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In recent experiments with goldfish (Ten- shadowing effect, independent of prior train-
nant & Bitterman, 1973), we failed to find ing with B, is due, presumably, to its "sa-
better intradimensional than extradimen- lience," while the blocking effect is defined
sional transfer, a phenomenon which has as an outcome of such prior training. In
been found in other veterbrates (pigeons, the present experiment, we used a design
rats, and monkeys) and which constitutes which permitted both effects to operate, a
some of the strongest evidence for the op- conservative strategy justified by the fact
eration of an attentional process in those that the only previous study of the problem
animals (Sutherland & Mackintosh, 1971). (an unpublished blocking experiment by
In the present experiments with carp and Ingle) had produced negative results. In-
goldfish, we looked for blocking and over- gle's results led Gleitman and Rozin (1971),
shadowing, two Pavlovian phenomena who cited them, to the conclusion that "at-
which, like the dimensional transfer effect, tentional processes in fish are weak and
are well known in other vertebrates and labile" (p. 259).
have been explained in attentional terms,
although alternative interpretations (which Method
do not predict the dimensional transfer ef- Subjects. The subjects were 13 10-cm- goldfish
fect) are available. The results bear on the (Carassius auratus) supplied by a local dealer.
phyletic generality of blocking and over- They were maintained on a 24-hr feeding sched-
ule in a temperature-controlled laboratory room.
shadowing as well as on their functional All had extensive previous training in a series of
relation to the dimensional transfer effect. free-operant red-green discrimination reversals.
Apparatus. Each subject was trained in its in-
dividual living tank which was carried to an ex-
EXPERIMENT 1: A HETEROMODAL perimental chamber located in a larger sound-
MIXED EFFECT IN GOLDFISH attenuating enclosure. ' The training apparatus,
mounted on a chassis of black Plexiglas, covered
A common feature of blocking and over- the top and front end of the tank. The manipu-
shadowing is that learning about A, a com- landum was a circular target, 4 cm in diameter,
ponent of the compound stimulus AB, is presented at the front end of the tank. The tar-
reduced by the presence of B. The over- get was fixed to a rod, the other end of which
was inserted into the needle holder of a phono-
This research was supported by Grant MH- graph cartridge, and any contact of the animal
23294 from the U.S. Public Health Service. with the target produced a voltage across the car-
Requests for reprints should be sent to M. E. tridge which was used to operate a response relay;
Bitterman, Laboratory of Sensory Sciences, Uni- the circuit has been described elsewhere (Wood-
versity of Hawaii, 1993 East-West Road, Hono- ard & Bitterman, 1974). At the center of the
lulu, Hawaii 96822. target was a Plexiglas nipple through which liquid
22
BLOCKING AND OVERSHADOWING IN FISH 23
for half). Thus, the blocking group could con- terms of mean discrimination ratios for the
tinue to respond in terms of the previously estab- blocking and control animals. The results
lished color discrimination, while the control group
could not. For both groups, of course, the two for Stage A (Sessions 1-12) show that the
angles were differentially reinforced. two groups were closely matched for per-
Stage C. In Session 19, all animals had an formance in the original color discrimina-
extinction test with white horizontal and vertical tion. The results for Stage B (Sessions 13-
lines. (As usual, the stimuli were presented in 18) suggest that the blocking group was
Gellermann orders, but response to neither was
reinforced.) unaffected by the introduction of the con-
Stage D. In Session 20, all animals were re- founded angle cues; the performance of the
trained on the confounded color-angle problem as control group was substantially affected by
in Stage B. the reversal of the color cues, although re-
Stage E. In Sessions 21-30, all animals were
trained to discriminate the white horizontals and covery was complete by Session 17-18.
verticals used in Stage C. For animals whose The results for the extinction test of Stage
previously positive angle was 0°, white-0° was C (Session 19) provide clear evidence of
positive, while for animals whose previously posi- blocking—the performance of the blocking
tive angle was 90°, white-90° was positive.
