郝玉杰，武林楠，馮建榮，白瑪多吉，王富霞，榮新民

(1. 石河子大學(xué)農(nóng)學(xué)院/特色果蔬栽培生理與種質(zhì)資源利用兵團(tuán)重點(diǎn)實(shí)驗(yàn)室，新疆石河子 832003；2. 石河子農(nóng)業(yè)科學(xué)研究院葡萄研究所/葡萄北疆試驗(yàn)站，新疆石河子 832000)

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NaCl脅迫對(duì)兩個(gè)葡萄品種葉片光合特性的影響

郝玉杰1，武林楠1，馮建榮1，白瑪多吉1，王富霞2，榮新民2

(1. 石河子大學(xué)農(nóng)學(xué)院/特色果蔬栽培生理與種質(zhì)資源利用兵團(tuán)重點(diǎn)實(shí)驗(yàn)室，新疆石河子 832003；2. 石河子農(nóng)業(yè)科學(xué)研究院葡萄研究所/葡萄北疆試驗(yàn)站，新疆石河子 832000)

【目的】研究NaCl脅迫對(duì)兩個(gè)葡萄新品種火洲紅玉、火洲黑玉光合特性的影響，為篩選耐鹽新品種提供理論依據(jù)?！痉椒ā坎捎盟喾?，對(duì)兩個(gè)葡萄品種進(jìn)行不同濃度NaCl脅迫(0、2、3、4 g/L)處理，8 d后測(cè)定葉片中的葉綠素相對(duì)含量(SPAD)值、初始熒光(Fo)、最大熒光(Fm)、PSⅡ?qū)嶋H光化學(xué)效率(ФPSⅡ)、PSⅡ最大光化學(xué)效率(Fv/Fm)、凈光合速率(Pn)、氣孔導(dǎo)度(Gs)、胞間二氧化碳濃度(Ci)和蒸騰速率(Tr)?！窘Y(jié)果】在NaCl≤3 g/L時(shí)，兩品種SPAD值較對(duì)照顯著升高，至4 g/L時(shí)，SPAD值均下降，火洲紅玉SPAD值顯著低于對(duì)照；隨NaCl濃度升高，火洲黑玉的Fo值升高但較對(duì)照無(wú)顯著性差異，F(xiàn)m、ФPSⅡ、Fv/Fm值呈下降趨勢(shì)但均與對(duì)照無(wú)顯著性差異；火洲紅玉Fo值在NaCl≥3 g/L時(shí)較對(duì)照顯著升高，F(xiàn)m、ФPSⅡ、Fv/Fm值較對(duì)照顯著下降；火洲黑玉Ci在2 g/L時(shí)，較對(duì)照顯著降低，在NaCl≥3 g/L時(shí)Ci升高但與對(duì)照無(wú)顯著性差異，Gs在各處理下較對(duì)照均無(wú)顯著性差異，Pn和Tr較對(duì)照均顯著降低；火洲紅玉Ci均顯著高于對(duì)照，Gs、Pn、Tr均顯著低于對(duì)照?！窘Y(jié)論】火洲黑玉在4 g/L NaCl處理時(shí)，SPAD值較對(duì)照顯著下降，Pn、Tr較對(duì)照顯著降低，其他指標(biāo)均與對(duì)照無(wú)顯著性差異，火洲紅玉在NaCl≥3 g/L時(shí)各項(xiàng)指標(biāo)均與對(duì)照存在顯著差異，火洲黑玉耐鹽性較火洲紅玉好。

