盐度对中华鲟生长的影响机制及中华鲟的等渗点分析
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摘要
中华鲟(Acipenser sinensis)为溯河洄游型鱼类,是国家一级重点保护动物。在迁地保护下,淡水环境中成熟的中华鲟亲鱼面临个体变小,繁殖力下降的问题。为复壮中华鲟个体大小,本文主要分析了盐度对中华鲟生长影响的机制；通过对5月龄,9月龄,12-15龄中华鲟进行实验,构建了“循环养殖系统中特定收集器和计数法计算饲料转化率的方法”和“分级稀释建立高连续梯度盐度选择实验的装置及方法”；探讨了中华鲟幼鱼盐度适应机制；分析了5月龄,9月龄,12-15龄中华鲟的等渗点；构建了叉长、体重和年龄与血清渗透压和等渗点的关系模型；对比了等渗环境和淡水环境下中华鲟幼鱼在与生长相关的行为、激素、酶含量以及组织和卵质成分方面的差异；荧光定量RT-PCR分析了等渗组和淡水组PRL(prolactin), GH (growth hormone), IGF-1(insulin-like growth factor1)在组织的表达差异；同时比较了淡水环境下中华鲟再成熟卵和海水卵的主要成分差异。得出主要结论：盐度并不影响中华鲟幼鱼个体的生长速率大小,但是渗透压调节能耗需要通过增加摄食来实现；全人工繁殖中华鲟幼鱼和野生幼鱼一样具备盐度适应性,且在淡水中已经形成盐度适应机制,喜好盐度位于等渗点区域；等渗点盐度有利于中华鲟幼鱼的钙质积累和酶耗能的节约；等渗组和淡水组相比,淡水组幼鱼需要显著较多的摄食实现和等渗组幼鱼同步生长；等渗点盐度和中华鲟年龄最相关,等渗点随年龄发生变化；盐度和溯河洄游习性对卵质有较大的影响；自幼鱼到性成熟建议中华鲟养殖盐度为8.3-11.5psu。具体方法及得到结论如下：
1.构建了“循环养殖系统中特定收集器和计数法计算饲料转化率的方法”和“分级稀释建立高连续梯度盐度选择实验的装置及方法”,前者为计算饵料系数和计算早、午、晚摄食节律和日摄食节律提供了方便和准确性；后者为盐度喜好实验提供了设备系统及方法论。
2.5月龄中华鲟在淡水中已经形成盐水适应机制且喜好盐度位于等渗盐度区域。5月龄中华鲟分别在淡水(FW,0.41psu)和咸水(BW,12.5psu)下养殖28天,前10天为适应期,后18天为适应后期,在第28天采集左右侧第二鳃弓鳃丝进行扫描电镜和透射电镜检查。淡水(0.41psu)和咸水(12.5psu)下,鳃丝中均出现浆细胞、扁平细胞和氯细胞。氯细胞方面,在淡水中氯细胞有两种,一种是球形氯细胞,高度高于临近的扁平细胞；另一种是三角形氯细胞,该氯细胞在鳃丝的高度低于扁平细胞,呈三角形。咸水中则仅出现一种氯细胞—球形氯细胞；且淡水和咸水下,球形氯细胞分布密度和大小没有差异。同时在淡水中发现从三角形氯细胞到球形氯细胞的过渡细胞。通过在适应后期,血清渗透压,血清PRL,GH和IGF-1水平在淡水和各盐水组间没有差异,也是淡水中已经形成盐水适应机制的一种反映。通过“分级稀释建立高连续梯度盐度选择实验的装置及方法”,分析得出5月龄中华鲟的喜好盐度区间为7.5-8.5psu,该盐度接近于等渗点9.52psu；
3.盐度改变了中华鲟血清离子的组成。9月龄中华鲟等离子点为：K+：9.65±1.01psu；Cl-：6.82±0.37psu；Na+：9.40±0.58psu； Ca2+：1.62±0.71psu。血清钠离子浓度随盐度增加没有显著变化(P=0.250),但血清氯离子随着盐度的增加而增加(P<0.05),NaCl理论贡献值和实际贡献百分比在各盐度没有差异(OsmNaCl=224.27±3.17-233.41±2.63rnOsm/kg,93.39±2.22%-96.79±4.17%)。在各个盐度,血清Mg2+的水平远远低于水样Mg2+浓度。但是血清P浓度却是远远高于水样。血清K+,Mg2+浓度随盐度升高而显著升高,但血清Ca2+则没有变化,Mg:Cl比随环境盐度升高而增加,Ca：Cl比和K：Cl比随盐度升高则没有变化。对比分析淡水组和等渗组血清电解质,发现血清K+, Mg2+, Ca2+浓度,Mg:Cl, Ca:Cl和K：C1比值没有显著差异。
4.盐度改变了中华鲟的行为。对于9月龄中华鲟而言,在所有盐度组中鳃盖运动频率最低(74±2min-1)和最低的摆尾频率(51±3min-1)均出现在IW(isosmotic water)组。饲料转换率FCR(1.14-1.46)和特定生长率SGR(0.85-1.31)在所有盐度组中没有显著性差异(包括等渗点)(P FCR>0.05和PSGR>0.05)。各盐度下早中晚摄食节律方面,0.25,6.32和25psu组均没有差异,在12psu下晚20：00摄食量显著高于15：00,在19psu下,8：00摄食量显著多于20：00。日摄食方面,在与等渗盐度差异较大的盐度水体(0.27psu和25psu)中华鲟依靠更多的摄食,补充用于渗透压调节消耗的能量。
5.等渗(IW)和淡水组(FW)对比实验中,盐度影响中华鲟血清PRL激素和GH/IGF-1mRNA的表达。对于9月龄中华鲟而言,在适应期实验中,PRL在12.7psu时略高于至5.98,6.32和9.69psu,但在19.1和25psu又下降到低盐度先前水平。生长激素随盐度增加而增加,在25psu时最高。IGF-1的趋势和GH相反,IGF-1的水平最低发生在25psu。在适应后实验中,三种激素水平盐度随盐度变化没有表现出显著差异性。在等渗组和淡水组比较实验中,淡水中PRL水平显著高于等渗组(P<0.01)但GH和IGF-1水平在等渗组和淡水组则没有显著性差异。
6.酶含量方面,淡水中华鲟相比于等渗盐度下中华鲟,消耗较大的消化力为渗透压调节提供能量。对于9月龄中华鲟而言,中肠,后肠和肾的Na+/K+-ATP酶含量在IW组和FW(freshwater)组相似,而在鳃丝(P<0.05),前肠(P<0.01)和螺旋瓣(P<0.01)FW明显高于IW高约7%。前肠,螺旋瓣和胃在FW中脂肪酶含量比IW组高出10%(P<0.05)。此外,前肠ALT含量在FW显著高于IW(P<0.05)。在肝脏中淀粉酶的含量也是FW组高于IW组(P<0.05)。另外,在螺旋瓣的蛋白酶含量在FW高于IW约18.1%,而在中肠,则是在IW高于FW约13.2%(P<0.05)。
7.等渗(IW)和淡水组(FW)对比实验中机体组成成分的差异显著。对于9月龄中华鲟而言,在等渗(IW)和淡水组(FW)对比实验中,脂肪及其脂肪酸组成,粪便蛋白质、脂肪及其氨基酸和脂肪酸组成没有差异,但是肌肉钙离子组成差异显著。经过52天的实验,等渗组肌肉钙离子含量显著高于淡水组；全骨板和头部软骨钙、磷含量等渗组低于淡水组,但是镁离子含量则是等渗组高于淡水组。
8.盐度造成中华鲟卵质的差异。海水卵较之于淡水卵蛋白含量明显较高,脂肪含量则明显偏低。脂肪酸含量有极大地差异。两种卵均含17种氨基酸,包括10种鱼类必需氨基酸。总氨基酸量和各氨基酸没有差异。卵黄蛋白没有差异。磷元素、镁离子和维他命A在海水卵中含量明显多于淡水卵,维他命E则是淡水卵多于海水卵。钙离子和锌离子没有差异。在淡水卵中检测到镉离子,海水卵中则未检测到。
9.中华鲟血清渗透压和等渗点和年龄最相关,自幼鱼到性成熟,中华鲟等渗点为8.3-11.5psu。5月龄中华鲟的喜好盐度位于等渗盐度点,血清渗透压与年龄最相关。9月龄子一代中华鲟幼鱼在0.27,6,12,19和25psu盐度下适应后,水渗透压和盐度的关系通过线性拟合确定。二阶多项式拟合血清渗透压和盐度之间的关系,通过解方程求得等渗点为9.57±0.06psu。在该盐度下,血清渗透压等于咸水渗透压,鱼体无渗透压力。五月龄子二代中华鲟在淡水中氯细胞已经形成咸水适应机制,等渗点盐度为9.52psu。以“分级稀释建立高连续盐度梯度实现水生动物盐度选择实验的装置和方法”为新的实验体系,分析得出5月龄子二代中华鲟喜好盐度为8.5-9.5psu,这和5月龄子二代中华鲟等渗点9.52psu保持一致。对5月龄和9月龄中华鲟血清渗透压分析发现：盐水环境和淡水环境下渗透压没有显著差异,淡水环境下渗透压可作为盐水环境渗透压；等渗点下血清渗透压和与中华鲟相适应的淡水条件下血清渗透压在同一个水平。据此,该结果提供了迁地保护下大型中华鲟亚成体等渗点的简易计算方法。测得淡水环境下中华鲟血清渗透压,即作为等渗点盐度渗透压。将此渗透压带入盐度和渗透压关系式,即可计算出等渗点盐度。血清渗透压和年龄的二次多项式模拟最吻合,相关系数最高。因此认为年龄和中华鲟血清渗透压最相关：Y=278.061-11.3127X+0.6587X2,Y表示血清渗透压X表示年龄。对该二次多项式求导得出导数为0时,年龄为8.6年,即从出生到9龄时,中华鲟血清渗透压逐渐降低；从9龄开始,中华鲟血清开始随年龄增加而逐渐升高。同理等渗点也和年龄最为相关：Y=9.79431-0.35711X+0.0213X2,Y表示血清渗透压,X表示年龄。一阶求导得零点横坐标为8.38龄,即第9龄,这和渗透压结果一致。即从出生到9龄时,中华鲟等渗点逐渐降低；从9龄开始,中华鲟血清开始随年龄增加而逐渐升高。
