Riedesel, M. L., and J. M. Steffen. 1980. Protein metabolism and urea recycling in rodent hibernators. Federation Proc.

39: 2959-2963. Invited paper.

Protein metabolism and urea recycling 1n rodent hibernators'

M. L. RIEDESEL AND J. M. STEFFEN
DepaltmtnL oJ'Biolou, Unit,ersitj of'1,{tiu Mcxico, Albuqrtcrque, i{eiu Mexico B7 1}1

L,1,.,."nt in thc sizc of'roclcnts ancl olhcl srnall rnarnnr:rls is a lelativell, high rnetabolic rate ancl a linritccl capar:itl' firr eneruy stor:lgc. In adclition the lalgc surface al'ea to r,olunre ratio results in elevalecl r:rtcs ol'hcat loss firllorvinq exposur-c to l()rv enr.'ir<ltrnre ntal temperaturcs. The metalrolit: dernancls irnposecl bv srnall l;ocll size alc arnplified n'lien pcriocls oi' lccluccd fbocl and fi'ee rvater av:ril:rbilitl' coincicle lr'ith seasonallv krrvcrccl ambient temper:rtLrles. Hibcrrration rcpresents a biolosical aclaptafion that le:rclily meets thc energ)' cletrurncls of small animals in colcl
clinrutes.

ABSTRACT A rt:r'iel of the nretabolism of hibernators supports the argument that proteitr utilization is
:rrr cssenti:rl aspect ol hibelnation. Ser,cral lines of'evidetr<:e in support of this hvJrothesis are rlist:Lrsserl, inclucling (1) the ficld prefererrt:es of some hibernators lor a rliet rich in protcirr

prior lo hibernation, 12) thc glur:oncopenic rnainlenance ol gll'cogcn stores during hibenration, (i) thc nraintcnarrr:e ol pl:rsrna amino acicl ler,cls drrrinc hibelnation and their
ol arousal, (4) changes irr lcvcls of nonllrotein nitrogen lvith hibelnation, (5) the potential for urea rccl'cling bv intestinirl rnicloflora in active fastecl aninrals, lruci (6) thc lack of'corrclalion bctrveen Ioss ol lean bodv rnass ancl ure:r recovery clrrring tlre hiberrn:rting season. Thcsc d:rta scrvc to enrphasize the contribution of protein rnctalrolism to the o\er:rll encrgetic acliustlltcnts re<lrrired fbr sulvival at a lorver body tcnrpclrtule.-Riedesel, M. L.; Steffen,J. M. Prot.ein niet:rbolism and urea reclcling in roclcnt lriberrralors. Federatian Prot. i\l): 2959-2963: 1980.
elcr,ation drrring the l)rocess

Wanu (30) has recenf.lv evaluatecl tltc cnelgetics firr seasonal torpol of Spermophilus richtLrdsonil in the fielcl.
These clata agree clrrite favorably rvith Kayscr's (16) classic revierv of the clirt:r c:ollected on r::rptivc animals. lt is apparent fronr these stuclies th:rt I at'ge nr:rm tnalian h iller'n:rtors, suclt :rs the bear, irre in a vel'y cliff-erent energy balance situation than r.trdent

scrved not to eat for sever:rl u,eeks follorvins the ternrinal irrotrsal from hibcrnation in the sprinn. Despite the eyidence of arlple enel.gy rcservcs r,I r)r:rlrrrrraliarr lrihernltols, irrreslisal.ors tencl to emphasize stored lipicl energv as thc kel'tiutriti<lnal factot'rerer:oqnizccl as being essential f or structural, enzvmatic, irnd osmotic functions. Certzrinly these basic re quirements must bc nrct in the hiberclifferer-rt strategies
sponsible firr the sucr:ess o{ hi}ret'nation. I rr :rr r intn I n r rt rit ir,n. proleit t is

high in protein (22, 28, 29). 'fhis pref erence is cspecially apparent clurinq thc 1-:rll prior to hibernation. fungi high in protcin. 'l'he
recent

scribes a distin<:t preference in the solden-mantlecl ground squirrel and several chipmunk species for a diet

