Serial list linking by macaque monkeys (Macaca mulatta): list property limitations

Four sophisticated macaque monkeys (Macaca mulatta) learned 6 different, 15-item ordinal lists (via conditional, 2-choice discriminations) as part of a study assessing some properties of serial list memory in monkeys. After assuring that the first 3 lists were well retained, the researchers attempted to link these by training only the 2 end-item pairs that ordered all 45 items into an inclusive series. A 20-day test of possible pairings among these 45 items was then conducted. Subsequently, the other 3 lists were trained and tested for retention, but no link training was provided. Then, a test like the one that had followed linking was administered. Unlike previous outcomes with three 5-item lists, linking did not yield immediate merger of three 15-item lists into a 45-item list, although 45-item lists were acquired after sufficient exposure to the testing/training regimen under both linking and nonlinking conditions. List length as a limiting factor in linking suggested processing restrictions analogous to those observed in human list memory. Results supported further investigation of list-linking characteristics.     

An experimental analysis of memory processing

Rhesus monkeys were trained and tested in visual and auditory list-memory tasks with sequences of four travel pictures or four natural/environmental sounds followed by single test items. Acquisitions of the visual list-memory task are presented. Visual recency (last item) memory diminished with retention delay, and primacy (first item) memory strengthened. Capuchin monkeys, pigeons, and humans showed similar visual-memory changes. Rhesus learned an auditory memory task and showed octave generalization for some lists of notes--tonal, but not atonal, musical passages. In contrast with visual list memory, auditory primacy memory diminished with delay and auditory recency memory strengthened. Manipulations of interitem intervals, list length, and item presentation frequency revealed proactive and retroactive inhibition among items of individual auditory lists. Repeating visual items from prior lists produced interference (on nonmatching tests) revealing how far back memory extended. The possibility of using the interference function to separate familiarity vs. recollective memory processing is discussed.     

Premise-pair training for valid tests of serial list organization in macaques

This study evaluated the role of several different training procedures on (1) efficiency of acquisition and (2) organizational characteristics of memory for lists that could be serially ordered. Five macaque monkeys were trained via two-choice object discriminations in a formboard apparatus on several five-item-series tasks that provided different levels of intrasession conditionality. Although ease of acquisition differed for subsets of the constituent pairs, concurrent inclusion of the four premise pairs that defined a list required equivalent amounts of training on every task. All training procedures yielded similar retention-test performances and showed common organizational properties (on both error and latency measures) consistent with the view that lists were retained as internally represented ordered series. Test outcomes emphasized the need for integrated exposition of all concurrent conditional relationships to allow appropriate tests of serial organization. However, if given such training, the monkeys revealed integrated serial memory even though they had never seen many of the possible novel combinations of list items. In overview, their performances offered further definition of the procedures required for valid assessment of inferential properties in comparative cognition. Electronic Publication     

Representation of serial order in monkeys (Cebus apella)

Cebus monkeys were trained on a five-item serial learning task, symbolized as ABCDE; the initial stages of training were on the shorter subseries AB, ABC, and ABCD. To assess the monkeys' knowledge of the sequential position of each item, pair-wise tests were given to 2 subjects after acquisition of the ABCD series and to 4 subjects after reaching criterion on the ABCDE series. In both tests, the monkeys performed at high levels on the interior pairs, which were BC for the ABCD series, and BC, BD, and CD for the ABCDE series. These results, as well as the orderly relations observed in the pair-wise tests between first-item response latency and first-item position and between second-item response latency and number of missing items, indicated that the monkeys had developed a well-organized internal representation of the four- and five-item series. Although pigeons are also capable of learning four-item and five-item series, they apparently do not develop a comparable representational structure. The disparity between the monkeys' and pigeons' representational competence for serial order is predictable from the difference in their capacities for associative transitivity.     

Sequential planning in rhesus monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

In the current study, we examined the planning abilities of rhesus monkeys (Macaca mulatta) by training them on a five-item list composed of coloured photographs and then testing them on switch and mask trials. In contrast to previous studies where monkeys made responses using a joystick, in the current study, monkeys made responses directly to a touch screen. On switch trials, after a response to the first list item, the on-screen positions of two list items were exchanged. Performance on trials in which the second and third list items were exchanged was poorer compared to normal (non-switch) trials for all subjects. When the third and fourth items were exchanged, however, only one subject continued to show performance deficits. On mask trials, following a response to the first item, the remaining items were covered by opaque white squares. When two items were masked, all four subjects responded to each masked item at a level significantly above chance. When three items were masked, however, only one subjected was able to respond to all three masked items at a level significantly above chance. The results of the present study indicate that three of our four monkeys planned one response ahead while a single monkey planned two responses ahead. The significance of these findings is discussed in relation to previous studies on planning in chimpanzees and monkeys.     

