The role of temporal generalization in a temporal discrimination task

Two experiments trained rats to discriminate two or three stimulus durations using a temporal discrimination task. A standard peak shift effect was observed when training was administered with short versus long signals in Experiment 1. Both discrimination accuracy scores and the generalization gradients revealed that shorter intervals were discriminated more accurately, which may be due to the scalar property of timing. In Experiment 2, three signals (short, medium, and long) were associated with three different responses, or two of the intervals were associated with one response (short and long or short and medium) and the other interval with a different response. Here, the short/medium versus long discrimination was learned most readily of the three tasks. The results of both experiments indicated a strong contribution of learning of individual durations combined with scalar generalization gradients, but Experiment 2 indicated that categorical encoding of durations may have also been operating.     

Prospective timing in pigeons: Isolating temporal perception in the time-left procedure

In the time-left procedure, a task used to study prospective timing, animals choose between two stimuli that signal different delays to reinforcement. Trials begin with one stimulus signaling C seconds to reinforcement and, at different moments since its onset, another stimulus, signaling S seconds to reinforcement, with C > S, is introduced. Optimal performance consists in choosing the stimulus signaling the shorter time to reinforcement. Animals have been found to perform in this optimal way. However, this procedure is complex and variables other than time may be responsible for the results. In two experiments with pigeons we sought to improve the time-left procedure to better isolate the effect of time in the animals’ behavior. We attempted to control for two confounding variables, the asymmetry in the time markers from training to testing and the cost of switching between the two response alternatives. We conclude that in the time-left task pigeons seem indeed to regulate their behavior based on time because, with our improved procedure, they still chose the stimulus associated with the shorter time to food. However, our version of the procedure created new interpretative difficulties, strengthening the idea that the time-left procedure may be too complex to study timing.     

Feature-short and feature-long discrimination learning in the pigeon: conditional control of a two-event temporal map

Eight pigeons were trained to peck an illuminated target key on discrete-trial fixed-interval schedules of reinforcement by food. Four birds were exposed to a feature-short (FS) task where a feature light signaled shortening of the forthcoming target-outcome interval from 30 to 15s, while the other four birds were exposed to a feature-long (FL) task where a feature light signaled extension of the forthcoming target-outcome interval from 15 to 30s. The discrimination performance measured by differential temporal distributions of pecks between featured and non-featured target trials suggested that the target-food temporal map was under conditional control of the feature light in both groups. The FS discrimination was more difficult to learn than the FL discrimination. This FS inferiority implies that our birds did not resort on the simple temporal discrimination by timing from the trial onset. The simple temporal discrimination account was also negated by the finding that increasing the feature-target gap did not have a predicted effect on the response distribution.     

Disruptive effects of prefeeding and haloperidol administration on multiple measures of food-maintained behavior in rats

Four rats responded under a choice reaction-time procedure. At the beginning of each trial, the rats were required to hold down a center lever for a variable duration, release it following a high- or low-pitched tone, and press either a left or right lever, conditionally on the tone. Correct choices were reinforced with a probability of .95 or .05 under blinking or static houselights, respectively. After performance stabilized, disruptive effects of free access to food pellets prior to sessions (prefeeding) and intraperitoneal injection of haloperidol were examined on multiple behavioral measures (i.e., the number of trials completed, percent of correct responses, and reaction time). Resistance to prefeeding depended on the probability of food delivery for the number of trials completed and reaction time. Resistance to haloperidol, on the other hand, was not systematically affected by the probability of food delivery for all dependent measures.     

