茶樹變異的分子機理研究正文

摘要茶樹(Camelliasinensis(L.)O.Ktze.)是一種世界廣泛種植且具有經(jīng)濟效益的葉用作物。因為具有特定葉色的茶葉的特征化學(xué)成分與茶葉質(zhì)量密切相關(guān)，因而茶樹葉色多樣性的研究一直以來被茶業(yè)界廣為關(guān)注。本研究以黃山群體中花白茶樹單株為研究材料，發(fā)現(xiàn)花白茶樹不同于溫敏型白化茶樹和光敏型白化茶樹隨著溫度和光照強度變化而變化，經(jīng)過修剪發(fā)現(xiàn)花白性狀依然存在。于是展開花白茶樹中內(nèi)含成分、組織結(jié)構(gòu)、基因表達差異和蛋白表達差異等研究，取得以下研究結(jié)果：（1）通過內(nèi)含物質(zhì)的含量比較分析發(fā)現(xiàn)，相對于正常葉片，花白葉片的葉綠素含量顯著降低，咖啡堿含量略有增長；相較于正常葉片，花白葉片中總兒茶素含量減少了52.66%，而總游離氨基酸含量增加了52.15%，其中花白葉片的茶氨酸含量增長了62.73%。通過石蠟切片發(fā)現(xiàn)花白葉片柵欄組織細胞排列疏松，組織結(jié)構(gòu)的疏松程度顯著低于正常葉片，并且觀察花白葉片的葉綠體超微結(jié)構(gòu)發(fā)現(xiàn)花白葉片的葉綠體發(fā)育異常，沒有明顯的類囊體且存在空腔。（2）通過對花白葉片與正常葉片轉(zhuǎn)錄組學(xué)分析，共獲得3431個差異表達基因。針對茶樹特征性代謝中茶氨酸和多酚生物合成相關(guān)基因進行分析，發(fā)現(xiàn)花白葉片在兒茶素代謝途徑中的相關(guān)基因低于正常葉片，而在茶氨酸代謝途徑中的基因趨勢相反。通過分析還發(fā)現(xiàn)，花白葉片中葉綠素生物合成代謝途徑、光合作用、葉綠體發(fā)育和生物合成中差異基因的表達模式受到影響?；ò兹~片中光合作用的光捕獲天線蛋白基因（LHC）表達均顯著低于正常葉片，合成葉綠素途徑相關(guān)的差異基因表達也低于正常。（3）通過對花白茶樹中花白葉片和正常葉片的蛋白組分析，一共獲得95個差異蛋白。經(jīng)過KEGG富集分析，結(jié)果發(fā)現(xiàn)花白茶樹在氧化磷酸化，類黃酮生物途徑和苯丙素生物合成等重要通路發(fā)生了顯著變化，同時在光合作用、嘌呤代謝等代謝途徑受到影響。（4）我們在對差異基因進行篩選時發(fā)現(xiàn)了Clp蛋白酶復(fù)合體與葉綠體發(fā)育、植物葉色有關(guān)。以花白葉片和正常葉片為模板，分別擴增CsClpP5基因。經(jīng)序列分析發(fā)現(xiàn)，從花白葉片中克隆的CsClpP5基因，全長序列為909bp，編碼303個氨基酸。然而正常葉片中擴增的CsClpP5基因全長903bp，編碼301個氨基酸。經(jīng)過序列比對發(fā)現(xiàn)了兩個氨基酸位點的突變，而經(jīng)過磷酸位點的分析發(fā)現(xiàn)該突變位點的改變了ClpP5的磷酸位點。由此我們推測CsClpP5突變，可能與茶樹產(chǎn)生花白表型有關(guān)。關(guān)鍵詞：茶樹；白化；轉(zhuǎn)錄組；蛋白組；CsClpP5基因AbstractAsaleafyplant,teaplant(Camelliasinensis(l.)o.Ktze.)isanimportanteconomiccropinChina.Becausethecharacteristicchemicalcompositionofteawithspecificleafcoloriscloselyrelatedtothequalityoftea,thestudyonthecolordiversityoftealeaveshasbeenwidelyconcernedbytheteaindustry.Inthisstudy,singleplantofwhiteteatreeinHuangshanpopulationwasusedastheresearchmaterial,anditwasfoundthatthewhiteteatreewasdifferentfromthetemperature-sensitivealbinoteatreeandlight-sensitivealbinoteatree,whichchangedwiththechangeoftemperatureandlightintensity.Therefore,studieswerecarriedoutonthecomponents,tissuestructure,geneexpressiondifferencesandproteinexpressiondifferencesinthewhiteteaplant,andthefollowingresultswereobtained:(1)Itwasfoundthroughthecomparativeanalysisofthecontentsofthecontainedsubstancesthatcomparedwiththenormalleaves,thechlorophyllcontentofthewhiteleavesdecreasedsignificantly,andthecaffeinecontentincreasedslightly.Comparedwithnormalleaves,thetotalcatechincontentinthewhiteleavesdecreasedby52.66%,whilethetotalfreeaminoacidcontentincreasedby52.15%,inwhichthetheaninecontentinthewhiteleavesincreasedby62.73%.Throughparaffinsection,itwasfoundthatpalisadetissuecellsofthepallidleaveswerearrangedloosely,andthedegreeoftissuestructurewassignificantlylowerthanthatofthenormalleaves.Theultrastructuralanalysisofthepallidleavesrevealedthatthechloroplastsofthepallidleaveswereabnormallydeveloped,withnoobviousthylakoidsandcavities.