Third International Symposium on Natural Mammalian Hibernation

Third International Symposium on Natural Mammalian Hibernation

CRYOIIIOI,OGY Vol. 2, N o . 2, 1965 THIRD INTERNATIONAL SYMPOSIUM ON NATURAL MAMMALIAN HIBERNATION Ramscy-Wright Zoological Laboratories, University...

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Vol. 2, N o . 2, 1965

THIRD INTERNATIONAL SYMPOSIUM ON NATURAL MAMMALIAN HIBERNATION Ramscy-Wright Zoological Laboratories, University of Toronto, Toronto, Ontario, Canada September 13-16, 1965 SI'ONSORS : National Roseavch Council (Canada) Nntional Aoronauties and Space Administration (U. S. A.) University of Toronto (Canndti) Ofli(:e of Nnval ltes(":trch (Chi(:'lgo Brnnch Oltiee) O I I G : \ N I Z I N G COM M I T T E E : DR. KZNXETrl C. FISHER, University of Toronlo~Chairman 1)1~. EDtrAm) S(:'Jff:ixl)au.~L University of Toronto DR. C'.)~AI~L),:S P. ],¥.~tax, Harvard Medical S(:hool Dl~. FRANK E. SotrTtl, ,IR., Ihliversity of Missouri 1)1¢. ALm.:R'r R. ])aWE, Office of Naval Resear(:h Review Papers 1. lq~ysiologi(.al variation of the central theme of to{'pidity Reviewer: ])r. Ja(.k W. Hudson, Delmrtment of Biology, t{icc l;niversitv, Houston, q c.. s, U. S. A. Monitor: Dr. A. Petrovic, University of StrasI:)oura'... St r~,sbo~n'g, Fr.mce 2. Relation of external conditions to onset arid cessa t i on of h lbc rn.~(.ion Reviewer: Dr. Erie T. Pengelley. Division of iAfc. Sciences. University of California, Riversid.e, California, U. S. A. Monitor- Dr. J. S. flart, NRC, Ottawa, Ontario, Cana(la. 3. Belmviour eapabilit,ies of i lie centr'd l:tervous system during hibm'nation Reviewer" Dr. Nicholas Mrosovsky, Depart;ment of Psychology, University College London, Gower Street,, London W.C.1, England Monitor: Dr. R. K. Andjus, b'm "x.'(.rslt.v , " . of Belgrade, Be]grqdel Yugoslavia 4, Internal rhythms of hibernators lteviewer • Dr. Felix Strumwasser, ' " Departrnent of Biology, California Institute of *,'1hfornm, .... Technology, Pasadena, .C, U. S . i A. Monitor: Dr. Curt L.-Riehter,: Johns ttopkins University, Baltimore, Maryland, U. Si A. 5, Control of body temperature b y t h e central nervous system, a'lld its relat.ion to hibernat ion Reviewer: Dr. H. T. Hammel, John B. Pierce Foundation Labs., New H.aven, Connecticut, U. S. A.

Monitor" Dr. E. Sellers, University of Torord~, Ontario, C,~ nada. Host-lntrasitic, lelatmn, and pathology in hilmrnating animals • f ' V " m~(l" ~ " ~ " R. Dr. Jerome P. ,.S,' "" Infectious (.mn,ut., Processes " r S.:hool ",¢ . . . . Unit,. "USA1 of Aerospace Medicine, Aerosl)ace Medical Division, Air Force Systems Comman~l, Brooks Air For(.(.,. Base, q cxas, U. S. A. Monitor: Dr. D. E. Davis, Pennsylvania State I 7ni ve/'si ty, [!ennsyl vania Still: and water })ala.nce dtn'ing hibernation Reviewer: Dr. Kenneth C. Fisher, Departmen|; of Zoology, University of Toronto, Ont'l rio, Canada Monitor: Dr. P. Suonmlainen, University of Helsinki, Helsinki, Finl.'md Iteart and circulation ,, Reviewer: Dr. Bengt Johansson, Cardtologlcal Laboratory, Department of Medicine, General Hospitall Mahn6, Sweden Monitor- Dr. C. P. Lyman, Harvard Medical School, Boston Ma~sachusett.s, U. S. A RelalAonsMp of sleep to manlmalian hibernation Reviewer: Dr. William C. Dement., Depart:merit of Psychiatry, Stanford ' University School of Medicine, Palo Alto, California, " L. S.A. Monitor: Dr. B. R. Lan(tau, University: of Washington Medical School, Seattle, Washington, L. S. A. Metabolism of fat, particularly of brown fat, and thermogenesis in hibernation, including "I].






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'FI 11 I{ I) I N T E I { N-VI'i ON A I, SYMI'OSI UM ON It 1BI,H~ N A T I O N the (it~e.ev(. G. ],intll)erg, Bioasl,ron'mtivs ],al)., Nm'tl~rop Slmee'ttories, II'twl horn+,, California, U. S. A. M e a l i e r : I)1". A. R. D'tw(,, Ofli~'e of Naval R(.s+,areh, Chi(,:tgo, illin<)is, U. S. A.



