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Satiation causes within-session decreases in instrumental responding

Article in Journal of Experimental Psychology Animal Behavior Processes · November 1998

DOI: 10.1037//0097-7403.24.4.439 · Source: PubMed

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Journal of Experimental Psychology: Animal Behavior Processes © 1998 by the American Psychological Association
October 1998 Vol. 24, No. 4, 439-452 For personal use only--not for distribution.

Satiation Causes Within-Session Decreases in Instrumental Responding

Lewis A. Bizo
Department of Psychology Arizona State University
Sergei V. Bogdanov
Department of Psychology Arizona State University
Peter R. Killeen
Department of Psychology Arizona State University
ABSTRACT

The amount of food necessary to cause within-session decreases in response rates was investigated
by varying reinforcer duration (Experiment 1) and grain size (Experiment 2). Within-session
response rates quickly increases to a peak, and the decrease throughout the rest of the session was
greater for conditions with larger amounts of food. The decreases were greatest for the pigeons that
ate the least when food was freely available. Previous studies of within-session changes in
responding are reviewed and reinterpreted. It was concluded that satiation causes the decrease in
responding and that this effect is not restricted to extreme manipulations of the amount of food
consumed.

Lewis A. Bizo is now at the Department of Psychology, University of Southampton, Highfield, Southampton,
United Kingdom This research was supported by National Science Foundation Grants IBN 9408022 and BNS
9021562 and by National Institute of Mental Health Award MH01293 K05. We thank David Baker, Josey Chu, and
Anders Eld, who helped to collect the data, and K. Geoffrey White, who commented on an earlier draft of the
manuscript.
Correspondence may be addressed to Lewis A. Bizo, Department of Psychology, University of Southampton,
Highfield, Southampton, United Kingdom, S0171BJ.
Electronic mail may be sent to lewisb@psy.soton.ac.uk
Received: March 25, 1997
Revised: March 30, 1998
Accepted: April 6, 1998

Holz and Azrin (1963) noted four methods for decreasing the frequency of operant responses: "punishment, satiation,
extinction and stimulus change" (p. 399). Of these, the decreases related to the ingestion of food are of primary
interest in the present article. Skinner (1938) has shown that (a) prefeeding rats depresses their subsequent
responding within a session, (b) increased deprivation increases response rates, and (c) rates of eating and drinking
decrease as time progresses in an experimental session. Reese and Hogenson (1962) reported satiation curves
showing that pigeons' response rates were relatively constant in the early part of a session and then decreased toward
the end of a session after repeated feedings.
In the same tradition, a series of experiments by McSweeney and colleagues has displayed orderly within-session
changes in the rate of operant responding (e.g., McSweeney, Hatfield, & Allen, 1990 ; McSweeney & Hinson, 1992
; McSweeney, Roll, & Cannon, 1994 ). Response rates increased from the beginning of the session to a peak and
then decreased monotonically through the middle and later part of the session.

The generality of this phenomenon has theoretical and methodological implications for the study of animal behavior
processes ( McSweeney, 1992 ). First, response rates that change during a session may not be well represented by
their average (e.g., Campbell & Dougan, 1995 ; Dougan, Kuh, & Vink, 1993 ). Second, theories accounting for
monotonic response-reinforcer relations (e.g., Herrnstein, 1970 ) also need to be expanded to explain and predict
within-session bitonic response functions to be complete and valid. Third, experimenters need to be mindful that
manipulations that produce between-session effects on behavior may also interact with bitonic patterns of responding
within a session in ways that are difficult to interpret ( McSweeney, 1992 ).

These bitonic patterns of responding have been thoroughly investigated by McSweeney and colleagues. They have
been replicated (a) for rats and pigeons responding on levers and keys on variable interval (VI), fixed interval (FI),
and variable ratio (VR) schedules (e.g., McSweeney, Roll, & Weatherly, 1994 ; McSweeney, Weatherly, Roll, &
Swindell, 1995 ); (b) for rats and pigeons with different instrumental response requirements in the different
components of concurrent (VI VI) schedules and multiple (VI VI) schedules (e.g., McSweeney, Roll, & Weatherly,
1994 ; McSweeney, Weatherly, & Roll, 1995 ; McSweeney, Weatherly, & Swindell, 1996b ); (c) during autoshaping
with pigeons ( McSweeney, Swindell, & Weatherly, 1996b ) and differential reinforcement of low rates (DRL;
McSweeney, Roll, & Weatherly, 1994 ); (d) for adjunctive and instrumental behavior on simple FI schedules with
rats ( McSweeney, Swindell, & Weatherly, 1996a ); (e) for instrumental behavior on simple VI schedules with rats
and pigeons where the variability of the VI series and the probability of reinforcement was manipulated ( McSweeney,
Weatherly, & Swindell, 1996c ); (f) for rats reinforced with water, Noyes pellets, and sweetened milk; (g) for
pigeons reinforced with mixed grain ( McSweeney, 1992 ; McSweeney, Weatherly, & Swindell, 1995b , 1995c );
and (h) for rats' running times in a T maze ( Melville & Weatherly, 1996 ) and pigeons on delayed
matching-to-sample and on successive discriminations ( McSweeney, Weatherly, & Swindell, 1996a ), and with
different types of reinforcers delivered in different components of a concurrent VI VI schedules with both rats and
pigeons ( McSweeney, Swindell, & Weatherly, 1996c ). Many of these and other studies are reviewed by
McSweeney and Roll (1993) , and McSweeney, Hinson, and Cannon (1996) .

An account of the bitonic response functions within a session in terms of sensitization and habituation has been
offered by McSweeney, Hinson, and Cannon (1996) , who argued that the increase in response rates early in the
session is caused by sensitization of responding to the reinforcer. McSweeney, Hinson, and Cannon reasoned that
"the empirical characteristics of within-session changes in operant responding are sufficiently similar to the empirical
characteristics of sensitization-habituation to suggest that common factors produce both phenomena" (p. 256). They
also argued that alternative explanations for the subsequent decrease, such as satiation, recovery from or anticipation
of handling, changes in arousal, fatigue, muscular warm-up, and anticipation of postsession feedings, are inadequate
to explain the data.

