T HE QUART ERLY JOURNAL OF EXPERIMENT AL PSYCHOL OGY, 1998, 51B (3), 193± 217

Peak Proced u re Perform an ce in You n g Ad u lt

an d Aged Rats:
Acq u isition an d Ad ap tation to a Ch an gin g
Tem p oral Criterion

Helga Lejeune and Andre Ferrara

Univ ersity of LieÁ ge, LieÁ ge, Belgium
Monique Sof® e and Marie Bronchart
Catholic Univ ersity of Louv ain, Louv ain-la-Neuv e, Belgium
John H. Wearden
Univ ersity of Manchester, Manchester, U.K .

Twenty-four-month-old and 4-month-old rats were trained on a peak-interval procedure,

where the time of reinforcement was varied twice between 20 and 40 sec. Peak times from the
old rats were consistently longer than the reinforcement time, whereas those from younger
animals tracked the 20- and 40- sec durations more closely. Different measures of perfor-
mance suggested that the old rats were either (1) systematically misremembering the time of
reinforcement or (2) using an internal clock with a substantially greater latency to start and
stop timing than the younger animals. Old rats also adjusted more slowly to the ® rst transi-
tion from 20 to 40 sec than did the younger ones, but not to later transitions. Correlations
between measures derived from within-trial patterns of responding conformed in general to
detailed predictions derived from scalar expectancy theory. However, some correlation values
more closely resembled those derived from a study of peak-interval performance in humans
and a theoretical model developed by Cheng and Westwood (1993), than those obtained in
previous work with animals, for reasons that are at present unclear.

T he temporal regulation of behaviour in animals has long been a major concern of

comparative psychology. As well as results collected with the Differential Reinforcement
of Low rate (DRL ) and Fixed- Interval (FI) schedules, a number of articles published
since the early 1980s have presented data obtained with the Peak I nterval (PI) procedure.

Requests for reprints should be sent to Helga Lejeune, Psychobiology of Temporal Processes Unit (PT PU),
University of LieÁ ge, Faculty of Psychology, 5 Bd. du Rectorat B32, Sart- T ilman 4000 LieÁ ge, Belgium. E-mail:
helga.lejeune@ulg.ac.be
T he experiment reported here was supported by an ``Action concerte e’ ’ grant 92/ 97± 158 from the Ministry of
the Belgian French Community to the University of Louvain.
We would like to thank Dr. A. El Ahmadi for advice on statistical matters and F. S imons for supervising the
software and hardware used in the control of the experiment and data analysis.

q 1998 T he Experimental Psychology S ociety

194 LEJEUNE ET AL.

T his schedule, developed initially by Catania (1970) and then later by Roberts (1981),
derives from the discrete-trial FI procedure. In the usual FI procedure, a food reinforcer
is delivered for the ® rst response that occurs t sec after the start of the interval, and
reinforcer delivery initiates the next interval. On discrete-trial FI, on the other hand,
intervals are signalled by the onset of a stimulus (usually light or sound) which ends at
reinforcer delivery. T he discrete trials also are separated by stimulus-free inter-trial
intervals (IT Is).
Two procedural changes transform discrete-trial FI into a PI procedure: ® rst, some of
the trials (``peak trials’ ’ ) last much longer than do the FI trials; second, these longer trials
end automatically, without reinforcer delivery. A PI session is thus typically a mixture of
some proportion of FI trials (during which a reinforcer is delivered for the ® rst response
t sec after the onset of the trial) and the longer peak trials, where responding is unrein-
forced. Data are usually only taken from peak trials, and, generally, the average response
rate function obtained from these trials increases progressively and smoothly up to a
maximum close to reinforcement time on the FI trials (t) and decreases thereafter with
a slight asymmetry, following a bell-shaped pro® le. Two behavioural indices are usually
derived from this response rate function: one is the peak rate, the highest response rate
from the function, and the other is peak time, the elapsed time in the trial at which peak
rate occurs, which is usually close to the usual time of reinforcement, t.
Behaviour collected with the PI procedure has been used to evaluate the predictions of
current theories of animal timing, particularly Scalar Expectancy Theory (SET, Gibbon,
1977). To be consistent with SET, timing must exhibit two properties. T he ® rst of these is
mean accuracy, the requirement that average estimates of some real time, t, are close to t.
Data from the PI procedure are consistent with this requirement, as the peak time very
closely matches the time when reinforcement is available on the FI trials of the PI
sessions. T he second requirement of SET is the scalar property itself, the requirement
that standard deviations of time estimates increase linearly with the mean. One way of
testing this is to construct a coef® cient of v ariation (cv) statistic by dividing the standard
deviation of behavioural measures by their mean, and the scalar property requires that the
cv remains constant as the time of reinforcement changes. In PI procedure data, the scalar
property is usually tested by ® nding the mean of the response rate versus elapsed time
curve (which corresponds to the peak time) and the standard deviation, derived from the
spread of the curve (i.e. how sharply it rises and falls around its peak). T he ratio of the
standard deviation to the mean is the cv. In practice, this cv is usually close to a
constant fraction of the peak time value as the time of reinforcement is varied (e.g.
Church, Miller, Meck, & Gibbon, 1991). Another way of testing the scalar property is
by superimposition (called superposition in the American literature)Ð the requirement that
measures of timed behaviour superimpose on the same relative scale as absolute dura-
tions timed are varied. Data from the PI procedure show a close approximation to
superimposition, as averaged response rate versus elapsed time functions obtained
with different reinforcement time FI values can be superimposed when relative
response rates are plotted against relativ e elapsed timeÐ that is when elapsed time is
plotted as a fraction of the time of reinforcement.
According to SET, the functional architecture underlying performance on timing tasks
including the PI procedure is composed of three interacting levels: a clock, a memory, and
 PEAK PROCEDURE IN AGED RATS 195

a decision level (Church, 1984). At the clock level, impulses generated by a pacemaker are
sent to an accumulator through a switch, which can be closed (so that impulses pass) or
open (impulses are stopped). T he closure of the switch is triggered by incoming signi® -
cant temporal information, its opening by the end of the temporal episode to be estimated
(Church, 1984; Gibbon, Church, & Meck, 1984). T he closure and opening latencies of
the switch can be modulated by variables among which are the nature of the stimulus and
attention (Roberts, 1981, 1983). T he pacemaker rate can also be in¯ ucenced, for instance
by drugs (Maricq, Roberts, & Chuch, 1981). T he accumulator is a short-term store
similar to an ``up’’ counter incremented by impulses that have passed the switch.
T he memory level includes a short-term store (which brie¯ y retains information
loaded from the accumulator) and a long-term store (reference memory) where reinforced
durations are transferred at the end of a trial. Durations stored in long-term memory can
be modulated by a memory constant K*, which can be equal to, greater than, or less than
1, depending on between-subject differences, or on various experimental manipulations
such as the administration of drugs (Meck, 1983, 1996). If K* is equal to 1, remembered
time is equal to reinforced time, whereas if K* is less than 1, remembered time is shorter
than the reinforced duration. If K* is greater than 1, the remembered time of reinforce-
ment is greater than the real reinforced time.
Finally, the decision level involves a comparator that compares the number of pulses
currently in the short-term store, n, and a sample from reference memory, n*. A decision
as to whether or not to respond depends on the comparison of the absolute difference
between n and n*, expressed as a fraction of n*, and a threshold b (Church, Meck, &
Gibbon, 1994). If this normalized difference is less than the threshold, responding is
initiated, and the response state persits until the threshold is exceeded. T his model of PI
responding has been used to account for response patterns within individual PI trials, as
well as patterns of responding averaged from whole experimental sessions. T he descrip-
tion given above refers to a ``one-threshold’’ model, where the threshold for starting and
stopping responding is the same. One obvious variant mentioned in Gibbon and Church
(1990) and explored in a number of subsequent articles (e.g. Cheng & Miceli, 1996;
Church et al., 1994) is a ``two-threshold model’’ , where the thresholds for starting and
stopping responding are not necessarily the same. T hese models of PI responding are
close to the original one invented by Gibbon and Church (1990), which in terms of its
application in published work has become the ``standard model’’. However, some other
models involving different decision processes have been shown to account for obatined
data equally well (Cheng & Miceli, 1996).
T he present article uses the PI procedure, and the analytical mechanisms of SET, to
examine any potential effects of ageing on timing in rats. T he effect of age on timing in
humans and animals has long given rise to speculation (e.g. that the pacemaker of a
hypothesized internal clock might ``slow down’’ with age), but, as Wearden, Wearden
and Rabbitt (1997) show, there is a paucity of experimental evidence as to what the effects
of ageing on timing really are. Furthermore, the evidence that exists is contradictory and
dif® cult to interpret, at least for studies with humans. Comparison of the behaviour of old
and younger members of a species on operant tasks is complicated by the fact that old
subjects frequently respond at lower rates than do younger ones (an effect that will be
seen later in our study), so some timing tasks like DRL, where temporal regulation and
196 LEJEUNE ET AL.

