LEARNING

AND MOTIVATION

17, 40-58 (1986)

Observational Learning of a Conditional Hue Discrimination

in Pigeons
DAVID

E. HOGAN

Northern Kentucky University

Three experiments examined a relationship between cooperation and observational learning in pigeons using procedures similar to Skinner (1962). In Experiment 1, dyads cooperated by searching and pecking nearly simultaneously
on the correct pair of adjacent keys of a 2 x 2 matrix. The dyads learned to
search for the correct keys in a coordinated fashion, but only if they were
permitted to watch one anothers performance. Experiment 2 disconfirmed the
hypothesis that the coordinated performance of cooperating pigeons reflects an
unIearned response tendency to peck at locations close to where the other bird
is pecking. Experiment 3 demonstrated that prior cooperation training involving
an observer and demonstrator pigeon facilitated subsequent observational learning
of a conditional hue discrimination. Cooperation training appeared to facilitate
observational learning by establishing the demonstrators peck response as an
instructional stimulus which indicated the reward significance of the discriminative
stimuli. GI I986 Academic Pres, Inc.

Skinner (1962) reported what appears to be the first account of generalized

imitative behavior in pigeons. In the demonstration experiment, a pair
of pigeons worked side by side in separate, bilaterally symmetrical compartments which contained a column of three keys and a grain feeder.
During a so-called cooperation training phase, a discrete trial procedure
was used in which only one row of keys was designated correct on any
given trial, but the correct row was unsignaled and changed randomly
across trials (i.e., on some trials the pair of keys comprising the top row
were designated as correct and on other trials the pair of keys comprising
the middle row were correct, etc.). Both birds were rewarded when they
pecked keys on the correct row within 0.1 s of one another.
This research was supported by a Project Grant and a Summer Fellowship from Northern
Kentucky University. I thank Anna Stegman, Teresa Landers, Kim Gregory, Holly Riffe.
Becky Schrand, Douglas Herald, Donna Hendrix, Kathleen Due, Lissa Knue, and Cheryl
Tornabene for their expert technical assistance, Reprints requests may be sent to David
E. Hogan, Psychology Department, Northern Kentucky University, Highland Heights, KY
41076.
40
0023-9690/86 $3.00
Copyright Q 1986 by Academic Press, Inc.
All rights of reproduction in any form reserved.

OBSERVATIONAL

LEARNING

IN

PIGEONS

41

After extensive training, a leader-follower

relationship developed in
which the leader pigeon pecked the keys in its compartment in a random
order, and the follower learned to immediately peck the adjacent key.
Eventually, the within-trial,
search-and-peck behavior of the two birds
became so imitative that they appeared as one pigeon seen in a mirror
(Skinner, 1962, p. 533). A more intriguing observation by Skinner, however,
was that the pigeons imitative behavior was not specific to just pecking
at keys simultaneously.
After the cooperation training, when the birds
were interchanged in their compartments, the follower pigeon appeared
to imitate other responses, such as drinking from a cup of water, head
bobbing, and exploring new areas of the chamber when such responses
were emitted by the leader pigeon.
The hypothesis that Skinners pigeons exhibited imitation which generalized across responses can be questioned, however, because generalization across responses refers to the case in which an observer imitates
a variety of topographically
unique responses which are emitted in a
series by the demonstrator (Baer & Sherman, 1964). The novel instances
of imitative behavior which Skinner reported, however, were very similar
in topography to the search-and-peck responses which were reinforced
during the cooperation training phase. The head bobbing behavior, for
example, closely resembled the up-and-down head movements which the
two birds emitted during the cooperation task, and the response of bending
down and drinking closely resembled the well-practiced approach response
downward from the keys to the feeder aperture near the floor. Perhaps
it is more parsimonious to conclude that cooperation training established
the leaders peck response as a visual instruction to peck novel objects
in novel locations.
The present experiments examined whether cooperation training would
facilitate subsequent observational learning of a visual discrimination
which was exhibited by a demonstrator pigeon. Skinners (1962) results
suggest that a follower pigeon acquires a strong tendency to look and
peck at novel stimuli which are similar to those which the leader pecks.
It is therefore reasonable to predict that if a leader pigeon exhibited a
well-learned visual discrimination in the followers presence, then the
attentive
follower may learn about the reward significance of the visual
stimuli which control differential responding in the leader and thus come
under immediate control of the same discriminative stimuli during a test.
Several organisms, including monkeys (Darby & Riopelle, 1959; Riopelle,
1960), rats (Kahn & Dennis, 1972), pigeons (Edwards, Hogan, & Zentall,
1980), and domestic chicks (Suboski &L Bartashunas, 1984) apparently
can learn a visual discrimination
by merely observing a well-trained
demonstrator perform discriminatively, but the research has been concerned
with merely demonstrating that observational learning of a discrimination

