Symbolic,identity, and probe delayed matching of sounds by the bottlenosed dolphin

The short-term memory for sounds of the bottlenosed dolphin was tested using symbolic, identity, and probe forms of the delayed matching-to-sample (DMS) task. The forms differed in the number (one or two) or nature (symbolic or identity matches of sample sounds) of postdelay test stimuli available as memory retrieval cues. Although symbolic DMS was difficult to learn, the final performance level was approximately equal to that for identity or probe DMS. On all tasks, the dolphin’s responses were above 80% correct through to delays of 90 sec and, in some cases, through to delays of 180 and 240 sec, the “limits” being governed mainly by the dolphin’s reluctance to continue being tested at long delays. Encoding of sample stimuli into their learned symbolic representation was hypothesized to have reduced symbolic DMS to a recognition memory task, resulting in the observed equivalence of performance with the other two recognition memory tasks. The probe DMS results, unlike those for identity or symbolic DMS, showed no significant proactive interference effects from samples of prior trials. Instead, proactive interference was traceable to the probe value of the prior trial. Overall, the auditory DMS data for the dolphin were functionally similar to results reported for monkeys tested on symbolic, identity, and probe visual DMS tasks.     

Short-term memory for visual and auditory stimuli in pigeons

Separate groups of pigeons were trained to perform symbolic delayed matching to sample with auditory and visual sample stimuli. For animals in the auditory group, ambient tones that varied in frequency served as sample stimuli; for animals in the visual group, ambient red and green lights served as sample stimuli. In both cases, the sample stimuli were mapped onto the yellow and blue comparison stimuli presented on left and right pecking keys. In Experiments 1 and 2, it was found that visual and auditory delayed matching were affected in the same ways by several temporal variables: delay, length of exposure to the sample stimulus, and intertrial interval. In Experiments 3, 4A, and 4B, a houselight presented during the delay interval strongly interfered with retention in both visual and auditory groups, but white noise presented during the delay had little effect in either group. These results seem to be more in line with a prospective memory model, in which visual and auditory sample stimuli are coded into the same instructional memories, than with a model based on concepts of retrospective memory and modality specificity.     

The role of test stimuli in matching to compound samples by pigeons

Two pigeons matched to sample in a three-key operant conditioning chamber. In Experiment I, two different kinds of samples were presented on the center key.Element samples were members of one of two sample sets — colors (a red or blue disk) or lines (a vertical or horizontal orientation of a set of white lines). These samples were followed by their respective sample sets on the side keys as comparison stimuli.Compound samples consisted of a set of lines superimposed on a colored disk. Following these samples, either sample set could appear as comparison stimuli. Matching to compound samples was less accurate than matching to element samples. One interpretation is that sharing of attention among elements of a compound sample weakened stimulus control by each element. A different interpretation is that an element sample controlled matching better because it was physically identical to a comparison stimulus whereas a compound sample was not. Experiments II–IV evaluated this “generalization decrement” alternative by testing element- vs. compound sample control with both element and compound comparison stimuli. Irrelevant elements were added to form compound comparison stimuli, some of which were physically identical to a preceding compound sample, but never identical to an element sample. In all experiments, the addition of irrelevant elements of comparison stimuli reduced sample control. However, the generalization decrement hypothesis failed to predict how differences in performance maintained by element and compound samples were affected by different tests of sample control. Matching accuracy appeared to be independently determined by the number of elements in a sample and whether irrelevant elements were present during tests of sample control.     

Interference and auditory short-term memory in the bottlenosed dolphin

Interference in auditory short-term memory in the bottlenosed dolphin,Tursiops truncatus (Montagu), was studied using a delayed matching-to-sample task. At each trial, one of two sample sounds, chosen randomly, was projected underwater for 4 sec and then, after a variable delay interval, both sounds were presented. A response to the sound matching the initial sample was reinforced. Correct matching was significantly reduced following short intervals between trials in combination with long delays after the sample (proactive interference), or when a near continuous irrelevant sound was inserted into the delay interval (retroactive interference). There was rapid habituation to interference if the irrelevant sound was short in duration relative to the delay interval. For both proactive and retroactive interference, the errors were predominantly responses to the sample sound appropriate to the prior trial rather than to the current trial, indicating that memory for the relative recency of events (temporal memory) was degraded by interference. When interference was deleted or minimized, temporal memory remained nearly perfect over 30-sec delay intervals, the longest tested. The importance of distinguishing between temporal memory and nontemporal, or event, memory in different forms of the delayed matching task was emphasized.     