response rate are wholly or partially confounded, are unsuitable for between-age compar-
isons. T he advantage of the PI procedure over some others is that it allows the dissocia-
tion of response timing and response rate; in fact, peak rate can vary without changing
peak time (Roberts, 1981), so the precision of timing can be compared under conditions in
which the absolute rate of response emitted varies, as, for example, between subjects of
different ages.
Another procedure that dissociates overall response rate and measures of timing is the
two-leverl DRL schedule (deriving from Mechner & Guevrekian’s Fixed Minimum Inter-
v al schedule, 1962). On this schedule, the temporal criterion for reinforcement is framed
by successive A± B lever presses. T hus, pressing Lever A starts a timer, whereas pressing
Lever B stops it, and reinforcement is contingent on the A± B interval exceeding some
criterion value. T his critical A± B interval is separated by the non-critical B± A interval, so
that the latter can act as a buffer ``absorbing’’ various behaviours or repetitive lever
presses that are, as in DRL, a major impediment to using DRL -type schedules as an
assay of ``pure’’ timing.
Using the two-lever DRL procedure, Sof® e and Lejeune (1991) compared aged (24-
and 25-month-old) and mature (7-month-old) Wistar male rats on acquisition and long-
term retention of a 5- sec time requirement between successive presses on Levers A and
B. T heir data showed that, relative to adults, senescent rats were slower to reach the
performance criterion, emitted fewer A± B sequences, and had longer B± A intervals.
However, B± B and A± A rates were equivalent at both ages, ruling out an explanation
of age effects in terms of motor impairment or hypoactivity. T he quality of response
timing was estimated by the median and the cv of the distribution of A± B intervals. No
age-related impairment in timing could be found, except for an increase of the cv during
the ® rst 10 minutes of the ® ve training sessions performed after the 5- sec criterion was
reached, which was interpreted as a temporary disruption of the ability to remember the
temporal criterion. However, long-term retention after a 21-day gap was not impaired in
either age group. No increase in the cv, as obtained in senescent rats over the initial
training sessions, was found.
T hese results were similar to those obtained on FI or DRL schedules, where response
timing in senescent, adult, and weanling rats did not differ at the end of a massed training
procedure (L ejeune, 1989; Lejeune, Jasselette, Nagy, & PereÂe, 1986). However, long-term
retention of FI (after 2 weeks) showed that senescent rats were temporarily unable to
suppress their responses during the ® rst segments of the FI (see also Campbell & Har-
outunian, 1981). Overall, these results indicate that the ability to form an accurate repre-
sentation of time, as well as the ability to translate such a representation into response
timing, was well preserved in aged rats.
T he only previous clear evidence for an age- related timing de® cit has been provided
by Meck, Church, and Wenk (1986) using the PI procedure. Mature (10± 13 month old)
rats were trained every other month on a PI 20- sec schedule, and they exhibited a shift of
peak time towards durations exceeding the reinforced time by about 5 sec when they were
27± 30 months old, whereas peak time had matched reinforcement time nearly perfectly
when the rats were 20 months old. T he 27± 30-month-old rats also showed a signi® cant
increase in the variability of the response rate function. Meck et al. do not provide
quantitative data on this variability increase, but the fact that a model based on SET
 PEAK PROCEDURE IN AGED RATS 197

® tted data well at all ages implies that the increase in variablity was scalarÐ that is, the
variability in the older rats was greater only because their mean peak time was greater.
T he overall effect of ageing on PI performance was interpreted in terms of an age- related
increase in the rememberd time of reinforcement (i.e. an increase in the memory constant
K*), which, according to some pharmacological manipulations used in the same study,
was probably due to an alteration of the cholinergic activity in the frontal cortex.
Why was there an effect of ageing on behaviour under the PI procedure, when there
was no effect on two-lever DRL in Sof® e and Lejeune’s (1991) study? As well as the
difference in the durations in the different conditions (5 vs. 20 sec), a further possibility is
that in two-lever DRL subjects were highly attentive to time on each trial, because they
themselves initiated trials by pressing L ever A, as well as controlling other events, such as
the intervals between emission of the critical A± B responses. On the other hand, trials and
the duration of the intervals between them are automatically controlled on PI, so attention
lapses might thus be much more frequent and reponsible for the apparent ``memory’ ’
de® cits of old animals. Indeed, it has been suggested that attentional defects might explain
age-related impairments both in humans (Craik, 1977) and in animals (Jones, Barnes,
Kirby, & Higgins, 1995; review in Kubanis & Zornetzer, 1981).
T he present experiment compares old (24 months) and young (4 months) adult rats on
the PI procedure. One of our aims was to replicate the results of Meck et al. (1986), and
another was to investigate whether timing behaviour in old rats was congruent with
detailed predictions of the scalar model about sources of variance operating in the dif-
ferent information processing stages. Some recent work (Cheng & Westwood, 1993;
Church et al., 1994; Gibbon & Church, 1990) has suggested that measures of performance
on individual trials of the PI procedure might be particularly diagnostic of the contribu-
tion to behaviour made by the different underlying parts of the timing mechanism posited
by SET. On most peak trials of the PI session, the behaviour of both rats and pigeons can
be characterized in terms of a 3- state process. For some time after trial onset, the animal
either pauses or responds only very sporadically. At some later time (the start time) the
animal begins to respond at a high rate, and this high rate continues for a period of time
(the spread), only to be followed at some later time (the stop time) by a return to zero or
low-rate responding for the remainder of the trial. As will be seen later, not only can the
start, stop, and spread values be related to both possible age effects and effects of the time
of reinforcement, but correlations between these measures can provide quantitative
insight into different underlying sources of variance.
As data obtained from aged animals and humans often reveal a reduced ¯ exibility or
ability to adapt quickly to changing conditions (Stephens, Weidmann, Quartermain &
Sarter, 1985; Reese & Rodeheaver, 1985), the ® nal aim of the study was to test temporal
¯ exibility by shifting rats from a 20- to a 40- sec reinforcement time, then back again to 20
and ® nally 40- sec, in an ABAB design.
For all the data, two sets of analyses were conducted. At the macrolevel, session-by-
session data were computed (such as peak time, peak rate, etc.). At the microlevel, within-
session data were analysed using results derived from individual peak trials. T he ® rst set
of data yielded information about the learning process, whereas the second allowed us to
investigate the mechanisms of the interval timer as described in the scalar model and its
information processing implementation (Gibbon et al., 1984).
198 LEJEUNE ET AL.