Reaction times identify a Pavlovian component in a two-choice discrimination

Six pigeons discriminated on discrete trials between two colors. In Experiment 1, two luminous spots were both either blue or green and the reinforced responses were “peck left” for blue and “peck right” for green. In Experiment 2, the hue of a center spot controlled subsequent choice pecks to left or right. In both experiments response bias was manipulated in two ways. During stimulus frequency (“SF”) sessions correct responses brought food on 40% of trials; in “imbalanced” blocks of sessions one hue appeared on 80% of trials and the other on 20%. During reinforcement probability (“RNF”) sessions the hues appeared equally often, but in imbalanced blocks the hues signaled different reinforcement probabilities, either 64% or 16%. In “balanced” control blocks the hues appeared equally often and were both reinforced at 40%. The experiments gave similar results. When bias was computed from choice percentages the imbalanced conditions yielded substantial response bias, and the amount of bias was about the same under RNF and SF treatments. However, reaction times (RTs) gave a different outcome. RNF imbalance slowed responses directed at the less reinforced stimulus, but SF imbalance had little RT effect (Experiment 1) or no effect (Experiment 2). These results suggest that choice was controlled by an instrumental stimulus-response-reinforcement association, whereas RTs were controlled by a Pavlovian stimulus-reinforcement association.     

Transference effects of prior non-contingent reinforcement on the acquisition of temporal control on fixed-interval schedules

In two experiments we examined the influence of response and time factors on the speed of acquisition of temporal control on FI schedules. In Experiment 1, prior exposure to FT accelerated the development of temporal control on FI schedules of the same temporal value. It was also found that the slower acquisition on FI with prior RT was similar to that of rats with prior standard training. In Experiment 2, prior exposure to FT accelerated the development of temporal control on a FI schedule with a threefold increase in temporal value. Additionally, it was found that with prior FI 30s training, acquisition of temporal control on FI 90s was even faster than with prior FT 30s. Measures of head-entries into the feeder along the experiments indicated that temporal control was already developed during the periodic but not during the non-periodic histories and that this control transferred to lever press during FI testing phase.     

Serial conditional discrimination and temporal bisection in rats selectively lesioned in the dentate gyrus

In positive serial conditional discrimination, animals respond during a target stimulus when it is preceded by a feature stimulus, but they do not respond when the same target stimulus is presented alone. Moreover, the feature and target stimuli are separated from each other by an empty interval. The present work aimed to investigate if two durations (4 or 16 s) of the same feature stimulus (light) could modulate the operant responses of rats to different levers (A and B) during a 5-s target stimulus (tone). In the present study, lever A was associated with the 4-s light, and lever B was associated with the 16-s light. A 5-s empty interval was included between the light and the tone. In the same training procedure, the rats were also presented with the 5-s tone without the preceding light stimuli. In these trials, the responses were not reinforced. We evaluated the hippocampal involvement of these behavioral processes by selectively lesioning the dentate gyrus with colchicine. Once trained, the rats were submitted to a test using probe trials without reinforcement. They were presented with intermediate durations of the feature stimulus (light) to obtain a temporal bisection curve recorded during the exposure to the target stimuli. The rats from both groups learned to respond with high rates during tones preceded by light and with low rates during tones presented alone, which indicated acquisition of the serial conditional discrimination. The rats were able to discriminate between the 4- and 16-s lights by correctly choosing lever A or B. In the test, the temporal bisection curves from both experimental groups showed a bisection point at the arithmetic mean between 4 and 16 s. Such processes were not impaired by the dentate gyrus lesion. Thus, our results showed that different durations of a feature stimulus could result in conditional properties. However, this processing did not appear to depend on the dentate gyrus alone.     

Contingency discriminability and the generalized matching law describe choice on concurrent ratio schedules of wheel-running reinforcement

Belke (2010) showed that on concurrent ratio schedules, the difference in ratio requirements required to produce near exclusive preference for the lower ratio alternative was substantively greater when the reinforcer was wheel running than when it was sucrose. The current study replicated this finding and showed that this choice behavior can be described by the matching law and the contingency discriminability model. Eight female Long Evans rats were exposed to concurrent VR schedules of wheel-running reinforcement (30s) and the schedule value of the initially preferred alternative was systematically increased. Two rats rapidly developed exclusive preference for the lower ratio alternative, but the majority did not - even when ratios differed by 20:1. Analysis showed that estimates of slopes from the matching law and the proportion of reinforcers misattributed from the contingency discriminability model were related to the ratios at which near exclusive preference developed. The fit of these models would be consistent with misattribution of reinforcers or poor discrimination between alternatives due to the long duration of wheel running.     

