Prelims-P373677 3/16/07 4:42 PM Page i COGNITION, BRAIN, AND CONSCIOUSNESS This page intentionally left blank Prelims-P373677 3/16/07 4:42 PM Page iii COGNITION, BRAIN, AND CONSCIOUSNESS Introduction to Cognitive Neuroscience Edited by B J. B ERNARD AARS N M. G ICOLE AGE AMSTERDAM • BOSTON (cid:127) HEIDELBERG (cid:127) LONDON (cid:127) NEW YORK (cid:127) OXFORD PARIS (cid:127) SAN DIEGO (cid:127) SAN FRANCISCO (cid:127) SINGAPORE (cid:127) SYDNEY (cid:127) TOKYO Academic Press is an imprint of Elsevier Prelims-P373677 3/16/07 4:42 PM Page iv Academic Press is an imprint of Elsevier 84 Theobald’s Road, London WC1X 8RR, UK 30 Corporate Drive, Suite 400, Burlington, MA01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA First edition 2007 Copyright © 2007 Elsevier Ltd. 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Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made British Library Cataloguing in Publication Data Acatalogue record for this book is available from the British Library Library of Congress Cataloging in Publication Data Acatalog record for this book is available from the Library of Congress ISBN: 978-0-12-373677-2 For information on all Academic Press publications visit our web site at http://books.elsevier.com Typeset by Charon Tec Ltd (AMacmillan Company), Chennai, India www.charontec.com Printed and bound in Hong Kong 07 08 09 10 11 10 9 8 7 6 5 4 3 2 1 Front cover:For centuries, epileptic episodes have been known to evoke striking visual experiences. The outside cover art shows an acrylic painting by artist Craig Getzlaff, called Brainstorms #20. The artist writes, “In these paintings, I am trying to communicate to the public and to myself what an immense experience an epileptic seizure really is” (p. 27). The painting comes from a collection called Visions: Artists living with epilepsy(Source:Schachter, 2003, with kind permission). Prelims-P373677 3/16/07 4:42 PM Page v Contents Preface xiii 2.1 The ‘inner senses’ 33 2.2 Output functions 34 List of contributors xvii 2.3 Only a fleeting moment ... 34 2.4 Understanding Clive Wearing Chapter 1 Mind and brain in the functional framework 37 2.5 The importance of immediate memory 38 Bernard J. Baars 3.0 Limited and large-capacity functions 39 1.0 Introduction 3 3.1 Dual task limits 39 2.0 An invitation to mind-brain science 3 3.2 Some very large brain capacities 41 3.0 Some starting points 4 3.3 Why are there such narrow 3.1 Distance: seven orders of magnitude 4 capacity limits? 42 3.2 Time: ten orders of magnitude 6 3.4 Measuring working memory 42 3.3 The need to make inferences – 4.0 The inner and outer senses 44 going beyond the raw observations 7 4.1 The mind’s eye, ear and voice 45 3.4 The importance of convergent measures 10 4.2 The imagery sketchpad may use visual 3.5 Major landmarks of the brain 10 regions of cortex 46 4.0 Some history, and ongoing debates 13 4.3 Is inner speech like outer speech? 47 4.1 The mind and the brain 13 4.4 Is there only one working memory? 47 4.2 Biology shapes cognition and emotion 15 5.0 The central executive 48 4.3 Cajal’s neuron doctrine: the working 5.1 Executive effort and automaticity 49 assumption of brain science 16 5.2 Executive and spontaneous attention 51 4.4 Pierre-Paul Broca and the localization of 6.0 Action 52 speech production 16 7.0 Consolidation of short-term events 4.5 The conscious and unconscious mind 23 into long-term memory 53 5.0 The return of consciousness in the sciences 24 7.1 Is working memory just re-activated 5.1 How conscious and unconscious permanent memory? 