Method
Subjects
NaõÈ ve male Wistar-derived rats bred in the laboratory colony were kept on a 12/ 12 light/ dark
cycle (lights on: 7.30 am) and a temperature of 21 6 18 C. Senescent 24- month-old (n = 10) and 4-
month- old young (n = 10) adults were used. For convenience, the young adults are labelled ``young’ ’
rats. Ten days before the beginning of the experiment, all rats were progressively food-deprived by
reducing their daily food supply. T hey were kept at about 85% of free-feeding body weights
throughout the experiment.

Apparatus
T he subjects were tested in 20 identical experimental cubicles (40 3 40 3 45 cm). T he sides were
transparent Plexiglas. T he bottom and the top were made of aluminium grid with square perforations
(1 3 1 cm). A pellet dispenser delivered 45-mg Noyes food pellets through an opening located on the
left front wall, 6 cm over the ¯ oor and 6 cm from the front panel. A retractable lever requiring a 20- g
force was located 10 cm to the right of the food cubicle, 5 cm above the ¯ oor. No water was available
in the experimental cubicle. Sounds (40 dB and between 3800 and 4200 Hz, depending on the
cubicle) could be delivered by a loudspeaker ® xed above the top of each cubicle. T hese sounds
were within the range of those well perceived by rats (Kelly & Masterton, 1977). Each experimental
cubicle was located in a larger ventilated, heated (20± 218 C), lighted (25-W bulb), and sound- isolated
enclosure (70 3 60 3 160 cm), which ensured that the sounds could not be heard from one cubicle to
the next. Each of these enclosures had a small observation window (22 3 22 cm) located on the front
door. Monitoring of the experiment and recording of the data were controlled by IBM compatible PC
computers.

Procedure
Each subject received two peak procedure sessions every day, at approximately the same time each
day and in the same operant chamber. From 9 a.m. until the end of the second daily session, rats
working in the same laboratory room were group-caged. T he ® rst session of the day started at about 1
p. m. After the end of this session, the subjects were returned to the ``group’ ’ cages for a 30-min
recess, during which they could drink freely (no water was available in the experimental cubicles,
preliminary observations having shown that some subjects were prone to develop polydipsia). At the
end of the between-session break, a second session identical to the ® rst one was run. Session
durations ranged between about 90 min and 2 hours, depending on the subject and the duration
criterion. Two relatively short sessions separated by a pause during which the animals could drink
were preferred to a long one that might have strained the old rats. Subjects were thereafter returned
to their quarters and separated in individual ``feeding’ ’ cages, where they received supplementary
food about 30 min after the end of the second daily session. T he subjects remained in the ``feeding’ ’
cages until the next morning before returning to the bigger ``group’’ cages.
Preliminary to peak procedure training, each rat was hand-shaped with a sound (60 dB, 4000 Hz)
permanently on and the response lever permanently inserted into the experimental cubicle. After
magazine training and successful hand shaping, the subjects were switched to an automatic discrete-
trial Continuous Reinforcement schedule (CRF) and allowed to eat 25 more food pellets. For this
session, the sound was brie¯ y interrupted after the delivery of each reinforcer while the lever was
retracted and immediately inserted again (the complete cycle lasted about 2 sec). On the second day,
two CRF sessions limited to 25 reinforcers each were run, with an interval between them of about
 PEAK PROCEDURE IN AGED RATS 199

30 min. T he ® rst of these sessions was identical to the previous one. T he second featured a 6-sec IT I
during which the lever was retracted and the sound switched off. On the third day, two sessions were
run, separated by at least 30 min. During the ® rst one, the IT I between successive reinforcement
opportunities was increased to 20 sec. A discrete-trial FI 10 sec came next, with the session limited to
25 reinforcers. From this session on and until the end of the complete experiment, IT Is lasted 30 sec
on average (range 15± 45 sec) and inter-session intervals were kept at about 30 minutes. On the
fourth day, an FI 10-sec session was followed by the ® rst discrete-trial FI 20-sec session, both limited
to 90 reinforcers.
A series of 14 discrete-trial FI 20-sec sessions then followed, before the PI procedure began.
T hese PI sessions were limited to 90 trials, 30% of which were non- reinforced peak trials (n = 27).
During the PI sessions, reinforcement was available according to the FI 20-sec schedule on rein-
forced trials (the whole procedure being referred to as PI 20 sec). 30 PI 20-sec sessions were run
(Phase 1) before the subjects were abruptly switched to a PI 40-sec schedule (Phase 2) for 20 sessions
limited to 80 trials each and with 30% of the trials in the session being non-reinforced peak trials (n =
24). With the 20-sec criterion, the duration of peak intervals was on average equal to 60 sec (range
55± 65 sec); they lasted on average for 85 sec (range: 80± 90 sec) in the case of the 40- sec criterion.
Replication sessions came next. Ten PI 20-sec sessions (Phase 3) were followed by 10 PI 40-sec
sessions (Phase 4). In all cases, sessions took place 7 days a week. Whatever the criterion, peak trials
were semi-randomly intermixed within FI trials. T he only restriction was that no more than 3 peak
trials could occur in succession.

Behavioural Measures and Data Analyses

During peak procedure sessions, responses were recorded in successive 1-sec segments. Measures
were obtained for each session and each rat. Data from the ® rst trial of each session were dropped. All
remaining trials were taken into account for data analysis.

Macroanalysis. T hree measures were derived from data obtained on peak trials: peak time, peak
rate, and the cv of the response- rate distribution. In every case, an interval equal to 2t was taken into
account for the computation of behavioural measuresÐ that is, 40 and 80 sec in the case of a 20- and
40-sec criterion, respectively.
T he peak times of the response rate functions were medians obtained by iteration from the
response rate versus elapsed time in the interval functions. First, a median was obtained from the
® rst 40 or 80 sec of the peak trial (depending on the time of reinforcement during the FI trials of
the PI session). A new median was then computed, taking the ® rst median as the centre. For
instance, if the ® rst median obtained from the 40-sec interval was 18 sec, the second median took
a 36-sec interval into account. If the second median was located within 6 0.5 sec of the ® rst
median, computation stopped. If not, a third median was computed from the interval de® ned by
the second median, and so on.
Peak rate was de® ned as the response rate at peak time, computed by linear interpolation between
the centres of the two nearest segments. For instance, if peak time was equal to 18 sec, and if the
segments centred at 17.5 and 18.5 sec had response rates equal, respectively, to 45 and 50 responses
per minute, then peak rate would be equal to 47.5 responses per minute, i.e. ( .5 3 45) + ( .5 3 50)
(cf. Roberts, 1981, p. 244).
T he cv was obtained by computing the ratio between the standard deviation and the mean of each
response rate function. Here, a low cv value re¯ ects a response rate distribution that is narrowly
spread around its mean.
200 LEJEUNE ET AL.

PI data were analysed by two-way ANOVAs with repeated measures. Peak rate, peak time, and cv
data were pooled into 2-sesion blocks before statistical analysis.

