温度驯化对刺参（Apostichopus japonicus）生长及耐热性的影响及生理生态学机制
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摘要
本文通过一系列实验研究了温度驯化和应激对刺参生长以及其耐热性的影响，结果总结如下：
1.热历史对刺参耐热性的影响：致死温度上限和热休克蛋白70基因表达
本研究探讨了不同的热历史对刺参致死温度和热休克蛋白70基因表达的影响。设置两个温度梯度：12（CA）和22℃（WA），将刺参在两个温度下驯化30d后，然后在相同环境17℃下恢复一周。通过实验发现，在12和22℃下驯化的刺参的半数致死温度（ULTs50）分别为30.9℃（30.6~31.3℃）和31.8℃（31.5~32.1℃），致死温度分别为32℃和33℃。将两组刺参在30℃热激2h后于17℃恢复，测定hsp70的表达时序。在热激实验中，将经过驯化的刺参在不同温度（25、27、29、31和32℃）热激2h，然后在17℃下恢复2h。使用半定量RT-PCR的方法测定hsp70表达水平，比较两组刺参基因表达最高值的诱导温度。CA和WA组刺参的表达最高值分别出现在29℃和31℃。结果表明，热历史能改变刺参的耐热上限，这种改变与hsp70的表达模式有关。
2.驯化温度对刺参生长、代谢及耐热性的影响
本实验探讨了不同的驯化温度(16、21和26℃)对刺参（Apostichopusjaponicus）生长、代谢及耐热性的影响。实验持续40d，期间进行刺参生长、代谢、摄食以及能量收支的测定。结果表明，16℃下饲养的刺参体重（末体重11.96±0.35g）明显高于21℃（10.33±0.41g）和26℃（8.31±0.19g）处理组（P<0.05），并且16℃下刺参生长能明显高于其它两组（P<0.05）。在16、21和26℃下饲养的刺参临界热极值分别为33.1、34.1和36.6℃，并且26℃下刺参热休克蛋白基因表达量明显高于其它两组。这一结果表明：温度驯化能改变刺参的耐热性，并且能通过临界热极值和热休克蛋白基因表达这两种指标进行反映。
3.温度和盐度驯化对刺参生长及热休克蛋白基因表达的影响：
本实验研究了温度和盐度驯化对刺参（Apostichopusjaponicus）幼参生长对热休克蛋白基因表达的影响。设置4个温度梯度（16、20、24和28℃），3个盐度梯度（22、27和32），按双因子4×3的组合分为12个不同的温度盐度驯化组。经过40d的驯化，发现温度和盐度对刺参的体重和特定生长率影响显著，并且高温下的刺参对低盐的耐受性较低温下降低。不同组的刺参热休克蛋白基因表达量不同，说明经过高温驯化，刺参对盐度胁迫的适应性增强，应对高温及低盐环境的能力得到提高。
4.高温刺激对刺参浮游期幼体生长、存活率及Hsp70表达量的影响
设计不同温度（21.5、26、28和30℃），对刺参（Apostichopusjaponicus）囊胚期、原肠期和耳状幼体期幼体进行热激处理，热激45min后，将幼体在21.5℃条件下继续培养至耳状幼体后期，每日测量幼体的体长，计算刺参幼体的存活率，并用蛋白杂交方法测定幼体Hsp70的表达量。结果表明，高温刺激对幼体的存活影响显著，各时期幼体的存活率均随热激温度的升高而降低，原肠期的存活率高于囊胚期和耳状幼体时期。原肠期幼体Hsp70表达量比囊胚期和耳状幼体时期高，这表明在发育过程中Hsp70表达量与幼体存活率具有相关性。
5.浮游期热应激对刺参幼参生长及耐热性的影响
设计不同温度（21.5、26、28和30℃），对刺参（Apostichopusjaponicus）囊胚期、原肠期和耳状幼体期幼体进行热激处理，热激45min后，将幼体在21.5℃条件下继续培养至幼参。然后分别在17和24℃驯化60天，期间每10d称量体重。驯化结束后，实验刺参在17℃下恢复12天，从31.5℃的2h预热激中恢复4天后，再遭受2h的34℃热激，然后恢复7天，计算死亡率。结果表明，17℃驯化下的刺参体重大于24℃下的个体。同一驯化温度下，囊胚期热激的个体体重大于其它处理组。浮游期经过热激处理的幼参耐热性强于未经热激的个体。这表明在刺参幼体发育的浮游时期进行热激处理可能是一种选择较强耐热性个体的有效方法。
6.刺参南方育苗以及度夏的初步研究
本实验对刺参南方（福建）育苗以及幼参高温期培育进行了初步研究。通过夜间提水、提高水位、投喂多维等方法，对刺参度夏有一定的帮助。在福建省春季开展刺参育苗，翌年春天就可收获成品参，生长周期明显缩短。
1. Thermal resistance in sea cucumber (Apostichopus japonicus Selenka) with different thermal history:Upper thermal limit and hsp70gene expression
Thermal history can affect organisms'thermal resistance, however, effects of thermal history on thermal resistance and roles of heat shock protein70(Hsp70) in sea cucumber Apostichopus japonicus are not well understood. In the present study, the lethal thermal limits and expressions of Hsp70of sea cucumbers that were acclimated at different temperatures (22℃or12℃) for30days were studied. The upper lethal temperatures (ULT50) for sea cucumbers acclimated at22and12℃were31.8℃(31.5～32.1℃) and30.9℃(30.6～31.3℃), respectively. In heat shock experiments, sea cucumbers acclimated at different temperatures (22℃or12℃) were exposed to several high temperatures (25,27,29,31and32℃) for2h, followed by2h recovery at17℃, and the levels of hsp70were measured using semi-quantitative RT-PCR. The maximum induction temperatures for sea cucumbers in22℃and12℃acclimation treatments were31℃and29℃, respectively. These results showed that the temperature acclimation could affect sea cucumbers'upper thermal limit, which was related to the changed expression pattern of Hsp70in the sea cucumber. In practice, sea cucumbers that overwinter in southern China could acquire higher thermal resistance than those reared in the north.
2. Effects of rearing temperature on growth, metabolism and thermal tolerance of juvenile sea cucumber, Apostichopus japonicus Selenka:Critical thermal maximum and hsps gene expression
Effects of different rearing temperatures (16,21and26℃) on growth, metabolic performance and thermal tolerance of juvenile sea cucumber Apostichopus japonicus (initial body weight7.72±0.96g, mean±SD) were investigated in the present study. During the40-day experiment, growth, metabolic performance, food intake and energy budget at different reared temperatures were determined. Sea cucumbers rearing at16℃obtained better growth (final body weight11.96±0.35g) than those reared at21(10.33±0.41g) and26℃(8.31±0.19g)(P<0.05), and more energy was allocated for growth at16℃(162.73±11.85J g-1d-1) than those at21(79.61±6.76J g-1d-1) and26℃(27.07±4.30J g-1d-1)(P<0.05). Critical thermal maxima (CTmax) values of juvenile sea cucumbers reared at16,21and26℃were33.1,34.1and36.6℃, respectively. The upregulation of hsps in sea cucumbers reared at26℃was higher than those acclimated at lower temperatures (16and21℃), indicating that temperature acclimation could change the thermal tolerance of the sea cucumber, and CTmax and hsps were sensitive indicators of the sea cucumber's thermal tolerance.
3. Effects of temperature and salinity acclimation on growth and hsps gene expression of juvenile sea cucumber, Apostichopus japonicus Selenka
Effects of temperature and salinity acclimation on growth and hsps gene expression of juvenile sea cucumber were investigated in this study. There were12treatments (temperature:16,20,24,28℃; salinity:22,27,32psu), each treatment had3replicates. After40-d culture, the body weight and specific growth rate was different from each other. At high temperature, salinity tolerance of sea cucumber was lower than those rearing at ralatively low temperature. Also, hsps gene expression was different among treatments. Levels of hsps increased with temperature increasing. After acclimation at high temperature, the adaptation to salinity stress of sea cucumber was strengthen, and the ability to cope with high temperature and low salinity was improved.
