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

3, 226-243

Copyright 1982 by the American Psychological Association, Inc. 0097-7403/82/0803-0226S00.75

Abstraction of Temporal Attributes

Warren H. Meek and Russell M. Church Brown University
Cross-modal transfer of classification rules for temporal intervals was demonstrated in four experiments. In Experiment 1, rats learned a temporal discrimination between a 2-sec and an 8-sec signal in one modality (vision or audition). There was positive transfer to a temporal discrimination between a 2-sec and an 8-sec signal in the other modality when the response rule was maintained, and negative transfer when the response rule was reversed. Experiment 2 demonstrated positive cross-modal transfer in a temporal generalization procedure. Experiment 3 demonstrated cross-modal transfer of both duration arid temporal location in a procedure in which rats were exposed to three successive signal durations. Experiment 4 demonstrated cross-modal transfer of both duration and temporal order in a procedure in which rats were exposed to simultaneously presented signal durations, one auditory and one visual. The conclusion is that rats can abstract temporal attributes from modality-specific aspects of a signal.

Some animal information processing may be mediated by modality-specific systems that are relatively independent. Other processing may disregard the specific qualities of a sense modality (e.g., audition, vision) in favor of a higher level analysis of nonspecific components. This involves, in part, the abstraction from the stimulus of certain basic attributes common to all stimuli, e.g., duration, intensity, location, and number. There are many examples of sensory equivalence in human subjects in which different stimulus energies (e.g., acoustical, chemical, and radiant) can carry the same stimulus information and control the same response (e.g., Freides, 1974). Although some of these cross-modal equivalences cannot be due to verbal coding since they are found in infants, language may be influential in many cases of human cross-modal transfer (e.g., Blank & Bridger, 1964; Bryant, Jones, Claxton, & Perkins, 1972; Koen, 1971). The evidence for cross-modal perforThis investigation was supported by National Science Foundation Grant BNS 79-04792. A report based on this work was presented at the Third Harvard Symposium on Quantitative Analysis of Behavior: Acquisition, Cambridge, Massachusetts, June 1980. The authors thank Fariba Komeily-Zadeh for her assistance in conducting the experiments. Requests for reprints should be sent to Warren H. Meek, Department of Psychology, Brown University, Providence, Rhode Island 02912.
226

mance in animals comes primarily from transfer studies in which a specific change of performance trained to some attribute of one sense modality is immediately evident during testing to the same attribute of another sense modality. Although cross-modal equivalence in animals has been a much studied topic, relatively few investigators have reported successful cross^modal transfer (e.g., Ettlinger, 1977; MacDonall & Marcucella, 1976; Over & Mackintosh, 1969; Seraganian & Popova, 1976; Ward, Silver, & Frank, 1976; Yeterian, Waters, & Wilson, 1976). Church and Meek (in press) reviewed the relevant literature and concluded that it is uncertain whether any experiment has unambiguously demonstrated cross-modal transfer in animals. The results from all of the studies examined may be accounted for in terms of mediation by experimenter-controlled bridging stimuli which are present during both training and transfer testing. Our interest in this study was to determine unequivocally whether rats are capable of abstracting temporal attributes from visual and auditory stimuli. The proposal is that stimulus information about such basic stimulus attributes as duration, intensity, location, and number may be identical when stimulus modalities are different. A simple discrete-trial choice procedure was used in Experiment 1 to demonstrate the

ABSTRACTION OF TEMPORAL ATTRIBUTES

227

cross-modal transfer of duration between intosh, 1969; Wilson & Shaffer, 1963). If vision and audition. To extend the generality there is cross-modal transfer of the attribute of the results, we used the basic design of of duration from one modality to another, Experiment 1 again in Experiment 2 with the percentage of correct responses of the a temporal generalization procedure. In Ex- nonreversal group would be above chance periment 3, rats were exposed to three suc- (50%), and the percentage of correct recessive signal durations and classified the sponses of the reversal group would be below sequence according to a complex combina- chance. The experiment employed random-duration rule. A cross-modal test was employed to demonstrate the abstraction of temporal tion intertrial intervals and a signal-omission location as well as duration. In Experiment test, since previous work has shown that 4, rats were exposed to simultaneously pre- fixed-duration intertrial intervals may parsented signal durations, one auditory and tially account for apparent cross-modal one visual. A cross-modal test was then em- transfer (Meek & Church, in press). Three ployed to demonstrate the abstraction of final test days were included to determine temporal order as well as duration. In over- the extent to which the classifications were view, these experiments focus on duration independent of modality. as a basic attribute. The results indicate that Method animals that have learned a particular temporal discrimination in one modality can Subjects and Apparatus abstract the temporal attributes from the The subjects were 20 experimentally naive male almodality-specific aspects of the signal. bino Norway rats (Charles River CD), about 10 mo old Experiment 1: Simple Choice Discrimination (Transfer of Signal Duration)
when the experiment began. Animals were individually housed in metal cages (Wahmann Mfg. Co., Baltimore, Maryland) in which water was continuously available. The rats were fed a daily ration of about 15 g of Charles River Rat Formula mixed with water immediately after the daily session. A light/dark cycle of 16:8 hr was maintained in the animal colony. All testing was conducted during the light phase of the cycle. The rats worked in 10 similar lever boxes (23 X 20 X 22 cm). Noyes Precision food pellets (45 mg) were delivered through an opening in the front wall to a food cup. A glass water bottle hung from the back wall of the chamber. Each box contained two retractable stainless steel levers, one on each side of the food cup. The levers in Boxes 1-6 were 1.6 X 4.6 cm, located 3.8 cm above the grid floor (Gerbrands Model 6311). The levers in Boxes 7-10 were 2.5 X 5.0 cm, located 5.0 em above the floor (BRS/LVE Model 123-07). Each lever box was housed in a large insulation-board chamber designed to minimize outside light or sound. Six boxes had a 7.5-W lamp attached to the middle of the back wall of the chamber; four boxes had a 6-W lamp attached to the outside of the roof of the lever box. A noise generator could be used to deliver white noise of about 80 dB (re 20 pN/m2; General Radio Sound Level Meter, Model 1565-D, A scale) above background level through a 4-in. (10.2-cm) speaker mounted inside each chamber. In Boxes 7-10, electric shock could be delivered to the grid floor through an autotransformer, a power transformer, and d 150-kfl resistor in series with the animal. In each insulation-board chamber, there was a fan for ventilation. A time-shared PDP-12 computer controlled the experimental equipment and recorded the responses.

