虾夷扇贝橘红色闭壳肌产生的原因及其在育种中的应用
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摘要
1橘红色闭壳肌虾夷扇贝的选择与繁育
虾夷扇贝自1982年从日本引进我国,目前是我国北方最重要的海水经济养殖贝类之一。在对虾夷扇贝开展选优策略育种工程的过程中,在野生虾夷扇贝群体中发现了一种比例极低,约占群体总数0.2%的颜色突变体虾夷扇贝,其闭壳肌呈鲜艳的橘红色而不是普通的白色,这种现象在贝类包括扇贝中从未见过报道。本研究对颜色突变体虾夷扇贝的性别比、壳色等进行了观察和统计,与对照组相比无差异;筛选性腺饱满的突变体闭壳肌虾夷扇贝雌雄各500只,及等量的白色闭壳肌虾夷扇贝对照,分别进行常规育种,养成扇贝在七月龄时便可观察到其闭壳肌颜色呈橘红色,比例为100%,对照组为0,结果表明闭壳肌颜色是稳定遗传的质量性状,可应用于育种。对22月龄的虾夷扇贝进行生长性状统计,结果表明在性状总重、软体重、柱重与对照组存在极显著差异,在性状壳长存在显著差异,橘红色闭壳肌虾夷扇贝生长速度较对照组具有明显优势;抗逆性实验证明,色素突变体虾夷扇贝的抗逆性较对照组高,表现出良好的抗逆性。我们把该品种命名为“海大金贝”。
2虾夷扇贝橘红色闭壳肌中主要色素的分离提纯与结构鉴定
通过紫外-可见光全波长光谱扫描,在450nm处有明显吸收峰并呈现较明显的“三指峰”现象,初步确定橘红色闭壳肌中的色素为脂溶性类胡萝卜素。通过柱层析、高效液相等手段对橘红色闭壳肌中的主要类胡萝卜素进行了提取分离,分离提纯得到两种类胡萝卜素,并通过MS、~1H-NMR和~(13)C-NMR对其进行结构鉴定,结果表明(3S,3'R)-3,3'-dihydroxy-7',8'-didehydro-β,β-Caroten-4-one和(3S,3'S) -7,7',8,8'-tetradehydro-β,β-Carotene-3,3'-diol,俗称pectenolone和pectenoxanthin是橘红色闭壳肌中的主要类胡萝卜素,这是国内外首次对虾夷扇贝闭壳肌中的类胡萝卜素进行提纯及结构鉴定。
3虾夷扇贝类胡萝卜素的种类组成和累积变化规律
建立了橘红色闭壳肌中类胡萝卜素的HPLC分析定量检测技术:色普柱为Agilent公司的Eclipse XDB-C18柱(200mm×4.6mm,i.d.5μm);进样体积,5μl ;流速1ml/min;检测波长450nm;柱温25℃;流动相为乙腈:甲醇:二氯甲烷(V/V/V)=40:56:4并加入0.1%的BHT;分离时间为8min。利用上述方法对7、10、12、15、18、22月龄的虾夷扇贝橘红色闭壳肌进行了追踪分析,结果表明在快速生长的幼贝期,其闭壳肌类胡萝卜素的含量一直呈上升趋势;在性腺发育时期闭壳肌内类胡萝卜素的含量达到最高值,生殖期过后类胡萝卜素的含量随之降低;在11月份-次年2月份以积累pectenoxanthin为主,而在4月份-10月份以累积pectenolone为主;闭壳肌内的类胡萝卜素含量与扇贝的大小无相关性,与生长阶段相关。利用HPLC法对橘红/白色闭壳肌虾夷扇贝各组织中含有的类胡萝卜素种类和含量,结果表明,虾夷扇贝在性腺、肝胰腺、外套膜、鳃丝中均含有类胡萝卜素pectenolone和pectenoxanthin,白色闭壳肌中未检测到类胡萝卜素,其中肝胰腺除上述两种类胡萝卜素外,还含有其他多种未鉴定的类胡萝卜素;在各组织中,雌性性腺、肝胰腺、鳃丝中类胡萝卜素的含量较高;橘红色闭壳肌虾夷扇贝各组织中的类胡萝卜素含量高于白色闭壳肌虾夷扇贝;雌性虾夷扇贝中的类胡萝卜素的含量高于雄性。
4累积类胡萝卜素对虾夷扇贝闭壳肌的形态结构和化学成分的影响
本文对橘红色闭壳肌进行了常规HE染色切片观察,并未观察到异常现象;超薄切片透射电镜观察发现橘红色闭壳肌在肌纤维类型及基质密度等方面有别于白色闭壳肌,在橘红色闭壳肌中有三种肌纤维,其中疏松型肌纤维为橘红色闭壳肌特有。另外,通过索氏提取法对橘红色闭壳肌和白色闭壳肌中的脂肪及脂肪酸进行了提取,以气相色谱-质谱联用仪对其中脂肪酸的种类和含量进行分析,结果表明橘红色闭壳肌中的总脂肪含量略高于白色闭壳肌,脂肪酸中亚油酸、油酸在橘红色闭壳肌中的含量远大于白色闭壳肌中的含量,可能与类胡萝卜素的吸收代谢有关。氨基酸中除甘氨酸及脯氨酸外,橘红色闭壳肌中的氨基酸含量均高于普通的白色闭壳肌且富含牛磺酸。
5橘红色闭壳肌虾夷扇贝选育群的遗传结构分析
对养成的F1群体通过SSR技术利用6对引物进行遗传结构分析,结果表明该群体并未发生大的遗传结构改变,具有丰富的遗传多样性。采用24对引物组合对橘红/白色闭壳肌虾夷扇贝群体进行了MSAP分析,结果表明,甲基化程度存在差异的位点46个,其中18个位点为甲基化或半甲基化位点,推测橘红色闭壳肌的发生可能与这些位点有关,它们调控了该性状的表达。
1 The selection and breeding of Yesso scallop with orange muscle
The Yesso scallop (Patinopecten yessoensis) is one of most important commercial aquacultural shellfish in the north of China and since it was introduced from Japan in 1982. During the course of a research project aimed at optimizing a breeding strategy for this scallop, we identified a rare color variant that occurred at about 0.2% in the natural population. This muscle tissue of this variant was orange as opposed to the common white. Orange variants have not been reported before in shellfish, including scallops. In this study, gender ratio and shell color of the orange variant were investigated and it showed no difference with the common. 