S3307對(duì)始花期和始粒期淹水綠豆光合作用及產(chǎn)量的影響

于 奇 馮乃杰 王詩(shī)雅 左官?gòu)?qiáng) 鄭殿峰,2,*

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S3307對(duì)始花期和始粒期淹水綠豆光合作用及產(chǎn)量的影響

于 奇1馮乃杰1王詩(shī)雅1左官?gòu)?qiáng)1鄭殿峰1,2,*

1黑龍江八一農(nóng)墾大學(xué)農(nóng)學(xué)院, 黑龍江大慶 163319;2黑龍江八一農(nóng)墾大學(xué)國(guó)家雜糧工程技術(shù)研究中心, 黑龍江大慶 163319

淹水脅迫是作物生長(zhǎng)發(fā)育過(guò)程中遭受的主要非生物脅迫之一, 探究提高綠豆耐淹性的機(jī)制對(duì)綠豆抗?jié)吃耘嗑哂兄匾饬x。本文在2017—2018年以耐淹性不同的綠豆品種綠豐2號(hào)和綠豐5號(hào)為試驗(yàn)材料, 采用盆栽方式探究了烯效唑(S3307)對(duì)淹水脅迫下綠豆葉片生理、光合作用及產(chǎn)量的影響。結(jié)果表明, 在不同生育時(shí)期淹水脅迫下, 綠豆葉片的葉綠素含量(SPAD)及光合特性參數(shù)均顯著下降, 丙二醛(MDA)含量顯著增加, 始花期(R1期)淹水脅迫下綠豆的減產(chǎn)率為24.70%~33.63%, 始粒期(R5期)減產(chǎn)率為18.07%~28.87%。2個(gè)綠豆品種均表現(xiàn)為R1期受淹水脅迫危害程度大于R5期, 綠豐2號(hào)耐淹性強(qiáng)于綠豐5號(hào)。噴施S3307后顯著提高淹水脅迫下綠豆葉片的SPAD、凈光合速率(n)、蒸騰速率(r)和氣孔導(dǎo)度(s), 并顯著降低了MDA含量。綠豆在R1期淹水脅迫下的緩解率為28.91%~52.34%, R5期緩解率為13.77%~27.36%。表明葉面噴施S3307可有效提高淹水脅迫下綠豆葉片的生理功能及光合能力, 進(jìn)而降低減產(chǎn)幅度, 但不同淹水時(shí)期和綠豆品種對(duì)S3307的調(diào)控響應(yīng)存在差異。

綠豆; 烯效唑(S3307); 淹水脅迫; 光合作用; 產(chǎn)量

根據(jù)聯(lián)合國(guó)食品與農(nóng)業(yè)組織統(tǒng)計(jì), 世界范圍內(nèi)的綠豆產(chǎn)量從1990年到2014年增加了4倍, 達(dá)到2140萬(wàn)噸, 其中亞洲約占85.4%[1]。綠豆是中國(guó)重要的豆科作物之一, 具有生育期短、播種范圍廣和適應(yīng)性強(qiáng)等特點(diǎn), 成為國(guó)際市場(chǎng)的高附加值產(chǎn)品[2-3]。同時(shí)綠豆通過(guò)根瘤菌的大氣固氮作用在恢復(fù)土壤肥力方面起到了關(guān)鍵作用, 有利于可持續(xù)農(nóng)業(yè)生產(chǎn)[4]。淹水脅迫是農(nóng)業(yè)生產(chǎn)中的主要非生物脅迫之一, 制約作物生長(zhǎng)、降低產(chǎn)量, 成為主要農(nóng)業(yè)制約因素[5-6]。據(jù)估計(jì), 全球范圍內(nèi)12%的作物區(qū)受淹水脅迫影響[7]。淹水脅迫造成了約40%~80%的作物產(chǎn)量損失, 面積超過(guò)1700萬(wàn)公頃[8-9]。中國(guó)綠豆產(chǎn)區(qū)主要在黃淮河流域及東北地區(qū), 該地區(qū)持續(xù)性強(qiáng)降雨大多發(fā)生在6月中旬至7月中旬, 正值綠豆生殖生長(zhǎng)期, 極易造成淹水危害[10-11]。

