本文選題：大口黑鱸(Micropterus + salmoides)�。� 參考：《上海海洋大學(xué)》2016年碩士論文

【摘要】：大口黑鱸(Micropterus salmoides)是鱸形目(Perciformes)中的一種,在我國的養(yǎng)殖品種里,也是養(yǎng)殖面積較大的魚類之一。大口黑鱸是典型的強(qiáng)肉食性魚類,其傳統(tǒng)的養(yǎng)殖方式主要是投喂冰鮮下雜魚,這不僅使其養(yǎng)殖業(yè)受海洋漁業(yè)資源不足的限制,還存在冰鮮魚體引入病原體污染水體的風(fēng)險(xiǎn),導(dǎo)致難以開展工廠化養(yǎng)殖和無公害水產(chǎn)品的生產(chǎn)。近年來,使用人工配合飼料投喂大口黑鱸取得了一定的成效,但人工配合飼料養(yǎng)殖大口黑鱸在中后期生長速度遲緩的現(xiàn)象普遍發(fā)生,這也嚴(yán)重限制到了我國大口黑鱸養(yǎng)殖業(yè)的進(jìn)一步發(fā)展。培育出能較好攝食并利用人工配合飼料、具有優(yōu)良生長性狀的大口黑鱸新品系,是現(xiàn)階段大口黑鱸選育工作的重要目標(biāo)。分子標(biāo)記輔助育種技術(shù)是魚類遺傳改良的重要手段,借助分子標(biāo)記對候選基因和目標(biāo)性狀進(jìn)行關(guān)聯(lián)和選擇分析,能很大程度提高相關(guān)經(jīng)濟(jì)性狀選擇的效率,并可有效縮短育種時(shí)間。根據(jù)已有研究報(bào)道,魚類的攝食受多種內(nèi)在和外在因素的影響,其中食欲狀況是魚類主要的內(nèi)在影響因素,當(dāng)養(yǎng)殖魚類處于食欲興奮狀態(tài)時(shí),它們的飼料攝入量以及利用率都會(huì)相應(yīng)的得到改善;魚類攝食的調(diào)控主要通過大腦和神經(jīng)外周信號(hào)的相互作用完成,食欲相關(guān)基因是參與魚類攝食調(diào)控的主效基因,該類基因的突變會(huì)對生長性狀產(chǎn)生影響;本研究選取Ghrelin、PACAP和NPY等攝食相關(guān)基因作為大口黑鱸生長性狀相關(guān)的候選基因,分析其單核苷酸多態(tài)性(single nucleotide polymorphism,SNP)與生長性狀的關(guān)聯(lián)性,以期為大口黑鱸的分子標(biāo)記輔助育種提供有效的標(biāo)記,促進(jìn)食性改良大口黑鱸新品系的培育。主要研究內(nèi)容如下:1.大口黑鱸Ghrelin基因SNPs的篩選及與生長性狀關(guān)聯(lián)性分析Ghrelin是脊椎動(dòng)物的一種腦腸肽,有促進(jìn)攝食功能,并能促進(jìn)生長激素(GH)釋放,參與能量平衡調(diào)控和糖類代謝。為探索ghrelin基因多態(tài)性與大口黑鱸生長性狀的相關(guān)性,實(shí)驗(yàn)針對大口黑鱸ghrelin基因的啟動(dòng)子序列,采用直接測序法獲得了2個(gè)SNPs位點(diǎn):S1(A-642C)和S2(A-639C)。選取327尾同批繁殖、同塘養(yǎng)殖的大口黑鱸,然后采用Sna Pshot方法進(jìn)行SNPs位點(diǎn)檢測和分型。結(jié)果顯示實(shí)驗(yàn)群體在ghrelin基因兩個(gè)SNPs位點(diǎn)上均處于哈溫平衡,S1和S2共組成了5種雙倍型(D1、D2、D3、D4和D5)。關(guān)聯(lián)分析的結(jié)果表明,S1位點(diǎn)上的AC基因型和S2位點(diǎn)上的AA基因型分別是兩個(gè)突變位點(diǎn)的優(yōu)勢基因型,雙倍型分析表明,雙倍型D1在各生長性狀上具有顯著優(yōu)勢。2.大口黑鱸PACAP基因SNPs的篩選及與生長性狀關(guān)聯(lián)分析垂體腺苷酸環(huán)化激酶多肽(pituitary adenylate cyclase-activating polypeptide,PACAP)是一種能夠促進(jìn)生長激素、促性腺激素和催乳素分泌的多效激素,屬于胰高血糖素/分泌肽家族的成員。為探索PACAP基因多態(tài)性對大口黑鱸生長性狀的影響,使用直接測序法在PACAP基因上篩選到一個(gè)SNP位點(diǎn):A-2282C,存在3種基因型:AA、AC和CC,其基因型頻率分別為0.034,0.645和0.321,A等位基因頻率為0.3563,C等位基因頻率為0.6437。選取同批繁殖、同塘養(yǎng)殖的327尾大口黑鱸用Sna Pshot方法進(jìn)行SNPs位點(diǎn)檢測和分型,構(gòu)建最小二乘分析模型,分析突變位點(diǎn)基因型與生長性狀的相關(guān)性。卡方檢測結(jié)果顯示,哈溫-平衡常數(shù)為0.0615,即該位點(diǎn)在所檢測大口黑鱸群體中基本處于Hardy-Weinbery平衡狀態(tài),有效等位基因數(shù)為1.8465,期望雜合度和觀測雜合度分別為0.5409和0.6442。方差分析結(jié)果顯示:AC基因型群體在各生長性狀上的平均表型值比AA和CC基因型都要高出很多;且AC基因型群體在體質(zhì)量、體高和全長上顯著高于AA和CC基因型群體(P0.05)。AA基因型群體的平均體質(zhì)量、體高和全長相對于CC基因型群體分別高出了18.1%、5.8%和5.4%,但AA型群體和CC群體在各生長性狀上沒有顯著差異(P0.05)。推測該位點(diǎn)可能是影響大口黑鱸生長性狀的的主效數(shù)量性狀核苷酸(quantitative trait nucleotides,QTN)或與之緊密連鎖。3.大口黑鱸北方亞種和佛羅里達(dá)亞種NPY基因的DNA和c DNA克隆及序列分析神經(jīng)肽Y(Neuropeptide Y,NPY)在機(jī)體的攝食活動(dòng)中發(fā)揮重要作用,是哺乳動(dòng)物最重要的一種內(nèi)源性促食欲因子。為了解大口黑鱸NPY基因的結(jié)構(gòu)及進(jìn)一步研究該基因在大口黑鱸中的功能作用,采用RT-PCR和RACE技術(shù),克隆了大口黑鱸北方亞種(M.salmoides salmoides)和佛羅里達(dá)亞種(M.salmoides floridanus)NPY基因c DNA序列,兩亞種NPY c DNA均包括一個(gè)編碼99個(gè)氨基酸的ORF框和長度為52bp的5′非編碼區(qū)(5′-UTR);結(jié)合PCR和基因組步移技術(shù)獲得了長度分別為3561bp和3565bp的大口黑鱸北方亞種和佛羅里達(dá)亞種NPY基因DNA序列。