There are stable individual differences in performance variability, both from moment to moment and from day to day

Individual differences in decision speed have been regarded as direct reflections of a "primitive" functional neurophysiological characteristic, which affects performance on all cognitive tasks and so may be regarded as the "biological basis of intelligence", or of age-related changes in mental abilities. More detailed analyses show that variability within an experimental session (WSV) is a stable individual difference characteristic and that mean choice reaction times (CRTs) are gross summary statistics that reflect variability, rather than maximum speed of performance. A total of 98 people aged from 60 to 80 years completed 36 weekly sessions on six different letter categorization tasks. After effects of practice and of circadian variability had been eliminated, individuals with lower scores on the Cattell Culture Fair intelligence test had slower CRTs and greater WSV on all tasks. A simulation study showed that the greater WSVs of low Cattell scorers led directly to the significantly greater variability of their mean CRTs from session to session. However because CRTs on tasks co-varied from session to session it was apparent that, besides being affected by WSV, individuals' between-session variabilities (BSVs) also vary because of state changes that affect their performance from day to day. It seems that both variability in performance from trial to trial during a session and variability in average performance from day to day are correlated, stable, individual difference characteristics that vary inversely with intelligence test performance. Methodological consequences of these results for interpretations of age-related cognitive changes, for variability between as well as within individuals, for individual differences in decision speed, and for circadian variability in performance are discussed.     

Number and time in acquisition,extinction and recovery

We measured rate of acquisition, trials to extinction, cumulative responses in extinction, and the spontaneous recovery of anticipatory hopper poking in a Pavlovian protocol with mouse subjects. We varied by factors of 4 number of sessions, trials per session, intersession interval, and span of training (number of days over which training extended). We find that different variables affect each measure: Rate of acquisition [1/(trials to acquisition)] is faster when there are fewer trials per session. Terminal rate of responding is faster when there are more total training trials. Trials to extinction and amount of responding during extinction are unaffected by these variables. The number of training trials has no effect on recovery in a 4-trial probe session 21 days after extinction. However, recovery is greater when the span of training is greater, regardless of how many sessions there are within that span. Our results and those of others suggest that the numbers and durations and spacings of longer-duration “episodes” in a conditioning protocol (sessions and the spans in days of training and extinction) are important variables and that different variables affect different aspects of subjects' behavior. We discuss the theoretical and clinical implications of these and related findings and conclusions—for theories of conditioning and for neuroscience.     

Repeated acquisition and performance chamber for mice: a paradigm for assessment of spatial learning and memory

Molecular genetic manipulation of the mouse offers the possibility of elucidating the function of individual gene products in neural systems underlying learning and memory. Many extant learning paradigms for mice rely on negative reinforcement, involve simple problems that are relatively rapidly acquired and thus preclude time-course assessment, and may impose the need to undertake additional experiments to determine the extent to which noncognitive behaviors influence the measures of learning. To overcome such limitations, a multiple schedule of repeated acquisition and performance was behaviorally engineered to assess learning vs rote performance within-behavioral test session and within-subject utilizing an apparatus modified from the rat (the repeated acquisition and performance chamber; RAPC). The multiple schedule required mice to learn a new sequence of door openings leading to saccharin availability in the learning component during each session, while the sequence of door openings for the performance component remained constant across sessions. The learning and performance components alternated over the course of each test session, with different auditory stimuli signaling which component was currently in effect. To validate this paradigm, learning vs performance was evaluated in two inbred strains of mice: C57BL/6J and 129/SvJ. The hippocampal dependence of this measure was examined in lesioned C57BL/6J mice. Both strains exhibited longer latencies and higher errors in the learning compared to the performance component and evidenced declines in both measures across the trials of each session, consistent with an acquisition phenomenon. These same measures showed little or no evidence of change in the performance component. Whereas three trials per session were utilized with C57BL/65 mice in each component, behavior of 129/SvJ mice could only be sustained for two trials per component per session, demonstrating differences in testing capabilities between these two strains under these experimental conditions and thus precluding the ability to make systematic strain comparisons of learning capabilities. Hippocampal lesions in C57BL/6J mice resulted in substantially longer latencies and increased errors in the learning but not the performance component, demonstrating the importance of this region to spatial learning as measured in the RAPC. In aggregate, this positive reinforcement-based operant paradigm to evaluate murine spatial learning detects strain differences and hippocampal dependence and permits explicit differentiation of the impact of noncognitive contributions to learning measures on a within-subject, within-session basis.     

