cain shyt permet practily verses of he iran physio omy of mammals riven by the SCN ean acon primary regular Never, {hima ining mar, vein el wor thts ostaine by natal even and change sre food spp, dangers ps J stor sheer ees, tracery of cing mate ah demands of dependent oping Wiha ofthese contingencies met the “> So eels sche of natural election, he fen oniitnginractions of s0 ary lip vives makes some typeof prtizationesental for survive "Nowhere this masking of biological clock gna seen more cel than in ‘humane Not only do humans disegard clock signals, bu the create ther own vonment i places s bizare as underground research caves in the aac {ce shuet or pgantc wotktatonsin outer space, The resultant fred fom the ral stitute of ecolgia niches and topic pyramids makes the story of human chronobiology unique For thes reser like the burst ofthe most dae ng reworks atthe er of nccebratry display, ‘onsderation of ura Goomeisineinaetots pL ookbegenngwiie tmaltythmitinction a Chapters andemchng ge wanternievnein EG Grape i 7 =Human Circadian Organization aids me for wasting tine wel Nigh — Wiliam Shakespeste, Introduction: Humans Are Intensely Curious about Their Own Daily Rhythms” “Most humars in Wester industaliza nations today liven highly manages en ronment driven by the clock more than by any oer factor Weekdays re dom ated by early alarm lok jolts, a dash fo School or the ofc, anda weary urn home in evening for recuperation and sleep Aldhugh weekend are more raked, scheduling tl highly smportant Nata les bate ne of day pay» mich Tess important ole for uate than nthe fey of ruta cultures orf ener civilizations Relax naw fora few moments and imagine living for several weeks avsay fom the bustle a normal sy lena very deep cave, onan underground bomb shor: Thinkabout he lethle nsuch constant environments pecially te Ince ofswrisnentch orl Ina deep cave, complete siznce and darkness reigns, and the empersture remains a cl 50° for the ese yz Water condenses onthe walls a dips ‘onotanously The fact that a henlampbestn penetrates te ploom ony afew fect restricts many activites (Figure9.1) The assignment i tolive ons nommal dally Shedale without time lus anc estimate the passage of ne, Life sucha tne Tess worlds hard to imagine ding the passage o dys, ‘when to ge up and start the day's seine are chaleging sk ‘Now try 3 ourney trough ine and space to tiny apartment seven sores underground beneath the main hospital of Mach, Germany. Called simply the Benker from the German wor: for "bomb shelter thi structure dates Back to Word har The sed fait is accessed vi sree of ily lit step, musty292 Chapter 9 stsreaes and then a seemingly endless walk along, teindingcoridor suggestive of catacomt, High above Inthe busy ety trolly ear rumblealong the wees, fall tf pasengur They hurry to wotk ey each morning td tues home each evening just as sunset announces trenng mealtime and approaching bedtime No cies ‘bout the outside world penetrate he Bunker to ant st the time of day. The assignment it ive here for 8 ‘month in oder to estimate perception oF re without Inatchora radio ora elepone. The Bunker nspasay FRrmished, but plentifal food i provided daly by unseen caretakers who dept goceis and other nec esses ian outer chamber. Te question of ners how someone might react to these creumatances, both Payehologialy and physiological. Perhaps surprising these scenarios are real, an they willbe deed inthis chapters pat ofthe story fof Buman chronabiology research Prvious chapters ‘have made clear he importance of biological tmeheep ing in the physiology and behavior of many living ‘organisms Then of thischaper sto explore human ‘kesdan tiekeeping systems in conta ote wealth ‘of informaton on cada regation a mas, very litle is known abou he longer infadian yt, such s menetraal and reannual cht, or about poste photoperiod regulation of reproduction, For {at reson, this chaplr will eal almost exclusively with cieadan shyt After starting with the early history of human inter cetin timekeeping the chapter vil progress fo modern ‘Sady methods then give an overview of reunaing Prods and enranment in humans fllowed by ay [Ena consderation of the major ccadan shyt in humans, The final section wil consider the special developmental sepests of human chronobiology in infants an nthe eldesy. The stage wil lussbesetfor Chap 1 to survey mal functions ef this timekeeping system, not fonly in 2 mesial content Sut alo it Felt inva ring the abr forking nigh. Interest in Time Measurement Has a Long History" Human avareness of dal, monthly, snd annual eyctes stretches back (2 Drehistonc ehillzations™ Enter