S e a s o n a l V a r i a t i o n in the l l i b e r n a t i n g B e h a v i o r of Citellus I o t e r a l i s . ~ JOHN ~V. TWENT~ AN;) ,I..xX~:T A. "I'wv,XTt.: ( H i b e r n a t i o n I'hy~ioh)gy ]',:tl.)oratory, University of Utah, S'tlt l,ake City, Utah). Hil)ernating pat, terns for golden-mantled grotmd squirrels (Cilcllu~ lab:rails) ,Áre described qtmntitatively from ('.ontinuous temperature reror(lings from !15 animals with p e r m a n e n t l y intphmted thermoeouph~s. ]'eriods of hil)ernution in the aul,umn gradually in(,.rease in dur,dion until the rift ll or sixth perio(l. Core temper:d,ures (luring She first, perio(l average 4 ° t,o 5°C above the m i c r o e n v i r o n m e n t and decre'tse tmtil the temperaÁurge of the mieroenvir0nmen(, is approximated during the fifth or sixth period. T h e time of activity between periods progressiveh' decreases. ad'tptation apis, Kt'aduai and continues after ]hi>era.titan" " has begun. The winter pattern is established t33" the sixth

' S u p p o r t e d in part by the University of Utah Research C o m m i t t e e and by U. S. Public Health Service Gran~, AM-5942.

period, llil)erna|ing p~riotls of mldist.urln:d iu,lividunls art, rt,l'ltiv(,ly ¢:~)Itslltlll ft))" oat.ll total,Á.'mlure level r(.'ronh,d. 1,1) th(.: sitting, lh(, l)ntl+.rll :q)l)(,:t)'s to I)t; lh(' revprst: of (.1,~ :inllun)). "l'l.: l,st ft)tlr periods l)rogressiv(,ly decren.-:t: in durali(,n: (:ore tentl)erature..s tt'nd to l)c,conm cloy, it:Ál; llw lime of activity incrensf,s; a),l hil)ernaiion (',eases :,ln'ul)tly. For atoll's, ll)(,s(, c'))nnlzos s(,+,nl tu l)p :)ssod.ted witll (lesc('n(lin~ lesl(,s. TI,: a(lal)l:tti())l f()l' hil:)ertmlion nltty 1>:~gr:)thlttlly It)st as ~| l'f'sUlt ¢)f i1,(')'en:(.
Temperature Regulation in tile I.itth: Brown Bat, Myotis lucifugus. I,hmt-'wr C. 8'ro.xl-:s .xNo .l.acou E. Wxr:m~rs ( D e l m r ( m e n t of 13iological Sciences, Miehignn Technological University, l h m g l , t o n , Midfig~m, nnd IAfe Sciences Department, Purdue University, Lafayette, Indiana). Enrlhu. laboratory st.udies suggest that lnoernathlg Slmeies of Imts c'm m:tinlnin ho,neothermism only when moving, flying, or feeding. Bats used in these earlier studies were fnste(I, frequently had low body weight, or we,'e refrigerate


!ortmr was re~dily imluced in winltu" Iw foo, l ,lel~)'ivali(m. Winler resistance to h y l m t h e r m i a was }):~si,,qlly a nh,tal)olic a(ljustment rather than an insuhtlivc' n¢ijuslment. Seasonal differences in tlm (ollowing homeoll,ertnic characteristics were d e tinetl: 1) ztme of l lmrmal neutrality, 2) level of -;~tlwul:ant,ous I,o~ly teml)erature, 3) peak m e t a b o lisnh ,1) ".critical" ambient tellq)orattlr(:, ,.'5) l.lpI}(~'r :tn~l lower crilical ambien{, temperatures, and {i) ~'ritical imdy temperalure. Teml~evature-nml/~holjsm ~;elationshil~.,i were: estaMisl,!~d to illustrate these differences. /:Iil.wnalin~ spet:ies o17 /ml,s' shouht not be consi,lere~l m a m m a l s wilh n, highly develol)ed thermor,,gll]alory mechanism. This investigation I)res{.!lliS m-i~lence, however, lhnt /lIl[{el" given ('il'l:llII'lS|~.lll~teS a i: least one hil)ern'tl,ing species, 3 1 y o l i s]ug.,a, van mainlain a lm~neothennic level of body temlWmlure ;tml met,'.~t)olisn:~ sin~ilar to nonhibernaling Sl)eeies of Imts an(I olher nlal~lll"tls, eVCll when i~metive. Normal Seasonal and Experintculally Induced C h a n g e s in K i d n e y s of S u m m e r A e l i v e trod W i n l e r l l i l ) e r n a t i n g l | a l s ( M y o t i s luci]ttgtts) : C'ytoehemical a n d E h : e t r o n Microscopic Observaliohs.'-' ROIH:RT h.'l. " I{OSENBAUM, I.tAROId.~ SOlU.;l,, :xNI) AIINOI,I~ MI.:I,M::.N (.[)eparlment of ih~lhology', Allu.wt ii:insloin C,ollege of M e d M n e , New York, New Y o r k ) . t?vt(wlmmical visua.lization mc, tho~ls with the light microscope based on mM ext:ending earlier sl=u~lies ( M e l m a n , A., and I:h.)sm~baum, 1t. M. Anal;. lice., 1..'~5" 401, 1963; R o s e n b a u m , It. M., qn~l Melman, A. J. C,ell Biol., 21" 3 2 5 ( 1 9 6 4 ) showed a decrease in ribonucleie ac " .i d basophilia in proximal ccmvoluted tubules in hibernating anilnals as compared wi/,h active s u m m e r l)ats. ]further, a p p a r e n t expansmns of "u'eas o17 eytol.)h~sln form va¢.'uoh~s or vesicles having inc'reased aetivikv for several differenl~ nueleosMe pho.-,l-)hat,t'' se'~ .... These structures are minimal o r al~sent in proximal tubules from st~m~ner animals t)tlt, show enormotls increase i n both n u m b e r of vacttoles and enzymic ,mtivity in winter hibernators. These al)rupt changes arc evident ma.inly Jn the Golgi zone. " W i n t e r - l i k e " changes ean be produced in act, ive s m n m e r bat, iubuhw epilhclia, when animals are subject to 'lesiecation , placed in '.'toM ~,.tot •age, " or betas. l'hese " w m " t e r - h "k e " changes can also be altered hy anlidiuretic factors frolil p i l u i t a w . While light microscope prepar'~tions show ac~ivit,v for adenosine triphosphatase and other im,'leoside plmsi)h~{tases in what. a p p e a r to be Oolgi =Supported by a contract with the Office of Naval Research and grants from the United State,. Public H e a l t h Service ( G M 03605).