Killeen (1995) has, alternatively, suggested that the initial increase in response rates is caused by an accumulation of
arousal following repeated presentation of a reinforcer, which can become conditioned to the experimental context (
Killeen, 1997 ), and that the decrease in response rates later in a session is caused by satiation. Satiation may be
defined as "reinforcers losing their ability to increase, or maintain, behavior when too many have been consumed" (
Roll, McSweeney, Johnson, & Weatherly, 1995 , p. 324). Likewise, Morgan (1974) suggested that satiation be
defined "as a state in which the animal will no longer engage in a particular consummatory response, (eating), even in
the presence of the appropriate incentive (food)" (p. 449). Morgan also noted that instrumental behavior that precedes
the consummatory response will also cease when an organism is satiated. Killeen's interpretation can account for
much of the published data on within-session changes in responding. McSweeney and associates (e.g., McSweeney
& Roll, 1993 ) have also spoken of the increase in responding early in a session as possibly being due to arousal,
although they more formally attributed this increase to sensitization to the experimental context (e.g., McSweeney,
Hinson, & Cannon, 1996 ). The issue that is of primary interest in the present article is the cause of the decrease in
responding through most of the session, rather than the distinction between arousal and sensitization.

The Arguments Against Satiation

The satiation explanation for the decrease in response rates has been rejected when tested in McSweeney's laboratory.
"Manipulations of several variables, which are traditionally thought to contribute to satiation, produce little or no
effect on the within-session pattern of operant responding" ( McSweeney, Hinson, & Cannon, 1996 , p. 265), except
with extreme reinforcer amounts. For example, Cannon and McSweeney (1995) varied the rate of reinforcement and
reinforcer duration from 2- to 20-s access to mixed grain for pigeons and found that the within-session response
decrement was greater for the 20-s reinforcer duration when reinforcers were delivered at a high rate (VI 30 s), but
not at a lower rate (VI 60 s). Cannon and McSweeney concluded that although "satiation sometimes contributes to the
within-session patterns of responding, . . . satiation is not necessary to produce these changes" (p. 291).

McSweeney, Roll, and Cannon (1994) , however, have acknowledged that "a factor related to reinforcement (e.g.,
'satiation') plays a role in producing the within-session changes" (p. 265). They thought it "unlikely," however, "that
the present decreases in responding were produced by 'satiation', interpreted simply" (p. 265), because they found
similar decreases in normalized response rates when they varied factors such as caloric value, size, kind,
stomach-loading properties, and sweetness of the reinforcer. (See Figure 5 and the General Discussion section.)

Qualifications
This ensemble of literature appears contradictory because responding does not always decrease during the course of a
session. When moderate amounts of food are delivered at moderate rates, no decreases are seen (e.g., Schaal, 1996 ):
Responding has often been traced through the course of a session without a within-session decrement ( Killeen &
Bizo, 1998 ). Roll et al. (1995) , for example, found no decrease in responding by rats who were reinforced with 0.2
ml of diluted, sweetened condensed milk on VI 60-s schedules, over a range of deprivation levels. Melville, Rybiski,
and Kamrani (1996) delivered sixty 45-mg Noyes pellets to rats on VI 60-s schedules, and they did not observe a
systematic decrease in responding. Palya and Walter (1997) varied the number of reinforcers delivered so as to
maintain individual pigeons at 80% ± 2% of their ad lib weights; they found no systematic decreases in within-session
responding for VI 60 s and 120 s, but they did find decreases for VI 15 s and 30 s, which disappeared when the
hopper duration was reduced. The general finding that there is no decrease in response rates during the session when
moderate amounts of food are delivered at moderate rates is consistent with a satiation account.

When nonnutritive reinforcers (printed messages) were provided to human participants who responded on a VI 60-s
schedule, their responding did not decrease through the session ( Roll, McSweeney, Cannon, & Johnson, 1996 ):
Roll et al. (1996) found relatively constant rates across a 60-min session and increasing rates across a 30-min session.

McSweeney and Johnson (1994) gave pigeons two consecutive 50-min sessions and delayed the beginning of the
second session by 3, 10, or 30 min. When the animals spent the delay outside of the chamber, responding during the
second session followed the decreasing trend established in the first session, but rates during the first 5-10 min were
depressed below that trend line. The results were more varied for these animals when they were allowed to remain in
the experimental chamber during the intersession interval. In one condition, subjects' rate of responding increased
over the entire 100-min session.

Present Experiment
Variables that were expected to affect satiation were manipulated in the present article in a further attempt to examine
the role satiation plays in mediating changes in responding within a session. If satiation causes within-session
decreases in responding, then responding should decrease more when the birds consume larger amounts of food
within a session. Consequently, in Experiment 1, reinforcer duration was manipulated, and in Experiment 2,
reinforcer quality (and amount) was manipulated. In Experiment 1, the decrease in key pecking was expected to be
steeper for the longer (4.5 s) than for the shorter (1.5 s) reinforcer duration. In Experiment 2, the decrease in key
pecking was expected to be steeper for a larger grain (popcorn) than for a smaller grain (milo). We predict that the
magnitude of the decrease in responding within a session should be greater when subjects consume larger amounts of
food within a session than when they consume smaller amounts of food.

Additionally, the present experiments also made use of two additional dependent measures-rates of general activity
(e.g., Killeen, 1975 ) and hopper-approach speed. Although we expect activity and key-pecking rate to be highly
correlated, this is not always the case ( Killeen & Bizo, 1998 ), and a measure assumed to be correlated with arousal
seemed a worthwhile redundancy. This is also the case with hopper-approach speeds. DiBattista and Bedard (1987)
have shown that golden hamsters' latency to begin eating was an inverse function of deprivation. If satiation to the
reinforcer causes the decrease in operant responding over the course of a session, decreases in general activity and
hopper-approach speed should also be observed.
At a methodological level, the present experiments make a useful contribution because a Lehigh Valley feeder was
used. The amount eaten from this brand of feeder is a linear function of reinforcer duration ( Epstein, 1985 ). Some
investigators have manipulated reinforcer duration using a Gerbrands hopper ( Cannon & McSweeney, 1995 ).
Unfortunately, grain does not resettle into where the pigeon has depleted it from a Gerbrands hopper, and food
consumed is a "negatively accelerated function of magazine-cycle duration" ( Epstein, 1981 , p. 64). Consequently,
the amount eaten by pigeons in experiments that used Gerbrands feeders could have been relatively modest and
relatively constant over a range of durations that might at first glance appear extreme. The use of a Lehigh rather than
Gerbrands feeder despite our more restricted manipulation of reinforcer duration, and attendant manipulation checks,
should provide a more effective test of the satiation hypothesis.

Experiment 1
Method Animals.

