Glossary

SYMBOLS (For Explanations, see below)

T Period of a biological rhythm (oscillation).

T Period of a zeitgeber.

a; p Activity time; rest time (two fractions of an activity rhythm).

4> Phase angle of biological oscillation. <I> Phase angle of zeitgeber.

if; Phase-angle difference between zeitgeber and biological oscillation.

LD Light-dark cycle. A light-dark cycle is composed of light time (L) and dark

time (D); the term photoperiod is used synonymously with light time.

LL; DD Continuous illumination; continuous darkness.

C.t. Circadian time. A time scale covering one full circadian period; the zero

point is defined arbitrarily.

Z.t. Zeitgeber time. A time scale covering one full zeitgeber period; the zero point is defined arbitrarily.

PRC Phase-response curve.

DEFINITIONS OF OSCILLATING SYSTEMS

1. Classification referring to internal features of the oscillating system:

a. Active system: capable of self-sustaining oscillations (endogenous rhythms).

b. Passive system: capable only of forced oscillations (exogenous rhythms) or

damped oscillations.

2. Classifications referring to external conditions:

a. Autonomous system: not under the influence of a periodic input (self-sustaining and damped oscillations).

b. Heteronomous (nonautonomous) system: under the influence of a periodic input (forced oscillations, whether in an active or in a passive system).

547

548

GLOSSARY

Period: time after which a definite phase of the oscillation recurs.

Frequency: reciprocal of period.

Mean value: arithmetic mean of all instantaneous values of an oscillating variable within

one period. Amplitude: difference between maximum (or minimum) and mean value in a sinusoidal

oscillation; often used in a looser sense for other oscillations, too.

Range of oscillation: difference between maximum and minimum value (independent of

shape of oscillation).

Phase: instantaneous state of an oscillation within a period, represented by the value of the

variable and all its time derivatives.

Phase angle: value of the abscissa corresponding to a phase of the oscillation, given in degree

or any other fraction of the whole period with reference to an arbitrarily set zero point.

It can be given in units of time, if the length of the period is stated.

EXPLANATION OF FURTHER TERMS

Acrophase: phase angle of the crest (maximal value) of a sine function fitted to the raw

data of a rhythm (time series).

Circarhythms: classes of rhythms that are capable of freerunning in constant conditions

with periods approximating that of the environmental cycle to which they are nor­

mally synchronized, and that are entrainable by zeitgebers (circadian, circatidal, cir­

calunar, circannual rhythms).

Desynchronization: (a) External: loss of synchronization between rhythm and zeitgeber.

(b) Internal: loss of synchronization between two rhythms within one organism.

Entraining agent: synonymous with zeitgeber. Entrainment: synchronization of a self-sustaining oscillation by a forcing oscillation (zeit­

geber). During entrainment, the frequencies of the two oscillations are the same or

integral multiples (entrainment by frequency demultiplication). Freerun: state of a nonentrained circarhythm, either in constant conditions or after loss of

entrainment by a zeitgeber that is still present but too weak. Infradian: biological rhythm with a period longer than that of a circadian rhythm. Pacemaker: localizable functional entity capable of self-sustaining oscillations and of syn­

chronizing other rhythms. Phase-angle difference: difference between corresponding phase angles in two coupled

oscillations, given either in degree or in units of time. Often, "corresponding" phase

angles have to be defined arbitrarily.

Phase-response curve: indicates how the amount and the sign of a phase shift, induced by

a single stimulus, depends on the phase at which the stimulus is applied.

Phase shift: single displacement of an oscillation along the time axis; may occur instanta­

neously or after several transients. Photoperiod: synonymous with light time (see symbol LD).

Range of entrainment: range of period (T) within which a self-sustaining oscillation can

be entrained by a zeitgeber. Transients: temporary oscillatory states between two steady states. Ultradian: biological rhythm with a period shorter than that of a circadian rhythm. Zeitgeber: the forcing (external) oscillation that entrains a biological (self-sustaining)

oscillation.

Index

Abraxas miranda, 412, 416 Acetabularia cell fragments, 62 N-Acetyltransferase, circadian rhythm and, 65,

250, 262-263 Acheta domesticus, 150 Acoustic displays, in insects, 131 hronycta rumicis, 414, 415, 418, 420

lcrophase. See also Phase maps defined, 24 long-term trends in, 479-480

phase shifts in, 222 ACTH

mammalian activity and, 193-194

ovarian cycle and, 532 sleep-waking rhythm and, 514

Actinia equina, 361 Active systems, self-sustaining oscillations in, 4 Activity

behavioral. See Behavioral activity locomotor. See Locomotor activity short-term rhythms in, 491-497

Activity rhythms, in mammals, 185-193 Actographs, locomotor activity in, 125 Adaptive behavior

daily movements and migration in, 287 -289 foraging and food intake in, 285-287 habit strategy and, 294-296 individual daily habits and, 291-296 periodic food anticipation and, 293 reproduction and life history in, 289-291 temporal niche shifts in, 282-283

Adaptive daily routines, constraints in, 276 Adrenal rhythmicity, in rat, 264 Aedes atropalpus, 416, 418, 432-433, 441 Aedessp.,176 Aftereffects, 85

549

Aging, circadian rhythm and, 84, 268-270, 322

Alligator mississiftpiensis, 198 Ambystoma tigrinum, 200 Ammodytes dubius, 203 Amphibians, behavioral rhythms in, 189-200 Amplitude, in developing rhythm, 261 Androtonus australis, 162 Anesthesia, human circadian rhythms and, 328 Anguilla vulgaris, 354 Angular velocity, pacemaker period and, 60 Animal behavior. See also Adaptive behavior;

Behavioral activity daily routines in, 275-276 daily time and energy allocation in, 283-291 diurnal, 279-283

foraging and food intake in, 285-287 nocturnal 279-283

Annual rhythms. See also Behavioral rhythms; Biological rhythms; Circadian rhythms; Circannual rhythms; Rhythms

in man, 475-486 mortality and, 478-482 perspective on, 381-388 proximate factors controlling, 384-388 ultimate factors in, 383-384

Anolis carolinensis, 246 Anolis lizards, 282

photoperiodism in, 465 Anopheles funestus, 134 Anopheles gambiae, 128 -134 Anopheles superpictus, 131 Antheraeapernyi, 128, 148-149, 165,414,419-

420, 424 Anthrenus verbasci, 392, 411 Aotus trivirgatus, 191, 369 Apis mellifica, 291

550

INDEX

Aplysis cali/ornica, 61, 65-66,146-147,154,156, 166-168

Apodemus sylvaticus, 189 Arctosa variana, 302 Army ants, flight times of, 290 Arrhythmic species, 279 Arrhythmicity, surgical lesions and, 160-161 Artibeus liturtatus, 189,369 Aschoff's rule, 47, 66, 118 Aspis cerastes, 198 Astro-orientation, biological clocks in, 299-306 Autocorrelation function, 27 Avian pineal organ, 247-252, 463-464 Avian reproduction, rainfall distribution and, 382.

