COGNITIVE NEUROSCIENCE PSYC 6252 3.0 (F)

Fall 2008 Wed. 11:30-2:30

Ross Building, S501 Instructor: R. Shayna Rosenbaum E-mail: shaynar@yorku.caTel: 416-736-2100 x20449 Website: http://silver.yorku.ca/2008f-gspsyc6252a-03 PURPOSE This course examines a variety of higher cognitive functions, such as memory, attention, imagery, language, spatial cognition, and executive function. Evidence for the involvement of specific brain areas in these functions from lesion and neuroimaging studies is addressed. DESCRIPTION How does the brain enable such abilities as recognizing friends, remembering how to navigate home, and fleeing from fearful situations? To gain a better understanding of these and other complex brain-behaviour relations, the focus of investigations need not be limited to the neuronal level, and may also include the study of larger systems and networks that involve the concerted effort of many millions of neurons. This course surveys issues concerning the localization and representation of higher-order cognitive functions in the healthy brain, and examines experimental and clinical studies illustrating behavioural effects of brain damage. Reference is also made to research involving the use of animal models and the growing use of brain-imaging techniques to study the neural basis of various cognitive processes. Students have the opportunity to discuss and critique current research in cognitive neuroscience within an interactive seminar format. The majority of the course focuses on the following topics: face and object recognition, attention, episodic and semantic memory, memory distortions, language, executive function, emotion, social cognition, and consciousness. The course begins with a brief overview of basic neuroanatomy, followed by discussion of lesion and neuroimaging techniques. This is followed by a comparison of object and face recognition, which serves as an introduction to the overarching theme of modular vs. distributed organization of brain function. We then consider whether attention is a single process or a collection of processes, how these process(es) are informed by lesion studies, and whether neuroimaging can tell us about how attention operates or only about what areas are involved. Current theories and data on memory are then presented, with a focus on the processes and systems involved at encoding, storage, and retrieval, as well as the errors of memory and the importance of memory in our everyday lives. Semantic memory is also considered within the framework of theories on language processing, with an emphasis on category-specific organization, as well as models of reading. The last few classes are devoted to examination of the many different functions in which the frontal lobes are involved, ranging from working memory to social cognition, and what these varied functions might have in

common. In particular, we address the nature of interactions between emotion and cognition, taking into account both developmental and evolutionary perspectives. BACKGROUND Students should be familiar with basic principles of brain function to appreciate fully the nature of neural models of complex cognitive and behavioural processes. COURSE FORMAT Students will have the opportunity to discuss and critique current research in cognitive neuroscience within an interactive seminar format. Each student will be required to submit a “thought” paper at the beginning of each of 6 classes of the student’s choice, in which they evaluate readings from that class and generate ideas for theory or experiments. Together with the instructor, students will serve as seminar leaders for a topic of their choice from the syllabus. The student will introduce the week’s topic by providing a clear, concise presentation of one of the assigned articles, including the critical questions asked, the methods, the findings, and the main conclusion. This will include extracting the important issues from the readings, discussing alternative interpretations of the findings, and proposing discussion questions for class. Seminars will also involve supplemental material, such as articles, patient videos, and case studies, designed to promote discussion of relevant topics and to challenge students to apply knowledge acquired in class to real-life clinical and research situations. Students will be encouraged to participate in discussions about theoretical and experimental issues raised in the presentations, readings, and thought papers. At the end of the term, students will be required to submit a more lengthy paper on the topic selected for the class presentation that incorporates a creative or applied component. READINGS There is no assigned text. Readings for each topic consist of empirical articles and/or book chapters selected by the instructor and are available for download from the course website. Supplementary readings are provided for background and to help students with their class presentations and written assignments. The readings are intended to acquaint students with current issues and debate in the field of cognitive neuroscience and serve to elaborate on topics discussed in class. Students are expected to read the required readings for each topic prior to class and are also encouraged to sample background and supplementary readings (see below). A list of recommended textbooks and online resources in Cognitive Neuroscience and Neuroimaging is also provided. The major journals relevant to the course include: Brain Cerebral Cortex Cognitive Neuropsychology Current Opinion in Neurobiology Hippocampus

Journal of Cognitive Neuroscience Journal of Neuroscience Nature Nature Neuroscience Nature Reviews Neuroscience Neuroimage Neuropsychologia Neuropsychology Neuron Philosophical Transactions of the Royal Society of London B Proceedings of the National Academy of Sciences Science Trends in Cognitive Sciences BASIS OF EVALUATION In addition to building knowledge of Cognitive and Behavioural Neuroscience, this course facilitates the development of skills that will allow students to become critical thinkers in this area of research. Accordingly, students will serve as seminar leaders for a topic of their choice from the syllabus and are encouraged to participate in discussions about theoretical and experimental issues raised in the presentations, readings, and thought papers (described below). Lectures will involve supplemental material, such as articles, patient videos, and case studies, designed to promote discussion of relevant topics and to challenge students to apply knowledge acquired in class to real-life clinical and research situations. 1. Weekly Thought Papers (8/9): 40% Each student is required to submit a one page, double-spaced, 12-point (250-300 words) “thought” paper on the readings at the beginning of each of 8 classes of the student’s choice. The purpose of the thought paper is to present the student’s view of the readings in at least one of the following ways: describe the interesting or main questions and how well the student believes they were addressed by the reading(s); evaluate the experimental design and/or the authors’ interpretation of the findings; generate ideas for theory or experiments that the paper(s) inspired; discuss how the papers complemented or contradicted each other. Importantly, the thought paper is NOT meant to summarize the readings but rather to serve as a stimulus for class discussion. Students may submit a total of 9 papers, and the best 8 will be counted towards the final grade (each paper will be graded on a 5-point scale). Please note that thought papers are not to be on the same topic area as the student’s presentations. 2. Class Presentation: 30% Each student will be responsible for two clear, concise (no longer than 20 min.) presentation of one of the assigned articles, including the critical questions asked, the methods, the findings, and the main conclusion. The student should extract the important issues from the readings, discuss alternative interpretations of the findings, and propose 2 discussion questions for class. The questions can be points of confusion, issues for further consideration, follow-up research ideas,

and so on (see sample discussion questions provided with the reading list). To do this well, the student may need to incorporate an additional article or two related to the topic. 3. Class Participation: 30% Students are required to participate in class discussions by presenting the opinions, comments, or views they expressed in their thought papers and by offering answers to questions posed by others.

SCHEDULE OF TOPICS

Date Topic Readings Presenter

Sept. 10 Course Overview, Review of Functional Neuroanatomy

No readings Shayna

Sept. 17 History and Methods in Cognitive Neuroscience Research

1. Rosenbaum et al. (2005); Holdstock et al. (2008)

2. Shallice (2003); Rorden & Karnath (2004)

3. Friston et al. (2006); Saxe et al. (2006)

1. 2. Steven 3.