42

DAVID

E. HOGAN

is within the species capacity. Experiment 3 of the present series examined

whether observational learning of a hue discrimination can be influenced
by a history of cooperative key pecking. Experiments 1 and 2 examined
task variables which affect learning to cooperate in pigeons.
EXPERIMENT

The present experiment used a 2 x 2 matrix of keys, which was simpler

to program electronically, instead of the 2 x 3 matrix used by Skinner.
As in Skinners experiment, however, both birds were rewarded for
exploring and simultaneously pecking (i.e., 0.1 s) the correct pair of keys
of the matrix (the keys comprising the top row had to be pecked simultaneously on a randomly selected half of the trials, and those comprising
the bottom row had to be pecked simultaneously on the remaining trials,
but the correct pair was unsignaled).
Group A-V dyads were separated by a thin transparent partition, so
the follower could see and hear which key the leader was pecking. Group
A-i7 dyads were separated by an opaque partition and were --permitted
only to listen to which key their partner was pecking. Group A-V dyads
performed in separate boxes, each containing an identical 2 x 2 matrix
(one bird had access to the left column of keys in one box and the other
bird had access to the right column of keys in the other box), and provided
a performance baseline under conditions in which the followers spatial
choice could not be influenced by visual or auditory cues associated with
the leaders
choice. Based on Skinners (1962) results, Group A-V
was expected to perform most efficiently, since the follower could both
listen and watch where the leader pecked.
A reversal test further assessed how well the dyads (especially Group
A-V) learned to coordinate their search-and-peck behavior. Testing included
a random mixture of (a) the familiar training trials in which keys on the
same row were correct and (b) novel reversal trials in which keys on
different rows were correct (on some trials the reinforcement contingency
required one bird to peck the top left location when its partner pecked
the lower right location. On other trials the bottom left location and
upper right location had to be pecked simultaneously). If the leaders
peck response served as a visual instruction for the follower to peck an
adjacent key in Group A-V, as Skinner suggests, then the latency to
terminate novel reversal trials should be long, relative to the latency to
terminate the familiar same-row trials in which adjacent keys were correct.
Method

Subjects
Twenty-six loft-reared pigeons approximately
at 75% of their free-feeding weight.

I year old were maintained

OBSERVATIONAL

LEARNING

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43

Apparatus

Two modified pigeon chambers were used, each measuring 36 cm high,

30 cm wide, and 36 cm deep. The chambers consisted of two compartments,
each measuring 28 cm high, 30 cm wide, and 18 cm deep. The compartments
were separated by a floor-to-ceiling
partition running perpendicular to
the test panel. Two interchangeable partitions were constructed for each
chamber, one transparent (clear PlexigIas) and the other opaque (Masonite
board). Each partition measured approximately 27.5 cm high, 30 cm wide,
and 0.6 cm thick. The partitions were hinged to the floor and could be
pulled down for easy access to the compartment which was farther from
the chamber door. A narrow strip of wood (0.6 cm wide x 0.6 cm thick
x 27.5 cm long) was screwed to the vertical midline of the panel and
served as a stop when the partition was standing upright. The opaque
partition overlapped the wooden strip by approximately 0.6 cm and prevented the birds from seeing activity in the adjacent compartment.
The test panel of both chambers was painted flat black and contained
a 2 x 2 matrix of clear plastic keys which measured 2.5 cm in diameter.
The keys were rear illuminated with white light by Industrial Electronics
Engineers projectors. The keys on the bottom row were centered 14 cm
from the floor and 3.2 cm from the panels midline, and those comprising
the top row were centered 21 cm from the floor and 3.2 cm from the
midline. Both compartments also contained a rear-mounted grain feeder
with a 5 x 5-cm access hole centered 5.5 cm from the floor and 10 cm
from the midline and a houselight centered 26 cm from the floor and 15
cm from the midline. The houselights were illuminated continuously
throughout the sessions of every phase. Programming equipment was
located in a separate room.
Procedure
Shaping. The key peck response was shaped via successive approximations in a single-key chamber to avoid conditioning a preference for
one of the two keys on the matrix. On 10 sessions, each of IO pecks to
a white light was reinforced with 2-s access to mixed grain. The latency
to emit each response was recorded. Two pigeons, which later served
as dummies,
were not shaped and remained in their home cage.
Preliminary training. The 24 trained birds were randomly assigned to
one of four compartments (6 per compartment) such that the peck latency,
averaged across the 10 shaping sessions, was balanced within and between
chambers. During the initial 16 preliminary training sessions, the top and
bottom keys were illuminated 16 times each, and a single peck produced
reinforcement.
The compartments were separated by the transparent
partition on eight sessions and the opaque partition on eight sessions
(randomly selected).

