光照和温度波动对凡纳滨对虾（Litopenaeus vannamei）蜕皮和生长的影响及机制的初步研究
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摘要
本研究以凡纳滨对虾为实验材料,通过一系列实验室养殖实验,初步研究了光照波动对凡纳滨对虾蜕皮、生长的影响,并对光照和温度波动对凡纳滨对虾生长影响的机制进行了初步探讨。主要结果如下：
一、光照波动对凡纳滨对虾蜕皮和生长的影响及机制
1、光照强度日节律波动对凡纳滨对虾蜕皮和生长的影响及机制
本文在实验室条件下,研究了光照强度日节律波动对凡纳滨对虾蜕皮和生长的影响。实验设2700±600lx、2700±1200 lx、2700±1800 lx和2700±2400 lx四个不同的光照强度日节律性波动组,以2700 lx恒定光照强度为对照组,测定了不同处理组对虾的蜕皮、生长、能量收支及体组成。实验对虾的初始湿体重为(2.736±0.002)g,养殖周期为45天。主要实验结果如下：(1)光照强度日节律性波动组2700±600 lx、2700±1200 lx和2700±1800 lx对虾的相对增重率和特定生长率均显著高于恒定光照组(P<0.05)；而光照强度日节律性波动组2700±2400lx对虾虽然相对增重率显著高于恒定光照组(P<0.05),但其特定生长率与恒定光照组没有显著性差异(P>0.05)。(2)光照强度日节律性波动组2700±600 lx、2700±1200lx和2700±1800 lx对虾的摄食率显著高于恒定光照组(P<0.05),但2700±2400lx组对虾的摄食率与恒定光照组没有显著性差异(P>0.05)；在食物转化效率方面,除了2700±600 lx组对虾的食物转化效率显著低于恒定光照组外(P<0.05),其他各光照变化组对虾的食物转化效率与恒定光照组均无显著性差异(P>0.05)。(3)光照强度日节律性变化组2700±1200 lx和2700±1800 lx对虾用于生长的能量比例均显著高于恒定光照组(P<0.05),而其用于呼吸的能量比例显著低于恒定光照组(P<0.05)。(4)对虾的蜕皮率及用于蜕皮的能量比例在恒定光照组和四个光照强度日节律性波动组间均没有显著性差异(P>0.05)。(5)光照强度日节律性波动对凡纳滨对虾的体组成有显著的影响；与恒定光照相比,光照强度的波动显著降低了对虾体内粗脂肪的含量。
2、光照强度周期变化对凡纳滨对虾蜕皮和生长的影响及机制
在实验室条件下研究了光照强度周期性变化对凡纳滨对虾蜕皮和生长的影响。实验设计了60lx恒定光照组和600-60 lx、1500-60lx、3000-60 lx、6000-60 lx四个不同的光照强度周期性变化组。实验虾的初始湿体重为(2.733±0.017)g,养殖周期为48天。主要实验结果如下：(1)光照强度变化组1500-60 lx和3000-60 lx对虾的相对增重率和特定生长率均显著高于恒定光照组(P<0.05)；光照强度变化组600-60 lx对虾的相对增重率显著高于恒定光照组(P<0.05),但其特定生长率与恒定光照组没有显著性差异(P>0.05)；光照强度变化组6000-60 lx对虾的相对增重率与恒定光照组没有显著性差异(P>0.05),但其特定生长率却显著高于恒定光照组(P<0.05)。(2)各个处理组之间对虾的摄食率没有显著性差异(P>0.05),但光照强度变化1500-60 lx、3000-60 lx和6000-60 lx组对虾的食物转化效率显著高于恒定光照组(P<0.05)。(3)光照强度变化组1500-60 lx、3000-60 lx和6000-60 lx对虾的呼吸能和排泄能分配比例均显著低于恒定光照组(P<0.05),而用于生长的能量分配比例均显著高于恒定光照组(P<0.05)。(4)对虾蜕皮率和用于蜕皮的能量比例以光照强度变化组6000-60 lx最低,显著低于其他各处理组(P<0.05),但光照强度变化组1500-60 lx和3000-60 lx与恒定光照组间没有显著性差异(P>0.05)。(5)实验结束后对虾的灰分含量基本上与对虾的末干体重成正比,光照强度周期性变化提高了对虾粗脂肪的含量。
3、光色日节律性波动对凡纳滨对虾蜕皮和生长的影响及机制
本文在实验室条件下,研究了光色的日节律性变化对凡纳滨对虾蜕皮和生长的影响。实验设计了黄光(Y)、绿光(G)、蓝光(B)三个恒定光照组,蓝光变黄光(BY)、蓝光变绿光(BG)、绿光变黄光(GY)三个光色变化组。实验虾的初始湿体重为(1.212±0.010)g,养殖周期为45天。主要实验结果如下：(1)BG组和BY组对虾无论相对增重率还是特定生长率均显著高于蓝光组(P<0.05),GY组对虾的相对增重率显著高于蓝光组(P<0.05),但其特定生长率与蓝光组未出现显著性差异(P>0.05)。(2)对虾的摄食率在五个处理组之间未出现显著性差异(P>0.05),GY组对虾的食物转化效率与蓝光组也未出现显著性差异(P>0.05),但BG组和BY组对虾的食物转化效率却显著高于蓝光组(P<0.05)。(3)虽然对虾用于呼吸的能量分配比例没有显著性差异(P>0.05),但BG组和BY组对虾用于排泄的能量比例显著低于蓝光组(P<0.05),而用于生长的能量比例均显著高于蓝光组(P<0.05)。(4)光色的节律性变化显著提高了对虾用于蜕皮的能量,对虾的蜕皮率显著高于恒定光照组(P<0.05)。(5)实验结束后对虾的体组成中灰分含量基本上与对虾的末干体重成正比,粗蛋白没有明显的相关性,脂肪含量与水分含量基本呈负相关。
4、光色周期性变化对凡纳滨对虾蜕皮和生长的影响及机制
采用实验生态学方法研究了光色的周期性变化对凡纳滨对虾稚虾蜕皮和生长的影响。实验设计了黄光(Y)、绿光(G)、蓝光(B)三个恒定光照组,蓝光变黄光(BY)、蓝光变绿光(BG)、绿光变黄光(GY)三个光色变化组。实验虾的初始湿体重为(1.212±0.011)g,养殖周期为45天。主要结果如下：(1)BG组对虾的相对增重率和特定生长率均为最高,与Y组、B组和GY组均有显著性差异(P<0.05)。GY组对虾仅相对增重率显著高于Y组和B组(P<0.05),而其特定生长率与Y组和B组没有显著性差异(P>0.05)。(2)对虾的摄食率在五个处理组之间未出现显著性差异(P>0.05),GY组对虾的食物转化效率与Y组和B组也未出现显著性差异(P>0.05),但BG组和BY组对虾的食物转化效率却显著高于Y组和B组(P<0.05)。(3)BG组对虾用于呼吸的能量比例虽然与其他组在统计学上没有差异,但其比生长最差的B组低了大约4%；因而BG组对虾用于生长的能量比例显著高于Y组和B组(P<0.05)。(4)对虾的蜕皮率以GY组最高,各组对虾的蜕皮率与生长之间未出现明显的相关性。(5)实验结束后对虾体组成中灰分含量基本上与对虾的末干体重成正比,粗蛋白没有明显的相关性；光色的周期性波动降低了对虾体组成中粗脂肪的含量。
二、光照波动对凡纳滨对虾生长影响机制的初步研究
1、光照强度及周期性变化对凡纳滨对虾乳酸含量、丙酮酸激酶和乳酸脱氢酶活力的影响
实验室条件下研究了光照强度及周期性变化对凡纳滨对虾乳酸含量、丙酮酸激酶和乳酸脱氢酶活力的影响。实验设600-60 lx、1500-60 lx、3000-60 lx和6000-60 lx四个光照强度周期性突变组及60 lx、600 lx、1500 lx、3000 lx和6000lx五个恒定光照组。光强突变组采用先高光强照射6天后再低光强照射2天的方式进行周期性波动,光照周期为14L:10D。所有处理组对虾经过16天的驯化后开始实验。恒定光照组取样的时间点分别为：光照0 h、0.5 h、1 h、3 h、6 h、9 h、12 h、14 h；光强突变组取样的时间点为：一个光照周期开始后的0.5 h、1 h、3 h、6 h、12 h、24 h、48 h、72 h、144 h、144.5 h(即光强改变后0.5 h)、145 h、147 h、150 h、156 h、168 h、192 h。每个取样点取5尾蜕皮间期的对虾,分别测定对虾肝胰脏中丙酮酸激酶(PK)活力、肌肉中乳酸脱氢酶活(LDH)活力和乳酸(LD)含量。主要实验结果如下：(1)光照强度周期性突变整体上降低了对虾肝胰脏中PK和LDH的活力,而对虾肌肉中的LD含量没有明显变化,这可能与光照强度周期波动提高了对虾机体有关酶的功效有关。(2)、恒定600 lx组和光强突变1500-60 lx组对虾的各种指标变化幅度较小,而6000 lx组和光照突变600-60 lx组对虾的各种指标波动幅度较大。(3)6000-60 lx组对虾肌肉中乳酸含量在高光照阶段一直维持在一个较高的水平,而600-60 lx组对虾在高光照后期和低光照开始阶段的肌肉中乳酸含量很高,说明这两种光照强度的波动给凡纳滨对虾造成了胁迫影响。(4)本实验中高光强向低光强波动时,对虾肌肉中LDH活力的波动比低光强向高光强波动时大,这可能与对虾对不同光照强度的适应不同以及光强波动周期中高光强照射时间较长有关。结果表明适宜的光照强度周期性波动使对虾机体内酶的功效提升,且其体内酶的波动幅度较小,有利于对虾的生长。
2、光色及其日节律性变化对凡纳滨对虾葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的影响