4. Effects of heat-shock in pelagic stages on growth, survival and Hsp70expression of juvenile sea cucumber, Apostichopusjaponicus Selenka
The effects of heat-shock in pelagic stages on growth, survival and Hsp70s expression of juvenile sea cucumber, Apostichopus japonicus Selenka were investigated. Larvae at the stage of blastula, gastrula and auricularia were heat-shocked at selected temperatures (21.5, control;26,28and30℃) for45min and returned to21.5℃for continuous rearing. After the heat-shock, there were significant differences in survival among larvae in the four heat-shocked treatments. The survival rate of larvae decreased with the increasing of heat-shock temperature, and was higher in the stage of gastrula than those in blastula and auricularia. The Hsp70s expression of larvae in the stage of gastrula was higher than those in the stages of blastula and auricularia, suggesting that the survival of larvae partly correlates with the expression of Hsp70s.
5. Effects of heat-shock selection during pelagic stages on growth and thermal sensitivity of juvenile sea cucumber, Apostichopus japonicus Selenka
The effects of heat-shock selection during pelagic stages on growth and survival of juvenile sea cucumber Apostichopus japonicus were investigated to test the possibility to acquire high thermal tolerance individuals after the heat-shock. Larvae at the stages of blastula (BLA), gastrula (GAS) and auricularia (AUR) were heat-shocked at selected temperatures (21.5, control;26,28and30℃) for45min and returned to21.5℃for continuous rearing. There was significant difference in thermal sensitivity among different developmental stages. Juveniles after the heat-shock selection at pelagic stages showed higher induced thermotolerance than those without heat-shock. Therefore, heat-shock application at early pelagic development stages is potentially an effective way to select sea cucumbers with a high thermotolerance.
6. Preliminary study in seeding and summer spending of sea cucumber (Apostichopus japonicus Selenka) in Southern China
Seeding and summer spending of sea cucumber(Apostichopus japonicus Selenka) in Southern China was investigated in this study. There were some ways to help sea cucumbers to spend summer, such as pumping water at night, raising water level, feeding multiple vitamin. If we carry out seeding in the spring of Fujian province, we can harvest adult sea cucumbers next spring, so the growth cycle could be shorten obviously.
1.热历史对刺参耐热性的影响：致死温度上限和热休克蛋白70基因表达
本研究探讨了不同的热历史对刺参致死温度和热休克蛋白70基因表达的影响。设置两个温度梯度：12（CA）和22℃（WA），将刺参在两个温度下驯化30d后，然后在相同环境17℃下恢复一周。通过实验发现，在12和22℃下驯化的刺参的半数致死温度（ULTs50）分别为30.9℃（30.6~31.3℃）和31.8℃（31.5~32.1℃），致死温度分别为32℃和33℃。将两组刺参在30℃热激2h后于17℃恢复，测定hsp70的表达时序。在热激实验中，将经过驯化的刺参在不同温度（25、27、29、31和32℃）热激2h，然后在17℃下恢复2h。使用半定量RT-PCR的方法测定hsp70表达水平，比较两组刺参基因表达最高值的诱导温度。CA和WA组刺参的表达最高值分别出现在29℃和31℃。结果表明，热历史能改变刺参的耐热上限，这种改变与hsp70的表达模式有关。
