Article(id=1277239985942168097, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277239982603502113, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.05.008, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1730908800000, receivedDateStr=2024-11-07, revisedDate=null, revisedDateStr=null, acceptedDate=1736179200000, acceptedDateStr=2025-01-07, onlineDate=1782447342620, onlineDateStr=2026-06-26, pubDate=1748102400000, pubDateStr=2025-05-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782447342620, onlineIssueDateStr=2026-06-26, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782447342620, creator=13701087609, updateTime=1782447342620, updator=13701087609, issue=Issue{id=1277239982603502113, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='5', pageStart='1025', pageEnd='1277', issueExtLink='null', onlineDate='null', pubDate='1748102400000', pubDateStr='2025-05-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782447341824, creator='13701087609', updateTime=1782447947315, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1277242522292319215, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277239982603502113, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1277242522292319216, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277239982603502113, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1094, endPage=1106, ext={EN=ArticleExt(id=1277239987632472611, articleId=1277239985942168097, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Characteristics and Transcriptome Analysis of High Temperature Albino Tea Variety Yaoqiu, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

In order to explore the characteristics and molecular regulation mechanism of high temperature albinism of tea variety Yaoqiu (YQ) and its molecular regulation mechanism, the effects of low temperature albinism on the shoot with one bud and two leaves of tea variety YQ in spring (YQC), summer (YQX), autumn albinism (YQQ) and autumn green (YQQL), and Baiye No. 1 (BY1H) in spring albinism (BY1HC), their biochemical components, and transcriptome sequencing were carried out on the new shoots in summer (BY1HX) and autumn (BY1HQ). The results of biochemical analysis showed that the total free amino acid content and tea polyphenols content of YQQ albino fresh leaves were lower than those of YQC green fresh leaves, and the difference of total free amino acid content was significant; while the total free amino acid content of BY1HC albino fresh leaves was significantly higher than that of BY1HQ green fresh leaves, and the tea polyphenols content was significantly lower than that of BY1HQ green fresh leaves. Cluster analysis revealed that the gene expression profiles of YQQ albino fresh leaves formed a distinct cluster. The comparison between YQQ and BY1HQ exhibited the highest number of differentially expressed genes. The new shoot leaves of YQ showed an albino phenotype induced by high-temperature stress in late summer and early autumn. This stress affected the structure of microtubules, cells, and subcellular components, leading to a reduction in the contents of amino acids and polyphenols. The three functional categories exhibiting the most significant gene enrichment were microtubule binding, movement of cellular or subcellular components, and non-membrane-bound organelles. Similarly, the top three pathways with the highest significance in gene enrichment were the flavonoid biosynthesis pathway, the starch and sucrose metabolism pathway, and the amino sugar and nucleotide sugar metabolism pathway. Through gene co-expression network analysis, modules exhibiting high expression levels were identified. Subsequently, the ten genes demonstrating the highest total connectivity (K value) were selected for further analysis and annotation. Notably, the functions of genes TGY042732 and the novel gene designated as novel.276 could not be predicted. These findings would provide a theoretical foundation for understanding the phenomenon of albinism in tea resources under conditions of elevated temperature.

, authors=null, authorsList=Haitao HUANG, Yi DING, Xiaojun NIU, Yun ZHAO, Jizhong YU, authorCompany=null, correspAuthors=Jizhong YU, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1277239997300343349, articleId=1277239985942168097, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=高温白化茶树品种曜秋的特性和转录组分析, columnId=1236256430517440904, journalTitle=热带作物学报, columnName=组学与生物技术, runingTitle=null, highlight=null, articleAbstract=

为探究茶树品种曜秋(YQ)高温白化的特性及其分子调控机制,以低温白化茶树品种白叶1号(BY1H)为对照,对茶树品种曜秋的春季(YQC)、夏季(YQX)、秋季白化(YQQ)和秋季绿色(YQQL)的一芽二叶新梢,以及白叶1号的春季白化(BY1HC)、夏季(BY1HX)和秋季(BY1HQ)的新梢为材料进行生化成分检测、转录组测序和分析。生化分析结果表明,YQQ白化鲜叶游离氨基酸总量和茶多酚含量均低于YQC绿色鲜叶,且游离氨基酸总量差异达显著水平;而BY1HC白化鲜叶游离氨基酸总量显著高于BY1HQ绿色鲜叶,茶多酚含量显著低于BY1HQ绿色鲜叶。转录组主成分分析结果表明,YQQ白化鲜叶基因表达信息独立于第二象限,明显与其他样品不同;聚类分析结果表明,YQQ白化鲜叶基因表达信息单独聚为一类;YQQ和BY1HQ的差异表达基因数量最多,曜秋在夏末秋初高温响应下新梢叶片白化,微管、细胞、亚细胞等结构受到影响,氨基酸、多酚等主要功能性理化成分含量均降低,差异表达基因富集显著性最高的3个功能类依次为微管结合、细胞或亚细胞成分转移和非膜结合细胞器,差异表达基因富集显著性排名前3的通路依次为类黄酮生物合成通路、淀粉和蔗糖代谢通路以及氨基糖和核苷酸糖代谢通路;基因共表达网络分析筛选出高表达模块,并对总连通性K值最高的10个基因进行分析和注释,其中TGY042732和未知基因novel.276的功能未能预测。本研究结果为了解茶树资源高温白化提供一定的理论基础。

, authors=

黄海涛(1981—),男,硕士,高级农艺师,研究方向:茶树遗传育种。

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* 余继忠(YU Jizhong),E-mail:
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黄海涛(1981—),男,硕士,高级农艺师,研究方向:茶树遗传育种。

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黄海涛(1981—),男,硕士,高级农艺师,研究方向:茶树遗传育种。

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(in Chinese), articleTitle=Phenotypic characterization and gene mapping of a thermo-sensitive albino leaf mutant tsa1 in rice, refAbstract=null), Reference(id=1277240045971047066, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, doi=null, pmid=null, pmcid=null, year=2000, volume=36, issue=4, pageStart=300, pageEnd=304, url=null, language=null, rfNumber=[18], rfOrder=27, authorNames=成浩, 陈明, 虞富莲, 李素芳, journalName=植物生理学通讯, refType=null, unstructuredReference=成浩, 陈明, 虞富莲, 李素芳. 茶叶片阶段性返白过程中色素蛋白复合体的变化[J]. 植物生理学通讯, 2000, 36(4): 300-304., articleTitle=茶叶片阶段性返白过程中色素蛋白复合体的变化, refAbstract=null), Reference(id=1277240046365311643, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, doi=null, pmid=null, pmcid=null, year=2000, volume=36, issue=4, pageStart=300, pageEnd=304, url=null, language=null, rfNumber=[18], rfOrder=28, authorNames=CHENG H, CHEN M, YU F L, LI S F, journalName=Plant Physiology Communications, refType=null, unstructuredReference=CHENG H, CHEN M, YU F L, LI S F. The variation of pigment-protein complexes in the albescent stage of tea[J]. Plant Physiology Communications, 2000, 36(4): 300-304. (in Chinese), articleTitle=The variation of pigment-protein complexes in the albescent stage of tea, refAbstract=null), Reference(id=1277240046759576220, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, doi=null, pmid=null, pmcid=null, year=2015, volume=10, issue=10, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[19], rfOrder=29, authorNames=LI C F, YAO M Z, MA C L, MA J Q, JIN J Q, CHEN L, journalName=PLoS One, refType=null, unstructuredReference=LI C F, YAO M Z, MA C L, MA J Q, JIN J Q, CHEN L. Differential metabolic profiles during the albescent stages of ‘Anji Baicha’ (Camellia sinensis)[J]. PLoS One, 2015, 10(10): e0139996., articleTitle=Differential metabolic profiles during the albescent stages of ‘Anji Baicha’ (Camellia sinensis), refAbstract=null)], funds=[Fund(id=1277240033379746427, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, awardId=2021C02067-4, language=CN, fundingSource=浙江省“十四五”农业新品种选育重大科技专项(2021C02067-4), fundOrder=null, country=null), Fund(id=1277240033799176828, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, awardId=CARS19, language=CN, fundingSource=国家茶叶产业技术体系项目(CARS19), fundOrder=null, country=null), Fund(id=1277240034298299005, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, awardId=2022HNCT-04, language=CN, fundingSource=杭州市农科院科技创新与示范推广基金项目(2022HNCT-04), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1277239998558634551, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, xref=null, ext=[AuthorCompanyExt(id=1277239998919344696, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, companyId=1277239998558634551, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Tea Research Institute, Hangzhou Academy of Agricultural Sciences, Hangzhou, Zhejiang 310024, China), AuthorCompanyExt(id=1277240000303465017, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, companyId=1277239998558634551, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=杭州市农业科学研究院茶叶研究所,浙江杭州 310024)])], figs=[ArticleFig(id=1277240019953779295, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 1, caption=Shoot color of YQ and BY1H at different time