See also Birds Aythya sp., 281

Background, periodogram and, 35 Basic rest-activity cycle, 8, 512-513

Bees. See also Honeybees rhythmicity of learning in, 136 Zeitgediichtnis of, 62

Beetle, independent pacemakers in, 156-158 Behavior. See also Adaptive behavior; Animal

behavior; Behavioral activity adaptive daily strategies in, 275-296 circadian organization of, 137-140

Behavioral activity in birds, 196-197 defined, 184 in fish, 201-206

Behavioral adaptation, temporal specialization and, 280-281

Behavioral categories activity in, 184 daily frequency variations in, 284

Behavioral rhythms. See also Circadian rhythms in amphibians, 199-200 behavioral categories and, 184 in birds, 194-197 feeding rhythms and, 130 in fish, 201-204 gated behavior and, 133-134 in invertebrates, 125-140 in mammals, 186-187 mating rhythms and, 130-132 in orientation behavior, 134-135 reproductive behavior and, 132 in reptiles, 198-200 responsiveness changes and, 136-137 time sense and, 135-136 in tsetse fly, 139 in vertebrates, 183-205

Behavioral variability, in vertebrates, 184 Bicoordinate sun navigation, 303-304 Biological clocks

in astro-orientation, 299-306 environmental time measurement by, 57-58

Biological noise, synchronization in, 52

Biological rhythms, 6-7, 14. See also Circadian rhythms

complications in interpretation of, 14-19 components in, 35 data collection methodology in, 11-19 endogenous, 4 entrained phase relationships in, 24-26 entrainment in, 16 estimating period of, 26-32 feedback loop in, 13 freerunning, 12-16 frequency range of, 3 functions served by, 8 interaction among, 7-8 long-term oscillatory trends in, 13 null hypothesis in, 34-35 persistent, 14 phase in, 23 phase shift in, 14 plasticity of rhythm properties and, 18-19 sleep as, 499-510 spectrum of, 3-5 statistical inference in, 33, 36 survey of, 3-8 sustained freerunning rhythms in, 14-16 teleonomy of, 7-8 true, 35 ultradian and infradian, 7 "waveform" of, 21

Biphasic circadian rhythm, 126 Birds

behavioral rhythms in, 194-197 circadian organization in, 251-252 endocrine aspects of photoperiodism in, 464-465 light perception in, 463 photoperiodism in, 461-465 pineal organ in, 247-252, 463-464 sun compass reactions of, 304 testicular development related to photoperiodism

in, 462 Blaps gigas, 155-158 Blennius pholis, 204, 359 Body temperature, work curve and, 335-336 Bombyx mori, 412, 414 Bonasa umbellus, 196 BRAC. See Basic rest-activity cycle Brain photoreceptors, in vertebrates, 246 Bruce eifect, 532 Bunning hypothesis, in plant photoperiodism, 421-

422, 429 Bunning oscillator, 401-402

Caching frequency, in adaptive behavior, 286 Calanus helgolandicus, 288 Calidris alpina, 383 "Calling" behavior, 131 Cambarus, 152 Capra ibex, 288 Carcinus, 154, 162-163

Cardioderma cor, 191 Carpocapsa pomonella, 419, 427, 442

Castor canadensis, 188 CEC. See Circumesophageal connectives

Central nervous system circadian modulation of, 159~ 162 circadian pacemakers in, 146~ 158

circarhythms in,S locomotor activity and, 126 sleep rhythms and, 515~517

Cephaloscyllium ventriosum, 201 Cervus nippon, 394, 398 Chickens, circadian pacemaker in, 251 Children, growth rate variation in, 4 7 6~4 77

Chionactis occipilalis, 198 Chorthippus curtipennis, 165 Circadian "Bunning" oscillator, 401 ~402 Circadian organization

in birds, 251 ~ 252 "new" functions for, 71

Circadian organization of behavior, 137 ~ 140 Circadian oscillations, in insect photoperiods, 432~

441

Circadian oscillator, as pacemaker, 96 Circadian pacemakers. See also Pacemaker(s)

in CNS, 146~ 158

defined, 44 different oscillators in, 51, 96 dynamic models of, 44~47

feedback loop in, 70 homeostasis of, 116 in vitro, 146~ 148

localization of via lesions, 150 ~ 155 localization versus transplantation of, 148~ 150 models of, 41 ~42 multiple, 155~ 158

optic lobes in, 157

period T of, lIS phase-response curves of, 98~ 102

photoperiodic time measurement for, 63 primary, 247 in region around singularity, 45 in subesophageal ganglion, 150 Zeitgediichtnis and, 62 zeitgeber and, 114~ 115

Circadian period, darkness and, 82 Circadian programs

model for, 73 transition from exogenous temporal order to

endogenous temporal organization in, 60~ 61 Circadian rhythmicity

control of in invertebrates, 145~ 168 neural basis of, 145 as postnatal phenomenon, 258 sensorimotor integration and, 159~ 163

Circadian rhythms. See also Circadian systems; Daily rhythms; Rhythms

N -acetyltransferase and, 250 aging and, 268~270, 322

Circadian rhythms (Cont.) amplitude in, 261 versus behavioral rhythms, 243

biphasic, 126 cortisol in synchronizing of, 228~229

damped oscillators and driven rhythms in, 253 food and water deprivation in, 191~193 freerunning, 6, 81~85, 90~91, 98~99, 315 frequency demultiplication of, 89, 106,322,401

hypothalamic injury and, 250 internal environment and, 268 internal synchrony between, 221 ~222 of locomotor activity, 82,127 neural and endocrine control of in vertebrates,

243~253

in neural and endocrine systems, 227 ~228 in neuronal firing rate, 227 ontogeny and, 173, 257~270 origin of term, 6

phase plasticity in, 223 of potassium flux, 237 regression line and, 27 in sensory response, 162~ 163 temperature and, 82~83 waveform determinants for, 216~217 zeitgeber and, 215

Circadian subsystem, analysis of, 234~235 Circadian surface, model of, 76 Circadian system

convergence in, 76~ 77

development of, 265 entrainment and, 9 5~ 123 general perspective in,S 7 ~ 77 human, 311 ~329 multioscillatory structure of, 86, 225~227 ontogeny of, 257 ~270 ovarian cycle and, 538~S40 phase maps of, 217~218

phase plasticity in, 220 photoreceptor input to, 244~ 246 short-term activity rhythms and, 494~49S transmeridian flights and, 32S~ 326 unity in diversity of, 76~ 77 zeitgeber shifts and, 32S~326

Circadian time REM sleep and, S09~S10 zone corresponding to, 44

Circadian time memory, in mammals, 194 Circadian timing system

light-dark cycles in, 232~234 lunar rhythms and, 375 models of, 225 pacemakers in, 230 qualitative models of, 231~235 subsystems in, 231 ~232

Circadian waveform, alterations in, 222 Circadian zeitgeber, 215, 32S~326. See also

Zeitgeber(s) in birds and reptiles, 66

551

INDEX

552

INDEX

Circalunar rhythms, 373 Circannual clocks, existence of, 397 Circannual functions, relation with individual

organisms, 396-397 Circannual period lengths, range of, 395 Circannual rhythms, 7-8, 386-387

adaptive significance of, 404-408 behavior with range of entrainment, 399-400 circadian rhythms as possible components of,

