Edited by GERHARD ROTH and MARIO F. WULLIMANN University of Bremen, Brain Research Institute and Center for Cognitive Science, Bremen, Federal Republic of Germany A JOHN WILEY & SONS, INC New York 2001 FROM THE BACK COVER: Can one fully understand the mechanisms of brain evolution? Why have some brains become large andcomplex, and others small and simple, while still others have remained the same for hundreds of millions of years? Where, how, and why did cognition evolve? Is there any definable relationship between cognitive function and brain structure and function? These are just some of the questions posed by the contributing authors of this unique book. By bringing together two intrinsically related topics-the structural evolution of the brain and the concomitant evolution of cognitive functions-Brain Evolution and Cognition addresses the much debated topic of brain evolution, cognitive functions, and the relationship between them. Uniting information on structural brain variability and cognitive aspects in one text, this book provides a survey of the current status of what is known about animal cognition and its relationship to the underlying diversity of brain structures. This volume cuts across boundaries by introducing data on various species and also: o Reviews developmental and adult brain organization in an evolutionary context o Presents case studies of vertebrate brain evolution o Offers an overview of cognition from neural basis to behavior o Is accessible to a broad range of readers with its nontechnical writing style o Includes lavish illustrations for a clear and concise presentation of concepts With contributions by world-renowned leaders in their fields, Brain Evolution and Cognition is an indispensable reference for neuroanatomists, neuroethologists, evolutionary biologists, developmental biologists, neurologists, neuroscientists, zoologists, and cognitive and behavioral psychologists. CONTENTS INTRODUCTION Problems in the study of brain evolution and cognition 1 Mario F. Wullimann and Gerhard Roth Handicaps in the search for an evolutionary history of vertebrate brains 1 Silent revolutions emerging 2 The amniote telencephalon as a case in point 3 A special position for humans? 5 What is cognition? 6 References 7 PART I DEVELOPMENTAL AND ADULT BRAIN ORGANIZATION IN EVOLUTION 1 Brain phenotypes and early regulatory genes: The Bauplan of the metazoan central nervous system 11 Mario F: Wullimann Introduction 11 Comparative phenotypic analysis of metazoan central nervous characters 13 The cladistic framework 13 A can of worms: Plathelminths, nemathelminths, nemertines 14 The molluscan controversy 20 The arthropod CNS, rather than being ancestral to the vertebrate CNS, is equally remote from the basic bilaterian Bauplan as the craniate brain 22 Deuterostome nervous systems 25 Conclusion 28 Early genes in neural development - Do they tell a different story? 30 Development and Bauplan of the vertebrate CNS 30 Early regulatory genes and neuromeres in the vertebrate brain 32 Early regulatory genes and the insect CNS 32 Phylogenetic interpretation of molecular genetic and phenotypic data 34 Conclusion 36 References 37 2 The echinoderm nervous system and its phylogenetic interpretation 41 Thomas Heinzeller and Ulrich Welsch Introduction 41 Description of Nervous Systems 44 Larval nervous system 44 Postmetamorphotic nervous system: Common features 45 Postmetamorphotic nervous system: Group-specific features 51 Cryptosyringida 54 Inter-class comparison of sensory versus motor function 60 Questions of Symmetry 60 Central part of the body 60 Bilateral symmetry and segmentation of the arms 61 Are echinoderm arms homologous with bilaterian trunks? 62 Echinoderm ectoneural cord versus chordate neural plate 62 Additional body axes 64 Hydrocoel and notochord - are they convergent or homologous? 65 Locomotion of Echinoderms 66 Phylogenetic Aspects 68 Consistency versus flexibility of regulatory genes 68 Garstang's hypothesis 68 Systematics of echinoderms 69 Missing brain 69 References 69 3 Evolution of vertebrate motor systems 77 Hans J. ten Donkelaar Introduction 77 Basics of vertebrate locomotion 80 Prehensile extremities 84 Features of the ancestral vertebrate motor system 85 Neural control of quadrupedal locomotion 91 Supraspinal control 93 Descending supraspinal pathways 93 The cerebellorubrospinal limb control system 98 The special case for birds 104 Summary 106 References 107 4 Sensory system evolution in vertebrates 113 William Hodos and Ann B. Butler How many senses? 113 How many cranial nerves? 