摄食、力竭性运动及其交互作用对鲇鱼幼鱼代谢及酸碱状态的影响
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摘要
本研究以鲇鱼(Silurus asotus Linnaeus)幼鱼为实验对象,在24.0±1.0℃下,首先研究了摄食(摄食水平为8%鱼体重)(摄食组)对动脉血酸碱状态的影响,以检验餐后“碱潮”现象是否在该鱼中出现;随后,分别考察了力竭性运动(禁食后运动组)和摄食与运动的交互作用(摄食后6 h运动,即摄食后运动组)对力竭性运动后过量耗氧(Excess post-exercise oxygen consumption, EPOC)和酸碱状态的影响,以检验当摄食与运动同时发生时,由摄食导致的餐后“碱潮”现象是否有利于阻止力竭性运动诱导的血液酸化,并对力竭性运动后的恢复过程产生影响。此外,还测定了摄食、运动及摄食与运动交互过程中耗氧率(Oxygen consumption, VO2)、呼吸频率(Ventilation frequency, Vf)、血红蛋白(Hemoglobin, Hb)、血糖、血乳酸、肌乳酸及肌糖原等生理生化指标的变化。
     研究结果:
     1.鲇鱼幼鱼摄食前动脉血pH(pHa)为7.74±0.02,摄食后3 h内,pHa无显著变化,摄食后6 h,pHa显著增加了0.14,并达到最大值(7.88±0.02)(P<0.05),直到摄食后12-24 h,pHa才逐渐恢复到摄食前水平;
     2.鲇鱼幼鱼摄食前动脉血HCO3-浓度([HCO3-]pl)为5.42±0.09 mmol L-1,摄食后[HCO3-]pl伴随着pHa的显著增加而增加,并在摄食后6 h达到最大值(7.83±0.37)(P<0.05),直到摄食后12 h与摄食前水平无显著差异;
     3.鲇鱼幼鱼摄食后VO2显著升高,摄食代谢峰值时VO2是摄食前的2.24倍,摄食后52 h恢复到摄食前水平;摄食对呼吸频率、动脉血CO2分压(PCO2)、血糖、血乳酸、血红蛋白、白肌乳酸和糖原均无显著影响;
     4.力竭性运动后鲇鱼幼鱼pHa下降了0.46±0.06,[HCO3-]pl浓度下降了约3 mmol L-1。血乳酸含量显著上升,并在运动后1 h到达最大值。pHa、[HCO3-]pl及血乳酸均在运动后4 h恢复到运动前水平。在力竭性运动后的整个恢复过程中,动脉血PCO2无显著变化;
     5.力竭性运动后即刻,鲇鱼幼鱼VO2显著升高并达到最大值,随后逐渐恢复到运动前水平。运动后恢复期内VO2(Y, mgO2h-1kg-1)和恢复时间(X, min)的关系可以用如下方程描述:Y=a±b e-cx;
     6.力竭性运动前鲇鱼幼鱼肌肉乳酸水平为4.8±0.26μmol g-1,运动后2h到达峰值(18.26±0.55μmol g-1),直到运动后8h恢复到与运动前水平无显著性差异;运动前鲇鱼幼鱼白肌糖原水平为18.59±1.36μmol g-1,运动后白肌糖原下降了61%,直到运动后16h仍没完全恢复到运动前水平;
     7.对于摄食后运动组,由于餐后“碱潮”的存在,鲇鱼幼鱼在力竭运动前具有较高的pHa和[HCO3-]pl,因此力竭性运动后pHa和[HCO3-]pl下降幅度(分别下降了0.36±0.05和2 mmol L-1)显著低于禁食后运动组;力竭性运动后的整个恢复过程中,动脉血PCO2无显著变化;
     8.对于摄食后运动组,鲇鱼幼鱼VO2在力竭性运动后立即显著上升,随后逐渐下降。由于SDA的存在,摄食后运动组运动前的VO2水平显著高于禁食后运动组运动前水平。摄食后运动组和禁食后运动组力竭性运动后VO2峰值、VO2上升倍率均存在显著差异。然而两个组EPOC没有显著差异;
     9.对于摄食后运动组,力竭性运动后鲇鱼幼鱼呼吸频率、血红蛋白、血液乳酸、肌乳酸和糖原等指标变化趋势与禁食运动组无显著差异。
     以上结果表明:
     1.鲇鱼幼鱼摄食后表现出显著的餐后“碱潮”现象;该现象为除虹鳟外的硬骨鱼类中的首次报道;
     2.与其它动物不同,鲇鱼餐后“碱潮”现象的补偿(缓解)可能主要通过鳃的作用将过多的代谢碱排泄到周围水环境中而不是依靠血浆PCO2的改变;
     3.鲇鱼幼鱼餐后“碱潮”现象明显地阻止了力竭性运动带来的动脉血液pHa和[HCO3-]pl浓度的立即下降,但是对力竭性运动后的恢复过程并没有显著的影响。
Feeding and exhaustive exercise are known to elevate metabolism. However, acid-base status may be oppositely affected by the two processes. In this study, we first investigated the acid-base response of Chinese catfish to feeding (the meal size was about 8% of body mass) to test whether an alkaline tide (a metabolic alkalosis created by gastric HCl secretion after feeding) would occur. We then determined the combined effects of feeding and exhaustive exercise on excess post-exercise oxygen consumption (EPOC) and acid-base status to determine whether the alkaline tide induced by