葡萄；鹽脅迫；光合特性；葉綠素?zé)晒?/p>

0 引 言

【研究意義】土壤鹽漬化已經(jīng)成為一個(gè)世界性的生態(tài)問(wèn)題，園藝作物生產(chǎn)也深受其害[1]。鹽害往往對(duì)作物造成滲透脅迫、離子毒害、營(yíng)養(yǎng)不平衡等危害，從而影響植物光合作用，抑制植物的生長(zhǎng)，甚至導(dǎo)致其死亡[2]。新疆鹽堿地占總耕地的31%[3]，葡萄作為在新疆種植面積較大的樹(shù)種之一[4]，鹽害已成為影響葡萄生產(chǎn)的主要問(wèn)題之一[5-7]。因此，研究葡萄的耐鹽性，對(duì)培育、推廣適宜鹽堿地種植的葡萄品種有重要的意義?！厩叭搜芯窟M(jìn)展】前人對(duì)葡萄在鹽脅迫下光合特性做了許多研究，如秦嶺等[8]認(rèn)為在100 mmol/L的NaCl脅迫下葉綠素含量降低，最終導(dǎo)致PSⅡ潛在活性中心受損，進(jìn)而影響CO2的固定和同化，光能轉(zhuǎn)化效率和凈光合效率明顯降低。李學(xué)孚等[9]研究表明，在NaCl>0.4%時(shí),‘鄞紅’葡萄葉綠素含量、葉綠素?zé)晒馓匦?、氣體交換參數(shù)均呈現(xiàn)下降趨勢(shì)。但馬帥等[10]研究顯示，隨鹽濃度的升高，葡萄的氣體交換參數(shù)除胞間CO2濃度(Ci)逐漸上升外，其余參數(shù)均呈現(xiàn)下降的趨勢(shì)。邢慶振等[11]對(duì)‘紅地球’研究表明，隨鹽脅迫的加劇，光合特性中除Ci呈現(xiàn)先下降后上升趨勢(shì)，其余各項(xiàng)指標(biāo)均呈現(xiàn)下降趨勢(shì)。秦紅艷等[12]認(rèn)為，隨鹽濃度的升高山葡萄初始熒光(Fo)顯著升高，其他熒光參數(shù)均顯著降低，PSⅡ反應(yīng)中心被破壞?！颈狙芯壳腥朦c(diǎn)】研究選取兩個(gè)葡萄新品種火洲黑玉和火洲紅玉，采用水培方法，從幼苗光合作用和葉綠素?zé)晒鈩?dòng)力學(xué)特征入手，探究其對(duì)不同濃度NaCl脅迫對(duì)兩品種的葉綠素相對(duì)含量、葉綠素?zé)晒鈪?shù)及光合特性影響?！緮M解決的關(guān)鍵問(wèn)題】測(cè)定兩個(gè)品種在鹽脅迫下光合指標(biāo)，了解兩個(gè)葡萄品種光合特性對(duì)鹽脅迫的響應(yīng)，為兩個(gè)葡萄品種在北疆地區(qū)進(jìn)一步培育和推廣提供理論依據(jù)。

1 材料與方法

1.1 材 料

參試的2個(gè)葡萄品種火洲黑玉，火洲紅玉是吐魯番葡萄研究所選育的歐亞種，由石河子葡萄研究所引入的2年生苗。2015年5月30日，選取生長(zhǎng)健壯的火洲黑玉和火洲紅玉葡萄幼苗各36株，從田間移出，用清水沖洗根，先置于塑料桶(上口徑20 cm，下口徑15 cm，高25 cm)用清水中培養(yǎng)1 d，用1/2Hoagland營(yíng)養(yǎng)液[13]培養(yǎng)4 d，最后用全營(yíng)養(yǎng)液進(jìn)行培養(yǎng)，每個(gè)塑料桶中3株，作為1個(gè)重復(fù)，并用苯板固定，3次重復(fù)，培養(yǎng)51 d，于7月21日開(kāi)始進(jìn)行鹽脅迫試驗(yàn)。

1.2 方 法

1.2.1 試驗(yàn)材料

設(shè)3個(gè)NaCl濃度梯度，對(duì)照為Hoagland營(yíng)養(yǎng)液；處理Ⅰ為Hoagland營(yíng)養(yǎng)液+2 g/L NaCl；處理Ⅱ?yàn)镠oagland營(yíng)養(yǎng)液+3 g/L NaCl；處理Ⅲ為Hoagland營(yíng)養(yǎng)液+4 g/L NaCl。試驗(yàn)期間營(yíng)養(yǎng)液用充氣泵連續(xù)通氣，每天補(bǔ)充塑料桶中散失的水分至5 L刻度線，并且每4 d更換一次營(yíng)養(yǎng)液。鹽脅迫第8 d，從每個(gè)重復(fù)中隨機(jī)選取1株，共選3株，每株從頂部向下數(shù)第3～4片功能葉用來(lái)測(cè)定葉片的葉綠素相對(duì)含量、氣體交換參數(shù)及葉綠素?zé)晒鈪?shù)。

1.2.2 測(cè)定項(xiàng)目1.2.2.1 葉綠素含量

利用SPAD-502+PLUS葉綠素儀參照李學(xué)孚等[8]方法于10：00～12：00對(duì)葉綠素含量進(jìn)行測(cè)定。

1.2.2.2 葉綠素?zé)晒鈪?shù)

