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        不同生育時(shí)期遮陰對(duì)大豆葉片光合和葉綠素?zé)晒馓匦缘挠绊?/h1>
        2016-07-18 07:44:14楊文鈺蘇本營
        中國農(nóng)業(yè)科學(xué) 2016年11期
        關(guān)鍵詞:大豆玉米

        王 一,張 霞,楊文鈺,孫 歆,蘇本營,崔 亮

        (1四川農(nóng)業(yè)大學(xué)農(nóng)學(xué)院/農(nóng)業(yè)部西南作物生理生態(tài)與耕作重點(diǎn)實(shí)驗(yàn)室,成都 611130;2四川省大英縣農(nóng)業(yè)技術(shù)信息推廣站,四川遂寧 629000;3重慶市潼南中學(xué)校,重慶 404100)

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        不同生育時(shí)期遮陰對(duì)大豆葉片光合和葉綠素?zé)晒馓匦缘挠绊?/p>

        王一1,2,張霞1,3,楊文鈺1,孫歆1,蘇本營1,崔亮1

        (1四川農(nóng)業(yè)大學(xué)農(nóng)學(xué)院/農(nóng)業(yè)部西南作物生理生態(tài)與耕作重點(diǎn)實(shí)驗(yàn)室,成都 611130;2四川省大英縣農(nóng)業(yè)技術(shù)信息推廣站,四川遂寧 629000;3重慶市潼南中學(xué)校,重慶 404100)

        摘要:【目的】研究在不同生育時(shí)期遮陰處理下,不同大豆品種植株葉片葉面積、比葉重、葉綠素含量、光合參數(shù)、葉綠素?zé)晒鈪?shù)和產(chǎn)量構(gòu)成因素的變化規(guī)律,為中國南方丘陵和山區(qū)大面積推廣的玉米大豆間套作種植技術(shù)提供理論依據(jù)?!痉椒ā坎捎猛腹饴?0%的遮陽網(wǎng)對(duì)生育期和生育時(shí)期總天數(shù)不同的3個(gè)參試大豆品種桂夏2號(hào)、南豆12和C103進(jìn)行遮陰,設(shè)置4個(gè)處理,分別為不遮陰(CK)、出苗至盛花期遮陰(VER2)、出苗至鼓粒期遮陰(VER5)和盛花期至完熟期遮陰(R2R8),測(cè)定葉面積、比葉重、葉綠素含量、光合參數(shù)、熒光參數(shù)和產(chǎn)量構(gòu)成因素。【結(jié)果】VER2處理下大豆品種葉面積、葉綠素(a+b)含量和表觀量子效率分別比對(duì)照高15.50%、12.95% 和74.13%,比葉重、光補(bǔ)償點(diǎn)和最大光合速率分別比對(duì)照低15.78%、26.16%和26.52%,R2R8處理下大豆品種葉面積、葉綠素(a+b)和表觀量子效率平均分別比對(duì)照高0.3%、10.53%和28.07%,比葉重、光補(bǔ)償點(diǎn)和最大光合速率平均分別比對(duì)照低10.15%、20.34%和12.13%;盛花期復(fù)光PSⅡ最大量子產(chǎn)量平均比對(duì)照低3.01%,非光化學(xué)熒光淬滅系數(shù)平均比對(duì)照高26.80%,鼓粒期復(fù)光PSⅡ最大量子產(chǎn)量平均比對(duì)照低8.47%,非光化學(xué)熒光淬滅系數(shù)平均比對(duì)照高40.79%;VER2、VER5和R2R8處理下,桂夏2號(hào)單株粒重分別比對(duì)照低40.84%、48.67%和59.16%,百粒重分別比對(duì)照低23.69%、39.31%和26.39%,南豆12單株粒重分別比對(duì)照低46.67%、54.16%和21.19%,百粒重分別比對(duì)照低3.91%、19.93%和26.14%,C103單株粒重分別比對(duì)照低69.8%、74.85%和73.89%,百粒重分別比對(duì)照低68.8%、69.55%和71.64%?!窘Y(jié)論】出苗至盛花期遮陰對(duì)參試大豆品種葉片光合及葉綠素?zé)晒馓匦缘挠绊懘笥谑⒒ㄆ谥镣晔炱谡陉帲蠖怪仓暾陉幒笤谑⒒ㄆ趶?fù)光,葉片光合能力有一定程度的恢復(fù),在鼓粒期復(fù)光,則表現(xiàn)為受到強(qiáng)光脅迫,因此,從減小遮陰對(duì)大豆葉片光合作用不良影響的角度考慮,在中國南方丘陵和山區(qū)玉米大豆間作優(yōu)于玉米大豆套作,在玉米大豆套作模式下選擇品種、播期及種植技術(shù)時(shí),應(yīng)確保大豆在盛花期前恢復(fù)光照,避免遮陰超過大豆鼓粒期。前期遮陰和后期遮陰對(duì)大豆產(chǎn)量的影響大小因大豆品種而異,但遮陰時(shí)間越長對(duì)大豆產(chǎn)量構(gòu)成影響越大。

