摘要
大菱鲆Scophthalmus maximus是目前我国工厂化养殖海水鱼产量最大的鱼种,也是支撑我国海水养殖业稳定发展的主要养殖种,产值巨大。但大菱鲆养殖业面临着地下海水资源枯竭、良种缺乏、养殖成本上升、病害频发等问题,制约着大菱鲆养殖产业的健康、可持续发展。本文围绕大菱鲆养殖产业缺乏的良种问题,以培育耐高温品系和快速生长品系为目标进行家系选育。在耐高温研究上,通过对我国养殖群体不同发育期的耐热性和仔、稚、幼鱼的热休克hsp70蛋白的RNA-cDNA表达规律研究,摸清了其耐高温能力的本底数据,通过选择耐高温亲鱼群体、胚胎热激筛选及比对不同家系的耐高温养殖试验结果,初步获得了性状优良的耐高温品系。通过对家系的存活率、白化率、畸形率和生长性能的比较,以及估算生长相关的幼鱼数量性状遗传力,对大菱鲆生长性状选育进行了研究。主要研究结果如下:
1、引进了法国、丹麦、西班牙群体亲鱼,收集了国内莱州、海阳、日照养殖群体,并获得了耐高温亲鱼群体,共同组成大菱鲆育种基础群体。形态分析表明法国与日照群体形态具有较大差异,日照、莱州、海阳群体形态趋同,丹麦群体具高体型特征,同其它5个群体形态差异较大。主成分分析的第一、二、三主成分贡献率分别为47.07%、28.10%、15.45%,三者累积贡献率为90.62%。
2、使用14对微卫星标记进行遗传多样性分析表明,6个亲本群体的等位基因数(A)为3~8,平均等位基因数为5.5,有效等位基因数(Ne)为1.07~5.54,平均有效等位基因数为3.03,各位点的杂合度观测度(Ho)为0.440~1.000,杂合度期望度(He)为0.509~0.824。各群体之间的无偏倚杂合度期望值由小到大依次为西班牙群体、法国群体、莱州群体、海阳群体、丹麦群体和日照群体;遗传多样性差异不显著,而国内累代养殖群体间遗传多样性差异较小,同进口群体间存在一定差异,但不显著(P>0.05)。群体间平均遗传分化指数(Fst)为0.11088,各群体间存在中度遗传分化。NJ法聚类结果:莱州和海阳群体首先聚为一类,二者聚类后复与法国群体聚为一类,表明这3个群体遗传较为接近;这3个群体随后逐次与西班牙群体、日照群体和丹麦群体聚类。结合Hardy-Weinberg平衡和遗传偏离参数(d),6个群体都不同程度的偏离平衡。适合作为良种选育的基础群体。
3、不同发育期大菱鲆胚胎对高温热激的耐受性存在显著差异,原肠期和细胞分裂期的耐受能力显著较低;初步确定器官形成期、尾芽期、出膜前期为适宜热激操作的时期。探明了大菱鲆胚胎期、卵黄囊期、前弯曲期和弯曲期仔鱼、稚鱼期经受2h热刺激的亚致死温度为24℃、23℃、<22℃、28℃、28℃,半致死温度为26℃、28℃、22℃、29℃、29℃,致死温度为30℃、29℃、27℃、30℃、30℃。耐热能力由弱到强依次为10日龄、胚胎、0日龄、25日龄、50日龄,总体上呈现随生长发育逐渐增强的趋势,10日龄可能为大菱鲆热刺激耐受力低谷。
4、半定量RT-PCR测定大菱鲆仔、稚、幼鱼高温热激后热休克蛋白hsp70基因表达,表明相同热激温度后的不同取样时间的表达量,在热激后1h最高、24h陡然降低、48h出现小幅度回升;hsp70表达量明显呈现幼鱼>稚鱼>仔鱼的趋势,表达量随热激温度升高而上升,且与个体发育时间显著呈正相关。
5、使用高温海水养殖淘汰大菱鲆成鱼,自150尾候选成鱼中筛选出耐高温个体39尾(筛选强度为26.0%),组成耐高温亲鱼群体。以验证选择育种“胚胎期选择法”为目的,使用短时间高温刺激的方法,比较和分析了卵裂期、原肠期、器官形成期、尾芽期和孵化前期5个胚胎发育时期的高温耐受能力:5个胚胎发育期的高温耐受性存在显著差异;随着热激温度的提高、热激时长的延长,胚胎的正常仔鱼率下降,并呈现较强的相关性和临界性;器官形成期、尾芽期、出膜前期适宜进行热激操作。通过家系幼鱼热刺激和高温水养殖试验结果,筛选出可能具有耐高温特性的亲鱼11尾和家系18个,成为核心育种群体的重要组成部分。
6、家系间的生长性状差异较大,前、后期的全重生长上表现出较显著的一致性。家系幼鱼存活率总体较低,不同家系之间差异极大,存活率可能主要由亲本的精、卵质量决定。共观察到较明显的幼鱼发育畸形现象9个,将各家系的畸形性状同亲本比对,发现白化、腹部畸形、脊柱弯曲、下颌上屈同亲本可能存在一定的相关性,鳃盖缺失、转眼不彻底、苹果臀、逆向转眼同亲本可能不存在明显的相关。半同胞家系组合的白化率差别较大,认为白化可能受到母系遗传或卵子质量的影响。多数家系的白化与正常幼鱼的生长和可比性状差异不显著(P>0.05)。白化与原色幼鱼的生长和身体形态整体上无显著差异。筛选出速生候选家系1个,高体型候选家系3个。
7、估算了大菱鲆幼鱼可量性状的遗传力。110日龄幼鱼全重、全长、体长、头长、上颌长、眼间距、总高、体高、尾柄高、尾柄长性状的遗传力分别为0.219、0.461、0.366、0.341、0.717、0.669、0.317、0.207、0.511和0.216;185日龄幼鱼全重、全长、体长、头长、总高、体高的遗传力分别为0.383、0.269、0.286、0.322、0.319、0.352。依据父系和母系半同胞方差组分估计的110dah的全长、总高、上颌长、体高和185dah的全部性状的遗传力达到显著水平;依据全同胞方差组分估计的110dah和185dah所有性状的遗传力都达到极显著水平。本研究中的多数性状都属中度遗传力,具有较大的选育潜力。
