曹小闖, 李曉艷，朱練峰，張均華，禹盛苗，吳良?xì)g，金千瑜,*

1 中國(guó)水稻研究所，水稻生物學(xué)國(guó)家重點(diǎn)實(shí)驗(yàn)室, 杭州　310006

2 浙江大學(xué)環(huán)境與資源學(xué)院，教育部環(huán)境修復(fù)與生態(tài)健康重點(diǎn)實(shí)驗(yàn)室，杭州　310058

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水分管理調(diào)控水稻氮素利用研究進(jìn)展

曹小闖1, 李曉艷2，朱練峰1，張均華1，禹盛苗1，吳良?xì)g2，金千瑜1,*

1 中國(guó)水稻研究所，水稻生物學(xué)國(guó)家重點(diǎn)實(shí)驗(yàn)室, 杭州310006

2 浙江大學(xué)環(huán)境與資源學(xué)院，教育部環(huán)境修復(fù)與生態(tài)健康重點(diǎn)實(shí)驗(yàn)室，杭州310058

水分管理；根際溶氧量；氮代謝；光合氮素利用率；水稻

水稻是我國(guó)種植面積最大、灌溉用水量最多的作物, 全國(guó)種植面積達(dá)3000萬hm2[1]。作為一個(gè)需氧的有機(jī)體，水稻雖然有發(fā)達(dá)的通氣組織以保證根系的正常需氧活動(dòng)，但半水生性特點(diǎn)要求其生長(zhǎng)過程中需要大量的灌溉水，因此土壤供氧狀況仍是限制水稻根系生長(zhǎng)的關(guān)鍵因子[2]。長(zhǎng)期處于淹水狀態(tài)的稻田，空氣很難進(jìn)入到土壤中，水稻根系及微生物的呼吸作用消耗大量氧氣，導(dǎo)致土壤中的氧濃度極低。持續(xù)的低氧環(huán)境會(huì)引起稻田還原性有毒物質(zhì)的積累，根系細(xì)胞能量代謝失衡、細(xì)胞質(zhì)酸化及低氧的生理生化反應(yīng)，對(duì)水稻根系形態(tài)結(jié)構(gòu)、根系活力和生長(zhǎng)發(fā)育等都會(huì)產(chǎn)生嚴(yán)重影響[3-4]。研究發(fā)現(xiàn)，濕潤(rùn)灌溉、干濕交替、好氧栽培等水分管理能通過調(diào)控根際溶氧量促進(jìn)水稻生長(zhǎng)[5-8]，誘導(dǎo)水稻的生理特性及改善根系的吸收功能增強(qiáng)其抗旱性能，達(dá)到以不犧牲光合產(chǎn)物積累而大幅度提高氮素利用效率的目的[9-10]。增加根際溶氧量還能刺激土壤氮的礦化作用，且氮素形態(tài)顯著影響水稻的光合速率[11]。隨著我國(guó)氮肥用量的不斷增加，水稻的氮素利用率呈不斷降低趨勢(shì)；灌水過多、不合理的水分管理也顯著增加稻田氮素?fù)p失，因此尋找提高氮肥利用率的新途徑是目前急需解決的突出問題[12]。本文通過國(guó)內(nèi)外最新文獻(xiàn)綜述了稻田水分管理對(duì)水稻根際氮素形態(tài)、氮吸收利用、光合速率及其氮環(huán)境效應(yīng)的影響等方面研究進(jìn)展，以期為相關(guān)研究提供參考。

