油茶种子发育过程组分及脂类代谢相关基因表达变化研究
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摘要
油茶泛指山茶科(Theaceae)山茶属（Camellia）中具有生产价值的油用物种，是中国重要木本油料树种，其中普通油茶（Camellia oleifera Abel）栽培面积最广、产量最高，在生产中应用最广泛。自“六五”以来，研究者们在普通油茶果实产量、果实性状及采后加工、茶油营养成分、茶油储藏等方面开展了大量研究，并已获得了巨大进展。但对油茶种子发育过程中果实形状、种子的物质组成及重要性状控制基因的表达变化规律等研究较少。研究油茶果实和种子的生长发育规律，探索种子发育过程中重要营养成分的合成贮存规律，将为采用分子手段调控油茶种子重要营养成分合成，进一步提高茶油的营养保健价值奠定理论基础。本文分析了油茶果实和种子发育过程中表型性状、物质组成、重要性状基因表达量等指标的变化规律，具体研究内容如下：
(1)分析了普通油茶长林系列4号、40号和166号3个无性系5-10月份果实果形、果重和种子的重量、鲜果出籽率等指标在果实生长过程中的变化规律。分析表明，3个无性系5月份果实大部分为圆橄榄形，8月份果形变化较大，到9月份基本分化成各无性系的特征果形；果实6-8月份间果实为迅速膨大期，果实重量和纵、横径大小在此时增长量最大，9月份之后果实重量和大小基本保持恒定；种皮厚度各月变化不大。种子重量和大小6-10月持续增长，其中6-8月份生长速度较快。鲜果出籽率6-8月迅速提高，9月份后基本稳定。因此6-8月是油茶果实及种子迅速生长期。
(2)检测了长林系列4号、40号和166号3个无性系7-10月种子的主要物质组成，结论如下：3个无性系的种子含水率5-10月均处于逐渐下降的趋势，其中8-10月水分含量迅速下降。鲜籽含油率7-10月持续增加，而8-10月份增长幅度最大。蛋白质、淀粉含量与油脂含量的变化规律基本一致，可溶性糖含量各月份变化不大。茶皂素含量随种子成熟不断提高。可见8-10月份是普通油茶种子主要物质积累期。
(3)采用solexa技术对长林4号无性系7-10月份种子进行转录组测序，获得大于200bp的unigene77050条。其中包括脂肪酸合成相关13个酶的99条unigene、脂肪酸链延长相关5个酶的8条unigene、7个脂肪酸去饱和酶的77条unigene，及与角鲨烯和茶皂素合成相关的4个重要酶的34条unigene等。并比较了7-10月份unigenes的RPKM值大小。
(4)筛选了6个主要油茶物种的相对稳定的内参基因。参考其他物种的内参基因，并以RNA-Seq数据为基础，选择种子发育过程中表达量相对稳定的15个基因，用qRT-PCR方法分析15个基因在浙江红花油茶、宛田红花油茶、广宁红花油茶、攸县油茶、小果油茶和普通油茶等6个油茶物种的根、茎、叶、花、种子中表达量的稳定性，结果表明不同物种和不同组织最稳定的内参基因不完全一致，其中TUBα-3、CESA及ACT7α为各物种和组织的表达量相对稳定的内参基因。
(5)以筛选出的CESA为内参，用qRT-PCR方法分析了4个脂类代谢途径29个相关酶基因不同时期的表达量，并进一步研究了在不同时期这些酶基因表达量与其相关代谢产物含量的相关性。比较采用qRT-PCR和RNA-Seq方法分析基因的表达量，结果表明两种方法检测的结果基本一致，但存在细微的差异。例如脂肪酸合成过程中BCCP基因、脂肪酸链延长的ECH基因和脂肪酸去饱和酶FAD8基因。基因表达与代谢相关产物含量变化相关性分析表明脂肪酸合成相关基因表达量变化提前于产物含量变化，而类固醇生物合成代谢中如角鲨烯合酶（SQS）和鲨烯环氧酶（SQE）基因表达量与角鲨烯和甾醇含量变化趋势一致。
(6)克隆了油茶籽油中重要营养成分角鲨烯合成的两个关键酶SQS和SQE基因。采用RACE方法，分离的普通油茶SQS基因cDNA全长1490bp，开放阅读框1245bp，编码414个氨基酸，SQE基因cDNA全长2058bp，开放阅读框1605bp，编码534个氨基酸。采样同源克隆方法，也分离出浙江红花油茶、宛田红花油茶、广宁红花油茶、攸县油茶、小果油茶的SQS和SQE基因，6个油茶物种中SQS和SQE基因cDNA同源性分别达到99.44%和99.47%
(7)利用pMDC32真核表达载体验证了油茶SQS和SQE的功能。采用Gateway技术分别构建了油茶SQS和SQE基因超表达载体pMDC32-SQS和pMDC32-SQE，表达载体通过农杆菌EHA105介导转化到粘红酵母中，利用PCR检测筛选阳性转化子，培养阳性转基因酵母菌株，提取酵母油脂，分析油脂中角鲨烯和甾醇含量。结果表明，转pMDC32-SQS的粘红酵母油脂中角鲨烯和甾醇含量分别低于对照26.26%和38.8%，而转pMDC32-SQE的粘红酵母油脂中甾醇含量比野生型提高202.62%，角鲨烯含量降低了41.09%。但相同的培养时间内，单位体积的培养基中两组转基因粘红酵母培养相同时间后，pMDC32-SQS和pMDC32-SQE的粘红酵母油脂酵母的产量比对照分别提高了81.96%和183.52%，总油脂产量也相应分别提高了69%和175%。
本研究基本明确了油茶果实和种子发育过程中形态、组分及脂类代谢相关基因表达量的变化规律，分析了脂类代谢部分基因的表达量与其相关产物含量变化的关系，为今后更加系统深入的研究油茶种子发育调控打下基础，也为改进育种和栽培措施，提高油茶效益提供理论参考。