44

DAVID

E. HOGAN

Twelve matched pairs were formed, based on the latency averaged

over the initial 16 sessions. The dyads then were assigned to three groups
(4 dyads per group), which were balanced for the mean latency. The
dyads of Group A-V and Group A-v had received training
in adjacent
-compartments of the same chamber. The dyads of Group A-V had received
training in noncorresponding compartments of different chambers (a pigeon
trained in the left-hand compartment of one chamber was paired with a
pigeon in the right-hand compartment of the other chamber, etc.). The
groups were counterbalanced over chambers.
During the next 18 sessions, all dyads were trained under their respective
conditions of observation. Group A-V dyads were separated by the transparent
partition, and Group A-v dyads by the opaque partition. Group
-A-V dyads performed in separate chambers; but the compartment adjacent
to the performing pigeon contained an untrained dummy
pigeon to
control for nonspecific visual and aural effects --of a conspecifics mere
presence (Zajonc, 1965). For two dyads (Group A-V-T), the transparent
partition separated
the performer and dummy, and for the other two
-dyads (Group A-V-O) the partititon was opaque. The keys were illuminated
and the food hopper was raised in the dummys compartment when each
respective event occurred in the performers compartments. Training
sessions consisted of a random mixture of forced-choice trials in which
only the top row of keys was illuminated (16 trials) or only the bottom
row was illuminated (16 trials). Reinforcement occurred when one bird
pecked the illuminated key within 1.0 s of the other (either bird could
lead).
They key lights were turned off during reinforcement, and a
new trial began immediately. Eighteen sessions ensured extinction of
combative behaviors, such as wing flapping and pecking at the bird which
was visible through the transparent partition.
Training Phases 1, 2, 3, und 4. Phase 1 (36 sessions) sessions consisted
of a random mixture of 16 forced-choice trials, structured identically to
the forced choice trials of the immediately preceding phase, and 16 freechoice trials in which all four keys were illuminated simultaneously. The
top row was designated correct on 8 free-choice trials, and the bottom
row was designated correct on the other 8 trials. The correct row on
free-choice trials was unsignaled. Half of the free-choice trials which
were correct on the top row were immediately preceded by a trial in
which the top row was also correct (four positive transitions) and the
remaining ones by a trial in which the bottom row was correct (four
negative transitions). Similarly, half of the bottom-row, free-choice trials
were preceded by a trial which was correct on the bottom row (four
positive transitions) and the remaining ones by a trial which was correct
on the top row (four negative transitions). Two different trial orders were
used randomly over days.

OBSERVATIONAL

LEARNING

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45

The cooperative response requirement was 1.0 s on free-choice and

forced-choice trials during Phase 1. As in Skinners experiment pecks
on an incorrect pair of keys on free-choice trials did not terminate the
trial (e.g., on a trial in which, say, the keys comprising the top row had
to be pecked within 1.0 s of each other, pecks which occurred within
1.0 s of one another on the two bottom keys, or the upper left and lower
right, or lower left and upper right keys, did not terminate the trial. The
trial continued until the birds pecked the correct pair within 1.0 s of one
another.) Since there was no contingency that one bird always lead
the other, either bird could be a leader on any given trial. However,
the number of reinforcers which a given bird produced by pecking at
most 1 s before its partner (i.e., the number of leader pecks) were
recorded.
The cooperative response requirement was reduced to 0.5 s during
Phase 2 (6 sessions), 0.2 s during Phase 3 (24 sessions), and 0.1 s during
Phase 4 (72 sessions). During the final 24 sessions of Phase 4, the cooperative response on the correct row was reinforced on a partial schedule
in anticipation of a test phase. On Sessions 49-50, twenty-five percent
of all trials were nonreinforced;
on Sessions 51-56, fifty percent were
nonreinforced,
and on Sessions 57-72, seventy-five percent were nonreinforced. On nonreinforced trials the key lights were turned off for 2
s when a cooperative response occurred on the correct row. The nonreinforced trials were counterbalanced across the top and bottom rows
of the matrix. Immediately following Session 56, a subject in Group AV-T became lame and was retired from the experiment, The plots and
data analyses beyond Session 56 reflect the missing dyad.
Testing. During test sessions (six sessions), all dyads were presented
a random mixture of the 16 forced-choice trials of Phase 4 and two types
of free-choice trials. On Same-row free-choice trials (8 trials) keys on
the same row of the matrix were designated as correct. The correct row
was counterbalanced over the matrix (i.e., on 4 Same-row trials the keys
comprising the top row were correct, and on the other 4 trials the keys
comprising the bottom row were correct). On Different-row, free-choice
trials (8 trials), the correct pair of keys involved the upper left and lower
right locations on 4 trials, and the lower left and upper right locations
on 4 trials. As on Same-row, free-choice trials, the correct row of keys
on Different-row,
free-choice trials was unsignaled. Seventy-five percent
of all test trials were nonreinforced.
Results and Discussion

Figure 1 depicts the mean latency to peck the correct pair of keys on
the free-choice trials for each group. The mean latency of each group
improved (i.e., became shorter) at roughly the same gradual rate in Phase
1 but reached slightly different asymptotic levels within each phase. A

46

DAVID

PHASE

GROUP A-J

GROUP A-v

GflOUP &ii-T

GROUP ii-v-0

E. HOGAN

.
.