Visual and auditory conditional position discrimination in chimpanzees (Pan troglodytes)

Chimpanzee cognition has been studied predominantly through the visual modality, and much less through the auditory modality. The aim of this study was to explore possible differences in chimpanzees’ processing of visual and auditory stimuli. We developed a new conditional position discrimination (CPD) task requiring the association between a stimulus (from either the auditory or the visual modality), and a spatial position (left or right). The stimuli consisted of the face and voice of two individuals well known to the subjects (one chimpanzee and one human). Six chimpanzees participated in both the visual and the auditory conditions. We found contrasting results between the two conditions: the subjects acquired the CPD more easily in the visual than in the auditory condition. This supports previous findings on the difficulties encountered by chimpanzees in learning tasks involving auditory stimuli. Our experiments also revealed individual differences: the chimpanzee with the most extensive experience in symbolic visual matching tasks showed good performance in both conditions. In contrast, the chimpanzee expert in an auditory-visual intermodal matching task showed no sign of learning in either condition. Future work should focus on finding the most appropriate procedure for exploring chimpanzees’ auditory-visual cognitive skills.     

Context effect in a temporal bisection task with the choice keys available during the sample

In a symbolic matching to sample task, six pigeons learned to associate different sample durations with different comparison stimuli. On “Short” trials, choice of Red and Green keylight comparisons were reinforced following 3-s and 9-s samples, respectively; on “Long” trials, Blue and Yellow keylight comparisons were reinforced following 9-s and 27-s samples, respectively. In contrast with previous studies, the comparison keys were available during the samples. After the temporal discriminations were learned, new pairs of comparison keys were presented and the preference for each was assessed during 27-s samples. One pair in particular, Green and Blue, was critical because it tested the predictions of two timing models, Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model. The results showed that preference for Green increased during the sample, a result consistent with LeT but not with SET. Other test results, however, were predicted by neither model.     

Comparison of Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model in a successive temporal bisection task

The present research tested the generality of the "context effect" previously reported in experiments using temporal double bisection tasks [e.g., Arantes, J., Machado, A. Context effects in a temporal discrimination task: Further tests of the Scalar Expectancy Theory and Learning-to-Time models. J. Exp. Anal. Behav., in press]. Pigeons learned two temporal discriminations in which all the stimuli appear successively: 1s (red) vs. 4s (green) and 4s (blue) vs. 16s (yellow). Then, two tests were conducted to compare predictions of two timing models, Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model. In one test, two psychometric functions were obtained by presenting pigeons with intermediate signal durations (1-4s and 4-16s). Results were mixed. In the critical test, pigeons were exposed to signals ranging from 1 to 16s and followed by the green or the blue key. Whereas SET predicted that the relative response rate to each of these keys should be independent of the signal duration, LeT predicted that the relative response rate to the green key (compared with the blue key) should increase with the signal duration. Results were consistent with LeT's predictions, showing that the context effect is obtained even when subjects do not need to make a choice between two keys presented simultaneously.     

Interstimulus interval as a discriminative stimulus: Evidence of the generality of a novel asymmetry in temporal discrimination learning

Three appetitive conditioning experiments with rats examined temporal discrimination learning within Pavlovian conditioning trials. In all experiments, the duration of a feature white noise stimulus signaled whether or not a subsequent 10-s target tone would be reinforced. In Experiment 1, the feature durations were 4 and 1 min. For one group of rats (Group 4+/1−), 4 min of noise signaled that the tone would be reinforced and 1 min of noise signaled that the tone would not be reinforced. A second group (Group 1+/4−) was trained with the reverse contingency. The results showed a clear asymmetry in temporal discrimination learning: rats trained with 4+/1− (Long+/Short−) learned the discrimination readily (responding more in the tone on reinforced than on nonreinforced trials), whereas rats trained with 1+/4− (Short+/Long) did not. In Experiment 2, the feature durations were shortened to 60 and 15 s. Due to strong excitatory conditioning of the 15-s feature, the reverse asymmetry was observed, with the Short+/Long− discrimination learned more readily than the Long+/Short− discrimination. However, Experiment 3 demonstrated that the original Long+/Short− advantage could be recovered while using 60− and 15-s feature durations if the excitatory conditioning of the feature was reduced by including nonreinforced feature trials. The results support previous research involving the timing of intertrial intervals and are consistent with the temporal elements hypothesis which holds that the passage of time is encoded as a series of hypothetical stimulus elements.     