Oil-tea Camellia is including important woody species which can product edible oil.Oil-tea Camellia is one of the most important woody oil species in China. The Camelliaoleifera Abel is planted in the most area and has the highest production. Until now, greatprogress were made on production, characters, post processing, oil nutrient content and oilstorage of Camellia oleifera Abel, However, the research on changes of fruits shape, theingredients in seeds and genes expressing of the important characters during development offruit and seeds were insufficient. Studying on the characters of fruits and seeds, thesynthesizing and storing law of the important nutritional ingredient during the development ofseeds can not only help us to promote the important cultivation measure of the Oil-tea Camellia,but also lay the foundation of improving the nutrition value by controlling the ingredients inOil-tea Camellia with molecular techniques. To seek the law of oil-tea Camellia fruits andseeds development, the changes of apparent traits, ingredients, genes expressions amongdifferent development stages of oil-tea Camellia fruits and seeds were studied, and thefollowing were the results.
The fruits and seeds of Changlin No.4, Changlin No.40and Changlin No.166of Camelliaoleifera Abel were collected from May to October, and the changing rules of fruit shape, theweight of fruits and seeds, rate of seeds during the development of the fruits. The resultsshowed that most of the fruits shapes of the three varieties were round olive in May; andchanged greatly in August, and became the characters fruit shape of the variety until September.Fruit grew fastest from June to August, the weight, the diameter and the transverse diameterhad the greatest growing. After September, the weight and volume of the fruits kept constantly.The peel of fruits was changed slightly during the fruit development. The seeds grew continualbigger from June to October unsteadily. The fastest growing months were from June to August.Seed production percentage increased the fastest from June to August, too, and seeds kept steady in September. So it is concluded that the volume and weight of Camellia oleifera Abelfruits and seeds increase fastest during June and August.