Turbot Scophthalmus maximus is a commercially valuable and tasty flatfish withhighest farming yield in northern China, it has been the main farmed species inmarine fish culture industry in northern China since2002. However, the sustainabledevelopment of turbot farming industry was constrained by more and more arisingproblems such as depletion of underground sea water, germplasm degeneration,ascended culture cost and disease etc. Thus, works on selective breeding should becarried out urgently to foster strains with superior traits especially with fast growthtraits and thermotolerant traits, and so on. In the present study, we aim at selectivebreeding of thermotolerant strains and its possible mechanism was investigated bystudying the expressions pattern of hsp70gene through temperature stimulation toembryos at different developmental stages. In addition, we obtained a thermotolerantstain of turbot by selecting high-temperature tolerant broodstocks, high temperaturetolerance of embryos and families culture under high temperature. Meanwhile, thesurvival rate, albino rate, malformation rate and growth related traits recorded fromdifferent families was compared and analyzed. Also, the growth traits of differentfamilies of turbot were studied.
1. We constructed the basic reference breeding population of turbot byintroducing France strain, Danmark strain and Spanish strain combined with theChinese culture populations from Lai-zhou, Hai-yang and Ri-zhao. Thethermotolerant population was also selected. The morphological analysis showed thatthere was notable difference between France and Ri-zhao strain, Ri-zhao andHai-yang strains were similar in morphology, and Denmark strain has the best bodyheight, which was obviously different from the other strains. The basic referencebreeding population was suitable for selective breeding study. Using principalcomponents analysis, three principal components were constructed, the contribution rate from the first, second and third principal component was47.07%,28.10%and15.45%, respectively.
