Preface ........................................................ XV
List of Contributors .......................................... XIX
Part I Systems
1 On the Role of Stress in Evolution ........................... 3
Lilach Hadany
1.1 Introduction ............................................ 3
1.2 Stress Through the Gene's Eye: the Evolution of
Stress-Induced Genetic Mixing ........................... 3
1.2.1 Stress-Induced Recombination ..................... 4
1.2.1.1 Classic Models of the Evolution of
Recombination ........................... 4
1.2.1.2 The Evolution of Stress-Induced
Recombination ........................... 4
1.2.1.3 Evidence for Stress-Induced
Recombination ........................... 6
1.2.2 Stress and Sex ................................... 6
1.2.3 Stress and Outcrossing ........................... 7
1.2.4 Stress and Dispersal ............................. 8
1.3 The Effect of Stress-Induced Variation on the
Evolvability of Complex Traits .......................... 9
1.4 Stress-Induced Variation and Pathogen Evolution ........ 10
1.5 Stress-Induced Mortality ............................... 10
Summary ..................................................... 12
References .................................................. 13
2 Catecholamines and Stress ................................... 19
Esther L. Sabban
2.1 Rapid Stress-Triggered Changes in Catecholamines ....... 19
2.2 Catecholamines and Stress-Related Disorders ............ 20
2.2.1 Cardiovascular Disease .......................... 20
2.2.2 Post-Traumatic Stress Disorder .................. 21
2.2.3 Depression ...................................... 21
2.2.4 Immune Disorders ................................ 21
2.2.5 Pain ............................................ 22
2.3 Stress-Triggered Regulation of Catecholamine
Biosynthetic Enzymes in Different Locations ............ 22
2.3.1 Pathway of Catecholamine Biosynthesis ........... 22
2.3.2 Adrenomedullary Hormonal System ................. 23
2.3.3 Sympathetic Nervous System ...................... 27
2.3.4 Noradrenergic Systems in the Brain .............. 28
Summary ................................................ 30
References ............................................. 30
3 Stress and the Cholinergic System ........................... 37
Marietta De Biasi
3.1 Acetylcholine and Stress ............................... 37
3.1.1 Cholinergic Innervation of the Brain ............ 38
3.1.2 Brain Cholinergic Receptors ..................... 38
3.1.3 AChR Distribution in the CNS .................... 39
3.1.4 The Septohippocampal Pathway and Stress ......... 40
3.1.5 Stress-Induced Molecular Adaptations in the
Cholinergic System
3.1.5.1 The Nicotinic Cholinergic System and
Stress ................................. 41
3.2 Contribution of Genetically Engineered Mouse Models
to the Understanding of the Role of Cholinergic
Receptors in Stress .................................... 42
Summary ................................................ 43
References ............................................. 44
Part II Cells and Circuits
4 Effects of Stress on the Function of Hippocampal Cells ...... 55
Marian Joels and Henk Karst
4.1 Introduction ........................................... 55
4.2 Non-Genomic Effects of Corticosterone .................. 57
4.3 Genomic Effects of Corticosterone ...................... 58
4.3.1 Ion Currents .................................... 59
4.3.2 Amino Acid Responses ............................ 61
4.3.3 Aminergic Responses ............................. 62
4.3.4 Implications for Hippocampal Function ........... 64
Summary ..................................................... 66
References .................................................. 66
5 Stress and Adult Neurogenesis in the Mammalian Central
Nervous System .............................................. 71
Elizabeth D. Kirby and Daniela Kaufer
5.1 Introduction ........................................... 71
5.2 Adult Neurogenesis: a Brief Primer ..................... 71
5.3 Measuring Neurogenesis: How to Find New Neurons ........ 72
5.3.1 Using DNA Replication to Detect New Cells ....... 72
5.3.2 Endogenous Markers of Cell Cycle ................ 73
5.3.3 Retroviral Labeling of New Cells ................ 73
5.3.4 Determining Cell Fate ........................... 73
5.4 Stress-Induced Alteration in Cell Proliferation ........ 74
5.4.1 Acute Stress .................................... 74
5.4.2 Chronic Stress .................................. 78
5.4.3 Cell Cycle Arrest Versus Progenitor Death ....... 78
5.5 Stress-Induced Alteration of New Cell Survival ......... 79
5.6 Stress-Induced Alteration of Cell Fate Choice .......... 79
5.7 Mechanism of Stress-Induced Changes in Adult