400-403 demonstration and distribution of, 392-394 evidence of, 391 innateness of, 396 mechanisms of, 400-404 in ovarian cycle, 533 persistence of, 393-395 properties of under constant envirpnmental

conditions, 394-397 synchronization of, 397 -400 zeitgebers and, 397-399

Circannual systems, 391-408 Circannuq.1 zeitgebers, 397-399. See also

Zeitgeber(s) "Circa-" oscillators, clocklike properties of, 58-60 "Circa-" pacemakers, 59. See also Circadian

pacemakers; Pacemaker(s) "Circa," prefix, 58 Circarhythms, 5-6. See also Circadian rhythms Circasemilunar rhythms, 373 Circa-solar-dian clock, 305 Circa tidal pattern, in fiddler crabs, 365 Circatidal rhythms

circadian oscillator and, 361 defined,356

Circhoral rhythm, 6 Circumesophageal connectives, arrhythmicity and,

160-161 Citellus lateralis, 392, 394, 403-404 Citellus mohavenis, 404 Citellus tereticaudus, 404 Clethrionymus gapperi, 190 Clock

circa-lunar-dian, 305 circa-solar-dian, 305 insect, 125,421-441

Clock-controlled orientation in space, 299-307 Clock-shift experiments, with wolf spiders, 302 Clunio marin us, 132, 177,365,371-374 Cockroach

circadian rhythmicity in, 157 compound eye of, 157 freerunning circadian locomotor rhythm in,

129 mutually coupled pacemakers in, 157 photo receptors in, 164

Coeloides brunneri, 414 College students, sleep-length studies in, 502 Colobus badius tephrosceles, 191

Color change, tidal rhythm and, 356 Columba domestica, 304 Community, diurnal and nocturnal behavior in,

279-283 Complete photoperiod

defined, 117 entrainment by, 110-114

Complex pacemakers, 119-121. See also Pacemaker(s)

PRC of, 120 Component, in biological rhythm, 35 Computer programs, power spectrum in, 28 Conception rates

latitude and, 482-483 long-term trends in, 479-480 seasonal rhythms in, 483 sunshine duration and, 484

Confidence limits, in data analysis, 35 Constant darkness, models dealing with, 47 Continuous illumination

complete photoperiod and, 110 versus constant darkness, 47 models dealing with, 47-49

Convergence, in circadian systems, 76-77 Convoluta roscoffensis, 132, 134, 355, 365 Corticosteroids, responsiveness and, 328 Corticotropin-releasing factor rhythm, 265 Cortisol concentration, in circadian rhythm

synchronizing, 228-229 Corvus corax, 289 Corvus frugilegus, 197 Cottus poecilopus, 204 "Coupling strengthener," in oscillator interactions,

51 Crabs, tidal activity in, 154 Crayfish

CEC axons in, 161-162 entrainment in, 165 ERG amplitude rhythm of, 166 extraretinal photoreception in, 165 pacemaker location in, 152-153

Crayfish optic lobe, DPLH in, 159 Crepuscular species, 279 CRF. See Cortic(,tropin-releasing factor Cricket

circadian rhythm in stridulation of, 150-151 optic nerve entrainment in, 164 tactic rhythm in, 135

Critical pulse experiment, 46 Crocodiles, behavior rhythm in, 198 Crocodilus niloticus, 198, 283 Crustacea

neuroendocrine control of rhythmic locomotor activity in, 161

pacemaker location in, 152 sun compass reactions in, 304

Cryptoblepharus, 356 Cyprinodon macularius, 203

Dacus fruitflies, 136, 173 Dacus tryoni, 131 Daily habits, individual behavior and, 291-296 Daily movements of migration, in adaptive

behavior, 287-289. See also Migration; Migratory birds

Daily rhythms, selection pressures for, 275-279. See also Circadian rhythms

Daily routines, adaptive, 276. See also Adaptive behavior

Daily strategies, in behavior adaptation, 275-296 Daphnia magna, 134, 136 Dark-dark cycle, entrainment and, 98-99 Dark pulses, defined, 48 Data analysis, 21-38

confidence limits in, 35-36 descriptive versus inferential procedures in, 22 Lamprecht-Weber procedure in, 29 least-squares spectrum analysis and, 30 power spectrum in, 28-30 subjective probability in, 37 -38

Data collection automatic or semiautomatic, 11 in biological rhythm studies, 11-19 schematic example of, 12-14

Data series average period in, 27 Fourier components and, 28-29 "rhythmic" quality of, 26

Day-neutral insects, 413 DCMU. See Nt(Dichlorophenyl)-Nrdimethylurea DD. See Constant darkness Descriptive statistics, 22-32

versus inferential procedures, 22 Diapause

induction and termination of, 411, 416-417 insect photoperiod and, 414 model for, 416-417

Diapause-inhibiting effects, in insect photoperiodism, 414-415

N t-(Dichlorophenyl)-N ,-dimethylurea, photosynthesis rhythm suppression by, 63

Dictyota dichotoma, 372 Didelphis marsupialis, 184 Diel pattern, of activity changes, 126 Digit span, memory and, 336 Dipodomys microps, 189 Dipodomys nitratoides, 189 Dipodomys spectabilis, 189 Direction, periodic change of, 306-307 Distal retinal pigment light-adapting hormone, in

crayfish optic lobe, 159 Diurnal species, 279 Diurnal tides, 352-353, 358 Diurnal ways of life, 279-283 DNA, circadian rhythm encoding in, 173 DPLH. See Distal retinal pigment light-adapting

hormone

Driven rhythms, in vertebrates, 253 Drosophila melanogaster, 100-101, 129, 132, 165,

174-180 Drosophila pseudoobscura, 44, 46-49, 51, 66, 69-

70,72,74,77,82,174,176,178-180, 422-423,429-430,437-438

"bistability" phenomenon in, 109 eclosion rhythm in, 97-99, 112-113 entrained steady states of, 109 entrainment in, 97-98 gated behavior in, 133 light continuum in, 111 pacemaker for, 103-104 phase relation and, 105 skeleton zeitgeber and, 107

Drosophila sp., 71, 165 Drosophila victoria, 175 Drug responsiveness rhythms, 328-329

Eclosion, defined, 125 Eclosion hormone, silkmoth behavior and, 159-160 Eclosion rhythm, of Drosophila pseudoobscura, 97-

99,112-113 Ecosystem, isochrone concept in, 45-46 Electromyogram changes, in sleep, 501 Embryonic stage, in insects, 175 Endogenous pacemaker, 61 Endogenous rhythms

circarhythms and, 5-6 defined,3-4

Entrainability, in freerunning circadian rhythms, 86-90

Entrained phase relationships, 24-26 cross-treatment comparisons in, 25-26 description of, 24-25 zeitgebers in, 25

Entrained steady states, stability of, 114-117 Entraining signals, perception of in vertebrates,

244-246 Entrainment

by artificial zeitgebers, 317 -319 biological rhythms and, 16 circadian systems and, 95-123 in cockroaches, 163-164 by "complete" photoperiods, 110-114 by exotic-light cycles, 104-106 via extraretinal photoreception, 165-166 general features of, 95-96 hierarchical, 68 by light-dark cycles, 96-97 limits of, 106 in model circadian program, 73 neural mechanisms in, 167-168 non parametric, 88 optic nerve in, 163-164 pacemaker, 59, 121-122 parametric, 88 partial, 89, 319-325

553

INDEX

554

INDEX

Entrainment (Cont.) photoreception and, 163-168 relative, 89 by "skeleton" photoperiods, 106-110 systemic versus pacemaker, 59, 121-122 zeitgeber and, 16, 95