        關(guān)鍵詞:大豆;玉米;生育時(shí)期;遮陰;光合;熒光特性

        聯(lián)系方式:王一,Tel:15908301340;E-mail:wy625265289@126.com。通信作者楊文鈺,Tel:0835-2882004;E-mail:wenyu.yang@263.net。通信作者孫歆,Tel:13658091923;E-mail:sunxin529@163.com

        0 引言

        【研究意義】大豆間套作栽培能提高復(fù)種指數(shù),增加大豆總產(chǎn)量,有效緩解大豆供需矛盾[1]。但高位作物玉米對(duì)低位作物大豆會(huì)產(chǎn)生遮陰抑制效果,降低其葉片光合同化能力[2],導(dǎo)致減產(chǎn)。間套作大豆減產(chǎn)幅度與遮陰程度[3]、遮陰時(shí)間長短[2]和遮陰生育時(shí)期[4]等因素相關(guān),在中國南方丘陵和山區(qū)大面積推廣的玉米大豆間套作種植模式下,套作大豆受遮陰的生育時(shí)期自出苗開始,至初花期或盛花期結(jié)束[5-6],間作大豆受到遮陰程度最大的生育時(shí)期自初花期或盛花期開始,至完熟期結(jié)束[7-8],且因玉米和大豆品種不同,大豆受遮陰的生育時(shí)期不盡相同。因此,研究不同生育時(shí)期遮陰對(duì)大豆光合特性的影響,可為大豆間套作種植提供理論依據(jù)?!厩叭搜芯窟M(jìn)展】王竹等[3]研究表明,玉米大豆套作,遮陰導(dǎo)致大豆葉面積指數(shù)增加,比葉重減小,且大豆葉片光合能力隨光照強(qiáng)度減小而減?。凰纹G霞等[9]在苗期對(duì)大豆遮陰提高了大豆幼苗葉片葉綠素含量、PSⅡ有效量子產(chǎn)量,降低了葉綠素a與葉綠素b比值;李植等[10]研究結(jié)果表明大豆玉米間作模式下,大豆葉片葉綠素含量、表觀量子效率和 CO2補(bǔ)償點(diǎn)上升,光補(bǔ)償點(diǎn)和光飽和點(diǎn)下降?!颈狙芯壳腥朦c(diǎn)】已有研究多集中于單一生育時(shí)期遮陰[9-11],但間套作大豆受遮陰生育時(shí)期不同,不同生育時(shí)期遮陰對(duì)大豆光合特性影響的報(bào)道較少,且不同生育時(shí)期遮陰對(duì)大豆光合特性的影響差異尚不清楚。【擬解決的關(guān)鍵問題】本研究選用3個(gè)生育時(shí)期不同的大豆品種,通過研究不同生育時(shí)期遮陰處理下,不同大豆品種葉片光合特性表現(xiàn)的共性規(guī)律,明確不同生育時(shí)期遮陰對(duì)大豆葉片光合及葉綠素?zé)晒馓匦缘挠绊懖町?,為完善大豆間套作栽培技術(shù)提供理論依據(jù)。

        1 材料與方法

        1.1 供試材料

        試驗(yàn)選取桂夏2號(hào)、南豆12和C103為供試大豆品種。桂夏2號(hào)2004年通過廣西省品種審定委員會(huì)審定,植株莖稈粗壯,株型收斂,是適宜南方間套作種植的早熟、高產(chǎn)大豆品種;南豆12于2008年通過四川省品種審定委員會(huì)審定,連續(xù)6年被國家農(nóng)業(yè)部確定為南方地區(qū)主導(dǎo)品種,是中國南方地區(qū)推廣面積最大的大豆品種,在四川、重慶占大豆播種面積50%以上;C103是由農(nóng)業(yè)部西南作物生理生態(tài)與耕作重點(diǎn)實(shí)驗(yàn)室提供的四川省南充市地方品種,該品種晚熟高產(chǎn),耐陰抗倒,適宜間套作種植。各品種生育時(shí)期見表1。

        表1 供試大豆品種的生育時(shí)期(月/日)和生育期Table 1 Growth stages (month/day) and maturity (d) of experimental cultivars

        1.2 試驗(yàn)方法

        試驗(yàn)于2012年6—11月在四川農(nóng)業(yè)大學(xué)教學(xué)科研園區(qū)進(jìn)行。為準(zhǔn)確控制遮陰生育時(shí)期和光照環(huán)境等處理?xiàng)l件,采用盆栽方式,3個(gè)品種各栽種60盆,其中,單個(gè)品種的一個(gè)遮陰處理12盆,單個(gè)品種的對(duì)照24盆,每盆栽3株;供試紫色土養(yǎng)分含量為有機(jī)質(zhì)8.96 g·kg-1、全氮1.21 g·kg-1、全磷0.61 g·kg-1、全鉀11.44 g·kg-1、速效氮62.35 mg·kg-1、速效磷24.34 mg·kg-1和速效鉀65.72 mg·kg-1,pH為6.55。

