运动训练对中华倒刺鲃幼鱼生长的影响及其机理研究
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摘要
适当流速下的持续游泳运动通常能够促进鱼类生长率的提高。本研究以我国长江中上游重要的经济鱼类中华倒刺鲤(Spinibarbus sinensis)幼鱼为实验对象,在25±0.5℃水温条件下,进行了五个系列的实验。实验一考察力竭追赶训练(1次/d,历时21d)对中华倒刺鲤幼鱼生长和摄食代谢特征的影响；实验二考察不同水流速度下[3cm/s,1b1/s(体长/秒),2bl/s和4bl/s]的有氧运动训练(20h/d,历时56d)对中华倒刺鳃幼鱼生长和摄食代谢特征的影响；实验三研究有氧运动训练对中华倒刺鲃幼鱼肠道排空率和消化酶活性的影响；实验四研究有氧运动训练对中华倒刺鲤幼鱼肌纤维结构和肌肉质构特性的影响；实验五研究有氧运动训练对中华倒刺鲤幼鱼肌肉一般营养成分、氨基酸和脂肪酸含量的影响。通过以上系列实验,确定中华倒刺鲃幼鱼最适生长流速范围,探讨运动训练对其生长产生影响的内在机理,进一步揭示鱼类摄食代谢的调节机制,丰富鱼类生理生态学理论,为流水养殖和增殖放流等实践提供重要的基础资料。
主要研究结果：
(1)力竭训练组中华倒刺鲃幼鱼摄食率、饲料效率、特定体重生长率和特定体长生长率都显著小于对照组(P<0.05)；力竭训练组中华倒刺鲤幼鱼的静止代谢率、摄食代谢峰值时间和SDA(特殊动力作用,Specific dynamic action)时间与对照组无显著性差异,但摄食代谢峰值、峰值比率、SDA总耗能和SDA系数显著低于对照组(P<0.05)。
(2)1bl/s和2b1/s训练组中华倒刺鳃幼鱼生长率显著高于对照组(P<0.05)；中华倒刺鳃幼鱼摄食率随着水流速度的增加而显著提高(P<0.05)；1bl/s和2b1/s训练组饲料效率与对照组无显著性差异,但4b1/s训练组显著低于其它三个实验组(P<0.05)；所有训练组中华倒刺鲤幼鱼静止代谢和摄食代谢峰值都显著高于对照组(P<0.05),其峰值时间显著小于对照组(P<0.05),但其SDA时间、峰值比率、SDA总耗能和SDA系数与对照组无显著性差异。
(3)所有实验组中华倒刺鲃幼鱼肠道排空均呈现先快后慢的变化特征,其最优数学模型均为平方根模型；1b1/s和2b1/s训练组中华倒刺鲃幼鱼的肠道排空率显著快于对照组,肠道排空时间显著小于对照组(P<0.05)；但4bl/s训练组肠道排空率和肠道排空时间与对照组无显著性差异；所有训练组中华倒刺鳃幼鱼胰蛋白酶和脂肪酶活性显著高于对照组(P<0.05)。
(4)1b1/s和2b1/s训练组的红肌纤维直径和密度与对照组无显著性差异,但4b1/s训练组的红肌直径显著小于对照组,密度显著大于对照组(P<0.05)；1b1/s和2b1/s训练组的白肌纤维直径和密度与对照组无显著性差异,但4b1/s训练组的白肌直径显著小于对照组,密度显著大于对照组(P<0.05)；随着水流速度的增加,中华倒刺鲃幼鱼肌肉各质构参数值呈现增大趋势(P<0.05),但各个训练组肌肉颜色参数值与对照组无显著性差异；各个训练组肌肉pH值都显著高于对照组(P<0.05)。
(5)有氧运动训练对中华倒刺鲃幼鱼肌肉水分和灰分含量没有显著性影响,但2b1/s训练组肌肉蛋白质含量显著高于对照组(P<0.05),4b1/s训练组肌肉脂肪含量显著低于1b1/s和2bl/s训练组(P<0.05)；1bl/s和2b1/s训练组肌肉氨基酸总含量和必须氨基酸总含量显著高于对照组,4b1/s训练组肌肉氨基酸总含量显著低于对照组(P<0.05)；1b1/s和2b1/s训练组肌肉脂肪酸含量与对照组无显著性差异,但4bl/s训练组肌肉n-6系列多不饱和脂肪酸含量显著低于对照组(P<0.05)。
主要研究结论：
(1)摄食率和饲料效率降低是力竭训练导致中华倒刺鳃幼鱼生长率下降的主要原因。
(2)适当流速(1-2b1/s)有氧运动训练通过刺激中华倒刺鳃幼鱼的食欲达到提高生长率的目的,其最适生长流速约为2.4b1/s；高流速(4b1/s)有氧运动训练虽然促进其摄食率提高,但游泳耗能增加可能是其生长率没有显著变化的主要原因。
(3)在不同运动训练制度下,中华倒刺鲤幼鱼采用的摄食代谢适应策略不同；中华倒刺鲃幼鱼采用维持静止代谢和降低摄食代谢峰值的策略适应力竭训练环境,通过提高静止代谢和摄食代谢峰值的策略适应有氧运动训练环境。
(4)有氧运动训练并没有对中华倒刺鲃幼鱼的肠道排空特征(先快后慢)产生显著影响,平方根模型是其肠道排空的最优数学拟合模型。
(5)适当流速(1-2b1/s)有氧运动训练导致中华倒刺鲃幼鱼肠道排空率加快、肠道排空时间缩短和消化酶活性提高可能是促进其生长的重要原因。
(6)适当流速(1-2b1/s)有氧运动训练导致中华倒刺鲃幼鱼肌肉蛋白质含量明显提高,表明在有氧运动训练条件下肌肉蛋白质合成率的增加可能是其生长率显著提高的原因之一。
(7)适当流速(1-2b1/s)有氧运动训练不仅有利于中华倒刺鲤幼鱼肌肉物理品质的提高,而且还能够改善其肌肉营养成分。因此,有氧运动训练可能是改善中华倒刺鲃肌肉品质的有效途径之一。