A, C: New shoots; B, D: The second leaf of new shoots.

, figureFileSmall=lqCgysXzZdLTmnocgoC2tg==, figureFileBig=ptuzKPoclAXeYpmX8e/eeA==, tableContent=null), ArticleFig(id=1277240020331266656, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图1, caption=曜秋和白叶1号不同时间的新梢颜色

A、C:一芽二叶新梢;B、D:一芽二叶新梢的第二叶。

, figureFileSmall=lqCgysXzZdLTmnocgoC2tg==, figureFileBig=ptuzKPoclAXeYpmX8e/eeA==, tableContent=null), ArticleFig(id=1277240021203681889, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 2, caption=BY1HC leaves turn green in late spring and YQQ leaves apoptosis in early autumn

A: White leaves turning green in spring in BY1HC; B: Top shoots turning green in autumn in YQ; C: White leaves apoptosis in autumn in YQQ.

, figureFileSmall=k3OwWamLY8bCMIHUFed00g==, figureFileBig=6y4CWfrEGJCIOhiyz/Q3rQ==, tableContent=null), ArticleFig(id=1277240022860431970, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图2, caption=白叶1号春末白化叶片转绿和曜秋秋初白化叶片凋亡

A:白叶1号春季白化叶片转绿;B:曜秋秋季顶梢转绿;C:曜秋秋季白化叶片凋亡。

, figureFileSmall=k3OwWamLY8bCMIHUFed00g==, figureFileBig=6y4CWfrEGJCIOhiyz/Q3rQ==, tableContent=null), ArticleFig(id=1277240023279862371, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 3, caption=Principal component analysis, figureFileSmall=0fEyzKLELUbsms4UZUaiJg==, figureFileBig=pA47+sI3FVlXUgOFJ6nBsw==, tableContent=null), ArticleFig(id=1277240023359554148, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图3, caption=主成分分析, figureFileSmall=0fEyzKLELUbsms4UZUaiJg==, figureFileBig=pA47+sI3FVlXUgOFJ6nBsw==, tableContent=null), ArticleFig(id=1277240023674126949, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 4, caption=Differential gene expression, figureFileSmall=D1/DtJMC3UYn3szBYxhQVw==, figureFileBig=cq/yhbhfNSEo415iiYg36g==, tableContent=null), ArticleFig(id=1277240023749624422, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图4, caption=差异基因表达量, figureFileSmall=D1/DtJMC3UYn3szBYxhQVw==, figureFileBig=cq/yhbhfNSEo415iiYg36g==, tableContent=null), ArticleFig(id=1277240024160666215, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 5, caption=Cluster diagram of differential genes, figureFileSmall=aX2YfE9HiUrhCOJ5rEeSSw==, figureFileBig=P59UG2gEQ1cOwSXnn5gjrA==, tableContent=null), ArticleFig(id=1277240024504599144, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图5, caption=差异基因聚类图, figureFileSmall=aX2YfE9HiUrhCOJ5rEeSSw==, figureFileBig=P59UG2gEQ1cOwSXnn5gjrA==, tableContent=null), ArticleFig(id=1277240024596873833, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 6, caption=GO (functional classification) enrichment analysis of differential genes

A: YQC vs BY1HC; B: YQX vs BY1HX; C: YQQ vs BY1HQ; D: YQQ vs YQQL.

, figureFileSmall=/aZ0ysCsoT72X3XJ9M1MlA==, figureFileBig=wyG4FhJ30Nk0qBwYChRgaA==, tableContent=null), ArticleFig(id=1277240024940806762, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图6, caption=差异基因GO(功能分类)富集分析

A:YQC与BY1HC;B:YQX与BY1HX;C:YQQ与BY1HQ;D:YQQ与YQQL。

, figureFileSmall=/aZ0ysCsoT72X3XJ9M1MlA==, figureFileBig=wyG4FhJ30Nk0qBwYChRgaA==, tableContent=null), ArticleFig(id=1277240025356042859, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 7, caption=Enrichment analysis of KEGG (metabolic pathway)

A: YQC vs BY1HC; B: YQX vs BY1HX; C: YQQ vs BY1HQ; D: YQQ vs YQQL.

, figureFileSmall=4JxyW4IYeIPI7/Bb9u02xA==, figureFileBig=QoZ0R3Fj3bMQ0S9AdcCSeg==, tableContent=null), ArticleFig(id=1277240025779667564, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图7, caption=差异基因KEGG(代谢通路)富集分析

A:YQC与BY1HC;B:YQX与BY1HX;C:YQQ与BY1HQ;D:YQQ与YQQL。

, figureFileSmall=4JxyW4IYeIPI7/Bb9u02xA==, figureFileBig=QoZ0R3Fj3bMQ0S9AdcCSeg==, tableContent=null), ArticleFig(id=1277240027461583469, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Fig. 8, caption=Network heatmap plot for selected genes, figureFileSmall=LHVDp6Paa01yt041niCrEg==, figureFileBig=8g033J8VMfDnPIOZkOfjNw==, tableContent=null), ArticleFig(id=1277240027524498030, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=图8, caption=模块基因聚类热图, figureFileSmall=LHVDp6Paa01yt041niCrEg==, figureFileBig=8g033J8VMfDnPIOZkOfjNw==, tableContent=null), ArticleFig(id=1277240027876819567, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 1, caption=

Characteristics of YQ and BY1H

, figureFileSmall=null, figureFileBig=null, tableContent=
品种Variety春季新梢颜色Color of new shoots in spring夏初新梢颜色Color of new shoots in early summer夏季中后期至秋初新梢颜色Color of new shoots from mid summer to early autumn秋季新梢颜色Color of new shoots in autumn性状Characteristic
YQ黄绿色紫绿色黄色绿色中等椭圆、叶长7.8 cm、叶宽3.2 cm、浅绿、上表皮隆起、叶缘波状、叶身平、叶齿密浅锐、叶尖渐尖、叶基楔形
BY1H白色绿色绿色绿色长椭圆、叶长8.8 cm、叶宽3.4 cm、绿色、上表皮平、叶缘平、叶身平、叶齿稀浅钝、叶尖渐尖、叶基楔形
), ArticleFig(id=1277240027952317040, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表1, caption=

曜秋和白叶1号的特性

, figureFileSmall=null, figureFileBig=null, tableContent=
品种Variety春季新梢颜色Color of new shoots in spring夏初新梢颜色Color of new shoots in early summer夏季中后期至秋初新梢颜色Color of new shoots from mid summer to early autumn秋季新梢颜色Color of new shoots in autumn性状Characteristic
YQ黄绿色紫绿色黄色绿色中等椭圆、叶长7.8 cm、叶宽3.2 cm、浅绿、上表皮隆起、叶缘波状、叶身平、叶齿密浅锐、叶尖渐尖、叶基楔形
BY1H白色绿色绿色绿色长椭圆、叶长8.8 cm、叶宽3.4 cm、绿色、上表皮平、叶缘平、叶身平、叶齿稀浅钝、叶尖渐尖、叶基楔形
), ArticleFig(id=1277240028296249969, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 2, caption=