        試驗(yàn)為雙因素隨機(jī)區(qū)組設(shè)計(jì),因素A為遮陰,因素B為大豆品種。試驗(yàn)參照FEHR等[12]的生育時(shí)期劃分法,設(shè)置 50%透光率[3]的遮陽網(wǎng)對(duì)不同品種大豆不同生育時(shí)期遮陰。處理與對(duì)照隨機(jī)排列。處理設(shè)置為不遮陰(CK)、出苗至盛花期遮陰(shaded from emergence to full-bloom,VER2)、出苗至始粒期遮陰(shaded from emergence to pod-fill,VER5)、盛花期至完熟期遮陰(shaded from full-bloom to full-ripe,R2R8)。

        1.3 測(cè)定項(xiàng)目與方法

        1.3.1 測(cè)定時(shí)期與部位 前期遮陰處理(VER2和VER5)在遮陰期間的最后一天(改變光強(qiáng)前)和恢復(fù)光照后第15天(改變光強(qiáng)后)測(cè)定或取樣;后期遮陰處理(R2R8)在自然光照條件下的最后一天(改變光強(qiáng)前)和遮陰后第15天(改變光強(qiáng)后)測(cè)定或取樣,對(duì)照(CK)與處理同步測(cè)定或取樣,3次重復(fù),每重復(fù)3株,取樣部位為植株倒3葉[13-14]。

        1.3.2 葉面積測(cè)定 掃描儀掃描葉片后用軟件Image J 1.44p計(jì)算葉面積。

        1.3.3 比葉重測(cè)定 采用打孔稱重法測(cè)定[15]。

        1.3.4 光合色素測(cè)定 參照舒展等[16]的方法,將葉片切成長約5 mm,寬約2 mm的細(xì)絲,用80%丙酮溶液浸提12 h。將浸提液在663 nm和645 nm波長下比色,所得光密度(OD)值代入公式計(jì)算溶液葉綠素a、葉綠素b和葉綠素(a+b)含量,公式如下:

        1.3.5 光合參數(shù)測(cè)定 選擇晴朗無風(fēng)天氣測(cè)定光合參數(shù),采用美國Li-COR公司生產(chǎn)的Li-6400光合測(cè)定系統(tǒng),選擇紅藍(lán)光源葉室,設(shè)置開放式氣路,CO2濃度為380 μmol·L-1,分別于光量子密度(photosynthetically available radiation,PAR)為0、50、100、200、400、600、800、1 000、1 200、1 400、1 600、1 800和2 000 μmol·m-2·s-1時(shí)測(cè)定葉片凈光合速率并繪制曲線,同時(shí),利用Li-COR公司研發(fā)的Photosyn Assistant軟件對(duì)光響應(yīng)曲線進(jìn)行擬合,計(jì)算并得出響應(yīng)光合參數(shù):光飽和點(diǎn)(light saturation point,LSP)、光補(bǔ)償點(diǎn)(light compensation point,LCP)、表觀量子效率(apparent quanta efficiency,AQE)、暗呼吸速率(dark respiration rate,Rd)和最大光合速率(maximum photosynthetic rate,Pnmax)。

        1.3.6 葉綠素?zé)晒鈪?shù)測(cè)定 用Mini-PAM(Walz,德國)便攜式脈沖調(diào)制式葉綠素?zé)晒鈨x測(cè)定暗適應(yīng)最大熒光(maximalfluorescence,F(xiàn)m)、暗適應(yīng)初始熒光(minimalfluorescence,F(xiàn)o),光下最大熒光(maximalfluorescence under light,F(xiàn)m′)和光下初始熒光(minimalfluorescence under light,F(xiàn)o′),根據(jù)公式計(jì)算出PSⅡ最大量子產(chǎn)量(maximal quantum yield of PSⅡ,F(xiàn)v/Fm)、PSⅡ?qū)嶋H量子產(chǎn)量(actual photochemical efficiency of PSⅡ,Yield)、光化學(xué)淬滅系數(shù)(photochemical quenching,qP)、非光化學(xué)淬滅系數(shù)(non-photochemical quenching,NPQ)和光合電子傳遞相對(duì)速率(relative photosynthetic electron transfer rate,rETR)。公式如下:Fv/Fm=(Fm-Fo)/Fm qP=(Fm′-F)/(Fm′-Fo);Yield=(Fm′-F)/Fm′;NPQ=(Fm-Fm′)/Fm′;rETR=Yield*PAR*0.5*0.84。

        2 結(jié)果

        表2 遮陰對(duì)大豆倒三葉葉面積和比葉重的影響Table 2 Effect of shading on leaf area and specific leaf weight of soybean leaves