Sept. 24 Object Recognition and Imagery 1. Kanwisher et al. (1997); Bukach et al. (2006)

2. Haxby et al. (2001); Grill-Spector et al. (2006)

3. Behrmann et al. (1992); O’Craven et al. (2000)

Oct. 1 No Class

Oct. 8 Language and Lexical-Semantic Organization

Overview: Rohrer et al. (2008) 1. Gil-da-Costa et al. (2007) 2. Martin (2007) 3. Caramazza & Mahon

(2003); Tyler & Moss (2001)

Oct. 15 Implicit Memory and Encoding/Perception

Overview: Schacter (1987) 1. Wig et al. (2005) 2. Ryan et al. (2000); Lee et

al. (2005) 3. Wagner et al. (1998);

Shott et al. (2006)

Oct. 22 Memory Retrieval and Distortion Overview: Buckner & Wheeler (2001) 1. Eichenbaum et al. (2007);

Bowles et al. (2007) 2. Schnider (2003); Gilboa et

al. (2006)

Oct. 29 Remote Memory Overview: Squire (1992) 1. Moscovitch et al. (2006);

Vargha-Khadem et al. (1997); Westmacott et al. (2004)

2. Tse et al. (2007); Ji et al. (2007)

Nov. 5 Future Imagining Overview: Wheeler et al. (1997) 1. Addis et al. (2007);

Hassabis et al. (2007); Buckner & Carroll (2007)

2. Tulving (2005); Raby et al. (2007)

Nov. 12 Spatial Cognition Overview: Aguirre & D’Esposito (1999) 1. Epstein (2008) 2. Rosenbaum et al. (2000);

Maguire et al. (2006); Shrager et al. (2008)

Nov. 19 Attention Overview: Posner & Petersen (1990) 1. Hilgetag et al. (2001);

Danckert & Ferber (2006) 2. O’Craven & Kanwisher

(1999); Shomstein & Behrmann (2006)

Nov. 26 Working Memory and Cognitive Control

Overview: Stuss & Levine (2002) 1. Baddeley et al. (2003);

Postle (2006) 2. Sayala et al. (2006);

Petrides (2005) 3. Badre (2008); Koechlin &

Hyafil (2007)

Dec. 3 Emotion and Social Cognition Overview: Phelps & LeDoux (2006) 1. Adolphs (2008); LaBar &

Cabeza (2006) 2. Saxe et al. (2004);

Mitchell (2008); Shamay-Tsoory et al. (2007)

*see below for detailed references

RECOMMENDED TEXTBOOKS (OPTIONAL) Cognitive Neuroscience and Neuropsychology Ward, J. (2006). The Student’s Guide to Cognitive Neuroscience, 1st Ed., Psychology Press. Gazzaniga, M.S. (2004). The Cognitive Neurosciences III. Cambridge, MA: MIT Press. Feinberg, T. E. and Farah, M. J., Editors (2003). Behavioral Neurology and Neuropsychology, 2nd Edition. New York: McGraw-Hill. Banich, M.T. (2004). Cognitive Neuroscience and Neuropsychology, 2nd edition. Boston: Houghton-Mifflin, Co. Neuroimaging Huettel SA, Song AW, McCarthy G (2004). Functional Magnetic Resonance Imaging. Sunderland, MA: Sinauer. R. Cabeza & A. Kingstone (Eds.), Handbook of functional neuroimaging of cognition (pp.331-377). Cambridge, MA: MIT Press, 2001. Luck, SJ (2005). An Introduction to the Event-related Potential Technique. Cambridge, MA: MIT Press. General Guides to Writing and Presentation Burchfield, R.W. (2004). Fowler’s Modern English Usage. Kosslyn, S.M. (2006). Graph Design for the Eye and Mind. New York, NY: Oxford University Press. Kosslyn, S.M. (2007). Clear and to the Point: 8 Psychological Principles for Compelling PowerPoint Presentations. New York, NY: Oxford University Press. ONLINE RESOURCES The Brain from Top to Bottom: http://thebrain.mcgill.ca/flash/index_d.html (Author: Canadian Institute of Health Research): Material is presented for three type of learners (beginner, intermediate, advance) and topics are organized in five levels (social, psychological, neurological, cellular, molecular). This is an excellent website. Dana Foundation Brain Web: http://www.dana.org/default.aspx (Author: Dana Foundation): provides information and links to validated sites about brain diseases and disorders, as well as webcasts & podcasts. *See below for resources on specific topics.

READING LIST FUNCTIONAL NEUROANATOMY Online Resources University of Washington Brain Atlas: http://www9.biostr.washington.edu/da.html: An excellent resource for anyone trying to learn human neuroanatomy. Harvard Medical School Brain Atlas: www.med.harvard.edu/AANLIB/home.html – visit “Top 100 Brain Structures” under “Normal Brain” University of Toronto Interactive Functional Neuroanatomy website (Author: P. Stewart): http://video.med.utoronto.ca/neuronotes/ HISTORY AND METHODS Case Study Approach Rosenbaum, R.S., Köhler, S., Schacter, D. L., Moscovitch, M., Westmacott, R., Black, S.E., Gao, F., & Tulving, E. (2005). The case of K.C.: Contributions of a memory-impaired person to memory theory. Neuropsychologia, 43, 989-1021. • This paper provides a framework for dealing with patients who are interesting from a

psychological perspective but complicated from a neurological one due to widespread brain damage. It also provides an overview that will help guide the other readings on memory. Read the Introduction and General Discussion and skim the rest.

Holdstock JS, Parslow DM, Morris RG, Fleminger S, Abrahams S, Denby C, Montaldi D, Mayes AR. (2008). Two case studies illustrating how relatively selective hippocampal lesions in humans can have quite different effects on memory. Hippocampus, 18, 679-91. • This article is important because it points out the difficulties in evaluating patients with

presumably selective hippocampal damage, who can exhibit highly variable memory impairments. The authors argue that more sophisticated imaging and volumetric analyses, together with functional imaging studies, will be needed to better evaluate brain-damaged patients with amnesia.

Patient vs. Neuroimaging Studies Rorden, C. & Karnath, H.-O. (2004). Using human brain lesions to infer function: a relic from a past era in the fMRI age? Nature Reviews: Neuroscience, 5, 813-819. Shallice, T. (2003). Functional imaging and neuropsychology findings: how can they be linked? NeuroImage, 20, S146-S154.

