Brief Communication 29
How birds use their eyes: Opposite left-right specialization for
the lateral and frontal visual hemifield in the domestic chick
Giorgio Vallortigara*†, Claudio Cozzutti‡, Luca Tommasi‡,
and Lesley J. Rogers§
Recent evidence has demonstrated that, in animals
with laterally placed eyes, functional cerebral
asymmetry is revealed by preferential use of either
the left or right eye in a range of behaviors (birds:
[1, 2, 3]; fish: [4, 5]; reptiles: [6, 7]). These findings
pose a theoretical problem. It seems that there
would be disadvantages in having a substantial
degree of asymmetry in the use of the two eyes; a
deficit on one side would leave the organism
vulnerable to attack on that side or unable to
exploit resources appearing on one side. We here
report a possible solution to the problem. We have
found that domestic chicks show selective use of
the lateral visual field of the left eye and of the
right hemifield in the binocular, frontal visual field
when they peck at strangers but not at cagemates.
Thus, during social recognition, there seems to be
opposite and complementary left-right
specialization for the lateral and frontal visual fields
of the two eyes. These findings can reconcile the
computational advantages associated with
asymmetry of the left and right sides of the brain
with the ecological demands for an animal to
perceive and respond equally well to the left and
right sides of its midline.
Addresses: *Department of Psychology, Animal Cognition and
Comparative Neuroscience Laboratory, University of Trieste, and
†B.R.A.I.N. Centre for Neuroscience, Trieste 34123, Italy. ‡Department
of General Psychology, University of Padova, Padova 35131, Italy.
§School of Biological Sciences, Division of Zoology, University of New
England, Armidale NSW 2351, Australia.
Correspondence: Giorgio Vallortigara
E-mail: vallorti@univ.trieste.it
Received: 6 October 2000
Revised: 8 November 2000
Accepted: 21 November 2000
Published: 9 January 2001
Current Biology 2001, 11:29–33
0960-9822/01/$ – see front matter
 2001 Elsevier Science Ltd. All rights reserved.
Results and discussion
Chicks were tested, on day 3 after hatching, in pairs com-
posed of companions (cagemates) or strangers. Social
pecking by each chick at its test partner was scored. Since
previous work had revealed that exposure of the embryo
to light during the final days of incubation determines
lateralization of attack and some other visually guided
responses in the chick [8, 9], we used both chicks hatched
from eggs maintained in the dark until hatching (D-chicks)
and chicks hatched from eggs exposed to light during
the last three days before hatching (L-chicks). For light
exposure, a 60 W lamp provided 300 Lux within the
incubator. We reared 40 L-chicks and 36 D-chicks in pairs
at a controlled temperature (30–35⬚C) in metal cages (30 ⫻
30 ⫻ 40 cm) that were illuminated from above by fluores-
cent lamps. For each pair of companion chicks, another
pair was selected, and the two pairs were tested simultane-
ously (either as pairs of companions or pairs of strangers)
in two separate cages (50 ⫻ 60 ⫻ 30 cm). The behavior
of the chicks was videorecorded for 5 min. Since data for
an individual subject could not be considered to be en-
tirely independent of the other chick in the pair, data for
each pair were considered together as one score (light
companions, N ⫽ 10; light strangers, N ⫽ 10; dark com-
panions, N ⫽ 8; dark strangers, N ⫽ 10). The numbers
of pecks directed to the conspecific and to the environ-
ment (at the floor or walls of the cage) were measured
from the videotapes. In addition, we measured the fre-
quency with which the chick used its lateral and frontal
fields of vision to fixate before pecking. We made these
measurements by using transparencies to superimpose
protractors on the monitor screen (Figure 1). We classified
angles between ⫾ 15 degrees of the midline as use of the
frontal (binocular) visual field and the remaining angles
as use of the lateral visual field [1]. Note that, although
the chick pecks by a ballistic movement of its head for-
ward from its midline, it fixates and decides whether to
peck or not prior to this by turning its head to view the
stimulus and using the high-acuity regions of its retina.