Ingredients of three Camellia oleifera Abel varieties seeds were measured from July toOctober. The results showed: The water content decreased gradually among seed developmentfrom May to October, especially from August to October. Oil content increased from July toOctober, especially from August to October. Changes of protein and starch were almost thesame with that of oil content. The soluble sugar showed little change in each month. The teasaponin content increased steadily with the mature of seeds. So it is concluded that theaccumulating stages of carbohydrate in Camellia oleifera Abel seeds were from August toOctober.
RNA-Seq of Camellia oleifera Abel seeds from July to October were sequenced by usingsolexa technology. And in total,77050unigenes longer than200bp were got. There were99unigenes of13enzymes related to4fatty acid synthesis pathways,8unigenes of5enzymesrelated to fatty acid chain elongation,77unigenes of7unsaturated enzymes related tounsaturated fatty acid synthesis pathway and34unigenes of4important enzymes associatedwith squalene and saponin in steroid biosynthesis pathways. The gene expressions wereanalyzed during July and October according to RPKM values of each unigene.
Relatively stable reference genes from6major species of oil-tea camellia were screened.According to reference genes of other species,15relatively stable genes which had stableexpressions in the seeds development were found out from RNA-Seq database. The stability ofthose genes which were expressed in roots, stems, leaves, flowers and seeds of Camelliaoleifera Abel., Camellia meiocarpa Hu, Camellia yuhsienensis Hu, Camellia chekiangoleosaHu, Camellia polyodonta F.C, How and Camellia semiserrata C.W. Chi were analyzed withqRT-PCR. The results showed that the most stable reference genes were different in differenttissues and species. But TUBα-3、CESA and ACT7α were relatively stable among all the tissuesand species.
CESA was used to be reference gene. The expressions of29genes in4lipid metabolismwere analyzed by using qRT-PCR technology. And the relationship between expression ofgenes and their related metabolic prodction in different stages was studied. The expression ofthese genes by using qRT-PCR showed only a little difference compared with the result byusing RNA-Seq technique, such as BCCP, ECH and FAD8. The correlation between geneexpressions and variation of related metabolic production showed that expressions changes ofgenes in fatty acid synthesis, fatty acid chain elongation and unsaturated fatty acid synthesispathways were in advance to the changes of product contents. But the expression changes weresynchronous with that of metabolic production in the steroid biosynthesis pathways.
Squalene synthase (SQS) gene and squalene epoxidase (SQE) gene were cloned fromCamellia oleifera Abel base on RACE technology. Full length of SQS was1490bp and thecoding sequence of SQS gene included2058bp, and encoding422amino acids. The full lengthof SQE genes was2058bp and the coding sequence of SQE gene included1605bp, encoding534amino acids. Then primers were designed according to the sequence of SQS and SQE genenucleotide sequence, and the SQS and SQE genes of Camellia meiocarpa Hu, Camelliayuhsienensis Hu, Camellia chekiangoleosa Hu, Camellia polyodonta F.C. How and Camelliasemiserrata C.W. Chi. were cloned using homology cloning technology. Relationship of SQSgenes and SQE genes of those six species were blasted. The result showed that the cDNAhomology of SQS and SQE in the six oil-tea Camellia species reached up to99.44%and99.47%respectively.
The function of SQS and SQE was verified by pMDC32eukaryotic expression vector. Thegateway over-expression vector pMDC32of SQS and SQE gene in Camellia oleifera Abel wasconstructed. Then they were transformed to the Rhodotorula glutinis（Fres.）Harrison mediatedby agrobacterium EHA105. Then they were detected whether those transformed yeasts werepositive by PCR. Oil of yeasts was extracted after being cultivated, and the amount of squaleneand sterol in the oil were compared with those of wild type. The results showed that thesqualene and sterol content in oil of pMDC32-SQS Rhodotorula glutinis（Fres.）Harrison were reduced by26.26%and38.8%respectively compared with those in wild type. However, thesterol content in oil of the pMDC32-SQE Rhodotorula glutinis (Fres.) Harrison increased by202.62%, the amount of the squalene was reduced by41.09%. The result still showed that aftercultivated in the unit volume substrate in the same period, the production of the yeasts in thetwo kinds of transformed Rhodotorula glutinis (Fres.) Harrison increased by81.96%and183.52%respectively, and the total oil content increased by69%and175%respectively tothose of the wild type.