feeding protects against acid-base disturbance during exhaustive exercise and affects subsequent recovery. At the same time, we investigated arterial blood acid–base status (pHa and [HCO3-]pl), oxygen consumption (VO2), ventilation frequency (Vf,), hemoglobin (Hb), blood glucose, blood lactate, muscle glycogen and muscle lactate concentrations after feeding, exhaustive exercise and both feeding and exhaustive exercise.
     The results as follows:
     1. In fasting fish, the pHa was 7.74±0.02. After feeding, the pHa showed no significant change within 3 h of feeding. However, pHa significantly increased by 0.14 units and reached a peak at 6 h after feeding (7.88±0.02). This transient increase slowly returned to the fasting pHa levels at 12—24 h after feeding.
     2. In fasting fish, the [HCO3-]pl was 5.42±0.29 mmol L-1. The increase in pHa was accompanied by a significant increase in [HCO3-]pl, which also peaked at 6 h (7.83±0.37 mmol L-1) after feeding and then slowly returned to fasting levels at 12 h.
     3. After feeding, the VO2 increased significantly, peaking at about 2.24 times higher than that of the pre-feeding level, and the increased metabolism persisted 52 h after ingestion. Feeding had no significant effect on Vf,, PCO2, [glucose]pl, [lactate]pl, Hb, white muscle lactate and white muscle glycogen.
     4. Exhaustive exercise led to a significant reduction in pHa by 0.46±0.06 units and a reduction in [HCO3-]pl by approximately 3 mmol L-1. After exhaustive exercise, the [lactate]pl significantly increased and peaked at 1 h. The pHa, [HCO3-]pl and [lactate]pl returned to resting levels at 4 h after exercise.
     5. The VO2 increased significantly after exhaustive exercise and gradually returned to pre-exercise level in fasting fish. The relationship between post-exercise VO2 (Y, mg O2 h-1kg-1) and time after exercise (X, min) was described by the following equation: Y=a±becx.
     6. The lactate concentration of white muscle rose acutely from a basal value of 4.80±0.26μmol g-1 to a peak level of 18.26±0.55μmol g-1 at 2 h after exercise in fasting fish and returned to pre-exercise level by 8 h. Exhaustive exercise caused a 61% decrease in glycogen, which remained low for at least 16 h post-exercise.
     7. In both feeding and exhaustive exercise fish, which had a higher pHa and [HCO3-]pl before exercise, exhaustive exercise only caused a reduction in pHa by 0.36±0.05 units and a reduction in [HCO3-]pl by about 2 mmol L-1.