chapters hve orphan that all living clocks depay the seme Basic tinekerping properties, although the siructuralcomponents may differ wide ly. The mojor reasons for considering, Fsnans separately will emerge a the pier proceeds, Tne stating ex universally on ‘ur plat. messued in days as Earth tre ont, months a the moon ort Earth or yeas as Earth ‘orbits the sun Inthe pst mane must Rave repo ‘sto thee cles hen came fo hunting to secking Safety during hows of darkness, and to cosng net Fer irgating land and planting ood. Records of asrologeal and ealendelcal tables, together with aligunetof some ef the stonehenge inocthern Europe See Chapter I) andthe pyramids in "Egypall point oa amen awareness in erly viz. tions of these eternal eles (igure 93). Ancin! peo Pes, however bad itlespprecstion for internal Ing Some ofthe atest ations of internal ning ‘nan cme om esaement of human body te ptr, A Bish surgenn in 1836 noted dally ane. ure flituations not asain with fever However Fisteas and oer datastrongly supporting temper ture hytims in humens were negated for many ‘decades bythe dens of two very prominent European physiologists a will be daca hort The minds time sense has important implications for time perception ard use by humans Subjective time adie memory sre wique spect of human time perepton. The great joy of tea inthis book concer plysiolopil tine mensureren, In humans however tie awatenes ea itl aspect of de se, The capacty fr concept formation slew nes, en fhe higher ling proceszes tends gra Iy modify the appieate of Phystlogica ime mess rement to Behavioral output in humans. Walk an increne inthe pace of moder industrialized societies,wi a quanttive measurement bu rather with pro ducviy Times worth money Asa esl the sana istheamountot work acomplshed, sot the absolute Aime spent ons ak Humana se owe often concerned therefor, with subjective awareness ofthe pesage of time than ith eal ine Teasvely itl s known about brn egulationo ‘he mind's tie. In some fashion the human brain Uniguely endowed to onganze and priori exper tees nto cronlagy of memory events and to eal them at will The lat bran ats 2a see fo magni tr delete cera evens and then rept these ether hareterm or Ing erm memory. The ihm poe "eaaly clear ined individuals A Ward War et fran may real with vivid etal the inateby minute evens of D-Day 30 years ago, ut fal orca what he Se or break ost a hor enti, The baa acai Nppacomps, an teporl be areimportant proces for ine memory Most Current ivi comes os bar damage pets Human Circadian © ganization 293, ‘Slnyon (8) Solar tytn wise over Hawa ‘When the bas feral ju neon of te event emai bu the conology of eens may be loa the crcl ppc pus sda ‘ge, serous amnesia may ensue, wth os of tility to store new memeres Damage tthe ‘eypeale adie Seirpampss pe texof the temporal oe especialy wes Se inary may permanesy delete years ten decades of eltmenoties, Very lide ion about ne tamping bythe ran, Te races by which he Bat acne specie Hime and place to an ever sort and then reeves ton command, bil pooty under Interest inthe physiological functions of ‘humans grew inthe ninsteenth century famsan set an Pao (40-95) Indie associated withthe iow cept of readianthythmein humans, Pavlov as tran the Nobel Prac in physiology and medicine in 90 for his work on neural eg Tatiana digestion in aromas cleat tiated they cnr itn hd immense impact on the sdentie word of is day an contin ounce rang hry Inthe tel of paycology even toy Hl they of 0 cile clase! conning stated that cet innate Tesponsescaled unconditioned exes could be mod ley repeated sosocaton wih an iil sinus toachieve a elabl ne spans For example, the ‘ation dogo the savory snl of mea could be co {tone tour in sponse a sul Sane thence : : te atmosphere of lakeninetet-century phyla ogy was charge wit daserson oer waters Versus Tengo the bass overeat bhava esponees. Pavlov tipped the Blane strongly in favor oeeming ‘wih reg to nat cicada relation in ane vows ae tributed ny ternpeatre hy anicy to acquired condoned respontes and profoundly delayed aceptance ofthese tomperature ths a tmanilestatonsof tue edogenouscieadian timing,294 Chapter 9 Ignoring of marked daily body temperature ehh in ‘Ruma was de uleneof the mest prominent French Bysilogst f his day Claude Bernard (1813-1878). eznad noted that and marasmus del with ge Sule eavcemontal changes in emperatr, ater eal dbl groviy and wind. He documented any YSio~ ogi mechanioms Ut protect animals from exeeme ‘yscland bok forces Laer the concept wa expand fly Arneian pilot Walle Caren, who coined the em homers for this monsmenta concep com ‘sming physlagcl protective