Whitening stage and temperature of YQ

, figureFileSmall=null, figureFileBig=null, tableContent=
观测年份Observation year白化起始日期Albinism start date最高气温高于35 ℃日期Date of maximum daily temperature above 35 ℃新梢转绿日期Albinism end date最高气温低于30 ℃日期Date of maximum daily temperature below 30 ℃
20208月1日7月12日9月20日9月11日
20217月10日7月5日8月6日7月24日
20227月10日7月3日9月2日8月27日
20238月10日8月5日9月4日8月29日
), ArticleFig(id=1277240028401107570, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表2, caption=

曜秋白化阶段与气温

, figureFileSmall=null, figureFileBig=null, tableContent=
观测年份Observation year白化起始日期Albinism start date最高气温高于35 ℃日期Date of maximum daily temperature above 35 ℃新梢转绿日期Albinism end date最高气温低于30 ℃日期Date of maximum daily temperature below 30 ℃
20208月1日7月12日9月20日9月11日
20217月10日7月5日8月6日7月24日
20227月10日7月3日9月2日8月27日
20238月10日8月5日9月4日8月29日
), ArticleFig(id=1277240028782789235, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 3, caption=

Comparison of biochemical compositions

, figureFileSmall=null, figureFileBig=null, tableContent=
样品Sample水浸出物含量Water extract content/%茶多酚含量TP content/%游离氨基酸总量Total AA content/%咖啡碱含量Caffeine content/%GC/%EGC/%
YQC41.71±0.33d13.55±0.35c4.89±0.02b2.57±0.00c0.59±0.10c0.83±0.02d
YQX45.89±0.03b15.36±0.22a3.82±0.04d2.54±0.03c1.59±0.09a2.64±0.21a
YQQ39.04±0.20e13.41±0.18c3.73±0.01d2.12±0.03d0.50±0.01c0.58±0.02e
YQQL37.87±0.10f14.8±0.09b4.41±0.04c1.94±0.02e0.31±0.01d0.75±0.04d
BY1HC49.08±0.15a12.06±0.09d5.31±0.08a3.11±0.00b1.65±0.00a2.33±0.10a
BY1HX49.24±0.03a15.76±0.04a3.25±0.01e3.27±0.01a1.27±0.00b2.01±0.01b
BY1HQ43.06±0.01c14.47±0.13b3.12±0.01f1.95±0.01e1.27±0.03b1.65±0.04c
样品SampleC/%EGCG/%EC/%GCG/%ECG/%CG/%
YQC0.18±0.01c5.22±0.01d0.34±0.00d2.51±0.00c0.77±0.00e0.24±0.00b
YQX0.42±0.11b6.22±0.01b1.09±0.13b2.64±0.28b1.64±0.14b0.43±0.13a
YQQ0.21±0.00c3.24±0.02g0.29±0.01d0.73±0.03f0.84±0.02d0.11±0.01c
YQQL0.15±0.00d4.77±0.07f0.33±0.01d0.76±0.16f0.85±0.01d0.09±0.01c
BY1HC0.59±0.01a5.02±0.01e1.46±0.02a1.86±0.00d1.38±0.00c0.29±0.00a
BY1HX0.53±0.02a8.37±0.01a1.19±0.00b3.16±0.00a1.92±0.01a0.58±0.00a
BY1HQ0.41±0.01b5.61±0.05c0.82±0.00c1.52±0.03e1.61±0.01b0.22±0.00b
), ArticleFig(id=1277240029139305076, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表3, caption=

生化组成比较

, figureFileSmall=null, figureFileBig=null, tableContent=
样品Sample水浸出物含量Water extract content/%茶多酚含量TP content/%游离氨基酸总量Total AA content/%咖啡碱含量Caffeine content/%GC/%EGC/%
YQC41.71±0.33d13.55±0.35c4.89±0.02b2.57±0.00c0.59±0.10c0.83±0.02d
YQX45.89±0.03b15.36±0.22a3.82±0.04d2.54±0.03c1.59±0.09a2.64±0.21a
YQQ39.04±0.20e13.41±0.18c3.73±0.01d2.12±0.03d0.50±0.01c0.58±0.02e
YQQL37.87±0.10f14.8±0.09b4.41±0.04c1.94±0.02e0.31±0.01d0.75±0.04d
BY1HC49.08±0.15a12.06±0.09d5.31±0.08a3.11±0.00b1.65±0.00a2.33±0.10a
BY1HX49.24±0.03a15.76±0.04a3.25±0.01e3.27±0.01a1.27±0.00b2.01±0.01b
BY1HQ43.06±0.01c14.47±0.13b3.12±0.01f1.95±0.01e1.27±0.03b1.65±0.04c
样品SampleC/%EGCG/%EC/%GCG/%ECG/%CG/%
YQC0.18±0.01c5.22±0.01d0.34±0.00d2.51±0.00c0.77±0.00e0.24±0.00b
YQX0.42±0.11b6.22±0.01b1.09±0.13b2.64±0.28b1.64±0.14b0.43±0.13a
YQQ0.21±0.00c3.24±0.02g0.29±0.01d0.73±0.03f0.84±0.02d0.11±0.01c
YQQL0.15±0.00d4.77±0.07f0.33±0.01d0.76±0.16f0.85±0.01d0.09±0.01c
BY1HC0.59±0.01a5.02±0.01e1.46±0.02a1.86±0.00d1.38±0.00c0.29±0.00a
BY1HX0.53±0.02a8.37±0.01a1.19±0.00b3.16±0.00a1.92±0.01a0.58±0.00a
BY1HQ0.41±0.01b5.61±0.05c0.82±0.00c1.52±0.03e1.61±0.01b0.22±0.00b
), ArticleFig(id=1277240029218996853, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 4, caption=

Amino acid composition

, figureFileSmall=null, figureFileBig=null, tableContent=
氨基酸Amino acid含量Content/%
YQCYQQBY1HCBY1HQ
天冬氨酸0.25±0.01b0.38±0.02a0.37±0.00a0.09±0.01c
苏氨酸0.03±0.000.02±0.000.02±0.00
丝氨酸0.11±0.00a0.06±0.00b0.13±0.00a0.07±0.00b
天冬酰胺0.05±0.000.05±0.00
谷氨酸0.33±0.01c0.59±0.04a0.44±0.00b0.18±0.02d
谷氨酰氨0.08±0.00b0.65±0.01a0.10±0.00b
茶氨酸2.20±0.05b1.15±0.03d2.47±0.02a1.66±0.02c
脯氨酸0.02±0.000.01±0.00
甘氨酸0.01±0.000.01±0.00
丙氨酸0.06±0.00b0.04±0.00b0.10±0.00a0.03±0.00b
缬氨酸0.04±0.000.04±0.00
亮氨酸0.02±0.00
γ-氨基丁酸0.01±0.00c0.01±0.00c0.04±0.00b0.20±0.01a
组氨酸0.02±0.00
精氨酸0.29±0.03b0.30±0.02b0.82±0.02a0.07±0.00c
), ArticleFig(id=1277240029957194358, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表4, caption=