本文在实验室条件下研究了光色及其日节律性变化对凡纳滨对虾葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的影响。实验设蓝光向绿光日节律变化组、恒定蓝光和绿光共三个光色处理组,分别在实验开始的第1、3、5、10、20和40天,于每天的7：00、9：00、13：00、14：00、16：00、20：00从各处理组中挑选蜕皮间期的对虾5尾用于葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的测定。主要实验结果如下：(1)开灯刺激引起各组对虾葡萄糖含量有小幅度提升,但对对虾乳酸含量没有显著的影响；蓝光和绿光组对虾肝胰脏中己糖激酶活力在开灯后有所下降,光色变化组则有所增加,而对虾丙酮酸激酶活力不受开灯刺激的影响。(2)在实验开始阶段,光色的日节律性变化对对虾血清中的葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力都有一定的影响,随着实验进行,这种影响在20天内逐步消失。(3)蓝光组对虾肝胰脏中己糖激酶和丙酮酸激酶活力在一天内波动的幅度较大,且这种波动趋于稳定的时间较长；而绿光组对虾肝胰脏中己糖激酶和丙酮酸激酶活力在一天内相对平稳。
三、温度波动对凡纳滨对虾生长影响机制的初步研究
1、温度周期性波动对凡纳滨对虾生长、糖酵解酶及HSP70的影响
本实验以水温25℃为恒温对照,设计4个不同幅度的变温组,分别为25℃±1℃、25℃±2℃、25℃±3℃、25℃±4℃,研究了五种不同温度处理下凡纳滨对虾生长、摄食率及饵料转化率；并在实验结束时取样测定了对虾血清中的葡萄糖含量、肝胰脏中的己糖激酶(HK)和丙酮酸激酶(PK)及肌肉组织中的热休克蛋白70的相对含量。结果表明：在平均温度为25℃时,变温幅度为±2℃和±3℃组对虾的特定生长率均高于恒温对照组,且以±2℃组最高；而变温幅度为±4℃组对虾的特定生长率显著低于其他各处理组。±4℃组摄食率最低,且其食物转化效率显著低于生长较好的±2℃和±3℃组。±2℃和±3℃组对虾的摄食率与其他处理组没有显著性差异,但对虾的食物转化效率显著高于其他组。由此可以推断：在平均温度为25℃时,适宜的变温幅度促进对虾生长的原因是提高了对虾的食物转化效率；而±4℃组对虾的摄食率和食物转化效率均低,这是导致对虾生长不良的原因。相关生理指标分析发现,±4℃组对虾血液中葡萄糖含量最低而丙酮酸激酶活力最高,说明对虾一直处于环境的胁迫下,需通过糖酵解代谢产生更多的能量用于抵抗不良的环境,影响了对虾用于生长的能量。±3℃组对虾血液中葡萄糖含量最高而已糖激酶的活力最低,表明机体没有出现对胁迫的响应机制,温度的波动并没有对凡纳滨对虾的生长产生不良的影响,对虾丙酮酸激酶的活力适中也进一步说明了此问题。相对于恒温组,各变温组对虾热休克蛋白没有显著性增高,可能与±4℃的温度波动幅度并没有诱导对虾热休克蛋白的高效表达有关。
2、温度周期性波动过程中凡纳滨对虾糖酵解酶和热休克蛋白70的变化
本实验以水温25℃为恒温对照,设计4个不同幅度的变温组,分别为25℃±1℃、25℃±2℃、25℃±3℃、25℃±4℃,研究了五种不同温度处理下凡纳滨血清中的葡萄糖含量、肝胰脏中的已糖激酶(HK)和丙酮酸激酶(PK)及肌肉组织中的热休克蛋白70的相对含量在一天内的变化。结果表明：在恒温组中葡萄糖含量和热休克蛋白70均存在昼夜变化规律,但在温度波动组中这种规律随着波动幅度的增大逐渐消失。恒温组的PK活力变化幅度不大,随着温度波动幅度的增大,其变化逐渐剧烈,以25℃±4℃组尤为明显。HK活力受到葡萄糖流量和糖酵解作用的影响,随着温度变化幅度的增加,其变化也有所趋于平稳,但没有葡萄糖明显。
A series of indoor experiments were conducted to investigate the effects of light fluctuations on the molting and growth and of Litopenaeus vannamei. Based on the results and the resrarches of temperature flucyuations, the mechanism for the different characters of L. vannamei under fluctuating light and temperature treatments was discussed. The main results were presented as the followings:
1. The effects of light fluctuations on the molting, growth of Litopenaeus vannamei and preliminary researches on its mechanisms
1.1 The effect of rhythmic light intensity fluctuation on the molting and growth of Litopenaeus vannamei and its mechanisms
The method of experimental ecology was used to investigate the effect of rhythmic light intensity fluctuation on the molting and growth of Litopenaeus vannamei. Molting, growth performance, feeding and body composition of the shrimps were tested in one constant light intensity (2700 lx) treatment and four rhythmic fluctuating light intensity treatments (2700±600 lx; 2700±1200 lx; 2700±1800 lx; 2700±2400 lx). The initial wet body weight of shrimp was 2.736±0.002 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) of shrimps in 2700±600 lx,2700±1200 lx and 2700±1800 lx treatments were significantly higher than those in 2700 lx treatment (P< 0.05); the WG in 2700±2400 lx treatment was significantly higher than those in 2700 lx treatment (P<0.05), but SGRd was not (P>0.05). (2) The feed intake (FId) in 2700 ±600 lx,2700±1200 lx and 2700±1800 lx treatments were significantly higher than those in 2700 lx treatment (P<0.05), but 2700±2400 lx treatment was not (P>0.05). The food conversion efficiency (FCEd) in 2700±600 lx treatment was significantly lower than those in 2700 lx treatment (P<0.05), but the other three rhythmic fluctuating light intensity treatments were not (P>0.05). (3) The energy allocation to growth in 2700±1200 lx and 2700±1800 lx were significantly higher than those in 2700 lx treatment (P<0.05) and the energy allocation in respiration in 2700±1200 lx and 2700±1800 lx were significantly lower than those in 2700 lx treatment (P<0.05). (4) There was no significantly difference of molting rate (MF) and energy allocation in exuviate between constant light intensity and four rhythmic fluctuating light intensity treatments (P>0.05). (5) The body composition of the shrimps was significantly affected by the different light treatments (P<0.05). Compared to the constant light intensity treatment, the value of crude lipid decreased because of the effect of the fluctuating light intensity.
