The problem of using correlation methods in brain function]

In a series of publications, the author used the correlation hypothesis to explain the main mechanisms of functioning of bat and dolphin echo locators. A good fit of calculated parameters to experimental data was shown. In this work, by the example of generalization of the recognition problem, the use by the brain of correlation methods for solving more general problems independently on the modality of sensor signals was analyzed. In favor of the hypothesis that correlation methods are widely used in brain functioning are the data presented in the paper, which prove that an associative neuron is a suitable analog computer. It is suited for rapid "computation" of the intercorrelation function of discrete input and reference signals. The set of weighting coefficients of neuron synaptic inputs serve as such signals. The pool of associative neurons determines the values on the correlation function in the required range by changing the "numbers" of inputs from neuron to neuron at which discrete signals arrive.     

The formation in the northern fur seal (Callorhinus ursinus L.) of flexible behavioral programs determined by the structure of the complex visual stimulus

In Callorhinus ursinus L. generalized rule was elaborated of perception of two-stimuli complex visual stimulus (CVS) presented in random and formed from a set of three differentiated stimuli. The animal was trained to perceive the visual information in linear sequence according to the rule of "reading" from top to bottom and make in conformity with this rule labile two-link visual-motor chains of actions signalled by CVS stimuli. Such behavioural strategies were called by authors ability to operate with CVS structure.     

Interaction of Different Types of Temporary Connections in the Course of Conditioning to Stimulus Ratio

Our studies have shown that the physiological basis of the generalization function is the development of systemic activity. From this point of view, conditioned reflexes in response to ratios of stimuli are of interest. These reflexes are one of the simplest forms of the integrating function in which the reflection of reality in generalized form takes place without the interposition of the spoken word. Here the conditioned stimulus is not the individual stimulus, but the ratio of two stimuli, which establishes a certain degree of abstraction from reality (Kleshchov [3], Dolin [2]).     

Intensity generalization: physiology and modelling of a neglected topic

I briefly review empirical data about the generalization of acquired behaviour to novel stimuli, showing that variations in stimulus intensity affect behaviour differently from variations in characteristics such as, for instance, visual shape or sound frequency. I argue that such differences can be seen already in how the sense organs react to changes in intensity compared to changes in other stimulus characteristics. I then evaluate a number of models of generalization with respect to their ability to reproduce intensity generalization. I reach three main conclusions. First, realistic stimulus representations, based on knowledge of the sense organs, are necessary to account for intensity effects. Models employing stimulus representations too remote from the sense organs are unable to reproduce the data. Second, the intuitive notion that generalization is based on similarities between stimuli, possibly modelled as distances in an appropriate representation space, is difficult to reconcile with data about intensity generalization. Third, several simple models, in conjunction with realistic stimulus representations, can account for a wide array of generalization phenomena along both intensity and non-intensity stimulus dimensions. The paper also introduces concepts which may be generally useful to evaluate and compare different models of behaviour.     

Invariant recognition of feature combinations in the visual system

 The operation of a hierarchical competitive network model (VisNet) of invariance learning in the visual system is investigated to determine how this class of architecture can solve problems that require the spatial binding of features. First, we show that VisNet neurons can be trained to provide transform-invariant discriminative responses to stimuli which are composed of the same basic alphabet of features, where no single stimulus contains a unique feature not shared by any other stimulus. The investigation shows that the network can discriminate stimuli consisting of sets of features which are subsets or supersets of each other. Second, a key feature-binding issue we address is how invariant representations of low-order combinations of features in the early layers of the visual system are able to uniquely specify the correct spatial arrangement of features in the overall stimulus and ensure correct stimulus identification in the output layer. We show that output layer neurons can learn new stimuli if the lower layers are trained solely through exposure to simpler feature combinations from which the new stimuli are composed. Moreover, we show that after training on the low-order feature combinations which are common to many objects, this architecture can – after training with a whole stimulus in some locations – generalise correctly to the same stimulus when it is shown in a new location. We conclude that this type of hierarchical model can solve feature-binding problems to produce correct invariant identification of whole stimuli. Received: 4 August 1999 / Accepted in revised form: 11 October 2000     