Stage F. In Sessions 31-38, all animals were group fell to chance, while that of the con-
trained on a confounded color-angle problem with trol group showed substantial discrimina-
orange and purple horizontals and verticals— tion of angle alone. The difference is sta-
either orange-0° vs. purple-90" or purple-00 vs. tistically reliable, F (1, 10) = 7.14, p < .05.
orange-90°. The former control group (e.g., Sub- It is interesting to note that in Stage D, when
ject 2) now became the blocking group and for
these animals the previously positive angle re- both groups were returned for a single ses-
mained positive (orange for half and purple for sion to the confounded problem of Stage B,
half). For the former blocking group (e.g., Sub- the performance of the control group gave
ject 1), now the control group, the previously some indication of disturbance by the prior
negative angle was positive (orange for half and
purple for half). That is, the blocking group extinction test with angle, while that of the
could continue to respond in terms of the pre- blocking group (apparently entirely under
viously acquired angle discrimination, while the the control of color) did not, F (1, 10) =
control group could not. 16.13, p < .01. Further evidence of block-
Stage G. In Session 39, all animals had an ex-
tinction test with orange and purple 45° lines ing is to be found in the superior perform-
(Gellermann orders, neither reinforced). ance of the control group in Stage E when
Stage H. In Session 40, both groups were both groups were trained with the white an-
trained to discriminate between the orange and gles, F (1, 10) =6.59, p< .05.
purple 45° lines. The positive color of Stage F As may be seen from Figure 2, the per-
was positive again in Stage H.
formance of both groups in the angle dis-
Results crimination of Stage E soon became very
Performance in each session of each stage good. When, in Stage F, a confounded
of the experiment is plotted in Figure 2 in color difference was introduced, the per-
1.0
o
0 .7
'5
.3
10 15 2O 25 30 35 40
SESSIONS
formance of the blocking group (for which mensional transfer effect, which had failed
the previously positive angle remained posi- to appear in our previous work with gold-
tive) was unaffected, while the performance fish, led us to look also for the dimensional
of the control group (for which the pre- transfer effect in carp.
viously positive angle now became negative)
was disrupted for a time, although it Method
equalled that of the blocking group by the Subjects. The subjects were 12 experimentally
end of Stage F. Evidence of blocking ap- naive 10-cm carp (Cyprinus carpio) supplied by
a local dealer. They were maintained on a 24-hr
peared again in the extinction test of Stage feeding schedule in a temperature-controlled lab-
G (when the stimuli differed only in color). oratory room.
While the performance of the two groups Apparatus. The apparatus was the same as in
was equally good at the end of training, per- Experiment 2. The colors used in this experi-
formance based on color alone in the ex- ment were red (Wratten filter No. 25), yellow
(three thicknesses of No. 8), green (No. 58),
tinction test was significantly better in the and blue (No. 47). The angles were 0" (ver-
control group than in the blocking group tical), 45°, 90°, 120°, 135°, and 150°, all mea-
F (1, 10) =67.90, p < .01. Retraining sured clockwise.
was rapid, however, and the performance Procedure. The discriminative training method
was the same in general as in the previous ex-
of the two groups did not differ significantly periments. Each session began with a warm-up
in Stage H (F < 1). on the white light and was followed by 3-min
We conclude that the blocking of angle presentations (in modified Gellermann orders) of
by color and of color by angle has been dem- the various stimuli to be discriminated, S+ on
VI 1-min reinforcement and S—• on extinction.
onstrated in goldfish. Part 1. The first part of the experiment was
concerned with overshadowing. In Table 3, the
EXPERIMENT 3: OVERSHADOWING AND procedure is illustrated in terms of the treatment
DIMENSIONAL TRANSFER IN CARP of two sample subjects.
In Stage 1 of Part 1, all animals were trained
In this experiment, we set out to deter- for five sessions with a yellow 135° line on VI
mine if a more salient color difference would 1-min reinforcement. There were 12 presenta-
overshadow a less salient angle difference, tions of this stimulus in each session. The pur-
salience being defined in terms of difficulty pose of the training was to familiarize the ani-
mals with the yellow color to be used in the over-
of discrimination as established in indepen- shadowing tests (Stages 3 and 4).
dent tests. Although we had begun our In Stage 2 of Part 1, six animals (overshadow-
work on attention in fishes with goldfish, ing group) were trained for: 20 sessions on a
we found it necessary here to use carp be- confounded color and angle discrimination. For
three of the fish red-120° was positive and green-
cause goldfish were for a considerable time ISO0 was negative, while for the other three,
in very short supply. Interest in the rela- green-ISO" was positive and red-120° was nega-
tion between overshadowing and the di- tive. The remaining six animals (control group)
were trained for 20 sessions on a pure angle
TABLE 3 discrimination; red-120" was positive and red-ISO0
was negative for three fish, while green-ISO0 was
DESIGN OF PART 1 OF EXPERIMENT 3 (OVER- positive and green-120° was negative for the other
SHADOWING) ILLUSTRATED IN TERMS OF three. In each session, there were 6 3-min pres-
TREATMENT OF Two SAMPLE SUBJECTS entations of S+ and 6 3-min presentations of S—.
