高溫和干旱對水稻的影響及其機(jī)制的研究進(jìn)展

段驊 佟卉 劉燕清 許慶芬 馬駿 王春敏

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高溫和干旱對水稻的影響及其機(jī)制的研究進(jìn)展

段驊*佟卉 劉燕清 許慶芬 馬駿 王春敏

(天津市農(nóng)作物研究所 天津市農(nóng)作物遺傳育種重點(diǎn)實(shí)驗(yàn)室，天津 300384；*通訊聯(lián)系人，E-mail: duanhua2004@gmail.com)

高溫和干旱是影響水稻生長、發(fā)育、產(chǎn)量和品質(zhì)的兩個(gè)最重要的環(huán)境因子，全面理解高溫和干旱脅迫對評價(jià)氣候變化對水稻生產(chǎn)的影響至關(guān)重要。概述了高溫、干旱及其復(fù)合脅迫對水稻生長發(fā)育、產(chǎn)量形成和稻米品質(zhì)的影響；從光合作用、抗氧化系統(tǒng)、內(nèi)源激素、蔗糖-淀粉代謝途徑關(guān)鍵酶活性、分子機(jī)制等方面闡述其生理機(jī)制；提出減輕水稻高溫干旱脅迫的調(diào)控措施；對未來深入開展水稻高溫干旱逆境的研究提出建議。

水稻；高溫；干旱；高溫干旱復(fù)合脅迫；產(chǎn)量；品質(zhì)；生理機(jī)制

氣候變化是21世紀(jì)中國乃至全球農(nóng)業(yè)面臨的嚴(yán)峻挑戰(zhàn)之一，對農(nóng)業(yè)的影響直接關(guān)系到糧食安全和經(jīng)濟(jì)安全。水稻是我國最主要的糧食作物之一，生殖生長期遇35℃以上的高溫就會對水稻產(chǎn)生危害[1]。全球氣候變化報(bào)告[2]指出，近130年(1880－2012年)全球平均地表溫度上升了0.85℃，預(yù)計(jì)2016－2035年全球平均地表溫度將繼續(xù)升高0.3℃~0.7℃，水稻遭受干旱脅迫的面積將擴(kuò)大一倍。而氣候變化造成糧食產(chǎn)量和品質(zhì)的降低，主要原因在于高溫和干旱脅迫[2]。近年來國內(nèi)外學(xué)者針對高溫或干旱單一因子影響水稻的特征和機(jī)理進(jìn)行了大量研究，涉及生長發(fā)育[3-5]、生理生態(tài)[6, 7]、產(chǎn)量與品質(zhì)[8, 9]等方面。但在大田生產(chǎn)中，高溫和干旱同時(shí)發(fā)生的幾率逐年增加[10]，加重了高溫或干旱對水稻產(chǎn)量和品質(zhì)形成的危害。筆者就高溫和干旱對水稻產(chǎn)量和品質(zhì)的影響及其生理機(jī)制進(jìn)行綜述，以期進(jìn)一步認(rèn)識高溫和干旱與水稻產(chǎn)量和品質(zhì)形成的關(guān)系，為指導(dǎo)水稻高溫干旱育種、抗熱節(jié)水栽培，保障我國糧食安全提供理論參考。

1 高溫與干旱對水稻生長發(fā)育及產(chǎn)量和品質(zhì)的影響

1.1 對生長發(fā)育的影響

在不同發(fā)育時(shí)期，水稻對高溫的響應(yīng)表現(xiàn)不同，敏感程度依次為抽穗開花期＞幼穗發(fā)育期＞灌漿期[11, 12]。在抽穗開花期遇到高溫，能使開花期提前[13, 14]，致使花藥開裂不良、花粉萌發(fā)率低和花粉活力下降，最終造成水稻籽粒敗育[15-17]；幼穗發(fā)育期遇到高溫，會抑制穎花分化，導(dǎo)致穎花退化；在灌漿期遇到高溫，會縮短灌漿期，阻礙籽粒充實(shí)[18]。營養(yǎng)生長期遇到高溫會促進(jìn)水稻的生長，株高、莖蘗數(shù)、葉面積和地上部干物質(zhì)量明顯增加[19]。

水稻營養(yǎng)生長期遇到干旱脅迫，光合作用受到限制，葉面積減小、分蘗減少[20]，葉片易卷曲[21]；水稻生殖生長期對干旱脅迫高度敏感，干旱不僅導(dǎo)致葉面積減小，株高和收獲指數(shù)降低，還會阻礙水稻的生殖器官發(fā)育，如降低可育花粉數(shù)量、延長開花期、致使花藥異常開裂等[12, 22]；灌漿中后期干旱導(dǎo)致葉片早衰，灌漿持續(xù)時(shí)間縮短，同化物供應(yīng)受限，粒重降低。