Entrainment mechanisms, discrete versus continuous, 96-98

Environmental factors, in adaptive daily routines, 276

Environmental time, biological clocks in measurement of, 57-58

Environmental zeitgebers differential coupling of oscillators to, 230 temporal order of, 221

Eocrepuscular distribution of activity, in insects, 119

Ephippiger, 167 Equilibrium, points of, 42 Estrous patterns, 538 Eudiapause insects, 414 Euglena gracilis, 134 Eunice viridis, 371 Euphagus cyanocephalus, 293 Eurydice sp., 359 Evolutionary radiation, night and day in, 280 Exclrolana chiltoni, 132,357,359-360,367 External coincidence model, in insect

photoperiodism, 422, 429-430, 434-440 External day, seasonal changes in, 117-121

Falco eleonarae, 383 Falco tinnunculus, 293, 295 Feedback loop, as slave oscillator, 70 Feeding

adaptive behavior in, 285-287 rhythm of in relation to locomotion, 130

Female "calling" behavior, male responsiveness and, 290

Field mole, wheel-running activity in, 492-493 Fish

behavioral rhythms in, 201-204 daily migration and feeding in, 201-203 environmental influences on, 203-204 sun compass reactions of, 304

Follicle-stimulating hormone in mammalian photoperiodism, 460-461 in ovarian cycle, 526 in vertebrate photoperiodism, 459-460

Folsomia candida, 7 Food deprivation, mammalian activity and, 191-

192 Food intake

adaptive behavior and, 285-287 locomotion rhythms and, 130 optimal foraging and, 286

Fourier components, in data analysis, 28-29 Fox-rabbit populations, periodic trajectories in,

42-43

Freerunning activity rhythms, properties of, 84 Freerunning circadian rhythms, 81-85

constant darkness cycle and, 98-99 entrainability and phase relationships in, 86-90 experimental conditions and, 83 external and internal factors in, 81-82 frequency dependence in, 81-82 frequency variability and, 84 history dependence in, 85 phase-angle difference in, 86-87 photoperiod in, 88 stability and, 84 zeitgeber diversity and, 85-86 zeitgeber phase shift and, 90-91

Freerunning circannual rhythms, T period of, 395 Freerunning period, environmental effects of, 17.

See also Freerunning circadian rhythms Freerunning period lability, 50

arrhythmic state and, 52 Freerunning rhythms. See also Freerunning

circadian rhythms in man, 315-316 medulla-lobula complex in, 151 zeitgeber in, 12-13

Freerunning stochastic pattern, in tidal timing, 363-364

Freerunning tidal rhythms, 356 Frequency

entrainment limits and, 106 in freerunning circadian rhythms, 81-82

Frequency demultiplication, 106 Fringilla coelebs, 6, 84, 90 Fruit fly, rhythm studies in, 173-180. See also

Dacus; Drosophila FTG. See Gigantocellular tegmental field Fundulus heteroclitus, 204

Gadus morhua, 203 Galago senegalensis, 191 Gallus domesticus, 287 Gated behavior, 133 Gating rhythm, 133 Gazella dorcas, 191 Gecarcinus, 154 Generation time, pacemaker period and, 61-62 Gigantocellular tegmental field, 517 Glossina morsitans, 139 Gnathonemus macrolepidotus, 202 Gonadal steroids, in vertebrate photoperiodism, 458 Gonadotropins

cyclic release of, 523 in vertebrate photoperiodism, 458-459

Gonyaulax polyedra, 63-64, 100 Gopherus polyphemus, 198 Grapholitha molesta, 419, 442 Growth rate, in children, 476-477 Guillemots, daily distribution of jumping in, 278-

279 Gymnorhamphichthys hypostomus, 204

Habits, strategy and, 294-296 Haemotopus ostralegus, 285 Halisidota argentata, 129 Hamsters, photoperiodic refractoriness in, 451-452 Hantzschia virgata, 134 Hemoglobin levels, increase of in spring, 477 Heterodontus francisci, 201 Hierarchical entrainment, 68 Hipposideros speoris, 86 Homing ability, tidal rhythm and, 355-356 Honeybees. See also Bees

"dance" of, 292 time memory in, 291-294 Zeitgediichtnis of, 62, 291

Hormonal rhythms, sleep and, 513-515 Hormone cycle

follicular phase, 526-527 luteal phase, 527 -528 periovulatory phase, 527 seasonal changes in, 534-535

Horseshoe crab, pacemaker-rhythm relationships in, 162

Hourglasses, in insect photoperiodism, 421, 424-429,442

House fly, as day-neutral insect, 413 Huechys sanguinea, 281 Human circadian rhythms

physiological and psychological functions in, 312-314

zeitgeber in, 314 Human circadian system, 311-329

and entrainment by artificial zeitgebers, 317 - 319 freerunning rhythms and, 315-316 internal desynchronization and partial

entrainment in, 319-325 medical application of, 328- 329 medical problems and, 328-329 reentrainment and, 326-328 rhythm patterns and, 311-315 zeitgeber shifts and, 325-326

Human entrainment, by artificial zeitgebers, 317-319. See also Entrainment; Zeitgeber(s)

Human infant, sleep-wake rhythmicity in, 259 Human mortality, annual rhythms and, 478-482 Human rhythm patterns, reproducibility and

dependence in, 311-315 Humans. See also Man

overt rhythmicity development in, 260-261 sleep-wake cycle in, 259, 314, 501-502

Hyalophora cecropia, 148-149, 165 Hyemoschus aquaticus, 188 Hypothalamic lesions, in sparrows, 249

Inferential methods, use and abuse of, 32-35 Inferential statistics, rhythm-specific issues in, 34-

37 Infradian rhythms, 7 Innate temporal programs, 57-58 Insect clocks, 125

Insect photoperiodism, 411-443 Bunning hypothesis in, 421-422, 429 circadian oscillations in, 432,-441 "clock" in, 421-441 control of in brain, 420 external coincidence model in, 422, 429-432,

434-440 hourglasses in, 421, 424-429 internal coincidence model in, 422-423, 440-

441 latitude/longitude effects in, 418-419 lunar-monthly rhythms in, 370 Megoura viciae "clock" and, 424-427 models of, 421-432 Nasonia vitripennis and internal coincidence

model in, 440-441 nature of, 441-443 Pectinophora gossypiella and external

coincidence model in, 429-432 resonance model in, 423-424 Sarcophaga argyrostoma and external

coincidence model in, 434-440 temperature and, 415-417 time measurement in, 421-432

Insects acoustic displays in, 131 circadian organization of behavior in, 137-140 day-neutral, 413 diapause of, 414 eocrepuscular activity in, 119 long-day and short-day species of, 412 oligopause and, 414

Instances, models and, 42 Interacting oscillators, populations of, 50-52 Internal circadian system, circannual variations in,

402. See also Circadian rhythms; Circadian system

Internal coincidence model, 73-74, 440-441 Internal de synchronization

in man, 319-325 reasons for, 322 in squirrel monkey, 225 strong zeitgeber and, 319

Internal environment, in circadian rhythm development, 268

Internal organization abstract models of, 224-231 single or multioscillator system in, 225

Internal phase-angle shifts, in circadian system, 222

Internal synchrony, between circadian rhythms, 221-222

Internal temporal order, 215-237, 311 in absence of environmental time cues, 221-223,