序列分析結(jié)果表明,大口黑鱸北方亞種和佛羅里達(dá)亞種NPY基因均由4個(gè)外顯子和3個(gè)內(nèi)含子組成。經(jīng)MATINSPECTOR軟件預(yù)測,在北方亞種和佛羅里達(dá)亞種啟動(dòng)子序列分布有TATA框、CAAT框、CCAAT-Box、GATA-Box等基本轉(zhuǎn)錄調(diào)控元件。實(shí)驗(yàn)在大口黑鱸兩亞種NPY DNA序列間發(fā)現(xiàn)了6個(gè)單個(gè)位點(diǎn)堿基差異,與大口黑鱸北方亞種相比佛羅里達(dá)亞種啟動(dòng)子區(qū)域出現(xiàn)一個(gè)4個(gè)堿基的插入。不同物種間NPY基因的序列同源性分析表明大口黑鱸與鱖魚和石斑魚的NPY基因核苷酸同源性最高,達(dá)90%和88%,氨基酸同源性分別為93%和95%。研究結(jié)果旨在為進(jìn)一步研究大口黑鱸神經(jīng)肽Y基因功能作用奠定基礎(chǔ)。4.大口黑鱸NPY基因的SNPs與生長性狀相關(guān)性的初步研究在獲得大口黑鱸NPY基因DNA序列的基礎(chǔ)之上,選取該基因作為研究大口黑鱸生長性狀的候選基因,鑒于神經(jīng)肽Y是魚類攝食調(diào)控網(wǎng)絡(luò)中最強(qiáng)有力的促進(jìn)食欲的攝食因子,推測該基因上的單核苷酸多態(tài)性會(huì)影響魚類的攝食以及相關(guān)激素的分泌,進(jìn)而對大口黑鱸的生長性狀產(chǎn)生影響。實(shí)驗(yàn)根據(jù)克隆得到的大口黑鱸NPY基因的DNA序列,在NPY基因5′非編碼區(qū)篩選獲得了1個(gè)SNP位點(diǎn):S1(A-85C)。根據(jù)實(shí)驗(yàn)所選取的分型群體,采樣Sna Pshot方法對實(shí)驗(yàn)群體的NPY基因進(jìn)行SNPs位點(diǎn)分型。對分型結(jié)果進(jìn)一步處理分析后結(jié)果表明,實(shí)驗(yàn)群體在突變位點(diǎn)處于哈溫平衡,該位點(diǎn)位點(diǎn)共有3種基因型:AA、AC和CC。AC基因型群體的平均體質(zhì)量、體寬、全長和體高分別高于AA基因型群體6.51%、2.56%、2.18%和3.34%;相對于CC基因型分別高出了15.37%、7.97%、4.09%和5.12%,但在統(tǒng)計(jì)上不同基因型在各項(xiàng)生長性狀間均沒有達(dá)到顯著相關(guān)性(P0.05)。但實(shí)驗(yàn)發(fā)現(xiàn)的SNPs位點(diǎn)對以后進(jìn)一步研究大口黑鱸NPY基因與其他性狀的聯(lián)系提供了參考。5.三個(gè)基因多態(tài)性及合并基因型對大口黑鱸生長性狀的遺傳效應(yīng)為進(jìn)一步探討將Ghrelin、PACAP和NPY基因作為影響大口黑鱸生長性狀的候選基因的可能性,在采用直接測序法檢測了這三個(gè)基因的單核苷酸多態(tài)性并分別分析了這3個(gè)基因SNPs與大口黑鱸生長性狀的關(guān)聯(lián)性的基礎(chǔ)之上,分析這三個(gè)基因間合并基因型對大口黑鱸生長性狀的影響。結(jié)果表明:Ghrelin基因的兩個(gè)突變位點(diǎn)與PACAP基因上的SNP位點(diǎn)分別有4種和5種合并基因型,與NPY基因也分別形成了4種和5種合并基因型,PACAP和NPY基因的合并基因型有4種,三個(gè)基因一起組成的合并基因型分別有9種和6種。關(guān)聯(lián)分析表明:Ghrelin-PACAP基因優(yōu)勢基因型聚合體ACAC和AAAC個(gè)體在各生長性狀上基本顯著優(yōu)于其它合并基因型個(gè)體(P0.05);以Ghrelin基因的S1位點(diǎn)與PACAP基因進(jìn)行合并基因型分析時(shí),ACAC基因型在體質(zhì)量、體寬和全長上比劣勢基因型CCCC分別高出了14.1%,4.3%和3.6%,以Ghrelin基因的S2位點(diǎn)與PACAP基因進(jìn)行合并基因型分析時(shí),AAAC基因型在體質(zhì)量、體寬和全長上比劣勢基因型ACAC分別高出了16.2%,4.5%和3.0%。Ghrelin-NPY基因優(yōu)勢基因型聚合體的生長性狀均值也優(yōu)于其它合并基因型,但在統(tǒng)計(jì)上沒有達(dá)到顯著相關(guān)性(P0.05)。PACAP-NPY基因的合并基因型中,ACAC基因型個(gè)體在體高和全長上顯著優(yōu)于CCAA基因型個(gè)體,在體質(zhì)量、體寬和全長上分別高出了12.4%,4.3%和3.3%。三個(gè)基因組成的合并優(yōu)勢基因型中,以S2位點(diǎn)參與合并而組成的AAACAC為最優(yōu)基因型,且大于任意兩基因間的最優(yōu)合并基因型在各生長性狀上的均值,以S1位點(diǎn)參與合并而組成的CCCCAA為劣勢基因型,AAACAC合并基因型個(gè)體在體質(zhì)量、體寬和全長上比CCCCAA基因型個(gè)體分別高出了26.8%,8.5%和4.2%。結(jié)果顯示:兩基因合并后對大口黑鱸生長性狀的影響效應(yīng)優(yōu)于任意單個(gè)基因優(yōu)勢基因型的影響效應(yīng),三個(gè)基因合并后的優(yōu)勢基因型在各生長性狀上也優(yōu)于任意兩個(gè)的合并優(yōu)勢基因型;據(jù)此我們做出以下判斷:在符合遺傳機(jī)理的前提下,可能合并的生長相關(guān)基因的優(yōu)勢單基因型越多,對大口黑鱸生長性狀的影響效應(yīng)就越大。