1.2 The effect of periodic abrupt change in light intensity on the molting and growth of Litopenaeus vannamei and its mechanisms
The molting and growth of Litopenaeus vannamei were tested in one constant light intensity (60 lx) treatment and four periodic abrupt changing light intensity treatments (600-60 lx; 1500-60 lx; 3000-60 lx; 6000-60 lx). The initial body weight of shrimp was 2.733±0.017 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) in 1500-60 lx and 3000-60 lx treatments were significant higher than those in 60 lx treatment (P<0.05); the WG in 600-60 lx treatment was significantly higher than those in 60 lx treatment (P<0.05), but the SGRd was not (P>0.05). Opposite to what was observed in the 600-60 lx treatment, the WG in 6000-60 lx had no difference from those in 60 lx treatment (P>0.05), but the SGRd in 6000-60 lx treatment was significantly higher than those in 60 lx treatment (P<0.05). (2) There is no significant difference in FId for all regimes (P>0.05), but the FId of shrimps in 1500 -60 lx,3000-60 lx and 6000-60 lx treatments were significant higher than those in 60 lx treatment (P<0.05). (3) The energy allocation in respiration and excretion of shrimps in 1500-60 lx,3000-60 lx and 6000-60 lx treatments were significantly lower than those in 60 lx treatment (P<0.05), the energy allocation in the growth were significantly higher than those in 60 lx treatment (P<0.05). (4) The MF and energy allocation in exuviate in 6000-60 lx treatment were lowest, and significantly lower than those in other treatments (P<0.05), but the MF and energy allocation in exuviate in 1500-60 lx or 3000-60 lx treatment was not significantly different from those in 60 lx treatment (P>0.05). (5) The ash had a positive correlation with the body weight, but the crude protein did not, and the abrupt changing light enhanced the content of crude lipid of shrimps.
1.3 The effect of rhythmic light color fluctuation on the growth and molting of Litopenaeus vannamei and its mechanisms
The molting and growth of Litopenaeus vannamei were tested under two kinds of color treatments:three constant light color treatments (yellow light, Y; green light, G; blue light, B) and three rhythmic fluctuating light color treatments (blue light was changed to yellow light, BY; blue light was changed to green light, BG; green light was changed to yellow light, GY). The initial body weight of shrimp was 1.212±0.010 g, the period of this experiment was 45 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) of shrimp in BG and BY treatments were significantly higher than those in B treatment (P>0.05). The WG in GY treatment was significantly higher than those in B treatment (P>0.05), but the SGRd was not (P>0.05). (2) There is no significant difference in FId for all regimes (P>0.05), the FCEd of shrimps in GY treatment was no significantly different from those in B treatment (P>0.05), but the FCEd of shrimps in BY and BG treatments were significantly higher than those in B treatment (P<0.05). (3) However, the energy allocation in respiration of shrimps in all treatment did not have significant difference (P >0.05), but the energy allocation to excretion of shrimps in BG and BY treatments were significantly lower than those in B treatment (P<0.05), the energy allocation to growth were significantly higher than those in B treatment (P<0.05). (4) The molting rate (MF) and energy allocation in exuviate of shrimps in three rhythmic fluctuating light color treatments were generally higher than that in three constant light color treatments (P<0.05), there was a clear correlation between MF and growth in every regime. (5) The ash of shrimps had a positive correlation with the body weight, but the crude protein did not, and the abrupt changing light enhanced the content of crude lipid of shrimps. The lipid level of shrimps increased with the decrease of moisture level.
