A Special Issue of Cognitive Neuropsychology The organisation of conceptual knowledge in the brain: Neuropsychological and neuroimaging perspectives Edited by Alex Martin National Institute of Mental Health, Bethesda, MD, USA and Alfonso Caramazza Harvard University, Cambridge, MA, USA HOVE AND NEW YORK Published in 2003 by Psychology Press Ltd 27 Church Road, Hove, East Sussex, BN3 2FA www.psypress.co.uk This edition published in the Taylor & Francis e-Library, 2005 “To purchase your own copy of this or any of Taylor & Francis or Routledge’s collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk.” Simultaneously published in the USA and Canada by Taylor & Francis Inc 29 West 5th Street, New York, NY 10001, USA Psychology Press is part of the Taylor & Francis Group © 2003 by Psychology Press Ltd All rights reserved No part of this book may be reprinted or reproduced or utilised in any form or by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying and recording, or in any information storage or retrieval system, without permission in writing from the publishers British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 0-203-50122-5 Master e-book ISBN ISBN 0-203-59549-1 (Adobe eReader Format) ISBN 1-84169-947-0 (hbk) ISSN 0264–3294 (Print Edition) This book is also a special issue of the journal Cognitive Neuropsychology and forms Issues 3, 4, 5, and of Volume 20 (2003) Cover design by Joyce Chester COGNITIVE NEUROPSYCHOLOGY Volume 20 Issue 3/4/5/6 May-September 2003 Contents Neuropsychological and neuroimaging perspectives on conceptual knowledge: An introduction A.Martin and A.Caramazza What are the facts of semantic category-specific deficits? A critical review of the clinical evidence E.Capitani, M.Laiacona, B.Mahon, and A.Caramazza 19 A case series analysis of “category-specific” deficits of living things: The HIT account G.W.Humphreys and M.Jane Riddoch 81 Semantic dementia with category specificity: A comparative case-series study M.A.Lambon Ralph, K.Patterson, P.Garrard, and J.R.Hodges 130 Category specificity and feature knowledge: Evidence from new sensory-quality categories F.Borgo and T.Shallice 152 The selective impairment of fruit and vegetable knowledge: A multiple processing channels account of fine-grain category specificity S.J.Crutch and E.K.Warrington 184 A case of impaired knowledge for fruit and vegetables D.Samson and A.Pillon 204 Genetic and environmental influences on the organisation of semantic memory in the brain: Is “living 401 things” an innate category? M.J.Farah and C.Rabinowitz 235 iv Neural correlates of conceptual knowledge for actions D.Tranel, D.Kemmerer, R.Adolphs, H.Damasio, and A.R.Damasio 244 Constraining questions about the organisation and representation of conceptual knowledge B.Z.Mahon and A.Caramazza 270 The similarity-in-topography principle: Reconciling theories of conceptual deficits W.K.Simmons and L.W.Barsalou 290 Three parietal circuits for number processing S.Dehaene, M.Piazza, P.Pinel, and L.Cohen 330 The influence of conceptual knowledge on visual discrimination I.Gauthier, T.W.James, K.M.Curby, and M.J.Tarr 352 Role of mental imagery in a property verification task: fMRI evidence for perceptual representations of conceptual knowledge I.P.Kan, L.W.Barsalou, K.O.Solomon, J.K.Minor, and S.L.Thompson-Schill 373 Do semantic categories activate distinct cortical regions? Evidence for a distributed neural semantic system L.K.Tyler, P.Bright, E.Dick, P Tavares, L.Pilgrim, P.Fletcher, M.Greer, and H.Moss 390 How is the fusiform gyrus related to category-specificity? 412 v C.J.Price, U.Noppeney, J.Phillips, and J.T.Devlin Neural foundations for understanding social and mechanical concepts A.Martin and J.Weisberg 428 Subject index 442 COGNITIVE NEUROPSYCHOLOGY This journal promotes the study of cognitive processes from a neuropsychological perspective Cognition is understood very broadly, as including for example perception, attention, object recognition, planning, language, thinking, memory, and action It covers neuropsychological work bearing on our understanding of normal cognitive processes as well as neuropsychological disorders of cognition arising at any stage of life span All manuscript submissions should be addressed to Sophie Forster, Journals Editorial Assistant, Psychology Press Ltd, 27 Church Road, Hove, East Sussex BN3 2FA, UK Notes for contributors are available from the publisher on request Subscription information Cognitive Neuropsychology is published by Psychology Press Ltd, a member of the Taylor & Francis Group New subscriptions and changes of address should be sent to: Psychology Press, c/o Taylor & Francis Ltd, Rankine Road, Basingstoke, Hampshire, RG24 8PR, UK Send change of address notices at least six weeks in advance, and include both old and new addresses Subscription rates to Volume 20, 2003 (8 