• These papers provide different viewpoints on the utility of using neuroimaging and lesion studies to infer cognitive function.

fMRI: Region of Interest (ROI) vs. Whole-Brain Analysis Friston, K.J., Rotsthein, P., Geng, J.J., Sterzer, P., & Henson, R.N.A. (2006). A critique of functional localizers, NeuroImage, 30, 1077-1087 Saxe, R., Brett, M., & Kanwisher, N. (2006). Divide and conquer: a defense of functional localizers. Neuroimage, 30, 1088-1096. • These papers represent a detailed exchange that describes differences of opinion (and

possibly naivety) regarding the utility of region-of-interest analysis of fMRI data (‘functional localizers’), which is an extension of single-unit recording methods, over whole-brain or network analyses.

Supplementary Raichle (1998). Behind the scenes of functional brain imaging: A historical and physiological perspective. Proceedings of the National Academy of Sciences, 95, 765-772. Fellows, L.K., Heberlein, A.S., Morales, D., Shivde, G., Waller, S., & Wu, D.H. (2005). Method matters: An empirical study of impact in Cognitive Neuroscience. Journal of Cognitive Neuroscience, 17, 850-858. Haynes, J-D, & Rees, G. (2006). Decoding mental states from brain activity in humans. Nature Reviews Neuroscience, 7, 523-534. Henson, R. (2005). What can functional neuroimaging tell the experimental psychologist? The Quarterly Journal of Experimental Psychology, 58A, 193-233. McIntosh, A.R. (1999). Mapping cognition to the brain through neural interactions. Memory, 7, 523-548. Hallett, M. (2007). Transcranial magnetic stimulation: A primer. Neuron, 55, 187-199. Stark, C.E.L., & Squire, L.R. (2001). When zero is not zero: The problem of ambiguous baseline conditions in fMRI. Proceedings of the National Academy of Sciences, 98, 12760-12766. Online Resources TMS demo http://www.telegraph.co.uk/earth/main.jhtml?view=DETAILS&grid=&xml=/earth/2008/05/16/scibrain216.xml Virtual EEG

http://brainserver.psych.indiana.edu/ (Author: The Mind Project, Indiana University): This website illustrates ERP and EEG methodology Discussion Questions 1. What are the relative strengths and weaknesses of the case study approach? 2. How do functional neuroimaging and lesion methods complement each other? 3. Can the two methods inform our understanding of basic cognitive processes? OBJECT RECOGNITION AND IMAGERY Object (Face) Recognition Kanwisher, N., McDermott, J., & Chun, M.M. (1997). The fusiform face area: a module in human extrastriate cortex specialized for face perception. Journal of Neuroscience, 17, 4302-4311. Bukach, C.M., Gauthier, I., & Tarr, M.J. (2006). Beyond faces and modularity: the power of an expertise framework. Trends in Cognitive Sciences, 10, 159-166. Grill-Spector, K., Sayres, R., & Ress, D. (2006) High-resolution imaging reveals highly selective nonface clusters in the fusiform face area. Nature Neuroscience, 9, 1177-1185. Haxby, J.V., Gobbini, M.I., Furey, M.L., Ishai, A., Schouten, J.L., & Pietrini, P. (2001). Distributed and overlapping representations of faces and objects in ventral temporal cortex. Science, 293, 2425-2430. • These papers represent an ongoing debate in the face recognition literature and provide

insight into the more general issue of modularity of function vs. distributed processes (see also Haxby et al. in supplementary readings).

Visual Imagery Behrmann, M., Winocur, G., & Moscovitch, M. (1992). Dissociations between mental imagery and object recognition in a brain-damaged patient. Nature, 359, 636-637. O’Craven, K.M., & Kanwisher, N. (2000). Mental imagery of faces and places activates corresponding stimulus-specific brain regions. Journal of Cognitive Neuroscience, 12, 1013-1023. • These papers address whether imagery draws on overlapping neural and functional

mechanisms used during perception. The first paper places this issue within a theoretical context; the other papers are examples of investigations of the dissociability of imagery and perception using different techniques.

Supplementary

Object Recognition Overview: Farah, M.J. (2003). Prosopagnosia. In Feinberg, T.E., & Farah, M.J. (Eds.), Behavioral neurology and neuropsychology, 2nd Ed. (pp. 239-241), McGraw Hill. Moscovitch, M., Winocur, G., & Behrmann, M. (1997). What is special about face recognition? Nineteen experiments on a person with visual object agnosia and dyslexia but normal face recognition. Journal of Cognitive Neuroscience, 9, 555-604. Gauthier, I., Skudlarski, P., Gore, J.C., & Anderson, A.W. (2000). Expertise for cars and birds recruits brain areas involved in face recognition. Nature Neuroscience, 3, 191-197. Visual Imagery Overview: Kosslyn, S. M. (2005). Mental images and the brain. Cognitive Neuropsychology, 22, 333-347. Mechelli, A., Price, C.J., Friston, K.J., & Ishai, A. (2004). Where bottom-up meets top-down: Neuronal interactions during perception and imagery. Cerebral Cortex, 14, 1256-1265. Online Resources Visual Image Database: http://titan.cog.brown.edu:8080/TarrLab/stimuli (Author: Michael Tarr): An excellent source of object images (including faces and Greebles!). Discussion Questions 1. Is the brain organized into ‘modules’? What are the advantages/disadvantages of a modular

vs. distributed system? 2. Are faces a special class of objects? 3. Is it possible to reconcile the domain-specific vs. distributed views of object recognition? 4. Is it possible to account for the contradictory findings reported in Behrmann et al. vs.

O’Craven & Kanwisher ? LANGUAGE AND LEXICAL-SEMANTIC ORGANIZATION Overview: Rohrer, J.D., Knight, W.D., Warren, J.E., Fox, N.C., Rossor, M.N., & Warren, J.D. (2008). Word-finding difficulty: a clinical analysis of the progressive aphasias. Brain, 131, 8-38. • A comprehensive review that compares acute focal with progressive aphasias and provides

insight into the organizational structure of language. The authors include useful figures depicting the levels at which language is disrupted by the various forms of aphasia and the aspects of language that should be considered in diagnosis.

Gil-da-costa, R., Martin, A., Lopez, M. A., Munoz, M., Fritz, J. B. & Braun, A. R. (2006). Species-specific calls activate homologs of Broca’s and Wernicke’s areas in the macaque. Nature Neuroscience, 9, 1064-1070. • This PET study of macaques demonstrates that non-human primates possess brain

mechanisms similar to those implicated in human language to process socially significant vocalizations.

