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Article(id=1237814983392818009, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1237814978405790425, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2025.10.004, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1745683200000, receivedDateStr=2025-04-27, revisedDate=null, revisedDateStr=null, acceptedDate=1749744000000, acceptedDateStr=2025-06-13, onlineDate=1773047689531, onlineDateStr=2026-03-09, pubDate=1761321600000, pubDateStr=2025-10-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1773047689531, onlineIssueDateStr=2026-03-09, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1773047689531, creator=13701087609, updateTime=1773047689531, updator=13701087609, issue=Issue{id=1237814978405790425, tenantId=1146029695717560320, journalId=1235980609244409860, year='2025', volume='46', issue='10', pageStart='2287', pageEnd='2547', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1773047688342, creator=13701087609, updateTime=1773049212967, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1237821373213635442, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1237814978405790425, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1237821373213635443, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1237814978405790425, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2323, endPage=2334, ext={EN=ArticleExt(id=1237814983627699045, articleId=1237814983392818009, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Identification and Expression Analysis of the CAD Gene Family in Tea Plant (Camellia sinensis) and Their Response to Infestation by Empoasca onukii, columnId=1236256430337085821, journalTitle=Chinese Journal of Tropical Crops, columnName=Omics & Biotechnology, runingTitle=null, highlight=null, articleAbstract=

During the biosynthesis process of lignin in plants, cinnamyl alcohol dehydrogenase (CAD) plays a crucial role, catalyzing the final step reaction in the entire metabolic pathway. To explore the potential functions of the CAD gene family in tea plants, CAD gene family members were identified in the genome of Huangyan tea plants and a series of bioinformatics analyses were conducted. Based on transcriptome data, the gene expression of the members in different organs of tea plants and after damage by the Empoasca vitis were studied. A total of 36 members of HD-CsCADs were identified, which were unevenly distributed on 9 chromosomes and encoded amino acid lengths ranging from 300 aa to 621 aa and protein molecular weights ranging from 321.40 kDa to 666.49 kDa. HD-CsCADs had 0 to 3 introns and the promoters containing 79 types of cis-acting elements, among which the number of elements related to stress response was the highest. Combined with the phylogenetic tree, HD-CsCADs could be divided into 4 subfamilies. There were significant differences in the expression levels of HD-CsCADs in different organs of tea plants. HD-CsCAD-15 was highly homologous to CAD genes involved in lignin biosynthesis in other plants and was highly expressed in stem tissues. In addition, the expression of the genes after damage by E. vitis on tea seedlings was analyzed. The expression of HD-CsCAD-11 and HD-CsCAD-15 was relatively obvious and could be used as important CAD genes related to lignin metabolism and tea plant defense against pests. The results would provide theoretical basis for the defense mechanism of tea plants against the small green leaf hopper and the utilization of CAD gene functions.

, correspAuthors=Shan JIN, authorNote=null, correspAuthorsNote=
*JIN Shan，E-mail：。
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肉桂醇脱氢酶（cinnamyl alcohol dehydrogenase，CAD）是木质素生物合成途径的一类关键酶，主要负责催化木质素单体生物合成途径的最后一步反应。为了探究茶树CAD基因家族的潜在功能，本研究利用生物信息学技术，在茶树黄棪基因组中对CAD基因进行鉴定，并基于转录组数据研究它们在茶树不同组织中及小贯小绿叶蝉为害后其基因表达的情况。研究结果表明：鉴定获得36个CAD基因家族成员，分别命名为HD-CsCAD-1~HD-CsCAD-36，不均匀地分布于9条染色体上，编码氨基酸长度为300~621 aa，蛋白分子量为321.40~666.49 kDa。HD-CsCADs有0~3个内含子，启动子中含有79种顺式作用元件，其中与逆境胁迫响应相关的元件数量最多。进化树分析结果表明，HD-CsCADs可分为4个亚家族。HD-CsCADs在茶树的不同组织中表达量存在明显差异，HD-CsCAD-15与其他植物参与木质素生物合成的CAD基因高度同源，并在茎组织中高度表达。此外，还分析它们在茶苗遭受小贯小绿叶蝉为害后的表达情况，其中HD-CsCAD-11和HD-CsCAD-15的表达水平显著上升，可作为研究茶树抗虫防御机制的关键候选基因。本研究结果可为茶树CAD基因功能的验证及抗虫防御分子机制的研究提供一定的理论依据。

, correspAuthors=金珊, authorNote=null, correspAuthorsNote=
*金珊，E-mail：。
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杨欢（2001—），女，硕士研究生，研究方向：茶树抗虫分子机制。

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杨欢（2001—），女，硕士研究生，研究方向：茶树抗虫分子机制。

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杨欢（2001—），女，硕士研究生，研究方向：茶树抗虫分子机制。

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Different lowercase letters indicate significant difference (P<0.05).

