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    嫁接對番茄抗逆性影響機制的研究進展

    2018-02-03 16:51:23邢佳毅
    中國蔬菜 2018年2期
    關(guān)鍵詞:抗逆性嫁接苗砧木

    邢佳毅 劉 偉

    (北京市農(nóng)林科學(xué)院蔬菜研究中心,農(nóng)業(yè)部都市農(nóng)業(yè)華北重點實驗室,北京 100097)

    嫁接作為一種園藝技術(shù)最早起源于2 000多年前的東亞,當(dāng)時是用于解決以有限的耕地集約化種植蔬菜,20世紀末傳入歐洲,之后傳入北美(Kubota et al.,2008)。現(xiàn)今,嫁接主要用于提高作物產(chǎn)量、減少病蟲害發(fā)生以及降低種植成本等(Rivard &Louws,2008;Mudge et al.,2009;Rivard et al.,2010;Haroldsen et al.,2012)。嫁接的優(yōu)勢主要體現(xiàn)在以下幾方面:提高優(yōu)良品種的產(chǎn)量潛力(Oztekin et al.,2009)、提高作物適應(yīng)不良環(huán)境的能力(Rivard et al.,2010)、減少化學(xué)物質(zhì)(藥劑和肥料)的使用(Louws et al.,2010)、提高作物對水分和土壤等資源的利用率(Schwarz et al.,2012)、通過表觀遺傳學(xué)產(chǎn)生有益的基因型變異(Albacete et al.,2015)、提高果實品質(zhì)等(Lee et al.,2010;S á nchez-Rod rí guez et al.,2013)。

    番茄(Solanum lycopersicumL.)作為重要的園藝作物之一,隨著其在設(shè)施生產(chǎn)中的發(fā)展,連作障礙、次生鹽漬化、青枯病以及根結(jié)線蟲等土傳病害日益嚴重(Thompson et al.,2007)。選擇適宜的砧木嫁接不僅能夠提高番茄果實產(chǎn)量、改善品質(zhì)(Flores et al.,2010),還能夠提高植株抵御病毒、細菌、真菌和線蟲等生物脅迫的能力(Kunwar et al.,2015),以及增強植株的耐熱性、耐寒性、耐鹽性、耐旱性、耐弱光能力等(Venema et al.,2008;Colla et al.,2010;Schwarz et al.,2010)。目前,關(guān)于嫁接番茄的研究主要集中在砧木對于接穗的影響,砧木能夠通過韌皮部和木質(zhì)部為接穗的生長提供水分、養(yǎng)分、激素、代謝產(chǎn)物、多肽、有機小分子以及核酸等物質(zhì)(Esta? et al.,2005;Villalta et al.,2008;Dun et al.,2009)。砧木中的 mRNA能夠通過韌皮部傳遞到接穗中,調(diào)控葉片形態(tài)的發(fā)育(Kim et al.,2001)。

    雖然嫁接在番茄上應(yīng)用廣泛,但對于嫁接如何提高番茄抗逆性的機制仍然不太清楚。本文就近年來番茄嫁接中砧木與接穗之間有關(guān)養(yǎng)分吸收與運輸、激素調(diào)控、基因與蛋白表達等方面的研究進行簡要綜述,以期為嫁接在番茄抗逆栽培和品種改良方面提供理論參考。

    1 養(yǎng)分吸收與運輸

    優(yōu)良的番茄砧木往往根系發(fā)達,具有更強的水分和養(yǎng)分吸收能力,能夠促進作物的生長發(fā)育,為作物抗逆性的提高提供生理基礎(chǔ)(Martinez-Rodriguez et al.,2008;Flores et al.,2010)。眾所周知,砧木對于養(yǎng)分和水分的攝取能夠影響植株的表型,但是這其中的生理機制還知之甚少。

    研究表明,與Florida47自根苗相比,以Beaufort和Multifort為砧木的番茄嫁接苗對水分和N素的利用率均有所提高,并且產(chǎn)量也分別提高了近27%和30%(Djidonou et al.,2013)。砧木LA1777能夠增加接穗Moneymaker葉片中C素和N素的積累,其中C素含量增加主要是由于葉片中淀粉的累積,從而提高了番茄植株的耐寒性(Venema et al.,2008)。Khah等(2006)也證實,番茄嫁接后植株能夠通過提高P素的利用率來提高產(chǎn)量。在干旱條件下,由于植株缺乏蒸騰作用,導(dǎo)致作物根系對N素的吸收和轉(zhuǎn)運出現(xiàn)問題(Robredo et al.,2011);干旱脅迫還能抑制N代謝過程中酶的活性,從而降低植株對N的吸收(Li & Lascano,2011)。S á nchez-Rod rí guez等(2013)研究表明,以Zarina為砧木進行嫁接能夠提高干旱條件下番茄植株對N的吸收和光呼吸作用,增加番茄葉片對的光合同化產(chǎn)物。Cristina等(2017)采用4種不同的嫁接組合來研究低P脅迫對番茄生長發(fā)育的影響,通過分析韌皮部汁液和葉片中的離子種類以及各激素含量變化,發(fā)現(xiàn)低P脅迫能夠影響番茄植株對P、Ca、S和Mn元素的吸收并引起乙烯和細胞分裂素的含量變化,最后篩選出在低P環(huán)境下能夠穩(wěn)定生長的砧木Hp-type。此外,砧木還能增強番茄接穗對于 K+、Ca2+和 Mg2+的吸收(Savvas et al.,2010)。雖然養(yǎng)分吸收和同化看起來對于砧木提高植株的活力非常重要,但是除了Albacete等(2009)關(guān)于75 mmol·L-1NaCl處理下嫁接能夠通過提高番茄植株韌皮部汁液中的K+/Na+比值來提高植株的抗鹽性以及增加產(chǎn)量的報道外,鮮有關(guān)于嫁接植株體內(nèi)養(yǎng)分濃度與產(chǎn)量的相關(guān)性研究。