500 male and female individuals of color variant were screened for conventional breeding separately and a control was set with same amount of white ones. The proportion of F1 generation with orange muscle was 100% when it was 7 months old, while there was no orange muscle appeared in the control group. It demonstrated that the color of muscle can be steadily inherited and it can be used for breeding. Statistic analysis of growth traits of the 22-month Yesso scallops showed that there was significant difference between the the group with orange muscle and the control group in total weight, soft weight, muscle weight and shell length, it indicated that the growth rate of the Yesso scallops with orange muscle was faster than the control group significantly. The new variety of the Yesso scallop was named“Hai da jin bei”.
2 Isolation and identification of the main pigment from the rare orange muscle of the Yesso scallop
The wavelength spectrum scanning by UV-visible of the extract showed that the peaks were determined upslope, apex and downlope, and there was an obvious peak value at 450nm, the pigments were therefore identified as carotenoids preliminary. Two main carotenoids were isolated and purified by silica gel chromatography(TLC) and high performance liquid chromatography(HPLC), and were characterized by mass spectrometry(MS) and ~1H- and ~(13)C-nuclear magnetic resonance(NMR) methods. Its structure was identified as (3S,3'R)-3,3'-dihydroxy-7',8'-didehydro-β,β-Caroten-4-one and (3S,3'S)-7,7',8,8'-tetra-dhydro-β,β-Carotene-3,3'-diol, with common name pectenolone and pectenoxanthin. This is the first report of pectenolone and pectenoxanthin, two types of carotenoid, in the muscle of the Yesso scallop.
3 Roles of the carotenoids composition and accumulation in Yesso scallop
The method for detecting carotenoids using reverse phase high performance liquid chromatography (RP-HPLC) was developed. The conditions of RP-HPLC analysis were an analytical scale (200×4.6 mm, inner diameter 5μm) Eclipse XDB-C18 column of Agilent corporation; 10μl injection volume; 25°C column temperature; acetonitrile, methanol and methylene dichloride mobile phase (40:56:4, V/VV) with 0.1% BHT; 1.0 ml per minute flow rate; 450 nm UV-VIS detector; 8 minutes elution.