     8. In both feeding and exhaustive exercise fish, the VO2 recovery profile was similar to those of fasting fish. However, there were significant differences in pre-exercise VO2, peak VO2 and the VO2 factorial scope between fasting and post-feeding fish. The EPOC between fasting and post-feeding fish was not significantly different.
     9. In both feeding and exhaustive exercise fish, the elimination of the lactate and the restoration of glycogen were similar to those of fasting fish.
     In conclusion:
     1. As anticipated, we detected a significant alkaline tide in Chinese catfish after feeding. To our knowledge, this is the first report of an alkaline tide in a teleost other than rainbow trout.
     2. Unlike other vertebrate classes, the alkaline tide in Chinese catfish may be compensated by excretion of metabolic base to the environment, so the PCO2 of the arterial blood did not change during digestion.
     3. The alkaline tide did dampen the reduction in pHa and [HCO3-]pl after exhaustive exercise, but recovery from the exhaustive exercise was not affected by digestion.
引文
[1]陈湘遴.我国鲇科鱼类总述[M].水生生物学集刊,1977,6: 197-218.
    [2]付世建,曹振东,谢小军.鱼类摄食代谢和运动代谢研究进展[J].动物学杂志, 2008, 43: 150-159
    [3]付世建.营养状况对南方鲇摄食代谢和能量分配的影响[M].博士论文.湖北武汉:中国科学院水生生物研究所,2004:1-18
    [4]林浩然,刘晓春.鱼类生理学实验技术[M].广州,广东高等教育出版社,2006,27-29
    [5]李黎,曹振东,付世建(2007)力竭性运动后鲇鱼幼鱼的乳酸、糖原和葡萄糖水平的变动[J].水生生物学报, 31(6): 880-885.
    [6]谢小军,孙儒泳.鱼类的特殊动力作用的研究进展[J].水生生物学报,1991,15(1):82-90.
    [7]彭姜岚,曹振东,付世建.鲇鱼力竭性运动后的过量耗氧及其体重的影响[J].水生生物学报, 2008, 32(3): 87-93.
    [8]施白南.嘉陵江鲇鱼的生物学简介[J].西南师范大学学报,1980,2:53-59.
    [9]郑保珊,陈庆泰.中国鱼类志[M].科学出版社, 1987, 210.
    [10]陶义训等.临床生化检验[M].上海科学出版社,上海, 1979, 274-275
    [11] Alsop DH and Wood CM. The interactive effects of feeding and exercise on oxygen consumption, swimming performance and protein usage in juvenile rainbow trout (Oncorhynchus mykiss) [J]. J Exp Biol., 1997, 200:2337-2346
    [12] Andersen JB and Wang T. Cardiorespiratory effects of forced activity and digestion in toads [J]. Physiol Biochem Zool, 2003, 76: 459-470
    [13] Bahr R, Hansson TT and Sejersted, OM. Triglyceride/fatty acid cycling is increased after exercise [J]. Metabolism, 1990, 39: 993-999
    [14] Bahr R. Excess post-exercise oxygen consumption magnitude, mechanisms, and practical implication [J]. Acta Physiological Scandia, 1992, 605:1-70
    [15] Bennett AF and Hicks JW. Postprandial exercise: prioritization or additivity of the metabolic response? [J]. J Exp Biol., 2001, 204:2127-2132
    [16] Blaikie HB and Kerr SR. Efect of activity level on apparent heat increment in Atlantic cod, Gadus morhua [J]. Can J Fish Aquatic Sci, 1996, 53:2093-2099
    [17] Booth RK and Kieffer JD, Davidson K et al. Effects of late season“catch and release”angling on anaerobic metabolism, acid-base status, survival and reproduction in“wild”Altlantic salmon [J]. Can. J Fish Aquat Sci, 1995, 52: 283-290
    [18] Bonen A, McDermott, JC, Tan MH. Glycogenesis and glyconeogenesis in skeletal muscle: effects of pH and hormones [J]. Am J Physiol, 1990, 21:693-700
    [19] Brown, JR and Cameron, JN. The relation between specific dynamic action and protein synthesis rates in channel catfish [J]. Physiol Zool, 1991, 64, 298-309
    [20] Bucking C and Wood CM. Water dynamics in the digestive tract of the freshwater rainbow trout during the processing of a single meal [J]. J Exp Biol, 2006a, 209:1883-1893
    [21] Bucking C and Wood CM. Gastrointestinal processing of Na+, Cl-, and K+ during digestion: implications for homeostatic balance in freshwater rainbow trout [J]. Am J Physiol. R, 2006b, 291:1746-1772