Serial learning by rhesus monkeys: I. Acquisition and retention of multiple four-item lists.

Two rhesus monkeys were trained to learn eight 4-item lists, each composed of 4 different photographs. Lists were trained in successive phases: A, A----B, A----B----C, and A----B----C----D. After List 4, retention, as measured by the method of savings, was, on average, 66% (range: 44-84%). Indeed, all 4 lists could be recalled reliably during a single session with neither a decrement in accuracy nor an increase in the latency of responding to each item. Response latencies on a subset test employing all possible 2- and 3-item subsets of each 4-item list support the hypothesis that monkeys form linear representations of a list. Latencies to Item 1 of a subset varied directly with the position of that item in the original list. On List 1, latencies to Item 2 varied directly with the number of intervening items between Item 1 and Item 2 in the original list. During the acquisition of Lists 5-8, both Ss mastered the A----B and A----B----C phases of training in the minimum number of trials possible.     

The generation effect in monkeys

How well one retains new information depends on how actively it is processed during learning. Active attempts to retrieve information from memory result in more learning than passive observation of the same information (the generation effect). Here, we present evidence for the generation effect in monkeys. Subjects were trained to respond to five-item lists of photographs in a particular order. On some lists, they could request "hints" to guide their behavior; on others, they had to generate the correct order from memory. Training with hints resulted in high levels of initial performance, but accuracy dropped precipitously when the hints were removed on the criterion test. Training without hints led to relatively poor initial performance, but accuracy increased steadily and remained high on the criterion test.     

Monkey memory: same/different concept learning, serial probe acquisition, and probe delay effects

Three rhesus monkeys were trained and tested in a same/different task with six successive sets of 70 item pairs to an 88% accuracy on each set. Their poor initial transfer performance (55% correct) with novel stimuli improved dramatically to 85% correct following daily item changes in the training stimuli. They acquired a serial-probe-recognition (SPR) task with variable (1-6) item list lengths. This SPR acquisition, although gradual, was more rapid for the monkeys than for pigeons similarly trained. Testing with a fixed list length of four items at different delays between the last list item and the probe test item revealed changes in the serial-position function: a recency effect (last items remembered well) for 0-s delay, recency and primacy effects (first and last list items remembered well) for 1-, 2-, and 10-s delays, and only a primacy effect for the longest 30-s delay. These results are compared with similar ones from pigeons and are discussed in relation to theories of memory processing.     

Discrimination of functionally appropriate and inappropriate throwing tools by captive tufted capuchins (<Emphasis Type="Italic">Cebus apella</Emphasis>)

A tool-throwing task was used to test whether capuchin monkeys understand the difference between functionally appropriate and functionally inappropriate tools. A group of monkeys was trained to obtain a sticky treat from a container outside their enclosure using a projectile attached to one end of an anchored line. Subsequently, these monkeys were given choice tests between functional and nonfunctional versions of tools used in training. A different feature of the tool was varied between alternatives in each choice test. The monkeys chose to use functional tools significantly more often than nonfunctional tools in early exposures to each choice test. A second experiment tested whether these subjects, as well as a second group of minimally trained participants, could distinguish between functional and nonfunctional tools that appeared different from those used in training. A new set of design features was varied between tools in these choice tests. All participants continued to choose functional tools significantly more often than nonfunctional tools, regardless of their tool-throwing experience or the novel appearance of the tools. These results suggest that capuchin monkeys, like chimpanzees studied in similar experiments, are sensitive to a variety of functionally relevant tool features.     

Mechanisms of inferential order judgments in humans (Homo sapiens) and rhesus monkeys (Macaca mulatta)

If A > B, and B > C, it follows logically that A > C. The process of reaching that conclusion is called transitive inference (TI). Several mechanisms have been offered to explain transitive performance. Scanning models claim that the list is scanned from the ends of the list inward until a match is found. Positional discrimination models claim that positional uncertainty accounts for accuracy and reaction time patterns. In Experiment 1, we trained rhesus monkeys (Macaca mulatta) and humans (Homo sapiens) on adjacent pairs (e.g., AB, BC, CD, DE, EF) and tested them with previously untrained nonadjacent pairs (e.g., BD). In Experiment 2, we trained a second list and tested with nonadjacent pairs selected between lists (e.g., B from List 1, D from List 2). We then introduced associative competition between adjacent items in Experiment 3 by training 2 items per position (e.g., B?C?, B?C?) before testing with untrained nonadjacent items. In all 3 experiments, humans and monkeys showed distance effects in which accuracy increased, and reaction time decreased, as the distance between items in each pair increased (e.g., BD vs. BE). In Experiment 4, we trained adjacent pairs with separate 9- and 5-item lists. We then tested with nonadjacent pairs selected between lists to determine whether list items were chosen according to their absolute position (e.g., D, 5-item list > E, 9-item list), or their relative position (e.g., D, 5-item list < E, 9-item list). Both monkeys' and humans' choices were most consistent with a relative positional organization.     