316 anatomical and physiological basis of, 223-235 defined,215 failures in, 236-237 importance of, 235-237, 329

555

INDEX

556

INDEX

Internal temporal order (Cont.) inherited features of, 219-220 in steady-state-entrained conditions, 215-221 zeitgeber and, 220-221, 316

Intertidal conditions, temporal adaptations to, 353-356

Invertebrates. See also Insects behavioral rhythms in, 125-140 circadian rhythm genetics and development in,

173-180 neural and endocrine control of circadian

rhythmicity in, 145-168 Isochrone, defined, 45 Isoperiodic conservative oscillators, 46

Jet lag, phase shifts and, 269, 325

Kestrel, movements of, 295-296

Lacerta sicula, 66, 246 Lampetra jluviatillis, 354 Lamprecht-Weber procedure, in data analysis,

29-30 Lampyris noctiluca, 132 laTUS argentatus, 197 LaTUS atricilla, 197 LaTUS hyperboreus, 197, 277-278 Latitude, death rates and suicides in relation to,

480-482 LD cycle. See Light-dark cycle Learning, rhythmicity in, 136 Least-squares spectrum analysis, in data analysis,

30 Lee-Boot effect, 532-533 Lemna perpusilla, 438 Lepomis macrochiTUs, 202 Leptinotarsa decemlineata, 412, 420 Leucophae maderae, 100, 129, 151, 157, 164 Leuresthes tenuis, 371 LH. See Luteinizing hormone LHRH. See Luteinizing-hormone-releasing

hormone Light compass reaction, 300 Light-dark cycles

in circadian rhythm development, 266-267 in circadian timing system models, 232-233 entrainment by, 96, 317 in ovarian cycle, 534 sleep and, 500 thermoperiods and, 415

Light sensitivity, lunar-monthly rhythms in, 370 Light stimuli, parametric entrainment and, 48 Limax jlavus, 392 Limit-cycle behavior, 103 Limit-cycle oscillator, 43 Limit-cycle system, circadian pacemaker as, 44 Limits of entrainment, frequency and, 106. See

also Entrainment Limulus polyphemus, 162

Littoral zone, distribution of organisms in, 353 Lizards

brain photoreception in locomotor rhythm entrainment by, 246

photoperiodism in, 465 Local time, recognition of, pacemaker entrainment

and, 59-60 Locomotion. See also Locomotor activity rhythms

feeding rhythm and, 130, 285-287 periodic change of, 307

Locomotor activity, neuroendocrine control of, 160 Locomotor activity rhythms, 126-130

circadian, 82 tidal, 127, 354-356

Lata lata, 354 Lower vertebrates, photoperiodism in, 465-466 Loxia curvirostra, 385 Luciola lusitanica, 132 Lunar-monthly rhythms, in insect populations, 370 Lunar-rhythmic behavior adaptations, 368-370 Lunar rhythms, 367-376

clock period and, 373 persisting, 372-373 zeitgeber experiments and, 373-374

Lunar-semimonthly rhythms, tidal zeitgeber and, 375

Lunar terminology, 367-368 Luteinizing hormone

circadian systeni and, 538 mammalian ovary and, 523, 527 in mammalian photoperiodism, 459-460 nutrition and, 532 in ovarian cycle, 527 seasonal changes in, 534-535 in vertebrate photoperiodism, 459-460

Luteinizing-hormone-releasing hormone mammalian ovary and, 523 in mammalian photoperiodism, 457

Lycosa jluviatilis, 304

Macaca mulatta, 188 Macaca nemestrina, 81 Macaque monkeys, social synchronization in, 188 Mallotus villosus, 203 Mammalian activity. See also Mammalian

photoperiodism; Mammals food and water restriction in, 191-192 learning and memory in, 193-194 seasonal cycle in, 190-192

Mammalian behavior. See also Mammalian activity

human interference in, 189 seasonal cycles in, 190-191

Mammalian mother, rhythmicity in, 188-189 Mammalian ovary, cyclic function of, 523-540.

See also Ovarian cycle Mammalian photoperiodism

annual cycles and, 449 endocrine aspects of, 458-461

Mammalian photoperiodism (Cont.) follicle-stimulating hormone and, 460-461 gonadotropins in, 458-459 light pathway in, 452-453 luteinizing hormone in, 459-460 melatonin in, 454-457 organs and physiological processes involved in,

452-458 pineal gland in, 453 prolactin in, 460-461 puberty and, 451 spontaneous processes and, 451-452 suprachiasmatic nuclei in, 453

Mammals activity behavior in, 185-193 annual cycles in, 449 behavioral rhythms in, 185-194 circadian time memory in, 194 environmental influences in, 185-193 lunar cycle and, 189 photoperiodic signal in, 450-451 sexual cyclicity in, 188 social stimuli in, 185-189 suprachiasmatic nuclei in, 252-253

Man. See also Human circadian rhythms; Human circadian system; Human entrainment; Humans

annual rhythms in, 475-486 sleep-wake cycle in, 315, 318-324 sleep patterns and, 502

Marathon dancers, new nest site and, 135 Marinogammarus, 366 Marmota monax, 190, 397 Masking, 13, 86, 90, 314 Mathematical models, 41-53

dynamic model instances and, 49 population phenomena and, 50

Mating rhythms, 130-132 Meal timing, optimal, 286 Measurement, objectivity in, 22-23 Medicine, drug responsiveness rhythms and, 328 Medulla-lobula complex, in freerunning

rhythmicity, 151 Megoura viciae, 174,419-421,424-426,438,442 Melanocyte-stimulating hormone, 193 Melatonin, 247-252, 454-457 Membrane model, 49 Memory

circadian time, 197 short-term, 336-337

Menarche, seasonal variation in, 477 Menidia audens, 203 Mesocricetus auratus, 98, 102, 108, 11 0, 120, 184 Methodology, in data collection, 11-19 Metriocnemus knabi, 414 Mice, short-term activity rhythms in, 491-497 Microtus agrestis, 495 Microtus arvalis, 495, 497 Microtus califomicus, 189, 191

Migration. See also Bird migration; Birds temporal and directional components of, 406-

407 tidal rhythms and, 354-355

Migratory birds, Zugunruhe in, 307, 392-397, 405

"Minimum tolerable night," pacemaker and, 116 Mixed tides, 358 Model(s)

continuous light and, 47-50 defined, 41 instances of, 42 mathematical. See Mathematical model topological, 44

Mole, wheel-running activity of, 492-493 Monestrous cycles, 538 Monkeys

behavioral rhythms in, 183-184 social synchronization in, 188

Monophasic activity, defined, 491 Moon, as orientational cue, 305-306 Moonlight

mammalian behavior and, 189-190 tides and, 367

Moon phases, environmental conditions related to, 367 -368. See also Lunar rhythms

Morning advances, interaction with evening delays, 114

Mortality long-term trends in, 478-482 seasonal rhythms in, 483

Mosquito, flight activity rhythm in, 176. See also Aedes sp.