Microanalysis. A more re® ned analysis focused on data from individual peak trials. Responding
in these peak trials generally followed a ``break± run± break’ ’ patternÐ that is, successive low± high±
low response rates. Four measures were computed from these individual peak trials, as de® ned in
Church et al. (1994): the time at which the rat started high-rate responding (S 1 ), the time at which
responding stopped (S 2 ), the spread of the interval between S 1 and S 2 (d = S 2 2 S 1 ) and the middle of
the spread located halfway between the two within-trial changes in response rate [m = (S 1 + S 2 )/ 2].
S 1 and S 2 were obtained by an exhaustive search of possible locations within the peak trial, so as to
maximize differences between high and low response rates, weighted by the relative time spent in the
low or high response state. To locate S 1 and S 2 , an index was computed on each individual peak trial.
T his index was t1 (r 2 r1 ) + t2 (r2 2 r) + t3 (r 2 r3 ), where t1 , t2 , and t3 were times from the beginning
of the peak trial until S 1 , the time between S 1 and S 2 , and the time from S 2 until the end of the trial,
respectively; r1 , r2 , and r3 were mean response rates during these time periods, and r was the overall
mean response rate on that trial. T he index was computed with S 1 at the time of each response except
the last one, and S 2 at each subsequent response (r1 includes the responses at S 1 , r2 included those at
S 2 and r3 included a hyopthetical response at the end of the trial). For each trial, S 1 and S 2 were the
times at which this index was maximized, thus effectively dividing the interval into three distinct
parts (low response rate± high rate± low rate) in the clearest way possible.
S 1 can be considered as similar to a ``breakpoint’ ’ as de® ned by Schneider (1969) and S 2 to the
``giving-up time’ ’ described in studies on foraging (Kacelnik, Brunner, & Gibbon, 1990). Data on
starts and stops were taken from all trials of the experimental sessions, without the selection used by
Church et al. (1994), who rejected trials where S 1 was higher than the temporal criterion (``bad
starts’’ ) and S 2 lower than the temporal criterion or higher than the shorter of two scheduled trial
durations (``bad stops’ ’).
Interest in within-interval patterns of responding on the PI procedure was stimulated by the
possibility that correlations between measures derived from such patterns would be diagnostic of
underlying sources of variance. Gibbon and Church (1990) ® rst demonstrated this, but other rele-
vant work includes Cheng and Westwood (1993) with pigeons, Church et al. (1994) with rats, and
Rakitin et al. (1998) with humans. As mentioned earlier, the SET analysis of PI trials assumes that
subjects are comparing the absolute difference between the number of pulses in the accumulator (n),
which continuously changes as time in the interval passes, with n*, a sample derived from the
reference memory of the time of reinforcement, t. T his difference (divided by n*) is compared
with a threshold, b, and when the difference falls below the threshold (i.e. n and n* are ``close
enough’ ’), the response state is initiated. As n continues to increase with increasing elapsed time
the absolute difference eventually exceeds b again, and responding ceases. Among potential sources
of variance in the model are variance in clock speed, variance in the memory representation of t,
variance in the threshold, b, and all these can vary from one trial to another (Gibbon & Church, 1990).
Particular theoretical interest has focused on the contributions of variance either in reinforcement
memory, in the threshold, or both. Suppose that the only source of variance in the model is memory
variance, with the threshold being constant. In this extreme case, the start time will vary from trial to
trial, but the start and stop times will be perfectly correlated, producing a constant response period
(i.e. constant spread). Furthermore, the position of the middle point of the response period will vary
from one trial to the next. At the other extreme, suppose that the memory representation of the time
of reinforcement is invariant, and variance exists only in the threshold. Now, the start and stop times,
and start and spread values, will be negatively correlated, but all other correlations between measures
will be zero. Between these two extreme cases are more plausible ones where both memory and
threshold variance is present, but to varying degrees (e.g. high memory variance± low threshold
 PEAK PROCEDURE IN AGED RATS 201

variance, low memory variance± high threshold variance, etc.), and Gibbon and Church (1990)
provide predictions about a wide range of such cases. A range of different correlation patterns is
possible with the different variance contributions. For example, positive correlations between start
and stop times indicate variance in memory of the time of reinforcement, or clock speed, whereas
negative correlations between the start time and the length of the response period (the spread)
indicate that the threshold for initiating and ceasing responding on the trial also varies from trial
to trial. Previous data from studies with rats and pigeons (Cheng & Westwood, 1993; Church et al.,
1994; Gibbon & Church, 1990) indicate that both sources of variance are present in peak procedure
performance.
In general, we used analyses derived from Gibbon and Church’ s work to attempt to discover the
underlying sources of variance present in our data (and to investigate whether these sources changed
with age), as well as to address other questions, such as whether the animals are using two thresholds
(e.g. a different threshold for starting and stopping responding) or only one (see Church et al., 1994,
p. 153, for the method of testing this).
Finally, start, stop, middle, and spread values from individual rats were taken to compute medians
for each successive peak trial in the session. T hese median points were then smoothed to obtain a
graph describing the trial-by-trial evolution of performance. T he smoothing process used a 5R
® lterÐ that is, a running median of 5, which repeatedly smoothed the data until resmoothing no
longer yielded further changes (see Tryon, 1983, for examples of the use of this method with
behavioural data).

Results
Peak Procedure Training

Macroanalysis. Average peak rate, peak time, and cv values from the successive
phases of the experiment are displayed in Figure 1. Peak rates from senescent rats (shown
at the top of Figure 1) were signi® cantly lower than those from the young subjects in each
phase of the experiment, except Phase 1Ð Phases 1 to 4: F(1, 18) = 3.74, p = .07; F(1, 18)
= 4.88, p < .05; F(1, 18) = 9.39, p < .01; F(1, 18) = 7.94, p < .02, respectively. T he
effects of the session blocks was limited to Phase 1Ð Phases 1 to 4: F(14, 252) = 7.67, p <
.0001; F(9, 162) = 1.44, p = .17; F(4, 72) = .75, p = .56; F(4, 72) = .50, p = .74,
respectively. Peak rates increased progressively in both groups before stabilization, which
appeared earlier for old subjects. At the ® rst transition from a 20-sec to a 40- sec time of
reinforcement, peak rates from old rats tended to drop, whereas those of adult subjects
displayed no clear-cut change, as can be seen in Figure 1. Interactions between age and
session blocks were not signi® cantÐ Phases 1 to 4: F(14, 252) = 1.42, p = .14; F(9, 162) =
1.01, p = .43; F(4, 72) = .50, p = .73; F(4, 72) = 1.40, p = .24, respectively. Further
transitions did not produce any ¯ uctuations of peak rates, whatever the age of the
subjects.
T he middle panel of Figure 1 shows that peak times from rats of both ages were close
to the criterion value over the ® rst phase of the experiment. Indeed, neither age, F(1, 18)
= 1.13, p = .30, session blocks, F(14, 252) = .92, p = .53, nor interaction effects, F(9, 162)
= .81, p = .65, were found for this phase. From PI 40- sec on, however, some age- related
effects were present. In particular, the old rats produced peak times that were higher than
the criterion time, whereas peak times from adults remained accurate. A signi® cant effect
FIG. 1. From top to bottom, mean peak rates, peak times and coef® cients of variation of young and old rats on
20-sec (Phases 1 and 3) and 40-sec (Phases 2 and 4) PI procedures. Successive phases of the experiment are
separated by vertical lines. Abscissa: successive blocks of 2 sessions.