Martin, A. (2007). The representation of object concepts in the brain. Annual Review of Psychology, 58, 25-45. Caramazza, A., & Mahon, B.Z. (2003). The organization of conceptual knowledge: the evidence from category-specific semantic deficits. Trends in Cognitive Sciences, 7, 354-361. • These reviews summarize competing theories of modality-specific and category-specific

representations of object knowledge. Tyler, L.K., & Moss, H. (2001). Towards a distributed account of conceptual knowledge. Trends in Cognitive Sciences, 5, 244-252. • It is suggested here that a semantic structure centred around intercorrelated vs. distinctive

features of concepts might better capture patterns of semantic memory loss in Semantic Dementia and Alzheimer’s disease.

Supplementary Background Alexander, M.P. (2003). Aphasia: Clinical and anatomic issues. In Feinberg, T.E., & Farah, M.J. (Eds.), Behavioral neurology and neuropsychology, 2nd Ed. (pp. 147-164), McGraw Hill. Saffran, E.M. (2003). Aphasia: Cognitive and neuropsychological issues. In Feinberg, T.E., & Farah, M.J. (Eds.), Behavioral neurology and neuropsychology, 2nd Ed. (pp. 165-177), McGraw Hill. Ungerleider LG, Mishkin M (1982) Two cortical visual systems. In: Ingle DJ, Goodale MA, Mansfield RJW (eds): Analysis of Visual Behavior. Cambridge: MIT Press, pp 549-586. Warrington, E. K., & Shallice, T. (1984). Category specific semantic impairments. Brain, 107, 829-853. Alternative Theoretical Frameworks Sharon L. Thompson-Schill, S.L. (2003). Neuroimaging studies of semantic memory: inferring “how” from “where”. Neuropsychologia, 41, 280-292. • This paper discusses the structure of semantic memory based on neuroimaging evidence.

Patterson, K., Nestor, P.J., & Rogers, T.T. (2007). Where do you know what you know? The representation of semantic knowledge in the human brain. Nature Reviews Neuroscience, Noppeney, U., Patterson, K., Tyler, L.K., Moss, H., Stamatakis, E.A., Bright, P., Mummery, C., & Price, C.J. (2007). Temporal lobe lesions and semantic impairment: a comparison of herpes simplex virus encephalitis and semantic dementia. Brain, 130, 1138-1147. • This paper examines the view that semantic memory is structured as a widely distributed

brain network that contains information regarding modality-specific features. The latter suggests, based on neuropsychological and neuroimaging data and connectionist modelling, that conceptual knowledge also requires an amodal hub.

McClelland, J.L., & Rogers, T.T. (2003). The parallel-distributed processing approach to semantic cognition. Nature Reviews Neuroscience, 4, 310-322. • This paper integrates predictions of classic theories of semantic organization within a

parallel distributed processing (PDP) connectionist framework to accommodate patterns of semantic memory acquisition, restructuring, and disintegration that are not readily accounted for by hierarchical feature-based (e.g., Quillian’s spreading activation model) or pure modality-based models.

Online Resources Language and Aphasia: http://psych.rice.edu/mmtbn/ (Author: Psychology Dept, Rice U., supported by NSF): This is multimedia textbook in behavioral neuroscience. The chapter on language includes video examples of patients with aphasia. Discussion questions 1. Is semantic knowledge organized according to category or modality? To what extent is this

organization similar to perceptual and action systems? 2. In what way do neuroimaging studies inform category-specific deficits in patients? 3. Do patterns of semantic loss observed in Semantic Dementia and Alzheimer’s disease fit

within a strict modality-based framework? IMPLICIT MEMORY AND EXPLICIT MEMORY (ENCODING) Overview: Schacter, D.L. (1987). Implicit memory: History and current status. Journal of Experimental Psychology: Learning, Memory, and Cognition, 13, 501-518. • This article provides some historical context and background for experimental and lesion

studies of implicit memory. Read pages 1001-1005 of Rosenbaum et al. (2005) for specific investigations of implicit memory in the case K.C. and pages 1005-1007 for studies of memory acquisition in K.C.

Wig, G.S., Grafton, S,T., Demos, K. E. & Kelley, W.M. (2005). Reductions in neural activity underlie components of repetition priming. Nature Neuroscience, 8, 1228-1233. (commentary: Martin, A., & Gotts, S.J. (2005). Making the causal link: Frontal cortex activity and repetition priming. Nature Neuroscience, 8, 1134-1135.) • This paper describes the neural basis of implicit memory (i.e., deactivation of a relevant

brain region rather than the increased activation that characterizes explicit memory). Ryan, J.D, Althoff, R.R., and Cohen, N.J. (2000). Amnesia is a deficit in relational processing. Psychological Science, 11, 454-461. • This paper presents findings on implicit memory in amnesia within a novel theoretical

framework that views the hippocampus as having a primary role in relational memory. See Davachi & Wagner (2002) and Eichenbaum (1999) in supplementary readings for related views.

Lee, A.C., Bussey, T.J., Murray, E.A., Saksida, L.M., Epstein, R.A., Kapur, N., Hodges, J.R., & Graham, K.S. (2005). Perceptual deficits in amnesia: challenging the medial temporal lobe ‘mnemonic’ view. Neuropsychologia, 43, 1-11. • Findings reported in this paper suggest that the role of the hippocampus may extend beyond

explicit memory and include perceptual discrimination. Wagner, A. D., Schacter, D. L., Rotte, M., Koutstaal, W., Maril, A., Dale, A. M., Rosen, B. R., & Buckner, R. L. (1998). Building memories: remembering and forgetting of verbal experiences as predicted by brain activity. Science, 281, 1188-1191. (with commentary by Rugg, M.D. (1998). Memories are made of this. Science, 281, 1151-1152.) • This paper was one of the first to use neuroimaging as a technique to predict items that are

remembered vs. those that are forgotten, a technique that has since been used in over 40 studies. Another study along the same lines was published in the same issue of Science by Brewer et al.

Shott, B.H., Richardson-Klavehn, A., Henson, R.N., Becker, C., Heinze, H.J., & Düzel, E. (2006). Neuroanatomical dissociation of encoding processes related to priming and explicit memory. Journal of Neuroscience, 26, 792-800. • This study provides evidence on the dissociability of regions involved in explicit versus

implicit memory processes by capitalizing on the paradigm made popular by Wagner et al. Supplementary Grill-Spector, K., Henson, R.N.A., & Martin, A. (2006). Repetition and the brain: neural models of stimulus-specific effects. Trends in Cognitive Sciences, 10, 14-23.

• This review provides a detailed analysis of three different mechanistic models that have been proposed to account for repetition-related reductions in neural activity. The models are evaluated in terms of their ability to account for data from neurophysiological and neuroimaging studies.

