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水稻染色質(zhì)重塑因子CHR729對(duì)激素代謝的影響

2015-03-20 08:38李健健
湖北農(nóng)業(yè)科學(xué) 2015年2期
關(guān)鍵詞:干旱激素水稻

李健健

摘要:CHR729是水稻(Oryza sativa L.)CHD3類染色質(zhì)重塑因子,廣泛參與水稻生長發(fā)育進(jìn)程,并影響全基因組明顯表達(dá)和組蛋白修飾,測定了多種植物內(nèi)源激素在chr729突變體中的含量,探討了CHR729與激素代謝之間的聯(lián)系。結(jié)果表明,在苗期地上部分和成熟期劍葉中,chr729突變體內(nèi)源生長素(IAA),脫落酸(ABA),茉莉酸(JA)含量明顯降低,生長素合成以及ABA合成基因的表達(dá)在chr729突變體中均呈下調(diào)表達(dá)。樹脂切片結(jié)果顯示,chr729突變體中莖稈成熟組織細(xì)胞明顯變小,居間分生組織細(xì)胞大小無明顯變化。苗期檢測細(xì)胞分裂素氧化酶/脫氫酶(CKX)基因表達(dá)發(fā)現(xiàn),chr729突變體中CKX1、CKX2、CKX4基因表達(dá)量升高,伴隨一些位點(diǎn)組蛋白變體H2A.Z富集程度的上升,表明CHR729可能抑制H2A.Z在基因組上的裝載,廣泛參與水稻激素代謝調(diào)控。

關(guān)鍵詞:水稻(Oryza sativa L.);染色質(zhì)重塑因子;激素;干旱

中圖分類號(hào):Q756 ? ? ? ?文獻(xiàn)標(biāo)識(shí)碼:A ? ? ? ?文章編號(hào):0439-8114(2015)02-0468-06

DOI:10.14088/j.cnki.issn0439-8114.2015.02.055

Effects of Rice Chromatin Remodeler CHR729 on Hormone Metabolism

LI Jian-jian

(College of Life Science and Technology, Huazhong Agricultural University, Wuhan 430070, China)

Abstract:CHR729 is a CHD3 chromatin remodeler playing multiple roles in growth and development of rice. Previous studies indicated that CHR729 modulate a subset of gene expression and histone modifications. The contents of multiple endogenous phytohormones were measured. The correlation of CHR729 with hormone dynamics was investigated. A significant decrease in endogenous auxin, abscisic acid and jasmonic acid was observed in chr729 seedlings and mature flag leaves. Genes involved in auxin or abscisic acid biosynthesis were down-regulated in chr729 mutant. The cell length in mature tissue and intercalary meristem in stem was examined. A significant decrease in cell length of mature tissue of chr729 was found. Cell size of intervening meristem had no significant changes. Some cytokinin oxidase/dehydrogenase genes were up-regulated in chr729 mutant, with a relatively high enrichment of histone variant H2A.Z at some loci, indicating the involvement of CHR729 in the deposition/removal of H2A.Z. It is implied that CHR729 was involved in multiple hormone metabolisms of rice.

Key words: rice; chromatin remodeler; phytohormone; drought

植物激素是植物合成產(chǎn)生的一類小分子化合物。截至目前,多種分子被認(rèn)定為植物激素,包括生長素或吲哚乙酸(IAA)、細(xì)胞分裂素、脫落酸(ABA)、赤霉素、乙烯、油菜素甾醇、茉莉酸、水楊酸等,均對(duì)植物生長發(fā)育起著重要的調(diào)控作用。生長素調(diào)控植物向性運(yùn)動(dòng)、頂端優(yōu)勢(shì)、根的發(fā)生等一系列生理過程[1-3]。細(xì)胞分裂素參與頂端優(yōu)勢(shì)、莖分生組織形成、葉片衰老、營養(yǎng)物質(zhì)的運(yùn)輸、種子萌發(fā)以及植物抗病應(yīng)答等[4]。脫落酸調(diào)控種子休眠,并在植物非生物脅迫響應(yīng)中發(fā)揮重要作用[5]。茉莉酸參與植物抗病,影響植物發(fā)育[6]。在不同的環(huán)境條件下,植物激素含量是高度動(dòng)態(tài)的。