1.4 The effect of periodic light color fluctuation on the molting and growth of Litopenaeus vannamei and its mechanisms
The method of experimental ecology was used to investigate the effect of periodic light color fluctuation on the molting and growth of Litopenaeus vannamei. Molting and growth performance of the shrimps were tested under two kinds of color treatments: three constant light color treatments (yellow light, Y; green light, G; blue light, B) and three periodic fluctuating light color treatments (blue light was changed to yellow light, BY; blue light was changed to green light, BG; green light was changed to yellow light, GY). The initial body weight of shrimp was 1.212±0.018 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) the weight gain (WG) and the specific growth rate (SGRd) of shrimp in BG treatments were highest and significantly higher than those in Y or B treatment (P>0.05). The WG in GY treatment was significantly higher than those in Y or B treatment (P>0.05), but the SGRd was not (P>0.05). (2) There is no significant difference in FId for all regimes (P>0.05), the FCEd of shrimps in GY treatment was no significantly different from those in Y or B treatment (P>0.05), but the FCEd of shrimps in BY and BG treatments were significantly higher than those in Y or B treatment (P<0.05). (3) Although there was no significant difference in energy allocation in respiration under different light color regimes (P>0.05); but the energy allocation in respiration in BG treatment was lower about 4% than those in B treatment, so the energy allocation to growth of shrimps in BG treatment was significantly higher than those in Y or B treatment (P<0.05). (4) The molting rate in GY treatment was highest, but there was no correlation between MF and growth. (5) The ash had a positive correlation with the body weight, but the crude protein did not, and the periodic light color fluctuation decreased the content of crude lipid of shrimps.
2. Effects and mechanism of light fluctuations on growth of Litopenaeus vannamei
2.1 The variation of LD, PK and LDH of Litopenaeus vannamei in periodic abrupt change in light intensity and constant light intensity
Four periodic fluctuation light intensity treatments (600-60 lx,1500-60 lx,3000-60 lx and 6000-60 lx) and five constant light intensity treatments (60 lx,600 lx,1500 lx,3000 lx,6000 lx) were arranged to investigate the effect of periodic abrupt change in light intensity and constant light intensity on the enzymes relating to respiratory metabolism. An alternation of six days high light intensity and two days low light intensity was adopted in the periodic fluctuation light intensity treatment. After two cycles of light intensity periodic fluctuation, sampling five inter-molt stage shrimps in the five constant light intensity treatments to assay the PK activities, the LDH activities and the lactic-acid at 0 h,0.5 h,1 h,3 h,6 h,9 h,12 h,14 h after the light on, respectively; and doing that in four periodic fluctuation light intensity treatments at 0.5 h,1 h,3 h,6 h,12 h,24 h,48 h,72 h,144 h,144.5 h,145 h,147 h,150 h,156 h,168 h, 192 h of the third cycle of light intensity periodic fluctuation, respectively. The main results were as follow:(1) A decrease of total PK and LDH activities was found in periodic fluctuation light intensity treatments, but LD been not; the reason for the phenomenon might be that the efficacy of some enzymes was enhanced in periodic fluctuation light intensity treatments. (2) The fluctuations of enzymes in 600 lx and 1500-60 lx treatments were stable, but those in 6000 lx and 6000-60 lx treatments were acute. (3) The lactic-acid content of 6000-60 lx treatment in the phase of high light was high and this phenomenon was also found in the later stages of high light and the beginning of low light in 600-60 lx. These results might indicate that the shrimp may be in a bad environment. (4) The LDH fluctuation when the light changed from high intensity to low intensity was more acute than those when the light changed from low intensity to high intensity. This may due to the difference types of shrimps acclimate to different light intensities and the longer time of high light intensity in our experiment. The results suggested that the suitable periodic abrupt changing light intensity enhanced the effectiveness of the enzyme in shrimps and its fluctuation of enzyme was stable, so the growth of shrimps was better.
2.2 The variations of GLU, LD, HK and PKof Litopenaeus vannamei under a rhythmic fluctuating color light and two constant color lights
A rhythmic fluctuating light color treatment (Blue light was changed to green light, BG) and two constant light color treatments (green light, G; blue light, B) were selected from the last experiment to investigate the effect of rhythmic fluctuations in light color on the enzymes relating to respiratory metabolism. Sampling five inter-molt stage shrimps to assay the hemolymph glucose, lactic-acid, the HK and PK activities of hepatopancreas at 7:00,9:00,13:00,14:00,16:00,20:00 on the following days:the first day, third day, fifths day, tenth day, twentieth day and fortieth day of the experiment, respectively. The main results were as follow:(1) The hemolymph glucose content had a small increase after the light on, but the hemolymph lactic-acid content did not. The HK activities of shrimps in B and G treatments decreased after switching on the light, but those in BG treatment increased. And the PK activities in all treatment did not have an obvious change. (2) A significant effect of rhythmic fluctuating light color on glucose, lactic-acid, HK and PK activities was found in the beginning of the experiment, but this effect disappear gradually when the experiment went on. (3) The fluctuations of HK and PK activities of shrimps in B treatment were acute, and the time of HK and PK activities became stable was late; but those in G treatment was comparatively stable.
3. Effects and mechanism of temperature fluctuations on growth of Litopenaeus vannamei
3.1 The effects of cyclical temperature changes on growth and physiological status of Litopenaeus vannamei
The purpose of this study was to investigate the growth and physiological status of Litopenaeus vannamei subjected to one constant temperature (25℃) and four cyclical temperature change regimes (25℃±1℃,25℃±2℃,25℃±3℃and 25℃±4℃). The growth rates of shrimp at 25℃±2℃or 25℃±3℃were significantly higher than that at a constant temperature of 25℃. On the other hand, the growth rate in 25℃±4℃regime was significantly lower than those in other regimes. The daily feed intake rate of shrimp at 25℃±4℃was the lowest and the food conversion efficiency was also significantly lower than those at 25℃±2℃and 25℃±3℃, respectively. The food conversion efficiency at 25℃±2℃or 25℃±3℃was significantly higher than those in other regimes. Thus, it can be inferred that the growth enhancement in the test shrimp at the suitable diel fluctuating temperatures was due to high food conversion efficiency. Studies of the physiological parameters showed that at 25℃±4℃, the hemolymph glucose content of the test shrimp was the lowest, while the activity of PK in hepatopancreas was the highest, which indicated that the test shrimp at 25℃±4℃was in a stressed condition. The hemolymph glucose content of the test shrimp at 25℃±3℃was the highest and the activity of HK in hepatopancreas was the lowest. These results indicated that the test shrimps at 25℃±3℃were not in a stressed condition. Compared with the constant temperature regime, the expression of HSP70 in any of the four cyclical temperature change regimes was not significantly increased. The reason for this might be that the fluctuation amplitude of±4℃did not induce the increased expression of HSP70.
3.2. The variations of enzymes involved glycolysis and HSP70 of Litopenaeus vannamei during the temperature cyclical change
The rhythmic fluctuations of hemolymph glucose content, HK and PK activities in hepatopancreas and HSP70 expression in Litopenaeus vannamei were measured under one constant temperature (25℃) and four cyclical temperature change regimes (25℃±1℃,25℃±2℃,25℃±3℃and 25℃±4℃). The main results were as follows: a day/night rhythmicity was found in the rhythmic fluctuations of hemolymph glucose content and HSP70 expression under 25℃, but the rhythmicity disappeared gradually with the increased fluctuation amplitude of temperature. The PK activities varied gently under 25℃. Due to the increase of temperature fluctuation amplitude, the fluctuation of PK activities became acute, especially in 25℃±4℃regime. Because of the effect of glucose flux and glycolysis, the fluctuation of HK activity became stable with the increased fluctuation amplitude of temperature.