淹水脅迫影響植物生長(zhǎng), 降低葉片光合作用效率, 最終降低作物產(chǎn)量, 影響農(nóng)業(yè)生產(chǎn)[12]。凈光合速率(n)、蒸騰速率(r)、氣孔導(dǎo)度(s)和胞間CO2濃度(i)是植物在淹水脅迫下光合生理響應(yīng)的重要指標(biāo)[13]。淹水脅迫會(huì)影響北美鵝掌楸[14]、灰化苔草幼苗[15]和玉米[16]的正常生長(zhǎng), 降低葉綠素含量和光合效率, 并使細(xì)胞膜質(zhì)過(guò)氧化程度加深, 丙二醛(MDA)含量增加。研究表明, 小麥淹水超過(guò)5 d會(huì)使其乳熟時(shí)間延后并降低產(chǎn)量, 玉米苗期受淹水脅迫3~7 d后其產(chǎn)量下降58.8%~69.8%, 印度木豆受淹后會(huì)使單株產(chǎn)量下降27.4%~61.5%[17-19]。通過(guò)化控技術(shù)提高作物的光合效率、調(diào)控作物的生理代謝、增加作物產(chǎn)量[20]。烯效唑(S3307)是一種新型高效三唑類(lèi)植物生長(zhǎng)延緩劑, 具有前控后促的作用, 在抗倒伏和抗逆等方面均有明顯效果[21-22]。噴施S3307能緩解淹水脅迫下大豆葉片凈光合速率(n)、蒸騰速率(r)、氣孔導(dǎo)度(s)和胞間CO2濃度(i)的下降, 且在恢復(fù)正常水分管理后能促進(jìn)光合能力的快速恢復(fù)[23]。葉噴S3307或S3307浸種后可提高大豆和綠豆的光合速率, 促進(jìn)生育后期植物葉片的水分利用效率[24-26]。Yan等[27]、曾紅等[28]、楊文鈺等[29]和Gawad等[30]研究表明, 在大豆、玉米和小麥等作物上應(yīng)用S3307后均有顯著的增產(chǎn)效應(yīng)。然而關(guān)于淹水脅迫對(duì)植物影響的研究多集中于大豆、棉花、玉米等作物, 較少涉及綠豆, 且關(guān)于S3307緩解綠豆淹水傷害的相關(guān)研究也較少。本試驗(yàn)探究不同淹水時(shí)期對(duì)耐淹性不同的2個(gè)綠豆品種光合作用及產(chǎn)量的影響以及S3307在始花期(R1期)和始粒期(R5期)淹水脅迫過(guò)程中對(duì)綠豆生長(zhǎng)、光合特性參數(shù)及產(chǎn)量的影響, 以期為綠豆抗?jié)吃耘嗟於ɑA(chǔ)。

1 材料與方法

1.1 試驗(yàn)材料

供試綠豆品種為綠豐2號(hào)和綠豐5號(hào), 由國(guó)家雜糧工程技術(shù)研究中心種植資源室提供。供試50 mgL–1烯效唑母液由黑龍江八一農(nóng)墾大學(xué)農(nóng)學(xué)院化控研究室提供。

1.2 試驗(yàn)設(shè)計(jì)

本試驗(yàn)于2017—2018年在黑龍江八一農(nóng)墾大學(xué)國(guó)家雜糧工程技術(shù)研究中心盆栽基地, 采用盆栽方式, 自然光照培養(yǎng)綠豆綠豐2號(hào)和綠豐5號(hào)。樹(shù)脂花盆上口徑∶底徑∶高 = 32 cm∶23 cm∶31 cm, 下部帶有排水口, 底部有隔水層。試驗(yàn)用土按過(guò)篩栽培土∶腐殖土∶酒糟土= 6∶1∶1比例均勻混合組成, 每盆裝土17 kg, 施肥量為純N 0.125 g kg–1, P2O50.25 g kg–1, K2O 0.1 g kg–1。挑選大小一致飽滿的綠豆種子, 用10%次氯酸鈉消毒15 min, 蒸餾水反復(fù)沖洗干凈并晾干后用于播種。每盆等距播種15粒, 于真葉期定苗5株, 試驗(yàn)期間用自動(dòng)定時(shí)滴灌裝置澆水, 使每盆土壤水分一致并保持在田間最大持水量的80%左右, 定期除草并噴灑農(nóng)藥防治病蟲(chóng)害。采用完全隨機(jī)區(qū)組設(shè)計(jì), 培養(yǎng)至始花期(R1期)和始粒期(R5期)開(kāi)始處理, 每個(gè)品種設(shè)R1期對(duì)照(R1期噴施蒸餾水后正常水分管理; R1CK)、R1期淹水處理(R1期噴施蒸餾水后淹水脅迫處理, 淹水時(shí)間持續(xù)5 d; R1W)、R1期S3307+淹水處理(R1期噴施S3307后淹水脅迫處理, 淹水時(shí)間持續(xù)5 d; R1W+S)、R5期對(duì)照(R5期噴施蒸餾水后正常水分管理; R5CK)、R5期淹水處理(R5期噴施蒸餾水后淹水脅迫處理, 淹水時(shí)間持續(xù)5 d; R5W)、R5期S3307+淹水處理(R5期噴施S3307后淹水脅迫處理, 淹水時(shí)間持續(xù)5 d; R5W+S) 6個(gè)處理。在R1期和R5期噴施等量的蒸餾水或S3307于綠豆葉片的正反面, 直至綠豆葉尖滴水時(shí)停止噴施, S3307噴施濃度為50 mg L–1。于噴藥5 d 后同一時(shí)間點(diǎn)套盆進(jìn)行淹水處理(以水面高于土面3 cm為準(zhǔn)), 淹水時(shí)間持續(xù)5 d, 5 d后放水恢復(fù)正常水分管理直至收獲。分別于R1期(R1-0, 即葉面噴施蒸餾水或S3307)、R1期后5 d (R1-5, 即噴藥5 d后)、R1期后10 d (R1-10, 即淹水5 d后)、R1期后15 d (R1-15, 即淹水脅迫解除5 d后)及R5期(R5-0)、R5期后5 d (R5-5)、R5期后10 d (R5-10)、R5期后15 d (R5-15)測(cè)定各個(gè)處理的葉綠素相對(duì)含量(SPAD)、丙二醛(MDA)含量及光合特性參數(shù)等指標(biāo), 測(cè)定完成后隨機(jī)選取每品種每處理長(zhǎng)勢(shì)一致的4盆綠豆植株正常水分管理直至收獲, 用于后期測(cè)定產(chǎn)量。

