Journal of Experimental Psychology: Animal Behavior Processes 1982, Vol. 8, No.

4, 313-328

Copyright 1982 by the American Psychological Association, Inc. 0097-7403/82/0804-03i3$00.75

Overshadowing and Summation in Compound Stimulus Conditioning of the Rabbit's Nictitating Membrane Response
E. James Kehoe
University of New South Wales, Kensington, New South Wales, Australia The present experiments examined acquisition of the rabbit's nictitating membrane response to a light + tone simultaneous compound stimulus and its components as a function of the intensity of the tone. In Experiment 1, the tone intensity was varied across the values of 85, 89, and 93 dB, and the CS-US (conditioned stimulus-unconditioned stimulus) interval'was 400 msec. In Experiment 2, the tone intensities were 73, 85, and 93 dB, and the CS-US interval was 800 msec. Experiments 3 and 4 further examined the effects of the 73-dB tone at CS-US intervals of 400 and 800 msec, respectively. All experiments included control groups, which were trained with either a light or a tone CS. In brief, the experiments revealed repeated instances of overshadowing, i.e., the impairment of conditioned response (CR) acquisition to one or both of the components of a compound. Moreover, two types of summation were obtained: within-subjects summation, in which rabbits trained with a compound showed a higher level of responding to the compound than to either of its component CSs (Experiments 2, 3, and 4), and between-groups summation, in which a group trained with a compound showed faster CR acquisition than either of its corresponding control groups trained with a single CS (Experiments 1 and 2). The results are discussed in terms of perceptual and distributive processing models of compound stimulus conditioning.

Any organism in even the most sterile en- cally possible to observe both the interaction vironment is faced with a continual and and integration of the component stimuli. multifaceted stream of stimulus events. To The delineation of an interaction concerns discover the laws governing behavioral ad- the extent to which conditioned response justments to the exigencies of an environ- (CR) acquisition to a given conditioned stimment, Pavlov (1927, pp. 110-113) and sub- ulus (CS) is influenced by the other comsequent investigators used compounds of two ponent in a compound, A prototypical exseparable stimuli (e.g., tone + light) as a lab- ample of an interaction is "overshadowing." oratory model for the array of innocuous In the overshadowing phenomenon, reevents that antedate a biologically significant sponse acquisition to one CS is reduced stimulus (e.g., Hull, 1943; Kehoe & Gor- through compound training with another CS, mezano, 1980; Razran, 1965, 1971; Wick- which often is highly "salient" in that it proens, 1954, 1959, 1965). In any compound duces rapid response acquisition (Kamin, stimulus conditioning procedure, it is logi- 1969; Mackintosh, 1976; Pavlov, 1927, p. 14L). While the study of interaction concerns the relation between the components of a This research was supported by the Australian Re- compound, the study of integration concerns search Grants Committee. The author expresses his grat- the relation of the components to the comitude to Nancy Amodei and Ann Topple for collecting the data. The author also thanks Linda A. Kehoe for her pound as a whole. The study of integration critical reading of the manuscript and J. C. Clarke, has been conducted both on a within-subjects C. R. Gustavson, and R. F. Westbrook for their sug- basis, by periodically testing the subject with gestions concerning interpretation of the data. individual presentations of the components Requests for reprints should be sent to E. James Kehoe, School of'Psychology, University of New South during compound training, and on a beWales, P.O. Box 1, Kensington, New South Wales 2033, tween-groups basis, by comparing the responding of a group trained with a comAustralia.
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pound to control groups trained with each of the components. The most common experimental outcome has been "summation," in which the level of responding to the compound exceeds that to the components (Kehoe & Gormezano, 1980). Although the label summation carries the connotation of an atomistic process of combination, the term is intended strictly as a descriptive one without prejudice to alternative theories. Many conditioning theories address overshadowing and summation. These can be roughly grouped into two classes, which will be denoted "perceptual" and "distributive." The most extreme perceptual hypotheses assume that the separate components of a compound fuse into a single functional stimulus, i.e., a "configuration" (Razran, 1965, 1971). Less extreme perceptual hypotheses assume that the component stimuli of a compound either modify each other's sensory encoding (Hull, 1943)'or contribute independently to a joint signal (Heinemann & Chase, 1975; Kehoe & Gormezano, 1980, pp. 370-373). For perceptual hypotheses, the results of tests with components outside the context of the compound (i.e., overshadowing and withinsubjects summation) presumably reflect the degree of generalization (or lack thereof) from the compound to the functional stimuli of the separate components. In contrast to the perceptual hypotheses, the distributive hypotheses assume that the components of a compound remain functionally the same in all contexts but are processed in such a way that there is a trade-off jn the amount of associative strength ultimately gained by each of the contending stimuli. Distributive hypotheses postulated a fixed processing capacity (Sutherland & Mackintosh, 1971) or a fixed associative strength (Rescorla & Wagner, 1972; cf. James & Wagner, 1980). However, Mackintosh (1975) argued that tradeoffs can be achieved by a process in which the stimulus with the greatest relative associative strength on a trial gains a proportional increment in its learning rate parameter while all other stimuli suffer a decrement in their respective learning rate parameters (cf. Moore & Stickney, 1980). To account for summation results, the distributive accounts all assume a relatively simple addition of associative strengths (Mackintosh, 1975; Rescorla & Wagner, 1972).

The variety of alternative formulations concerning compound conditioning betrays the fragmented nature of the available data. Although there is a growing body of research concerning stimulus interaction (e.g., James & Wagner, 1980; Mackintosh, 1976; Mackintosh & Reese, 1979) and stimulus integration (e.g., Bellingham & Gillette, 1981; Gillette & Bellingham, 1982; Kehoe & Gormezano, 1980), no single study has sought to establish the relation between overshadowing, within-subjects summation, and between-groups summation. Moreover, there are major unresolved empirical questions concerning each of these phenomena. Reciprocal Overshadowing Reciprocal overshadowing is said to occur when acquisition to both components of a compound show some impairment relative to corresponding single-stimulus controls (cf. Mackintosh, 1976). The issue of reciprocal overshadowing has been raised in connection with distributive hypotheses which postulate a fixed processing capacity. According to this postulate, which has been designated the inverse hypothesis, acquisition to one stimulus occurs always at the expense of other concurrent stimuli (Mackintosh, 1975, 1976). When two CSs are of equal salience, reciprocal negative interactions would be expected, because the two CSs would evenly divide the limited resource. On an empirical level, overshadowing effects have been largely unidirectional in nature (e.g., Kamin, 1969; Mackintosh, 1976). However; Mackintosh (1976) found that reciprocal overshadowing effects were detectable but only when equisalient stimuli were weak or moderate in absolute intensity (cf. James & Wagner, 1980). The Constancy of WithinSubjects Summation Although within-subjects summation has been demonstrated repeatedly, there have been relatively few attempts to determine whether a fixed empirical rule can be found to describe the quantitative relation between responding to a compound and to its components (Kehoe & Gormezano, 1980). Among the available theories, the configural hypotheses generally argue against any stable