总之,考虑到在等渗盐度组较之于淡水组明显较低的鳃盖运动频率,摆尾频率,鳃丝Na+/K+-ATPase含量以及所观察到的低脂肪酶,ALT和淀粉酶含量,显著降低的日摄食量和不变的特定生长率,肌肉中显著升高的Ca2+含量和增加的骨板和软骨中Mg2+含量,以及盐度对卵质产生的影响,等渗盐度是中华鲟的较理想生长环境盐度。此外,在淡水养殖中,Ca2+和Mg2+应该进行适当补偿。同时,中华鲟血清渗透压和等渗点盐度随年龄而变化,随着生长的进行,中华鲟的等渗养殖盐度需要进行调整。
Populations of A.sinensis, an anadromous fish species with Class-I state protection in China, grow in coastal waters (seawater) and enter into the Yangtze River (freshwater) to spawn after reaching sexual maturity, IUCN assessed as CR (Critically Endangered) status. A.sinensis subjecting to ex-situ conservation no longer live in brackish water, but spend their entire life cycle in freshwater (FW). The small size of the adults matured in freshwater was associated with decreased fecundity, resulting in more challenge to the conservation work. To restore the body size, this paper aimed to argue whether salinity affect sturgeon body size growth.5-month-old,9-month-old and12-15-year-old sturgeon were tested; Therefore, we constructed "specific collector combined count method in recirculating aquaculture system for feed conversion ratio determination" and the 'establishment of high grade continuous gradient salinity experiment apparatus and method for salinity preference in aquatic animal"; Explored the juvenile Chinese sturgeon salinity adaptation mechanisms; analyzed isosmotic points of5-month-old,9-month-old and12-15-year-old sturgeon; constructed relationships between fork length, body weight and age and serum osmolality, relationships between fork length, body weight and age and isosmotic points; Compared sturgeon behaviors, hormones, enzymes activities and tissue compositions between isosmotic environment and freshwater environment; Quantified fluorescence RT-PCR analysis of GHmRNA and IGF-1mRNA in tissues between isosmotic water (IW) and FW groups; Alalyzed the salinity and migration loss effects on the compositions of freshwater eggs and seawater eggs of Chinese sturgeon; The following conclusions were obtained:
1. Establish two methods:'specific collector combined count method in recirculating aquaculture systems for feed conversion ratio determination' and 'establishment of high grade continuous gradient dilution salinity experiment apparatus and method for salinity preference'; Count method could reduce physical, biochemical and mechanical error of quantifying uneaten feed collected from holding water. Simultaneously, it saved the disposals uneaten pellets after the uneaten pellets are collected, replaced siphoning by special designed collector and saved time and labors. In addition, the effects of the feces can be ignored. The former one was for calculation of feed conversion and provides a convenient feeding rhythm, meanwhile improved accuracy of FCR; the latter provide the equipment system and methodology for salinity preference experiments. The advantages lies in, on the one hand, the appetas promises long-time salinity gradients than former appatases. On the other hand, tested animals can swim free and rechoose their location.