115 Trends in sensory system evolution 116 Ascending sensory pathways 119 Neuroembryology and the evolution of sensory systems 120 Evolution of new sensory receptors 122 Evolution of new primary, secondary, and higher order sensory nuclei 122 The evolution of sensory specialists 123 Evolution of central sensory pathways 124 The evolution of new central sensory nuclei 125 The evolution of sensory maps 126 Loss of sensory receptors and central pathways 127 Mechanisms of sensory system evolution 130 References 131 5 Evolution of the forebrain in tetrapods 135 Toru Shimizu Introduction135 Evolution of tetrapods 137 Early tetrapods - Ancestral amphibians 137 Early amniotes - Ancestral reptiles 137 Synapsids - Mammals 140 Sauropsids - Reptiles 142 Sauropsids - Birds 144 Conclusion 145 Forebrain organization of living tetrapods 145 Amphibian pattern 146 Mammalian pattern 152 Sauropsid pattern 156 Conclusion 161 Evolutionary history of the tetrapod forebrain 162 Early tetrapods 163 Early amniotes 163 Synapsids and sauropsids 165 Conclusion 169 Environmental pressures on the tetrapod forebrain 170 Anamniote pattern versus amniote pattern 171 Sauropsid pattern versus mammalian pattern 172 Conclusion 175 References 176 6 Neocortical macrocircuits 185 Rudolf Nienwenhuys Introduction 185 Major sensorimotor projections 186 Control systems 189 Reticular, greater limbic and general modulatory inputs to neocortical circuitry 192 The ascending reticular system 192 The greater limbic system 193 Monoaminergic and cholinergic modulatory systems 194 Cortico-subcortico-cortical association systems 195 The thalamic association system 196 The striatal association system 198 The cerebellar association system 199 Summary 201 References 202 7 Hunting in barn owls: Peripheral and neurobiological specializations and their general relevance in neural computation 205 Hermann Wagner Introduction 205 Evolutionary position and geographical distribution of the barn owl 206 Hunting as a complex behavior 206 General comments 206 Formal description of the hunting situation 208 Adaptations of barn owls to hunting in the night 209 The barn owl's brain 212 Morphological adaptations of the owl's brain to hunting and life at night 213 Physiological adaptations 216 Coincidence detection 218 Further brain adaptations subserving sound-localization behavior 225 Differences in the representation of acoustic space in diurnal and nocturnal owls 228 General meaning of coincidence detection and across-frequency integration 228 Conclusions 231 References 232 8 Evolution and devolution: The case of bolitoglossine salamanders 237 Gerhard Roth and David B. Wake Introduction 237 The Bolitoglossini 238 The brain of salamanders and frogs 244 The visual system of bolitoglossines 249 Retina and retinofugal system 249 Tectum 250 The fate of other sensory systems 253 Causes and consequences of simplification in the context of paedomorphosis 254 What do bolitoglossines tell us about evolution in general and brain evolution in particular? 258 References 260 9 Evolutionary constraints of large telencephala 265 Gerd Rehkamper, Heiko D. Frahm, and Michael D. Mann Evolution -What does that mean? 265 Brain and brain part size as a heuristic tool 266 What factors influence brain size or brain part size? 268 There is no brain size alteration 269 Brain size alterations are epiphenomena 269 Brain size is influenced by individual learning 270 Brain size and brain part size reflect adaptation 271 Definition of "large telencephala" 271 Mammals 272 Telencephala enlarged because of dominance of olfactory orientation 272 Telencephala enlarged because of superior spatial cognition 272 Telencephala enlarged because of a voluminous isocortex 275 Isocortex (and therefore, telencephalon) enlarged because of elaborated somatosensory areas together with a necessity of motor coordination in a subterranean life 276 Telencephala enlarged because of multimodal integration 278 Birds 278 Telencephala enlarged because of olfaction 278 Telencephala enlarged because of spatial cognition 279 Telencephala enlarged because of isocortical equivalents 280 Domesticated animals 283 Again: Theories of brain size and brain composition 285 Conclusions 288 References 289 PART II COGNITION: FROM NEURAL BASIS TO BEHAVIOR 10 Brain and cognitive function in teleost fishes 297 Leo S. Demski and Joel A. Beaver Introduction 297 Studies on cognition in fishes 298 Brain lesions and cognitive behavior in fishes 300 Telencephalon: Nonspatial learning 300 Telencephalon: Spatial learning 301 Cerebellum 302 Tectum 303 Relative brain size and development: Implications for cognitive function in fishes 304 Studies in minnows (Cypriniformes) 304 Blind and sighted characins 306 The cichlids of the African great lakes 306 Coral reef percomorphs 307 Microcircuitry of telencephalic enhancements in selected percomorphs 311 Area dorsalis telencephali pars lateralis (dorsal part) 313 Area dorsalis