where x is the FI value and m, b, k, and n are parameters

whose values were based on the results of a program

in SigmaStat® that uses a Marquardt–Levenberg algorithm
to determine the parameter values that minimize
the sum of the squared differences between the data and
predictedwait time durations. The results are presented
in Table 1. Both functions fit the data from individual
subjects well, with R2 values ranging from 0.946 to
0.992 for a linear function and from 0.958 to 0.994 for
a power function. For a majority of fish (LB01, LB08,
and LB10) a power equation resulted in a higher R2
value, however the difference was not statistically significant.
Table 1
Parameter and R2 values from best-fit linear and power functions to
individual and group break point for the different FI reinforcement
schedules
Subject Linear Power
R2 m b R 2 n k
LB01 0.974 0.637 − 14.609 0.994 1.428 0.0569
LB06 0.992 0.905 − 26.087 0.990 1.375 0.103
LB08 0.955 0.276 5.739 0.958 0.819 0.798
LB10 0.946 0.655 − 21.913 0.992 1.805 0.007
Group 0.988 0.618 − 14.217 0.999 1.369 0.076
J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509 507
Summarizing, both wait time and break point increased
with increases in the FI value, with more
consistent changes in break point. Furthermore, the
break point pattern was well described by both a linear
and power function. Still, for three of four subjects,
the best-fit linear equation for the break points
resulted in a sizeable negative y-intercept, which underestimates
the results from the smaller FI schedules,
hence it may be the case that a power function
is a better description of the break point pattern. Interestingly,
Zeiler and Powell (1994) found similar results
in a study examining temporal performance of
pigeons tested using a peak procedure with FI values
ranging from 7.5 to 480 s. A possible explanation for
our results is that the FI values chosen did not produce
consistent and large enough changes in the pattern
of swimming responses. Specifically, we may be
seeing a floor effect in response latencies. Two fish,
LB06 and LB10, often producedwait times longer than
30 s during the FI 30 s schedule, which indicates that
the actual interval duration was longer than the programmed
requirement. It may be that larger FI values
(longer than 240 s) are necessary when studying interval
timing effects in Bettas responding by swimming
through a hoop for visual reinforcement. An
even wider range of intervals (we currently tested up
to an eight fold change in the interval duration) may
be necessary for characterizing performance during FI
schedules.
2.3. Scalar timing
Another common timing result is a constant coefficient
of variation (CV) across a range of interval values,
a form ofWeber’s law referred to as scalar timing (e.g.,
Gibbon, 1977; see also Staddon and Higa, 1999 for discussion
of differences between proportional and scalar
timing), which is observed under most steady state conditions,
although there are some data showing that it
is sometimes violated at relatively shorter (e.g., milliseconds)
and longer interval durations (e.g., Crystal,
2003; Gibbon et al., 1997; Zeiler and Powell, 1994).
We checked for the scalar property in wait times using
a method described by Lowe and Wearden (1981): we
plotted the standard deviation of the wait times against
the mean wait time and conducted a linear regression
analysis (Fig. 5). Each point is based on the data
from a session. Although overall levels of each measure
varied across subjects (note x- and y-axis scales),
all showed a positive relation between standard deviation
and mean wait time. The regression analysis indicates
that a line fit the data fairly well, with R2 values
ranging from 0.76 to 0.83 and slope values varying
from 0.61 to 0.93. Furthermore, while a majority of
the intercepts were negative, none were significantly
different from zero, as confirmed by a t-test (LB01,
p = 0.59; LB06, p = 0.60; LB08, p = 0.67; LB10,
p = 0.52). Intercept values that are near zero are excepted
if wait time is a constant fraction of the mean,
which is a form of scalar timing. Lowe and Wearden
(1981) reported similar results from a regression analysis
on data from rats and pigeons: Slope values ranged
from 0.45 to 0.98 with negative intercepts for three
of four data sets that did not differ significantly from
zero. Thus, like some birds and mammals, the wait
times of Bettas also appear to follow a scalar timing
rule.
Prior studies have examined learning processes
in Bettas using a method that involves visual reinforcement
and a swimming response (e.g., Lattal and
Metzger, 1994;Wirth et al., 2003). These studies show,
for example that the stimulus control of swimming
is similar to that found in other fish (e.g., goldfish,
Yarczower and Switalski, 1969) and other species (e.g.,
pigeons, Reynolds, 1961). The present study extends
what is known about Betta behavior to interval timing.
Overall, our results suggest that Bettas were sensitive
to the different times at which a reinforcer was available.
In particular, we observed cumulative patterns of
responding that were, in many cases, positively accelerating
as the interval elapsed. This result supports the
results from other species, including other fish species,
in which an FI scallop was reported (e.g., Dews, 1970;
Talton et al., 1999). Furthermore, although wait times
did not differ significantly during the smaller FI values,
wait times were much longer during the largest
FI schedule (FI 240 s). Break point in an interval, too,
increased with increases in the FI schedule. Although
studies with fish (e.g., Talton et al., 1999) have reported
proportional timing effects with goldfish responding
for food, we found only provisional evidence of a linear
relation betweenwait time and break point and changes
in the FI value. Our analysis suggest that linear and
power functions both provide suitable descriptions of
the results. Lastly, we found evidence of a scalar timing
effect when comparing mean wait times against the
508 J.J. Higa, L.A. Simm / Behavioural Processes 67 (2004) 501–509
Fig. 5. The relation between the mean and standard deviation of wait times for individual subjects. The data are from each session and all
conditions. The straight line represents the best-fit line from a linear regression analysis. Also shown are R2, slope, and intercept values for each
line.
standard deviation, which is comparable to results on
wait time and FI schedules in other species (e.g., Lowe
and Wearden, 1981).
Summarizing, the present study demonstrates that
it is possible to study timing behavior in Bettas, using
mirror presentation reinforcers for swimming through
a hoop. Bettas showed timing effects consistent with
other timing studies that use different procedures and
species. However, it is not yet clear whether a proportional
or power function best characterizes changes in
wait time or break point. It may be that our results
represent species differences or is limited to the particular
method used (i.e., the type or duration of the
reinforcer or the response requirement). In addition,
unlike most studies with rats and birds, our conditions
lasted for only a few sessions at a time, and it may be
that chronic exposure is necessary for both properties
of interval timing. The present study provides evidence
supporting a method that researchers may use to investigate
these and other factors involved in the timing
behavior of fish and evaluate theories about the underlying
mechanism.