Episodic-like memory: pigeons can report location pecked when unexpectedly asked

Pigeons were tested for their ability to report the location they recently pecked, without prior experience having to do so. They were first pretrained to report the location that they had just pecked. They were then trained on a conditional discrimination to associate yellow and blue samples with vertical and horizontal comparisons, respectively, independent of comparison location. On probe trials in testing, when after choosing a vertical or horizontal line following the yellow or blue sample, the pigeons were ‘asked’ which location they had just pecked, they showed a significant tendency to choose correctly in spite of the fact that location of the correct comparison was incidental to the task. Performing on probe trials is analogous to asking the pigeons an unexpected question about their recent behavior and it is similar to the episodic memory question asked of humans, “What did you have for breakfast this morning?”.     

Oscillations following periodic reinforcement

Three experiments examined behavior in extinction following periodic reinforcement. During the first phase of Experiment 1, four groups of pigeons were exposed to fixed interval (FI 16 s or FI 48 s) or variable interval (VI 16 s or VI 48 s) reinforcement schedules. Next, during the second phase, each session started with reinforcement trials and ended with an extinction segment. Experiment 2 was similar except that the extinction segment was considerably longer. Experiment 3 replaced the FI schedules with a peak procedure, with FI trials interspersed with non-food peak interval (PI) trials that were four times longer. One group of pigeons was exposed to FI 20 s PI 80 s trials, and another to FI 40 s PI 160 s trials. Results showed that, during the extinction segment, most pigeons trained with FI schedules, but not with VI schedules, displayed pause-peck oscillations with a period close to, but slightly greater than the FI parameter. These oscillations did not start immediately after the onset of extinction. Comparing the oscillations from Experiments 1 and 2 suggested that the alternation of reconditioning and re-extinction increases the reliability and earlier onset of the oscillations. In Experiment 3 the pigeons exhibited well-defined pause-peck cycles since the onset of extinction. These cycles had periods close to twice the value of the FI and lasted for long intervals of time. We discuss some hypotheses concerning the processes underlying behavioral oscillations following periodic reinforcement.     

Successive two-item same-different discrimination and concept learning by pigeons

Four pigeons were trained in a successive same/different procedure involving the alternation of two stimuli per trial. Using a go/no-go procedure, two different or two identical color photographs were alternated, with a brief, dark, inter-stimulus interval, on a computer screen for 20s. Pigeons learned to discriminate between same (S+) and different (D-) sequences with moderate to large contrasts between successive pictures. Analyses of pecking behavior within single trials revealed this discrimination emerged at the earliest possible point in the sequence (i.e. by the presentation of the second item). Pigeons transferred to novel color and gray-scale pictures, and showed savings in tests with novel video stimuli. These results suggest that same/different discrimination and concept formation can be acquired with successively presented pairs of stimuli by pigeons. When combined with results using simultaneous same/different presentations, these findings further support a qualitative similarity among birds and primates in their capacity to judge certain types of stimulus relations.     

Associative symmetry and stimulus-class formation by pigeons: The role of non-reinforced baseline relations

Two experiments tested the assumption of Urcuioli's (2008) theory of pigeons’ equivalence-class formation that consistent non-reinforcement of certain stimulus combinations in successive matching juxtaposed with consistent reinforcement of other combinations generates stimulus classes containing the elements of the reinforced combinations. In Experiment 1, pigeons were concurrently trained on symbolic (AB) and two identity (AA and BB) successive tasks in which half of all identity trials ended in non-reinforcement but all AB trials were reinforced, contingent upon either responding or not responding to the comparisons. Subsequent symmetry (BA) probe trials showed evidence of symmetry in one of four pigeons. In Experiment 2, pigeons learned three pair-comparison tasks in which left versus right spatial choices were reinforced after the various sample-comparison combinations comprising AB, AA, and BB conditional discriminations. Non-differentially reinforced BA probe trials following acquisition showed some indication of symmetrical choice responding. The overall results contradict the theoretical predictions derived from Urcuioli (2008) and those from Experiment 2 challenge other stimulus-class analyses as well.     

