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Glial Physiology and Pathophysiology

by ;
Edition:
1st
ISBN13:

9780470978528

ISBN10:
047097852X
Format:
Hardcover
Pub. Date:
4/15/2013
Publisher(s):
Wiley-Blackwell
List Price: $154.95

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Summary

This comprehensive advanced text on the biology and pathology of glial cells--the most numerous cells in the brain and an emerging field in neuroscience--offers detailed coverage of the morphology and interrelationships between glial cells and neurones in different parts of the nervous system. An accompanying website offers downloadable figures and slides.

Table of Contents

Preface xvii

About the Authors xxi

Abbreviations xxv

About the Companion Website xxxii

1 History of Neuroscience and the Dawn of Research in Neuroglia 1

1.1 The miraculous human brain: localising the brain functions 1

1.2 Cellular organisation of the brain 10

1.3 Mechanisms of communications in neural networks 14

1.3.1 Electrical/ionic nature of excitability 14

1.3.2 Chemical signalling between neural cells 26

1.4 The concept of neuroglia 27

1.5 Beginning of the modern era 47

1.6 Concluding remarks 49

References 49

2 General Overview of Signalling in the Nervous System 59

2.1 Intercellular signalling: wiring and volume modes of transmission 59

2.2 Cellular signalling: receptors 62

2.3 Intracellular signalling: second messengers 67

2.4 Calcium signalling 67

2.4.1 Cellular Ca2þ

regulation 69

2.5 Concluding remarks 72

3 Neuroglia: Definition, Classification, Evolution, Numbers,

Development 73

3.1 Definition of neuroglia as homeostatic cells of the nervous system 74

3.2 Classification 75

3.3 Evolution of neuroglia 76

3.3.1 Evolution of astrocytes 79

(i) Nematoda: neuroglia in Caenorhabditis elegans 79

(ii) Annelida: astroglia in leech 81

(iii) Arthropoda: astrocytes in Drosophila and other insects 83

(iv) Neuroglia in early Deuterostomia (Hemichordata and

Echinodermata) 85

(v) Neuroglia in low vertebrates 85

(vi) Glial advance in higher vertebrates 86

3.3.2 Evolution of myelination 89

3.3.3 Evolution of microglia 92

3.4 Numbers: how many glial cells are in the brain? 93

3.5 Embryogenesis and development of neuroglia in mammals 96

3.5.1 Macroglial cells 96

3.5.2 Astroglial cells are brain stem cells 98

3.5.3 Peripheral glia and schwann cell lineage 98

3.5.4 Microglial cell lineage 99

3.6 Concluding remarks 99

References 100

4 Astroglia 105

4.1 Definition and heterogeneity 107

4.2 Morphology of the main types of astroglia 113

4.3 How to identify astrocytes in the nervous tissue 119

4.4 Astroglial syncytial networks 120

4.4.1 Gap junctions, connexons and connexins 120

4.4.2 Astroglial networks 123

4.5 Physiology of astroglia 125

4.5.1 Membrane potential and ion distribution 125

4.5.2 Ion channels 126

(i) Potassium channels 126

(ii) Voltage-operated sodium channels (Nav) 130

(iii) Calcium channels 131

(iv) Transient receptor potential or TRP channels 131

(v) Anion/chloride channels 131

(vi) Aquaporins 132

4.5.3 Receptors to neurotransmitters and neuromodulators 133

(i) Glutamate receptors 137

(ii) Purinoceptors 141

(iii) g-aminobutiric acid receptors (GABA) receptors 146

(iv) Glycine receptors 148

(v) Acetylcholine receptors 148

(vi) Adrenergic receptors 148

(vii) Serotonin receptors 149

(viii) Histamine receptors 149

(ix) Cannabinoid receptors 149

(x) Neuropeptide receptors 149

(xi) Cytokine and chemokine receptors 150

(xii) Complement receptors 151

(xiii) Platelet-activating factor receptors 151

(xiv) Thrombin receptors 151

(xv) Ephrin receptors 151

(xvi) Succinate receptors 152

4.5.4 Astroglial membrane transporters 152

(i) ATP-dependent transporters 153

(ii) Secondary transporters 154

4.5.5 Calcium signalling in astroglia 156

(i) Endoplasmic reticulum provides for Ca2+ excitability

of astrocytes 156

(ii) Store-operated Ca2þ

entry in astrocytes 158

(iii) Ionotropic Ca2þ

permeable receptors in astrocytes 158

(iv) Sodium/calcium exchanger in astroglial Ca2þ

signalling 159

(v) Mitochondria in astroglial Ca2þ