    [22] Bucking C and Wood CM. The alkaline tide and ammonia excretion after voluntary feeding in the freshwater rainbow trout [J]. J Exp Biol, 2008, 211:2533-2541
    [23] Bucking C, Fitzpatrick JL and Nadella S R, Wood CM. Post-prandial metabolic alkalosis in the seawater-acclimated trout: the alkaline tide comes in [J]. J Exp Biol, 2009, 12:2159-2166
    [24] Busk M, Jensen FB and Wang T. Effects of feeding on metabolism, gas transport, and acid-base balance in the bullfrog Rana catesbeiana. [J] Am J Physiol, 2000, 278:R185-195
    [25] Cooper CA and Wilson RW. Post-prandial alkaline tide in the freshwater rainbow trout: effect of meals anticipation on recovery from acid-base and ion regulatory disturbances [J]. J Exp Biol., 2008, 211:2542-2550
    [26] Dobson GP and Hochachka PW. Role of glycolysis in adenylate depletion and repletion during work and recovery in teleost white muscle [J]. J Exp Biol., 1987, 129:125-140
    [27] Ferguson PA, Kieffer JD and Tufts BL. The effects of body size ob the acid-base and metabolite status in the white muscle of rainbow trout before and after exhaustive exercise [J]. J Exp Biol., 1993, 180: 195-207
    [28] Farrell A P, Thorarensen H, Axelsson M, Crocker C E, Gamperl A K, Cech, J J Jr. Gut blood flow in fish during exercise and severe hypercapnia[J]. Comp Biochem Physiol A. 2001, 128: 549-561.
    [29] Fu SJ, Xie XJ and Cao ZD. Effect of meal size on specific dynamic action in southem catfish (Silurus meridionalis Chen)[J]. Comp Biochem Physiol A, 2005a, 140:445-451
    [30] Fu SJ, Xie XJ and Cao ZD. Effect of fasting on resting metabolic rate and postprandial metabolic response in southern catfish (Silurus meridionalis Chen)[J]. J Fish Biol. 2005b, 67:279-285
    [31] Fu SJ, Xie XJ and Cao ZD. Effect of dietary composition on specific dynamic action in southern catfish, Silurus meridionaliss[J]. Aqua Res, 2005c, 36:1384-1390
    [32] Fu SJ, Xie XJ and Cao ZD. Effect of feeding level and feeding frequency on specific dynamicaction in southern catfish Silurus meridionalis Chen [J]. J Fish Biol. 2005d, 67:l7l-l81
    [33] Fu SJ, Cao ZD, Peng JL. Effects of meal size on postprandial metabolic response in Chinese catfish (Silurus asotus Linnaeus) [J]. J Comp Physiol B, 2006, 176:489-495
    [34] Fu SJ, Cao ZD, Peng JL Effect of feeding and fasting on excess post-exercise oxygen consumption in southern catfish (Silurus meridionalis) [J]. Comp. Biochem. Physiol. A, 2007, 146: 435-439
    [35] Fu SJ, Cao ZD, Peng JL, Wang YX. Effects of meal size on excess post-exercise oxygen consumption in fishes with different locomotive and digestive performance [J]. J Comp Physiol B, 2009, 179:509-517
    [36] Gaesser GA and Brooks GA. Metabolic base of excess post-exercise oxygen consumption: a review [J]. Med Sci Sports Exerc,1984, 16:29-43
    [37] Gleeson TT. Pattern of metabolic recovery from exercise in amphibians and reptiles [J]. J Exp Biol., 1991, 160:187-195
    [38] Gleeson TT. Post-exercise lactate metabolism: a comparative review of sites, pathway, and regulation [J]. Ann Rev Physiol, 1996, 58: 565-581
    [39] Ginneken VV. Blood plasma substrate and muscle lactic-acid response after exhaustive exercise in common carp and trout:indications for a limited lactate-shuttle [J]. Animal Biol., 2004, 54(12) : 119-130
    [40] Graham J B, Dewar H, Lai N C, Lowell W R, Arce S M. Aspects of shark swimming performance determ ined using a large water tunne1[J]. J Exp Biol., 2004, 151: 175-192.