1.3 測(cè)定項(xiàng)目與方法

1.3.1 葉綠素相對(duì)含量(SPAD)測(cè)定 選取每處理4株長(zhǎng)勢(shì)一致的植株, 采用SPAD-502型葉綠素儀(Japan)測(cè)定每株綠豆主莖頂部倒數(shù)第3片復(fù)葉中間小葉上、中、下部的SPAD值, 取平均值。

1.3.2 丙二醛(MDA)含量測(cè)定 取綠豆主莖頂部倒數(shù)第3片復(fù)葉, 液氮速凍30 min后置–40℃冰箱中保存,待樣品全部收集完畢后統(tǒng)一測(cè)定。取0.5 g混合后的新鮮葉片置預(yù)冷的研缽中, 分3次加入10 mL 50 mmol L–1預(yù)冷的磷酸緩沖液(pH 7.8)在冰浴上研磨成勻漿, 在4℃、9000 r min-1下離心25 min, 取上清液(即粗提液)保存于4℃中備用, 采用硫代巴比妥酸(TBA)法[31]測(cè)定丙二醛(MDA)含量。

1.3.3 葉片光合特性參數(shù) 在R1期和R5期選擇晴朗無(wú)風(fēng)的天氣, 于上午8:30–11:00, 隨機(jī)選取每個(gè)品種每個(gè)處理長(zhǎng)勢(shì)一致的4盆植株, 用CI-340光合儀(CID, USA)測(cè)定每盆中5株綠豆主莖頂部倒數(shù)第3片復(fù)葉中間小葉的凈光合速率(n)、蒸騰速率(r)、氣孔導(dǎo)度(s)和胞間CO2濃度(i), 取其平均值。測(cè)定時(shí)溫度為28~34℃, 大氣CO2濃度為430~530 μmol L-1, 相對(duì)濕度為70%~80%, 光合有效輻射為800~1600 μmol m-2s-1。

1.3.4 產(chǎn)量及產(chǎn)量構(gòu)成因素的測(cè)定 于收獲期對(duì)每品種每處理的4盆綠豆植株進(jìn)行室內(nèi)考種, 統(tǒng)計(jì)每盆中5株植株中各株的單株莢數(shù)、單莢粒數(shù)、百粒重和單株產(chǎn)量, 取其平均值。

1.4 數(shù)據(jù)處理

采用Microsoft Excel 2016進(jìn)行數(shù)據(jù)的錄入、整理及繪圖, 用SPSS 23.0對(duì)數(shù)據(jù)進(jìn)行方差分析。

