Contents. Acknowledgments

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1 Contents Preface Acknowledgments vii ix 1 The Study of Biological Rhythms Introduction... 1 A Time for Everything... 4 Three Rhythm Domains... 6 Implications of Body Clocks Essay 1.1: Accidents and Catastrophes Chronobiology: An Integrating Discipline Chapters in This Book Take-Home Message General Features of Rhythms: Terminology and Characteristics Introduction Details of a Rhythm Period and Frequency Circadian Domain Essay 2.1: Use of the term Circadian Other Domains Essay 2.2: Time on Earth as We Know It Amplitude Phase Genetics and Inheritance Period Amplitude and Phase Primary Circadian Clocks The Brain SCN Identification SCN as Synchronizer Retinal Melanopsin Retinohypothalamic Tract xi

2 xii Contents Characteristics of the Period Stability and Free-Running Frequency Multiplication/Demultiplication Light Quality Temperature Chemicals Characteristics of the Amplitude Damping Characteristics of the Phase Synchronizers Phase-Shifting by Light Phase Shifts by Chemicals or Temperature Phase Response Curve Masking Need for a Cyclic Environment In Darwin s Footsteps Exercise 2.1: An Ultradian Experiment: Circumnutation Exercise 2.2: A Circadian Experiment: Leaf Movements Mechanisms Take-Home Message Physical and Biological Time Introduction Rotations and Revolutions of the Earth and Moon The Day The Year The Month Clocks and Calendars: Ancient Times The Month and Year The Day The Week Clocks and Calendars: Middle Ages to Now Longitude and Clocks Springs to Atoms Essay 3.1: Wristwatches Time Zones Daylight Saving Time Recording Date and Time Recording Biological Time The 24-h Biological Clock Concept Early Studies Considerations... 89

3 Contents xiii Ultradian and Infradian Clocks Endogenous vs. Exogenous Comparison with Manufactured Clocks Evolution of the Clock Molecular Building Blocks Geological History and Rhythmic Components Adaptation to Avoid Harmful Light Ancestral Traits and Convergent Evolution Take-Home Message Photoperiodism Introduction Daylight and Seasons Photoperiodism: The Process Response Types Critical Daylength Diversity of Responses Early Studies Latitude Light and Photoreceptive Regions Extraretinal Photoreceptors Pigments Cryptochromes Spectra Rhythmic Association Endogenous Oscillators Bünning s Hypothesis Phase Response Curves Ultradian Cycles Circannual Cycles Bird Migration Deer Antlers Vernalization Photoperiodism and Humans Birth Patterns Indoor vs. Outdoor Light Disorders Take-Home Message Biological Oscillators and Timers: Models and Mechanisms Introduction Approaches to Models and Mechanisms Mechanical Models Pendulum

4 xiv Contents Hourglass External Coincidence Hands of a Clock Mathematical Models Differential Equations Limit Cycles and Topography Chaos Spatiotemporal Systems Biochemical and Metabolic Models Chemical Systems Biochemical Systems Essay 5.1: Selected Biochemical Notes Glycolytic Oscillations Nucleotides and Enzymes Membrane Models Essay 5.2: Membranes and the Phospholipid Bilayer Lipids and Proteins Transport and Feedback Molecular Models Essay 5.3: From Genes to Proteins and Mutants Genes and Nomenclature Clock Mutations Circadian: System and Clock Transcription/Translation Feedback Loops Light Temperature Five Circadian Clocks Neurospora Circadian Clock Advantages Overt Rhythms Genetic Highlights Feedback Loops and Components Drosophila Circadian Clock Advantages Overt Rhythms Genetic Highlights Feedback Loops and Components Mammalian Circadian Clock Advantages Overt Rhythms Genetic Highlights Feedback Loops and Components Arabidopsis Circadian Clock Advantages Overt Rhythms...187