05483 and R 0 1 - A M -


l,',m',,=li.te.- and -vesicles, v.4c, o' nIt,• microscopy, sh nvs no such no| iv/ty in (] olgi st ruct ures pc:~3so. I nsl o;,i, this area. is eonltmsed Of large numlmr,.s of-i!~tcrdigitaling vaelmles forlucd fronl th0 microlulmles O *' " I)..m.a, lh lhe brush t mrder. "!• his v a o u o l a l " s vstOlll slmws m'livil:y for tile nueh:,osi~te phosplmtases visualize~l by light m]"~.!. . os~ . . . Olne ' evloehenlieal . . me.tide(is. AcJ.l phosphai,,'lse is not preseut in th( ..... slvuclures, and hence they ave not the. kv.ldeal ,,,> calle~l "re,~o,'l~tion droldel-d' or Idmgosomes norreally f0rmea in vm't~lm~tc.- proximal kidney luImles. TImt a speeialize~! resorpii0n l}lay ocellr -in t1,,.~, vesMe:..s.', lmwever, can be d e p m n s l r a t e d by injection of. emlogenous lUWoxhlase or hmnoglobin LI



llS tlllll'k {H'8.

This subcellular p i c t u r e is eonq,le/ely allered 12 hr nfl(,r hil)ernaling trois have I)(-,eolno t'll'OllSf!ll, t:c) t)roduce a picture similar to tlmt, of a<:tive s u m m e r bats. D,'il]l |.hose experin~cntgll methods hMu(,iug "winter-like"-chang:es in slimmer bat ki~lneys, the subeeihflar morphology is htentical in kim], lm(. not ahvays in (,,)",,with (hat seen in the hil)t,rnaling winler ,mimal. C e s i u m - t 3 7 D i s l r i l ) u t i o n in T i s s u e s o f A c t i v e a n d H i h c r n a l i n g C i t e i h t s la'teralis, a Ma,~wix L. I~IEDI.*:SEIq V l '

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(Departmenl, of BhHogy, University of New M ex i e o, A i J~u elue rq 1ue, N e w M ex ie o ). -#.~ [,xpcvtments were conducted to define the. extent to which lowered body. l.cmtcl,ltutc' ~'" ':' all,rages' " " biological half-time an~.! dislrilmlion of cesium-137 in tissues. Daily whole belly counts were m a d e on animals, given a single intral)eritoneal, injpetmn" of ihe rqdioisotope, 0.48 ,ue in 0.3 ml of buffered Ringer's sohition per 100 g of. l m d y weight. A n i mals were sacnhecd after first., seeoiid, third, and fom'ih bi~dogical half-times, and ra~lioaetivity of tiss1~es was measured. Tissue.s examined inch.Me blood, henri,, liver, spleen, stomach, small intestine, kMney, liver, ...l,clet,d muscle, skin, an I bone. T h e data, cxt ;c..sed as tasstle i n d e x , " lhe ratio of activity per gram of tissue i,o ,~.c.t,~ , " "-"it-3- per gram of body weight, allow coral)arisen of tim distribution of ees/um in a n i m a l s with varii~ble arllounl,s 0f whole bod~,- a c t i v i t y , i.e., first, ~econd, ete,, biologie,tl h'df-l;imes. Test eondi(:,ions included deprivation n,{ drinking w a t e r and cell'~ exposure, with and without hitmrnation. Animals sacrificed during h i b e r t m t i o n had been in hibernation a m i n i m u m of 2 days, Dep,'ival, ion of drinking w a t e r and hibernation prolonged the retention of cesmm-13~ " " 5- and 10foM, respectively. N e i t h e r test Condition effeetod a redistribution of the i s o t o p e a m o n g b o d y i.i~ams

with the ex,.,eption of an increase in the eesiurn " Supported by Atomic E n e r g y Commission C o ~ t rac t, A T ( 2 9 - 2 ) ' 162q.

90 (,()n(.tqil.

'I'ItII1D .1NTI'~ltNATIONAI, SY~II'OS11rM ON I I I B I , ; I 1 N A T I O N ()t" (.ar(li't(. lissues during hil.)crn,ltion. ".['lie

(lliil), lliity })e exl)laine(t by the following liypothosty,s: 1) lh(: lneehallism reslTonsible for reiention of vesiuln-137 is ieinperllture-ill(tependent,; 2 ) ' e i r m i htlion in lisstles (hiring hiberllatioli is inadequate fO roiliov(' cesitiin lost fl'olli yells. I n f l u e n c e of l,ight Ul)On C i r c a d i a n R h y t h m s in I l i i ) e r n a t i o n . l"fvn,~ray l)oHt, (5'lax-Planel~ Insiitut fiir Verhaltensphysiologit; Seewiesen und Erling-Andechs, 813l Erling-Andeehs, G e r m a n y ) . It has I)cen rel')orled (.hat, a ('ireadian r h y t h m of Sl)onlan(:ous locomolor aelivily l)ersisled atruler c()tlsianl (,on(liiions of ligh,t .rod lemt)end,ure during hi)),~rnntion in the c o m m o n dortnouse ((~'h's oils 1,3 (Pohl, H. Natm-wiss., 1905, in press). The length of the free r u n n i n g period rein'fined almost, (,oilsllll}l (ll])Olit~ 26.5 hr) ov(:r tt 5-month t)erio(l of hil)er)),)lion ~nlerrtlpled I)3" Sl)onlnnemls :irousltl~.