Continual swimming exercise usually promotes growth in fish at a moderate water velocity. Five series of the experiment were conducted in juvenile Spinibarbus sinensis, one of the commercial fish species in the middle and upper reaches of the Yangtze River of China, at25±0.5℃, In experiment Ⅰ, the growth performance and postprandial metabolic response were examined after exhaustive chasing training once daily for21days in juvenile S. sinensis. In experiment Ⅱ, the growth performance and postprandial metabolic response were examined after aerobic exercise training20hours daily for56days at a water velocities of control (3cm/s),1,2and4body length (bl)/s in juvenile S. sinensis. In experiment Ⅲ, the effects of aerobic exercise training on the gut evacuation characteristics and digestive enzyme activities were examined in juvenile S. sinensis. In experiment Ⅳ, the effects of aerobic exercise training on the muscle structure and texture properties were investigated in juvenile S. sinensis. In experiment V, the effects of aerobic exercise training on the nutritional components were assessed in juvenile S. sinensis. The objectives of this study are to (1) determine the optimal water velocity for growth in juvenile S. sinensis,(2) investigate the mechanism of exercise training on growth performance in juvenile S. sinensis,(3) reveal the underlying regulation mechanism of fish feeding metabolism,(4) enrich related theory about the physiological ecology of fish and (5) provide practical information for application of running water fish culture and fishery enhancement and release. The main results of present study as follows:
(1) Exhaustive chasing training resulted in significant decrease in feed intake, feed efficiency, weight specific growth rate, length specific growth rate, the peak metabolic rates, the energy expended on specific dynamic action (SDA) and SDA coefficients compared with those of control group in juvenile S. sinensis (P<0.05). Resting metabolic rates, times of peak metabolic rate and durations of SDA were not significantly different between control and trained groups in juvenile S. sinensis.