signalling 159

(vi) Calcium waves in astrocytes 159

4.5.6 Sodium signalling in astrocytes 164

4.5.7 Release of neurotransmitters and neuromodulators from astroglia 165

(i) Exocytotic release of neurotransmitters from astrocytes 167

(ii) Diffusional release of neurotransmitters from astrocytes 172

(iii) Transporter-mediated neurotransmitter release from

astrocytes 173

(iv) Astrocytes as a main source of adenosine in the CNS 174

(v) Physiological role of astroglial release

of neurotransmitters 174

4.6 Functions of astroglia 175

4.6.1 Developmental function: neurogenesis and gliogenesis 176

(i) Embryonic neurogenesis and gliogenesis 176

(ii) Neurogenesis and gliogenesis in the adult brain 178

4.6.2 Neuronal guidance 179

4.6.3 Regulation of synaptogenesis and control of synaptic

maintenance and elimination 182

4.6.4 Structural function: astrocytes define the micro-architecture

of the grey matter and create neurovascular units 185

4.6.5 Structural function: astrocytes and the brain-blood barrier 186

4.6.6 Astrocytes regulate brain microcirculation 192

4.6.7 Brain energetics and neuronal metabolic support 195

4.6.8 Astroglia and neuroimaging 200

4.6.9 Ion homeostasis in the extracellular space 200

(i) Astrocytes and extracellular potassium homeostasis 200

(ii) Astrocytes and chloride homeostasis 204

(iii) Astrocytes and extracellular Ca2þ

204

(iv) Astrocytes and regulation of pH 205

(v) Astrocytes and zinc homeostasis 206

4.6.10 Astrocytes and homeostasis of reactive oxygen species 206

4.6.11 Water homeostasis and regulation of the extracellular

space volume 207

(i) Regulation of water homeostasis 207

(ii) Regulatory volume decrease in astrocytes 208

(iii) Redistribution of water during neuronal activity and

dynamic regulation of the extracellular space 208

4.6.12 Neurotransmitters homeostasis 209

(i) Astroglia control glutamate homeostasis and

glutamatergic transmission in the CNS 209

(ii) Astroglia and GABA-ergic transmission 213

(iii) Astroglia and adenosine homeostasis 214

4.6.13 Astroglia in synaptic transmission 215

(i) The astroglial synaptic compartment: concept of

the tripartite synapse 215

(ii) The astroglial synaptic compartment: concept of

the astroglial cradle 219

(iii) Morphological plasticity of the astroglial synaptic

compartment 221

(iv) What is the role of astroglia in regulation of synaptic

transmission? 222

4.6.14 Astroglia and central chemoception of pH and CO2 224

4.6.15 Astrocytes in regulation of systemic sodium homeostasis 224

4.6.16 Astroglia and glucose sensing 225

4.6.17 Astroglia and circadian rhythms 226

4.6.18 Astroglia and sleep 227

4.6.19 Astroglia and control of reproduction 228

4.6.20 M€uller glial cells as light guides in retina 228

4.6.21 Astroglia in ageing 228

4.6.22 Astrocytes as a cellular substrate of memory and

consciousness? 230

4.7 Concluding remarks 231

References 231

5 Oligodendrocytes 245

5.1 Oligodendrocyte anatomy 247

5.1.1 The generalised structure of a myelinating oligodendrocyte 247

5.1.2 Subtypes of myelinating oligodendrocytes 249

5.1.3 Non-myelinating oligodendrocytes 251

5.2 Myelin structure and function 252

5.2.1 Myelin and saltatory conduction 252

5.2.2 Oligodendrocyte-axon interactions and nodes of Ranvier 256

5.2.3 Myelin structure and metabolism 257

5.2.4 Myelin biochemistry 258

(i) Lipids 258

(ii) Proteins 261

5.2.5 Myelin transport 265

5.3 Physiology of oligodendrocytes 266

5.3.1 Voltage-operated ion channels 270

(i) Outwardly rectifying potassium channels 270

(ii) Inward rectifier potassium channels (Kir) 271

(iii) Voltage-operated sodium channels (Nav) 272

(iv) Voltage-operated calcium channels (VOCC, Cav) 272

(v) Chloride and acid-sensing ion channels (ASIC) 273

5.3.2 Glutamate receptors 273

(i) Ionotropic glutamate receptors (iGluRs) 273

(ii) Metabotropic glutamate receptors (mGluRs) 275

5.3.3 Purinergic receptors 275

(i) P1 purinergic receptors 275

(ii) P2X receptors 276

(iii) P2Y receptors 276

5.3.4 GABA receptors 277

5.3.5 Other neurotransmitter receptors 278

5.3.6 Transporters and exchangers 279