202
 PEAK PROCEDURE IN AGED RATS 203

of subject age on peak time was obtained in Phases 2, 3, and 4Ð (F(1, 18) = 13.8, p <
.002; F(1, 18) = 9.24, p < .01; F(1, 18) = 16.22, p < .001, respectively. T he overshoot of
criterion time by peak time at 40 sec (phases 2 and 4) was about twice as great as the
overshoot at 20 sec in the senescent rats (on average, mean peak times computed over the
4 last session blocks at Phases 1 to 4 were 21.23, 41.23, 20.70, and 40.92 sec for young rats
and 22.46, 47.17, 23.41, and 46.25 sec for old animals, respectively). After transition to a
new criterion value, peak time continued to increase or decrease for a few sessions, before
a new baseline value was reached. Signi® cant session block effects were found in Phases 2,
3, and 4Ð Phases 1 to 4: F(14, 252) = .92, p = .53; F(9, 162) = 12.71, p < .0001; F(4, 72)
= 21.49, p < .0001; F(4, 72) = 12.46, p < .0001, respectively. Furthermore, during the
® rst transition from 20 to 40 sec, old rats needed more sessions before stable performance
was reachedÐ signi® cant Age 3 Session Block interaction, F(9, 162) = 5.25, p < .001.
Other interactions were not signi® cantÐ Phases 1, 3, and 4: F(14, 252) = .81, p = .65;
F(4, 72) = .68, p = .61; F(4, 72) = 1.83, p = .13, respectively.
Average cvs are displayed at the bottom of Figure 1. T hey were not affected by ageÐ
Phases 1 to 4, respectively: F(1, 18) = .05, p = .82; F(1, 18) = 1.39, p = .25; F(1, 18) =
1.80, p = .20; F(1, 18) = .00, p = .99Ð but, as a consequence of continued training,
decreased progressively over the successive two-session blocks of Phases 1, 2, and 4Ð
Phases 1 to 4: F(14, 252) = 15.70, p < .0001; F(9, 162) = 19.06, p < .0001; F(4, 72) = .89,
p = .48; F(4, 72) = 15.43, p < .0001, respectively. Local increases in cv, if any, were found
only during the ® rst sessions after transition from one criterion time to the next. Inter-
actions between session block and age were signi® cant for Phases 1 and 2Ð Phases 1 to 4:
F(4, 252) = 2.88, p < .001; F(9, 162) = 2.63, p < .01; F(4, 72) = 1.53, p = .20; F(4, 72) =
1.29, p = .28, respectively. As can be seen in Figure 1, average cvs of old rats were in some
instances lower than those of the young controls. However, this non-signi® cant effect was
due to three out of the ten old subjects having much lower cvs than the other rats of the
group. Median data would have shown that at the end of training at each criterion value
(over the last 5 sessions of Phases 3 and 4), the cvs of old rats were equal to or higher than
those of the younger subjects.
Scalar properties of timing were investigated by plotting response rate functions on a
relative scale, with maximum response rate and reinforcement criterion, t, equal to 1Ð
that is, all elapsed time in interval values were divided by the criterion time. According to
SET, accuracy and sensitivity to time should remain constant, whatever the value of t. If,
for instance, the FI duration increased from 10 to 20 sec, peak time as well as the spread
of the response rate function should be rescaled accordinglyÐ that is, their absolute value
should be multiplied by 2, which means that the relative values should remain unchanged.
Accordingly, response rate functions can be superimposed when peak rate and peak time
are expressed in relative terms. Data were taken from the last 5 sessions of Phases 3 and 4,
and the top of Figure 2 displays the superimposition of response rate functions in young
(left) and old (right) subjects for the 20- and 40-sec reinforcement criterion times. T he
smoother appearance of the function at 40 sec is probably due to the fact that each bin was
derived from about twice as much data as at 20 sec. Inspection of the response rate
functions suggests that there were no striking differences in the shape of the function
at the 20- and 40- sec reinforcement times in the old or younger rats. Furthermore, data
from the 20- and 40- sec reinforcement times superimposed well.
204 LEJEUNE ET AL.

FIG. 2. Response rate functions of young and old rats, plotted on relative time scales with peak rate (ordinate)
and criterion time (abscissa) equal to 1. Top: Performance of young (left) and old rats (right) at PI 20- and 40- sec
(within-age comparisons). Bottom: between- age comparison at the 20- sec (left) and 40-sec (right) criteria.
Arrows point to the respective values of peak time.

Overall, superposition testing, as well as cv data, suggested that the behaviour of both
young and old rats exhibited the scalar property (i.e., standard deviations of time esti-
mates were a constant fraction of the mean, leading to a constant cv) and that mean
accuracy held up reasonably well, as peak times were close to the reinforcement times.
T he gradual decline in the values of the cv over the course of training is probably just a
consequence of the overtraining (to be discussed later) due to the ABAB experimetnal
 PEAK PROCEDURE IN AGED RATS 205

design. T his decline does not violate SET because this model ® ts performance collected
when different groups of animals (and not only one as here) are subjected to a similar
amount of training, each at a different value of t. SET makes no predictions about the
absolute value the cv should take. However, comparing the performance of old and young
rats at the same reinforcement criterion value showed that the peak times of old rats were
relatively shifted to the right, as can be seen in the bottom of Figure 2 and in the middle
part of Figure 1.

Microanalysis. Group median Start (S 1 ), Stop (S 2 ), middle (m), and spread (d) values
from the individual peak trials are displayed in Figure 3. T he leftmost part of the ® gure
presents the data points from the last 20 sessions of the ® rst 20-sec PI training. T he other
panels display the complete data from the other phases of the experiment. As can be seen,
the within-trial responding of old rats started and stopped later, compared to that of the
younger rats. Consequently, their middle values were also shifted to the right. However,
the spread of responding was similar at both ages, as data points matched perfectly. T hese
features were statistically con® rmed, as t-tests computed over the data points from the last
5 sessions of each phase of the experiment yielded signi® cant between-age differences for
the S 1 , S 2 , and m variables. Age- related differences in spread values (d) were never
signi® cant, as can be seen from Table 1, which presents group means, standard deviations,
and coef® cients of variation for each dependent variable. Comparison of Table 1 with the
same measures obtained on various PI schedules in another research (15, 30, and 60 sec,
Church et al., 1994, Table 1, p. 144) highlights some similarities between our data and
those collected previously. For example, in both our data and those of Church et al., cv
values of S 1 were at least twice as high as those obtained for S 2 , m, and d. Some other
points of comparison between our data and those of Church et al. (1994) are discussed
further.
Correlations between start, stop, middle, and spread were computed using data from
the last 5 sessions at Phases 3 (20 sec) and 4 (40 sec) of the experiment. T hey are
displayed in Table 2. As mentioned earlier, according to models of PI performance that
follow S ET principles (Church et al., 1994; Gibbon & Church, 1990), such correlations
are diagnostic of the sources of variance underlying performance on the PI task. Correla-
tion values obtained by Gibbon and Church (1990), Cheng and Westwood (1993), Church
et al. (1994), and Rakitin et al. (in press) are given for comparison. All our correlations but
one (between middle and spread) showed the same positive or negative pattern as the
other reports of data from animals, although in some respects, such as the high S 1 ± S 2
correlation that we found, our data were more similar to those obtained from the human
analogue of the PI task employed by Rakitin et al. (in press) than previous data from
animals. How these patterns of correlations might come about will be discussed further.
T he behaviour of our rats was thus generally congruent with the pattern of correla-
tions encountered when behaviour ® ts the prescriptions of scalar timing about the sources
of variance modulating the memory and decision levels of the temporal information
processor. Most important, T able 2 also showed no age- related differences: whatever
pattern of correlations was found in the old rats was also found in the younger ones.
A measure speci® cally designed to test the one- versus two-thresholds hypothesis was
proposed by Church et al. (1994). If only one response threshold is used, a plot of the
FIG. 3. From top to bottom, start, stop, middle, and spread values (in seconds, ordinate) on successive peak
trials (abscissa). Each data point is the group median from young or old rats. T he left column displays data from
the last 20 sessions of the ® rst Phase of the experiment. T he other columns present data from the complete
Phases 2, 3, and 4. T he signi® cance of between- age differences is given for each behavioural index and each
Phase of the experiment (** p < .01; *** p < .001; **** p < .0001; NS: nsÐ nonsigni® cant).

206
 TABLE 1
Mean, Standard Deviation, and Coef® cient of Variation of Start, Stop, Middle, and Spread of Individual Peak
Trials, as a Function of the Age of the Subjects

Start (s1) Stop (s2) Middle (m) Spread (d)

M SD CV M SD CV M SD CV M SD CV

20A Y 11.45 8.68 .76 32.51 7.56 .23 21.98 7.58 .34 21.06 5.91 .28
O 15.72 10.52 .67 37.21 9.9 .27 26.47 9.58 .36 21.49 7.08 .33
p < .01 < .009 < .005 = .73

40A Y 25.92 14.3 .55 58.79 11.18 .19 42.35 11.63 .27 32.87 10.88 .33
O 31.91 15.13 .47 64.62 14.05 .22 48.27 13.47 .30 32.71 11.27 .34
p < .01 < .0002 < .0001 = .50

20B Y 12.34 8.15 .66 31.82 7.07 .22 22.09 7.02 .32 19.48 5.99 .31
O 17.29 10.96 .63 37.74 9.17 .24 27.52 9.54 .35 20.45 6.66 .33
p < .006 < .00006 < .0002 = .94

40B Y 29.17 13.53 .46 59.07 10.81 .18 44.12 11.44 .26 29.9 8.76 .29
O 34.47 16.25 .47 63.97 14.02 .22 49.02 14.27 .29 29.1 10.35 .36
p < .009 < .0005 < .0007 = .65

M = mean; SD = standard deviation; CV = coef® cient of variation; s1 = start; s2 = stop; m = middle; d = spread;
Y = young; O = old.
Note: From top to bottom, data from the last 5 sessions of the successive phases of the experiment (20A, 40A, 20B and
40B). T he signi® cance (p level) of between- age differences is provided for each experimental condition.

