Manipulating decision processes in the human scalar timing system

Two experiments attempted to manipulate the decision processes used in a temporal generalisation task with humans. In Experiment 1, payoffs (points awarded or deducted) were used to try to alter behaviour when the standard duration was 400ms, and the comparison durations ranged from 100 to 700ms in 100ms steps. Two conditions which either encouraged or discouraged the subject to identify a comparison duration as the standard were compared with a neutral condition. Encouraging identifications of the standard increased the proportion of identifications of the standard, whereas the discouraging manipulation had more ambiguous effects. Using the "modified Church and Gibbon" model, it was shown that the effect of the encourage manipulation was an increase in the response threshold, consistent with the information-processing version of scalar timing theory. A second experiment compared encourage and discourage manipulations with a more difficult discrimination (comparison durations spaced in 50ms steps around the 400ms standard), and with more distinct payoff differences for the different conditions. Behavioural effects were much more marked in Experiment 2, with the encourage condition producing more identifications of a comparison duration as the standard for all comparison durations except the shortest, compared with the discourage condition. Modelling showed that the main theoretical difference between the two conditions was in a change in the response threshold, in a manner consistent with the scalar timing model.     

Applications of timing theories to a peak procedure

This article describes applications of scalar expectancy theory (SET), learning-to-time theory (LeT), and Packet theory to data from a peak procedure. Twelve rats were trained in a multiple cued-interval procedure with two fixed intervals (60 and 120 s) signaled by houselight and white noise. Twenty-five percent of the cycles were nonfood cycles, which were 360 s long and had no reinforcement. Mean and individual response rates on nonfood cycles were fitted with explicit solutions of SET, LeT and Packet theory. Applications of the three timing theories were compared in terms of goodness of fit and complexity.     

Pavlov as a psychologist. A reappraisal

American psychologists are informed on Pavlov's work on conditional reflexes but not on the full development of his theory of higher nervous activity. This article shows that Pavlov's theory of higher nervous activity dealt with concepts that concerned contemporary psychologists. Pavlov used the conditioning of the salivary reflex for methodological purposes. Pavlov's theory of higher nervous activity encompassed overt behavior, neural processes, and the conscious experience. The strong Darwinian element of Pavlov's theory, with its stress on the higher organisms' adaptation, is described. With regard to learning, Pavlov, at the end of his scholarly career, proposed that although all learning involves the formation of associations, the organism's adaptation to the environment is established through conditioning, but the accumulation of knowledge is established by trial and error.     

Counting in a time discrimination task in children and adults

The present study investigated in 5- and 8-year-olds, as well as in adults, the effect of verbal counting on temporal discrimination behavior in a generalization task with two duration ranges in order to test the scalar timing property. The results showed that counting improved temporal sensitivity in all age groups, although sensitivity to time remained lower in the younger children. Furthermore, in the 5-year-olds, the temporal generalization behavior conformed well to the scalar property of variance both in the counting and the non-counting condition. However, this conformity to the scalar timing property disappeared when counting was used in the 8-year-olds and the adults. The development of the ability to count time at a constant rhythm is discussed as the major reason for this departure of temporal behavior from the scalar property of variance when counting is employed.     

How rats combine temporal cues

The procedures for classical and operant conditioning, and for many timing procedures, involve the delivery of reinforcers that may be related to the time of previous reinforcers and responses, and to the time of onsets and terminations of stimuli. The behavior resulting from such procedures can be described as bouts of responding that occur in some pattern at some rate. A packet theory of timing and conditioning is described that accounts for such behavior under a wide range of procedures. Applications include the food searching by rats in Skinner boxes under conditions of fixed and random reinforcement, brief and sustained stimuli, and several response-food contingencies. The approach is used to describe how multiple cues from reinforcers and stimuli combine to determine the rate and pattern of response bouts.     