In Stage 3 of Part 1, both groups had a single
No. of
Stage sessions Subject 1 Subject 2 extinction session in which yellow-120" and yel-
low-ISO" were presented in a balanced order (six
3-min presentations of each).
1 5 Y,135° + In Stage 4 of Part 1, both groups were trained
R,120°+G,150°~ for a single session on a pure yellow-angle dis-
2 20
R,120°+ R,150°- crimination with the angle which was positive in
3 1 Y,120°- Y,150°- Stage 2 positive again in Stage 4. For animals
4 1 Y,120° + Y,150°- whose previously positive stimulus was red-1200,
yellow-1200 was positive and yellow-ISO0 was
Note, Abbreviations: Y = yellow, R = red, and G = green; negative. For animals whose previously positive
120, 133, and ISO are angular orientations; + = reinforced, stimulus was green-150", yellow-150" was posi-
— = unreinforced. Subject 1 is in the overshadowing group
and Subject 2 is in the control-group. tive and yellow-1200 was negative.
BLOCKING AND OVERSHADOWING IN FISH 27
sions 16-22) also was the same in the two tion that components of a compound stimu-
groups—that is, performance was the same lus independently acquire excitatory and
whether the relevant dimension had been inhibitory properties by association with
relevant or irrelevant in the prior stage. reinforcement and nonreinforcement—or
Furthermore, performance with the red- whether some principle of stimulus domi-
yellow horizontal-vertical set in Stage 3 nance or stimulus selection is required.
(Session 23) was not impaired by extra- Even where (as here) the experimental de-
dimensional training in Stage 2; nor in sign makes it possible to rule out explana-
Stage 4 (Session 24) was concurrent per- tions in terms of stimulus generalization or
formance with the two sets of stimuli any afferent neural interaction, the possibility
better when the same dimension was rele- must be considered that different experience
vant in both sets than when the relevant with the critical components in the training
dimension of one set was the irrelevant di- stage is responsible for the outcome. Con-
mension of the other. Like goldfish, then, sider, for example, the present experiments:
carp fail to show dimensional transfer un- Although in each case the two groups had
der these conditions. equal exposure to the blocked or over-
shadowed components, and the same number
DISCUSSION of reinforcements for response to S+ (with
The work reported here is significant pri- VI 1-min training, a minimal rate of re-
marily because it extends the range of ver- sponding to S+ ensures the delivery of all
tebrates in which blocking and overshadow- scheduled reinforcements), the blocking or
ing have been demonstrated, lending weight overshadowing group made more unrein-
to the common assumption that they are forced responses to S+ and fewer unrein-
general phenomena of vertebrate learning forced responses to S— than did the control
(Bitterman & Woodard, 1975). The re- group. There simply is no way in the train-
sults bear also on the theory of blocking ing stage of a blocking or overshadowing
and overshadowing. experiment to equate the experience of the
The first question to be asked about the two groups both with respect to dura-
two phenomena, of course, is whether they tion of exposure to the critical components
can be dealt with in terms of conventional and with respect to frequency of respond-
continuity theory—that is, on the assump- ing in the presence of those components.
1.O
0 9
? .8
- R-Y
Q .7
1,
I 5 INTRADIMENSIONAL
or '
EXTRADIMENSIONAL
.3L
1 1O 15 20
SESSIONS
FIGURE 4. Performance of the two groups in the initial training with
the red-yellow (R-Y) set, in the subsequent training with the blue-green
(B-G) set, and in subsequent tests with the red-yellow set and with both
sets combined (Experiment 3, Part 2).