    [41] Hancock T V and Gleeson TT. Metabolic recovery in the desert iguana (Dipsosaurus dorsalis) following activities of varied intensity and duration[J]. Functional Ecology, 2002, 16: 40-48
    [42] Hicks JW, Wang T, Bennett AF. Patterns of cardiovascular and ventilatory response to elevated metabolic states in the lizard Varanus exanthematicus[J]. J Exp Biol., 2000, 203:2437-2445
    [43] Hicks JW, Bennett AF. Eat and run: prioritization of oxygen delivery during elevated metabolic states. Res Physiol Neurol, 2004, 144:215-224
    [44] Hunt von Herbing I, White L. The effects of body mass and feeding on metabolic rate in small juvenile Atlantic cod [J]. J Fish Biol., 2002, 61:945-958
    [45] Jobling M and Davies P S. Effects of feeding on metabolic rate, and the specific dynamic action in plaice, Pleuronectes platessa L [J]. J Exp Biol. 1980, 16:629-638.
    [46] Jobling M. The influences of feeding on the metabolic rate of fishes: a short review [J]. J Fish Biol. 1981, 18:385-400
    [47] Kieffer JD, Currie S and Tufts BL. Effects of environmental temperature on the metabolic andacid–base responses of rainbow-trout to exhaustive exercise [J]. J Exp Biol., 1994, 194:299–317
    [48] Kieffer JD. Limits to exhaustive exercise in fish [J]. Comp Biochem Physiol A, 2000, 126:161-179
    [49] Kieffer JD, Wakefield AM and Litvak MK Juvenile sturgeon exhibit reduced physiological responses to exercise [J]. J Exp Biol., 2001, 204:4281-4289
    [50] Lee CG, Farrell AP, Lotto A, Hinch SG, Healey MC. Excess post-exercise oxygen consumption in adult sockeye (Oncorhynchus nerka) and coho (O.kisutch) salmon following critical speed swimming [J]. J Exp Biol., 2003a, 206:3253-3260
    [51] Lee CG, Devlin RH and Farrell AP. Swimming performance, oxygen consumption and excess post-exercise oxygen consumption in adult transgenic and ocean-ranched coho samon [J]. J Fish Biol.,2003b, 62:753-766
    [52] Liu Y, Cao ZD, Fu SJ, Peng JL, Wang YX. The effect of exhaustive chasing training on swimming performance in juvenile darkbarber catefish (Peltebagrus vachelli) [J]. Comp Physiol B. 2009, 179:847-855.
    [53] Luo YP and Xie XJ. Effects of temperature on specific dynamic action of the southern catfish Silurus meridionalis [J].Comp Biochem Physiol A. 2008, 149:150-156
    [54] Milligan CL and Farrell AP. Lactate utilization by an in situ perfused trout heart: effects of workload and blockers of lactate transport [J]. J Exp Biol., 1991, 155: 357-373
    [55] Milligan CL and Girard SG. Lactate metabolism in rainbow trout [J]. J Exp Biol., 1993, 180: 175-193
    [56] Milligan CL. Metabolic recovery from exhaustive exercise in rainbow trout [J]. Comp Biochem Physiol A , 1996, 113:51-60
    [57] Milligan CL, Hooke GB and Johnson C. Sustained swimming at low velocity following a bout of exhaustive exercise enhances metabolic recovery in rainbow trout [J]. J Exp Biol., 2000, 203:921-926.