The carotenoids from orange muscle of Yesso scallop of 7, 10, 12, 15, 18, 22 months old was detected using the method refered above. It showed that the carotenoids content was increasing during juvenile stage when the scallop grow rapidly, and reached plateau in the breeding reason, and then decreased after reproduction. It can also be concluded that the muscle mainly accumulated pectenoxanthin in November to February, while mainly pectenolone in April to October. The carotenoids content of orange muscle had no relationship with the size of the scallop, but the growth stage.
The composition and content of carotenoids of various tissues in Yesso scallop with orange/white muscles was detected by RP-HPLC. It was showed that pectenolone and pectenoxanthin were all found in Gonads, hepatopancreas, mantle and gill, among of them, ovary, hepatopancreas and gill possessed more carotenoid than other tissues, but no carotenoids were found in the white muscles of Yesso scallops. Hepatopancreas contains a variety other carotenoids some kind of other not identified beside pectenolone and pectenoxanthin. The result also showed that the Yesso scallop with orange muscle had more the carotenoids in a variety of tissues than the common, and the female more than the male.
4 Impacts of carotenoid accumulation on morphological structure and chemical composition in orange muscle of Yesso scallop
There was no difference between the orange and white muscle obserced by conventional HE staining, while musclular fiber type and matrix density in the orange muscle was different from white ones by the TEM observation of ultra-thin slices, and there were three musclular fiber types, among of them, the loose muscular fiber was orange muscle specifically. The fat and fat acid of orange and white muscle were ectracted through Soxhlet extraction method, the varieties and content were detected by GC-MS. The result showed that the fat of orange muscle was a little higher than the white ones, and the linoleic acid, oleic acid content in the orange muscle was more than white ones, which may be related to the carotenoids metabolism. Amino acids content of orange muscle were higher than common white ones except glycine and proline, and taurine was rich in orange muscle.
5 Genetic structure of breeding population with orange muscle
We surveyed the level of genetic variation and the pattern of genetic structure in the breeding population with orange muscle using SSR markers. Based on 6 primers, the result showed that the breeding population has high genetic diversity and was no significant change with respect to the wild population. And MSAP with 24 primer was used for analysis in Yesso scallops with orange/white muscle, it was showed that there were 46 loci which had differences in the degree of methylation between the population with orange muscle and the white, among of which 18 were methylated or semi-methlated loci, which may be related to the appearance of orange muscle.
虾夷扇贝自1982年从日本引进我国,目前是我国北方最重要的海水经济养殖贝类之一。在对虾夷扇贝开展选优策略育种工程的过程中,在野生虾夷扇贝群体中发现了一种比例极低,约占群体总数0.2%的颜色突变体虾夷扇贝,其闭壳肌呈鲜艳的橘红色而不是普通的白色,这种现象在贝类包括扇贝中从未见过报道。本研究对颜色突变体虾夷扇贝的性别比、壳色等进行了观察和统计,与对照组相比无差异;筛选性腺饱满的突变体闭壳肌虾夷扇贝雌雄各500只,及等量的白色闭壳肌虾夷扇贝对照,分别进行常规育种,养成扇贝在七月龄时便可观察到其闭壳肌颜色呈橘红色,比例为100%,对照组为0,结果表明闭壳肌颜色是稳定遗传的质量性状,可应用于育种。对22月龄的虾夷扇贝进行生长性状统计,结果表明在性状总重、软体重、柱重与对照组存在极显著差异,在性状壳长存在显著差异,橘红色闭壳肌虾夷扇贝生长速度较对照组具有明显优势;抗逆性实验证明,色素突变体虾夷扇贝的抗逆性较对照组高,表现出良好的抗逆性。我们把该品种命名为“海大金贝”。
