玉米弱勢(shì)粒庫(kù)活性與籽粒內(nèi)源激素及多胺含量的關(guān)系

王志剛梁紅偉高聚林于曉芳孫繼穎蘇治軍胡樹(shù)平余少波李雅劍魏淑麗楊 哲

1內(nèi)蒙古農(nóng)業(yè)大學(xué)農(nóng)學(xué)院,內(nèi)蒙古呼和浩特 010019;2內(nèi)蒙古自治區(qū)農(nóng)牧業(yè)科學(xué)院,內(nèi)蒙古呼和浩特 010031

玉米弱勢(shì)粒庫(kù)活性與籽粒內(nèi)源激素及多胺含量的關(guān)系

王志剛1,**梁紅偉1,2,**高聚林1,*于曉芳1孫繼穎1蘇治軍1胡樹(shù)平1余少波1李雅劍1魏淑麗1楊 哲1

1內(nèi)蒙古農(nóng)業(yè)大學(xué)農(nóng)學(xué)院,內(nèi)蒙古呼和浩特 010019;2內(nèi)蒙古自治區(qū)農(nóng)牧業(yè)科學(xué)院,內(nèi)蒙古呼和浩特 010031

玉米籽粒形成期的庫(kù)活性是弱勢(shì)粒敗育或灌漿受限的核心限制因子,明確弱勢(shì)粒中內(nèi)源激素及多胺水平對(duì)其庫(kù)活性的調(diào)控機(jī)制,對(duì)探索密植條件下玉米弱勢(shì)粒調(diào)控途徑具有重要意義。本研究以典型玉米雜交種鄭單958和先玉335為材料,在控制授粉條件下(不完全授粉IcP、完全授粉CP),比較分析了成功發(fā)育弱勢(shì)粒(IcP處理)和發(fā)育不良弱勢(shì)粒(CP處理)的內(nèi)源激素及多胺水平差異及其與庫(kù)活性的關(guān)系。結(jié)果表明,品種和年度對(duì)籽粒庫(kù)活性、內(nèi)源激素和多胺水平整體無(wú)顯著影響。IcP處理下弱勢(shì)粒的可溶性酸性蔗糖轉(zhuǎn)化酶(SAI)活性顯著高于CP處理,平均差異和最大差異分別達(dá)13.5%和21.8%。在玉米籽粒形成期,弱勢(shì)粒中玉米素和玉米素核苷(Z+ZR)、生長(zhǎng)素(IAA)、赤霉素(GA3)和脫落酸(ABA)含量在兩種控制授粉處理間無(wú)顯著差異。弱勢(shì)粒中多胺含量表現(xiàn)為IcP處理顯著高于CP處理,而乙烯釋放速率則恰恰相反。弱勢(shì)粒中SAI活性與多胺含量顯著正相關(guān),而與乙烯釋放速率顯著負(fù)相關(guān),且多胺含量與乙烯釋放速率顯著負(fù)相關(guān)?？梢?jiàn),在玉米籽粒形成期,其弱勢(shì)粒中Z+ZR、IAA、GA3和ABA與其庫(kù)活性即SAI活性無(wú)關(guān);弱勢(shì)粒庫(kù)活性主要受多胺和乙烯含量影響,多胺促進(jìn)SAI活性而乙烯則抑制其活性,二者的平衡關(guān)系決定了弱勢(shì)粒成功發(fā)育與否;多胺和乙烯平衡關(guān)系受同化物質(zhì)供應(yīng)水平的調(diào)控。

