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Time is a universal psychological dimension, but time perception has often been studied and discussed in relative isolation. Increasingly, researchers are searching for unifying principles and integrated models that link time perception to other domains. In this review, we survey the links between temporal cognition and other psychological processes. Specifically, we describe how subjective duration is affected by nontemporal stimulus properties (perception), the allocation of processing resources (attention), and past experience with the stimulus (memory). We show that many of these connections instantiate a "processing principle," according to which perceived time is positively related to perceptual vividity and the ease of extracting information from the stimulus. This empirical generalization generates testable predictions and provides a starting-point for integrated theoretical frameworks. By outlining some of the links between temporal cognition and other domains, and by providing a unifying principle for understanding these effects, we hope to encourage time-perception researchers to situate their work within broader theoretical frameworks, and that researchers from other fields will be inspired to apply their insights, techniques, and theorizing to improve our understanding of the representation and judgment of time. (PsycINFO Database Record

Five experiments were conducted to determine the effects of hippocampal damage on timing and the memory for temporal events. In Experiments 1-3, rats were trained to discriminate between auditory signals that differed in both duration (2 or 8 s) and rate (2 or 16 cycles/s). Half of the rats were trained to discriminate duration, and half were trained to discriminate rate. After rats acquired the relevant discrimination, signals with intermediate durations and rates were presented to obtain psychophysical functions that related signal duration and/or rate to response choice. Rats then received either lesions of the fimbria-fornix or control operations. Postoperatively, the accuracy of duration and rate discriminations as measured by the difference limen (DL) was unaffected by the lesion, but the point of subjective equality (PSE) was shifted to a shorter duration and a slower rate by the lesion in Experiment 1. Both rats with lesions and rats with control operations showed cross-modal transfer of duration and rate from the auditory signals used in training to visual signals used in testing in Experiment 2. A 5-s delay was imposed between the end of a signal and the opportunity to respond in Experiment 3. The delay served as a retention interval for the rats trained in the rate discrimination, and the rats with fimbria-fornix lesions were selectively impaired by the addition of the delay as measured by an increase in the DL. The delay did not serve as a retention interval for rats trained in the duration discrimination because they were able to continue timing through the delay. A peak procedure was employed in Experiment 4. The maximum response rate of control rats was approximately at the time of scheduled reinforcement (20 s), but the maximum response rate of rats with fimbria-fornix lesions was reliably earlier than the time of scheduled reinforcement. When a 5-s gap was imposed in the signal, control rats summed the signal durations before and after the gap, whereas rats with fimbria-fornix lesions showed no retention of the signal duration prior to the gap. Experiment 5 continued the testing of the rats used in Experiments 1-4 and showed that rats with lesions had an impairment in a test of spatial working memory in an eight-arm radial maze. Taken together, these results demonstrate that a fimbria-fornix lesion interferes with temporal and spatial working memory, reduces the remembered time of reinforcement stored in reference memory, and has no effect on the animal's sensitivity to stimulus duration.

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.

The authors hypothesized that during a gap in a timed signal, the time accumulated during the pregap interval decays at a rate proportional to the perceived salience of the gap, influenced by sensory acuity and signal intensity. When timing visual signals, albino (Sprague-Dawley) rats, which have poor visual acuity, stopped timing irrespective of gap duration, whereas pigmented (Long-Evans) rats, which have good visual acuity, stopped timing for short gaps but reset timing for long gaps. Pigmented rats stopped timing during a gap in a low-intensity visual signal and reset after a gap in a high-intensity visual signal, suggesting that memory for time in the gap procedure varies with the perceived salience of the gap, possibly through an attentional mechanism.

The roles that psychostimulant sensitization and tolerance play in temporal perception in the seconds-to-minutes range were assessed in rats. Cocaine (20 mg/kg/day) was administered for 2 weeks either intermittently via daily injections (induces sensitization) or continuously via an osmotic minipump (induces tolerance). Interval timing was evaluated throughout administration and withdrawal. Injections of cocaine caused immediate, proportional, leftward shifts in peak times, indicating an increase in the speed of an internal clock. These shifts grew progressively larger with repeated administration, indicating that stimulant-induced increases in clock speed can be sensitized. Continuous cocaine administration produced no reliable effects. These results suggest that the mechanisms of sensitization may play a considerable role in drug-induced alterations of the perception of time.