In this experiment a discrete-trials choice procedure was used to measure any crossmodal transfer of specific learning between vision and audition. Rats were first trained to classify the duration of a signal in one modality as short (left response) or long (right response) with a procedure that leads to rapid temporal discrimination learning (e.g., Church & Deluty, 1977; Maricq, Roberts, & Church, 1981). Following this training with signals in one modality, animals were retrained with signals in a different modality (vision or audition). In order to obtain a sensitive indication of cross-modal transfer, half of the animals were given nonreversal cross-modal retraining with signals in the new modality (i.e., reinforcement of a left response following a short signal and reinforcement of a right response following a long signal), and half of the animals were given reversal cross-modal retraining with signals in the new modality (i.e., reinforcement of a left response following a long signal and reinforcement of a right response Procedure following a short signal). This same design Pretraining. Each rat received at least two sessions has been used by others (e.g., Over & Mack- of continuous reinforcement training. During these ses-

228

WARREN H. MECK AND RUSSELL M. CHURCH both modalities remained the same as during crossmodal training. Test Day 2 was identical to Test Day 1 except all response rules were reversed. Thus, the original nonreversal groups received their first reversal condition, and the original reversal groups received their second reversal condition. (Table 1 presents the conditions of reinforcement for each group in each phase of the experiment through Test Day 2.) Test Day 3 was identical to the first 7 days of retraining except (a) only eight subjects were tested, (b) signals were either a 2or an 8-sec duration of constant .15-mA electric footshock (rather than light or sound), and (c) the response rules were reversed from those used in retraining for half of the rats in each original modality group.

sions a pellet of food was delivered each minute for 30 min, and, in addition, each lever press produced food. The session began with the insertion of the left lever, and 10 responses were reinforced, after which the left lever was retracted and the right lever inserted. Ten right lever responses were then reinforced, followed by retraction of the right lever and insertion of the left lever. This alternation between levers continued until the rat had pressed each lever 30 times or 30 min had passed, whichever came first. The houselight illuminated the chamber at all times during the session. Training (Days 1-21 j. Half of the rats were trained to press the left lever (short response) following a light termination signal of 2-sec duration and to press the right lever (long response) following a light termination signal of 8-sec duration; the remaining half of the rats were trained in the same manner with sound onset as the signal. On each trial one of the two signal durations was randomly presented with a probability of .5. The houselight went off (or the sound went on) for the selected duration; at the end of this period of time, the houselight was turned back on (or the sound was turned off), and both levers were inserted into the box. If the animal made the correct response, a pellet of food was immediately delivered; if it made the incorrect response, no pellet was delivered. When either lever was pressed, there was a .5-sec delay and then both levers were retracted. After an intertrial interval of 5 sec plus a geometrically distributed duration with a minimum of .1 sec and a mean of 40 sec, another trial was begun. On Days 1-5, if an incorrect response had been made on the previous trial, the same stimulus duration was presented again on the next trial (correction procedure). From Day 6 until the end of the experiment, there were no correction trials. A daily session began within 30 min of the same time each day and lasted for 1 hr 50 min. A record was kept of the number of left and right responses following each of the two signals. Cross-modal training (Days 22-28), Each group of rats (light and sound) was evenly divided into nonreversal and reversal groups in terms of their originally trained response rules. The first 20 trials of the first day of cross-modal training consisted of the same procedure used on Days 6-21. No data were collected on these trials. Thereafter, rats were retrained with 2-sec and 8sec signals from a new modality (light or sound). Half of the rats were tested with the same response rule (reinforcement of a left response following a 2-sec signal and reinforcement of a right response following an 8sec signal), and half were tested with a reversed response rule (reinforcement of a right response following an 8sec signal and reinforcement Of a left response following a 2-sec signal). Signal-omission training (Days 29-35). The procedure was the same as that used in cross-modal training except the signal was omitted. Retraining (Days 36-42). The rats received further signal training for 7 days under the same conditions as cross-modal training. Then there were 3 test days. Tests (Days 43-45). These 3 days are referred to as Test Days 1, 2, and 3. On Test Day 1, half of the rats had light signal training during the first 1.5 hr and sound signal trials during the last 1.5 hr of a 3-hr session. The remaining half of the rats had the order of light and sound signal training reversed. The response rule for

Results The performances of the nonreversal and reversal light groups were similar throughout training (see Table 1). During the last seven sessions of training, the mean percentage of correct responses of rats in the nonreversal group trained with light was not significantly different from the mean percentage correct of rats in the reversal group trained with light. The performances of the nonreversal and reversal groups trained with sound were also similar. The percentage of correct responses of rats in the groups trained with sound, however, was significantly higher than the percentage of correct responses of rats in the groups trained with light, r(18) = 4.35, p<. 001. Cross-Modal Training The second column of data in Table 1 shows the mean percentage of correct responses during the first session of crossmodal training. The nonreversal group transferred to sound (and trained with light) had a mean percentage correct significantly greater than chance, t(4) = 6.0, p < .01. The nonreversal group transferred to light (and trained with sound) also had a mean percentage correct significantly greater than chance, r(4) = 3.03, p <, .05. In both cases the transfer was incomplete since the rats had a significantly higher mean percentage correct on the previous seven sessions, f(4) = 2.77, p< .05 and *(4) = 11.9, p < .001. During the same session the reversal group transferred to sound (and trained with light) had a mean percentage correct significantly less than 50%, f(4) = 7.4, p < .01. The reversal group transferred to light (and trained

ABSTRACTION OF TEMPORAL ATTRIBUTES

229

with sound) had a mean percentage correct not significantly less than 50%, t(4) = 1.5, p < .10, but significantly different from the comparable nonreversal group, f(8) == 3.0, p < .05. Figure 1 shows the percentage correct for individual subjects in each group averaged over responses to both 2-sec and 8-sec durations for the first session of cross-modal training. Figure 2 shows that the difference between nonreversal and reversal groups was maintained for at least five sessions of retraining, fs(8) > 2,36, ps < .05. The reversalnonreversal differences for the seven sessions of cross-modal training were greater for rats transferred to sound than for rats transferred to light. An analysis of variance of the percentage of correct response during crossmodal training, with factors reversal-nonreversal, signal (light or sound), and session,
Table 1

showed all three main effects to be reliable: reversal-nonreversal, F(l, 120) = 77.0, p < .001; signal, F(l, 120) = 13.5, p< .001; and session, F(6, 96) = 52.2, p < .001. The session effect was more pronounced for the reversal group than the nonreversal group, F(6, 96)= 17.1, p<.001, and the signal difference was more pronounced for the nonreversal group than the reversal group, F( 1, 120) = 6.8, p< .025. During the first session of cross-modal training, rats transferred to light had a higher proportion of correct responses following short signals than long signals (79 5 vs. 36 6), t(9) = 5.91, p< .001. No such difference was obtained for rats transferred to sound. The proportions of correct responses following short and long signals were 66 7 and 72 6, respectively, a nonsignificant difference, t(9) < 1. The absolute difference between accuracy to short and

Mean (SE) Percentage Correct Responses as a Function of Group and Phase

Phase
Group Nonreversal lightsound Condition % correct Nonreversal soundlight Condition % correct Reversal light-sound Condition % correct Reversal sound-light Condition % correct Training (Days 15-21) Crossmodal (Day 22) Signal omission (Day 29) Retraining (Days 36-42) Test Day 1 (Day 43) Test Day 2 (Day 44)

light (LR)
83 2

sound (LR)
74 4

-(LR)
51 2

sound (LR)
95 2

light (LR) sound (LR) light 84 2 sound 97 3

light (RL) sound (RL) light 44 2 sound 39 2

sound (LR)
93 1

light (LR)
60 3

-(LR)
49 2

light (LR)
86 3

light (LR) sound (LR) light 88 .3 sound 97 2 light (RL) sound (RL) light 51 2 sound 95 3

light (RL) sound (RL) light 39 2 sound 35 2 light (LR) sound (LR) light 68 2 sound 37 2

light (LR)
84 4

-sound (RL)
36 2

-(RL)
48 2

sound (RL)
90 2

sound (LR)
93 2

light (RL)
44 4

-(RL)
51 2

light (RL)
83 + 3

light (RL) light (LR) sound (RL) sound (LR) light 88 2 light 41 2 sound 47 2 sound 70 2

Note. LR refers to reinforcement of left response after 2-sec signal and reinforcement of right response after 8sec signal; RL refers to reinforcement of right response after 2-sec signal and reinforcement of left response after 8-sec signal.