2 結(jié)果與分析

2.1 S3307對(duì)不同時(shí)期淹水綠豆生理特性參數(shù)的影響

2.1.1 葉片SPAD值 由圖1可知, 在R1期和R5期淹水脅迫后, 綠豆葉片SPAD值整體呈下降趨勢(shì)。與CK相比, 綠豐2號(hào)和綠豐5號(hào)均表現(xiàn)為R1期淹水脅迫(W)處理的SPAD降低幅度高于R5期, 并且兩時(shí)期淹水脅迫下綠豐5號(hào)W處理的SPAD降低幅度高于綠豐2號(hào)。噴施S3307后均提高了淹水后及脅迫解除后綠豆葉片SPAD值。2個(gè)時(shí)期淹水后(R1-10和R5-10), 綠豆W處理葉片SPAD較CK均降低, W+S處理在R1期較W處理分別增加了3.49%和28.27%, R5期分別增加了11.25%和9.64%。脅迫解除后(R1-15和R5-15), 綠豆W處理的葉片SPAD較各自CK均顯著降低, 綠豐2號(hào)和綠豐5號(hào)W+S處理在R1期較W處理分別增加了10.06%和35.22%, R5期分別增加了5.45%和32.59%。

圖1 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片SPAD的影響

R1CK: 始花期對(duì)照; R1W: 始花期噴施蒸餾水后淹水脅迫處理; R1W+S: 始花期噴施S3307后淹水脅迫處理; R5CK: 始粒期對(duì)照; R5W: 始粒期噴施蒸餾水后淹水脅迫處理; R5W+S: 始粒期噴施S3307后淹水脅迫處理。R1-0: 始花期噴施蒸餾水或S3307; R1-5: 始花期后5 d (即噴藥5 d后); R1-10: 始花期后10 d (即淹水5 d后); R1-15: 始花期后15 d (即脅迫解除5 d后), R5-0: 始粒期噴施蒸餾水或S3307; R5-5: 始粒期后5 d (即噴藥5 d后); R5-10: 始粒期后10 d (即淹水5 d后); R5-15: 始粒期后15 d (即脅迫解除5 d后)。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

R1CK: the beginning bloom control; R1W: exposed to waterlogging stress after spraying water at the beginning bloom; R1W+S: exposed to waterlogging stress after spraying S3307 at the beginning bloom; R5CK: the beginning seed control; R5W: exposed to waterlogging stress after spraying water at the beginning seed; R5W+S: exposed to waterlogging stress after spraying S3307 at the beginning seed.R1-0: spayed water or S3307 at the beginning bloom; R1-5: 5 d after the beginning bloom (5 d after spayed water or S3307); R1-10: 10 d after the beginning bloom (5 d after waterlogging stress); R1-15: 15 d after the beginning bloom (recovered 5 d), R5-0: spayed water or S3307 at the beginning seed; R5-5: 5 d after the beginning seed (5 d after spayed water or S3307); R5-10: 10 d after the beginning seed (5 d after waterlogging stress); R5-15: 15 d after the beginning seed (recovered 5 d). Values of SPAD within the same day with different letters are significantly different at the 0.05 probalility level.

2.1.2 葉片丙二醛(MDA)含量 由圖2可知, 除綠豐2號(hào)在R1期淹水后W處理MDA含量較CK顯著降低了21.29%外, 其余淹水時(shí)期綠豆的W處理MDA含量較CK顯著增加。R1期淹水后(R1-10), 綠豐2號(hào)W+S處理較W處理增加了9.44%, 綠豐5號(hào)降低了8.81%; 脅迫解除后(R1-15), 綠豐2號(hào)和綠豐5號(hào)W+S處理較W處理分別顯著降低了7.49%和33.43%。R5期淹水后(R5-10), 綠豐2號(hào)和綠豐5號(hào)W+S處理較W處理分別顯著降低了9.72%和10.33%, 脅迫解除后(R5-15)分別顯著降低了13.17%和12.68%。

2.2 S3307對(duì)不同時(shí)期淹水綠豆葉片光合特性參數(shù)的影響

2.2.1 對(duì)葉片凈光合速率(n) 由圖3可知, 綠豆在淹水及緩解后其凈光合速率整體變化趨勢(shì)為CK>W+S處理>W處理。綠豐2號(hào)W處理的凈光合速率淹水后(R1-10和R5-10)在R1期和R5期較CK分別顯著降低52.21%和43.06%, W+S處理凈光合速率較W處理顯著增加74.43%和42.83%; 綠豐2號(hào)W處理凈光合速率緩解后(R1-15和R5-15)在R1期和R5期較CK分別降低18.17%和7.53%, W+S處理凈光合速率在R1期較W處理顯著增加26.09%, R5期降低0.29%且差異不顯著。綠豐5號(hào)在不同時(shí)期受淹及緩解后的凈光合速率變化趨勢(shì)與綠豐2號(hào)基本一致, 但其下降幅度大于綠豐2號(hào)。