1　稻田水分管理對(duì)土壤和根際氮素形態(tài)的影響

增加根際溶氧量還能顯著提高根際氧化還原電位及根系活力，這不僅有利于根系對(duì)營(yíng)養(yǎng)物質(zhì)的吸收，還使根際土壤磷有效性增加、還原性有害物質(zhì)(Fe2+、Mn2+、H2S等)減少，為水稻生長(zhǎng)、氮吸收代謝提供有利的外部環(huán)境[19-20]。干濕交替及好氧栽培模式下土壤團(tuán)聚體破碎釋放出的閉蓄態(tài)有機(jī)物、微生物細(xì)胞溶解及滲透調(diào)節(jié)物質(zhì)的分泌等都將提高基質(zhì)的有效性及碳、氮礦化率[21]；還提高了有機(jī)質(zhì)有效性并能抵制微生物進(jìn)入半休眠狀態(tài)，這也增加微生物生物量碳、氮[18]。碳源是微生物生長(zhǎng)繁殖的限制因素，高量的溶解性有機(jī)碳提升了微生物生物量和酶活性，隨之土壤結(jié)構(gòu)得到改善，碳氮有效性將進(jìn)一步提高[22]。

2　稻田水分管理對(duì)水稻氮吸收利用的影響調(diào)控研究

2.1稻田水分管理對(duì)水稻氮吸收的影響

2.2稻田水分管理對(duì)水稻氮吸收的調(diào)控機(jī)制研究

根際是植物、土壤和微生物相互作用的特殊區(qū)域，是植物、土壤、微生物進(jìn)行物質(zhì)和能量交換及信息傳遞的門戶。根際缺氧或無氧環(huán)境下水稻根系細(xì)胞迅速?gòu)挠醒鹾粑袚Q為以乙醇發(fā)酵途徑為主的無氧呼吸，ATP合成量?jī)H為正常時(shí)3%—5%，能量嚴(yán)重虧缺，不利于水稻對(duì)氮的吸收[41]。短期缺氧脅迫時(shí)，水稻根系能通過增加通氣組織功能提高氧氣向根尖的轉(zhuǎn)運(yùn)效率, 減少吸收營(yíng)養(yǎng)物質(zhì)的根表面積, 但提高著生在主根上的側(cè)根數(shù)目以增加對(duì)營(yíng)養(yǎng)物質(zhì)的吸收[42]。通過調(diào)控水分管理增加根際溶氧量可顯著提高稻田氧化還原電位和硝態(tài)氮含量，這兩個(gè)因素都會(huì)影響水稻的根系形態(tài)建成和氮素吸收[43]。增加根際溶氧量對(duì)水稻氮吸收代謝有顯著的促進(jìn)作用，這可能是由于隨著根際溶氧量增加，根系呼吸作用加強(qiáng)，為氮吸收提供更多的能量；也可能受增氧條件下根系形態(tài)變化的影響，主要表現(xiàn)為最長(zhǎng)根增長(zhǎng)、孔隙度減小、側(cè)根和不定根數(shù)減少、外皮層厚壁組織細(xì)胞疏松，根系活力增強(qiáng)[6,44]。楊菲等[45]發(fā)現(xiàn)干濕交替能明顯提高根際氧濃度，增加水稻根系生物量、根毛數(shù)、根系表面積，并促進(jìn)水稻4級(jí)次生側(cè)根的發(fā)生。Jackson等[46]發(fā)現(xiàn)增氧可顯著提高水稻根長(zhǎng)及其生物量，且能顯著增加深層根的比例，說明增加根際氧濃度可促進(jìn)根系向下層生長(zhǎng)，提高土壤中水分和養(yǎng)分的有效性。富氧環(huán)境可提高水稻總根長(zhǎng)和下層根系所占的干物質(zhì)比例，且提高了根系自根基到根尖10—20cm、>20cm部分所占的生物量比例[47]。但根際氧濃度并非越高越好，水稻對(duì)根際溶氧量需求存在一定的闕值。研究發(fā)現(xiàn)水分含量達(dá)到田間持水量的70%—75%時(shí)，最有利于水稻根系生長(zhǎng)，在此基礎(chǔ)上增加或降低根際溶氧量，水稻的產(chǎn)量及相關(guān)氮代謝均會(huì)受到顯著影響[48]。總之，根系形態(tài)結(jié)構(gòu)的改變必將導(dǎo)致水稻對(duì)土壤氮素吸收利用能力的改變，根系生物量大、直徑大、空隙高度及不定根數(shù)量高的水稻品種在氮素吸收及產(chǎn)量上均有明顯的優(yōu)勢(shì)，這將為今后選育理想根系品種提供重要參考。