選取健壯植株，使用FMS-2脈沖調(diào)制式熒光儀參照邢慶振等[11]方法，于10:00～12:00測(cè)定熒光參數(shù)，光適應(yīng)下，打開(kāi)內(nèi)源光化光(600 μmol/(m2·s))測(cè)定光適應(yīng)后最小熒光(Fs)、最大熒光(Fm’)、PSⅡ?qū)嶋H光化學(xué)效率ФPSⅡ，暗適應(yīng)30 min后先照射檢測(cè)光(<0.05 μmol/(m2·s))，測(cè)定初始熒光(Fo)、再照射飽和脈沖光(12 000μmol/(m2·s))，測(cè)定 PSⅡ最大光化學(xué)量子效率Fv/Fm。

1.2.2.3 光合指標(biāo)

利用LI-6400便攜式光合作用測(cè)定系統(tǒng)參照徐煥文等[14]方法于10:00～12:00測(cè)定光和參數(shù)，凈光合速率(Pn)、氣孔導(dǎo)度(Gs)、蒸騰速率(Tr)、胞間CO2濃度(Ci)由儀器自動(dòng)給出[14]。測(cè)定時(shí)溫度為(27±2)℃，光照強(qiáng)度為1 300 μmol/(m2·s)，CO2濃度(380±10) μmol/mol。

1.3 數(shù)據(jù)統(tǒng)計(jì)

試驗(yàn)原始數(shù)據(jù)處理和繪圖采用Excel軟件完成，差異顯著分析采用SPSS 19.0軟件。

2 結(jié)果與分析

2.1 鹽脅迫對(duì)兩個(gè)葡萄品種葉片中葉綠素相對(duì)含量的影響

研究表明，不同濃度NaCl處理后，火洲黑玉和火洲紅玉的葉綠素相對(duì)含量(SPAD)值隨著NaCl濃度升高呈現(xiàn)先升高后降低的趨勢(shì)。在NaCl≤3 g/L時(shí)兩品種SPAD值較對(duì)照均顯著升高且在3 g/L時(shí)達(dá)到峰值，分別為31.2和28.2，在4 g/L處理時(shí)，火洲黑玉SPAD值下降但與對(duì)照差異不顯著，而火洲紅玉SPAD值較對(duì)照顯著降低。鹽脅迫下火洲黑玉的SPAD值顯著高于火洲紅玉。圖1

圖1 鹽脅迫下葡萄葉片中葉綠素相對(duì)含量變化
Fig.1 Effects of NaCl stress on the chlorophyll relative content in grape leaves

2.2 鹽脅迫對(duì)不同葡萄品種葉片中熒光參數(shù)的影響

研究表明，隨著NaCl濃度的升高，兩個(gè)葡萄品種Fo呈現(xiàn)上升趨勢(shì)，F(xiàn)m、ФPSⅡ、Fv/Fm均呈現(xiàn)下降趨勢(shì)?；鹬藓谟馞o值在各處理時(shí)較對(duì)照無(wú)顯著性差異；火洲紅玉在NaCl≥3 g/L時(shí)Fo值較對(duì)照均顯著升高，分別升高了55.6%、79.5%?；鹬藓谟馞m值在各處理時(shí)較對(duì)照無(wú)顯著性差異；火洲紅玉在NaCl≥3 g/L時(shí)Fm值較對(duì)照顯著下降，分別下降了38.2%和44%。火洲黑玉在各處理時(shí)ФPSⅡ值較對(duì)照無(wú)顯著性差異；火洲紅玉在NaCl≥3 g/L時(shí)ФPSⅡ值較對(duì)照顯著下降，分別下降了14.4%和19.3%。火洲黑玉Fv/Fm值在各處理時(shí)較對(duì)照無(wú)顯著性差異；火洲紅玉在NaCl≥3 g/L時(shí)Fv/Fm值較對(duì)照顯著下降，分別下降了14.5%和23.2%?；鹬藓谟馞o值上升幅度顯著低于火洲紅玉，火洲黑玉Fm、ФPSⅡ、Fv/Fm值下降幅度顯著低于火洲紅玉，火洲紅玉在NaCl≥3 g/L處理時(shí)較火洲黑玉PSⅡ受損程度更高。圖2

圖2 鹽脅迫下葡萄葉片熒光參數(shù)變化
Fig. 2 Effects of NaCl stress on fluorescence parameters of grape leaves