Figure 2 shows that, as expected, both L- and D-chicks
pecked more at strangers than at companions. Pecks at
the environment, in contrast, were higher in D-chicks
than in L-chicks and tended to be higher among strangers
in L-chicks and among companions in D-chicks.
Use of the lateral and frontal visual fields before pecking
shows opposite directions of lateralization in D-chicks
(Figure 3): D-chicks preferentially used the left, lateral
monocular field and the right, frontal hemifield before
pecking at strangers. L-chicks showed significant use of
the left, lateral monocular field but, in contrast to the
D-chicks, no differences with respect to the frontal hemi-
fields. A more detailed analysis of the different targets of
pecking (Figure 4) confirmed these data and showed that
30 Current Biology Vol 11 No 1
Figure 1
Figure 2
Number of pecks directed at a conspecific and at the environment in
pairs of companion and stranger chicks. There were more pecks at
strangers than at companions in both L- and D-chicks [test condition:
F(1, 34) ⫽ 16.313, p ⬍ 0.001; hatching condition: F(1, 34) ⫽ 0.050
(not significant, or “ns”); test ⫻ hatching: F(1, 34) ⫽ 0.300 (ns)].
Separate analyses showed that the companions/strangers difference
was evident (see bottom row) only for pecks at the head [F(1, 34) ⫽
10.108, p ⫽ 0.003], and the body [F(1, 34) ⫽ 19.804, p ⬍ 0.001]
Schematic representation of a chick’s use of
the frontal or lateral field of vision before
pecking (␣ indicates the angle used to peck
at the target in this case).
but not for pecks at the feet [F(1, 34) ⫽ 0.286 (ns)]. Pecking at the
environment was the same among pairs of companions and
strangers [F(1, 34) ⫽ 0.470, p ⫽ 0.498]; however, pecking at the
environment was higher among D- than among L-chicks [F(1, 34) ⫽
6.087, p ⫽ 0.019], and there was a tendency [interaction: F(1,34) ⫽
3.815, p ⫽ 0.059] in L-chicks for higher pecking among strangers
and in D-chicks for higher pecking among companions.

Figure 3
Chicks’ use of the lateral field before pecking at a conspecific showed
a significant effect of testing conditions [F(1, 34) ⫽ 7.387, p ⫽ 0.010]
in both L- and D-chicks; there were no other statistically significant
effects [hatching: F(1,34) ⫽ 0.18 (ns); test ⫻ hatching: F(1,34) ⫽
0.003 (ns)]. Chicks showed a significant preference (one-sample
t-test; two-tailed; see figure) for using the left lateral field when they
were tested in pairs of strangers. Use of the frontal field before pecking
at a conspecific showed a significant effect of testing conditions
[F(1,34) ⫽ 4.175, p ⬍ 0.05], but not of hatching [F(1,34) ⫽ 0.165
the effects were largely confined to hemifield use before
pecks at the head. No significant preferences in hemifield
use were apparent before pecking at the environment
(Figure 3).
These findings provide a solution to a puzzling phenome-
non. A number of independent reports have provided
convincing evidence that animals with laterally placed
eyes show preferential left and right use in different tasks
(reviewed in [10]). For instance, chicks use the left eye
to look at an aerial predator [2] and the right eye to look
at a familiar imprinting stimulus [1, 11]. Anolis lizards [6,
7] and toads [12, 13] use the left eye during agonistic
encounters, and fish use the right eye to look at a danger-
ous stimulus [14] and the left eye to look at conspecifics
[5]. Results of studies that use selective occlusion of one
eye complement these results by showing that birds’ per-
formance in a variety of tasks depends on which eye they
use (reviews in [15, 16, 17]). However, having one eye
that is better at responding to a predator or recognizing
a partner appears to be a disadvantage on ecological
Brief Communication 31
(ns)]; the test ⫻ hatching interaction was, however, significant [F(1,34) ⫽
5.335, p ⬍ 0.05]. D-chicks, but not L-chicks, showed a significant
preference (one-sample t-test, two-tailed; see figure) for using the right
frontal field when tested as pairs of strangers.
There were no statistically significant effects associated with eye use
before pecking at the environment [lateral field — hatching, F(1,34) ⫽
0.023; test, F(1,34) ⫽ 0.066; test ⫻ hatching, F(1, 34) ⫽ 0.137.