5.3.7 Gap junctions 279

5.3.8 Intracellular calcium 280

5.4 Oligodendrocyte development 283

5.4.1 Developmental origins of oligodendrocytes 284

5.4.2 Stages of oligodendrocyte differentiation 284

5.4.3 Trophic factors and oligodendrocyte differentiation 286

5.4.4 Regulation of oligodendrocyte differentiation 288

5.4.5 Axoglial interactions regulating oligodendrocyte

differentiation and myelination 293

5.4.6 Downstream signalling cascades that regulate

oligodendrocyte differentiation and myelination 297

5.5 Concluding remarks 299

References 299

6 NG2-glial Cells 321

6.1 Definition of NG2-glia 321

6.2 Structure of NG2-glia 324

6.2.1 Identification 324

6.2.2 Morphology and distribution 325

6.2.3 Relationship of NG2-glia with neuroglial domains 326

6.2.4 NG2-glia and synapses 326

6.3 Physiology of NG2-glia 327

6.3.1 Membrane properties 328

6.3.2 Gap junctional coupling 328

6.3.3 Voltage-operated ion channels 330

6.3.4 Neurotransmitter receptors 330

6.3.5 Neurone-NG2-glial cell signalling at synapses 331

6.4 Proliferation of NG2-glia and generation of oligodendrocytes 332

6.4.1 Normal adult brain 332

6.4.2 Are NG2-glia multipotent stem cells? 333

6.4.3 Response of NG2-glia to injury and demyelination 333

6.5 Relationship between NG2-glia and CNS pericytes 333

6.5.1 Identification of pericytes 333

6.5.2 Developmental origin of pericytes 334

6.5.3 Pericytes are multipotent stem cells in the adult brain 336

6.6 Evolution of NG2-glia 336

6.7 Concluding remarks 337

References 337

7 Microglia 343

7.1 Definition of microglia 344

7.2 Microglial origin and development 345

7.3 Morphology of microglia 345

7.3.1 Morphology in the healthy tissue: resting or survelliant phenotype 345

7.3.2 Morphology in pathological tissue: activated phenotype 349

7.3.3 Morphology in the dish 350

7.3.4 Identification of microglial cells in neural tissues 351

7.4 General physiology of microglia 351

7.4.1 Membrane potential and ion distribution 351

7.4.2 Ion channels in microglia 352

(i) Sodium channels 352

(ii) Calcium-permeable channels 352

(iii) Potassium channels 357

(iv) Anion channels 358

(v) Proton channels 358

7.4.3 Calcium signalling in microglia 358

7.4.4 Neurotransmitter receptors 360

(i) Purinoceptors 360

(ii) Glutamate receptors 363

(iii) GABA receptors 363

(iv) Acetylcholine receptors 364

(v) Adrenergic receptors 364

(vi) Dopamine receptors 364

(vii) Serotonin receptors 364

7.4.5 Receptors for neurohormones and neuromodulators 364

7.4.6 Cytokines and chemokines receptors 367

7.4.7 Pattern-recognition receptors 369

7.4.8 Other receptor systems 370

7.4.9 Microglial plasmalemmal transporters 371

7.5 Microglial migration and motility 372

7.6 Physiological functions of microglia: role in synaptic transmission

and plasticity 373

7.7 Microglia in ageing 375

7.8 Concluding remarks 375

References 376

8 Peripheral Glial Cells 381

8.1 Peripheral nervous system 382

8.1.1 Basic structure 382

8.1.2 Development 385

8.1.3 The CNS-PNS interface 387

(i) Structure of the CNS-PNS interface 387

(ii) Development of the CNS-PNS interface 389

(iii) CNS-PNS interface in degeneration and regeneration 390

8.2 Schwann cells 390

8.2.1 Schwann cell subtypes 391

8.2.2 Development of Schwann cells 394

(i) Stages of Schwann cell differentiation 394

(ii) Regulation of Schwann cell differentiation 396

(iii) Control of myelination 397

8.2.3 Axoglial interactions and myelination 397

(i) The Schwann cell basal lamina 399

(ii) Organisation of nodes of Ranvier in the PNS 399

(iii) Schwann cell perinodal microvilli 399

8.2.4 PNS myelin structure and biochemistry 400

(i) Lipids 401

(ii) Proteins 401

8.2.5 Physiology of Schwann Cells 403

(i) Ion channels and neurotransmitter receptors 403

(ii) Ca2þ

signalling in Schwann cells 406

(iii) Schwann cells and pain 407

8.3 Satellite glial cells 407

8.3.1 Organisation of sensory and autonomic ganglia 407

8.3.2 Satellite glia in sensory and autonomic ganglia 408

8.3.3 Physiology of satellite glia 408

(i) Electrical properties 408

(ii) Homeostatic function 409

