Stimulus compounding in interval timing: The modality–duration relationship of the anchor durations results in qualitatively different response patterns to the compound cue.

We have previously demonstrated that rats trained on a two-duration peak procedure in which two modal signals (i.e., tone and houselight) predicted probabilistic reinforcement availability at two times (10 s and 20 s) would respond in a scalar manner at a time between the trained durations in response to the simultaneous compound cue (tone + houselight). In these experiments, we evaluated whether this scalar response pattern would remain with greater relative separation between the anchor durations. Results revealed an effect of the modality–duration relationship, such that scalar responding was seen on compound trials in rats trained that the auditory stimulus signaled the shorter duration, whereas the visual stimulus signaled the longer duration, but not in the reverse condition. In rats showing scalar responding on compound trials, post hoc analyses demonstrated that the peak time of compound responding was most accurately predicted by the reinforcement probability weighted average of anchor peak times. In contrast, rats trained that the visual stimulus signaled the shorter duration, whereas the auditory stimulus signaled the longer duration, responded in a highly rightward skewed manner. In these rats, initiation of responding to the compound stimulus appeared to be controlled by the visual stimulus only, whereas response terminations reflected control by both modal stimuli. These latter data provide evidence of separate determinants of response initiation and termination. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Nucleus accumbens dopamine modulates response rate but not response timing in an interval timing task.

While previous work has demonstrated that systemic dopamine manipulations can modulate temporal perception by altering the speed of internal clock processes, the neural site of this modulation remains unclear. Based on recent research suggesting that changes in incentive salience can alter the perception of time, as well as work showing that nucleus accumbens (NAc) shell dopamine (DA) levels modulate the incentive salience of discriminative stimuli that predict instrumental outcomes, we assessed whether microinjections of DA agents into the NAc shell would impact temporal perception. Rats were trained on either a 10-s or 30-s temporal production procedure and received intra-NAc shell microinfusions of sulpiride, amphetamine, and saline. Results showed that NAc DA modulations had no effect on response timing, but intra-NAc shell sulpiride microinfusions significantly decreased response rates relative to saline and amphetamine. Our findings therefore suggest that neither NAc shell DA levels, nor the resultant changes in incentive salience signaled by this structure, impact temporal control. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

The pattern of responding in the peak-interval procedure with gaps: An individual-trials analysis.

Humans and lower animals time as if using a stopwatch that can be “stopped” or “reset” on command. This view is challenged by data from the peak-interval procedure with gaps: Unexpected retention intervals (gaps) delay the response function in a seemingly continuous fashion, from stop to reset. We evaluated whether these results are an artifact of averaging over trials, or whether subjects use discrete alternatives or a continuum of alternatives in individual-trials: A Probability-of-Reset hypothesis proposes that in individual gap trials subjects stochastically use discrete alternatives (stop/reset), such that when averaged over trials, the response distribution in gap trials falls in between “stop” and “reset.” Alternatively, a Resource Allocation hypothesis proposes that during individual gap trials working memory for the pregap duration decays, such that the response function in individual gap trials is shifted rightward in a continuous fashion. Both hypotheses provided very good fits with the observed individual-trial distributions, although the Resource Allocation hypothesis generated reliably better fits. Results provide support for the usefulness of individual-trial analyses in dissociating theoretical alternatives in interval timing tasks. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of the NMDA receptor antagonist MK-801 on short-interval timing in rats.

Effects of MK-801, an N-methyl-D-aspartate antagonist, on short-interval timing were examined using the peak-interval (PI) and PI-gap procedures. Fisher 344 rats were given daily injections of 0.025 mg/kg, 0.05 mg/kg, and 0.2 mg/kg MK-801. The main results were (a) 0.2 mg/kg MK-801 produced an immediate overestimation of the criterion time; (b) MK-801 increased peak rate of responding; (c) 0.2 mg/kg MK-801 produced an increase in variability; (d) during the PI-gap procedure, a reset pattern was observed for all rats (MK-801 and saline). Results suggest that MK-801 has at least 2 effects. First, MK-801 interferes with short-interval timing by producing an overestimation of time and a nonscalar increase in variability. Second, MK-801 increases response rate, suggesting a decrease in response inhibition. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Magnitude estimation reveals temporal binding at super-second intervals.