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relation (Razran, 1965, 1971). Other perceptual hypotheses postulate a stable relation but contend that the quantitative relation between responding to the compound and responding to its components will vary with factors affecting the degree of generalization between the compound and its nominal components (Bellingham & Gillette, 1981; Heinemann & Chase, 1975; Hull, 1943). In contrast, the distributive accounts appear to predict a relatively fixed quantitative relation between responding to a compound and responding to its components (Mackintosh, 1975; Rescorla & Wagner, 1972). While they do not specify an exact rule for mapping associative strength into responding, they do assume that there is a fixed monotonic relation between the associative strengths of current stimuli and overt performance. Between-Groups Summation and Overshadowing

.(1976) observed between-groups summation in conjunction with substantial overshadowing of one component. In order to provide additional evidence relating to these issues, the present experiments were designed to examine acquisition of the rabbit's nictitating membrane response to a light + tone compound and its components as a function of tone intensity. Test trials for the individual components were interspersed among the compound training trials to permit a determination of withinsubjects summation throughout acquisition. Moreover, a complete set of single-stimulus controls were included in order to assess both overshadowing and between-groups summation effects. Experiment 1 - The design of Experiment 1 was modeled on that of Mackintosh (1976, Experiment 1). In order to detect overshadowing, a fixed visual CS (flashing house light) was compounded with a auditory CS (1000-Hz tone) which was varied in intensity across groups. The values of the tone were selected on the basis of previous research in the University of New South Wales laboratory. An 85-dB (SPL) tone had been found to produce a CR acquisition rate comparable with that of the light CS. According to the convention that defines salience in terms of demonstrated conditionability, the 85-dB tone CS and the light CS would be regarded as having equal salience. If an inverse law is correct, mutual negative interactions would be expected for animals trained with a compound of the light and 85-dB tone. In order to determine whether unidirectional overshadowing of the light could be obtained, two more intense values were used, namely, 89 and 93 dB.

Between-groups summation occurs whenever CR acquisition in a group trained with a compound is faster than that in corresponding groups trained with only one of the components. The only formulation that predicts that between-groups summation will occur under all circumstances is that of Hull (1943). Hull assumed that the associative strengths of the component stimuli functional inside a compound will summate to yield betweeq-groups summation even if one or both of the nominal stimuli suffer substantial generalization decrements when tested outside the context of the compound (Kamin, 1969, p. 55). All other formulations, both perceptual (e.g., Heinemann & Chase, 1975) and distributive (Mackintosh, 1975; Rescorla & Wagner, 1972), generally predict between-groups summation, but they also expect it to disappear when profound overshadowing occurs. Under the distributive Method hypotheses, overshadowing represents a defSubjects. The subjects were 56 naive male and female icit in acquisition of associative strength. If albino rabbits (Oryctolagus cuniculus). On arrival, each was 70-80 days old and weighed about 1.5 kg. one component of a compound can be en- rabbit Each rabbit had free access to food and water in its home tirely prevented from acquiring associative cage. strength, then between-groups summation Apparatus. The apparatus and recording procedure disappears. The available evidence is ambig- for the nictitating membrane response were patterned uous: In conditioned suppression paradigms, after those of Gormezano (1966) and were described in detail by Kehoe, Schreurs, and Amodei (1981). During Kamin (1969, pp. 53-55) found that be- training sessions, each rabbit was held in a Perspex retween-groups summation and overshadow- strainer and placed in one of eight sound-attenuating, ing were mutually exclusive, but Mackintosh ventilated chambers. On the wall of the chamber in front

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E. JAMES KEHOE A response was defined as any extension of the nictitating membrane exceeding .5 mm (1 mm of pen deflection). A CR on reinforced trials was defined as any response initiated after the onset of the CS(s) but before US onset, i.e., a response with a latency of less than 400 msec following CS onset. On the test trials, a CR was defined as any response occurring with a latency less than 1,000 msec after CS onset. For analysis of the data, a set of contrasts was written by using the Bonferroni technique (Miller, 1966). The rejection level was set according to a Type I error rate (.05) for a family of contrasts.

of the subject was located a stimulus panel. A speaker was mounted at the midpoint of the stimulus panel, 8 cm anterior to and 16 cm above the subject's head. The speaker provided both white noise and an auditory CS, which was a 1000-Hz tone of 85, 89, or 93 dB (SPL) superimposed on an 82-dB background provided by the white noise and exhaust fans. Mounted on the stimulus panel 4 cm above the speaker was an 8-W frosted neon light tube. To provide a visual CS, the light was flashed at a rate of 20 Hz. The unconditioned stimulus (US) was a 50-msec, 3-mA, 50-Hz AC shock delivered via stainless steel Autoclip wound clips positioned 10 mm apart and 10-15 mm posterior to the dorsal canthus of the right eye. The sequence and timing of stimulus events on each trial were controlled by solid-state circuitry. Each rabbit's right external eyelids were held open by No. 3 tailor-hooks mounted on a Velcro strap which fitted about the head. A muzzle-like head-set, fitted securely about the snout, supported a photosensitive transducer for monitoring movements of the nictitating membrane. A small hook was attached to a silk loop sutured in the nictitating membrane of the rabbit's right eye. The hook was connected by a thread to one end of an L-shaped piano-wire lever, which mechanically transmitted the movement of the nictitating membrane to a photoelectric transducer. The variations in electrical signal from the transducer were amplified and recorded on a four-channel, ink-writing oscillograph operating at a speed of 50 mm/sec. Procedure. All animals received 1 day of preparation, 1 day of recovery, 1 day of adaptation, and 6 days of acquisition training. On the preparation day, a small loop of silk (000 Dynex) was sutured into the rabbit's right nictitating membrane, the surrounding hair was removed, and the animals were returned to their home cages for 1 day of recovery. On the adaptation day, the animals were placed in the conditioning apparatus for a period equal to the length of the subsequent training sessions, but neither the CSs nor the US was presented. Following preparation, the animals were assigned randomly to one of seven groups (n = 8). Three groups were trained with a compound of light and tone. Across the three groups, the light was fixed, but the tone was varied over the values of 85, 89, and 93 dB. These compound groups were designated as LT85, LT89, and LT93, which denotes light, tone, and intensity of the tone. The compound groups received 54 reinforced trials per day, which were interspersed with 3 unreinforced test trials each to the light (L) and tone (T). The test trials were presented on Trials 10, 20, 30, 40, 50, and 60. On oddnumbered days, the sequence of test trials was LTTLLT, and on even days, the sequence was TLLTTL. In addition to the compound stimulus groups, there were four single-CS control groups: One was trained with the light and the others with the 85-, 89-, and 93-dB tones, respectively. The control groups were designated L, T85, T89, T93. The light group received 54 reinforced trials per day, 3 unreinforced light test trials, and 3 unreinforced test trials with the 85-dB tone. Similarly, the tone control groups received 54 reinforced trials per day, 3 unreinforced tone test trials, and 3 unreinforced light test trials. For all groups, the CS duration and the CSUS interval were 400 msec. The intertrial intervals were 50, 60, and 70 sec (M = 60 sec).