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.     

Manipulating pre-feed, density of reinforcement, and extinction produces disruption in the Location variation of a temporal discrimination task in pigeons

When pharmacological and non-pharmacological agents are used to disrupt temporal discrimination, two major findings have emerged in the literature. One result reveals lateral shifts of the psychophysical curve for time due to disruptors, while the other is a decrease in accuracy for classifying short and long intervals and a flattening of the psychophysical curve. These results represent a discrepancy within the timing literature that requires clarification. The current study determined the effects of pre-feed, increased density of reinforcement during session, and extinction on the Location variation of a temporal discrimination procedure. The results showed that extinction and pre-feed (at higher levels), when presented in an acute fashion, led to right-ward shifts in the psychophysical curve. Our results, when compared to similar studies in the literature, suggest that lateral shifts are more likely to be found due to disruptors when the Location variation is being used and when procedures are less complicated.     

Effects of a parasitic nematode on male mate choice in a livebearing fish with a coercive mating system (western mosquitofish, Gambusia affinis)

Studies of choice in steady state have shown that sensitivity to reinforcement increases with increasing fixed-ratio changeover (FR CO) requirements. We assessed the generality of this finding with choice in transition. Food deliveries were programmed according to concurrent variable-interval (VI) schedules. Seven different VI pairs arranged ratios of food deliveries (left/right) of 27:1, 9:1, 3:1, 1:1, 1:3, 1:9, and 1:27 at a constant overall rate across components. Within sessions, all seven ratios were presented in random order. Each component lasted for 10 food deliveries; components were separated by 60-s blackouts. A changeover lever required 1, 2, 4, 8, 16, 32, and 64 responses to alternate between two main levers. Redeterminations to all FR COs, but 64 responses, were obtained in descending order. Choice adjusted rapidly to rapid changes in the reinforcer ratio, tracking the lever associated with the highest probability of reinforcer. Sensitivity to reinforcement increased with increasing FR CO, replicating the negatively accelerated function found in our earlier study. With successive reinforcers in components, however, sensitivity reached asymptote values sooner with the largest (8, 16, and 32 responses), than with the smallest (1, 2, and 4 responses), FR CO requirements.     

Temporal generalization accounts for response resurgence in the peak procedure

The peak interval (PI) procedure is commonly used to evaluate animals' ability to produce timed intervals. It consists of presenting fixed interval (FI) schedules in which some of the trials are replaced by extended non-reinforced trials. Responding will often resume (resurge) at the end of the non-reinforced trials unless precautions are taken to prevent it. Response resurgence was replicated in rats and pigeons. Variation of the durations of the FI and the non-reinforced probe trials showed it to be dependent on the time when reinforcement is expected. Timing of both the normal time to reinforcement, and the subsequent time to reinforcement during the probe trials followed Weber's law. A quantitative model of resurgence is described, suggesting how animals respond to the signaling properties of reinforcement omission. Model results were simulated using a stochastic binary counter.     

Effect of clozapine on interval timing and working memory for time in the peak-interval procedure with gaps

Previous research indicates that dopamine controls both the speed of an internal clock [Maricq, A.V., Church, R.M., 1983. The differential effects of haloperidol and methamphetamine on time estimation in the rat. Psychopharmacology 79, 10-15] and sharing of resources between the timer and other cognitive processes [Buhusi, C.V., 2003. Dopaminergic mechanisms of interval timing and attention. In: Meck, W.H. (Ed.), Functional and Neural Mechanisms of Interval Timing. CRC Press, Boca Raton, FL, pp. 317-338]. For example, dopamine agonist methamphetamine increases the speed of an internal clock and resets timing after a gap, while dopamine antagonist haloperidol decreases the speed of an internal clock and stops timing during a gap [Buhusi, C.V., Meck, W.H., 2002. Differential effects of methamphetamine and haloperidol on the control of an internal clock. Behav. Neurosci. 116, 291-297]. Using a 20-s peak-interval procedure with gaps we examined the acute effects of clozapine (2.0mg/kg i.p.), which exerts differential effects on dopamine and serotonin in the cortex and striatum, two brain areas involved in interval timing and working memory. Relative to saline, clozapine injections shifted the response functions leftward both in trials with and without gaps, suggesting that clozapine increased the speed of an internal clock and facilitated the maintenance of the pre-gap interval in working memory. These results suggest that clozapine exerts effects in different brain areas in a manner that allows for the pharmacological separation of clock speed and working memory as a function of peak trials without and with gaps.     