2.3 鹽脅迫對(duì)兩個(gè)葡萄品種葉片氣體交換參數(shù)的影響

研究表明，隨NaCl濃度的升高兩品種Pn、Tr、Gs均呈下降趨勢(shì)，火洲黑玉Ci先下降后上升，而火洲紅玉Ci則呈上升趨勢(shì)?；鹬藓谟窈突鹬藜t玉在各處理下Pn值較對(duì)照均顯著降低，分別降低了12.8%、25.6%、43.6%和21.4%、42.9%、61.9%。火洲黑玉和火洲紅玉Tr值在各個(gè)處理時(shí)較對(duì)照均顯著降低，分別降低了22.7%、57.1%、64.2%和32.3%、44.6%、67.5%?；鹬藓谟裨诟魈幚硐翯s值與對(duì)照相比均未達(dá)到顯著水平，較對(duì)照分別降低了10.6%、12.9%、14.8%；火洲紅玉在各處理下Gs值均顯著低于對(duì)照，分別降低了28.1%、24.3%、50.6%。火洲黑玉在2 g/L時(shí)Ci值顯著低于對(duì)照，下降幅度為19.2%，在3 g/L、4 g/L時(shí)Ci較對(duì)照相比上升幅度未達(dá)顯著水平；火洲紅玉的Ci值在2、3 g/L時(shí)與對(duì)照相比無(wú)顯著性差異，在4 g/L時(shí)Ci值較對(duì)照顯著升高，升高了35.4%?；鹬藓谟竦腜n、Tr、Gs下降幅度顯著低于火洲紅玉。圖3

圖3 鹽脅迫下葡萄葉片氣體交換參數(shù)變化
Fig.3 Effects of NaCl stress on gas exchange parameters in grape leaves

3 討 論

植物葉片中葉綠素含量是衡量植物抗逆性的一個(gè)重要生理指標(biāo)[15]。李學(xué)孚等[8]對(duì)‘鄞紅’葡萄光合特性研究結(jié)果表明，低鹽脅迫下(NaCl≤0.4%)，SPAD值無(wú)顯著變化，而高鹽脅迫下(NaCl>0.4%)SPAD值顯著下降，鹽濃度(NaCl=0.4%)成為‘鄞紅’葡萄耐鹽性的臨界閾值。試驗(yàn)研究結(jié)果表明，火洲黑玉和火洲紅玉在(NaCl≤0.3%)時(shí)，SPAD值均表現(xiàn)升高的趨勢(shì)，在(NaCl>0.3%)時(shí)，兩個(gè)葡萄品種SPAD值表現(xiàn)下降的趨勢(shì)，鹽濃度(NaCl=0.3%)成為兩品種耐鹽性的臨界閾值，這與李學(xué)孚等[8]研究結(jié)果不完全一致，李學(xué)孚[8]使用的‘鄞紅’葡萄是歐美雜交種群，而試驗(yàn)使用的火洲黑玉和火洲紅玉是由吐魯番葡萄研究所選育的歐亞種，說(shuō)明試驗(yàn)利用的火洲黑玉和火洲紅玉的耐鹽性沒(méi)有歐美雜交的葡萄品種強(qiáng)。

NaCl脅迫導(dǎo)致植物光合作用的降低既有氣孔因素，也有非氣孔因素[18]。根據(jù)Farquhar等[19]提出的光合作用氣體交換模型中，若光合速率的下降并且Ci的上升，那么影響光合作用的主要限制因素為非氣孔因素，若光合速率下降伴隨著Ci同時(shí)下降，那么主要的限制光合作用的因素則為氣孔因素。試驗(yàn)研究表明，2 g/L處理下火洲黑玉葡萄葉片Pn、Gs下降，且Ci呈下降趨勢(shì)(3-D)，3 g/L、4 g/L處理下Pn和Gs下降的同時(shí)Ci呈現(xiàn)上升的趨勢(shì)，表明在低鹽濃度(NaCl<3 g/L)脅迫時(shí)氣孔限制成為降低光合作用的主要因素，而高濃度(NaCl≥3 g/L)脅迫時(shí)則轉(zhuǎn)為非氣孔限制所致。房玉林等[20]對(duì)8804品系葡萄的研究認(rèn)為隨鹽濃度升高葡萄光合作用的影響逐漸從氣孔限制轉(zhuǎn)變?yōu)榉菤饪紫拗?。試?yàn)與房玉林等[19]研究結(jié)果一致?；鹬藜t玉的Pn和Gs在整個(gè)脅迫過(guò)程中均呈現(xiàn)下降的趨勢(shì)，Ci呈現(xiàn)上升趨勢(shì)，表明火洲紅玉對(duì)鹽脅迫較為敏感，其光合作用下降都由非氣孔限制所致。這與馬帥等[10]對(duì)‘摩爾多瓦’研究結(jié)果一致。