Auditory matching-to-sample in monkeys (Cebus apella)

Four of 8 monkeys were successfully trained on an identity matching-to-sample task employing two acoustic stimuli. In five subsequent tests with different pairs of auditory stimuli, their performances were at levels that provided strong evidence for concept-mediated transfer. Thus, despite past failures to demonstrate the matching concept in the auditory modality, the present results indicate that the matching concept is not limited in monkeys to the visual modality. On the other hand, the failure of 4 subjects to learn the initial matching task constitutes additional evidence of cognitive asymmetry in monkeys with regard to the visual and auditory modalities.     

Limited capacity information processing and pigeon matching-to-sample: Testing alternative hypotheses

The limited capacity hypothesis explains the element superiority effect observed in pigeons’ element-compound matching-to-sample performance as the result of a central information processing overload occurring at the time of sample exposure. Major alternative hypotheses offered in the literature to date argue that element superiority is due to a difference in element- and compound-sample memory codes or to a peripheral sensory limitation during sample exposure. These alternative factors weresimultaneously prevented from influencing matching performance in the present experiment, but the element superiority effect remained. A central information processing account of the element superiority effect is supported by the strong tests of alternative hypotheses provided herein. The discussion addresses remaining challenges to the hypothesis that information overload for compound samples occurs at the time of sample exposure.     

Restricted processing of simultaneously presented brightness and pattern stimuli in pigeons

Performance during simultaneous matching-to-sample was assessed in pigeons presented with element and compound visual samples. In Experiment 1, pigeons were trained with a symbolic matching procedure, in which different pairs of colored comparison cues presented on side keys were mapped onto a bright or dim houselight as one pair of sample stimuli and onto vertical and horizontal lines on the center key as a second pair of sample stimuli. They were then tested with houselight-line compound samples. It was found that matching accuracy for lines was significantly diminished with compound samples relative to element samples. Conversely, house-light intensities were matched as well with compound samples as with element samples. In Experiment 2, a similar effect was found with pigeons that had been trained to match only line samples. In Experiment 3, it was discovered that sample duration had no influence on the matching deficit found with lines following compound samples in birds either trained or not trained to match houselight intensities. These results, taken in combination with recent findings from experiments with auditory-visual compounds, suggest a restricted processing account of pigeon processing of simultaneously presented stimuli from different sources.     

Generalization of visual matching and delayed matching by a California sea lion (Zalophus californianus)

Only a limited number of species have been found capable of generalized matching-to-sample (MTS) after exposure to relatively few training exemplars. We trained a juvenile, experimentally naive California sea lion (Zalophus californianus) in MTS, using a pair of three-dimensional objects as samples. Successful matching to a criterion of 90% correct or better over 2 successive sessions was attained in 12 sessions (269 trials and 70 errors). Two subsequent “partial” transfer tests, in which each of the two training objects was paired with a novel test object, and four additional transfer tests, all with novel objects, were presented following training. An 80% performance criterion over 2 successive sessions was reached, or closely approximated, in from 2 to 4 transfer sessions for all transfer tests; errors to criterion tended to be reduced across the successive novel transfer tests and were as few as five during the final two tests; and performance on the first 48 trials of the last two novel transfers was not significantly different from a near-ceiling level baseline performance measure. Neophobic responses of the sea lion to new objects precluded an unbiased evaluation of immediate (Trial 1) transfer. The sea lion’s short-term memory for sample objects was also measured. Matching performance was maintained at a level of 78% correct responses or better for delays through to 45 sec after removal of the sample object. At a 58-sec delay, the longest tested, performance declined to 69% correct responses. These retention levels are only somewhat below levels reported for dolphins and nonhuman primates tested on visual delayed MTS, but they are above levels typically reported for pigeon subjects.     

Object recognition through eavesdropping: Passive echolocation in bottlenose dolphins

A bottlenose dolphin (Tursiops truncatus) demonstrated the ability to select the matching object in a matching-to-sample task after listening to another dolphin inspect the sample object via echolocation. The listener was prevented from inspecting the sample himself. In Experiment 1, with objects familiar to both dolphins, the listener’s performance was significantly better than chance. In Experiment 2, objects familiar to only one of the dolphins were used. On these trials, the listener’s performance was significantly better than chance only when the inspecting dolphin made a correct choice. Analysis of the listener’s responses when the inspector made an error demonstrated that this contingency was not due to the listener’s matching the inspector’s response, but was apparently due instead to inadequate information in the echo. The results suggest that the listener was able to “eavesdrop” on echoes produced by the inspector’s clicks and derive characteristics of the sample object.     

Simultaneous processing of visual and spatial stimuli in pigeons

Pigeons were trained to symbolically match comparison stimuli to either visual sample stimuli presented on a center key or to spatial sample stimuli presented on side keys. Tests were carried out in which visual and spatial cues were simultaneously presented in compound and short-term memory was probed for either visual or spatial information. Symmetrical interference with the matching of visual and spatial components of compounds was found when the visual and spatial cues were presented on separate keys. However, when visual and spatial cues were superimposed on the same side key, no interference was observed relative to element control tests. Discussion of these findings focuses on accounts in terms of limited processing capacity, coding decrement, and receptor orientation mechanisms.     

