Relationship between food availability, glycerol and glycogen levels in lowtemperature challenged rainbow smelt Osmerus mordax

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1 866 The Journl of Experimentl Biology, Published by The Compny of Biologists 7 doi:.4/jeb.749 Reltionship between food vilbility, glycerol nd glycogen levels in lowtemperture chllenged rinbow smelt Osmerus mordx Willim R. Driedzic nd Connie E. Short Ocen Sciences Centre, Memoril University of Newfoundlnd, St John s, Newfoundlnd, AC 5S7, Cnd Author for correspondence (e-mil: wdriedzic@mun.c) Accepted June 7 Rinbow smelt Osmerus modx ccumulte glycerol in winter tht serves s n ntifreeze. Fish were held t 8 C, or subjected to decrese in wter temperture to C over 9 dy period, nd subsequently mintined t C from 5 Jnury to My 4. Strved fish did not survive the chllenge of temperture decrese, with deth ensuing bove the typicl freeze point for mrine teleosts (.8 C). A decrese in temperture ctivtes the glycerol ccumultion mechnism t bout 5 C with pek plsm levels exceeding mol ml. Glycerol levels begin to decrese in lte Februry even t wter tempertures below C, suggesting either n inherent circnnul or photoperiod trigger, possibly in ssocition with sufficiently high levels of ntifreeze protein. Glycogen levels in liver did not chnge significntly in strved fish mintined t 8 C. However, liver glycogen ws depleted in fish subjected to Summry the low-temperture chllenge nd t fster rte in strved thn in fed fish. Stored glycogen in liver nd other tissues cn ccount for only smll mount of the totl glycerol production, suggesting strong requirement for food during ccelerted glycerol production. Liver glycogen levels incresed in April nd My in ssocition with the decrese in glycerol. Levels of glycerol in liver, kidney, spleen, gill, intestine, hert, muscle nd brin follow the sme pttern s tht in plsm. During the erly prt of the glycerol ccumultion phse, ll tissues except for liver hve lower levels of glycerol in the intrcellulr spce thn the levels in plsm. In liver, glycerol is in equilibrium between the two comprtments. Key words: rinbow smelt, Osmerus mordx, glycerol, low temperture, freeze resistnce. Introduction Rinbow smelt Osmerus mordx depress the freezing point of their body fluids through combintion of ntifreeze protein (Ewrt nd Fletcher, 99) nd elevted glycerol levels (Rymond, 99). These fish remin ctive nd continue to feed in icy sewter, but fce the chllenge of continul loss of glycerol cross the epitheli (Rymond, 99), resulting in the necessity for n ongoing synthesis of glycerol. Studies involving injection of rdioisotopes nd hevy isotopes revel tht glycerol is produced from glucose nd mino cids, which would be obtined from the diet nd/or on-bord reserves (Rymond, 995; Rymond nd Driedzic, 997; Wlter et l., 6). Liver is primry site of glycerol production, s indicted by enzyme complement, gene expression nd synthesis by isolted liver preprtions (Driedzic nd Ewrt, 4; Driedzic et l., 6; Liebscher et l., 6). Glycogen in liver is importnt s metbolic source of glycerol. Glycogen in liver decresed by 75 h post cpture in fish held without feeding t C nd muscle glycogen content ws only smll percentge of the level in liver (Rymond, 995). Liver glycogen declined significntly more t C thn + C following 4 or dys without food (Rymond et l., 996) but even in fed fish, under lbortory conditions, liver glycogen decresed s winter progressed from December to April (Treberg et l., ). The picture tht emerges is tht dietry glucose, mino cids nd glycogen reserves support glycerol synthesis for ntifreeze protection. In the present study, detils of the interply between temperture, food vilbility, onbord fuel reserves nd tissue glycerol content re ddressed. Glycerol content in plsm of rinbow smelt begins to increse when the temperture decreses to bout 4 C, nd my rech levels pproching 5 mmol l (Lewis et l., 4; Driedzic et l., 6). Glycerol levels in tissues other thn plsm re not well understood. In rinbow smelt smpled t one time point in winter t C, the glycerol content in number of different tissues ws similr to tht in plsm (Rymond, 99). In prtil sesonl study, glycerol levels in muscle followed similr profile s in plsm; however, t the pek of plsm glycerol levels the content of glycerol in liver ppered to be lower thn in plsm (Treberg et l., ). A similr reltionship ws noted in fish tht were force-chilled (Driedzic et l., 6). Plsm glycerol content peks in Februry nd therefter decreses, even though wter temperture is still below C. Plsm glycerol content reches 5 mmol l in My, similr to levels in the fll. As plsm glycerol decreses, ntifreeze protein becomes the dominnt freeze protection mechnism (Lewis et l., 4). Although decreses in plsm glycerol