Changes of morphology, ingredients and gene expression related to lipid metabolismduring development of Camellia oleifera Abel fruits and seeds were defined. and also made therelationship of genes expression and contents of their production clear accroding this paper,which provide a usful theory to find better cultivation methods and breed better varieties.
(1)分析了普通油茶长林系列4号、40号和166号3个无性系5-10月份果实果形、果重和种子的重量、鲜果出籽率等指标在果实生长过程中的变化规律。分析表明，3个无性系5月份果实大部分为圆橄榄形，8月份果形变化较大，到9月份基本分化成各无性系的特征果形；果实6-8月份间果实为迅速膨大期，果实重量和纵、横径大小在此时增长量最大，9月份之后果实重量和大小基本保持恒定；种皮厚度各月变化不大。种子重量和大小6-10月持续增长，其中6-8月份生长速度较快。鲜果出籽率6-8月迅速提高，9月份后基本稳定。因此6-8月是油茶果实及种子迅速生长期。
(2)检测了长林系列4号、40号和166号3个无性系7-10月种子的主要物质组成，结论如下：3个无性系的种子含水率5-10月均处于逐渐下降的趋势，其中8-10月水分含量迅速下降。鲜籽含油率7-10月持续增加，而8-10月份增长幅度最大。蛋白质、淀粉含量与油脂含量的变化规律基本一致，可溶性糖含量各月份变化不大。茶皂素含量随种子成熟不断提高。可见8-10月份是普通油茶种子主要物质积累期。
(3)采用solexa技术对长林4号无性系7-10月份种子进行转录组测序，获得大于200bp的unigene77050条。其中包括脂肪酸合成相关13个酶的99条unigene、脂肪酸链延长相关5个酶的8条unigene、7个脂肪酸去饱和酶的77条unigene，及与角鲨烯和茶皂素合成相关的4个重要酶的34条unigene等。并比较了7-10月份unigenes的RPKM值大小。
(4)筛选了6个主要油茶物种的相对稳定的内参基因。参考其他物种的内参基因，并以RNA-Seq数据为基础，选择种子发育过程中表达量相对稳定的15个基因，用qRT-PCR方法分析15个基因在浙江红花油茶、宛田红花油茶、广宁红花油茶、攸县油茶、小果油茶和普通油茶等6个油茶物种的根、茎、叶、花、种子中表达量的稳定性，结果表明不同物种和不同组织最稳定的内参基因不完全一致，其中TUBα-3、CESA及ACT7α为各物种和组织的表达量相对稳定的内参基因。
(5)以筛选出的CESA为内参，用qRT-PCR方法分析了4个脂类代谢途径29个相关酶基因不同时期的表达量，并进一步研究了在不同时期这些酶基因表达量与其相关代谢产物含量的相关性。比较采用qRT-PCR和RNA-Seq方法分析基因的表达量，结果表明两种方法检测的结果基本一致，但存在细微的差异。例如脂肪酸合成过程中BCCP基因、脂肪酸链延长的ECH基因和脂肪酸去饱和酶FAD8基因。基因表达与代谢相关产物含量变化相关性分析表明脂肪酸合成相关基因表达量变化提前于产物含量变化，而类固醇生物合成代谢中如角鲨烯合酶（SQS）和鲨烯环氧酶（SQE）基因表达量与角鲨烯和甾醇含量变化趋势一致。
(6)克隆了油茶籽油中重要营养成分角鲨烯合成的两个关键酶SQS和SQE基因。采用RACE方法，分离的普通油茶SQS基因cDNA全长1490bp，开放阅读框1245bp，编码414个氨基酸，SQE基因cDNA全长2058bp，开放阅读框1605bp，编码534个氨基酸。采样同源克隆方法，也分离出浙江红花油茶、宛田红花油茶、广宁红花油茶、攸县油茶、小果油茶的SQS和SQE基因，6个油茶物种中SQS和SQE基因cDNA同源性分别达到99.44%和99.47%
(7)利用pMDC32真核表达载体验证了油茶SQS和SQE的功能。采用Gateway技术分别构建了油茶SQS和SQE基因超表达载体pMDC32-SQS和pMDC32-SQE，表达载体通过农杆菌EHA105介导转化到粘红酵母中，利用PCR检测筛选阳性转化子，培养阳性转基因酵母菌株，提取酵母油脂，分析油脂中角鲨烯和甾醇含量。结果表明，转pMDC32-SQS的粘红酵母油脂中角鲨烯和甾醇含量分别低于对照26.26%和38.8%，而转pMDC32-SQE的粘红酵母油脂中甾醇含量比野生型提高202.62%，角鲨烯含量降低了41.09%。但相同的培养时间内，单位体积的培养基中两组转基因粘红酵母培养相同时间后，pMDC32-SQS和pMDC32-SQE的粘红酵母油脂酵母的产量比对照分别提高了81.96%和183.52%，总油脂产量也相应分别提高了69%和175%。
本研究基本明确了油茶果实和种子发育过程中形态、组分及脂类代谢相关基因表达量的变化规律，分析了脂类代谢部分基因的表达量与其相关产物含量变化的关系，为今后更加系统深入的研究油茶种子发育调控打下基础，也为改进育种和栽培措施，提高油茶效益提供理论参考。