一、光照波动对凡纳滨对虾蜕皮和生长的影响及机制
1、光照强度日节律波动对凡纳滨对虾蜕皮和生长的影响及机制
本文在实验室条件下,研究了光照强度日节律波动对凡纳滨对虾蜕皮和生长的影响。实验设2700±600lx、2700±1200 lx、2700±1800 lx和2700±2400 lx四个不同的光照强度日节律性波动组,以2700 lx恒定光照强度为对照组,测定了不同处理组对虾的蜕皮、生长、能量收支及体组成。实验对虾的初始湿体重为(2.736±0.002)g,养殖周期为45天。主要实验结果如下：(1)光照强度日节律性波动组2700±600 lx、2700±1200 lx和2700±1800 lx对虾的相对增重率和特定生长率均显著高于恒定光照组(P<0.05)；而光照强度日节律性波动组2700±2400lx对虾虽然相对增重率显著高于恒定光照组(P<0.05),但其特定生长率与恒定光照组没有显著性差异(P>0.05)。(2)光照强度日节律性波动组2700±600 lx、2700±1200lx和2700±1800 lx对虾的摄食率显著高于恒定光照组(P<0.05),但2700±2400lx组对虾的摄食率与恒定光照组没有显著性差异(P>0.05)；在食物转化效率方面,除了2700±600 lx组对虾的食物转化效率显著低于恒定光照组外(P<0.05),其他各光照变化组对虾的食物转化效率与恒定光照组均无显著性差异(P>0.05)。(3)光照强度日节律性变化组2700±1200 lx和2700±1800 lx对虾用于生长的能量比例均显著高于恒定光照组(P<0.05),而其用于呼吸的能量比例显著低于恒定光照组(P<0.05)。(4)对虾的蜕皮率及用于蜕皮的能量比例在恒定光照组和四个光照强度日节律性波动组间均没有显著性差异(P>0.05)。(5)光照强度日节律性波动对凡纳滨对虾的体组成有显著的影响；与恒定光照相比,光照强度的波动显著降低了对虾体内粗脂肪的含量。
2、光照强度周期变化对凡纳滨对虾蜕皮和生长的影响及机制
在实验室条件下研究了光照强度周期性变化对凡纳滨对虾蜕皮和生长的影响。实验设计了60lx恒定光照组和600-60 lx、1500-60lx、3000-60 lx、6000-60 lx四个不同的光照强度周期性变化组。实验虾的初始湿体重为(2.733±0.017)g,养殖周期为48天。主要实验结果如下：(1)光照强度变化组1500-60 lx和3000-60 lx对虾的相对增重率和特定生长率均显著高于恒定光照组(P<0.05)；光照强度变化组600-60 lx对虾的相对增重率显著高于恒定光照组(P<0.05),但其特定生长率与恒定光照组没有显著性差异(P>0.05)；光照强度变化组6000-60 lx对虾的相对增重率与恒定光照组没有显著性差异(P>0.05),但其特定生长率却显著高于恒定光照组(P<0.05)。(2)各个处理组之间对虾的摄食率没有显著性差异(P>0.05),但光照强度变化1500-60 lx、3000-60 lx和6000-60 lx组对虾的食物转化效率显著高于恒定光照组(P<0.05)。(3)光照强度变化组1500-60 lx、3000-60 lx和6000-60 lx对虾的呼吸能和排泄能分配比例均显著低于恒定光照组(P<0.05),而用于生长的能量分配比例均显著高于恒定光照组(P<0.05)。(4)对虾蜕皮率和用于蜕皮的能量比例以光照强度变化组6000-60 lx最低,显著低于其他各处理组(P<0.05),但光照强度变化组1500-60 lx和3000-60 lx与恒定光照组间没有显著性差异(P>0.05)。(5)实验结束后对虾的灰分含量基本上与对虾的末干体重成正比,光照强度周期性变化提高了对虾粗脂肪的含量。
3、光色日节律性波动对凡纳滨对虾蜕皮和生长的影响及机制
本文在实验室条件下,研究了光色的日节律性变化对凡纳滨对虾蜕皮和生长的影响。实验设计了黄光(Y)、绿光(G)、蓝光(B)三个恒定光照组,蓝光变黄光(BY)、蓝光变绿光(BG)、绿光变黄光(GY)三个光色变化组。实验虾的初始湿体重为(1.212±0.010)g,养殖周期为45天。主要实验结果如下：(1)BG组和BY组对虾无论相对增重率还是特定生长率均显著高于蓝光组(P<0.05),GY组对虾的相对增重率显著高于蓝光组(P<0.05),但其特定生长率与蓝光组未出现显著性差异(P>0.05)。(2)对虾的摄食率在五个处理组之间未出现显著性差异(P>0.05),GY组对虾的食物转化效率与蓝光组也未出现显著性差异(P>0.05),但BG组和BY组对虾的食物转化效率却显著高于蓝光组(P<0.05)。(3)虽然对虾用于呼吸的能量分配比例没有显著性差异(P>0.05),但BG组和BY组对虾用于排泄的能量比例显著低于蓝光组(P<0.05),而用于生长的能量比例均显著高于蓝光组(P<0.05)。(4)光色的节律性变化显著提高了对虾用于蜕皮的能量,对虾的蜕皮率显著高于恒定光照组(P<0.05)。(5)实验结束后对虾的体组成中灰分含量基本上与对虾的末干体重成正比,粗蛋白没有明显的相关性,脂肪含量与水分含量基本呈负相关。
4、光色周期性变化对凡纳滨对虾蜕皮和生长的影响及机制
采用实验生态学方法研究了光色的周期性变化对凡纳滨对虾稚虾蜕皮和生长的影响。实验设计了黄光(Y)、绿光(G)、蓝光(B)三个恒定光照组,蓝光变黄光(BY)、蓝光变绿光(BG)、绿光变黄光(GY)三个光色变化组。实验虾的初始湿体重为(1.212±0.011)g,养殖周期为45天。主要结果如下：(1)BG组对虾的相对增重率和特定生长率均为最高,与Y组、B组和GY组均有显著性差异(P<0.05)。GY组对虾仅相对增重率显著高于Y组和B组(P<0.05),而其特定生长率与Y组和B组没有显著性差异(P>0.05)。(2)对虾的摄食率在五个处理组之间未出现显著性差异(P>0.05),GY组对虾的食物转化效率与Y组和B组也未出现显著性差异(P>0.05),但BG组和BY组对虾的食物转化效率却显著高于Y组和B组(P<0.05)。(3)BG组对虾用于呼吸的能量比例虽然与其他组在统计学上没有差异,但其比生长最差的B组低了大约4%；因而BG组对虾用于生长的能量比例显著高于Y组和B组(P<0.05)。(4)对虾的蜕皮率以GY组最高,各组对虾的蜕皮率与生长之间未出现明显的相关性。(5)实验结束后对虾体组成中灰分含量基本上与对虾的末干体重成正比,粗蛋白没有明显的相关性；光色的周期性波动降低了对虾体组成中粗脂肪的含量。
二、光照波动对凡纳滨对虾生长影响机制的初步研究
1、光照强度及周期性变化对凡纳滨对虾乳酸含量、丙酮酸激酶和乳酸脱氢酶活力的影响
实验室条件下研究了光照强度及周期性变化对凡纳滨对虾乳酸含量、丙酮酸激酶和乳酸脱氢酶活力的影响。实验设600-60 lx、1500-60 lx、3000-60 lx和6000-60 lx四个光照强度周期性突变组及60 lx、600 lx、1500 lx、3000 lx和6000lx五个恒定光照组。光强突变组采用先高光强照射6天后再低光强照射2天的方式进行周期性波动,光照周期为14L:10D。所有处理组对虾经过16天的驯化后开始实验。恒定光照组取样的时间点分别为：光照0 h、0.5 h、1 h、3 h、6 h、9 h、12 h、14 h；光强突变组取样的时间点为：一个光照周期开始后的0.5 h、1 h、3 h、6 h、12 h、24 h、48 h、72 h、144 h、144.5 h(即光强改变后0.5 h)、145 h、147 h、150 h、156 h、168 h、192 h。每个取样点取5尾蜕皮间期的对虾,分别测定对虾肝胰脏中丙酮酸激酶(PK)活力、肌肉中乳酸脱氢酶活(LDH)活力和乳酸(LD)含量。主要实验结果如下：(1)光照强度周期性突变整体上降低了对虾肝胰脏中PK和LDH的活力,而对虾肌肉中的LD含量没有明显变化,这可能与光照强度周期波动提高了对虾机体有关酶的功效有关。(2)、恒定600 lx组和光强突变1500-60 lx组对虾的各种指标变化幅度较小,而6000 lx组和光照突变600-60 lx组对虾的各种指标波动幅度较大。(3)6000-60 lx组对虾肌肉中乳酸含量在高光照阶段一直维持在一个较高的水平,而600-60 lx组对虾在高光照后期和低光照开始阶段的肌肉中乳酸含量很高,说明这两种光照强度的波动给凡纳滨对虾造成了胁迫影响。(4)本实验中高光强向低光强波动时,对虾肌肉中LDH活力的波动比低光强向高光强波动时大,这可能与对虾对不同光照强度的适应不同以及光强波动周期中高光强照射时间较长有关。结果表明适宜的光照强度周期性波动使对虾机体内酶的功效提升,且其体内酶的波动幅度较小,有利于对虾的生长。
2、光色及其日节律性变化对凡纳滨对虾葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的影响
本文在实验室条件下研究了光色及其日节律性变化对凡纳滨对虾葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的影响。实验设蓝光向绿光日节律变化组、恒定蓝光和绿光共三个光色处理组,分别在实验开始的第1、3、5、10、20和40天,于每天的7：00、9：00、13：00、14：00、16：00、20：00从各处理组中挑选蜕皮间期的对虾5尾用于葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力的测定。主要实验结果如下：(1)开灯刺激引起各组对虾葡萄糖含量有小幅度提升,但对对虾乳酸含量没有显著的影响；蓝光和绿光组对虾肝胰脏中己糖激酶活力在开灯后有所下降,光色变化组则有所增加,而对虾丙酮酸激酶活力不受开灯刺激的影响。(2)在实验开始阶段,光色的日节律性变化对对虾血清中的葡萄糖、乳酸含量及肝胰脏中己糖激酶和丙酮酸激酶活力都有一定的影响,随着实验进行,这种影响在20天内逐步消失。(3)蓝光组对虾肝胰脏中己糖激酶和丙酮酸激酶活力在一天内波动的幅度较大,且这种波动趋于稳定的时间较长；而绿光组对虾肝胰脏中己糖激酶和丙酮酸激酶活力在一天内相对平稳。
三、温度波动对凡纳滨对虾生长影响机制的初步研究
1、温度周期性波动对凡纳滨对虾生长、糖酵解酶及HSP70的影响