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WARREN H. MECK AND RUSSELL M. CHURCH

SOUND

GROUP

Figure 1. Percentage correct for individual subjects during the first session of cross-modal testing for subjects with nonreversal and reversal classification rules. (Figures were drawn with a computer program developed by Hayes, 1981.)

The results from Test Day 2 are given in the last column of Table 1. All original nonreversal groups had levels of performance below chance for both light and sound, ts(9) ;> 2.41, p < .05. The original reversal groups also had a level of performance below chance for the retrained signal modality, ?(9) = 2.87, p < .02. In contrast, performance on the originally trained signal modality was significantly above chance, t(9) = 2.93, p < .02. With the exception of an initial period in which no responding occurred, the results with electric shock as a signal (Test Day 3) were similar to those for previous crossmodal training. Combining over rats originally trained with light or sound signals, the nonreversal groups were 64% 3% correct, a level of performance significantly above chance, r(3) = 6.37, p < .01; the reversal groups were 36% 2% correct, a level of performance significantly below chance, f ( 3 ) = 11.2, p < .01. Discussion The major finding of Experiment 1 was the positive and negative cross-modal transfer of a specific duration discrimination. This result replicates previous reports from our laboratory (Meek & Church, in press; Rob-

accuracy to long signals of rats transferred to light was reliably greater than for rats transferred to sound, f(18) = 3.72, p< .01. The third column of data in Table 1 shows the mean percentage of correct responses on the first day of signal omission for the four groups of rats. The percentage of correct responses during the 7 days of signal omission was not reliably different from chance on any of the days, ts(9) < 1During retraining, the pattern of results observed during training was replicated, i.e., the percentage correct for sound was greater than for light. The results from Test Day 1 are given in the next to last column of Table 1. The nonreversal groups had a percentage of correct responses well above chance for both light and sound, ts(4) > 3.77, p < .02. In addition, performance to sound signals was better than performance to light signals, *(9) = 4,1, /x.OOl. The reversal groups also had a percentage of correct responses for the retrained signal modality that was significantly greater than chance ts(4) ^. 3.78, p < .02. The major result was that the performance of the reversal groups on the originally trained signal modality was not significantly different from chance for either light or sound, fs(4) < 1.

NONREVERSAL

SOUND LIGHT 0---0- SOUND A---A LIGHT

DAY

Figure 2. Percentage correct as a function of days of cross-modal transfer testing for groups with nonreversal and reversal classification rules. (The stimulus used in the cross-modal transfer test [sound or light] is listed in the legend.)

ABSTRACTION OF TEMPORAL ATTRIBUTES

231

erts, 1982). Both positive and negative transfer of duration were demonstrated during the first session of cross-modal training, and the difference between nonreversal and reversal groups persisted for five sessions. The relatively long random intertrial in^ terval eliminated the possibility that the intertrial interval mediated the transfer effect since no control by time was observed when the signal was omitted. The amount of transfer was greater, and the transfer lasted longer than the cross-modal transfer by animals previously reported (e.g., Over & Mackintosh, 1969). The fact that the positive transfer was somewhat greater than the negative transfer probably reflects some new learning that occurred during the first session. The results with sound and light signals differed in three ways: (a) The asymptotic temporal discrimination with sound was significantly greater than the asymptotic temporal discrimination with light during both training and retraining, (b) transfer from light to sound was greater than from sound to light, and (c) rats transferred to light, but not those to sound, made more erroneous short responses than long responses. Similar differences between light and sound signal durations have been obtained with human subjects (e.g., Goldstone & Goldfarb, 1966). Posner (1978), in describing human information processing, proposed that under some conditions auditory signals will have automatic access to information analyzers and visual signals will not. If automaticity contributes to a high level of performance, the asymptotic level of performance might generally be higher for sound than light. Of course, the intensity of stimulus change is also a relevant variable. According to Posner's proposal, to initiate visual processing, subjects must learn to direct attention to the visual stimulus; no such learning is required for auditory processing. Thus subjects transferred from light to sound would have greater transfer than subjects transferred from sound to light. Finally, according to Posner's proposal, the time to process a visual signal is longer and more variable than the time to process an auditory signal. If this were the case in the rats trained with sound signals and transferred to light signals, the

light would be slower to initiate the timing process. This would produce a shorter subjective time, which results in the observed asymmetrical error rates for the sound-light group, i.e., more short than long errors. Other work (Meek, Note 1) has used different tests to demonstrate that there are fundamental differences in the temporal processing of light and sound signals by rats. The results of Test Days 1 and 2 for the nonreversal groups are simply replications. On Test Day 1, the rats continued to perform the trained responses; on Test Day 2, they had negative transfer when the response rule was reversed. The important results from the test days were obtained from the original reversal groups. During Test Days 1 and 2, these groups differentially responded to light and sound signals, although the same reinforcement contingency applied to both signals. Performance could be predicted by the signal modality and response rule originally trained. Such performance indicates that sensory-specific memories were established by the original training conditions and continued to persist even following cross-modal transfer. This suggests that sensory-specific information is maintained in conjunction with an attribute abstraction system. Animals may utilize this information network by matching sensory-specific and nonspecific associations with the stimuli currently available. This finding is consonant with work in human cognition wherein people can recall sensory-specific information although the emphasis of the task is on abstracting information that is equivalent between modalities (e.g., Broadbent, 1958; Posner, 1978). The results from Test Day 3 with electric shock provided another example of crossmodal transfer. The main purpose of this test was to determine whether rats that experienced negative transfer (rats in the reversal groups) learned not to transfer a response rule to a new modality. The result was that the animals continued to use the response rule last trained when transferred to the new electric shock signal. In summary, the major finding from this experiment was cross-modal transfer of a specific duration discrimination. In addition, (a) discrimination performance was better for sound than for light signals, (b) transfer

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WARREN H. MECK AND RUSSELL M. CHURCH

the rat had received 60 reinforcements or 30 min had passed, whichever came first. For the next 2 days, rats were given discrete trials of a signal presentation (houselight off) followed immediately by lever insertion and a random-interval schedule of reinforcement with a 3-sec limited hold. Reinforcement intervals were 1 sec plus a geometrically distributed duration with a minimum of. 1 sec and a mean of 5 sec. The lever retracted following the reinforced response. The reinforced signal duration was 4 sec for half of the rats (4-sec group) and 12 sec for the others (12-sec group). Intertrial intervals were 10 sec plus a geometrically distributed duration with a minimum of . 1 sec and a mean of 30 sec. The rats were tested in two sessions of 10 rats each. They were randomly assigned; 5 in each session were from the 4-sec group and 5 from the 12-sec group. Sessions began within 30 min of the same time each day and lasted 1 hr'50 min. Training with unreinforced signal durations (Days 1-40). Training was the same as the final condition of pretraining except that unreinforced signal durations of 2 and 6 sec for the 4-sec group and 6 and 18 sec for the 12-sec group were also presented. The reinforced duration was presented on a random half of the trials, and the two unreinforced durations were randomly presented with equal probability on the remaining trials. The response lever was withdrawn following nonreinforced signal trials according to the rule used for reinforced signal trials except that no reinforcement was given. Beginning on Day 6 of training, the response lever was present at all times during the session. Cross-modal training (Day 41). Rats trained with a light-oft* signal were tested with a sound-on signal. Except for the change in modality, the procedure remained the same as during original training. Data taken. Responses were recorded as a function of time for the first 5 sec of the random-interval reinforcement schedule. A mean response rate was calculated for these 5-sec intervals.

from light to sound was greater than from sound to light, and (c) although the attribute of duration may be used independently of stimulus modality, modality-specific information is also available for use. Experiment 2: Temporal Generalization (Transfer of Duration) To demonstrate the generality of crossmodal transfer of duration, we attempted to replicate the positive transfer results of the previous experiment with a different timing procedure, a temporal generalization task. In this procedure only one lever was used, and except for early stages of training, it remained in the box. Rats were given training with three durations of a light signal. Half of the rats received signals of 2, 4, or 6 sec, and half received signals of 6, 12, or 18 sec. If the signal was the intermediate one, which occurred on a random half of the trials, reinforcement was primed after a random interval. If the signal was either of the extreme durations, there was no reinforcement. The measure was the response rate during the first 5 sec after signal termination. A similar procedure has been used extensively in our laboratory (e.g., Church & Gibbon, 1982), arid it leads to slow temporal discrimination learning. This procedure differs from the one used in Experiment 1 in several respects. In the choice discrimination procedure, the animal reports which of two remembered signal durations is closer to the presented signal by making one of two discrete responses; in the temporal generalization procedure, the animal reports whether the remembered signal duration is close enough to the presented signal by varying its response rate. Method Subjects and Apparatus
Twenty rats, similar to those used in Experiment 1, were used in this experiment. The apparatus described in Experiment 1 was used.

