Signal Modality Interactions Reveal Developmental/Aging Changes in Interval Timing

Interval-timing paradigms have recently attracted interest as a method of studying age-related differences in cognition (e.g., Craik & Hay, 1999; McCormack, Brown, Maylor, Darby, & Green, 1999; Vanneste & Pouthas, 1999; Wearden, Wearden & Rabbit, 1997; for review see Block, Zakay, & Hancock, 1998). Models of timing and time perception often include components of interest to many cognitive aging researchers, such as attention, memory, and decision processes (e.g., Gibbon, Church, & Meck, 1984; Thomas & Weaver, 1975). Furthermore, temporal cognition is reliably affected by a wide range of behavioral (e.g., Lejeune, 1998; Macar, Grondin, & Casini, 1994) and neurobiological (Gibbon, Malapani, Dale, & Gallistel, 1997; Ivry, 1996; Meck & Benson, in press) manipulations of attention and memory.

For example, valid and invalid cues to the modality of an upcoming to-be-timed stimulus can affect the latency to begin timing (Meck, 1984), and the efficiency of these cues is influenced by drug-induced changes in attention (e.g., Penney, Holder & Meck, 1996). Internal clock and memory processes involved in time perception are also affected by manipulations of brain dopamine levels in both normal adults (Rammsayer, 1997) and patients with Parkinson's disease (Malapani, Rakitin, Meck, Deweer, Dubois, & Gibbon, 1998). Other clinical disorders such as Alzheimer's disease and amnesia also impact time perception in a manner consistent with the cognitive disruptions accompanying these conditions (e.g., Carrasco, Guillem, & Redolat, 2000; Nichelli, Venneri, Molinari, Tavani, & Grafman, 1993). The sensitivity of interval timing procedures to variables affecting attention and memory makes these paradigms a useful method of studying age-related changes at both behavioral and physiological levels (e.g., Meck, 1996).

The temporal bisection task (Allan & Gibbon, 1991) has been used in conjunction with an information-processing model of timing (Gibbon et al., 1984) to investigate the roles of attention and memory in time perception. In this model, time is marked by pacemaker pulses being gated to an accumulator when attention to a stimulus closes a switch. During training for the bisection task, participants learn to call one duration (e.g., 3 s) "short" and another duration (e.g., 6 s) "long". The accumulated pacemaker pulses associated with each label are then passed into reference memory. Because of variability in the encoding and decoding of durations, each label is associated with a distribution of time values rather than a single value (e.g., Gibbon et al., 1984).

At test, participants are presented with stimuli that last for durations corresponding to either the short anchor, the long anchor, or an intermediate value and asked to indicate whether the presented duration is closer to the "short" or "long" anchor. The dependent variable is the proportion of trials in which the participant classifies each test duration as "long". The decision whether to call a particular stimulus item "short" or "long" is made by comparing the number of pacemaker pulses accumulated during the presented stimulus with the number of pulses associated with "short" and "long" in reference memory.

Recently, the bisection task has been used in combination with the classic finding that "sounds are judged longer than lights" to examine how attention and reference memory act together in temporal discrimination. When both modalities are used within an experimental session, an auditory stimulus is typically judged as longer than a visual stimulus of the same physical duration (e.g., Goldstone & Goldfarb, 1964; Walker & Scott, 1981; Wearden, Edwards, Fakhri, & Percival, 1998). Penney, Allan, Meck, and Gibbon (1998) suggested this difference may occur because auditory signals capture and hold attention relatively easily, whereas attending to visual stimuli requires more attentional control (Meck, 1984). Because auditory stimuli are better able to capture and hold attention, they are more efficient than visual stimuli in closing the attentional switch that allows pacemaker pulses to accumulate. As a result, more pacemaker pulses accumulate for an auditory stimulus than for a visual stimulus of the same physical duration. If a test stimulus from either modality is compared to "short" and "long" distributions in reference memory that are comprised of values from both modalities then an auditory test stimulus will be more likely judged "long" than a visual stimulus of the same physical duration.

In an extensive series of experiments using college students as participants, Penney and associates (e.g., Penney et al., 1998; Penney, Gibbon, & Meck, 2000) tested participants with the temporal bisection procedure using both auditory and visual signals. The results of these studies indicated that when the same anchor durations were used for both modalities, the auditory and visual test stimuli were compared to a reference memory distribution comprised of both signal modalities.

Under these "memory mixing" conditions the classic finding was obtained: Auditory stimuli were more likely to be judged "long" than visual stimuli of the same physical duration. However, this was not the case when separate reference memory distributions were obtained for the two modalities (for example, if the modalities were used in separate experimental sessions rather than within one session or if different anchor points were used for each modality). See Wearden et al. (1998) for additional data relating modality differences to the attention and memory components of the internal clock model.