The clevated level of lipid de position in the fall coincides rvith a diet of
demonstration by Montoya et al. (24) that protein deprivzrtion in the garden dt,rm,rusc indur ed t()rpol irr Jn)

hiberrrators. Bears I'rave typically

bcerr firuncl to lose l5 to 257o of thcir bocly r.veiuht ovcr thc period of'con-

nating :lnimal. .|ust ils there

tinuous r,vinter hibernation rvhereas rcrdents nray lose 40 tc> 50% of the bocly wcieht. Ninety' percent of tire erlersv utilizecl during the hibernation seasrrrr <tf S. rir:ho,rdsonil is associatecl rvith the 30 C elevation of bodl' temperature experienced during perioclic alousals ancl the follorvine phase of normothermia (30).
acl:rptations to both recluced body t.emperature and prokrngecl Ihsting. Northern species may spencl up to 8 or 9 months in hibenration (23) rvith little or no food intake (13). Energy reserves are not consiclered to be a limiting factor in rodent hibernation. In fact rnany rodents have been ob0014-9446/80/0039-2959/$01.25.

in meeting the

are

basic neecl fbr protein among anirn:ils

irr general, it is to bc cxpected that there :rre also clill'crcrlt strateqies for
accluisition, utilization, and conserva-

tion of protein among hibelnators. The ,rhjct tire ,,f this plescrrtatiort is
to summarize evidence that iclcntifies adaptive acliustments in protein metabolism that are as essential to the hibernating nrdent as is the accumulation of'body fat prior to hibernation and oxidation of lipicl as an cnersy soulce d rrring lrihcln:rl ion.
DIET AND BODY COMPOSITION

Hibernators have devektpecl marked

season emphasizcs the f'act that pr<itein may bc a limitins factor in the metabolism ol hibernators. Both lipirl and protein lcserves may inf luence the onset, duration, and success of hibernation. Changes re ported in lean body mrtss irrdit ate thal prorcin nlay a( count for approximately l0%: of the cnetgy expenditure during the hibernatins season ( 12) Jame son and Mead ( 14) noted conservation of fat-free dry rveieht both in animals hoardins loocl for consumption during periodic arousals, as do chip1 Fronr tlre Symposium Protein and Fat Metabolism during Mammalian Hlpophag'ta and Llibernation presented at the 30th Annual Fall

Fielcl observation, inclucling the exarnination of' stomach c()ntents. dc-

Meeting

of The American

Physiological So-

r:iety, New Orleans, October 19, 1979.

O FASEB

2959

munks, and in those that do not ezrt, such as ground squirrels. Holr,et,er,
even though chipmunk spccies cache

nonrodent heclgehog, although

as the

period of hiberrration

concentrations, horvever, there

is

r'vas extendecl

ccinsiclerable contrr)versy in the lirnited

food prior to hibernation, they

undergo

these values increased slightly. Plzrsnra

prehibe rnation fattenins that is as extensivc as the fattening in the ground squirrel (14) The cachir-rg of food reserves may result primarily from the demand {br protein rather than the neecl for energy. Although the storage of f'at as an energy sourcc f or utilization during hibernation is well-established in ground squirrels, there are n<r studies that examine the minimum body protein recluired prior to hibernation. Carpenter (7) has reported that the declinc in nitrosetr elirnina-

lvsine, and ornithine \{ere elevatecl

levels of' valine, leucine, isoleucine,

thtrughout hibernation.'l'his

same

study reported no changes in plasma arnino acid conce nttatiolr in the

tridecemLineatus (8), S. coLutnbianus (20), and Malmotu, .flaaiuentri.s (15).

literature. Plasma ure a concerltrati<ins have been reporte d not to change durins hibernation in S.
However, Passrnore et al. (27) rep()rt increased plasma urea in S. columbi-

hamster cluring hibernati<irt. Klair-r irrrd Whitten (17) exanrined S. trirlecemlirteatus and

reported little change in plasrna :rmino acid concentrations rvith hiberr.ration. Plasma lcucine, :rreinine, and alanine rvere the ortly

amino acid concer)tr:ltions that intle:rscd u ith lribelrr:rtit,n.

nnzs while Fisher (10) and Galster and M<lrrison (l l) have noted reductions in S. Lateralis and S. undulcttus, respectively. Obviously there is no consistent pattern that may be deduced from these clata on urea. Analysis of tissue content of non-

tion cluring hibernation paralleled the decline in oxygen c()nsumption. Catabolism of protein when thc renal

rorrle of ttitrogett ex( rcl ion is c\\etttially nonluncli,rrtal mav nt t estit:tl<' the stclrage or recvcling cif nitrogerr.