Oil-tea Camellia is including important woody species which can product edible oil.Oil-tea Camellia is one of the most important woody oil species in China. The Camelliaoleifera Abel is planted in the most area and has the highest production. Until now, greatprogress were made on production, characters, post processing, oil nutrient content and oilstorage of Camellia oleifera Abel, However, the research on changes of fruits shape, theingredients in seeds and genes expressing of the important characters during development offruit and seeds were insufficient. Studying on the characters of fruits and seeds, thesynthesizing and storing law of the important nutritional ingredient during the development ofseeds can not only help us to promote the important cultivation measure of the Oil-tea Camellia,but also lay the foundation of improving the nutrition value by controlling the ingredients inOil-tea Camellia with molecular techniques. To seek the law of oil-tea Camellia fruits andseeds development, the changes of apparent traits, ingredients, genes expressions amongdifferent development stages of oil-tea Camellia fruits and seeds were studied, and thefollowing were the results.
The fruits and seeds of Changlin No.4, Changlin No.40and Changlin No.166of Camelliaoleifera Abel were collected from May to October, and the changing rules of fruit shape, theweight of fruits and seeds, rate of seeds during the development of the fruits. The resultsshowed that most of the fruits shapes of the three varieties were round olive in May; andchanged greatly in August, and became the characters fruit shape of the variety until September.Fruit grew fastest from June to August, the weight, the diameter and the transverse diameterhad the greatest growing. After September, the weight and volume of the fruits kept constantly.The peel of fruits was changed slightly during the fruit development. The seeds grew continualbigger from June to October unsteadily. The fastest growing months were from June to August.Seed production percentage increased the fastest from June to August, too, and seeds kept steady in September. So it is concluded that the volume and weight of Camellia oleifera Abelfruits and seeds increase fastest during June and August.
Ingredients of three Camellia oleifera Abel varieties seeds were measured from July toOctober. The results showed: The water content decreased gradually among seed developmentfrom May to October, especially from August to October. Oil content increased from July toOctober, especially from August to October. Changes of protein and starch were almost thesame with that of oil content. The soluble sugar showed little change in each month. The teasaponin content increased steadily with the mature of seeds. So it is concluded that theaccumulating stages of carbohydrate in Camellia oleifera Abel seeds were from August toOctober.
