Reward magnitude and timing in pigeons

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

Within-session modulation of timed anticipatory responding: When to start responding

The peak procedure is widely used in the study of interval timing with animals. Multiple timing measures can be derived from peak responding. These measures are typically presented as averages across many trials based on the implicit assumption that peak responding is stable throughout the session. We tested this assumption by examining whether peak responding changed over the course of the session in 45 mice that were trained on a fixed-interval 30-s schedule. All common measures of peak responding, except stop times, changed over the course of the session: start times increased, response rates and spreads decreased, and, although less reliably, peak times also shifted rightward. These results are congruent with a motivational interpretation, whereby increased satiety leads to the observed behavioral signature of within-session modulation of timed anticipatory responding.     

Acquisition of peak responding: What is learned?

We investigated how the common measures of timing performance behaved in the course of training on the peak procedure in C3H mice. Following fixed interval (FI) pre-training, mice received 16 days of training in the peak procedure. The peak time and spread were derived from the average response rates while the start and stop times and their relative variability were derived from a single-trial analysis. Temporal precision (response spread) appeared to improve in the course of training. This apparent improvement in precision was, however, an averaging artifact; it was mediated by the staggered appearance of timed stops, rather than by the delayed occurrence of start times. Trial-by-trial analysis of the stop times for individual subjects revealed that stops appeared abruptly after three to five sessions and their timing did not change as training was prolonged. Start times and the precision of start and stop times were generally stable throughout training. Our results show that subjects do not gradually learn to time their start or stop of responding. Instead, they learn the duration of the FI, with robust temporal control over the start of the response; the control over the stop of response appears abruptly later.     

Choice and timing in concurrent chains: effects of initial-link duration

Theories of timing have been applied to choice between delayed rewards by assuming that delays are represented in memory and that subjects sample from memory when choosing between alternatives. To search for covariation in single-trial measures of performance that might confirm this assumption, we used a procedure that allowed for convergent measurement of choice and timing behavior. Four pigeons responded in a concurrent chains/peak procedure in which the terminal links were fixed-interval (FI) 8s and FI 16s, across conditions the duration of the initial-link schedule was either short or long, and one quarter of the terminal links lasted for 48 s and ended without reinforcer delivery. Preference for the FI 8-s alternative was stronger with shorter initial links, replicating the 'initial-link effect'. Responding on no-food trials was unaffected by initial-link duration, and aggregated across trials, was typical of the peak procedure: response distributions were approximately Gaussian, with modes near the FI schedule durations, and variance was greater for the FI 16-s terminal link. Analysis of local measures of initial-link performance (e.g., pause to begin responding, time spent responding, number and duration of visits to each alternative, etc.) found that the initial-link effect was associated with an increase in the number and duration of visits per cycle to the nonpreferred alternative. Regression analyses showed that local initial-link measures contributed relatively little additional variance in predicting performance on individual no-food trials beyond that accounted for by FI schedule. Our results provide no clear evidence that initial- and terminal-link responding in concurrent chains are mediated by a common representation of terminal-link delays.     

The effects of concurrent task and gap events on peak time in the peak procedure

The effect of a concurrent task on timing performance of pigeons was investigated with the peak interval procedure. Birds were trained to peck a side key on a discrete-trial schedule that included reinforced fixed-interval (FI) 30-s trials and nonreinforced extended probe trials. Then, in separate sessions, birds were trained to peck a 6-s center key for food. In a subsequent test phase, the FI procedure was in effect along with dual-task probe test trials. On those test trials, the 6-s center key (task cue) was presented at 3, 9, or 15s after probe trial onset. During another test phase, a 6-s gap (the FI keylight was extinguished) was presented at 3, 9, or 15s after probe trial onset. Peak time increased with center key time of onset, and was greater under task than gap conditions. Moreover, peak time under task conditions exceeded values predicted by stop and reset clock mechanisms. These results are at variance with current attentional accounts of timing behavior in dual-task conditions, and suggest a role of nontemporal factors in the control of timing behavior.     