Visuo-spatial path learning,stress, and cortisol secretion following military cadets’ first parachute jump: the effect of increasing task complexity

The present field experiment examined how multi-trial visuo-spatial learning and memory performance are impacted by excessive arousal, instigated by a potentially life-threatening event (i.e., a first parachute jump). Throughout a parachute training activity, subjective and neuroendocrine (i.e., cortisol) stress levels were assessed of 61 male military cadets who were randomly assigned to a control (n = 30) or a jump stress condition (n = 31). Post-stress learning and memory capacity was assessed with a 10-trial path-learning task that permitted emergence of learning curves. Pre-activity cortisol concentrations indicated a significant neuroendocrine anticipatory stress response in the stress group. Following parachuting, subjective stress levels and salivary cortisol reactivity differed significantly between groups. Visuo-spatial path-learning performance was impaired significantly after jump stress exposure, relative to the control group. Moreover, examination of the learning curves showed similar learning and memory performance at onset of the trials, with curves bifurcating as the task became more complex. These findings are in accordance with leading theories that acknowledge a moderating effect of task complexity. In sum, the present study extends knowledge concerning anticipatory stress effects, endogenously instigated cortisol reactivity, and the influence of extreme arousal on visuo-spatial path learning.     

Chronic stress modulates the use of spatial and stimulus-response learning strategies in mice and man

Acute stress modulates multiple memory systems in favor of caudate nucleus-dependent stimulus-response and at the expense of hippocampus-dependent spatial learning and memory. We examined in mice and humans whether chronic stress has similar consequences. Male C57BL/6J mice that had been repeatedly exposed to rats ("rat stress") used in a circular hole board task significantly more often a stimulus-response strategy (33%) than control mice (0%). While velocity was increased, differences in latency to exit hole, distance moved or number of holes visited were not observed. Increased velocity and performance during retention trials one day later indicates altered emotionality and motivation to explore in rat stressed mice. Forty healthy young men and women were split into "high chronic stress" and "low chronic stress" groups based on their answers in a chronic stress questionnaire ("Trier Inventory of Chronic Stress"-TICS) and trained in a 2D task. A test trial immediately after training revealed that participants of the "high chronic stress" group used the S-R strategy significantly more often (94%) than participants of the "low chronic stress" group (52%). Verbal self-reports confirmed the strategy derived from participants' choice in the test trial. Learning performance was unaffected by the chronic stress level. We conclude that one consequence of chronic stress is the shift to more rigid stimulus-response learning, that is accompanied by changes in motivational factors in mice.     

Oculomotor evidence of sequence learning on the serial reaction time task

Manual and oculomotor measures of sequence learning were examined on the serial reaction time (SRT) task Participants were assigned into four groups differing on response modality (manual, oculomotor) and trial type (sequence, pseudorandom). The pattern of manual RTs replicated previous studies. Frequency of anticipatory eye movements followed similar patterns as RTs. Participants made many anticipations, even in pseudorandom blocks, and frequency of anticipations did not depend on presence of concurrent manual responses. Excluding participants with explicit awareness did not change results. Anticipations were negatively related to RTs in both incidental and intentional learning. Anticipations were positively related to sequence recall in intentional, but not incidental, learning. Results suggest that (1) anticipatory eye movements reflected sequence learning and (2) participants made overt and covert shifts of visuospatial attention to likely stimulus locations prior to stimulus onset, whether or not they made manual responses and whether or not there was a sequence.     

Effects of cocaine and d-amphetamine on the repeated acquisition and performance of conditional discriminations.