2. Five-month-old A.sinensis reared respectively in FW (0.41psu) and brackish water (BW,12.5psu) for26days. The first10days was adaptation period, the later16days were post acclimated period. In the26th day, one of filaments both in left and right second branchial arch was sampled for scanning electron microscopy and transmission electron microscopy examination, respectively. The gill filaments appeared in the mucous cells, pavement cell and MRC on the surface of filament. In terms of MRCs, there were two subtypes of MRCs in FW, one was the ball-shaped MRC, which was higher above the neighboring pavement cells, and the other one was triangular MRC, which was triangle and flat, but it depressed between pavement cells. Only ball-shaped MRCs emerged in filament of BW-acclimated fish. Furthermore, the density and distribution of ball-shaped MRCs did not differ between freshwater and brackish group. It was also found that transitional cells, from triangle to ball-shaped, existed in FW-acclimated filament. Serum PRL, GH and IGF-1levels did not differ between FW and BW groups, which also reflected fish developeded BW adaptation mechanism in FW. Five-month-old A.sinensis had formd adapted MRCs (mitochondria-rich cell) to BW, whose isosmotic point was9.52psu. Taken "establishment of high grade continuous gradient dilution salinity experiment apparatus and method for salinity preference" as a new experimental system and revealed that five-month-old A.sinensis prefer7.5-8.5psu. This agreed to that isosmotic point of5-month-old was9.52psu. As salinity increased, food intake decreased significantly at30psu and fish ceased feeding at38psu compared to food intake at12.5psu.
3. For9-month-old sturgeon, isoionic points were analyzed in the same way as the isosmotic point was determined: K+:9.65±1.01psu; Cl-:6.82±0.37psu; Na+:9.40±0.58psu; Ca2+:1.62±0.71psu. The trend of Na+concentration as salinity increased was not pronounced (P=0.250), but Cl-tended to pronouncedly higher levels as salinity increased (P=0.021). NaCl contributed to total osmolality and the real NaCl percentages had the same levels in all salinity treatments (OsmNaCl=224.27±3.17-233.41±2.63mOsm/kg;93.39±2.22%-96.79±4.17%) as it did at the isosmotic point (OsmNaCl=227.45±8.04mOsm kg-1;89.93±4.09%). Serum Mg2+level were far lower than the water Mg2+level at any salinity treatment. Nevertheless, serum P was much higher than water at any salinity treatment. Serum Mg2+and K+levels were significant higher in as salinity increased.
4. Salinity changed the gill ventilation rate, tail beat frequency and daily feeding rhythm.
For9-month-old A.sinensis, the lowest gill ventilation rate (74±2min-1) and the lowest tail beat frequency (51±3min-1) were observed in IW. The feed intake had no difference at8:00,15:00and20:00in0.25,6.32and25psu groups. FCRs (1.14-1.46) and SGRs (0.85-1.31) were comparable at all salinity groups, including the isosmotic point (PFCR>0.05and PSGR>0.05), but FCR was smaller at25psu and IW groups. In long term salinity tolerance, the condition factor was markedly higher after22d for all15sturgeon compared to the values before the test (P<0.05). However, the feeding rhythm at20:00of12psu-acclimated fish was significantly more than15:00. In19psu group, the feed intake at8:00was significantly more than at20:00. The difference resulted from individual difference. In general, there was no difference in feeding rhythm of each feeding at any salinity group. But the daily feed intake in25and0.25psu groups were significantly higher than6.32and12psu groups.