(2)Throughtranscriptomeanalysisofwhiteandnormalleaves,atotalof3431differentialgeneswereobtained.Basedontheanalysisofthegenesrelatedtotheanineandpolyphenolsbiosynthesisinthecharacteristicmetabolismofteatree,itwasfoundthatthegenesrelatedtocatechinmetabolismpathwayinthewhiteleaveswerelowerthanthoseinthenormalleaves,whilethegenesrelatedtotheaninemetabolismpathwaywereopposite.Theanalysisalsoshowedthattheexpressionpatternsofdifferentgenesinchlorophyllbiosynthesisordegradation,photosynthesis,chloroplastdevelopmentandbiosynthesiswereaffected.Theexpressionoflightcaptureantennaprotein(LHC)inphotosyntheticleaveswassignificantlylowerthanthatinnormalleaves,andtheexpressionofdifferentgenesrelatedtothesynthesisofchlorophyllpathwaywasalsolowerthanthatinnormalleaves.(3)Atotalof95differentiallyexpressedproteinswereobtainedbyproteomicanalysisofthewhiteandnormalleavesofthewhiteteaplant.ThroughKEGGpathwayenrichmentanalysis,theresultsshowedthattheOxidativephosphorylation(Oxidativephosphorylation),flavonoidsbiologicalpathways(Flavonoidbiosynthesis)andstyreneacrylicbiosynthesis(Phenylpropanoidbiosynthesis)andotherimportantpathwayschangeddramatically,butinphotosynthesis,purinemetabolismandothermetabolicpathwayisaffected.(4)Clpproteasecomplexwasfoundtoberelatedtochloroplastdevelopmentandleafcolorinthescreeningofdifferentialgenes.TheCsClpP5genewasamplifiedinwhiteandnormalleaves.SequenceanalysisshowedthattheCsClpP5geneclonedfromthewhiteleaveswas909bplongandencoded303aminoacids.However,theCsClpP5geneamplifiedinthenormalleaveswasfulllength903bp,encoding301aminoacids.Mutationoftwoaminoacidsiteswasfoundbysequencealignment,andanalysisofthephosphoricacidsiterevealedthatthemutationsitechangedthephosphoricacidsiteofClpP5.Therefore,wespeculatedthatthemutationofCsClpP5mightberelatedtotheproductionofwhitephenotypeinteatrees.Keywords:Camelliasinensis;albinophenotype;RNA-seq;Proteomics;CsClpP5gene1文獻綜述1.1白化茶樹的研究進展作為茶樹的發(fā)源地，中國的茶樹資源豐富，其中白化茶樹因獨特茶味，包括高鮮味和低澀味，使得白化茶樹從茶樹(Camelliasinesnis)中脫穎而出。根據(jù)不同的環(huán)境因素，可以將白化茶樹分為光敏感型白化茶樹、溫度敏感型白化茶樹和生態(tài)不敏感型白化茶樹ADDINEN.CITE<EndNote><Cite><Author>王開榮</Author><Year>2008</Year><RecNum>1</RecNum><DisplayText>[1]</DisplayText><record><rec-number>1</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">1</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>王開榮</author><author>梁月榮</author><author>張龍杰</author><author>李明</author></authors></contributors><titles><title>白化茶種質(zhì)資源的分類及特性</title><secondary-title>中國茶葉</secondary-title></titles><periodical><full-title>中國茶葉</full-title></periodical><pages>9-11</pages><volume>30</volume><number>8</number><dates><year>2008</year></dates><urls></urls></record></Cite></EndNote>[\o"王開榮,2008#1"1]。1.1.1光敏型白化茶樹黃金芽、白雞冠、御金香為光敏型白化茶樹的主要品種，葉片在強光照(>15000lx)條件下黃化，但隨著光照強度的降低或遮蔭條件下逐漸恢復(fù)綠色ADDINEN.CITEADDINEN.CITE.DATA[\o"Zhang,2017#2"2-4]。溫敏型茶樹的白化一般出現(xiàn)在晚春寒潮期，這種特殊的氣候現(xiàn)象導(dǎo)致氣溫明顯低于年平均氣溫，有時甚至?xí)抵?0℃以下ADDINEN.CITE<EndNote><Cite><Author>Sharma</Author><RecNum>5</RecNum><DisplayText>[5]</DisplayText><record><rec-number>5</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">5</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Sharma,R.R.</author><author>V.B.Patel</author><author>HareKrishna</author></authors></contributors><titles><title>Relationshipbetweenlight,fruitandleafmineralcontentwithalbinismincidenceinstrawberry(FragariaxananassaDuch.)