[Abstract]:Micropterus salmoides, a kind of bass (Perciformes), is one of the large aquaculture fishes in our country. The big mouth bass is a typical strong carnivorous fish. The traditional way of breeding is to feed the icy fresh fish, which not only makes the aquaculture limited by the shortage of marine fishery resources. There are also the risk of the ice fish body introducing the pathogen to the water body, which is difficult to carry out the production of the plant culture and the harmless aquatic products. In recent years, the use of artificial feed to feed the big mouth bass has achieved certain results. It is seriously restricted to the further development of Chinese big mouth bass breeding industry. It is an important goal for the breeding of big mouth bass that can be better feeding and using artificial mix feed and good growth traits. The molecular marker assisted breeding is an important means of genetic improvement of fish, with the help of molecular markers. The correlation and selection analysis of the candidate genes and target traits can greatly improve the efficiency of the selection of the related economic traits and shorten the breeding time effectively. According to the previous study, the feeding of fish is influenced by many internal and external factors, in which the appetite condition is the main internal influence factor of fish, and it is used as a fish fish. When they are in the state of appetite, their feed intake and utilization are improved correspondingly. The regulation of fish feeding is done mainly through the interaction of brain and nerve peripheral signals, and appetite related genes are the main genes involved in the regulation of fish feeding. Select Ghrelin, PACAP and NPY as the candidate genes related to the growth traits of the large mouth bass, analyze the association between the single nucleotide polymorphism (single nucleotide polymorphism, SNP) and the growth traits, in order to provide effective marker for the molecular marker assisted breeding of the big mouth bass, and promote the food to improve the new product of the big mouth bass. The main research contents are as follows: 1. the screening of the Ghrelin gene SNPs of the big mouth bass and the correlation analysis with the growth traits, Ghrelin is a kind of brain gut peptide of vertebrates, which can promote the feeding function, promote the release of growth hormone (GH), participate in the regulation of energy balance and carbohydrate metabolism. In order to explore the polymorphism of ghrelin gene and the large mouth bass The correlation of long characters, 2 SNPs loci, S1 (A-642C) and S2 (A-639C) were obtained by direct sequencing by direct sequencing of the promoter sequence of the ghrelin gene of large mouth bass. The SNPs loci were detected by the same batch of 327 tails, and the Sna Pshot method was used to detect and classify the SNPs loci. The results showed that the experimental population was in ghrelin gene. The two SNPs loci were in the HW equilibrium, and S1 and S2 were composed of 5 double types (D1, D2, D3, D4 and D5). The results of correlation analysis showed that the AC genotypes and AA genotypes on the S1 loci were the dominant genotypes of two mutation sites respectively. Screening