2.驯化温度对刺参生长、代谢及耐热性的影响
本实验探讨了不同的驯化温度(16、21和26℃)对刺参（Apostichopusjaponicus）生长、代谢及耐热性的影响。实验持续40d，期间进行刺参生长、代谢、摄食以及能量收支的测定。结果表明，16℃下饲养的刺参体重（末体重11.96±0.35g）明显高于21℃（10.33±0.41g）和26℃（8.31±0.19g）处理组（P<0.05），并且16℃下刺参生长能明显高于其它两组（P<0.05）。在16、21和26℃下饲养的刺参临界热极值分别为33.1、34.1和36.6℃，并且26℃下刺参热休克蛋白基因表达量明显高于其它两组。这一结果表明：温度驯化能改变刺参的耐热性，并且能通过临界热极值和热休克蛋白基因表达这两种指标进行反映。
3.温度和盐度驯化对刺参生长及热休克蛋白基因表达的影响：
本实验研究了温度和盐度驯化对刺参（Apostichopusjaponicus）幼参生长对热休克蛋白基因表达的影响。设置4个温度梯度（16、20、24和28℃），3个盐度梯度（22、27和32），按双因子4×3的组合分为12个不同的温度盐度驯化组。经过40d的驯化，发现温度和盐度对刺参的体重和特定生长率影响显著，并且高温下的刺参对低盐的耐受性较低温下降低。不同组的刺参热休克蛋白基因表达量不同，说明经过高温驯化，刺参对盐度胁迫的适应性增强，应对高温及低盐环境的能力得到提高。
4.高温刺激对刺参浮游期幼体生长、存活率及Hsp70表达量的影响
设计不同温度（21.5、26、28和30℃），对刺参（Apostichopusjaponicus）囊胚期、原肠期和耳状幼体期幼体进行热激处理，热激45min后，将幼体在21.5℃条件下继续培养至耳状幼体后期，每日测量幼体的体长，计算刺参幼体的存活率，并用蛋白杂交方法测定幼体Hsp70的表达量。结果表明，高温刺激对幼体的存活影响显著，各时期幼体的存活率均随热激温度的升高而降低，原肠期的存活率高于囊胚期和耳状幼体时期。原肠期幼体Hsp70表达量比囊胚期和耳状幼体时期高，这表明在发育过程中Hsp70表达量与幼体存活率具有相关性。
5.浮游期热应激对刺参幼参生长及耐热性的影响
设计不同温度（21.5、26、28和30℃），对刺参（Apostichopusjaponicus）囊胚期、原肠期和耳状幼体期幼体进行热激处理，热激45min后，将幼体在21.5℃条件下继续培养至幼参。然后分别在17和24℃驯化60天，期间每10d称量体重。驯化结束后，实验刺参在17℃下恢复12天，从31.5℃的2h预热激中恢复4天后，再遭受2h的34℃热激，然后恢复7天，计算死亡率。结果表明，17℃驯化下的刺参体重大于24℃下的个体。同一驯化温度下，囊胚期热激的个体体重大于其它处理组。浮游期经过热激处理的幼参耐热性强于未经热激的个体。这表明在刺参幼体发育的浮游时期进行热激处理可能是一种选择较强耐热性个体的有效方法。
6.刺参南方育苗以及度夏的初步研究
本实验对刺参南方（福建）育苗以及幼参高温期培育进行了初步研究。通过夜间提水、提高水位、投喂多维等方法，对刺参度夏有一定的帮助。在福建省春季开展刺参育苗，翌年春天就可收获成品参，生长周期明显缩短。
1. Thermal resistance in sea cucumber (Apostichopus japonicus Selenka) with different thermal history:Upper thermal limit and hsp70gene expression
Thermal history can affect organisms'thermal resistance, however, effects of thermal history on thermal resistance and roles of heat shock protein70(Hsp70) in sea cucumber Apostichopus japonicus are not well understood. In the present study, the lethal thermal limits and expressions of Hsp70of sea cucumbers that were acclimated at different temperatures (22℃or12℃) for30days were studied. The upper lethal temperatures (ULT50) for sea cucumbers acclimated at22and12℃were31.8℃(31.5～32.1℃) and30.9℃(30.6～31.3℃), respectively. In heat shock experiments, sea cucumbers acclimated at different temperatures (22℃or12℃) were exposed to several high temperatures (25,27,29,31and32℃) for2h, followed by2h recovery at17℃, and the levels of hsp70were measured using semi-quantitative RT-PCR. The maximum induction temperatures for sea cucumbers in22℃and12℃acclimation treatments were31℃and29℃, respectively. These results showed that the temperature acclimation could affect sea cucumbers'upper thermal limit, which was related to the changed expression pattern of Hsp70in the sea cucumber. In practice, sea cucumbers that overwinter in southern China could acquire higher thermal resistance than those reared in the north.