BLOCKS
FIG.

1. Acquisition

OF SIX SESSIONS

of the cooperation

task in Experiment

I.

comparison of the asymptotic levels reveals that the dyads

which were
-separated by a transparent partition (Groups A-V and A-V-T) averaged
slightly longer latencies
than the dyads separated by an opaque partition
-(Groups A-v and A-V-O). The latency data of each dyad were averaged
over sessions, rank ordered, and analyzed with a Mann-Whitney
U test.
(The significance
level
was
p
c
.05).
The
analyses
confirmed
that
Groups
-A-V and
-- A-V-T performed significantly longer latencies than Groups Av and A-V-O combined during Phase 2 [U(6, 6) = 21, Phase 3 [U(6, 6)
= 11, and Phase 4 [U(6, 5) = 21. The fact that Group A-V did not exhibit
the shortest latencies relative to all of the other groups is inconsistent
with Skinners (1962) conclusion that the cooperation reinforcement contingency produces well-coordinated,
or imitative
search-and-peck
behavior.
The general similarity in the acquisition functions suggests that the
performance of all dyads was under the control of some factor which
was common across all conditions of observation. A comparison of performance on the positive and negative transition trials (trial n) indicated
that the dyads of all three groups exhibited win-stay
behavior during
the training sessions (i.e., a preference for pecking the location on trial
IZ which was correlated with reinforcement on trial n - l), which gradually
dissipated across the phases. Transition performance, averaged over sessions and groups, is presented in Table I. Difference t tests revealed
significantly better performance on positive transition trials during Phase

OBSERVATIONAL

LEARNING

47

IN PIGEONS

TABLE 1
Mean Latency (in Seconds) on Positive and Negative Transition
Transition
Phase

Postive

Negative

1
2
3
4

I.1
5.0
1.4
7.5

9.4
6.1
8.4
7.8

Trials
Difference
scores
- 1.7
-1.1
- I.0
- .3

Nofe. Difference scores were obtained by subtracting the mean latency on negative
transition trials from positive transition trials.

1 [t(ll) = 3,60)] and Phase 3 [f(ll) = 2.351 and marginal significance

during Phase 2 [r(ll) = 2.101. Transition performance was equivalent
during Phase 4, however [r(lO) = .538].
Perhaps the short intertrial interval (O-s ITI) is related to learning the
win-stay behavior, because the pigeons could easily remember which
key had been associated with reinforcement on trial n - 1. Skinner also
used a O-s ITI, and this may be why extensive training was needed to
produce stable performance (Gilbert & Beaton, 1967; Hake, Donaldson,
&L Hyten, 1983). The subsequent experiments minimize the potentially
interfering carryover effect of the prior trial by using a longer intertrial
interval (10 s).
The test results, which appear in Fig. 2, strongly suggest that the
Group A-V dyads ultimately learned to coordinate their choice behavior
once the win-stay behavior extinguished. Note that performance on
Same-row trials was comparable among all groups and that only Group
A-V was profoundly disrupted on Different-row trials. Experimenter observations revealed that the disruption on Different-row trials was due
to Group A-V dyads pecking adjacent keys persistently. For example,

El

SAME ROW
DIFFERENT

GROUP A-v

FIG. 2.

Performance

ROW

GROUP h-j

on Same-row and Different-row

test trials in Experiment

I.

48

DAVID

E. HOGAN

on a Different-row
trial in which, say, the upper left and lower right
locations had to be pecked simultaneously? Group A-V dyads would
spend several seconds pecking the keys comprising the top row, then
would shift together to the bottom row of keys and resume pecking for
several seconds, then would shift together and resume pecking the top
row again, etc. Group A-v, on the other hand, was utterly unaffected
by the Different-row
trials, which indicates that the sound of a key being
pecked by one bird did not control the others tendency --to peck an
adjacent location. Apparently, the dyads of Groups A-v and A-V simply
pecked the keys rapidly and independently of one another and were
reinforced when they fortuitously pecked the correct pair of keys.
Kruskal-Wallis tests indicated a nonsignificant difference among groups
on Same-row trials [H(4, 4, 3) = 0.931, but a significant difference on
Different-row
trials [JZ(4, 4, 3) = 7.211. Mann-Whitney
tests verified
that Group A-V was significantly worse on Different-row trials relative
to the other groups [U(4, 7) = 01.
The mean percentage of trials in which one pigeon led the other was
quite close to 50% for all dyads throughout training and testing. Skinner
(1962) reported that an important determinant of leadership in this task
is the relative difference in motivational levels of the two birds (i.e., the
hungrier pigeon leads). Perhaps one pigeon did not consistently lead the
other in the present study because the dyads were maintained at equivalent
motivational levels.
EXPERIMENT