Defoliation causes parallel temporal responses in a host tree and its fungal symbionts

Growth of the host and its symbiont is often closely linked and so host damage may negatively affect the symbiont. While negative effects of aboveground herbivory on belowground fungal symbionts have been reported in several woody and herbaceous plants, here we report, for the first time, on differential effects of the timing of foliar damage on ectomycorrhizal (ECM) fungal symbionts. The phenologies of host trees and their ECM symbionts differ; the growth of the latter mainly occurs later in the season than that of the host. By removing Scots pine foliage on three occasions during the growing season (early, middle and late season defoliation) in one, two or three successive years, we demonstrate that, despite the differences in the seasonal growth dynamics of the tree and the symbionts, ECM fungi follow the host’s response patterns to defoliation. Early season defoliation was most detrimental to the host and resulted in an increased proportion of low-biomass ectomycorrhizae which are presumed to require less carbon from the host tree. This may improve the recovery of the host, as most roots remained mycorrhizal in spite of the defoliation treatments repeated in successive years.     

Elevated temperature alters the ionic dependence of amine-induced pacemaker activity in a conditional burster neuron

Summary The anterior burster neuron of the lobster (Panulirus interruptus) stomatogastric ganglion is a conditional burster that functions as the primary pacemaker for the pyloric motor network. When modulatory inputs to this cell are blocked, it loses its bursting properties and becomes quiescent. Applications of the monoamines, dopamine, octopamine or serotonin restore rhythmic bursting in this cell (Flamm and Harris-Warrick 1986). At 15 °C, serotonin- and octopamine-induced oscillations depend critically upon sodium entry (blocked by low sodium saline or tetrodotoxin); dopamine-induced oscillations depend upon calcium entry (blocked by reduced extracellular calcium; Harris-Warrick and Flamm 1987). We show here that the ionic dependence of amine-induced oscillations in the anterior burster cell differs at 15 and 21 °C. At 21 °C, all amines have the potential to induce rhythmic oscillations in saline containing tetrodotoxin. At the elevated temperature and in tetrodotoxin, both calcium and sodium currents are essential for the maintenance of dopamine-induced oscillaions; serotonin-induced oscillations do not depend upon either calcium or sodium alone; octopamine-induced oscillations do not depend upon calcium and show a variable dependence upon sodium. Thus, multiple ionic mechanisms, which vary with both the modulator and the ambient temperature, can be recruited to support rhythmic activity in a conditional burster neuron.Abbreviations AB anterior burster - PD pyloric dilator - PY pyloric constrictor - DA dopamine - 5HT serotonin - Oct octopamine - STG stomatogastric ganglion - TTX tetrodotoxin - GSP graded synaptic potential     

Optimisation of a screening procedure for house dust mite numbers in carpets and preliminary application to buildings

The relative performances of the mobility and heat escape methods for measuring house dust mite numbers in carpet has been evaluated. The optimum method was found to be the mobility method for 24 hours at ambient temperatures which exhibited a mite collection efficiency of approximately 30%. Measurements in three dwellings showed that the method should be applied to carpet at several locations in the living room and/or bedrooms as a general sampling procedure, or adjacent to loungeroom seating to determine the worst case scenario. Carpets using different types of fibre within the same dwelling should be assessed separately. For the limited number of dwellings investigated, wool carpets were found to exhibit higher mite numbers than nylon carpets, even when the former had been insect-resist treated. No house dust mites were found in the wool carpets of an office building which was mechanically ventilated and heated and achieved low indoor humidities in winter.