of the PACAP gene SNPs of the oral bass and its association with the growth traits, the pituitary adenylate cycled kinase polypeptide (pituitary adenylate cyclase-activating polypeptide, PACAP) is a multi effect hormone that promotes growth hormone, gonadotropin and prolactin secretion. It is a member of the glucagon / secretory peptide family. It is to explore PACAP. The effect of genetic polymorphism on the growth traits of big mouth bass was found by direct sequencing. The PACAP gene was screened by direct sequencing: A-2282C, with 3 genotypes: AA, AC and CC, the genotype frequencies were 0.034,0.645 and 0.321, the A allele frequency was 0.3563, C allele frequency was 0.6437. selected for the same batch reproduction and 327 tail in the same pond culture. The Sna Pshot method was used to detect and classify the SNPs loci, and the least square analysis model was constructed to analyze the correlation between the genotypes of the mutant loci and the growth traits. The ksquare test results showed that the hash equilibrium constant was 0.0615, that is, the site was basically in the equilibrium state of Hardy-Weinbery in the detected large mouth bass population, and the effective allele was effective. The number of genes was 1.8465, and the expected heterozygosity and observed heterozygosity were 0.5409 and 0.6442. variance analysis showed that the average phenotypic values of AC genotypes were much higher than those of AA and CC genotypes, and the AC genotypes were significantly higher in body mass, in body height and in full length than in AA and CC genotypes (P0.05).AA genes. The average body mass of the type population, body height and full length were 18.1%, 5.8% and 5.4% higher for CC genotypes, but there was no significant difference in the growth traits between AA and CC populations (P0.05). It is presumed that this locus may be the main quantitative trait nucleotide (quantitative trait nucleotides, QTN) affecting the growth traits of large mouth bass. The cloning and sequence analysis of the DNA and C DNA of the northern subspecies and subspecies of Florida subspecies of Florida subspecies NPY, neuropeptide Y (Neuropeptide Y, NPY) plays an important role in the feeding activities of the organism. It is the most important endogenous appetite promoting factor of mammalian. To understand the structure and further of the NPY gene of the large mouth bass. To study the function of the gene in the big mouth bass, RT-PCR and RACE techniques were used to clone the DNA sequence of the northern subspecies (M.salmoides salmoides) and the NPY gene C of Florida subspecies (M.salmoides floridanus) of the large mouth bass. The two subspecies NPY C DNA included a 99 amino acid coded frame and the 5 'non coding region (5'). -UTR); PCR and genomic step technique were used to obtain the DNA sequence of NPY gene of the northern