氨基酸组成

, figureFileSmall=null, figureFileBig=null, tableContent=
氨基酸Amino acid含量Content/%
YQCYQQBY1HCBY1HQ
天冬氨酸0.25±0.01b0.38±0.02a0.37±0.00a0.09±0.01c
苏氨酸0.03±0.000.02±0.000.02±0.00
丝氨酸0.11±0.00a0.06±0.00b0.13±0.00a0.07±0.00b
天冬酰胺0.05±0.000.05±0.00
谷氨酸0.33±0.01c0.59±0.04a0.44±0.00b0.18±0.02d
谷氨酰氨0.08±0.00b0.65±0.01a0.10±0.00b
茶氨酸2.20±0.05b1.15±0.03d2.47±0.02a1.66±0.02c
脯氨酸0.02±0.000.01±0.00
甘氨酸0.01±0.000.01±0.00
丙氨酸0.06±0.00b0.04±0.00b0.10±0.00a0.03±0.00b
缬氨酸0.04±0.000.04±0.00
亮氨酸0.02±0.00
γ-氨基丁酸0.01±0.00c0.01±0.00c0.04±0.00b0.20±0.01a
组氨酸0.02±0.00
精氨酸0.29±0.03b0.30±0.02b0.82±0.02a0.07±0.00c
), ArticleFig(id=1277240030062051959, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 5, caption=

Transcriptome sequencing data quality

, figureFileSmall=null, figureFileBig=null, tableContent=
样品Sample总读数Number of raw reads过滤读数(占比)Number of clean reads (percentage/%)比对读数(占比)Number of reads on the genome (proportion/%)Q30占比Q30 ratio/%GC占比GC percentage/%
YQC3_147 809 65647 056 592 (98.4)40 761 590 (86.62)91.7043.91
YQC3_245 531 73245 012 482 (98.9)39 856 326 (88.55)92.3044.02
YQC3_345 153 64644 572 166 (98.7)38 794 378 (87.04)91.8343.67
BY1HC3_146 340 04445 821 166 (98.9)40 437 081 (88.25)92.4543.55
BY1HC3_247 036 20846 570 078 (99.0)41 134 343 (88.33)92.4443.65
BY1HC3_345 568 54045 114 320 (99.0)39 529 694 (87.62)91.6943.35
YQX3_147 737 61447 248 972 (99.0)41 875 021 (88.63)92.3644.00
YQX3_246 464 07446 094 356 (99.2)40 840 242 (88.60)91.8043.74
YQX3_346 179 54445 764 578 (99.1)40 520 396 (88.54)92.1544.08
BY1HX3_148 921 21248 304 384 (98.7)43 308 202 (89.66)92.6244.30
BY1HX3_245 791 15845 167 186 (98.6)39 925 265 (88.39)92.6144.01
BY1HX3_347 029 02446 407 440 (98.7)40 932 320 (88.2)92.1143.70
YQQ3_146 266 16645 650 008 (98.7)40 463 720 (88.64)92.2243.55
YQQ3_246 001 42645 534 716 (99.0)40 718 255 (89.42)91.9543.64
YQQ3_346 191 54645 708 558 (99.0)40 598 857 (88.82)92.3743.6
BY1HQ3_145 75340245 277 448 (99.0)40 168 038 (88.72)92.2943.96
BY1HQ3_246 565 81646 155 276 (99.1)40 993 795 (88.82)92.0944.03
BY1HQ3_346 689 87046 382 642 (99.3)41 149 806 (88.72)91.6544.00
YQQL3_146 431 85045 994 034 (99.1)40 863 964 (88.85)92.6243.84
YQQL3_247 464 88847 053 100 (99.1)41 994 693 (89.25)92.2543.85
YQQL3_345 226 90644 681 450 (98.8)39 832 883 (89.15)91.9543.92
), ArticleFig(id=1277240032104678008, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表5, caption=

转录组测序数据质量

, figureFileSmall=null, figureFileBig=null, tableContent=
样品Sample总读数Number of raw reads过滤读数(占比)Number of clean reads (percentage/%)比对读数(占比)Number of reads on the genome (proportion/%)Q30占比Q30 ratio/%GC占比GC percentage/%
YQC3_147 809 65647 056 592 (98.4)40 761 590 (86.62)91.7043.91
YQC3_245 531 73245 012 482 (98.9)39 856 326 (88.55)92.3044.02
YQC3_345 153 64644 572 166 (98.7)38 794 378 (87.04)91.8343.67
BY1HC3_146 340 04445 821 166 (98.9)40 437 081 (88.25)92.4543.55
BY1HC3_247 036 20846 570 078 (99.0)41 134 343 (88.33)92.4443.65
BY1HC3_345 568 54045 114 320 (99.0)39 529 694 (87.62)91.6943.35
YQX3_147 737 61447 248 972 (99.0)41 875 021 (88.63)92.3644.00
YQX3_246 464 07446 094 356 (99.2)40 840 242 (88.60)91.8043.74
YQX3_346 179 54445 764 578 (99.1)40 520 396 (88.54)92.1544.08
BY1HX3_148 921 21248 304 384 (98.7)43 308 202 (89.66)92.6244.30
BY1HX3_245 791 15845 167 186 (98.6)39 925 265 (88.39)92.6144.01
BY1HX3_347 029 02446 407 440 (98.7)40 932 320 (88.2)92.1143.70
YQQ3_146 266 16645 650 008 (98.7)40 463 720 (88.64)92.2243.55
YQQ3_246 001 42645 534 716 (99.0)40 718 255 (89.42)91.9543.64
YQQ3_346 191 54645 708 558 (99.0)40 598 857 (88.82)92.3743.6
BY1HQ3_145 75340245 277 448 (99.0)40 168 038 (88.72)92.2943.96
BY1HQ3_246 565 81646 155 276 (99.1)40 993 795 (88.82)92.0944.03
BY1HQ3_346 689 87046 382 642 (99.3)41 149 806 (88.72)91.6544.00
YQQL3_146 431 85045 994 034 (99.1)40 863 964 (88.85)92.6243.84
YQQL3_247 464 88847 053 100 (99.1)41 994 693 (89.25)92.2543.85
YQQL3_345 226 90644 681 450 (98.8)39 832 883 (89.15)91.9543.92
), ArticleFig(id=1277240032494748281, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=EN, label=Tab. 6, caption=

Core gene analysis

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基因ID Gene ID总连通性K total基因位置Gene chr基因起点Gene start基因终点Gene end基因链Gene strand基因长度Gene length基因功能描述Gene description基因家族Gene family
TGY0427322799.45GWHASIV0000000533 390 55633 397 5681195
TGY1098582776.93GWHASIV0000001390 289 55490 301 362+2571Alkaline/neutral invertase E, chloroplastic;
TGY0468802755.22GWHASIV00000005137 642 465137 652 3273224CRM-domain containing factorCFM3, chloroplastic/mitochondrial;
TGY1074492751.79GWHASIV0000001330 855 42630 881 948+2182GTP-binding protein TypA/BipA homologGTP_EFTU
TGY0146232739.66GWHASIV0000000264 692 44364 698 1291247Ribulose-phosphate 3-epimerase, chloroplastic;
TGY0964572734.73GWHASIV0000001173 083 01673 086 372+1723Ribulose bisphosphate carboxylase/oxygenase activase 1, chloroplastic;AAA
novel.2762734.52GWHASIV00000001182 916 565182 925 699+849
TGY1232902727.31GWHASIV0000001553 970 37153 989 385+3334ATP-dependent Clp protease ATP-binding subunit ClpA homolog CD4B, chloroplastic;Clp_N
TGY0501752724.96GWHASIV00000005212 083 578212 087 229+1343Pyruvate dehydrogenase (acetyl-transferring)] kinase, mitochondrial;HATPase_c
TGY0882522724.36GWHASIV0000001062 857 06562 907 8802643Uncharacterized aarF domain-containing protein kinase At4g31390, chloroplastic;ABC1
), ArticleFig(id=1277240032582828666, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277239985942168097, language=CN, label=表6, caption=