本实验以水温25℃为恒温对照,设计4个不同幅度的变温组,分别为25℃±1℃、25℃±2℃、25℃±3℃、25℃±4℃,研究了五种不同温度处理下凡纳滨对虾生长、摄食率及饵料转化率；并在实验结束时取样测定了对虾血清中的葡萄糖含量、肝胰脏中的己糖激酶(HK)和丙酮酸激酶(PK)及肌肉组织中的热休克蛋白70的相对含量。结果表明：在平均温度为25℃时,变温幅度为±2℃和±3℃组对虾的特定生长率均高于恒温对照组,且以±2℃组最高；而变温幅度为±4℃组对虾的特定生长率显著低于其他各处理组。±4℃组摄食率最低,且其食物转化效率显著低于生长较好的±2℃和±3℃组。±2℃和±3℃组对虾的摄食率与其他处理组没有显著性差异,但对虾的食物转化效率显著高于其他组。由此可以推断：在平均温度为25℃时,适宜的变温幅度促进对虾生长的原因是提高了对虾的食物转化效率；而±4℃组对虾的摄食率和食物转化效率均低,这是导致对虾生长不良的原因。相关生理指标分析发现,±4℃组对虾血液中葡萄糖含量最低而丙酮酸激酶活力最高,说明对虾一直处于环境的胁迫下,需通过糖酵解代谢产生更多的能量用于抵抗不良的环境,影响了对虾用于生长的能量。±3℃组对虾血液中葡萄糖含量最高而已糖激酶的活力最低,表明机体没有出现对胁迫的响应机制,温度的波动并没有对凡纳滨对虾的生长产生不良的影响,对虾丙酮酸激酶的活力适中也进一步说明了此问题。相对于恒温组,各变温组对虾热休克蛋白没有显著性增高,可能与±4℃的温度波动幅度并没有诱导对虾热休克蛋白的高效表达有关。
2、温度周期性波动过程中凡纳滨对虾糖酵解酶和热休克蛋白70的变化
本实验以水温25℃为恒温对照,设计4个不同幅度的变温组,分别为25℃±1℃、25℃±2℃、25℃±3℃、25℃±4℃,研究了五种不同温度处理下凡纳滨血清中的葡萄糖含量、肝胰脏中的已糖激酶(HK)和丙酮酸激酶(PK)及肌肉组织中的热休克蛋白70的相对含量在一天内的变化。结果表明：在恒温组中葡萄糖含量和热休克蛋白70均存在昼夜变化规律,但在温度波动组中这种规律随着波动幅度的增大逐渐消失。恒温组的PK活力变化幅度不大,随着温度波动幅度的增大,其变化逐渐剧烈,以25℃±4℃组尤为明显。HK活力受到葡萄糖流量和糖酵解作用的影响,随着温度变化幅度的增加,其变化也有所趋于平稳,但没有葡萄糖明显。
A series of indoor experiments were conducted to investigate the effects of light fluctuations on the molting and growth and of Litopenaeus vannamei. Based on the results and the resrarches of temperature flucyuations, the mechanism for the different characters of L. vannamei under fluctuating light and temperature treatments was discussed. The main results were presented as the followings:
1. The effects of light fluctuations on the molting, growth of Litopenaeus vannamei and preliminary researches on its mechanisms
1.1 The effect of rhythmic light intensity fluctuation on the molting and growth of Litopenaeus vannamei and its mechanisms
The method of experimental ecology was used to investigate the effect of rhythmic light intensity fluctuation on the molting and growth of Litopenaeus vannamei. Molting, growth performance, feeding and body composition of the shrimps were tested in one constant light intensity (2700 lx) treatment and four rhythmic fluctuating light intensity treatments (2700±600 lx; 2700±1200 lx; 2700±1800 lx; 2700±2400 lx). The initial wet body weight of shrimp was 2.736±0.002 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) of shrimps in 2700±600 lx,2700±1200 lx and 2700±1800 lx treatments were significantly higher than those in 2700 lx treatment (P< 0.05); the WG in 2700±2400 lx treatment was significantly higher than those in 2700 lx treatment (P<0.05), but SGRd was not (P>0.05). (2) The feed intake (FId) in 2700 ±600 lx,2700±1200 lx and 2700±1800 lx treatments were significantly higher than those in 2700 lx treatment (P<0.05), but 2700±2400 lx treatment was not (P>0.05). The food conversion efficiency (FCEd) in 2700±600 lx treatment was significantly lower than those in 2700 lx treatment (P<0.05), but the other three rhythmic fluctuating light intensity treatments were not (P>0.05). (3) The energy allocation to growth in 2700±1200 lx and 2700±1800 lx were significantly higher than those in 2700 lx treatment (P<0.05) and the energy allocation in respiration in 2700±1200 lx and 2700±1800 lx were significantly lower than those in 2700 lx treatment (P<0.05). (4) There was no significantly difference of molting rate (MF) and energy allocation in exuviate between constant light intensity and four rhythmic fluctuating light intensity treatments (P>0.05). (5) The body composition of the shrimps was significantly affected by the different light treatments (P<0.05). Compared to the constant light intensity treatment, the value of crude lipid decreased because of the effect of the fluctuating light intensity.
1.2 The effect of periodic abrupt change in light intensity on the molting and growth of Litopenaeus vannamei and its mechanisms
The molting and growth of Litopenaeus vannamei were tested in one constant light intensity (60 lx) treatment and four periodic abrupt changing light intensity treatments (600-60 lx; 1500-60 lx; 3000-60 lx; 6000-60 lx). The initial body weight of shrimp was 2.733±0.017 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) in 1500-60 lx and 3000-60 lx treatments were significant higher than those in 60 lx treatment (P<0.05); the WG in 600-60 lx treatment was significantly higher than those in 60 lx treatment (P<0.05), but the SGRd was not (P>0.05). Opposite to what was observed in the 600-60 lx treatment, the WG in 6000-60 lx had no difference from those in 60 lx treatment (P>0.05), but the SGRd in 6000-60 lx treatment was significantly higher than those in 60 lx treatment (P<0.05). (2) There is no significant difference in FId for all regimes (P>0.05), but the FId of shrimps in 1500 -60 lx,3000-60 lx and 6000-60 lx treatments were significant higher than those in 60 lx treatment (P<0.05). (3) The energy allocation in respiration and excretion of shrimps in 1500-60 lx,3000-60 lx and 6000-60 lx treatments were significantly lower than those in 60 lx treatment (P<0.05), the energy allocation in the growth were significantly higher than those in 60 lx treatment (P<0.05). (4) The MF and energy allocation in exuviate in 6000-60 lx treatment were lowest, and significantly lower than those in other treatments (P<0.05), but the MF and energy allocation in exuviate in 1500-60 lx or 3000-60 lx treatment was not significantly different from those in 60 lx treatment (P>0.05). (5) The ash had a positive correlation with the body weight, but the crude protein did not, and the abrupt changing light enhanced the content of crude lipid of shrimps.