Davachi, L. (2006). Item, context, and relational episodic encoding in humans. Current Opinion in Neurobiology, 16, 693-700. Eichenbaum, H. (2004). Hippocampus: Cognitive processes and neural representations that underlie declarative memory. Neuron, 44, 109-120. • These theoretical reviews build on the notion that the hippocampus is primarily involved in

processing relations among memories, whereas regions of parahippocampal gyrus are involved in maintaining individual memories.

Shrager, Y., Gold, J.J., Hopkins, R.O., & Squire, L.R. (2006). Intact visual perception in memory-impaired patients with medial temporal lobe lesions. Journal of Neuroscience, 26, 2235-2240. Goshen-Gottstein, Y., & Moscovitch, M. (1995). Repetition priming for newly formed and preexisting associations: Perceptual and conceptual influences. Journal of Experimental Psychology: Learning, Memory and Cognition, 21, 1229-1248. Verfaellie, M., Martin, E., Page, K., & Keane, M.M. (2006). Implicit memory for novel conceptual associations in amnesia. Cognitive and Affective Behavioral Neuroscience, 6, 91-101. • These two studies attempt to reconcile debate on the integrity of priming of novel

associations in amnesia. The latter study relates findings to relational memory theory as described in Ryan et al. (2000) and Eichenbaum (2004).

Brewer, J.B., Zhao, Z., Desmond, J.E., Glover, G.H., & Gabrieli, J.D. (1998). Making memories: brain activity that predicts how well visual experience will be remembered. Science, 281, 1185-1187. • Companion to Wagner et al. paper. Turk-Browne, N.B., Yi, D.J., & Chun, M.M. (2006). Linking implicit and explicit memory: common encoding factors and shared representations. Neuron, 49, 917-927. • Findings are opposite to that reported in Shott et al. (2006). Discussion questions 1. How has the conceptualization of implicit memory changed since Schacter’s 1987 review?

Is this due to methodological advances? 2. How might the finding of repetition suppression during priming be applied to areas of

cognition beyond implicit memory?

3. What are the behavioral and neural differences between explicit and implicit memory? How might neuroimaging distinguish between the two? How could neuroimaging be used to determine explicit contamination during priming?

4. Do findings of ‘implicit memory’ and perceptual deficits in amnesia contradict the view that the hippocampus plays a primary role in explicit memory acquisition? How can these two views be reconciled?

5. How is encoding represented in the brain? What do neuroimaging studies tell us about the cognitive processes needed to remember information well?

MEMORY RETRIEVAL AND RECONSTRUCTION Memory Retrieval Overview: Buckner, R.L., & Wheeler, M.E. (2001). The cognitive neuroscience of remembering. Nature Reviews: Neuroscience, 2, 624-634. • This review summarizes the neural basis of different aspects of retrieval and is particularly

informative in its description of prefrontal contributions. Eichenbaum, H., et al. (2007). The medial temporal lobe and recognition memory. Annual Review in Neuroscience, 30, 123-152. • This review deals with a major controversy in the cognitive neuroscience of memory relating

to whether a single mechanism supports recognition memory or whether there are distinct processes for recollection and familiarity, both of which contribute to recognition. It provides a comprehensive summary of a host of recent animal and human studies that have distinguished between brain regions involved in recollection vs. familiarity.

Bowles B, Crupi C, Mirsattari SM, Pigott SE, Parrent AG, Pruessner JC, Yonelinas AP, Köhler S. (2007). Impaired familiarity with preserved recollection after anterior temporal-lobe resection that spares the hippocampus. Proceedings of the National Academy of Sciences, 104, 16382-16387. • This study provides new and unique evidence based on a rare case of selective damage to

perirhinal cortex that supports the view that recollection and familiarity have distinct representations in the medial temporal lobe. Relate to theories proposed by Eichenbaum (2004) and Davachi (2006) in last week’s Encoding section.

Memory Reconstruction Schnider, A. (2003). Spontaneous confabulations and the adaptation of thought to ongoing reality. Nature Reviews: Neuroscience, 4, 662-671. Gilboa, A., Alain, C., Stuss, D.T. Melo, B., Miller, S., & Moscovitch, M. (2006). Mechanisms of spontaneous confabulation: A strategic retrieval account. Brain, 129, 1399-1414.

• These papers describe experiments designed to test different theories of confabulation. Supplementary Rugg, M.D. (2004). Retrieval Processes in Human Memory: Electrophysiological and fMRI Evidence. In Gazzaniga, M.S. (Ed), The Cognitive Neurosciences III, pp. 727-739. Cambridge, MA: MIT Press. Aggleton, J.P., & Brown, M.W. (1999). Episodic memory, amnesia and the hippocampal-anterior thalamic axis. Behavioral and Brain Sciences, 22, 425-489. • An influential article that reviews evidence that the hippocampal-anterior thalamic system

underlies episodic memory and argues that recognition memory depends on two processes supported by distinct structures.

Sauvage MM, Fortin NJ, Owens CB, Yonelinas AP, Eichenbaum H. (2008). Recognition memory: opposite effects of hippocampal damage on recollection and familiarity. Nature Neuroscience, 11, 16-18. • This study investigates performance of rats with hippocampal lesions on a novel odor-

association task in rats to demonstrate that, as in humans, recollection and familiarity are separable.

Gilboa, A., Winocur, G., Rosenbaum, R.S., Poreh, A., Gao, F., Black, S.E., Westmacott, R., & Moscovitch, M. (2006). Hippocampal contributions to recollection in retrograde and anterograde amnesia. Hippocampus, 16, 966-980. • This study demonstrates that the dissociation between recollection and familiarity extends to

remote (retrograde) memory. Schacter, D.L. (1999). The seven sins of memory. American Psychologist, 54, 182-203. • This article presents a theory to account for the occurrence of memory distortions. Brainerd, C.J. and Reyna, V.F. (2002) Fuzzy trace theory and false memory. Current Directions in Psychological Science, 11, 164-168. • This paper presents alternative accounts of memory distortions/reconstruction to the ones

proposed by Schacter. Discussion questions 1. What roles do the medial temporal lobe and prefrontal cortex play during memory retrieval?

How might they interact? 2. How do recollection and familiarity differ at a behavioural level? What are the neural

correlates of this distinction? 3. How do the factors that promote accurate memory also result in memory distortion?

4. What is the relationship between memory distortion/reconstruction and confabulation? REMOTE MEMORY Overview: Squire, L. R. (1992). Memory and the hippocampus: A synthesis from findings with rats, monkeys and humans. Psychological Review, 99, 195-231. Moscovitch, M., Nadel, L., Winocur, G., Gilboa, A., & Rosenbaum, R. S. (2006). The cognitive neuroscience of remote memory: A focus on functional neuroimaging. Current Opinion in Neurobiology, 16, 179-190. • These articles focus on a current controversy regarding the representation of episodic and

semantic memory in the brain and the theories that may account for differences or similarities between them.