Disruption of short-term memory for reinforcement by ambient illumination

Pigeons were trained on a delayed conditional discrimination (DCD) in which choice of one of two simultaneously presented stimuli was reinforced if the trial had been initiated by presentation of a food sample. On trials in which no sample was presented, choice of the other colour was reinforced. Illumination of the houselights during the retention interval was provided in an attempt to interfere with retention of information about the food sample which served as a conditional cue. In two experiments, retention interval illumination produced a greater disruption of DCD performance on no sample trials than on food sample trials. The finding that retention interval illumination disrupts DCD performance on no sample trials suggests that this manipulation does not affect memorial processes since there was good evidence that performance on no sample trials did not depend on remembering what happened at the outset of the trial. Furthermore, the magnitude of the disruption was larger if the illumination immediately preceded the choice stimuli than if it followed presentation of the sample stimuli. These results support the hypothesis that retention interval illumination disrupts DCD performance by interfering with discriminative control of the choice response rather than with memorial processes. In neither study did retention interval illumination impair discriminative autoshaping to keylight stimuli that immediately preceded the food sample and no sample DCD trials.     

Linking of serially ordered lists by macaque monkeys (Macaca mulatta): list position influences

Five monkeys (Macaca mulatta) were trained on 2 sets of 3 5-item serially ordered lists. Then, each set was either linked or not in a counterbalanced, within-subject design. Linking entailed training on the 2 pairs that ordered the 3 5-item lists into a single overall 15-item series. Choices on novel pairings after linking conditions attempted to define the unique contributions of knowledge of within-list ordinal position and between-lists link training. With linkage, the series was immediately treated as a 15-item ordered list. Without linkage, choices reflected list positions from initial learning, but continued testing with directional reward yielded gradual ordering into a 15-item list. Apparently, monkeys remembered and used initial list-position information, but linkage allowed inference of an integrated serial relationship among items. Results supported primate list memory as an organizational process.     

Auditory List Memory in Rhesus Monkeys

Auditory memory of 2 rhesus monkeys was tested in a serial probe recognition task. Lists of four environmental or natural sounds were followed by a retention interval and a test. The test matched one of the list items on half of the trials. The retention interval was varied across sessions. Six experiments showed similar results and changes in the serial position function. At short retention intervals, there was good memory for first list items (primacy effect) and poor memory for last list items. At intermediate retention intervals, memory improved for last list items (recency effect). At long retention intervals (20 s and 30 s), the recency effect was strong, and the primacy effect had dissipated. These auditory primacy and recency effects and their changes with retention interval were opposite to those for visual memory. Implications for processes and mechanisms of memory are discussed.     

Infants' memory processing of a serial list: list length effects

Six-month-olds, trained with a three-mobile serial list, exhibit a primacy effect 24 h later. In three experiments, we demonstrated that increasing list length impairs their memory for serial order. In all experiments, 6-month-olds were trained with a five-mobile list. In Experiment 1, infants failed to exhibit a primacy effect on a 24-h delayed recognition test, recognizing mobiles from all serial positions. In Experiment 2, infants did exhibit a primacy effect on a reactivation (priming) test, suggesting that they may originally have encoded serial-order information. Experiment 3 confirmed that serial-order information was represented in infants' training memory. After the reactivation treatment, infants were precued with one list member and tested for recognition of another. When precues specified valid order information, infants recognized test mobiles from the later serial positions. The memory dissociation for serial order on delayed recognition and reactivation tests adds to the growing evidence that young infants possess two functionally distinct memory systems.     

Development of conditional and equivalence relations without differential consequences

Two experiments were conducted to establish conditional stimulus relations without differential consequences and to test for the emergence of other relations. In Experiment 1, 3 adults responded to match-to-sample displays in which sample-comparison pairs were constant while the second comparison presented with each pair changed periodically across trials. No differential consequences followed any comparison selections. All subjects learned conditional relations between constant samples and comparisons, but results of tests for transitivity in those relations were equivocal. In Experiment 2, 4 children were given unreinforced training and testing similar to that provided to the adults in Experiment 1, with procedural refinements. One child learned conditional relations and demonstrated emergent relations that confirmed the development of two four-member equivalence classes. Another child learned the conditional relations but did not demonstrate any emergent relations reliably. A 3rd child, after reinforced training on two conditional relations, learned four new conditional relations without differential consequences and demonstrated symmetry but not equivalence in the trained relations. The 4th child did not learn the conditional relations. These findings emphasize the importance of careful construction of tests for stimulus equivalence and suggest a need for critical analyses of the apparent emergence of untrained stimulus relations on unreinforced tests that has been observed in several stimulus equivalence studies.     