Bayesian calibration of simultaneity in audiovisual temporal order judgments

After repeated exposures to two successive audiovisual stimuli presented in one frequent order, participants eventually perceive a pair separated by some lag time in the same order as occurring simultaneously (lag adaptation). In contrast, we previously found that perceptual changes occurred in the opposite direction in response to tactile stimuli, conforming to bayesian integration theory (bayesian calibration). We further showed, in theory, that the effect of bayesian calibration cannot be observed when the lag adaptation was fully operational. This led to the hypothesis that bayesian calibration affects judgments regarding the order of audiovisual stimuli, but that this effect is concealed behind the lag adaptation mechanism. In the present study, we showed that lag adaptation is pitch-insensitive using two sounds at 1046 and 1480 Hz. This enabled us to cancel lag adaptation by associating one pitch with sound-first stimuli and the other with light-first stimuli. When we presented each type of stimulus (high- or low-tone) in a different block, the point of simultaneity shifted to "sound-first" for the pitch associated with sound-first stimuli, and to "light-first" for the pitch associated with light-first stimuli. These results are consistent with lag adaptation. In contrast, when we delivered each type of stimulus in a randomized order, the point of simultaneity shifted to "light-first" for the pitch associated with sound-first stimuli, and to "sound-first" for the pitch associated with light-first stimuli. The results clearly show that bayesian calibration is pitch-specific and is at work behind pitch-insensitive lag adaptation during temporal order judgment of audiovisual stimuli.     

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).     

Auditory pre-experience modulates classification of affect intensity: evidence for the evaluation of call salience by a non-human mammal,the bat <Emphasis Type="Italic">Megaderma lyra</Emphasis>

Introduction

Immediate responses towards emotional utterances in humans are determined by the acoustic structure and perceived relevance, i.e. salience, of the stimuli, and are controlled via a central feedback taking into account acoustic pre-experience. The present study explores whether the evaluation of stimulus salience in the acoustic communication of emotions is specifically human or has precursors in mammals. We created different pre-experiences by habituating bats (Megaderma lyra) to stimuli based on aggression, and response, calls from high or low intensity level agonistic interactions, respectively. Then we presented a test stimulus of opposite affect intensity of the same call type. We compared the modulation of response behaviour by affect intensity between the reciprocal experiments.

Results

For aggression call stimuli, the bats responded to the dishabituation stimuli independent of affect intensity, emphasising the attention-grabbing function of this call type. For response call stimuli, the bats responded to a high affect intensity test stimulus after experiencing stimuli of low affect intensity, but transferred habituation to a low affect intensity test stimulus after experiencing stimuli of high affect intensity. This transfer of habituation was not due to over-habituation as the bats responded to a frequency-shifted control stimulus. A direct comparison confirmed the asymmetric response behaviour in the reciprocal experiments.

Conclusions

Thus, the present study provides not only evidence for a discrimination of affect intensity, but also for an evaluation of stimulus salience, suggesting that basic assessment mechanisms involved in the perception of emotion are an ancestral trait in mammals.

     

The effect of study-test modalities on the remembrance of subjective duration from long-term memory

It was examined whether stimulus modality (auditory vs. visual) affects the retrieval of subjective duration from memory. In two experiments the temporal generalization paradigm was used. Participants had to decide whether the previously learned standard duration (400 ms) occurred in the context of comparison stimuli. Two major results were found. (1) Discrimination was more accurate if the training and testing stimuli were of the same modality than if they were of opposite modalities. (2) If both modality of learning and modality of testing were different, subjects systematically underestimated the test durations, i.e. temporal generalization gradients (the proportion of identifications of a stimulus as the standard, plotted against stimulus duration) shifted to the right. The observed shift is interpreted as a result of a delayed timing process.     

Discrimination of phase spectra in complex sounds by the bullfrog (Rana catesbeiana)

1. Male bullfrogs at two different natural calling sites were presented with playbacks of synthetic advertisement calls differing in phase spectra. Sounds were presented in a ABA design to analyze the ability of the animals to perceive changes in repeated series of stimuli.
2. The number of individual croaks in an answering call significantly increased over repeated presentations of two of the three stimulus phase types in condition A1. There were significantly fewer croaks to the third stimulus. These data suggest that two stimuli were perceived in a similar manner.
3. Latency of calling to stimuli presented in conditions A and B changed in response to shifts in phase spectrum at a low density calling site. These differences were significant when comparing latency to playbacks where shifts in the phase spectrum changed the temporal fine-structure and waveform periodicity of the stimulus.
4. The increase in number of croaks and decrease in response latency across condition A1 and the increase in latency in condition B suggest that discrimination may take the form of stimulus-specific sensitization. In this context, sensitization might reflect an increase in arousal due to repeated presentation of a salient stimulus.
5. The operation of a hypothetical ‘mating call detector,’ based on linear summation of temporal responses from the eighth nerve, provides output similar to the behavioral results.

