Physiology & Behavior, Vol. 34, pp. 575-577. Copyright ©Pergamon Press Ltd., 1985. Printed in the U.S.A. 0031-9384/85 $3.00 + .00
Lateralization of Visually Controlled
Behavior in Pigeons
ONUR GUNTORKON
Experimentelle Tier.psychologie, Psychologisches lnstitut, Ruhr-Universitiit Bochum, D 4630 Bochum I, FRG
Received 16 August 1983
GONTORKON, O. Lateralization of visually controlled behavior in pigeons. PHYSIOL BEHAV 34(4) 575-577, 1985.-
Pigeons were trained binocularly on two different successive pattern discrimination tasks. After reaching criterion they
performed the tasks with either the left or the right eye occluded or with both eyes unobstructed. The number of responses
emitted was greater when the fight rather than the left eye was uncovered. Also the percent correct discrimination scores
were better when seeing with the right eye than with the left one. Some pigeons originally acquired the discrimination
monocularly, generally using the right eye, although both eyes were uncovered. The results are discussed in relation to
brain structures that may mediate this performance.
Lateralization Visual system Visual discrimination learning Pigeon
DESPITE an early report of left-handedness in parrots [4],
lateraiization has been thought to be an exclusive attribute of
higher primates. Nottebohm [15] however, showed that le-
sions of a part of the left hypoglossal nerve reduced the song
repertoire of male canaries by a greater amount than did
lesions of the right nerve. Later studies also demonstrated
that the left telencephalic vocalizational areas have a greater
influence on the song production in this bird than the right
ones [16]. Howard et al. [10] showed that injection of gluta-
mate, a putative amino acid transmitter, into the left fore-
brain hemisphere of chicks enhanced the amount of sexual
and aggressive behavior and impeded auditory habituation
and the visual discrimination of grain from pebbles. Similar
results were also obtained with cycloheximide, an inhibitor
of ribosomal protein synthesis [19]. Andrew et aL [1] found
marked differences in the visual habituation of chicks de-
pending on whether the stimuli were seen with the right or
the left eye. In an elegant study by Rogers [19], the laterali-
zation of visually guided behavior in chicks was shown to be
due to the differential exposure to light of the two eyes dur-
ing a critical embryonic period.
The visual system of birds is particularly appropriate
for studying information processing in the separate brain
hemispheres. The axons of the optic nerves decussate
completely at the chiasma and project to the contralateral
tectum and thalamus. A tecto-thalamo-telencephalic and a
thalamo-telencephalic pathway ascend mainly to the ipsilat-
eral forebrain [2]. Birds do not have a corpus callosum but
have instead a number of smaller commissures in which the
two visual pathways recross only to a small extent to the
contralateral brain hemisphere. The degree of transfer of
monocularly learned information to the contralateral brain
depends on the specific experimental conditions [7,20].
The aim of this study was to investigate whether laterali-
zation occurs in the context of complex visual discrimination
tasks.
METHOD
Eight experimentally naive adult homing pigeons (Col-
umba livia) of local stock were used. They were maintained
at 80% of their normal body weight throughout the experi-
ments. A conventional Skinner-box with a single response key
on which the visual stimuli were back-projected with a multi-
channel microprojector was used. Rewards were delivered
with a solenoid activated food-hopper, situated below the
response key. A reinforcement light accompanied food re-
ward. The box, which was located in a sound-attenuated
chamber, was illuminated by a houselight. All relevant
events were programmed with digital modular equipment,
which also counted the key-pecking responses. The pecking
of the white-light illuminated key was shaped using an au-
toshaping procedure. When the animals had learned to
associate key pecking with food reward discrimination train-
ing began. Two stimuli, shown in Fig. 1, were successively
projected onto the response key in a quasi-random sequence
[3]. For half of the pigeons the first of the two stimuli was
deemed to be correct and for the other half the second. Pecks
to the correct stimulus yielded 4 sec food access. Each peck
on the incorrect stimulus extended the projection time for 2
sec thus ensuring extinction of pecking to the incorrect pat-
tern. Without such extension each trial lasted 20 sec. To
assess the frequency of pecking to the correct and the incor-
rect stimuli only the responses during the standard first 20
sec were considered. Each session consisted of 40 trials.
At the beginning of the discrimination the pigeons were
reinforced on a VR 4 schedule. After reaching 85% of correct
responses in one session the ratio was increased to VR 8,
then to VR 16 finally to VR 32. After reaching 85% of correct
responses in three consecutive sessions with a VR 32
schedule the acquisition phase was completed and the pi-
geons underwent surgery. While the animals were anaesthe-
tized (Equithesin, 0.5 ml per 100 g bodyweight) the scalp was
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