Dopamine receptor antagonists reverse amphetamine-induced behavioral alteration on a differential reinforcement for low-rate (DRL) operant task in the rat

Previous studies have shown that amphetamine significantly alters operant responding on the behavior maintained on a schedule of differential reinforcement of low-rate (DRL). As such, behavioral deficiency of DRL responding has been observed by the drug-induced increase of non-reinforced responses and a leftward shift of inter-response time (IRT) curve on DRL responding in the rat. However, the neurochemical basis for amphetamine-induced DRL behavioral alternations remain to be elucidated. The present study was then designed to examine whether the effects of amphetamine were dependent on dopamine-subtyped receptors, this was carried out by the co-administration of the selective D1 and D2 receptor antagonists, SCH23390 and raclopride respectively. Rats were first trained to perform on DRL 10-sec task and then divided into four groups, which received separate types of double injections before the behavioral session. The four groups were the saline control group, the amphetamine alone group, the dopamine antagonist alone group, and the combination of [corrected] amphetamine and dopamine antagonist group. The saline control group performed DRL responding in an efficient manner with a major index for the peak time of the IRT curve, which was fairly localized within the 10-sec bin throughout the test phase. The subjects injected with amphetamine (1 mg/kg) significantly shortened IRT that led to a leftward shift of IRT curve, which was further revealed by a decreased peak time without significant effectiveness on the peak rate and burst response. Even though the group given SCH23390 or raclopride alone showed profound disruption on DRL behavior by flattening the IRT curve, the co-administration of amphetamine with SCH23390 or raclopride reversed the aforementioned amphetamine-induced behavioral deficiency on DRL task. Together, these results suggest that the dopamine D1 and D2 receptors are involved and important to the temporal regulation of DRL response under psychostimulant drug treatment. Furthermore, this highlights the involvement of the brain dopamine systems in the temporal regulation of DRL behavior performance.     

The effect of intruded events on peak time: the role of reinforcement history during the intruded event

Pigeons were studied in an extension of a study by Aum et al. [Aum, S., Brown, B.L., Hemmes, N.S. 2004. The effects of concurrent task and gap events on peak time in the peak procedure. Behav. Process. 65, 43-56] on timing behavior under a discrete-trial fixed-interval (FI) procedure during which 6-s intruded events were superimposed on peak-interval (PI) test trials. In Aum et al., one event consisted in termination of the timing cue (gap trial); the other was a stimulus in the presence of which subjects had been trained to respond under an independent random-interval (RI) schedule of reinforcement (concurrent task trial). Aum et al. found a disruption of timing on concurrent task trials that was greater than that on gap trials. The present study investigated history of reinforcement associated with intruded events as a possible explanation of this earlier finding. After training to peck a side key on a 30-s PI procedure, discrimination training was conducted on the center key in separate sessions; red or green 6-s stimuli were associated with RI 24s or EXT (extinction) schedules. During testing under the PI procedure, three types of intruded events were presented during probe trials--the stimulus associated with the RI (S+) or EXT (S-) schedule during discrimination training, or a gap (termination of the side-keylight). Intruded events occurred 3, 9, or 15s after PI trial onset. Effects of reinforcement history were revealed as substantial disruption of timing during the S+ event and relatively little disruption during the S- event. Intermediate effects were found for the gap event. Results indicate that postcue effects are at least partially responsible for the disruptive effects of the S+ event.     

Stepping back to improve sprint performance: a kinetic analysis of the first step forwards

Using a step backward to initiate forward movement can increase force and power at push-off and improve sprint performance over short distances. However, it is not clear whether the benefit provided by this paradoxical step influences the mechanics of the first step forwards. Twenty-seven men of an athletic background performed maximal effort 5-m sprints from a standing start and employed a step forwards (parallel and split stance) or backwards (false) to initiate movement. Each sprint was started with an audio cue that also activated the timing gates. Three trials of each starting style were performed and movement (0 m), 2.5-, and 5-m times were recorded. An in-ground force plate placed at the 0-m mark measured the kinetic and temporal characteristics of the first step. Sprint times to 2.5 and 5 m were slower (p < 0.05) when a parallel start was used. No differences were seen in the normalized peak forces (vertical and horizontal) or the vertical impulse between starts, but the vertical mean force was 11 and 12% higher for the false and split starts, respectively. Surprisingly, the parallel start's impulse was significantly greater than that of the false (24%) and split (22%) styles, a consequence of the additional time spent in contact with the ground. The ground contact time, time to peak force, and time from peak force to toe-off (vertical and horizontal) were significantly longer for the parallel start. These temporal variables were also better correlated with sprint performance than any kinetic measure (0.42 ≤ r ≤ 0.75). The false start appears to be advantageous over short distances by improving push-off and the temporal characteristics of the first step.     