Several recent studies (e.g., Haggard, Aschersleben, Gehrke, & Prinz, 2002; Haggard & Clark, 2003; Haggard, Clark, & Kalogeras, 2002) have demonstrated a “Temporal Binding” effect in which the interval between an intentional action and its consequent outcome is subjectively shorter compared to equivalent intervals that do not involve intentional action. The bulk of the literature has relied on the “Libet Clock” (Libet, Gleason, Wright, & Pearl, 1983; but see also Engbert & Wohlschl?ger, 2007; Engbert, Wohlschl?ger, Thomas, & Haggard, 2007; Engbert, Wohlschl?ger, & Haggard, 2008). Here we demonstrate that Temporal Binding is a robust finding that can also be reliably achieved with a Magnitude Estimation procedure, and that occurs over intervals far greater than those previously explored. Implications for the underlying mechanisms are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Intertrial interval as a contextual stimulus: Further analysis of a novel asymmetry in temporal discrimination learning.

Four experiments investigated discrimination learning when the duration of the intertrial interval (ITI) signaled whether or not the next conditional stimulus (CS) would be paired with food pellets. Rats received presentations of a 10-s CS separated half the time by long ITIs and half the time by short ITIs. When the long ITI signaled that the CS would be reinforced and the short interval signaled that it would not be (Long+/Short?), rats learned the discrimination readily. However, when the short ITI signaled that the CS would be reinforced and the long interval signaled that it would not (Short+/Long?), discrimination learning was much slower. Experiment 1 compared Long+/Short? and Short+/Long? discrimination learning with 16-min/4-min or 4-min/1-min ITI combinations. Experiment 2 found no evidence that Short+/Long? learning is inferior because the temporal cue corresponding to the short interval is ambiguous. Experiment 3 found no evidence that Short+/Long? learning is poor because the end of a long ITI signals a substantial reduction in delay to the next reinforcer. Long+/Short? learning may be faster than Short+/Long?because elapsing time involves exposure to a sequence of hypothetical stimulus elements (e.g., A then B), and feature-positive discriminations (AB+/A?) are learned quicker than feature-negative discriminations (A+/AB?). Consistent with this view, Experiment 4 found a robust feature-positive effect when sequentially presented CSs played the role of elements A and B. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Pavlovian contingencies and temporal information.

The effects of altering the contingency between the conditioned stimulus (CS) and the unconditioned stimulus (US) on the acquisition of autoshaped responding was investigated by changing the frequency of unsignaled USs during the intertrial interval. The addition of the unsignaled USs had an effect on acquisition speed comparable with that of massing trials. The effects of these manipulations can be understood in terms of their effect on the amount of information (number of bits) that the average CS conveys to the subject about the timing of the next US. The number of reinforced CSs prior to acquisition is inversely related to the information content of the CS. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Timing During Interruptions in Timing.

Duration and location of breaks in time interval production were manipulated in various conditions of stimulus presentation (Experiments 1-4). Produced intervals shortened and then stabilized as break duration lengthened, suggesting that participants used the break as a preparatory period to restart timing as quickly as possible at the end of the break. This interpretation was supported in Experiment 5, in which similar results were obtained with a reaction time response executed at the end of the break. In all experiments, produced intervals lengthened as the break occurred later during the interval. The authors conclude that varying break location and duration reveal, respectively, the influence of attentional time-sharing before the interruption and of preparatory processes taking place during the break. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Interval timing with gaps: Gap ambiguity as an alternative to temporal decay.

C. V. Buhusi, D. Perera, and W. H. Meck (2005) proposed a hypothesis of timing in rats to account for the results of experiments that have used the peak procedure with gaps. According to this hypothesis, the introduction of a gap causes the animal's memory for the pregap interval to passively decay (subjectively shorten) in direct proportion to the duration and salience of the gap. Thus, animals should pause with short, nonsalient gaps but should reset their clock with longer, salient gaps. The present authors suggest that the ambiguity of the gap (i.e., the similarity between the gap and the intertrial interval in both appearance and relative duration) causes the animal to actively reset the clock and prevents adequate assessments of the fate of timed intervals prior to the gap. Furthermore, when the intertrial interval is discriminable from the gap, the evidence suggests that timed intervals prior to the gap are not lost but are retained in memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Disrupted temporal control in the R6/2 mouse model of Huntington’s disease.