Previous data suggest that rats (Rattus norvegicus) and pigeons (Columba livia) use different intervaltiming strategies when a gap interrupts a to-be-timed signal: Rats stop timing during the gap, and pigeons reset their timing mechanism after the gap. To examine whether the response rule is controlled by an attentional mechanism dependent on the characteristics of the stimuli, the authors manipulated the intensity of the signal and gap when rats and pigeons timed in the gap procedure. Results suggest that both rats and pigeons stop timing during a nonsalient gap and reset timing after a salient gap. These results also suggest that both species use similar interval-timing mechanisms, influenced by nontemporal characteristics of the signal and gap. Clearly it is advantageous for all organisms to be able to anticipate and predict the occurrence of an event in space and time. Many types of behavior (e.g., Pavlovian and instrumental conditioning, risk-sensitive foraging, and skilled motor learning) show sensitivity to the temporal arrangements of events in a manner suggesting that organisms are able to do the “right thing at the right time.” Consequently, it is not surprising that a wide range of species (e.g., bees, fish, turtles, birds, rodents, and monkeys, as well as human infants and adults) show that the ability to use temporal information in the seconds-to-hours range is important and that similar psychophysical properties hold across species (e.g., Bateson & Kacelnik, 1997, 1998; Brodbeck, Hampton, & Cheng, 1998; Gallistel, 1990; Lejeune & Wearden, 1991; Matell &

The effects of signal modality on duration classification in college students were studied with the duration bisection task. When auditory and visual signals were presented in the same test session and shared common anchor durations, visual signals were classified as shorter than equivalent duration auditory signals. This occurred when auditory and visual signals were presented sequentially in the same test session and when presented simultaneously but asynchronously. Presentation of a single modality signal within a test session, or both modalities but with different anchor durations did not result in classification differences. The authors posit a model in which auditory and visual signals drive an internal clock at different rates. The clock rate difference is due to an attentional effect on the mode switch and is revealed only when the memories for the short and long anchor durations consist of a mix of contributions from accumulations generated by both the fast auditory and slower visual clock rates. When this occurs auditory signals seem longer than visual signals relative to the composite memory representation.

Contrary to data showing sensitivity to nontemporal properties of timed signals, current theories of interval timing assume that animals can use the presence or absence of a signal as equally valid cues as long as duration is the most predictive feature. Consequently, the authors examined rats' behavior when timing the absence of a visual or auditory stimulus in trace conditioning and in a "reversed" gap procedure. Memory for timing was tested by presenting the stimulus as a reversed gap into its timed absence. Results suggest that in trace conditioning (Experiment 1), rats time for the absence of a stimulus by using its offset as a time marker. As in the standard gap procedure, the insertion of a reversed gap was expected to "stop" rats' internal clock. In contrast, a reversed gap of 1-, 5-, or 15-s duration "reset" the timing process in both trace conditioning (Experiment 2) and the reversed gap procedure (Experiment 3). A direct comparison of the standard and reversed gap procedures (Experiment 4) supported these findings. Results suggest that attentional mechanisms involving the salience or content of the gap might contribute to the response rule adopted in a gap procedure. When an ongoing timed interval is interrupted by a break or gap, animals seem to suspend their temporal processing and choose a response rule that falls between two extremes: They may restart the entire timing process, a phenomenon usually called "reset" (Church, 1978; W. A. Roberts, Cheng, & Cohen, 1989) or they may "stop" timing for the duration of the gap and resume it after the gap (Church, 1978; Meek, Church, & Olton, 1984; S. Roberts

The properties of the internal clock, temporal memory, and decision processes used to time short durations were investigated. The peak-interval procedure was used to evaluate the timing of 8-, 12-, and 21-s intervals, and analyses were conducted on the mean response functions and on individual trials. A distractor task prevented counting, and visual feedback on accuracy and precision was provided after each trial. Mean response distributions were (a) centered at the appropriate real-time criteria, (b) highly symmetrical, and (c) scalar in their variability. Analysis of individual trials indicated more memory variability relative to response threshold variability. Taken together, these results demonstrate that humans show the same qualitative timing properties that other animals do, but with some quantitative differences.