The acute and chronic effects of cocaine and d-amphetamine on food-reinforced behavior were investigated in pigeons responding on a two-component multiple schedule. In one component, the behavioral task consisted of the same chain of conditional discriminations each session (performance). In the other component, the chain of conditional discriminations was changed from session to session (learning). In comparison to control sessions, both acute cocaine and d-amphetamine increased errors in each component of the multiple schedule. Responding in the learning component, however, was generally disrupted at lower doses than those that affected responding in the performance component. At high doses, both drugs produced pauses in responding in each component in three of the four subjects. Pausing engendered by d-amphetamine was approximately twice as long as that under cocaine. Upon chronic administration, both the pausing and error-increasing effects of each drug diminished. Drug-induced changes in timeout responding, however, did not decrease during chronic administration. Redeterminations of the d-amphetamine dose-effect curves following chronic cocaine administration suggested the existence of cross-tolerance between cocaine and d-amphetamine. Both the acute and chronic data are consistent with the view that conditions of stimulus control may modulate the behavioral effects of drugs.     

Bouts of responding on variable-interval schedules: effects of deprivation level

Rats obtained food pellets on a variable-interval schedule of reinforcement by nose poking a lighted key. After training to establish baseline performance (with the mean variable interval set at either 60, 120, or 240 s), the rats were given free access to food during the hour just before their daily session. This satiation operation reduced the rate of key poking. Analysis of the interresponse time distributions (log survivor plots) indicated that key poking occurred in bouts. Prefeeding lengthened the pauses between bouts, shortened the length of bouts (less reliably), and had a relatively small decremental effect on the response rate within bouts. That deprivation level affects mainly between-bout pauses has been reported previously with fixed-ratio schedules. Thus, when the focus is on bouts, the performances maintained by variable-interval schedules and fixed-ratio schedules are similarly affected by deprivation.     

Responding changes systematically within sessions during conditioning procedures.

When the procedure is held constant within an experimental session, responding often changes systematically within that session. Many of these within-session changes in responding cannot be dismissed as learning curves or by-products of satiation. They have been observed in studies of positive reinforcement, avoidance, punishment, extinction, discrimination, delayed matching to sample, concept formation, maze and alley running, and laboratory analogues of foraging, as well as in the unconditioned substrates of conditioned behavior. When aversive stimuli are used, responding usually increases early in the session. When positive reinforcers are used, responding changes in a variety of ways, including increasing, decreasing, and bitonic functions. Both strong and minimal reinforcement procedures produce within-session decreases in positively reinforced behavior. Within-session changes in responding have substantial theoretical and methodological implications for research in conditioning.     

On rotating to smooth functions

Tucker has outlined an application of principal components analysis to a set of learning curves, for the purpose of identifying meaningful dimensions of individual differences in learning tasks. Since the principal components are defined in terms of a statistical criterion (maximum variance accounted for) rather than a substantive one, it is typically desirable to rotate the components to a more interpretable orientation. Simple structure is not a particularly appealing consideration for such a rotation; it is more reasonable to believe that any meaningful factor should form a (locally) smooth curve when the component loadings are plotted against trial number. Accordingly, this paper develops a procedure for transforming an arbitrary set of component reference curves to a new set which are mutually orthogonal and, subject to orthogonality, are as smooth as possible in a well defined (least squares) sense. Potential applications to learning data, electrophysiological responses, and growth data are indicated.Portions of this research were supported by the National Research Council of Canada, Grant A8615 to the second author. We thank Jagdeth Sheth for supplying his raw data.     

Pigeons perform poorly on a midsession reversal task without rigid temporal regularity

Animals make surprising anticipatory and perseverative errors when faced with a midsession reversal of reinforcer contingencies on a choice task with highly predictable stimulus–time relationships. In the current study, we asked whether pigeons would anticipate changes in reinforcement when the reinforcer contingencies for each stimulus were not fixed in time. We compared the responses of pigeons on a simultaneous choice task when the initially correct stimulus was randomized or alternated across sessions. Pigeons showed more errors overall compared with the typical results of a standard midsession reversal procedure, and they did not show the typical anticipatory errors prior to the contingency reversal. Probe tests that manipulated the spacing between trials also suggested that timing of the session exerted little control of pigeons’ behavior. The temporal structure of the experimental session thus appears to be an important determinant for animals’ use of time in midsession reversal procedures.     