Huntington’s disease is characterized by corticostriatal dysfunction and degeneration of the striatum with progressive loss of the medium spiny neurons. These circuits are important for instrumental responding, interval timing, and temporal control over motor output. We investigated the acquisition of timed operant responding in two R6/2 Huntington’s Disease models, differing in CAG repeat length and genetic background (115 and 250 CAG repeats, and a mixed CBAxC57 or pure C57 background) and their corresponding wild type controls using the peak procedure. Both mouse lines exhibited similar response control deficits. In unreinforced peak trials, mice either did not learn to terminate an ongoing response past reinforcement time or required more trials to acquisition compared to the wild type mice. While transgenic and wild type mice did not exhibit differences in temporal accuracy, response curves were flatter in transgenic mice, suggesting decreased temporal control over operant responding. The results are discussed in terms of the neurobiology of interval timing, instrumental responding, and the neuropathology of HD and R6/2 mice. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Interval timing accuracy and scalar timing in C57BL/6 mice.

In many species, interval timing behavior is accurate—appropriate estimated durations—and scalar—errors vary linearly with estimated durations. Whereas accuracy has been previously examined, scalar timing has not been clearly demonstrated in house mice (Mus musculus), raising concerns about mouse models of human disease. The authors estimated timing accuracy and precision in C57BL/6 mice, the most used background strain for genetic models of human disease, in a peak-interval procedure with multiple intervals. Both when timing 2 intervals (Experiment 1) or 3 intervals (Experiment 2), C57BL/6 mice demonstrated varying degrees of timing accuracy. An important finding was that, both at the individual and group levels, their precision varied linearly with the subjective estimated duration. Further evidence for scalar timing was obtained using an intraclass correlation statistic. This is the first report of consistent, reliable scalar timing in a sizable sample of house mice, thus validating the peak-interval procedure as a valuable technique, the intraclass correlation statistic as a powerful test of the scalar property, and the C57BL/6 strain as a suitable background for behavioral investigations of genetically engineered mice modeling disorders of interval timing. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Interval timing with gaps and distracters: Evaluation of the ambiguity, switch, and time-sharing hypotheses.

Gaps and distracters were presented during the timed signal to examine whether the stop/reset mechanism is activated by (a) changes in the timed signal (switch hypothesis), (b) ITI-like events (ambiguity hypothesis), or (c) processes concurrent with the timing process (time-sharing hypothesis). While the switch and ambiguity hypotheses predict that rats should time through (ignore) distracters, the time-sharing hypothesis predicts that extraneous events (e.g., gaps and distracters) delay timing by causing working memory to decay in proportion to the events' salience. The authors found that response functions were displaced by both gaps and distracters, in accord with the time-sharing hypothesis. Computer simulations show that the time-sharing and memory-decay hypotheses can mechanistically address present data, and reflect different levels of the same model. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of methamphetanine on duration discrimination.

Experiments 1 and 2 address the controversy regarding the reliability of methamphetamine effects on interval timing. A temporal discrimination procedure was used, in which the rats were reinforced for pressing the left or the right levers after short and long signals, respectively. Methamphetamine (0.5 mg/kg sc) severely disrupted operant performance at 20-100 min after injection, which disabled the measurement of drug effects on temporal perception (Experiment 1). The same dose of methamphetamine shifted the psychometric function to the left at 100-180 min after injection, indicating an increase in subjective durations (Experiment 2). Although these results confirm the role of dopamine in interval timing, that a change in the speed of a neural clock mediates the methamphetamine-induced change in temporal perception is still a working hypothesis. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Time and stimulus specificity in extinction of the conditioned nictitating membrane response in the rabbit (Oryctolagus cuniculus).

The present experiments demonstrated that, in the rabbit nictitating membrane preparation, a conditioned response (CR) can be selectively eliminated in one portion of a conditioned stimulus (CS) while it is still paired with the unconditioned stimulus (US). Rabbits were initially trained with two stimuli (tone, light). Each was paired with the US by using a mixture of two CS-US interstimulus intervals (ISIs): 200 ms and 1,200 ms in Experiment 1; 150 ms and 500 ms in Experiment 2. The CRs showed double peaks, one for each ISI. Subsequently, one CS (A) was trained with only the longer ISI, whereas the other CS (B) continued to be trained with both ISIs. Consequently, the CR peak based on the shorter ISI disappeared for CSA but not for CSB. The later CR peaks during both CSA and CSB were maintained. These results support time-based models of conditioning. Implications for proposed mechanisms of extinction are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Perception of anticipatory time in temporal discounting.