2 Glycerol production in rinbow smelt 867 content while wter tempertures re still very low re documented, it is not cler if other tissues follow this pttern nd if the decrese in plsm glycerol is triggered by smll upswings in temperture towrds C or some other fctor such s photoperiod. Also, it is not known if one of the ftes of plsm glycerol is reincorportion into the glycogen pool. The current experiment ddresses number of questions relevnt to glycogen nd glycerol mngement in freezeresistnt rinbow smelt. Foremost, we tested the hypothesis tht continul feeding is requirement to survive low tempertures. The initil glycogen content in number of tissues is reported. Therefter, glycogen levels in liver nd glycerol level in numerous tissues were determined in fish mintined t low temperture in winter through to the spring period. The quntittive importnce of exchnge between these two metbolic pools nd the reltionship between plsm glycerol nd tissue glycerol levels s the former chnges ws ssessed. Mterils nd methods Animls nd experimentl protocols Rinbow smelt Osmerus mordx Mitchill 84 were collected by seine netting from Long Hrbour, Plcenti By, Wter temperture ( C) Cumultive mortlities Jn A B Feb Mr Apr My Jun Force chilled strved Force chilled fed Ambient fed Smpling dtes Force chilled strved Force chilled fed Newfoundlnd, Cnd in lte October nd 5, trnsported to the Ocen Sciences Centre, Memoril University of Newfoundlnd, nd trnsferred to l () or 8 l tnks (5) with flow-through sewter. Fish were kept on nturl photoperiod with fluorescent lights set on n outdoor photocell nd fed diet of chopped herring unless otherwise indicted. During the /4 period, fish were fed twice week; during the 5/6 seson, they were fed dily. Some fish were held t 8±.5 C for the durtion of the experiments in both the /4 nd the 5/6 studies. Most of the experiments were conducted over the /4 seson with fish subjected to controlled decrese in wter temperture. In these studies, rinbow smelt were trnsferred to 5 l tnk set t 8 C, weeks prior to reduction in wter temperture. On 5 Jnury 4, temperture ws decresed over 9-dy period to C nd subsequently mintined t pproximtely C until the finl smpling point on My 4. In this experiment, fish either continued to receive food or were strved from the initition of the wter temperture decrese. A popultion of fish ws lso llowed to trck mbient wter temperture during the /4 seson. A group of fish from the October 5 collection ws fed nd mintined t 8 C until smpled on 6 Jnury 6. Therefter, this popultion ws strved for 6 dys nd smpled gin. All experiments involved both mle nd femle fish. At smpling times, fish were rndomly selected, weighed, mesured for length, nd blood drwn vi cudl vessel. Fish were then killed with blow to the hed, tissues removed nd stored t tempertures below 65 C for lter glycerol nd/or glycogen nlysis. Blood ws centrifuged t 9 g immeditely fter smpling, plsm ws collected nd frozen in liquid nitrogen. Biochemicl ssys nd wet weight determintions Glycerol level in the plsm ws determined directly using colorimetric detection kit (F648, Sigm-Aldrich, St Louis, MO, USA). Smples were red t 54 nm fter 5 min incubtion t room temperture. Tissues were homogenized in 9 vol. % perchloric cid, the homogente centrifuged t 5 g nd the superntnt ssyed for glycerol. Glycogen ws mesured by the Fig.. (A) Temperture profiles. Wter temperture ws decresed by cooling, beginning on 5 Jnury 4, nd fish were either strved (filled circles) or fed (open circles). Control fish (squres) were mintined under mbient wter temperture conditions (4) nd fed. Filled inverted tringles indicte smpling dtes. (B) Cumultive mortlities for fed nd strved rinbow smelt Osmerus mordx subjected to controlled decrese in wter temperture. The dte t which the temperture decrese ws initited (i.e. 5 Jnury 4) is used s dy. Initil popultion size ws for both groups.