5 Contents xv Genetic Highlights Feedback Loops and Components Cyanobacteria Circadian Clock Advantages Overt Rhythms Genetic Highlights Feedback Loops and Components Models in Perspective Generalized Schematic Model for Biological Rhythms Take-Home Message Tidal and Lunar Rhythms Introduction Moon and Light Moon and Tides High and Low Tides Spring and Neap Tides Earth Tide Essay 6.1: Earth Tides Marine Organisms Circatidal Rhythms Crab Activity Circadian vs. Circatidal Other Organisms Reproduction Color Change Terrestrial Organisms The Menstrual Cycle Atmospheric Tides Insects Plants Lunar/Tidal Clock Hypotheses Circadian vs. Circalunidian Interacting Oscillators? Take-Home Message Sexuality and Reproduction Introduction Essay 7.1: Parasexuality Nuclear Division and Genetics Essay 7.2: Mitosis, Meiosis, and the Punnett Square Sex and Reproduction: The Difference

6 xvi Contents Essay 7.3: Artificial Hybridization and How Sex Produces both Lunch and an Embryo Asexual Reproduction Essay 7.4: An Abbreviated Life History of Neurospora crassa Courtship and Mating Photoperiodism and Sexuality Diet Flowers Rhythmic Phases of Sexual Behavior in Humans Activity Disease The Menstrual Cycle Duration and Phase Essay 7.5: Brief Physiology of Menstrual Cycle Events and Phases Social Synchronization Sexual Activity and Birth Control Primary and Secondary Sex-Related Rhythms in Men Ultradian and Circadian Cycles Infradian Cycles Essay 7.6: 17th Century Notes of Monthly Rhythms in Males Body Weight Grip Strength Cutaneous Pain Hormones Essay 7.7: More About Infradians in Male Hormones Emotions Facial Sebum Beard Growth and Body Hair Sexual Activity Take-Home Message Natural Resources and Agriculture Introduction Photoperiodism Essay 8.1: Photoperiodism as a Basic Principle of Biology and Its Applications Thermoperiodism and Temperature Cycles Vernalization Temperature Compensation Migration

7 Contents xvii Birds Butterflies Pest Management and Agents of Stress Herbicides Pest Control Plant Responses to Injury Plant Diseases Production of Produce Fisheries and Aquaculture Weather Patterns and Agriculture Gardens Outdoor Hobbies Birding Fishing Essay 8.2: Fly-fishing for Trout Rural and Urban Development Telemetry Tracking Systems Muskrats Squirrels and Foxes Hare Ruffed Grouse The Outdoor Laboratory Temporal Agroecosystems Light Pollution Aquatic Animals Vertical Migration Drift Trout Turtles Insects Birds Take-Home Message Veterinary Medicine Introduction Body Temperature and Activity Diurnal vs. Nocturnal Timing of Food Masking Environment Cattle Dogs and Cats Poultry Hematology and Urology Sampling Blood

8 xviii Contents Multiple Rhythms Peak Times Collecting Urine Excretion Rates Urinary Rhythms Interpreting a Sample A Primary Circadian Oscillator The Suprachiasmatic Nucleus The Pineal Gland Diseases, Pests, and Stress Parasites Bacterial Infections Seasonal Diseases Flies Fleas Reproduction and Photoperiodism Photoperiod Melatonin Domestic Fowl Sheep Horses Pigs and Goats Artificial Insemination Semen Quality and Season Implications Take-Home Message Society Introduction Past and Present The Natural Day vs. 24/ Time Schedules Social Synchronization Essay 10.1: Social Synchrony in Animals Circadian Events Ultradian and Infradian Events Aggression and Violence Night and Shiftwork Time for Sleep Problems with Shiftwork Adjusting to Shiftwork The Global Workplace Communication Work Schedules and Outsourcing