The generality of empirical and theoretical explanations of behavior

For theoretical explanations of data, parameter values estimated from a single dependent measure from one procedure are used to predict alternative dependent measures from many procedures. Theoretical explanations were compared to empirical explanations of data in which known functions and principles were used to fit only selected dependent measures. The comparison focused on the ability of theoretical and empirical explanations to generalize across samples of the data, across dependent measures of behavior, and across different procedures. Rat and human data from fixed-interval and peak procedures, in which principles (e.g., scalar timing) are well known, were described and fit by a theory with independent modules for perception, memory, and decision. The theoretical approach consisted of fitting closed-form equations of the theory to response rate gradients calculated from the data, simulating responses using parameter values previously estimated, and comparing theoretical predictions with dependent measures not used to estimate parameters. Although the empirical and theoretical explanations provided similar fits to the response rate gradients that generalized across samples and had the same number of parameters, only the theoretical explanation generalized across procedures and dependent measures.     

An application of the active time model to multiple concurrent variable-interval schedules

The current experiment investigates whether an active time model can account for anomalous results that have emerged from multiple schedule, concurrent variable-interval (VI) VI experiments. The model assumes that (1) during concurrent VI VI training pigeons learn a function that relates time since the most immediate response, i.e., active time, to changeover probabilities and (2) that molar preference is the result of an interaction between inter-response time frequencies and the learned active time changeover functions. Pigeons were trained under a concurrent VI 30-s VI 30-s schedule and a concurrent VI 60-s VI 60-s schedule. Probes were conducted in which VI 30-s and VI 60-s stimuli were paired. During these probes, birds allocated choices equally to the stimuli. The active time model accurately fit individual subject data. In contrast data were not fit by a variant of scalar expectancy theory proposed by Gibbon [Gibbon, J., 1995. Dynamics of time matching: arousal makes better seem worse. Psychon. Bull. Rev. 2, 208-215].     

Implicit, predictive timing draws upon the same scalar representation of time as explicit timing

It is not yet known whether the scalar properties of explicit timing are also displayed by more implicit, predictive forms of timing. We investigated whether performance in both explicit and predictive timing tasks conformed to the two psychophysical properties of scalar timing: the Psychophysical law and Weber's law. Our explicit temporal generalization task required overt estimation of the duration of an empty interval bounded by visual markers, whereas our temporal expectancy task presented visual stimuli at temporally predictable intervals, which facilitated motor preparation thus speeding target detection. The Psychophysical Law and Weber's Law were modeled, respectively, by (1) the functional dependence between mean subjective time and real time (2) the linearity of the relationship between timing variability and duration. Results showed that performance for predictive, as well as explicit, timing conformed to both psychophysical properties of interval timing. Both tasks showed the same linear relationship between subjective and real time, demonstrating that the same representational mechanism is engaged whether it is transferred into an overt estimate of duration or used to optimise sensorimotor behavior. Moreover, variability increased with increasing duration during both tasks, consistent with a scalar representation of time in both predictive and explicit timing. However, timing variability was greater during predictive timing, at least for durations greater than 200 msec, and ascribable to temporal, rather than non-temporal, mechanisms engaged by the task. These results suggest that although the same internal representation of time was used in both tasks, its external manifestation varied as a function of temporal task goals.     

Behavioral pharmacology and timing

Drug effects on temporally patterned behavior are often described under the rubric of rate dependency: the effect of a drug on behavior is related to the rate of behavior in the absence of the drug. Specifically, drugs increase low rate behavior and decrease high rate behavior. These same types of effects are interpreted in the timing literature, however, as selective changes in temporal discrimination. The present series of experiments arrange situations that allow divergent predictions based on the two interpretations. In one component of a multiple schedule, when the response key is lit blue, food is available after the houselight is presented for a short duration (5 s). In the other component of the multiple schedule, when the response key is lit green, food is available after the houselight is presented for a long duration (30 s). No food is available after intermediate durations. Specific focus is given to a neuropharmacological information-processing model of timing. Predictions were compared for drugs that are thought to affect the clock and memory stages in the model. The results do not generally lend support for the neuropharmacological interpretation of the scalar expectancy theory, but emphasize the need for an explanatory mechanism that is consistent with the empirical generalization of rate dependency.     

