Post on 23-Apr-2022
INTRODUCINGBIOLOGICAL RHYTHMS
A Primer on the Temporal Organization of Life,with Implications for Health, Society, Reproduction
and the Natural Environment
Willard L. Koukkari and Robert B. SothernCollege of Biological Sciences
University of MinnesotaSt. Paul, MN, USA
4y Springer
Contents
Preface viiAcknowledgments ix
1 The Study of Biological RhythmsIntroduction 1A Time for Everything 4Three Rhythm Domains 6Implications of Body Clocks 11
Essay 1.1: Accidents and Catastrophes 11Chronobiology: An Integrating Discipline 12Chapters in This Book 14Take-Home Message 15
2 General Features of Rhythms: Terminology and CharacteristicsIntroduction 19Details of a Rhythm 20
Period and Frequency 21Orcadian Domain 21Essay 2.1: Use of the term "Orcadian" 23Other Domains 25Ev.vav 2.2: Time on Earth as We Know It 26Amplitude 29Phase 29
Genetics and Inheritance 29Period 30Amplitude and Phase 30
Primary Orcadian Clocks 32The Brain 33SCN Identification 34SCN as Synchronizer 36Retinal Melanopsin 36Retinohypothalamic Tract 37
XI
xii Contents
Characteristics of the Period 38Stability and Free-Running 38Frequency Multiplication/Demultiplication 39Light Quali ty 40Temperature 41Chemicals 42
Characteristics of the Amplitude 42Damping 43
Characteristics of the Phase 44Synchronizers 44Phase-Shifting by Light 44Phase Shifts by Chemicals or Temperature 45Phase-Response Curve 46
Masking 47Need for a Cyclic Environment 48In Darwin's Footsteps 49
Exercise 2.1: An Ultradian Experiment: 52Circumnutation
Exercise 2.2: A Orcadian Experiment: 54Leaf Movements
Mechanisms 56Take-Home Message 56
3 Physical a n d Biological TimeIntroduction 66Rotations a n d Revolutions of the Earth and Moon 67
The Day 68The Year 69The Month 71
Clocks and Calendars: Ancient Times 72The Month and Year 72The Day 77The Week 78
Clocks and Calendars: Middle Ages to Now 79Longitude and Clocks 8 0
Springs to Atoms 8 2
Essay 3.1; Wristwatches 83Time Zones 8
Daylight Saving Time 85Recording Date and Time 8 5
Recording Biological Time 86The 24-h Biological Clock Concept 8 7
Early Studies 8 7
Considerations 8^
Contents xiii
Ultradian and Infradian Clocks 89Endogenous vs. Exogenous 90Comparison with Manufactured Clocks 90
Evolution of the Clock 92Molecular Building Blocks 93Geological History and Rhythmic Components 95Adaptation to Avoid Harmful Light 99Ancestral Traits and Convergent Evolution 101
Take-Home Message 102
4 PhotoperiodismIntroduction 107Daylight and Seasons 107Photoperiodism: The ProcessResponse Types 113
Critical DaylengthDiversity of Responses.
Early StudiesLatitude.
s.Light and Photoreceptive RegionsExtraretinal Photoreceptors .PigmentsCryptochromesSpectra
Rhvthmic Association.Endogenous Oscillators..Biinning's HypothesisPhase-Response Curves .Ultradian CyclesCircannual CyclesBird MigrationDeer AntlersVernalization
Photoperiodism and HumansBirth PatternsIndoor vs. Outdoor Light.Disorders
Take-Home Message
14151616171819
2123232525252626282829293030•p
5 Biological Oscillators and Timers: Models and MechanismsIntroduction 13SApproaches to Models and Mechanisms 139Mechanical Models 141
Pendulum 142
xiv Contents
Hourglass 142External Coincidence 143Hands of a Clock 144
Mathematical Models 145Differential Equations 148Limit Cycles and Topography 149Chaos 153Spatiotemporal Systems 153
Biochemical and Metabolic Models 155Chemical Systems 155Biochemical Systems 155Essay 5.1: Selected Biochemical Notes 156Glycolytic Oscillations 157Nucleotides and Enzymes 158
Membrane Models 158Essay 5.2: Membranes and the Phospholipid Bilayer 159Lipids and Proteins 163Transport and Feedback 165
Molecular Models 167Essay 5.3: From Genes to Proteins and Mutants 167Genes and Nomenclature 171Clock Mutations 173Circadian: System and Clock 173Transcription/Translation Feedback Loops 174Light 177Temperature 178Five Circadian Clocks 179Neurospora Circadian Clock 179Advantages 179Overt Rhythms 179Genetic Highlights 181Feedback Loops and Components 181Drosophila Circadian Clock 181Advantages 182Overt Rhythms 183Genetic Highlights 183Feedback Loops and Components 183Mammalian Circadian Clock 184Advantages 184Overt Rhythms 184Genetic Highlights 184Feedback Loops and Components 186Arabidopsis Circadian Clock 187Advantages 187Overt Rhythms 187
Contents xv