2虾夷扇贝橘红色闭壳肌中主要色素的分离提纯与结构鉴定
通过紫外-可见光全波长光谱扫描,在450nm处有明显吸收峰并呈现较明显的“三指峰”现象,初步确定橘红色闭壳肌中的色素为脂溶性类胡萝卜素。通过柱层析、高效液相等手段对橘红色闭壳肌中的主要类胡萝卜素进行了提取分离,分离提纯得到两种类胡萝卜素,并通过MS、~1H-NMR和~(13)C-NMR对其进行结构鉴定,结果表明(3S,3'R)-3,3'-dihydroxy-7',8'-didehydro-β,β-Caroten-4-one和(3S,3'S) -7,7',8,8'-tetradehydro-β,β-Carotene-3,3'-diol,俗称pectenolone和pectenoxanthin是橘红色闭壳肌中的主要类胡萝卜素,这是国内外首次对虾夷扇贝闭壳肌中的类胡萝卜素进行提纯及结构鉴定。
3虾夷扇贝类胡萝卜素的种类组成和累积变化规律
建立了橘红色闭壳肌中类胡萝卜素的HPLC分析定量检测技术:色普柱为Agilent公司的Eclipse XDB-C18柱(200mm×4.6mm,i.d.5μm);进样体积,5μl ;流速1ml/min;检测波长450nm;柱温25℃;流动相为乙腈:甲醇:二氯甲烷(V/V/V)=40:56:4并加入0.1%的BHT;分离时间为8min。利用上述方法对7、10、12、15、18、22月龄的虾夷扇贝橘红色闭壳肌进行了追踪分析,结果表明在快速生长的幼贝期,其闭壳肌类胡萝卜素的含量一直呈上升趋势;在性腺发育时期闭壳肌内类胡萝卜素的含量达到最高值,生殖期过后类胡萝卜素的含量随之降低;在11月份-次年2月份以积累pectenoxanthin为主,而在4月份-10月份以累积pectenolone为主;闭壳肌内的类胡萝卜素含量与扇贝的大小无相关性,与生长阶段相关。利用HPLC法对橘红/白色闭壳肌虾夷扇贝各组织中含有的类胡萝卜素种类和含量,结果表明,虾夷扇贝在性腺、肝胰腺、外套膜、鳃丝中均含有类胡萝卜素pectenolone和pectenoxanthin,白色闭壳肌中未检测到类胡萝卜素,其中肝胰腺除上述两种类胡萝卜素外,还含有其他多种未鉴定的类胡萝卜素;在各组织中,雌性性腺、肝胰腺、鳃丝中类胡萝卜素的含量较高;橘红色闭壳肌虾夷扇贝各组织中的类胡萝卜素含量高于白色闭壳肌虾夷扇贝;雌性虾夷扇贝中的类胡萝卜素的含量高于雄性。
4累积类胡萝卜素对虾夷扇贝闭壳肌的形态结构和化学成分的影响
本文对橘红色闭壳肌进行了常规HE染色切片观察,并未观察到异常现象;超薄切片透射电镜观察发现橘红色闭壳肌在肌纤维类型及基质密度等方面有别于白色闭壳肌,在橘红色闭壳肌中有三种肌纤维,其中疏松型肌纤维为橘红色闭壳肌特有。另外,通过索氏提取法对橘红色闭壳肌和白色闭壳肌中的脂肪及脂肪酸进行了提取,以气相色谱-质谱联用仪对其中脂肪酸的种类和含量进行分析,结果表明橘红色闭壳肌中的总脂肪含量略高于白色闭壳肌,脂肪酸中亚油酸、油酸在橘红色闭壳肌中的含量远大于白色闭壳肌中的含量,可能与类胡萝卜素的吸收代谢有关。氨基酸中除甘氨酸及脯氨酸外,橘红色闭壳肌中的氨基酸含量均高于普通的白色闭壳肌且富含牛磺酸。
5橘红色闭壳肌虾夷扇贝选育群的遗传结构分析
对养成的F1群体通过SSR技术利用6对引物进行遗传结构分析,结果表明该群体并未发生大的遗传结构改变,具有丰富的遗传多样性。采用24对引物组合对橘红/白色闭壳肌虾夷扇贝群体进行了MSAP分析,结果表明,甲基化程度存在差异的位点46个,其中18个位点为甲基化或半甲基化位点,推测橘红色闭壳肌的发生可能与这些位点有关,它们调控了该性状的表达。
1 The selection and breeding of Yesso scallop with orange muscle
The Yesso scallop (Patinopecten yessoensis) is one of most important commercial aquacultural shellfish in the north of China and since it was introduced from Japan in 1982. During the course of a research project aimed at optimizing a breeding strategy for this scallop, we identified a rare color variant that occurred at about 0.2% in the natural population. This muscle tissue of this variant was orange as opposed to the common white. Orange variants have not been reported before in shellfish, including scallops. In this study, gender ratio and shell color of the orange variant were investigated and it showed no difference with the common. 500 male and female individuals of color variant were screened for conventional breeding separately and a control was set with same amount of white ones. The proportion of F1 generation with orange muscle was 100% when it was 7 months old, while there was no orange muscle appeared in the control group. It demonstrated that the color of muscle can be steadily inherited and it can be used for breeding. Statistic analysis of growth traits of the 22-month Yesso scallops showed that there was significant difference between the the group with orange muscle and the control group in total weight, soft weight, muscle weight and shell length, it indicated that the growth rate of the Yesso scallops with orange muscle was faster than the control group significantly. The new variety of the Yesso scallop was named“Hai da jin bei”.