issues) are as follows (prices inclusive of postage and packing): To individuals: £176.00 (UK); $290.00 (Rest of world) To institutions: £544.00 (UK); $897.00 (Rest of world) Cognitive Neuropsychology (USPS permit number 016265) is published eight times per year in February, March, May, June, July, September, October, and December The 2003 US Institutional subscription price is $897.00 Periodicals postage paid at Champlain, NY, by US Mail Agent IMS of New York, 100 Walnut Street, Champlain, NY US Postmaster: Please send address changes to pCGN, PO Box 1518, Champlain, NY 12919, USA Copyright The material published in this journal is copyright No part of this journal may be reproduced in any form, by photostat, microform, retrieval system, or any other means, without the prior written permission of the publisher Information about Psychology Press journals and other publications is available from http://www.psypress.co.uk Go to http://www.tandf.co.uk/journals/pp/02643294.html for current information about this journal, including how to access the online version or to register for the free table of contents alerting service Cognitive Neuropsychology is covered by the following abstracting, indexing, and citation services: Current Contents (ISI); ASSIA; APA PsycINFO; Sociological Abstracts; Biosciences Information Service; MLA International Bibliography; EMBASE; LLBA; Neuroscience Citation Index (ISI); Research Alerts (ISI); Social SciSearch (ISI); Social Science Citation Index (ISI); Social Services Abstracts; SciSearch (ISI); CDAB; UnCover; Linguistics Abstracts vii Printed in the United Kingdom by Henry Ling Limited, at the Dorset Press, Dorchester, DTl 1HD This publication has been produced with paper manufactured to strict environmental standards and with pulp derived from sustainable forests © 2003 Psychology Press Ltd NEUROPSYCHOLOGICAL AND NEUROIMAGING PERSPECTIVES ON CONCEPTUAL KNOWLEDGE: AN INTRODUCTION Alex Martin National Institute of Mental Health, Bethesda, USA Alfonso Caramazza Harvard University, Cambridge, USA The modern era of study of the representation of object concepts in the human brain began in 1983 with a report by Warrington and McCarthy of a patient with preserved knowledge for animals, foods, and flowers, relative to inanimate objects (Warrington & McCarthy, 1983) This was followed the next year by a report of four patients with the opposite pattern of preserved and impaired category knowledge (Warrington & Shallice, 1984) Specifically, these patients presented with a relatively selective impairment for knowing about living things and foods Since publication of these seminal case studies, over 100 patients have been reported with a category-specific deficit for biological categories (living things, especially four-legged animals), relative to inanimate objects (especially tools and other artifacts), and more than 25 cases with the opposite pattern of deficit (Figure 1) Heightened appreciation of the importance of these clinical cases for understanding the organisation of conceptual knowledge, as well as for object recognition, the organisation of the lexicon, and the storage of long-term memories, has also motivated an increasing number of functional brain-imaging studies of object category representation in the normal human brain The goal of this special issue of Cognitive Neuropsychology is to provide a forum for new findings and critical, theoretical analyses of existing data from patient and functional brainimaging studies THE THEORIES OF CONCEPT ORGANISATION A number of different theoretical positions have been advanced to explain category-specific deficits However, as described by Capitani, Laiacona, Mahon, and Caramazza (2003-this issue), much of the current debate centres on whether concepts are organised by property or by category.¹ Most investigators assume that the deficits are a direct consequence of the organisation of object properties in the brain The best known property-based model of semantic category-specific deficits is the sensory/functional theory (S/ FT), proposed by Warrington, Shallice, and McCarthy (Warrington & McCarthy, 1987; Warrington & Shallice, 1984) Although there are important differences among them, similar accounts have been Note, however, that the two types of organisation need not be mutually exclusive It is possible that concepts are organised into domains and within domains the organisation may very well be by property type or correlation (Caramazza, 1998; see also Mahon & Caramazza, 2003-this issue) Requests for reprints should be addressed to Alex Martin, PhD, Laboratory of Brain and Cognition, National Institute of Mental Health, Building 10, Room 4C-104, 10 Center Drive MSC 1366, Bethesda, Maryland 20892-1366, USA (Email: alex@ codon nih.gov) © 2003 Psychology Press Ltd http://www.tandf.co.uk/journals/pp/02643294.html