</title><secondary-title>ScientiaHorticulturae</secondary-title></titles><periodical><full-title>ScientiaHorticulturae</full-title></periodical><pages>0-70</pages><volume>109</volume><number>1</number><dates></dates><urls></urls></record></Cite></EndNote>[\o"Sharma,#5"5]。黃金芽的白化葉片大大減少了葉綠素的含量，經(jīng)檢測黃金芽白化葉片的葉綠素a含量減少了96%以及葉綠素b降低了77%。黃金芽的超微結(jié)構(gòu)顯示白化葉片的葉綠體在前質(zhì)體期被抑制，沒有基粒結(jié)構(gòu)，取而代之的是普遍存在的嗜鋨顆粒。黃金芽的白化葉片比非白化葉片含有茶氨酸、谷氨酸、谷氨酰胺、天冬氨酸分別升高31.0、39.8、50.8、62.2%ADDINEN.CITE<EndNote><Cite><Author>Zhang</Author><Year>2017</Year><RecNum>2</RecNum><DisplayText>[2]</DisplayText><record><rec-number>2</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">2</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Zhang,Qunfeng</author><author>Liu,Meiya</author><author>Ruan,Jianyun</author></authors></contributors><titles><title>IntegratedTranscriptomeandMetabolicAnalysesRevealsNovelInsightsintoFreeAminoAcidMetabolisminHuangjinyaTeaCultivar</title><secondary-title>Frontiersinplantscience</secondary-title><alt-title>FrontPlantSci</alt-title></titles><periodical><full-title>Frontiersinplantscience</full-title><abbr-1>FrontPlantSci</abbr-1></periodical><alt-periodical><full-title>Frontiersinplantscience</full-title><abbr-1>FrontPlantSci</abbr-1></alt-periodical><pages>291-291</pages><volume>8</volume><keywords><keyword>Camelliasinensis</keyword><keyword>chloroticmutation</keyword><keyword>freeaminoacid</keyword><keyword>metabolism</keyword><keyword>nitrogenmetabolism</keyword></keywords><dates><year>2017</year></dates><publisher>FrontiersMediaS.A.</publisher><isbn>1664-462X</isbn><accession-num>28321230</accession-num><urls><related-urls><url>/28321230</url><url>/pmc/articles/PMC5337497/</url></related-urls></urls><electronic-resource-num>10.3389/fpls.2017.00291</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[\o"Zhang,2017#2"2]。葉綠體類囊體膜在白雞冠的白化葉片中受到損壞。研究發(fā)現(xiàn)白雞冠的白化葉片（不遮蔭）與對照（遮蔭）相比，兒茶素含量顯著降低，主要集中在表沒食子兒茶素、兒茶素和表兒茶素等非酯型兒茶素ADDINEN.CITE<EndNote><Cite><Author>Wu</Author><Year>2016</Year><RecNum>3</RecNum><DisplayText>[3]</DisplayText><record><rec-number>3</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">3</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wu,Quanjin</author><author>Chen,Zhidan</author><author>Sun,Weijiang</author><author>Deng,Tingting</author><author>Chen,Mingjie</author></authors></contributors><titles><title>DenovoSequencingoftheLeafTranscriptomeRevealsComplexLight-ResponsiveRegulatoryNetworksinCamelliasinensiscv.Baijiguan</title><secondary-title>Frontiersinplantscience</secondary-title><alt-title>FrontPlantSci</alt-title></titles><periodical><full-title>Frontiersinplantscience</full-title><abbr-1>FrontPlantSci</abbr-1></periodical><alt-periodical><full-title>Frontiersinplantscience</full-title><abbr-1>FrontPlantSci</abbr-1></alt-periodical><pages>332-332</pages><volume>7</volume><keywords><keyword>Camelliasinensis</keyword><keyword>antioxidantenzymes</keyword><keyword>chlorophyllsynthesis</keyword><keyword>chloroplastdevelopment</keyword><keyword>differentiallyexpressedgenes</keyword><keyword>light-shading</keyword><keyword>yellowphenotype</keyword></keywords><dates><year>2016</year></dates><publisher>FrontiersMediaS.A.