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不同小写字母表示差异显著（P<0.05）。

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Sequence characteristics of tea CAD gene family members

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基因
Gene		基因
IDGene ID		长度
Length/aa		分子量
MW/kDa		等电点
PI		不稳定系数
Instability		脂肪数
AI		亲水性
GRAVY		亚细胞定位
Subcellular localization
HD-CsCAD-1HD.02G0030910.t1365395.116.8331.1789.450.013细胞质
HD-CsCAD-2HD.02G0030870.t1367393.705.7031.1587.870.093细胞质
HD-CsCAD-3HD.02G0030900.t1367394.215.6633.2786.810.097细胞质
HD-CsCAD-4HD.01005610.t1356379.446.7522.1688.400.174细胞质外质
HD-CsCAD-5HD.01005605.t1359383.767.5021.4890.920.173叶绿体
HD-CsCAD-6HD.05G0007720.t1361389.616.3128.6393.930.057细胞质
HD-CsCAD-7HD.04G0022770.t1361390.336.3128.8393.380.064细胞质
HD-CsCAD-8HD.04G0026700.t1359384.756.6327.0097.410.162细胞质
HD-CsCAD-9HD.04000767.t1364398.006.4526.3188.87-0.124过氧化物酶体
HD-CsCAD-10HD.1209294.t1356383.076.6323.9394.690.108细胞质
HD-CsCAD-11HD.03G0033730.t1356383.076.6323.9394.690.108细胞质
HD-CsCAD-12HD.02G0008450.t1360386.876.9427.1290.170.007细胞质
HD-CsCAD-13HD.10G0022480.t1440470.457.9048.4685.80-0.019叶绿体
HD-CsCAD-14HD.04G0022660.t1361389.526.6328.4994.210.049细胞质
HD-CsCAD-15HD.13G0019690.t1357389.515.3527.2891.88-0.029细胞骨架
HD-CsCAD-16HD.01G0026210.t1300321.406.7830.9993.830.089细胞质
HD-CsCAD-17HD.02G0008490.t1360389.997.2025.7492.61-0.012细胞质
HD-CsCAD-18HD.04G0022810.t1356382.426.8230.4897.720.154细胞质
HD-CsCAD-19HD.05G0028420.t1357389.085.5319.3793.220.043细胞质
HD-CsCAD-20HD.02G0008470.t1621666.496.6029.1791.80-0.014细胞质
HD-CsCAD-21HD.01G0026200.t1360390.495.9028.0794.440.055细胞质
HD-CsCAD-22HD.06G0017630.t1356387.566.2624.9986.46-0.017叶绿体
HD-CsCAD-23HD.04008754.t1360395.546.4627.8386.31-0.053细胞质
HD-CsCAD-24HD.07012446.t1359384.886.3728.2787.940.096细胞质外质
HD-CsCAD-25HD.04008762.t1351384.896.0829.1086.87-0.049细胞骨架
HD-CsCAD-26HD.14G0016950.t1379407.306.7828.4486.890.063细胞质
HD-CsCAD-27HD.06G0024180.t1386419.725.9731.2196.420.097细胞质
HD-CsCAD-28HD.06G0024190.t1387416.775.6332.3592.610.106细胞质
HD-CsCAD-29HD.10G0013100.t1380415.656.4334.2685.08-0.060细胞质
HD-CsCAD-30HD.10G0013120.t1380415.626.4335.9384.32-0.073细胞质
HD-CsCAD-31HD.10G0013080.t1380414.666.8629.7285.08-0.035细胞质
HD-CsCAD-32HD.09006357.t1389426.847.9233.8884.40-0.060细胞质
HD-CsCAD-33HD.10G0013070.t1382417.426.2332.6583.14-0.086细胞质
HD-CsCAD-34HD.04G0010500.t1381410.636.1243.1590.030.089叶绿体
HD-CsCAD-35HD.02G0027290.t1428456.906.2826.8388.53-0.050细胞质
HD-CsCAD-36HD.14G0003760.t1334361.326.7524.4294.490.064细胞质
), ArticleFig(id=1237814992226021866, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1237814983392818009, language=CN, label=表1, caption=