5. For9-month-old A.sinensis, in the acclimation period, PRL increased slightly at12.7psu compared to5.98,6.32and9.69psu groups, but it declined again at19.1and25psu. In terms of GH, this hormone gradually increased as salinity increased and was the highest at25psu. The trend of IGF-1was contrary to that of GH, with the lowest IGF-1level observed at25psu. In long term salinity tolerance, the three hormone levels exhibited no significant differences among salinities. As time went on, no obvious changes were observed in any hormone. In IW group and FW group comparison test, the PRL level was significantly higher at the isosmotic point than in freshwater (P<0.01). However, GH and IGF-1exhibited no significant differences. For5-month-old A.sinensis, PRL, GH and IGF-1showed no trend in both acclimated period and post-acclimatedion. RT-PCR analysis of related expression of pituitary GH and IGF-1in different issues showed: pituitary GH mRNA and liver IGF-1mRNA expression in FW-acclimated group were significantly higher than IW-acclimated group; Muscle and kidney IGF-1mRNA levels had the same level between FW and IW groups.
6. The digestive enzymes and Na+/K+-ATPase consumed more energy instead of energy for growth group in FW than in IW group. The in the gill (P<0.05), foregut (P<0.01) and spiral valve (P<0.01) in FW were approximately7%higher than in IW. The lipase activities in the foregut, spiral valve and stomach in FW were10%higher than in the IW group (P<0.05), and the activities were similar in the other examined issues. In addition, ALT (alamine transaminase) activity in the foregut was significantly higher in FW than in IW, showing a difference of approximately8%compared to IW group (P<0.05). Amylase activity in the liver was also higher in FW than in IW (P<0.05), while these values were equal in other tissues. Furthermore, the protease activity in the spiral valve was18.1%lower in IW than in FW, while in the mid-gut, it was13.2%higher in IW than in FW (P<0.05).
7. Comparisons of feces protein and amino acids composition showed no difference, as well fatty acids. Muscle protein and amino acids had the similar levels between IW-acclimated fish and FW-acclimated fish. Both groups had the same level fat content (1.92±2.69) g and similar fatty acids compositions. However, in terms of minerals of muscle, Ca content in IW acclimated fish was significantly higher than in in FW-acclimated fish, P and Mg contents had the same levels. Different conditions about Ca, P, Mg contents were present in scutes and cartilage. In abdominal scute and cartilage, Ca levels were higher in FW-acclimated fish than IW-acclimated fish, but in lateral scute and back scute, Ca, Mg and P levels in IW group were higher than in FW group. In cartilage, Mg was not detected both in IW and IW groups. Overall, Ca and P levels in the total scutes and head cartilage in FW-acclimated fish were higher than in IW-acclimated fish, but Mg level in IW (2019.7±183.4mg/100g) was higher than in FW (1827.8±107.8mg/100g). The liver lipid contents (60.96±5.30mg/100g,45.29±18.34mg/100g), VA contents (31.22±9.41mg/100g,24.43±5.15mg/100g) had similar level in the two groups (FW was former, IW was latter). VE values were not detected in liver.
8. Eggs biochemical contents from wild origin and remature A.sinensis in freshwater were compared, which were related to induce spawning success. The results included dry weight, lipids, fatty acids, protein, amino acids, vitellin (Vn), phosphorus (P), vitamin A (VA) and vitamin E (VE) as well as the concentrations of relevant trace elements. The dry weights of the eggs did not differ between the sturgeons. The wild sturgeon eggs had significantly higher protein content but lower lipid content than the remature sturgeon eggs. There were multiple and significant differences in the fatty acids. Vn showed no difference. The P, magnesium (Mg) and VE contents in seawater eggs were significantly higher than in freshwater eggs. The calcium (Ca) and zinc (Zn) contents showed no difference between both batches. Cadmium (Cd) was only detected in one sample of freshwater eggs. These results helped to discriminate between remature (farmed) and wild origin Chinese sturgeon eggs. It is important to take measures to decrease lipid percentages and to increase protein contents of eggs for remature Chinese sturgeon or even sturgeon matured in freshwater.
9. A.sinensis were acclimated in0.27,6,12,19and25psu, relationship between water osmolality and salinity was determined by linear fitting. Second-order polynomial fitting were used to assess relationship between serum osmolality and salinity. Isosmotic point was got (9.57±0.06psu) through solution of the two equations. At isosmotic salinity environment, serum osmolality equaled to environment osmolality, sturgeon faced no osmotic pressure. The results of analyzing serum osmolality of5-month and9-month-old A.sinensis showed: there were no significant differences of serum osmolality under FW or BW, including IW. Accordingly, the results provide method to assess isosmotic point of larger ex-situ conservation subadult sturgeon in FW. Regard as the value of serum osmolality in FW as serum osmolality in BW. Serum osmolality in FW-acclimated fish had the same level as serum osmolality in IW-acclimated fish. Simultaneously, we could also take serum osmolality in FW-acclimated fish as serum osmolality in IW-acclimated fish. Take the value of serum osmolality in FW group into the equation of relationship between salinity and water osmolality, and then get the salinity i.e. isosmotic point. The order-two polynomial fitted best in relationship between age and serum osmolality, the correlation coefficients was highest. Therefore age was considered as the most relevant factor to serum osmolality:Y=278.061-11.3127x+0.6587X^2. Derived the equation and the X corresponding to value of derivative equled to zero was8.6years, i.e. from birth to age9the serum osmolality decreased and serum gradually began to increase with age since9-year-old. Similarly, isosmotic osmolality and age were most relevant:Y=9.79431-0.35711X+0.0213X2. Derived the equation and the X corresponding to value of derivative (0) was8.6years. This was similar to serum osmolality equation, i.e. from birth to age9-years, the Chinese sturgeon isosmotic point decreases, from9years on, the isosmotic point began to gradually increase with age.