mechan. Homeostasis the maintenance of the iternal sriliew ote ody within vor arco brit in ope of the trdeney fora individu ssuroundinge and act ‘isto change For example, reflx responses predsely regulate bo temperature esmoic pressure an com pln ofthe blood, and oer responses. The concept [Sf internal constancy of waimn blooded vertebrates ‘hoeed the minds of centist fo aceepting changes body tempersture a3 normal, endogenously pro- |fommed response in eal humans. ‘canckS IN THE HOMEOSTASS THEORY. Acceptance of ferperatre dythms in humane asa nocmal even as ‘very slow. The medial fled tended fo view temper {ore deviations from a fixed norm, such t fever or Iypothermia asa sign oflnes: Even today most home {hermometershavea red arow at SBT OFC deg ating rolled normal temperature The oustanding, research of Jargn Aschlf on human ternperatre ro "ation was veeyefecive in gaining ecepance st of ndogenou temperature vy and eventually of human ethic in gener Study of human circadian rhythms accelerated very quickly after 1950" (Few BXAMPLSS OF HUMAN guTYTIS. Many dally chavs physiologic and biochemical hts ‘humans atin synchrony withthe daly light-dark eles (Figure 93}. few representative cases suffice bere wo Inwoduce human ‘hythms, The cbserved ‘thas tend to fal into one of ta groupe The Rest ‘roup contains shyt that peak during the day and Sreansocintd withthe activity paseo the individ Core temperature, mental performance, physical ee ‘ency gastrointestinal activities, blood pressure and heart rate and hormone adrenaline eeetion al tio this group. The second group concerns rhythms that shows peak during nocturnal seep. This group is Salle and inludes growth hormone and melatonin eta of human tydicfunction wll be considered EARLY CAVE AND BUNKER EERMENTS. An endogenous Component of ythmic varaons in physiological and w reader) © nT) FIGURE ®2 Repreerative sy hth in an LD 168 bed fr lhe Sulpes wee eee ght ke {Eig te day (A) Seeding fests (Coe sap Limpert (©) Growin utmane tercton (0) Cor ‘Slane e)Usery pots Thetime ator) [Fercslantour whe he nas ra ime ous fmm Mon Be ndSasmon 98) ‘behavioral variables in animals such as hamsters and rats ha bon demonstrated by stdin of act pt terns under conditions of constant darkness nd con ‘stant ambient ternperatue. The best way’ to provide inrexpertnents Because no tne cus are present in Such experiment Utemethod i deserbed as eeu ‘lng protocl Before early work onthe endogenouscomponent of human circadian shythms becve feasible, spel environments and experimental protcls ad tbe crete > eliminate the elects of surroundings of, | lest a theses on te al bs irl experimenters aed some very £2) unasol worker iabortres Unde Bost ‘ground caves uclnedsounproofed chan Ber, or the Arce In summer provided | relatively constant conditions Inch oe Hons, much of the complexty due to 6 Ihumantnteacione and innown iting 8 conulions was reduced or eliminated In SOF the 830s the mous Americans phys? sa fologist, Nathaniel Kletman, and isco non-24hsleeprvake cycle for up to 4 ‘werk monet to messur thet boy tem perature rythms. The ain question was Inet the yt was inate or leaned ‘The data were somevshat ambiguous bt provided evidence for an inate nature beceuse body femperature cycles showed independence fom seep sake cycles ‘Aroha ave experiment was cared outa 1962 by young ave enthusiast named Michel Sif He ved B7sfootdeep cavein the Freeh Alpe whoa watch tnd recorded hit sleep-wake ohy- thus for 62 days. The cavern nae tollydakexept forthe light fom 12)- hshaadlamp, andthe environment . Wwasdankand bone chiling during Fiventie ny By rar ofan pe ‘ibe telephone Site nformed is Fred very ime he went tobe or woke The rest was an seo (ERE Se Ee ae) Sleep-wake rhythm (Figure 94). Remarkably he lst rack of 25 ays ‘because he did't coun presumed ‘napping periods and thse tue outtobesinalghs tn 1957, Mary Lobbaa and her collegues lived in Spitsbergen, 2 Norwegian arpelgo inthe Acie (Ocean ring the ime ofthe su ‘mer midnight oun. Phone cues Inthe continuous light were min. ‘al cring the sbjestive day ard at ight the subject sep in Backed tutes Without detest jects Knowledge watches had ben doctored provide either 2 cr 2? I days instead of 24h days Nine solar (24h ays ere equivalent to ny days with 2 27h perio Human Creadian Organization 295, FIGURE 94 Body ompertre hy of anvil vng ina deep Tighter ove fore dap. ae Fale 8 Such protocol, st employed by Kleiman in his cavestudies, s noe called forced desynchrontzation, Bnd tis widely used in human experiments The ‘arabes measured showed diferences in the relative importance of exogenous nd endogenous