茶树CAD基因家族成员的序列特征

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基因
Gene		基因
IDGene ID		长度
Length/aa		分子量
MW/kDa		等电点
PI		不稳定系数
Instability		脂肪数
AI		亲水性
GRAVY		亚细胞定位
Subcellular localization
HD-CsCAD-1HD.02G0030910.t1365395.116.8331.1789.450.013细胞质
HD-CsCAD-2HD.02G0030870.t1367393.705.7031.1587.870.093细胞质
HD-CsCAD-3HD.02G0030900.t1367394.215.6633.2786.810.097细胞质
HD-CsCAD-4HD.01005610.t1356379.446.7522.1688.400.174细胞质外质
HD-CsCAD-5HD.01005605.t1359383.767.5021.4890.920.173叶绿体
HD-CsCAD-6HD.05G0007720.t1361389.616.3128.6393.930.057细胞质
HD-CsCAD-7HD.04G0022770.t1361390.336.3128.8393.380.064细胞质
HD-CsCAD-8HD.04G0026700.t1359384.756.6327.0097.410.162细胞质
HD-CsCAD-9HD.04000767.t1364398.006.4526.3188.87-0.124过氧化物酶体
HD-CsCAD-10HD.1209294.t1356383.076.6323.9394.690.108细胞质
HD-CsCAD-11HD.03G0033730.t1356383.076.6323.9394.690.108细胞质
HD-CsCAD-12HD.02G0008450.t1360386.876.9427.1290.170.007细胞质
HD-CsCAD-13HD.10G0022480.t1440470.457.9048.4685.80-0.019叶绿体
HD-CsCAD-14HD.04G0022660.t1361389.526.6328.4994.210.049细胞质
HD-CsCAD-15HD.13G0019690.t1357389.515.3527.2891.88-0.029细胞骨架
HD-CsCAD-16HD.01G0026210.t1300321.406.7830.9993.830.089细胞质
HD-CsCAD-17HD.02G0008490.t1360389.997.2025.7492.61-0.012细胞质
HD-CsCAD-18HD.04G0022810.t1356382.426.8230.4897.720.154细胞质
HD-CsCAD-19HD.05G0028420.t1357389.085.5319.3793.220.043细胞质
HD-CsCAD-20HD.02G0008470.t1621666.496.6029.1791.80-0.014细胞质
HD-CsCAD-21HD.01G0026200.t1360390.495.9028.0794.440.055细胞质
HD-CsCAD-22HD.06G0017630.t1356387.566.2624.9986.46-0.017叶绿体
HD-CsCAD-23HD.04008754.t1360395.546.4627.8386.31-0.053细胞质
HD-CsCAD-24HD.07012446.t1359384.886.3728.2787.940.096细胞质外质
HD-CsCAD-25HD.04008762.t1351384.896.0829.1086.87-0.049细胞骨架
HD-CsCAD-26HD.14G0016950.t1379407.306.7828.4486.890.063细胞质
HD-CsCAD-27HD.06G0024180.t1386419.725.9731.2196.420.097细胞质
HD-CsCAD-28HD.06G0024190.t1387416.775.6332.3592.610.106细胞质
HD-CsCAD-29HD.10G0013100.t1380415.656.4334.2685.08-0.060细胞质
HD-CsCAD-30HD.10G0013120.t1380415.626.4335.9384.32-0.073细胞质
HD-CsCAD-31HD.10G0013080.t1380414.666.8629.7285.08-0.035细胞质
HD-CsCAD-32HD.09006357.t1389426.847.9233.8884.40-0.060细胞质
HD-CsCAD-33HD.10G0013070.t1382417.426.2332.6583.14-0.086细胞质
HD-CsCAD-34HD.04G0010500.t1381410.636.1243.1590.030.089叶绿体
HD-CsCAD-35HD.02G0027290.t1428456.906.2826.8388.53-0.050细胞质
HD-CsCAD-36HD.14G0003760.t1334361.326.7524.4294.490.064细胞质
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茶树CAD基因家族的鉴定与表达分析及小贯小绿叶蝉为害对其表达的影响
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杨欢 , 阮玲玲 , 闫佳伟 , 任卫威 , 柳紫琼 , 金珊 *
热带作物学报 | 组学与生物技术 2025,46(10): 2323-2334
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热带作物学报 | 组学与生物技术 2025, 46(10): 2323-2334
茶树CAD基因家族的鉴定与表达分析及小贯小绿叶蝉为害对其表达的影响
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杨欢, 阮玲玲, 闫佳伟, 任卫威, 柳紫琼, 金珊*
作者信息
  • 福建农林大学园艺学院/茶学福建省高校重点实验室，福建福州　350002

    • 杨欢（2001—），女，硕士研究生，研究方向：茶树抗虫分子机制。

通讯作者:

*金珊，E-mail：。
Identification and Expression Analysis of the CAD Gene Family in Tea Plant (Camellia sinensis) and Their Response to Infestation by Empoasca onukii
Huan YANG, Lingling RUAN, Jiawei YAN, Weiwei REN, Ziqiong LIU, Shan JIN*
Affiliations
  • College of Horticulture, Fujian Agriculture and Forestry University / Key Laboratory of Tea Science of Fujian Provincial University, Fuzhou, Fujian 350002, China
出版时间: 2025-10-25 doi: 10.3969/j.issn.1000-2561.2025.10.004
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摘要
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肉桂醇脱氢酶（cinnamyl alcohol dehydrogenase，CAD）是木质素生物合成途径的一类关键酶，主要负责催化木质素单体生物合成途径的最后一步反应。为了探究茶树CAD基因家族的潜在功能，本研究利用生物信息学技术，在茶树黄棪基因组中对CAD基因进行鉴定，并基于转录组数据研究它们在茶树不同组织中及小贯小绿叶蝉为害后其基因表达的情况。研究结果表明：鉴定获得36个CAD基因家族成员，分别命名为HD-CsCAD-1~HD-CsCAD-36，不均匀地分布于9条染色体上，编码氨基酸长度为300~621 aa，蛋白分子量为321.40~666.49 kDa。HD-CsCADs有0~3个内含子，启动子中含有79种顺式作用元件，其中与逆境胁迫响应相关的元件数量最多。进化树分析结果表明，HD-CsCADs可分为4个亚家族。HD-CsCADs在茶树的不同组织中表达量存在明显差异，HD-CsCAD-15与其他植物参与木质素生物合成的CAD基因高度同源，并在茎组织中高度表达。此外，还分析它们在茶苗遭受小贯小绿叶蝉为害后的表达情况，其中HD-CsCAD-11和HD-CsCAD-15的表达水平显著上升，可作为研究茶树抗虫防御机制的关键候选基因。本研究结果可为茶树CAD基因功能的验证及抗虫防御分子机制的研究提供一定的理论依据。