Results Figure 1 depicts the mean CR percentage shown by the seven groups across successive 2-day blocks of training. Panels A, B, and C show the CR acquisition curves for the light + tone compound and its components for Groups LT85, LT89, and LT93, respectively. Panel D shows the CR acquisition curves as measured on reinforced trials for the single-stimulus control groups, namely, T85, T89, T93, and L. Overshadowing. Inspection of Figure 1 reveals clear evidence of overshadowing in that CR acquisition to the fixed light CS was impaired by training in compound with the more intense tones. Across the three compound groups, the rate and level of acquisition to the light decreased as tone intensity increased. Overall, there was a significant downward linear trend in the means for responding to the light, which were 82%, 47%, and 34% for Groups LT85, LT89, and LT93, respectively, F(\, 28) = 19.54. More important, inspection of panel D of Figure 1 reveals that CR acquisition in Group L, which was given reinforced training with only the light, rapidly reached an asymptote near 100% and showed an overall mean of 74%. In comparison with Group L, Groups LT89 and LT93 showed significantly lower levels of responding to the light CS, Fs(\, 28) = 8.97 and 17.74, but Group LT85's responding to the light failed to differ from that of Group L (F<\). Since CR acquisition to light was impaired by compound training with the more intense tones, it is possible that what level of responding there was to the light component may have represented only crossmodal generalization from the intense tone component. In order to ascertain the degree of crossmodal generalization from tone to light, the level of

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responding on light test trials in the tone control groups was examined. The overall mean levels of responding to the light in Groups T85, T89, and T93 were 29%, 18%, and 9%, respectively. These levels were, respectively, 53,29, and 25 percentage points below those obtained with light in the corresponding compound groups, smallest f\l, 42) == 6.89. Accordingly, crossmodal generalization from tone accounts only for a portion of the responding to the light in groups trained with a compound. Nevertheless, the differences in responding to the light between the tone and compound groups were larger at the lower intensities, for the interaction between the linear trend across tone intensity and the

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Figure 1. Mean percentage of conditioned responses (CRs) in Experiment 1 plotted as a function of 2-day blocks. (Panels A, B, and C depict CR acquisition to the compound and its components for Groups LT85, LT89, and LT93, respectively. Panel D shows the CR acquisition curves for Groups T85, T89, T93, and L.)

compound versus tone training conditions was significant, F(\, 42) = 4.14. Since the level of responding to light in both the compound and the tone control groups was inversely related to tone intensity, different levels of crossmodal generalization may have contributed to the differences among compound groups in their respective levels of responding to the light component. Inspection of the acquisition curves for tone, as shown in Figure 1, reveals that the level of responding to tone in both the tone control and the compound groups was generally high across all three intensities. Despite the large effects of tone intensity on CR acquisition to light, there was little evidence of pronounced differences in salience among the tone intensities as measured in the control groups. In connection with the issue of reciprocal overshadowing, there was little suggestion of impairment in CR acquisition to tone in any of the compound groups. In the tone groups, the mean levels of responding shown to the tone by Groups T85, T89, and T93 were 79%, 87%, and 75%, respectively. The means of the corresponding compound groups were 78%, 83%, and 85%, respectively. As may be apparent, there is one disparity in the pattern of means between the tone groups and the compound groups: Group T89 showed a higher level of responding than groups T85 or T93, while in the compound groups, the level of responding to the tone appeared to be a positive function of intensity. These differences were most pronounced early in training and were confirmed by a significant three-way interaction between the tone versus compound conditions (Factor 1), linear trend across tone intensity (Factor 2), and linear trend across blocks of trials (Factor 3), F(l, 42) = 4.20. Befween-groups summation. Even when overshadowing occurred, CR acquisition to the compound appeared to proceed more rapidly than in the single-stimulus control groups. In Groups LT85, LT89, and LT93, the overall mean levels of responding to the compound were 91%, 89%, and 91%, respectively. In Groups T85, T89, T93, and L, the overall means for reinforced trials were 77%, 85%, 73%, and 74%, respectively. Inspection of Figure 1 reveals that differences were most apparent in the first block of train-

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ing, after which the performance of all groups converged. Statistical analysis of the overall means confirmed that all three compound groups were individually superior to Group L, smallest F(l, 49) = 12.24. Group LT85 was superior to Group T85, f\l, 49) = 10.72, and Group LT93 was superior to Group T93, F(l, 49) = 18.30. However, Group LT89 was not superior to Group T89 (F < 1). An analysis of differences in linear trends across training blocks revealed an identical pattern of results. Within-subjects summation. Inspection of panels A, B, and C of Figure 1 reveals some evidence for within-subjects summation. In Group LT85, responding to the compound was initially higher than responding to either component, but responding on all three types of trials converged to a mean asymptotic level near 100%. The initial difference in the acquisition curves for the compound and its components was confirmed in the statistical comparison between the linear trends for compound versus tone trials, F(l, 21) = 16.19, but not for compound versus light trials, F( 1,21) = 7.25, p < . 10. In Groups LT89 and LT93, the level of responding to compound was only marginally higher than responding to the tone throughout acquisition, but responding to the compound was substantially higher than responding to the light. In both groups, there was no significant difference between responding to the compound and responding to the tone, but there was a significant overall difference between responding to the compound and responding to the light, Fs(lt 21) = 22.97 and 43.55 for Groups LT89 and LT93, respectively. In order to determine whether the group mean curves reflected the performance of the individual subjects, the relative levels of responding to the compound and its components over Days 1 and 2 were examined for each subject. Out of eight subjects in each group, Groups LT85, LT89, and LT93 contained, respectively, six, seven, and three subjects that showed a higher level of responding to the compound than to either component.

summation in Groups LT85 and LT93, and (c) weak evidence of within-subjects summation, which was most apparent at the lower tone intensities. The pattern of results lies between that of Mackintosh (1976) and that of Kamin (1969): The observation that overshadowing occurred in conjunction with between-groups summation agrees with the findings of Mackintosh (1976, Experiment 1). However, with respect to the question of reciprocal overshadowing, the present results resemble those of Kamin, i.e., unidirectional overshadowing of the light was observed with the 89- and 93-dB tones, but unimpeded CR acquisition to both light and tone apparently occurred with the 85-dB tone. Mackintosh (1976, Experiment 2) gave reinforced training with equally salient light and tone stimuli to three compound groups which differed in the absolute intensities of both stimuli. Reciprocal overshadowing was found in the group trained with the least intense CSs, whereas the other two groups trained with more intense CSs showed no evidence of overshadowing. In the present experiment, the CSs used were highly salient, as demonstrated by the rapid CR acquisition seen in all the single-CS control groups. Accordingly, the unimpeded CR acquisition to the tone and light in Group LT85 can be construed as consistent with Mackintosh's (1976, Experiment 2) findings. In the assessment of both overshadowing and within-subjects summation, the interval for scoring of responses on light and tone test trials, 1,000 msec, was longer than that of reinforced compound trials, 400 msec. Consequently, the level of responding on test trials may have provided a higher estimate of CR strength than reinforced compound trials, which would attenuate the apparent magnitude of overshadowing and withinsubjects summation effects. However, other experimentation with the rabbit nictitating membrane response (NMR) preparation failed to reveal any differences in the percentage CR measure for observation intervals of 400 msec on CS-US trials and 1,000 msec on CS test trials (Tait, Kehoe, GorDiscussion mezano, Note 1). Yet, in order to avoid any The results of Experiment 1 revealed (a) potential biases in subsequent experiments, unidirectional overshadowing of light as a the interval of measurement on all trials was function of tone intensity, (b) between-groups equal to the CS-US interval.