有關(guān)高溫和干旱復(fù)合脅迫對水稻生長發(fā)育的影響，已有學(xué)者進(jìn)行了初步研究[17, 23]。多數(shù)研究認(rèn)為，大氣溫度和土壤溫度的升高會提高作物蒸騰耗水量和農(nóng)田蒸散量，增加作物總耗水量，造成干旱缺水或進(jìn)一步加劇干旱脅迫的危害[24-26]，如Lawas等[10]發(fā)現(xiàn)，水稻抽穗灌漿期間，高溫干旱復(fù)合脅迫顯著降低株高和生物量；但Rang等[17]利用不同基因型水稻品種為材料，在開花期設(shè)置高溫、干旱和高溫干旱復(fù)合脅迫，通過花藥開裂、花粉萌發(fā)和主穗結(jié)實(shí)率等指標(biāo)，觀察不同品種對高溫、干旱的響應(yīng)。結(jié)果表明，雖然高溫、干旱和高溫干旱雙重脅迫均導(dǎo)致主穗敗育，但高溫脅迫造成的空粒率在不同處理中最高。高溫干旱復(fù)合脅迫對主穗發(fā)育的影響并沒有遠(yuǎn)大于高溫或干旱單一脅迫的影響，這體現(xiàn)了高溫和干旱復(fù)合脅迫的獨(dú)特性和復(fù)雜性。

1.2 對產(chǎn)量及其構(gòu)成因素的影響

水稻減數(shù)分裂期和抽穗開花期是對高溫和水分脅迫最敏感的時(shí)期[27-29]。在水稻開花當(dāng)日，如果田間遭遇高溫或水分脅迫，就會導(dǎo)致花藥異常開裂[30, 31]，傳粉受阻[32, 33]和花粉發(fā)育異常[34]，造成小穗不育，易形成空秕粒，從而導(dǎo)致結(jié)實(shí)率和粒重降低，進(jìn)而使水稻減產(chǎn)[35]。營養(yǎng)生長期高溫或干旱影響分蘗發(fā)生，進(jìn)而降低有效穗數(shù)[21, 36]。

現(xiàn)有較多報(bào)道證明[37-39]，水稻抽穗開花期高溫干旱雙重脅迫會導(dǎo)致產(chǎn)量下降，高溫干旱雙重脅迫的影響大于單一高溫或干旱脅迫。在產(chǎn)量構(gòu)成要素中，結(jié)實(shí)率降幅最為明顯，而對于千粒重，則有不同的結(jié)論。多數(shù)研究認(rèn)為[33, 37, 39]，高溫干旱會顯著降低千粒重，但也有研究發(fā)現(xiàn)[38]，在脅迫環(huán)境下光合產(chǎn)物集中供應(yīng)少數(shù)籽粒，會引起千粒重在一定幅度上的增加，但粒重之增遠(yuǎn)不足以彌補(bǔ)結(jié)實(shí)率和實(shí)粒數(shù)之失，最終還是會導(dǎo)致產(chǎn)量大幅下降。

1.3 對稻米品質(zhì)的影響

稻米品質(zhì)形成是品種遺傳特性和環(huán)境條件綜合作用的結(jié)果[11]。在環(huán)境因子中，溫度升高對稻米品質(zhì)影響的研究和進(jìn)展最多[11, 35, 40]。雖然對水稻品質(zhì)的影響尚存在諸多不確定性，這可能與品質(zhì)分析和研究比較復(fù)雜有關(guān)，但研究結(jié)果多為不利的影響。灌漿結(jié)實(shí)期是環(huán)境影響稻米品質(zhì)的關(guān)鍵時(shí)期，特別是灌漿前、中期高溫對品質(zhì)的影響最大[41]。高溫通過縮短灌漿持續(xù)期，降低光合產(chǎn)物積累和運(yùn)轉(zhuǎn)、籽粒中蔗糖-淀粉代謝酶活性以及胚乳細(xì)胞發(fā)育和淀粉體的充實(shí)等生理過程，造成整精米率降低和直鏈淀粉含量降低[42, 43]、堊白粒率和堊白度增加、蛋白質(zhì)含量和糊化溫度升高[11]，導(dǎo)致稻米品質(zhì)變劣[11, 35, 44]。

干旱對稻米品質(zhì)的影響與基因型和水分脅迫程度有關(guān)。結(jié)實(shí)期土壤適度干旱，可以顯著提高籽粒內(nèi)蔗糖-淀粉代謝途徑中關(guān)鍵酶活性和灌漿速率，降低內(nèi)源乙烯水平，顯著提高稻米的最高黏度和崩解值，降低堊白度和消減值，改善品質(zhì)，而重度干旱的結(jié)果則相反[45]。