207
208 LEJEUNE ET AL.

TABLE 2
Correlations Between S1 (start), S 2 (stop), m (middle) and
d (spread) from the Present Study and from Other Work

Correl atio n v alu e

D ata S1 ± S2 S1 ± m S 1± d S 2± m S2 ± d m± d

Y(20 sec) .67 .92 2 .44 .91 .35 2 .05

Y(40 sec) .73 .95 2 .57 .92 .13 2 .27
O(20 sec) .80 .95 2 .42 .95 .19 2 .13
O(40 sec) .78 .94 2 .49 .95 .15 2 .21
G&C90 .34 .74 2 .34 .89 .77 .38
C&W93 .38 .77 2 .36 .88 .71 .31
C&W93S IM .54 .88 2 .51 .87 .45 2 .04
CMG94 .31 .70 2 .33 .89 .78 .41
RETAL(1) .87 2 .13 .11
RETAL(3) .85 .02 .23

Note: Data from the present study come from young (Y) and old (O)
rats at the 20- and 40- sec criterion times. Other studies shown are
Gibbon and Church (1990: G&C90), Cheng and Westwood (1993:
C &W 9 3 ) , Ch en g an d We s tw ood G au s s ian s i mu l ati on ( 19 93 :
C&W93S IM), Church et al. (1994: CMG94), and Rakitin et al. [1998:
RETAL, Experiments 1 and 3 indicated by (1) and (3)]. Data from the
latter study were averaged over the 8, 12, and 21-sec criterion times.

correlation between the spread and the middle of the peak trials, r(d,m) against the ratio of
the coef® cients of variation of the middle and the spread, cv(m)/ cv(d), should fall on a
diagonal straight line. If two thresholds were used, points should fall below this line.
Indeed, a low positive or negative r(d, m) versus a positive cv(m)/ cv(d) can be obtained
only if the ``peak’ ’ of individual trials does not occur at the middle of the high rate
segmentÐ that is if different thresholds were used to start and stop responding. Figure
4 presents the appropriate data plot of old and young rats, computed over the last 5
sessions of Phases 3 (criterion 20 sec) and 4 (criterion 40 sec). As was the case in Church
et al. (1994, Figure 14, p. 150), all points of the scattergram were located below the
diagonal. Furthermore, differences between old and young rats at 20 and 40 sec were
never signi® cantÐ r(d, m) : t(18) = .93, p = .36 and t(18) = .74, p = .46, respectively; cv(m)/
cv(d): t(18) = .04, p = .96 and t(18) = .30, p = .76, respectively.

Discussion
T he experiment described above had three aims: ® rst, to try to replicate data described by
Meck et al. (1986); second, to verify whether performance of old rats was congruent with
the scalar timing model developed to explain the within-interval timing performance of
adult animals; and third, to test the hypothesis of a reduced ¯ exibility or adaptability of
old rats with regard to young controls. T hese issues are discussed in turn.
In Meck et al. (1986), rats trained at the age of 10 months on PI 20 sec and retrained
periodically up to the age of 30 months showed peak times shifted to the right when they
 PEAK PROCEDURE IN AGED RATS 209

FIG. 4. S cattergram of correlations between middle and spread r(d,m), as a function of the ratio between the
coef® cients of variation of the middle and the spread cv(m)/ cv(d) for young and old rats on PI 20 and 40 sec.

were 30 months old. T his shift amounted to about 21% of the value produced when the
same rats were young (from about 20.7 to 25 sec). T he effect was interpreted as an age-
related alteration in the content of reference memoryÐ that is, as an alteration of the
memory constant K* associated with an age-related alteration of cholinergic activity in
neurons of the frontal cortex. Psychologically, this interpretation means that the time of
reinforcement was systematically misremembered as being longer than it really was.
Another effect was that the variability of the function relating response probability to
elapsed time in the interval was signi® cantly increased with age. However, this increase
remained congruent with the scalar property and could be accounted for by the proposed
change in K*.
T he performance pattern obtained in our study con® rmed these results. At the 20- and
40-sec reinforcement times, 24-month-old rats displayed a signi® cant increase in esti-
mated time of reinforcement availability, as was the case in Meck et al. (1986), when peak
time values were calculated. For example, compared with the young controls, old rats
increased their peak times by about 15% at 20 and 40 sec (Figure 1). T he increase
appeared to be proportional to the time of reinforcement (i.e. about twice as large at
40 sec as at 20 sec). In addition, as in Meck et al. (1986), the rightward shift of peak time
was not accompanied by any non-scalar change in the variability of the response rate
function, as the cvs were not signi® cantly different in the old and younger rats, even when
210 LEJEUNE ET AL.

peak times were different. Such an apparently proportional shift in peak times is con-
sistent with Meck et al.’ s suggestion that the older rats remembered the time of reinfor-
cement as longer than it actually was; in the framework of SET, this is represented by an
increase in the memory constant, K*.
T he middles (Table 1 and Figure 3) calculated from the within-trial patterns of
responding likewise showed a signi® cant rightward shift in the old rats compared with
controls, but this time the shift appeared to be an absolute, rather than proportional
displacement. Indeed, shifts computed from middle values were, on average, equal to 7
and 8.65 sec at 20- and 40-sec peak trials, respectively, whereas they were equal to 2.93
and 6.72 sec when computed from peak time data (Figure 1). S uch an effect is not
consistent with a change in K*, which would be multiplicative, but with other possibi-
lities, particularly a difference in latency to close and open the switch connecting the
pacemaker and accumulator (i.e. to start and stop timing) between the old and young rats.
Suppose, for example, that the old rats start and stop timing later on FI trials than do the
younger rats. Such an effect would displace the middles to the right and would also
produce a similar displacement of start and stop values, as observed in Table 1.
Such a change in latency to start and stop timing might be a consequence of attentional
de® cits consequent on ageing and, consistent with this idea, some apparent changes in
switch closure latencies in other sorts of studies with rats (e.g. Meck, 1984) have also been
interpreted in attentional terms. For example, when the onset of a light stimulus at the
start of a PI 30-sec trial was made unexpected by being preceded by a sound cue, peak
time was displaced to the right by about 1 sec compared with control conditions, as if the
latency to begin timing was delayed by 1 sec. T his explanation cannot, however, be
directly applied to our data, as the displacement appears to be much greater than that
previously obtained, and in addition there was no stimulus event to make trial onset
unexpected.
However, the notion of a general attentional impairment in old age was recently
supported by data recorded in ® ve- choice serial reaction time task (Jones et al., 1995).
According to this report, the attentional de® cit of old rats might be mediated by an age-
related change in cholinergic mechanisms. In fact, a similar biochemical change was
proposed by Meck et al. (1986) to support their hypothesis that reinforcement time
was misremembered in their old rats. In their study, a bioechemical treatment intended
to stimulate the cholinergic system in fact ``inoculated’’ a group of rats against the shift in
peak time that was otherwise observed with ageing. Finally, some recent data obtained
with 6-month-old rats on the PI procedure showed a rightward shift in peak time after the
administration of an a2 noradrenergic agonist, clonidine, which reduces brain norepi-
nephrine levels (Penney, Holder, & Meck, 1996). T hese data were interpreted as consis-
tent with an increase of the latency to start the temporal integration process at signal
onsetÐ in other words, with an increase in the switch closure latency. T his interpretation
is, of course, similar to that proposed here. However, further research is needed to
evaluate the implications of Penney et al.’ s results with regard to aged rats and to under-
stand further the interconnections between neurotransmitter systems subserving the
processing of time.
T he data from the average response rate functions and the measures calculated from
within-trial patterns of responding thus seem to be in contradiction as to whether the
 PEAK PROCEDURE IN AGED RATS 211