真核生物的遺傳物質(zhì)存在于染色質(zhì)中,染色質(zhì)是經(jīng)多級(jí)組裝形成的高度動(dòng)態(tài)的結(jié)構(gòu)。染色質(zhì)的不同狀態(tài),影響轉(zhuǎn)錄機(jī)器對(duì)DNA序列的接近與識(shí)別。染色質(zhì)結(jié)構(gòu)的動(dòng)態(tài)變化主要受到兩類復(fù)合物調(diào)控,一類是組蛋白共價(jià)修飾酶類;另一類是ATP依賴的染色質(zhì)重塑復(fù)合物。SWI2/SNF2家族的ATP酶是染色質(zhì)重塑復(fù)合物的核心成員,其通過水解ATP釋放能量,驅(qū)動(dòng)染色質(zhì)構(gòu)象發(fā)生改變,增加或者降低DNA結(jié)合蛋白對(duì)DNA序列的可接近程度,進(jìn)而調(diào)控基因的表達(dá)[7]。擬南芥(Arabidopsis thaliana)中一些SWI2/SNF2家族成員已被鑒定參與調(diào)控?cái)M南芥生長發(fā)育的多個(gè)進(jìn)程。CHD3類染色質(zhì)重塑因子PICKLE(PKL)參與擬南芥種子萌發(fā),主根和側(cè)根的發(fā)育,根分生組織活性以及心皮的分化[8-15];另一個(gè)染色質(zhì)重塑因子BRAHMA(BRM)調(diào)控?cái)M南芥莖和葉片的發(fā)育以及開花等過程[16-19]。此外,PKL和BRM也與多種激素調(diào)控相關(guān),如生長素、細(xì)胞分裂素、脫落酸等[12,20-23]。

水稻(Oryza sativa)的染色質(zhì)重塑因子CHR729也已被鑒定出來,CHR729功能缺失造成水稻多種生長缺陷,例如植株矮小、近軸面葉片白化、開花期推遲等[5,24]。CHR729基因缺失影響全基因組水平大量基因的表達(dá)和組蛋白修飾[24]。然而,CHR729是否影響激素代謝仍不清楚,本研究通過測定多種內(nèi)源植物激素的含量,發(fā)現(xiàn)IAA、ABA、JA含量在chr729突變體中顯著降低,激素代謝相關(guān)基因的表達(dá)也受到CHR729功能突變的影響,CHR729影響水稻多種激素代謝。

1 ?材料與方法

1.1 ?材料

本試驗(yàn)使用的亞洲栽培稻粳稻品種分別為中花11(Oryza sativa L. subsp. japonica cv. Zhonghua 11),HY(Oryza sativa L. subsp. japonica cv. Hwayoung)和SSBM(Oryza sativa L. subsp. japonica cv. Ishikari-shiroge)。CHR729基因的兩個(gè)等位突變體分別為T-DNA插入突變體chr729,背景為HY,EMS誘變突變體oschr4,背景為SSBM。

1.2 ?幼苗處理

1)CHR729表達(dá)模式驗(yàn)證時(shí),正常成長的野生型中花11在不同生長時(shí)期進(jìn)行取樣并誘導(dǎo)愈傷取樣,取樣部位包括愈傷、幼苗、葉片、劍葉、莖和穗子。

2)聚乙二醇和脫落酸處理試驗(yàn)時(shí),將在生根培養(yǎng)基上無菌狀態(tài)下發(fā)芽13 d的幼苗移出無菌環(huán)境煉苗1 d后,分別用20% PEG 6000和2 μmol/L ABA浸泡幼苗并在不同的時(shí)間點(diǎn)取樣。

1.3 ?基因表達(dá)檢測

野生型HY和chr729突變體水稻種子去殼并經(jīng)過表面消毒后,接種至生根培養(yǎng)基上,正常光照條件下生長11 d后取地上部分,按照TRIzol試劑(Invitrogen公司)說明書提取水稻RNA。取4 μg總RNA,用1 μL DNase I (invitrogen) 37 ℃ 消化15 min去除 DNA污染,加1 μL 25 mmol/L EDTA后,冰上放置5 min,65 ℃滅活10 min;在冰上冷卻2 min后,加1 μL Oligo dT15(100 ng)和1 μL 10 mmol/L dNTPs 65 ℃靜置5 min;冰上放置2 min,加入4 μL 5×Buffer、1 μL ?M-MLV、1 μL HPRI(RNA 酶抑制劑)、2 μL 0.1 mol/L DTT,37 ℃反轉(zhuǎn)錄1.5 h;100 ℃滅活10 min,加入140 μL ddH2O。Real-time PCR使用ABI SYBR green試劑盒,儀器為ABI 7500 Real time PCR system。每個(gè)樣品以肌動(dòng)蛋白(actin)為內(nèi)參,重復(fù)3次。

1.4 ?組蛋白突變體H2A.Z在基因位點(diǎn)的富集檢測

染色質(zhì)免疫沉淀(ChIP)操作方法參照文獻(xiàn)[25]。取正常條件下生根培養(yǎng)11 d的水稻幼苗約2 g置于1%甲醛中真空下交聯(lián)后液氮中研磨,染色質(zhì)被超聲波打成200~750 bp的片段,以H2A.Z抗體進(jìn)行免疫沉淀。抗體沉淀前后的DNA以基因特異性引物進(jìn)行熒光定量PCR檢測,分析H2A.Z在基因位點(diǎn)的富集情況。