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.     

Behavioral contrast of a complex operant

Two experiments were conducted in which pigeons were trained to perform a complex operant consisting of a peck to one key followed by a peck to another key. In the first experment this performance was reinforced on a variable-interval schedule and the birds were then subjected to a multiple schedule in which the variable-interval was alternated with extinction. The first two pigeons trained showed some deterioration of the cohesiveness of the response sequence. After measures were taken to correct this the pigeon gave evidence of behavioral contrast in a condition where sessions with discrimination were alternated rapidly with sessions with no discrimination (variable-interval only). Another two birds were trained on variable-interval followed by discrimination. One of these birds showed evidence of contrast but the results of the other were not clear.Experiment II was a three phase experiment in which animals were first trained to make the complex response. In the second phase they were extinguished by providing noncontingent reinforcement in each of two components. In the third phase reinforcement was removed from one component, providing for a stimulus-reinforcer contingency. In the third phase the complex response reappeared temporarily.The results of these two studies indicate that a complex response may show behavioral contrast and that it may be enhanced by stimulus-reinforcer contingencies if it has already been trained.     

Individual differences in motor skills ability affect the self-regulation of heart rate

Twenty males who scored relatively high on the rotor-pursuit motor skills task (High performance group) were given seven 2-minute trials to increase heart rate and seven 2-minute trials to decrease heart rate, as were 20 males who scored relatively low on the rotor-pursuit task (Low performance group). Visual analogue feedback was not provided during the first and last acceleration and deceleration trials but was presented during all other trials. Both groups of subjects were able to decrease heart rate significantly with and without feedback. Subjects in the High performance group were able to increase heart rate significantly with feedback and could generalize this increase to a no-feedback trial following feedback trials. Subjects in the Low performance group could not increase heart rate with or without feedback. Changes in respiration rate paralleled those noted for heart rate, but changes in chin electromyographic activity generally did not parallel the heart rate results. The heart rate data are discussed in terms of motor skills theories of self-regulation of heart rate.Some of the results reported in this article are based on a doctoral dissertation submitted to the Graduate School of Northern Illinois University by the second author. The heart rate data were presented at the Convention of the American Psychological Association, Anaheim, 1983.     

The effect of competition on choice by pigeons: foraging rate,resource availability and learning

The effect of competition on food choice by pigeons was examined in a within-subject laboratory study. Pigeons foraged on maize (the more preferred item) and wheat (the less preferred item) in two conditions: alone and with a competitor. Over the course of the experiment an effect of competition on individual choice developed: the pigeons came to choose less maize before the first grain of wheat was taken, and to show a higher proportion of choices of wheat. The effect of competition was entirely due to resource availability: no effect of the competitor per se was found once the number of grains of maize and wheat available for any one choice was taken into account. Choice behaviour changed over time because the birds learned to forage faster in competition, which in turn led to faster depletion rates.     

Sub-Optimal Allocation of Time in Sequential Movements

The allocation of limited resources such as time or energy is a core problem that organisms face when planning complex actions. Most previous research concerning planning of movement has focused on the planning of single, isolated movements. Here we investigated the allocation of time in a pointing task where human subjects attempted to touch two targets in a specified order to earn monetary rewards. Subjects were required to complete both movements within a limited time but could freely allocate the available time between the movements. The time constraint presents an allocation problem to the subjects: the more time spent on one movement, the less time is available for the other. In different conditions we assigned different rewards to the two tokens. How the subject allocated time between movements affected their expected gain on each trial. We also varied the angle between the first and second movements and the length of the second movement. Based on our results, we developed and tested a model of speed-accuracy tradeoff for sequential movements. Using this model we could predict the time allocation that would maximize the expected gain of each subject in each experimental condition. We compared human performance with predicted optimal performance. We found that all subjects allocated time sub-optimally, spending more time than they should on the first movement even when the reward of the second target was five times larger than the first. We conclude that the movement planning system fails to maximize expected reward in planning sequences of as few as two movements and discuss possible interpretations drawn from economic theory.     