Six pigeons ( Columba livia ) with various experimental histories were maintained at 85% ± 10 g of their ad lib
weights. Each subject was housed individually with free access to water and grit in its home cage and was maintained
on a 12-hr light-dark cycle with dawn at 7 a.m. The birds were fed mixed grain when necessary at the end of the day
to maintain their weights within the specified range. The birds' 85% weights ranged from 294 g to 418 g ( M = 376
g, SD = 47 g).

Apparatus.

A standard Lehigh Valley Electronics (Laurel, MD) sound-attenuating chamber housed the experimental enclosure,
which was 31 cm high, 31 cm wide (front panel), and 36 cm deep. A Plexiglas Gerbrand's (Arlington, MA) response
key, 2.5 cm in diameter, was mounted centrally, 20 cm above the chamber floor and could be illuminated white. A
force of at least 0.23 N was required to signal a response. General illumination in the chamber was provided by a
houselight that was located 28 cm above the floor and 17 cm from the left wall. A centrally located 6- ? 5-cm hopper
opening provided timed access to the magazine that contained milo grain; the hopper was illuminated when activated.
A photocell installed in the opening signaled when a pigeon's head entered the hopper. Masking noise was provided
by a speaker mounted behind the interface panel and by a ventilation fan mounted on the wall opposite the interface
panel (combined 75 dB). A Lafayette Instrument Animal Activity Monitor (#18060 Lafayette, IN) was built into
the floor of the chamber and reported the physical activity of the bird inside the chamber. The gain and activity
controls of the activity monitor were set permanently at "4" and "slow." Programs written in Microsoft Quick Basic
4.5 controlled and recorded experimental events through an IBM-compatible 386 DX2 computer located atop the
sound attenuating chamber.

Procedure.

Key pecking for all subjects was reinforced according to a VI 60-s schedule. Interreinforcement intervals were
scheduled according to Catania and Reynolds's (1968) series.

For Birds 65, 93, and 95, the reinforcer duration was 1.5 s for 13 sessions, and then 4.5 s for 25 sessions. For Birds
50, 62, and 94, the reinforcer duration was 4.5 s for 13 sessions, and then 1.5 s for 25 sessions. Birds were run in a
condition until their session response rates appeared stable to visual inspection. A session ended after 65 min elapsed
or after the delivery of 60 reinforcers.

The number of key pecks and amount of general activity were recorded in both conditions. Speed to the approach
hopper was also measured as the reciprocal of the time between the peck and hopper entry. If the pigeon did not
approach the illuminated hopper within 15 s, hopper presentation was aborted, the keylight was illuminated, and the
next VI interval began timing. Pigeons were weighed before and after a session to determine the amount of food
consumed. Because this measure doesn't take excretion into account, this will in fact be a conservative estimate of the
amount eaten. The level of significance for all statistical tests was p < .05, one-tailed.

Results

To summarize the experimental effects, we averaged key-pecking rates, activity rates, and speed to approach the
hopper across birds for the different reinforcer durations in Experiment 1. Reinforcer presentation was not included in
the time base for key pecking or activity-rate calculations. Figure 1 presents the mean key-pecking rate (top panel),
mean activity rate (bottom left panel), and mean hopper approach speed (bottom right panel) for the last five sessions
of each condition as a function of successive 5-min bins for the 1.5-s and 4.5-s reinforcer durations. The solid curves
through the data points in the top panel of Figure 1 are of the form proposed by Killeen (1995) for satiation effects.

The decreases produced by the 4.5-s reinforcer duration in key pecking were steeper than those produced by the 1.5-s
reinforcer duration. (Because hopper duration was timed from when the pigeon's head entered the hopper opening,
hopper-approach latencies did not affect reinforcer duration.) Changes in the dependent measures across a session
were quantified by subtracting the median value for the last three 5-min bins from the median value for the first three
5-min bins for both conditions. Response rates decreased significantly more for the 4.5-s reinforcer duration ( M =
15.3/min) than for the 1.5-s condition ( M = 1.5/min; Wilcoxon's matched-pair signed-ranks test, T = 1), whereas
hopper approach speeds decreased significantly more for the 1.5-s reinforcer duration ( M = 0.05/s) than for the
4.5-s condition ( M = -0.05/s; Wilcoxon's matched-pair signed-ranks test, T = 2). The decreases in activity rates
were not significantly different for the 4.5- ( M = 11.9/min) than for the 1.5-s reinforcer durations (M = 5.1/min;
Wilcoxon matched-pair signed-ranks test, T = 5). One should note that there were large individual differences in
response rates between subjects, with some animals actually showing a within-session increase in the 4.5-s reinforcer
condition. Figure 3 (left panels) summarizes response rate data for individual subjects.

The pigeons gained significantly more weight over the course of the sessions with the 4.5-s reinforcer ( M = 14.1 g)
than with the 1.5-s reinforcer ( M = 3.2 g), t (5) = 6.3. In general, the longer reinforcer duration produced steeper
decreases in responding as a function of time through the session than did the shorter reinforcer duration. Consistent
with our manipulation of the independent variable (reinforcer duration), we believe that it was the difference in food
consumption (inferred from hopper duration and weight gain) that caused steeper decreases in key pecking.

Discussion

Satiation is sufficient for these within-session decrements in response rates, where such decreases are found: The
response-rate decreases were significantly greater when the pigeons ate more (weight gain = 14 g) than when they ate
less (weight gain = 3.2 g). However, there were substantial within-subject differences in the magnitude of the
within-session decreases (see Figure 3 ), which we analyze later in concert with the data from Experiment 2.

The change in activity rates was consistent with the change in response rates; 5 of the 6 animals became less active as
they became more satiated. The hopper-approach speeds, however, were faster for the longer reinforcer and showed a
small but significant increase across a session. This suggests an incentive effect, such that the larger reinforcer
engendered faster approaches to the hopper. Why didn't hopper-approach speeds decrease with increased reinforcer
duration, as one might expect if the animals were becoming satiated? Fischer and Fantino (1968) have shown that the
behavior that is closest to reinforcer delivery is least affected by manipulations that affect satiation. In the present
experiment, hopper approach is the closest behavior to reinforcer delivery. It is therefore not surprising that
hopper-approach speed was apparently not affected by satiation, unlike the target response that is more distant from
reinforcer delivery.