Neurogenesis ........................................... 81
5.7.1 Direct Effects of Glucocorticoids on Adult
Neurogenesis .................................... 82
5.7.2 Indirect Effects of Glucocorticoids on Adult
Neurogenesis .................................... 82
5.7.2.1 Excitatory Amino Acids ................. 82
5.7.2.2 Serotonin .............................. 83
5.7.2.3 Growth Factors ......................... 83
5.7.3 Intracellular Mechanisms of Glucocorticoids
Effects ......................................... 84
5.7.4 An Overall Picture of Mechanism: Putting the
Pieces Together ................................. 84
5.7.5 Function of Regulation of Adult Neurogenesis
by Stress and Glucocorticoids ................... 84
Summary ................................................ 85
References .................................................. 86
6 Individual Differences in Reactivity to Social Stress in
the Laboratory and Its Mediation by Common Genetic
Polymorphisms ............................................... 93
Idan Shalev, Elad Lerer, Salomon Israel, Fiorina
Uzefovsky, Inga Gritsenko, David Mankuta, Marsha Kaitz,
and Richard P. Ebstein
6.1 Stressors and HPA Axis Regulation: Naturalistic
Studies ................................................ 93
6.2 Trier Social Stress Test ............................... 94
6.3 Genes and Regulation of the HPA Axis ................... 95
6.4 Jerusalem Studies ...................................... 97
6.4.1 Experimental Design ............................. 97
6.4.2 Genes Associated with TSST Response ............. 98
6.5 Growth Factors ......................................... 98
6.5.1 BDNF Gene ....................................... 98
6.6 Neuropeptides ......................................... 103
6.6.1 Arginine Vasopressin la Receptor (AVPRla) ...... 103
6.7 Serotonin Transmission ................................ 104
6.7.1 Serotonin Transporter (SLC6A4) ................. 104
6.7.2 Monoamine Oxidase A (MAOA) ..................... 104
6.8 Sex Steroids .......................................... 105
6.8.1 Estrogen Receptor Alpha (ESR1) ................. 105
6.8.2 Genes Not Associated with TSST Response ........ 106
Summary .................................................... 107
References ................................................. 108
Part III Cognition and Behavior
7 Corticosteroid Hormones in Stress and Anxiety - Role of
Receptor Variants and Environmental Inputs ................. 779
Roel H. DeRijk, Efthimia Kitraki, and E. Ronald de Kloet
7.1 Introduction .......................................... 119
7.2 Corticosteroid Hormones, Fear and the Stress
Response .............................................. 121
7.2.1 Neuroanatomical Basis .......................... 121
7.2.2 Role of Corticosteroids and Other Stress
Mediators ...................................... 122
7.2.3 Cognitive and Emotional Aspects of Stress ...... 124
7.3 Gene Variants and Early Life Experiences .............. 125
7.4 MR and GR Gene Structure .............................. 126
7.5 Genetic Variants of the MR and GR Genes ............... 127
7.5.1 MR Gene ........................................ 129
7.5.2 GR Gene ........................................ 129
7.6 HPA Axis Regulation ................................... 131
7.6.1 Basal Cortisol Levels .......................... 131
7.6.2 Feedback Sensitivity ........................... 131
7.6.3 Psychosocial Challenges ........................ 132
7.7 Epigenetic Modifications Affecting GR and MR
Actions ............................................... 133
7.8 Stress-Related Psychopathology ........................ 135
7.8.1 Corticosteroid-Controlled Pathways as Targets
for New Therapeutic Interventions .............. 135
7.9 Early Experience and Psychopathology .................. 136
7.10 Genetic Variants and Psychopathology .................. 237
7.11 Concluding Remarks .................................... 140
Summary .................................................... 141
References ................................................. 142
8 Corticotropin-Releasing Factor (CRF) and CRF-Related
Peptides - a Linkage Between Stress and Anxiety ............ 757
Thomas Blank and Joachim Spiess
8.1 The CRF Family, Its Receptors and Ligands ............. 151
8.2 The Role of CRF and CRF-Related Peptides in
Neuroendocrine Aspects of Stress ...................... 153
8.3 The Role of CRF and CRF-Related Peptides in
Behavioral Aspects of Stress .......................... 154
8.4 The Role of CRF and CRF-Related Peptides in Anxiety ... 156
8.5 Crosstalk Between the CRF and Serotonin Systems ....... 158
8.6 Future Directions ..................................... 159
Summary .................................................... 159