Moth, rhythm studies in, 175-176. See also Silkmoth

Mother, as zeitgeber, 267 -268 Mouse, phase map of circadian system for, 218 Multicellular circadian systems, pacemaker

localization in, 64-65. See also Circadian systems

Multigene analysis, pacemaker genetics and, 177-178

Multiple oscillations, in circadian systems, 226 Multiple oscillator systems, organization of, 230-

231 Multiple pacemakers, 66-68,155-158. See also

Circadian pacemakers; Pacemaker(s) Musca domestica, 413 Mus musculus, 110, 188-189 Myotis myotis, 189 Myotis mystacinus, 283 Myrmeleon obscurus, 369 Mytilus califomicus, 361 Mytilus edulis, 130, 361

Nasonia vitripennis, 74-75, 414-415, 420, 428-429,432,440-441

Natrix sipedon, 198 Navanax sp., 155

557

INDEX

558

INDEX

Neural/endocrine systems, circadian rhythms in, 227-228

Neuronal firing rate, circadian rhythms in, 227 Neurosecretory activity, circadian modulation of,

159-162 Neurospora, 180 Nocturnal species, 279 Nocturnal ways of life, 279-283 Nonparametric entrainment, 88. See also

Entrainment NREM sleep, 517 Null hypothesis, in data-generating model, 34-35 Numerical signal averaging, 31 Nutrition, ovarian cycle and, 532 Nycticebus coucang, 191

Oligopause insects, 414 Oncopeltus fasciatus, 130, 132 Ondatra zibethicus, 191 Ontogeny

of circadian rhythms, 257 -270 defined,257

Optic lobes as circadian pacemaker, 157-158 DPLH band in, 159 in pacemaker function, 151-152, 157-158

Optimal foraging, in meal timing, 286 Orchestia, 354 Organization, emergence of, 258-265 Orientation angle, as behavioral rhythm, 134-135 Oriented activities, periodic change in, 306-307 Oryctolagus cuniculus, 287 Oscillation amplitude, determination of, 26 Oscillator coupling, mechanism of, 227 Oscillators

damped,253 isoperiodic conservative, 46 multiplicity of, 66 populations of, 50-52

Oscillator theory, basic concepts from, 42-43 Ostrinia nubilalis, 74, 415, 424, 428, 441 Ovarian cycle, 523-540

aggression and emotionality in, 529-530 altered characteristics of by species, 530-533 behavioral aspects of, 528-530 circadian system and, 538-540 follicular phase of, 524-525 ingestive behavior and, 529 interspecific differences in temporal occurrence

of, 533-538 light and, 530-531 light-dark cycle and, 534 luteal phase of, 525-526 nutrition and, 532 peri ovulatory phase of, 525 rhythmic variables associated with, 524-530 seasonality in, 533 sexual behavior and, 528-529, 535-536

Ovarian cycle (Cont.) social factors in, 532-533 suprachiasmatic nuclei and, 540 temperature and, 531-532 variety in, 538

Overt rhythmicity. See also Rhythm(s); Rhythmicity

circadian as postnatal phenomenon, 258 development of in humans, 260-262 genetics and,.177 -180

Ovulation, induced, 527 Oxymycterus rutilans, 191

Pacemaker(s) circadian. See Circadian pacemakers as circadian oscillator, 96 in circadian timing system, 230 complex, 119-121 cycle-to-cycle variability in period of, 36 defined,96 developmental ontogeny of, 173-177 discrete entrainment of by exotic light cycles,

104-106 endogenous, 61 genetics of, 177 -180 model limitations and, 41 multiple, 66-68, 155-158 mutually coupled, 157 "nonredundant," 156 oscillator and, 96, 173 phase relation of, 104-106 versus program, 63-71 slave oscillations and, 68-71, 74 sun-compass orientation and, 62-63 true phase of, 36 two-oscillator structure of, 120 zeitgeber and, 114-115

Pacemaker clock functions, origin and diversification of, 60-63

Pacemaker entrainment. See also Entrainment local time and, 59-60 versus system entrainment, 121-122

Pacemaker functions diversification of, 62-63 optic lobes in, 151-152

Pacemaker genetics, single-gene analysis in, 177-180 Pacemaker localization

in multicellular systems, 64-65 zeitgeber pathways and, 65-66

Pacemaker period generation time and, 61-62 homeostasis of with angular velocity, 60

Pacemaker phase shifts, kinetics of, 103-104 Pacemaker-slave phase relationship, 74 Panonychus ulmi, 441-442 Papaver rhoeas, 291 Paradoxical sleep, 504 Paramecium aurelia, 130

Parametric entrainment. See also Entrainment defined, 88 light stimuli and, 48

Parietovisceral ganglion, 148 Partial entrainment, and internal

desynchronization in man, 319-325 Parus major, 286, 289 Passer domesticus, 83, 244, 247 Patterns of sleep, defined, 501 Pectinophora gossypiella, 61, 66, 74, 175-177,

414-415,427,429-432,438,442 Perca fiavescens, 202 Performance efficiency

individual differences in, 343-346 sleep-waking cycle and, 339

Performance measurement, time-of-day effects in, 336-337

Performance rhythms, 333-347. See also Circadian rhythms

measurement of, 333-338 motivational factors in, 343-346 round-the-clock studies of, 338-340 situational factors in, 343-346 zeitgeber phase shifts and, 340-343

Periodic change, of oriented activities, 306-307 Periodic food, anticipation of in bees, 293- 294 Periodic internal system, advantages of, 236 Periodic trajectories, population dynamics and, 42 Periodogram

background of, 35 defined, 30 Lamprecht-Weber procedure with, 31

Periplaneta americana, 149-151 Perognathus longimembris, 84 Peromyscus californicus, 190 Peromyscus eremicus, 190 Peromyscus leucopus, 97, 100, 108, 116 Peromyscus maniculatus, 190 Personality factors, in performance efficacy, 343-346 Phase, in biological rhythms, 23 Phase-angle difference, in entrained circadian

rhythms, 86-87 Phase maps, of circadian system, 217-218 Phase plane, defined, 43 Phase plasticity, in circadian systems, 220 Phase-reference points

defined, 23 determination of, 23-26 during freerun, 23-24 inconsistency in, 36

Phase relation defined, 87 between eclosion peak and pacemaker, 122 between pacemaker and zeitgeber, 104-105

Phase relationships entrained, 24-26 in man, 316 in rat, 265

Phase-response curves, 44 of circadian pacemakers, 98-102 continuous light and, 47 of large amplitude, 49 normal, 48 shape of, 45 smoothness of, 52 symmetric, 117 for unicellular systems, 63

Phase shift in biological rhythm, 14 kinetics of, 103 measurement of, 17 -18

Phase transition curves, 101 Phormia terraenovae, 84 Photinus pyralis, 132 Photomenotaxis, 300 Photoperiod. See also Photoperiodism

in circannual rhythms, 395-396 "complete," 110-114 "skeleton," 97

Photoperiodic induction defined, 411 hourglasses in, 421, 442 models of, 421-424 physiology of, 419-443 rationale for, 441-443

Photoperiodic oscillation, 442 Photoperiodic phenomena, zeitgeber and, 122 Photoperiodic reactions, internal coincidence model

and,402 Photoperiodic refractoriness, spontaneous processes

and, 451-452, 463 Photoperiodic response, 412-419

photoreceptor and spectral sensitivity of, 419-- 421 temperature and, 415-417, 465