consitun's (igure 95). The slep-wake eye showed 8 27 8 di ‘ins, ight may stil be aeting ae 25ental agent nity though melstonin suppres Son. Excision a al photic mul therefore eel before a conclusion si nonphotic synchronization in Bn indivi canbe aceted, any nonphetic entra agent ext in humans, {hei rare eran iene I ght subjets bt not inblind individu naps taken cing vacuo day cause pose site oppostet light pulse fic at ‘hesame tne of day an thee hfe ae probably dae toa change nigh exposure nsociated with shutting ofthe eyes. The postural changes associated withthe sleep-wake cycle ae accompanies by major changes in Sympathetic activation and os pay a rle Te fet af motor act partly pica execs has Been Investigated by several groupe A feof the stad sup est very small phase shifing due to exer, ‘One ofthe mast every studied nonphote ym chroizes is periodic administration ofthe hormone tneatonin: The SCN conlins elton eeptors snd {ima shies have demons thet exognons mel tenia can phasechiftcreadan shyt in vivo an in ito. Adminiration of melt fo renuning blind Individuals can symone the sleep-wake ye ed the “endogenous nelatoin ryt in some but natal ‘ius In sighted intvcuais melatonin anton ‘an faite recnainment ef endogenous cadion ‘yh to an lined stacy ard ight-drk cycle [APR hasbeen measured for melatonin ingestion The Shits ten tobe the inverse ol those pred by ight ‘Melatonin ingestion in the aftmcon and eat evening advances theceadian cack and inte nights ay ‘ming dels it Beene of ese phases fs, melatonin hasbeen cle an teal syntwonten “Theroleof the endogenciaty generated rythan of ‘meltonin inthe human cieadan eee cern bu the phase shifting elles a ligtand melatonin appese to minforce each ther Melati normally seed ‘overnight ind hors darkness nd ight suppres melatonin secretion by an amount proportional 0 ight intent. Bright ight dnetly advances the phase ofthe cicadianelockin teeny morning at afte the enero in heigl ih ‘ippreses melatonin seretor and prevents he phase Aelaying effect that melatoni woula have exerted 3 ths me vin itsown PRC, Temporal Regulation of Several Physiological Functions illustrates the Importance of Circadian Rhythms in Humans?» Humans are more aware of he sleep-wake rhythn than anyother circadian rhythm’ 36 CCENERAL PATER, Humans ae acutely aware oftheir Slegp-wake ehythme. Atypical person Boe fo lee at 2800 to 2600 local time and ies at 070) to 0800, Human Cicadian Organization 309 Marked vasauons in bstimes and ake ines eit both within anderen age goup Formos Sling in igh nds aed deen fac ke shew scl clus an etal schedules oF hilren ae major ators ne sop nae sched ‘On weekends and holidays, seep reels may hanger some people ty tae sap the ae neon i adaion he major octal dep epcode Thus the socal eveonert ea ma deena of slp wake schdle, een oghitis net eceaany insynchrny withthe cxerelmatral ight cycle Sendra expenence conn calor tronzing te sagpvake sched het Yel secu The seep-sake cyclen stud and described ty vasous teagan Suess sep ad ‘wake meson be dn seep des Suc can locement the timing of nope op welas estimates of Step soni Alltec pride werd normation on the oul of sep Reoring oy etc can Se conducted anbuelyfr paged pred of time Theda nde seep dan Sd Sep ef Sine. Atel are bing ed ince il Sto nd lbortory sees of human seeps platen The sand thease sao is palo nogeaphic recording a slep. Date collted include tlectoeneephslograns (ECS tard ran waves, ‘leo arog tert ye movanents and dee tromyograns to recor musts tone On te bas of these vale the eords on be ated ar wake ‘ss norapi je movement on REM) ee, ‘pid aye movesen (REM ssp Now REM seeps “Sully further subd to ape through fon theta theampiude and reoeneyof he EEC roves which conte ase veal or ay Spel anata Sager Sand fae cacy Kaown tion ware dep. Such methods ae edt quan he ine couse ftp features of he sleep EEG such towne chercy sow wavealow-have ep 02-40, and Ecepnenomens such ashe 1208 Re slecp Sn des & poof thse apse calles hypogean ‘Atypilatomaten ofthe sep ape ean ose dur noral actual Sep gu 9.0) Now REM Snd REM separate witha ltndanpesity approximately 701090 reg Teds of EM spire aswell a te doe of pieye movements SEEM, nese ne Cou esta seep. Sowa epi prominent ne ft per of he slop peved sr amon aba in th econ Pall Contdanbesnge? sep present nthe sco al ofa seep pei Seep ind he ltadan aeration ‘eveen ni REMC and REM sep reatonpared by many changes in pyilogy,oning tom heat rate rather ne varity to flosmn ren concenaons310 chapter 9 Lalla {area Et = ste at ia FIGURE S16 Baseline sleep. Tine coir of spade acy (A) EEG slow-wave seit see sg nd em itoinconcstratons (0) ing 9 m9 istalseepeptode fae yenead mate Couey oD} Dyk) € Dliond ‘Anentraine young aul ing in society go © exp shout Oh before ts tty temperate mini, hic Daring freeraning conditions, conta sleep fated lone othe ternperatre mimi and termina ‘hyn, The change inthe interval phase rafonahip betwen sleep ae core lemperature provides some clues about the cicadan regulation of sep structure. The tine course of low-wave lep reas unallred, but the time course of REM sleep changes during free ‘tng conden The ronal crear the duration sfsuccasve REM epiaode over the courte of nights Slop salted, and REMsleep sis award the begin ring of sees. uring prolonged temporal isolation a change in the phase ratonhip bevween the slew and the body temperature cyles may precede spontaneot= intemal dexyrehronizsion The core temperature “yt continues th eri slighty greter tan 2. In While the sleep wakecycle might be about 161 oemare frequently about 32 (Figure 9:17). As Figure 9.17 Indiates internal eynchrony preva! dag days 1 to with sett and empermire pestods toh egal 0 257. Alter the star of desynchroizatin on day 15, the activity eyele had pean periow of 334 hand the temperature period was 251 8 "The simplest interpretation ofthese data shat the Sleeprvake cele andthe cor temperate syn are {ovemed by separate ociltrs, Normals the to ar {oupled in the presence of an external ig-dack cycle But become uneoupie in the absence of in LD eyee ‘The period ofthe oclator driving the temperature cle is close 1024 hand stable the plod of ‘Seep-wakeosailator sable and can deviate con ‘ably from the cid ange. Notall cents ‘gre. Some researchers have agued tha he apparent Spontaneous cesymehronizatin lean aac of how Sibjects designate sep episodes alter major seep ial iin = * = » — ‘Ste wate is open Br lee ner angle ‘eopecivelytamperstire asim snd mina plated Irie to Show decent sa angles [ther explnaon seth om ee 308) Human Cieadian Organization 311 episodes or naps: However, spontansousintral des {hronization benvoen the slegp-wake and temperate {eles has been ported insti in ehh al major ind minor slep episodes were ecored and report ‘Others have pointed aut ta spontaneoe deny cht ‘zation sot observed in nonhuman sia but fe data are available to support that contention Sleeps rorly recorded in animale athe ecg of le vanabes ever oer Spontancous itera desychronzation maybe on artfctin humans of the profound influence co ‘ious thought during wakefulness about when 091 Slep. Every corel functions change dug spor "ancous intemal desyncronation. When inital are asked to etimate the passage of hourly time inter Vals astng ereation etwoen duration of the hourly ‘estimates and duration ofthe ovake episode i seen Possibly along estimate of hourly time oes lang with Along ime tine sp. However, te estimation Fmt intervals dung spontaneous intra dey “hronzation continues to covsry withthe core temper ature rythm. The neobolglea bass of hee nt Ing phenomens emaine unkown Spontaneous interaldesynhvonizaton mean thst, sleep episodes are initate st many phases he core body temperature cycle The distribution of phases which slog initiated, however, snot uniform. Very few sleep episodes are initated during the interval of about to Gh before the temperature minimum This phase has ben called therefore the wake mainesanee one: In contrast, most sleep episodes ae inated appronimately 380 Shafer the temperature min, Under entrained conditions these wo phases ofthe ‘ore tmperature shyt oecur clove to habitual bd ime and wake time, respectively. The duration ofa ‘asles from approximately 4h to about to hThe lenges sleep eptodes are inated inthe hours before the temperature minimum: the sorta episodes after minimum. Wake duration mist nso vay geil but they donot appear to havea major influence on sue {quent slep duration. Instead, the phase ofthe cad fn pacemaker as Indexed by the core temperature ‘yt, appears tobe the major determinant of sees cverlfeatres of sleep structute are als fected by theccaian pase which ep occurs, Sloe sleep ad REM sleep ae affected very diferenly Sh Wave sleep declines during seep episodes inated a alleeadin phases In contrast the ime course of REM, Step is mode by te ccadian ps at which sep fccurs The peak of the circadian rhythm of REM propensity is oated ator shorly ater, the mim the cor terperatre yt Figure 918). These data provide compeling eviene thatthe circadian clock312 Chapter ee z a8 s ‘Grea pine Sieepceendent \ \ — ctvingthe temperate shythmexets a mujra {nce on sleep dation and REM sep. Sep length depen lo on th eof init ingslecp and maintaining sep. Experiment ave ‘tn prior osvesgate hon eay Histo fal asleep at diferent times ofthe day. One prota onited of dividing the 24 hol he day i 725g trens of minterech nad of se sen thevelinter waned 7 mst fl aleep inthe comot ofa slep nborvory, and a econo lec activity nbn ated the oun af sleep obtained The sbjea was avakened Ssiep the en the? mead weed toremain awa forthe next minut Te net minute segment wo tested nthe sae wy, ne the protocol coined ul the endo the at Segment The eas indiatehat falingaskep was ‘tera rightand more difcul uring te day The ‘Bilt Bl snap nvrored he eon ton ‘ey temperature Low body tempera dang the later prt of the might mas aes th rater ee of geting t seep. The higher the tee. Prraure the arderif was fl eslep andthe es ‘Sp nas bain ina mine segment WAKECCLE. The slep- wake gee and the compo sii of sleep are ot determined soley by yt te factors sleep ning depends fo some extent Sls onthe degree of stimulation Sleep wil be start derbi the subjects bored or er exciting or Interesting ceumstances preva ‘Sleep physiologists have knovn for many decades iat sep structre sn slepines elf ‘depend in parton the duratonet wakefulness The ‘esulls have provided evidence that homeortatie processes contribute oslesp elation and operate ‘bith eave slow tne constnts A subjet en be Seprived of slow-wave seep by couse stimulation frat does not induce wakettnes Aer cept Slowsrave sleep abounds in the ndsttbed pare ofthe night the sleeve sleep dep ‘ation lasts teenie slep ese, then rebound ‘cecurs during the subsequent sleep epiaode Ap RGUKE 9a Ile of din pa ead ie antes esrbing scp Wileuloes sng Sep {Epo (hy REM se (5) on Rt scp) me ii spe activi (Dy soars hone te pert) Natit he non tae Srablep ve, Zr degre (pene ng ‘like led diol heed fon of ety ‘appronatly G63 raf ay webown nt Al Ae ‘okt Caer 1985)parently slow save slap segue very accurately. ‘Quantitative aaa ofthe slasp EEC shows tat slow wave sleep provides a quantitative measure lp omeostae Siow-waveslep seems tobe very tense seep, te ‘normaly ccentated in the i part the sleep pe {Asleep drive deceases the moan! of slow wave Sleep lo decnes Asa result exten othe sou In tum shoul obliterate the need fore major irae insleep duration ‘Suppor forthe notion of an ines component sleep comes rn dose response stds ofthe elec cutecurtallmenof sleep duration on sbsequent ale ‘ess and performance. Resulls india hat the fist 210 ‘Thos aleep contribute the mst tothe neovey of ler. ‘ness and eleny. Conversely, pin the Ite fer. ooa wil emporriy improve perform, 2 wel 5 reduce slow-wave lep inthe subsequent nochuroal sleep episode. The homeo facet of REM soe re Ulaton snot os easily quantified, bu he presence fuicha mechanism undisputed ‘these ests ply that he ep oscilstor tracks how longa petsn as been aa or slep, This tharacteriste makes the oclatr suse for fling thehineionof seep hamessasis hats, cntraling the 2eragelev! of siep debt Sleep-wake belvior ae 2 malo determinant of slap propensity and sleeps ture Tt is pare ofthe carl loop that genertes the sleep-wake cation srl a hangeobehavioral sae willimmeditely reset the ocilation. The sleep hore tetas hction, ofcourse. very eiferen fromthe le Afsleep-vake behavior contbutng the oilltons body temperate or miata fhe ts pein the absence of sleep ‘TC EACTORS. The sleep-wake cle segue by he ieraction betwen enilstory and homeosai pro This concep hasbeen very sucess ncorporat. ‘6 nto matheratcal model ofthe shep-ake Cycle These models have eto many preston relat 0 sleep duration an slp struc in brary expe rents and fel studies in which the slep-wake syle was desynchronized from endogenous cecaian eth tity: They have ls provided insighs into the eps Taton of waking pertrmance Finally negation ofthe concep of sleep homeowtns and ciation fythisy offers a functional explanation forthe interactions betwen the wo proces an thelr pase lationship during entrainment Under normal conditions, ciredian time and sleep-wake histories change in synchrony. Humans ‘wake up inthe marin es hous ater the core fe perature minimum and stay awake url apposite Human Circadian Organization 313 ly 6 befor the minimum, Dusing the waking dy. te awake increases progressively ang during sep, homeostatislep pressure disiptes in a mote ‘Slcp propensity consistent highest at theca lan phate that would normally orc inthe late ‘evening hours and lowest tte cada phase not mnally occurring inthe morning hours For example, the daytime and then shorter afte he ost f noc ral melatonin secretion to rech a minimum sea he time ofthe minimum ofthe core temperate shyt, Thus humans normaly goo slep shorty afer the pet in the