RNA-Seq of Camellia oleifera Abel seeds from July to October were sequenced by usingsolexa technology. And in total,77050unigenes longer than200bp were got. There were99unigenes of13enzymes related to4fatty acid synthesis pathways,8unigenes of5enzymesrelated to fatty acid chain elongation,77unigenes of7unsaturated enzymes related tounsaturated fatty acid synthesis pathway and34unigenes of4important enzymes associatedwith squalene and saponin in steroid biosynthesis pathways. The gene expressions wereanalyzed during July and October according to RPKM values of each unigene.
Relatively stable reference genes from6major species of oil-tea camellia were screened.According to reference genes of other species,15relatively stable genes which had stableexpressions in the seeds development were found out from RNA-Seq database. The stability ofthose genes which were expressed in roots, stems, leaves, flowers and seeds of Camelliaoleifera Abel., Camellia meiocarpa Hu, Camellia yuhsienensis Hu, Camellia chekiangoleosaHu, Camellia polyodonta F.C, How and Camellia semiserrata C.W. Chi were analyzed withqRT-PCR. The results showed that the most stable reference genes were different in differenttissues and species. But TUBα-3、CESA and ACT7α were relatively stable among all the tissuesand species.
CESA was used to be reference gene. The expressions of29genes in4lipid metabolismwere analyzed by using qRT-PCR technology. And the relationship between expression ofgenes and their related metabolic prodction in different stages was studied. The expression ofthese genes by using qRT-PCR showed only a little difference compared with the result byusing RNA-Seq technique, such as BCCP, ECH and FAD8. The correlation between geneexpressions and variation of related metabolic production showed that expressions changes ofgenes in fatty acid synthesis, fatty acid chain elongation and unsaturated fatty acid synthesispathways were in advance to the changes of product contents. But the expression changes weresynchronous with that of metabolic production in the steroid biosynthesis pathways.
Squalene synthase (SQS) gene and squalene epoxidase (SQE) gene were cloned fromCamellia oleifera Abel base on RACE technology. Full length of SQS was1490bp and thecoding sequence of SQS gene included2058bp, and encoding422amino acids. The full lengthof SQE genes was2058bp and the coding sequence of SQE gene included1605bp, encoding534amino acids. Then primers were designed according to the sequence of SQS and SQE genenucleotide sequence, and the SQS and SQE genes of Camellia meiocarpa Hu, Camelliayuhsienensis Hu, Camellia chekiangoleosa Hu, Camellia polyodonta F.C. How and Camelliasemiserrata C.W. Chi. were cloned using homology cloning technology. Relationship of SQSgenes and SQE genes of those six species were blasted. The result showed that the cDNAhomology of SQS and SQE in the six oil-tea Camellia species reached up to99.44%and99.47%respectively.
The function of SQS and SQE was verified by pMDC32eukaryotic expression vector. Thegateway over-expression vector pMDC32of SQS and SQE gene in Camellia oleifera Abel wasconstructed. Then they were transformed to the Rhodotorula glutinis（Fres.）Harrison mediatedby agrobacterium EHA105. Then they were detected whether those transformed yeasts werepositive by PCR. Oil of yeasts was extracted after being cultivated, and the amount of squaleneand sterol in the oil were compared with those of wild type. The results showed that thesqualene and sterol content in oil of pMDC32-SQS Rhodotorula glutinis（Fres.）Harrison were reduced by26.26%and38.8%respectively compared with those in wild type. However, thesterol content in oil of the pMDC32-SQE Rhodotorula glutinis (Fres.) Harrison increased by202.62%, the amount of the squalene was reduced by41.09%. The result still showed that aftercultivated in the unit volume substrate in the same period, the production of the yeasts in thetwo kinds of transformed Rhodotorula glutinis (Fres.) Harrison increased by81.96%and183.52%respectively, and the total oil content increased by69%and175%respectively tothose of the wild type.
Changes of morphology, ingredients and gene expression related to lipid metabolismduring development of Camellia oleifera Abel fruits and seeds were defined. and also made therelationship of genes expression and contents of their production clear accroding this paper,which provide a usful theory to find better cultivation methods and breed better varieties.
引文
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