        2.1 不同遮陰處理大豆葉片的葉面積和比葉重

        表2表明,VER2和VER5遮陰期間,參試大豆品種葉面積均高于對(duì)照,比葉重均低于對(duì)照,VER5受遮陰影響程度大于VER2,桂夏2號(hào)VER2和VER5葉面積分別比對(duì)照高 15.28%和 28.26%,比葉重分別比對(duì)照低15.77%和19.56%,南豆12 VER2和VER5葉面積分別比對(duì)照高 15.84%和 28.25%,比葉重分別比對(duì)照低15.78%和19.55%,C103 VER2和VER5葉面積分別比對(duì)照高 15.28%和 28.25%,比葉重分別比對(duì)照低15.80%和19.55%;復(fù)光后,VER2比葉重較對(duì)照的差異比在遮陰期間小,但VER5比葉重較對(duì)照的差異比在遮陰期間大,桂夏2號(hào)VER2和VER5比葉重分別比對(duì)照低14.06%和24.94%,南豆12 VER2和VER5比葉重分別比對(duì)照低14.05%和24.93%,C103 VER2和 VER5比葉重分別比對(duì)照低 14.05%和24.94%;各大豆品種R2R8遮陰后比葉重顯著低于對(duì)照,桂夏2號(hào)分別比對(duì)照低10.16%,南豆12分別比對(duì)照低10.15%,C103分別比對(duì)照低10.14%。

        2.2 遮陰對(duì)大豆葉片葉綠素及其組分含量的影響

        不同遮陰處理對(duì)葉綠素及其組分的含量的影響程度不同(表3),VER2和VER5遮陰期間,參試大豆品種葉綠素(a+b)和葉綠素b含量高于對(duì)照,葉綠素a/b比值低于對(duì)照,桂夏2號(hào)VER2和VER5葉綠素(a+b)含量分別比對(duì)照高2.25%和35.97%,葉綠素b含量分別比對(duì)照高29.07%和91.35%,葉綠素a/b比值分別比對(duì)照低 24.12%和 34.81%,南豆 12 VER2和VER5葉綠素(a+b)含量分別比對(duì)照高 32.21%和1.34%,葉綠素b含量分別比對(duì)照高63.01%和19.81%,葉綠素a/b比值分別比對(duì)照低24.64%和22.86%,C103 VER2和VER5葉綠素(a+b)含量分別比對(duì)照高3.13% 和 3.78%,葉綠素 b含量分別比對(duì)照高 41.88%和39.64%,葉綠素 a/b比值分別比對(duì)照低 33.81%和30.69%;復(fù)光后,各大豆品種 VER2葉綠素(a+b)和葉綠素a/b比值低于對(duì)照,桂夏2號(hào)分別比對(duì)照低15.41%和 4.29%,南豆 12分別比對(duì)照低 4.79%和17.37%,C103分別比對(duì)照低4.8%和17.91%,同時(shí),葉綠素(a+b)含量高于對(duì)照,葉綠素a/b比值低于對(duì)照,桂夏2號(hào)葉綠素(a+b)含量比對(duì)照高36.51%,葉綠素a/b比值比對(duì)照低34.15%,南豆12葉綠素(a+b)含量比對(duì)照高 1.38%,葉綠素 a/b比值比對(duì)照低23.33%,C103葉綠素(a+b)含量比對(duì)照高3.88%,葉綠素a/b比值比對(duì)照低30.96%;R2R8遮陰15d后,各大豆品種葉綠素(a+b)和葉綠素b含量均高于對(duì)照,葉綠素a/b比值均低于對(duì)照,桂夏2號(hào)葉綠素(a+b)和葉綠素b含量分別比對(duì)照高19.33%和46.78%,葉綠素a/b比值比對(duì)照低25.08%,南豆12葉綠素(a+b)和葉綠素b含量分別比對(duì)照高6.58%和44.17%,葉綠素a/b比值比對(duì)照低33.53%,C103葉綠素(a+b)和葉綠素b含量分別比對(duì)照高6.66%和42.28%,葉綠素a/b比值比對(duì)照低33.13%。

        表3 遮陰對(duì)大豆葉片葉綠素含量的影響Table 3 Effect of shading on chlorophyll content in soybean leaves

        2.3 不同遮陰處理下大豆葉片的光合參數(shù)