Interval timing by an invertebrate, the bumble bee Bombus impatiens

Sensitivity to temporal information and the ability to adjust behavior to the temporal structure of the environment should be phylogenetically widespread. Some timing abilities, such as sensitivity to circadian cycles, appear in a wide range of invertebrate and vertebrate taxa [1,2]. Interval timing--sensitivity to the duration of time intervals--has, however, only been shown to occur in vertebrates [3,4]. Insect pollinators make a variety of decisions that would appear to require the ability to estimate elapsed durations. We exposed bumble bees to conditions in which proboscis extension was reinforced after a fixed duration had elapsed or after either of two fixed durations had elapsed. Two groups of bees were trained with a short duration (either 6 s or 12 s) and a long duration (36 s) in separate experimental phases (independent timing groups), whereas two other groups were trained with a short duration (either 6 s or 12 s) and long duration (36 s) always intermixed unpredictably (multiple timing groups). On long intervals, independent timing groups waited longer than mixed timing groups to generate the first response and responded maximally near the end of the interval. Multiple timing groups waited the same amount of time on average before generating the first response on both long and short intervals. On individual trials, multiple timing groups appeared to time either the long duration only or both the short and long durations: most trials were characterized by a single burst of responding that began between the short and long duration values or by two bursts of responding with the first burst bracketing the short value and the second burst beginning in anticipation of the long value. These results show that bumble bees learn to time interval durations and can flexibly time multiple durations simultaneously. The results indicate no phylogenetic divide between vertebrates and invertebrates in interval timing ability.     

Time estimation by pigeons on a fixed interval: the effect of pre-feeding

Pigeons well trained on a fixed interval 10-s schedule of reinforcement were tested on the peak procedure. In a successive conditions design, they were either pre-fed or not in the experimental setting. Pre-feeding decreased the rate of responding. It also led to a maximum rate of responding that occurred 2-3 s later than in the control condition, where the maximum occurred at the usual time of reinforcement. The shift in peak time in response to pre-feeding shows that peak time may not be a pure measure of timing. The results are also interpreted in terms of timing theories.     

The proportion of fixed interval trials to probe trials affects acquisition of the peak procedure fixed interval timing task

A common procedure for studying the ability of animals to time is the peak procedure. With the peak procedure, animals are first trained on a fixed interval schedule (i.e., 30s). After the animals have been well trained on the fixed interval schedule, probe trials are introduced. On probe trials, the stimulus is presented longer (i.e., 90s) and the animal does not receive reinforcement for responding. When animals are first presented with probe trials responding remains flat following the point that reinforcement normally occurs on fixed interval trials. The descending slope that eventually emerges is acquired with experience with probe trials. The present experiments manipulated the percentage of probe trials compared to FI trials across groups of rats. It was hypothesized that the descending limb of peak responding would be acquired more quickly when there were many probe trials per session as this might facilitate extinction of responding beyond the interval that reinforcement normally occurs. It was found, however, that acquisition of peak responding occurred best when there were few probe trials per session.     

The effect of an intruded event on peak-interval timing in rats: isolation of a postcue effect

The present experiment employed the peak-interval (PI) procedure to study the effect of an intruded cue on timing behavior. Rats were trained on a 30-s PI procedure with a tone cue. Subsequently, a 6-s flashing light was paired off-baseline with foot shock (Experiment 1) or presented alone (Experiment 2). Then, in test trials, the light cue was presented 9s prior to (before) or 3s after (during) the onset of the timing cue, or the light was omitted (probe). Results showed rightward shifts in peak time occurring on both before and during trials in both experiments. Peak shifts on during trials exceeded the reset prediction in Experiment 1. When PI functions for before and probe trials were normalized in peak rate and peak time, they superimposed better than when functions were adjusted additively along the time axis, suggesting that the light cue may engender a decrease in functional clock rate. The findings suggested that the intruded cue produced both intracue and postcue interference with timing that was enhanced by fear conditioning.     