In conclusion, the Chinese sturgeon developed seawater-entry readiness in freshwater and the fish preferred isosmotic salinity compared to other salinities. Considering the obviously lower gill ventilation rate, tail beat frequency, Na+/K+-ATPase activity in the gills as well as the lower lipase, ALT and amylase activities observed, lower daily feeding rhythm but same SGR levels, significantly higher Ca content in muscle and increased Mg content in scutes and cartilage in IW compared to FW a salinity corresponding to the IW is ideal for the growth of naive A. sinensis. A. sinensis is suggested to rear in8.3-11.5psu. Besides, Ca2+and Mg2+should be compensated in FW culture. Meanwhile, the plasma osmolality and isosmotic point may vary with the size and age of fish, and salinity may need to be adjusted as naive A. sinensis grow.
1.构建了“循环养殖系统中特定收集器和计数法计算饲料转化率的方法”和“分级稀释建立高连续梯度盐度选择实验的装置及方法”,前者为计算饵料系数和计算早、午、晚摄食节律和日摄食节律提供了方便和准确性；后者为盐度喜好实验提供了设备系统及方法论。
2.5月龄中华鲟在淡水中已经形成盐水适应机制且喜好盐度位于等渗盐度区域。5月龄中华鲟分别在淡水(FW,0.41psu)和咸水(BW,12.5psu)下养殖28天,前10天为适应期,后18天为适应后期,在第28天采集左右侧第二鳃弓鳃丝进行扫描电镜和透射电镜检查。淡水(0.41psu)和咸水(12.5psu)下,鳃丝中均出现浆细胞、扁平细胞和氯细胞。氯细胞方面,在淡水中氯细胞有两种,一种是球形氯细胞,高度高于临近的扁平细胞；另一种是三角形氯细胞,该氯细胞在鳃丝的高度低于扁平细胞,呈三角形。咸水中则仅出现一种氯细胞—球形氯细胞；且淡水和咸水下,球形氯细胞分布密度和大小没有差异。同时在淡水中发现从三角形氯细胞到球形氯细胞的过渡细胞。通过在适应后期,血清渗透压,血清PRL,GH和IGF-1水平在淡水和各盐水组间没有差异,也是淡水中已经形成盐水适应机制的一种反映。通过“分级稀释建立高连续梯度盐度选择实验的装置及方法”,分析得出5月龄中华鲟的喜好盐度区间为7.5-8.5psu,该盐度接近于等渗点9.52psu；
3.盐度改变了中华鲟血清离子的组成。9月龄中华鲟等离子点为：K+：9.65±1.01psu；Cl-：6.82±0.37psu；Na+：9.40±0.58psu； Ca2+：1.62±0.71psu。血清钠离子浓度随盐度增加没有显著变化(P=0.250),但血清氯离子随着盐度的增加而增加(P<0.05),NaCl理论贡献值和实际贡献百分比在各盐度没有差异(OsmNaCl=224.27±3.17-233.41±2.63rnOsm/kg,93.39±2.22%-96.79±4.17%)。在各个盐度,血清Mg2+的水平远远低于水样Mg2+浓度。但是血清P浓度却是远远高于水样。血清K+,Mg2+浓度随盐度升高而显著升高,但血清Ca2+则没有变化,Mg:Cl比随环境盐度升高而增加,Ca：Cl比和K：Cl比随盐度升高则没有变化。对比分析淡水组和等渗组血清电解质,发现血清K+, Mg2+, Ca2+浓度,Mg:Cl, Ca:Cl和K：C1比值没有显著差异。
4.盐度改变了中华鲟的行为。对于9月龄中华鲟而言,在所有盐度组中鳃盖运动频率最低(74±2min-1)和最低的摆尾频率(51±3min-1)均出现在IW(isosmotic water)组。饲料转换率FCR(1.14-1.46)和特定生长率SGR(0.85-1.31)在所有盐度组中没有显著性差异(包括等渗点)(P FCR>0.05和PSGR>0.05)。各盐度下早中晚摄食节律方面,0.25,6.32和25psu组均没有差异,在12psu下晚20：00摄食量显著高于15：00,在19psu下,8：00摄食量显著多于20：00。日摄食方面,在与等渗盐度差异较大的盐度水体(0.27psu和25psu)中华鲟依靠更多的摄食,补充用于渗透压调节消耗的能量。
5.等渗(IW)和淡水组(FW)对比实验中,盐度影响中华鲟血清PRL激素和GH/IGF-1mRNA的表达。对于9月龄中华鲟而言,在适应期实验中,PRL在12.7psu时略高于至5.98,6.32和9.69psu,但在19.1和25psu又下降到低盐度先前水平。生长激素随盐度增加而增加,在25psu时最高。IGF-1的趋势和GH相反,IGF-1的水平最低发生在25psu。在适应后实验中,三种激素水平盐度随盐度变化没有表现出显著差异性。在等渗组和淡水组比较实验中,淡水中PRL水平显著高于等渗组(P<0.01)但GH和IGF-1水平在等渗组和淡水组则没有显著性差异。
6.酶含量方面,淡水中华鲟相比于等渗盐度下中华鲟,消耗较大的消化力为渗透压调节提供能量。对于9月龄中华鲟而言,中肠,后肠和肾的Na+/K+-ATP酶含量在IW组和FW(freshwater)组相似,而在鳃丝(P<0.05),前肠(P<0.01)和螺旋瓣(P<0.01)FW明显高于IW高约7%。前肠,螺旋瓣和胃在FW中脂肪酶含量比IW组高出10%(P<0.05)。此外,前肠ALT含量在FW显著高于IW(P<0.05)。在肝脏中淀粉酶的含量也是FW组高于IW组(P<0.05)。另外,在螺旋瓣的蛋白酶含量在FW高于IW约18.1%,而在中肠,则是在IW高于FW约13.2%(P<0.05)。
7.等渗(IW)和淡水组(FW)对比实验中机体组成成分的差异显著。对于9月龄中华鲟而言,在等渗(IW)和淡水组(FW)对比实验中,脂肪及其脂肪酸组成,粪便蛋白质、脂肪及其氨基酸和脂肪酸组成没有差异,但是肌肉钙离子组成差异显著。经过52天的实验,等渗组肌肉钙离子含量显著高于淡水组；全骨板和头部软骨钙、磷含量等渗组低于淡水组,但是镁离子含量则是等渗组高于淡水组。
8.盐度造成中华鲟卵质的差异。海水卵较之于淡水卵蛋白含量明显较高,脂肪含量则明显偏低。脂肪酸含量有极大地差异。两种卵均含17种氨基酸,包括10种鱼类必需氨基酸。总氨基酸量和各氨基酸没有差异。卵黄蛋白没有差异。磷元素、镁离子和维他命A在海水卵中含量明显多于淡水卵,维他命E则是淡水卵多于海水卵。钙离子和锌离子没有差异。在淡水卵中检测到镉离子,海水卵中则未检测到。