Genetic Highlights 187Feedback Loops and Components 187Cyanobacteria Circadian Clock 189Advantages 190Overt Rhythms 191Genetic Highlights 191Feedback Loops and Components 191Models in Perspective 192Generalized Schematic Model for 192
Biological RhythmsTake-Home Message 193
6 Tidal and Lunar RhythmsIntroduction 207Moon and Light 208Moon and Tides 210
High and Low Tides 210Spring and Neap Tides 214Earth Tide 214Essay 6.1: Earth Tides 214
Marine Organisms 216Circatidal Rhythms 216Crab Activity 217Circadian vs. Circatidal 218Other Organisms 219Reproduction 220Color Change 224
Terrestrial Organisms 226The Menstrual Cycle 227Atmospheric Tides 227Insects 228
Plants 228Lunar/Tidal Clock Hypotheses 229
Circadian vs. Circalunidian 229Interacting Oscillators'.' 230
Take-Home Message 231
7 Sexuality and ReproductionIntroduction 237
Essay 7.1: Parasexuality 238Nuclear Division and Genetics 240
Es.sav 7.2: Mitosis. Meiosis. and 240the Punnett Square
Sex and Reproduction: The Difference 250
xvi Contents
Essay 7.3: Artificial Hybridization and 251How Sex Produces both "Lunch"and an Embryo
Asexual Reproduction 255Essay 7.4: An Abbreviated Life History of 256
Neurospora crassaCourtship and Mating 257
Photoperiodism and Sexuality 259Diet 259Flowers 260
Rhythmic Phases of Sexual Behavior in Humans 261Activity 261Disease 261
The Menstrual Cycle 262Duration and Phase 262Essay 7.5: Brief Physiology of Menstrual Cycle 262
Events and PhasesSocial Synchronization 268Sexual Activity and Birth Control 268
Primary and Secondary Sex-Related Rhythms in Men 272Ultradian and Circadian Cycles 272Infradian Cycles 273
Essay 7.6: 17th Century Notes of Monthly 273Rhythms in Males
Body Weight 274Grip Strength 274Cutaneous Pain 277Hormones 277Essay 7.7: More About Infradians 277
in Male HormonesEmotions 278Facial Sebum 279Beard Growth and Body Hair 280Sexual Activity 282
Take-Home Message 284
8 Natural Resources and AgricultureIntroduction 293Photoperiodism 294
Essay S.I: Photoperiodism as a Basic 294Principle of Biology and Its Applications
Thermoperiodism and Temperature Cycles 297Vernalization 298Temperature Compensation 300
Migration 300
Contents xvii
Birds 301Butterflies 302
Pest Management and Agents of Stress 304Herbicides 304Pest Control 306Plant Responses to Injury 307Plant Diseases 308
Production of Produce 309Fisheries and Aquaculture 310Weather Patterns and Agriculture 313
Gardens 313Outdoor Hobbies 317
Birding 317Fishing 318Essay 8.2: Fly-fishing for Trout 318
Rural and Urban Development 322Telemetry Tracking Systems 324Muskrats 324Squirrels and Foxes 324Hare 324Ruffed Grouse 326The Outdoor Laboratory 327Temporal Agroecosystems 327
Light Pollution 328Aquatic Animals 328Vertical Migration 329Drift 330Trout 330Turtles 331Insects 331Birds 332
Take-Home Message 332
9 Veterinary MedicineIntroduction 341Body Temperature and Activity 342
Diurnal vs. Nocturnal 343Timing of Food 344Masking 346Environment 346Cattle USDogs and Cats 348Poultry 349
Hematology and Urology 349Sampling Blood 349
xviii Contents
Multiple Rhythms 349Peak Times 351Collecting Urine 351Excretion Rates 353Urinary Rhythms 354Interpreting a Sample 354
A Primary Circadian Oscillator 355The Suprachiasmatic Nucleus 356The Pineal Gland 356
Diseases, Pests, and Stress 357Parasites 357Bacterial Infections 358Seasonal Diseases 358Flies 359Fleas 360
Reproduction and Photoperiodism 360Photoperiod 361Melatonin 361Domestic Fowl 362Sheep 363Horses 364Pigs and Goats 365Artificial Insemination 365Semen Quality and Season 365
Implications 366Take-Home Message 366
10 SocietyIntroduction 376Past and Present 377
The Natural Day vs. 24/7 377Time Schedules 378
Social Synchronization 380Essay 10.1: Social Synchrony in 380
AnimalsCircadian Events 381Ultradian and Infradian Events 384
Aggression and Violence 384Night and Shiftwork 385
Time for Sleep 385Problems with Shiftwork 386Adjusting to Shiftwork 387
The Global Workplace 388Communication 388Work Schedules and Outsourcing 389
Contents xix
Sports and Performance 389Body Temperature and Performance Variables 391A Time to Train or Win 391Jet Lag and Professional Sports 392Allowing for Jet Lag 392
Travel on the Earth's Surface 393Driver Fatigue and Vehicle Accidents 394Alcohol, Driving, and Fatigue 395The Post-Lunch Dip 396Animal Activity and Vehicle Accidents 396
Travel Above the Earth's Surface 396Jet Lag 397Life in Space 398
Travel Beneath the Seas 399Mealtimes and Health 401