    砧木不僅能夠增強番茄植株對養(yǎng)分的吸收,在有些情況下還能減少植株對有毒離子的吸收和轉(zhuǎn)運(Savvas et al.,2010)。例如,在硼(B)元素過量的環(huán)境下,以Arnold為砧木的番茄嫁接苗中編碼B轉(zhuǎn)運蛋白的基因表達量下降,從而減少根系對B的吸收以及該元素在莖部的積累(Gioia et al.,2017)。

    砧木不但可以調(diào)節(jié)番茄植株對營養(yǎng)元素的吸收和運輸,而且能夠通過提高植株體內(nèi)化合物的含量和活性來調(diào)節(jié)生長發(fā)育(Fernaandez-Garciaa et al.,2004),并因此增強植株的抗逆性(Ghanem et al.,2008;S á nchez-Rod rí guez et al.,2012b)。在鹽分脅迫條件下,嫁接能夠提高番茄過氧化氫酶(CAT)、抗壞血酸過氧化物酶(APX)、脫氫抗壞血酸還原酶(DHAR)和谷胱甘肽還原酶(GR)等抗氧化酶的活性(He et al.,2009)。而在高溫脅迫條件下,嫁接番茄植株體內(nèi)谷胱甘肽過氧化物酶(GPX)和CAT的活性增強,導(dǎo)致各組織中的H2O2含量增加,從而提高番茄的耐熱性(Rosa et al.,2003)。Angelika和Dietmar(2013)研究表明,嫁接能夠緩解弱光對于番茄生長發(fā)育的抑制,與自根苗相比,以Piccolino和Classy為砧木的嫁接番茄果實中類胡蘿卜素(包括番茄紅素和β-胡蘿卜素)以及糖、酸的含量更高。在干旱脅迫條件下,采用Zarina作砧木能夠提高植株的抗壞血酸、酚類、黃酮類、番茄紅素和β-胡蘿卜素等抗氧化物質(zhì)的含量,從而提高櫻桃番茄的產(chǎn)量和果實品質(zhì)(S á nchez-Rod rí guez et al.,2012a)。

    到目前為止,關(guān)于番茄嫁接的大部分研究還是集中在篩選養(yǎng)分利用率高的砧木,或者與嫁接苗栽培配套的灌溉與施肥措施上,而從生理水平和分子水平上揭示這種高養(yǎng)分利用率的發(fā)生機制的研究還有待加強。

    2 植物激素調(diào)控

    在嫁接番茄中,砧木可以通過激素調(diào)節(jié)來影響接穗的生長(Holbrook et al.,2002;P é rez-Alfocea et al.,2010)。例如,以耐鹽野生品種為砧木嫁接栽培品種,可以提高番茄的產(chǎn)量和品質(zhì),原因在于砧木能夠增強調(diào)節(jié)葉面積與葉片衰老的根源性離子和激素因子的供應(yīng)能力(Santa et al.,2002;Fern á ndez-Gar cí a et al.,2004)。在 75 mmol·L-1NaCl處理下,與對照相比,番茄嫁接苗葉片木質(zhì)部中反玉米素(t-Z)、脫落酸(ABA)和吲哚乙酸(IAA)的濃度均發(fā)生變化(Albacete et al.,2008,2010)。此外,在鹽分脅迫條件下,將野生型番茄嫁接到一個能夠持續(xù)表達異戊烯基腺苷轉(zhuǎn)移酶(IPT)基因(調(diào)控細胞分裂素合成)的砧木(35S∶∶IPT)上,結(jié)果發(fā)現(xiàn)與野生型自根苗相比,嫁接苗的產(chǎn)量提高了30%,這種產(chǎn)量上的差異很可能是由于嫁接促進了番茄植株莖部的發(fā)育并減少了花的敗育,因為嫁接苗中t-Z的濃度比對照高1.5~2.0倍,而這種激素能夠促進發(fā)育的果實中細胞的分化和膨大,從而提高單果質(zhì)量(Ghanem et al.,2011)。Ghanem等(2011)還指出,鹽分脅迫能夠通過降低番茄莖部細胞分裂素(CK)的濃度來減少果實產(chǎn)量,而采用35S∶∶IPT砧木嫁接后,嫁接苗根部產(chǎn)生的CK能夠通過韌皮部傳遞到植株地上部分,從而緩解鹽分脅迫帶來的危害。還有報道指出,在嫁接番茄中CK等激素能夠通過調(diào)節(jié)庫源活力來提高植株的抗逆性(Albacete et al.,2014),而 Dodd等(2009)將ABA缺失突變體(fl acca)嫁接到正常的野生型番茄砧木上,發(fā)現(xiàn)根部產(chǎn)生的ABA能夠調(diào)控葉片氣孔的發(fā)育。