The stimulus control hypothesis (Millard, 1979; Skinner, 1962), which

holds that the spatial choice of the leader was a visual discriminative
stimulus which controlled the spatial response of the follower, provides
an adequate account of the apparent imitative behavior displayed by
Skinners pigeons and by Group A-V dyads. However, the imitative
search behavior may also reflect an unlearned response tendency to
approach and peck near one another, a process which underlies a phenomenon called local enhancement (Thorpe, 1963). Such instinctive
behavior would have clear survival value for hungry pigeons, since food
is likely to be found in the vicinity of another pecking pigeon. Since the
keys comprising the same row of the matrix were separated by only a
few centimeters, local enhancement may account for the followers tendency to peck a key which is adjacent to that pecked by the leader.
The dyads of Experiment 2 were required to solve the multiple-choice
problem under one of three conditions. Group Same dyads were trained
under conditions similar to Group A-V dyads of Experiment 1 (i.e., the
keys comprising the top row were correct on some trials and those
comprising the bottom row were correct on others). Group Different
pigeons, however, were required to peck keys on different rows of the

OBSERVATIONALLEARNINGINPlGEONS

49

matrix for reinforcement

(i.e., the upper left and lower right locations
were correct on some trials, and the lower left and upper right locations
were correct on others). Group Mixed pigeons received a random mixture
of Same-row trials (i.e., the trials presented to Group Same) and Differentrow trials (i.e., the trials presented to Group Different) within each
session. If an unlearned response tendency induces pigeons to orient and
peck close to one another, then Group Same should perform better than
Group Different, and Group Mixed should perform better on Same-row
trials than on Different-row trials. On the other hand, the stimulus control
hypothesis would predict comparable learning rates for Groups Same
and Different (and comparable performance on Same-row and Differentrow trials for Group Mixed), since it does not assume that an unlearned
response tendency governs performance in the cooperation task.
Finally, all dyads were transferred to the mixed-row problem to compare
the strength of conspecific stimulus control in Groups Same and Different.
The stimulus control hypothesis predicts that Group Same dyads should
be disrupted on novel Different-row
test trials, and Group Different
should be equivalently disrupted on novel Same-row test trials.
Method
Subjects

The subjects were 24 food-deprived

Carneaux pigeons.
naive and 20 had prior red/green match to sample training.

Four were

Apparatus

Only one cooperation chamber, which was equipped with a transparent

partition, was used in Experiment 2.
Procedure
Shaping and preliminary
training. All birds were magazine trained
then autoshaped to peck a white key light in the single-key apparatus.
Sessions included IO pairings of the illuminated key (10-s duration) and
reinforcement
(2-s access to grain). The intertrial interval averaged 1
min. Pecks on the illuminated key had no scheduled consequence. Autoshaping continued until all pigeons pecked on at least 90% of the trials
within a session. During the next eight sessions, each of 10 pecks produced
reinforcement and a 10-s ITI. The response latency was recorded on
each trial. Following shaping, 12 subjects were assigned to each compartment of the cooperation chamber such that the mean latency was
balanced over the two compartments.
During the next eight sessions each pigeon was trained in isolation to
peck the top and bottom keys 12 times each in the cooperation chamber.
A single peck produced reinforcement and a 10-s ITI. Twelve matched

50

DAVID

E. HOGAN

pairs were formed, based on latencies pooled over the eight sessions.
Four matched pairs were then assigned to each of three groups, which
were also balanced.
During the following nine sessions, all dyads received forced-choice
training. Group Same dyads received 12 trials with only the top row
illuminated and 12 trials with only the bottom row illuminated. Reinforcement occurred when the dyads pecked the illuminated keys within
1.5 s of each other. The IT1 was IO s. Group Different dyads received
12 trials with the upper left and lower right locations illuminated and 12
trials with the lower left and upper right locations illuminated. Group
Mixed dyads received a random mixture of 6 Same-row trials (i.e., 3
trials in which the top row was correct and 3 trials in which the bottom
row was correct) and 6 Different-row
trials (i.e., 3 trials in which the
upper left and lower right keys were correct, etc.) The trial order was
randomized with the constraint that a particular key location was not
correct on more than 3 consecutive trials.
TruCzg Phuses l-5. Phase I (12 sessions) involved a random mixture
of 12 forced-choice trials and 12 free-choice trials. The forced-choice
trials were structured identically to those of the immediately preceding
phase. On free-choice trials all four keys on the panel were illuminated.
For Group Same, 6 free-choice trials were correct on the top row and
6 were correct on the bottom row. For Group Different, there were 6
free-choice trials with the upper left and lower right locations correct
and 6 with the lower left and upper right correct. Group Mixed dyads
received 3 of each kind of free-choice trial presented to Groups Same
and Different. Reinforcement and a 10-s IT1 occurred when the dyads
pecked on the correct row within 1.5 s of each other. Pecks on the
incorrect row did not terminate the trial. The performance measure was
the latency to peck the correct pair of keys on free-choice trials within
I .5 s. As in Experiment 1, there was no contingency that one bird always
lead the other.
The cooperative response requirement was reduced to 0.5 s during
Phase 2 (9 sessions), 0.2 s during Phase 3 (18 sessions), and 0.1 s during
Phase 4a (12 sessions). During Phase 4b (Sessions 13-36) and Phase 4d
(Sessions 49-54) two cooperative responses on the correct row were
required for each reward. DuringPhase4c (Sessions 37-48) four cooperative
responses were required. The purpose of increasing the number of correct
responses to terminate the trial was to increase perseverative responding
during the subsequent test phase. Since four responses (Phase 4c) caused
a ratio strain in some dyads, the response requirement was reset to
two (Phase 4d). During Phase 5, all dyads were exposed to the mixedrow condition for 6 sessions. Reward occurred after two cooperative
responses on the correct row.