Arnino acids have been recognized as an irnportant s()rlrce ol e ne rey durins arousal frclm hibernation; this is inf'errecl trorn both increased levels of plasnra amino acids (18) and
clevated rates ()f':rmino acid oxiclation clurins the arousal pr(xress (6, 3l, 32). Alanine has irnportant roles in both nitrogen rnetabolism and gluconeosenesis. In the glucose-alanine cyclc, skelet.al muscle scl-ves as the sourcc o{'alanine. Following uptake o{' this arnino acicl by the liver, the anrino group may be {urrrreled to either urca amino acicls. The carbotr skelettin may subsecluently be c<invertecl to glucosc. Plasma lcvels clf'alanine rise markedly cluring arous:rl of the l3-lined sround

protein nitrogen has resulted

ir.r

several interesting observatiot-ts. Kristofferson (18) rep<lrts an elevated urea content of both diaphragm and

GLYCOGEN RESERVES -l'he maintenance of ulvcogen st()res in both muscle and liver throughout hibernation serves as zr clue to the irnportance of gluconeogenesis during hibernzrtion (16, 34). -fhc interpretation of data collected on animals hibernatirrg irr the lab<,ralory is r,rntplicated by the recent report o1'Bintz et al. (5) on fielcl anirnals. 'I'he fielclkilled animals had live r glycoecn values less than 8% oI' labolatory acclimated anirnals. Grouncl squirrels housed in the laboralory fbr I year arrcl f:rsted firr 8 days to 30% rveipJht loss had 4.5 times the liver gl,vcoecn <lbserved in the field anirnals. Bintz's data support the hypothesis thzrt gluconeogenesis may be a rnore important source of blood glucose than is glycrigenolysis firr the fielcl animuls. Hrrrrevet thcrt' is tl() ( ()llclusive evidence regarding the exte-nt to rvhich gluconeogenesis in the hibernating rodent rcsults f rtlm trtilization of'- anrino acicls <ir glycerol

production ()r synthesis ol other

squirrcl (17), as cloes the rate of alanine conversion to glycogen (32). As mentioned earlier, the cost of arousal ancl periodic norrnothermia accounts lirr 90% of the e ncrsy

creasing arnmonia c()ntent in both cerebrum and celebellum with increasing duration of hibernation in S. pygmacus. At the same time that amrnonia content increased, urea concerltration fell. The recent demonstration that methylated arnino acids n'ray serve as an index <ll skeletal muscle protein tLrl'nover shoulcl be applied to hibernators (33). Deterrnination of the amount of rnethylated amino acids accumulated during hibernaticxr would identify the extent of uluconeogenesis fiom skeletal muscle proteins. Data we have collected on summer anirnals indicate urine- and bloodTABLE 1. Calculated water balance in

small intestine durins hibernation in the hedeel'rog. In addition, Err-rirbekov and Mukailov (9) report in-

expenditure during the winter hibernatins se:lson (30); thus alanine ancl other amino acids nray represent a large portiotr ofthc cnergy expendiIrrle rturirtrr thc Iribernating seasort.
NONPROTEIN NITROGEN

a rodent hibernator
Sourus oJ'uater Frec water (adipose

]'he

hypethesis

6f' nitr',rgep()1s \\astes and the

that

accunrulati.n
re-

are cagsal factgrs ip the perioclic derived frorn the breakdown oi' altusals {}rm hibernatiol has reneutral fats. ceived attention for a number of
years. The evidcnce that renal func-

sultant to*i.iry alcl osrn.tic intbalance

- 25%; muscle - 75%) Adipose, l0 kcal : I . 1 g yields 0.4 ml rvater Muscle, 10 kcal : 2.5 g yields 7.5 ml water Metaboli<: u'ater (fat : 1.07 ml/g; muscle - 0.4 rnl/g) Aclipose, 10kcal - 1.1 eyields 1.2 ml water Muscle, 10 kcal : 2.5 g yields i.0 ml water Iter uatt-r pius meltb()lic watet' Fat
Protein

