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The Vestibular System
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I. Introduction Chapter 1- The Vestibular System in Everyday Life 1.1 Overview of the vestibular system 1.2 Visual acuity and the vestibulo-ocular reflex 1.3 Air-righting reflex in the cat 1.4 Post-rotational reactions 1.5 Positional alcohol nystagmus (PAN) 1.6 Motion sickness 1.7 Vection illusions 1.8 The subjective visual vertical 1.9 Adaptive plasticity 1.10 Path finding and spatial orientation 1.11 Postural control 1.12 Summary 1.13 Selected readings II. Peripheral Vestibular System Chapter 2 - Structure of the Vestibular Labyrinth 2.1 Gross and microscopic anatomy 2.2 Fine structure of the sensory regions Hair cells Supporting cells Transitional regions 2.3 Regional variations in cellular architecture and afferent innervation Cristae ampullares Utricular macula Saccular macula 2.4 Efferent innervation 2.5 Summary 2.6 Selected readings Chapter 3- Hair Cell Transduction 3.1 Mechanoelectric transduction 3.2 Basolateral currents 3.3 Neurotransmitter release and presynaptic calcium channels Calcium channels Neurotransmitter release. 3.4 Postsynaptic mechanisms 3.5 Synaptic transmission involving type I hair cells 3.6 Spike encoding 3.7 Efferent neurotransmission 3.8 Summary 3.9 Selected readings Chapter 4- Physiology of the Vestibular Organs 4.1 General features of the vestibular organs Vestibular organs are inertial sensors Resting discharge Discharge regularity Information transmission 4.2 Semicircular canals Directional properties Macromechanics and the torsion-pendulum model Interspecies variations and canal dimensions Afferent response dynamics Variations in gain and phase Afferent morphology and physiology Dynamic range of afferent discharge 4.3 Otolith organs Directional properties Macromechanics and the otoconial membrane Afferent response dynamics Dynamic range of afferent discharge Variations in gain and phase Afferent morphology and physiology 4.4 Summary 4.5 Selected readings Chapter 5- The Efferent Vestibular System 5.1 Comparative anatomy of central efferent pathways 5.2 Responses of afferents to electrical stimulation of EVS Mammals. Non-mammals 5.3 Responses of efferents to natural stimulation 5.4 Efferent-mediated responses of afferents 5.5 Possible functions of efferents in mammals 5.6 Summary III. Central Vestibular System Chapter 6 - Neuroanatomy of Central Vestibular Pathways 6.1 Introduction 6.2 The vestibular nuclei: subdivisions and anatomical organization Medial vestibular nucleus (MVN) Lateral vestibular nucleus (LVN) Superior vestibular nucleus (SVN) Descending vestibular nucleus (DVN) y group Interstitial nucleus of the vestibular nerve (INT8) Associated cell groups (z, x, f, l,m) Projection and intrinsic neurons Connections with the ipsilateral vestibular nerve Commissural pathways 6.3 Vestibulo-ocular and optokinetic systems Semicircular canal projections to oculomotor neurons Otolith projections to oculomotor neurons Nucleus prepositus hypoglossi (NPH) Interstitial nucleus of Cajal (INC) Reticular formation Optokinetic pathways 6.4 Vestibulospinal systems Medial vestibulospinal tract (MVST) Lateral vestibulospinal tract (LVST) Vestibulo-ocular cervical pathways (VOC) Other vestibulospinal tracts Spinal projections to the vestibular nuclei 6.5 Vestibulocerebellar relations Basic circuitry Vestibular projections to the cerebellum Prepositus nucleus Projections from the deep cerebellar nuclei to the vestibular nuclei Projections from the cerebellar cortex to the vestibular nuclei Cerebellar cortical modules Lateral reticular nucleus Vestibulo-paramedian tract projections 6.6 Vestibulo-autonomic connections 6.7 Vestibular connections with the neocortex 6.8 Pathways involving the hippocampal formation 6.9 Summary 6.10 Selected readings 6.11 List of abbreviations Chapter 7 -Synaptic Mechanisms in the Vestibular Nuclei 7.1 Historical perspective 7.2 Basic circuitry of the vestibular nuclei Ipsilateral vestibular nerve