Geographical variation in blood parasites in feral pigeons: the role of vectors

Prevalence and intensity of blood parasites are known to vary in space within a same species, yet the causes underlying such variation are poorly known. Theoretically, blood parasites variation can be attributed to differences to exposure to parasite vectors and/or to differences in host susceptibility. Here, we show that prevalence of Haemoproteus columbae in feral pigeons Columba livia varied among five near-by populations (range 15%-100%), paralleled by variation in the abundance of its main vector, the louse flies Pseudolynchia canariensis. Geographic variation in intensity of blood parasites did not covary with abundance of vectors. Within populations, older individuals had a higher probability of being parasitized than younger ones, whereas younger birds, when infected, suffered higher intensities. Furthermore, we found no evidence of sex-related differences neither in prevalence nor in intensity of blood parasite infections. To demonstrate that geographical variation in prevalence was actually due to differences in vector exposure, we conducted two experiments based on translocation of unparasitized pigeons from a vector-free area to an area where both the parasite and vector were abundant. With the first experiment, we demonstrated that unparasitized pigeons were not resistant to the parasite because when transmission was possible pigeons became parasitized in a few months. With the second experiment, in which half of the pigeons were prevented from contacts with the vector, we ruled out the posibility that pigeons we considered as unparasitized would have suffered from latent infections. Therefore, both observational and experimental evidence supports the view that vector abundance is the major factor influencing the spatial variation in prevalence of H. columbae in pigeons.     

Lexical Planning in Sentence Production Is Highly Incremental: Evidence from ERPs

The scope of lexical planning, which means how far ahead speakers plan lexically before they start producing an utterance, is an important issue for research into speech production, but remains highly controversial. The present research investigated this issue using the semantic blocking effect, which refers to the widely observed effects that participants take longer to say aloud the names of items in pictures when the pictures in a block of trials in an experiment depict items that belong to the same semantic category than different categories. As this effect is often interpreted as a reflection of difficulty in lexical selection, the current study took the semantic blocking effect and its associated pattern of event-related brain potentials (ERPs) as a proxy to test whether lexical planning during sentence production extends beyond the first noun when a subject noun-phrase includes two nouns, such as “The chair and the boat are both red” and “The chair above the boat is red”. The results showed a semantic blocking effect both in onset latencies and in ERPs during the utterance of the first noun of these complex noun-phrases but not for the second noun. The indication, therefore, is that the lexical planning scope does not encompass this second noun-phrase. Indeed, the present findings are in line with accounts that propose radically incremental lexical planning, in which speakers plan ahead only one word at a time. This study also provides a highly novel example of using ERPs to examine the production of long utterances, and it is hoped the present demonstration of the effectiveness of this approach inspires further application of ERP techniques in this area of research.     

Reward magnitude and timing in pigeons

We investigated the interaction of motivation and timing by manipulating the expected reward magnitude during a peak procedure. Four pigeons were tested with three different reward magnitudes, operationalized as duration of food access. Each stimulus predicted a different reward magnitude on a 5 s fixed-interval schedule. Trials with different reward magnitudes were randomly intermingled in a session. Most pigeons responded less often and started responding later on peak trials when a smaller reward was expected, but showed no differences in response termination or peak times. Reward magnitude was independently corroborated through unreinforced choice trials, when pigeons chose between the three stimuli presented simultaneously. These results contribute to a growing body of evidence that the expected reward magnitude influences the decision to start anticipatory responding in tasks where the reward becomes available after a fixed interval, but does not alter peak times, nor the decision to stop responding on peak trials.     