An Operant Conditioning Method for Studying Auditory Behaviors in Marmoset Monkeys

The common marmoset (Callithrix jacchus) is a small New World primate that has increasingly been used as a non-human model in the fields of sensory, motor, and cognitive neuroscience. However, little knowledge exists regarding behavioral methods in this species. Developing an understanding of the neural basis of perception and cognition in an animal model requires measurement of both brain activity and behavior. Here we describe an operant conditioning behavioral training method developed to allow controlled psychoacoustic measurements in marmosets. We demonstrate that marmosets can be trained to consistently perform a Go/No-Go auditory task in which a subject licks at a feeding tube when it detects a sound. Correct responses result in delivery of a food reward. Crucially, this operant conditioning task generates little body movement and is well suited for pairing behavior with single-unit electrophysiology. Successful implementation of an operant conditioning behavior opens the door to a wide range of new studies in the field of auditory neuroscience using the marmoset as a model system.     

Comparison of interval timing behaviour in mice following dorsal or ventral hippocampal lesions with mice having δ-opioid receptor gene deletion

Mice with cytotoxic lesions of the dorsal hippocampus (DH) underestimated 15 s and 45 s target durations in a bi-peak procedure as evidenced by proportional leftward shifts of the peak functions that emerged during training as a result of decreases in both ‘start’ and ‘stop’ times. In contrast, mice with lesions of the ventral hippocampus (VH) displayed rightward shifts that were immediately present and were largely limited to increases in the ‘stop’ time for the 45 s target duration. Moreover, the effects of the DH lesions were congruent with the scalar property of interval timing in that the 15 s and 45 s functions superimposed when plotted on a relative timescale, whereas the effects of the VH lesions violated the scalar property. Mice with DH lesions also showed enhanced reversal learning in comparison to control and VH lesioned mice. These results are compared with the timing distortions observed in mice lacking δ-opioid receptors (Oprd1^−/−) which were similar to mice with DH lesions. Taken together, these results suggest a balance between hippocampal–striatal interactions for interval timing and demonstrate possible functional dissociations along the septotemporal axis of the hippocampus in terms of motivation, timed response thresholds and encoding in temporal memory.     

Quantitative analysis of activation and inactivation of asymmetry currents in biological membranes,based on a conformational transition model

Summary A basic voltage-dependent conformational transition mechanism is proposed. It comprises one relatively fast conversion between two individual states which are comparatively slowly coupled with a third state. Having introduced voltage as an additional parameter of state, standard methods of thermodynamics and rate theory are employed to describe the equilibrium and kinetic behavior of the system. In particular, a quantitative discussion is given regarding the asymmetrical displacement currents generated by switching on and off a voltage pulse. Effects of temperature pulse duration, and application of a conditioning prepulse are examined. The results provide a comprehensive basis for a quantitative analysis of pertinent experimental work. The so far presented measuring data can indeed be very well described along these lines.     

When to respond? And how much? : Temporal control and response output on mixed-fixed-interval schedules with unequally probable components

Rats were trained on mixed-fixed-interval (FI) schedules, with component FIs of 30 and 60s. The probability of reinforcement according to FI 30s varied between conditions, across values of 0.1, 0.3, 0.5, 0.7 and 0.9. When response rate in the 60s intervals was measured, separate response peaks, one close to 30s, the other at 60s, could be identified when the probability of reinforcement at 30s was 0.3 or greater. Nonlinear regression found that the location of the earlier peak was always close to 30s, that the coefficient of variation of the response functions at 30 and 60s were unaffected by reinforcement probability, but that the 30s component appeared to be timed slightly more precisely than the 60s one. Response rate at around 30s increased with increasing probability of reinforcement according to FI 30s, but responding at 60s was unaffected by reinforcement probability. The data are discussed with respect to a number of contemporary models of animal timing (scalar expectancy theory, the Behavioural Theory of Timing and the Learning to Time model), and a recent account of response output on FI-like schedules.     