</publisher><isbn>1664-462X</isbn><accession-num>27047513</accession-num><urls><related-urls><url>/27047513</url><url>/pmc/articles/PMC4801010/</url></related-urls></urls><electronic-resource-num>10.3389/fpls.2016.00332</electronic-resource-num><remote-database-name>PubMed</remote-database-name><language>eng</language></record></Cite></EndNote>[\o"Wu,2016#3"3]。御金香白化葉片的葉肉細胞中葉綠體少于遮蔭后的葉片，且葉綠體的形態(tài)異常。白化葉片中EGCG、EGC、ECG與EC均低于遮蔭后的葉片，其中槲皮素與山柰醇在御金香白化葉片中有所積累，而茶氨酸在遮蔭后的更高含量ADDINEN.CITEADDINEN.CITE.DATA[\o"Liu,2017#4"4]。1.1.2溫敏型白化茶樹安吉白茶、千年雪、小雪芽以及黃山白茶1號是溫敏型白化茶樹的主要扦插對象，它們在低溫(<20℃)下萌發(fā)出雪白嫩葉，在高溫(22至25℃)下恢復(fù)標(biāo)準(zhǔn)葉色ADDINEN.CITE<EndNote><Cite><Author>Du</Author><Year>2006</Year><RecNum>6</RecNum><DisplayText>[6,7]</DisplayText><record><rec-number>6</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">6</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Du,Y.Y.,Liang,Y.R.,Wang,H.,Wang,K.R.,Lu,J.L.,Zhang,G.H.,Lin,W.P.,Li,M.,Fang,Q.Y</author></authors></contributors><titles><title>Astudyonthechemicalcompositionofalbinoteacultivars.</title><secondary-title>J.Hortic.Sci.Biotechnol</secondary-title></titles><volume><styleface="normal"font="default"charset="134"size="100%">81</style></volume><edition><styleface="normal"font="default"charset="134"size="100%">812</style></edition><section><styleface="normal"font="default"charset="134"size="100%">809</style></section><dates><year><styleface="normal"font="default"charset="134"size="100%">2006</style></year></dates><urls></urls></record></Cite><Cite><Author>Wang</Author><Year>2010</Year><RecNum>7</RecNum><record><rec-number>7</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">7</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Wang,KR</author><author>Shao,SH</author><author>Lu,JL</author><author>Liang,YR</author></authors></contributors><titles><title>DevelopmentofspecificRAPDmarkersforidentifyingalbinoteacultivars'Qiannianxue'and'Xiaoxueya'</title><secondary-title>AfricanJournalofBiotechnology</secondary-title></titles><periodical><full-title>AfricanJournalofBiotechnology</full-title></periodical><volume>9</volume><number>4</number><dates><year>2010</year></dates><urls></urls></record></Cite></EndNote>[\o"Du,2006#6"6,\o"Wang,2010#7"7]。安吉白茶在白化期間葉綠體的發(fā)育不完全，類囊體結(jié)構(gòu)中的基粒模糊。白化嚴(yán)重時期的葉肉細胞中并未發(fā)現(xiàn)葉綠體。安吉白茶在白化期的茶氨酸和胡蘿卜素高于綠葉期ADDINEN.CITEADDINEN.CITE.DATA[\o"Li,2016#8"8]。千年雪的抗逆性比安吉白茶的強ADDINEN.CITE<EndNote><Cite><Author>王開榮</Author><Year>2007</Year><RecNum>111</RecNum><DisplayText>[9]</DisplayText><record><rec-number>111</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">111</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">王開榮,林偉平,方乾勇,李明,梁月榮,秦嶺,俞茂昌.