關(guān)于高溫干旱復(fù)合脅迫下稻米品質(zhì)的變化，相關(guān)研究報(bào)道較少。筆者[33, 39]曾觀察到，抽穗灌漿早期高溫、干旱或高溫干旱雙重脅迫顯著降低了稻米的精米率、整精米率和崩解值，增加了堊白米率、堊白度和消減值，但在品種間存在很大差異，高溫及高溫干旱雙重脅迫對高溫干旱敏感型品種的影響大于耐熱耐旱型品種，稻米品質(zhì)在高溫干旱雙重脅迫下變劣的幅度大于單一脅迫。高煥曄等[46]研究也表明，在相同脅迫時(shí)間內(nèi)，高溫干旱復(fù)合脅迫導(dǎo)致稻米直鏈淀粉含量降低和蛋白質(zhì)含量增高的效應(yīng)遠(yuǎn)大于單一高溫脅迫和干旱脅迫。表1總結(jié)了高溫和干旱對水稻不同生育階段生長發(fā)育及產(chǎn)量和品質(zhì)的影響。

2 高溫與干旱影響水稻生長發(fā)育與產(chǎn)量形成的生理機(jī)制

2.1 光合作用

水稻遭遇高溫或干旱單一脅迫會導(dǎo)致光合作用速率下降[35]，一種原因是氣孔限制，即脅迫促使氣孔關(guān)閉，降低氣孔導(dǎo)度，導(dǎo)致CO2供應(yīng)受阻，進(jìn)而降低光合作用和物質(zhì)生產(chǎn)[54]；另一種是非氣孔限制，即脅迫通過影響植株內(nèi)Rubisco活性和光系統(tǒng)PSⅡ結(jié)構(gòu)而抑制光合作用[55]?，F(xiàn)有研究均表明，水稻在高溫干旱復(fù)合脅迫下光合作用速率較單一脅迫下大幅降低[56-60]。然而，水稻在高溫干旱復(fù)合脅迫下光合作用速率降低的原因，究竟是由氣孔因素還是非氣孔因素引起，尚無定論。趙鳳云等[60]認(rèn)為，葉綠素含量及其比例是高溫干旱復(fù)合脅迫下水稻光合作用降低的原因；劉照等[59]發(fā)現(xiàn)，氣孔因素和非氣孔因素均會導(dǎo)致高溫干旱復(fù)合脅迫下水稻光合速率下降；高煥曄等[56]雖然比較了水稻在高溫、干旱單一脅迫與雙重脅迫間的光合生理差異，包括葉綠素含量和氣孔導(dǎo)度等參數(shù)的變化，但是不同脅迫下影響光合作用的氣孔限制與非氣孔限制的比重如何，并未詳細(xì)闡述；Perdomo等[58]研究觀察到，水稻在干旱下光合速率降低的原因是氣孔因素，而在高溫下以及在高溫干旱復(fù)合脅迫下，光合降低的原因與水稻植株內(nèi)Rubisco活性降低顯著相關(guān)。

表1 高溫和干旱對水稻不同生育階段生長發(fā)育及產(chǎn)量和品質(zhì)的影響

2.2 抗氧化系統(tǒng)

活性氧(reactive oxygen species，ROS)在調(diào)控生物進(jìn)程(如生長、發(fā)育和響應(yīng)逆境脅迫)中作為信號分子起關(guān)鍵作用[61]。在干旱或高溫等逆境脅迫下，活性氧的積累會導(dǎo)致大量細(xì)胞氧化損傷，抑制水稻光合作用[62]。為防止植物體受損，活性氧會被抗氧化機(jī)制清除。植物內(nèi)的抗氧化系統(tǒng)可分為酶促系統(tǒng)和非酶促系統(tǒng)兩大類，酶促系統(tǒng)主要包括超氧化物歧化酶(SOD)、谷胱甘肽還原酶(GR)、過氧化物酶(POD)和過氧化氫酶(CAT)等，非酶促系統(tǒng)主要由抗壞血酸(ASA)和還原性谷胱甘肽(GSH)構(gòu)成?？寡趸到y(tǒng)也會受到逆境脅迫的影響[61]。

現(xiàn)有研究均表明，高溫脅迫或干旱脅迫下，水稻抗氧化酶活性下降，清除能力降低，活性氧積累增加，膜脂過氧化作用加劇，細(xì)胞膜結(jié)構(gòu)和功能受到破壞，質(zhì)膜透性增加，劍葉生理生化機(jī)能受到傷害，光合速率降低[21, 49, 63, 64]。而高溫或干旱脅迫對抗氧化系統(tǒng)的影響程度因品種的耐熱性不同而存在差別[52, 65]。最近，Lai等[66]發(fā)現(xiàn)生物鐘調(diào)控基因在晝夜節(jié)律條件下表現(xiàn)出特定的表達(dá)階段，白天ROS清除系統(tǒng)的效率較高。Byeon和Back[67]觀察到，夜間高溫對水稻的影響與褪黑素的產(chǎn)生密切相關(guān)，褪黑素作為一種有效的抗氧化劑，能有效清除夜間高溫下植物細(xì)胞中的ROS。因此，夜間ROS清除系統(tǒng)效率較低可能是水稻生產(chǎn)對夜間溫度升高更敏感的原因之一。