Results Figure 3 (left panel) shows mean response rate (responses/min) for subjects in the 4sec group during the last 5 days of training (light signal) and the single test day (sound signal) at each signal duration. The response rate following the reinforced signal (/?) and the mean of the response rates following the two unreinforced signals (/) were combined in a discrimination index, (JR. U)/R. A discrimination index of 0 would reflect chance performance; higher values would reflect better discrimination. During the last 5 days of training, the mean discrimination ratio for subjects in the 4-sec group was .23% .03%; during the 1 day of cross-modal training, this discrimination index for the nine subjects that responded was also .23% .03%, an unreliable difference between training and testing.

Procedure
Pretraining. Each rat received at least two sessions of continuous reinforcement training. During these sessions a pellet of food was delivered each minute for 30 min, and, in addition, each lever press produced food. Only the left lever was used. The session ended when

ABSTRACTION OF TEMPORAL ATTRIBUTES j

233

Figure 3 (right panel) shqws mean re- the maximum remained at the reinforced sponse rate for rats in the 12-sec group dur- value with no change in the accuracy of the ing the last 5 days of training (light signal) discrimination. The reduction in response and the single test day (sound signal) at each rate shows that there was some influence of signal duration. During the last 5 days of modality, but the forpi of the generalization training, the mean discrimination index for functions indicates that rats that learned a subjects in the 12-sec group 'was .21% temporal criterion (4 or 12 sec) when the .04%. During the 1 day of cross-modal train- internal clock was driven by a light signal ing, this discrimination index for the seven used the same criterion when the clock was subjects that responded was .23% .03%, an driven by a noise signal. For this temporal unreliable difference between training and generalization task, as for the previous task, any stimulus change (e.g., light or sound) testing. During training, the discrimination mea- was sufficient to start an internal clock, and sure for both 4-sec and 12-sec groups was the duration registered by the clock was a reliably greater than chance, ts(9) s= 7,0, major determinant of performance. ps < .05, but the difference between these two groups was not significant, f(18)<l. Experiment 3: During cross-modal training, the accuracy Conditional Discriminations (Transfer of subjects that responded in all groups was of Duration and Temporal Location) reliably greater than chance, *s(15) a 2.36, In the previous two experiments, the anps < .05, and there were no reliable differences in accuracy between training and test- imals had only to form a simple association ing, ?s(15) < 1. However, the response rate between a duration and a response. In Exof both groups was lower during cross-modal periment 1, a "long" response was associated training than during original training, f(6) = with signal durations greater than some criterion; in Experiment 2, rapid response rates 2.64, p < .05 and f(8) = 2.33, p < .05. were associated with signal durations close enough to some criterion/These simple asDiscussion sociations between signal durations and reDuring testing with the new modality, sponses transferred from one modality to there was a reduction in response rate, but another (e.g., vision to audition).

80

60

5
UJ
-
o

TRAIN (LIGHT) TEST (SOUND)

20

-----

12TIME ( S E C )

18

Figure 3. Average number of responses per minute as a function of signal duration under conditions of original training and cross-modal testing. (Left panel shows data for the group trained with responses reinforced after 4-sec signals; the right panel shows data for the group trained with responses reinforced after 12-sec signals.)

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WARREN H. MECK AND RUSSELL M. CHURCH

Table 2 Signal Duration, Reinforced Response, and Reinforcement Rule

Signal duration (in sec): Reinforced response Larger (A/C) or (CIA)

Method Subjects and Apparatus

Twenty rats similar to those used in Experiment 1 were used in this experiment. The apparatus described in Experiment 1 was used.

Rule \A-C\ + 1

c 1
3 3 3

Procedure
Pretraining. The pretraining described in Experiment 1 was used. Same-different training (Days 1-50). Half of the rats were trained on a same-different conditional discrimination. Signals consisted of three segments with durations, A, B, and C. A white noise signal was on for duration A, off for duration B, and on again for duration C. The durations A, B, and C could be 1, 2, or 3 sec, making a total of 27 possible signal combinations. For this experiment, only 18 of the 27 possible cases were used; those signals having segment C = 2 were omitted (see Table 2.) On each trial a signal combination was presented. Then both levers were inserted into the box. If the rat pressed the correct lever, food was delivered; when either lever was pressed, there was a .5-sec delay and then both levers were retracted. After a 30-sec intertrial interval, another trial was begun. Signals were randomly presented with equal probability. The houselight was on at all times during the session which began within 30 min of the same time each day and lasted for 1 hr 50 min. A record was kept of the number of left and right responses following each signal. In the same-different conditional discrimination, the left (same) response was reinforced if \AC\ - 0 sec, and the right (different) response was reinforced if \A-C\ = 1 sec. Intermediate values, in which \AC\ = 1, were not reinforced. (Table 2. states the experimenter's classification rule with the addition of a constant, 1, to constrain the variable between 1 and 3 sec.) During the training phase, a left response was reinforced if A and C were both 1 or 3 sec; a right response was reinforced if A = 3 and C = 1 or if A = 1 and C = 3. The duration of B was irrelevant. If the rat made the incorrect response, no pellet was delivered, and the same signal was presented again on the next trial (correction method). Testing on a new subset of signals (Days 51-60). The procedure was similar to that previously described except that a new subset of 11 unreinforced signals was added, with duration B = 2 sec. From Day 51 to the end of the experiment, there were no correction trials. Table 2 shows the signal presentations and experimenter's classification rules used for this stage of testing. Retraining (Days 61-65). Training was the same as on Days 1 -50 except that no correction trials were given. Oddity training (Days 1-65). Half of the rats were trained on an oddity conditional discrimination. The training procedure was similar to that of the same-different discrimination except as described below. In the oddity conditional discrimination, the segment classification rules were such that if A = B, classify C; ifA = C, classify B; if B = C, classify A. If the segment to be classified (the odd segment) was 1 sec, the left response (short) was reinforced; if the segment was 3 sec, the right response (long) was reinforced. Intermediate values, in which the odd segment equaled 2 sec,

Same-different training (Days 1-50 and 61-65)

1 2 3 1 2 3

X X X
X X X

1 2 3 3 2 1

X X X X X X

\ 2 3
3 1.5 1

New subset training (Days 51-60) 2 4 5 4 2 5 3 5 2 6 2 2 2 2 2 2 2 2 2 2 2 2 2

4

5 2
4

3 5 2 5 2 6

1 1 1 3 3 3 3 4 4 5 5

2 1 1 2 2 1 2 2 2 2 2 1.67 2 , 1.67 2 2.5 2 2.5 2 3 2 3

Cross-modality training (Days 66-75)

1 2 3 1 2 3 1 X 3
3 X 1

1 1 1 3
3

L R R L

1 2 3 3 2 1

1 X 3 3
X 1

1 2 3 3
1.5 1

Note. X = 1, 2, or 3 sec.