BLOCKING AND OVERSHADOWING IN FISH 29
The suggestion of Wagner (1969) that clas- might be some objection, furthermore, to
sical conditioning is more appropriate in our use of experienced animals, although
such experiments than instrumental condi- there is nothing in attention theory which
tioning is acceptable only on the assumption suggests that the dimensional transfer ef-
that differences in responding are of less fect occurs only in naive animals or is mani-
importance in classical conditioning. It re- fested less readily by experienced animals.
mains, however, to be determined whether It should be noted that the subjects of Ex-
the entire pattern of results obtained in the periment 1 were not naive, nor were the
blocking and overshadowing experiments subjects that showed blocking of color by
which have been reported thus far can in angle in Experiment 2 (having previously
fact be derived rigorously from continuity been used to demonstrate blocking of angle
theory. by color). Given our negative results, it
Two other interpretations of blocking and seems reasonable for the moment to assume
overshadowing are available, one which may that the dimensional transfer effect is not
be phrased in attentional terms (Sutherland to be found in goldfish or carp and there-
& Mackintosh, 1971.) and the other in in- fore to entertain the hypothesis that the
formational . terms (Rescorla & Wagner, processes responsible for dimensional trans-
1972; Wagner & Rescorla, 1972). The fer are different from those which produce
change in the associative strength of some blocking and overshadowing.
component of a compound stimulus pro- REFERENCES
duced by reinforcement or nonreinforcement
is related, in the one view, to the amount Bitterman, M. E., & Woodard, W. T. Vertebrate
learning; Common processes. In R. B. Master-
of attention paid that component and, in the ton, M. E. Bitterman, C. B. G. Campbell, &
other, to the predictability of reinforcement N. Hotten (Eds.), The evolution of brain and
or nonreinforcement from the compound. behavior in vertebrates. Potomac, Md.: Erl-
In the evaluation of these alternatives, the baum, 1975.
Gleitman, H., & Rozin, P. Learning and mem-
negative results reported here on dimen- ory. In W. S. Hoar & D. J. Randall (Eds.),
sional transfer in carp, along with our pre- Fish physiology (Vol. 6) : Environmental rela-
vious results for goldfish (Tennant & Bit- tions and behavior. New York: Academic Press,
terman, 1973), may be of some assistance. 1971.
Rescorla, R. A., & Wagner, A. R. A theory of
Since attention theory predicts better intra- Pavlovian conditioning: Variations in the ef-
dimensional than extradimensional transfer fectiveness of reinforcement and nonreinforce-
as well as blocking and overshadowing, the ment. In A. H. Black & W. F. Prokasy (Eds.),
informational theory, which does not predict Classical conditioning II. New York: Appleton-
Century-Crofts, 1972.
the dimensional transfer effect, might appear Sutherland, N. S., & Mackintosh, N. J. Mecha-
superfluous if the dimensional transfer ef- nisms of discrimination learning in animals,
fect were found in all animals that show New York: Academic Press, 1971.
blocking and overshadowing. With our Tennant, W. A., & Bitterman, M. E. Some com-
failure to find dimensional transfer in carp parisons of intra- and extradimensional transfer
in discriminative learning of goldfish. Journal
and goldfish, however, the demonstration of of Comparative and Physiological Psychology,
blocking and overshadowing in the same ani- 1973, 83, 134-139.
mals suggests that the three phenomena Wagner, A. R. Incidental stimuli and discrimina-
may not have a common explanation and tion learning. In R. M. Gilbert & N. S. Suther-
land (Eds.), Animal discrimination learning.
emphasizes the usefulness of the Wagner- New York: Academic Press, 1969.
Rescorla model. Wagner, A. R., & Rescorla, R. A. Inhibition in
How seriously this argument is taken de- Pavlovian conditioning: Application of a theory.
pends, of course, on how much weight is In R. A. Boakes & M. S. Halliday (Eds.), In-
given to the negative results of our dimen- hibition and learning. New York: Academic
Press, 1972.
sional transfer experiments. The possibil- Woodard, W. T., & Bitterman, M. E. Improved
ity certainly must be considered that dimen- techniques for the measurement of consumma-
sional transfer is a more subtle effect than tory behavior in fishes. Behavior Research
blocking or overshadowing, one which our Methods and Instrumentation, 1974, 6, 321-324.
technique is inadequate to detect. There (Received March 1, 1974)