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Behavioural Brain Research
Volume 198, Issue 1, 2 March 2009, Pages 214-223
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Copyright © 2008 Elsevier B.V. All rights reserved.

Research report

Neuronal encoding of meaning: Establishing category-selective response patterns in the avian ‘prefrontal cortex’

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Janina A. Kirscha, c, Corresponding Author Contact Information, E-mail The Corresponding Author, Ioannis Vlachosc, Markus Hausmannb, Jonas Rosea, Man Yi Yimc, Ad Aertsenc and Onur Güntürküna

aDepartment of Biopsychology, Institute of Cognitive Neuroscience, Faculty of Psychology, Ruhr-University Bochum, 44780 Bochum, Germany

bDepartment of Psychology, Durham University, Durham DH1 3LE, England, United Kingdom

cBernstein Center for Computational Neuroscience, Albert-Ludwigs University Freiburg, 79104 Freiburg, Germany

Received 30 July 2007; 
revised 28 October 2008; 
accepted 2 November 2008. 
Available online 12 November 2008.


Forebrain association areas interweave perceived stimuli with acquired representations of own actions and their outcome. Often, relevant stimuli come in a bewildering variety of shapes and sizes and we slowly have to learn to group them into meaningful categories. Therefore, the aim of the present study was twofold: First, to reveal how single units in the pigeon's nidopallium caudolaterale (NCL), a functional analogue of the mammalian prefrontal cortex (PFC), encode stimuli that differ in visual features but not in behavioral relevance. The second aim was to understand how these categorical representations are established during learning. Recordings were made from NCL neurons while pigeons performed a go–nogo categorization paradigm. Responses during presentation of the two S+ stimuli and non-responding during presentation of the two S− stimuli were followed by reward. We recorded from two pigeons at different learning stages. In the beginning of the learning process, neurons were active during and shortly before reward, but only in go trials. These data suggest that during the early phase of learning avian ‘prefrontal’ neurons code for rewards associated with the same behavioral demand, while ignoring feature differences of stimuli within one category. When learning progressed, (1) category selectivity became stronger, (2) responses selective for nogo stimuli appeared, and (3) reward-related responses disappeared in favor of category-selective responses during the stimulus phase. This backward shift in time resembles response patterns assumed by the temporal difference (TD) model of reinforcement learning, but goes beyond it, since it reflects the neuronal correlate of functional categories.

Keywords: Pigeon; Columba livia; Forebrain; Nidopallium caudolaterale (NCL); Multiple single-unit activity; Functional categorization

Article Outline

1. Introduction
2. Materials and methods
2.1. Behavioral task
2.2. Recording procedures
2.3. Data analysis
2.3.1. Behavior
2.3.2. Neuronal responses Spike detection and spike sorting Classification of neuronal responses ROC analysis of categorical neurons
2.4. Histology
3. Results
3.1. Behavior
3.2. Electrophysiology
3.2.1. Responses of categorical neurons
3.2.2. Responses of non-categorical neurons
3.2.3. Histological reconstruction of the recordings sites
3.2.4. ROC analysis of categorical neurons
4. Discussion

Corresponding Author Contact InformationCorresponding author. Tel.: +49 761 203 9575; fax: +49 761 203 9559.
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Behavioural Brain Research
Volume 198, Issue 1, 2 March 2009, Pages 214-223
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