54 brain events are studied today 24 8.0 Summary 55 5.2 History hasn’t stopped 26 9.0 Study questions and drawing practice 56 6.0 Summary 27 9.1 Study questions 56 7.0 End of chapter exercises 28 9.2 Drawing exercises 56 7.1 Study questions 28 7.2 Drawing exercise 29 Chapter 3 Neurons and their connections Bernard J. Baars Chapter 2 Aframework 1.0 Introduction 59 Bernard J. Baars 1.1 Real and idealized neurons 60 1.0 Introduction 31 1.2 Excitation and inhibition 61 2.0 Classical working memory 32 1.3 Neural computation 63 v Prelims-P373677 3/16/07 4:42 PM Page vi vi CONTENTS 2.0 Working assumptions 63 5.1 Why we need multiple tests of 2.1 Starting simple: receptors, brain function 116 pathways and circuits 65 5.2 Brain damage and causal inferences 117 3.0 Arrays and maps 67 6.0 Summary 118 3.1 Maps flow into other maps 69 7.0 Chapter review 118 3.2 Neuronal arrays usually have two- 7.1 Drawing exercises and study questions 118 way connections 71 3.3 Sensory and motor systems work together 72 Chapter 5 The brain 3.4 Temporal codes: spiking patterns and Bernard J. Baars brain rhythms 73 3.5 Choice-points in the flow of information 74 1.0 Introduction 121 3.6 Top-down or expectation-driven 1.1 The nervous system 122 processing 75 1.2 The geography of the brain 123 4.0 How neural arrays adapt and learn 76 2.0 Growing a brain from the bottom up 126 4.1 Hebbian learning: ‘Neurons that fire 2.1 Evolution and personal history are together, wire together’ 76 expressed in the brain 126 4.2 Neural Darwinism: survival of the 2.2 Building a brain from bottom to top 127 fittest cells and synapses 79 3.0 From ‘where’ to ‘what’: the functional 4.3 Symbolic processing and neural nets 80 roles of brain regions 132 5.0 Coordinating neural nets 82 3.1 The cerebral hemispheres: the left-right 5.1 Functional redundancy 83 division 132 6.0 Summary 84 3.2 Output and input: the front-back 7.0 Study questions and drawing exercises 85 division 136 7.1 Study questions 85 3.3 The major lobes: visible and hidden 138 7.2 Drawing exercises 85 3.4 The massive interconnectivity of the cortex and thalamus 142 3.5 The satellites of the subcortex 144 Chapter 4 The tools: Imaging the living brain 4.0 Summary 146 Bernard J. Baars and Thomas Ramsøy 5.0 Chapter review 146 5.1 Study questions 146 1.0 Introduction 87 5.2 Drawing exercises 146 1.1 Brain recording: more and less direct measurements 88 1.2 The time-space tradeoff 91 Chapter 6 Vision 2.0 Arange of useful tools – measuring Frank Tong and Joel Pearson electric and magnetic signals 93 2.1 Single-unit recording 93 1.0 Introduction 149 2.2 Animal and human studies cast 1.1 The mystery of visual experience 149 light on each other 96 1.2 The purpose of vision: knowing 2.3 Electroencephalography (EEG) 96 what is where 150 2.4 Magnetoencephalography 101 1.3 Knowing what: perceiving features, 2.5 Zapping the brain 101 groups and objects 151 3.0 fMRI and PET: indirect signals for 1.4 Knowing where things are 152 neural activity 106 2.0 Functional organization of the visual system 152 3.1 Pros and cons of PET and fMRI 106 2.1 The retina 152 3.2 Regions of interest 107 2.2 Lateral geniculate nucleus (LGN) 155 3.3 The resting brain is not silent 113 2.3 Primary visual cortex (V1) 156 3.4 Empirically defining cognitive 2.4 Extrastriate visual areas – outside of V1 158 functions: the creative key 114 2.5 Area MT 158 4.0 Conscious versus unconscious brain events 115 2.6 The ventral and dorsal pathways: 5.0 Correlation and causation 115 knowing what and where 159 Prelims-P373677 3/16/07 4:42 PM Page vii CONTENTS vii 2.7 Areas involved in object recognition 161 4.4 The link between speech perception and 2.8 Lateral occipital complex (LOC) 162 production 212 2.9 Fusiform face area (FFA) 162 4.5 Damage to speech perceptual systems 213 2.10 Parahippocampal place area (PPA) 162 4.6 Aworking model for speech perception 3.0 Theories of visual consciousness: in the brain 215 where does it happen? 162 5.0 Music perception 216 3.1 Hierarchical and interactive 5.1 Stages of music processing 216 theories of vision 163 5.2 Aseparate system for music perception? 