Interaction between Purkinje cells and inhibitory interneurons may create adjustable output waveforms to generate timed cerebellar output

We develop a new model that explains how the cerebellum may generate the timing in classical delay eyeblink conditioning. Recent studies show that both Purkinje cells (PCs) and inhibitory interneurons (INs) have parallel signal processing streams with two time scales: an AMPA receptor-mediated fast process and a metabotropic glutamate receptor (mGluR)-mediated slow process. Moreover, one consistent finding is an increased excitability of PC dendrites (in Larsell's lobule HVI) in animals when they acquire the classical delay eyeblink conditioning naturally, in contrast to in vitro studies, where learning involves long-term depression (LTD). Our model proposes that the delayed response comes from the slow dynamics of mGluR-mediated IP3 activation, and the ensuing calcium concentration change, and not from LTP/LTD. The conditioned stimulus (tone), arriving on the parallel fibers, triggers this slow activation in INs and PC spines. These excitatory (from PC spines) and inhibitory (from INs) signals then interact at the PC dendrites to generate variable waveforms of PC activation. When the unconditioned stimulus (puff), arriving on the climbing fibers, is coupled frequently with this slow activation the waveform is amplified (due to an increased excitability) and leads to a timed pause in the PC population. The disinhibition of deep cerebellar nuclei by this timed pause causes the delayed conditioned response. This suggested PC-IN interaction emphasizes a richer role of the INs in learning and also conforms to the recent evidence that mGluR in the cerebellar cortex may participate in slow motor execution. We show that the suggested mechanism can endow the cerebellar cortex with the versatility to learn almost any temporal pattern, in addition to those that arise in classical conditioning.     

Conditioning and learning in relation to disease

Of the two generally recognized processes through which learning occurs--imprinting and conditioning--only the latter with its two paradigms, classical and operant, has both practical and heuristic implications for disease. From the classical conditioning experiments of Pavlov's laboratory over 100 years ago to the later work in operant conditioning by Skinner and others in the past four decades has evolved much of the basis of modern learning theory and its applications to disease in the form of behavior therapy. Variants of behavior therapy have been employed in the treatment of wide variety of medical and psychiatric illnesses. Recent developments in the study of brain function and biochemistry have led to renewed interest in the conditioning paradigm and its value as tool in these areas of research.     

Operant conditioning in invertebrates

Learning to anticipate future events on the basis of past experience with the consequences of one's own behavior (operant conditioning) is a simple form of learning that humans share with most other animals, including invertebrates. Three model organisms have recently made significant contributions towards a mechanistic model of operant conditioning, because of their special technical advantages. Research using the fruit fly Drosophila melanogaster implicated the ignorant gene in operant conditioning in the heat-box, research on the sea slug Aplysia californica contributed a cellular mechanism of behavior selection at a convergence point of operant behavior and reward, and research on the pond snail Lymnaea stagnalis elucidated the role of a behavior-initiating neuron in operant conditioning. These insights demonstrate the usefulness of a variety of invertebrate model systems to complement and stimulate research in vertebrates.     

Cooperative activity of motor and frontal cortex neurons in trained cats systemically administered M-cholinoreceptor blockers]