1.5 ?激素測定

植物內(nèi)源激素IAA、ABA和JA的含量測定參照文獻(xiàn)[26]進(jìn)行。取正常條件下生根培養(yǎng)11 d的水稻幼苗地上部分約0.1 g,液氮研磨后加入到750 μL預(yù)冷的提取緩沖液中(甲醇∶水∶醋酸=80∶19∶1),緩沖液中加入內(nèi)標(biāo)(10 ng 2H6 ABA,10 ng DHJA, 5 ng D2-IAA,3 μg NAA),4 ℃搖床300 r/min避光提取16 h以上。4 ℃,13 000 r/min離心15 min后轉(zhuǎn)移上清液,沉淀繼續(xù)以400 μL緩沖液重懸,搖床提取4 h以上。離心后合并上清液,用1 mL注射器吸取合并的上清并過0.22 μm濾膜(津騰公司,尼龍66)至另一離心管中,在通風(fēng)櫥中用氮?dú)獯蹈?,?00 μL甲醇顛倒幾次后4 ℃溶解3~6 h。4℃、13 000 r/min離心15 min,輕輕吸取上清液150~180 μL至內(nèi)插管中,放置于質(zhì)譜專用上樣瓶中用于質(zhì)譜分析。

2 ?結(jié)果與分析

2.1 ?聚乙二醇和脫落酸對(duì)CHR729基因表達(dá)的影響

為了解CHR729基因的功能,對(duì)水稻多個(gè)組織或器官中CHR729基因表達(dá)進(jìn)行了分析。定量PCR結(jié)果表明,CHR729基因廣泛表達(dá)于多個(gè)組織或器官,包括愈傷、幼苗、葉片、劍葉、莖和穗子等(圖1A)。其中,在葉片和愈傷中,CHR729基因的表達(dá)量相對(duì)較高,表明CHR729在葉片和愈傷中可能發(fā)揮重要功能,并且chr729突變體葉片近軸面白化、葉片變窄,愈傷褐化(圖1B、圖1C)也驗(yàn)證了這一點(diǎn)。CHR729基因突變導(dǎo)致多種生長缺陷,為了說明CHR729是否在增強(qiáng)水稻對(duì)環(huán)境的適應(yīng)性方面發(fā)揮功能,分別檢測了幼苗經(jīng)聚乙二醇(PEG)處理或者脫落酸(ABA)處理后CHR729基因的表達(dá)情況。結(jié)果(圖1D、圖1E)表明,CHR729基因的表達(dá)受到PEG和ABA的誘導(dǎo)。

2.2 ? CHR729基因突變對(duì)植物激素的影響

植物激素在植株正常生長發(fā)育中發(fā)揮重要作用,為了說明chr729突變體產(chǎn)生的多種表型是否與激素水平有關(guān),分別對(duì)苗期地上部分和成熟期劍葉中IAA、ABA和JA的含量進(jìn)行了分析。由圖2可以看出,CHR729基因突變后,IAA、ABA和JA的含量均明顯下降。為了說明CHR729是否影響激素合成,在幼苗期地上部分對(duì)生長素合成基因表達(dá)進(jìn)行檢測,發(fā)現(xiàn)生長素合成相關(guān)的大部分YUCCA類基因的表達(dá)量在突變體中均下降。此外,對(duì)苗期ABA合成相關(guān)基因的表達(dá)也發(fā)現(xiàn),OsZEP、OsLCY、OsZDS、OsPDS的表達(dá)在CHR729基因的兩個(gè)等位突變體(chr729和oschr4)中表達(dá)量均下降,說明CHR729基因影響了ABA合成。

2.3 ?干旱對(duì)chr729突變體的影響

CHR729基因的表達(dá)受到PEG和ABA的誘導(dǎo),且chr729突變體中內(nèi)源ABA含量降低,推測CHR729可能參與抗旱應(yīng)答。通過測定野生型與chr729突變體離體葉片失水速率,發(fā)現(xiàn)chr729突變體中失水速率比野生型快。進(jìn)一步對(duì)野生型和chr729突變體植株進(jìn)行干旱處理,發(fā)現(xiàn)chr729突變體在干旱條件下葉片更容易表現(xiàn)出卷曲的表型,而正常澆水條件下,葉片均表現(xiàn)為自然伸展,說明CHR729基因突變后,植株對(duì)干旱敏感性增強(qiáng)(圖3)。