核心基因分析

, figureFileSmall=null, figureFileBig=null, tableContent=
基因ID Gene ID总连通性K total基因位置Gene chr基因起点Gene start基因终点Gene end基因链Gene strand基因长度Gene length基因功能描述Gene description基因家族Gene family
TGY0427322799.45GWHASIV0000000533 390 55633 397 5681195
TGY1098582776.93GWHASIV0000001390 289 55490 301 362+2571Alkaline/neutral invertase E, chloroplastic;
TGY0468802755.22GWHASIV00000005137 642 465137 652 3273224CRM-domain containing factorCFM3, chloroplastic/mitochondrial;
TGY1074492751.79GWHASIV0000001330 855 42630 881 948+2182GTP-binding protein TypA/BipA homologGTP_EFTU
TGY0146232739.66GWHASIV0000000264 692 44364 698 1291247Ribulose-phosphate 3-epimerase, chloroplastic;
TGY0964572734.73GWHASIV0000001173 083 01673 086 372+1723Ribulose bisphosphate carboxylase/oxygenase activase 1, chloroplastic;AAA
novel.2762734.52GWHASIV00000001182 916 565182 925 699+849
TGY1232902727.31GWHASIV0000001553 970 37153 989 385+3334ATP-dependent Clp protease ATP-binding subunit ClpA homolog CD4B, chloroplastic;Clp_N
TGY0501752724.96GWHASIV00000005212 083 578212 087 229+1343Pyruvate dehydrogenase (acetyl-transferring)] kinase, mitochondrial;HATPase_c
TGY0882522724.36GWHASIV0000001062 857 06562 907 8802643Uncharacterized aarF domain-containing protein kinase At4g31390, chloroplastic;ABC1
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高温白化茶树品种曜秋的特性和转录组分析
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黄海涛 , 丁一 , 牛小军 , 赵芸 , 余继忠 *
热带作物学报 | 组学与生物技术 2025,46(5): 1094-1106
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热带作物学报 |组学与生物技术 2025 , 46 (5) : 1094 -1106
高温白化茶树品种曜秋的特性和转录组分析
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黄海涛(1981—),男,硕士,高级农艺师,研究方向:茶树遗传育种。

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黄海涛(1981—),男,硕士,高级农艺师,研究方向:茶树遗传育种。

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黄海涛, 丁一, 牛小军, 赵芸, 余继忠*
作者信息
  • 杭州市农业科学研究院茶叶研究所,浙江杭州 310024
通讯作者:
* 余继忠(YU Jizhong),E-mail:
Characteristics and Transcriptome Analysis of High Temperature Albino Tea Variety Yaoqiu
Haitao HUANG, Yi DING, Xiaojun NIU, Yun ZHAO, Jizhong YU*
Affiliations
  • Tea Research Institute, Hangzhou Academy of Agricultural Sciences, Hangzhou, Zhejiang 310024, China
出版时间: 2025-05-25 doi: 10.3969/j.issn.1000-2561.2025.05.008
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为探究茶树品种曜秋(YQ)高温白化的特性及其分子调控机制,以低温白化茶树品种白叶1号(BY1H)为对照,对茶树品种曜秋的春季(YQC)、夏季(YQX)、秋季白化(YQQ)和秋季绿色(YQQL)的一芽二叶新梢,以及白叶1号的春季白化(BY1HC)、夏季(BY1HX)和秋季(BY1HQ)的新梢为材料进行生化成分检测、转录组测序和分析。生化分析结果表明,YQQ白化鲜叶游离氨基酸总量和茶多酚含量均低于YQC绿色鲜叶,且游离氨基酸总量差异达显著水平;而BY1HC白化鲜叶游离氨基酸总量显著高于BY1HQ绿色鲜叶,茶多酚含量显著低于BY1HQ绿色鲜叶。转录组主成分分析结果表明,YQQ白化鲜叶基因表达信息独立于第二象限,明显与其他样品不同;聚类分析结果表明,YQQ白化鲜叶基因表达信息单独聚为一类;YQQ和BY1HQ的差异表达基因数量最多,曜秋在夏末秋初高温响应下新梢叶片白化,微管、细胞、亚细胞等结构受到影响,氨基酸、多酚等主要功能性理化成分含量均降低,差异表达基因富集显著性最高的3个功能类依次为微管结合、细胞或亚细胞成分转移和非膜结合细胞器,差异表达基因富集显著性排名前3的通路依次为类黄酮生物合成通路、淀粉和蔗糖代谢通路以及氨基糖和核苷酸糖代谢通路;基因共表达网络分析筛选出高表达模块,并对总连通性K值最高的10个基因进行分析和注释,其中TGY042732和未知基因novel.276的功能未能预测。本研究结果为了解茶树资源高温白化提供一定的理论基础。

高温  /  白化  /  茶树  /  生化成分  /  转录组  /  表达基因

In order to explore the characteristics and molecular regulation mechanism of high temperature albinism of tea variety Yaoqiu (YQ) and its molecular regulation mechanism, the effects of low temperature albinism on the shoot with one bud and two leaves of tea variety YQ in spring (YQC), summer (YQX), autumn albinism (YQQ) and autumn green (YQQL), and Baiye No. 1 (BY1H) in spring albinism (BY1HC), their biochemical components, and transcriptome sequencing were carried out on the new shoots in summer (BY1HX) and autumn (BY1HQ). The results of biochemical analysis showed that the total free amino acid content and tea polyphenols content of YQQ albino fresh leaves were lower than those of YQC green fresh leaves, and the difference of total free amino acid content was significant; while the total free amino acid content of BY1HC albino fresh leaves was significantly higher than that of BY1HQ green fresh leaves, and the tea polyphenols content was significantly lower than that of BY1HQ green fresh leaves. Cluster analysis revealed that the gene expression profiles of YQQ albino fresh leaves formed a distinct cluster. The comparison between YQQ and BY1HQ exhibited the highest number of differentially expressed genes. The new shoot leaves of YQ showed an albino phenotype induced by high-temperature stress in late summer and early autumn. This stress affected the structure of microtubules, cells, and subcellular components, leading to a reduction in the contents of amino acids and polyphenols. The three functional categories exhibiting the most significant gene enrichment were microtubule binding, movement of cellular or subcellular components, and non-membrane-bound organelles. Similarly, the top three pathways with the highest significance in gene enrichment were the flavonoid biosynthesis pathway, the starch and sucrose metabolism pathway, and the amino sugar and nucleotide sugar metabolism pathway. Through gene co-expression network analysis, modules exhibiting high expression levels were identified. Subsequently, the ten genes demonstrating the highest total connectivity (K value) were selected for further analysis and annotation. Notably, the functions of genes TGY042732 and the novel gene designated as novel.276 could not be predicted. These findings would provide a theoretical foundation for understanding the phenomenon of albinism in tea resources under conditions of elevated temperature.