2.2.2 對(duì)葉片蒸騰速率(r) 由圖4可知, 在R1期和R5期綠豐2號(hào)W處理在淹水后(R1-10和R5-10)葉片蒸騰速率均顯著低于CK, W+S處理葉片蒸騰速率均顯著高于W處理; 而在緩解后(R1-15和R5-15)各個(gè)處理變化趨勢(shì)不同, W處理均高于CK, 而W+S處理在R1期高于W處理, 在R5期低于W處理, 但均高于各自CK。綠豐5號(hào)在淹水及緩解后, 除R5期的W+S處理在淹水后(R5-10)略高于CK外, 其他處理變化趨勢(shì)均為CK>W+S處理>W處理。

2.2.3 對(duì)葉片氣孔導(dǎo)度(s) 由圖5可知, 綠豐2號(hào)在R1和R5期噴施S3307 5 d后(R1-5和R5-5)葉片氣孔導(dǎo)度較CK顯著增加; 綠豐2號(hào)W處理在淹水后(R1-10和R5-10)葉片氣孔導(dǎo)度均顯著低于CK, W+S處理葉片氣孔導(dǎo)度均顯著高于W處理; 在緩解后(R1-15和R5-15) W處理在R1期高于CK, 在R5期低于CK, 差異均不顯著, W+S處理葉片氣孔導(dǎo)度在兩時(shí)期均高于CK和W處理。綠豐5號(hào)除在R5期緩解后(R5-15)W+S處理葉片氣孔導(dǎo)度大于CK外, 在R1期和R5期淹水及緩解后的變化趨勢(shì)均為CK>W+S處理>W處理。

圖2 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片MDA含量的影響

縮寫(xiě)同圖1。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of MDA content within the same day with different letters are significantly different at the 0.05 probalility level.

圖3 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片凈光合速率的影響

縮寫(xiě)同圖1。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of the photosynthetic rate within the same day with different letters are significantly different at the 0.05 probalility level.

圖4 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片蒸騰速率的影響

縮寫(xiě)同圖1。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of the transpiration rate within the same day with different letters are significantly different at the 0.05 probalility level.

圖5 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片氣孔導(dǎo)度的影響

縮寫(xiě)同圖1。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of the stomatal conductance within the same day with different letters are significantly different at the 0.05 probalility level.

2.2.4 對(duì)葉片胞間CO2濃度(i) 由圖6可知, 綠豐2號(hào)和綠豐5號(hào)在R1和R5期噴施S3307 5 d后(R1-5和R5-5)W+S處理葉片胞間CO2濃度較CK均增加; 綠豐2號(hào)W處理的葉片胞間CO2濃度在淹水后(R1-10和R5-10)均顯著低于CK, W+S處理葉片胞間CO2濃度均高于W處理, 但差異不顯著; 綠豐2號(hào)在R1期緩解后(R1-15和R5-15)的變化趨勢(shì)為W+S處理>CK>W處理, R5期的變化趨勢(shì)為CK>W+S處理>W處理。綠豐5號(hào)在兩時(shí)期淹水后的變化趨勢(shì)均為W+S處理>CK>W處理;在兩時(shí)期緩解后的變化趨勢(shì)均為CK>W+S處理>W處理。

2.3 S3307對(duì)不同時(shí)期淹水綠豆籽粒產(chǎn)量的影響

由表1可知, R1期綠豐2號(hào)和綠豐5號(hào)R1W單株產(chǎn)量較R1CK分別顯著降低了24.70%和29.54%; R1W+S較R1W分別顯著增加了32.40%和28.91%。兩品種綠豆單株莢數(shù)表現(xiàn)為R1W+S>R1CK>R1W, 單莢粒數(shù)表現(xiàn)為R1CK>R1W+S>R1W。綠豐2號(hào)百粒重在處理之間差異不顯著, 綠豐5號(hào)百粒重表現(xiàn)為 R1WR1W+S>R1W, 除綠豐2.號(hào)R1W+S和R1CK差異不顯著外, 其余處理之間差異均顯著。R5期綠豐2號(hào)和綠豐5號(hào)R5W單株產(chǎn)量較R5CK分別降低21.03%和25.35%, R5W+S較R5W分別增加27.36%和20.86%。綠豐2號(hào)和綠豐5號(hào)單株莢數(shù)在處理之間差異均不顯著。綠豐2號(hào)單莢粒數(shù)R5W+S 和R5CK分別與R5W差異顯著, 綠豐5號(hào)單莢粒數(shù)在處理之間差異不顯著。兩品種綠豆百粒重表現(xiàn)為 R5CK>R5W+S>R5W, R5CK與R5W差異顯著。綠豐2號(hào)單株產(chǎn)量表現(xiàn)為R5W+S>R5CK>R1W, R5CK與R5W差異顯著, 與R5W+S差異不顯著, 綠豐5號(hào)單株產(chǎn)量表現(xiàn)為R5CK>R5W+S>R5W, 并且處理之間差異均顯著。

圖6 S3307對(duì)R1期和R5期淹水脅迫下綠豆葉片胞間CO2濃度的影響

縮寫(xiě)同圖1。同天內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of the intercellular CO2concentration within the same day with different letters are significantly different at the 0.05 probalility level.