Frontal field — hatching, F(1,34) ⫽ 0.035; test, F(1,34) ⫽ 1.820;
test ⫻ hatching, F(1,34) ⫽ 0.810].
grounds; other things being equal, these stimuli might
happen to be located on either side at random. The answer
that has been provided so far is that the computational
advantages associated with possession of an asymmetric
brain [18, 19, 20] should compensate for the ecological
disadvantages of perceiving and responding with less effi-
ciency to one or the other side of the body. However, our
results suggest a different answer. The two eyes of the
domestic chick seem to provide visual information that
can be used for social recognition, but they utilize differ-
ent parts of the visual field; of the two lateral fields, the
left is used preferentially, whereas in the binocular field
the area to the right of the midline (right binocular hemi-
field) is used preferentially. Interestingly, previous evi-
dence had shown that chicks wearing monocular eye
patches were unable to discriminate between companions
and strangers when they were using their right eye [21,
22] and that, in contrast, chicks with blinkers covering
the right frontal field (but leaving the lateral field unob-
structed) showed difficulty in inhibiting pecks at compan-
ions [23]. This is now understandable because use of eye
32 Current Biology Vol 11 No 1
Figure 4
Chicks’ use of the lateral field before pecking at different parts of the
conspecific revealed a significant difference between companions
and strangers for pecks at the head [F(1, 34) ⫽ 12.226, p ⫽ 0.001],
whereas use of the frontal field revealed a significant difference for
patches prevents (or disturbs) use of the frontal visual
field and thus reveals only part of the true specialization
of the two eyes.
Unlike the pigeon, the chick has only one region of high
ganglion-cell density, providing high-acuity vision, that is
directed primarily to the lateral monocular field [24, 25].
This area of the retina of only the left eye is used before
a peck is directed toward a stranger. An area of high but
somewhat lower ganglion-cell density extends away from
this central region in the horizontal plane and just into
the temporal retina (i.e., it receives input from the frontal
field). This provides acuity vision into the binocular fields,
but there is no overlap across the midline of the high-
acuity regions of each eye (see Figure 10 of [24]) unless
the eyes are converged. It is, therefore, likely that when
a chick fixates binocularly before pecking, it may be using
only the right half of the binocular field. We might there-
fore conclude that, before pecking a stranger, the chick
uses either the lateral field of its left eye (right hemi-
sphere) or the right frontal field (left hemisphere or per-
haps both hemispheres if the eyes are converged). Thus,
it appears that there is a form of complementary specializa-
tion of the eye fields, at least in the case of chicks incu-
bated in the dark. Chicks exposed to light also show
preferential use of the left lateral field for pecking a
both pecks at the head [F(1, 34) ⫽ 5.510, p ⫽ 0.025] and pecks at
the feet [F(1, 34) ⫽ 0.047]. There were no other statistically
significant effects.
stranger, but they use both the left and right binocular
fields. These findings may indicate better integration of
the hemispheres, which could result because early experi-
ence with light allows better coordination of visual input
from the two eyes.
It is interesting to note that, in both D- and L-chicks,
the left eye sends its input via the thalamofugal pathway
more strongly to the right hemisphere than to the left
hemisphere and via the fast relay system of the tecto-
fugal pathway also only to the right hemisphere (i.e., via
the rotundal-ectostriatal connections without collateral
branches [26]). (Note that it is likely that the slower sys-
tem of the tectofugal pathway, via the unmyelinated ro-
tundal-ectostriatal projections with collateral branches
that cross the midline, is not used for attack pecking [26].)
Inputs from the right eye differ in D- and L-chicks. In
L-chicks, the right eye sends inputs to both hemispheres
via the thalamofugal pathway and, again, via the fast sys-
tem of the tectofugal pathway to only the left hemisphere
[27]. In contrast, the right eye of D-chicks sends inputs
almost entirely to the left hemisphere via both systems.
The left and right frontal fields of D-chicks are, therefore,
less integrated than those of L-chicks, and that could
explain why the frontal-field asymmetry is revealed only
in D-chicks.
 Brief Communication 33

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