2 Isolation and identification of the main pigment from the rare orange muscle of the Yesso scallop
The wavelength spectrum scanning by UV-visible of the extract showed that the peaks were determined upslope, apex and downlope, and there was an obvious peak value at 450nm, the pigments were therefore identified as carotenoids preliminary. Two main carotenoids were isolated and purified by silica gel chromatography(TLC) and high performance liquid chromatography(HPLC), and were characterized by mass spectrometry(MS) and ~1H- and ~(13)C-nuclear magnetic resonance(NMR) methods. Its structure was identified as (3S,3'R)-3,3'-dihydroxy-7',8'-didehydro-β,β-Caroten-4-one and (3S,3'S)-7,7',8,8'-tetra-dhydro-β,β-Carotene-3,3'-diol, with common name pectenolone and pectenoxanthin. This is the first report of pectenolone and pectenoxanthin, two types of carotenoid, in the muscle of the Yesso scallop.
3 Roles of the carotenoids composition and accumulation in Yesso scallop
The method for detecting carotenoids using reverse phase high performance liquid chromatography (RP-HPLC) was developed. The conditions of RP-HPLC analysis were an analytical scale (200×4.6 mm, inner diameter 5μm) Eclipse XDB-C18 column of Agilent corporation; 10μl injection volume; 25°C column temperature; acetonitrile, methanol and methylene dichloride mobile phase (40:56:4, V/VV) with 0.1% BHT; 1.0 ml per minute flow rate; 450 nm UV-VIS detector; 8 minutes elution.
The carotenoids from orange muscle of Yesso scallop of 7, 10, 12, 15, 18, 22 months old was detected using the method refered above. It showed that the carotenoids content was increasing during juvenile stage when the scallop grow rapidly, and reached plateau in the breeding reason, and then decreased after reproduction. It can also be concluded that the muscle mainly accumulated pectenoxanthin in November to February, while mainly pectenolone in April to October. The carotenoids content of orange muscle had no relationship with the size of the scallop, but the growth stage.
The composition and content of carotenoids of various tissues in Yesso scallop with orange/white muscles was detected by RP-HPLC. It was showed that pectenolone and pectenoxanthin were all found in Gonads, hepatopancreas, mantle and gill, among of them, ovary, hepatopancreas and gill possessed more carotenoid than other tissues, but no carotenoids were found in the white muscles of Yesso scallops. Hepatopancreas contains a variety other carotenoids some kind of other not identified beside pectenolone and pectenoxanthin. The result also showed that the Yesso scallop with orange muscle had more the carotenoids in a variety of tissues than the common, and the female more than the male.
4 Impacts of carotenoid accumulation on morphological structure and chemical composition in orange muscle of Yesso scallop
There was no difference between the orange and white muscle obserced by conventional HE staining, while musclular fiber type and matrix density in the orange muscle was different from white ones by the TEM observation of ultra-thin slices, and there were three musclular fiber types, among of them, the loose muscular fiber was orange muscle specifically. The fat and fat acid of orange and white muscle were ectracted through Soxhlet extraction method, the varieties and content were detected by GC-MS. The result showed that the fat of orange muscle was a little higher than the white ones, and the linoleic acid, oleic acid content in the orange muscle was more than white ones, which may be related to the carotenoids metabolism. Amino acids content of orange muscle were higher than common white ones except glycine and proline, and taurine was rich in orange muscle.
5 Genetic structure of breeding population with orange muscle
We surveyed the level of genetic variation and the pattern of genetic structure in the breeding population with orange muscle using SSR markers. Based on 6 primers, the result showed that the breeding population has high genetic diversity and was no significant change with respect to the wild population. And MSAP with 24 primer was used for analysis in Yesso scallops with orange/white muscle, it was showed that there were 46 loci which had differences in the degree of methylation between the population with orange muscle and the white, among of which 18 were methylated or semi-methlated loci, which may be related to the appearance of orange muscle.
引文
毕克,包振民,黄晓婷,王珏,赵洋,胡景杰:栉孔扇贝♀×华贵栉孔扇贝♂受精及早期胚
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