     

Behavioral determination of stimulus pair discrimination of auditory acoustic and electrical stimuli using a classical conditioning and heart-rate approach

Acute animal preparations have been used in research prospectively investigating electrode designs and stimulation techniques for integration into neural auditory prostheses, such as auditory brainstem implants and auditory midbrain implants. While acute experiments can give initial insight to the effectiveness of the implant, testing the chronically implanted and awake animals provides the advantage of examining the psychophysical properties of the sensations induced using implanted devices. Several techniques such as reward-based operant conditioning, conditioned avoidance, or classical fear conditioning have been used to provide behavioral confirmation of detection of a relevant stimulus attribute. Selection of a technique involves balancing aspects including time efficiency (often poor in reward-based approaches), the ability to test a plurality of stimulus attributes simultaneously (limited in conditioned avoidance), and measure reliability of repeated stimuli (a potential constraint when physiological measures are employed). Here, a classical fear conditioning behavioral method is presented which may be used to simultaneously test both detection of a stimulus, and discrimination between two stimuli. Heart-rate is used as a measure of fear response, which reduces or eliminates the requirement for time-consuming video coding for freeze behaviour or other such measures (although such measures could be included to provide convergent evidence). Animals were conditioned using these techniques in three 2-hour conditioning sessions, each providing 48 stimulus trials. Subsequent 48-trial testing sessions were then used to test for detection of each stimulus in presented pairs, and test discrimination between the member stimuli of each pair. This behavioral method is presented in the context of its utilisation in auditory prosthetic research. The implantation of electrocardiogram telemetry devices is shown. Subsequent implantation of brain electrodes into the Cochlear Nucleus, guided by the monitoring of neural responses to acoustic stimuli, and the fixation of the electrode into place for chronic use is likewise shown.     

Lateralized visual behavior in bottlenose dolphins (Tursiops truncatus) performing audio-visual tasks: the right visual field advantage

Analyzing cerebral asymmetries in various species helps in understanding brain organization. The left and right sides of the brain (lateralization) are involved in different cognitive and sensory functions. This study focuses on dolphin visual lateralization as expressed by spontaneous eye preference when performing a complex cognitive task; we examine lateralization when processing different visual stimuli displayed on an underwater touch-screen (two-dimensional figures, three-dimensional figures and dolphin/human video sequences). Three female bottlenose dolphins (Tursiops truncatus) were submitted to a 2-, 3- or 4-, choice visual/auditory discrimination problem, without any food reward: the subjects had to correctly match visual and acoustic stimuli together. In order to visualize and to touch the underwater target, the dolphins had to come close to the touch-screen and to position themselves using monocular vision (left or right eye) and/or binocular naso-ventral vision. The results showed an ability to associate simple visual forms and auditory information using an underwater touch-screen. Moreover, the subjects showed a spontaneous tendency to use monocular vision. Contrary to previous findings, our results did not clearly demonstrate right eye preference in spontaneous choice. However, the individuals' scores of correct answers were correlated with right eye vision, demonstrating the advantage of this visual field in visual information processing and suggesting a left hemispheric dominance. We also demonstrated that the nature of the presented visual stimulus does not seem to have any influence on the animals' monocular vision choice.     

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.     

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.     

Study of lever-holding phenomenon during the escape reaction in rats

A "preparatory" hypothesis according to which the bar-holding develops as a preparatory response placing an animal in an advantageous position for the reaction on the subsequent action of the electric current has been experimentally examined. Comparing the avoidance training under the condition allowing the bar-holding until the next conditioned and unconditioned stimuli, and under conditions excepting such long holding it has been established that in first case the number of bar-holding responses is more and the avoidance responses are less than in the second one. It contradicts the "preparatory" hypothesis. It has been shown that the dynamics of inter-trail and the bar-holding responses results from stimulus generalization and discrimination.     