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Experiment 2

The present experiment was conducted to (a) further test Mackintosh's (1976) hypothesis that reciprocal overshadowing occurs only when the salience of the two components is equal and low and (b) examine summation effects when the overall rate of CR acquisition is slower than that observed in Experiment 1. In Experiment 1, CR acquisition on reinforced trials in both the compound and the control groups was relatively rapid and reached asymptotes near 100% CRs. With the exception of the overshadowing results, differences between groups and within subjects were largely confined to the first 2 days of training. Accordingly, it could be argued that the failure to observe reciprocal overshadowing in Group LT85 and the weakness of within-subjects summation could be attributed partly to the ceiling on measurement imposed by the rapid approach to 100% CRs. In fact, Kamin (1969, pp. 54-56) found that even unidirectional overshadow-, ing varied with the overall rate of acquisition and could be eliminated if the overall rate of conditioning was raised by increases in the intensity of a shock US. Consequently, in the ^present study, the overall rate of CR acquisition was reduced by lengthening the CSUS interval to 800 msec, a value that produces a modest but reliable rate of CR acquisition (Kehoe, 1979; Smith, Coleman, & Gormezano, 1969). While the available theories (Hull, 1943; Mackintosh, 1975; Rescorla& Wagner, 1972) differ regarding questions of reciprocal overshadowing, they all agree that overshadowing is a function of relative CS salience, Thus, it should be possible to reduce tone intensity to a point at which the tone will be overshadowed by the light used in the present experiments. Consequently, the present experiment extended the range of the tone intensity manipulation by including a tone of 73 dB, which was demonstrably less salient than the Results and Discussion light CS. Method
Subjects. The subjects were 44 naive male and female albino rabbits, 70-90 days old and weighing about 2 kg on arrival. Apparatus and procedure. Unless otherwise specified, the apparatus, recording technique, and procedure were

identical to those described for Experiment 1. The timing of stimulus presentations was controlled by an Apple II computer with interfaces and software described by Scandrett and Gormezano (1980). The only change in procedure caused by the introduction of computer control was in the intertrial interval (ITI) values. With the control system used for Experiment 1, the ITIs were randomized among the values of 50,60, and 70 sec (M = 60), whereas the computer system denominated the ITI values in 1-sec units over the range 50-70 sec (M = 60). For recording purposes, the signals from the movement transducer were sent to the 8-bit analog-to-digital converter of the computer as well as to the oscillograph. During trials, the computer sampled the signal from each transducer every 5 msec. The rabbits were randomly assigned to one of seven groups, all of which received 8 days of training. Groups (n = 8) labeled LT73, LT85, and LT93 received reinforced training with a light + tone compound, in which the tone intensities were 73,85, and 93 dB, respectively. Groups (n = 4) labeled T73, T85, and T93 received reinforced training with a tone CS, for which the intensities were 73, 85, and 93 dB, respectively. Finally, a group (n = 8) labeled L received reinforced training with the light CS. During training, four animals died, one in Group LT85, one in Group LT93, and two in Group L. Their incomplete data were not included in the description or analysis of the results. For all groups, both the CS duration and the CS-US interval were 800 msec. Moreover, the response scoring interval for all trials was 800 msec measured from CS onset. Comparison of computer versus human data. Preliminary statistical analyses compared CR frequency data collected by the computer with those collected by the human observer. Separate analyses of the data from tone and from light test trials failed to reveal any significant discrepancies between the two methods of response counting. However, the analysis of data from reinforced trials revealed a small but significant difference of four percentage points between the overall mean CR frequencies collected by the computer (55%) and those collected by the human (59%), f\l, 29) = 5.48, p < .05. There were no significant interactions of data collection method with the tone intensity variable, compound versus single stimulus manipulation, or trends across training. A trial-by-trial examination indicated that the human observer counted as responses some movements that, according to the computer, were just under the nominal .5 mm (1 mm of pen movement) criterion for a response. The human observer's criterion appeared to be equivalent to 3/8ths mm (6/8ths mm of pen movement). Since the two methods of data collection were in good agreement, the results reported for Experiments 2, 3, and 4 are based on the computer-collected data.

Figure 2 depicts the mean CR percentage shown by the seven groups across successive 2-day blocks of training. Panels A, B, and C show the CR acquisition curves to the light + tone compound and its components for Groups LT73, LT85, and LT93, respectively.

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Panel D shows the CR acquisition curves on reinforced trials for the single-stimulus control groups, T73, T85, T93, and L. Overshadowing. Inspection of Figure 2 reveals that all three compound groups displayed overshadowing of light. The level of responding to the light in the compound groups rose to terminal levels less than 50% CRs, which diverged significantly from the high terminal level shown by Group L (M = 88% CRs). Statistical comparisons confirmed that there were significant differences in the linear trends across training for each compound group relative to Group L, smallest F(\, 24) = 12.23. Comparisons between the
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Figure 2. Mean percentage of conditioned responses (CRs) in Experiment 2 plotted as a function of 2-day blocks. (Panels A, B, and C depict CR acquisition to the compound and its components for Groups LT73, LT85, and LT93, respectively. Panel D shows the CR acquisition curves for Groups T73, T85, T93, and L.)

compound groups failed to reveal any significant differences in their respective levels of responding to light. To determine whether the observed responding to the light in the compound groups could be attributed to crossmodal generalization from the tone component, I compared the levels of responding on light test trials in the tone control groups with those of the compound groups. Specifically, Groups T73, T85, and T93 showed overall mean levels of 5%, 11%, and 12% CRs, respectively. Collectively, these levels of generalized responding to light were significantly lower than the overall mean levels of responding to the light shown by Groups LT73, LT85, and LT95, which were 32%, 16%, and 26%, respectively, F(\, 33)= 5.29. Examination of responding to the tone CS revealed some evidence of reciprocal overshadowing in Groups LT85 and LT93. Both Groups LT85 and LT93 showed terminal levels of responding (45% and 64%) that were substantially lower than those of their respective tone control groups^ T85 and T93 (90% and 90%). In comparisons between linear trends across training, Group LT85 versus Group T85 failed to attain the declared level of significance, F( 1,28) = 6.18, p > .05, but Groups LT93 and T93 did differ significantly, F(l, 28) = 8.14. As shown in panel D of Figure 2, Groups T93 and T85 both showed rapid CR acquisition which followed largely the same course as that of Group L. Thus, the salience of the 93- and 85-dB tones appeared close to that of the light, creating the conditions for reciprocal overshadowing under the inverse hypothesis. While the reciprocal overshadowing evident in Groups LT85 and LT93 was consistent with an inverse law, Group LT73's results were perplexing both theoretically and empirically. In examining the control groups, it is clear that the light was more salient than the 73-dB tone. Specifically, Group L showed more rapid CR acquisition and a higher overall level of responding than Group T73, F(\, 14) = 13.74. However, in Group LT73, CR acquisition to the light but not the tone was impeded. Although the rate of CR acquisition to the tone in Group LT73 was slow, its course of acquisition failed to differ from the equally slow acquisition of Group T73 (F <