inputs Commissural connections 7.3 Neurotransmitters in the vestibular nuclei Transmission between the vestibular nerve and secondary neurons Transmission within the vestibular nucleus Output pathways of the vestibular nuclei 7.4 Properties of individual neurons Resting discharge 7.5 Central projections of regular and irregular afferents Electrophysiological studies Functional ablation of irregular afferents 7.6 Convergence from separate vestibular organs Convergence from separate vestibular organs Canal-canal convergence Otolith-otolith convergence Spatio-temporal convergence Canal-otolith convergence Convergence from somatosensory receptors 7.7 Summary 7.8 Selected readings IV. Vestibulo-ocular and Vestibulopinal Mechanisms Chapter 8 - An Oculomotor Tutorial 8.1 Overview and classification of eye movement types 8.2 Ocular structure and functional implications The extraocular eye muscles. Mechanics of the oculomotor plant Oculomotor motoneuron discharge. Plant mechanics and premotor control. 8.3 Gaze Redirection Saccades Smooth pursuit Vergence 8.4 Gaze Stabilization Vestibulo-ocular reflexes. Optokinetic system. 8. 5 Interactions between eye and head movements 8.6 Summary 8.7 Selected readings Chapter 9 -Vestibulo-ocular Reflexes 9.1. Semicircular-canal related angular VOR (AVOR) General properties of the canal-related AVOR. AVOR during high frequency rotations. AVOR at low frequencies - velocity storage. AVOR-visual interactions: the optokinetic system. AVOR in three-dimensions. 9.2. Otolith-ocular reflexes Tilt VOR. Otolith influences on the AVOR during off-vertical axis rotations (OVAR). Otolith influences on the AVOR during canal/otolith conflict. Translational VOR (TVOR). Optic flow during translation. Visual mechanisms for short latency visual compensation during translation. Distinguishing tilts from translations. Differences between the AVOR and the TVOR. Functional differences: Foveal rather than full-field image stabilization. Dependence on viewing distance and eye position. Response latency and neural pathways. Comparative adaptation 9.3 Summary 9.4 Selected readings Chapter 10-The Vestibulospinal System and Postural Control 10.1. Reflexes versus multisensory strategies 10.2 Multisensory strategies 10.3. Vestibular reflexes: general considerations 10.4 Vestibulocollic reflexes The angular VCR. The linear VCR evoked by translation and tilts. The cervicocollic reflex 10.5. Control systems analysis of the head-neck plant. Head plant. Vestibulocollic reflex. The cervicocollic reflex Reflex interactions. Use of control systems models 10.6. Vestibulospinal and neck reflexes acting on the limbs Spatial and temporal properties of the reflexes. Afferent origin of the reflexes Neural substrate of the reflexes. Vestibulospinal actions on hindlimb motoneurons. Vestibulospinal actions on forelimb motoneurons. Tonic neck reflexes. 10.7 Summary 10.8 Selected readings V. Signal Processing in Alert Animals Chapter 11- Signal Processing in Vestibular Nuclei of Alert Animals During Natural Behaviors 11.1 Introduction 11.2 Classes of neurons in head-restrained, alert monkeys Position-vestibular-pause (PVP) neurons. Vestibular-only (VO) and vestibular-pause cells. Eye-head (EH) neurons. Burst-tonic (BT) neurons. 11.3 Dynamics of neuronal responses Frequency response during sinusoidal rotations Response linearity Velocity storage 11.4 Response to linear translations in alert head-restrained monkeys Distinguishing translational from tilt. 11.5 Interactions with the oculomotor pathways that control pursuit eye movements 11.6 Integration of inputs from vestibular and optokinetic pathways VN modulation during the OKR Optokinetic pathways to the VN. 11.7 Integration of vestibular and proprioceptive inputs 11.8 Differential processing of active versus passive head movements Neuronal responses during active versus passive head movement. Mechanisms for the differential processing of actively-generated versus passive head movement. 