Previous research with human subjects has typically obtained cross-modal transfer in tasks involving the processing of simple unidimensional discriminations but modality-specific learning in tasks involving complex pattern information processing (e.g., Freides, 1974), In the present experiment, the animals had a much more difficult task than in the first two experiments. A sequence of three signals of various durations was presented. The animals had to select which elements of the sequence to time, and then to use a conditional classification rule. The question was whether, rats would transfer a complex classification rule involving several sequentially presented durations from one modality to another.

ABSTRACTION OF TEMPORAL ATTRIBUTES were not reinforced. The durations of A, B, or C could be relevant. If the rat made the incorrect response, no pellet was delivered, and the same signal was presented again on the next trial (correction method). From Day 51 to the end of the experiment, there were no correction trials. Cross-modality training with a new modality (Days 66-75). Five subjects from each of the two groups (same-different and oddity) continued and were identically treated in this phase of the experiment. There were two major changes from previous training: (a) The modality of the signal was changed from sound onset to light termination, and (b) there were two classification rules that would work. The rats could use the previously described same-different rule based on a comparison of the duration of A and C, or, more simply, they could use the duration of B. If B = 1 sec, a left response rule was reinforced; if B = 3 sec, a right response was reinforced. Intermediate values, in which B = 2 sec, were not reinforced for either response. Table 2 shows the signals and the experimenter's classification rules used for cross-modal training.

235

.01; but there was no relation between the percentages of right responses as a function of the duration of B, which was an irrelevant stimulus, F(2, 4) < 1.

Testing With a New Subset of Signals During the previous phase the animals were responding to a comparison between duration A and duration C, but were the animals dealing with differences or ratios? The data from the previous phase do not distinguish between these alternative classification rules since a left response was reinforced following an absolute difference of 0 (or a ratio of 1) and a right response was reinforced following an absolute difference of 2 (or a ratio of 3). The new subset, however, unconfounds the ratio and the difference. During the first 5 days of testing with Results these additional signals, the median percentage right (different) when \AC\ + 1 = Same-Different Training 1 sec was 3 + 2; when |/4-C| + 1 = 3 sec, The same-different discrimination was not it was 25 3; when \A-C\ + 1 = 4 sec, it easy to learn. After 50 sessions, only 3 of the was 53 1; and when \A-C\ + 1 = 5, it was 10 rats learned the task. The results from 71 1. These results indicate a significant these 3 subjects for the last 5 days of re- effect of the value of \A-C\, F(3, training are shown in the left panel of 6) = 215, p < .01. Figure 4. The data may also be analyzed in terms A single factor (within-subjects) analysis of the ratio of the longer segment to the of variance indicated a significant relation shorter segment. Figure 5 displays the mebetween the percentages of right (different) dian percentage of right (different) reresponses as a function of the absolute dif- sponses as a function of this ratio. The best ference between A and C, F(2, 4) = 49, p < fitting straight line accounts for 95% of the
,
100
LIGHT LIGHT

. LU u

50

s?

TIME ( SEC )
Figure 4. Percentage of right ("different") responses as a function of the value of two classification rules, a comparison of the first and third interval (\AC\ + 1 ) and the duration of the second interval (B). (The left panel shows data for original training with sound; the middle panel shows data for the first five sessions of cross-modal transfer with light; the right panel shows data for the next five sessions of cross-modal transfer.)

236
100
I CD t 50

WARREN H. MECK AND RUSSELL M. CHURCH

2
RATIO

Figure 5. Percentage of right responses as a function of the ratio of segments A and C: Longer to shorter. (Open circles represent the training cases; filled circles, the subset of testing cases.)

variance in the median animal. If the ratio of the shorter segment to the longer segment is used, the relation is less linear, so the best fitting straight line accounts only for 75% of the variance in the median animal. The Oddity Training ratio of the percentage variance unacNone of the 10 rats showed evidence of counted for by a straight line with the long/ learning the oddity conditional discriminashort measure was significantly less than the tion. The last 5 days of their training are percentage variance unaccounted for by a straight line with the short/long measure, shown in the left panel of Figure 6. The relation between the percentage right response t(2) = 21.6, p < .01. The critical comparison between the ab- and the duration of the odd segment or of solute difference and the ratio accounts re- segment B was unreliable, Fs(2, 18) < 1. lating the temporal values of A and C to the response function is whether there is a sig- Cross-Modality Training nificant difference between performances at The oddity group's results for the first and signal ratios of 1.67 and 2.0. The values of second set of 5 days of transfer to light are A and C, which distinguish between these shown in the center and right panels of Figtwo ratio values, are not separable by the ure 6, respectively. In this phase the rats absolute difference account since 5 3 + 1 could classify the signal either by \A-C\ or and 4 2 + 1 both equal 3 sec, but the ratios by B. For the first set of 5 days, the relation of 5:3 and 4:2 are 1.67 and 2.0, respectively. between the signal classification \A C\ and A comparison of these two ratios revealed percentage right response was unreliable, a significantly greater percentage of right (different) responses for the 4:2 ratio than F(2, 8) < 1; In contrast, the relation between the duration of segment B and percentage the 5:3 ratio, f(2) = 6.23, p < .05. right response was reliable, F(2, 8) = 42, p<.0l. Cross-Modality Training The mean percentage right response durThe performance of the same-different ing the second set of 5 days had a pattern group during the first and second set of 5 similar to that of the first set of 5 days. There days of transfer to light are presented in the was again an unreliable relation between the center and right panels of Figure 4. In this percentage right response and the signal phase the rats could use the same-different classification \A C\, F(2, 8)<1, whereas rule with A and C, or they could use the there continued to be a reliable relation beduration of B to determine whether a left or tween the percentage right response and

right response would be reinforced. When transferred to the light signal, they continued to use the same-different rule in light during the first five sessions, even though the simple classification of B also provided the necessary information. The results from the three subjects that learned the same-different rule showed a reliable relation between percentage right (different) response and the classification of \A-C\, F(2, 4) = 25, p < .01. In contrast, there was an unreliable relation between responding aitd the durations of segment B, F(2, 4) < 1. During the second set of 5 days, the relation between the percentage right response and the signal classification \AC\ declined, F(2, 4) = 1.37, p > .05. In contrast, the relation between the duration of segment B and right responses increased, F(2, 4) = 39,

ABSTRACTION OF TEMPORAL ATTRIBUTES

237

,
X O

'10.0
B
-

SOUND

LIGHT
B
s'

LIGHT

a:
1 - 50
1 It

- -

~
ODD

/'
s 0

yr '

IA-CI+I '
s

u a: 1i 1
L-U

LU

&
. 1 1

D_

_
1 ' 1 1
1

TIME (SEC)
Figure 6. Percentage of right ("long") responses as a function of the value of two classification rules, a comparison of the "odd" interval during training with sound and the duration of the second interval (B). (The left panel shows data for original training with sound; the middle panel shows data for the first five sessions of cross-modal transfer with light where the duration of [|/1-C| + 1 ] and [B] were now both relevant; the right panel shows data for the next five sessions of cross-modal transfer.)