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1). If salience is to be anchored to the rate and level of CR acquisition produced by a stimulus when trained by itself, then Group LT73 appears to be a case in which a less salient stimulus can unidirectionally overshadow a more salient stimulus. Between-groups summation. Whereas Experiment 1 revealed consistent betweengroups summation, only Group LT93 in the present experiment showed facilitation of GR acquisition to the compound: Group LT93's overall level of responding to the compound (79%) was significantly greater than that of Group L (52%), F(\, 33) = 8.78, and there was a significant divergence between the linear trends across training for Groups LT93 and T93 (overall M = 55%), F(\, 33) = 8.49. Among the other groups, Group LT85's overall mean level of performance (69%) failed to differ from that of either Group T85 (52%) or Group L, while Group LT73's overall mean (59%) differed from that of Group T73 (17%), F(\, 33) = 16.10, but not that of Group L. The inconsistent nature of between-groups summation in the present experiment is particularly damaging to contentions that the associative strengths of stimuli functional inside a compound should always summate to facilitate CR acquisition to a compound (Hull, 1943). Within-subjects summation. Inspection of panels A, B, and C of Figure 2 reveals strong evidence for within-subjects summation, especially in comparison with the weak effects seen in Experiment 1. In all three compound groups, the level of responding to the compound significantly exceeded the level of responding to either the tone or the light component throughout training, smallest F( 1, 19) = 11,14. To determine whether the group mean curves reflected the performance of the individual subjects, I examined the mean levels of responding to the compound and its components over all days for each subject. In Groups LT73, LT85, and LT93, 7/8, 7/7, and 6/7 subjects, respectively, showed a higher level of responding to the compound than to either component. Evaluation of a combination rule. One of the key issues dividing the available theories concerns the constancy of within-subjects summation: Configure! hypotheses expect an unstable relation between the level of overt

responding to the compound and that to its components. The remaining perceptual hypotheses expect a stable relation but would include a correction for generalization decrements from compound training to component testing. Likewise, the distributive hypotheses would also expect a relatively stable quantitative relation between performance to the compound and its components. Since nonconfigural hypotheses, both perceptual and distributive, specify the combination rule as a summation of theoretical quantities (i.e., associative strength), the exact form of the quantitative relation for overt responding cannot be specified. However, a fair test for the stability of within-subjects summation can be accomplished by comparing observed levels of responding to a compound with predicted levels derived from a fixed rule for combining the observed levels of responding on tone and light test trials. In particular, the percentage CRs to a compound (PAE) can be compared with predicted levels (PAS') based on the percentage CRs to the separate components (PA, PB) combined according to the formula PAS' = PA + PB- (PA X PB). This formula is, of course, the general formula for the probability of at least one "hit" when sampling simultaneously from two independent sources. Informal examination of compound conditioning data in a variety of procedures has indicated that predicted levels from the independent combination rule approximate the observed levels of responding to the compound (Kehoe & Gormezano, 1980, pp. 360-361). To assess the stability and accuracy of the independent combination rule, I compared the obtained levels of responding to the compounds with the predicted levels in a set of orthogonal comparisons. In the analysis of Experiment 1's results, the only comparison involving a difference between the obtained and the predicted performances was in an interaction of the linear trend across days with the linear trend across tone intensity, F(l, 21) = 5.17. Examination of the means used in testing the interaction revealed some modest discrepancies between obtained and predicted performances. As the most notable, in Group LT93, there was a 7% overprediction for the first 2-day block and a 7% underprediction for the last 2-day block. For all

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90 80 70 60 50 40 30 20 10

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Figure 3. A scattergram of the obtained percentage of conditioned responses (CRs) on reinforced compound trials (abscissa) versus the predicted percentage of CRs based on the independent combinations of CR probabilities estimated from light and tone test trials (ordinate). (Each point represents the datum of a single subject from the compound groups in Experiment 1 [Days 1-2] and Experiment 2 [Days 3-4].)

other points, the largest discrepancy between the obtained and the predicted means was 2%. However, in the analysis of Experiment 2's results, there was a significant main effect comparison between the overall mean of the predicted performance (48%) and the mean of the obtained performance (69%). Moreover, there were no interactions involving the comparison between predicted and obtained levels. Thus, there was a consistent, 21 percentage point underproduction by the combination rule under consideration. To depict the subject-by-subject correspondence between obtained and predicted levels of responding, Figure 3 is a scattergram of the actual percentage CRs on reinforced compound trials and the predicted percentage CRs based on the independent combination of CR probabilities estimated from tone and light test trials. For Experiment 1, the data shown are from Days 1-2 of training, during which time the levels of performance across subjects varied most widely. After the first 2 days, the range of variation became constricted as the level of responding to both the compound and the components

in many subjects reached 100% CRs. For comparable data from Experiment 2, the data from Days 3-4 are shown. Figure 3 also shows the best fitting straight lines for the regression of the predicted performance on the observed performance in Experiments 1 and 2. Inspection of Figure 3 reveals that there was moderate correspondence between the actual and the predicted levels of performance, which was statistically confirmed by significant product-moment correlation coefficients. For Experiment 1, r = .68, n = 24, and for Experiment 2, r = .79, n = 22. As indicated by the statistical analyses, the results of Experiment 1 showed some overprediction, particularly in subjects that showed a moderate level of responding to the compound, and the results of Experiment 2 showed a consistent underprediction. Despite the discrepancies between the predicted and the obtained compound performance levels in both experiments, the outcome of the independent combination rule indicates that there are stable relations between responding to a compound and responding to its components. Thus, the evaluation of a combination rule supports nonconfigural hypotheses. If the underpredictions are taken seriously, they particularly support arguments by perceptual hypotheses that generalization decrements in the level of responding to components outside the compound underestimate the associative strength of the stimuli functional inside the compound (Heinemann & Chase, 1975; Hull, 1943). The underpredictions can be handled by distributive hypotheses but only at the expense of adding a perceptual postulate. Specifically, the Rescorla-Wagner model has been appended with a configural, "unique stimulus" hypothesis. This hypothesis contends that the observed responding to a compound represents the summation of not only the associative strengths of the components but also that of an additional, configural stimulus that arises during each presentation of a compound stimulus (Rescorla, 1972, 1973; Whitlow & Wagner, 1972). Accordingly, a prediction of responding to the compound based on only the levels of responding to the separate components would not include the associative strength of the unique stimulus.

OVERSHADOWING AND SUMMATION

323
SOO-mnec CS-US Interval

Experiments 3 and 4 The most startling result of Experiment 2 was the observation that a demonstrably weak 73-dB tone was able to impair CR acquisition to a more salient light without the tone itself suffering any noticeable reciprocal impairment. Experiment 3 was conducted to confirm the overshadowing of light by the 73-dB tone under an 800-msec CS-US interval. Subsequently, Experiment 4 was conducted to determine whether the same result would be obtained when the overall rate of CR acquisition was increased by shortening the CS-US interval to 400 msec. Method
The subjects were 32 albino rabbits, 70-90 days old and weighing 2 kg on arrival. In each experiment, three groups of rabbits were trained, which consisted of Groups LT73 (n = 8), T73 (n = 4), and L ( = 4). These three groups were trained and tested in the same fashion as described for the corresponding groups in Experiment 2, with the exception that Group L was tested with the 73-dB tone three times in each block of 60 trials. The procedure was identical to that of Experiment 2, with the following exceptions: In Experiment 3, training was conducted in 3 days, each composed of 180 trials, which corresponded in sequence to three successive sessions of 60 trials used in Experiment 2. In Experiment 4, the CS-US interval was 400 msec, and training was conducted across 5 days, each composed of 120 trials corresponding in sequence to two successive sessions in Experiment 2. The reduction in the number of trials per session from Experiment 3 to Experiment 4 was necessitated by observations that the 180 trials per session in Experiment 3 produced within-session decrements in responding by some animals, which increased the withincell variance. The reduction in the number of trials per session from Experiment 3 to Experiment 4 was expected to have a positive effect on the rate of CR acquisition along with the reduction in the CS-US interval (Kehoe & Gormezano, 1974).