Photoperiodic response curves, 412-413 Photoperiodic signal, in mammals, 450-451 Photoperiodic time measurement, 63 Photoperiodism, 71-7 6

in birds, 461-465 bird testicular development and, 462 in insects. See Insect photoperiodism in lower vertebrates, 465-466 in mammals, 449-461

Photoperiod refractoriness. See Photoperiodic refractoriness

Photoreception entrainment and, 163-168 extraretinal, 165-166, 463, 465

Photoreceptors bypassing of, 163 localization of, 163-167 organized, 420

Photosynthesis rhythm, suppression of, 63 Phototactic rhythms, 134 Phoxinus phoxinus, 204, 293 Phylloscopus collybita, 405-406

559

INDEX

560

INDEX

Phylloscopus trochilus, 406 Phyllostomus hastatus, 189 Pieris brassicae, 416, 418-419, 427, 442 Pieris napi, 418 Pieris rapae, 418 Pineal gland

in birds, 247-252 ovarian cycle and, 536-537 in vertebrate photoperiodism, 453-455

Pineal rhythmicity, development of, 262-263 Pittendrigh's empirical model, 44 Plankton, tide-related drift of, 354-355 Plant photoperiodism, Blinning's hypothesis in,

421-422. See also Photoperiodism Platynereis dumerilii, 371-372 Poikilothermic vertebrates, annual cycles in, 465 Points of equilibrium, stable versus unstable, 42 Polyestrous cycles, 535 Polyphasic activity, defined, 491 Population dynamics, in theory of oscillators, 42 Population phenomena, in mathematical models,

50 Populations of oscillators, general features of, 50-

52 Porcupines, visual patterns in, 281 Potamobius, 152 Potassium flux, circadian rhythms of, 237 Power spectrum

defined,28 range of values in, 29-30

PRC. See Phase-response curve Presby tis john ii, 183 Probability, subjective, 37-38 Procambarus bouveri, 153 Procambarus clark ii, 152-153, 163, 166 Prolactin, in mammalian photoperiodism, 460-461 Protophormia terraenovae, 138 Proximate factors

in annual rhythm control, 384-385 hierarchical organization and, 387 -388 separation of from ultimate factors, 386

PS. See Paradoxical sleep .." (psi). See Phase relation .." Psi jump phenomenon, 116 Pterodroma, 196 Pterostichus nigrita, 432-433 Puberty, photoperiodic effects on, 451

Rainfall distribution, ovarian reproduction and, 382 Rapid eye movements, in sleep. See REM sleep Rat

adrenal rhythmicity in, 264 maternal behavior in, 267 paradoxical or activated sleep in, 504 physiology of emerging rhythmicity in, 262-265 pineal rhythmicity in, 262-263 short-term activity rhythms in, 491

Rat estrous cycle, as endogenous rhythm, 5

Reciprocal interaction hypothesis, 517 Rectal temperature

desynchronized rhythm and, 323-324 freerunning rhythms and, 315 sleep-wake cycle and, 318-324

Reentrainment, time needed for, 91, 325-327. See also Entrainment

Relative coordination, 13, 16, 90, 226, 317 REM sleep. See also Sleep; Sleep-wake cycle

ACTH and, 514 BRAC and, 512-513 central nervous system and, 516 changes in, as function of age, 506 circadian time and, 509-510 defined, 501 hormonal rhythms and, 513 real-time-locked, 511 ultradian rhythms and, 7-8, 510-511 wakefulness and, 509 for young adults, 503-504

Reproductive behavior, rhythms in, 132 Reptiles

behavioral rhythms in, 198-200 photoperiodism in, 465-466 rhythmic changes in, 136-137

Resynchronization, need for, 51 Retinal photoreception, circadian system and, 245 Retinohypothalamic tract, 252 Rhythm(s)

behavioral. See Behavioral rhythms circadian. See Circadian rhythms driven, 253 hormonal,513-515 in learning, 136 locomotor activity, 82,126-130,160,354-356 in orientation behavior, 134-135 postfeeding drop-off, 134 short-term, 491-497 tidal. See Tidal rhythms ultradian, 7-8, 510-513

Rhythmic locomotor activity, 82, 126-130, 160-161,354-356

Rhythmic ontogeny of circadian rhythms, 257-270 defined,257 environment role in, 266-268

Rhythm transients, following pacemaker phase shift, 72

Rhythm waveform, determinants of, 216-217 Rissa tridactyla, 197 Rousettus aegyptiacus, 189

Salamandra salamandra, 200 Salmo trutta, 119, 203 Sandhoppers, correction for sun's movement in, 135 Sarcophaga argyrostoma, 74-75,100,414-417,

428-430,434-440,442 Scarus croicensis, 202

ScelopoTUs olivaceus, 246 Schuler's constant, 513 SCiUTUS carolinensis, 190 Seasonal activities, timing of, 404-406 Seasonality, 381-383

neuroendocrine correlates of, 534-538 SEG. See Subesophageal ganglion Semidiurnal tides, 352, 358 Sensorimotor integration, circadian rhythmicity

and, 159-163 Sensory input, circadian modulation of, 162-163 Sequence-of-stages approach, 403-404 Serinus canarius, 87 -88 Serotonin

in Aplysia eye, 167 rhythmicity and, 262

Sesarma haemotocheir, 371 Sesarma intermedium, 371 Sexual behavior, seasonality in, 335-336 Short-day insects, 412-413 Short-day thermoperiods, 415-416 Short-term activity rhythms, 491-497

causal considerations in, 492-495 functional considerations in, 495-497 oxygen availability and, 496-497

Silkmoth, eclosion behavior in, 159-160 Single-gene analysis, in pacemaker genetics, 178-

180 Singularity, defined, 42 Skeleton photoperiods, 97

entrained steady-state stability and, 114-117 entrainment by, 106-110

Skeleton zeitgeber, 107 Skunks, visual patterns in, 281 Slave oscillators. See also Oscillators

feedback loop as, 70 pacemaker and, 68-69, 74

Sleep as biological rhythm, 499-510 college population studies in, 502 dimensions of, 501 electromyogram changes in, 501 light-dark schedules and, 500 ontogenetic aspects of, 506 paradoxical, 504 phase control by, 314-315 phylogeny and, 502, 504 prolactin secretion and, 514-515 rapid-eye-movement. See REM sleep slow-wave, 504 temporal characteristics of, 499-517 ultradian rhythms in, 7-8, 510-513

Sleep length, in young adults, 502 Sleep patterns

defined, 501 naps and, 502

Sleep rhythms, CNS and, 515-517 Sleep structure, defined, 501

Sleep-wake cycle ACTH and, 514 human, 259, 314, 316 performance efficiency and, 339 rectal temperature and, 315, 318, 320-324 time schedules and, 508-510 ultradian rhythms and, 512-513

Slow-wave sleep age factor in, 507 CNS and, 516 in rat, 504

Snakes, behavioral rhythms in, 198-200 Social stimuli, in mammals, 185-189 Sorex araneus, 495 Space, clock-controlled orientation in, 299-307 Sparrow

freerunning rhythmicity in, 248 hypothalamic lesions in, 249 photoreceptive input to circadian system in,