3 868 W. R. Driedzic nd C. E. Short method of Wls nd Wls (Wls nd Wls, 95) s described by Driedzic et l. (Driedzic et l., 998). Percentge wter content ws clculted from the difference in mss between fresh tissue nd fter drying to constnt mss t 9 C. Dt nlysis Unless otherwise indicted, vlues (mens ± s.e.m.) were compred with one-wy nlysis of vrince (ANOVA) for ll mesurements followed by Duncn s post-hoc test. In some situtions involving comprisons between two vlues, t-test ws pplied; P<.5 considered s sttisticlly significnt. Results Reltionship between temperture nd vibility in fed nd strved fish Temperture profiles re presented in Fig. A. Most of the experiments were conducted with fish mintined t 8 C from the time of cpture on 6 October nd subsequently subjected to controlled decrese in temperture on 5 Jnury 4. Wter temperture ws decresed over 9 dys to C nd, with the exception of -dy spike to. C on 9 Mrch 4, mintined under C until My 4. A control popultion of fish ws mintined t norml sesonl tempertures. On 5 Jnury 4 mbient wter temperture ws C nd decresed to. C by 6 Februry 4. Additionl experiments were conducted on smll number of nimls tht were mintined t 8 C in Jnury/Februry 4 nd 6. The cumultive mortlities ssocited with the controlled decrese in temperture re shown in Fig. B. The figure is set with dy s the strt of temperture decrese. Ech group [Glycogen] (µmol glucosyl units g ) A B Brin c (.9) c (8),b,c Gill Gut Gond Hert Kidney Liver Muscle Spleen (.5) (.) (.),b () (.) ,b b (.),b Strved Fed Ambient fed b,c (.4),b,c (.) strted with fish, of which six helthy fish were removed on ech of the dtes shown in Fig. A. There is cler difference in the spontneous deth rte between fed nd strved rinbow smelt. Fish tht were fed survived well. By dy t. C, there were nine recorded deths with more between dy nd dy 5 t tempertures of C or lower. In contrst, in the group of strved fish, ll individuls hd succumbed by dys (4 Februry 4), when the temperture hd reched. C. Mny of the fish died t tempertures well bove the nticipted freezing point of body fluids. For instnce, 5 fish died by dy ( Februry 4), when the wter temperture ws still bove.8 C. A typicl teleost should be ble to survive these tempertures even in the bsence of ntifreeze mechnisms. In the /4 experiment, there were no inexplicble mortlities in fish tht were fed nd mintined t either mbient or elevted tempertures over the time window of the bove described experiment. In the 5/6 study, eight pprently helthy fish tht hd been mintined on heted wter, were food deprived beginning 6 Jnury 6. One fish died over the next 6 dys, when ll hd been smpled. Glycogen levels Tissue-specific glycogen levels were determined in rinbow smelt held t 8 C (Fig. A). Fish were smpled on 5 Jnury 4. Glycogen content ws highest in liver t 7±64 mol glucosyl unit g, followed by hert t 5± mol glucosyl unit g. Muscle nd gut levels were pproximtely 7 mol glucosyl unit g, with lower levels being noted in the other tissues. In fish tht were fed nd subjected to decrese in temperture, there ws significnt decrese in liver glycogen by dy 5 (Fig. B). The glycogen level remined low until dy 4 (9 April 4;.4 C temperture) nd dy 6 ( My 4;. C temperture), t which time glycogen levels incresed to mounts not significntly different from the initil vlue. Liver glycogen in strved fish decresed to minimum level of 7.7±5. mol glucosyl unit g by dy nd remined low until the lst smple ws tken on dy. The glycogen level on dy ws significntly lower in strved thn in fed fish (t-test; P=.). Fig.. Glycogen levels in rinbow smelt Osmerus mordx. (A) Glycogen levels in vrious tissues of fish mintined t 8 C nd smpled on 5 Jnury 4, immeditely prior to the temperture decrese. Note the brek in xis tht seprtes liver from other tissues. Vlues re mens ± s.e.m., N=4. Different letters over brs indicte sttisticlly significnt difference between tissues. (B) Glycogen levels in liver of fish subjected to controlled decrese in wter temperture with either feeding (open circles) or food deprivtion (filled circles). The dte t which the temperture decrese ws initited (i.e. 5 Jnury 4) is used s dy. Filled squre, liver glycogen content in fish mintined t mbient temperture. Vlues re mens ± s.e.m., N=4. Different letters indicte sttisticlly significnt difference between time points for the fed group. Wter temperture ( C) is shown in prentheses.

