to the change-shift strategy, if the symbol changes, the monkeys should shift to a different choice. We recorded the activity of prefrontal cortex neurons while monkeys chose responses according to these two strategies. Many neurons had activity selective for the strategy used. In a subsequent block of trials, the monkeys learned fixed stimulus-response mappings with the same stimuli. Some neurons had activity selective for choosing responses based on fixed mappings, others for choosing based on abstract strategies. These findings indicate that the prefrontal cortex contributes to the implementation of the abstract response strategies that monkeys use during trial-and-error learning.     

Attention, the presolution period, and choice accuracy in pigeons

Six pigeons were trained first on a color then on a form discrimination; four other pigeons were trained first on form and then on color. One of two colors or one of two forms (sample stimuli) appeared in the center of a touch sensitive monitor for 5 pigeons and in the center and in 16 other locations for 5 other pigeons. A peck anywhere within the region in which the sample stimuli appeared produced two white disks (comparison stimuli), one on the left and one on the lower right corners of the screen. Correct left-right choices provided food. Although of no consequence, the location of pecks in presence of the sample was predictive of the pigeon's subsequent choice. Accuracy, choice of the correct comparison stimulus, was greater when the sample stimuli appeared in the center as well as 16 other locations than when it appeared only in the center. The presolution period, the period of chance accuracy prior to evidence of discrimination learning, was decreased on each task following training on the other task. This evidence of facilitation following an extra-dimensional shift was attributed to continued relevance of the conditions under which the first task was learned. The duration of the presolution period was inversely related to asymptotic accuracy-data accounted for by Heinemann's (1983) theory of information processing during the presolution period.     

Effects of stimulus size and spatial organization on pigeons’ conditional same-different discrimination

In two experiments, we explored the effects of varying the size and the spatial organization of the stimuli in multi-item arrays on pigeons’ same-different discrimination behavior. The birds had previously learned to discriminate a simultaneously presented array of 16 identical (Same) visual items from an array of 16 nonidentical (Different) visual items, when the correct choice was conditional on the presence of another cue: the color of the background (Castro et al., in press). In Experiment 1, we trained pigeons with 7-item arrays and then tested them with arrays containing the same item, but in a variety of sizes. In Experiment 2, we tested the birds with the items grouped in novel locations: the top, the bottom, the left, or the right portions of the display area, which generated different vertical and horizontal alignments. Accuracy scores revealed virtually perfect stimulus generalization across various item sizes and spatial organizations. Reaction times revealed that the birds perceived different sizes of a single icon as the same stimulus (Experiment 1) and that the birds processed vertical arrangements faster than horizontal arrangements (Experiment 2). These results suggest that the pigeons noticed both physical and spatial changes in the stimuli (as shown by their reaction times), but that these changes did not disrupt the birds’ discriminating the sameness or differentness of the multi-item arrays (as shown by their accuracy scores).     

Analysis of the effect of a training regime for the correct choice of matched learning in gray crows

It was shown that a large set of training stimuli promotes abstract concept learning. These experiments were designed to assess whether an application of a large set of training stimuli would facilitate matching learning in crows. Four hooded crows were trained with a set of 72 unique combinations of stimuli in two-alternative simultaneous matching tasks with stimuli of three different categories: achromatic color (white, light-grey, dark-grey, and black), shape (Arabic numerals from 1 to 4 used as visual shapes only), and number of elements (heterogeneous graphic arrays from 1 to 4 items). Although the performance of all crows was significantly above chance (p < 0.01) in some 72-trial blocks, birds were unable to establish matching and to reach the criterion of learning 80% correct or better over 72 consecutive trials) in 5184 trials. Thus, the modified training procedure was less efficient than the training technique previously used (successive cyclic repetition of three small sets of training stimuli), which allowed four of six crows to acquire the matching rule after 1780, 2360, 3830, and 5260 trials [4,9].     

Long-term Retention of Many Visual Patterns by Pigeons

Using a simultaneous discrimination procedure it was shown that pigeons were capable of learning to discriminate 100 different black and white visual patterns from a further 625 similar stimuli, where responses to the former were rewarded and responses to the latter were not rewarded. Tests in which novel stimuli replaced either the rewarded or nonrewarded stimuli showed that the pigeons had not only learned about the 100 positive stimuli but also about the 625 negative stimuli. The fact that novel stimuli enhanced discrimination performance when they replaced the many negative stimuli indicated that the pigeons had categorized the stimuli into two classes, familiar and less familiar. Long-term retention was examined after a 6-month interval. To begin with it seemed poor but a recognition test performed after the subjects had been retrained with a subset of the stimuli after an interval of 7 months confirmed that pigeons are capable of retaining in memory several 100 visual items over an extended period. It is proposed that the initial retrieval weakness was due to a forgetting of the contingencies between stimulus categories and response outcomes. Further tests involving variously modified stimuli indicated that while stimulus size variations had a negative effect on performance, orientation changes did not interfere with recognition, supporting the view that small visual stimuli are memorized by pigeons largely free of orientation labels. The experiment generally confirms that pigeons have the capacity of storing information about a large number of visual stimuli over long periods of time.