由表2可知, R1期綠豐2號(hào)和綠豐5號(hào)淹水處理的單株產(chǎn)量較對(duì)照分別降低27.16%和33.63%, R5期分別降低18.07%和28.87%, R1期S3307處理較淹水處理分別增加34.08%和52.34%, R5期分別增加13.77%和21.18%。由表1~表3可知, 在淹水脅迫和S3307處理下, 2017—2018年綠豆產(chǎn)量及產(chǎn)量構(gòu)成因素的變化趨勢(shì)基本一致, 均表現(xiàn)為2個(gè)綠豆品種在R1期減產(chǎn)幅度高于R5期, 同時(shí)期下綠豐2號(hào)的減產(chǎn)幅度低于綠豐5號(hào)。R1期淹水脅迫影響了綠豆的單株莢數(shù)、單莢粒數(shù)和百粒重, R5期影響了綠豆單莢粒數(shù)和百粒重。噴施 S3307后使淹水脅迫下綠豆單株產(chǎn)量顯著增加。

表1 S3307對(duì)R1期和R5期淹水脅迫下綠豆產(chǎn)量及產(chǎn)量構(gòu)成因素的影響(2017年)

縮寫(xiě)同圖1。同欄內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of yield and yield components within the same column with different letters are significantly different at the 0.05 probalility level.

表2 S3307對(duì)R1期和R5期淹水綠豆產(chǎn)量及產(chǎn)量構(gòu)成因素的影響(2018年)

縮寫(xiě)同圖1。同欄內(nèi)標(biāo)以不同字母的值在= 0.05水平上差異顯著。

Abbreviations are the same as those given in Fig. 1. Values of yield and yield components within the same column with different letters are significantly different at the 0.05 probalility level.

表3 2017–2018年R1期和R5期淹水脅迫下綠豆單株產(chǎn)量的減產(chǎn)率及緩解率

R1: 始花期; R5: 始粒期; 減產(chǎn)率 = (W – CK)/CK×100%; 緩解率 = (W+ S3307 – W)/W×100%, 以每個(gè)品種每個(gè)處理單株產(chǎn)量計(jì)算。

R1: the beginning bloom; R5: the beginning seed; Yield reduction rate = (waterlogging treatment – control)/control×100%; Remission rate = (waterlogging+S3307 treatment – waterlogging treatment)/waterlogging treatment×100%. The calculation in laced on yield per plant per treatment of each variety.

3 討論

淹水脅迫會(huì)對(duì)植物的生長(zhǎng)及生理代謝造成嚴(yán)重影響, 隨著脅迫時(shí)間延長(zhǎng)或程度加深, 植物受害越嚴(yán)重。葉綠素的含量變化是植物對(duì)水分脅迫響應(yīng)較為敏感的生理指標(biāo)之一, 反映了植物葉片的光合性能及衰老程度[14]。丙二醛(MDA)作為膜質(zhì)過(guò)氧化指標(biāo), 其含量的高低代表了植物在逆境條件下受害程度的深淺[32]。綠豆在開(kāi)花期和鼓粒期維持較高的葉綠素含量及光合速率對(duì)籽粒產(chǎn)量形成具有重要意義[33]。本試驗(yàn)結(jié)果表明, R1和R5期淹水脅迫會(huì)影響綠豆的正常生長(zhǎng), 降低葉綠素含量, 使細(xì)胞膜質(zhì)過(guò)氧化程度加深, 丙二醛(MDA)含量增加。但不同綠豆品種對(duì)淹水脅迫的響應(yīng)有所不同。綠豐2號(hào)在R1期受淹及緩解后, 淹水處理的MDA含量較對(duì)照相比出現(xiàn)先降低后增加的變化趨勢(shì), 綠豐5號(hào)淹水處理較對(duì)照則呈現(xiàn)顯著上升趨勢(shì)。說(shuō)明在一定淹水脅迫時(shí)間內(nèi), 綠豐2號(hào)對(duì)淹水脅迫具有一定的適應(yīng)性, 這與曹旖旎等[34]研究結(jié)果相似。在逆境脅迫條件下應(yīng)用S3307后, 會(huì)緩解脅迫對(duì)植物造成的危害。有研究發(fā)現(xiàn)[35-36], 葉面噴施S3307后能夠提高鹽脅迫下大豆的葉綠素含量, 在R1期噴施S3307后可降低大豆葉片MDA含量, 緩解脅迫對(duì)大豆葉片細(xì)胞膜的傷害。本試驗(yàn)發(fā)現(xiàn), 淹水脅迫條件下, 噴施S3307增加了受淹綠豆葉片的葉綠素含量并降低MDA含量, 減輕淹水脅迫下綠豆葉片的細(xì)胞膜質(zhì)過(guò)氧化作用, 保證綠豆幼苗在淹水逆境下的生長(zhǎng)及生理代謝, 這也是S3307增強(qiáng)綠豆植株耐淹性的原因之一。