2. Effects of rearing temperature on growth, metabolism and thermal tolerance of juvenile sea cucumber, Apostichopus japonicus Selenka:Critical thermal maximum and hsps gene expression
Effects of different rearing temperatures (16,21and26℃) on growth, metabolic performance and thermal tolerance of juvenile sea cucumber Apostichopus japonicus (initial body weight7.72±0.96g, mean±SD) were investigated in the present study. During the40-day experiment, growth, metabolic performance, food intake and energy budget at different reared temperatures were determined. Sea cucumbers rearing at16℃obtained better growth (final body weight11.96±0.35g) than those reared at21(10.33±0.41g) and26℃(8.31±0.19g)(P<0.05), and more energy was allocated for growth at16℃(162.73±11.85J g-1d-1) than those at21(79.61±6.76J g-1d-1) and26℃(27.07±4.30J g-1d-1)(P<0.05). Critical thermal maxima (CTmax) values of juvenile sea cucumbers reared at16,21and26℃were33.1,34.1and36.6℃, respectively. The upregulation of hsps in sea cucumbers reared at26℃was higher than those acclimated at lower temperatures (16and21℃), indicating that temperature acclimation could change the thermal tolerance of the sea cucumber, and CTmax and hsps were sensitive indicators of the sea cucumber's thermal tolerance.
3. Effects of temperature and salinity acclimation on growth and hsps gene expression of juvenile sea cucumber, Apostichopus japonicus Selenka
Effects of temperature and salinity acclimation on growth and hsps gene expression of juvenile sea cucumber were investigated in this study. There were12treatments (temperature:16,20,24,28℃; salinity:22,27,32psu), each treatment had3replicates. After40-d culture, the body weight and specific growth rate was different from each other. At high temperature, salinity tolerance of sea cucumber was lower than those rearing at ralatively low temperature. Also, hsps gene expression was different among treatments. Levels of hsps increased with temperature increasing. After acclimation at high temperature, the adaptation to salinity stress of sea cucumber was strengthen, and the ability to cope with high temperature and low salinity was improved.
4. Effects of heat-shock in pelagic stages on growth, survival and Hsp70expression of juvenile sea cucumber, Apostichopusjaponicus Selenka
The effects of heat-shock in pelagic stages on growth, survival and Hsp70s expression of juvenile sea cucumber, Apostichopus japonicus Selenka were investigated. Larvae at the stage of blastula, gastrula and auricularia were heat-shocked at selected temperatures (21.5, control;26,28and30℃) for45min and returned to21.5℃for continuous rearing. After the heat-shock, there were significant differences in survival among larvae in the four heat-shocked treatments. The survival rate of larvae decreased with the increasing of heat-shock temperature, and was higher in the stage of gastrula than those in blastula and auricularia. The Hsp70s expression of larvae in the stage of gastrula was higher than those in the stages of blastula and auricularia, suggesting that the survival of larvae partly correlates with the expression of Hsp70s.
5. Effects of heat-shock selection during pelagic stages on growth and thermal sensitivity of juvenile sea cucumber, Apostichopus japonicus Selenka
The effects of heat-shock selection during pelagic stages on growth and survival of juvenile sea cucumber Apostichopus japonicus were investigated to test the possibility to acquire high thermal tolerance individuals after the heat-shock. Larvae at the stages of blastula (BLA), gastrula (GAS) and auricularia (AUR) were heat-shocked at selected temperatures (21.5, control;26,28and30℃) for45min and returned to21.5℃for continuous rearing. There was significant difference in thermal sensitivity among different developmental stages. Juveniles after the heat-shock selection at pelagic stages showed higher induced thermotolerance than those without heat-shock. Therefore, heat-shock application at early pelagic development stages is potentially an effective way to select sea cucumbers with a high thermotolerance.
6. Preliminary study in seeding and summer spending of sea cucumber (Apostichopus japonicus Selenka) in Southern China
Seeding and summer spending of sea cucumber(Apostichopus japonicus Selenka) in Southern China was investigated in this study. There were some ways to help sea cucumbers to spend summer, such as pumping water at night, raising water level, feeding multiple vitamin. If we carry out seeding in the spring of Fujian province, we can harvest adult sea cucumbers next spring, so the growth cycle could be shorten obviously.
引文
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