1.3 The effect of rhythmic light color fluctuation on the growth and molting of Litopenaeus vannamei and its mechanisms
The molting and growth of Litopenaeus vannamei were tested under two kinds of color treatments:three constant light color treatments (yellow light, Y; green light, G; blue light, B) and three rhythmic fluctuating light color treatments (blue light was changed to yellow light, BY; blue light was changed to green light, BG; green light was changed to yellow light, GY). The initial body weight of shrimp was 1.212±0.010 g, the period of this experiment was 45 days. The main results were as follows:(1) The weight gain (WG) and the specific growth rate (SGRd) of shrimp in BG and BY treatments were significantly higher than those in B treatment (P>0.05). The WG in GY treatment was significantly higher than those in B treatment (P>0.05), but the SGRd was not (P>0.05). (2) There is no significant difference in FId for all regimes (P>0.05), the FCEd of shrimps in GY treatment was no significantly different from those in B treatment (P>0.05), but the FCEd of shrimps in BY and BG treatments were significantly higher than those in B treatment (P<0.05). (3) However, the energy allocation in respiration of shrimps in all treatment did not have significant difference (P >0.05), but the energy allocation to excretion of shrimps in BG and BY treatments were significantly lower than those in B treatment (P<0.05), the energy allocation to growth were significantly higher than those in B treatment (P<0.05). (4) The molting rate (MF) and energy allocation in exuviate of shrimps in three rhythmic fluctuating light color treatments were generally higher than that in three constant light color treatments (P<0.05), there was a clear correlation between MF and growth in every regime. (5) The ash of shrimps had a positive correlation with the body weight, but the crude protein did not, and the abrupt changing light enhanced the content of crude lipid of shrimps. The lipid level of shrimps increased with the decrease of moisture level.
1.4 The effect of periodic light color fluctuation on the molting and growth of Litopenaeus vannamei and its mechanisms
The method of experimental ecology was used to investigate the effect of periodic light color fluctuation on the molting and growth of Litopenaeus vannamei. Molting and growth performance of the shrimps were tested under two kinds of color treatments: three constant light color treatments (yellow light, Y; green light, G; blue light, B) and three periodic fluctuating light color treatments (blue light was changed to yellow light, BY; blue light was changed to green light, BG; green light was changed to yellow light, GY). The initial body weight of shrimp was 1.212±0.018 g (mean±S.E.), the period of this experiment was 48 days. The main results were as follows:(1) the weight gain (WG) and the specific growth rate (SGRd) of shrimp in BG treatments were highest and significantly higher than those in Y or B treatment (P>0.05). The WG in GY treatment was significantly higher than those in Y or B treatment (P>0.05), but the SGRd was not (P>0.05). (2) There is no significant difference in FId for all regimes (P>0.05), the FCEd of shrimps in GY treatment was no significantly different from those in Y or B treatment (P>0.05), but the FCEd of shrimps in BY and BG treatments were significantly higher than those in Y or B treatment (P<0.05). (3) Although there was no significant difference in energy allocation in respiration under different light color regimes (P>0.05); but the energy allocation in respiration in BG treatment was lower about 4% than those in B treatment, so the energy allocation to growth of shrimps in BG treatment was significantly higher than those in Y or B treatment (P<0.05). (4) The molting rate in GY treatment was highest, but there was no correlation between MF and growth. (5) The ash had a positive correlation with the body weight, but the crude protein did not, and the periodic light color fluctuation decreased the content of crude lipid of shrimps.