4 Glycerol production in rinbow smelt 869 Tble. Liver glycogen, nd plsm nd liver glycerol levels in rinbow smelt Osmerus mordx mintined t 8 C nd either fed or food deprived Fed Strved [Glycogen] ( mol glucosyl unit g ) 5±. 9±6 [Glycerol] Plsm ( mol ml ).4±.6.6±.9 Liver ( mol ml ) 9.8±5. 6.5±5. Fed fish were smpled on 6 Jnury 6 (N=). Strved fish were smpled on Februry 6 (i.e. following 6 dys of without food) (N=6). Asterisks indicte vlues tht re significntly different between fed nd strved fish. Fish tht were fed nd held under mbient temperture conditions hd liver glycogen level of 7.±.5 mol glucosyl unit g (smpled 6 Februry 4; C) (Fig. B; squre symbol). This vlue tended to be lower thn the levels of 7±64 nd 5± mol glucosyl unit g noted in liver of fish held t 8 C nd smpled on 5 Jnury 4 (Fig. ) or 6 Jnury 6 (Tble ), respectively but significntly (ANOVA; P=.) higher thn the.±.9 mol glucosyl unit g in fish subjected to the forced decresed in temperture (Fig. B; dy ). In the 6 experiment, the liver glycogen content in rinbow smelt held t 8 C nd fed ws no different thn in fish subjected to 6 dys of strvtion t this temperture (Tble ). Plsm glycerol levels Plsm glycerol in fish tht were force chilled nd fed incresed from 5 mol ml to levels in excess of mol ml t dy 4 (7 Februry 4; C) (Fig. A). Therefter, plsm glycerol concentrtion decresed to 7.4±9.7 mol ml on the finl smple dte ( My 4;. C). Rinbow smelt subjected to the chllenge of decrese in wter temperture, without food, showed n increse in plsm glycerol content 4 5 A (8) B similr to fed fish for the first dys of the experiment (Fig. B). Glycerol levels reched 5 7±7.4 mol ml by the finl smpling point (6 Februry 4;. C). These dt re bsed only on those fish tht survived the temperture decrese chllenge. Rinbow smelt mintined t elevted tempertures hd plsm glycerol levels of less thn mol ml (Fig. A). Food deprivtion for 6 dys t 8 C resulted in significnt decrese in plsm glycerol level (Tble ). Tissue glycerol levels The content of glycerol in vrious tissues in fish tht were subjected to forced decrese in wter temperture nd fed is presented in Fig. 4. These vlues follow the generl pttern of tht in plsm with n increse to dy 4 (7 [Glycerol] (µmol ml plsm) 5 (.9) () Tble. Percentge wter content in tissues of rinbow smelt Osmerus mordx % Wter N=6 in ech cse. Februry 4; C), followed by continul decrese. This pttern occurred in ll tissues smpled. The level of tissue glycerol t the pex of the curves ws highest in kidney t 6±7 mol g nd lowest in gill t 5± mol g. Glycerol levels in fish tht were deprived of food, under the sme therml conditions, followed the sme pttern up to dy, t which time the study ended (dt not shown). The level of glycerol in plsm, expressed s mol ml, ws generlly higher thn in tissue, expressed s mol g, over the first 4 dys of the temperture decrese. As plsm glycerol levels decresed, tissue levels did s well. This prompted first pproximtion nlysis of the intrto extrcellulr glycerol grdient. The percentge wter (.) (.9) (.5) (.5) (.9) (4.5) (7.9) (8.) (.6) (.) (.7) Liver 7.±. Kidney 8.±.5 Spleen 7.5±.8 Gill 84.±.6 Intestine 5.±.4 Hert 78.9±.5 Muscle 77.6±. (.) (.) (.) (.6) (.) (.4) (.) (.8) Force chilled fed Ambient fed Heted fed (.) (.) 5 (8) () (4.5) Fig.. Plsm glycerol levels in rinbow smelt Osmerus mordx. The dte t which controlled temperture decrese ws initited (i.e. 5 Jnury 4) is used s dy. Wter temperture ( C) is shown in prentheses. Vlues re mens ± s.e.m., N=5 or 6 for ll points. (A) Open circles, fish subjected to controlled decrese in wter temperture; filled squres, fish mintined t nturl mbient temperture; filled tringles, fish mintined t elevted temperture. All fish were fed. (B) Fish subjected to controlled decrese in wter temperture without feeding.