Vargha-Khadem, F., Gadian, D. G., Watkins, K. E., Connelly, A., Van Paesschen, W., & Mishkin, M. (1997). Differential effects of early hippocampal pathology on episodic and semantic memory. Science, 277, 376-380. Westmacott R, Black SE, Freedman M, Moscovitch M. (2004). The contribution of autobiographical significance to semantic memory: evidence from Alzheimer's disease, semantic dementia, and amnesia. Neuropsychologia, 42, 25-48. • These papers have implications for the relation between episodic and semantic memory

based on studies of patients with developmental amnesia and semantic dementia. Tse, D., Langston, R.F., Kakeyama, M., Bethus, I., Spooner, P.A., Wood, E.R., Witter, M.P., & Morris, R.G. (2007). Schemas and memory consolidation. Science, 316, 76-82. Ji, D., & Wilson, M.A. (2007). Coordinated memory replay in the visual cortex and hippocampus during sleep. Nature Neuroscience, 10, 100-107. • The first study reports that the duration of memory consolidation depends on the extent to

which new memories can be incorporated into an existing schema by hippocampal processing. The second study uses repeated simultaneous multi-unit recordings in rats to demonstrate how neocortex and hippocampus may interact during sleep in order to consolidate memories.

Supplementary Tulving (1972). Episodic and semantic memory. In E. Tulving & W. Donaldson (Eds.), Organization of Memory (pp. 381-403). New York: Academic Press. Reed, J.M., & Squire, L.R. (1998). Retrograde amnesia for facts and events: findings from four new cases. Journal of Neuroscience, 18, 3943-3954.

Squire, L. R., & Bayley, P. J. (2007). The neuroscience of remote memory. Current Opinion in Neurobiology, 17, 185-196. Nadel, L., & Moscovitch, M. (1997). Memory consolidation, retrograde amnesia and the hippocampal complex. Current Opinion in Neurobiology, 7, 217–227. Rosenbaum, R.S., Moscovitch, M., Foster, J.K., Schnyer, D.M., Gao, F.Q., Kovacevic, N., Verfaellie, M., Black, S.E., & Levine, B. (2008). Patterns of autobiographical memory loss in medial temporal lobe amnesic patients. Journal of Cognitive Neuroscience, 20, 1490-1506. Gardiner, J.M., Brandt, K.R., Baddeley, A.D., Vargha-Khadem, F., & Mishkin, M. (2008). Charting the acquisition of semantic knowledge in a case of developmental amnesia. Neuropsychologia, 46, 2865-2868. Discussion questions 1. What are the arguments for and against of traditional consolidation theory? 2. How are episodic and semantic memory related? Are they dependent on each other? How

do they interact? 3. What happens in the brain to determine whether we remember something for a long time? 4. What neural events during sleep impact long-term memory consolidation? FUTURE IMAGINING Overview: Wheeler, M.A., Stuss, D.T., & Tulving, E. (1997). Toward a theory of episodic memory: the frontal lobes and autonoetic consciousness. Psychol Bulletin, 121, 331-54. • This paper extends the ideas described in Tulving (1985) regarding the different levels of

self-awareness and gives the highest level a neural correlate. Addis, D.R., Wong, A.T., & Schacter, D.L. (2007). Remembering the past and imagining the future: Common and distinct neural substrates during event construction and elaboration. Neuropsychologia, 45,1363-1377. Hassabis D, Kumaran D, Vann SD, Maguire EA. (2007). Patients with hippocampal amnesia cannot imagine new experiences. Proceedings of the National Academy of Sciences, 104, 1726-1731. Buckner, R.L., & Carroll, D.C. (2007). Self-projection and the brain. Trends Cognitive Sciences, 11, 49-57. • These papers represent renewed interest in the relationship between memory for past

experiences and the generation of imagined and future events (and related non-mnemonic abilities). Note the differences in the explanations provided for this correspondence.

Tulving, E. (2005). Episodic memory and autonoesis: Uniquely human? In: The missing link in cognition: Origins of self-reflective consciousness, In Terrace, H.S., & Metcalfe, J. (Eds.), pp. 3–56. Oxford University Press. Raby, C.R., Alexis, D.M., Dickinson, A., & Clayton, N.S. (2007). Planning for the future by western scrub-jays. Nature, 445, 919-921. • Here, the ongoing debate of whether non-human animals are capable of episodic re-

experiencing and future “pre-experiencing” is addressed. The second paper presents a clever investigation of the ability to plan ahead in scrub jays.

Supplementary Conway, M.A. (2001). Sensory-perceptual episodic memory and its context: autobiographical memory. Philosophical Transactions of the Royal Society of London, 356, 1375-1384. • This paper describes a model of autobiographical memory and its components. Tulving, E. (1985). Memory and consciousness. Canadian Psychology, 26, 1-12. Atance, C.M., & O’Neill, D.K. (2001). Episodic future thinking. Trends in Cognitive Sciences, 5, 533-539. Clayton, N. S., Bussey, T. J. & Dickinson, A. (2003) Can animals recall the past and plan for the future? Nature Reviews Neuroscience, 4, 685-691. Suddendorf, T., & Corballis, M.C. (2007). The evolution of foresight: What is mental time travel, and is it unique to humans? Behavioural and Brain Sciences, 30, 299-313; discussion 313-51. Spreng, R.N., Mar, R.A., & Kim, A.S. (2008). The Common Neural Basis of Autobiographical Memory, Prospection, Navigation, Theory of Mind and the Default Mode: A Quantitative Meta-analysis. Journal of Cognitive Neuroscience, in press. Discussion questions 1. What are the advantages of having a flexible episodic memory system? 2. Which of the theories accounts best for the overlap in brain regions supporting such disparate

abilities as future imagining, theory of mind, perception, and discourse? 3. Is mental time travel uniquely human? SPATIAL COGNITION Theoretical Framework and Neocortical Component

Overview: Aguirre, G.K., and D’Esposito, M. (1999). Topographical disorientation: a synthesis and taxonomy. Brain, 122, 1613-1628. Epstein, R.A. (2008). Parahippocampal and retrosplenial contributions to human spatial navigation. Trends in Cognitive Sciences, in press. • The first paper provides a taxonomy of brain regions/processes believed to support different

aspects of real-world navigation. The second paper focuses on two aspects of this framework.