2. Effects and mechanism of light fluctuations on growth of Litopenaeus vannamei
2.1 The variation of LD, PK and LDH of Litopenaeus vannamei in periodic abrupt change in light intensity and constant light intensity
Four periodic fluctuation light intensity treatments (600-60 lx,1500-60 lx,3000-60 lx and 6000-60 lx) and five constant light intensity treatments (60 lx,600 lx,1500 lx,3000 lx,6000 lx) were arranged to investigate the effect of periodic abrupt change in light intensity and constant light intensity on the enzymes relating to respiratory metabolism. An alternation of six days high light intensity and two days low light intensity was adopted in the periodic fluctuation light intensity treatment. After two cycles of light intensity periodic fluctuation, sampling five inter-molt stage shrimps in the five constant light intensity treatments to assay the PK activities, the LDH activities and the lactic-acid at 0 h,0.5 h,1 h,3 h,6 h,9 h,12 h,14 h after the light on, respectively; and doing that in four periodic fluctuation light intensity treatments at 0.5 h,1 h,3 h,6 h,12 h,24 h,48 h,72 h,144 h,144.5 h,145 h,147 h,150 h,156 h,168 h, 192 h of the third cycle of light intensity periodic fluctuation, respectively. The main results were as follow:(1) A decrease of total PK and LDH activities was found in periodic fluctuation light intensity treatments, but LD been not; the reason for the phenomenon might be that the efficacy of some enzymes was enhanced in periodic fluctuation light intensity treatments. (2) The fluctuations of enzymes in 600 lx and 1500-60 lx treatments were stable, but those in 6000 lx and 6000-60 lx treatments were acute. (3) The lactic-acid content of 6000-60 lx treatment in the phase of high light was high and this phenomenon was also found in the later stages of high light and the beginning of low light in 600-60 lx. These results might indicate that the shrimp may be in a bad environment. (4) The LDH fluctuation when the light changed from high intensity to low intensity was more acute than those when the light changed from low intensity to high intensity. This may due to the difference types of shrimps acclimate to different light intensities and the longer time of high light intensity in our experiment. The results suggested that the suitable periodic abrupt changing light intensity enhanced the effectiveness of the enzyme in shrimps and its fluctuation of enzyme was stable, so the growth of shrimps was better.
2.2 The variations of GLU, LD, HK and PKof Litopenaeus vannamei under a rhythmic fluctuating color light and two constant color lights
A rhythmic fluctuating light color treatment (Blue light was changed to green light, BG) and two constant light color treatments (green light, G; blue light, B) were selected from the last experiment to investigate the effect of rhythmic fluctuations in light color on the enzymes relating to respiratory metabolism. Sampling five inter-molt stage shrimps to assay the hemolymph glucose, lactic-acid, the HK and PK activities of hepatopancreas at 7:00,9:00,13:00,14:00,16:00,20:00 on the following days:the first day, third day, fifths day, tenth day, twentieth day and fortieth day of the experiment, respectively. The main results were as follow:(1) The hemolymph glucose content had a small increase after the light on, but the hemolymph lactic-acid content did not. The HK activities of shrimps in B and G treatments decreased after switching on the light, but those in BG treatment increased. And the PK activities in all treatment did not have an obvious change. (2) A significant effect of rhythmic fluctuating light color on glucose, lactic-acid, HK and PK activities was found in the beginning of the experiment, but this effect disappear gradually when the experiment went on. (3) The fluctuations of HK and PK activities of shrimps in B treatment were acute, and the time of HK and PK activities became stable was late; but those in G treatment was comparatively stable.
3. Effects and mechanism of temperature fluctuations on growth of Litopenaeus vannamei
3.1 The effects of cyclical temperature changes on growth and physiological status of Litopenaeus vannamei
The purpose of this study was to investigate the growth and physiological status of Litopenaeus vannamei subjected to one constant temperature (25℃) and four cyclical temperature change regimes (25℃±1℃,25℃±2℃,25℃±3℃and 25℃±4℃). The growth rates of shrimp at 25℃±2℃or 25℃±3℃were significantly higher than that at a constant temperature of 25℃. On the other hand, the growth rate in 25℃±4℃regime was significantly lower than those in other regimes. The daily feed intake rate of shrimp at 25℃±4℃was the lowest and the food conversion efficiency was also significantly lower than those at 25℃±2℃and 25℃±3℃, respectively. The food conversion efficiency at 25℃±2℃or 25℃±3℃was significantly higher than those in other regimes. Thus, it can be inferred that the growth enhancement in the test shrimp at the suitable diel fluctuating temperatures was due to high food conversion efficiency. Studies of the physiological parameters showed that at 25℃±4℃, the hemolymph glucose content of the test shrimp was the lowest, while the activity of PK in hepatopancreas was the highest, which indicated that the test shrimp at 25℃±4℃was in a stressed condition. The hemolymph glucose content of the test shrimp at 25℃±3℃was the highest and the activity of HK in hepatopancreas was the lowest. These results indicated that the test shrimps at 25℃±3℃were not in a stressed condition. Compared with the constant temperature regime, the expression of HSP70 in any of the four cyclical temperature change regimes was not significantly increased. The reason for this might be that the fluctuation amplitude of±4℃did not induce the increased expression of HSP70.
3.2. The variations of enzymes involved glycolysis and HSP70 of Litopenaeus vannamei during the temperature cyclical change
The rhythmic fluctuations of hemolymph glucose content, HK and PK activities in hepatopancreas and HSP70 expression in Litopenaeus vannamei were measured under one constant temperature (25℃) and four cyclical temperature change regimes (25℃±1℃,25℃±2℃,25℃±3℃and 25℃±4℃). The main results were as follows: a day/night rhythmicity was found in the rhythmic fluctuations of hemolymph glucose content and HSP70 expression under 25℃, but the rhythmicity disappeared gradually with the increased fluctuation amplitude of temperature. The PK activities varied gently under 25℃. Due to the increase of temperature fluctuation amplitude, the fluctuation of PK activities became acute, especially in 25℃±4℃regime. Because of the effect of glucose flux and glycolysis, the fluctuation of HK activity became stable with the increased fluctuation amplitude of temperature.
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