9.中华鲟血清渗透压和等渗点和年龄最相关,自幼鱼到性成熟,中华鲟等渗点为8.3-11.5psu。5月龄中华鲟的喜好盐度位于等渗盐度点,血清渗透压与年龄最相关。9月龄子一代中华鲟幼鱼在0.27,6,12,19和25psu盐度下适应后,水渗透压和盐度的关系通过线性拟合确定。二阶多项式拟合血清渗透压和盐度之间的关系,通过解方程求得等渗点为9.57±0.06psu。在该盐度下,血清渗透压等于咸水渗透压,鱼体无渗透压力。五月龄子二代中华鲟在淡水中氯细胞已经形成咸水适应机制,等渗点盐度为9.52psu。以“分级稀释建立高连续盐度梯度实现水生动物盐度选择实验的装置和方法”为新的实验体系,分析得出5月龄子二代中华鲟喜好盐度为8.5-9.5psu,这和5月龄子二代中华鲟等渗点9.52psu保持一致。对5月龄和9月龄中华鲟血清渗透压分析发现：盐水环境和淡水环境下渗透压没有显著差异,淡水环境下渗透压可作为盐水环境渗透压；等渗点下血清渗透压和与中华鲟相适应的淡水条件下血清渗透压在同一个水平。据此,该结果提供了迁地保护下大型中华鲟亚成体等渗点的简易计算方法。测得淡水环境下中华鲟血清渗透压,即作为等渗点盐度渗透压。将此渗透压带入盐度和渗透压关系式,即可计算出等渗点盐度。血清渗透压和年龄的二次多项式模拟最吻合,相关系数最高。因此认为年龄和中华鲟血清渗透压最相关：Y=278.061-11.3127X+0.6587X2,Y表示血清渗透压X表示年龄。对该二次多项式求导得出导数为0时,年龄为8.6年,即从出生到9龄时,中华鲟血清渗透压逐渐降低；从9龄开始,中华鲟血清开始随年龄增加而逐渐升高。同理等渗点也和年龄最为相关：Y=9.79431-0.35711X+0.0213X2,Y表示血清渗透压,X表示年龄。一阶求导得零点横坐标为8.38龄,即第9龄,这和渗透压结果一致。即从出生到9龄时,中华鲟等渗点逐渐降低；从9龄开始,中华鲟血清开始随年龄增加而逐渐升高。
总之,考虑到在等渗盐度组较之于淡水组明显较低的鳃盖运动频率,摆尾频率,鳃丝Na+/K+-ATPase含量以及所观察到的低脂肪酶,ALT和淀粉酶含量,显著降低的日摄食量和不变的特定生长率,肌肉中显著升高的Ca2+含量和增加的骨板和软骨中Mg2+含量,以及盐度对卵质产生的影响,等渗盐度是中华鲟的较理想生长环境盐度。此外,在淡水养殖中,Ca2+和Mg2+应该进行适当补偿。同时,中华鲟血清渗透压和等渗点盐度随年龄而变化,随着生长的进行,中华鲟的等渗养殖盐度需要进行调整。
Populations of A.sinensis, an anadromous fish species with Class-I state protection in China, grow in coastal waters (seawater) and enter into the Yangtze River (freshwater) to spawn after reaching sexual maturity, IUCN assessed as CR (Critically Endangered) status. A.sinensis subjecting to ex-situ conservation no longer live in brackish water, but spend their entire life cycle in freshwater (FW). The small size of the adults matured in freshwater was associated with decreased fecundity, resulting in more challenge to the conservation work. To restore the body size, this paper aimed to argue whether salinity affect sturgeon body size growth.5-month-old,9-month-old and12-15-year-old sturgeon were tested; Therefore, we constructed "specific collector combined count method in recirculating aquaculture system for feed conversion ratio determination" and the 'establishment of high grade continuous gradient salinity experiment apparatus and method for salinity preference in aquatic animal"; Explored the juvenile Chinese sturgeon salinity adaptation mechanisms; analyzed isosmotic points of5-month-old,9-month-old and12-15-year-old sturgeon; constructed relationships between fork length, body weight and age and serum osmolality, relationships between fork length, body weight and age and isosmotic points; Compared sturgeon behaviors, hormones, enzymes activities and tissue compositions between isosmotic environment and freshwater environment; Quantified fluorescence RT-PCR analysis of GHmRNA and IGF-1mRNA in tissues between isosmotic water (IW) and FW groups; Alalyzed the salinity and migration loss effects on the compositions of freshwater eggs and seawater eggs of Chinese sturgeon; The following conclusions were obtained:
1. Establish two methods:'specific collector combined count method in recirculating aquaculture systems for feed conversion ratio determination' and 'establishment of high grade continuous gradient dilution salinity experiment apparatus and method for salinity preference'; Count method could reduce physical, biochemical and mechanical error of quantifying uneaten feed collected from holding water. Simultaneously, it saved the disposals uneaten pellets after the uneaten pellets are collected, replaced siphoning by special designed collector and saved time and labors. In addition, the effects of the feces can be ignored. The former one was for calculation of feed conversion and provides a convenient feeding rhythm, meanwhile improved accuracy of FCR; the latter provide the equipment system and methodology for salinity preference experiments. The advantages lies in, on the one hand, the appetas promises long-time salinity gradients than former appatases. On the other hand, tested animals can swim free and rechoose their location.