</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">白茶新品種“千年雪”選育研究報告[J]</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">中國茶葉</style></secondary-title></titles><periodical><full-title>中國茶葉</full-title></periodical><pages><styleface="normal"font="default"charset="134"size="100%">24-26</style></pages><number><styleface="normal"font="default"charset="134"size="100%">02</style></number><dates><year><styleface="normal"font="default"charset="134"size="100%">2007</style></year></dates><urls></urls></record></Cite></EndNote>[\o"王開榮,2007#111"9]，氨基酸和多酚含量與安吉白茶所含的差異不大ADDINEN.CITE<EndNote><Cite><Author>王開榮.</Author><Year>2006</Year><RecNum>112</RecNum><DisplayText>[10]</DisplayText><record><rec-number>112</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">112</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">王開榮.</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">白化茶種質(zhì)資源綜合性狀研究[D]</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">浙江大學(xué)</style></secondary-title></titles><periodical><full-title>浙江大學(xué)</full-title></periodical><dates><year><styleface="normal"font="default"charset="134"size="100%">2006</style></year></dates><urls></urls></record></Cite></EndNote>[\o"王開榮.,2006#112"10]。與小雪芽的綠葉組織比較，發(fā)現(xiàn)小雪芽的白化葉片所含的葉綠素和類胡蘿卜素顯著不足，葉綠體嚴(yán)重破壞。此外，小雪芽的白化葉片中與品質(zhì)有關(guān)的次生代謝物的積累也發(fā)生了改變，總氨基酸、茶氨酸和谷氨酸含量顯著增加，生物堿、兒茶素和多酚含量顯著降低ADDINEN.CITE<EndNote><Cite><Author>Na-NaLi</Author><Year>2019</Year><RecNum>9</RecNum><DisplayText>[11]</DisplayText><record><rec-number>9</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">9</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Na-NaLi,Jian-LiangLu</author></authors></contributors><titles><title><styleface="normal"font="default"size="100%">DissectionofChemicalCompositionandAssociatedGeneExpressioninthePigment-DeficientTeaCultivar</style><styleface="normal"font="default"charset="134"size="100%">‘Xiaoxueya’RevealsanAlbinoPhenotypeandMetaboliteFormation</style></title><secondary-title>Frontiersinplantscience</secondary-title></titles><periodical><full-title>Frontiersinplantscience</full-title><abbr-1>FrontPlantSci</abbr-1></periodical><dates><year><styleface="normal"font="default"charset="134"size="100%">2019</style></year></dates><urls></urls></record></Cite></EndNote>[\o"Na-NaLi,2019#9"11]。黃山白茶1號在白化時期所含的葉綠素含量低而游離氨基酸含量增加。其中黃山白茶1號白化階段總氨基酸的含量是復(fù)綠階段的2.6倍ADDINEN.CITE<EndNote><Cite><Author>ZhougaoShen</Author><Year>2020</Year><RecNum>118</RecNum><DisplayText>[12]</DisplayText><record><rec-number>118</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">118</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>ZhougaoShen,YanHe,YeyunLi,TianyuanYang,XiaoniuXu,ChangjunJiang</author></authors></contributors><titles><title>Insightsintotheprofilingchangesofaminoacidcontentinanalbinomutant(Camelliasinensiscv.Huangshanbaicha)duringthealbinosticstage[J]</title><secondary-title>ScientiaHorticulturae</secondary-title></titles><periodical><full-title>ScientiaHorticulturae</full-title></periodical><volume><styleface="normal"font="default"charset="134"size="100%">260</style></volume><dates><year><styleface="normal"font="default"charset="134"size="100%">2020</style></year></dates><urls></urls></record></Cite></EndNote>[\o"ZhougaoShen,2020#118"12]。