Simulation of habituation to simple and multiple stimuli

Within the psychological literature there are a number of models that reproduce the defining properties of habituation to a single stimulus. However, most of them do not reproduce the phenomenon of dishabituation shown in empirical studies, consisting in the recovery of a stimulus previously habituated upon the appearance of a novel stimulus. The present work offers a model of habituation which, in addition to reproducing the basic properties of habituation to a stimulus, also does so when more than one stimulus is presented, and thus includes the dishabituation phenomenon. This model consists of two functions, one called "activation" and the other "availability", and is tested by means of simulation of the responses in the context of different stimulus patterns. The results of the simulation show a good qualitative fit to the empirical results on the phenomena of habituation, including dishabituation. In addition, the model is suitable for inclusion in associative models that reproduce classical conditioning, which will make it possible in the future to incorporate into these in a simple way the influence that the habituation of each stimulus may have on its association with other stimuli.     

Hill Coefficients of a Polymodal Monod-Wyman-Changeux Model for Ion Channel Gating

Allosteric transitions of ion channels can be driven by multiple sources of free energies. One class of model for describing such transitions is the multistimulus Monod-Wyman-Changeux model, in which each stimulus interacts with a specific sensor on the protein and activation of the sensor is allosterically coupled to conformational changes of the protein. In general, when a protein is stressed by multiple stimuli, one stimulus can influence the response to another, which can result in both a shift of the midpoint of the dose-response curve and a change of the slope of the curve. Here I show that, for a Monod-Wyman-Changeux model with independent sensors, the different dose-response curves of open probability for one stimulus have the same slope at the same agonist concentration. In the other words, the slope of the dose-response curve for one stimulus is an intrinsic property of the sensors for that stimulus; it is independent of other stimuli or their sensor properties. As the dose-response curve for many receptors can be fit to a Boltzmann or Hill equation, this property provides a practical, usable test for applicability of such models.     

Common marmosets are sensitive to simple dependencies at variable distances in an artificial grammar

Recognizing that two elements within a sequence of variable length depend on each other is a key ability in understanding the structure of language and music. Perception of such interdependencies has previously been documented in chimpanzees in the visual domain and in human infants and common squirrel monkeys with auditory playback experiments, but it remains unclear whether it typifies primates in general. Here, we investigated the ability of common marmosets (Callithrix jacchus) to recognize and respond to such dependencies. We tested subjects in a familiarization-discrimination playback experiment using stimuli composed of pure tones that either conformed or did not conform to a grammatical rule. After familiarization to sequences with dependencies, marmosets spontaneously discriminated between sequences containing and lacking dependencies (‘consistent’ and ‘inconsistent’, respectively), independent of stimulus length. Marmosets looked more often to the sound source when hearing sequences consistent with the familiarization stimuli, as previously found in human infants. Crucially, looks were coded automatically by computer software, avoiding human bias. Our results support the hypothesis that the ability to perceive dependencies at variable distances was already present in the common ancestor of all anthropoid primates (Simiiformes).     

Visual artificial grammar learning: comparative research on humans, kea (Nestor notabilis) and pigeons (Columba livia)

Artificial grammar learning (AGL) provides a useful tool for exploring rule learning strategies linked to general purpose pattern perception. To be able to directly compare performance of humans with other species with different memory capacities, we developed an AGL task in the visual domain. Presenting entire visual patterns simultaneously instead of sequentially minimizes the amount of required working memory. This approach allowed us to evaluate performance levels of two bird species, kea (Nestor notabilis) and pigeons (Columba livia), in direct comparison to human participants. After being trained to discriminate between two types of visual patterns generated by rules at different levels of computational complexity and presented on a computer screen, birds and humans received further training with a series of novel stimuli that followed the same rules, but differed in various visual features from the training stimuli. Most avian and all human subjects continued to perform well above chance during this initial generalization phase, suggesting that they were able to generalize learned rules to novel stimuli. However, detailed testing with stimuli that violated the intended rules regarding the exact number of stimulus elements indicates that neither bird species was able to successfully acquire the intended pattern rule. Our data suggest that, in contrast to humans, these birds were unable to master a simple rule above the finite-state level, even with simultaneous item presentation and despite intensive training.