References ................................................. 160
9 Stress, Emotion and Memory: the Good, the Bad and
the Intriguing ............................................. 767
Marie-France Marin, Tania Elaine Schramek, Françoise
S. Maheu, and Sonia J. Lupien
9.1 Introduction .......................................... 167
9.2 The Relativity of Stress .............................. 168
9.3 Important Characteristics of Stress Hormones .......... 169
9.3.1 Stress Hormone Receptors ....................... 169
9.3.2 The Impact of Stress Hormones on Cognition ..... 170
9.4 Effects of GCs on Learning and Memory ................. 171
9.4.1 Direct Effects of GCs .......................... 172
9.4.2 Modulatory Effects of GCs ...................... 173
9.4.3 Chronic Effects of GCs on Cognition and Brain
Integrity ...................................... 174
9.5 Stress, Emotion and Memory ............................ 176
9.5.1 Central Focus Phenomenon ....................... 176
9.5.2 Retention Intervals and Emotional Memory ....... 177
9.5.3 Stress Hormones and Emotional Memory ........... 178
9.5.4 Stress Hormone Effects of the Encoding and
Consolidation of Emotional Material ............ 178
9.5.5 Stress Hormone Effects of the Retrieval of
Emotional Material ............................. 179
9.6 Contextual Effects .................................... 180
9.7 New Directions ........................................ 181
9.8 Conclusion ............................................ 182
Summary .................................................... 183
References ................................................. 183
10 Contribution of Early Life Stress to Anxiety Disorder ...... 789
Maria Weinstock
10.1 Introduction .......................................... 189
10.2 Organization of the Stress Response ................... 190
10.2.1 Glucocorticoid Receptors ....................... 190
10.2.2 CRH and Its Receptors .......................... 191
10.2.3 Role of the Amygdala and CRH in Mediating
Anxiogenic Behavior ............................ 192
10.3 Alterations in Circulating Hormonal Levels Induced
by Chronic Gestational Stress ......................... 192
10.3.1 Experimental Animals ........................... 192
10.3.2 Human Subjects ................................. 193
10.4 Alterations in the Regulation of the HPA Axis as
a Result of Prenatal Stress ........................... 194
10.4.1 Experimental Animals ........................... 194
10.4.1.1 Effects on Basal HPA Axis Activity .... 194
10.4.1.2 Effect of Prenatal Stress on
the Response to Stress of the HPA
Axis .................................. 194
10.4.2 Human Subjects ................................. 195
10.4.2.1 Effects of Prenatal Stress on Basal
HPA Axis Activity ..................... 195
10.4.2.2 Effect of Prenatal Stress on the
Response to Stress of the HPA Axis .... 196
10.5 Alterations in the Regulation of the HPA Axis as
a Result of Postnatal Stress .......................... 196
10.5.1 Experimental Animals ........................... 196
10.5.2 Human Subjects ................................. 197
10.6 Association Between Prenatal Stress and Anxiety
Disorder .............................................. 197
10.6.1 Experimental Animals ........................... 197
10.6.2 Human Subjects ................................. 199
10.7 Association Between Early Life Stress and Anxiety
Disorder .............................................. 200
10.7.1 Experimental Animals ........................... 200
10.7.2 Human Subjects ................................. 202
Summary .................................................... 202
References ................................................. 203
Part IV Immune Responses
11 Stress Effects on Immunity in Vertebrates and
Invertebrates .............................................. 209
Michael Shapira
11.1 Introduction .......................................... 209
11.2 The Neuroendocrine Stress Response in Vertebrates
and Invertebrates ..................................... 210
11.2.1 Stress Responses and Hormones in
Invertebrates .................................. 211
11.3 The Immune System in Vertebrates and Invertebrates .... 212
11.3.1 Effects of Acute Stress on Immunity ............ 216
11.3.1.1 Vertebrates ........................... 216
11.3.1.2 Invertebrates ......................... 218
11.3.2 Effects of Chronic Stress ...................... 219
11.3.2.1 Vertebrates ........................... 219
11.3.2.2 Invertebrates ......................... 220
Summary and Future Prospects ............................... 221
References ................................................. 222
12 Immunity to Self Maintains Resistance to Mental Stress:
