presents an illustration of the test apparatus.

An infrared light and detector were each placed

J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509 503
Fig. 1. Illustration of test apparatus. Two glass tubes with cross-bars formed a square aperture or “hoop” through which the fish swam. The
tubes contained an infrared light source and detector that detected movement through the hoop. A reflective mirror was mounted on a stand and
located outside the tank.
in individual 2.5 cm diameter glass tubes, separated
by three glass cross-bars: two at the base forming a
3.75 cm square aperture (called a “hoop”) and one at
the top stabilizing the tubes. The infrared light and detector
created a beam that extended across the middle
of the hoop. A mirror-box controlled delivery of
a reinforcer (a reflective mirror). The box contained
two 25W bulbs, arranged side-by-side, that back illuminated
a one-way mirror attached to the front.
The mirror measured 20 cm × 7.5 cm and was located
on the long side of the tank, centered horizontally
and 3.0 cm from the top of the tank. Turning
the light bulbs off and on made the mirror reflective
and transparent, respectively. An IBM-compatible
computer and a program written in MED-PC controlled
the experimental events and recorded breaks in the infrared
beam as the fish swam through the hoop. To
prevent recording one response more than once as
the fish swam through the hoop, the program waited
one second before detecting subsequent breaks in the
beam.
1.3. Procedure
The fish received training in preliminary study and
were already trained to swim through a hoop 1. We exposed
the fish to four FI schedules in the following
order: FI 30, 120, 60, and 240 s. A program made available
a reinforcer (20 s of a reflective mirror) for the first
response after the FI requirement elapsed since completion
of the previous reinforcer. Reinforcers were given
immediately after the fish broke the infrared beam. Sessions
consisted of 30 intervals and began with a 20 s
presentation of the reflective mirror, marking the start
of the first interval. Each condition was in effect for
five sessions.
1 Ina preliminary study, the fish were given training with a reflective
mirror until it consistently evoked the aggressive response.
Next, the hoop was placed into the tank, just below the water level.
The fish did not require specific training as they all readily swam
through the hoop for mirror presentation reinforcers.
504 J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509
2. Results and discussion
We analyzed the data and looked at several different
measures of timing behavior. In all cases we used data
from all intervals and all sessions for a total of 150
intervals for each condition, for each subject.
2.1. Cumulative records
Fig. 2 presents cumulative records for individual
subjects. Each record represents the mean time of a response
in an interval. In some places one response (n)
appears to occur at an earlier time on the x-axis than
the preceding response (n − 1). Arrows in Fig. 2 point
to examples of these instances. Backward excursions
in the cumulative record occur because the data point
represents an average of several responses. Hence, in
some cases the average time of say the fifth response
in an interval is shorter overall than the fourth response
(e.g., see record for LB06). The first noticeable result
is that some fish (LB01 and LB08) produced more responses
than others (LB06 and LB10). Still, in most
cases the fish responded at a higher rate towards the
end of the interval than at the beginning, as indicated
by changes in the slope of the records. Below, we report
estimates of the point at which response rates change in
an interval. Next, responding either increased gradually
throughout an interval resembling an FI scallop pattern
(e.g., LB08, FI 240 s) or remained high throughout an
interval (e.g., LB10, FI 30 s). When comparing cumulative
records across conditions, increases in the FI requirement
produced lower levels of responding during
the start of each interval. Similar patterns of responding
during FI schedules have been reported in other species
(e.g., Barnes and Keenan, 1993; Dews, 1970; Ferster
and Skinner, 1957; Zeiler and Powell, 1994) as well as
fish (e.g., Gee et al., 1994; Lejeune andWearden, 1991;
Rozin, 1965; Talton et al., 1999).
2.2. Wait time and break point
Fig. 3 (top) presents the meanwait time for each fish
and the group across the different FI schedules. Overall,
wait time increased with increases in the FI requirement.