2. Genetic diversity within and between6turbot broodstocks strains wereassessed by using14microsatellite loci. The results showed that the number of alleles(A) and effective alleles (Ne) ranged from3to8and1.07to5.54, respectively. Theobserved heterozygosity value (H0) and unbiased expected heterozygosity value (He)ranged from0.440to1.000and0.509to0.824, respectively. The order of the Hevalues for the6broodstocks strains was: Spanish 3. The thermotolarant ability of turbot embryos to heat stimulates notably variedat different development stages. Embryos at gastrula stage and cell division stagewere eligible to heat-harden treatment, whereas embryos at the organic forming stage,tail-bud stage and hatching stages were suitable for heat-harden treatment. Thesub-lethal temperature,50%mortality (LT50) temperature, lethal temperature forembryo, yolk-sac larvae (0days after hatching, dah), preflexion larvae (10dah),flexion larvae (25dah) and juveniles (50dah) of turbot which experienced2hheat-harden treatment were examined in this thesis. The order of thermotolerantability of such5developmental stages was:10dah<embryo<0dah<25dah<50dah,which showed a trend of gradual increase with development generally.10dah stagehave the most fragile response to heat treatment.
4. The expressions pattern of hsp70mRNA of turbot larvae and juveniles whichexperienced2h heat treatment were determined using semi-quantitative real-time PCR.After the same temperature drop of the same developmental stage, expression level ofhsp70mRNA was highest at1h post heat treatment, then decreased sharply at24h,and rebounded slightly at48h. Based on the expression level of hsp70, the young fish expressed highest level of hsp70, followed by juveniles and larvae. The hightemperature could stimulate higher expression level of hsp70in turbot.
5. A total of39individual parent turbot were selected by culturing the adult fishunder high temperature condition, then the thermotolerant spawner population wasconstructed. In order to test the mothed of sellection at embryo stage, short timeheat-harden treatment was used to compare the high temperature tolerance of embryosat five different development stages: cell division stage, gastrula stage, organicforming stage, tail-bud stage and hatching stage. The results showed that thedifference of high temperature tolerance between5embryo developmental stages wassignificant. Tolerance of heat stimulates of embryos at gastrula stage and cell divisionstage were weak than that of embryos at the other3stages. The rate of normal larvaewas declined with the elevated temperature and prolonged heat-harden time, andshowed a strong correlation. The three embryonic development stages which suitablefor heat shock operation were organic forming, tail-bud and hatching stage. A total of11thermotolerant spawner fish strains and18full-sub families were selected throughheat shock and high temperature culture test. They became important part of the corebreeding population of turbot.
6. The difference of growth related traits among full-sib families was notable. Itshowed a remarkable consistency between the total weight growth of110dah and185dah youngfish. The survival rate of youngfish from different full-sub families wascontrasting, and it was low overall. The survival rate may be decided by the quality ofembryo and sperm to a large extent. There were9abnormal traits observed in culturedyoungfish. By the comparison between parent fish and family with abnormalmorphology, some obvious morphological deformation were observed: albino,abdominal deformity, spine curvature and low-jaw upward, all these deformationtraits may be associated with parent, but gill cover missed, eye transformation failure,apple-buttock and inversus metamorphosis may be not associated with parents.Significant difference of albino rates among half-sibs families were observed, whichmight be caused by maternal inheritance and ova quality. There were no significantdifferences in growth rates and body sharp between albino and normal juveniles in the present study. Additionally, a fast growing family and three high-body type familieswere selected as candidates for selective breeding program from all the F2families.
7. The heritabilities of10growth related traits were estimated by intra-groupcorrelation of full-sibs of turbot. The heritability of110dah youngfish in total weight,total length, body length, head length, length of up-jaw, length between two eyes, totalheight, body height, caudal peduncle height and caudal peduncle length were0.219,0.461,0.366,0.341,0.717,0.669,0.317,0.207,0.511and0.216, respectively. Theheritability of185dah youngfish in total weight, total length, body length, head length,total height and body height were0.383,0.269,0.286,0.322,0.319and0.352,respectively. The difference was significant in components variance of full-sibs withinsires and dams. From components of variance of full-sibs within sires and dams wereprecise and unbiased for the quantitative character traits. The present study shows thatthe heritability in the traits of turbot is moderate.
引文
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