2. Five-month-old A.sinensis reared respectively in FW (0.41psu) and brackish water (BW,12.5psu) for26days. The first10days was adaptation period, the later16days were post acclimated period. In the26th day, one of filaments both in left and right second branchial arch was sampled for scanning electron microscopy and transmission electron microscopy examination, respectively. The gill filaments appeared in the mucous cells, pavement cell and MRC on the surface of filament. In terms of MRCs, there were two subtypes of MRCs in FW, one was the ball-shaped MRC, which was higher above the neighboring pavement cells, and the other one was triangular MRC, which was triangle and flat, but it depressed between pavement cells. Only ball-shaped MRCs emerged in filament of BW-acclimated fish. Furthermore, the density and distribution of ball-shaped MRCs did not differ between freshwater and brackish group. It was also found that transitional cells, from triangle to ball-shaped, existed in FW-acclimated filament. Serum PRL, GH and IGF-1levels did not differ between FW and BW groups, which also reflected fish developeded BW adaptation mechanism in FW. Five-month-old A.sinensis had formd adapted MRCs (mitochondria-rich cell) to BW, whose isosmotic point was9.52psu. Taken "establishment of high grade continuous gradient dilution salinity experiment apparatus and method for salinity preference" as a new experimental system and revealed that five-month-old A.sinensis prefer7.5-8.5psu. This agreed to that isosmotic point of5-month-old was9.52psu. As salinity increased, food intake decreased significantly at30psu and fish ceased feeding at38psu compared to food intake at12.5psu.
3. For9-month-old sturgeon, isoionic points were analyzed in the same way as the isosmotic point was determined: K+:9.65±1.01psu; Cl-:6.82±0.37psu; Na+:9.40±0.58psu; Ca2+:1.62±0.71psu. The trend of Na+concentration as salinity increased was not pronounced (P=0.250), but Cl-tended to pronouncedly higher levels as salinity increased (P=0.021). NaCl contributed to total osmolality and the real NaCl percentages had the same levels in all salinity treatments (OsmNaCl=224.27±3.17-233.41±2.63mOsm/kg;93.39±2.22%-96.79±4.17%) as it did at the isosmotic point (OsmNaCl=227.45±8.04mOsm kg-1;89.93±4.09%). Serum Mg2+level were far lower than the water Mg2+level at any salinity treatment. Nevertheless, serum P was much higher than water at any salinity treatment. Serum Mg2+and K+levels were significant higher in as salinity increased.
4. Salinity changed the gill ventilation rate, tail beat frequency and daily feeding rhythm.
For9-month-old A.sinensis, the lowest gill ventilation rate (74±2min-1) and the lowest tail beat frequency (51±3min-1) were observed in IW. The feed intake had no difference at8:00,15:00and20:00in0.25,6.32and25psu groups. FCRs (1.14-1.46) and SGRs (0.85-1.31) were comparable at all salinity groups, including the isosmotic point (PFCR>0.05and PSGR>0.05), but FCR was smaller at25psu and IW groups. In long term salinity tolerance, the condition factor was markedly higher after22d for all15sturgeon compared to the values before the test (P<0.05). However, the feeding rhythm at20:00of12psu-acclimated fish was significantly more than15:00. In19psu group, the feed intake at8:00was significantly more than at20:00. The difference resulted from individual difference. In general, there was no difference in feeding rhythm of each feeding at any salinity group. But the daily feed intake in25and0.25psu groups were significantly higher than6.32and12psu groups.
5. For9-month-old A.sinensis, in the acclimation period, PRL increased slightly at12.7psu compared to5.98,6.32and9.69psu groups, but it declined again at19.1and25psu. In terms of GH, this hormone gradually increased as salinity increased and was the highest at25psu. The trend of IGF-1was contrary to that of GH, with the lowest IGF-1level observed at25psu. In long term salinity tolerance, the three hormone levels exhibited no significant differences among salinities. As time went on, no obvious changes were observed in any hormone. In IW group and FW group comparison test, the PRL level was significantly higher at the isosmotic point than in freshwater (P<0.01). However, GH and IGF-1exhibited no significant differences. For5-month-old A.sinensis, PRL, GH and IGF-1showed no trend in both acclimated period and post-acclimatedion. RT-PCR analysis of related expression of pituitary GH and IGF-1in different issues showed: pituitary GH mRNA and liver IGF-1mRNA expression in FW-acclimated group were significantly higher than IW-acclimated group; Muscle and kidney IGF-1mRNA levels had the same level between FW and IW groups.