    [58] McDonald DG, McFarlane W J and Milligan C L.Anaerobic capacity and swim performance of juvenile salmonids[J]. Can. J.Fish. Aquat. Sci., 1998, 55: 1198-1207
    [59] Moyes CD, Schulte PM and West TG. Burst exercise recovery metabolism in fish white: the role of the mitochondria. Am. J. Physiol, 1993, 262: R295-R304
    [60] Niv Y and Fraser GM. Esophageal and gastric diseases: the alkaline tide phenomenon [J]. J Clin Gastroenterol., 2002, 35:5-8
    [61] Pagnotta A and Milligan CL. The role of blood glucose in the restoration of muscle glycogen during recovery from exhaustive exercise in rainbow trout (Oncorhynchus mykiss) and winter flounder (Pseudopleuronectes americanus) [J]. J Exp Biol., 1991, 161:489-508
    [62] Pagnotta A, Brooks L and Milligan L.The potential regulatory role of cortisol in the recovery from exhaustive exercise in rainbow trout [J]. Can J Zool. 1994, 72:2136-2146
    [63] Paton, KR., Cake, MH and Potter, IC. Muscle glycogen, lactate and glycerol-3-phosphate concentrations of larval and young adult lampreys in response to exercise [J].comp. Biochem .Physiol. B, 2004,129, 759-766
    [64] Pearson MP, Spriet LL and Stevens ED. Effect of sprint training on swim performance and white muscle metabolism during exercise and recovery in rainbow trout (Salmo gairdneri) [J]. J Exp Biol., 1990, 149:45-60
    [65] Rune SJ. The metabolic alkalosis following aspiration of gastric acid secretion. [J]. Scand J Clin Lab Invest, 1965, 17:305-310
    [66] Reidy SP, Nelson JA, Tang Y, Kerr SR. Postexercise metabolic rate in Atlantic cod and its dependence upon the method of exhaustion [J]. J Fish Biol, 1995, 47:377-386
    [67] Reidy SP, Kerr SR, Nelson JA. Aerobic and anaerobic swimming performance of individual Altantic cod.[J]. J Exp Biol., 2000, 203:347-357
    [68] Scarabello M, Heigenhauser GJ, Wood CM. Gas exchange, metabolite status, and excess post-exercise oxygen consumption after repetitive bouts of exhaustive exercise in juvenile rainbow trout [J]. J Exp Biol., 1992, 167:155-169
    [69] Schulte PM, Moyes CD and Hochachka PW. Integrating metabolic pathways in post-exercise recovery of white muscle [J]. J Exp Biol., 1992, 166:181-195
    [70] Schwalme K and Mackay WC. The influence of angling-induced exercise on the carbohydrate metabolism of northern pike (Esoxlucius L.) [J]. J. Comp. Physiol. B, 1985, 156:67-75
    [71] Secor SM and Diamond J. Determinants of the postfeeding metabolic response of Burmese pythons, Python molurus [J]. Physiol Zoology. 1997, 70:202-212
    [72] Secor SM, Hicks JW, Bennett AF. Ventilatory and cardiovascular responses of a python (Python molurus) to exercise and digestion [J]. J Exp Biol, 2000, 203:2447-2454
    [73] Secor S M. Regulation of digestive performance: a proposed adaptive response [J]. Comp Biochem Physiol A , 2001, 128:565-567
    [74] Secor S M and Faulker A C. Effect of meal size, meal size, meal type, body temperature, and body size on the specific dynamic action of the marine toad, Bufo marinus[J]. Physol Biochem Zool. 2002, 75:557-571.