Hippocampus Rosenbaum, S.R., Priselac, S., Kohler, S., Black, S.E. Gao, F., Nadel, L., & Moscovitch, M. (2000). Studies of remote spatial memory in an amnesic person with extensive bilateral hippocampal lesions. Nature Neuroscience, 3, 1044-1048. Maguire EA, Nannery R, Spiers HJ. (2006). Navigation around London by a taxi driver with bilateral hippocampal lesions. Brain, 129, 2894-2907. • These patient studies speak to debate regarding the widely held view that the hippocampus

plays an essential role in supporting allocentric spatial representations, whether formed recently or long ago.

Shrager, Y., Kirwan, C.B., & Squire, L.R. (2008). Neural basis of the cognitive map: path integration does not require hippocampus or entorhinal cortex. Proceedings of the National Academy of Sciences, 105, 12034-12038. • The results presented here may be considered controversial in challenging the view that the

hippocampus is needed for spatial computations involved in simple path integration (contrast with Maaswinkel et al. paper listed below).

Supplementary Morris, R. G. M., Garrud, P., Rawlins, J. N. P., & O'Keefe, J. (1982). Place navigation impaired in rats with hippocampal lesions. Nature, 297, 681-683. • A seminal animal lesion study that provided strong evidence in favour of Cognitive Map

Theory, which states that the primary function of the hippocampus is in learning allocentric spatial relationships within the environment.

Burgess, N. (2008). Spatial cognition and the brain. Annals of the New York Academy of Sciences, 1124, 77-97. • This paper describes a theory of the neural network associated with spatial perception,

imagery, and navigation as informed by neurophysiology and neuroimaging, derived from ‘Cognitive Map’ and ‘Scene Construction’ theories.

Shapiro, M.L., & Eichenbaum, H. (1999). Hippocampus as a memory map: synaptic plasticity and memory encoding by hippocampal neurons. Hippocampus, 9, 365-384. • Discusses the idea that ‘place cells’ form associations between various stimuli, whether

spatial or non-spatial (i.e., in rodents, these associations happen to be mainly spatial, but they are not specialized for coding space per se).

Maguire, E.A., & Spiers, H.J. (2007). The neuroscience of remote spatial memory: a tale of two cities. Neuroscience, 149, 7-27. • A recent survey of the growing literature on the cognitive structure and neural correlates of

remote spatial memory, addressing debate about hippocampal involvement in supporting memories of environments learned long ago.

Maaswinkel, H., Jarrard, L.E., & Whishaw, I.Q. (1999). Hippocampectomized rats are impaired in homing by path integration. Hippocampus, 9, 553-561. • A study showing the dependence of path integration on hippocampus in rats with selective

lesions. Bohbot, V.D., Lerch, J., Thorndycraft, B., Iaria, G., & Zijdenbos, A.P. (2007). Gray matter differences correlate with spontaneous strategies in a human virtual navigation task. Journal of Neuroscience, 27, 10078-10083. • A volumetric study demonstrating the neural basis of individual differences in navigational

strategy (spatial vs. stimulus-response). Ino, T., Doi, T., Hirose, S., Kimura, T., Ito, J., & Fukuyama, H. (2007). Directional disorientation following left retrosplenial hemorrhage: a case report with fMRI studies. Cortex, 43, 248-254. • A combined patient-fMRI study demonstrating evidence of dissociations in retrosplenial

involvement in heading direction and mechanisms underlying recovery of function. Wilson, B.A., Berry, E., Gracey, F., Harrison, C., Stow, I., Macniven, J., Weatherley, J., & Young, A.W. (2005). Egocentric disorientation following bilateral parietal lobe damage. Cortex, 41, 547-554. • A paper discussing the dependence of egocentric spatial representations on posterior

parietal cortex. Keep this in mind for next week’s class on attention. Discussion Questions: 1. What are the components of spatial navigation according to Aguirre & D’Esposito and

Burgess? How are some of these components integrated in the brain? 2. To what extent do these models stand the test of experimental investigation?

3. Does Cognitive Map Theory provide a viable explanation of hippocampal function? 4. What might distinguish adept from poor navigators? ATTENTION AND NEGLECT Overview: Posner, M., & Petersen, S.E. (1990). The attention system of the human brain. Annual Review of Neuroscience, 13, 25-42. • Classic paper proposing a posterior-to-anterior organization of attention processes, and the

brain region(s) and/or networks underlying various component processes, grounded in evidence from cognitive paradigms, neurophysiology, and patients.

Neglect Hilgetag, C.C., Théoret, H., Pascual-Leone, A. (2001). Enhanced visual spatial attention ipsilateral to rTMS-induced 'virtual lesions' of human parietal cortex. Nature Neuroscience, 4, 953-957. • A fascinating demonstration in support of the ‘hemispheric rivalry account’ of spatial

attention/neglect through the application of rTMS to a region of parietal cortex opposite to the side of a ‘virtual lesion.’

Danckert, J., & Ferber, S. (2006) Revisiting unilateral neglect. Neuropsychologia, 44, 987-1006. • This paper offers a refreshing perspective on unilateral neglect and its neural basis. Note

how it attempts to account for the Hilgetag et al. findings. Selective Attention O’Craven, K.M., Downing, P.E., & Kanwisher, N. (1999). fMRI evidence for objects as the units of attentional selection. Nature, 401, 584-587. Shomstein, S., & Behrmann, M. (2006). Cortical systems mediating visual attention to both objects and spatial locations. Proceedings of the National Academy of Sciences, 103, 11387-11392. • Clever fMRI studies testing object- vs. space-based attention. Supplementary Driver, J. (2001). A selective review of selective attention research from the past century. British Journal of Psychology, 92, 53-78. Kastner, S., & Ungerleider, L.G. (2000). Mechanisms of visual attention in the human cortex. Annual Review of Neuroscience, 23, 315-341.

Corbetta, M. & Schulman, G. L. (2002). Control of goal-directed and stimulus-driven attention in the brain. Nature Reviews Neuroscience, 3, 201-215. • Excellent reviews of top-down and bottom-up influences on attention and their interactions. Mesulam, M-M. (1981). A cortical network for directed attention and unilateral neglect. Annals of Neurology, 10, 309-325. • A classic paper that describes mechanisms of neglect based on neurological patients. Nico D, Piccardi L, Iaria G, Bianchini F, Zompanti L, & Guariglia C. (2008). Landmark based navigation in brain-damaged patients with neglect. Neuropsychologia, 46, 1898-1907. • This article demonstrates an interesting link between representational vs. perceptual neglect

and navigation in a water-maze-like task and inform models of parietal-hippocampal interactions.