玉米;弱勢(shì)粒;庫(kù)活性;激素;多胺

密植條件下弱勢(shì)粒不能正常發(fā)育或灌漿不充實(shí)造成的玉米產(chǎn)量損失在10%以上[1-2],促進(jìn)弱勢(shì)粒建成與灌漿是玉米產(chǎn)量挖潛的重要途徑之一[2-4]。作物籽粒能否有效灌漿與作物籽粒庫(kù)特征即籽粒庫(kù)對(duì)同化物質(zhì)的容納和吸收能力有關(guān)[5]。作物籽粒庫(kù)特征可分為庫(kù)容量和庫(kù)活性?xún)蓚€(gè)方面[6-7]。庫(kù)容量主要決定于胚乳細(xì)胞數(shù)目,是物理性限制因子,與胚乳細(xì)胞分裂過(guò)程有關(guān)[8];庫(kù)活性則是庫(kù)器官卸載向其輸入的同化物質(zhì)的能力,在玉米籽粒形成期,主要與可溶性酸性蔗糖轉(zhuǎn)化酶(SAI)活性有關(guān)[9-12]。梁紅偉等[13]在控制授粉條件下比較分析成功發(fā)育弱勢(shì)粒和敗育弱勢(shì)粒的庫(kù)特征表明,玉米籽粒形成期的庫(kù)活性(SAI活性)是弱勢(shì)粒敗育或灌漿受限的核心限制因子;徐云姬等研究則表明[14-15],在灌漿期玉米弱勢(shì)粒灌漿差、粒重低與其較低的庫(kù)活性(淀粉合成相關(guān)酶活性)密切相關(guān)。籽粒中激素含量及多胺水平高低對(duì)籽粒的灌漿充實(shí)起重要調(diào)控作用,但前人的研究主要聚焦在不同粒位籽粒間的差異上,對(duì)敗育弱勢(shì)籽粒和成功發(fā)育弱勢(shì)籽粒間的內(nèi)源激素及多胺含量差異研究甚少[9-19],且玉米籽粒形成期的庫(kù)活性是否受激素和多胺水平的調(diào)控并最終決定其能否成功發(fā)育也尚不明確。因此,本試驗(yàn)通過(guò)控制授粉創(chuàng)造成功發(fā)育弱勢(shì)粒和敗育弱勢(shì)粒,比較二者間內(nèi)源激素及多胺含量的差異,并分析其與籽粒庫(kù)活性的關(guān)系,以期為探討玉米弱勢(shì)粒調(diào)控途徑提供理論參考。

1 材料與方法

1.1 試驗(yàn)地概況

包頭市土默特右旗薩拉齊鎮(zhèn)(Salaqi,40°28′N(xiāo), 110°28′E)試驗(yàn)基地,前茬為玉米,土質(zhì)為沙壤土。2014年0~30 cm土壤含有機(jī)質(zhì)24.5 g kg–1、堿解氮21.2 mg kg–1、速效磷26.7 mg kg–1、速效鉀120.4 mg kg–1,pH值為7.8;生育期內(nèi)總降雨量為417.1 mm,≥10℃活動(dòng)積溫2896.5℃,太陽(yáng)輻射總量3450.1 MJ m–2。2015年0~30 cm土壤含有機(jī)質(zhì)26.5 g kg–1、堿解氮20.5 mg kg–1、速效磷24.2 mg kg–1、速效鉀90.8 mg kg–1,pH值為7.4;生育期內(nèi)總降雨量為398.5 mm,≥10℃活動(dòng)積溫 2934.5℃,太陽(yáng)輻射總量3503.8 MJ m–2(圖1)。

1.2 試驗(yàn)設(shè)計(jì)

圖1 2014年和2015年玉米生育期內(nèi)的降雨量、溫度和日輻射量Fig.1 Precipitation,daily average temperature and solar radiation during the growth period of maize in 2014 and 2015

以典型玉米單交種鄭單958(ZD958)和先玉335 (XY335)為材料,種植密度7.5萬(wàn)株 hm–2,行距60cm,完全隨機(jī)排列,3次重復(fù),共6個(gè)小區(qū),小區(qū)面積112 m2(8 m×14 m)。以P2O5105 kg hm–2、純N 40 kg hm–2(磷酸二銨)、K2O 40 kg hm–2(硫酸鉀)為底肥于春播前旋耕入土,于拔節(jié)期追施純N 150 kg hm–2(尿素),其他管理同當(dāng)?shù)卮筇锷a(chǎn)。

采用馮漢宇等[20]的方法,玉米抽絲前在各小區(qū)內(nèi)選取生長(zhǎng)整齊一致、有全田代表性的植株80~100株掛牌標(biāo)記并對(duì)其雌穗套袋,待到花絲完全抽出后,選取一半植株的果穗做不完全授粉處理(IcP,果穗頂部和中部小花正常授粉,通過(guò)剪掉外圍花絲使果穗下部花絲不授粉),另一半植株的果穗做完全授粉處理(CP,玉米果穗全部花絲同步授粉),分別建成弱勢(shì)籽粒和敗育(或發(fā)育不良)弱勢(shì)籽粒。