telencephali pars centralis 317 Area dorsalis telencephali pars medialis 319 Behavioral studies on the enlarged telencephalon of percomorphs 321 Summary and conclusions 323 References 325 11 Cognition in insects: The honeybee as a study case 333 Randolf Menzel, Martin Giurfa, Bertram Gerber, and Frank Hellstern Introduction: Brain, behavior, and biology of honeybees 333 Behavior and biology of honeybees 333 Design of an insect brain 335 Elementary and configural forms of learning in classical conditioning 338 The preparation: Classical conditioning of the proboscis extension reflex 338 A cognitive approach to memory dynamics 340 The elementary-configural distinction 342 Cognitive aspects of elementary forms of conditioning? 342 Configural forms of conditioning 345 Learning in the natural context 346 Context-dependent learning and retrieval 346 Serial order in a spatiotemporal domain 348 The representation of space in navigation 350 Visual discrimination learning in honeybees: Generalization, categorization, and concept formation 354 Conclusion 359 Basic cognition with a small brain 359 The ecological niche and basic cognition 360 References 362 12 Insect brain 367 Nicholas J. Strausfeld Introduction 367 General features of segmental ganglia 368 The protocerebrum and the preoral brain 372 Evolutionary considerations 372 General organization of the protocerebrum 373 The mushroom bodies 375 Structure 375 Evolution of mushroom bodies in insects 377 Relationship to primary sensory neuropils 378 Mushroom body physiology 381 Roles of mushroom bodies 382 The central complex 384 Evolutionary considerations 384 Organization of the central complex 384 Central complex function 385 Comparisons of brain regions amongst arthropods 389 Mushroom bodies 389 The central complex 390 Insect and vertebrate brains compared 393 Equivalence of insect and vertebrate embryonic forebrain 392 The adult brain 393 References 395 13 Conservation in the neurology and psychology of cognition in vertebrates 401 Euan M. Macphail Introduction: Complexity in brains and behaviour 401 Species differences in intelligence 402 Birds and mammals compared 403 The basal ganglia 405 Paleostriatal lesions and classical conditioning in the pigeon 407 The archistriatum 409 Posteromedial archistriatum: Fear and avoidance 409 Anterior and intermediate archistriatum: Parallels with isocortex 411 Olfactory cortex 411 Hippocampal complex 412 Isocortical analogues/homologues 416 Conclusions 426 References 427 14 Multimodal areas of the avian forebrain-Blueprints for cognition? 431 Onur Gunturkun and Daniel Durstewitz The theme 431 Working memory and prefrontal cortex 432 Avian brain and cognition 434 Details of the machine 437 The decline of a memory store 439 Simulation of the machine 442 Looking inside 449 References 450 15 Cognition of birds as products of evolved brains 451 Juan D. Delius, Martina Siemann, Jacky Emmerton and Li Xia Introduction 451 Categorization 456 Concepts 461 Transitivity 467 Numerosity 472 Epilogue 477 References 483 16 What can the cerebral cortex do better than other parts of the brain 491 Almut Schaz Introduction 491 Basic connectivity of the isocortex 492 The cerebral cortex and cognition 494 Comparative aspects 497 Brain size and connectivity 497 Allocortex and reptilian cortex 498 References 499 17 Evolution and complexity of the human brain: Some organizing principles 501 Michel A. Hofman Introduction 501 Evolution of brain size 502 Encephalization in primates 505 General constraints on brain evolution 507 Evolution and geometry of the cerebral cortex 510 Design principles of neuronal organization 514 Biological limits to information processing 515 Concluding remarks 518 References 519 18 The evolution of neural and behavioral complexity 523 Harry J. Jerison Introduction 523 Vigilance and attention: An old-fashioned view 525 Costs and attention 526 Costs and brains 531 Brain size in living vertebrates: Allometry and encephalization 533 Exceptions 535 Early avian and mammalian encephalization 539 Progressive encephalization in mammals 542 More on neural information 544 Neural and behavioral complexity 546 Why some brains are big: What do big brains do? 547 References 551 19 The evolution of consciousness 555 Gerhard Koth Phenomenology of consciousness 556 The neurobiological basis of the different states and appearances of consciousness 558 Cognition and consciousness in animals 564 Animal brains and human brain 568 Consciousness and language 577 Conclusions 579 References 580 Index 583 WHERE TO ORDER: John Wiley and Sons, Inc. 1 Wiley Drive Somerset, NJ 08875-1272 Tel: 1-800-225-5945 URL: http://www.wiley.com ISBN 0-471-33170-8 (cloth: elk. Paper) Price: $125.00
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