6. The digestive enzymes and Na+/K+-ATPase consumed more energy instead of energy for growth group in FW than in IW group. The in the gill (P<0.05), foregut (P<0.01) and spiral valve (P<0.01) in FW were approximately7%higher than in IW. The lipase activities in the foregut, spiral valve and stomach in FW were10%higher than in the IW group (P<0.05), and the activities were similar in the other examined issues. In addition, ALT (alamine transaminase) activity in the foregut was significantly higher in FW than in IW, showing a difference of approximately8%compared to IW group (P<0.05). Amylase activity in the liver was also higher in FW than in IW (P<0.05), while these values were equal in other tissues. Furthermore, the protease activity in the spiral valve was18.1%lower in IW than in FW, while in the mid-gut, it was13.2%higher in IW than in FW (P<0.05).
7. Comparisons of feces protein and amino acids composition showed no difference, as well fatty acids. Muscle protein and amino acids had the similar levels between IW-acclimated fish and FW-acclimated fish. Both groups had the same level fat content (1.92±2.69) g and similar fatty acids compositions. However, in terms of minerals of muscle, Ca content in IW acclimated fish was significantly higher than in in FW-acclimated fish, P and Mg contents had the same levels. Different conditions about Ca, P, Mg contents were present in scutes and cartilage. In abdominal scute and cartilage, Ca levels were higher in FW-acclimated fish than IW-acclimated fish, but in lateral scute and back scute, Ca, Mg and P levels in IW group were higher than in FW group. In cartilage, Mg was not detected both in IW and IW groups. Overall, Ca and P levels in the total scutes and head cartilage in FW-acclimated fish were higher than in IW-acclimated fish, but Mg level in IW (2019.7±183.4mg/100g) was higher than in FW (1827.8±107.8mg/100g). The liver lipid contents (60.96±5.30mg/100g,45.29±18.34mg/100g), VA contents (31.22±9.41mg/100g,24.43±5.15mg/100g) had similar level in the two groups (FW was former, IW was latter). VE values were not detected in liver.
8. Eggs biochemical contents from wild origin and remature A.sinensis in freshwater were compared, which were related to induce spawning success. The results included dry weight, lipids, fatty acids, protein, amino acids, vitellin (Vn), phosphorus (P), vitamin A (VA) and vitamin E (VE) as well as the concentrations of relevant trace elements. The dry weights of the eggs did not differ between the sturgeons. The wild sturgeon eggs had significantly higher protein content but lower lipid content than the remature sturgeon eggs. There were multiple and significant differences in the fatty acids. Vn showed no difference. The P, magnesium (Mg) and VE contents in seawater eggs were significantly higher than in freshwater eggs. The calcium (Ca) and zinc (Zn) contents showed no difference between both batches. Cadmium (Cd) was only detected in one sample of freshwater eggs. These results helped to discriminate between remature (farmed) and wild origin Chinese sturgeon eggs. It is important to take measures to decrease lipid percentages and to increase protein contents of eggs for remature Chinese sturgeon or even sturgeon matured in freshwater.
9. A.sinensis were acclimated in0.27,6,12,19and25psu, relationship between water osmolality and salinity was determined by linear fitting. Second-order polynomial fitting were used to assess relationship between serum osmolality and salinity. Isosmotic point was got (9.57±0.06psu) through solution of the two equations. At isosmotic salinity environment, serum osmolality equaled to environment osmolality, sturgeon faced no osmotic pressure. The results of analyzing serum osmolality of5-month and9-month-old A.sinensis showed: there were no significant differences of serum osmolality under FW or BW, including IW. Accordingly, the results provide method to assess isosmotic point of larger ex-situ conservation subadult sturgeon in FW. Regard as the value of serum osmolality in FW as serum osmolality in BW. Serum osmolality in FW-acclimated fish had the same level as serum osmolality in IW-acclimated fish. Simultaneously, we could also take serum osmolality in FW-acclimated fish as serum osmolality in IW-acclimated fish. Take the value of serum osmolality in FW group into the equation of relationship between salinity and water osmolality, and then get the salinity i.e. isosmotic point. The order-two polynomial fitted best in relationship between age and serum osmolality, the correlation coefficients was highest. Therefore age was considered as the most relevant factor to serum osmolality:Y=278.061-11.3127x+0.6587X^2. Derived the equation and the X corresponding to value of derivative equled to zero was8.6years, i.e. from birth to age9the serum osmolality decreased and serum gradually began to increase with age since9-year-old. Similarly, isosmotic osmolality and age were most relevant:Y=9.79431-0.35711X+0.0213X2. Derived the equation and the X corresponding to value of derivative (0) was8.6years. This was similar to serum osmolality equation, i.e. from birth to age9-years, the Chinese sturgeon isosmotic point decreases, from9years on, the isosmotic point began to gradually increase with age.
In conclusion, the Chinese sturgeon developed seawater-entry readiness in freshwater and the fish preferred isosmotic salinity compared to other salinities. Considering the obviously lower gill ventilation rate, tail beat frequency, Na+/K+-ATPase activity in the gills as well as the lower lipase, ALT and amylase activities observed, lower daily feeding rhythm but same SGR levels, significantly higher Ca content in muscle and increased Mg content in scutes and cartilage in IW compared to FW a salinity corresponding to the IW is ideal for the growth of naive A. sinensis. A. sinensis is suggested to rear in8.3-11.5psu. Besides, Ca2+and Mg2+should be compensated in FW culture. Meanwhile, the plasma osmolality and isosmotic point may vary with the size and age of fish, and salinity may need to be adjusted as naive A. sinensis grow.
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