光合作用是作物生長(zhǎng)發(fā)育過(guò)程中重要的能量轉(zhuǎn)化代謝系統(tǒng), 光合能力的提高為籽粒干物質(zhì)的積累奠定了基礎(chǔ), 是作物產(chǎn)量形成的物質(zhì)基礎(chǔ)。耐淹植物通?？梢员３州^高的光合特性及光合速率, 這些光合氣體交換參數(shù)在一定程度上可以作為判定植物耐淹性的指標(biāo)[37]。研究表明, 噴施S3307后可提高蕓豆[38]和大豆[39]鼓粒后期不同冠層葉片的凈光合速率、蒸騰速率和氣孔導(dǎo)度。在干旱脅迫條件下, 經(jīng)S3307處理后百日草幼苗的凈光合速率、氣孔導(dǎo)度和蒸騰速率有所提高, 說(shuō)明S3307能夠提高脅迫下百日草幼苗的光合能力[40]。本研究發(fā)現(xiàn), 2個(gè)綠豆品種在淹水脅迫后光合氣體交換參數(shù)均有不同程度的降低, 噴施S3307后能顯著提高淹水脅迫下綠豆的凈光合速率、蒸騰速率和氣孔導(dǎo)度。但不同品種對(duì)淹水脅迫及S3307的調(diào)控響應(yīng)有所不同, 在脅迫解除后, 綠豐2號(hào)淹水處理和S3307處理的凈光合速率、蒸騰速率和氣孔導(dǎo)度均有所緩解, 并且蒸騰速率和氣孔導(dǎo)度均高于對(duì)照, 然而綠豐5號(hào)淹水處理在脅迫解除后凈光合速率仍低于對(duì)照。說(shuō)明綠豐2號(hào)在脅迫解除后具有較強(qiáng)的恢復(fù)能力, 耐淹性高于綠豐5號(hào)。這與Chaves等[41]和Antonio等[42]關(guān)于不同耐淹性品種對(duì)淹水脅迫響應(yīng)不同的研究結(jié)果基本一致。此外, 在R1期淹水脅迫下綠豆淹水處理的葉綠素含量、凈光合速率和氣孔導(dǎo)度的降低幅度高于R5期淹水。說(shuō)明綠豆在R1期受淹水脅迫危害程度高于R5期。

綠豆受淹后根系處于低氧環(huán)境, 使地上部光合參數(shù)值下降, 影響作物正常的光合作用, 進(jìn)而影響綠豆的干物質(zhì)積累, 對(duì)綠豆單株莢數(shù)、單莢粒數(shù)和百粒重等產(chǎn)量構(gòu)成因素造成嚴(yán)重影響, 最終降低綠豆的產(chǎn)量。本試驗(yàn)結(jié)果表明, R1期淹水脅迫使綠豆單株莢數(shù)和單莢粒數(shù)顯著降低, 產(chǎn)量顯著降低24.70%~33.63%; R5期淹水脅迫使綠豆單莢粒數(shù)顯著降低, 產(chǎn)量顯著降低18.07%~28.87%。這與前人研究結(jié)果[17-19]相似。有研究發(fā)現(xiàn)[43], 在大豆始花期和盛花期噴施S3307后會(huì)增加大豆的單株莢數(shù)和單株粒數(shù)并提高產(chǎn)量。本試驗(yàn)在噴施S3307后均能顯著提高R1和 R5期淹水脅迫下綠豆的單株產(chǎn)量, 同時(shí)S3307對(duì)不同時(shí)期淹水脅迫下綠豆的產(chǎn)量構(gòu)成因素調(diào)控不同, R1期噴施S3307后能提高脅迫下綠豆的單株莢數(shù)、單莢粒數(shù)和百粒重; R5期噴施S3307后能顯著提高脅迫下綠豆的單株莢數(shù)和單莢粒數(shù)。2年試驗(yàn)結(jié)果均顯示, R1期淹水脅迫下綠豆的減產(chǎn)率高于R5期, 再次說(shuō)明綠豆在R1期受淹水脅迫的危害程度大于R5期, 同時(shí) S3307 對(duì) R1 期淹水脅迫的緩解率高于R5期, 調(diào)控效應(yīng)較好。這與Linkemer等[44]研究結(jié)果基本一致。有關(guān)噴施S3307對(duì)淹水脅迫下綠豆葉片及根系抗氧化酶系統(tǒng)、碳氮代謝、活性氧代謝系統(tǒng)及相關(guān)耐淹性基因表達(dá)調(diào)控等影響有待進(jìn)一步研究。