EXPERIMENT 3,
100

GROUP
90 80

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Figure 4. Mean percentage of conditioned responses (CRs) in Experiment 3 (800-msec CS-US interval) plotted as a function of days. (Left panel depicts CR acquisition to the compound and its components for Group LT73. Right panel shows the CR acquisition curves for Groups T73 and L.)

Results and Discussion Experiment 3. In Figure 4, the left panel shows the CR acquisition curves for Group LT73's compound and its components, and the right panel shows the CR acquisition curves for the single-stimulus control groups, T73 and L. Inspection of Figure 4 reveals that there was clear overshadowing of the light in that the mean level of responding to the light in Group LT73 (27%) was significantly lower than in Group L (70%), F(l, 10) = 31.12. With respect to the tone, the

control group, T73, showed a relatively low overall level of responding (17%), and thus there was little room to observe whether CR acquisition to the tone in Group LT73 (M = 10%) suffered any impairment indicative of reciprocal overshadowing. Although both groups showed some acquisition to the tone across days, F(\, 10) = 10.17, any tendency of Group LT73's responding to the tone to diverge to a lower level than that of Group T73 was not significant, F(\, 10) = 3.91, p > .05. Finally, a comparison between the levels of responding in the control groups confirmed that the light was more salient than the 73-dB tone, F(\, 13) = 9.79. To determine whether the low level of responding to the tone in Group LT73 may have reflected generalization from the light component of the compound rather than the tone's own associative strength, I estimated the degree of crossmodal generalization from light to tone by examining responding on the tone test trials given to Group L. In fact, none of the four subjects in Group L ever responded to the tone over 27 test presentations, whereas in Group LT73, six subjects showed at least one response to the tone. Since there was no within-cell variance for Group L, I decided to use the nonparametric

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EXPERIMENT 4,
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E. JAMES KEHOE
4OO-msec CS-US Interval CONTROLS

LT73
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oo GROUP L

ONE-DAY BLOCKS

Figure 5. Mean percentage of conditioned responses (CRs) in Experiment 4 (400-msec CS-US interval) plotted as a function of days. (Left pafcel depicts CR acquisition to the compound and its components for Group LT73. Right panel shows the CR acquisition curves for Groups T73 and L.)

Mann-Whitney test, which revealed a significant difference between Group L and Group LT73 (C/ = 4, one-tailed, p = .024). Consequently, crossmodal generalization from light to tone does not account for the responding to tone in Group LT73. Consonant with the results of Experiment 2, there was little evidence of between-subjects summation. In fact, the level of responding to the compound in Group LT73 fell at a level that was intermediate to those of Groups L and T73. However, statistical comparisons failed to confirm that Group LT73 differed significantly from either Group T73, F(l, 13) = 4.82, p > .05, or Group L, F(\, 13) = 2.01, p>.05. .With regard to within-subjects summation, Group LT73 displayed a level of responding to the compound that was significantly higher than the level of responding to either the tone, F(\, 7) = 24.73, or the light, F(\, 7) = 14.76. Examination of the individual subjects in Group LT73 revealed that all eight showed a higher level of responding to the compound than to either component. Moreover, an assessment of the independent combination rule revealed that the mean overall level of responding to the compound (49% CRs) was underpredicted by the combined levels of respond-

ing to the components (26%), F(\, 7) = 19.03. Experiment 4. Figure 5 shows the CR acquisition curves for Group LT73 in the left panel and Groups T73 and L in the right panel. Inspection of Figure 5 reveals that in Group LT73, CR acquisition to both tone and light appeared to show some impairment relative to the respective control groups. However, statistical analyses failed to reveal a significant difference between Group LT73 and Group L in their overall level of responding to light, F(l, 10) = 2.66. Examination of the individual subjects' responding revealed that while Group L showed relatively homogeneous levels of responding (range in overall means = 27 percentage points), Group LT73 showed highly variable levels of responding (range = 73 percentage points). In comparisons between Group LT73 and Group T73, their linear trends across days of training differed significantly, F(l, 10) = 12.82. Specifically, Group LT73 initially showed a higher level of responding to the the tone than Group T73 but subsequently crossed over and diverged to a lower level. In summary, there was some evidence of reciprocal overshadowing of the tone and light. This pattern of results represents a near reversal of that obtained in Experiments 2 and 3, in which the overall rate of CR acquisition was slower. Furthermore, the performances of Groups L and T73 showed similar, rapid CR acquisition. Thus, it appears that the increases in the overall rate of acquisition in Experiment 4 effectively erased the disparity in salience between the light and the 73-dB tone. Although the pattern of overshadowing results in Experiment 4 was altered from the patterns seen in Experiments 2 and 3, the remaining results generally agreed with those found in Experiments 2 and 3. First, there was little evidence that crossmodal generalization from light to tone contributed in any major way to responding to the overshadowed tone component of the compound group. Specifically, in Group L, the overall mean level of responding to the tone was only 2% as compared with the mean level of 49% CRs to tone in Group LT73, F(l, 10) = 15.09. Second, there was little evidence of between-groups summation: The overall level

OVERSHADOWING AND SUMMATION

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of responding to the compound in Group LT73 (77%) failed to differ significantly differ from that of either Group L (71%) or Group T73 (56%). However, Group LT73's performance over days did diverge to a higher level than Group T73's, as indicated by a significant difference in their respective linear trends, F(l, 13) = 7.56. Third, Group LT73 did show within-subjects summation, in that the overall level of responding to the compound (77%) was significantly higher than the overall level of responding to either the tone (49%), F(l, 13) = 20.41, or the light (46%), F(l, 13) - 12.70. Individually, seven of the eight subjects showed a higher level of responding to the compound than to either component. Finally, an assessment of the independent combination rule revealed that the mean overall level of responding to the compound (76% GRs) was underpredicted by the combined levels of responding to the components (64%), However, the apparent underprediction failed to attain the declared level of significance, F(l, 6) = 5.09, .05 < p < .10. There are two key findings in Experiments 3 and 4. First, the unidirectional overshadpwing of a salient light by a less salient, i73dB tone can be duplicated at an 800-msec interval, but when the overall rate of acquisition is raised primarily by shortening the CS-US interval, the difference in salience between the light and the tone is reduced, and the pattern of overshadowing appears to be reciprocal. Second, the results of withinsubjects and between-groups summation comparisons were consistent with those obtained in Experiment 2. General Discussion With respect to the key issues outlined in the introduction, the major findings of the four experiments are now discussed. Between-Groups Summation and Overshadowing Only three of the eight compound-trained groups in the present experiments showed statistically significant between-groups summation, namely, Groups LT85 and LT93 in Experiment 1 and Group LT93 in Experi-