1.3 測(cè)定指標(biāo)及方法

從授粉后開(kāi)始每隔6 d選不完全授粉和完全授粉果穗各5個(gè),取2~10環(huán)籽粒(從果穗最頂端數(shù)起)為上部弱勢(shì)籽粒,脫100粒左右混勻后,隨機(jī)挑出20粒左右稱(chēng)其鮮重后放入15 mL的玻璃試管,并在試管中放入一條浸濕的濾紙條以保持濕度,管口用配套的翻口皮塞塞緊,25℃條件下避光放置12 h后采氣,采用GC-2014型氣相色譜儀測(cè)定乙烯含量并計(jì)算乙烯釋放速率[21]。

隨機(jī)選取10粒左右參照高俊鳳[22]的方法測(cè)定可溶性酸性蔗糖轉(zhuǎn)化酶活性。剩余部分用于內(nèi)源玉米素(Z)、玉米素核苷(ZR)、吲哚-3-乙酸(IAA)、脫落酸(ABA)和赤霉素(GA3)及精胺(Spm)、亞精胺(Spd)、腐胺(Put)含量的測(cè)定。提取、純化和定量分析各內(nèi)源激素參照徐云姬等[13]的方法,采用Agilent-1200型液相色譜儀檢測(cè)分析。色譜條件為Dubhe C18 4.6×250,5 μm,流動(dòng)相體積比為 5%乙腈、50%甲醇、45%的0.6%冰乙酸,流速為0.8 mL min?1,采用梯度洗脫法,檢測(cè)波長(zhǎng)為254 nm;柱溫30℃,進(jìn)樣量10 μL。樣品回收率為(86.6±2.3)%,每一個(gè)樣品重復(fù)3次,外標(biāo)法定量。

提取、純化和定量分析籽粒中多胺參照劉華英等[23]的方法并加以改進(jìn)。采用Agilent-1200型液相色譜儀(帶600控制器和雙波長(zhǎng)吸光度檢測(cè)器),流動(dòng)相為甲醇∶水=60∶40,流速0.7 mL min–1,等度洗脫;進(jìn)樣量為10 μL,采用C18反相柱測(cè)試,柱溫30℃,在波長(zhǎng)254 nm下檢測(cè)。

1.4 數(shù)據(jù)分析

采用Microsoft Excel 2003、Origin 8.0統(tǒng)計(jì)分析數(shù)據(jù),SPSS17.0中One-way ANOVA進(jìn)行方差分析, SigmaPlot 12.0作圖。

2 結(jié)果與分析

2.1 不同控制授粉處理下弱勢(shì)粒庫(kù)活性比較

由圖2可見(jiàn),兩年度不同品種玉米弱勢(shì)粒的可溶性酸性蔗糖轉(zhuǎn)化酶(SAI)活性隨著生育進(jìn)程呈單峰曲線變化,在18~24 d間達(dá)到最大值。弱勢(shì)粒的SAI活性都表現(xiàn)為IcP處理(成功發(fā)育弱勢(shì)粒)顯著高于CP處理(發(fā)育不良弱勢(shì)粒),平均差異達(dá)13.5%,最大差異達(dá)21.8%。

圖2 不同控制授粉處理下玉米弱勢(shì)?？扇苄运嵝哉崽寝D(zhuǎn)化酶(SAI)活性變化Fig.2 Dynamics of soluble acid invertase(SAI)activity of inferior kernels of maize under different pollination treatmentsXY335:先玉335;ZD958:鄭單958;CP:完全授粉;IcP:不完全授粉。XY335:Xianyu 335;ZD958:Zhengdan 958;CP:complete pollination;IcP:incomplete pollination.