Rossi, A.F., Bichot, N.P., Desimone, R., & Ungerleider, L.G. (2007). Top down attentional deficits in macaques with lesions of lateral prefrontal cortex. Journal of Neuroscience, 27, 11306-11314. • This study clearly demonstrates the necessity of the prefrontal cortex in achieving rapid

switches of selective attention based on an instructional cue and separates this ability from general selective attention. The results have implications for frontal lobe patients who exhibit complex behavioral deficits.

Discussion questions 1. How can neglect inform our understanding of attention? Is this disorder restricted to

attention? Does neglect permit an understanding of all aspects of attention? 2. Is attention a multi-component set of processes? 3. Is selective attention object-based, space-based, or feature-based? How does it relate to

perceptual organization? WORKING MEMORY AND COGNITIVE CONTROL Overview: Stuss, D.T., & Levine, B. (2002). Adult clinical neuropsychology: Lessons from Studies of the Frontal Lobes." Annual Review of Psychology, 53, 401-433. • This article provides an overview of dissociation of function within the frontal lobes based on

lesion studies in humans. This model has since been updated but continues to provide a useful taxonomy.

Baddeley, A. (2003). Working memory: Looking back and looking forward. Nature Reviews: Neuroscience, 4, 829-839.

Postle, B. R. (2006.) Working memory as an emergent property of the mind and brain. Neuroscience 139, 23-38. • The paper by Baddely provides an update to a standard theory of working memory. The

paper by Postle provides a critique of the standard Baddeley model. Sayala, S., Sala, J.B., & Courtney, S.M. (2006). Increased neural efficiency with repeated performance of a working memory task is information-type dependent. Cereb Cortex, 16, 609-617. Petrides, M. (2005). Lateral prefrontal cortex: architectonic and functional organization. Philosophical Transactions of the Royal Society of London B, 360, 781-795. • Both papers touch on a controversy relating to the precise roles of dorsal vs. ventral frontal

regions in working memory, with Sayala et al. in support of a domain-based organization and Petrides a proponent of process-based organization.

Badre, D. (2008). Cognitive control, hierarchy, and the rostro-caudal organization of the frontal lobes. Trends in Cognitive Sciences, 12, 193-200. Koechlin, E., & Hyafil, A. (2007). Anterior prefrontal function and the limits of human decision-making. Science, 318, 594-598. Supplementary

Milner, B. (1982). Some cognitive effects of frontal lobe lesions in man. Philosophical Transactions of the Royal Society of London, B298, 211-226. D’Esposito, M. From cognitive to neural models of working memory. Philosophical Transactions of the Royal Society of London B, 362, 761-772. Stuss, D.T., & Alexander, M.P. (2007). Is there a dysexecutive syndrome? Philosophical Transactions of the Royal Society of London B, 362, 901-915. Norman, D.A. and Shallice, T. (1986) Attention to action: willed and automatic control of behaviour. In Consciousness and Self Regulation: Advances in Research and Theory (Vol. 4) (Davidson, R. et al., eds), pp. 1–18, Plenum. Passingham, R.E. (1993). The frontal lobes and voluntary action, Oxford University Press Stuss, D.T. and Benson, D.F. (1987) The frontal lobes and control of cognition and memory. In The Frontal Lobes Revisited (Perecman, E., ed.), pp. 141–158, The IRBN Press. Bunge, S.A., & Wright, S.B. (2007). Neurodevelopmental changes in working memory and cognitive control. Current Opinion in Neurobiology, 17, 243-250.

Discussion questions 1. Can the process-oriented and stimulus-oriented views fit with Baddeley’s multi-component

model of memory? 2. What is the function of the frontal poles? EMOTION AND SOCIAL COGNITION Emotion Overview: Phelps, E.A., & LeDoux, J.E. (2005). Contributions of the amygdala to emotion processing: from animal models to human behavior. Neuron, 48, 175-187. Adolphs, R. (2008). Fear, faces, and the human amygdala. Current Opinion in Neurobiology, in press. • These papers demonstrate the utility of the comparative approach in understanding the role

of the amygdala beyond processing fear. LaBar, K.S., & Cabeza, R. (2006). Cognitive neuroscience of emotional memory. Nature Reviews Neuroscience, 7, 54-64. • A detailed review of lesion and neuroimaging studies of emotional memory. Theory of Mind and Self-Referential Processing Saxe R, Xiao D, Kovacs G, Perrett DI, Kanwisher N. (2004). A region of right posterior superior temporal sulcus responds to observed intentional actions. Neuropsychologia, 42, 1435-1446. Mitchell, J.P. (2008). Activity in right temporo-parietal junction is not selective for theory-of-mind. Cerebral Cortex, 18, 262-271. • This paper challenges the view that the right TPJ is specialized for theory of mind. Relate

this paper back to Corbetta and Posner articles on attention and to Stuss & Levine (2002) review.

Shamay-Tsoory SG, Aharon-Peretz J. (2007). Dissociable prefrontal networks for cognitive and affective theory of mind: a lesion study. Neuropsychologia, 45, 3054-67. • This paper is one of a number of large group focal lesion studies to show that medial

prefrontal cortex is specialized for emotional theory of mind. Supplementary Belova, M.A., Paton, J.J., Morrison, S.E., & Salzman, C.D. (2007). Expectation modulates neural responses to pleasant and aversive stimuli in primate amygdala. Neuron, 55, 970-984.

Rowe G, Hirsh JB, Anderson AK. (2007). Positive affect increases the breadth of attentional selection. Proceedings of the National Academy of Sciences, 104(1):383-8. Susskind, J. Lee, D., Cusi, A., Feinman, R. & Grabski, W. Anderson, A.K. (2008). Expressing fear enhances sensory acquisition. Nature Neuroscience, 11(7):843-50 Amodio, D.M., & Frith, C.D. (2006). Meeting of minds: the medial frontal cortex and social cognition. Nature Reviews Neuroscience, 7, 268-277. Jenkins AC, Macrae CN, Mitchell JP. (2008). Repetition suppression of ventromedial prefrontal activity during judgments of self and others. Proceedings of the National Academy of Sciences, 105, 4507-4512. Greene, J.D., et al. (2001) An fMRI investigation of emotional engagement in moral judgment. Science, 293, 1971-1972. Blair, R.J. (2007). The amygdala and ventromedial prefrontal cortex in morality and psychopathy. Trends in Cognitive Sciences, 11, 387-392. Discussion questions 1. Does emotion perception require “simulation” of sensory experiences? 2. How do emotions influence memory? 3. Is there a specialized neural system that supports human “theory of mind”? Does theory of

mind necessarily involve emotional processing? 4. Is empathy simply an extension of theory of mind? To what extent do these abilities rely on

episodic memory?