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克氏原螯虾繁育机制及养殖生态学研究
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摘要
克氏原螯虾(Procambarus clarkii)是世界性淡水经济虾类,原产地北美洲南部,上世纪三十年代引入我国,目前己广泛分布于江苏、湖北、安徽和江西等省市,成为我国重要的水产经济虾类。近年来该虾经济价值飞涨,全国各地掀起了一股龙虾养殖的热潮,是目前我国水产业中发展最为迅速、最具特色、最有潜力的养殖品种之一。克氏原螯虾的人工养殖,虽然发展势头迅猛,但养殖技术仍处于初级阶段,苗种、成虾规格及产量成为限制克氏原螯虾产业进一步发展的瓶颈。因此本文对克氏原螯虾的繁殖生物学及养殖生态学进行研究,以期为解决限制该产业进一步规模化发展的关键问题提供思路。主要研究结果如下:
     1.冬季加温刺激克氏原螯虾抱卵繁殖
     本文通过在秋季收集雌虾,在冬季利用加温刺激越冬克氏原螯虾抱卵繁殖,研究了克氏原螯虾冬季抱卵、孵化及幼虾的生长情况,为克氏原螯虾规模化育苗提供了一个新思路。结果显示:冬季提高水温可以刺激克氏原螯虾雌虾抱卵。经过38d的加温刺激,在26℃温度下陆续观察到有虾抱卵,18℃以下未观察到抱卵虾,但该水温能促进雌虾性腺发育。雌虾经加温刺激后所抱卵在18℃,21℃和26℃温度下均能正常孵化出幼虾。18℃,21℃和26℃下孵化时间分别为(39.67±3.06)d,(28.33±2.52)d和(15.00±2.00)d,孵化所需时间随着温度的提升而减少(P<0.05)。孵化的幼虾经45d养殖,18℃,21℃和26℃下体长分别为(1.90±0.25)cm,(2.73±0.38)cm和(3.78±0.70)cm,体重分别为(0.223±0.100)g,(0.688±0.331)g和(2.145±1.272)g。18℃,21℃水温下的幼虾在此生长阶段为非异速生长(b=3),26℃水温下的幼虾为异速生长(b>3)。在3种温度下,幼体生长速度随温度的升高明显加快(P<0.05)。
     2.克氏原螯虾卵离体孵化研究
     本试验对不同时期剥离的虾卵孵化及剥离后不同孵化方式对孵化率的影响作了研究,结果显示:静水孵化,抱卵当天、72h、144h、216h剥离的卵的孵化率都很低,无显著差异(P>0.05),288h剥离的卵孵化率较高,达到65.00±10.00%,与抱卵当天、72h、144h、216h剥离的卵孵化率差异极显著(P<0.01)。充气孵化,抱卵当天、72h、144h剥离的卵孵化率均较低,无显著差异(P>0.05),216h剥离的卵的孵化率明显升高(P<0.05),达到46.67±7.64%,288h剥离的卵的孵化率达到75.00±13.23%,与抱卵当天、72h、144h剥离的卵孵化率差异极显著(P<0.01),与216h剥离的卵的孵化率差异显著(P<0.05)。充气孵化的卵的孵化率均高于静水孵化,0,72h剥离的差异显著(P<0.05),216h剥离的差异极显著(P<0.01),144h,288h剥离的无显著差异。并且充气孵化,216h剥离的卵即开始出现孵化率的显著提升,静水孵化在288h剥离的卵才开始出现孵化率的显著提升。
     3.雌虾对幼体生长及成活率的影响
     克氏原螯虾有抱卵孵化的习性,即使在卵孵化后,能独立游动的虾仍不时会吸附在母虾的腹部,因此本试验研究了雌虾对幼体生长及成活率的影响。结果显示:母体对仔虾成活率有显著影响。有母虾的试验组成活率仅为34.00±11.14%,显著低于没有母虾的试验组(P<0.01)。稚虾阶段剥离的仔虾的成活率与幼虾阶段剥离的仔虾的成活率无显著差异(P>0.05),分别为66.66±4.32%,74.00±11.14%。经培养后,有母虾的试验组,仔虾的体长体重分别为2.11±0.32cm,0.31±0.13g;幼虾阶段剥离的仔虾的体长体重分别为2.09±0.39cm,0.31±0.21g;稚虾阶段剥离的仔虾的体长体重分别为1.67±0.27cm,0.15±0.07g。有母虾的试验组与幼虾阶段剥离的试验组差异不显著(P>0.05),但这两组的体长体重显著高于稚虾阶段剥离的仔虾的体长体重(P<0.01)。有母虾组、稚虾剥离组和幼虾剥离组的虾的体长体重方程分别为y=0.030x2.991,R2=0.970,y=0.029x3.030,R2=0.947,y=0.028x3.083,R2=0.965,均为等速生长(b≈3)。
     4.养殖条件下克氏原螯虾体长体重和螯长螯宽的关系
     目前,养殖条件下,克氏原螯虾的体长体重及螯长螯重关系尚无详细研究,因此本试验的目的就是确定养殖条件下的克氏原螯虾体长体重和螯长螯重的关系,并分析其两性差异。测量了678只虾(雄性337只,雌性341只)的体长和体重,并建立了关系式。雄虾、雌虾和所有个体的体长体重的方程分别为:WWT=0.014×LB3.603,WWT=0.020×LB3.350和WWT=0.017×LB3.467,均为异速增长(b>3)。雌雄虾的体长体重关系有显著差异(P<0.001)。所有个体的左螯肢和右螯肢的螯长(Lch)、螯肢长(Lcp)和螯宽(Wch)均无显著差异(p>0.05)。我们选择左螯肢来建立螯长螯宽的关系式并确定幂函数为最佳方程式。雄虾雌虾的螯长螯宽方程分别为WCh=3.083×Lch0.847andWCh=0.284×LCh1.232,雌雄差异显著,雄虾的螯宽显著大于雌虾(P<0.001)。678只虾的体长范围为1.8-11.1cm,尾长范围为1.1-6.1cm,体重范围为0.18-83.43g。雄虾比雌虾重,雌虾的尾长要比雄虾长,差异极显著(P<0.001)。相同体长的雄虾的螯长、螯肢长和螯宽明显比雌虾长(P<0.001)。无螯肢的雄虾和雌虾的体长体重关系分别为WBw=0.074×上B2638andWBw=0.033×LB3.023,相同体长的雌雄虾,体重无显著差异(P>0.05)。