Three experiments assessed the ability of male Sprague-Dawley rats to organize the spatial locations of different food types in a hierarchical manner to maximize the efficiency of working memory. Independent groups were exposed, on a 12-arm radial maze, to baiting arrangements varying in the stability of the pattern and type of food used as bait. Training rats with stable, differentiable baiting arrangements produced increased accuracy in choice performance, hierarchically ordered patterns of choice selection, slower growth of proactive interference when trials were massed, and the learning of the geometrical relations among food types independent of other extramaze cues. Such findings are strong evidence of the rat's ability to encode and use local cues for navigation, based on properties of the reinforcer. The application of a chunking strategy may provide for more efficient use of working memory by facilitating information storage, recall, or resetting mechanisms.

Seven experiments assessed the ability of rats to process temporal information from two internal clocks simultaneously and independently. In the first six experiments a light stimulus signalled an overall interval between the beginning of a trial and the availability of food reinforcement (e.g., a 50-s fixed interval). During the overall interval a sound stimulus was used to signal shorter intervals that divided the overall interval into equal segments. When there was a fixed temporal relation between the final segment signal and the availability of reinforcement, there was a double-scallop pattern of responding throughout the segmented overall interval; the function relating response rate to time during segment intervals was similar to the function relating response rate to time in unsegmented overall intervals; a change in response rate occurred at the time that a normally presented segment signal was omitted. Taken together, the results indicate that rats timed the overall interval and the segment intervals simultaneously and independently without interference. In Experiment 7 a light stimulus was used on some trials, and a sound stimulus was used on other trials to signal a discrete-trial 50-s peak procedure. When these two signals were presented in compound, there was a leftward shift of the response function, which suggests that rats timed both signals simultaneously. For all of the experiments a scalar timing model with specific stimulus integration rules is used to explain the results. The stimulus integration rule used in the first six experiments, in which there were two signals for the same reinforcement, was to respond if both the segment and the overall interval had exceeded a response threshold. The stimulus integration rule used in Experiment 7, in which there were two signals for different reinforcements, was to respond if the response threshold for either interval had been exceeded.

ion of Temporal Attributes Warren H. Meek and Russell M. ChurchBrown University Cross-modal transfer of classification rules for temporal intervals was demon-strated in four experiments. In Experiment 1, rats learned a temporal discrim-ination between a 2-sec and an 8-sec signal in one modality (vision or audition).There was positive transfer to a temporal discrimination between a 2-sec andan 8-sec signal in the other modality when the response rule was maintained,and negative transfer when the response rule was reversed. Experiment 2 dem-onstrated positive cross-modal transfer in a temporal generalization procedure.Experiment 3 demonstrated cross-modal transfer of both duration arid temporallocation in a procedure in which rats were exposed to three successive signaldurations. Experiment 4 demonstrated cross-modal transfer of both duration andtemporal order in a procedure in which rats were exposed to simultaneouslypresented signal durations, one auditory and one visual. The conclusion is thatrats can abstract temporal attributes from modality-specific aspects of a signal.