        參試大豆品種不同遮陰處理的光合參數(shù)存在顯著差異(表4),遮陰期間,VER2處理下各大豆品種光飽和點(diǎn)、光補(bǔ)償點(diǎn)和最大光合速率均顯著低于對(duì)照,桂夏2號(hào)分別比對(duì)照低13.23%、34.5%和17.56%,南豆12分別比對(duì)照低34.09%、15.02%和33.46%,C103分別比對(duì)照低7.98%、14.36%和28.07%,同時(shí)VER2表觀量子效率均顯著高于對(duì)照,桂夏 2號(hào)比對(duì)照高39.34%,南豆 12比對(duì)照高 87.8%,C103比對(duì)照高112.2%;復(fù)光后,VER2光飽和點(diǎn)、光補(bǔ)償點(diǎn)、最大光合速率和表觀量子效率顯著低于對(duì)照,桂夏2號(hào)分別比對(duì)照低19.31%、11.52%、16.42%和26.67%,南豆12分別比對(duì)照低14.52%、12.87%、16.82%和8%,C103分別比對(duì)照低 21.03%、20.27%、34.99%和29.41%,說明前期遮陰處理遮陰后復(fù)光各大豆品種對(duì)光量子利用率卻明顯降低;R2R8遮陰15d后光飽和點(diǎn)、光補(bǔ)償點(diǎn)和最大光合速率均低于對(duì)照,桂夏2號(hào)分別比對(duì)照低17.91%、23.3%和8.39%,南豆12分別比對(duì)照低13.87%、15.9%和9.04%,C103分別比對(duì)照低18.94%、22.49%和18.36%,同時(shí)R2R8表觀量子效率高于對(duì)照,桂夏2號(hào)比對(duì)照高26.67%,南豆12比對(duì)照高24%,C103比對(duì)照高35.29%,說明后期遮陰處理大豆植株由自然光照環(huán)境轉(zhuǎn)入遮陰環(huán)境后對(duì)光量子利用率顯著提高了。

        表4 遮陰對(duì)大豆葉片主要光合參數(shù)的影響Table 4 Effect of shading on the photosynthetic parameters of soybean leaves

        2.4 不同遮陰處理對(duì)大豆熒光參數(shù)的影響

        熒光參數(shù)是反映遮陰對(duì)大豆葉片光合作用過程影響的重要指標(biāo)(表5),VER2和VER5遮陰期間,參試大豆品種 PSⅡ最大量子產(chǎn)量和非光化學(xué)熒光淬滅系數(shù)均低于對(duì)照,PSⅡ?qū)嶋H量子產(chǎn)量、光化學(xué)淬滅系數(shù)和光合電子傳遞相對(duì)速率均高于對(duì)照,桂夏2號(hào)PSⅡ最大量子產(chǎn)量和非光化學(xué)熒光淬滅系數(shù)在VER2處理下分別比對(duì)照低3.9%和39.66%,在VER5處理下分別比對(duì)照低5.95%和22.73%,PSⅡ?qū)嶋H量子產(chǎn)量、光化學(xué)猝滅系數(shù)和光合電子傳遞相對(duì)速率在VER2處理下分別比對(duì)照高11.67%、8.51%和8.68%,在VER5處理下分別比對(duì)照高12.9%、22.78%和18.03%,南豆12 PSⅡ最大量子產(chǎn)量和非光化學(xué)熒光淬滅系數(shù)在VER2處理下分別比對(duì)照低3.85%和37.78%,在VER5處理下分別比對(duì)照低4.82%和58.06%,PSⅡ?qū)嶋H量子產(chǎn)量、光化學(xué)淬滅系數(shù)和光合電子傳遞相對(duì)速率在VER2處理下分別比對(duì)照高11.86%、11.63%和11.5%,在VER5處理下分別比對(duì)照高7.81%、4.49%和9.32%, C103 PSⅡ最大量子產(chǎn)量和非光化學(xué)熒光淬滅系數(shù)在VER2處理下分別比對(duì)照低5.33%和41.38%,在VER5處理下分別比對(duì)照低7.32%和29.31%,PSⅡ?qū)嶋H量子產(chǎn)量、光化學(xué)淬滅系數(shù)和光合電子傳遞相對(duì)速率在VER2處理下分別比對(duì)照高38.64%、36.99%和39.6%,在VER5處理下分別比對(duì)照高1.54%、11.11%和1.8%;復(fù)光后,VER2 PSⅡ最大量子產(chǎn)量低于對(duì)照,桂夏 2號(hào)比對(duì)照低3.61%,南豆12比對(duì)照低2.41%,C103比對(duì)照低 3.57%,非光化學(xué)熒光淬滅系數(shù)高于對(duì)照,桂夏2號(hào)比對(duì)照高21.88%,南豆12比對(duì)照高56%,C103比對(duì)照高12.5%,R2R8遮陰后PSⅡ最大量子產(chǎn)量低于對(duì)照,桂夏2號(hào)比對(duì)照低4.82%,南豆12比對(duì)照低6.02%,C103比對(duì)照低3.57%,光化學(xué)淬滅系數(shù)高于對(duì)照,桂夏2號(hào)比對(duì)照高8.51%,南豆12比對(duì)照高8.51%,C103比對(duì)照高9.57%。