The l)r(-,sent paper l)rovi~ies (,vidence l.h,d light~lark cyeh:s of 12:12 nnd 6:18 lit' (Zeitgvbcr) were al)lo Io entrain tlm 1'ree rmmimz period of l~co-( ' m()lor ,lelivily of dormi('e even ilurinl.;'lfil)ern'alion a( 8"C amt)ien~ (emlmrature. Timing of Sl)Onlanoous arousals w~us also dr, l.o.rmin(~d I)y light(l'trk eyries. Changing ligh-t, intensity Slel)wise from 0.004 ((t 1 ]ux varied the l)eriod of aetivi!6" r h y t h m from a minimun: r of 23A 1o ~t m a x i m u m r of 28 hr in the :~ciire (Iornmuse at. 18"(3 and etmstant, illutnination, following Asehoff's rule for (htt'k-n(2tive animuls. It was imliealed lh,'tt, this ('or(elation might: be llresenl: in hil)e.rn,';ling (lot= mice as well. In anoll{er seri(:s of expt:rinmnls0 a .rhythnt of (feel) l)o(ly Ienal)ern.iure (re('or(letl from itnplanted therhlo(:oul/le,~), s.~:nchronized with a 12:12-1n" light-dark eyck,, wqs found in a lighi,-aetive ani'real, the 13-lined ground s.quirrei (Cilellus lridec(:mli)leat.s). The .l~""cr, i g•' .O-- range of oseillidion WaS Ill)otil: 3°C Ill. fi~C ;iinl)ient tornperlil,ure. l i , is t.onchided that. 11 hil)erii/ll.or exposed to it light'dark. Zeitg"eber, eilher (hiring hilmrmltion or during the active s l a t e i)ct.weeif .periods of d o f mlmcy, will be synchronized wilh its enviromneut, thus providing exacl, iiming of its internal rhythms, which l/lay be diurnal or annilal. The significance of eir(:a(tian rhyt}lnis in (,lie hibernaling animal will be (liscussed,

1Meltabolism and" Body Temperature in Active at.! Torpid Eastern Chipmunks (Tamias striatus lysteri). Rzcl{aRo L. N~:t:.~tax~" (DeparttnenZ'of Zoology, Syracuse' University, Syracuse, New York). Thirty eastern chipmunks, 7'amias stria(us lltsteri, were caught-on the Syracuse Universii,y emnpus

aml studie(l fi'om the sumnu.'r of 1063 lhrouglL lilt, winlc, r of 10(i5. Specimens ol./quint-,d during the s u n ) n w r had a zone of lherxnoneulrnliiy from :qq)roximalcly 30: to 3(P'C. Metalmlism wilhin tim zone of (I,~llnone//Iralily was 1.24 ve (): per g per hr :m,l invrmlso{I nt a rate of "d~out. 0.11 (,~' 0-: per g hr p¢,r "(?. I)¢,l(m" 30°C. This mmabolic ira:(ease is similar i~) 'that uf :i, (rol)ieal animal wilh only mode)'alely (hwelol)(,(t insulation. Body telnpe.rnlllres v(ero t)olw(,en 3S' aml 41.5°C at am!ileal tmnlnq"dllres rnnging fr~nn

--6 ° to 37~'C. T o r p i d i t y was ol)s(,,rvp(l (In'ovlgho'u( lh(, winl(,r :iS

qml,imil lomperaiures fn;nt 0 '~ Io 22°C in (.J~il)mimks nmin{aim~(I in the lal,ora(ory aml in (ultsi~h. eag,,s. Willie iorl)i(I :tt (I~ It) 10°C. only :l st,lali number, of individuals ;lllow(,~t their tmdy (('llll)Ol':lllll'eS 1o al)Proximale lho umlfitrnt Imnl..'t'nlure: inslo'ul, m a n y of the chipmunl~s exl)erimv'e,I somilorl)i~lily; i.e., Ilmir h(.nly lemper.Huro o n l y t h , d i n e d I. 15 ~ 1o 2(1':C,. ]towover, indivi, luals iorl,i,I Ill. I(T~ to 22~( ~ h:ul body IOlli])/,rllllli'es ,losoly al)l)roxihl:llillK lh(, 'llul.)ioDI i e l l l l m l ' a l m ' p . ('hil,muni
inla ehilm~unks :it llmslt same nmlfient lt,!)ll)orqltlr¢:s. The mel:d)olism of imlividuals tm'l)i~l nt hint)lent leml)eral).lros from If}~ lo -,~9""¢'.. r:mge{l from 0.0,1 It) 0A ce 0.2 per g l)m" hr. A gn-ulimil ~)f 2<' to 7°C exisled .Imt ween body l(.qFll)('l':tllll'e llltll ambient, leml.)et'aiurc in those chipt,~tniks lorl)i~l ,'it 0 ° to 6°C. l..lnfortunntely it .wits not possil)10 1o •lsoortaill '1113' ()xygell consumlffion vnhn,s for l]l(,.s(, individuals. 11. w o u h l nlqmar l hnl= the eastern chipmunk 'illows i(self Io.he(,onm deeply lorpid :tl. less sever,, lemlmral,ares, bttt lends to r,:sisl, eoJnplete {o1"1)(.)1" nt those lower tempt.'ralures at whivh other grounii squirrels (Marmotini). experienc(, dee.l) lt)rl)or. L y m u n and Brinks (.I. Cell. C,oml). l)hysiol., ;kl: 53, 1959) h a v e shown that the isohlte(l hem'( of 7'. striatm~stops i)(,'tiiIlg at, hi)out 7<' i o 8°C', while those of oilmr grotintl'squirrels coal into, 1o belt( •).t from ~1 ° t.o. 2°C. ])epfll of torl)i(lity in t h e oasiel.ll c}lipilitlllk in,'ly llills lm gmT(:!!'ned I)3" tile inat)ility of ils hearl to fum~lion nl tow lemlmrntur~.s. A profound winle)" Iorpor and its consf;rv,ilion of energy m a y (hus l)e eompensaied 'for l).v short, or shallow bouts of torpidity supplernenl(~ll will) ll)e hoar(li!Ig of food in the eastern ehipn)mlk. Tii(,, winler torpor eharnelerisli~.,, of lhese ehipm u n k s tufty Ills() be partially exl)l'dn,(t b y tim animal's rehltiffnship 1.o the nonhibernating t.)'ve squirrels and lhe hibernating ground squirrels.