茶树  /  肉桂醇脱氢酶基因  /  基因家族  /  木质素  /  小贯小绿叶蝉

During the biosynthesis process of lignin in plants, cinnamyl alcohol dehydrogenase (CAD) plays a crucial role, catalyzing the final step reaction in the entire metabolic pathway. To explore the potential functions of the CAD gene family in tea plants, CAD gene family members were identified in the genome of Huangyan tea plants and a series of bioinformatics analyses were conducted. Based on transcriptome data, the gene expression of the members in different organs of tea plants and after damage by the Empoasca vitis were studied. A total of 36 members of HD-CsCADs were identified, which were unevenly distributed on 9 chromosomes and encoded amino acid lengths ranging from 300 aa to 621 aa and protein molecular weights ranging from 321.40 kDa to 666.49 kDa. HD-CsCADs had 0 to 3 introns and the promoters containing 79 types of cis-acting elements, among which the number of elements related to stress response was the highest. Combined with the phylogenetic tree, HD-CsCADs could be divided into 4 subfamilies. There were significant differences in the expression levels of HD-CsCADs in different organs of tea plants. HD-CsCAD-15 was highly homologous to CAD genes involved in lignin biosynthesis in other plants and was highly expressed in stem tissues. In addition, the expression of the genes after damage by E. vitis on tea seedlings was analyzed. The expression of HD-CsCAD-11 and HD-CsCAD-15 was relatively obvious and could be used as important CAD genes related to lignin metabolism and tea plant defense against pests. The results would provide theoretical basis for the defense mechanism of tea plants against the small green leaf hopper and the utilization of CAD gene functions.