        2.5 不同遮陰處理下大豆產(chǎn)量的差異

        不同遮陰處理對(duì)不同大豆品種產(chǎn)量構(gòu)成因素的影響不同(表6),VER2、VER5和R2R8處理下,桂夏 2號(hào)單株粒數(shù)分別比對(duì)照低 22.46%、30.23%和32.66%,單株粒重分別比對(duì)照低 40.84%、48.67%和 59.16%,百粒重分別比對(duì)照低 23.69%、39.31%和26.39%,南豆 12單株粒數(shù)分別比對(duì)照低 25.33%、52.29%和 5.01%,單株粒重分別比對(duì)照低 46.67%、54.16%和21.19%,百粒重分別比對(duì)照低3.91%、19.93% 和 26.14%,C103單株粒數(shù)分別比對(duì)照低 3.22%、17.34%和 7.92%,單株粒重分別比對(duì)照低 69.8%、74.85%和 73.89%,百粒重分別比對(duì)照低 68.8%、69.55%和71.64%,說明前期遮陰和后期遮陰對(duì)大豆產(chǎn)量的影響大小因大豆品種而異,但遮陰時(shí)間越長對(duì)大豆產(chǎn)量構(gòu)成影響越大。

        表5 遮陰對(duì)大豆葉片主要熒光參數(shù)的影響Table 5 Effect of shading on chlorophyll fluorescence parameters of soybean leaves

        表6 遮陰對(duì)大豆產(chǎn)量性狀的影響Table 6 Effect of shading on yield characters of soybean

        3 討論

        3.1 遮陰對(duì)大豆葉片光合特性的影響

        遮陰影響作物光合作用[17]、干物質(zhì)積累分配[18],最終影響產(chǎn)量[19]。董鉆[2]研究表明,不同大豆品種光合作用受遮陰的影響程度不同,本試驗(yàn)選用3個(gè)生育時(shí)期長短不同的大豆品種,按生育時(shí)期進(jìn)行遮陰,3個(gè)品種不同處理間變化規(guī)律基本一致,試驗(yàn)的結(jié)果更具普適性。研究表明,葉片是大豆植株主要的光合器官,其形態(tài)生理性狀能反映大豆植株受遮陰影響時(shí)的光合特性[3,9-10]。PAUSCH等[20]研究表明,遮陰提高大豆倒3葉葉面積;GHASSEMI-GOLEZANI等[21]研究結(jié)果表明,遮陰降低大豆功能葉單葉干重;本試驗(yàn)結(jié)果表明,參試大豆品種前期遮陰(VER2和VER5)葉面積增加,比葉重減少,且遮陰時(shí)間越長,影響越大,本試驗(yàn)研究還發(fā)現(xiàn)參試大豆品種后期遮陰(R2R8)對(duì)葉面積和比葉重的影響程度小于前期遮陰。XU等[22]研究表明,小麥花期受遮陰影響,旗葉葉綠素(a+b)含量增加,葉綠素 a/b比值減少,本試驗(yàn)研究結(jié)果與XU的結(jié)果一致,遮陰導(dǎo)致參試大豆品種葉片葉綠素(a+b)和葉綠素b的含量增加,葉綠素a/b比值減少,但葉綠素a含量變化因參試大豆品種不同而不同。馬德華等[23]研究發(fā)現(xiàn),遮陰時(shí)間過長會(huì)導(dǎo)致黃瓜葉片葉綠素總含量下降,本研究結(jié)果表明,前期遮陰時(shí)間超過R2期后(VER5)葉綠素(a+b)含量相對(duì)于對(duì)照增幅有所下降。且后期遮陰對(duì)參試大豆品種葉綠素(a+b)的影響小于前期遮陰。李冬梅等[24]研究表明,遮陰降低玉米葉片的光補(bǔ)償點(diǎn)、光飽和點(diǎn)和最大光合速率,提高表觀量子利用效率。本研究結(jié)果表明,不同生育時(shí)期遮陰均降低大豆葉片的光補(bǔ)償點(diǎn)、光飽和點(diǎn)和最大光合速率,提高表觀量子利用效率,且后期遮陰對(duì)參試大豆品種葉片光合參數(shù)的影響小于前期遮陰。遮陰導(dǎo)致光合電子傳遞性能的改變,葉綠素?zé)晒鈪?shù)能準(zhǔn)確反映其變化[9-10]。徐彩龍等[25]研究表明,遮陰6 d后,小麥旗葉PSⅡ最大量子產(chǎn)量顯著低于對(duì)照,PSⅡ?qū)嶋H量子產(chǎn)量顯著高于對(duì)照,本研究結(jié)果表明,受遮陰影響,VER2和VER5 的PSⅡ最大量子產(chǎn)量減少,PSⅡ?qū)嶋H量子產(chǎn)量增加,且 VER5 PSⅡ最大量子產(chǎn)量減少幅度大于 VER2,VER5 PSⅡ?qū)嶋H量子產(chǎn)量增加幅度小于 VER2,說明遮陰至R2期,隨遮陰時(shí)間延長,PSⅡ反應(yīng)中心結(jié)構(gòu)與功能受損程度增加,原初光能轉(zhuǎn)化效率相對(duì)于對(duì)照增幅減小。