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1 N'I'I':II N A ' I ' I O N . ~ I ,


l~llick (l:hlll. ~XItis. ('olnl.). Zoo],, 1,70: 113, 1963) ft:.i~ls ililii ihc ,'hil)niililk is ,+in inll)rllit~llilil(' I}("lwl;~'ii Jim ll'~l~ an~t 7rouiid sql.iirrois, i.e., ihlil l}ie gl'~llilill StlUii'l'l'l. ~ ~volw,~! frolii ;i "t~hilnlilnik-likt; fi)l'lii. '~ Ttiis l l r i l i l i t i v o chil)n/liilk liiily liavo llOss~,ss(~l ltit:~ iiteit)¢,nl stag(,s of mlAi)iiv~. , t:orpidiiy wllil'h worr > ltll(~r l~(:l'ft~'i~:~l I)y l ltl:' liilll:,ri/aling and v.-qiw!iilig iiiOliilJl,rs (if Jim ii'il/~, i%larnlothii. T h e ~.~llll¢,liil;~Ol'iir ), c h i l l l i i l i l i k lii~iv lll)ssoss ~)liliorils of i(n'l,i(liiy ~iliiilllr IO lira lll(n'l, il'iiil)l(! lilil,t,st.oi's of flu:, liilmrnal ing groulid S{lllirrol:

Non-Shivl'rilig Ill,ill ]'rlldiiclhln during Arousal frolli llillt~rnlllhln lind Evhh:ili:l, f o r lhl; COilIrihulhtn of llrown Foi. J. S. ]];tYWAliD AND (". t I. I,'rMAN (1)l,ltlirliil?~llt. of AlllitOlity, i t a i ' r l i r d ;Mt'~li('lil Svh()0l; ]h)sl()il, ~:lassili'hilsl' Ri'('('nt t,Vi(ll,llvl' iil(ii{'lil{'s thlit tn'(iwn lillil)osi) tissll~' i s a l l l a j o r .~t)lll'l'l' tJf lil~, li(,at lliq~('~ill'y i'()l" ai'()lisa] frolii hi!iornalioli. Slu(lios ill oilr liillin':lt~)l'v fllrthor inttivalv thM Iji'owli flil is nltire iili1}oi.i.~lnt. hi l(,rln~ ()f i'ol:il/v{, ~ heat l)rtlihl{"lioli in lials lliail in r~l(h,iil hilloriialoi's. Siill,(: locinii~lliOs for l~artitioning tile i~ i.,/t,o ll~,<~ll I~rolh.lvlioll ~)f V~,l'lii/n lissm,s suvh lis In'owli flit, (hn'ing ill'l)llSii} cff lli(' Iml. ar~,'lllcking, :i Stel~ ill l.ll~ d i r m ' i i o n (,f ~ll~'ii l m r l l i i i n i ix Io del(~>riilillt , i.ho liOiiSliiveriilg li~iil i~l'Odllcl i()ii. We h:ivl~ t'Oilill;li'~,d llic • t.olnll¢~i'~ilill'l: (hofll'l and I)i-owli f a l ) ~il'OllSiil i)iitlorns of ('llrai'iztq! lllill liorlllilJ I)iltS (J'2tl~({.SiclIS SUsCi/.~), JlillllSll'l'S t,I)..~<,cr/ct~ItIU /lt/r+lltl,,~), itllll lifirliiii,i., (Oli.'~'91i~). (;~llrlll'iZl.qi, arlilh;ially !'CSllirilt(:~l ll/ils Ciili ].)O al'Ollv('(I I,O poitk I,o~ly i(qlil.ii~l~alili'Os Ill. li i'litt~ lluli is insignilielinlly slitTorPllt+..frolil coll|,rol ili(lividual:, lt~)(l(~lilS, on liie ~ l i 0 r liali~l, lil'O tlnalile t()lil.Oll,We ill, ~1 llOrlll~ll rlllo ~;iit,ii shivt~rillg is l)lockod. Tiit,,~t" thltlings lii'(~ supi Jelled I~V Ctillll)itrillivo (.~l(:ei i'OlllVtJtLl'ltlll rlwor/Is dtll'illg li()iiein'ar/zeit aroiis:iis w h i c h doin(silslr:lto. nl/ich I(:)Ss s h i r o r i l l g in lJle l)iit. Olir tolill)(,rlll.lll-e rt~('.ords on l lm 1.mis shclw l.hti,t t)rowll flit is (,x)]y w a r i l i ~ r l.]itlii tlie heart l h r o i l g h o l l l lll'OUSa] in f;il.iic, r :l eohl (iF i.o 8"(D) or ;t Wfli'lll ( 2 2 ~ io 23'-'C) Olll'ii'Olllll('.-[ll. ?i]:lxinillin t(:,lilt~oraillr~,s ill l.liO h r o w n fal roli('h very high \'Mlit~s ( l l l ) l ) r o × i n u i l e l y 42~C). Thl; very" ral)ilt t']tililll;s o/)scrve~l in I)rowli flit. rel,qiive i(l COFe. toinl:~ei' iridical.e l|litl i:hc i)rowii hit h(>.l/l, ln"o,tuelion is un(lor l)i'l>(.;iso t:)ltysioh:lgical coliiro]. The liow tlTchnhlu ~ of therlliogrlll)]iy has l:)el;n utilized 1o obtahi till ii}teresting iJictorial eXaml)h_' ,,Jr lhe tmtil lJro(hietion by hrown fal in tin fir()llsifig [)at. Tile l.hc;'mogral)h s('ilns llia infi'arod radi
ON I t i I ~ E I { N A T I O N