(iii) Ca2þ

signalling 409

(iv) Other receptors in SGCs 409

(v) Neurotrophic function of SGC 409

8.3.4 Injury response of satellite glia 410

8.3.5 Sensory satellite glia and pain 411

8.4 Enteric glia 412

8.4.1 Organisation of the enteric nervous system 412

8.4.2 Development of enteric glia 413

8.4.3 Structure of enteric glia 413

8.4.4 Physiology of enteric glia 415

(i) Electrical properties 415

(ii) Ion channels and neurotransmitter receptors 415

(iii) Ca2þ

signalling 416

8.4.5 Functions of EGCs 416

(i) Homeostatic functions 416

(ii) Barrier function 417

(iii) Immune functions 418

(iv) Enteric glia in intestinal diseases 418

8.5 Olfactory ensheathing cells (OECs) 418

8.5.1 Organisation and structure of OECs 418

8.5.2 Physiology of OECs 419

(i) Electrical properties 419

(ii) Ca2þ

signalling 420

8.5.3 OECs facilitate olfactory neurogenesis throughout life 420

8.5.4 OECs and regeneration 421

8.5.5 OECs and remyelination 422

8.6 Concluding remarks 422

References 423

9 General Pathophysiology of Neuroglia 431

9.1 Neurological disorders as gliopathologies 431

9.2 Reactive astrogliosis 433

9.3 Wallerian degeneration 439

9.4 Excitotoxic vulnerability of oligodendrocytes: the death of white matter 442

9.5 Activation of microglia 444

9.5.1 Pathological potential of activated microglia 449

9.6 Concluding remarks 449

References 450

10 Neuroglia in Neurological Diseases 453

10.1 Introduction 454

10.2 Genetic astrogliopathology: Alexander disease 456

10.3 Stroke and ischaemia 458

10.3.1 Glial cell death during ischaemia 460

10.3.2 Astroglia protect the brain against ischaemia 463

10.3.3 Astrocytes may exacerbate brain damage in ischaemia 465

10.3.4 Oligodendrocytes and microglia in stroke 467

10.4 Migraine and spreading depression 467

10.5 CNS oedema 469

10.5.1 Traumatic oedema 470

10.5.2 Ischaemic oedema 470

10.5.3 Oedema in hepatic encephalopathy 471

10.5.4 Hyponatremia 471

10.6 Metabolic disorders 471

10.6.1 Hepatic encephalopathy 471

10.6.2 Congenital glutamine deficiency with glutamine

synthetase mutations 472

10.6.3 Pyruvate carboxylase deficiency 472

10.6.4 Niemann-pick type C disease 473

10.6.5 Aceruloplasminemia 473

10.7 Toxic encephalopathies 473

10.7.1 Methylmercury toxic encephalopathy 473

10.7.2 Lead toxic encephalopathy 474

10.7.3 Manganese neurotoxicity 474

10.7.4 Aluminium toxic encephalopathy 474

10.8 Neurodegenerative diseases 474

10.8.1 Post-stroke dementia 475

10.8.2 Amyotrophic lateral sclerosis 477

10.8.3 Wernicke encephalopathy 479

10.8.4 Fronto-temporal, thalamic, HIV-associated and other

non-Alzheimer’s type dementias 479

10.8.5 Alzheimer’s disease (AD) 480

(i) Astrogliosis and astroglial degeneration in AD 482

(ii) Astroglia and b-amyloid 483

(iii) The neuro-vascular unit in AD: role for astrocytes 483

(iv) Metabolic remodelling of astroglia in AD 484

(v) Microglia in AD 484

10.8.6 Parkinson’s disease 485

10.8.7 Huntington’s disease 486

10.8.8 Infantile neuroaxonal dystrophy 486

10.8.9 Nasu-Hakola disease: microglial pre-senile dementia 486

10.9 Leukodystrophies 487

10.9.1 Megalencephalic leukoencephalopathy with

subcortical cysts 487

10.9.2 Vanishing white matter disease 487

10.10 Epilepsy 488

10.11 Psychiatric diseases 490

10.12 Autistic disorders 491

10.12.1 Autism 491

10.12.2 Fragile X syndrome 491

10.12.3 Rett syndrome 491

10.13 Neuropathic pain 492

10.14 Demyelinating diseases 494

10.14.1 Multiple sclerosis 494

10.14.2 Neuromyelitis optica 496

10.15 Infectious diseases 496

10.15.1 Bacterial and viral infections 496

10.15.2 Human immunodeficiency virus (HIV) infection 497

10.15.3 Human T-lymphotropic virus type-1 498

10.15.4 Human herpes virus-6 499

10.16 Peripheral neuropathies 499

10.16.1 Hereditary neuropathies 499

10.16.2 Acquired inflammatory neuropathies 500

10.16.3 Diabetic neuropathies 500

10.16.4 Leprosy 501

10.17 Gliomas 501

10.17.1 Glial complications of glioma therapy 504

10.18 Concluding remarks 504

References 504

Author Index 513

Subject Index 517



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