O.4

7.5

+ 1.2 - l-6 rnl + 1.0 - 8.5 ml

EWL
0.70 g/liter O,

tion i.s inrpaired during hibernation (27) lent inclirect support f or the Plasma concentration ol amin<i acids hypclthesis, as did early measuremay serve as a useful indirect Ineasure ments OI'plasrna nonprotein nitrogen of the rate of proteiu tul'Ir()vcl'. concentrati<tns, lvhich revealed values Kristoffersson and Broberg (19) re- several-fbld higher in hibernators ported lor'r,ered plasrna levels for rnost than in mammals in seneral (10, 25).
AMINO

ACIDS

Eu't!,'rnlit t turtlcr 1,"' I EWLt Oxygen neecled

Fat - 2.2 liter Protein = 2.4 liter

Gain
less lo.ss

per 10 kcal

Fat
Protein
0.06 gain

: 1.54 g - I.68 g

Fat
Protein

amino acids during hibernation of the

2960

With

re spect

to pl:rsma ()r serurrr urea

1.6

8.5

- I.54 * 1.ti8 :

6.8 gain

FEDERATION PROCEEDINGS VOL.39, NO.

12

OCTOBER 1980

o
=

yields more water per calorie of energy than does catabolism of

600 fat (Table l).
UREA CATABOLISM AND NITROGEN RECYCLING Recycling of urea nitrogen in a large hibernator, the black bear, has been

500

UJ

3 F

4oo

UJ

e,
3oo

by Dr. Nelson in this symposium (26) and recycling might well be anticipated to take on a somewhat different form in smaller species
described

fi E,

2oo

100

DAYS
Figure l. Seasonal effects of f astins or Iirstins:rnd warer deprivation (FWD) on daily urea excretion. FWD rats (Dec.) a a FWD p;round scluirrels (May) n [l F-WD ground scluirrels (Dec.) I l fiisted ground squilrels (May-July) O O. Vertical lines represenr seivr. -

urea values are n()t g(x)d indicat()rs of'the extent of protein catabolism.
MUSCLE AS A SOURCE OF WATER

Whereas hiberrrati,rrr is

rrr energt'ti-

cally successf ul approach to cr()nserviiFigure 2. 'I-irrre course r>f bloorl rrrea charrqes
firlkrn'ing urea loading. Ve rtical lines lepreserrt
DISAPPEMANCE OF UREA INJECTION

5
E l o
J m

s o fl

tion of limited resources, the state of hibernation is also ass()ciated lvith a number of limitations clr disadvantages. Of paramount concern is the extended absence o{' water intake in the face ofa reduced, but continuous, evaporative water loss (EWL). 'I'he EWL is minimized in part by the hibernating posture, which promotes rebreathing of humidified air (4). Loss of' body water, much of'n,hich may be ascribed to pulnrontrry insensible water loss, arn()Llrlts t() one quarter of the total weight krss cluring the hibernation season (12). It rvould be of advantage, therefitre, to consider not only the energy available f rom catabolism of a specific tissue but also the potential f<rr procluction of'rnctabolic water or release Of' free water. In contrast to its relativc absence in stored lipid, free water represents 7 5% oI the mass of muscle tissue. As a result of' the elevated ventilation required to provide zrclequate oxygen fbr complete oxidatirtn of highly reduced lipid compounds, the total water available {'rom oxidation clf' fat is near the pulrnonary F,Wl- value. Catabolisnr of protein

capable of hibernation. There are rnajor differences in both physiological and metabolic aspects of hibernation between bears and smaller mammalian hibernators; for instance, capacity for energy storage, depth of torpor, and periodic arousals in small hibernators. 'I'he potential firr urea recycling has been studied in fasted and in fasted, water-deprived (FWD) ground squirrels (5. LateraLis). These conditions, lvhich are accompanied by a reduction in urine f<rrmation and excreti()n, mimic the situatic-rn faced annually during hibernation. Bintz et al. (2, 3) describe a pre{'erence fcrr protein catabolisrn in the foodand water-deprived golden-mantled ground squirrel. In terms of' water balance tl're advantage of protein over fat catabolism is evident from the above discussion. The 60 to 70% reduction in urea excretion associated with fasting or FWD in S, Lateralis (Fig. l) appears to be in conflict with Bintz's evidence of increased protein catabolism under the same conditions. Urea-loading experiments involving laboratory rats and ground
Figure 3. Tinrt tourrc ol urine urer exctetion urea loading. Vertical lines represent

l:11:"'"t
I
3 ! o o

Qcrcuuo

sutRlLs

l)

tae nars

5
U

r l
l

z
U

E
F

PROTEIN AND FAT METABOLISM DURING MAMMALIAN HYPOPHAGIA AND HIBERNATION

2961

squirrels injected intraperitoneally

with 0.3 g urea per 100 g body weight reveal an absence of plasma urea accumulation in the ground squirrel (Fig. 2). The same load results in a