ment 2. The lack of consistent between*groups summation clearly contradicts Hull's (1943) contention that the addition of the associative strengths of the functional stimuli within a compound should always facilitate CR acquisition to a compound. However, the pattern of results does not offer particularly strong support for the distributive accounts (Mackintosh, 1975; Rescorla & Wagner, 1972). They would expect between-groups summation to occur under conditions of equal salience and to disappear under conditions of strong unidirectional overshadowing. In fact, between-groups summation did not appear to be related to the degree of overshadowing. Specifically, Group LT85 in Experiment 1 did not evidence overshadowing, Group LT93 in Experiment 1 showed unidirectional overshadowing of light, and Group LT93 in Experiment 2 showed reciprocal overshadowing. From the perspective of perceptual formulations, the infrequent occurrence of between-groups summation indicates that a compound stimulus is generally processed in the same manner as an individual CS, i.e., as either a unitary event (Razran, 1971) or as the conjoint basis for a single response decision (cf. Blough, 1972, 1975; Heinemann & Chase, 1975; Kehoe & Gormezano, 1980). The Constancy of Within-Subjects Summation Statistically reliable within-subjects summation was obtained for six of the eight compound-trained groups in the present series of experiments. The observation of within-subjects summation confirms previous demonstrations of within-subjeets summation in rabbit eye-blink conditioning (Saavedra, 1975, Experiment 2) as well as in other conditioning paradigms (Kehoe & Gormezanb, 1980). Furthermore, an assessment was made of a descriptive combination rule for predicting the likelihood of responding to the compound from the likelihood of responding to the individual components. For the results of Experiments 2, 3, and 4, the proposed rule produced an underprediction of the level of \ responding to the compounds. While withinsubjects summation is generally consistent with the expectations of all the formulations

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under consideration, the tendency for the descriptive rule to underpredict responding to the compound tends to favor the generalization decrement postulates of perceptual formulations (cf. Bellingham & Gillette, 1981; Heinemann & Chase, 1975; Kehoe & Gormezano, 1980). Since the quantitative relation between responding to the components and responding to the compound was relatively stable, the within-subjects summation results can also be regarded as supporting the distributive accounts if supplemented by a configural, unique-stimulus hypothesis to account for underproductions of responding to the compound (cf. Rescorla, 1972, 1973; Whitlow & Wagner, 1972). Reciprocal Overshadowing The results of all four experiments provided repeated and systematic demonstrations of unidirectional overshadowing, which previously has been shown only indirectly in the NMR preparation (cf. Marchant & Moore, 1973). Moreover, reciprocal overshadowing was found in Group LT93 of Experiment 2 and perhaps Group LT85 of Experiment 2 and Group LT73 of Experiment 4. In terms of their theoretical implications, the present results contradicted the expected one-to-one relations between overshadowing, physical CS intensity, and CS salience as denned by the rate of CR acquisition. Under the 400-msec CS-US interval of Experiment 1, the level of responding to the light component of a compound declined as tone intensity increased, but, as measured in single-CS groups, the tone stimuli appeared to have equivalent salience. In Experiment 2, under the 800 msec CS-US interval, the opposite pattern of results appeared, namely, CR aquisition to the light was equally impaired across variations in tone intensity, which, in the control groups, proved to be differentially salient. More important, in Experiments 2 and 3, it appeared that a demonstrably weak, 73-dB, 800-msec tone impaired CR acquisition to the light, without CR acquisition to the tone itself appearing to suffer a reciprocal impairment. Although the correspondence between the degree of overshadowing and salience was not all what the available theories would pre-

dict, appeals to ceiling effects on measurement and other vagaries of comparing overshadowing and salience across different groups might be used to defend the assumption of a one-to-one relation between the degree of overshadowing and the relative CS salience. However, in Experiments 2 and 3, the unidirectional overshadowing of the light by the less salient 73-dB, 800-msec tone represents an apparent reversal of the relation between overshadowing and relative CS salience which defies any simple appeal to measurement difficulties. Since the overshadowing of light by the 73-dB, 800-msec tone was undeniable, the only way out of the theoretical paradox posed by the anomalous overshadowing lies in explaining why CR acquisition to the 73-dB, 800-msec tone failed to show a reciprocal impairment. In examining the total pattern of results, there are two pieces of evidence that CR acquisition to the 73-dB, 800-msec tone was subject to reciprocal overshadowing by the light. First, under the 800-msec CS-US interval in Experiment 2, CR acquisition to the 85-dB and 93-dB tones did show some reciprocal impairment. Second, under the 400-msec CS-US interval in Experiment 4, the 73-dB tone itself suffered impairment in the final days of acquisition training. On the basis of this converging evidence, it appears possible to infer that the associative strength of the 73-dB, 800-msec tone was reduced but that overt responding arose from any of three sources: (a) a within-compound association between the tone and light (Rescorla, 1981, 1982; Rescorla & Cunningham, 1978), (b) stimulus generalization from the light component of a compound, and (c) stimulus generalization from a configural stimulus. In considering the possible sources of responding to the 73-dB, 800-msec tone, the contribution of a within-compound association between the light CS and tone CS is difficult to assess. On the basis of results from other rabbit NMR studies, the evidence for associations between simultaneous stimuli is nonexistent: Conditioning between simultaneous CSs and USs has never been found (Kehoe, Feyer, & Moses, 1981, Experiment 3A; Smith et al., 1969). However, until it is determined whether second-order conditioning can be obtained with simulta-

OVERSHADOWING AND SUMMATION

327

neousCSs in the NMR preparation, a withincompound association remains a possible contributor. As a source of generalized responding to the 73-dB tone, it is possible to rule out direct generalization from the light itself: In Experiment 3, examination of the control groups revealed negligible levels of generalization from light to tone. Finally, in the compound groups, there is the additional possible source of generalized response strength, namely, from a configural stimulus. What is most notable is that the anomalous overshadowing of light but not tone occurred only under the 800-msec CS-US interval for which the independent combination rule significantly underpredicted the likelihood of responding to the compound. This underprediction was construed as supporting the general hypothesis that some perceptual integration (e.g., Bellingham & Gillette, 1981; Heinemann & Chase, 1975; Kehoe & Gormezano, 1980) between the components occurred over the CS-US interval. Under a configural hypothesis, it would be expected that the level of responding to the components would be largely attributable to stimulus generalization from the associative strength of the configural stimulus. In the case of the 73-dB, 800-msec tone in Experiments 2 and 3, the level of responding was generally low, and, thus, with even modest stimulus generalization from a configural stimulus, it would be possible to account for responding to the 73-dB, 800-msec tone. Despite the anomalies between overshadowing and salience measurements, the overshadowing results agree with Mackintosh's (1976) hypothesis that reciprocal overshadowing will be found only when the CSs are of relatively low salience. Thus, these results tend to challenge a strict inverse hypothesis of overshadowing (e.g., Rescorla & Wagner, 1972) and favor the more open-ended Mackintosh (1975, 1976) model or the perceptual formulations (Heinemann & Chase, 1975; Hull, 1943). Consequently, the theoretical burden now shifts toward detailing the exact conditions under which reciprocal overshadowing occurs. In connection with the Mackintosh (1975, 1976) model, it will be necessary to specify more precisely the initial learning rate parameters, the size of the decrements in them from trial to trial, and the

mapping of associative strength unto overt responding (cf. Moore & Stickney, 1980). For the perceptual formulations, it will be necessary to obtain independent scales of the generalization decrements between the compound and its components (cf. Kehoe & Gormezano, 1980). Reference Note
1. Tail, R. W., Kehoe, E. J., & Gormezano, I. Effects of US duration on classical conditioning of the rabbit's nictitating membrane response. Unpublished manuscript, University of Iowa, 1982.