OBSERVATIONAL

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51

Results and Discussion

The acquisition curves appear in Fig. 3. One salient finding was that
performance stabilized much faster than in Experiment 1, perhaps because
the longer IT1 value retarded the development of the interfering winstay behavior.
Also noteworthy, vis-a-vis the local enhancement hypothesis, was that
Groups Same and Different were virtually indistinguishable during each
phase, but both groups performed better than Group Mixed. A casual
inspection revealed that Group Mixed also performed equivalently on
Different- and Same-row trials during every phase. The latency data
were pooled over sessions within each phase and rank ordered. Differences
among groups were not significant during Phase I [H(4, 4, 4) = 4.001,
but they were significant during Phase 2 [H(4, 4, 4) = 9.401. MannWhitney tests indicated that Group Mixed was significantly inferior to
Groups Same and Different [U(4, 8) = 01. The three groups also differed
significantly during Phase 3 [H(4, 4, 4) = 5.921. Again, Group Mixed
was significantly inferior to Groups Same and Different [U(4, 8) = 3.001.
The three groups performed comparably during Phase 4a, but significant
differences were observed during Phases 4b, 4c, and 4d [H(4, 4, 4) =
8.56, 8.32, and 8.32, respectively]. Mann-Whitney
tests confirmed that
Group Mixed was inferior to Groups Same and Different during Phases
4b, 4c, and 4d [U(4, 8) = 1, 0, and 1, respectively).
The test results appear in Figure 4. The latency on familiar training
trials was comparably short for each group. However, performance on
PHASES
2

3
GROUP SAME

o---c
-

GROUP DIFFERENT

GROUP MIXED

40

4d

4~

OLuL-luu~uLl
1

FIG. 3.

131

Acquisition

61
41
BLOCKS OF THREE SESSIONS

of the cooperation

task in Experiment

12

2.

DAVID

52

E. HOGAN

SAME ROW
DIFFERENT

GROUP
SAME

FIG. 4.

Performance

ROW

GROUP
MIXED

GROUP
DIFFERENT

on Same-row and Different-row

test trials in Experiment

2.

the novel test trials was disrupted for both Groups Same and Different,
although more for Group Same. The latencies were pooled over sessions,
and difference scores were calculated by subtracting the latency of
Different-row
trials from that of Same-row trials. A Kruskal-Wallis
test
on the ranked difference scores revealed a significant difference among
groups [H(4, 4, 4) = I I .8]. Mann-Whitney
tests indicated that Group
Mixed was significantly different from Group Same [U(4, 4) = 01 and
also Group Different [U(4, 4) = 01. Mann-Whitney
tests on the absolute
value of the difference scores indicated that Group Same was not disrupted
significantly more than Group Different [U(4, 4) = 21, although the
difference approached significance.
In summary, the local enhancement hypothesis can be rejected because
the acquisition rates of Groups Same and Ditferent were quite comparable,
and Group Mixed performed comparably on Same- and Different-row
trials. Furthermore, the disruption on the reversal test trials was roughly
equivalent in Groups Same and Different. The alternative explanation is
that Groups Same and Different systematically approached and pecked
the appropriate locations because they were reinforced for doing so.
Group Mixed dyads presumably did not control one anothers choice
behavior, since it imparted minimal information about the potentially
correct location.
EXPERIMENT

Experiment 3 assessed the proactive influence of cooperative key pecking

on observational learning of a conditional hue discrimination. The essential
procedure consisted of an observation phase during which one member
of each dyad (a so-called observer) was permitted to watch its partner
(i.e., a so-called demonstrator) perform a well-learned successive con-

OBSERVATIONAL

LEARNING

IN

PIGEONS

53

ditional hue discrimination.