1.1.3生態(tài)不敏感型白化茶樹不同于光敏型與溫敏型白化茶樹受環(huán)境所影響，在不考慮環(huán)境的情況下，產(chǎn)生穩(wěn)定的、且可遺傳的雜色葉片稱為生態(tài)不敏感型白化茶樹。其中主要包括炎陵銀邊茶、黃金斑、銀紅九號等變異體ADDINEN.CITEADDINEN.CITE.DATA[\o"盧翠,2018#10"13-15]。變異的炎陵銀邊茶存在復(fù)色的表型，研究中發(fā)現(xiàn)白化部分中葉綠體的基粒發(fā)育不良，并有許多的空泡結(jié)構(gòu)存在。其內(nèi)含成分咖啡堿、表沒食子兒茶素沒食子酸酯和沒食子兒茶素沒食子酸酯以及表兒茶素沒食子酸酯比正常組織含量低ADDINEN.CITE<EndNote><Cite><Author>盧翠</Author><Year>2018</Year><RecNum>10</RecNum><DisplayText>[13]</DisplayText><record><rec-number>10</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">10</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">盧翠</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">炎陵銀邊茶斑錦變異的分子機理研究</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">湖南農(nóng)業(yè)大學(xué)</style></secondary-title></titles><periodical><full-title>湖南農(nóng)業(yè)大學(xué)</full-title></periodical><dates><year><styleface="normal"font="default"charset="134"size="100%">2018</style></year></dates><urls></urls></record></Cite></EndNote>[\o"盧翠,2018#10"13]。黃金斑的研究較少，是黃金芽的白化變種，黃金斑茶樹在視覺上構(gòu)成了黃綠相間的復(fù)色狀態(tài)ADDINEN.CITE<EndNote><Cite><Author>韓震</Author><Year>2013</Year><RecNum>12</RecNum><DisplayText>[14]</DisplayText><record><rec-number>12</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">12</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">韓震,王開榮,鄧隆,李明,張龍杰</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">白化茶樹新品系——黃金斑</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">茶葉</style></secondary-title></titles><periodical><full-title>茶葉</full-title></periodical><pages><styleface="normal"font="default"charset="134"size="100%">125-126</style></pages><volume><styleface="normal"font="default"charset="134"size="100%">39</style></volume><number><styleface="normal"font="default"charset="134"size="100%">03</style></number><dates><year><styleface="normal"font="default"charset="134"size="100%">2013</style></year></dates><urls></urls></record></Cite></EndNote>[\o"韓震,2013#12"14]。黃化的英紅九號中葉綠體缺乏結(jié)構(gòu)良好的類囊體膜，類囊體腫脹，且葉綠體的形態(tài)異常ADDINEN.CITEADDINEN.CITE.DATA[\o"Ma,2016#105"16]。1.1.4白化茶樹的研究前景與其他植物最大的不同在于，白化茶樹具有人類所追求的品質(zhì)特征。茶氨酸和兒茶素是茶樹中最重要的次生代謝物，它們不僅在調(diào)節(jié)碳氮代謝方面有顯著的作用，并且對茶的風(fēng)味也有貢獻，并與人體健康和藥用價值呈正相關(guān)ADDINEN.CITE<EndNote><Cite><Author>Chacko</Author><Year>2010</Year><RecNum>13</RecNum><DisplayText>[17,18]</DisplayText><record><rec-number>13</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">13</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Chacko,SabuM</author><author>Thambi,PriyaT</author><author>Kuttan,Ramadasan</author><author>Nishigaki,Ikuo</author></authors></contributors><titles><title>Beneficialeffectsofgreentea:Aliteraturereview</title><secondary-title>ChineseMedicine</secondary-title></titles><periodical><full-title>ChineseMedicine</full-title></periodical><volume>5</volume><number>1</number><dates><year>2010</year></dates><urls></urls></record></Cite><Cite><Author>Haqqi</Author><RecNum>14</RecNum><record><rec-number>14</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">14</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Haqqi,T.M.