有研究報(bào)道[37, 68]，高溫干旱復(fù)合脅迫下，水稻葉片中的抗氧化酶SOD和POD的活性降低，丙二醛、脯氨酸及超氧陰離子含量增加，降低了水稻的抗氧化能力；但也有研究發(fā)現(xiàn)[69]，在高溫脅迫、干旱脅迫及高溫與干旱的復(fù)合脅迫下，水稻葉片的SOD、POD、CAT的活性總體上均顯著高于對照，但SOD在高溫與嚴(yán)重干旱的長期復(fù)合脅迫下，其活性比對照顯著下降。

2.3 內(nèi)源激素

根據(jù)植物激素對植物生長發(fā)育的調(diào)控作用可以將其分為抑制型植物激素和促進(jìn)型植物激素兩類。通常將脫落酸(abscisic acid，ABA)和乙烯稱為抑制型植物激素，將生長素(indole-3-acetic acid，IAA)、細(xì)胞分裂素(cytokinins，CTK)和赤霉素(gibberellin，GA)稱為促進(jìn)型植物激素。楊建昌等[70]研究表明，在減數(shù)分裂期遭受水分脅迫，穎花中ABA、乙烯和1-氨基環(huán)丙烷-1-羧酸(ACC)濃度顯著增加。Yang等[71]發(fā)現(xiàn)，結(jié)實(shí)期適度土壤干旱，導(dǎo)致水稻籽粒激素平衡發(fā)生改變，特別是赤霉素減少和ABA增加，促進(jìn)了莖鞘中貯藏性14C的運(yùn)轉(zhuǎn)，加快了籽粒灌漿速率。還有一些研究表明，適度干旱后的復(fù)水灌溉可以顯著提高葉片[72]和籽粒[73]中的CTK含量，這將有助于提高作物的光合能力和對氮素的吸收利用[74]，加速胚乳細(xì)胞增殖，提高籽粒產(chǎn)量[75]。Tang等[76]發(fā)現(xiàn)，高溫降低水稻花藥中IAA、赤霉素含量，但增加ABA含量。

植物體內(nèi)的多胺，最常見的有腐胺(putrescine，Put)、亞精胺(spermidine，Spd)和精胺(spermine，Spm)，被普遍認(rèn)為是生長調(diào)節(jié)物質(zhì)或激素的第二信使，調(diào)節(jié)植物的生長、發(fā)育、形態(tài)建成和對環(huán)境逆境的響應(yīng)[77-79]。曹云英等[3, 80]報(bào)道，灌漿期高溫脅迫引起劍葉多胺積累，耐熱性強(qiáng)的品種積累得更多，說明多胺積累能增強(qiáng)水稻對高溫的適應(yīng)性。Yang等[81]觀察到，水稻具有較強(qiáng)的增強(qiáng)葉片多胺生物合成能力，以響應(yīng)水分脅迫。多胺在植物防御水分脅迫中的作用因多胺形式和脅迫階段而異。具有更高水平的游離型Spd/Spm和非溶性結(jié)合Put以及早期游離多胺積累的生理特性，對水稻適應(yīng)干旱更有利。

李鈺等[68]研究表明，高溫干旱復(fù)合脅迫下，水稻內(nèi)源激素中IAA、玉米素(ZT)和 GA 含量下降，而ABA和水楊酸(SA)含量上升，但作者并未比較高溫或干旱單一脅迫下的內(nèi)源激素變化。高溫干旱復(fù)合脅迫如何影響水稻籽粒內(nèi)源激素和多胺水平，目前尚缺乏充分的試驗(yàn)證據(jù)。