On some discrimination trials, both of the
demonstrators
keys were illuminated with red light, and a peck to the
top key produced reward. On other trials, both keys were illuminated
with green light, and a peck to the bottom key produced reward. The
observers keys were not illuminated during the observation phase, but
in order to maintain control over the observers orienting response by
the demonstrators
discriminative behavior, the observer was rewarded
when the demonstrator emitted a correct response. Phase 2 included a
stimulus control test in which the observer was required to learn the
same discrimination
task which it had seen the demonstrator perform.
Group Same observers were expected to perform above chance on
the initial test trials, because during the observation phase, they presumably
would (a) look at the same (would-be correct) stimuli which the demonstrator pecks (i.e., red in the top location and green in the bottom
location) and (b) associate those stimuli with reinforcement which they
received. However, Group Different observers were expected to perform
below chance because they may experience reinforcement for looking
at the key which was above or below the correct key which the demonstrator pecked (i.e,, the observer may look at red on the bottom key
when the demonstrator was pecking red on the top key, etc.). Chance
performance was anticipated in Group Mixed observers since the observers
orienting response was not likely to be systematically controlled by the
demonstrators
discriminative behavior.
Method
Subjects and Apparatus

The dyads were the same as those of Experiment 2. The apparatus

was the same as that used in Experiment 2, except that red and green
lamps were added to the projectors.
Procedure
Preliminary training. Since the test phase of Experiment 2 may have
weakened cooperative behavior, all of the dyads were retrained on their
original cooperation task approximately
2 weeks after Experiment 2.
During cooperation sessions, the dyads were rewarded when they emitted
two cooperative responses on the correct row within 0.2 s of each other.
Sessions involved 24 trials and were conducted during the morning hours
for 18 days.
During the afternoon hours, one of the dyads (the demonstrator) received
discrimination
training while its partner (the observer) remained in its
home cage. The demonstrator was trained in the outside compartment
of the chamber. On discrimination trials, both keys in the demonstrators
compartment were illuminated red on 12 trials or green on 12 trials. The
trial order was random. The top key was designated correct when the

54

DAVID

E. HOGAN

keys were lit red, and the bottom key was correct when they were green.
Five pecks were required to terminate a trial, and the location of the
fifth peck determined the trial outcome (five pecks were required so that
the observer would have sufficient time to observe the demonstrators
discriminative stimuli during the subsequent study phase of the experiment).
A correct response produced 1.5 s of reward and a 10-s ITI, during which
the key lights were turned off. An incorrect response produced only the
ITI. The performance measure was the percentage of trials which terminated
with reward. Discrimination training for the demonstrators was conducted
for 22 sessions.
Obsemztion und testing phuses. During Phase 1 (5 sessions) of the
experiment proper, the dyads of each group first received a block of 24
cooperation trials (the task parameters were the same as preliminary
training), followed immediately by a block of discrimination study trials.
On study trials the demonstrator was presented 24 discrimination trials
(the discrimination procedure was the same as preliminary training, i.e.,
there were 12 trials in which both keys were illuminated red and 12 trials
in which both keys were green). The observers keys were dark on study
trials, but the observer was rewarded when the demonstrator was rewarded
(i.e., both birds were rewarded if the demonstrators fifth peck occurred
to the top location when the two keys were red, or to the bottom location
when the two keys were green). If the demonstrators fifth response
occurred to the incorrect location, the trial was terminated without reward
and a 10-s IT1 commenced.
Phase 2 sessions (12 sessions) consisted of an initial block of 24 study
trials (which were procedurally identical to the study trials of Phase I)
followed immediately by a block of discrimination test trials. On discrimination test trials, both keys in the observers compartment were
illuminated either red (12 trials) or green (12 trials) and the demonstrators
keys were dark. Both birds were rewarded if the observers fifth response
occurred to the top location when its keys were both red, or to the
bottom location when its keys were both green. Reward consisted of
1.5-s access to grain and a 10-s ITI. If the observers fifth response
occurred to the incorrect location of a trial, the trial was terminated
without reward and a 10-s IT1 commenced. On discrimination test trials,
the performance measure was the percentage of trials which the observer
terminated with a correct response. Cooperation trials were omitted from
Phase 2, because the reward for correct cooperative responses could
have created a problem of satiation during the final discrimination test
period of each day.
Results and Discussion

Performance on cooperation trials, pooled across all sessions was comparable among groups [H(4, 4, 4) = 4.431. Furthermore, the demonstrators

OBSERVATIONAL

LEARNING

IN PIGEONS

55

of Groups Same, Different, and Mixed, did not differ in their discrimination
performance during the Phase 1 study trials. The mean percentage of
correct choices pooled across the Phase 1 study trials for Groups Same,
Different, and Mixed demonstrators was 99.0, 97.6, and 97.8, respectively.
Their performance remained nearly perfect during the Phase 2 study
trials also.
The observers discrimination performance appears in Fig. 5. As may
be seen, the percentage of correct responses of Group Same was superior
to Group Mixed, which in turned was superior to Group DiRerent, especially
during the first block of two sessions. The difference among groups on
the first block was significant [F(2,9) = 4.461. A one-tailed r test indicated
that Group Same was superior to the pooled performance of Groups
Mixed and Ditferent [f(lO) = 1.8381, and Group Different was significantly
inferior to Group Mixed [t(6) = 2.1501. The difference between Groups
Same and Mixed was not significant [f(6) = 0.8891 on the first block but
approached significance on the first session [f(6) = 1.6241. Analyses of
data pooled over all sessions also indicated a significant effect of Groups
[F(2, 9) = 31.301. Again, Group Same was significantly superior to Groups