2.4 ?CHR729對(duì)細(xì)胞分裂和伸長的影響

chr729突變體表現(xiàn)出株高變矮的表型,為了說明株高變矮與細(xì)胞分裂和細(xì)胞伸長的關(guān)系,對(duì)成熟莖稈進(jìn)行了縱向樹脂切片,并對(duì)居間分生組織和成熟組織細(xì)胞大小進(jìn)行了統(tǒng)計(jì)。結(jié)果表明,chr729突變體成熟組織中細(xì)胞大小比野生型細(xì)胞明顯變小,而居間分生組織突變體與野生型的細(xì)胞大小無明顯差異,說明chr729突變體株高變矮與成熟組織細(xì)胞伸長減弱有關(guān)。此外,通過細(xì)胞分裂素氧化酶/脫氫酶(CKX)的表達(dá)可以反映細(xì)胞分裂的快慢。對(duì)CKX基因的表達(dá)模式進(jìn)行了分析,發(fā)現(xiàn)CKX1、CKX2、CKX4、CKX5、CKX8、CKX11在苗期地上部分有表達(dá)。進(jìn)一步Real-time PCR驗(yàn)證發(fā)現(xiàn)在chr729突變體幼苗地上部分CKX1、CKX2、CKX4基因的表達(dá)明顯上升,同時(shí)染色質(zhì)免疫沉淀(ChIP)發(fā)現(xiàn)突變體中H2A.Z在CKX2、CKX4、CKX8、CKX11的基因位點(diǎn)富集程度明顯高于野生型,說明CHR729可能通過抑制H2A.Z在基因組一些位點(diǎn)的裝載進(jìn)而抑制基因的表達(dá)(圖4)。

3 ?小結(jié)與討論

3.1 ?CHR729在水稻激素代謝中的作用

植物激素在植物的正常生長過程中有著非常重要的作用。在擬南芥中,PKL和BRM與多種激素應(yīng)答有關(guān)[12,20-23],水稻chr729突變后會(huì)出現(xiàn)多種表型[5,24]。本研究分析結(jié)果也表明,CHR729突變會(huì)導(dǎo)致多種激素水平的下降,包括IAA、ABA和JA,并且伴隨著IAA和ABA合成相關(guān)基因表達(dá)量的下降以及細(xì)胞分裂素分解代謝相關(guān)基因表達(dá)量的上升。進(jìn)一步推測CHR729可以通過調(diào)整體內(nèi)不同激素水平或者調(diào)控激素應(yīng)答以調(diào)控水稻的正常生長。

3.2 ?CHR729在H2A.Z裝載過程中的抑制作用

組蛋白突變體H2A.Z通常會(huì)被SWR1類的ATP酶裝載到基因5末端,從而促進(jìn)相應(yīng)基因的轉(zhuǎn)錄[27-29],并被INO80類的ATP酶清除以抑制基因的表達(dá)[30]。擬南芥中與SWR1復(fù)合物組分的同源蛋白被報(bào)道參與了葉片發(fā)育、開花、DNA修復(fù)、體細(xì)胞重組、減數(shù)分裂以及系統(tǒng)獲得性抗性應(yīng)答等過程[31-34],擬南芥INO80蛋白則參與控制同源重組過程[35],CHD蛋白家族也被報(bào)道參與組蛋白H3突變體的裝載過程[36-39],但它與H2A.Z突變體之間的聯(lián)系還不是很清楚。本研究發(fā)現(xiàn)CHR729能夠抑制H2A.Z在一些基因位點(diǎn)的富集,進(jìn)而調(diào)控基因的表達(dá)。

3.3 ?CHR729影響ABA的應(yīng)答

多數(shù)植物染色質(zhì)重塑因子重塑機(jī)制并不清楚,但是也有報(bào)道相繼闡述了其在逆境應(yīng)答中的功能。在擬南芥中,染色質(zhì)重塑因子PKL和BRM并不受外源ABA誘導(dǎo),在沒有受到非生物逆境時(shí),它們通過組蛋白修飾H3K9me2和H3K27me2以及高密度的核小體抑制ABI3和ABI5的表達(dá),從而降低了非逆境條件下ABA的應(yīng)答[22,40]。與之相反,擬南芥SWI3同源蛋白AtSWI3B,屬于SNF2染色質(zhì)重塑復(fù)合物成員,卻可以通過維持ABA應(yīng)答基因RAB18和RD29B的表達(dá)促進(jìn)對(duì)ABA的響應(yīng)[41]。這些結(jié)果說明染色質(zhì)重塑因子對(duì)ABA的調(diào)控是雙向的,既可以促進(jìn),也可以抑制其作用。本研究發(fā)現(xiàn)染色質(zhì)重塑因子基因CHR729受到PEG和ABA誘導(dǎo),并且chr729突變體對(duì)干旱處理高度敏感,這些結(jié)果表明CHR729在水稻抗旱中發(fā)揮作用,但是具體的機(jī)制仍需進(jìn)一步研究。

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