2.4 蔗糖-淀粉代謝途徑關(guān)鍵酶活性

稻米胚乳中的淀粉約占糙米質(zhì)量的90%以上[82]，籽粒灌漿實(shí)質(zhì)上是淀粉合成與累積的過程。源器官光合同化物(含莖鞘儲存的非結(jié)構(gòu)性碳水化合物)以蔗糖的形式經(jīng)韌皮部運(yùn)輸?shù)阶蚜?，之后在一系列酶作用下形成淀粉[83, 84]。據(jù)報(bào)道[84]，水稻胚乳發(fā)育期參與籽粒蔗糖-淀粉代謝途徑的酶有33種，但5種酶在其中起關(guān)鍵作用[84-86]。這些酶包括蔗糖合酶(sucrose synthase，EC 2.4.1.13，SuS)、腺苷二磷酸葡萄糖焦磷酸化酶(ADP glucose pyrophosphorylase, EC 2.7.7.27, AGP)、淀粉合酶(starch synthase，EC 2.4.1.21, StS)、淀粉分支酶(starch branching enzyme，EC 2.4.1.18, SBE)和淀粉脫支酶(starch debranching enzyme，EC 3.2.1.70, DBE)。其中，淀粉合酶又分為可溶性淀粉合酶(soluble starch synthase，EC 2.4.1.18, SSS)和顆粒結(jié)合型淀粉合酶(granule bound starch synthase, EC 2.4.1.242, GBSS)，每組酶有幾種同工型[87]。在灌漿期水稻籽粒里，這5種酶活性與籽粒灌漿速率和淀粉積累速率正相關(guān)[83, 84, 88]。

高溫脅迫會影響這些酶的活性，進(jìn)而影響籽粒中淀粉的生成[43, 89]。Jiang等[89]觀察到，花后高溫下SBE和GBSS活性降低，SSS活性增加，導(dǎo)致支鏈淀粉的分支頻率下降，從而導(dǎo)致胚乳支鏈淀粉長鏈的比例增加。Cheng等[90]則發(fā)現(xiàn)，高溫下AGP活性和蔗糖濃度增加，而淀粉積累和SuS活性降低。

Yang等[88]研究表明，在適度土壤干旱條件下，水稻籽粒灌漿過程中SuS、SSS和SBE活性顯著增強(qiáng)，與淀粉積累速率呈正相關(guān)。土壤干旱使AGP活性增強(qiáng)，但與淀粉積累速率相關(guān)性較小。GBSS及酸性轉(zhuǎn)化酶活性受土壤干旱的影響較小。結(jié)果表明，水稻在籽粒灌漿期遭遇土壤干旱，可以通過調(diào)節(jié)蔗糖-淀粉代謝途徑關(guān)鍵酶活性增大庫強(qiáng)，從而加快籽粒灌漿速率。

針對水稻抽穗開花期遭遇高溫、干旱單因子脅迫影響籽粒淀粉積累和酶活性的研究已較多，但高溫干旱復(fù)合脅迫研究較少。筆者[91]發(fā)現(xiàn)，水稻遭遇高溫和干旱雙重脅迫時(shí)，SuS、AGP、StS、SBE和DBE活性降低，淀粉中長鏈比例上升，短鏈比例下降，說明高溫干旱脅迫通過淀粉合成關(guān)鍵酶而影響籽粒淀粉的合成與積累，最終影響水稻品質(zhì)。

2.5 分子機(jī)制

逆境脅迫會使植物改變自身的生理生化、分子細(xì)胞水平來順應(yīng)不利的生存環(huán)境。對不同逆境脅迫下植物的不同組織器官、不同生長發(fā)育階段、不同環(huán)境脅迫因子響應(yīng)時(shí)的差異表達(dá)的功能基因進(jìn)行分析篩選，獲取關(guān)鍵功能基因和抗性之間的聯(lián)系，將有助于從轉(zhuǎn)錄水平上了解脅迫因子的傷害機(jī)理及植物適應(yīng)逆境脅迫的分子機(jī)制[92]。

Shen等[93]最近報(bào)道，類受體蛋白激酶ERECTA基因(簡稱“ER基因”)可以通過調(diào)控細(xì)胞死亡，提高轉(zhuǎn)基因作物的耐熱性。研究還發(fā)現(xiàn)，ER基因在水稻和番茄中也有相同的功能，它能使轉(zhuǎn)基因植物在正常氣溫下保證不減產(chǎn)，高溫條件下，產(chǎn)量優(yōu)勢明顯。其主要原因在于ER基因能促進(jìn)植物細(xì)胞數(shù)量增多，細(xì)胞體積增大，導(dǎo)致各器官與生物量的增大。Li等[94]成功發(fā)掘出水稻抗熱數(shù)量性狀基因(Thermo-tolerance 1)，并揭示了作物抗熱新機(jī)制，能增強(qiáng)包括水稻、草坪草和十字花科等在內(nèi)的多種植物的抗熱性，在水稻、小麥、玉米、大豆和蔬菜等作物抗熱育種中有廣泛的應(yīng)用前景。Li等[95]研究了耐熱型水稻品種(N22)和熱敏感品種(Moroberekan)的花藥、授粉前雌蕊和授粉后雌蕊的代謝組學(xué)和轉(zhuǎn)錄組學(xué)變化，發(fā)現(xiàn)復(fù)合脅迫下N22編碼糖轉(zhuǎn)運(yùn)蛋白基因()和細(xì)胞壁轉(zhuǎn)化酶基因()的表達(dá)量增高，而敏感型品種表現(xiàn)出和基因表達(dá)量降低、CSA(Carbon Starved Anther)基因表達(dá)量增高。