Optimal timing of one-shot interventions for epidemic control

The interventions and outcomes in the ongoing COVID-19 pandemic are highly varied. The disease and the interventions both impose costs and harm on society. Some interventions with particularly high costs may only be implemented briefly. The design of optimal policy requires consideration of many intervention scenarios. In this paper we investigate the optimal timing of interventions that are not sustainable for a long period. Specifically, we look at at the impact of a single short-term non-repeated intervention (a “one-shot intervention”) on an epidemic and consider the impact of the intervention’s timing. To minimize the total number infected, the intervention should start close to the peak so that there is minimal rebound once the intervention is stopped. To minimise the peak prevalence, it should start earlier, leading to initial reduction and then having a rebound to the same prevalence as the pre-intervention peak rather than one very large peak. To delay infections as much as possible (as might be appropriate if we expect improved interventions or treatments to be developed), earlier interventions have clear benefit. In populations with distinct subgroups, synchronized interventions are less effective than targeting the interventions in each subcommunity separately.     

Fewer peak trials per session facilitate acquisition of peak responding despite elimination of response rate differences

It has been shown in previous research [Kaiser, D.H., 2008. The proportion of fixed interval trials to probe trials affects acquisition of the peak procedure fixed interval timing task. Behav. Process., 77 (1), 100-108] that rats acquired peak responding sooner when fewer peak trials were presented during sessions of training with the peak procedure timing task. One potential problem with that research was that there were large differences in response rates among the groups. The present experiment attempted to examine the effect of proportion of peak trials when differences in response rate were controlled. Two groups of rats were each simultaneously tested with two versions of the peak procedure. One group was tested with 10% peak trials per session, and the other group was tested with 50% peak trials per session. For both of the groups, one of the panel lights and levers was associated with the traditional peak procedure. The other panel light and lever was associated with a similar peak procedure; however, reinforcement was provided at the end of each peak trial. This manipulation eliminated differences in response rate among the groups, however, Group 10% acquired peak responding more quickly than Group 50%, effectively replicating previous work in the absence of a response bias.     

Gaussian modeling of the P-wave morphology time course applied to anticipate paroxysmal atrial fibrillation

This paper introduces a new algorithm to quantify the P-wave morphology time course with the aim of anticipating as much as possible the onset of paroxysmal atrial fibrillation (PAF). The method is based on modeling each P-wave with a single Gaussian function and analyzing the extracted parameters variability over time. The selected Gaussian approaches are associated with the amplitude, peak timing, and width of the P-wave. In order to validate the algorithm, electrocardiogram segments 2 h preceding the onset of PAF episodes from 46 different patients were assessed. According to the expected intermittently disturbed atrial conduction before the onset of PAF, all the analyzed Gaussian metrics showed an increasing variability trend as the PAF onset approximated. Moreover, the Gaussian P-wave width reported a diagnostic accuracy around 80% to discern between healthy subjects, patients far from PAF, and patients less than 1 h close to a PAF episode. This discriminant power was similar to those provided by the most classical time-domain approach, i.e., the P-wave duration. However, this newly proposed parameter presents the advantage of being less sensitive to a precise delineation of the P-wave boundaries. Furthermore, the linear combination of both metrics improved the diagnostic accuracy up to 86.69%. In conclusion, morphological P-wave characterization provides additional information to the metrics based on P-wave timing.     

Diel and lunar spawning periodicity of the hawkfish Paracirrhites forsteri (Cirrhitidae) on the reefs of Kuchierabu-jima Island, southern Japan

We observed spawning behavior of the hawkfish Paracirrhites forsteri on reefs of southern Japan. Spawning generally occurred after sunset, prior to the full and new moon with semi-lunar spawning peak periodicity. No egg predation was observed in spawning after sunset, and high tide often occurred at dusk in the lunar phases. Thus, diel timing and lunar synchronicity may increase larval survival. We found that the mating activity tended to start earlier in the day during early mating season than during mid-to-late mating season. The advantages of the earlier start of mating activity were also examined in relation to adult biology contexts.     