     5.不同性别的克氏原螯虾幼虾生长的研究
     淡水螯虾普遍具有两性差异,而克氏原螯虾的两性差异主要体现在雄虾比雌虾更为巨大的螯肢。为确定克氏原螯虾幼虾第二性征何时出现,本试验研究雌雄幼虾的生长情况。结果显示:经75天培养雌雄幼虾的增重率和增长率分别为2577.00±407.33%,171.75±10.25%;2982.40±605.97%,177.62±17.32%,差异不显著(P>0.05)。雌雄幼虾在不同阶段的体长体重均无显著差异(P>0.05)。以体长为协变量,体重为变量进行协方差分析,0d,15d,30d,45d雌雄体重无显著差异(P>0.05),第60,75天差异显著(P<0.05)。从生长情况来看,试验第60d,雌雄虾的体长分别为5.54±0.38cm和5.61±0.43cm,因此可以认为在幼虾体长达到5.5cm左右时,两性差异开始显现。
     6.不同性比对克氏原螯虾生长的影响
     为弄清更具攻击性的雄虾是否会对克氏原螯虾的成活率及生长速度有所影响,本试验进行了不同性比对克氏原螯虾生长的影响研究。结果显示:全雌,全雄及雌雄组的幼虾的成活率、增重率和增长率分别为54.17±7.22%,4996.00±1857.42%,258.33±39.83%;45.83±14.43%,4075.70±1620.98%,228.67±39.50%;37.5±12.50%,4972.70±479.65%,254.00±5.00%。全雌组在成活率、增重率和增长率上均好于其他两组,但差异并不显著(P>0.05)。全雌,全雄及雌雄组的成虾的成活率、增重率和增长率分别为91.67±8.34%,8.33±0.97%,1.60±0.25%;83133±8.34%,1.82±0.91%,1.41±0.20%;86.11±4.82%,24.84±2.06%,5.94±0.53%。各组成活率差异不显著p>0.05);雌雄组的增重率>全雌组>全雄组,差异极显著(P<0.01);增长率全雌组与全雄组差异不显著(P>0.05),雌雄组明显大于全雌组与全雄组(P<0.01)。
Procambarus clarkii is world freshwater economic shrimp, originates from south of North America. It was introduced to China in the1930s, and its distribution has expanded to Jiangsu, Hubei and Jiangxi Province. P. clarkii has become one of the most important freshwater products in China. In recent years, the value of P. clarkii is soaring and P. clarkii farming is booming throughout China. Its growing has become one of the fastest, most characteristical and potential variety of cultivation in Chinese aquaculture. But the technology of cultivation P. clarkii is poor, seeding, yields and specifications are the bottlenecks of the crayfish industry. Therefore, the reproductive biology and cultivation ecology was investigated in this article, in order to solve the limitation of the crayfish industry. The main results were present as follows:
     1. Warming up to stimulate Procambarus clarkii to breed in winter
     Warming up to stimulate Procambarus clarkii to breed in winter was researched. And the effect of temperature on hatching and larvae development of Procambarus clarkii was investigated. The results showed that the female crayfish could breed by warming up in winter. The female crayfish was oogenetic in the water tank of26℃temperature after38days. The female crayfish wasn't oogenetic in the water tank of18℃, but the ovary developed. The eggs carried by female crayfish were fertilized and could be hatching normally. The hatching time was (39.67±3.06)d,(28.33±2.52)d and (15.00±2.00)d respectively in the tank of18℃,21℃and26℃water temperature. During45d breeding, the body length of larvae was (1.90±0.25)cm,(2.73±0.38)cm and (3.78±0.70)cm respectively in the tank of18℃,21℃and26℃water temperature, and the weight was (0.223±0.100)g,(0.688±0.331)g and (2.145±1.272)g respectively. Larvae exhibited positive allometric growth (b>3) at26℃and showed isometric growth at18℃and21℃(b=3).The hatching speed and larvae growth were positively correlated with temperature in a certain range.