For example, mean wait time for LB01 increased
from roughly 16 to 38 s across the FI schedules. A twoway
repeated measures analysis of variance (ANOVA)
comparing the main effect of the FI requirement and
sessions indicated that the main effect of sessions was
not significant, however, there were significant differences
across the FI requirement, F(3, 36) = 23.726,
p < 0.001 (alpha level set at 0.05). Pair-wise multiple
comparisons (Tukey) indicated significant differences
between the FI 240 and 30 s (q = 11.046, p < 0.001),
FI 60 s (q = 9.269, p < 0.001), and FI 120 s (q = 8.006,
p < 0.001) schedules. Other FI combinations were not
statistically significant.
Historically, there has been discussion over whether
the relation between independent temporal variables
and dependent measures such as wait time duration
is best described by a proportional (e.g., Shull,
1971,1979; Starr and Staddon, 1974; Staddon and
Cerutti, 2003) or power rule (e.g., Catania, 1970; Lowe
et al., 1979; Zeiler and Powell, 1994). Given no significant
differences in wait times among the smaller FI
values, it is not possible to determine whether a linear
or power function best describes the pattern of wait
times because there are essentially two data points: one
at FI 240 s and the other clustered at the smaller FI
schedules. Another method is to look at wait time as a
proportion of the FI value, given in Fig. 3 (bottom). The
figure shows a pattern in whichwait time is a decreasing
proportion of the FI value, as would be the case with a
power function. However, although a one-way repeated
measures ANOVA comparing relative wait time across
changes in the FI requirement was significant, F(3, 9)
= 26.169, p < 0.001, pair-wise comparisons show no
significant differences between FI 60 and 120 s, and FI
120 and 240 s.
It may be the case that wait times from fish are a
less sensitive measure of timing than other measures
such as break point. Talton et al. (1999) reported relatively
small changes in wait time during different FI
values, and found more systematic changes in break
point times. Following the method reported by Talton
et al. (1999),we estimated break points by first normalizing
the pattern of responding in an interval, both in
terms of time in an interval (x-axis) and the frequency
of responses (y-axis). To do this we calculated the mean
number of responses in bins of 1/10 the FI requirement,
and normalized the overall level of responding based on
the maximum generated in the interval. This was done
for individual subjects. An advantage of this procedure
is that it results in an equivalent number of points (10)
over which to calculate a break point. Next, to calculate
the break point, we used a linear regression analysis by
J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509 505
Fig. 2. Cumulative records for each subject and condition. The data are based on all intervals and sessions. Each data point represents the average
time of a response in an interval.
which each normalized response-curve was fitted with
two lines that intersected at bins 3–8 as possible break
points. For example, we calculated the R2 of a line
through bins 1–3 and a line through bins 3–10. In the
next iteration we calculated the R2 of a line through
bins 1–4 and 4–10, and so forth. The combination of
lines that maximized the average R2 was used as an
estimate of the break point; the value (i.e., bin number)
was then multiplied by the bin duration that was used
to generate the normalized curve for each condition so
that we could represent the break point in terms of the
actual time in an interval. The results of this analysis
506 J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509
Fig. 3. Mean wait time for individual subjects and the group as a
function of the FI requirement (top). The data are also plotted as a
proportion of the FI requirement (below).
are presented in Fig. 4. As in the Talton et al. (1999)
study, break point increased with increases in the FI
requirement, and ranged from about 10 to 138 s, F(3,
9) = 20.403, p < 0.001, with significant differences between
FI 30 s versus FI 240 s (q = 10.029, p < 0.001),
FI 60 s versus FI 240 s (q = 9.032, p < 0.001), and FI
120 s versus FI 240 s (q = 6.804, p < 0.004).
We next determined which of two functions best
described changes in break point by calculating bestfit
linear and power functions for individual animals as
well as the group mean.
The equations were:
f (x) = mx + b (1)
Fig. 4. Break point in an interval as a function of the FI requirement.
Shown are the results for individual subjects as well as the group
mean.