subspecies and Florida subspecies of Florida subspecies, large mouth bass and 3565bp, respectively. The sequence analysis showed that the NPY genes of the northern subspecies and the subspecies of Florida subspecies were composed of 4 exons and 3 introns. The subspecies of northern subspecies and subspecies of Florida subspecies are distributed with TATA frame, CAAT frame, CCAAT-Box, GATA-Box and other basic transcriptional regulators. The experiment found 6 single site base differences between the two subspecies NPY DNA sequences of the large mouth bass, and a 4 base insertion in the subspecies of the subspecies of the northern subspecies of the big mouth bass. The sequence homology of NPY gene among different species showed that the nucleotide homology of the NPY gene of the large mouth bass and the mandarin fish and the grouper was the highest, reaching 90% and 88%. The amino acid homology was 93% and 95%., respectively, to lay the foundation for the further study of the Y gene function of the large mouth perch neuropeptide Y gene, which lay the foundation for the SNPs of the.4. large mouth bass NPY gene. A preliminary study on the correlation of growth traits on the basis of obtaining the DNA sequence of the NPY gene of the large mouth bass, selected the gene as a candidate gene for the study of the growth traits of large mouth bass. In view of the fact that neuropeptide Y is the most powerful feeding factor promoting appetite in the fish feeding regulation network, it is speculated that the single nucleotide polymorphisms on this gene will be affected. The feeding of fishes and the secretion of related hormones had an effect on the growth traits of the large mouth bass. According to the DNA sequence of the NPY gene of the cloned bass bass, 1 SNP loci were obtained in the 5 'non coding region of the NPY gene: S1 (A-85C). According to the selected group of the experiment, the Sna Pshot method was sampled for the experimental population. The NPY gene was typed at the SNPs locus. The results of further analysis showed that the experimental population was at the equilibrium at the mutation site, and there were 3 genotypes at the loci: the average body mass of the AA, AC and CC.AC genotypes, the body width, the total length and the body height were 6.51%, 2.56%, 2.18% and 3.34%, respectively, compared with the AA genotypes; relative to C. The C genotypes were 15.37%, 7.97%, 4.09% and 5.12% higher respectively, but there was no significant correlation between the different genotypes of the genotypes (P0.05). However, the SNPs loci found in the experiment further studied the relationship between the NPY gene and other traits of the big mouth bass, which provided the reference to three gene polymorphisms and the combination genotype of.5.. The genetic effect on the growth traits of large mouth bass is to further explore