100

3
BLOCKS

FIG. 5. Discrimination
Mixed in Experiment 3.

performance

GROUP

DIFFERENT

GROUP

MIXED

OF TWO SESSIONS

of the observers of Groups Same, Different, and

54

DAVID

E. HOGAN

Mixed and Different [t(lO) = 2.0661, and Group Mixed was superior to
Group Different [l(6) = 6.1871.
On study trials, experimenter observations revealed that some of the
Group Same observers stood beside the transparent partition and emitted
incipient pecks at the demonstrators
top key when the demonstrator
was pecking red on the top key and at the demonstrators bottom key
when the demonstrator was pecking green on the bottom key. On the
other hand, Group Different observers attempted to peck a key which
was above or below the key which the demonstrator pecked (i.e., if the
demonstrator was pecking red on the top key, the observer would orient
to red on the bottom key, etc.). Group Mixed observers did not display
consistent differential orienting behavior on study trials.
The demonstrators also appeared to prompt correct responses when
the observers performed on discrimination test trials. For example, the
demonstrators attempted to peck the observers top red key when the
observers keys were red and would attempt to peck the observers
bottom green key when the keys were green. Prompting probably was
not necessary for the observed facilitation and interference, however,
because observational learning can be expressed in performance when
the demonstrator is removed from the test situation (Epstein, 1984).
GENERAL DISCUSSION
The major result of Experiments 1 and 2 was that the spatial choice
of one pigeon controlled its partners tendency to peck an adjacent location
in Groups A-V and Same. Furthermore,
the cooperative performance
did not reflect an automatic, unconditioned tendency to peck close to
one another. Local enhancement underlies a form of imitative behavior
observed in neonatal chicks, which approach and peck where the mother
hen pecks (Stokes, 1971: Suboski & Bartashunas, 1984), and is one way
by which chicks learn to discriminate food from inedible objects of similar
size and shape. The locative preference of chicks must be governed by
an innate factor since it occurs so soon after hatching. However, the
locative preference of pigeons of Groups A-V and Same in the cooperation
task was apparently shaped by the positive reinforcement contingency.
Experiment 3 revealed that prior cooperation training affected subsequent
observational learning of a conditional discrimination. The strongest evidence for this claim is the initial beIow-chance performance of Group
Different. The negative transfer suggests that the demonstrators
discriminative behavior served as an instructional stimulus which induced
the observers to look at the would-be incorrect stimulus on study trials.
Furthermore, since the observer received reinforcement on study trials,
those stimuli may have become associated with reinforcement through
autoshaping (Brown & Jenkins, 1968; Boakes & Gaertner, 1977) and
elicited approach and pecking on test trials,

OBSERVATIONAL

LEARNING

IN PIGEONS

57

The slight facilitation exhibited by Group Same observers is also consistent with the instructional control plus autoshaping hypothesis. Presumably, the observers looked at the would-be correct stimuli which the
demonstrator pecked on study trials and associated those stimuli with
reinforcement.
However, since Group Same was not significantly better
than Group Mixed, this comparison does not provide the strongest available
evidence for a proactive influence of cooperation on observational learning.
Apparently, cooperation training did not establish the demonstrators
response as a sufficiently strong instruction in all of the Group Same
dyads.
Observational
learning research may provide some insight into the
learning mechanisms which underlie group foraging in pigeons. It is not
uncommon to observe that the presence of a few pigeons pecking on
the ground can entice other pigeons to approach and peck nearby. An
obvious explanation for why observers are attracted to the feeding site
is because they see the food itself, but in many instances the small grains
are concealed by grass or foliage and are barely discernible. The prevailing
ethological explanation is that the sight of other foraging pigeons unconditionally elicits approach and pecking in observers. The innate mechanism would more or less ensure that the observer would be drawn to
familiar or novel food sources and thus reduces its dependence on learning
to locate food sites by trial and error. However, the tendency of
pigeons to approach and peck where others are pecking may be less
dependent on an innate response disposition than the ethological perspective
suggests. Skinners (1962) experiment and Experiment 3 of the present
study suggest that a pigeons tendency to peck novel objects in novel
locations or to peck discriminatively at novel visual stimuli which signal
food reinforcement
may reflect the observers history of reinforcement
for pecking in the presence of other pecking pigeons.
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Analysis

Received March 3, 1985

Revised September 30, 1985; November

11, 1985