tea plant  /  CAD gene  /  gene family  /  lignin  /  Empoasca vitis
杨欢, 阮玲玲, 闫佳伟, 任卫威, 柳紫琼, 金珊. 茶树CAD基因家族的鉴定与表达分析及小贯小绿叶蝉为害对其表达的影响. 热带作物学报, 2025 , 46 (10) : 2323 -2334 . DOI: 10.3969/j.issn.1000-2561.2025.10.004
Huan YANG, Lingling RUAN, Jiawei YAN, Weiwei REN, Ziqiong LIU, Shan JIN. Identification and Expression Analysis of the CAD Gene Family in Tea Plant (Camellia sinensis) and Their Response to Infestation by Empoasca onukii[J]. Chinese Journal of Tropical Crops, 2025 , 46 (10) : 2323 -2334 . DOI: 10.3969/j.issn.1000-2561.2025.10.004
正文
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茶[Camellia sinensis（L.）O. Kuntze.]是一种具有悠久历史的山茶科（Theaceae Mirb.）山茶属（Camellia L.）的常绿灌木或小型乔木[1]，广泛分布在多个国家，是最古老的树种之一[2]。在传统的中医中，茶树被用作草药和兴奋剂，用于促进消化、解毒、调节血糖和体温以及伤口愈合等[3]。茶树芽和叶均可用于制作茶叶，其馥郁的香气和独特的滋味深受全世界的喜爱。中国是茶叶的原产地和最大生产国，其茶树的种植面积和总产量均居全球首位[4]。目前气候的不断变化、病虫害频发及土壤退化等问题严重威胁着茶叶的产量与品质[5]。
木质素作为植物细胞壁中的一种重要结构组分，在植物体内的含量仅次于纤维素，是一种重要的酚类聚合物[6]。木质素在细胞壁中的沉积不仅增强了植物整体的机械支撑能力，还在抵御病原体入侵、干旱胁迫等生物与非生物逆境中发挥重要防护作用[7]。木质素的形成是通过苯丙烷代谢途径实现的[8]，这一过程需要多种酶的协同作用。在植物体内木质素的生物合成过程中，肉桂醇脱氢酶（cinnamyl alcohol dehydrogenase，CAD）起到了至关重要的作用，它负责催化整个合成代谢途径中的最后一步反应。在氧化型辅酶NADP的作用下，CAD能够催化肉桂醛（如香豆醛、芥子醛、松柏醛等）转化为肉桂醇[9]。CAD基因家族已在多个植物中被鉴定，例如：在水稻（Oryza sativa）中有12个[10]，在拟南芥（Arabidopsis thaliana)中有9个[11]，在谷子（Setaria italica）中有13个[12]，在小麦（Triticum aestivum）有47个[13]等。研究表明，CAD基因可直接参与植物体内木质素的生物合成，例如：水稻中的OsCAD2[10]和高粱（Sorghum bicolor）中的SbCAD2[14]可直接催化木质素的前体物质转化，是木质素合成中的核心调控基因。在拟南芥中AtCAD5主要参与花茎的木质素生物合成[11]；梨（Pyrus spp.）的PpCAD2在番茄（Solanum lycopersicum）中的过表达显著提升了番茄植株中的木质素沉积[15]。除此之外，CAD基因也参与植物体内的非生物逆境的胁迫响应，例如：在水稻中，OsCAD2和OsCAD6的表达明显受到UV处理诱导，说明其对水稻抵御UV辐射有重要影响[16]；小麦的TaCAD12被发现能够增强小麦对纹枯病的抗性[17]。综上，CAD基因家族可能通过各自的调控机制介导植物体内木质素合成的不同通路，同时又参与植物体内非生物逆境的胁迫反应，这充分体现了木质素在植物生理过程中所表现出的多功能特性[18]。
小贯小绿叶蝉（Empoasca onukii Matsuda）（以下简称叶蝉）以其刺吸式口器刺吸茶树的嫩叶或嫩茎，导致茶树产生叶脉红变、叶缘卷曲，甚至引发“叶蝉烧”等症状。叶蝉种群呈高密度分布特征，且具备快速繁殖的能力，对茶树的正常生长发育造成严重威胁，进而也影响茶叶的产量和品质[19-21]。因此，研究茶树对叶蝉的防御机制，是利用茶树品种抗虫性、提升茶叶质量的重要基础。
青心大冇是中国台湾地区最具代表性、种植最广泛的茶树品种之一，后引入福建省，青心大冇茶树鲜叶是制作美人茶的重要原料[22]。然而，针对青心大冇的相关研究较少，也缺乏基因组数据的支撑。本课题组前期组装了高香型乌龙茶栽培品种“黄棪”的染色体级别基因组[23]，可为乌龙茶茶树品种的基因鉴定提供参考。另一方面，CAD基因对木质素的合成及调控研究主要体现在改良树木品种、抗倒伏和抗病害等方面，与植物抗虫关系的报道较少[24]。目前，只在茶树中克隆鉴定了3个与茶尺蠖（Ectropis obliqua Prout）取食诱导相关的CAD基因[25]。本研究利用课题组前期的基因组数据对茶树CAD基因家族进行鉴定并对其进行生物信息学分析，包括理化特性、系统发育分析、染色体定位、保守序列及顺式作用元件分析等特征分析，同时通过转录组数据分析在茶树不同组织中及在叶蝉胁迫处理下CAD成员的表达模式，为今后深入研究CAD基因在茶树抗虫防御以及茶品质形成中的分子机制提供理论依据。
1　材料与方法
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1.1　材料
试验材料来源于福建农林大学茶学福建省高校重点实验室温室，选取生长良好、无病虫害的一年半生青心大冇盆栽茶苗。试验所用的小贯小绿叶蝉（Empoasca onukii Matsuda）（以下简称叶蝉）成虫采自福建农林大学教学茶场，参考闫佳伟等[26]的方法，采用双层parafilm膜夹营养液法提取叶蝉唾液蛋白，并在液氮中冷冻，置于-80 ℃冰箱中保存备用。