Latent contextual inhibition of the cardiac component in the startle reaction to a sound stimulus in rats

Heart rate (HR), freezing score, and motor component were estimated during acoustic startle (ASR) habituation (two sessions with 24-hour interval, 10 trials in a session). It was shown that rats previously exposed to the experimental context (once for 5 min 24 h before training) demonstrated HR decrease in response to the first stimulus and tachycardia in response to the 4th-10th stimuli during the first session. The interstimulus HR declined from the 4th to the 6th trials. The same profile of cardiovascular response was observed in this group at the beginning of the second session with the following (to the 7th trial) habituation of tachycardia. These rats didn't demonstrate the intertrial HR decrease during the second session. Nonadapted rats responded by bradycardia to the 1st and 2nd trials of the first session. The response didn't change for tachycardia with continuation of stimuli presentation. Tachycardia appeared only in response to the 7th-10th trials of the second session and didn't habituate. The intertrial HR level decreased in this group only during the second session. The results are discussed in terms of contextual latent inhibition of the cardiac acoustic startle response.     

Shifts in the psychophysical function in rats

The primary goal was to compare results from a free-operant procedure with pigeons [Machado, A., Guilhardi, P., 2000. Shifts in the psychometric function and their implications for models of timing. J. Exp. Anal. Behav. 74, 25-54, Experiment 2] with new results obtained with rats. The secondary goal was to compare the results of both experiments with dependent variables that were not used in the original publication. As in the original study with pigeons, rats were trained on a two-alternative free-operant psychophysical procedure in which left lever press responses were reinforced during the first and second quarters of a 60-s trial, and right lever press responses were reinforced during the third and fourth quarters of the trial. The quarters were reinforced according to four independent variable interval (VI) schedules of reinforcement. The VI duration was manipulated in each quarter, and shifts in the psychophysical functions that relate response rate with time since trial onset were measured. The results obtained with rats were consistent with those previously obtained with pigeons. In addition, results not originally reported were also consistent between rats and pigeons, and provided insights into the perception, memory, and decision processes in Scalar Expectancy Theory and Learning-to-Time Theory.     

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.     

Within-trial contrast: The effect of probability of reinforcement in training

There is evidence that pigeons prefer conditioned reinforcers that are preceded by greater effort over those that are preceded by less effort (an effect that has been attributed to within-trial contrast). In past research the probability of reinforcement for correct choice of the conditioned reinforcer has been 100%, however, the high level of reinforcement for both alternatives in training may result in a performance ceiling when choice between those alternatives is provided on test trials. In the present study we tested this hypothesis by including a group for which the probability of reinforcement in training was only 50%. Pigeons were trained on two simultaneous discriminations, one that was preceded by a 30 peck requirement the other by a single peck requirement. On test trials, we found a significant preference for the S+ that required the greater effort in training for pigeons trained with 100% and a small but nonsignificant effect for pigeons trained with 50% reinforcement. Although the hypothesis that the within-trial contrast effect was constrained by a performance ceiling was not confirmed, we did find a reliable within-trial contrast effect with 100% reinforcement.     

Effect of a warm‐up sample on stabilizing the performance of untrained panelists in time–intensity evaluation

To improve the reliability of sensory evaluation, it is necessary to minimize response bias. In this study, we investigated the effect of a warm‐up sample on time–intensity evaluation of after‐flavor. Untrained panelists evaluated continuously perceived intensity of bitterness and retronasal aroma, using four types of samples (two brands of coffee beverages × two types of lids). Half of participants evaluated bitterness intensity in the first session and retronasal aroma intensity in the second session, whereas the other half evaluated them in the opposite order. Each session consisted of four trials, and we regarded the first trial sample of each session as a warm‐up. Results revealed that perceived intensity was significantly lower in the first trial than in the following trials, and that the time courses of the second, third, and fourth trials were similar except when participants evaluated bitterness intensity in the second session. In other words, the use of a warm‐up sample stabilized the performance of time–intensity evaluation of bitterness and retronasal aroma in the first session. Furthermore, no warm‐up sample was needed in the second session when participants evaluated bitterness intensity, but a warm‐up was needed when they evaluated retronasal aroma intensity.