        3.2 復(fù)光對(duì)大豆葉片光合特性的影響

        研究表明,作物葉片增加捕光色素復(fù)合體數(shù)量提高其對(duì)光能的捕獲能力以適應(yīng)弱光逆境,而恢復(fù)正常光照,傳遞到光反應(yīng)中心的激發(fā)能過剩,過剩能量導(dǎo)致光合系統(tǒng)適應(yīng)性調(diào)整[26],造成葉片光合特性改變。楊丹霞等[27]研究發(fā)現(xiàn)遮陰后復(fù)光,玉米葉片葉綠素a/b比值高于弱光下生長的處理,低于自然光照下處理。本試驗(yàn)研究結(jié)果表明,VER2和VER5遮陰期間葉綠素a/b均低于對(duì)照,且VER5下降幅度大于VER2下降幅度,復(fù)光后VER2和VER5葉綠素a/b仍低于對(duì)照,但相對(duì)于遮陰時(shí),與對(duì)照的差異程度均減小,且VER2與對(duì)照差異程度小于VER5與對(duì)照差異程度。說明R2期復(fù)光,大豆葉片葉綠素a/b比值恢復(fù)性調(diào)整效果明顯,而R5期復(fù)光,葉綠素a/b比值無明顯變化。前人研究結(jié)果表明,作物葉片遮陰后復(fù)光,通過調(diào)整其光合機(jī)構(gòu)性能以適應(yīng)光環(huán)境的變化,表現(xiàn)為葉綠素?zé)晒鈪?shù)的改變[27-28]。PSⅡ最大量子產(chǎn)量反映了植物的潛在最大光合能力,PSⅡ最大量子產(chǎn)量下降表示植物受到了脅迫[26],非光化學(xué)熒光淬滅系數(shù)反映植物耗散過剩光能為熱的能力,研究表明,植物光合器官適應(yīng)弱光環(huán)境后恢復(fù)自然光照,會(huì)通過提高耗散過剩光能為熱的能力以保護(hù)光合器官,表現(xiàn)為非光化學(xué)熒光淬滅系數(shù)上升[27]。本試驗(yàn)研究結(jié)果表明,VER2和VER5復(fù)光后,VER2 PSⅡ最大量子產(chǎn)量相對(duì)對(duì)照下降幅度比在遮陰期間小,VER5卻與之相反,VER2和VER5 非光化學(xué)熒光淬滅系數(shù)均高于對(duì)照且VER2上升幅度小于VER5,說明R2期復(fù)光,大豆葉片光合機(jī)構(gòu)受弱光的損害有一定恢復(fù),R5期復(fù)光,已適應(yīng)弱光環(huán)境的葉片對(duì)自然光照表現(xiàn)為強(qiáng)光脅迫,光合機(jī)構(gòu)進(jìn)一步受到強(qiáng)光脅迫破壞,且R2期和R5期復(fù)光,大豆葉片為保護(hù)光合組織均表現(xiàn)為耗散過剩光能的能力大幅提升。

        4 結(jié)論

        不同生育時(shí)期遮陰對(duì)大豆葉片光合作用有顯著影響,光合和葉綠素?zé)晒馓匦苑謩e在不同大豆品種遮陰處理間的差異上表現(xiàn)出共性規(guī)律:出苗至盛花期遮陰對(duì)大豆葉片光合及葉綠素?zé)晒馓匦缘挠绊懘笥谑⒒ㄆ谥镣晔炱谡陉?;大豆從出苗開始遮陰在盛花期前復(fù)光,有利于葉片光合能力的恢復(fù),但超過鼓粒期復(fù)光,已經(jīng)適應(yīng)弱光環(huán)境的葉片突然暴露在自然光下表現(xiàn)為受到強(qiáng)光脅迫,反而不利于大豆葉片光合作用。因此,從減小遮陰對(duì)大豆葉片光合作用不良影響的角度考慮,在中國南方丘陵和山區(qū)玉米大豆間作優(yōu)于玉米大豆套作;在玉米大豆套作模式下選擇品種、播期及種植技術(shù)時(shí),應(yīng)確保大豆在盛花期前恢復(fù)光照,避免遮陰超過大豆鼓粒期。另外,遮陰對(duì)大豆產(chǎn)量構(gòu)成也有顯著影響,且遮陰時(shí)間越長,大豆減產(chǎn)越嚴(yán)重,但大豆產(chǎn)量最終由光合作用和光合同化產(chǎn)物分配2個(gè)因素決定,且參試大豆品種間存在差異,所以不同遮陰處理對(duì)大豆籽粒產(chǎn)量影響的差異在本文中還不明確,有待進(jìn)一步試驗(yàn)研究。