Experiment 2
McSweeney, Hinson, and Cannon (1996) argued that habituation to the reinforcer, rather than satiation, accounts for
decreases in responding within a session. Habituation increases with exposure; thus, there is an accumulation of
exposure through the course of the session, which brings about the decrease in response rates. This is an interesting
hypothesis, which has some power to explain the intriguing differences in satiation as a function of food type that
McSweeney, Swindell, and Weatherly (1996c) have demonstrated. In Experiment 1, the 4.5-s condition gave three
times as much exposure to the illuminated hopper as the 1.5-s condition. If habituation increases with eating time, that
also could have been a causal factor. How can this potential causal variable be ruled out, because time of exposure to
the food is usually confounded with amount eaten? Our tactic was to provide seeds of different sizes.

Pigeons typically consume only one grain per bite (e.g., Killeen, Cate, & Tran, 1993 ; Zeigler & Feldstein, 1971 ;
Zeigler, Green, & Lehrer, 1971 ), and Ploog and Zeigler (1996) have shown that pecking rate is faster for larger
grains. The pigeons should therefore satiate more with the larger grain when given equal exposure to large and small
grain types.

Method Animals and apparatus.

The pigeons and apparatus were the same as used in Experiment 1, except that the center key could be illuminated red
instead of white.
Procedure.

The procedure was the same as in Experiment 1, except that maize (popcorn) and milo seed were used as reinforcers.
A popcorn seed weighed on average 0.28 g, and a milo seed weighed on average 0.14 g. Each reinforcer duration
was 3 s, timed from when the pigeon's head entered the hopper.

Birds 65, 93, and 95 were reinforced with milo for 25 sessions each and then with popcorn for at least 25 sessions.
Birds 50, 62, and 94 were reinforced with popcorn for at least 22 sessions and then with milo for 29 sessions each.
Birds were run in a condition until their session response rates appeared stable to visual inspection.

Results and Discussion

Key-pecking rates, activity rates, and speed to approach the hopper were averaged across birds for the different types
of reinforcer in Experiment 2. Figure 2 presents the mean key-pecking rate (top panel), mean activity rate (bottom left
panel), and mean hopper-approach speed (bottom right panel) for the last five sessions of each condition as functions
of successive 5-min bins for popcorn and milo. The solid curves through the data in the top panel of Figure 2 are
consistent with Killeen's (1995) satiation model.

The pigeons gained significantly more weight with popcorn ( M = 17.2 g) than with milo ( M = 7.4 g), t (5) =
6.4. Changes in the dependent measures across a session were quantified by subtracting the median value for the last
three 5-min bins from the median value for the first three 5-min bins for both conditions, separately for response and
activity rates, and hopper-approach speeds. The decreases in key pecking and activity produced by consumption of
popcorn ( M = 14.5/min; M = 18.9/min) were significantly greater than those produced by consumption of milo (
M = -3.2/min; M = 0.54/min; Wilcoxon's matched-pair signed-ranks test, T = 2 and T = 0), respectively. There
was a significant increase in hopper-approach speeds for milo ( M = -0.07/s) compared with popcorn ( M = 0.05/s;
Wilcoxon's matched-pair signed-ranks test, T = 0).

It is unlikely that responding decreased when popcorn was given as a reinforcer because the pigeons preferred milo to
popcorn: At the beginning of the session, response rates were higher for 5 of the 6 birds for popcorn than for milo
(see left panels, Figure 3 ). Furthermore, Killeen et al. (1993) showed that pigeons prefer larger grains, and in
particular, they prefer popcorn to milo. The steeper decrease in response rates for popcorn is consistent with a
satiation explanation, because the pigeons consumed considerably more food when popcorn (weight gain = 17.2 g)
rather than milo (weight gain = 7.4 g) was the reinforcer. Therefore, it is concluded that it was most likely that
satiation and not habituation caused the within-session decreases in responding.

General Results
In the present experiments, the amount eaten by the pigeons during a session was greater for the larger reinforcer
amounts in both experiments as inferred from changes in the pigeons' body weight. Some pigeons exhibited much
greater decreases in responding than others. Figure 3 shows that the decrease ranges from dramatic (e.g., Bird 94), to
nonexistent (e.g., Bird 93). Palya and Walter (1997) have also demonstrated such individual differences, which they
attributed to the birds' ad lib body weights, with smaller pigeons showing the steepest within-session decreases in
responding.

We hypothesized that these differences occurred because some pigeons could consume more food than others before
they became satiated. To test that hypothesis, we gave the pigeons free access to milo for 1 hr in the illuminated
experimental chamber. Their body weight was recorded before and after the feeding session and this difference gave
us a measure of how much food the pigeon could eat in a session. The pigeons were returned to their 85% weight,
and the process was repeated with popcorn. There was a significant correlation (Pearson's r = 0.95, N = 6)
between the weight gained under the milo and popcorn capacity tests, that is, those birds that could consume a large
amount of milo also consumed a large amount of popcorn. (Of interest, the relation between the pigeons' weight gain
and their ad lib weights was not significant for either milo, Pearson's r = - 0.11, N = 6, or popcorn, Pearson's r
= - 0.31, N = 6.) In Figure 3 , the order of presentation of the pigeons' data was ranked according to the amount of
food they consumed in our capacity tests. Animals that ate a small amount of food showed the steepest functions in
our Experiments 1 and 2, and animals that ate a large amount of food showed the flattest functions. Birds 94 and 95
had the smallest capacity (consuming only 11 g [milo and popcorn averaged] in the capacity tests) and had the steepest
slopes, whereas Birds 93 and 62 had the greatest capacity (22 and 32 g [milo and popcorn averaged] ), and had the
flattest slopes.

To quantify differences in response rates in the present experiments, we calculated the percentage change in response
rates from the median rate in the first three 5-min bins to the median rate in the last three 5-min bins for the 4.5-s
reinforcer-duration condition from Experiment 1 and for the popcorn condition from Experiment 2. These conditions
were chosen because the pigeons' body weights increased the most in these two conditions. Figure 4 shows the
change index plotted against capacity: The relation is significant (Pearson's r = 0.82, N = 12). The amount of food
(milo and popcorn) eaten in an hour of capacity testing accurately predicts the magnitude of decrease in responding
within the regular sessions.

General Discussion
The present experiments clearly demonstrated that longer reinforcer durations ( Figure 1 ), and larger grain types (
Figure 2 ) produce greater decreases in response rates within a session. The pigeons ate more when given longer
access to the reinforcer, and they ate more when larger grains were used, and in both cases response rates decreased
more than with the smaller reinforcers.