Essay 10.2: Preclinical Meal-Timing Studies 402Changes in Body Weight and Rhythms 404Food: What, How Much, and When 408
Light Pollution 408Effects on Melatonin Production 409Effects on Clinical Health 410Better Lighting Practices 411
Pseudoscience: Birthdate-Based Biorhythms 412Essax 10.3: Development of the Biorhythm 412
"Theory"Lack of Scientific Support 413Rigidity vs. Elasticity of Infradian Periods 414An Oversimplification of Rhythms 414
Take-Home Message 415
11 Clinical MedicineIntroduction 426Circadian Rhythms in Health 437
Essay 1 I.I: Adjusting Urinary Concentrations 438for Volume and Time
Overview of Rhythms in Body Systems 430What and When is Normal? 433Time-Specified Normal Limits 438
Circadian Rhythms in Symptoms and Disease 441Birth and Death 442Cardiovascular Disease 443
Circannual Rhythms in Health 445Orcannual Rhythms in Symptoms and Disease 446
Cardiovascular Disease 448The Coagulation System 449
xx Contents
Cholesterol 450Respiratory Illness 452Mental Disorders 452Seasonal Affective Disorder 452
The Menstrual Cycle 453Disorders 454Medical Procedures 455Male Cycles 455
Melatonin and Human Health 456Darkness and Melatonin 456Sexuality 458Immune Function 459Light, Melatonin and Cancer 460Light Leaks at Night 461
When to Sample? 462Diagnosing Normal Levels 463Diagnosing Infectious Agents 464Diagnosing Abnormal Levels 467Using Rhythm Characteristics in Diagnosis 467
Hours of Changing Resistance 470Early Pre-Clinical Findings 471Time-Related Responses to 471
Anti-Cancer DrugsStage of Rhythm vs. Time of Day 473Varying Positive or Negative Effects 473
Timing Treatment: Chronotherapy 475Three Times a Day? 476Constant Dosing 476Rhythm-Dependent Effects of Some Drugs 478Administering Chronotherapy 478
Examples of Applied Chronotherapy 480Asthma 481Cancer-Animal Studies 482Cancer-Human Trials 484
Cellular Clocks and Chronotherapy 486Time-Indicating Genes 487Molecular Machinery Underlies Physiology 487
Marker Rhythms 488The Medical Community and the Concept of Timing 489Take-Home Message 491
12 AutorhythmometryIntroduction 526Measuring Your Own Body Rhythms 527
Schoolchildren 528
Contents xxi
Adults 529Performance 529Ultradian Rhythms 529Self-help Health Care 529Monitoring Symptoms 530
Body Temperature 530Internal Marker Rhythm 531Measurement Site 532What and When is Normal? 533
Blood Pressure 534Monitoring Hypertension 534Ambulatory Monitoring 535
Morningness-Eveningness 535Questionnaires 535Morningness vs. Life Factors 536Endogenous Disposition 536Body Temperature Phase 536Cognitive Tasks 537
When and How Long to Measure? 537Self-Measurements During Travel 537Self-Measurements During Isolation 540Essay 12.1: Self-Measurements in 540
"Aschoff's Bunker"Long Self-Measurement Series 544
What can be Self-Measured? 545Equipment 545
Internal or External Body Temperature 546Temperature Devices 547Blood Pressure Devices 550Automatic Devices 550Other Equipment 552Saliva, Urine, and Blood 555
Procedures for Self-Measurements 555Keeping Records 555Sampling Sequence 555
Looking at the Data 556Making Graphs 556Testing for a Time-Effect 557
Take-Home Message 558Appendix
Item I: Sample sheet for recording oral 568temperature and other functions
Item 2: Sample sheet for recording and 569graphing temperature, pulse andblood pressure
xxii Contents
Item 3: Detailed instructions for performing 570self-measurements
Item 4: Random Number Adding Speed Test 572sample pages
Item 5: Random Number Memory Test 574sample pages
13 Chronobiometry: Analyzing for RhythmsIntroduction 577Data Collection 578
Number of Timepoints 579How Long? 580Sampling Often Enough 580Aliasing 580Decision Making 580
Data Preparation 582Graphs and Visual Inspection 582Editing or Transforming Data 583Essay 13.1: Standard Deviation and Error 584Normalizing Data 584Partitioning Data Spans 585
Statistical Detection of Time Effects 586Using Two Timepoints 586Using Three or More Timepoints 586
Statistical Detection of Rhythms 588Analyzing Time-Series by Standard Methods 588Limitations of Standard Methods 588Analyzing Time-Series by Curve Fitting 589The Least-Squares Technique 590The Best-Fitting Curve 591Statistical Significance 592Complex Waveforms 592Rhythm Parameter Comparisons 593Lack of Rhythm Detection 593
Descriptive Rhythm Parameters 595The Cosinor Illustrated 596
Example of a Cosinor Program 596Take-Home Message 600
Author Index 603Subject Index 649