217 4.0 Brain areas necessary for visual 6.0 Learning and plasticity 217 awareness: lesion studies 164 6.1 Plasticity due to deprivation 217 4.1 Consequences of damage to early 6.2 Plasticity due to learning 218 visual areas 164 6.3 Plasticity due to expertise 219 4.2 Extrastriate lesions – damage outside 7.0 Auditory awareness and imagery 219 area V1 165 7.1 Auditory awareness during 4.3 Damage to ventral object areas 166 sleep and sedation 219 4.4 Damage to dorsal parietal areas 168 7.2 Auditory imagery 221 5.0 Linking brain activity and visual experience 170 8.0 Summary 222 5.1 Multistable perception 170 9.0 Chapter review 222 5.2 Binocular rivalry: what you see is 9.1 Study questions 222 what you get activated 171 9.2 Drawing exercise 222 5.3 Visual detection: did you see it? 172 9.3 Exploring more 223 5.4 Constructive perception: more to vision than meets the eye... 173 5.5 Neural correlates of object recognition 175 Chapter 8 Attention and consciousness 6.0 Manipulations of visual awareness 175 Bernard J. Baars 6.1 Transcranial magnetic stimulation 176 6.2 Unconscious perception 177 Introduction 225 7.0 Summary 178 2.0 Adistinction between attention and 8.0 Study questions and drawing exercises 180 consciousness 225 2.1 Cortical selectionand integration 226 2.2 Selective attention: voluntary and Chapter 7 Hearing and speech automatic 228 Nicole M. Gage 3.0 Experiments on attention 228 3.1 Methods for studying selective 1.0 Introduction 183 attention 230 1.1 Amodel for sound processing 184 4.0 The brain basis of attention 233 1.2 Sound and hearing basics 186 4.1 Attention as biased competition among 2.0 The central auditory system 190 neuron populations 233 2.1 Auditory pathways 191 4.2 Guiding the spotlight 234 2.2 Auditory cortex 192 4.3 Salience maps help guide attentional 3.0 Functional mapping of auditory selection 234 processing 197 4.4 Executive (voluntary) attention 235 3.1 Primary auditory cortex 197 4.5 Visual attention may have evolved 3.2 The role of the planum temporale in from eye movement control 236 sound decoding 197 4.6 Maintaining attention against distraction 236 3.3 Cortical auditory ‘what’ and 4.7 Attention and consciousness 240 ‘where’ systems 199 5.0 The brain basis of conscious experience 240 4.0 Speech perception 207 5.1 Conscious cognition 240 4.1 Background and history 208 5.2 Unconscious comparisons 241 4.2 Early theories of speech perception 210 5.3 Binding features into conscious objects 243 4.3 Functional mapping of speech-specific 5.4 Visual feature integration in processes 211 the macaque 244 Prelims-P373677 3/16/07 4:42 PM Page viii viii CONTENTS 5.5 Conscious events recruit widespread brain 6.3 Different types of working memory 284 activation 246 6.4 Prefrontal cortex – storage or process 5.6 Fast cortical interactions may be needed for control? 285 conscious events 250 6.5 Combining prefrontal and MTLregions for 6.0 Asummary and some hypotheses 251 working memory 286 7.0 Study questions 253 7.0 Retrieval and metacognition 286 7.1 False retrieval 287 7.2 Hemispheric lateralization in retrieval 287 Chapter 9 Learning and memory 7.3 Theta rhythms may coordinate memory Morris Moscovitch, Jason M. Chein, Deborah Talmi, and retrieval 288 Melanie Cohn 8.0 Other kinds of learning 288 9.0 Summary 289 1.0 Introduction 255 10.0 Drawings and study questions 290 1.1 Afunctional overview 258 1.2 Learning and memory in the functional framework 258 Chapter 10 Thinking and problem-solving 1.3 Implicit and explicit memory 260 Bernard J. Baars 2.0 Amnesia 261 2.1 HM: the best-studied amnesia patient 261 1.0 Working memory 294 2.2 Asummary of amnesia 264 1.1 Working memory overlaps with attention, 2.3 Spared functions in amnesia: implicit and conscious events and episodic recall 295 procedural memory 264 2.0 Explicit problem-solving 296 2.4 Spared implicit learning 266 2.1 Executive control in problem-solving 298 3.0 Memories are made of this 267 3.0 Mental workload and cortical activity 301 3.1 Electrically evoked autobiographical 4.0 Using existing knowledge 303 memories 267 4.1 Practice and training may change 3.2 Long-term potentiation and long-term connectivities in the brain 304 depression: excitatory and inhibitory 4.2 Semantic memory 304 memory traces 269 4.3 Abstract concepts, prototypes, and 3.3 Consolidation: from temporary to networks 305 permanent storage 270 4.4 Knowledge comes in networks 306 3.4 Rapid consolidation: synaptic mechanisms, 4.5 Conceptual deficits 308 gene transcription, and protein synthesis 273 4.6 Judgments of