</author><author>Anthony,D.D.</author><author>Gupta,S.</author><author>Ahmad,N.</author><author>Lee,M.-S.</author><author>Kumar,G.K.</author><author>Mukhtar,H.</author></authors></contributors><titles><title>Preventionofcollagen-inducedarthritisinmicebyapolyphenolicfractionfromgreentea</title><secondary-title>ProceedingsoftheNationalAcademyofSciencesoftheUnitedStatesofAmerica</secondary-title></titles><periodical><full-title>ProceedingsoftheNationalAcademyofSciencesoftheUnitedStatesofAmerica</full-title></periodical><pages>4524-4529</pages><volume>96</volume><number>8</number><dates></dates><urls></urls></record></Cite></EndNote>[\o"Chacko,2010#13"17,\o"Haqqi,#14"18]。白化茶樹的白化期L-茶氨酸含量是常規(guī)茶樹品種的0.5到2倍，但總兒茶素含量只有常規(guī)茶樹品種的一半，導(dǎo)致多酚和氨基酸的比例被顯著降低，這使得白化茶樹加工的綠茶不含苦味，而具有非凡的鮮味ADDINEN.CITE<EndNote><Cite><Author>Guangyu</Author><RecNum>16</RecNum><DisplayText>[19]</DisplayText><record><rec-number>16</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">16</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Guangyu</author><author>Zou</author><author>Yanzhong</author><author>Xiao</author><author>Miaosen</author><author>Wang</author><author>Hongmei</author><author>Zhang</author></authors></contributors><titles><title>Detectionofbitternessandastringencyofgreenteawithdifferenttastebyelectronicnoseandtongue</title></titles><dates></dates><urls></urls></record></Cite></EndNote>[\o"Guangyu,#16"19]。目前，由于白化突變，白化茶樹的種植收入是普通品種的十倍ADDINEN.CITE<EndNote><Cite><Author>任富麗</Author><Year>2015</Year><RecNum>17</RecNum><DisplayText>[20]</DisplayText><record><rec-number>17</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">17</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">任富麗,王文龍,楊國慶.,</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">安吉白茶產(chǎn)業(yè)發(fā)展面臨的主要問題及其對策[J]</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">湖州師范學(xué)院學(xué)報</style></secondary-title></titles><periodical><full-title>湖州師范學(xué)院學(xué)報</full-title></periodical><pages>10-13+64</pages><volume>37</volume><number>9</number><dates><year><styleface="normal"font="default"charset="134"size="100%">2015</style></year></dates><urls></urls></record></Cite></EndNote>[\o"任富麗,2015#17"20]。與此同時，白化茶樹中白化色澤不一，存在白色、黃色、復(fù)色等獨特的色系。與天竺葵、灑金珊瑚等植株類似，都具有觀賞性價值。白化茶樹可以作為茶園觀賞品種，可應(yīng)用于園林設(shè)計、規(guī)劃等ADDINEN.CITE<EndNote><Cite><Author>王蔚</Author><Year>2017</Year><RecNum>119</RecNum><DisplayText>[21]</DisplayText><record><rec-number>119</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">119</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author><styleface="normal"font="default"charset="134"size="100%">王蔚,郭雅玲.