Step changes in the intertrial interval in the midsession reversal task: Predicting pigeons' performance with the learning-to-time model

Our goal was to assess the role of timing in pigeons' performance in the midsession reversal task. In discrete-trial sessions, pigeons learned to discriminate between 2 stimuli, S1 and S2. Choices of S1 were reinforced only in the first half of the session and choices of S2 were reinforced only in the second half. Typically, pigeons choose S2 before the contingency reverses (anticipatory errors) and S1 after (perseverative errors), suggesting that they time the interval from the beginning of the session to the contingency reversal. To test this hypothesis, we exposed pigeons to a midsession reversal task and, depending on the group, either increased or decreased the ITI duration. We then contrasted the pigeons' performance with the predictions of the Learning-to-Time (LeT) model: In both conditions, preference was expected to reverse at the same time as in the previous sessions. When the ITI was doubled, pigeons' preference reversal occurred at half the trial number but at the same time as in the previous sessions. When the ITI was halved, pigeons' preference reversal occurred at a later trial but at an earlier time than in the previous sessions. Hence, pigeons' performance was only partially consistent with the predictions of LeT, suggesting that besides timing, other sources of control, such as the outcome of previous trials, seem to influence choice.     

Demonstration of nondeclarative sequence learning in mice: development of an animal analog of the human serial reaction time task

In this paper, we demonstrate nondeclarative sequence learning in mice using an animal analog of the human serial reaction time task (SRT) that uses a within-group comparison of behavior in response to a repeating sequence versus a random sequence. Ten female B6CBA mice performed eleven 96-trial sessions containing 24 repetitions of a 4-trial sequence. During the 12th session, the repeating sequence was replaced with the random sequence halfway through the session. Reaction time (RT) to respond to an illuminated nose-poke was recorded, and performance was compared at the halfway point in each session to test for any change in behavior. For learning effect, RTs decreased over the no-switch repeating-sequence sessions. For interference effect, behavior did not change appreciably at the halfway point during the last repeating-sequence session. However, RTs deteriorated significantly after the switch from repeating to random sequences halfway through session 12. The mice demonstrated a robust interference effect when switched from repeating to random sequences. This pattern of behavior in humans performing the SRT is interpreted as evidence of nondeclarative sequence learning. The similarity between the human and mouse SRTs will enable more direct comparisons of mouse-human nondeclarative memory behavior and will provide a useful behavioral end-point in mouse-models of basal ganglia dysfunction.     

Within-session reversal learning in rhesus macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

In a midsession reversal (MSR) task, animals are typically presented with a simple, simultaneous discrimination (S1+, S2?) where contingencies are reversed (S1?, S2+) half-way through each session. This paradigm creates multiple, relevant cues that can aid in maximizing overall reinforcement. Recent research has shown that pigeons show systematic anticipatory and perseverative errors across the session, which increase as a function of proximity to the reversal trial. This behavior has been theorized to indicate primary control by temporal cues across the session, instead of the cues provided by recent reinforcement history that appear to control behavior shown by humans. Rats, however, appear to be guided by recent reinforcement history when tested in an operant context, thereby demonstrating behavior that parallels that seen in humans, but they appear to be guided by temporal cues when tested in an open-field apparatus, showing behavior more akin to that seen in pigeons. We tested rhesus macaques (Macaca mulatta) on the MSR with a computerized simultaneous visual discrimination to assess whether they would show errors indicative of control by time or by recent reinforcement history. When a single reversal point occurred midsession, rhesus macaques showed no anticipation of the reversal and a similar level of perseveration to rats tested in an operant setting. Nearly identical results also were observed when the monkeys were trained with a single, variable reversal point or with multiple, variable reversal points within a session. These results indicate that temporal cues are not guiding response flexibility in rhesus macaque visual discrimination.     

FUNCTIONAL ANALYSIS AND TREATMENT OF EYE POKING

In four studies we analyzed the eye poking of a youth with profound disabilities. In Study 1, a functional analysis showed that eye poking occurred during the no-attention condition, but not during demand, attention, or recreation conditions. The analysis did not identify socially mediated variables involved in the maintenance of eye poking; rather, eye poking may have been maintained by consequences produced directly by the response. In Study 2 we had the student wear goggles to prevent potential reinforcement from finger—eye contact. The results of Study 2 indicated that eye-poking attempts were reduced when the student wore goggles. We then tested in Study 3 the effects of two alternative topographies of stimulation. Study 3 demonstrated that eye poking was reduced when a video game was provided as a competing source of visual stimulation, and that music was less effective in reducing eye poking. In Study 4, a contingency analysis using the video game was conducted in an attempt to (a) reduce the frequency of eye poking and (b) study whether the video game functioned as a reinforcer. The results of Study 4 demonstrated substantive reductions in the frequency of eye poking, and suggested that the video game served as a reinforcer.     