Spatial and configural factors in compound stimulus processing by pigeons

Three matching-to-sample experiments examined whether spatial or configural factors determined how the element arrangement of compound sample stimuli influenced matching accuracy in pigeons. Seven types of compound stimuli were tested. The arrangement of color and line-orientation elements in these compounds varied in terms of the spatial separation between the elements, the degree of consistency in element spatial location, and the number of bounded areas containing the elements. Matching accuracy was examined upon initial exposure to the compounds, during asymptotic conditions of shared attention, and with variation of sample durations ranging from .04 to 5.935 sec. In all three experiments, when spatial proximity, locational certainty, and the number of lines were precisely controlled or equated, no evidence for the proposed configural processing of “unified” compounds was found (Lamb & Riley, 1981). Element spatial separation, and to a lesser degree perceptual limitations, determined compound performance. These results question our lab’s previous evidence for configural compound processing by pigeons (Lamb, 1988; Lamb & Riley, 1981). They suggest instead that pigeons independently and separately process the individual elements of color/line-orientation compounds, with element separation determining the distribution of processing between the elements.     

Stimulus modality and short-term memory in rats

Two experimental paradigms are presented aimed at determining the retention of auditory and visual information over brief delay intervals. First, a conditional delayed matching-to-sample procedure was used in which rats were required to symbolically match the modality of the sample stimulus with one of two comparison stimuli. In the second experiment, subjects were trained and tested using a Konorski-type procedure. Despite the conceptual and procedural differences between the two procedures, subjects in both experiments showed steeper forgetting functions for visual events than for auditory events, while performance levels at 0-sec delay intervals were equivalent for both stimuli. These results, when taken together with related research conducted with pigeons, suggest that content of memory may have important influences on the short-term retention abilities of animal subjects.     

Effects of signaled retention intervals on pigeon short-term memory

In three delayed matching-to-sample experiments, pigeons were given distinctive stimuli that were either correlated or uncorrelated with the scheduled retention intervals. Experiment 1 employed a single-key, go/no-go matching procedure with colors as the sample and test stimuli; lines of differing orientations signaled short or long delays for one group, whereas the lines and the delays were uncorrelated for the other group. The function relating discriminative test performance to delay length was steeper in the correlated group than in the uncorrelated group. In addition, the line orientation stimuli controlled differential rates of sample responding in the correlated group, but not in the uncorrelated group. In Experiment 2, subjects extensively trained with correlated line orientations were exposed to reversed cues on probe trials. Miscuing decreased discriminative test responding at the short delay, but enhanced it at the long delay. As in the correlated group of the first experiment, rates of sample keypecking were higher in the presence of the “short” time tag than in the presence of the ”long” time tag. Experiment 3 used a three-key choice-matching procedure and a within-subjects design, and equated reinforcement rate at the short and long delays. When auditory stimuli were correlated with delay length, the function relating choice accuracy to delay was steeper than when the stimuli and the delays were uncorrelated. The consistent effects of signaled retention intervals on memory performance may be understood in terms of differential attention to the sample stimuli.     

Spontaneous recovery from overexpectation

In three Pavlovian magazine approach experiments, rats received conditioning of auditory and visual stimuli by pairing with a pellet. Then the stimuli received additional conditioning while presented in simultaneous compound and were tested either immediately or after a delay. The compound conditioning resulted in a decrement in responding to the individual stimuli (overexpectation). However, there was recovery of responding with the passage of time. These results suggest that the decrements produced by an overexpectation procedure share some properties with those produced by nonreinforcement.     

Auditory delayed discriminations by the dolphin: Nonequivalence with delayed-matching performance

Working memory in a bottlenosed dolphin was tested in both indirect and direct auditory delayed-discrimination tasks in which a correct spatial response was conditional upon the nature of a preceding sound. In the indirect task, either one of two possible sounds was briefly presented. After a prescribed delay, the dolphin was cued to go either to a left-hand or right-hand paddle pair. Responses to the outer paddle of a pair were rewarded following sound A, and responses to the inner paddle of a pair were rewarded after sound B. In the direct delayed-discrimination task, only one paddle pair was used in each session. In both tasks, the delay interval between the discriminative sound stimulus and the opportunity for a spatial response was progressively increased over sessions until the animal failed to meet a specified performance criterion or self-terminated a session. Delay limits of about 30 and 60 sec were obtained in the indirect and direct tasks, respectively. The increase in delay limit in the latter task was attributable to the use of an overt mediational response during the longer delays. In both cases, however, the obtained delay limits fell considerably short of the 2- to 3-min limits obtained in auditory delayed-matching studies using the same test sounds and the same subject. The task differences indicate that working memory functions cannot depend upon memory of the predelay stimulus alone, but must be determined in part by additional processes.     