近年來，水稻抗逆分子機(jī)制的研究主要集中在轉(zhuǎn)錄因子及其分子調(diào)控機(jī)制方面。在水稻中，目前研究較多的轉(zhuǎn)錄因子類型主要有bZIP[96, 97]、MYB/MYC[98]、WRKY[99]、AP2/EREBP[100]和NAC[101]，它們的結(jié)構(gòu)通常由DNA結(jié)合結(jié)構(gòu)域、轉(zhuǎn)錄活化結(jié)構(gòu)域、寡聚化位點(diǎn)和核定位信號組成[102]。例如，當(dāng)、和在水稻中過表達(dá)時(shí)，其通過調(diào)節(jié)脫水蛋白、膜轉(zhuǎn)運(yùn)蛋白的表達(dá)，增強(qiáng)水稻抗旱性[103-105]。過表達(dá)能增強(qiáng)谷氨酰胺合成酶()、谷氨酰胺轉(zhuǎn)移酶()和谷氨酸脫羧酶3()等靶基因的表達(dá)，并調(diào)節(jié)其他氨基酸代謝基因，從而提高水稻的耐熱性[98]。如受熱處理誘導(dǎo)，在熱激誘導(dǎo)啟動(dòng)子的驅(qū)動(dòng)下，的超量表達(dá)顯著提高轉(zhuǎn)基因水稻幼苗的高溫和干旱抗性[106]；被絲裂原活化蛋白激酶(MAP)激活，過量表達(dá)顯著提高水稻的耐旱性[107]。ERF類轉(zhuǎn)錄因子SUB1A不僅能增強(qiáng)水稻的耐澇性，而且能提高水稻的抗旱性[108]。關(guān)于DREB類轉(zhuǎn)錄因子的研究報(bào)道較多，Chen等[109]從水稻中分離到3個(gè)與擬南芥同源的基因、和，其中過量表達(dá)和均可顯著提高水稻耐旱性。同樣，已證明NAC家族的轉(zhuǎn)錄因子在水稻的耐旱性中起關(guān)鍵作用，例如，、、和的過表達(dá)通過增加水稻根數(shù)和直徑來改善耐旱性[101, 110-113]。過表達(dá)的轉(zhuǎn)基因水稻通過調(diào)節(jié)氣孔減少水分流失，在營養(yǎng)和生殖階段均表現(xiàn)出抗旱性[114]。

3 減輕水稻高溫干旱脅迫的調(diào)控措施

3.1 選育耐熱耐旱新品種

現(xiàn)有研究已經(jīng)明確，不同品種間耐熱、耐旱的能力差異較大，選用耐高溫和抗旱品種可在一定程度上減輕高溫或干旱的危害。長期以來，我國的作物育種的方向以高產(chǎn)(含抗病蟲)和優(yōu)質(zhì)為主，但未來應(yīng)確立抗逆、廣適性的育種目標(biāo)，以此作為解決高溫、干旱等非生物逆境脅迫的主要技術(shù)途徑之一。由于耐熱和耐旱性狀是兩個(gè)有區(qū)別但又密切相關(guān)的性狀，至今雖已獲得一批具有較強(qiáng)耐熱或耐旱性的水稻品系，但既耐熱、又耐旱、還高產(chǎn)的品系較少，且尚未獲得商業(yè)用品種。因此，采用常規(guī)育種與分子育種緊密結(jié)合的技術(shù)路線可能是盡快獲得可在大面積上應(yīng)用的耐熱耐旱新類型品種的一個(gè)關(guān)鍵環(huán)節(jié)。

3.2 應(yīng)對高溫干旱的栽培調(diào)控技術(shù)

利用耕作、栽培和化控技術(shù)減輕高溫干旱等逆境對作物的傷害，特別是減少逆境對作物產(chǎn)量與品質(zhì)的不利影響，是水稻生產(chǎn)主要的措施之一。如根據(jù)高溫天氣規(guī)律和水稻高溫、干旱敏感期，通過調(diào)整播種期，使開花結(jié)實(shí)期避開高溫和干旱，已經(jīng)成為緩解開花期高溫和干旱危害的主要對策之一[115]。

合理施肥和適宜的水分管理也可有效提高水稻的抗熱抗旱能力。抽穗結(jié)實(shí)期遭受高溫脅迫，在穗分化期[116]和開花期[117]適當(dāng)施用氮肥，以及在抽穗期采用輕干濕交替灌溉[118]，可以提高根系性能和地上部植株生理活性，從而獲得較高的產(chǎn)量和較好的稻米品質(zhì)。也有研究報(bào)道，通過配合施用生物炭和磷肥，在高溫脅迫發(fā)生時(shí)，可以減輕或減緩高溫脅迫帶來的產(chǎn)量損失[119]。