McSweeney, Hinson, and Cannon (1996) claimed that habituation is what determines the rate of decrease of
responding within a session. This hypothesis was tested by holding reinforcer duration constant and varying grain
size (Experiment 2). Response rates decreased more steeply across a session for the larger grain than for the smaller
grain. Because reinforcer duration was constant, habituation should also have been constant, and therefore could not
have caused the decrease, unless one assumes post hoc that there is more habituation to popcorn than milo because
habituation to popcorn is greater than milo. Such an assumption is not needed for the satiation hypothesis that predicts
that larger grains (that weigh more) will be more satiating.

Satiation Defined

To understand the claim that satiation does not cause within-session decrements in response rates, we must first agree
on what is meant by satiation, cause, and pattern. Morgan (1974) suggested that satiation be defined "as a state in
which the animal will no longer engage in a particular consummatory response, (eating), even in the presence of the
appropriate incentive (food)" (p. 449). Instrumental behavior under the control of the consummatory response will
also cease when an organism is fully satiated (e.g., Dickinson & Balleine, 1995 ).

This commonsense definition immediately exposes a problem: Satiation describes a change in behavior, not an
independent variable. Furthermore, it is a continuous change in a propensity to engage in food-oriented behavior
(e.g., Ackroff & Sclafani, 1996 ), not an all-or-none state.

To clarify the situation, it is useful to distinguish between satiation operations, satiation states, and satiation effects.
Satiation operations are the provision of nutrients adequate to restore an organism to a nondeprived state. Satiation
effects are changes in dependent variables correlated with such repletion of a deprived organism. Internal states such
as satiation or hunger are inferred from correlations of operations and effects, and they are often ascribed a causal role
in bringing about the effects. Both operations and effects are necessary to support the ascription of states. When an
animal stops responding after being given 20 g of food, satiation may be given as a plausible reason: But when
another larger animal continues responding after being given 20 g of food, we may infer that it is still hungry.
Decreasing rates suggest progressive satiation. Such analysis is useful, if somewhat post hoc. The positive
contribution of the present article is the provision of an independent measure that predicts the rates at which different
animals will satiate-that is, one which predicts when a satiation operation will produce satiation effects.

Reanalysis of Previous Research

Why have some researchers not obtained differences in responding when subjects consumed different amounts of
food? In most of the studies cited above, McSweeney and colleagues manipulated variables related to satiation and
reported a failure to find changes in the resulting pattern of normalized response rates. It is our claim that these
satiation operations were sometimes ineffective (i.e., did not produce the relevant state); other times they were applied
in ways that should not have caused within-session decreases. We find problem with (a) the data analysis and (b) the
experimental manipulation.

Response rate normalization.

Graphing techniques may have obscured the differences in the slopes of the response functions. McSweeney and
colleagues graphed responses that occurred within a bin as proportions of the total number of responses within a
session, rather than absolute response rates, so that "the differences in the absolute rates at which animals responded
do not obscure the pattern of responding within the session" ( Cannon & McSweeney, 1995 , p. 88). If such
normalizing hides relationships, then unnormalizing should reveal them. Now it is shown that this is the case.

Data from two experiments by Roll et al. (1995) were unnormalized. They varied deprivation level and caloric density
of the reinforcer for rats' responding on a multiple VI 60-s VI 60-s schedule. Roll et al. (1995) reported that
deprivation level did not affect the slope of the normalized response rates (top-left panel, Figure 5 ) and argued that
the state of deprivation did not change the "within-session pattern of responding" (p. 338), and therefore that satiation
was not a contributing factor to within-session changes in responding. We disagree with their conclusion for three
reasons. First, overall response rates were clearly depressed by decreases in deprivation level, as shown in the
bottom-left panel of Figure 5 (slightly different data are reported in the cited article, McSweeney, Hinson, & Cannon,
1996 ). Surely, overall response rate-a first-order effect-is an important component of within-session patterns of
responding. When level is taken into account, within-session patterns of responding are therefore changed by
deprivation level. The satiation hypothesis, but not the habituation hypothesis, predicts these first-order effects. It also
predicts many of the second-order effects that McSweeney and associates attribute to habituation.

Second, we disagree with their conclusion because the slope of within-session responding should not be expected to
change when deprivation level is manipulated: All deprivation does is change the initial level of responding. The
amount of food delivered within a session at each deprivation condition was not varied, hence, within-session
repletion should occur at the same rate at all deprivation levels. The satiation hypothesis predicts parallel curves, and
that is what Roll et al. (1995) found. The slopes would only be expected to deviate from parallel if differential
amounts of food were consumed at the different levels of deprivation, or if the least-deprived animals still had food in
their stomachs, thus reducing their capacity. This is possible, but not necessary.
Finally, we disagree because Figure 5 illustrates the promethian nature of the dependent variable "pattern." This is not
a classic "McSweeney within-session pattern," as no decrease in rates was found through the session. Why is this?
This is because the rats were reinforced with restricted access to 0.2 ml of condensed milk, a procedure which was,
apparently, insufficient to satiate the animals. The satiation hypothesis is not embarrassed by this fact. Why is the
habituation hypothesis not embarrassed? Why are such data in general taken as evidence for a pattern of responding
that is due to habituation, not satiation?

Ineffective manipulations.

Several experiments have manipulated the amount of food without apparently affecting responding. McSweeney,
Hinson, and Cannon (1996) argued on the basis of these experiments that satiation is not necessary for within-session
decreases in responding. Each of these experiments will be addressed in turn.

Roll et al. (1995) varied caloric density by changing the amount of diluted, sweetened condensed milk per reinforcer,
and they found that there was a difference between the pattern of within-session responding for the largest amount
(1.0 ml), and the other amounts (baseline, 0.20, and 0.60 ml). In the 1.0-ml and 0.60-ml conditions, 0.20 ml was
delivered orally, and the remaining amount (0.80 ml and 0.40 ml) was intubated directly into the rats' stomachs. Roll
et al. found that there was no difference in the within-session normalized pattern of responding between the baseline
(0.20) and .40-ml conditions (top-right panel Figure 5 ), although the largest amount caused a radical decrease in
normalized rates. Reanalysis of their data tells a different story (bottom-right panel of Figure 5 ). Absolute response
rates were different for all three conditions. Response rates were so depressed for the largest amount that any
conclusions about the within-session pattern of responding are tenuous.

Although unnormalization of these data favors our view that deprivation manipulations have their major effect on
level, factors other than satiation may have been at work. Deutsch (1983) noted that undigested milk placed directly
into the stomach can produce strong taste aversion, whereas digested milk will not produce taste aversion. The
decrease in level of responding observed by Roll et al. (1995) may have been caused by nausea in addition to
satiation.