Boosting Immunity as a Complement to Psychological
Therapy .................................................... 229
Сil М. Lewitus, Osnat Schwartz-Stav, and Michal Schwartz
12.1 Introduction .......................................... 229
12.2 Stress and the Blood-Brain Barrier .................... 230
12.3 Lymphocyte Surveillance of the CNS .................... 230
12.4 Stress-Induced Lymphocyte Mobilization ................ 231
12.5 Stress Hormones and Lymphocyte Trafficking ............ 232
12.6 The Physiological Relevance of Lymphocytes in the
Brain ................................................. 233
12.7 Protective Autoimmunity ............................... 233
12.8 Behavioral Immunization ............................... 235
12.9 Immune Memory to Self Antigens Underlies Resilience
to Mental Stress ...................................... 235
Summary .................................................... 237
References ................................................. 238
13 Brain lnterleukin-1 (IL-1) Mediates Stress-Induced
Alterations in HPA Activation, Memory Functioning and
Neural Plasticity .......................................... 243
Inbal Goshen and Raz Yirmiya
13.1 Introduction .......................................... 243
13.2 The Bi-Directional Interaction Between IL-1 and the
HPA Axis .............................................. 243
13.3 Stress-Induced HPA Axis Activation is Regulated by
Brain IL-1 ............................................ 246
13.3.1 IL-1 Mediates Immunological Stress-Induced
HPA Axis Activation ............................ 246
13.3.2 Brain IL-1 Mediates Psychological Stress-
Induced HPA Axis Activation .................... 246
13.4 Stress-Induced Alterations in Memory and Neural
Plasticity are Regulated by Brain IL-1 ................ 249
13.4.1 Brain IL-1 Underlies Stress-Induced Memory
Impairment by Modulating Glucocorticoids
Secretion ...................................... 250
13.4.2 Stress-Induced IL-1 Reduces Hippocampal
Neurogenesis: Implications for Memory
Impairments .................................... 253
Summary .................................................... 254
References ................................................. 255
Part V Post-Traumatic Stress Disorder
14 Post-Traumatic Stress Disorder in Animal Models ............ 263
Hagit Cohen, Nitzan Kozlovsky, Gal Richter-Levin, and
Joseph Zohar
14.1 Introduction .......................................... 263
14.2 Animal Models of PTSD ................................. 265
14.2.1 Trauma/Stress-Based Models ..................... 265
14.2.2 Mechanism-Based Models ......................... 266
14.2.2.1 Enhanced Fear Conditioning ............ 267
14.2.2.2 Impaired Extinction ................... 267
14.2.3 Individual Differences in Response to an
Exposure to a Traumatic Experience ............. 268
14.2.4 Behavioral Assessments ......................... 268
14.2.5 Classification According to Cut-Off
Behavioral Criteria ............................ 269
14.3 Selected CBC-Based Studies ............................ 271
14.3.1 Behavioral Response Patterns Versus Time ....... 271
14.3.2 Physiological Correlates ....................... 272
14.3.3 Strain/Genetic Studies ......................... 272
14.3.3.1 HPA Axis Response in Lewis and
Fischer Rats .......................... 273
14.3.3.2 Stress-Induced Behavioral Responses
in Inbred Mouse Strains ............... 273
14.3.4 Molecular Neurobiological Correlates ........... 273
14.3.5 Drug Studies ................................... 274
14.3.5.1 Early Intervention with an SSRI
(Sertraline) .......................... 275
14.3.5.2 Early Intervention with
Corticosterone ........................ 275
14.3.5.3 Early Intervention with
Benzodiazepine (Alprazolam) ........... 276
Summary .................................................... 276
References ................................................. 277
15 The Cholinergic Model for PTSD: from Acute Stress to
PTSD, from Neuron to Network and Behavior .................. 283
Alon Friedman and Lev Pavlovsky
15.1 From Acute Stress to PTSD ............................. 283
15.2 Stress and Brain Cholinergic Pathways ................. 283
15.3 Anatomical Considerations ............................. 284
15.4 Neurophysiological Considerations ..................... 285
15.4.1 The Role of ACh in Behavior, Learning and
Memory ......................................... 288
15.5 Stress Induces Cholinergic Dysfunction ................ 288
15.5.1 Stress Event Affects Cholinergic Functions ..... 289
15.5.2 Cholinergic Dysfunction Persists After
Stress ......................................... 289
15.5.3 Cholinergic Dysfunction May Underlie Short-
and Long-Term Clinical Symptoms ................ 290