the possibility of Ghrelin, PACAP and NPY genes as candidate genes affecting the growth traits of large mouth bass. The single nucleotide polymorphisms of these three genes were detected by direct sequencing and the correlation between the 3 gene SNPs and the growth traits of large mouth bass was analyzed. On the basis of the analysis, the effects of the combined genotypes of the three genes on the growth traits of the big mouth bass were analyzed. The results showed that the two mutation sites of the Ghrelin gene and the SNP loci on the PACAP gene were 4 and 5 genotypes respectively. 4 and 5 genotypes were formed respectively with the NPY gene, and the combined genotype of the PACAP and NPY genes had the same genotypes. There were 9 and 6 types of combined genotypes composed of three genes. The correlation analysis showed that the Ghrelin-PACAP gene dominant genopolymer ACAC and AAAC were significantly better than other combined genotype individuals (P0.05) in the growth traits, and ACAC was combined with the S1 site of the Ghrelin gene and the PACAP gene, ACAC Genotypes were 14.1%, 4.3% and 3.6% higher in body mass, body width and length than inferior genotypes, respectively. When the S2 locus of Ghrelin gene and PACAP gene were combined with genotype analysis, the AAAC genotype was 16.2%, 4.5%, and 3.0%.Ghrelin-NPY gene dominant genotype in body mass, body width and full length than inferior genotype ACAC, respectively. The average growth traits of the polymers were also better than those of other combined genotypes, but in the statistics of P0.05.PACAP-NPY gene combination, the ACAC genotype was significantly better than the CCAA genotype in the body height and length, and the body mass, body width and the whole length were 12.4%, 4.3% and 3.3%. three genomes respectively. In the combined dominant genotypes, AAACAC is the best genotype with the S2 site involved in the combination, and it is larger than the average of the optimal combination genotype between any two genes. The CCCCAA is a inferior genotype with the S1 locus involved in the combination, and the AAACAC combined with the genotypes is in body mass, body width and total length than CCCCA The A genotype individuals were 26.8%, 8.5% and 4.2%. respectively. The results showed that the effect of the two gene combination on the growth traits of the big mouth bass was better than that of any single gene dominant genotype, and the dominant genotypes of the three genes were also superior to the two combined dominant genotypes in the growth traits. In accordance with the genetic mechanism, the more dominant single genotypes of the associated growth related genes, the greater the effect on the growth traits of the big mouth bass.
【學(xué)位授予單位】：上海海洋大學(xué)
【學(xué)位級別】：碩士
【學(xué)位授予年份】：2016
【分類號(hào)】：S917.4

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