Retardation and facilitation of matching acquisition by differential outcomes

Pigeons were trained on many-to-one matching-to-sample with food and no-food outcomes that were either differential or nondifferential with respect to the sample stimuli. In the differential condition, outcomes were correlated with the correct comparison-alternatives-for half of the subjects, and were uncorrelated with those alternatives for the remaining subjects. Relative to non-differential training, matching acquisition was facilitated in the correlated condition but retarded in the uncorrelated condition. These results clearly demonstrate that differential outcomes do not affect conditional discrimination learning merely by enhancing the discriminability or distinctiveness of the samples with which they are associated. Rather, they apparently give rise to another discriminative cue (viz., an outcome expectancy), which can either enhance or interfere with performance, depending on its predictive validity.     

Sample duration and type of stimuli in delayed matching-to-sample in rhesus monkeys

Two experiments were performed to determine the effect of sample duration (0.1, 2, and 4 sec), delay interval (.03, 4, 8, 16, and 32 sec), and type of stimulus (color and shape) on the matching performance of rhesus monkeys. In Experiment 1, the 15 possible delay-duration combinations were randomly presented in blocks of 15 trials. In Experiment 2, each duration was held constant and the five delays randomly presented. Then each delay interval was held constant with the three durations randomly varied. Matching performance increased as sample duration increased (ps < .01 and .005), while length of delay did not significantly affect performance. The type of stimuli paired in the matching test significantly affected performance (ps < .05 and .10) with the shape/shape choices leading to the poorest performance. Stimulus discriminability and amount of training with brief sample durations were implicated as significant determinants of matching performance.     

Do pigeons prefer information in the absence of differential reinforcement?

Prior research has indicated that pigeons do not prefer an alternative that provides a sample (for matching to sample) over an alternative that does not provide a sample (i.e., there is no indication of which comparison stimulus is correct). However, Zentall and Stagner (Journal of Experimental Psychology. Animal Behavior Processes 36:506?C509, 2010) showed that when delay of reinforcement was controlled, pigeons had a strong preference for matching over pseudomatching (i.e., there was a sample, but it did not indicate which comparison stimulus was correct). Experiment 1 of the present study replicated and extended the results of the Zentall and Stagner (Journal of Experimental Psychology. Animal Behavior Processes 36:506?C509, 2010) study by including an identity relation between the sample and one of the comparison stimuli in both the matching and pseudomatching tasks. In Experiment 2, in which we asked whether the pigeons would still prefer matching if we equated the two tasks for probability of reinforcement, we found no systematic preference for matching over pseudomatching. Thus, it appears that in the absence of differential reinforcement, the information provided by a sample that signals which of the two comparison stimuli is correct is insufficient to produce a preference for that alternative.     

Some determinants of response summation

Response summation in pigeons was examined in four experiments. In Experiment 1, summation was not found with a compound of two visual stimuli on a television screen after they had individually been used for instrumental conditioning. In this experiment, the training and test trials were separated by an interval during which the television screen was dark. Summation was found in Experiment 2 for which the television screen was permanently white during the interval between trials and in the region that was not occupied by the experimental stimuli. These results were replicated using a within-subject design (Experiment 3) and autoshaping (Experiment 4). Experiment 2 also revealed summation with compounds of auditory and visual stimuli, but not with compounds of two auditory stimuli or two diffuse lights. The results can be explained by a variety of theories of learning, if they take account of generalization between the stimuli.     

Differential effects of omitting comparison stimuli on symbolic and identity matching to sample: Evidence for altered instructional processes in pigeon short-term memory

Two experiments were performed to determine the effects of omitting the comparison stimuli in a matching-to-sample task. In Experiment 1, birds were trained initially on both symbolic and identity matching to sample. Comparison stimuli were then omitted following the presentation of a particular sample stimulus, and this decreased the number of sample (observing) responses. The reintroduction of the comparison stimuli on subsequent probe trials revealed that the accuracy of symbolic matching was reduced to chance levels, while identity matching accuracy was significantly below chance. In Experiment 2, a similar procedure was employed; however, observing responses to the comparison-omitted samples were maintained by direct reinforcement (fixed ratio 20). Matching accuracy during probe trials was again at chance levels for symbolic matching but, contrary to Experiment 1, was significantly above chance for identity matching. The differential effects of omitting comparison stimuli on symbolic and identity matching trials in these two experiments were interpreted within a framework which assumes that instructional processes are altered by comparison-omission procedures.