Evaluating timing in spontaneously hypertensive and Wistar-Kyoto rats using the peak procedure

A core deficit in timing may underlie the symptoms of attention-deficit/hyperactivity disorder (ADHD). Timing deficits have been observed in ADHD-diagnosed children but have yet to be fully explored in the spontaneously hypertensive rat (SHR), a purported model of ADHD. We asked whether SHRs demonstrate ADHD-like timing deficits using the peak procedure. Response rates across peak intervals were modeled using the sum of two Gaussian curves. Results showed that SHRs peaked earlier than Wistar-Kyotos based on 4 s intervals that contained the individuals’ maximum response rates but not based on model parameters. The strains showed approximately equal precision of timing based on Weber fractions derived from model parameters, a result that replicates previous findings and does not support the use of SHRs to model this aspect of ADHD.     

Exposure to stallion accelerates the onset of mares' cyclicity

Horses (Equus caballus) belong to the group of seasonally polyestrous mammals. Estrous cycles typically start with increasing daylight length after winter, but mares can differ greatly in the timing of onset of regular estrus cycles. Here, we test whether spatial proximity to a stallion also plays a role. Twenty-two anestrous mares were either exposed to one of two stallions (without direct physical contact) or not exposed (controls) under experimental conditions during two consecutive springs (February to April). Ovarian activity was monitored via transrectal ultrasound and stallion's direct contact time with each mare was determined three times per week for one hour each. We found that mares exposed to a stallion ovulated earlier and more often during the observational period than mares that were not exposed to stallions. Neither stallion identity nor direct contact time, mare age, body condition, size of her largest follicle at the onset of the experiment, or parasite burden significantly affected the onset of cyclicity. In conclusion, the timing of estrous cycles and cycle frequency, i.e., crucial aspects of female reproductive strategy, strongly depend on how the mares perceive their social environment. Exposing mares to the proximity of a stallion can therefore be an alternative to, for example, light programs or elaborated hormonal therapies to start the breeding season earlier and increase the number of estrous cycles in horses.     

Investigations of timing during the schedule and reinforcement intervals with wheel-running reinforcement

Across two experiments, a peak procedure was used to assess the timing of the onset and offset of an opportunity to run as a reinforcer. The first experiment investigated the effect of reinforcer duration on temporal discrimination of the onset of the reinforcement interval. Three male Wistar rats were exposed to fixed-interval (FI) 30-s schedules of wheel-running reinforcement and the duration of the opportunity to run was varied across values of 15, 30, and 60s. Each session consisted of 50 reinforcers and 10 probe trials. Results showed that as reinforcer duration increased, the percentage of postreinforcement pauses longer than the 30-s schedule interval increased. On probe trials, peak response rates occurred near the time of reinforcer delivery and peak times varied with reinforcer duration. In a second experiment, seven female Long-Evans rats were exposed to FI 30-s schedules leading to 30-s opportunities to run. Timing of the onset and offset of the reinforcement period was assessed by probe trials during the schedule interval and during the reinforcement interval in separate conditions. The results provided evidence of timing of the onset, but not the offset of the wheel-running reinforcement period. Further research is required to assess if timing occurs during a wheel-running reinforcement period.     

Impulsive responding on the peak-interval procedure

The pattern of responding on a peak-interval timing task allows one to make inferences regarding the sources of variation that contribute to interval timing behavior. Non-temporal factors such as impulsivity may impact the validity of these inferences. Rats were trained on a 15s peak-interval procedure (PI) or a mixed 15s behaviorally dependent variable-interval, 15s peak-interval procedure (bdVIPI) for an extended number of sessions. Extended training on the PI revealed a bi-modal distribution in the times at which subjects started responding for temporally predictable reinforcement, suggesting that multiple processes contribute to the behavioral pattern obtained in this procedure. Training on the bdVIPI eliminated the early mode of this bi-modal distribution, thereby decreasing the variation in start times. These results suggest that alternative response options can modulate the influence of impulsivity in timing tasks.