前人為了緩解逆境對水稻帶來的不利影響，利用化控技術(shù)做了大量的嘗試，如噴施維生素C、維生素E、油菜素內(nèi)酯、茉莉酸甲酯、水楊酸等植物生長調(diào)節(jié)劑[120-122]，以及噴施吡唑醚菌酯[123]等化學(xué)制劑，都取得了較好的效果。今后在深入闡明水稻耐高溫、耐干旱機(jī)理的基礎(chǔ)上，需從多途徑選擇栽培調(diào)控措施，突出作物生理調(diào)控作用，即利用作物本身或給作物創(chuàng)造環(huán)境發(fā)揮作物對逆境的適應(yīng)能力和抵抗能力，以促進(jìn)不同技術(shù)途徑的深入發(fā)展，在實(shí)際生產(chǎn)中得到更廣泛的應(yīng)用。

3.3 響應(yīng)高溫干旱的分子調(diào)控機(jī)制

隨著基因工程的深入，篩選和培育耐熱、耐旱新品種已從簡單的生理生化研究拓展到分子生物學(xué)領(lǐng)域當(dāng)中。目前可通過調(diào)節(jié)抗性蛋白表達(dá)、選擇耐性基因載體和遺傳改良等多種方法進(jìn)一步提高植物的抗逆性。中國科學(xué)院遺傳與發(fā)育生物學(xué)研究所薛勇彪研究組與程祝寬研究組合作，成功克隆了1個(gè)耐熱基因(Thermotolerant Growth Required 1)。該基因編碼細(xì)胞核定位的DEAD-box RNA解旋酶。作為pre-rRNA的分子伴侶保證了高溫下細(xì)胞分裂所需的rRNA有效加工，從而增強(qiáng)了水稻的耐熱能力[124]。該研究不僅闡明了水稻耐高溫的分子機(jī)制，而且為分子培育耐高溫水稻品種提供了基因資源。

隨著分子生物學(xué)和生物化學(xué)的不斷進(jìn)步和完善，利用現(xiàn)代生物技術(shù)，諸如生物信息學(xué)分析、蛋白互作分析、組學(xué)分析、基因芯片技術(shù)、高通量RNA測序(RNA-Seq)技術(shù)以及全基因組關(guān)聯(lián)分析(GWAS)技術(shù)等，對這些問題進(jìn)行深入地探索與分析，有望全面系統(tǒng)地理解水稻逆境相關(guān)轉(zhuǎn)錄因子的詳細(xì)調(diào)控機(jī)理，為水稻抗逆機(jī)理研究提供更多的理論依據(jù)。圖1總結(jié)了高溫干旱對水稻的影響及其調(diào)控模式。

4 展望

4.1 高溫和干旱影響水稻體內(nèi)生理代謝整體認(rèn)識

水稻對高溫和干旱脅迫的響應(yīng)是一個(gè)復(fù)雜的、但又是有序的生理生化過程，這一過程涉及水稻逆境下生長發(fā)育規(guī)律和養(yǎng)分吸收規(guī)律、根系形態(tài)建成和生理機(jī)制、酶學(xué)機(jī)制和激素機(jī)理、源庫協(xié)調(diào)機(jī)制和物質(zhì)運(yùn)轉(zhuǎn)分配機(jī)理、品質(zhì)形成特點(diǎn)與機(jī)理。要充分認(rèn)識其機(jī)理還要進(jìn)行大量艱苦的工作。今后建議從群體、個(gè)體、組織、器官、細(xì)胞和分子等不同水平上研究水稻對高溫和干旱及其復(fù)合脅迫的響應(yīng)機(jī)制，揭示水稻對單一及復(fù)合脅迫響應(yīng)和適應(yīng)性機(jī)理；將水稻的產(chǎn)量和品質(zhì)作為抗逆性的評定指標(biāo)。研究并應(yīng)用耕作、栽培、化控技術(shù)減輕高溫和干旱對作物的傷害，特別是減少高溫和干旱對作物產(chǎn)量與品質(zhì)的不利影響。突出作物生理調(diào)控作用，即利用作物本身或給作物創(chuàng)造環(huán)境發(fā)揮作物對逆境的適應(yīng)能力和抵抗能力。