4 結(jié) 論

在NaCl≤3 g/L處理下火洲黑玉和火洲紅玉，SPAD值顯著升高，NaCl>3 g/L處理下，SPAD值均下降，火洲紅玉SPAD值較對(duì)照顯著下降；NaCl脅迫下，兩品種Fo值上升、Fm、Fv/Fm和ФPSⅡ值下降，但火洲黑玉在各處理下較對(duì)照均無(wú)顯著性差異，火洲紅玉在NaCl≥3 g/L處理下較對(duì)照均存在顯著性差異；火洲黑玉NaCl脅迫下，Gs較對(duì)照無(wú)顯著性差異，Pn、Tr及2 g/L時(shí)Ci較對(duì)照顯著下降，在NaCl≥3 g/L時(shí)，Ci上升但與對(duì)照無(wú)顯著性差異；火洲紅玉Ci較對(duì)照顯著上升，Pn、Gs、Tr均較對(duì)照顯著下降。火洲紅玉各項(xiàng)指標(biāo)變化幅度均顯著高于火洲黑玉，火洲黑玉耐鹽性較火洲紅玉好。

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Fund project:Special funds for construction projects of modern agricultural industry technology system(CARS-30)

Effects of NaCl Stress on Photosynthetic Characteristics of Leaves of the Two Grape Cultivars

HAO Yu-jie1，WU Lin-nan1，F(xiàn)ENG Jian-rong1，Baimaduoji1，WANG Fu-xia2，RONG Xin-min2

(1.CollegeofAgronomy,ShiheziUniversity/XPCCKeyLaboratoryofSpecialFruitsandVegetablesCultivationPhysiologyandGermplasmResourcesUtilization,ShiheziXinjiang832003,China; 2GrapeTestStationofGrapeResearchInstituteofShiheziAcademyofAgriculturalSciences,ShiheziXinjiang832000,China)

【Objective】 The impact of salt stress on the photosynthetic characteristics of two new grape cultivars 'Huozhouheiyu' and 'Huozhouhongyu' were studied for providing a foundation for screening of new salt tolerant cultivars.【Method】Two cultivars in hydroponic cultures were treated with different concentrations of NaCl(0,2 g/L, 3 g/L, 4 g/L), and Chlorophyll content (SPAD), the minimal fluorescence (Fo), the maximal fluorescence(Fm), actual photochemical efficiency of PSⅡ(ФPSⅡ), the photochemical efficiency of PSⅡ(Fv/Fm), net photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (Gs) and Intercellular CO2concentration (Ci) were determined after eight days.【Result】SPADof two cultivars were significantly higher in the 3 g/L and 4 g/L treatment than in the control treatment. In the 4 g/L treatment,SPADof two cultivars decreased, andSPADof 'Huozhouhongyu' was significantly lower than that of the control treatment.Foof 'Huozhouheiyu' increased and had no significant difference than control treatment.Fm, ФPSⅡ,Fv/Fmof 'Huozhouheiyu' decreased and had no significant difference than that of the control treatment.Foof 'Huozhouhongyu' was significantly higher in the 3 g/L and 4 g/L treatment than in the control treatment, andFm, ФPSⅡ,Fv/Fmof 'Huozhouhongyu' was significantly lower in the 3 g/L and 4 g/L treatment than in the control treatment.Ciof 'Huozhouheiyu' was significantly lower in the 2 g/L treatment than in the control treatment.Ciof 'Huozhouheiyu' increased and was not significantly different than that of the control treatment.Gsof 'Huozhouheiyu' was not significantly different than that of the control treatment.PnandTrwere significantly lower than that of the control treatment.Ciof 'Huozhouhongyu' was significantly higher than that of the control treatment, andPn,Tr,Gswere significantly lower.【Conclusion】In the 4 g/L treatment,SPADof 'Huozhouheiyu' decreased.PnandTrwere significantly lower than those of the control treatment. Other indexes were not significantly different than those of the control treatment. In the 3 g/L and 4 g/L, all indexes of 'Huozhouhongyu' were not siginificantly different than those of the control treatment. 'Huozhouhongyu' was affected more seriously than 'Huozhouheiyu' by salt stress.

grapes; salt stress; photosynthetic characteristics; chlorophyll fluorescence

10.6048/j.issn.1001-4330.2016.10.004

2016-05-12

現(xiàn)代農(nóng)業(yè)產(chǎn)業(yè)技術(shù)體系建設(shè)專項(xiàng)資金項(xiàng)目(CARS-30)

郝玉杰(1993-)，女，新疆人，碩士研究生，研究方向?yàn)楣麡?shù)種質(zhì)資源與遺傳育種，(E-mail)1275185994@qq.com

馮建榮(1969-)，女，新疆人，教授，博士，研究方向?yàn)楣麡?shù)種質(zhì)資源與分子輔助育種，(E-mail)fengjr102@126.com

S663.1

A

1001-4330(2016)10-1794-07