4 結(jié)論

噴施烯效唑(S3307)減輕了淹水脅迫下綠豆葉片細(xì)胞膜質(zhì)過(guò)氧化作用, 提高了始花期(R1期)和始粒期(R5期)淹水脅迫下綠豆的葉綠素含量及光合性能, 進(jìn)而提高了單株產(chǎn)量。綠豆在R1期受淹水脅迫危害程度高于R5期, 綠豐2號(hào)的耐淹性強(qiáng)于綠豐5號(hào)。S3307對(duì)不同時(shí)期淹水脅迫下不同綠豆品種的調(diào)控效應(yīng)有所不同, 但可用于綠豆抗?jié)吃耘? 緩解淹水脅迫產(chǎn)生的危害。

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Effects of S3307 on the photosynthesis and yield of mung bean at R1 and R5 stages under waterlogging stress

YU Qi1, FENG Nai-Jie1, WANG Shi-Ya1, ZUO Guan-Qiang1, and ZHENG Dian-Feng1,2,*

1College of Agronomy, Heilongjiang Bayi Agricultural University, Daqing 163319, Heilongjiang, China;2National Coarse Cereals Engineering Research Center, Heilongjiang Bayi Agricultural University, Daqing 163319, Heilongjiang, China

Waterlogging stress is one of the main abiotic stresses during the growth and development of crops. It is of great significance to explore the mechanisms for improving the flood resistance and waterlogging resistance cultivation of mung bean under waterlogging stress. In this experiment, the effects of uniconazole (S3307) on physiology, photosynthesis and yield of mung bean leaves under waterlogging stress were investigated in pot culture with different flood resistance mung bean varieties Lufeng 2 and Lufeng 5 from 2017 to 2018. Under the stress of waterlogging at different growth stages, the chlorophyll content (SPAD) and photosynthetic characteristic parameters of mung beans leaves were significantly decreased, malondialdehyde (MDA) content was significantly increased. The yield reduction rate of mung beans under the stress of waterlogging was 24.70%–33.63% at the beginning bloom (R1 stage), and 18.07%–28.87% at the beginning seed (R5 stage). Both mung bean varieties showed that the effect of waterlogging stress at R1 stage was greater than that at R5 stage, and the flood resistance of Lufeng 2 was stronger than that of Lufeng 5. After S3307 sprayed, SPAD, net photosynthetic rate (n), transpiration rate (r), and stomatal conductance (s) in the leaves of mung beans could be significantly increased and MDA content could be significantly decreased. The remission rate of mung beans under waterlogging stress was 28.91%–52.34% at R1 stage, and 13.77%–27.36% at R5 stage. The results showed that S3307 sprayed on the leaf surface could effectively alleviate the physiological function and photosynthetic capacity of mung bean leaves under the stress of waterlogging, thus reduce the yield reduction, but there were differences in the regulatory response of mung bean varieties to S3307 in different waterlogging periods.

mung bean; uniconazole (S3307); waterlogging stress; photosynthesis; yield

2018-11-26;

2019-01-19;

2019-03-08.

10.3724/SP.J.1006.2019.84160

鄭殿峰, E-mail: byndzdf@126.com

E-mail: yrkiyrki@163.com

本研究由國(guó)家自然科學(xué)基金項(xiàng)目(31871576)和國(guó)家“十二五”科技支撐計(jì)劃項(xiàng)目(2014BAD07B05)資助。

This study was supported by the National Natural Science Foundation of China (31871576) and the National Key Technology Support Program of China for the 12th Five-Year Plan (2014BAD07B05).

URL: http://kns.cnki.net/kcms/detail/11.1809.S.20190307.1438.002.html