Another area of timing research that has recently attracted interest in the cognitive aging literature is the impact that circadian arousal patterns have on age differences in attention and memory (for a review of the internal clocks that mediate circadian and interval timing see Hinton & Meck, 1997). Scores on measures such as the Horne-Ostberg (1976) Morningness-Eveningness Questionnaire generally classify older adults as "morning" types and young adults as "neutral" or "evening" types, and these classifications are consistent with circadian rhythmicities on variety of physiological, psychological, and cognitive measures (e.g., Hoch, Reynolds, Jennings, Monk, Buysse, Machen, & Kupfer, 1992; May, Hasher, & Stolzfus, 1993; Mecacci, Zani, Rochetti, & Lucioli, 1986; see Yoon, May & Hasher, 1999 for a review).

For example, May et al. (1993) found that for participants tested in the afternoon (young adults' optimal and older adults' nonoptimal time of day), young adults showed better recognition memory performance than did older adults, replicating standard findings. However, there were no age differences in recognition memory for participants tested in the morning (young adults' nonoptimal and older adults' optimal time of day). Particularly for older adults, performance on measures of controlled attention such as the Stroop (1935) interference effect or the interference condition of the Trail-Making Test (Reitan, 1958) changes as a function of time of day, but performance of well-learned, relatively automatic tasks (such as vocabulary performance or Stroop color-naming) does not (Hasher, Zacks, & May, 1999; May & Hasher, 1998; for review see Yoon et al., 1999).

Our current investigations have built on the above findings and on a previous experiment examining circadian influences on interval timing in animals (Meck, 1991) to determine how the interaction of subject and task variables thought to affect attention and memory influence performance on a time perception task. Young and older adults participated in a temporal bisection experiment using auditory and visual signals and both single and compound (divided attention) trials during either the morning or the afternoon. It was hypothesized that older adults' performance would be worse than that of young adults under task conditions requiring more controlled attention, particularly when older adults were tested at their "non-optimal" time of day (e.g., Lustig & Meck, 1998, 2001a). The classic finding that "sounds are judged longer than lights" was obtained, with greater modality differences in the afternoon. Older adults showed greater disparities in their duration judgments for visual versus auditory stimuli than did young adults. In addition, while young adults performed equally well on the single and compound trials, older adults performed as well as young adults on the single trials, but much worse than young adults on the compound trials, which required divided attention. Finally, older adults tested in the morning showed unusually high sensitivity to the single visual trials and relative insensitivity to the other trial types, suggesting that reduced attentional resources may have caused them to focus their efforts on this trial type to the detriment of others. These results suggest that age and circadian differences in the perception of time are influenced by a slower clock rate for visual than for auditory signals and the reduction and allocation of attentional control. Our most recent work has also shown developmental changes in attention and retrieval of durations from reference memory in children (8-12 years, young adults (18-25 years) and aged adults (60-75 years) as a function of signal modality (e.g., Lustig & Meck, 2001b).

Annotated Bibliography

Allan, L.G., & Gibbon, J. (1991). Human bisection at the geometric mean. Learning and Motivation, 22, 39-58.

*This paper is the "classic" example of the temporal bisection in humans and the finding that subjects find the subjective middle between two durations to be at the geometric mean of those durations for a wide range of signal values.

Block, R.A., Zakay, D., & Hancock, P.A. (1998). Human aging and duration judgements: A meta-analytic review. Psychology and Aging, 13, 584-596.

Vanneste, S., & Pouthas, V. (1999). Timing in aging: The role of attention. Experimental Aging Research, 25, 49-67.

Wearden, J.H.; Wearden, A.J.; Rabbit, P.M.A. (1997). Age and IQ effects on stimulus and response timing. Journal of Experimental Psychology: Human Perception and Performance, 23, 962-979.

*These papers provide an excellent review of the effects of normal aging on timing and time perception.

Gibbon, J. Malapani, C., Dale, C.L., & Gallistel, C.R. (1997). Toward a neurobiology of temporal cognition: Advances and challenges. Current Opinion in Neurobiology, 7, 170-184.

Ivry, R. B. (1996). The representation of temporal information in perception and motor control. Current Opinion in Neurobiology, 6, 851-857.

Meck, W.H. (1996). Neuropharmacology of timing and time perception. Cognitive Brain Research, 3, 227-242.

*These papers provide an excellent overview of what is currently known about the neurobiological and pharmacological basis of interval timing in the seconds-to-minutes range and relates it to various theoretical models and constraints.

Penney, T.B., Allan, L.G., Meck, W.H., & Gibbon, J. (1998). Memory mixing in duration bisection. In D.A. Rosenbaum & C.E. Collyer (Eds.), Timing of Behavior: Neural, Psychological, and Computational Perspectives (pp.165-193). Cambridge, MA: MIT Press.

Penney, T. B., Gibbon, J., & Meck, W. H. (2000). Differential effects of auditory and visual signals on clock speed and temporal memory. Journal of Experimental Psychology: Human Perception and Performance, 26, 1770-1787.

Walker, J.T., & Scott, K.J. (1981). Perceived duration of lights, tones, and gaps. Journal of Experimental Psychology: Human Perception and Performance, 7, 1327-1339.

Wearden, J.H., Edwards, H., Fakhri, M., & Percival, A. (1998). Why "sounds are judged longer than lights": Application of a model of the internal clock in humans. Quarterly Journal of Experimental Psychology, 51B, 97-120.

*These papers provide the essential basis for understanding how and why auditory stimuli are perceived to be longer than visual stimuli.

References

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