c¢)lis l~i('iiously ~h.,lilll'::ilt'~! " h i l l "


llr,;li I hut ('olin'ida's

t,Xli~!l])' willi the ~hl.llit" of i til~ i.nlderiying I)l'l}wii flil.




I:IL-~.NK (7. |~.rt. IJ'l'.:X AND

lhe Iiibcrnaling Bat, ltaltOH) A. I(.L','TU(,U (l)e-

])ill'lllHqllS of A l i n | i ) l l l y , Sclmols of ~|(~dit'ille. Slalt~ l Tnivei:.~ilv of New Y o r k at |hlfl'al(}, Buffalo, Nmv h ork, Ztlll[ t]IC ~.~IilV('FSII'.V ()f |{oi'hi,sier. 1:{och(,s{Ol', .N~'('W Y o r k ) . l/il,,,rn,'~lil}g is)sits ]lavt~lmen ,Ioscril,l,~l in. l lm liter:llure as anuri,.. This view is llol SUlqmrl,.~,l I)y oui" stlldi,,s of urin{~ flmv l.{ii,,s in cailiei(,rized litih: br()wn I)nts (_ll!/oli.~ lu:ifufl.s), ltatll{-r, a stnali init ('(~llsl~llii: urine th)w lias invarial)ly tn,oi} folnul in hi)tit slililnl~!r and whiier I)als al 8.5°,t() 10"C, l)ody telnl~l~r<'lllll'CS eoliln|t)llly r,,(.t~gnizcd tls withiii tim utqmr rang~, ~)f din'l, hiln,rn,'ilicm. Flmv suflMcill to I~t, of signitic~lncc in l)l,M~ler ,'lislensi(m iuis t)(~cn d~m}¢mstrtii~,d at. l)~)dy li,nll)or:itur~,s dowii 1o 5~'(," hi whiioi'. Alilli'ili is sug!~cst.t~'d l..,h:lw these liniils, niltil.lllgh flow l'ales ill ihc' Ol'dl,r tlf 0.0005 l~1 I)er liiin l~l'l" i~ of hody w~dghi lnlty |ilivP (.,sl,iil)~;d dl;,tc'cihm with lilt' })rl:)i'i, fhtrP,s llS~:,~l. l~i'ili(, fh)w in ~lllieliy rtmslilig It;if>-.:, Siltije?lod neiliil:r 1o i,xt,ossiw;, wtilo.r loliding llor lo d o h y d r i ! litiii, ;il)l)rtixiriial{~s iI,sil)iilly oxi)Olil~lililil ~ie{'l'eflst: (Jr w(,ll over l(10-fohl froin fiighi, tc, nil)P.ratllres (ill I to 4!.5°(.?) l.o Ill(:, l'llllg(.', whc, rl; tli-in{.) l l l l t l t l l t is 11o ]oilger dtd.(:(.ltlllle. Siiliilil(~r lilld whlll~r flow rtl!l;,s tltll}~iir conllilirliliit'. This is hilcrltroli;d liS Jill lldfil)llilioll 1.o ali exlt~ll(h'(t ypiir-i'olnid lilOtiit)olic l'lillgO, which lliliy li("Sl i;,xlillihl 1.1i(, llior£, rillJil| (iem't.,liSe in i)ill, llrhlt, lt(~w ii~ I'oiliiJfil'lr,d t.() l.he ri/[ or gr(iillid ~lltliri.o] Ill r~dtico(l, b e l l y l.Olnl:iorlltliros, Oiir findings hl(lil'ale thal..t.ihi(Mc~r (lisiensioll c.lllllliJt, ),l~l. lJ{~ ruh_,(t olil. llS It c o i i i r i l } l l l o i . y sl, hilillils lo liroii~lil, ali([ silggosl. {hat o l l l i i n i i l iirillo, itow l'iti.l+ lnttv ].)1.~ li fiicli)r ill stdectioil (if;.liil)erii:.ll.iiig ll.,nll)Oral.ilr{'s i:iv t h i s Sliel:;i0~.