cieadian deve for wakefulness and eke up Shot after the peak the radian deve fo see oe Figure 98) The actors influencing the sleep-wake cycle have boon considered in some detail, na only because hey are well documented but also because they ent such 2 profound influence on behavior and the body a8 & whole. The slep-wake eyele produces important ‘hangs in poste sei igh exposure, a food Intake allot hich are important fen = chrono ial pint of views For many vate the fet of he sleep-wake cele contibte tothe exogenous masking fe of Ue creaian yh The vane that have Srongexogenous component appeat to aus in phase ier the sap wake cleus been phase shied. load pressure, bert at, the secretion of inalin and growth Formone and many components of rinay excretion fe eumples deed, the phases of these rhythms are often cexpeesed interme of he reltonship o micsleep ‘Separating the endogenous component tat aus Phase wit he lek markers, sch a plan eatin Sn cortisol om core temperate egies 3 costa ‘urine oceddesynehronzaion, or demasking othe Core body temparsture has boon studied extensively in omans=™"" Core body temperature reflects the balance between et loss and haat production mechanisms, The most reliable measurement comes rom 3 rectal thermistor Core temperature hasbeen the sljet of many ie sn studies, primarily becouse i relatively easy to measure and hae 9 clear endogenous componest Temperatare es rom ey daylight an then act decline afew hours before habitual bedtime as Figure 5:18, Typaly tals rapidly when person gestae and reaches a minimum nthe later half ie Brain emperature shows a similar ime cures “The range of body temperature is typically 36 to 575°C, though deviations outside this ange canbe314 Chapter 9 observed in sesponse to stenuous activity hot or cod Showers, and changes in posture. Core temperatures protected by svarsy of thermoregulatory relleses, Frciing the tne of citneour baod vee sweating, snutshiverng These post of thermoregulation S hgh get Sey of mythic inctonsim hme ese no wily Sexpidin ntl the ident cry Graal estilo ccadian Sing ebln! rom asec Fon organs were scepter hua phys ‘gy. Overt cyt was ecg othe ration ‘ofa feerunning nema selesutand pacemaker th Synchrntzes he external world Teche ofthese Syctronizes isthe igh-dark je of the sla dy Ely eszach used sts such as caves, underground rooms, or arte continuous daylight in summer to pro= ‘ce constant conitions for studying the endogenous ement of circadian rhythms. Later moder hitech human tries provided convenience, comfort or the ocarch volun atone of making in da er {ng ae the sled staff mde to cary Ou hua sd ies succesauly; Humans have unique advantages and lncvantoges for cacian research "Among the innovative prtocos developed for the ‘new faelites were the constant routine and freed Seaynehronization for aboratrystdies a well at ‘rious methods for subjects fe ving eld dion “The method coniomed the fundarertal properties of readonly un humans hat hd bee almost Universally documented for nanhutnan organs Humans fern constant conditions with pe od of approximately 24.3 h. Many endogenous and ‘exogenous factors play a ole inthe overt thm ou ute of humans: The environment act nt only 28 ‘masking influence but also spn entaining agent Entsinments bought about altnet excuse by the light-dark yee: Humana are very sensitive to phase stifling even by very dim light The phot phase "response curve documented for humans resembles tht tf other urna animals bt tends to show light ‘esporsiveress inthe subjective daytime. Very Few other {ntsining agents are known for humans one note ception is exogenously insted melatonin. ‘The primary pecoraher ithe suprachiasmalic cleus fe ventral ypothalamus. The elects of Ue SCN permeate the body beause they at on core te perature, plasma hormone concentrations, out of the sympathetic nervous system tothe pineal gland, and thesleep-wake cycle “The most widely sted human rhythms are core ‘by temperate shythm and the seep-rake cyse ‘The rnpeeture eel bas m srong endogenous come ponent buts naked by activi. The leprae yee inode by many exogenous factors, end appears tact in concot wih mechani thal ann cep homeostasis: Other important cecadian Functions in humane inch the alefnes nd metal performance ‘te the physi! activity eye the plasma corte ‘jl and fe endogenous melatonin yc Te expres- Son of each of these thythm shows considerable interindividual variation ‘Developmental progression inthe expression of sya een from bith ood age. Rhythms in newborn humans are largely ltradan with pei of ‘otuiness every Soh, Bt tis thm cones inthe fist yeor ino crac pattern