the duration of segment B, F(2, 8) =47, p < .01. Discussion The design of this experiment is the same as that of the well-known Luchins' water-jug problem which is used to demonstrate set formation in human problem solving (Luchins, 1942). Such a set may become habitual or mechanized, even to the extent of producing deleterious effects. To illustrate, a subject is first given a task that can be solved only with a complex rule, e.g., M-C|, and then given a series of problems that can be solved either by the same complex rule or by a simple rule, e.g., B. Subjects with the prior training continue to use the complex rule, evidence of set formation; subjects without the prior training use the simple rule. The additional factor in the present experiment is that there was a change in the modality of the signal from sound to light. Would a rat that learned a complex classification rule for a three-segment sound signal continue to use that rule for a three-segment light signal when it could use either the complex rule or a simple one? In the cross-modal training phase, the rat could either use the same-different rule on A and C, or it could use the duration of B to determine whether

a left or a right response would be reinforced. The results from the first five sessions of transfer show that the rats were using the complex rule \A-C\ in sound. When transferred to the light signal, they continued to use the same rule in light, even though the simple rule involving only the duration of B was sufficient. The results from the next five sessions of transfer show that with added retraining, the rats adopted the simple, rule, i.e., their behavior was controlled by the duration of fi, not the complex rule. In this case, the animals acquired by incidental learning the simple rule B while they were performing the more complex rule |/4-C|. We have typically found that rats eventually select the simplest reliable classification rule. A group of five subjects, which was unable to learn the complex oddity rule for sound, was transferred to a three-segment light signal that could be classified either by \A-C\ or by B. The duration of B controlled their behavior, and they showed no evidence of control by |/4-C|. Thus, the B rule is simpler than the \A-C\ rule. Our conclusion is that animals learned a complex timing classification with one modality (in this case, sound) and transferred it to a different modality (in this case, light). How the animals accomplished this task is

238

WARREN H. MECK AND RUSSELL M. CHURCH

uncertain. They could have been responding to a single clock value that was controlled by durations A and C, or they could have been responding to the relation between two remembered times, A and C. The single clock analysis could work for the data presented in Figure 5; for example, a clock that ran during A and C would register 4 sec when the "different" response was reinforced, and 2 or 6 sec when the "same" response was reinforced. Or a single clock that increased during A and decreased during C would register 0 sec when the "same" response was reinforced and a non-zero number when the "different" response was reinforced. This single clock reading, however, could not account for the results from the new subset of unreinforced test signals. The median percentage right (different) response was not ordered in the same way as the sum of A and C or consistently with the absolute difference between A and C; it was ordered in the same way as the ratio of the longer segment (A or C) to the shorter segment (A or C). This suggests that the animals were not using a single summing or differencing clock but were remembering two temporal intervals and using a ratio comparison rule. If equal changes in the ratio produce equal changes in performance, the rats were using the ratio of the larger to the smaller, rather than the smaller to the larger. The scalar timing theory applied to such tasks as timeleft (Gibbon & Church, 1981), temporal generalization (Church & Gibbon, 1982), and bisection (Gibbon, 1981) requires that comparisons are made by such ratios, rather than differences. Experiment 4: Simultaneous Temporal Processing (Transfer of Duration and Temporal Order) In the previous experiment, rats chose a response after exposure to three successive signal durations; in this experiment they had to choose a response after exposure to simultaneously presented signal durations. Rats were trained on the peak procedure, a modification of a fixed-interval schedule of reinforcement. In this procedure, a signal occurs, and on some trials food is primed after a fixed interval; on other trials no food

is primed, and the trial lasts for a relatively long time after the fixed interval. As a result, the response rate increases as a function of time since signal onset (as it does during normal fixed-interval training), but after the fixed interval has passed, the response rate decreases in a fairly symmetrical fashion. The interval during which the response rate is maximal is called the "peak time" and serves as a measure of when the food is maximally expected. The peak time occurs near the time that the food may be primed. The response rate at the peak time is called the "peak rate." Roberts (1981) showed that peak time and peak rate are independent measures of performance and that the symmetry of the response distributions when plotted in terms of seconds suggests that subjective time is linear in real time. In addition to the peak procedure, brief (1-sec) signals occurred at regular intervals during a trial. These signals produce a temporary decrease in response rate in a manner such that the overall positively accelerated function contains miniature segment functions of a similar form. We have previously described this "double scallop" effect (Meek & Church, Note 2) and have demonstrated by various signal manipulations that this response patterning is evidence for the simultaneous temporal processing ;of segment and overall intervals. In the present experiment, light onset was used to signal the 50-sec overall interval, and noise onset was used to signal the 10-sec segment intervals. In cross-modal testing, the signal modalities were reversed. If modalityspecific effects were dominant, the light would continue to signal a 50-sec interval, and noise would continue to signal a 10-sec interval. It is possible, however, that major control is from the signal duration and order. In this case, the first sustained signal (light or noise) would be used for a 50-sec temporal discrimination, and the later phasic signals (light or noise) would be used for a 10-sec temporal discrimination. Method Subjects and Apparatus
Five rats, similar to those used in Experiment 1 but with previous experience in a task similar to that of Experiment 3, were used in this experiment. The apparatus described in Experiment 1 was used.

ABSTRACTION OF TEMPORAL ATTRIBUTES

239

Procedure
Fixed-interval (FI) pretraining (Days 1-15). The rats were given discrete trials of FI 50-sec reinforcement signaled by houselight on. On each trial the houselight was turned on and remained on until food was primed 50 sec after trial onset. Once primed, food was given immediately following the rat's next response. There was an intertrial interval of 130 sec with houselight off between the delivery of food and the beginning of the next trial. The response lever was extended and operative at all times. All sessions lasted for a total of 3 hr, plus any further time needed to complete a trial in progress. Sessions were conducted 6 days a week and began within 30 min of the same time each day. Training (Days 16-40). The FI 50-sec trials were the same as those described for Days 1-15 except that intervals were segmented with 1-sec response-independent presentations of white noise occurring every 10 sec. Noise signals occurred during Seconds 1, 11, 21, 31, and 41, but not during Second 51 (so that the noise signal was not paired with food). This type of procedure is known as a second-order schedule (Kelleher, 1966). On a random half of the FI 50-sec trials containing brief stimuli, food was not primed at 50 sec, and the light remained on for 130 sec. Noise signals continued to occur during Seconds 61, 71,81, 91, 101, 111, and 121. Another trial began following a 130-sec intertrial interval. Cross-modal transfer of training (Days 41-45). This procedure was the same as the training procedure on Days 16-40 except that signal modalities were .reversed for a random half of the no-food trials. Rats trained with the onset of a sustained light signaling the FI 50sec schedule of reinforcement and segmented by brief 1-sec presentations of noise indicating an interval of at least 10 sec without food were now switched to noise signaling the FI 50-sec schedule of reinforcement and segmented by 1-sec presentations of light onset. Data taken. Responses were collected in 1-sec time intervals for both types of nonreinforced FI 50-sec trials containing brief stimuli. Data analysis. For each rat a response rate function was calculated for Seconds 1-100 after combining response intervals over the 5 days of testing. The peak time of this function was calculated with the method used by Roberts (1981). First, the "median;1 of the function was found, the median being the time where half of the responses were earlier and half later. Second, a new median was computed over the interval of which the first median is the center. If the first median was x sec, and x < 50, the next interval would be Sec 0 to Sec 2x; if x > 50, the interval would be Sec 2(x - 50) to Sec 100. If the second median was within .05 sec of the first, the calculation stopped; the second median is the peak time. If the second median was not within ,05 sec of the first, the calculation would continue. A third median would be computed over the interval defined by the second and so on. Peak rate was defined as the response rate at the peak time. The discrimination index was (75 percentile 25 percentile)/50 percentile, where the various percentiles weere computed over the interval defined by the peak time (=50 percentile). If the response-rate function is flat, this measure equals one; better discrimination corresponds to higher numbers. A

70

TIME ( S E G )
Figure 7. Average number of responses per minute as a function of time since trial began. (The top panel shows nonfood trials identical to those used in training [continuous light with food primed after 50 sec, and 1sec sound signals at 10-sec intervals shown by arrows]; the bottom panel shows nonfood trials of cross-modal testing [continuous sound with food primed after 50 sec; and 1-sec light signals at 10-sec intervals shown by arrows].) response decrement measure (RD) was used to index the effect of noise signals. This was calculated by the equation RD = L/H, where H is the highest response rate during the 10 sec prior to signal termination and L is the lowest response rate during the 10 sec following signal onset.