15.6 The Cholinergic Basis for PTSD: the Model ............. 290
Summary .................................................... 292
References ................................................. 292
Part VI Vulnerability to Disease
16 Stress and Neurodegeneration: Adding Insult to Injury? ..... 299
Amit Berson, Моr Наnаn, and Hermona Soreq
16.1 Abstract .............................................. 299
16.2 Alzheimer's Disease Shares Phenotypic Biomarkers
with Stress Reactions ................................. 299
16.3 Molecular Underpinnings of Stress-Induced Cognitive
Impairments ........................................... 300
16.4 Direct Evidence Linking Stress Reactions and AD ....... 303
16.5 The Cholinergic System Connects Stress Reactions and
AD .................................................... 304
16.6 Chronic Stress Affects Behavior Thorough Chromatin
Remodeling and Alternative Splicing ................... 307
16.7 GCs Regulate Neuronal Excitability by Modifying
Alternative Splicing .................................. 309
Summary .................................................... 310
References ................................................. 310
17 Stress and Neurotransmission: Clinical Evidence and
Therapeutic Implications ................................... 317
Hadar Shalev and Jonathan Cohen
17.1 Introduction .......................................... 317
17.2 Serotonin ............................................. 319
17.3 Adrenergic Antagonists ................................ 321
17.4 γ-Aminobutyric Acid ................................... 323
Summary .................................................... 325
References ................................................. 327
18 Metabolic Components of Neuroendocrine Allostatic
Responses: Implications in Lifestyle-Related Diseases ...... 331
Ronan M.C. Berg and Bente Klarlund Pedersen
18.1 Introduction .......................................... 331
18.2 Neuroendocrine Allostatic Responses and Fat
Metabolism ............................................ 332
18.2.1 Background ..................................... 332
18.2.2 Effects of GCs on Fat Metabolism and
Distribution ................................... 333
18.2.3 Effects of Catecholaminergic Signaling on
Lipolysis and Dyslipidemia ..................... 334
18.3 Neuroendocrine Allostatic Responses and Glucose
Metabolism ............................................ 335
18.3.1 Background ..................................... 335
18.3.2 The Effects of GCs on Gluconeogenesis and
Glycogen Synthesis ............................. 335
18.3.3 Effects of Catecholaminergic Signaling on
Hepatic Glucose Metabolism ..................... 337
18.3.4 Effects of GCs on Peripheral Insulin
Sensitivity .................................... 337
18.3.5 Effects of Catecholaminergic Signaling on
Peripheral Glucose Transport ................... 338
18.4 Discussion ............................................ 339
Summary .................................................... 342
References ................................................. 342
19 Environmental Stress is Not Always Vicious: a Lesson from
Heat Acclimation-Mediated Neuroprotection After Traumatic
Brain Injury ............................................... 349
Michal Horowitz and Esther Shohami
19.1 Introduction .......................................... 349
19.2 TBI Damage - Detrimental Signaling and Possible
Protective Pathways ................................... 349
19.2.1 Short Background ............................... 349
19.2.2 TBI-Induced Tissue and Cellular Stress
Response ....................................... 350
19.2.2.1 Glutamate ............................. 350
19.2.2.2 Reactive Oxygen Species ............... 351
19.2.2.3 Neuroinflammation ..................... 351
19.2.3 Treatments ..................................... 352
19.3 Preconditioning: Inherent Protective Mechanisms Are
Induced by Preceding Sub-Lethal (Multiple)
Stressors ............................................. 352
19.4 Heat Acclimation and Neuroprotection .................. 353
19.4.1 Heat Acclimation ............................... 353
19.4.2 Heat Acclimation Affords Neuroprotection
After TBI ...................................... 354
19.4.2.1 Physiological Evidence ................ 354
19.4.3 Underlying Signaling Pathways .................. 356
19.4.3.1 Oxidative Stress ...................... 356
19.4.3.2 HIF-1-Erythropoietin Pathway .......... 356
19.4.3.3 Acute Inflammatory Response ........... 357
Summary .................................................... 359
References ................................................. 359
Index ......................................................... 365
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