high temperature  /  albino  /  tea plant  /  biochemical components  /  transcriptome  /  expressed genes
黄海涛, 丁一, 牛小军, 赵芸, 余继忠. 高温白化茶树品种曜秋的特性和转录组分析. 热带作物学报, 2025 , 46 (5) : 1094 -1106 . DOI: 10.3969/j.issn.1000-2561.2025.05.008
Haitao HUANG, Yi DING, Xiaojun NIU, Yun ZHAO, Jizhong YU. Characteristics and Transcriptome Analysis of High Temperature Albino Tea Variety Yaoqiu[J]. Chinese Journal of Tropical Crops, 2025 , 46 (5) : 1094 -1106 . DOI: 10.3969/j.issn.1000-2561.2025.05.008
茶树新梢叶色变异广泛,有绿色、白色、黄色、红色、紫色等。白色和黄色茶树新梢表型统称为白化表型,其中新梢白化程度高低主要受温度和光照影响,其中受温度影响的类型又可根据白化响应温度不同分为高温白化型和低温白化型[1-2]。低温白化型茶树品种有白叶1号、小雪芽等[3-8],白叶1号白化的温度阈值在20~22 ℃之间,但该温度仅在春季越冬芽萌发的初期发挥作用,其白化现象主要与叶片色素含量的变化及其叶绿体超微结构的变化密切相关,叶片白化期叶绿体膜结构发育发生障碍,叶绿体退化解体,叶绿素合成受阻,质体膜上各种色素蛋白复合体缺失,导致叶片白化[9-10];LI等[4]对低温白化茶树品种小雪芽的表型和转录组研究发现,小雪芽白化叶片(芽下第三叶)叶绿体发育异常,表现为体积膨胀变大,出现囊状空泡,基粒片层和类囊体膜结构发育受阻,光合色素叶绿素类和类胡萝卜素类含量显著减少,氨基酸含量显著增加,茶多酚、咖啡因、总儿茶素类、表没食子儿茶素没食子酸酯(EGCG)和表儿茶素没食子酸酯(ECG)含量以及酚氨比值显著降低,转录组分析结果表明其白化叶片的1-脱氧-D-木酮糖-5-磷酸合成酶基因以及原叶绿素酸酯还原酶基因表达受到显著抑制,导致小雪芽白化叶片的叶绿素和类胡萝卜素含量较低。茶树高温白化型是指气温达到25 ℃以上或持续高温时的芽叶白化,后随气温下降逐步返绿,但综合性状不理想,因此,高温白化型茶树少有相关研究[1]。高温白化种质在其他作物上也仅有少量研究,李军等[11]报道的粳稻高温白化复绿突变体tcd52在高温(>24 ℃)条件下,二叶期叶色呈白色失绿,三叶期开始复绿,四叶期后与野生型无明显差异;而在低温(20 ℃)条件下,tcd52突变体苗期叶色与野生型一致呈绿色,无白化现象。
茶树品种曜秋因高温白化而得名,是从浙江省杭州市余杭区瓶窑镇鸠坑群体种自然实生后代中采用单株选择的方法系统选育而成的特色茶树品种,春季新梢为黄绿色、夏季新梢叶色为紫绿色,夏末至秋初新梢白化,白化叶片不返绿,凋亡脱落,秋季后期继续生长的顶梢呈绿色,长势较弱,是一种较好的高温白化茶树种质资源。本研究拟以低温白化茶树品种白叶1号为对照品种,对曜秋不同季节芽叶的理化性状进行检测,并结合转录组分析其高温白化时的主要成分和基因表达变化。
本研究于2022年度在杭州市余杭区瓶窑镇杭州市农业科学研究院茶叶研究所资源圃(30°23ʹE,119°53ʹN)选取2个茶树品种,试验组为高温白化茶树品种曜秋(YQ),分别取其春季(YQC)一芽二叶、夏季(YQX)一芽二叶、秋季白化(YQQ)一芽二叶和秋初返绿(YQQL)一芽二叶作为研究材料;对照组为低温白化茶树品种白叶1号(BY1H),分别取其春季白化(BY1HC)一芽二叶、夏季(BY1HX)一芽二叶和秋季(BY1HQ)一芽二叶,每个样品3个重复,用锡纸包裹后迅速投入液氮中速冻,–80 ℃低温保存备用,用于后续生理生化指标测定和总RNA提取。
采用国家标准《茶水浸出物测定》GB/T 8305—2013的方法检测样品水浸出物含量。
采用国家标准《茶叶中茶多酚和儿茶素类含量的检测方法》GB/T 8313—2018检测样品茶多酚含量。
采用HPLC法检测,儿茶素类包括没食子儿茶素(GC)、表没食子儿茶素(EGC)、儿茶素(C)、表儿茶素(EC)、表没食子儿茶素没食子酸酯(EGCG)、表儿茶素没食子酸酯(ECG)、没食子儿茶素没食子酸酯(GCG)和儿茶素没食子酸酯(CG),仪器为Agilent 1100,色谱柱为Hypsial ODS的C18柱(5 μm,4.6 mm×250 mm);流动相A为2%乙酸溶液,流动相B为乙腈,流速1 mL/min,柱温35 ℃,检测波长280 nm,进样量10 μL,0~12 min内流动相A由93.5%变为92.0%,12~16 min时流动相A由92.0%变为85.0%,16~20 min时流动相A由85.0%变为15.0%,20~30 min回到初始状态即流动相A为93.5%,平衡5 min[12]
采用领苯二酚(OPA)柱前衍生结合高效液相色谱(HPLC)荧光检测法对氨基酸组分进行检测,包括天冬氨酸(Asp)、谷氨酸(Glu)、天冬酰胺(Asn)、丝氨酸(Ser)、谷氨酰胺(Gln)、组氨酸(His)、苏氨酸(Thr)、精氨酸(Arg)、丙氨酸(Ala)、γ-氨基丁酸(Gaba)、茶氨酸(Thea)、缬氨酸(Val)、亮氨酸(Leu)。柱前衍生方法:先配制OPA溶液,方法为9 mL硼酸缓冲液(0.4 mol/L,pH 10.2)+0.01 g OPA+1 mL乙腈+100 μL巯基丙酸,混匀后过0.22 μm膜。将配制完成的OPA溶液50 μL与0.4 mol/L硼酸缓冲液(pH 10.2)500 μL、450 μL超纯水以及5 μL待测茶样溶液,混匀后进行检测。检测条件为Zorbax EclipseAAA色谱柱(3.5 μm,4.6 mm×150 mm),柱温箱40 ℃,发射波长340 nm,接收波长450 nm,进样体积10 μL,流速1.5 mL/min,流动相A为40 mmol/L Na2HPO4缓冲液(pH 7.8),流动相B为乙腈∶甲醇∶超纯水=45∶45∶10(V/V/V)。洗脱梯度为0~18 min,流动相B由5%(V)线性增加至60%(V),18~32 min流动相B由60%(V)线性增加至100%(V),32~37 min流动相B降至5%(V),并保持7 min[13]
基于Illumina技术测序平台,利用双末端测序(paired-end)的方法,分别对曜秋春季(YQC)与白叶1号春季白化(BY1HC)的一芽二叶新梢,曜秋夏季(YQX)与白叶1号夏季(BY1HX)的一芽二叶新梢,曜秋秋季白化(YQQ)与白叶1号秋季(BY1HQ)的一芽二叶新梢,曜秋秋季(YQQ)白化与曜秋秋季绿色(YQQL)的一芽二叶新梢进行转录组测序分析,原始测定数据(raw reads)进行过滤处理后得到高质量数据(clean reads),使用HISAT2将clean reads与茶树[14]的参考基因组进行比对。使用StringTie软件进行新转录本组装,采用DESeq2[15-16]根据|log2(fold change)|≥1且padj≤0.05的标准筛选差异基因,差异基因的表达丰度用FPKM值(fragments per kilobase of exon model per million mapped fragments)表示,采用clusterProfiler软件对差异基因集进行GO(gene ontology)功能富集分析和KEGG(kyoto encyclopedia of genes and genomes)通路富集分析,采用软阈值为6进行网络构建,使用相异度对网络中的基因进行层次聚类,并利用基因总连通性K值进行核心基因筛选。