(2) Aerobic exercise training induced a significant increase in feed intake compared with the control group, while the feed efficiency of the4bl/s group was significantly lower than that of the other three groups in juvenile S. sinensis (P<0.05). The1and2bl/s groups showed a significantly higher weight specific growth rate over the control group in juvenile S. sinensis (P<0.05). The resting metabolic rates and peak metabolic rates in the three training groups were significantly higher than that in the control group (P<0.05) in juvenile S. sinensis. Times of peak metabolic rate was significantly shorter in the1,2and4bl/s training groups compared with the control group, while exercise training showed no effect on SDA duration, factorial metabolic scope, energy expended on SDA and the SDA coefficient when compared to the control group in juvenile S. sinensis.
(3) As postprandial time increased, the gut evacuation showed a fast-to-slow change characteristic in all experimental groups. The best mathematical model fitted to gut evacuation data in juvenile S. sinensis was the square-root model. Gut evacuation rates of the1and2bl/s training groups were significantly higher than that of the control group, while gut evacuation times of the1and2bl/s training groups were significantly lower than that of the control group after aerobic exercise training (P<0.05). However, gut evacuation rate and duration were not significantly different between control and4bl/s trained group in juvenile S. sinensis. Aerobic exercise training elevated the activity of both trypsin and lipase in the hepatopancreas and intestinal tract of juvenile S. sinensis (P<0.05).
(4) There were no significant differences in the diameter and density of red muscle fibre for the1and2bl/s training groups and those of control group. However, the diameter of red muscle fibre was significantly smaller in the4bl/s training groups as compared to the control group,while the density of red muscle fibre was significantly larger in the4bl/s training groups as compared to the control group (P<0.05). The change trend for the diameter and density of white muscle fibre was similar as that of red muscle fibre after aerobic exercise training in juvenile S. sinensis. The hardness, springiness, adhesiveness, chewiness, cohesivenss and resilience of the muscle exhibited an increase trend with the increase of water velocity. However, there was not significant difference in the colour of muscle between control and trained groups in juvenile S. sinensis. The pHs of muscle in three training groups were significantly higher than that of control group (P<0.05).
(5) No significant differences in the moisture and ash contents were found among the different groups. The percentage of protein in the2bl/s training groups was significantly higher than the control group (P<0.05). However, the percentage of fat in the4bl/s training group was significantly lower than those of1and2bl/s training groups (P<0.05). The contents of total amino acid and essential amino acid in1and2bl/s training groups were significantly higher than respective values of control group (P<0.05). The content of total amino acid in4bl/s training group was significantly lower than that of control group (P<0.05). The contents of fatty acid in1and2bl/s training groups were not significantly different as compared to that of control group. However, The content of total n-6fatty acids in4bl/s training group was significantly lower than that of control group (P<0.05). The main conclusion as follows:
(1) The lower growth rate might be induced by poor feed intake and feed efficiency after exhaustive chasing training for21d in juvenile S. sinensis.
(2) The improvement of growth may be primarily due to an increase in the feed intake after long-term training in juvenile S. sinensis. The optimum water velocity for the growth of juvenile S. sinensis occurred at approximately2.4bl/s. The juvenile S. sinensis in the4bl/s training group might have allocated more energy from their increased food consumption to sustain swimming rather than growth.
(3) The juvenile S. sinensis adopt different feeding metabolic adaptation strategies under the different regimes of exercise training. The maintaining of resting metabolic rate and reduction of peak metabolic rate have been found after exhaustive training in juvenile S. sinensis. The up-regulated resting metabolic rate and peak metabolic rate were induced by aerobic exercise training in juvenile S. sinensis.
(4) There were no significant changes in the gut evacuation characteristic and best mathematical model under condition of aerobic exercise training in juvenile S. sinensis.