The Strategy-Specific Nature of Improvement: The Power Law Applies by Strategy Within Task

If strategy shifts speed up performance, learning curves should show discontinuities where such shifts occur. Relatively smooth curves appear consistently in the literature, however. To explore this incongruity, we examined learning when multiple strategies were used. We plotted power law learning curves for aggregated data from four mental arithmetic experiments and then plotted similar curves separately for each participant and strategy. We then evaluated the fits achieved by each group of curves. In all four experiments, plotting separately by strategy produced significantly better fits to individual participants' data than did plotting a single power function. We conclude that improvement of solution time is better explained by practice on a strategy rather than practice on a task, and that careful assessment of trial-by-trial changes in strategy can improve understanding of the effects of practice on learning.     

Monaural Auditory Cue Affects the Process of Choosing the Initial Swing Leg in Gait Initiation

The authors investigated the effect of an auditory cue on the choice of the initial swing leg in gait initiation. Healthy humans initiated a gait in response to a monaural or binaural auditory cue. When the auditory cue was given in the ear ipsilateral to the preferred leg side, the participants consistently initiated their gait with the preferred leg. In the session in which the side of the monaural auditory cue was altered trial by trial randomly, the probability of initiating the gait with the nonpreferred leg increased when the auditory cue was given in the ear contralateral to the preferred leg side. The probability of choosing the nonpreferred leg did not increase significantly when the auditory cue was given in the ear contralateral to the preferred leg side in the session in which the auditory cue was constantly given in the ear contralateral to the preferred leg side. The reaction time of anticipatory postural adjustment was shortened, but the probability of choosing the nonpreferred leg was not significantly increased when the gait was initiated in response to a binaural auditory cue. An auditory cue in the ear contralateral to the preferred leg side weakens the preference for choosing the preferred leg as the initial swing leg in gait initiation when the side of the auditory cue is unpredictable.     

The “pure-study” learning curve: The learning curve without cumulative testing

The customary assumption in the study of human learning using alternating study and test trials is that learning occurs during study trials and that test trials are useful only to measure learning. In fact, tests seem to play little role in the development of learning, because the learning curve is similar even when the number of test trials varies widely (Tulving, Journal of Verbal Learning and Verbal Behavior 6:175-184, 1967). However, this outcome seems odd, because other research has shown that testing fosters greater long-term learning than does studying. We report three experiments addressing whether tests affect the shape of the learning curve. In two of the experiments, we examined this issue by varying the number of spaced study trials in a sequence and examining performance on only a single test trial at the end of the series (a "pure-study" learning curve). We compared these pure-study learning curves to standard learning curves and found that the standard curves increase more rapidly and reach a higher level in both free recall (Exp. 1) and paired-associate learning (Exp. 2). In Experiment 3, we provided additional study trials in the "pure-study" condition to determine whether the standard (study-test) condition would prove superior to a study-study condition. The standard condition still produced better retention on both immediate and delayed tests. Our experiments show that test trials play an important role in the development of learning using both free-recall (Exps. 1 and 3) and paired-associate (Exp. 2) procedures. Theories of learning have emphasized processes that occur during study, but our results show that processes engaged during tests are also critical.     

The influence of passive preexposure on escape from a Morris pool

In 3 experiments rats were preexposed to the landmarks that surround a Morris pool by being placed on a submerged platform within the pool. They were then required to escape from the pool by swimming to the platform, which was in a location that had not been used during preexposure. Preexposure facilitated subsequent escape from the pool, provided that the platform was not moved during preexposure and the relative position of the landmarks to each other remained constant throughout preexposure. In contrast, if during preexposure the platform was moved from session to session (Experiment 1), or the array of landmarks was altered unsystematically from trial to trial (Experiments 2 and 3), then subsequent learning to escape from the pool was disrupted. These findings suggest that the effects of preexposure to the landmarks in a Morris pool is determined by whether or not they are of relevance for identifying the location of the platform. When they are relevant, then subsequent learning is facilitated, but when they are irrelevant, then subsequent learning is disrupted.