    [75] Secor S M. Specific dynamic action: a review of the postprandial metabolic response [J]. J Comp Physiol B, 2009, 179:1–56
    [76] Stainsby WN and Barclay JK. Exercise metabolism: O2 deficit, steady level O2 uptake and O2 uptake for recovery [J], Med Sci Sports, 1970, 16:177-181
    [77] Taylor JR and Grosell M. Feeding and osmoregulation: dual function of the marine teleost intestine [J]. J Exp Biol, 2006, 209:2939-2951
    [78] Taylor JR, Whittamore JM, Wilson RW, Grosell M. Postprandial acid-base balance and ion regulation in freshwater and seawater-acclimated European flounder, Platichthys flesus. [J]. J Comp Physiol B, 2007, 177:597-608
    [79] Thorarensen H, Farrell AP. Post-prandial intestinal blood flow, metabolic rate, and exercise in Chinook salmon (Oncorhynchus tshawytscha) [J]. Physiol Biochem Zool. 2006, 74:688-694
    [80] Tufts B L, Currie S and Kieffer JD. Relative effects of carbonic anhydrase infusion or in-hibition on carbon dioxide transport and acid–base status in the sea lamprey Petromyzon marinus following exercise [J]. J Exp Biol., 1996, 199:933–940
    [81] Turner JD, Wood CM and Clark D Lactate and proton dynamics in the rainbow trout (Salmo gairdneri) [J]. J Exp Biol., 1983, 62:950-963
    [82] Vaziri ND, Byrne C, Ryan G and Wilson A. Preservation of urinary postprandial alkaline tide despite inhibition of gastric acid secretion [J]. Am, J.Gastroenterol.1980, 74:328-331
    [83] Wang Y, Heigenhauser GJ and Wood CM. Integrated responses to exhaustive exercise and recovery in rainbow trout white muscle: acid-base, phosphogen, carbohydrate, lipid, ammonia, fluid volume and electrolyte metabolism [J]. J Exp Biol, 1994, 192:299-305
    [84] Wakefield AM, Cunjak RA and Kieffer JD. Metabolic recovery in Atlantic salmon fry and parr following forced activity [J]. J. Fish Biol., 2004, 65: 920-932
    [85] Wilkie MP, Brobbel MA, Forsyth L, Davidson K and Tufts BL.The influence of temperature on the post-exercise physiology and survival of Atlantic salmon (Salmo salar) [J]. Can. J. Fish. Aquat. Sci., 1997, 54:503-511.
    [86] Wang T, Busk M, Overgaard J. The respiratory consequences of feeding in amphibians and reptiles [J]. Comp Biochem Physiol A, 2001, 128:535-549
    [87] Wood CM, Perry SF, Walsh PJ andThomas S. HCO3- dehydration by the blood of an elasmobranch in the absence of a Haldane effect [J]. Respir. Physiol. 1994, 98:319-337
    [88] Wood CM, Part P, Wright PA. Ammonia and urea metabolism in relation to gill function and acid-base balance in a marine elasmobranch , the spiny dogfish (Squalus acanthias) [J]. J Exp Biol., 1995, 198:1545-1558
    [89] Wood CM, Kajimura M, Mommsen TP, Walsh PJ. Alkaline tide and nitrogen conservation after feeding in an elasmobranch (Squalus acanthias) [J]. J Exp Biol., 2005, 208:2693-2705
    [90] Wood CM, Kajimura M, Bucking C, Walsh PJ. Osmoregulation, ionoregulation and acid-base regulation by the gastrointestinal tract after feeding in the elasmobranch (Squalus acanthias) [J]. J Exp Biol, 2007a, 210:1335-1349
    [91] Wood CM, Bucking C, Fitzpatrick J, Nadella SR. The alkaline tide goes out and nitrogen stays in after feeding in the dogfish shark, Squalus acanthias [J]. Respir Physiol Neurobiol, 2007b, 159:163-170
    [92] Wood CM and Milligan CL. Adrenergic analysis of extracelluar and intracellular lactate and H dynamics after strenuous exercise in the starry flounder (Platichthys stellatus) [J]. Physiol. Zool., 1987, 60:69-81
    [93] Xie XJ and Sun RY. The bioenergetics of southern catfish (Silurus meridionalis Chen): growth rate as a function ration level, body weight and temperature [J]. J Fish Biol., 1992, 40, 719-730

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