    目前在豌豆(Pisum sativum)和擬南芥(Arabidopsis thaliana)中已經(jīng)證實,激素信號的傳遞往往與類胡蘿卜素等化合物的代謝有關(guān)(Dun et al.,2009;Sieburth & Lee,2010),鑒別出這些分子及其信號元件能夠更好地提高嫁接番茄適應(yīng)逆境的能力。

    3 基因與蛋白表達

    番茄作為模式植物已經(jīng)完成了全基因組測序,并得到了許多性狀鮮明的突變體,這為在嫁接中篩選優(yōu)質(zhì)砧木,以及研究在番茄根部特異表達的基因如何影響地上部的形態(tài)發(fā)育提供了有利條件(Asins et al.,2010;Harada,2010)。

    轉(zhuǎn)錄組學(xué)和蛋白組學(xué)的方法是研究嫁接番茄砧木與接穗之間基因和蛋白表達變化的重要方法(Vitale et al.,2014)。Turhan等(2016) 通 過SDS-PAGE和免疫印跡的方法發(fā)現(xiàn),與自根苗相比,以Beril、Logure和Valiant為砧木的番茄嫁接苗能夠通過提高葉片中熱激蛋白HSP23和HSP60的含量來抵抗高溫脅迫。Muneer等(2016)對以B-blocking為砧木的番茄嫁接苗分別進行低溫(15℃)和高溫(30 ℃)處理,發(fā)現(xiàn)溫度脅迫下植株能夠產(chǎn)生H2O2和兩種形式的活性氧,并且嫁接苗中超氧化物歧化酶(SOD)、CAT和APX的含量也更高,通過2-DE技術(shù)共鑒定出87個差異表達蛋白,這些蛋白參與了防御、應(yīng)激反應(yīng),離子結(jié)合、運輸,光合作用和蛋白質(zhì)合成等過程。Georgia等(2017)采用營養(yǎng)液膜技術(shù)(NFT)研究發(fā)現(xiàn),低溫脅迫條件下耐低溫砧木LA1777能夠提高番茄植株莖部的干、鮮質(zhì)量,葉面積,根干質(zhì)量,氣孔導(dǎo)度,胞間CO2濃度和過氧化物酶活性等;對番茄植株葉片進行轉(zhuǎn)錄組分析,共篩選出361個差異表達基因,對這些基因進行功能分類發(fā)現(xiàn)纖維素合成可能是植株在低溫下的響應(yīng)機制之一。

    近年來的研究發(fā)現(xiàn),一些特異的RNA分子能夠通過韌皮部傳遞來調(diào)控作物的器官發(fā)育(Harada,2010),而在模式植物煙草(Nicotiana benthamiana)中已經(jīng)證實,利用轉(zhuǎn)錄后基因沉默技術(shù)(PTGS)對砧木中的特定基因進行沉默后,這種siRNA同樣能夠傳遞到接穗中(Kasai et al.,2011)。在嫁接番茄中開展類似試驗,有利于更好地了解嫁接番茄的抗逆機制。

    4 展望

    雖然嫁接番茄在提高植株抗逆性方面的作用已經(jīng)被證實,但由于具有多重抗性的優(yōu)良砧木較少,目前在生產(chǎn)上嫁接番茄的應(yīng)用面積并不是很大。鑒于此,一方面要加強對于野生番茄資源的挖掘和選育,培育出更多的優(yōu)良砧木品種;另一方面,要加深嫁接番茄砧木與接穗之間關(guān)于信號傳遞和基因表達等方面的研究,闡明嫁接番茄的抗逆機制,更好地利用嫁接這一手段來鑒定、開發(fā)和創(chuàng)造新的遺傳多樣性,從而提高番茄適應(yīng)逆境的能力。

    另外,Velez-Ramirez等(2014,2105)在研究嫁接番茄耐持續(xù)光照的試驗中,將敏感型的A131接穗和耐受型的CLT接穗同時嫁接到A131上,結(jié)果發(fā)現(xiàn)與根部相比,提高植株莖部的耐光照性更能提高番茄的產(chǎn)量,因此培育耐光照的接穗比耐光照的砧木更加重要,這一結(jié)果為研究嫁接番茄植株響應(yīng)抗逆境的信號發(fā)生部位提供了新思路。

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