Results The results from peak trials obtained during the 5 days of the testing procedure are shown in pgure 7. In the top panel, the response rate (responses/min) is plotted as a function of the unreinforced sustained light duration signals with segment noise signals (training); in the bottorn panel, the response rate is plotted as a function of the unreinforced sustained sound duration signals with segment light signals (cross-modal testing). For training functions, the median value for peak time was 50 .5 sec, and the me-

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WARREN H. MECK AND RUSSELL M. CHURCH

dian peak rate was 64 4 responses/min. The median discrimination index for the overall 50-sec training functions was 1.93 .04, while the median response decrement (a measure of the effect of the regularly presented 1-sec signals) was 33% 2%. For testing functions, the median peak time was 50 .7 sec, not reliably different from the measure of peak time obtained for training functions, t(4)<l. The median peak rate for testing functions was 47 + 3 responses/min, reliably lower during testing than training, <(4) = 2.56, p < .05. The median discrimination index for the overall 50sec test functions was 1.97 .06, and the median response decrement was 29% 2%, neither measure reliably different from those obtained during training conditions, fs(4)< 1. Discussion The peak procedure used in this experiment results in an overall increase in response rate to a maximum near the time food was sometimes primed, 50 sec, followed by a fairly symmetrical decrease in rate. In addition, each of the 1-sec segment noise signals, which guaranteed an interval of at least 10 sec without food, led to a temporary decline in response rate. Thus, the rats appeared to be simultaneously processing two temporal signals, a light signal that indicated that food (if primed) would occur 50 sec after onset and a noise signal that indicated that food would not be primed for at least 10 sec. Similar results with a more extensive analysis of simultaneous temporal processing have been reported by Meek and Church (Note 2). During the five testing sessions in which signal conditions were reversed on a random half of the no-food trials, the overall response rate also increased to a maximum near 50 sec, and then declined in a fairly symmetrical fashion. In addition, there were local decreases in the response rate at the time of segment signals. Thus, it appears that the rats were still simultaneously processing two signal durations. The performance of rats in terms of accuracy (peak time, discrimination index, and response decrement) was relatively unaffected by the reversal of modality.

There was, however, a significant decline in peak rate during the reversal condition, which indicated that there was some modality-specific learning. Note that none of these reversed modality trials were reinforced, so that the rats had no experience with reinforcement after a 50-sec noise signal, or occasional reinforcement 10 sec after a 1-sec light signal. The consistent honreinforcement of these particular signals should eventually override the cross-modal transfer effect. Both before and after cross-modal transfer, the first sustained signal was used to define the 50-sec fixed interval, and the second brief signal was used to define the 10-sec interval in which no reinforcement occurs. In this experiment, the first compound signal (light and sound onset) may start the overall 50-sec clock and a 10-sec clock; subsequent compound signals may reset the 10sec clock and leave the 50-sec clock unchanged. The response rate is primarily determined by the 50-sec clock, but there are local decreases in response rate whenever another compound signal is presented. It is important to note that the performance cannot be determined solely by the compound signals; other experiments have shown that the 50-sec clock is stopped in the absence of the sustained signal (e.g., Roberts, 1981; Meek & Church, Note 2). General Discussion The ability of animals to abstract temporal attributes from sensory modalities implies that some internal stirnulus representations are not rigidly determined by the characteristics of the external stimulus. Pavlov (1927) considered the capacity of animals to respond to basic stimulus attributes irrespective of modality to be a rudimentary form of conceptual learning. Such an account emphasizes the information extraction and processing characteristics of sensory systems and deemphasizes sensory modalities per se. The capacity to respond differentially to basic stimulus attributes such as duration, intensity, location, and number without regard to sense modality would suggest that these are universal stimulus attributes,

ABSTRACTION OF TEMPORAL ATTRIBUTES

241

readily encoded in all principal sense modalities. Gibson (1969, ch. 11) proposed the existence of an amodal information capability dealing with basic attributes and suggested that cross-modal functions are mediated by this capability. If amodal capabilities exist, nearly equivalent performances in each modality for each type of problem would be expected as long as the discrimination was mediated by these basic attributes. Although this proposal is supported by the results from both the simple and relatively complex temporal processing tasks reported here, Freides (1974) pointed out that there are conditions under which it is difficult to transfer complex spatial and temporal patterns to "nonadept" modalities. Animals have physiological systems for the analysis of modality-specific stimuli; they may also have physiological systems for the analysis of attributes common to all stimuli. A person observing a stream of action potentials from various sensory neurons could make reasonable inferences regarding the duration, intensity, location, and number of the stimuli (e.g., Uttal, 1973). One.way the duration of a stimulus might be measured, for example, is by the time interval from the beginning to the end of the electrical activity above baseline; the number of sequential stimuli would be reflected in the number of bunches of potentials. The major physiological question is whether there is a neural mechanism that can act like the human observer to decode the stream of spike and other electrical potentials into attributes that can be numerically evaluted. There have been some reports of success in identifying higher neural units that respond to certain attributes, such as number (Thompson, Mayers, Robertson, & Patterson, 1970) or location (Newman & Hartline, 1981; Olton, Branch; & Best, 1978) without regard to modality, but some form of integrator would be required to bridge the long delays required to evaluate the temporal attributes. Our understanding of such central mechanisms is much inferior to our understanding of physiological mechanisms closely related to the physical stimulus or the motor response. In the case of time, it is not even certain where in the brain to search for relevant units, or for an integrator with the

properties of an internal clock. (For a review of the physiological basis of the internal clock, see Church and Meek, in press.) Much previous research on cross-modal transfer has been motivated by an interest in the brain mechanisms involved in such higher mental processes (e.g., Yeterian, 1977; Yeterian et al., 1976). Lesion studies have localized many sensory functions (e.g., Kliiver, 1933; Mountcastle, 1974), but are there lesions that do not affect any modality (e.g, vision or audition) but do affect the ability of an animal to transfer the value of an attribute from one modality to another? Previous work in this area has been ambiguous because of the difficulty in demonstrating convincing cross-modal transfer. The present experiments suggest that stimulus duration is a particularly good attribute to use in cross-modal transfer experiments aimed at identifying the brain mechanisms responsible for it. Previous studies have identified several properties of the internal clock (Church, 1978), and those properties apply with broad generality to results of timing the duration of stimuli from any modality. In the first two experiments, the animals apparently abstracted the temporal aspect of the signal from the modality of the signal, since they responded in a manner appropriate to the duration of the signal when the modality was changed. In these experiments, it is possible that any stimulus change might start an internal clock. In the third experiment, the animals demonstrated not only cross-modal transfer of the duration of a signal but also cross-modal transfer of the location of a sig-, nal. That is to say, they learned to time particular signals in a sequence regardless of the modality of the signals. Although information regarding temporal order might have been a factor in this experiment, its appreciation was not a prerequisite for acquiring the discrimination. The final experiment demonstrated the importance of temporal order in signal patterning, but in a rather different manner than previous studies in which discriminations were trained among various arrangements of two-event sequences (e.g., Weisman, Wasserman, Dodd, & Larew, 1980). When the temporal order and the modality of the stimulus were put