     2. In vitro incubation of P. clarkii ovum
     The effect of hatchability on different period ovum in vitro incubation and different incubation ways was researched. The results showed that the hatchability of ovum separate from crayfish after0h,72h,144h and216h were very low in standing water, no significant difference among them(P>0.05). In this way, the hatchability of ovum separate from crayfish after288h reached65.00±10.00%, was significant higher(P<0.01). The hatchability of ovum separate from crayfish after0h,72h and 144h were very low in inflating water, no significant difference among them(P>0.05). In this way, the hatchability of ovum separate from crayfish after216h reached46.67±7.64%, was significant higher(P<0.05). And the hatchability of ovum separate from crayfish after288h reached75.00±13.23%, was highest (P<0.05). The hatchability of in vitro ovum in inflating water was higher than in standing water.
     3. The effect of female crayfish on larval growth and survival rate.
     The female P. clarkii carry ovum on their abdomen. Even after the ovum hatching, the larval can be back to the crayfish's abdomen occasionally. This experiment was conducted to study the effect of female crayfish on larval growth and survival rate. The results showed that the female crayfish was significant influence the survival rate of larval. The survival rate of larval with mother is34.00±11.14%, significantly lower than which without mother. The survival rate of period Ⅰ and period Ⅱ separated from mother were66.66±4.32%and74.00±11.14%respectively, had no significantly difference (P>0.05). After cultivation, the body length and weight of the crayfish with mother and period Ⅱ separated from mother were2.11±0.32cm,0.31±0.13g, and2.09±0.39cm,0.31±0.21g respectively, has no significant difference (P>0.05). But the body length and weight of these two groups were significant higher than the group of crayfish period Ⅰ separated from mother, which were1.67±0.27cm,0.15±0.07g respectively. The length-weight relationships for the crayfish with mother, period Ⅰ and period Ⅱ separated from mother were y=0.030x2.991, R2=0.970, y=0.029x3.030, R2=0.947and y0.028x3.083, R2=0.965respectively, all growth were isometric(b≈3).