(5) Aerobic exercise training boosted the activity of digestive enzymes and gut evacuation rate, which could favor fast digestion and growth at a moderate water velocity in juvenile S. sinensis.
(6) Aerobic exercise training resulted in significant increase in muscle protein content at a moderate water velocity in juvenile S. sinensis. This finding may have resulted in an improvement of growth because it is possible that the increased growth rates resulted from the proportionately greater increase in the protein synthesis rate in trained fish.
(7) Aerobic exercise training may be one of effective ways to improve flesh quality in S. sinensis due prolonged swimming at moderate water velocity could improve texture properties and nutritional components of muscle in juvenile S. sinensis.
主要研究结果：
(1)力竭训练组中华倒刺鲃幼鱼摄食率、饲料效率、特定体重生长率和特定体长生长率都显著小于对照组(P<0.05)；力竭训练组中华倒刺鲤幼鱼的静止代谢率、摄食代谢峰值时间和SDA(特殊动力作用,Specific dynamic action)时间与对照组无显著性差异,但摄食代谢峰值、峰值比率、SDA总耗能和SDA系数显著低于对照组(P<0.05)。
(2)1bl/s和2b1/s训练组中华倒刺鳃幼鱼生长率显著高于对照组(P<0.05)；中华倒刺鳃幼鱼摄食率随着水流速度的增加而显著提高(P<0.05)；1bl/s和2b1/s训练组饲料效率与对照组无显著性差异,但4b1/s训练组显著低于其它三个实验组(P<0.05)；所有训练组中华倒刺鲤幼鱼静止代谢和摄食代谢峰值都显著高于对照组(P<0.05),其峰值时间显著小于对照组(P<0.05),但其SDA时间、峰值比率、SDA总耗能和SDA系数与对照组无显著性差异。
(3)所有实验组中华倒刺鲃幼鱼肠道排空均呈现先快后慢的变化特征,其最优数学模型均为平方根模型；1b1/s和2b1/s训练组中华倒刺鲃幼鱼的肠道排空率显著快于对照组,肠道排空时间显著小于对照组(P<0.05)；但4bl/s训练组肠道排空率和肠道排空时间与对照组无显著性差异；所有训练组中华倒刺鳃幼鱼胰蛋白酶和脂肪酶活性显著高于对照组(P<0.05)。
(4)1b1/s和2b1/s训练组的红肌纤维直径和密度与对照组无显著性差异,但4b1/s训练组的红肌直径显著小于对照组,密度显著大于对照组(P<0.05)；1b1/s和2b1/s训练组的白肌纤维直径和密度与对照组无显著性差异,但4b1/s训练组的白肌直径显著小于对照组,密度显著大于对照组(P<0.05)；随着水流速度的增加,中华倒刺鲃幼鱼肌肉各质构参数值呈现增大趋势(P<0.05),但各个训练组肌肉颜色参数值与对照组无显著性差异；各个训练组肌肉pH值都显著高于对照组(P<0.05)。
(5)有氧运动训练对中华倒刺鲃幼鱼肌肉水分和灰分含量没有显著性影响,但2b1/s训练组肌肉蛋白质含量显著高于对照组(P<0.05),4b1/s训练组肌肉脂肪含量显著低于1b1/s和2bl/s训练组(P<0.05)；1bl/s和2b1/s训练组肌肉氨基酸总含量和必须氨基酸总含量显著高于对照组,4b1/s训练组肌肉氨基酸总含量显著低于对照组(P<0.05)；1b1/s和2b1/s训练组肌肉脂肪酸含量与对照组无显著性差异,但4bl/s训练组肌肉n-6系列多不饱和脂肪酸含量显著低于对照组(P<0.05)。
主要研究结论：
(1)摄食率和饲料效率降低是力竭训练导致中华倒刺鳃幼鱼生长率下降的主要原因。
(2)适当流速(1-2b1/s)有氧运动训练通过刺激中华倒刺鳃幼鱼的食欲达到提高生长率的目的,其最适生长流速约为2.4b1/s；高流速(4b1/s)有氧运动训练虽然促进其摄食率提高,但游泳耗能增加可能是其生长率没有显著变化的主要原因。
(3)在不同运动训练制度下,中华倒刺鲤幼鱼采用的摄食代谢适应策略不同；中华倒刺鲃幼鱼采用维持静止代谢和降低摄食代谢峰值的策略适应力竭训练环境,通过提高静止代谢和摄食代谢峰值的策略适应有氧运动训练环境。
(4)有氧运动训练并没有对中华倒刺鲃幼鱼的肠道排空特征(先快后慢)产生显著影响,平方根模型是其肠道排空的最优数学拟合模型。
(5)适当流速(1-2b1/s)有氧运动训练导致中华倒刺鲃幼鱼肠道排空率加快、肠道排空时间缩短和消化酶活性提高可能是促进其生长的重要原因。
(6)适当流速(1-2b1/s)有氧运动训练导致中华倒刺鲃幼鱼肌肉蛋白质含量明显提高,表明在有氧运动训练条件下肌肉蛋白质合成率的增加可能是其生长率显著提高的原因之一。
(7)适当流速(1-2b1/s)有氧运动训练不仅有利于中华倒刺鲤幼鱼肌肉物理品质的提高,而且还能够改善其肌肉营养成分。因此,有氧运动训练可能是改善中华倒刺鲃肌肉品质的有效途径之一。