TABLE 2. Percentage of inlected taO collected in urine and expired air in control, fasted-water-deprived, and fasted-water-deprived-drug treated ground squirrels
Percentage of injected taC

collected in 24 h T'wenty-lour-hour urine volume (ml) Urine CO, trap

threefbld elevation of plasma urea in the rat. The absence of plasma urea accumulation is not a result of more efficient renal excretion of urea in the ground squirrel, as evidenced by the fact that urea excretion is maintained at or below the levels seen in the rat (Fig. 3). We may conclude from this evidence that a urea load sufficient to elevate blood urea threefold in the laboratory rat does

Control (N

9)

Fasted-water-deprived (N

8.9 + 0.7o
8)

f penicillin (N : 5) + sulfa (N : 4) * tetracycline (N : 3)

u

1.1

0.7

79 + 4.4 2.6 + 0.80

2.4 + 0.I5 26.5 + 0.80 7.3 * 0.9 4.4 + I.2

1.4

0-6

Mean I sr.

not cause short-term accumulation of urea in the blood or enhance urea excretion in the ground squirrel.

Figure 4. EfI'ect of lasting or f asting combined with drug treatment (FDT) on daily weight-specific urea excretion. Vertical lines represent sEM.

Catabolism

been examined in FWD S. lateralis. ln these animals, fasting for'4 days cclnrbined with 24 h of water deprivation

of

[laC]urea has

alscr

results in a lO-fold increase in ulea catabolism and a 30-fold reduction of labeled excretion (Table 2). Several monogastric mammalian species, including man, have been shown to

.9 o)

-c

= E

d)
ctr

harbor microorganisms capable of o urease activity (21). Treatment of FWD ground squirrels with several microflora-reducing antibiotics re- c' sults in return of urea catabolism 6 to near control levels. In addition, !t d, antibiotic treatme nt results in en- E o hanced excretion of urea, with drugx

treated squirrels excreting three times as much urea as fasted animals, thus

lrJ

indicating storage or recycling of E approximately 75% of the urea :)

produced in fasted animals (Fig. a). These data suggest that retention and catabolism of urea may be components of the metabolic response to food and water deprivation in the ground squirrel. Although there have been no direct studies of' the potential for urea recycling in the hibernating state. examination of the literature reveals evidence of urea recycling. Both in the hibernating arctic ground squirrel and in the hibernating marmot, loss
of lean body mass cannot be accounted REFERENCES

otr

45
Doys of Fostirg
crease body protein in excess of the am()Llnt of urea excreted durinq the hibernation season is the best evi-

fbr by excreted urea (l, I l, 12). Thus these animals exhibit charactelistics similar to fasted water-deprived active S. lateralis in which the potential Ibr urea recycling was high.

of energy during arousal, and de-

hibernatol to recycle urea nilrogen. utilize anrino acids as a major source

In summary, the ability of

the

of the vital role of adjustments in protein metabolism made for the survival of the rodent hibernator. E
dcnce in support

I.

2. Bintz, G. L. Oxidation of

Benedict, F. G,; Lee, R. C. Hibentation and ma,rmot physiology. Carnegie Inst. Washington Publ. 497, Baltimore: Waverley Press; 1938, table 36.
laC-labeled

lisrn by water-deprived laboratory rats

philns kueralis. Cornp. Biocherrt. P\siol.

and ground squirrels. Cryobiologl

8:

38A: l2l-127:1971.

carbohydrate, fatty acid, and amino acid as a cr-iterion of selective tissues catabo-

315;1971 (abstract). 3. Bintz, G. L.; Bintz, L. B.; Riedesel, M. L. Respiratory quotient as an index of selective tissue catabolism by rvater. cleprived laboratory rats and Spermo-

4. Bintz,

G. L.; Roesbery, H.W,Evaporative rv:rtel loss bv control and starved labtirat()ry r:rts and Spermophilus richord.sonii. CrmQ. Biothern. Phlsiol. 59A: 275-278 I 978.