244-246 pineal organ of, 247 SCN lesions in, 248-249

Species' daily repertoires, time and energy allocation in, 283-291

Spermophilus richardsoni, 190 Spiders

clock-shift experiments with, 302 sun compass reactions in, 304

"Splitting" phenomenon, 66-67, 121, 326 Spring, hemoglobin level increase in, 477 Squirrel monkey

acrophases for, 219 internal desynchronization in, 225

Starling, operant behavior versus rewards in, 293 State variables, defined, 43 Statistical inference, in biological-rhythm studies,

33, 36. See also Inferential statistics Statistics

descriptive, 22-32 inferential use of, 32-36

Steady-state-entrained conditions, internal temporal order in, 215-221

Sterna fuscata, 278, 382 Streptopelia risoria, 196 Stridulation

circadian rhythm in, 150-151 rhythmic control of, 131 transient cycles in rhythm of, 155

Sturnus vulgaris, 293, 304, 394, 398 Subesophageal ganglion, circadian pacemaker in,

150 Subjective circadian time, 44 Subjective probability, in biological-rhythm data,

37-38 Suicide

long-term trends in, 479-481 seasonal rhythms in, 485

Sun, as orientation cue, 300-305

561

INDEX

562

INDEX

Sun compass circadian pacemaker in orientation with, 62-63 time-compensated, 300-301 vertebrate reactions to, 304

Sun navigation, bicoordinate, 303-305 Sunshine, conception rate and, 484 Suprachiasmatic nuclei

in circadian-rhythm development, 268 mammalian, 252-253 multiple-unit neural activity in, 65 ovarian cycle and, 540 of rats, 68 sleep rhythms and, 516 of sparrows, 68, 248 in vertebrate photoperiodism, 453

Supraesophageal ganglion, retinal pigment migration rhythm and, 153

Sus scra/a, 283 Sustained freerunning rhythms, search for, 14-16.

See also Freerunning circadian rhythms; Freerunning rhythms

SWS. See Slow-wave sleep Sylvia atricapilla, 393, 406 Sylvia borin, 392, 406 Synchelidium, 358-359 Synchronization, in oscillator interactions, 52 System entrainment, versus pacemaker

entrainment, 121-122. See also Entrainment

Tactic rhythm, 135 Taeniopygia guttatta, 385 Talitrus saltator, 301, 354 Talorchestia martensi, 304 T (tau)

as circadian pacemaker period, 115-116 external conditions and, 395 of freerunning circannual rhythm, 395 PRC shape and, 117-119

Teleogryllus com modus, 131, 150, 164 Temperature, photoperiodic response and, 415-

417,465 Temporal information, from zeitgebers, 232-234 Temporal niche shifts, 282-283 Temporal order, internal. See Internal temporal

order Temporal patterns, programming of, 406-408 Temporal program

distortion of, 71 pacemaker-slave relationships in, 68-71 seasonal change in, 71-76

Temporal segregation, in adaptive behavior, 281-282

Temporal specialization, 280-281 Terrestrial animals, lunar-day rhythms in, 368-

369 Tetrao tetrix, 289 Thamnophis radix, 198

Thermoperiodic phenomena, zeitgeber and, 122 Tidal hourglass timing, 364-365 Tidal rhythms, 351-367

basic oscillator problem in, 359-360 circadian rhythms and, 356-361 daily modulation of, 366 freerunning, 356 homing ability and, 355-356 in locomotor activities, 127, 354-356 persistent, 357 phast response curve and, 360 semilunar modulations in, 366-367 zeitgeber experiments and, 358-359

Tidal stimuli, direct response to, 361 Tidal timing

stochastic principles in, 361-362 tidal stimuli and, 361

Tidal zeitgeber, 373-375 Tide-generating forces, 351-352 Tidemarks, environmental conditions between,

351-352 Tides

diurnal and semidiurnal, 332-333 mean high- and low-water, 353 mixed, 353

Time-compensated sun compass, 300-301 Time/energy allocation, in species' daily

repertoires, 283-291 Time series data

general properties of, 21-:22 statistical treatment of, 21

Tissue transplantation, pacemaker localization and, 148

Topological model, development of, 44 Trajectories, periodic, 42 Transmeridian flights, circadian system kinetics

and, 325-326 Trichoplusia ni, 137-138 Triturus vulgaris, 200 True rhythm, defined, 35. See also Circadian

rhythms; Rhythm(s) Tsetse fly

behavioral rhythmicity in, 139-140 locomotor activity in, 126 response to slow-moving stimuli, 137

Tupaia tree shrew, 66-67 Turdus migratorius, 196 Two-pulse experiment, 46

Uca annuli pis, 362 Uca minax, 357 Uca pugnax, 134, 357 Uca urvillei, 362 Ultradian rhythms

defined, 7-8 sleep and, 510-513

Unicellular systems, pacemaker/program relationships in, 63-64

Upward migration, timing of in Calanus helgolandicus, 288

Uria aalge, 197 Uria lomvia, 197, 277-278

Velocity-response curve approach, in continuous illumination studies, 110

Vertebrate behavioral rhythms, 183-205. See also Circadian rhythms; Rhythm(s)

learning and memory in, 193-194 Vertebrate photoperiodism. See also

Photoperiodism endocrine aspects of, 458-461 gonadal steroids in, 458 gonadotropins in, 458-460 light pathway in, 452-453 luteinizing hormone in, 459-460 melatonin in, 454-457 organs and physiological processes involved in,

452-458 pineal gland in, 453-454 suprachiasmatic nuclei in, 453

Vertebrates. See also Mammals circadian rhythms in, 243-253 damped oscillators and driven rhythms in, 253 entraining signals in, 244-246 photoperiodism in, 449-466 poikilothermic, 465 sun compass reactions in, 304

Voles, short-term rhythm in, 494 VRC approach. See Velocity-response curve approach

Warm-up period, in work curve, 335 Waveform determinants, for circadian rhythms,

216-217 Waveform estimates, procedure for, 30 Wheel-running activity, in field mole, 492-493 Wolf spiders, clock-shift experiments with, 302 Work curve

body temperature and, 335-336 warm-up period in, 335

World Health Organization, 475 Wyeomyia smithii, 414, 418

Zeitgeber(s) absence of in resynchronization, 51-52 artificial, 317 - 319 biological rhythm entrained by, 25 circadian rhythm and, 66, 215 common, 66 contrast in, 221 coupling strength and, 220-221 daily temperature cycle as, 96 defined, 95 diversity and, 86 in freerunning circadian rhythm, 12,85-86 in human circadian rhythms, 314 internal desynchronization and, 319 as light cycle, 114 masking effect of, 86 mother as, 267-268 oscillator and, 59, 104, 115,230 pacemaker and, 114-115 performance efficiency and, 340-343 phase relation of oscillator to, 59, 104-106 phase shift and, 13,90-91,325,340-343 "skeleton," 107 "strength" of, 86 strong versus weak, 16, 319 temporal information reception from, 232-

234 tidal rhythms of, 358-359 two-pulse, 116-117 in vertebrate behavioral rhythms, 185

Zeitgeber experiments, lunar rhythms and, 373-375

Zeitgeber pathways, pacemaker localization and, 65-66

Zeitgeber period change in, 88-89, 318 variance-free, 122-123

Zeitgediichtnis, of honeybee, 62, 291 Zero population growth, 42 Zonotrichia albicollis, 403 Zonotrichia leucophrys, 196 Zugunruhe, in migratory birds, 289, 307, 392-

394,396-397,405

563

INDEX