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        [27] 楊丹霞, 張亞田, 蔡敦江, 孫廣玉. 育苗溫室弱光下的玉米幼苗移栽到自然光下的光合和熒光特性研究. 玉米科學(xué), 2010, 18(5): 60-64. YANG D X, ZHANG Y T, CAI D J, SUN G Y. Studies on photosynthetic characteristics and chlorophyⅡ Fluorescence in maize seedlings transferred from lowlight-grown in greenhouse to sunlight. Journal of Maize Sciences, 2010, 18(5): 60-64. (in Chinese)

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        (責(zé)任編輯 楊鑫浩,李莉)

        Effect of Shading on Soybean Leaf Photosynthesis and Chlorophyll Fluorescence Characteristics at Different Growth Stages

        WANG Yi1,2, ZHANG Xia1,3, YANG Wen-yu1, SUN Xin1, SU Ben-ying1, CUI Liang1
        (1College of Agriculture, Sichuan Agriculture University/Key Laboratory of Crop Ecophysiology and Farming System in Southwest China, Ministry of Agriculture, Chengdu 611130;2Agricultural Technology Promotion Station of Daying, Suining 629000, Sichuan;3Chongqing Tongnan Middle School, Chongqing 404100)

        Abstract:【Objective】The present work studied the effect of shading on photosynthesis and chlorophyll fluorescencecharacteristics, including leaf area, lamina mass per unit area, chlorophyll content, yield characters, photosynthetic and fluorescence parameters of three soybean cultivars at different growth stages, in order to support the development of maize-soybean intercropping and relay-intercropping systems in hilly and mountainous area of southern China.【Method】Three cultivars named Guixia 2, Nandou 12 and C103, whose period and total days for growth were different with each other, were studied. Soybean plants were shaded by shading nets with 50% transmittance during growth stages of emergence to full-bloom (VER2), emergence to pod-fill (VER5), and full-bloom to full-ripe (R2R8), respectively. Plants grown under natural light were used as control (CK). Leaf area, lamina mass per unit area, chlorophyll content, photosynthetic and fluorescence parameters, as well as yield characters were measured and analyzed. 【Result】 Compared to the control (CK), leaf area, chlorophyll (a+b)content and apparent quanta efficiency at the VER2 stage increased by 15.5%, 13.0% and 74.1%, respectively; lamina mass per unit area, light compensation point and maximum photosynthetic rate were decreased by 15.8%, 26.2% and 26.5%, respectively,compared to CK. Leaf area, chlorophyll (a+b) content and apparent quanta efficiency increased by 0.3%, 10.5% and 28.1%,respectively at the R2R8 stage; while lamina mass per unit area, light compensation point and maximum photosynthetic rate decreased by 10.2%, 20.3% and 12.1%, respectively, compared to CK. When soybean plants were re-illuminated at full bloom stage, maximal quantum yield of PSII was decreased by 3.0% and non-photochemical quenching was increased by 26.8%,compared to CK. When plants were re-illuminated at pod-fill stage, maximal quantum yield of PSII was decreased by 8.5% and non-photochemical quenching was increased by 40.8%, compared to CK. In addition, at VER2, VER5 and R2R8 stages for Guixia 2, yields per plant decreased by 40.8%, 48.7% and 59.2%, and 100-seed weight decreased by 23.7%, 39.3% and 26.4%,respectively, compared to CK. For Nandou 12 at these stages, yields per plant decreased by 46.7%, 54.2% and 21.2%, and 100-seed weight were decreased by 3.9%, 19.9% and 26.1%, respectively, compared to CK. For C103, yields per plant decreased by 69.8%, 74.9% and 73.9%, and 100-seed weight decreased compared to CK by 68.8%, 69.6% and 71.6%, respectively. 【Conclusion】Shading to the VER2 stage had a greater impact than for R2R8 on the photosynthesis and chlorophyll fluorescence characteristics of soybean. Photosynthetic capacity of soybean leaves made some recovery from shading when re-illuminated at R2. However, high-light stress was observed for plants re-illuminated at R5. Therefore, we suggest that maize-soybean intercropping to be a more advantageous system than relay-intercropping in hilly and mountainous area of southern China, with the perspective of minimizing the negative effects of shading on soybean photosynthesis. We also suggest, it is necessary to re-illuminate the soybean plants before full-bloom stage, in order to avoid an excessively long shading duration, when making decisions about on cultivar use, sowing time, and planting technology. The effect of shading period can influence yield of soybean. Moreover, the longer the shading duration last, the more the expected yield loss.

        Key words:soybean; maize; growth stages; shading; photosynthesis; chlorophyll fluorescence

        收稿日期:2015-11-06;接受日期:2016-03-21

        基金項(xiàng)目:國家自然科學(xué)基金(31171476,31201169)、國家“973”計(jì)劃(2011CB100402)、農(nóng)業(yè)部公益性行業(yè)科研專項(xiàng)(201203096)

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