rightward displacement seen in the older rats was proportional (peak times) or absolute
(middles). Although at ® rst sight the middles might seem to be a more fundamental
measure than peak times, as they come from individual trials rather than aggregates of
many trials, in fact the two measures might be reasonably regarded as different. For
example, absolute response rates from one 1- sec bin to another affect the location of
the peak time, when it is calculated in the conventional manner (after Roberts, 1981) used
in our study, whereas the middles are calculated on the basis of start and stop times, and
local ¯ uctuations in response rate, if any, play no role. Although some other studies
present examples both of average response rate functions and measures from individual
trials (e.g. Church et al., 1994) only Rakitin et al. (1998) directly compare peak times from
response rate functions and middles quantitatively (their Tables 2 and 3). In their study,
like ours, peak times from averages and middle points were very close, but not exactly the
same. In Rakitin et al., for example, middles were always longer on average than peak
timesÐ between 2 and 4% longer, depending on the criterion time. For the purposes of
Rakitin et al.’ s article, these differences were unimportant, but if they had been important,
that study would also have encountered the dif® culty of deciding which measure of
central tendency of responding (peak time or middles) was the ``correct’’ one.
In general, measures of responding in the present study suggested that the behaviour
of both the young and old rats we used conformed well to the principles of scalar timing.
Response rate functions superimposed reasonably well at both ages, and the pattern of
correlations obtained from within-trial measures (Table 2) resembled that obtained from
some previous animal studies in most respects. However, there were also some differences.
Our S 1 ± S 2 correlations were much higher than in other animal studies and approached
those obtained from humans by Rakitin et al. (1998Ð see Table 2). Furthermore, our
middle-spread correlations were negative, whereas those from previous animal studies
had been positive, and although the correlations obtained from these measures in Rakitin
et al. generally remained slightly positive on average, some individuals showed negative
middle-spread correlations as our rats did (see their Figure 5).
Another similarity between our rats and Rakitin et al.’ s humans is that the spread
values were insensitive to changes in reinforcement time, at least compared with Church
et al.’ s (1994) rats. For example, Church et al. doubled reinforcement time from 15 to
30 sec, then doubled it again (although the comparisons were all between different subject
groups). Spreads were from 81 to 95% of the reinforcement time ratios; in other words,
they changed markedly, and nearly exactly proportionally, with reinforcement time ratios.
Spreads from our rats and Rakitin et al.’ s humans also changed with changes in reinfor-
cement time but were more insensitive: for example, our spread changes were from 71%
to 78% of the reinforcement time change, and Rakitin et al.’ s humans showed similar
values (spread ratios from 72% to 83% of reinforcement time ratios). T hus both our rats
and Rakitin et al.’ s humans showed relatively constant spread values as the times of
reinforcement changed.
As mentioned earlier, the correlations between measures derived from single-trial
performance are diagnostic of the sources of variance underlying performanceÐ assum-
ing, of course, that the model of within-trial responding proposed by Gibbon and Church
(1990) is correct. Two sources of potential variance have been the focus of interest. T he
® rst is variance in the representation of duration that arises either from variance in the
212 LEJEUNE ET AL.

memory of the time of reinforcement (the FI value used on FI trials) or variance in clock
speed from one trial to the next. T he second source is threshold variance, variability from
one trial to another in the threshold value used to initiate or cease responding in the trial.
Near maximal start± stop correlations indicate that timing variability (arising from mem-
ory or clock variance) makes a much larger contribution than does threshold variance.
Values from both our young and old rats, at both times of reinforcement, were much
higher (averaging 0.75) than those from previous animal studies (Cheng & Westwood,
1993; Church et al., 1994; Gibbon & Church, 1990) which averaged 0.34, and were closer
to the 0.80+ correlation values obtained from humans by Rakitin et al. (1998). However,
the fact that our start± spread correlations were negative indicated that threshold varia-
bility still played a role. Our start± spread correlations were close to the normal values for
animal experiments, and much more negative than those obtained by Rakitin et al. (1998).
T he most unusual correlation values in our study were the slightly negative middle-
spread correlations, averaging 2 0.16. Other animal studies have found these to be
substantially positive (the average from the three studies quoted above was 0.37). In
Rakitin et al. (1998) middle± spread correlations were on average positive (although
with smaller positive values than in animal studies), but some individual subjects showed
negative values, as our rats did.
Negative middle± spread correlations have no ready interpretation within the original
model of within-trial responding proposed by Gibbon and Church (1990), where in
general measures of response variablity (like spread) are positively related to measures
of central tendency of responding, like middles. If subjects are producing a constant
response period on each trial, on the other hand, then middles and spread will be
uncorrelated, and random variability may produce small negative or positive correlations
in some data samples. However, small-valued middle-spread correlations (both slightly
positive and slightly negative) are consistent with a model proposed by Cheng and West-
wood (1993). T his essentially assumed that the tendency to respond rose and fell within
individual trials of a PI session according to a Gaussian function of elapsed time in the
trial. However, the word ``tendency’ ’ is emphasized above, as predictions about how actual
response rate would vary within a trial depend on rules as to how this rising and falling
function is translated into response rate. Obviously, a Gaussian function is a continuous
series of real numbers, whereas response rate in some time interval (e.g. a 1- sec bin) can
only have a few discrete integer values. T he fact that some translation of the Gaussian
function into responding is necessary means that the Gaussian model does not necessarily
predict that response rates rise and fall during the response period itself, in spite of a rise
and fall in underlying ``excitation’’ .
Inspection of results produced by Cheng and Westwood’s simulation (Cheng & West-
wood, 1993; see C&W93S IM in Table 2) shows that their Gaussian model produced
higher start± stop correlations than found in other data (including Cheng & Westwood’s
own), more negative start± spread correlations, and very slightly negative middle± spread
correlations, unlike the positive values found in all studies but our own. T his pattern
means that their model actually ® ts the correlation patterns shown by our rats better than
the values obtained in their pigeons, and it was perhaps the lack of correspondence
between their model and their data that resulted in the Gaussian model being only
sketchily described in Cheng and Westwood’s article.
 PEAK PROCEDURE IN AGED RATS 213