Continual swimming exercise usually promotes growth in fish at a moderate water velocity. Five series of the experiment were conducted in juvenile Spinibarbus sinensis, one of the commercial fish species in the middle and upper reaches of the Yangtze River of China, at25±0.5℃, In experiment Ⅰ, the growth performance and postprandial metabolic response were examined after exhaustive chasing training once daily for21days in juvenile S. sinensis. In experiment Ⅱ, the growth performance and postprandial metabolic response were examined after aerobic exercise training20hours daily for56days at a water velocities of control (3cm/s),1,2and4body length (bl)/s in juvenile S. sinensis. In experiment Ⅲ, the effects of aerobic exercise training on the gut evacuation characteristics and digestive enzyme activities were examined in juvenile S. sinensis. In experiment Ⅳ, the effects of aerobic exercise training on the muscle structure and texture properties were investigated in juvenile S. sinensis. In experiment V, the effects of aerobic exercise training on the nutritional components were assessed in juvenile S. sinensis. The objectives of this study are to (1) determine the optimal water velocity for growth in juvenile S. sinensis,(2) investigate the mechanism of exercise training on growth performance in juvenile S. sinensis,(3) reveal the underlying regulation mechanism of fish feeding metabolism,(4) enrich related theory about the physiological ecology of fish and (5) provide practical information for application of running water fish culture and fishery enhancement and release. The main results of present study as follows:
(1) Exhaustive chasing training resulted in significant decrease in feed intake, feed efficiency, weight specific growth rate, length specific growth rate, the peak metabolic rates, the energy expended on specific dynamic action (SDA) and SDA coefficients compared with those of control group in juvenile S. sinensis (P<0.05). Resting metabolic rates, times of peak metabolic rate and durations of SDA were not significantly different between control and trained groups in juvenile S. sinensis.
(2) Aerobic exercise training induced a significant increase in feed intake compared with the control group, while the feed efficiency of the4bl/s group was significantly lower than that of the other three groups in juvenile S. sinensis (P<0.05). The1and2bl/s groups showed a significantly higher weight specific growth rate over the control group in juvenile S. sinensis (P<0.05). The resting metabolic rates and peak metabolic rates in the three training groups were significantly higher than that in the control group (P<0.05) in juvenile S. sinensis. Times of peak metabolic rate was significantly shorter in the1,2and4bl/s training groups compared with the control group, while exercise training showed no effect on SDA duration, factorial metabolic scope, energy expended on SDA and the SDA coefficient when compared to the control group in juvenile S. sinensis.