'l'elclncicrcd PlJysiologieal Measurements o f Arctic Bears during Natural Cohl Exposure. (]. ]i:m;m~ FOLK, Jm, l tlctt,~ru) C. SI,~IMO~DS, A.','i) lie(am S. H~:l)(m (Physioh}to-.Bran(4t, Arctic Aeronledit'tll ],~;tborlitor3", .Fort tiVainwrig}ll;, A hiska, an,i A retie .Rcseareh l,tll)ora[ ory, Barrow, Maska), M e t h o d s dcscribe(l earlier for l~hysiological to,leniel i t from iml:danicd (;apsules in larg~ arctic carnivores were inodified for use on bears, 'I:wo black b¢.~ars and two grizzly Imars were tstudied in outdoor enclosures. As cold w e a t h e r developed (as low as ,:-60~F), they gradually spent more time in dens. Recordings Iti 15- or 30-rain intervals were t~nmsmitted t h r o u g h h e a v y metal cages over singlc-idl)O tuned anu~nna~ or an a n t e n n a around Ibe c:~ge, l,o~s of d , i l a due to transmitt.m" failura



I N"T I .:R N .A. .I. . I O N A L S Y M P O S I U M ON 11II~ERN-VI"ION

w,s re{hwed by iml}lanting two or three capsules in ea{,h I)ear. All three transmitters fl'om :t single :mimnl were recorded on the stone antenna, l'l~ysiologi,'al observations were: 1) the l,eak of a(:tivity began l'tter (as the fall season (levelol)ed) at. "t rat,: of al.)ot~t 1 hr per v,'e(.'k (7 a.,'u, on October 1 and 10 a..~. on Noveml)er 1); 2) at first (in the early fall season) h e a r t rates r,mge(l from 40 lo 120 beats per r,~in ; 3) la(er in the fall season the range wns ,10 Io 100 I)eais per rain; 4) when.the l)eriod of winter rPst in dens began, (he range for all heats ('hang('(l (() .'~ m:w k)w level hts(ing 75% of each (lay, ranging from 8 to 36 i)eals per rain with occasional l~(,vio(ls of 70 beats per rain; 5) m.tivi(y l)(mks {'on[illll{?d t o ('Gllle n e I i r li~)ontillle. To 811111m:u'ize,. |he most. signilimmt ()l)s(:rvntion was |,lint l,e:u't r,l.(.,s of sleel)ing I)ears of medium size are al)out 40 })e:~Is l)[,r rain in Sllllllllur l)ut, ns (he state of winler res! i:rogre.
BRO{:K (Gerontology Br'meh, Baltimore City I:Iosldtais, Baltintore, M a r y l a n d ) . Various as.-.'t~ys of whole blood t)nd triehloroaeeti(" neid exlra(.ts (}f erythroeyt{:s and 1}lasma indic,tted that, aftt:r a few weeks' cohl exl)osure, gohien h,'tm8ters ( au)'atus) which eventually hil}ernate(l had increased hematocrits nnd markedly decreased e~Tthrocyte concentra(.ions of "aplmr{hit glucose" (glucose oxidase assay), t}oth characteristic of hil}ernators. Typical increases in inorganic phosphate (1P) oeeurre(l in erythrocytes mad plasma of hibernators. Blood from hamsters in all experimental groul)S was rapidly exi.raete(l, and the phOsl)horyhRed constituents were separated and (luantitated (~sing tt micro-ion exchange column ehronmtographic procedure followed by diagnosLie tests f o r various sugars, phosl)hates, and nucleotides. Most glycolytie intermediates were found in about, the sume steady state concentrations in control, cold-expose{i, and 3- and 4-d'ty hibernating hamsters, except for II }, which was signitieantly higher in hibernatot..s. Adenosine t ril)h~)sph'~te ( A T P ) , diph.o.-q}hoglycerate ( D P G ) , and ribose phosl}lmte eoncenlr:ttions were somewhat increased in ('old-exl)osed hamsters, whih: the same compounds were found in decreased nmounls in hibernators as compared to (:ontrols. In bloo(l from normotlmrmie hamsters centrifuged at 4°C for about 1 hr prior to e x t r a c t i o n , a signifieamt decline in A T P nnd rise in fructose diphosphate ( F D P ) was observed. The I ) P G concentration increased slightly, and an unidentified inosine-like material aplmare(l. In eontr.ast, none of these changes, exANN

eept for a slight in{.rease in FI)I', (}:.Ptlrre{l (hn'ing eentrifilgation of l}lood from hil:}{,rnators. Ery{ hro(:)'tes from hil}ernntors m:~int:lined higl~ A T I ' levels evt,n when the idootl w:,s warm(:{l 1}rietty before {.entl'ifu,,zalion nml extr:wti(m. ]~lo~}d fi'{ml normolhermi(" hamsters l}restwv{:(l at -t:~C in m'ideitrnto-{lt,xlrose for ,I {lnys is c{}ml)h'l(~ly unlike hh}od from ,l-{lay l~il}{,rtmt{}rs. : \ n as 3"o1 uni,lontifled inosim,-like mat{,rial, I}olyl,hosl}h(}ryl:t{('{l in re'y| hroeytes aml nl}lUtronlly n(mldmSl}lmryl:~te{I in t}insma, wns f{}umt in all ~.Xl}t'rinwnt:d grt}tq,s. "l't.:' r{,sults :u'o in|,,rl,ro{{'(I :t~ :t {'(}t,~l)hmih)n l}olh ~)1" 11,' effe('ls of the lmvere{I ('elluhu" l('nll]oratul'o u n { ' { l U i lil}rium {'ont'enlrations of {he intortu(,{linles :u~{I of '~ .q~ift in lira p:dh ~}f g[~w(}st, uliliz:~|ion al lira ons(q of hil)(wm~tion. Intestinal Absorption in l l a m s t e r and ( ; r o u n d S q u i r r e l : Exl}erimental Rclationshil~s b e t w e e n in V i t r o and in V i r o Ai}i}roaeh(.s. X. ,J. M USAC{'I{ta ( l)(:part merit of l~ti()logy, Saint Louis University, St. Louis, Missouri ). Und{'rstan{ling of fun(qicmrd fi'nlllros ()f ga.,:lrohltt,stinnl lrn(.t in hil)ernnl()rs is woi,fully innde{lll~II.e. F e a t tll'OS {}f ill{{'.st inuI :tl}soI'I}l iOll 'lll(l {'elhllar :wtive t rHliSl}or| ill (Trict'l~t.~ attral~l,s ,'111{1 Cil('ll,,m t r i d e c e m l i , c a t u,~ were stuclied. 3I'unster intestine in ripe showed n {'onsislenlly gre~flor nbilily to tdJst)rl) D-gltwose, 100 mg % in I(rel}s-I~inger I}i('.~rbon,ale 'tt 37°{'. A eompnrison, in terms of u.~t g]tlcose 'd)s{)rl}e{l per g {lry we|gill l}er 30 rain in various intestinnl :|teas, is: llamster