11.9 Vestibular processing depends on current gaze strategy. Vestibular processing during voluntary gaze shifts. Vestibular processing during visual tracking; VOR cancellation and eye-head pursuit. Vestibular processing during near versus far viewing. 11.10 Summary 11.11 Selected readings Chapter 12 - The Cerebellum and the Vestibular System 12. 1 Overview of signal processing in the cerebellum The basic cerebellar circuit Vestibular inputs are specific to localized regions of the cerebellum 12.2 Nodulus and Ventral Uvula Mossy fiber inputs. Climbing fiber inputs. Efferent connections. Neuronal responses Lesions and function. 12.3 Flocculus and ventral paraflocculus Mossy fiber inputs. Climbing fiber inputs Efferent projections of the flocculus. Differences between the flocculus and ventral paraflocculus Neuronal responses. Complex spikes: Simple spikes. Changes in neuronal responses following VOR learning Lesions and function Lesions studies emphasize the role of the flocculus in VOR motor learning. 12.4 The Vermis of the Anterior and Posterior Lobes 12.4 The Vermis of the Anterior and Posterior Lobes 12.5 The Deep Cerebellar Nuclei Fastigial Nucleus. Rostral fastigial nucleus Caudal fastigial nucleus The interposed nuclei. Dentate nuclei. 12.6 Summary 12.7 Selected Readings VI..Functional Considerations Chapter 13 - Learning and Compensation in the Vestibular System 13.1 Motor learning in the vestibulo-ocular reflex The adaptive capabilities of the VOR. Signal flow in the VOR network. Rules for the VOR and motor learning. Possible sites of motor learning: cerebellum versus brain stem Evidence for sites of learning and memory. Possible cellular mechanisms of synaptic plasticity Cerebellar mechanisms Brain stem mechanisms Consolidation of VOR motor memory Generalization: can learning be applied to new situations? 13.2. Compensation for vestibular damage Uninilateral labyrinthectomy Activity in the vestibular nuclei following labyrinthectomy Cellular mechanisms of compensation in the vestibular nuclei intrinsic mechanisms. synaptic mechanisms axonal reorganization stress The role of the cerebellum in compensation 13.3 Summary 13.4 Selected readings Chapter 14-Cortical Representations of Vestibular Information 14.1. Introduction 14.2. Historical Perspective 14.3. Multiple representations of vestibular signals in the cerebral cortex. Visuomotor areas in frontal cortex. Extrastriate visual cortex (MSTd). Ventral intraparietal (VIP) area Parieto-insular vestibular cortex (PIVC), area 2v and area 3a 14.4. Ascending vestibular pathways through the thalamus 14.5 Descending cortical information affecting vestibular responsiveness in the vestibular nuclei 14.6. Vestibular influences in the head direction circuit of the limbic system 14.7 Summary 14.8 Selected readings Chapter 15-Reference Frames Used in the Coding Vestibular Information 15.1. Definitions of coordinate systems and reference frames 15.2. Head- versus body-centered reference frames: Vestibular/neck proprioceptive interactions 15.3. Head- versus eye-centered reference frames for self-motion perception: vestibular/visual interactions in extrastriate visual cortex 15.4. Head- versus world-centered reference frames: Canal/otolith convergence for inertial motion detection Computational solution for the two ambiguities of peripheral vestibular sensor The linear acceleration problem: evidence for segregation of tilt and translation VII. Clinical Disorders Chapter 16- Clinical Manifestations of Vestibular Dysfunction 16.1 Prevalence and impact of vestibular disorders 16.2 Diagnosis of vestibular disorders 16.3 Planes of individual canals and direction of eye movements Benign paroxysmal positional vertigo Positional alcohol nystagmus Superior semicircular canal dehiscence syndrome 16.4 Recovery of the horizontal VOR after unilateral labyrinthectomy 16.5 Multisensory control of posture 16.6 Disorders of otolith function 16.7 Clinical tests of vestibular function Caloric test Rotational chair tests Quantitative evaluation of the VOR evoked by rapid head movements Vestibular-evoked myogenic potentials (VEMPs) 16.7 Future directions Hair-cell regeneration Vestibular prosthesis 16.8 Summary 16.9 Selected readings