The current article focuses on the role of anticipatory time perception in temporal discounting. We propose a perceived-time–based model and demonstrate that 2 aspects of time perception are relevant to hyperbolic discounting. Specifically, our model states that diminishing sensitivity to longer time horizons (i.e., how long individuals perceive short time horizons to be relative to long time horizons) and the level of time contraction overall (i.e., how long or short individuals perceive time horizons to be overall) contribute to the degree of hyperbolic discounting. We estimate individual differences in the degree of diminishing sensitivity to time and the degree of time contraction, and demonstrate that each significantly predicts the degree of hyperbolic discounting. These results empirically confirm two unique aspects of anticipatory time perception in determining individuals’ temporal discounting. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Recovery of fear memories in rats: Role of gamma-amino butyric acid (GABA) in infantile amnesia.

Infantile amnesia is a ubiquitous phenomenon, but its neural bases remain largely unknown. The authors identify a role for GABAergic transmission in suppressing retrieval of memories acquired in infancy. Eighteen-day-old rats received pairings of white noise and shock; considerable forgetting of this experience (assessed by freezing) occurred after 10 days. The memory was recovered by pretest administration of the GABAA inverse agonist FG7142 10 days, but not 2 months, after training. This effect of FG7142 generalized when a passive avoidance procedure was used. Also, FG7142 decreased fear of a latently inhibited conditioned stimulus, showing that the observed memory recovery effect was not due to a state-dependent process. It appears that GABA may be involved in infantile amnesia regardless of the emotional content of the memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Blinded by emotion? Effect of the emotionality of a scene on susceptibility to false memories.

Although distortion is commonly present in memory, the relation between the emotionality of a witnessed scene and susceptibility to mistaken memories is controversial. Participants (N = 90; aged 17-43 years) were recruited for research focusing on "emotional processing" and were not informed that their memories were being investigated. Then, they viewed either a highly positive, neutral, or highly negative emotional scene (e.g., graphic fatal accident) from the International Affective Picture System (e.g., Lang, Bradley, & Cuthbert, 1999). Half of participants were exposed to misleading questions--one of which included a major false suggestion (i.e., large animal in the scene). An hour later all participants were asked to recall the scene and asked 10 direct questions, five of which related to the misinformation provided earlier. Overall, misleading questions impaired recall accuracy by 37%. Further, negative emotion increased susceptibility to false memories for the major misinformation. Whereas 0% of nonmisled participants in any condition recalled seeing the major false detail, misled participants in the negative condition recalled seeing the major false detail more often (80%) than those in the positive (40%) and neutral (40%) conditions. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal ventriloquism in a purely temporal context.

This study examines how audiovisual signals are combined in time for a temporal analogue of the ventriloquist effect in a purely temporal context, that is, no spatial grounding of signals or other spatial facilitation. Observers were presented with two successive intervals, each defined by a 1250-ms tone, and indicated in which interval a brief audiovisual stimulus (visual flash + noise burst) occurred later. In “test“ intervals, the audiovisual stimulus was presented with a small asynchrony, while in “probe” intervals it was synchronous and presented at various times guided by an adaptive staircase to find the perceived temporal location of the asynchronous stimulus. As in spatial ventriloquism, and consistent with maximum likelihood estimation (MLE), the asynchronous audiovisual signal was shifted toward the more reliably localized component (audition, for all observers). Moreover, these temporal shifts could be forward or backward in time, depending on the asynchrony order, suggesting perceived timing is not entirely determined by physical timing. However, the critical signature of MLE combination—better bimodal than unimodal precision—was not found. Regardless of the underlying model, these results demonstrate temporal ventriloquism in a paradigm that is defined in a purely temporal context. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Examination of bilateral eyeblink conditioning in rats.

This experiment monitored eyelid responses bilaterally during delay eyeblink conditioning in rats. Rats were given paired or unpaired training with a tone or light conditioned stimulus (CS) and a unilateral periorbital shock unconditioned stimulus (US). Rats given paired training acquired high levels of conditioned responses (CRs), which occurred in both eyelids. However, acquisition was faster, and the overall percentage of CRs was greater in the eyelid that was ipsilateral to the US. CRs in the eyelid ipsilateral to the US also had shorter onset latencies and larger amplitudes than CRs in the contralateral eyelid. Both eyelids consistently showed high percentages of unconditioned responses (UR) to the US, and the UR amplitude decreased across training sessions in the paired group. The present study demonstrated that CRs occur robustly in both eyelids of rats given eyeblink conditioning, which is similar to previous findings in humans and monkeys. The results also showed that conditioning occurs more prominently in the eyelid that is ipsilateral to the US, which is similar to previous findings in humans, monkeys, dogs, and rabbits. (PsycINFO Database Record (c) 2010 APA, all rights reserved)