242

WARREN H. MECK AND RUSSELL M. CHURCH Freides, D. Human information processing and sensory modality: Cross-modal functions, information complexity, memory, and deficit. Psychological Bulletin, 1974, 81, 284-310. Gibbon, J. On the form and location of the psychometric bisection function for time. Journal of Mathematical Psychology, 1981, 24, 58-87. Gibbon, J., & Church, R. M. Time left: Linear vs. logarithmic subjective time. Journal of Experimental Psychology: Animal Behavior Processes, 1981, 7, 87108. Gibson, E. J. Principles of perceptual learning and development. New York: Appleton-Century-Crofts, 1969. Goldstone, S., & Goldfarb, J. L. The perception of time by children. In A. H. Kidd & J. L. Riviore (Eds.), Perceptual development in children. New York: International Universities Press, 1966. Hayes, W. GRAPH: A BASIC-11 graphics program for digital plotters. Behavior Research Methods & Instrumentation, 1981, 13, 367. Kelleher, R. T. Chaining and conditioned reinforcement. In W. K. Honig (Ed.), Operant behavior: Areas of research and application. Englewood Cliffs, N.J.: Prentice-Hall, 1966. Kltiver, H. Behavior mechanisms in monkeys. Chicago: University of Chicago Press, 1933. Koen, F. Verbal mediators in crossmodal form discrimination. Canadian Journal of Psychology, 1971, 25, 103-110. Luchins, A. S. Mechanization in problem solving. Psychological Monographs, 1942,54(6, Whole No. 248). MacDonall, J. S., & Marcucella, H. Cross-modal transfer of stimulus control in the albino rat: A stimulus delay procedure. Animal Learning & Behavior, 1976, 4, 341-346. Maricq, A. V., Roberts, S., & Church, R. M. Methamphetamine and time estimation. Journal of Experimental Psychology: Animal Behavior Processes, 1981, 7, 18-30. Meek, W. H., & Church, R. M. Discrimination of fixed intertrial intervals in cross-modal transfer of duration. Bulletin of the Psychonomic Society, in press. Mountcastle, V. B. (Ed.). Medical physiology (Vol. 1). St. Louis, Mo.: Mosby, 1974. Newman, E. A., & Hartline, P. H. Integration of visual and infrared information in bimodal neurons of the rattlesnake optic tectum. Science, 1981, 213, 789791. Olton, D. S., Branch, M., & Best, P.. J. Spatial correlates of hippocampal unit activity. Experimental Neurology, 1978, 58, 387-409. Over, R., & Mackintosh, N. J. Cross-modal transfer of intensity discrimination by rats. Nature, 1969, 224, 918-919. Pavlov, I. P. Conditioned reflexes (G. V. Anrep, trans.). London: Oxford University Press, 1927. Posner, M. I. Chronometric exploration of mind. Hillsdale, N.J.: Erlbaum, 1978. Roberts, S. Isolation of an internal clock. Journal of Experimental Psychology: Animal Behavior Processes, 1981, 7, 242-268. Roberts, S. Cross-modal use of an internal clock. Journal of Experimental Psychology: Animal Behavior Processes, 1982, 8, 2-22.

in conflict in a simultaneous temporal processing task, animals responded appropriately with respect to the temporal order rather than to the modality characteristics of the signal. All of these experiments are consistent with the proposal that the same internal clock can time signal durations from various modalities (Roberts & Church, 1978) and are inconsistent with the notion that time discriminations are mediated by peripheral mechanisms such as the degree of dark adaptation. We do not know whether animals have specialized systems for the analysis of other basic attributes, such as intensity, location, and number. Successful demonstration of substantial cross-modal transfer of such attributes would be a reasonable first step in exploring this possibility. Reference Notes
1. Meek, W. H. Directed attention and automatic activation of psychological pathways. Manuscript submitted in partial fulfillment of the requirements for the PhD degree at Brown University, 1981. 2. Meek, W. H., & Church, R. M. Simultaneous temporal processing. Paper presented at the meeting of the Eastern Psychological Association, Hartford, April 1980.

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Blank, M., & Bridger, W. H. Cross-modal transfer in nursery-school children. Journal of Comparative and Physiological Psychology, 1964, 58, 277-282. Broadberit, D. E. Perception and communication. London: Pergamon Press, 1958. Bryant, P. E., Jones, P., Claxton, V., & Perkins, G. M. Recognition of shapes across modalities by infants. Nature, 1972, 240, 303-304. Church, R. M. The internal clock. In S. H. Hulse, H. Fowler, & W. K. Honig (Eds.), Cognitive processes in animal behavior. Hillsdale, N.J.: Erlbaum, 1978. Church, R. M., & Deluty, M. Z. Bisection of temporal intervals. Journal of Experimental Psychology: Animal Behavior Processes, 1977, 3, 216-228. Church, R. M., & Gibbon, J. Temporal generalization. Journal of Experimental Psychology: Animal Behavior Processes, 1982, 8, 165-186. Church, R. M., & Meek, W. H. Acquisition and crossmodal transfer of classification rules for temporal intervals. In M. L. Commons, A. R. Wagner, & R. J. Herrnstein (Eds.), Quantitative analysis of behavior: Discriminative processes (Vol. 4). Cambridge, Mass.: Ballinger, in press. Ettlinger, G. Interactions between sensory modalities in nonhuman primates. In A. M. Schrier (Ed.), Behavioral primatology: Advances in research and theory (Vol. 1). Hillsdale, N.J.: Erlbaum, 1977.

ABSTRACTION OF TEMPORAL ATTRIBUTES Roberts, S., & Church, R. M. Control of an internal clock. Journal of Experimental Psychology: Animal Behavior Processes, 1978, 4, 318-337. Seraganian, P., & Popova, Y. I. Cross-modal transfer of a conditioned flexion response in dogs. Pavlavian Journal of Biological Science, 1976, //, 162-174. Thompson, R. F., Mayers, K. S., Robertson, R. T., & Patterson, C. J. Number coding in association cortex of the cat. Science, 1970, 168, 271-272. Uttal, W. R. The psychobiology of sensory coding. New York: Harper & Row, 1973. Ward, J. P., Silver, B. V., & Frank, J. Preservation of cross-modal transfer of a rate discrimination in the bushbaby (Galago senegalensis) with lesions of posterior neocortex. Journal of Comparative and Physiological Psychology, 1976, 90, 520-527. Weisman, R. G., Wasserman, E. A., Dodd, P. W. D.,

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' & Larew, M. B. Representation and retention of twoevent sequences in pigeons. Journal of Experimental Psychology: Animal Behavior Processes, 1980, 6, 312-325. Wilson, W. A., & Shaffer, O. C. Intermodality transfer of specific discriminations in the monkey. Nature, 1963, 197, 107. Yeterian, E. H. The effects of visual or auditory cortical lesions on specific cross-modal transfer in the rat. Neuropsychologia, 1977, 15, 517-527. Yeterian, E. H., Waters, R. S., & Wilson, W. A. Posterior cortical lesions and specific cross-modal transfer in the rat. Physiological Psychology, 1976, 4, 281284.

Received July 29, 1981