DOI: 10.1080/02643290342000050 THE ORGANISATION OF CONCEPTUAL KNOWLEDGE IN THE BRAIN Figure Cumulative number of patients with category-specific disorders for biological objects and artefacts reported in the literature since 1983 Based on the review provided by Capitani et al (2003-this issue) proposed by a number of other investigators (e.g., Damasio, 1989; Humphreys & Forde, 2001; Martin, Ungerleider, & Haxby, 2000) The central idea behind S/FT-like theories is that conceptual knowledge about objects is organised by sensory features (e.g., form, motion, colour, smell, taste) and functional properties (the motor habits related to their use, typical location where they may be found, their social value, etc.).2 Categories differ as to the importance or weight assigned to each of these properties In this view, categoryspecific (C-S) semantic disor ders occur when a lesion disrupts knowledge about a particular property or set of properties critical for defining that object category and for distinguishing among its members Thus damage to regions where information about object form is stored will produce a C-S disorder for animals This is because visual appearance is assumed to be a critical property for defining animals, and because the distinction between different animals is assumed to be heavily dependent on knowing about subtle differences in their visual form (e.g., distinguishing among four-legged animals) A critical component of these models is that the lesion should affect knowledge of all object categories with these characteristics, not only animals In a similar fashion, damage to regions where information about how an object is used Theories differ as to what is meant by “functional” properties In the early literature, “functional” was used together with “associative” (functional-associative) to distinguish sensory from nonsensory properties of objects (e.g., Farah & McClelland, 1991) When used in the context of S/FT it has generally been interpreted in this sense However, in some theories the term “functional” is restricted to the sense “use” and in others to the sense “motor habit.” Sensory/motor theories of the representation of objects havt tended to favour the latter sense (Martin et al., 2000) However, if we were to restrict “functional” to mean “motor habit” we would only be able to use the term “function” for a very small set of objects—primarily tools This can be easily appreciated when we consider the functions of various artefacts Thus, although some functions are associated with fairly specific motor patterns (e.g., scissors: used for cutting with a highly specific motor pattern), others are not associated with any specific motor pattern (e.g., car: used for transportation; house: used for shelter, shoes: used to protect feet; wedding ring: used to indicate a particular social status; etc.) These examples illustrate that “function” cannot be reduced to a specific sensorimotor system NEURAL FOUNDATIONS FOR UNDERSTANDING SOCIAL 433 spontaneously by all 12 subjects During scanning, subjects' response choices were highly accurate: (>97 % for both social and mechanical vignettes.) Imaging data Direct comparison of the social and mechanical vignettes revealed different and highly selective patterns of activity in a number of regions (Table 1) These differences were confirmed by the mixed-effects ANOVA of the time-series data, which also revealed differences between other conditions In ventral temporal cortex, vignettes interpreted as social elicited greater bilateral activity in the lateral portion of the fusiform gyrus, relative to mechanical vignettes, F=6.37, p.10 (Figure 2B) In contrast, vignettes interpreted as mechanical showed greater bilateral activity in the medial portion of the fusiform gyrus, relative to social animations, F=19.58; p< 001 (Figure 2A) Here, activity associated with interpreting social animations was no greater than when viewing the random motion control condition composed of the same elements, F Social L fusiform gyrus (medial) R fusiform gyrus (medial) L middle temporal gyrus Social = Mechanical Meaningful > Random L polar frontal cortex R polar frontal cortex R anterior temporal cortex Z value X Y Z 3.95 3.57 4.49 3.37 >7.0 >7.0 >7.0 5.14 5.14 3.76 5.12 –44 –40 41 40 56 –49 53 –56 19 –57 –38 –52 –33 –58 –57 –20 –21 –2 52 43 –23 –20 –15 –21 19 17 –4 –10 –11 –13 4.22 6.95 3.05 3.16 –25 27 –49 –34 –44 –57 –56 –58 –18 –16 –9 –7 >7.0 >7.0 >7.0 –28 35 42 40 50 –10 37 27 26 434 THE ORGANISATION OF CONCEPTUAL KNOWLEDGE IN THE BRAIN Vignettes interpreted as social or mechanical also produced differential activity along the lateral surface of posterior temporal cortex (Table 1) Relative to the mechanical vignettes, activity associated with interpreting the social vignettes was centred on the posterior region of STS, bilaterally: F=113.49; p