Cheng and Westwood’s model not only assumes a Gaussian pattern of response ten-
dency within a trial, but also supposes that the height of the peak of the Gaussian curve,
its peak location, and its standard deviation are determined independently on each trial.
In spite of the independence of these underlying values, some measures of responding
from analyses of single simulated trials exhibit substantial correlations. Using details
published in Cheng and Westwood (1993, e.g. p. 62) and further information provided
by Cheng (personal communication), we reconstructed Cheng and Westwood’s Gaussian
model as a computer program and explored its operation further. A detailed discussion of
this work is beyond the scope of the present article, but we will brie¯ y report a few results
here. First, we replicated the basic pattern of correlations that Cheng and Westwood
reported in their article and found that the pattern of strongly positive start± stop correla-
tions, strongly negative start± spread correlations, and low-valued middle± spread correla-
tions was readily obtained with many different parameter values. Second, by manipulating
parameters of their model, some of our correlation data could be simulated even more
closely than by Cheng and Westwood’s original parameter values. For example, one of our
simulations produced a start± stop correlation of 0.67, a start± spread correlation of 2 0.44,
and a middle± spread correlation of 2 0.03, values nearly identical to those found in our
young rats with a 20-sec reinforcement time (see Table 2). T hird, in spite of some success
with other correlation values, the substantially negative middle± spread correlations
obtained in most of our conditions remained elusive, and their explanation still poses a
considerable theoretical challenge. However, it may be that a model similar to that of
Cheng and Westwood (1993) can be developed further.
It is unclear why our correlation patterns were similar in some respects to those
obtained with humans and from Cheng and Westwood’s Gaussian model, but one spec-
ulative possibility is that the amount of training may change response patterns in some
subtle way. Our rats received considerably more training than did those of Church et al.
(1994), at least when measured in terms of the number of trials where reinforcement was
delivered. However, they also received fewer trials than Cheng and Westwood’s (1993)
pigeons, at least during the ® rst PI 20- sec phase. On the other hand, it may be that the
shift towards the particular correlation pattern we noted occurs with over-training relative
to the number of trials initially required to master the temporal discrimination. A recent
experiment with humans (Wearden et al., 1997) showed that even humans who were
elderly and of low general ability could acquire simple temporal discriminations to a
high degree of accuracy within a single-® gure number of trials with feedback, and other
studies show that even more complex procedures (e.g. Wearden, 1995) are mastered by
humans within a few tens of trials that provide relevant information. In Rakitin et al.’ s
(1998) Experiment 1, humans received in total more than 800 trials that either demon-
strated the criterion time of one of the three PI values in force or gave feedback as to
whether responding was accurate, and such a number of trials might, for humans, con-
stitute massive over-trainingÐ the equivalent of 20 or 50 times as many trials as were
needed to learn the criterion time values in the ® rst place. If pigeons learn more slowly
than rats, or are less temporally sensitive (Lejeune & Wearden, 1991), then even the large
number of reinforced trials given by Cheng and Westwood (1993) might constitute less
overtraining than our rats, and Rakitin et al.’ s humans, received. However, this explana-
tion remains speculative, and the differences between our correlation patterns and those
214 LEJEUNE ET AL.

of some other animal studies should not, perhaps, be over-interpreted. For example, not
only were the positivity and negativity of values usually in the same direction as other
studies (even though absolute correlation values might be different) but the analysis
intended to test the one- versus two-threshold model of responding yielded results
very similar to those of Church et al. (1994) when their SET- based model was applied.
T he ® nal aim of the experiment was to test any possible age- related differences in
adaptability to changing temporal criterion values. Indices derived from macroanalyses
showed that age- related differences were found only during the ® rst transition from the
20- to 40- sec reinforcement times. T hese results were in agreement with those from
spatial reversal studies reporting that old rats were slower than younger ones in learning
a reversal task, but that the age effect tended to vanish over subsequent reversal trials
(S tephens et al., 1985; Zornetzer, T hompson, & Rogers, 1982). T hus, as seen for spatial
learning, old rats rapidly acquired a strategy enabling them to cope with a new value of
the time of reinforcement.
Overall, our data show that timing of responding on the PI procedure, as well as the
ability to adjust to changing reinforcement times, is little impaired in old age in the rat.
T he only major difference between the old and younger rats, apart from slightly slower
adjustment to the ® rst reinforcement time transition (but not subsequent ones) in the old
rats, was that peak times and the middles of the within-trial response period were system-
atically displaced to the right in the old animals. Peak time locations suggested that this
was due to misremembering of reinforcement time, in agreement with Meck et al.’ s (1986)
conclusions, whereas values of middles were more consistent with a change in the latency
to start and stop timing in the older animals. As Meck et al. (1986) did not employ
individual-trial analysis, which did not come into general use until some years later, those
authors could not have examined this possibility in their data. Of course, contributions to
behaviour from both effects are possible. Older rats also tended to show more variability
in responding, although any change in variability was scalar (i.e. coef® cients of variation
were constant). Patterns of correlations from within-trial measures of behaviour showed
some features previously only observed in PI data from humans, for reasons that are
unclear but may be linked to the amount of training received.
Our rats, like the humans in Wearden et al. (1997), showed only small and subtle
differences in timing performance from that obtained in much younger subjects. T his
® nding suggests that the timing mechanism is largely independent of other age- related
behavioural changes, although there was some evidence that memory and/ or attentional
effects, possibly related to decreases in acetylcholine levels with age, may produce some
small age- related changes in an otherwise generally intact scalar timing mechanism.

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Manuscript receiv ed 21 July 1997

Accepted revision received 5 December 1997

Performance de rats jeunes et aÃge s en procedure du pic de

de bit de reÂponse (Peak Interval procedure, PI): Acquisition
et adaptation aÁ des changements du criteÁre temporel

Des rats aÃge s de 4 et 24 mois ont e te entraine s avec une proce dure PI dont le criteÁ re temporel
(le moment de distribution du renforcement) passait deux fois de 20 (A) aÁ 40 (B) secondes,
selon un plan ABAB. Le moment du pic de de bit de re ponse (peak time) des rats aÃge s a
systeÂmatiquement de passe le moment du renforcement, alors que celui des rats jeunes eÂtait
proche des dure es de 20 ou 40 secondes. Diverses mesures de performance suggeÁ rent 1) que
les rats aÃge s surestimaient le moment pre cis auquel le renforcement e tait distribue ou 2) que
les latences de de clenchement et d’ arreÃt de leur horloge interne e taient allonge es, par rapport
aÁ celles des jeunes animaux. Les rats aÃge s se sont adapte s plus lentement que les jeunes aÁ la
premieÁ re transition entre les criteÁres temporels de 20 et 40 secondes, mais cette diffeÂrence a
disparu lors des transitions ulteÂrieures. Les corre lations entre les indices de performance
de riveÂs des essais isoleÂs (peak trials) se conforment aux pre dictions de la the orie du temps
scalaire. Cependant, certaines d’ entre elles eÂtaient, pour des raisons non encore e lucide es,
plus proches de celles obtenues avec des sujets humains en proce dure PI et de celles de riveÂes
d’ un modeÁ le the orique de veloppe par Cheng et Westwood (1993), que de celles obtenues
anteÂrieurement chez l’ animal.
 PEAK PROCEDURE IN AGED RATS 217

ActuacioÂn de ratas joÂvenes y viejas bajo el procedimiento

de ``peak interval procedure’’ (PI): AdquisicioÂn y
adaptacioÂn a un criterio temporal variable

Se entreno a ratas de 4 y 24 meses de edad usando el procedimiento PI en el que el criterio

temporal (el momento de la distribucioÂn del reforzamiento) pasaba dos veces de 20 (A) a 40
(B) segundos, segu n un plan ABAB. El momento de ``peak time’ ’ de las ratas viejas depasaba
sistemaÂticamente el momento de renforzamiento, mientras que el de las ratas joÂvenes estaba
proÂximo de las duraciones de 20 o 40 segundos. Diversas medidas de actuacioÂn sugieren 1)
que las ratas viejas sobreestimaban el momento preciso en el que el renforzamiento era
administrado o 2) que las latencias de puesta en marcha y de parada de su reloj interno
estaban retrasados, en comparacioÂn con el de los animales maÂs joÂvenes. Las ratas viejas se
adaptaron maÂs lentamente que las joÂvenes a la primera transicio  n entre los criterios tempor-
ales de 20 y 40 segundos, pero esta diferencia desaparecõÂ a en las transiciones ulteriores. Las
correlaciones entre los Âõ ndices de actuacioÂn obtenidas de los ``peak trials’’ concuerdan con las
predicciones de la teorõÂ a escalar del tiempo. Sin embargo, algunas de esas correlaciones
estaban, por razones au n no dilucidas, maÂs proÂximas a las obtenidas con sujetos humanos
usando un procedimiento PI o a las derivadas del modelo teoÂrico desarrollado por Cheng y
Westwood (1993), que a las obtenidas anteriormente en el animal.