McSweeney, Roll, and Cannon (1994) observed that higher rates of reinforcement for rats led to steeper decreases in
response rates and earlier peaks, and they concluded that these effects support the "idea that a factor related to
reinforcement (e.g., "satiation") plays a role in producing the within session changes" (p. 265). They also argued,
however, "it is unlikely that the present decreases in responding were produced by 'satiation', interpreted simply" (p.
265), because of similar patterns in normalized response rates for different reinforcers (Noyes pellets and sweetened
condensed milk). This claim deserves examination. In their Experiments 1 and 2, there are two conditions where rats
were reinforced on a multiple VI 60-s VI 60-s schedule through a 60-min session. In one condition, lever pressing
was reinforced with a single 45-mg Noyes pellet, and in the other, key pressing was reinforced with 5-s access to a
0.25-ml dipper of diluted, sweetened condensed milk. Response rates were 14.4 (lever presses) and 43.8 (key
presses) per minute for these two conditions, respectively. The top panel of Figure 6 shows that the normalized
response rates overlap for these two conditions, which supports their claim. When these data were reanalyzed,
plotting real response rates against successive 5-min components, a different functional relation was revealed. Real
response rates change little for lever pressing for Noyes pellets, whereas the decrease in key pressing for diluted,
sweetened condensed milk is dramatic. How can this be? Consider the quantity of food that was delivered to the rats.
Sixty 45-mg pellets weigh approximately 2.7 grams, whereas 60 dippers filled with 0.25 ml of diluted, sweetened
condensed milk weigh approximately 15 grams. This is a substantial difference. The data of McSweeney, Roll, and
Cannon (1994) can be interpreted in terms of satiation, simply. The highest real response rates and largest decrease in
rate occurred when the subjects consumed the most food.

McSweeney, Roll, and Cannon (1994) data for pigeons were ambiguous, with the highest rate of reinforcement VI 15
s yielding approximately the same response rate (25.7/min) as the lowest rate of reinforcement VI 240 s (23.1/min),
both substantially lower than found at intermediate rates of reinforcement (VI 30 s, 60 s, and 120 s): 43/min. The
decrease in average response rate under VI 15 s is consistent with satiation, but one might have also expected a faster
decline in response rates within the session, which apparently did not happen.

There are data extant that support a satiation explanation for within-session decreases in responding (e.g., Palya &
Walter, 1997 ). McSweeney (1992) found with rats that as the rate of reinforcement was increased the pattern of
within-session responding steepened and peaked earlier. Rats received more food when reinforced at higher rates, so
these changes in responding are consistent with satiation. Weatherly, McSweeney, and Swindell (1996) found very
steep decreases in responding when pigeons and rats were delivered large numbers of reinforcers on a conjoint VI VT
schedule compared with when they were delivered small numbers of reinforcers.
There are, however, some notable data that seem to falsify the assertion that satiation is the major factor causing
within-session decreases in responding. Cannon and McSweeney (1995) systematically varied reinforcer duration
across a range from 2-s to 20-s access to mixed grain at two different reinforcement rates (VI 30 s and VI 60 s). They
observed a steeper decrease in responding across a session when the pigeons were given 20-s compared with 2-s
access to grain, but only for VI 30 s, not for VI 60 s. Why was there no big effect for the substantial 20-s feeding?
With just 4.5-s access, some of our pigeons required several days off to maintain their running weight at 85%-surely
20 s should have satiated the birds after a few feedings. But as Cannon and McSweeney noted, the amount the
pigeons get to eat is not proportional to hopper time: The grain does not resettle into where the pigeon has depleted it
from in a Gerbrands hopper ( Epstein, 1981 ). Despite Cannon and McSweeney's dramatic increase in reinforcer
duration, the amount eaten by their pigeons could have been relatively constant over the range of reinforcer durations
they reported. A similar problem also exists for the data of Weatherly, McSweeney, and Swindell (1995) . This is not
true of all feeders: For those designed by Lehigh Valley, the amount eaten is a linear function of reinforcer duration (
Epstein, 1985 ). A Lehigh Valley feeder was used in the present experiments; consequently, our more modest change
in reinforcer duration may have provided a more effective manipulation of reinforcer amount. Note that our
manipulation check confirmed that the increase in the birds' body weights in the present Experiment 1 was
significantly larger with a 4.5-s reinforcer ( M = 14.1 g) than with a 1.5-s reinforcer ( M = 3.2 g).

It is easy to draw the wrong conclusion from satiation studies. McSweeney, Weatherly, and Swindell (1996b) ran
pigeons on concurrent VI VI schedules and found that response rates varied inversely with reinforcement rates,
whereas body weights remained constant for the richest schedules. Because of this constancy, they concluded that
satiation could not have produced the inverse relationship. But as subjects satiate, they will neither eat much or
respond much, thus decreasing the overall response rates at higher feeding rates. If their birds (like our Birds 94 and
95) worked until they satiated and then stopped, they would stop sooner at higher rates of reinforcement, yielding the
observed relation-less responding at higher rates of feeding. The inverse relation between reinforcement rate and
response rate is thus exactly what is predicted if satiation is responsible for the decreases in responding within a
session. That the body weights of all of the subjects were roughly the same for the four highest rates of reinforcement
is also consistent with this argument, because it suggests that the pigeons had eaten to capacity. Because the
researchers only ran the subjects when they dropped back to 85%, the pattern they noticed is just the one predicted by
a satiation explanation.

Recently, Weatherly, McSweeney, and Swindell (1997) reported that response rates decreased steeply for rats and
pigeons reinforced on a simple VI 15-s schedule. They varied the probability of reinforcer delivery, such that in the
richest and leanest conditions, rats received 161 or 6 reinforcer deliveries (0.1-ml sweetened condensed milk) and
pigeons received 70 or 11 reinforcer deliveries (5-s access to mixed grain) in a 60-min session. Weatherly et al.
(1997) argued that the steep decreases in within-session normalized response rates for the richest schedules were
consistent with a satiation explanation, and they also argued that steep decreases in within-session response rate for
the leanest conditions were inconsistent with a satiation explanation. This is an important observation, given that the
rats consumed only 6 g versus 161 g of sweetened condensed milk, and the pigeons only had 55 s versus 350 s
access to grain, on average within a session in the leanest versus the richest condition. This is a serious challenge to
the satiation hypothesis, because it would not predict steep decreases in response rates when animals consumed a
small amount of food. Because we were concerned about the challenge raised by these data, we converted the
normalized rates to responses per minute, and then by using a linear regression, we calculated the slopes of the
within-session response rates. Contrary to the conclusions reached by Weatherly et al. (1997) , we discovered that
response rates did decrease steeply within a session for the richest schedules for both rats (slope = -0.30) and pigeons
(slope = -0.41); however, response rates did not decrease steeply for the leanest schedules for the rats (slope = -0.08)
or pigeons (slope = 0.19). Our reanalysis of Weatherly et al.'s data highlights the danger that analysis, solely in terms
of normalized rates, poses. (Weatherly et al. performed ANOVAs on normalized data and plotted their data in terms of
normalized rates; see Appendix .)