Practical application

Based on the results of this study, we concluded that the use of a warm‐up sample stabilized performance in time–intensity evaluation of after‐flavor by untrained panelists. We proposed that in order to obtain reliable performance in time–intensity evaluation of after‐flavor, untrained panelists should be provided a training trial using warm‐up sample before staring the test trials.     

Duration ratio discrimination in pigeons: a criterion-setting analysis

Pigeons received trials beginning with a sequence of two colors (blue-->yellow) on the center key of a three-key array. The colors lasted different lengths of time. At the end of the sequence pigeons chose between two keys based on a criterial ratio of the temporal sequence. One choice was reinforced if the time ratio was less than the criterion and the alternate choice was reinforced if the time ratio was greater than the criterion. The criterial ratios (first to second duration) were 1:1, 1.5:1, and 3:1. The same set of intervals was used for the different criterion ratios, producing a balanced distribution of time ratios for the 1.5:1 condition, and unbalanced distributions for the 1:1 and 3:1 conditions. That is, for the 1.5:1 condition half of the duration pairs were less than the criterion and half were greater. However, for the 1:1 and 3:1 conditions, more duration pairs were less than (3:1) or greater than (1:1) the criterion. Accuracy was similar across criterion ratios, but response bias was influenced by the asymmetries of time ratios in the 1:1 and 3:1 conditions. When these asymmetries were controlled, the response biases were reduced or eliminated. These results indicate that pigeons are flexible in establishing a criterion for discriminating duration ratios, unlike humans, who are less flexible and are bound to categorical distinctions in the discrimination of duration ratios.     

The effect of food deprivation on self-control

Two experiments examined the effect of food deprivation on choice in a discrete-trials self-control paradigm, choice between a larger, more-delayed reinforcer and a smaller, less-delayed reinforcer. In Experiment 1, four pigeons were each deprived to 65%, 80%, and 90% of their free-feeding weights, and the delay to the smaller reinforcer was varied. Deprivation level did not affect choice, but the rate of ineffective key pecks made during the reinforcer delays increased as deprivation increased. In Experiment 2, four pigeons were exposed to conditions in which they were fed up to their 80% free-feeding weights following experimental sessions, and in which they were given no postsession feedings. Both the pigeons' weights and their latencies to insert their heads into the food hopper when food was available were lower when the pigeons were not fed following experimental sessions. Choice showed no change. Deprivation level affects response rate and eating behavior in these procedures, but not choice.     

Late ERP components in visual and auditory Go/Nogo tasks

In an audio-visual Go/Nogo paradigm we studied whether the Go/Nogo difference, usually found in the time range of the visual N2, is also present after auditory stimuli, which bears on the common response inhibition hypothesis of this N2 effect. Moreover the possible presence and variation of P300 subcomponents were studied with the goal of clarifying the reasons for the commonly observed P300 topography changes between Go and Nogo trials. To disentangle possible P300 subcomponents we applied a crossmodal divided attention (DA) condition, in which the subcomponents are known to be separated after auditory stimuli in choice tasks.An N2 effect was found after visual but not after auditory stimuli, which is evidence against the response-inhibition hypothesis. After visual stimuli a positive complex (P400) was seen, whereas after auditory stimuli two dissociated components (P400 and P507) were found instead. The P507 had a parietal maximum for both Go and Nogo trials. It was larger and it peaked later in Go than in Nogo trials. The P400 showed topographic differences between Go and Nogo trials, which could be explained by the overlap of the two subcomponents. We assume that (i) both subcomponents have a stable topography across response type, and (ii) the first subcomponent is invariant with response type, whereas the second (which overlaps the first one) is larger and peaks later on Go than on Nogo trials.