2.3稻田水分管理對(duì)水稻光合速率的調(diào)控研究

3　稻田水分管理對(duì)氮素環(huán)境效應(yīng)的影響

氮素利用率是決定氮肥增產(chǎn)效果的主要因素，受氮肥用量、施肥方法、水分管理等因素影響。長(zhǎng)期淹水種植使稻田氮礦化速率極慢，還田秸稈等有機(jī)質(zhì)分解不充分，導(dǎo)致土壤積累大量的酚醛木質(zhì)素化合物，降低氮素有效性[75]，這將阻礙水稻對(duì)土壤背景氮的吸收但對(duì)肥料氮吸收沒有影響，加重水稻生產(chǎn)對(duì)肥料的依賴。以南太湖流域淹灌稻田為例，淹灌稻田中氮揮發(fā)、徑流、淋失等途徑是造成氮肥利用率低的主要原因，三者氮損失量分別占施氮量的20%—40%、1.4%—6.3%及1.0%—1.9%[76]。稻田氮素?fù)p失受溫度、水分、氮源、pH 值、C/N、氧化還原電位等諸多因素影響，其中土壤持水量能夠很好的反映土壤溶解氧的狀況，因而通過調(diào)控稻田水分含量將影響稻田氮素?fù)p失。在稻田生態(tài)系統(tǒng)中，CH4是造成全球溫室效應(yīng)的主要?dú)怏w。Liesack等[77]指出，稻田產(chǎn)生的CH4大約有高達(dá)36%—80%在根際微氧區(qū)域被甲烷氧化細(xì)菌消耗, 因此根際生物氧化在控制稻田溫室氣體排放過程中具有非常重要的作用[78]。甲烷氧化菌是一種嚴(yán)格的好氧菌, 而產(chǎn)生CH4、N2O等溫室氣體的甲烷細(xì)菌、反硝化細(xì)菌大部分都屬于厭氧菌[79]。稻田持續(xù)淹水導(dǎo)致土壤缺氧，抑制甲烷氧化菌的活性。當(dāng)頻繁干濕交替及增加根際溶氧量后土壤的氧化還原電位增加，土壤微生物群落結(jié)構(gòu)發(fā)生變化，甲烷氧化菌等好氧微生物活性顯著增加，可抑制溫室氣體的產(chǎn)生。李香蘭等[80]發(fā)現(xiàn)，稻田持續(xù)淹水處理CH4排放量是烤田處理的12—20倍。因此通過適宜的水分管理改善根系氮素、溶氧量狀況，可能會(huì)降低甲烷細(xì)菌、反硝化細(xì)菌等厭氧微生物的數(shù)量和活性，減少CH4、N2O等溫室氣體的生成。

4　問題與展望

氮素生理利用率與光合作用緊密相關(guān)，水稻氮肥利用率的降低必然與植物光合氮素利用率的降低有關(guān)。張?jiān)茦虻萚83]發(fā)現(xiàn)，高氮效水稻品種的葉綠素含量較低，但光合速率下降卻不明顯；且氮高效水稻基因型在各生育期均具有理想的株型、葉面積指數(shù)、光合勢(shì)和群體生長(zhǎng)速率來協(xié)調(diào)其氮素高效吸收利用[84]。當(dāng)前有關(guān)水稻光合同化物的形成與利用、氮素在體內(nèi)運(yùn)輸和轉(zhuǎn)運(yùn)、籽粒充實(shí)等對(duì)于根際氧濃度的響應(yīng)，以及相關(guān)吸收和轉(zhuǎn)運(yùn)蛋白的表達(dá)調(diào)控機(jī)制等方面報(bào)道較少。因此，加強(qiáng)這些方面的研究能夠更好完善水稻氧營(yíng)養(yǎng)代謝機(jī)制，將為高氮效水稻品種選育提供更好的參考價(jià)值。