Degree of learning,interpolated tests,and rate of forgetting

The purpose of the two experiments reported here was to observe the effects of degree of learning, interpolated tests, and retention interval, primarily on the rate of forgetting of a list of words, and secondarily on hypermnesia for those words. In the first experiment, all the subjects had one study trial on a list of 20 common words, followed by two tests of recall. Half of the subjects had further study and test trials until they had learned the words to a criterion of three correct consecutive recalls. Two days later, half of the subjects under each learning condition returned for four retention tests, and 16 days later, all the subjects returned for four tests. Experiment 2 was similar, except that all the subjects had at least three study trials followed by four recall tests on Day 1, intermediate tests were given 2 or 7 days later, and they all had final tests 14 days later. The results showed that rate of forgetting was attenuated by an additional intermediate set of tests but not by criterion learning. Hypermnesia was generally found over the tests that were given after a retention interval of 2 or more days. The best predictor of the amount of hypermnesia over a set of tests was the difference between overall cumulative recall and net recall on the first test of the set.     

Formation of the sameness-difference concept by Japanese monkeys from a small number of color stimuli

Japanese monkeys were trained to form the sameness-difference concept. In Experiment 1, four monkeys were trained with two colors to discriminate matching stimulus pairs from nonmatching pairs by reinforcing only lever-pressing responses to matching pairs with a variable-interval schedule. Three monkeys showed successful transfer of this discrimination to two new colors, thus demonstrating that some Japanese monkeys are able to form this relational concept from a minimum number of stimuli. In Experiment 2, two monkeys were trained, in a Yes/No procedure with three colors, to press one lever under matching pairs and another lever under nonmatching pairs. Poor transfer performances to three new colors suggest that simultaneously establishing two different response patterns to matching and nonmatching pairs is ineffective in forming the concept. In Experiment 3, the amount of transfer to three new colors after mastering a standard three-color matching-to-sample task was compared with that of a modified task in which correct responses were reinforced with a within-trial variable-interval schedule. All three monkeys showed greater transfer with the modified procedure. The results suggest that the variable-interval schedule adopted within trials is effective in forming the sameness-difference concept.     

Serial list combination by monkeys (Macaca mulatta): test cues and linking

This investigation assessed prospective bases of non-human primate cognitive operations that support serial list memory. Four macaques learned 3-, 5-item ordered lists of objects (as two-choice problems) and then either did or did not (in a within-subject design) receive training on pairs that linked the three original lists into a 15-item serial order. Next, subjects experienced selective exposure trials on object pairs that either maintained or contrasted to the serial position relationships seen during original learning. Subsequent comprehensive tests assessed the interactive effects of linking and exposure conditions on choosing in accord with a combined 15-item serial order. Linking readily induced monkeys to merge lists into a 15-item order, but restricting early exposure to pairs with the same positional relationships as original training slowed, but did not prevent, list combination. Exposure to positional relationships congruent with the combined (15-item) list and different from those of original 5-item training aided both expression of the linking effect and acquisition after no link training. Thus, list linking facilitated serial reorganization by inducing release from error derived from memory for prior learned positional relationships. The task was recommended as a prospective evaluator of continuity of cognitive processes among species.     

Spacing repetitions over 1 week

Does varying the spacing of repetitions over intervals as long as 1 week aftect recall? The answer from three experiments is yes. Subjects incidentally processed words repeated within a single list and words repeated in separate lists at list spacings of up to 1 week. Memory was tested by free recall shortly after the second presentations or after retention intervals of up to 1 week. Recall of the words repeated across separate lists conformed to a proportionality rule. When the retention interval is short relative to the spacing intervals, performance is inversely related to spacing. When the retention interval is a large proportion of the spacing intervals, performance is directly related to spacing. Does varying the spacing of repetitions within a single list affect recall after a retention interval of 2 weeks? The answer depends on the processing used while studying the words. Processing that generated interitem associations resulted in a within-list spacing effect even after a 2-week retention interval. Without the interitem associations, the effect was absent after a 1-day retention interval. Most of these findings were explained by examining the changing relationship between the retrieval context and the context stored during study.     

Some tests of response membership in acquired equivalence classes

Pigeons were trained on many-to-one matching in which pairs of samples, each consisting of a visual stimulus and a distinctive pattern of center-key responding, occasioned the same reinforced comparison choice. Acquired equivalence between the visual and response samples then was evaluated by reinforcing new comparison choices to one set of samples, and examining generalization of these choices to the other samples. Three separate experiments found no evidence of such generalization, as indexed by performance on class-consistent versus class-inconsistent tests. Other tests showed that the pigeons' center-key response patterns during training had indeed served as a conditional cue for choice. These results do not support the hypothesis that different defined responses can become members of acquired equivalence classes.