It was shown previously that peripherally administered antagonists of the central 1 M-cholinoreceptors led to a selective impairment of bar-pressing response in a food-reinforced operant conditioned task but did not alter contextual behavior and functions such as motivation, perception, and locomotion. To obtain information about the central mechanisms of the conditioning impairment, we recorded simultaneously the extracellular multiunit activity from the frontal and motor neocortical areas of five cats trained to acquisition criteria in a food-reinforced operant conditioning task. Multiunit recordings were performed drur 1) normal conditioning; 2) conditioning during subcutaneous administration of muscarinic antagonists scopolamine (0.03 mg/kg), trihexyphenidyl (1 mg/kg), and methylscopolamine (0.03 mg/kg). Autocorrelation analysis showed that scopolamine and trihexyphenidyl but not methylscopolamine led to a significant increase in the tendency of cortical cells to fire in a cyclic way (i.e., the shift of the firing pattern from a single-spike discharge to burst, rhythmic, or rhythmic-burst discharge) both in the motor and frontal areas. Cross-correlation analysis showed that the bursting and rhythmic-bursting cells synchronized their activity within and (in a number of cases) between the cortical areas. These changes in the neuronal activity within the motor cortex and frontal cortex were accompanied by a significant decrease in the functional connectivity both inside and between the cortical areas in parallel with selective impairment of the conditioned response.     

An introduction to the psychophysiology of breathing

Breathing can be viewed as an independent variable which affects emotion, cognition, and behavior as well as a dependent variable which reflects changes in emotion, cognition, and behavior. This bidirectional interaction is basic to an appreciation of the significance of breathing in terms of its relevance in research and application. The underlying premise of the present article is that since breathing is a behavior that is under voluntary as well as reflexive control, it can be modified according to the principles of both instrumental training (operant conditioning) and Pavlovian (classical) conditioning. The implications of this premise are relevant to theory, diagnosis, and treatment of stress and anxiety-related disorders (e.g., panic disorder, phobias, test anxiety, occupational strain, and related psychosomatic disorders), and to basic and applied research in the psychophysiology of breathing.     

Preschool-aged children’s jumps: Imitation performances

Imitative behavior underlaid by perception and action links during children’s development in complex locomotor skills has been the object of relatively few studies. In order to explore children’s motor coordination modes, 130 children divided into five age groups from 3.5 to 7.5 years were instructed to imitate jumping tasks in spontaneous motor situation and in various imitative contexts by an adult providing verbal orders and gestural demonstrations. Their conformity to the model, stability and variability scores were coded from a video analysis when they performed jumps with obstacles. To evaluate their postural-motor control level, the durations of the preparatory phase and jumping flights were also timed. Results showed that all age groups generated the demonstrator’s goal but not necessarily the same coordination modes of jumping. In imitation with temporal proximity, the model helped the youngest age groups to adopt his coordination modes and stabilized only the oldest age groups’ performances starting from 5.5 years old, without effect on learning imitation. Differences between the youngest and oldest children in the jump duration suggested that the reproduction of a complex motor activity such as jumping with a one foot take-off would require resolution and adjustment of main postural stability.     

Time representing cortical activities: two models inspired by prefrontal persistent activity

 Timing information in the range of seconds is significantly correlated with our behavior. There is growing interest in the cognitive behaviors that rely on perception, comparison, or generation of timing. However, little is known about the neural mechanisms underlying such behaviors. Here we model two different neural mechanisms to represent timing information in the range of seconds. In one model, a recurrent network of bistable spiking neurons shows a quasistable state that is initiated by a brief input and typically lasts for a few to several seconds. The duration of this quasistable activity may be regarded as the neural representation of internal time obeying a psychophysical law of time recognition. Another model uses synfire chains to provide the timing information necessary for predicting the times of anticipated events. In this model, the neurons projected to by multiple synfire chains are conditioned to fire synchronously at the times when an external event (GO signal) is expected. The conditioning is accomplished by spike-timing-dependent plasticity. The two models are inspired by the prefrontal activities of the monkeys engaging in different timing-information-related tasks. Thus, this cortical region may provide the timing information required for organizing various behaviors. Received: 12 March 2002 / Accepted in revised form: 26 November 2002 / Published online: 28 March 2003 Correspondence to: T. Fukai (e-mail: tfukai@eng.tamagawa.ac.jp, Tel.: +81-42-7398434, Fax: +81-42-7397135) Acknowledgements. K. Kitano was supported by Japan Society for the Promotion of Science.