基于以上的分析，可以認(rèn)為稻田生態(tài)系統(tǒng)中水稻氮素利用需要從以下幾方面進(jìn)行深入研究：

(1) 合理的水分管理能促進(jìn)水稻氮素吸收，提高氮素利用率。因此，通過控制稻田水氮互作協(xié)調(diào)土壤水-肥-氣的平衡，根據(jù)影響氮肥利用率和光合作用的限制因素探索和開發(fā)新型水氮集成管理技術(shù)，將有利于構(gòu)建理想水稻根系、健康稻田環(huán)境，提高水稻產(chǎn)量和氮素利用效率。

(2) 氮水平、氮形態(tài)能通過調(diào)控葉片光合速率影響水稻的生長(zhǎng)發(fā)育及氮素利用率。加強(qiáng)水氮互作對(duì)水稻不同生育期光合速率的影響研究，明確光合氮素利用率與根際氮形態(tài)特征及溶氧量的響應(yīng)關(guān)系，這將為我們?nèi)骊U述水氮互作提高水稻氮肥利用率的光合生理本質(zhì)奠定基礎(chǔ)。

(3)當(dāng)前根際氧調(diào)控根系吸收特性的相關(guān)研究較少，進(jìn)一步揭示不同根際氧濃度下水稻根系功能與養(yǎng)分利用間的內(nèi)在聯(lián)系是根際氧研究的發(fā)展趨勢(shì)。水稻根系及植株的衰老主要受激素調(diào)控，明確根際氧與激素在延緩衰老中的互作機(jī)制，進(jìn)而闡述溶氧量調(diào)控根系生長(zhǎng)發(fā)育和生理功能的機(jī)理，提高根系機(jī)能并帶動(dòng)外界營(yíng)養(yǎng)的高利用率以及光合產(chǎn)物的轉(zhuǎn)運(yùn)效率，對(duì)于水稻后期產(chǎn)量潛能的發(fā)揮和超高產(chǎn)栽培都具有重要科學(xué)意義。

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Effects of water management on rice nitrogen utilization: a review

CAO Xiaochuang1, LI Xiaoyan2, ZHU Lianfeng1, ZHANG Junhua1, YU Shengmiao1, WU Lianghuan2, JIN Qianyu1,*

1StateKeyLaboratoryofRiceBiology,ChinaNationalRiceResearchInstitute,Hangzhou310006,China

2MinistryofEducationKeyLaboratoryofEnvironmentalRemediationandEcosystemHealth,CollegeofEnvironmentalandResourceSciences,ZhejiangUniversity,Hangzhou310058,China

water management; rhizosphere dissolved oxygen; nitrogen metabolism; photosynthetic nitrogen-use efficiency; rice

10.5846/stxb201411202298

浙江省自然科學(xué)基金資助項(xiàng)目(LQ15C130004)；國(guó)家重點(diǎn)基礎(chǔ)研究發(fā)展973計(jì)劃資助項(xiàng)目(2015CB150502)；國(guó)家自然科學(xué)基金資助項(xiàng)目(31172032, 30900880)

2014-11-20; 網(wǎng)絡(luò)出版日期:2015-10-30

Corresponding author.E-mail: jinqy@mail.hz.zj.cn

曹小闖, 李曉艷，朱練峰，張均華，禹盛苗，吳良?xì)g，金千瑜.水分管理調(diào)控水稻氮素利用研究進(jìn)展.生態(tài)學(xué)報(bào),2016,36(13):3882-3890.

Cao X C, Li X Y, Zhu L F, Zhang J H, Yu S M, Wu L H, Jin Q Y.Effects of water management on rice nitrogen utilization: a review.Acta Ecologica Sinica,2016,36(13):3882-3890.