1.2　方法
1.2.1　茶苗处理
参考闫佳伟等[26]的方法，选取茶苗进行处理。（1）对照组（CK）：选取健康无病虫害的4盆茶苗，不做任何处理。（2）机械损伤组（MD）：选取健康无病虫害的4盆茶苗，在嫩芽处用00#昆虫针扎刺50~80针，造成叶片机械损伤。（3）叶蝉取食组（CH）：选取健康无病虫害的茶苗4盆，分别用透气尼龙网罩罩住，每盆放入约50头叶蝉成虫。（4）机械损伤+唾液蛋白组（SM）：选取4盆机械损伤茶苗，将提取的叶蝉唾液蛋白均匀地涂抹在机械损伤叶的叶片两面。（5）唾液处理组（SD）：选取健康无病虫害的茶苗4盆，将纯化的唾液蛋白均匀涂抹在健康叶片两面。以上处理在0 h时均用透气的尼龙网罩覆盖，并放在相同的条件下（温度26 ℃，相对湿度70%，光周期L∶D=16∶8）处理48 h。在处理后的12、24、48 h分别进行采样。每次采摘2~3片一芽二叶嫩茶稍后放入铝箔袋中，每个处理组均进行3次生物学重复，立即置于液氮中冷冻，并储存于-80 ℃冰箱中备用。
1.2.2　茶树CAD基因家族成员的鉴定
为了鉴定茶树中的CAD基因家族成员，分别从茶树网站（http://tpia.teaplants.cn/）和Pfam数据库（https//pfam.xfam.org/）中下载黄棪基因组数据和CAD保守域（PF00107和PF08240）的隐马尔可夫模型（HMM）配置文件，在TBtools中使用Simple HMM Search程序进行筛选，获得候选序列。利用SMART（https://smart.embl.de/smart/batch.pl）和NCBICCD（https://www.ncbi.nlm.nih.gov/Structure/bwrpsb/bwrpsb.cgi）网站进行结构域验证，去除结构域不完整的基因序列，最终确定茶树的CAD基因家族成员。
1.2.3　茶树CAD基因家族蛋白理化性质分析与系统发育分析
利用在线工具SignalP-4.1（https://services.healthtech.dtu.dk/services/SignalP-4.1/）和TMHMM（https://services.healthtech.dtu.dk/service.php?TMHMM-2.0）分别进行CAD蛋白的信号肽分析和跨膜结构预测，使用TBtools-Protein Paramter Calc进行氨基酸数量、分子量和等电点分析，同时在WOLF PSORT（https://wolfpsort.hgc.jp/）上进行成员亚细胞定位预测。通过MEGA软件，构建拟南芥[11]、水稻[10]、毛白杨（Populus tomentosa Carrière）[24]、小麦[13]、火炬松（Pinus taeda L.）[25]、油棕（Elaeis guineensis Jacq.）[27]的CAD基因与所鉴定得到的茶树CAD家族基因的系统发育树，利用网站（https://www.evolgenius.info/evolview/）进行进化树的美化，并对确定的CAD基因家族成员进行亚家族分类。
1.2.4　茶树CAD基因家族染色体定位及保守基序分析
使用TBtools软件中的Gene Location Visualize from GFF/GTF程序进行CAD基因染色体定位分析。利用在线工具MEME（https://meme-suite.org/meme/tools/meme）对CAD蛋白保守基序motif进行茶树CAD家族成员蛋白全长的保守基序（motif）分析，并利用TBtools-Gene Structure View（Advanced）对结果进行可视化。
1.2.5　茶树CAD基因启动子顺式元件分析
利用在线工具PlantCARE（http://bioinformatics.psb.ugent.be/webtools/plantcare/html/）对顺式作用元件的数目进行预测，并在TBtools中对结果进行可视化。
1.2.6　茶树CAD基因共线性分析和蛋白互作预测
使用TBtools工具进行茶树物种内的共线性分析，并使用MCScanX工具预测拟南芥和黄棪不同物种间的同源基因。利用TBtools软件确定拟南芥的CAD基因和茶树CAD基因之间的共线关系，并进行可视化。同时在STRING（https://cn.string-db.org/）上进行蛋白互作网络分析。
1.2.7　茶树CAD基因的组织特异性表达分析
下载黄棪的转录组数据，参照前人的方法[19]得到HD-CsCADs在芽、嫩叶、老叶、茎和根上的FPKM值，利用TBtools软件绘制基因在茶树不同组织上的特异表达热图。
1.2.8　转录组数据分析
将1.2.1准备的材料采用RNA提取试剂盒（FastPure Universal Plant Total RNA Isolation Kit，南京Vazyme公司）提取茶树叶片的总RNA，样品委托上海中科新生命生物科技有限公司进行转录组测序。从转录组数据中提取目的基因的FPKM值，经过转换计算后进行图表绘制。
2　结果与分析
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2.1　茶树CAD基因家族成员的鉴定及蛋白理化性质分析
使用CAD的隐马尔可夫模型文件（PF）并应用hmmer软件在黄棪蛋白文件中建立CAD蛋白序列（E≤e–5），经过CDD和SMART保守结构域分析，在黄棪基因组中共鉴定出36个HD-CsCAD（表1）。将确定的CAD基因家族成员命名为HD-CsCAD-1~HD-CsCAD-36。茶树CAD基因亚细胞定位预测分析表明，细胞质上有27个（75.00%）成员，叶绿体上有4个（11.11%）成员，细胞骨架和细胞外基质上分别有2个（0.06%）成员，过氧化物酶体上有1个（0.03%）成员。氨基酸序列长度介于300~621 aa之间，分子量介于321.40~666.49 kDa之间，理论等电点介于5.35~7.92之间，其中有32个是酸性蛋白，4个是碱性蛋白。不稳定系数介于19.37~48.46之间，平均亲水性介于-0.124~0.174之间，14个蛋白具有亲水性，其他均具有疏水性。预测跨膜螺旋数量均为0，均不含有信号肽。