About the Author

Jay M. Goldberg, The Department of Neurobiology, Pharmacology and Physiology, University of Chicago, Chicago, IL. Victor J. Wilson, Laboratory of Neurophysiology, Rockefeller University, New York, NY. Kathleen E. Cullen, Laboratory of Neurophysiology, McGill University, Montreal, Quebec, Canada. Dora E. Angelaki, Department of Anatomy and Neurobiology, Division of Biology and Biomedical Sciences School of Medicine, Washington University in St. Louis, St. Louis, MO. Dianne M. Broussard, Division of Cellular and Molecular Biology, Toronto Western Hospital, Toronto, Ontario Canada Jean A. Buttner-Ennever, Institute of Anatomy III, Ludwig-Maximilians University of Munich, Munich, Germany. Kikuro Fukushima, Department of Physiology, Hokkaido University School of Medicine, Kitaku, Sapporo, Japan. Lloyd B. Minor, Department of Otolaryngology, Head and Neck Surgery, Department of Neuroscience, The Johns Hopkins University School of Medicine, Baltimore, MD.

Reviews

"In this fine new treatise on the Vestibular System an introductory chapter overviews the vital roles the system plays in everyday life, while a final chapter reveals the devastating consequences of its clinical malfunction. Between these two chapters we are led systematically from mechano-neural transduction in the periphery, through an elaborate maze of central neural information processing to the behaviorally meaningful outputs of vestibulo-ocular and postural motor control. An appealing aspect of the book is that it reads like a series of living detective stories, written by scientists actively engaged in the tales they tell. In overview It emerges as a landmark achievement which will undoubtedly prove essential reading for a wide range of workers in the field." -- Geoffrey Melvill Jones, MB BCh, FRSC, FRS , Research Professor (Adjunct), Department of Clinical Neurosciences, University of Calgary, Alberta, Canada "The world of vestibular science has been waiting restlessly for this book for a very long time. Written by those who have made many of the most important contributions to the field, it will become the standard reference work for anyone wanting to know about the neurophysiology of the vestibular system." -- Michael Halmagyi, MD, Neurology Department, Royal Prince Albert Hospital, Camperdown NSW 2050, Sydney, Australia "At last, an 'update' of Vestibular Physiology by Wilson and Melvill Jones, which has been THE BOOK of the last three decades for all serious students of the vestibular system. One of the more attractive features of that book was that it emphasized function, and incorporated anatomical and physiological data into conceptual and mathematical models that made sense! The Vestibular System-A Sixth Sense has retained this elegance...using the many new techniques available to probe the signal processing in the brain that underlies vestibular behavior. Its unique style of multi authorship, in which all contribute to every chapter, has made the entire book attractive to specialists of all types who might want to delve into the vestibular system, from the bedside clinician to the membrane physiologist." -- David S. Zee, MD, Department of Neurology, Johns Hopkins Hospital, Baltimore, MD 21287 and co-author of The Neurology of Eye Movements, Fourth Edition (Oxford University Press) "This is an outstanding reference that will likely find its way into the offices of basic scientists and clinicians alike. It is an important contribution to vestibular science...The authors have done a wonderful job of consolidating a broad body of current knowledge into a readable book." -- Doody's "This book is one of those rare ones on the vestibular system that takes a broad and deep look into it. Its coverage is not only extensive but intensive as well... It is an excellent book on the subject." --Biz India

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