l)uo{lenu na Jejunum Jejunnn'l lleunl

90.,I l()l .2 {18.0 i 0 1 . t|

Ground Squirrel

64.5 87.8 65.2 -I 1.2

Corn pq risons between nonfasting, 12-hr, 2-l-hr, und 48-hr fasting lmm.stevs were made. After 12 !," of fasting there is a0% reduction in levels of sugqr "d)sort)tion, ,'lft(,,r 2-t hr the levels of ai>:orption are approxim'ltely.' equtfl with nonfnsling subject,-:, nml "lf(er 48 hr levels of absorption, 'llthough somewh,tL erratic, are increased 'd:,out 30,¢~ in jejunum and ileum. It,. vit, ro sugar absorption is inhibite{l by phlorizin; previous assessment of phlorizin action with in viuo prepa, rations in hamster or ground squirrel h'ls not been reported. In hamsters in vD.o, 5 × 10-" .~t phlorizin was strongly inhibitoI3"; 5 x 10-' .~I was less inhibitoIT, permitting low levels of glucose absorption; and 5 X 10 "'~ ,xt was noninhil)ilory and values were comparable to those of controls. In ground squirrels the intestine was divided lute two a r e a s : upper ((luodenuna :rod jejunum) and lower (jejunum ,~l{t ileum). Upper gut was •~




T H I R D 1 N T F I i N A T I O N A I , SY.MPOSII.TM ON H I B I ( I ~ N A T I O N filled wi(ll 100 mg'5~ ;)-gllw()se in Krel)s-Ringer biearl)onnle, nn(] l()wor gut with simil:u" solution ))lus 5 × 10 "4 .~1 phh:)rizin. In ea('h animnl glucose ~v'ts al)sorl:)ed only in till> nOn-l)hh)vizin-lrea1~!'d t:,ovlion of inl¢:stin('. Phlorizin inhibition of intesli)ml "if'tire tl'ttllS])Orl wt|;~ lhllS (temonsl;':tled. Al)so,'ption 17)]l(,l~oilmnn r(,lal.e(l IO hihern:dion and awakening were slu(lie(l. ],ow~'sl levels of ab:÷Orl)lion we,re ol:)lnin(,(l in animals un(ler lfil)ern'd ~Orl)()r; 3 l o 4 l)r afl('r awnk(ming fro)n hil)ernn= tion, s).)gn)" ;I] )s(:)rl)lion Wals <'rrnli(', i.e.., high in sonu, segmenls :m(l h)w in olh(,rs; :~ft(,r 20 hr g|l:ls.orpiion wns eleva(~,<] ill llltllly S('tZItlCIIIS. Call P r o l i f e r a t i o n K i n e t i c s in l:libcrnating Dorm i c e (Glis glis). S. .l. ADv,[,s'n~Ix, C. 1'. |,v.~tA.x, A.','I) I{).:GIXA C. ()'Bl~llC.x (1)el);trtment of Anatomy, tI:H'v:)rd M,',li('al S,'hool, Boston, Mass:> ('hnscq Is). Tlwre is eonsid(,rql)le evidence thal, eellul'u" prolifiw~,lion is redu('o,l (luvin~ nnl:ur:d mammalian hil:)ernnl,ion in both norm'tl nn(l neoplnstie tissues. ITtilizing the deoxyrihonueleic acid ( D N A ) precursor trili,'lte(t (hymi(line (I!P-TdR), the kinetics of this inhil:)ition hnve 1)een st,udied in (..1lis oli.s. ttibernnting (lormi('e were injected with H3-TdR


(4 #(' l)er g) lhrough intr,,~l:)eritoneal or intraarterial polye(h.vlene ennnulas. C'.ontinuous monitoring of body l eml~e,'ature enst, red that, only undisturl)ed hil)ernntors were employed in the experimenls. Tissue w.~s divided for histologieal-radio,'mto~r,',ldJie lWelmrat,ion nnd ehemie:d analysis. 'I'h,~ mueosnl aml lymphoid cells of l;iimrntd>ing dormice :u'e disiine(ly labeled following the adminislraiion of Ha-TdR although much more f.tinily l]mn in nciive tmimnls, qThe chemically measured upt:d~e of precursor into D N A is redu,'ed by 95% with ;t Q~,, of 2.9. LTp to 72 hr following the introduction of the isotope,-no labeled mitoses nre ol)served in (on(inuously hihernating Gli.s, wldle in aelive ro~lenis these" npl)ear in less than 2 hr. ']'wo hours following the I'el:Ul'n of body |eIn ° ])el'nlllre fronl hibernat, ion to 37°C, all mitoses are lnbele(t. There is n marl¢ed dep,'ession of tlm mitotic index in the iniesiine during hilmrnal,ion, with a flourishing of milosis observed on arousal. These data. suggest that the normally proliferating cell populations nre blocked in premitoses (G..,) (luring hibernt.ion while D N A synthesis continues, although at a markedly reduced rate. On arousnl the Hock is relieved and (:ell division is resumed.