2.2　茶树CAD基因家族系统发育分析
采用NJ法构建CAD基因家族成员氨基酸序列系统发育树（图1），根据进化关系分析可将茶树和其他物种的CAD基因分为4个亚家族，其中，亚家族Ⅰ中只有2个茶树成员，分别是HD-CsCAD-15和HD-CsCAD-19，还包括拟南芥AtCAD-4和AtCAD-5、水稻OsCAD-2和火炬松PtaCAD；而亚家族Ⅱ中含有17个茶树成员，只有1个水稻OsCAD-7；亚家族Ⅲ中含有3个茶树成员，即HD-CsCAD-22、23和HD-CsCAD-25，还含有AtCAD1、OsCAD1和OsCAD4；亚家族Ⅳ中含有14个成员，还包括了AtCAD7、AtCAD8和AtCAD6相关。
2.3　茶树CAD基因的染色体定位和保守基序分析
36个茶树CAD基因一共存在于9条染色体上（图2），其中Chr07染色体上存在8个成员，Chr02、Chr04和Chr09染色体上分别存在6个成员，Chr01染色体上存在4个成员，Chr03和Chr12染色体上分别存在2个成员，Chr13和Chr11均只有1个成员。
保守基序结果表明（图3），在所有的HD-CsCADs成员中均检测到Motif2、Motif3、Motif4、Motif 8及Motif 10，而Motif 19、Motif 20只存在于HD-CsCAD-1~5中。亚家族Ⅰ和亚家族Ⅲ、Ⅳ中包含的Motif类型相同，除HD-CsCAD-20外，均含有11个motif，亚家族Ⅱ中除HD-CsCAD-1~5和HD-CsCAD-13外，其他成员含有的motif类型均相同。除HD-CsCAD-4、HD-CsCAD-5和HD-CsCAD-30基因不含有外显子外，其余均含有1~3个内含子。
2.4　茶树CAD基因的启动子顺式作用元件分析
通过分析得到79种顺式作用元件（图4），主要包括光反应、生长发育调控、植物激素调控及逆境胁迫响应元件，其中大多数元件与逆境胁迫响应相关。所有的家族成员中，HD-CsCAD-27含有顺式作用元件种类最多（33种），HD-CsCAD-28含有的最少（14种）。光反应顺式作用元件中，Box4几乎存在于所有家族成员中，Gap-box和chs-CMA2c分别只存在于1个成员中。生长发育调控元件中存在10种类型，其中有2个家族成员不含有生长发育调控元件，HD-CsCAD-11和HD-CsCAD-20含有的元件数量最多（9个），其中AACA_motif只存在于HD-CsCAD-23中，MBSI存在于HD-CsCAD-9中，19个家族成员均含有O2-site元件。植物激素调控元件有6种类型，即生长素激素、乙烯激素、赤霉素激素、水杨酸激素、茉莉酸激素和脱落酸激素，其中乙烯激素ERE存在于27个家族成员中，26个家族成员含有ABRE响应元件，AuxRE只存在于HD-CsCAD-2中，AT~ABRE只存在于HD-CsCAD-35中。逆境胁迫响应元件中HD-CsCAD-27、HD-CsCAD-31和HD-CsCAD-34等含有的元件数量较多，34个家族成员含有MYB，与木质素相关的顺式作用元件AC-II只存在于HD-CsCAD-34中。
2.5　茶树CAD基因家族成员共线性分析
通过茶树物种内的共线性分析（图5），发现CAD基因家族中只有1对共线性基因，即HD-CsCAD-15和HD-CsCAD-19。此外，茶树CAD基因家族的成员与拟南芥之间有6对同源基因对（图6），因此茶树与拟南芥基因组之间也具有明显的共线性关系。
2.6　蛋白互作网络分析
在拟南芥中，使用STRING网站进行了HD-CAD基因编码蛋白相互作用网络，重点是PER、ADH和CAD之间的相互作用。在蛋白质互作图谱中（图7），HD-CsCAD-15、HD-CsCAD-19和HD-CsCAD-1~HD-CsCAD-5基因的编码蛋白连接性最高。
2.7　茶树CAD基因特异性表达分析
基于转录组数据分析HD-CsCADs在芽、嫩叶、茎、老叶和根中的表达模式（图8），发现HD-CsCAD-15和HD-CsCAD-26在5个器官中均高表达。HD-CsCAD-13、HD-CsCAD-7主要在根中表达；HD-CsCAD-11和HD-CsCAD-36主要在芽、嫩叶和茎中表达。上述表明，HD-CsCAD不同基因在不同部位的表达程度有明显差异，可能在不同的组织中发挥不同作用。
2.8　茶树CAD基因在不同处理下的表达情况
从转录组的数据中选取在茎中表达量较高的6个基因，在未处理（CK）、叶蝉取食（CH）、机械损伤（MD）、机械损伤+唾液蛋白（SM）和唾液处理（SD）处理后的12、24、48 h之内的表达量进行分析。根据相对表达情况（图9）可知，与健康组相比，叶蝉取食特异性诱导了HD-CsCAD-3和HD-CsCAD-15基因的持续上调表达，其表达量在48 h达到峰值，而HD-CsCAD-19和HD-CsCAD-26则呈现下降表达的趋势。在机械损伤处理中，HD-CsCAD-19呈下降表达的趋势，在12 h达到峰值。在机械损伤和唾液蛋白处理下，HD-CsCAD-11、HD-CsCAD-15、HD-Cs CAD- 19均呈先上升后下降的趋势，且在24h达到峰值，HD-CsCAD-3、HD-CsCAD-17、HD-CsC AD-26呈先下降后上升的趋势。在唾液处理组中，HD-CsCAD-3呈下降表达趋势。
3　讨论