</style></author></authors></contributors><titles><title><styleface="normal"font="default"charset="134"size="100%">白化茶品種的開發(fā)與應(yīng)用[J]</style></title><secondary-title><styleface="normal"font="default"charset="134"size="100%">食品安全質(zhì)量檢測學(xué)報</style></secondary-title></titles><periodical><full-title>食品安全質(zhì)量檢測學(xué)報</full-title></periodical><pages><styleface="normal"font="default"charset="134"size="100%">3104-3110</style></pages><volume><styleface="normal"font="default"charset="134"size="100%">8</style></volume><number><styleface="normal"font="default"charset="134"size="100%">08</style></number><dates><year><styleface="normal"font="default"charset="134"size="100%">2017</style></year></dates><urls></urls></record></Cite></EndNote>[\o"王蔚,2017#119"21]。因此，白化茶樹在飲用、食用、園林設(shè)計等方面中具有極高的開發(fā)潛力，廣泛的推廣培育并且充分的利用把握白化品種茶樹資源有著深遠的意義。1.2白化植物的分子研究白化(也稱為白化病)在植物界中是一種常見的變異，其中大量的白化突變體中在擬南芥ADDINEN.CITE<EndNote><Cite><Author>Naested</Author><RecNum>18</RecNum><DisplayText>[22]</DisplayText><record><rec-number>18</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">18</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Naested</author><author>H.</author></authors></contributors><titles><title>ArabidopsisVARIEGATED3encodesachloroplast-targeted,zinc-fingerproteinrequiredforchloroplastandpalisadecelldevelopment</title><secondary-title>JournalofCellScience</secondary-title></titles><periodical><full-title>JournalofCellScience</full-title></periodical><pages>4807-4818</pages><volume>117</volume><number>20</number><dates></dates><urls></urls></record></Cite></EndNote>[\o"Naested,#18"22]、玉米ADDINEN.CITE<EndNote><Cite><Author>Asakura</Author><RecNum>19</RecNum><DisplayText>[23]</DisplayText><record><rec-number>19</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">19</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Asakura</author><author>Y.</author></authors></contributors><titles><title>MaizeMutantsLackingChloroplastFtsYExhibitPleiotropicDefectsintheBiogenesisofThylakoidMembranes</title><secondary-title>PlantCell</secondary-title></titles><periodical><full-title>PlantCell</full-title></periodical><pages>201-214</pages><volume>16</volume><number>1</number><dates></dates><urls></urls></record></Cite></EndNote>[\o"Asakura,#19"23]、棉花突變體ADDINEN.CITE<EndNote><Cite><Author>Ladygin</Author><Year>2006</Year><RecNum>20</RecNum><DisplayText>[24]</DisplayText><record><rec-number>20</rec-number><foreign-keys><keyapp="EN"db-id="5dxdzvdpnx2p5veaavbx2ssozefptraerat5">20</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Ladygin,VG</author><author>Semenova,GA</author><author>Shega?,ID</author></authors></contributors><titles><title>Structural-functionalorganizationofchloroplastsinleavesofxantha-702mutantofGossypiumhirsutumL</title><secondary-title>Tsitologiia</secondary-title></titles><periodical><full-title>Tsitologiia</full-title></periodical><pages>537-553</pages><volume>48</volume><number>7</number><dates><year>2006</year></dates><urls></urls></record></Cite></EndNote>[\o"Ladygin,2006#20"24]和水稻突變體ADDINEN.CITEADDINEN.CITE.DATA[\o"Jiang,2007#21"25]以及番茄突變體ADDINEN.CITE<EndNote><Cite><Author>Ishiga</Author><Year>2009</Year><RecNum>22</RecNum><DisplayText>[26]</DisplayText><record><rec-number>22</rec