使用JMP 10.0软件进行统计分析,包括方差分析(ANOVA)和主成分分析(PCA)。测量的理化成分含量以平均值±标准偏差(SD)表示,采用t检验确定不同样品之间在5%的显著性水平,P≤0.05且|log2(fold change)|≥1视为显著差异表达的指标。
曜秋原始编号为J1615,来自浙江余杭,灌木型,半开张,生长势中等,萌发中等,茸毛少,春季新梢叶色为黄绿色,夏季初期新梢为紫绿,夏季中后期至秋初新梢白化(图1),秋季后期新梢呈绿色,成熟叶片为中等椭圆、叶长7.8 cm、叶宽3.2 cm、浅绿、上表皮隆起、叶缘波状、叶身平、叶齿密浅锐、叶尖渐尖、叶基楔形(表1)。按照规范2~3 d观测1次,以观测到变化的日期为其起始和转绿日期;最高气温高于35 ℃和低于30 ℃的日期来自杭州市气象局资料。根据2020—2023年的观测(表2),曜秋在夏季气温上升至33~35 ℃出现明显的白化现象,且白化程度随着温度的升高而增加,茎、叶、脉及腋芽均呈白化状,秋初温度降至30 ℃以下,新生长的新梢芽叶颜色转为正常绿色(图2B),原白化叶片逐渐成熟、凋亡和脱落(图2C)。
白叶1号又名安吉白茶,原产浙江安吉,灌木型,半开张,生长势中等,茸毛少,春季新梢叶色呈玉白色(图1),叶脉淡绿色,夏秋季新梢均为绿色,成熟叶片为长椭圆、叶长8.8 cm、叶宽3.4 cm、绿色、上表皮平、叶缘平、叶身平、叶齿稀浅钝、叶尖渐尖、叶基楔形。白叶1号茶树新梢在春季气温上升至20~22 ℃阶段性出现白化现象,至春末,白化叶片逐渐成熟并转绿,早期白化度较高的叶片逐渐脱落,后期白化度低的叶片形成绿白斑驳状(图2A),夏季和秋季新生长新梢的颜色均为正常的绿色。
茶多酚、氨基酸、咖啡碱、儿茶素与茶叶风味品质紧密相关,是茶叶风味化学成分的主体。2个品种不同季节的一芽二叶的水浸出物、茶多酚、氨基酸、咖啡碱和儿茶素含量见表3。2个品种的水浸出物含量、茶多酚含量均在夏季最高,春秋较低;氨基酸总量均为春季较高,夏秋季较低;咖啡碱含量春季、夏季较高,而秋季较低;曜秋品种的水浸出物含量在不同季节均低于白叶1号;曜秋品种春季的茶多酚含量高于白叶1号,秋季茶多酚含量则低于白叶1号;曜秋品种春季的氨基酸含量低于白叶1号,夏季和秋季的氨基酸含量均高于白叶1号;曜秋品种春季和夏季的咖啡碱含量均低于白叶1号;曜秋品种春季、秋季的GC、EGC、C和EC等简单儿茶素含量均低于同期的白叶1号,复杂儿茶素无明显规律。曜秋品种秋季白化鲜叶较春季绿色鲜叶的水浸出物、咖啡碱、EGC、EGCG、GCG、CG等含量,以及游离氨基酸总量等均显著降低,而ECG含量显著升高。白叶1号品种春季白化鲜叶较秋季绿色鲜叶茶多酚、EGCG含量显著降低,而水浸出物、咖啡碱、GC、EGC、C、EC、CG等含量,以及游离氨基酸总量均显著增升高。游离氨基酸总量显著增加和茶多酚含量显著降低是白叶1号低温白化产生的一个重要代谢特性,而曜秋高温白化鲜叶的游离氨基酸总量并未显著增加,这可能与2个茶树品种不同的白化机制相关。
氨基酸组成对于茶汤鲜爽度、滋味品质也具有重要影响。2个品种春季和秋季鲜叶的游离氨基酸含量见表4,不同样品间天冬氨酸、丝氨酸、谷氨酸、谷氨酰胺、茶氨酸、丙氨酸、γ-氨基丁酸和精氨酸含量差异达到显著水平。曜秋春季鲜叶的天冬氨酸、谷氨酸、茶氨酸、γ-氨基丁酸、精氨酸含量显著低于白叶1号品种,曜秋秋季鲜叶的茶氨酸、γ-氨基丁酸含量显著低于白叶1号,而天冬氨酸、谷氨酸、精氨酸含量显著高于白叶1号。曜秋秋季白化鲜叶样品相较于其春季绿色鲜叶样品,天冬氨酸、谷氨酸和谷氨酰胺含量显著升高,丝氨酸和茶氨酸含量显著降低。白叶1号春季白化鲜叶样品相较于其秋季绿色鲜叶样品,天冬氨酸、丝氨酸、谷氨酸、谷氨酰胺、茶氨酸、丙氨酸和精氨酸含量显著升高,而γ-氨基丁酸含量显著降低。综上,天冬氨酸、谷氨酸和谷氨酰胺3种氨基酸在品种白化过程中的变化趋势一致,而丝氨酸和茶氨酸2种氨基酸在品种白化过程中的变化趋势相反,这可能与2个茶树品种不同的白化机制相关。
曜秋和白叶1号各设3个生物学重复,共构建21个cDNA文库,基于Illumina Hiseq TM4000测序获得的数据详见表5。经过滤筛选后共获得144.85 Gb的clean data,每个样品获得6.9 Gb左右的clean data,占比均超过98%,Q30(一个碱基的识别可靠性等于99.9%)的占比超过91.00%,GC含量(鸟嘌呤和胞密啶的摩尔百分比)均在43.00%以上。说明试验取样合理,样品RNA-seq数据质量可靠。
为了进一步了解2个茶树品种白化现象的区别,对样品进行了主成分分析(图3)。结果表明,曜秋品种春季、夏季、秋季鲜叶基因表达信息集中在第三象限,曜秋秋季白化鲜叶基因表达信息独立在第二象限;而白叶1号品种春季、夏季、秋季鲜叶基因表达信息主要集中在第一、四象限。曜秋和白叶1号2个品种不同季节的鲜叶基因表达信息差异较大,曜秋秋季白化的鲜叶与其春季、夏季和秋初返绿的鲜叶基因表达信息之间差异明显,而白叶1号春季白化、夏季和秋季的鲜叶信息之间差异较小。
通过差异基因表达量分析(图4),曜秋春季鲜叶和白叶1号春季鲜叶之间以|log2(fold change)|≥1且padj≤0.05为标准,共筛选出7288个差异表达基因(DEGs),包含3732个上调表达基因和3556个下调表达基因;曜秋夏季鲜叶和白叶1号夏季鲜叶之间筛选出7793个差异表达基因,包含4137个上调表达基因和3656个下调表达基因;曜秋秋季鲜叶和白叶1号秋季鲜叶之间筛选出14 982个差异表达基因,包含7025个上调表达基因和7957个下调表达基因;曜秋秋季高温白化鲜叶和曜秋秋季新生绿色鲜叶样品之间筛选出7797个差异表达基因,包含2865个上调表达基因和4932个下调表达基因。其中曜秋茶树秋季白化新梢和白叶1号秋季新梢的差异表达基因最多,远超其他时期的差异表达基因。
通过差异表达基因聚类分析(图5),曜秋秋季白化新梢表达基因与白叶1号春季白化新梢相比区别较大,与其他季节的新梢表达基因也具有较大的差异。7个样本聚为2大类群,其中YQQ单独聚为第一大类群,而其余BY1HC、BY1HX、BY1HQ、YQC、YQX和YQQL聚为第二大类群,其中BY1HC、BY1HX和BY1HQ聚为第二大类群的第一小类,YQC、YQX和YQQL聚为第二大类群的第二小类。
对差异表达基因进行GO功能富集,曜秋春季鲜叶和白叶1号春季鲜叶之间,富集显著性最高的3个功能类依次为类囊体、RNA-DNA杂交核糖核酸酶活性和光合作用,而富集差异基因数最多的功能类则为RNA催化活性类(78个)(图6A)。曜秋夏季鲜叶和白叶1号夏季鲜叶之间,富集显著性最高的3个功能类依次为RNA-DNA杂交核糖核酸酶活性、过氧化物酶体膜和过氧化物酶体组织类,而富集差异基因数最多的功能类则为蛋白质二聚化活性类(107个)(图6B)。曜秋秋季鲜叶和白叶1号秋季鲜叶之间,富集显著性最高的3个功能类依次为微管结合、细胞或亚细胞成分转移和非膜结合细胞器,而富集差异基因数最多的功能类则为DNA结合转录因子活性(219个)(图6C)。曜秋秋季白化鲜叶和曜秋秋季白化后新生绿叶之间,富集显著性最高的3个功能类依次为转移酶活性,转移酰基、细胞或亚细胞组分转移和染色体部分,而富集差异基因数最多的功能类则为转移己糖基转移酶活性类(167个)(图6D)。