9 Contents xix Sports and Performance Body Temperature and Performance Variables A Time to Train or Win Jet Lag and Professional Sports Allowing for Jet Lag Travel on the Earth s Surface Driver Fatigue and Vehicle Accidents Alcohol, Driving, and Fatigue The Post-Lunch Dip Animal Activity and Vehicle Accidents Travel Above the Earth s Surface Jet Lag Life in Space Travel Beneath the Seas Mealtimes and Health Essay 10.2: Preclinical Meal-Timing Studies Changes in Body Weight and Rhythms Food: What, How Much, and When Light Pollution Effects on Melatonin Production Effects on Clinical Health Better Lighting Practices Pseudoscience: Birthdate-Based Biorhythms Essay 10.3: Development of the Biorhythm Theory Lack of Scientific Support Rigidity vs. Elasticity of Infradian Periods An Oversimplification of Rhythms Take-Home Message Clinical Medicine Introduction Circadian Rhythms in Health Essay 11.1: Adjusting Urinary Concentrations for Volume and Time Overview of Rhythms in Body Systems What and When is Normal? Time-Specified Normal Limits Circadian Rhythms in Symptoms and Disease Birth and Death Cardiovascular Disease Circannual Rhythms in Health Circannual Rhythms in Symptoms and Disease Cardiovascular Disease The Coagulation System

10 xx Contents Cholesterol Respiratory Illness Mental Disorders Seasonal Affective Disorder The Menstrual Cycle Disorders Medical Procedures Male Cycles Melatonin and Human Health Darkness and Melatonin Sexuality Immune Function Light, Melatonin and Cancer Light Leaks at Night When to Sample? Diagnosing Normal Levels Diagnosing Infectious Agents Diagnosing Abnormal Levels Using Rhythm Characteristics in Diagnosis Hours of Changing Resistance Early Pre-Clinical Findings Time-Related Responses to Anti-Cancer Drugs Stage of Rhythm vs. Time of Day Varying Positive or Negative Effects Timing Treatment: Chronotherapy Three Times a Day? Constant Dosing Rhythm-Dependent Effects of Some Drugs Administering Chronotherapy Examples of Applied Chronotherapy Asthma Cancer Animal Studies Cancer Human Trials Cellular Clocks and Chronotherapy Time-Indicating Genes Molecular Machinery Underlies Physiology Marker Rhythms The Medical Community and the Concept of Timing Take-Home Message Autorhythmometry Introduction Measuring Your Own Body Rhythms School Children

11 Contents xxi Adults Performance Ultradian Rhythms Self-help Health Care Monitoring Symptoms Body Temperature Internal Marker Rhythm Measurement Site What and When is Normal? Blood Pressure Monitoring Hypertension Ambulatory Monitoring Morningness Eveningness Questionnaires Morningness vs. Life Factors Endogenous Disposition Body Temperature Phase Cognitive Tasks When and How Long to Measure? Self-Measurements During Travel Self-Measurements During Isolation Essay 12.1: Self-Measurements in Aschoff s Bunker Long Self-Measurement Series What can be Self-Measured? Equipment Internal or External Body Temperature Temperature Devices Blood Pressure Devices Automatic Devices Other Equipment Saliva, Urine, and Blood Procedures for Self-Measurements Keeping Records Sampling Sequence Looking at the Data Making Graphs Testing for a Time-Effect Take-Home Message Appendix Item 1: Sample sheet for recording oral temperature and other functions Item 2: Sample sheet for recording and graphing temperature, pulse and blood pressure

12 xxii Contents Item 3: Detailed instructions for performing self-measurements Item 4: Random Number Adding Speed Test sample pages Item 5: Random Number Memory Test sample pages 13 Chronobiometry: Analyzing for Rhythms Introduction Data Collection Number of Timepoints How Long? Sampling Often Enough Aliasing Decision Making Data Preparation Graphs and Visual Inspection Editing or Transforming Data Essay 13.1: Standard Deviation and Error Normalizing Data Partitioning Data Spans Statistical Detection of Time Effects Using Two Timepoints Using Three or More Timepoints Statistical Detection of Rhythms Analyzing Time-Series by Standard Methods Limitations of Standard Methods Analyzing Time-Series by Curve Fitting The Least-Squares Technique The Best-Fitting Curve Statistical Significance Complex Waveforms Rhythm Parameter Comparisons Lack of Rhythm Detection Descriptive Rhythm Parameters The Cosinor Illustrated Example of a Cosinor Program Take-Home Message Author Index Subject Index...649

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