The similarity of animal counting and timing processes was demonstrated in four experiments that used a psychophysical choice procedure. In Experiment 1, rats initially learned a discrimination between a two-cycle auditory signal of 2-sec duration and an eight-cycle auditory signal of 8-sec duration. For the number discrimination test, the number of cycles was varied, and the signal duration was held constant at an intermediate value. For the duration discrimination test, the signal duration was varied, and the number of cycles was held constant at an intermediate value. Rats were equally sensitive to a 4:1 ratio of counts (with duration controlled) and a 4:1 ratio of times (with number controlled). The point of subjective equality for the psychophysical functions that related response classification to signal value was near the geometric mean of the extreme values for both number and duration discriminations. Experiment 2 demonstrated that 1.5 mg/kg of methamphetamine administered intraperitoneally shifted the psychophysical functions for both number and duration leftward by approximately 10%. Experiment 3 demonstrated that the magnitude of cross-modal transfer from auditory signals to cutaneous signals was similar for number and duration. In Experiment 4 the mapping of number onto duration demonstrated that a count was approximately equal to 200 msec. The psychophysical functions for number and duration were fit with a scalar expectancy model with the same parameter values for each attribute. The conclusion was that the same internal mechanism is used for counting and timing. This mechanism can be used in several modes: the "event" mode for counting or the "run" and the "stop" modes for timing.

Four experiments studied the scaling of time by rats. The purpose was to determine if internal clock and memory processes could be selectively adjusted by pharmacological manipulations. All of the experiments used a temporal discrimination procedure in which one response ("short") was reinforced following a 2-sec noise signal and a different response ("long") was reinforced following an 8-sec noise signal; unreinforced signals of intermediate duration were also presented. The proportion of "long" responses increased as a function of signal duration. All drugs were administered intraperitoneally (ip) and their effect on clock or memory processes was inferred from the observed pattern of change in the point of subjective equality of the psychophysical functions under training and testing conditions. Experiment 1 demonstrated that methamphetamine (1.5 mg/kg) can selectively increase clock speed and that haloperidol (.12 mg/kg) can selectively decrease clock speed. Experiment 2 demonstrated that footshock stress (.2 mA) can selectively increase clock speed during continuous administration but leads to a decrease in clock speed below control values when the footshock is abruptly terminated. Experiment 3 demonstrated that vasopressin (.07 pressor units/kg) and oxytocin (.02 pressor units/kg) can selectively decrease the remembered durations of reinforced times, which suggests that memory storage speed increased. Experiment 4 demonstrated that physostigmine (.01 mg/kg) can selectively decrease the remembered durations of reinforced times and that atropine (.05 mg/kg) can selectively increase these remembered durations, which suggests that memory storage speed was differentially affected. The conclusion is that internal clock and memory processes can be dissociated by selectively adjusting their speed of operation and that these changes can be quantitatively modeled by a scalar timing theory.

Temporal integration in duration and number discrimination by rats was investigated with the use ofa psychophysical choice procedure. A response on one lever ("short" response) following a l-s white-noise signal was followed by food reinforcement, and a response on the other lever ("long" response) following a 2-s white-noise signal was also followed by food reinforcement. Either response following a signal of one of five intermediate durations was unreinforced. This led to a psychophysical function in which the probability of a long response was related to signal duration in an ogival manner. On 2 test days, a white-noise signal wit h 5, 6, 7, 8, or 10 segments of either 0.5-s on and 0.5-s off or l-s on and l-s off was presented, and a choice response following these signals was unreinforced. The probability of a long response was the same function of a segmented signal and a continuous signal if each segment was considered equivalent to 200 ms. A quantitative fit of a scalar estimation theory suggested that the latencies to initiate temporal integration and to terminate the process are both about 200 ms, and that the same internal accumulation process can be used for counting and timing. Rats and other animals have the ability to discriminate between the number of events. The events may either be stimuli or responses. Davis and Memmott (1982) reviewed this evidence. Not all studies that attempted to train rats to discriminate between the number of events have been successful (Imada, Shuku, & Moriya, 1983), and in many cases the discrimination may have been stimulus durations rather than stimulus number. Even when all temporal cues are controlled, however, animals can learn to discriminate between the number of events (Fernandes & Church, 1982). There are many similarities between animal counting and timing processes. (In this article counting refers to number discrimination, and timing refers to duration discrimination; some authors prefer to restrict the term counting further, e.g., Davis & Memott, 1982, and Gelman & Gallistel, 1978.) Meck and Church