     4. Length-weight and chelae length-width relationships of the crayfish Procambarus clarkii under culture conditions
     Length-weight (body length; LB vs. wet weight; WWT) and chelae length-width (LCh vs. WCh) relationships were determined for male and female crayfish (Procambarus clarkii) under culture conditions. The length-weight relationships for males, females, and total individuals were described as WWT=0.014×LB3.603,WWT=0.020×LB3.350, and WWT=0.017×LB3.467, respectively. This relationship was significantly different (P<0.001) between males and females. Compared to females, males were heavier and their tail length was shorter (P<0.001). No statistical difference was observed in mean LCh, cheliped length (LCp), and WCh between the right and left sides within a gender (P>0.05), but significant differences were detected between males and females (P<0.001). The regressions for LCh vs. WCh for males and females were WCh=3.083×LCh0.847and WCh=0.284×LCh1.232, respectively. This regression differed significantly between males and females; males' chelae were wider than those of females (P<0.001). Lch,LCp, and Wch were highly related to LB;males had longer Lch and LCp and wider Wch than females. Individuals without chelipeds were measured to determine the length and body weight(WBW) relationship. The LB-WBW relationships for males and females were described as=0.074×LB2.638and WBW=0.033×LB3.023, respectively. Females exhibited positive allometric growth (b>3), and males showed negative allometric growth (b<3). No statistical difference was observed in mean WBW between males and females when LB was used as a covariate (P>0.05).
     5. The growth of juvenile crayfish of different gender
     Sexual dimorphism appears to be common in freshwater crayfish. Particularly obvious sexual dimorphism in crayfish is due largely to the disproportionately rapid growth of chelae in males compared with females. This experiment was in order to determine when the secondary sexual characteristics appeared. The results showed that the weight and length growth rate of female and male juvenile crayfish were2577.00±407.33%,171.75±10.25%and2982.40±605.97%,177.62±17.32%respectively after75d culture, has no significant difference(P>0.05). The body length and weight of male and female juvenile crayfish at different stages showed no significant difference(P>0.05). The weight of male and female has no significant difference at Od,15d,30d and45d(P>0.05), but·showed significant difference at60d and75d(P<0.05), when body length was used as a covariate. The length of female and male at60d were5.54±0.38cm and5.61±0.43cm respectively, so the secondary sexual characteristics emerged when the body length of crayfish reached around5.5cm.
     6. The effect of sex ratio on P. clarkii growth
     The effect of sex ratio on P. clarkii growth were researched in order to understand if the male shrimps affect the growth and survival of P. clarkii. The results showed that survival rate, weight and length growth rate of female, male and two genders juvenile crayfish groups were54.17±7.22%,4996.00±1857.42%,258.33±39.83%;45.83±14.43%,4075.70±1620.98%,228.67±39.50%and37.5±12.50%,4972.70±479.65%,254.00±5.00%respectively, had no significant difference among them(P>0.05). The survival rate, weight and length growth rate of female male and two genders adult crayfish groups were91.67±8.34%,8.33±0.97%, 1.60±0.25%;83.33±8.34%,1.82±0.91%,1.41±0.20%and86.11±4.82%,24.84±2.06%,5.94±0.53%respectively. The survival rate had no significant difference(P>0.05). But the growth rate of weight had significant difference(P<0.01), two genders>female>male. The growth rate of length had no significant difference between female and male(P>0.05), but both were significant lower than the group of two genders(P<0.01).
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