Behavioural Brain Research 222 (2011) 51–56
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Behavioural Brain Research
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Research report
Visual Wulst analyses “where” and entopallium analyses
“what” in the zebra finch visual system
Shigeru Watanabe
a,∗
, Uwe Mayer
b
, Hans-Joachim Bischof
b
a
Department of Psychology, Keio University, Mita 2-25-45, Minato-Ku, Tokyo, Japan
b
Department of Behavioural Biology, Bielefeld University, 33501 Bielefeld, Germany
article info
Article history:
Received 2 February 2011
Received in revised form 10 March 2011
Accepted 14 March 2011
Keywords:
Birds
Zebra finch
Visual system
Pattern discrimination
Spatial discrimination
Brain lesions
abstract
Quite a lot of studies have tried to elucidate the differences in function of the two telencephalic targets of
the avian visual system. We have tried to find out how the two systems are involved in orientation towards
a food tray which is either marked by a special pattern or has to be identified by its relation to spatial
cues. In this report, we compared in the zebra finch the effects of Wulst lesions on pattern discrimination
with Wulst lesion effects on spatial discrimination, and we examined the effect of entopallium lesions
on spatial discrimination. Birds with Wulst lesions showed deficits in spatial discrimination, but not in
pattern discrimination. Entopallial lesions caused no deficits in spatial discrimination tasks. Combining
the present results with a previous study revealing an impairment of pattern discrimination by such
entopallial lesions [19], we are able to demonstrate a double dissociation: namely, an impairment of
pattern discrimination by entopallial lesions and impairment of spatial discrimination by Wulst lesions,
but no effects of the opposite pairing of task and lesion site. The entopallium is thus involved if the food
source is identified by a pattern, and the Wulst if it has to be found by spatial cues.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
In birds and mammals, visual information is processed by three
visual pathways. The so-called accessory system processes reti-
nal image motion as produced by self motion of the subject. The
two other pathways, the lemnothalamic and the collothalamic one,
are thought to be involved in object detection and recognition. In
birds, the telencephalic terminal of the lemnothalamic projection is
called the “visual Wulst,” and consists of the hyperpallium apicale
(HA), the hyperpallium intercalatum (HI), and the hyperpallium
densocellulare (HD). It has been considered to be homologous to
the primary visual cortex in mammals [11], but the finding of more
than one retinotopic map in this region [8] may suggest that it also
corresponds to other areas of the mammalian visual neocortex. The
avian entopallium is the telencephalic terminal of the collotha-
lamic pathway and a part of the dorsal ventricular ridge (DVR).
In most mammals, the lateral geniculate–visual cortex projection
(lemnothalamic pathway) is believed to be the main visual path-
way, whereas in birds, the collothalamic pathway is the dominant
one [4].
After a pioneering study by Hodos and Karten [5], it has been
shown frequently that entopallial lesions cause deficits in pattern
∗
Corresponding author.
E-mail address: swat@flet.keio.ac.jp (S. Watanabe).
discrimination in pigeons [2,12–14]. Likewise, entopallial lesions
led to deficits in discriminating pseudoconcepts, in which four food
items and four nonfood items were classified into two arbitrary
(pseudo) groups. Interestingly, no such deficits were detected in
the case of food vs. non-food concept discrimination. The birds
had already natural concepts about food before the training while
they had to learn the pseudoconcepts in the course of the training
[12].
Information on the function of the entopallium in birds other
than pigeons is sparse. In a first attempt to investigate the
interplay of the two pathways with hippocampus, we have com-
pared the effects of entopallial lesions with those of hippocampal
lesions on pattern discrimination in zebra finches [19]. The birds
were trained to discriminate a feeder with a dot pattern from
feeders with a stripe pattern. When the birds had learned the
discrimination, either the entopallium or the hippocampus was
damaged. The entopallial lesions caused deficits in this visual pat-
tern discrimination, whereas hippocampal lesions did not. Effects
of the entopallium lesions on pattern discrimination agree with
effects shown in our previous experiments in pigeons. The avian
hippocampus, in contrast, is obviously not involved in pattern dis-
crimination learning, but, as we have shown previously, plays a
dominant role in spatial learning [18].
Wulst lesions do not cause deficits in visual discrimination in
pigeons [5,13,14]. Lesions of the principal optic thalamic nuclei of
the lemnothalamic projection had little or no effect on intensity
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doi:10.1016/j.bbr.2011.03.035