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木质素的形成是通过苯丙烷代谢途径实现的[8]，这一过程需要多种酶的协同作用。其中，CAD起到至关重要的作用，负责催化整个合成代谢途径中的最后一步反应[18]。于1992年，CAD基因从烟草（Nicotiana tabacum L.）的茎部分离获得[28]，后来又从火炬松中分离获得[29]。目前CAD基因已在多个植物中被鉴定出来。本研究共鉴定到36个CAD基因。有研究表明，根据系统发育分析可将CAD基因家族分为3大类，其中第一类被认为是真正的CAD[30]。本研究将鉴定到的茶树中的36个CAD基因和其他物种的CAD基因分为4个亚家族，聚类在亚家族Ⅰ的CAD成员，在多种植物中己被证实对木质素的生物合成起主要作用[31]，如水稻中的OsCAD-2、拟南芥中的AtCAD4和AtCAD5、小麦中的TaCAD1和火炬松PtaCAD，被认为与木质素生物合成有关。它们优先在活跃的木质化组织中表达[32]，主要参与松柏醇和新树醇的合成[33]。茶树中的HD-CsCAD-15和HD-CsCAD-19也归属这一亚家族，这意味着HD-CsCAD-15和HD-CsCAD-19也可能参与木质素生物合成。第二类和第三类的CAD主要在木质素中表达，可能在植物逆境调控或生长发育等其他方面具有功能[18]。亚家族Ⅱ只有1个水稻OsCAD-7，研究发现，水稻中OsCAD-7与LpCAD2的表达模式相似[10]，然而，LpCAD2蛋白质序列是独特的，并且具有叶绿体信号肽和独特的底物结合位点。研究认为LpCAD2是一个失去功能的基因[34]，因此推测亚家族Ⅱ中茶树的CAD基因家族成员也失去功能。亚家族Ⅲ中只有3个茶树CAD基因家族成员，还含有AtCAD1、OsCAD1和OsCAD4。在拟南芥中，AtCAD1对木质素合成的催化活性不高[11]。因此这3个基因是否催化木质素合成还未确定。亚家族Ⅳ中含有14个茶树CAD基因家族成员，还包括AtCAD7、AtCAD8和AtCAD9。在拟南芥中，AtCAD7和AtCAD8与植物抗性相关[11]。因此，我们假设Ⅳ家族的大多数基因成员参与茶树抗性，参与茶树防御机制或作为功能冗余的CAD基因发挥作用[35]。
在基因复制事件中，如果2个同源基因之间间隔的基因数不超过5个，则认为它们是串联重复基因；如果间隔基因数量超过5个或分布于不同的染色体上，则认为它们是节段性重复[36]。在茶树中只发现了1对基因复制，即HD-CsCAD-15和HD-CsCAD-19，因此可能是由于节段性重复而形成的。
启动子中含有79种顺式作用元件，其中逆境胁迫响应的元件含量最多，说明茶树CAD家族成员在植物的逆境胁迫中发挥着重要作用。在组织特异性表达分析中，HD-CsCAD-15基因在茶树的茎部呈高表达，这表明该基因可能特异性参与茶树茎秆木质化的生物合成过程。
结合转录组数据分析发现，CAD基因的表达调控受到生物和非生物胁迫的影响，叶蝉取食后，HD-CsCAD-3、HD-CsCAD-11和HD-CsCAD-15均随着时间表现出明显变化。尤其是HD-CsCAD-11在12 h时表达最高，而HD-CsCAD-17、HD-CsCAD-19和HD-CsCAD-26的表达量均低于CK组。在机械损伤后，HD-CsCAD-17和HD-CsCAD-26的表达量随着时间和CK组表达量的变化趋势相同，表明HD-CsCAD-17和HD-CsCAD-26在茎和老叶中表达量高与叶蝉取食和机械损伤无关。只有HD-CsCAD-3和HD-CsCAD-15是呈上升趋势。当植物在受到叶蝉取食或机械损伤等外界胁迫时，可能激发细胞壁木质化沉积的防御应答机制，进而提升植株的抗病和抗逆性[9,21]。综上所述，叶蝉取食和机械损伤处理均能诱导茶树CAD基因不同程度的表达。结合茶树不同组织及叶蝉胁迫处理下的表达模式分析，虫害胁迫显著诱导部分HD-CsCADs基因表达上调，可能通过调控木质素合成增强茶树机械防御能力。其中HD-CsCAD-11和HD-CsCAD-15基因的表达较明显，这2个基因可能通过调控木质素生物合成参与茶树抗虫防御反应，可作为分析茶树-昆虫互作机制的关键候选基因。本研究为茶树抗虫分子机制解析及后续开展CAD基因功能验证提供理论依据。
基金
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  • 中国乌龙茶产业协同创新中心开放研究基金项目(2024W02)
  • 福建张天福茶叶发展基金会科技创新基金项目(FJZTF01)
文献
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2025年第46卷第10期
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doi: 10.3969/j.issn.1000-2561.2025.10.004
  • 接收时间：2025-04-27
  • 首发时间：2026-03-09
  • 出版时间：2025-10-25
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  • 收稿日期：2025-04-27
  • 录用日期：2025-06-13
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中国乌龙茶产业协同创新中心开放研究基金项目(2024W02)
福建张天福茶叶发展基金会科技创新基金项目(FJZTF01)
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    福建农林大学园艺学院/茶学福建省高校重点实验室，福建福州　350002

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*金珊，E-mail：。
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
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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