5 87 W. R. Driedzic nd C. E. Short [Glycerol] ( mol g ) Liver Kidney Spleen Gill Intestine Hert Muscle Brin () (.5) () (.) content ws clculted for ech of the tissues (Tble ) nd rnged from 5% wter for gut to 84% wter for gill. Wter content did not chnge with temperture (dt not shown). Tissue levels were converted from mol g to mol ml tissue wter nd divided by mol ml plsm to obtin the rtio of glycerol from totl tissue wter to plsm (Fig. 5). A rtio of. indictes equilibrtion of glycerol between the intr- nd extrcellulr comprtments. Vlues less thn. indicte diffusion grdient from plsm to tissue, wheres vlues greter thn. indicte diffusion grdient from tissue to plsm. Precise intr- to extrcellulr grdients cnnot be clculted without knowledge of extrcellulr tissue wter. As plsm glycerol incresed to the pex on dy 4 the level of glycerol ws either close to equilibrium between the two comprtments or there ws grdient from the plsm spce to the tissue spce. Lte in the experiment (i.e. dys 4 nd 6), s plsm glycerol levels decresed, the verge rtio for ll tissues ws in excess of.. Although the vlue for ny specific tissue is not significntly different from., the overll picture shows them to be consistently positive, suggesting tht further study of this my be required in order to determine whether glycerol content in tissue wter might exceed tht in plsm. Discussion Rinbow smelt subjected to low temperture with ccess to food or mintined t high temperture without food survive quite well in cptivity. However, the combintion of food deprivtion nd trnsition to low temperture is lethl. This is most unusul s it is expected tht low temperture should prolong survivl in the bsence of food by reducing energy demnd. Moreover, deth ensued t tempertures well bove the freezing point of body fluids. This could hve importnt ecologicl consequences s it implies tht under field conditions, n bsence of usul food sources s the temperture decreses in the fll, could result in mssive mortlity. We do not know the cuse for the premture deth with respect to temperture nd food deprivtion; however, it is clerly not mtter of simple energy blnce, s rinbow smelt mintined t 8 C tolerte longer periods of strvtion thn fish chllenged with lower tempertures. An understnding of this phenomenon will be dependent upon detiled nlysis of both surviving nd dying individuls. An imposed temperture decrese leds to increses in glycerol levels in plsm nd other tissues, s hs been observed in erlier studies (Driedzic et l., 6). Even though the temperture ws held below C, glycerol in plsm nd other tissues begn to decrese in smples tken fter 7 Februry 4 (i.e. dy 4), with tissue levels of glycerol generlly following the sme pttern s plsm glycerol. This pttern of glycerol ccumultion nd subsequent decrese is strikingly similr to tht observed in two other experiments in which rinbow smelt were mintined under nturl therml nd light conditions (Treberg et l., ; Lewis et l., 4). A decrese in temperture lone is sufficient to ctivte the glycerol ccumultion mechnism nd occurs t bout 5 C, but the signl for reducing the level of glycerol resides elsewhere. Potentil cndites re photoperiod or n inherent circnnul rhythm, possibly in ssocition with sufficient levels of ntifreeze protein. The highest level of glycogen occurs in liver, followed by hert, with lower levels in number of other tissues. At 8 C, strvtion lone for 6-dy period did not result in significnt decrese in liver glycogen, lthough the men vlue decresed by %. This is common for fish, for exmple Atlntic cod held t 8 C showed bout 6% decrese in liver glycogen following 9 dys of food deprivtion (Hll et l., 6). In rinbow smelt, (.4) (.) Fig. 4. Glycerol levels in tissues of rinbow smelt Osmerus mordx subjected to controlled decrese in wter temperture. The dte t which controlled temperture decrese ws initited (i.e. 5 Jnury 4) ws used s dy. Wter temperture ( C) on ech smpling dy is shown on bottom pnel in prentheses. Vlues re mens ± s.e.m., N=4 for ll time points.