Fischer and Fantino (1968) have found that satiation operations differentially affect responses, depending on their
temporal relationship to food presentation: Responses that normally occur close to food delivery are more resistant to
satiation. For example, Robbins (1969) found that rats' running speed increased with larger durations of liquid
reinforcement on a runway when rats only received a single trial per day. With one trial per day, satiation could not
contribute to performance, and incentive effects predominated. Killeen (1995 , Equation 12) showed that there
generally will be an inverted-U relation between amount and response strength, with incentive effects dominating
when smaller amounts are consumed, and satiation effects dominating when larger amounts are consumed.

Although we believe that food consumption in relation to an individual's normal capacity to consume food accounts
for most or all of the data reviewed in this article, one should recognize that there can be important conditioning
effects of motivational states, as nicely shown by McSweeney, Swindell, and Weatherly (1996c) . Davidson (1993)
has shown that food-deprivation states in rats produce discriminable interoceptive stimuli. Morgan (1974) noted
several examples of "resistance to satiation," where the subject continues to produce the instrumental response despite
apparent satiation, that is, when the consummatory response has ceased. Dickinson and Balleine (1995) have argued
that motivational states engendered by different values of food are learned (for a review of learned satiation see,
Booth, Gibson, Toase, & Freeman, 1994 ):

Animals have to learn through experience with a particular food in the undeprived state that it has a low incentive
value when they are not hungry. Moreover, the role of such learning is not restricted to a downshift in hunger. If
animals are trained in the undeprived state, testing under food deprivation often elevates performance only if they have
previously experienced the food pellets when hungry ( Dickinson & Balleine, 1995 , p. 164).

In conclusion of this section reviewing evidence for and against an explanation of within-session response decreases,
it appears that satiation operations in some published research were not effective in modulating satiation states to the
degree necessary to produce noticeable differences in responding. Because manipulation checks (e.g., the amount of
weight change over a session) are seldom reported, no conclusion can be made concerning the degree of satiation
actually achieved in these studies (e.g., Cannon & McSweeney, 1995 ; Roll et al., 1995 ; Weatherly et al., 1995 ).
The present Experiments 1 and 2 show that if the animals eat enough food, then their response rates will decrease
across the course of a session as they get progressively more satiated. Furthermore, the amount of food it takes for
this to happen for some pigeons (4.5 s in Experiment 1) is nearly the same amount McSweeney and colleagues
routinely use in studies showing this within-session effect (5 s). McSweeney, Hinson, and Cannon (1996) argued
that satiation will "contribute little to the within-session patterns of responding under the relatively standard operant
procedures used in many experiments" (p. 266). The present experiments have shown that this amount of food will
cause satiation in some animals. But satiation will in fact contribute little to response patterns (or levels) in standard
operant procedures, because the standard hopper duration is less than 5 s. In most standard procedures, satiation is
not a serious threat, and response rates will not decrease.
Our Claim for Satiation

Satiation is sufficient to cause within-session decrements in response rates, and that is not in dispute. Satiation
manipulations have long been known to decrease the likelihood of responding within a session. Reese and Hogenson
(1962) reported satiation curves that showed that pigeons response rates were relatively constant in the early part of a
session and then decreased toward the end of a session after repeated presentations of 4-s access to grain. They also
noted that their pigeons would not respond for food when their weights were above 85% of free feeding.

In general, satiation is not necessary to bring about within-session decreases in rates: Any behaviorist could list many
factors (e.g., drugs, punishment, signaled decrease in reinforcement probability, etc.), which would bring about a
decrease (e.g., Holz & Azrin, 1963 ). However, those factors are not an issue here. The rejection of necessity
(because decreases occurred without satiation operations) is not flawed because of confounds: It is flawed because the
amount of food some researchers have typically fed animals, 5-s access to grain, is in fact a satiation operation, and it
is adequate to bring about significant changes in motivational level through the course of the session, as shown by
decreases in the general activity of our subjects.

Our analysis of individual subject's data revealed some interesting and novel effects; those subjects that had higher
capacities to consume food showed smaller decreases in response rates. The strongest evidence for the satiation
hypothesis is the covariation in the amount of free food adequate to satiate an animal and the decrease in responding
over the course of an experimental session (see Figures 3 and 4 ). There is no reason to expect such individual
differences on the basis of habituation. In conjunction with the experimental manipulation of Experiments 1 and 2,
one may conclude that satiation caused the within-session decrease in responding witnessed in these experiments. We
may infer that it is the most likely cause of within-session decreases in experiments of this type.

That in the present experiments the decreases in response rates across the course of a session were satiation effects
does not mean that within-session changes could not in some cases be due to other variables such as habituation (
Groves & Thompson, 1970 ). This is especially the case where the reinforcer is a nonconsumable stimulus change
(for a review, see McSweeney & Roll, 1993 ). For example, McSweeney and Roll (1993) listed seven studies in
which leverpressing for light change decreases within a session. Some of those authors (e.g., Kish & Baron, 1962 )
referred to those decreases as "satiation of sensory reinforcement," presumably because the stimulus change was
self-administered, and in that context it is not unreasonable to refer to habituation as satiation. But to likewise
explain the present results for consumable reinforcers in terms of habituation would require that habituation covary
with amount consumed, as inferred from the difference in body weights between pre- and post-session measurements
( Figure 3 ). This step would shift the claim from "habituation, not satiation" to "satiation as habituation." This may
be a brilliant step, but it is a reinterpretation of satiation, not its denial. In fact Swithers and Hall (1994) identified
habituation to the oral stimulus properties of food as a contributing factor to the onset of satiation, one that helps
control when animals stop eating. However, they also identified other variables that contribute to the termination of
eating, such as gastric fill. Some researchers have noticed stimulus specificity, which they call the "dessert effect,"
where apparently satiated subjects can be made to eat again when a new food is presented (e.g., Clifton, Burton, &
Sharp, 1987 ). Swithers and Hall (1994) also noted there was less oral habituation, and subjects did not decrease their
food-seeking behavior with more intense stimuli, for example, 15% sucrose versus 7.5% or water, when gastric load
was held constant. This is consistent with an incentive effect in the operant literature. In our experiments, when we
varied reinforcer amount we changed crop and stomach distension in the pigeon, that is, gastric load was probably the
major controlling variable that contributed to satiety.