Inthe elec the amplifede of csadion rhythms is commonly refusedCONTRIBUTORS James M. Witerhouse and Patric}. OsCoesey wish thank Torbjorn ARetsed Jonephine Arecl, Chaves A (ets, Dek Jan Dik David F Ding ted Kenneth P Wight J for hele acl eoiributecs Chapter 9. STUDY QUESTIONS AND EXERCISES ‘The following est adapted from Horr and Ontcrg, 197) asseae fan nv isa mom Spe or an evening ype or tach of question, choose the mos appropriate anne ‘Then chek he answer kay below 1) What ene would you choone og up if you Es00-0600 B-O60040790 «(790-1000 ‘41900-1100 2) For important busines mestings, scheduling 2 the peak of mental powers important ‘When would you prefer toeheletis bese? 2 ORONO b. 1300-1300 1500-1700 a Yom-2100 What time would you choose tog tobe if You were entirely fe to pan? Man00"2ta0"p.2100-2845 «215-0030 d.cns0-014 4) Aid wishes ogo jogging and suggests ‘Sarg ot 070-0860, How woul you ‘Seseibe th ine for you 5. Good 'b. Reasonable © Dial iit 5) Every individual performs physi work best at eran ties of day. What me would be = OROO-1000 b.1L00-1300 « 1500-1720 4 S9e0-2100 6 you have to go tobe 9 2300, tow woud youtel? E°Noat al red, and unable to ge to le ‘ulchy . BAe ted, but unlikely to gett sleep ‘ley Ay ted and aly to get to sleep quick A Nanya very likely to gato sep suey 7) Wher you have ben up for half an hour on a ogl working day, how do youre? Very ted Befay tied’ Fanly Fetesid deny refed 1) At eat tine of th day do you feel bet? 3080-1000" 00-1300 ¢ S800-1700 iom-2100 9) Another tend suggest jogging a 2200/3500 How would ou debe this 4.very Human Creadian Organization 321 8.Good Reasonable « Dif Very ‘iat Answer Key Scoe each qution with he follow Ing pints then total the points. For questions 1 tough Sand 8:a=1,b=2c=3.d=4 For ques tins 7,and a= 4.b=3,c~34=1 Yourscore fa range fom 9 1 36, Here ave the suggested Scores for detrmining activity types 9-1, de hitly a moming type: 1518, mogeraelyatoen Ing ype 20-25 an intermedi types 2-50 mod erately an evening types 3-36 definitely an evening ype Most individuals sore behseen 20 and 25 only bout 5% ofthe population sores Ito less, oF 3. ‘ormore. This tye of questions fe wdely ued experimental nd thas been tasted info several lngunpe, (One very important relationship pointed out in th chapter concerns alertness felatve tothe boy temperature cycle Design a siple experiment sermnstate ths lationtip Showa here ian Slernes sale that hasbeen widely sed in Alertness sale Reins Coon | 7 Filan somtgee Sd St | & Mosse fain ake moder fague 1 ene eat an nares | ‘rennet mae nigae Pana protocol or measuring your temperature t regular 2hineval thoughout your wake pid Many ways are avaliable for mesturng temper ature but the most valle for students probably» singal socalled fever thermometer From the lea drugstore. To gain an arate reading beep your mouth cloned and breathe rough your Hose forthe al'S minutes of the reading and tla rom aking drinking, o ting for 15 minutes before the sat of he temperature renaing,322 Chapter 3. Wht eft do naps and aivity have on alertness durng the day? Have sje take» nap 903 ther ake mensurements ater the subject ‘wakes. Compe results with those taken atthe ‘ne ime of day onda without a ap. 4. neni what are sore posible masking fetes in your dally route that could interfere srith te epee reading? How could these cor be educed during te exporoen? 5. This Chapter has emphasized repeatedly the problems of masking in human research, tcl in field sade How el do constant Foutine and forced desynehronization acount for asking eset? Quik review Chapters 2nd 3, Sha summarize the kins of mashing tht are seen inponkuman cieadian shes Why Nas 0 much Tee ffort been devoted fo demasking of nonhuman anna ala than to dematking of human data? 6. Eerie increases body temperature, but does it ddosoequslyadiferentceacn times of? Pan a shor Iboratory exercise fo est his {question remembering hat thi chapter has faporied about exogenous fect ofaviy on ody temperre REFERENCES 1 Akt nd Gere 1 Th ean vr of SST 2 SEES. PS Stew 1 Hae. Piper ml Ware 18 Ret te * Gitomer 5 fom FL er Pie Conn, and Ages ig rites Ranma ‘cipal sgn Seoer hen Semen oe Gee eee ee 1 Sowa CM ads Eh rg Pub Sinaia ft Par Mao CJ. Aa MS an a 1s Sg ng Te Sopestac kas yap" PNY 7 =e ae aa > Senn ma tt Se eee ie dee Bag Se “aie gig saammecss 2 Daa A 2 Renting wn SS. A 27 Se 25 Df andC A Eile 19M, Pn ig of the Shatinuibeltnaiar aa? Dut an il 2 Gat i Ce rR Se eeu Find) so © Rt pengeeme ee Eel epepe rons Pe ow eS on ss igen i Sel eew 2 5 Hoa, nd Yds, at egos cn ea Rese tpoyareetesetirssnat' aa Human Creadan Organization 323, Aye el ial ag at