2962

FEDERATION PROCEEDINGS VOL.39.

NO.12

OCTOBER 1980

Bintz, G. L.; Roesbery, H. W.; Bintz, L. B. (llycoscn levels in held ancl laboratorv:rcclimatecl Rit:har<lson uround stluir'rt:ls. Camp. 3'12; M79.

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l5 Kastner, P.

R;

Zatzman,

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Burlington, R. F.; Klain, G. J. (ih,rt:oneo1

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F. E,;Johnson" J.

A, Rertirr-;rrrtiulpr)s;rr-

gcncsis <lrrring hibet-rratiotr :rncl arrtrtsal trrrn lribet nat i ctn. ( ) o n p. [] i oc hen L. P I t1.s io!.

Rl82:1978.
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fbod protcins in a hibernator, the galclen dcrrmouse (ELiomys quercinus I'). Comp. Biochem. Physiol. 62 A: 37 I -37 6; 1979. Musacchia, X. J.; Wilber, C. G. Studies on the biochemistry of the arctic ground srluirrel../. Mamtnal. 33: 356-362; I952 2ti. Nelson, R. ,4. Protcin and fat met:rbolism

7.

22: 701-70t3; l9ti7. Carpenter, T. M. Thc par-titioir oI urin:rry nitlogen ol f:rsting ancl hibcrrr:rting
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343-3,17; 1938. Chilian, W. M.; Tollefson, D. Rlood urea

levcls anc[ cr'\,thr'ocvte fiagility to isos-

J.

Bir:tl.

Kayser, C.The phl.siobn of natural hibertur 1ioz. Ner, Yolk: Peleamon Press: l96l: 325 p. Klain, G. J.; Whitten, B. K. I'lasma fl-ee amino acids in hibernation and arnusal.
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Proc.

39: 2955-2958: 1980. Fassmore, J, C.; Pfeiffer, E. W.; Templeton,J. R. flrea excrction jn the hibcrnatirrg (lrrlurnbian grollnd sclrirrel (Spermo-

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:rnd

l8 Kristoffersson, R. Urei-lcve ls in blood and tissues of hibcrnating and nttn1r

lthilus col,umbian,us). J. I)xp. Z'ool 192: 83-86:1975. Tevis, L., Jr. Autumn foods of chiprrunks
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ibcr-rratin g hccl ge ho gs. N ah t r e 197:'102-403; 1963.

LLt

nd.ott)

l9 Kristoffersson, R.; Broberg, S.

Free and

29

Emirbekov, E. Z.; Mukailov, M. I. f)otrtent

ol anrmonia, glrrtanrine, ancl urea

tion.
B1zr11. Eh.sp.

and non-hibernalins Iiedgehogs

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33:198-205; I952. Tevis, L., Jr. Stt>m"lch contellts of chipnrunks and matrtled scluirrels in northeastern Calilbrn ia. J . Mam,nal. 31:.
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in

lrrain tissucs of strsliks rlrrrinc hibern:r-

Biol. X'ltd.69(4):

6'1 66:1970. l0 Fisher, K. C. On thc rncchanism of perioclic :rlousal in hibern:rting sroun(l

sqr-riltel. Srromal:rincn. P., cd. Mnn nulian IlibertLolitn II. ArLrt. Acarl. Sti. F-enrt. Scr..,\4 196.1: l,+.1 -156. Galster, W.; Morrison, P. R. Glrrconeosencsis in ar ctic grouncl scluin'els betrvecrr pcriods of hibelnatiort. Am. J.
Phy.siol.

130: 3-22: 1968. I-esser, R, W.i Moy, R,; Passmore, J. C.; Pfeiffer, E. W. Renal rcgulation of urea excrction in arousitrg :rncl httme'.rtherm ic ground scluirrels (C ite I l.us c o l.umbianu.ii. Conp. Biochern. PhysioL. 36:
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rLnd, th,ernngen.e.sjs. Ncrv

York: Academic I'ress; t978: i09-145. Whitten, B. K.; Burlington, R. F.; Posiacid c:rtabolism during arortsal from
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PROTEIN AND FAT METABOLISI\4 DURING MAMMALIAN HYPOPHAGIA AND HIBERNATION

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