quantity and number 309 3.5 System consolidation: interaction between the 5.0 Implicit thinking 310 medial temporal lobes and neocortex 273 5.1 Feelings of knowing 311 4.0 Varieties of memory 273 6.0 Summary and conclusions 313 4.1 Episodic and semantic memory: 7.0 Drawings and study questions 314 ‘Remembering’ versus ‘knowing’ 274 4.2 Episodic memories may turn into semantic memories over time 277 Chapter 11 Language 4.3 Episodic and semantic memory are often Bernard J. Baars combined 277 5.0 MTLin explicit learning and memory 278 1.0 Introduction 317 5.1 Divided attention interferes with 2.0 The nature of language 318 learning 278 2.1 Biological aspects 320 6.0 Prefrontal cortex, consciousness and working 2.2 Language origins 325 memory 279 2.3 Speech versus language 325 6.1 Working with memory: the frontal lobe 3.0 The sounds of spoken language 325 works purposefully withmemory 282 4.0 Planning and producing speech 328 6.2 Prefrontal cortex in explicit (conscious) and 5.0 Evolutionary aspects of speaking and implicit (unconscious) learning and listening 330 memory 283 6.0 Words and meanings 331 Prelims-P373677 3/16/07 4:42 PM Page x x CONTENTS 4.0 Summary 409 6.0 Other topics in neuron modeling 459 5.0 Chapter review 409 6.1 Hebbian learning 459 5.1 Study questions 409 6.2 Activation functions 459 5.2 Drawing exercises 409 7.0 More than one neuron 459 7.1 The perceptron 460 7.2 Limitations of the perceptron 460 Chapter 15 Development 7.3 The multilayer perceptron 461 Nicole M. Gage and Mark H. Johnson 7.4 Cognition and perceptrons? 461 8.0 Recursive or dynamic networks 462 1.0 Introduction 411 8.1 Asimple example of a recursive net 462 1.1 New techniques for investigating the 8.2 Hopfield nets and Boltzmann developing brain 412 machines 462 1.2 The mystery of the developing brain: old 8.3 Other recursive systems 463 questions and new 413 9.0 Looking back on Part l 465 2.0 Prenatal development: from blastocyst to baby 413 Part 2: Seeing is believing 465 2.1 Epigenesis 414 10.0 What is to be seen? 465 2.2 The anatomy of brain development 414 11.0 The NRM neuron 465 2.3 Neural migration 416 11.1 The activation and output computation 2.4 Nature and nurture revisited 420 algorithm 465 2.5 Prenatal hearing experience: voice and 11.2 An experiment with a single basic digital music perception before birth 422 neuron 466 3.0 The developing brain: a lifetime of change 423 11.3 An experiment with a layer of basic digital 3.1 The rise and fall of postnatal brain neuron 467 development 423 12.0 The NRM dynamic neural net 467 3.2 Regional differences in brain 12.1 The state as a label of the input 467 development 424 12.2 The state as an inner image (icon) of the 4.0 Developing mind and brain 425 input 468 4.1 The first year of life: an explosion of 12.3 The inner state as a repository of sensory growth and development 427 memories 469 4.2 Childhood and adolescence: dynamic and 12.4 The state space of the memory staged growth 437 network 469 5.0 Early brain damage and developmental 13.0 Acomplex cognitive system in NRM 470 plasticity 448 13.1 What does the model currently tell us? 471 6.0 Chapter Summary 450 13.2 Response to input changes 472 7.0 Chapter review 451 13.3 Response to simulated voice input 472 7.1 Study questions 451 13.4 Recall in the absence of stimulus 473 13.5 Summary of NRM work 473 13.6 Neural models of cognition: conclusion 473 Appendices 14.0 Self-test puzzles and some solutions 474 14.1 Exercises with NRM 474 A Neural models: Aroute to cognitive 14.2 Looking at a more complex system (visual brain theory awareness) 475 Igor Aleksander 14.3 Beyond the call of duty: stacking and self 476 Part 1: Traditional neural models 454 1.0 Why two parts? 454 2.0 What is a neural model? 454 B Methods for observing the living brain 3.0 The neuron 455 Thomas Ramsøy, Daniela Balslev, and Olaf Paulson 4.0 The basic artificial neuron (McCulloch and Pitts, 1943) 456 1.0 Historical background 477 5.0 Learning in a neuron – some basic 1.1 Correlating brain and mind 477 notions 457 1.2 Recording brain activation 478