(3) As postprandial time increased, the gut evacuation showed a fast-to-slow change characteristic in all experimental groups. The best mathematical model fitted to gut evacuation data in juvenile S. sinensis was the square-root model. Gut evacuation rates of the1and2bl/s training groups were significantly higher than that of the control group, while gut evacuation times of the1and2bl/s training groups were significantly lower than that of the control group after aerobic exercise training (P<0.05). However, gut evacuation rate and duration were not significantly different between control and4bl/s trained group in juvenile S. sinensis. Aerobic exercise training elevated the activity of both trypsin and lipase in the hepatopancreas and intestinal tract of juvenile S. sinensis (P<0.05).
(4) There were no significant differences in the diameter and density of red muscle fibre for the1and2bl/s training groups and those of control group. However, the diameter of red muscle fibre was significantly smaller in the4bl/s training groups as compared to the control group,while the density of red muscle fibre was significantly larger in the4bl/s training groups as compared to the control group (P<0.05). The change trend for the diameter and density of white muscle fibre was similar as that of red muscle fibre after aerobic exercise training in juvenile S. sinensis. The hardness, springiness, adhesiveness, chewiness, cohesivenss and resilience of the muscle exhibited an increase trend with the increase of water velocity. However, there was not significant difference in the colour of muscle between control and trained groups in juvenile S. sinensis. The pHs of muscle in three training groups were significantly higher than that of control group (P<0.05).
(5) No significant differences in the moisture and ash contents were found among the different groups. The percentage of protein in the2bl/s training groups was significantly higher than the control group (P<0.05). However, the percentage of fat in the4bl/s training group was significantly lower than those of1and2bl/s training groups (P<0.05). The contents of total amino acid and essential amino acid in1and2bl/s training groups were significantly higher than respective values of control group (P<0.05). The content of total amino acid in4bl/s training group was significantly lower than that of control group (P<0.05). The contents of fatty acid in1and2bl/s training groups were not significantly different as compared to that of control group. However, The content of total n-6fatty acids in4bl/s training group was significantly lower than that of control group (P<0.05). The main conclusion as follows:
(1) The lower growth rate might be induced by poor feed intake and feed efficiency after exhaustive chasing training for21d in juvenile S. sinensis.
(2) The improvement of growth may be primarily due to an increase in the feed intake after long-term training in juvenile S. sinensis. The optimum water velocity for the growth of juvenile S. sinensis occurred at approximately2.4bl/s. The juvenile S. sinensis in the4bl/s training group might have allocated more energy from their increased food consumption to sustain swimming rather than growth.
(3) The juvenile S. sinensis adopt different feeding metabolic adaptation strategies under the different regimes of exercise training. The maintaining of resting metabolic rate and reduction of peak metabolic rate have been found after exhaustive training in juvenile S. sinensis. The up-regulated resting metabolic rate and peak metabolic rate were induced by aerobic exercise training in juvenile S. sinensis.
(4) There were no significant changes in the gut evacuation characteristic and best mathematical model under condition of aerobic exercise training in juvenile S. sinensis.
(5) Aerobic exercise training boosted the activity of digestive enzymes and gut evacuation rate, which could favor fast digestion and growth at a moderate water velocity in juvenile S. sinensis.
(6) Aerobic exercise training resulted in significant increase in muscle protein content at a moderate water velocity in juvenile S. sinensis. This finding may have resulted in an improvement of growth because it is possible that the increased growth rates resulted from the proportionately greater increase in the protein synthesis rate in trained fish.
(7) Aerobic exercise training may be one of effective ways to improve flesh quality in S. sinensis due prolonged swimming at moderate water velocity could improve texture properties and nutritional components of muscle in juvenile S. sinensis.
引文
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