References
Bellingham, W. P., & Gillette, K. Spontaneous configuring to a tone-light compound using appetitive training. Learning and Motivation, 1981, 12, 416-428. Blough, D. S. Recognition by the pigeon of stimuli varying in two dimensions. Journal of the Experimental Analysis of Behavior, 1972, 18, 345-367. Blough, D. S. Steady state data and a quantitative model of operant generalization and discrimination. Journal of Experimental Psychology: Animal Behavior Processes, 1975, 1, 3-21. Gillette, K., & Bellingham, W. P. Loss of within-compound flavour associations: Configural preconditioning. Experimental Animal Behaviour, 1982, 1, 1-17. Gormezano, I. Classical conditioning. In J. B. Sidowski (Ed.), Experimental methods and instrumentation in psychology. New York: McGraw-Hill, 1966. Heinemann, E. G., & Chase, S. Stimulus generalization. In W. K. Estes (Ed.), Handbook of learning and cognitive processes: Vol. 2. Conditioning and behavior theory. Hillsdale, N.J.: Erlbaum, 1975. Hull, C. L. Principles of behavior. New York: AppletonCentury-Crofts, 1943. James, J. H., & Wagner, A. R. One-trial overshadowing: Evidence of distributive processing. Journal of Experimental Psychology: Animal Behavior Processes, 1980, 6, 188-205. Kamin, L. J. Selective association and conditioning. In N. J. Mackintosh & F. W. K. Honig (Eds.), Fundamental issues in associative learning. Halifax, Nova Scotia: Dalhousie University Press, 1969. Kehoe, E. J. The role of CS-US contiguity in classical conditioning of the rabbit's nictitating membrane response to serial stimuli. Learning and Motivation, 1979, 10, 23-38. Kehoe, E. J., Feyer, A., & Moses, J. L. Second-order conditioning of the rabbit's nictitating membrane response as a function of the CS2-CS1 and CS1-US intervals. Animal Learning & Behavior, 1981, 9, 304315. Kehoe, E. J., & Gormezano, I. Effects of trials per session on conditioning of the rabbit's nictitating membrane response. Bulletin of the Psychonomic Society, 1974, 2, 434-436. Kehoe, E. J., & Gormezano, I. Configuration and combination laws in conditioning with compound stimuli. Psychological Bulletin, 1980, 87, 351-378.

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E. JAMES KEHOE ing produces S-S learning in conditioned suppression. Journal of Experimental Psychology: Animal Behavior Processess, 1982, 8, 23-32. Rescorla, R. A., & Cunningham, C. L. Within-compound flavor associations. Journal of Experimental Psychology: Animal Behavior Processes, 1978,4,267275. Rescorla, R. A., & Wagner, A. R. A theory of Pavlovian conditioning: Variations in the effectiveness of reinforcement and nonreinforcement. In A. Black & W. F. Prokasy (Eds.), Classical conditioning II. New York: Appleton-Century-Crofts, 1972. Saavedra, M. A. Pavlovian compound conditioning in the rabbit. Learning and Motivation, 1975, 6, 314326. Scandrett, J., & Gormezano, I. Microprocessor control and A-D data acquisition in classical conditioning. Behavior Research Methods & Instrumentation, 1980, 12, 120-125. Smith, M. C., Coleman, S. R., & Gormezano, I. Classical conditioning of the rabbit's nictitating membrane response at backward, simultaneous, and forward CSUS intervals. Journal of Comparative and Physiological Psychology, 1969, 69, 226-231. Sutherland, N. S., & Mackintosh, N. J. Mechanisms of animal discrimination learning. New York: Academic Press, 1971. Whitlow, J. W., Jr., & Wagner, A. R. Negative patterning in classical conditioning: Summation of response tendencies to isolable and configural components. Psychonomic Science, 1972, 27, 299-301. Wickens, D. D. Stimulus-response theory as applied to perception. In Learning theory, personality theory, and clinical research: The Kentucky symposium. New York: Wiley, 1954. Wickens, D. D. Conditioning to complex stimuli. American Psychologist, 1959,14, 180-188. Wickens, D. D. Compound conditioning in humans and cats. In W. F. Prokasy (Ed.), Classical conditioning. New York: Appleton-Century-Crofts, 1965.

Kehoe, E. J., Schreurs, B. G,, & Amodei, N. Blocking acquisition of the rabbit's nictitating membrane response to serial conditioned stimuli. Learning and Motivation, 1981, 12, 92-108. Mackintosh, N. J. A theory of attention: Variation in the associability of stimuli with reinforcement. Psychological Review, 1975, 82, 276-298. Mackintosh, N. J. Overshadowing and stimulus intensity. Animal Learning & Behavior, 1916,4,186-192. Mackintosh, N. J., & Reese, B. One-trial overshadowing. Quarterly Journal of Experimental Psychology, 1979, 31, 519-526. Marchant, H. G., Ill, & Moore, J. W. Blocking of the rabbit's nictitating membrane response in Kamin's two-stage paradigm. Journal of Experimental Psychology, 1973, 101, 155-158. Miller, R. G. Simultaneous statistical inference. New York: McGraw-Hill, 1966. Moore, J. W., & Stickney, K. J. Formation of attentional-associative networks in real time: Role of the hippocampus and implications for conditioning. Physiology and Behavior, 1980, 8, 207-217. Pavlov, I. P. Conditioned reflexes: An investigation of the physiological activity of the cerebral cortex (G. V. Anrep, trans.). London: Oxford University Press, 1927. Razran, G. Empirical codification and specific theoretical implications of compound-stimulus conditioning: Perception. In W. Prokasy (Ed.), Classical conditioning. New York: Appleton-Century-Crofts, 1965. Razran, G. Mind in evolution. New York: HoughtonMifilin, 1971. Rescorla, R. A. "Configural" conditioning in discretetrial bar pressing. Journal of Comparative and Physiological Psychology, 1972, 79, 307-317. Rescorla, R. A. Evidence for "unique stimulus" account of configural conditioning. Journal of Comparative and Physiological Psychology, 1973, 85, 331-338. Rescorla, R. A. Simultaneous associations. In P. Harzem & M. D. Zeiler (Eds.), Advances in analysis of behavior: Vol. 2. Predictability, correlation, & contiguity. New York: Wiley-Interscience, 1981. Rescorla, R. A., Simultaneous second-order condition-

Received September 18, 1981

Correction to Timberlake, Wahl, and King

In the article "Stimulus and Response Contingencies in the Misbehavior of Rats" by William Timberlake, Glenda Wahl, and Deborah King (Journal of Experimental Psychology: Animal Behavior Processes, 1982, Vol. 8, No. 1, pp. 62-85), the abscissa of Figures 1, 2, 5, and 6 were incorrectly labeled. In each case, the word DAYS should replace the word TRIALS.