6 Glycerol production in rinbow smelt 87 however, decrese in temperture from 8 to. C over only dys resulted in liver glycogen levels close to zero. The decrese in liver glycogen occurs even in fed fish, but not to s gret n extent. As such, either strvtion or low temperture results in glycogen mobiliztion, with the combintion of both hving the most impct. Liver glycogen reserves cn only ccount for smll proportion of the glycerol tht ccumultes. For instnce, liver glycogen with n initil level of 75 mol glucosyl units g could produce 5 mol glycerol g. Of this bout 5 mol g would be retined in the liver with the reminder (i.e. 5 5= mol g ) vilble for export to other tissues. A 5 g fish with liver mss of.75 g could relese 5 mol of glycerol, which if distributed evenly cross ll tissues, would result in glycerol increment of only mol g. With the possible exception of hert, none of the other tissues hve dequte glycogen reserves to ccount for the increse in glycerol content. There is cler need for dietry intke of fuels or dditionl on-bord reserves such s mino cids from protein to support glycerol production. In this context it is importnt to note tht glycerol from triglycerides is not considered to be contributor to glycerol production (Rymond, 995). Tissue [glycerol] / Plsm [glycerol] Liver Kidney Spleen Gill Intestine Hert Muscle Brin ()()(.5) (.) (.4) The decrese in glycerol content in ll tissues observed lte in the study is ssocited with n increse in glycogen content in liver, nd this occurs while the temperture is below C. It is likely tht one fte of glycerol is incorportion into the glycogen pools. Between dys 89 nd 6 the content of glycogen in liver incresed on verge by bout 5 mol glucosyl units g, or the equivlent of 5 mol glycerol g. This could ccount for the decrese in liver glycerol level of bout 5 mol g nd still provide room to ccommodte some of the decrese of glycerol in the plsm nd other tissues. Glycogen levels my cycle in other tissues s well. The importnt point is tht glycogen synthesis cn occur t low tempertures nd could serve s sink for glycerol, suggesting tht glycerol need not be lost to the environment nor metbolized to CO during this period. Glycerol level in tissues follows the sme generl profile s tht in plsm, with n increse s temperture is reduced followed by decrese in level. As plsm glycerol incresed there ws either grdient from the plsm spce to the tissue spce or the level of glycerol ws close to equilibrium between the two comprtments. In liver, on dy 5, there is no significnt difference between glycerol in tissue wter nd in plsm, but for ll other tissues tested glycerol is lower in the tissue wter thn in plsm. This would be consistent with movement of glycerol from liver into plsm nd subsequently down concentrtion grdient from plsm into other tissues. This contention is supported by the relese of glycerol by isolted preprtions (Driedzic et l., 998; Driedzic nd Ewrt, 4). However, on dy 4, when plsm glycerol is t its highest level, glycerol in liver tissue wter is less thn in plsm. This sitution, of lower thn expected concentrtion of glycerol in liver t times of high plsm glycerol, hs been observed in two previous experiments (Treberg et l., ; Driedzic et l., 6). There re number of possible explntions to ccount for this. As previously suggested, bsed on findings with yest, there my be ctive pumping of glycerol out of liver into plsm (Driedzic et l., 6). An lterntive explntion is tht the process of glycerol sequestering into glycogen my be strting with the intrcellulr metbolic steps proceeding fster thn diffusion of glycerol into the liver. A third possibility is tht other tissues, such s kidney ( known gluconeogenic tissue), (.) Fig. 5. Rtio of [Glycerol] in tissue ( mol ml tissue wter)/plsm ( mol ml plsm) in rinbow smelt Osmerus mordx subjected to controlled decrese in wter temperture. The dte t which controlled temperture decrese ws initited (i.e. 5 Jnury 4) ws used s dy. The broken line represents equl vlues for glycerol in tissue wter nd plsm. Asterisks indicte vlues significntly different from.. Wter temperture ( C) on ech smpling dy is shown on bottom pnel in prentheses. Vlues re mens ± s.e.m., N=4 for ll points.