4.2 多脅迫因子對水稻生長發(fā)育的交互影響

目前的研究多集中在高溫或干旱單因子對水稻的影響，但雙因子乃至高溫、干旱、高CO2濃度或高O3濃度等多因子耦合作用的研究甚少。未來氣候變化對水稻產(chǎn)量和品質(zhì)的最終影響取決于所有環(huán)境因子間的協(xié)同作用，多因子的共同作用才能代表大田條件下水稻生長發(fā)育的真實(shí)情景。因此，水稻生殖生長期不同氣候變化因子單獨(dú)和耦合影響有待加強(qiáng)研究。另外，充分利用各種研究手段，如大田控制試驗(yàn)、遮雨棚、土培池、玻璃溫室、人工氣候室等模擬水稻的生長環(huán)境，綜合分析溫度、水分等多種環(huán)境因子的作用，是揭示水稻對高溫、干旱等多重脅迫響應(yīng)的基礎(chǔ)。

4.3 水稻抗高溫和干旱的分子設(shè)計(jì)育種與基因組編輯

近年來，我國在水稻生物學(xué)、進(jìn)化與基因組學(xué)和激素生物學(xué)等領(lǐng)域表現(xiàn)尤為突出，并取得了許多突破性的研究成果[125, 126]，標(biāo)志著中國在該領(lǐng)域居于引領(lǐng)地位。如李家洋團(tuán)隊(duì)率先提出并踐行“分子設(shè)計(jì)育種”的理念，在系統(tǒng)研究作物產(chǎn)量、品質(zhì)、耐逆性、營養(yǎng)高效、抗病蟲等復(fù)雜農(nóng)藝性狀調(diào)控機(jī)理的基礎(chǔ)上，挖掘關(guān)鍵基因的有利變異，通過品種設(shè)計(jì)進(jìn)行多基因的配組優(yōu)化，實(shí)現(xiàn)復(fù)雜性狀的定向改良，達(dá)到綜合性狀優(yōu)異的目標(biāo)，為我國水稻分子設(shè)計(jì)育種與生產(chǎn)的跨越式發(fā)展奠定了開創(chuàng)性基礎(chǔ)[127]?；蚪M編輯是近年來生命科學(xué)領(lǐng)域的突破性技術(shù)，能夠精確改造生物基因組DNA，從而在不改變其目標(biāo)基因組整體穩(wěn)定性的基礎(chǔ)上直接對目的基因進(jìn)行分子設(shè)計(jì)改良，其終產(chǎn)品無任何外源DNA成分，具有廣闊的應(yīng)用前景。我國科學(xué)家在作物基因組編輯技術(shù)領(lǐng)域取得多項(xiàng)突破性進(jìn)展，使我國成為作物基因組編輯研究的國際領(lǐng)跑者[128, 129]。未來開展水稻高溫、干旱抗性的改良研究，需要首先鑒定和了解控制抵抗高溫、干旱脅迫的關(guān)鍵基因，解析其表達(dá)調(diào)控機(jī)制、作用機(jī)理、信號途徑和不同基因組合形成的調(diào)控網(wǎng)絡(luò)。然后，根據(jù)需要改良的耐熱、抗旱等農(nóng)藝性狀進(jìn)行針對性的設(shè)計(jì)，選擇最適宜的等位基因組，最終培育出抗高溫、干旱以及高溫干旱復(fù)合脅迫的水稻新品種，是水稻育種的發(fā)展方向。

圖1 高溫和干旱對水稻的影響及其調(diào)控模式

Fig. 1. Mode chart in the effect of heat and drought on rice and its regulation.

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Research Advances in the Effect of Heat and Drought on Rice and Its Mechanism

DUAN Hua*, TONG Hui, LIU Yanqing, XU Qingfen, MA Jun, WANG Chunmin

(,,,;*Corresponding author, E-mail: duanhua2004@gmail.com*Corresponding author, E-mail: duanhua2004@gmail.com)

Heat and drought are two major environmental stresses that affect rice growth, productivity, and grain quality. a comprehensive understanding of which is critical to evaluate the impacts of climate change on crop production. We review the independent and combined effects of heat and drought on rice growth, yield, and grain quality, demonstrate the possible mechanisms involved from multiple perspectives, such as photosynthesis, antioxidant system, endogenous hormones, activities of the key enzymes involved in sucrose-to-starch conversion, and molecular profiling, proposes reasonable strategies to mitigate the stress of environmental heterogeneity, and provide considerable suggestions for future study.

rice; heat; drought; combined heat and drought; grain yield; quality; physiological mechanism

Q945.78; S511.01

A

1001-7216(2019)03-0206-13

10.16819/j.1001-7216.2019.8106

2018-09-20；

2019-01-28。

國家自然科學(xué)基金資助項(xiàng)目(31601248)；天津市應(yīng)用基礎(chǔ)與前沿技術(shù)研究計(jì)劃資助項(xiàng)目(15JCQNJC14800)；天津市科技支撐重點(diǎn)項(xiàng)目(18YFZCNC01150)。