KEGG通路富集分析显示,曜秋春季鲜叶和白叶1号春季鲜叶之间,富集显著性排名前3的通路依次为光合作用-天线蛋白,光合作用和光合生物的碳固定,分别富集到21、30、62个DEGs(图7A)。曜秋夏季鲜叶和白叶1号夏季鲜叶之间,富集显著性排名前3的通路依次为缬氨酸、亮氨酸和异亮氨酸生物合成、泛酸和CoA生物合成和2-氧代羧酸代谢通路,分别富集到21、22、35个DEGs(图7B)。曜秋秋季鲜叶和白叶1号秋季鲜叶之间,富集显著性排名前3的通路依次为类黄酮生物合成、淀粉和蔗糖代谢以及氨基糖和核苷酸糖代谢,分别富集到47、121、109个DEGs(图7C)。曜秋秋季白化鲜叶和曜秋秋初新生绿色鲜叶之间,富集显著性排名前3的通路依次为类黄酮生物合成、苯丙素类生物合成和脂肪酸延伸,分别富集到35、76、23个DEGs(图7D)。
软阈值设定为6,筛选基因表达数据,并建立基因表达模块层次聚类树以及模块基因聚类热图(图8),本研究结果发现绿松石色模块基因高表达。
对绿松石色模块中总连通性K值最高的10个基因进行分析和注释(表6),其中TGY042732和未知基因novel.276的功能未能预测,其余基因功能多数与叶绿体相关,这些在基因共表达网络中处于核心位置的新基因为今后进一步探索茶树高温白化机理提供重要参考。
刘钰龙等[17]报道一株水稻温敏型白化突变体tsa1在低温20~24 ℃下明显白化,叶绿素a和类胡萝卜素的含量显著降低,突变体白化组织中叶绿体发育停滞,出现了大量小型异常叶绿体结构,而高温28~32 ℃下突变体外观及光合色素含量趋于正常。白叶1号为低温白化茶树品种,其越冬芽萌发初始叶片为绿色,温度升至20~22 ℃白化启动,白化叶片的叶面为白色,叶脉为绿色,早期白化度较高的叶片后期会脱落,而白化度较低的叶片在春季末期形成白-绿斑驳形态,部分叶片的此种形态可持续全年,夏季和秋季新生的新梢叶片均为正常绿色。成浩等[18]研究发现白叶1号全白期间叶片中不存在P700叶绿素a蛋白复合体与捕光叶绿素a/b蛋白质复合体,但含有很少量的捕光叶绿素a/b蛋白质复合体的脱辅基蛋白和少量的游离色素,认为捕光叶绿素a/b蛋白质复合体的缺失影响到叶绿体基粒结构的形成是白叶1号白化重要原因之一。以上研究均表明叶绿体发育受阻和结构异常是植物低温白化表型的重要原因。
李军等[11]报道一株水稻高温敏感型白化突变体tcd52在20 ℃时叶绿素含量与野生型无显著差异,而在24 ℃时,tcd52叶绿素下降到野生型(WT)的一半;28 ℃时,下降至WT的1/4;而32 ℃时,tcd52中几乎检测不到叶绿素,电镜观察此时其叶片细胞中基本无完整的、成形的叶绿体结构。该研究表明叶绿体结构降解是植物高温白化表型的主要原因。
本研究中的材料曜秋为高温白化茶树品种,其春季、夏季初期叶片为正常绿色或紫绿色,温度为33~35 ℃启动白化,白化叶片的叶面、叶脉、腋芽均呈现黄色,随着温度升高白化程度可增加至乳白色,温度降低至30 ℃以下时,白化叶片逐渐凋亡,此后新梢继续生长的叶片为正常绿色。曜秋高温白化生物过程与tcd52类似,结合曜秋转录组基因注释,推测曜秋高温白化也是与叶绿体结构降解有关。
王新超等[9]对白叶1号正常叶片和白化叶片差异表达的基因进行了比较和功能推测,发现参与叶绿体/叶绿素合成的调控基因(A18-4C3),参与信号物质合成有关基因(G11-2C3)等与白叶1号阶段性白化有关。本研究表明曜秋秋季白化新梢表达基因与白叶1号春季白化新梢相比区别较大,与曜秋其他季节的新梢表达基因也具有较大的差异,其中曜秋秋季高温白化鲜叶与曜秋秋季新生绿色鲜叶相比下调表达基因较多,与白叶1号秋季绿色新梢相比也是下调表达基因较多,且差异表达基因数量最多,高达14 982个差异表达基因,表明曜秋秋季白化具有较为复杂的生物学机制。
LI等[19]对白叶1号不同阶段的代谢和转录研究表明,白叶1号在WII阶段(白化后期)的差异代谢产物主要参与光合生物的碳固定、苯丙素类化合物的生物合成、半乳糖代谢、类黄酮的生物合成以及精氨酸和脯氨酸的代谢。而在本研究中,曜秋秋季白化鲜叶和曜秋秋初新生绿色鲜叶之间,富集显著性排名前3的通路依次为类黄酮生物合成,苯丙素类生物合成和脂肪酸延伸,主要与类黄酮生物合成,苯丙素类生物合成和脂肪酸延伸等途径有关。推测曜秋高温白化途径与白叶1号低温白化后期代谢途径具有一定的共性。
同时,通过基因共表达网络分析筛选的处于核心位置的新基因为今后进一步开展茶树高温白化机理研究提供重要参考。
本研究通过对曜秋春季绿色、夏季绿色、秋季白化和秋季绿色,白叶1号春季白化、夏季绿色、秋季绿色等的鲜叶为材料通过表型、理化组成和转录组等联合分析解析了高温白化茶树曜秋的白化机制。曜秋在夏末秋初高温(33~35 ℃)响应下新梢叶片白化,温度降低至30 ℃左右时,白化叶片启动凋亡,而继续生长的新梢为正常绿色,与白叶1号低温白化新梢氨基酸含量显著增加不同,曜秋高温白化新梢游离氨基酸总量反而较春季绿色新梢显著降低。曜秋秋季白化新梢基因表达信息在主成分分析与聚类分析中均表现出独特性,通过基因共表达网络分析筛选出10个核心基因,包括TGY042732TGY109858TGY046880TGY107449TGY014623TGY096457novel.276TGY123290TGY050175TGY088252。研究结果为今后进一步开展茶树资源的高温白化研究奠定基础。
  • 浙江省“十四五”农业新品种选育重大科技专项(2021C02067-4)
  • 国家茶叶产业技术体系项目(CARS19)
  • 杭州市农科院科技创新与示范推广基金项目(2022HNCT-04)
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2025年第46卷第5期
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doi: 10.3969/j.issn.1000-2561.2025.05.008
  • 接收时间:2024-11-07
  • 首发时间:2026-06-26
  • 出版时间:2025-05-25
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  • 收稿日期:2024-11-07
  • 录用日期:2025-01-07
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浙江省“十四五”农业新品种选育重大科技专项(2021C02067-4)
国家茶叶产业技术体系项目(CARS19)
杭州市农科院科技创新与示范推广基金项目(2022HNCT-04)
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    杭州市农业科学研究院茶叶研究所,浙江杭州 310024

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* 余继忠(YU Jizhong),E-mail:
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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