7 87 W. R. Driedzic nd C. E. Short my become site of glycerol relese s the seson progresses. Finlly, when plsm glycerol returns to miniml levels there pper to be higher levels of glycerol in tissues thn would be expected if glycerol ws in equilibrium between the intr- nd extrcellulr spce. Generl conclusions Rinbow smelt must feed to live t low tempertures; in fooddeprived fish deth ensues t bout 5 C. Glycerol ccumultion is ctivted by low temperture lone but the decrese in glycerol tht occurs in lte winter/erly spring is tempertureindependent. The erly phse of glycerol production is ssocited with mobiliztion of liver glycogen; however, glycogen from liver nd in ll other tissues cn ccount for only smll frction of the totl glycerol tht ccumultes. This is consistent with the need to continue feeding t low temperture with constnt supply of ingested crbohydrte nd mino cids for glycerol synthesis. During lte winter/erly spring, even while the temperture is still subzero, glycerol levels decrese with concomitnt increse in glycogen. This work ws supported by Nturl Sciences nd Engineering Reserch Council of Cnd Discovery Grnt (W.R.D.), the Cndin Institutes of Helth Reserch through the Regionl Prtnership Progrm, nd Newfoundlnd nd Lbrdor Deprtment of Innovtion, Trde nd Rurl Development. W.R.D. holds the Cnd Reserch Chir in Mrine Bioscience. We thnk the Field Services Unit of the Ocen Sciences Centre for the collection of specimens nd we thnk Vny Ewrt (NRC IMB) for helpful discussion nd comments on the mnuscript. References Driedzic, W. R. nd Ewrt, K. V. (4). Control of glycerol production by rinbow smelt (Osmerus mordx) to provide freeze resistnce nd llow forging t low winter tempertures. Comp. Biochem. Physiol. 9B, Driedzic, W. R., West, J. L., Sephton, D. H. nd Rymond, J. A. (998). Enzyme ctivity levels ssocited with the production of glycerol s n ntifreeze in the liver of rinbow smelt (Osmerus mordx). Fish Physiol. Biochem. 8, 5-4. Driedzic, W. R., Clow, K. A., Short, C. E. nd Ewrt, K. V. (6). Glycerol production in rinbow smelt (Osmerus mordx) my be triggered by low temperture lone nd is ssocited with the ctivtion of glycerol-- phosphte dehydrogense nd glycerol--phosphtse. J. Exp. Biol. 9, 6-. Ewrt, K. V. nd Fletcher, G. L. (99). Isoltion nd chrcteriztion of ntifreeze proteins from smelt (Osmerus mordx) nd Atlntic herring (Clupe hrengus hrengus). Cn. J. Zool. 68, Hll, J. R., Short, C. E. nd Driedzic, W. R. (6). Sequence of Atlntic cod (Gdus morhu) GLUT4, GLUT, nd GPDH: developmentl stge expression, tissue expression nd reltionship to strvtion-induced chnges in blood glucose. J. Exp. Biol. 9, Lewis, J. M., Ewrt, K. V. nd Driedzic, W. R. (4). Freeze resistnce in rinbow smelt (Osmerus mordx): Sesonl pttern of glycerol nd ntifreeze protein levels liver enzyme ctivity ssocited with glycerol production. Physiol. Biochem. Zool. 77, Liebscher, R. S., Richrds, R. C., Lewis, J. M., Short, C. E., Muise, D. M., Driedzic, W. R. nd Ewrt, K. V. (6). Sesonl freeze resistnce of rinbow smelt (Osmerus mordx) is generted by differentil expression of glycerol--phosphte dehydrogense, phosphoenolpyruvte crboxykinse nd ntifreeze protein genes. Physiol. Biochem. Zool. 7, 4-4. Rymond, J. A. (99). Glycerol is colligtive ntifreeze in some northern fishes. J. Exp. Zool. 6, Rymond, J. A. (99). Glycerol nd wter blnce in ner-isosmotic teleost, winter-cclimted smelt. Cn. J. Zool. 7, Rymond, J. A. (995). Glycerol synthesis in the rinbow smelt Osmerus mordx. J. Exp. Biol. 98, Rymond, J. A. nd Driedzic, W. R. (997). Amino cids re source of glycerol in cold-cclimtized rinbow smelt. Comp. Biochem. Physiol. 8B, Rymond, J. A., Httori, H. nd Tsumur, K. (996). Metbolic responses of glycerol-producing Osmerid fishes to cold temperture. Fish. Sci. 6, Treberg, J. R., Wilson, C. E., Richrds, R. C., Ewrt, K. V. nd Driedzic, W. R. (). The freeze voidnce response of smelt, Osmerus mordx: initition nd subsequent suppression of glycerol, trimethylmine oxide nd ure ccumultion. J. Exp. Biol. 5, Wlss, O. nd Wlss, E. (95). Effect of epinephrine on rt diphrgm. J. Biol. Chem. 87, Wlter, J. A., Ewrt, K. V., Short, C., Burton, I. W. nd Driedzic, W. R. (6). Accelerted heptic glycerol synthesis in rinbow smelt (Osmerus mordx) is fuelled directly by glucose nd lnine: H nd C nucler mgnetic resonnce study. J. Exp. Zool. Prt A 5,

Accepted 10 January 2006

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