The role of gastric fill has long been recognized as a major contributor to satiety (e.g., Young, Gibbs, Antin, Holt, &
Smith, 1974 ). Animals will eat enormous quantities when they are "sham fed," as the food does not fill the gut to
provide stop signals ( Smith & Gibbs, 1979 ). "Oral satiety" also plays a role in cessation of eating, but it is
secondary in importance to gastric fill, and only occurs under restricted conditions ( Nissenbaum & Sclafani, 1987 ).

Conclusion
Satiation (states) must be defined in terms of both operations and effects. An example is the provision of greater
access to food that leads to decreased response rate. To avoid circularity, independent predictors of onset of satiation
should be identified. To provide an independent measure of propensity to satiate, we measured capacities to consume
food. To check the effectiveness of the satiation operation, we measured changes in body weight from the beginning
to the end of the session in the present experiment. We found that when animals consume more, response rates
decrease more within the session, and we attribute this to satiation; those animals with smaller capacities to consume
food satiate faster ( Figure 4 ).

Experiments that fail to show satiation effects may be (a) operating in a range of the independent variable where
incentive effects predominate; (b) using a dependent variable that is more sensitive to incentive effects than to satiation
effects (as was the case for hopper approach speed), (c) using ineffective satiation manipulations; (d) demotivating the
animals before the session, and therefore should not expect to see consequent within-session changes (other than level
changes); or (e) normalizing their data in a way that attenuates differential changes.
The most parsimonious explanation of this ensemble of data is that when animals begin to eat they begin to satiate,
and this will be reflected in within-session changes in response rates, until the animals are fully satiated when they
will stop responding. Whether or not the causal factors for within-session response decreases include oral
habituation, crop distension, stomach distension, intestinal distension, or changes in blood chemistry associated with
the ingestion of food, the mechanism is one of satiation.
APPENDIX A
Data Treatment The Pattern

McSweeney and colleagues typically analyze not response rate, but the proportion of the total number of responses in
a session through the session; that is, a normalized response rate. In many cases, this renormalization erases
information that bears on the claims concerning nonsatiation.

What does this normalization do to the changes in the response rates, the main dependent variable? In particular, how
does the normalized slope change with changes in the nonnormalized slope? A linear regression, y = mt + b ,
provides a good characterization of average within-session decreases, where y is the number of responses (or rate of
responding), b is the y intercept, and m is the slope (usually negative) of the response rates as a function of time
through the session ( t ). The normalized data result from dividing such a function by its integral over the session: y
Norm = mt + b &int; 0 T mt + b dt where T is the total duration of the session. This integrates to: y Norm =
mt + b T mT /2 + b T > 0, t < | b / m | (To avoid negative rates, the model requires that animals stop
responding when the regression crosses the x axis, that is, | mt | < b .)

The slope of this normalized function, m Norm , is: m Norm = m T mT /2 + b The normalized slope ( m Norm )
covaries with the real slope ( m ) as it should, but is greatly attenuated because the real slope appears in both the
numerator and the denominator. As the intercept b gets smaller, as it would with animals that are pre-fed or are run
closer to their ad lib weight, the relation between the two slopes is further attenuated (see left panels of Figure 5 ).

Session duration appears in the denominator, and if this varies, so will the normalized slope. In experiments where
session duration covaried with experimental conditions, the use of normalized rates thus introduces a confound.
Longer sessions (where satiation is more likely) have larger values of T , and this further attenuates the normalized
slopes.

Normalization

We found large differences in response rates for different animals ( Figure 3 ). A traditional average provides a
representative picture ( Figures 1 and 2 ). If, however, one wishes to normalize to account for individual differences,
there are better ways to do it ( Loftus, 1993 ; Loftus & Masson, 1994 ), ways which do not interact with the pattern
measured. Normalization that obscures one of the key effects of the experimental manipulation (level of responding)
and biases others (slope changes) is counterproductive.

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Figure 1. Mean key-pecking rate (top panel), vibration rate (bottom left panel), and hopper-approach speed (bottom
right panel), plotted as a function of successive 5-min bins. Data were averaged across pigeons and across the last
five sessions of each condition. The reinforcer duration was either 1.5 s (empty circles) or 4.5 s (filled circles).

Figure 2. Mean key-pecking rate (top panel), vibration rate (bottom left panel), and hopper-approach speed (bottom
right panel), plotted as a function of successive 5-min bins within a session. Data were averaged across pigeons and
across the last five sessions of each condition. The reinforcer was either milo (circles) or popcorn (disks).
Figure 3. Key-pecking rates for individual pigeons plotted as a function of successive 5-min bins within a session for
Experiment 1 (left panels) and Experiment 2 (right panels). Data were averaged across the last five sessions for each
individual bird. The panels are rank ordered according to the body-weight change (given in parentheses in grams)
averaged across eating capacity tests for milo and popcorn.

Figure 4. The percentage changes in response rates in the two richest conditions from Experiments 1 (disks) and 2
(squares) are plotted as a function of the crop capacity of the pigeons, as determined by the amount their body weight
changed over the course of 1 hr of free feeding.
Figure 5. Proportion of total session response rates (top panels) from J. M. Roll, F. K. McSweeney, K. S. Johnson,
and J. N. Weatherly (1995). Absolute response rates (bottom panels) were recovered from average response rates
reported by Roll et al. (1995).

Figure 6. Top panel: Proportion of total session response rates. From "The Generality of Within-Session Patterns of
Responding: Rate of Reinforcement and Session Length," by F. K. McSweeney, J. M. Roll, and C. B. Cannon,
1994, Animal Learning & Behavior, 22, p. 258 and 264. Bottom panel: Absolute response rates were recovered from
average response rates reported by McSweeney et al. (1994). The disks are for lever presses when the reinforcer was
a 45-mg Noyes pellet. The inverted triangles indicate key presses when the reinforcer was 5-s access to a 0.25-ml
dipper that contained diluted, sweetened condensed milk. The rats were reinforced according to a VI 60-s schedule,
and sessions lasted for 60 min.

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