2.2 不同控制授粉處理下玉米弱勢(shì)粒 Z+ZR、IAA、GA3、ABA含量比較

隨著籽粒發(fā)育進(jìn)程,籽粒各內(nèi)源激素含量變化差異明顯(表1)。細(xì)胞分裂素類(lèi)物質(zhì)(Z+ZR)及3-吲哚乙酸(IAA)含量呈單峰曲線變化,在授粉后18 d左右達(dá)到峰值;脫落酸(ABA)含量變化呈相反趨勢(shì),至授粉后12~18 d降至低谷,之后明顯回升;赤霉素(GA3)含量則隨灌漿進(jìn)程呈持續(xù)降低趨勢(shì)。除IAA含量在授粉后12~24 d間表現(xiàn)為不穩(wěn)定的交替上升趨勢(shì),Z+ZR含量在2014年授粉后6 d表現(xiàn)為IcP>CP, GA3含量在2014年授粉后6 d表現(xiàn)為CP>IcP外,各激素含量在CP和IcP處理間整體差異不顯著。

表1 不同授粉處理下玉米弱勢(shì)粒的Z+ZR、IAA、GA3和ABA含量Table 1 Z+ZR,IAA,GA3,and ABA content of inferior kernels of maize under different pollination treatments

2.3 不同控制授粉處理下弱勢(shì)籽粒的乙烯釋放速率比較

由圖3可見(jiàn),各處理玉米弱勢(shì)籽粒乙烯釋放速率變化趨勢(shì)表現(xiàn)一致,授粉后6 d弱勢(shì)籽粒乙烯的釋放速率最大,隨籽粒發(fā)育進(jìn)程乙烯釋放速率逐漸降低。CP處理弱勢(shì)粒的乙烯釋放速率顯著高于IcP處理,CP處理較高的乙烯釋放量可能是造成其弱勢(shì)籽粒敗育或灌漿停滯的重要因子。

2.4 不同控制授粉處理下玉米弱勢(shì)粒發(fā)育期間的多胺含量比較

由圖4可見(jiàn),控制授粉后,同年份間兩品種籽粒中多胺含量變化趨勢(shì)整體一致,品種間無(wú)顯著差異,但年際間存在明顯差異,且變化趨勢(shì)不一致,表現(xiàn)為峰值出現(xiàn)的時(shí)間不同。2014年精胺和腐胺含量峰值出現(xiàn)在授粉后12 d,2015年則出現(xiàn)在授粉后18~24 d,而亞精胺含量峰值出現(xiàn)在授粉后12~18 d,年際間相對(duì)穩(wěn)定。IcP處理籽粒多胺含量在12~24 d都顯著高于CP處理。

圖3 不同控制授粉處理下玉米弱勢(shì)粒乙烯釋放速率變化Fig.3 Dynamics of ethylene producing rate of inferior kernels of maize under different pollination treatments縮寫(xiě)同圖2。Abbreviations are the same as those given in Fig.2.

2.5 玉米弱勢(shì)粒乙烯、多胺含量與SAI活性的相關(guān)分析

玉米弱勢(shì)籽粒內(nèi)源乙烯釋放速率與SAI活性呈極顯著負(fù)相關(guān)(r=0.497**~0.850**)(圖5),而Spm含量和 Spd含量與 SAI活性呈極顯著正相關(guān)(r= 0.544*~0.627**),各年份、品種及處理間規(guī)律一致(圖6)。由圖7可見(jiàn),乙烯釋放率與Spm和Spd含量呈顯著或極顯著負(fù)相關(guān)(r=-0.489*~-0.562**)。但Put含量與SAI活性及乙烯釋放速率相關(guān)不顯著。

3 討論

圖4 不同控制授粉處理下玉米弱勢(shì)粒多胺含量變化

作物籽粒庫(kù)特征受庫(kù)容量和庫(kù)活性?xún)蓚€(gè)因素的影響[6-7]。梁紅偉等[13]在不同控制授粉處理下比較玉米成功發(fā)育弱勢(shì)粒和敗育弱勢(shì)粒籽粒形成期的庫(kù)特征后認(rèn)為,胚乳細(xì)胞分裂過(guò)程和最大胚乳細(xì)胞數(shù)即庫(kù)容量不是弱勢(shì)粒敗育或灌漿停滯的制約因子,弱勢(shì)粒敗育或灌漿受限主要受這一時(shí)期庫(kù)活性(SAI活性)控制。但玉米籽粒形成期SAI活性受何因素調(diào)控則并不清楚。綜合一些學(xué)者在不同位勢(shì)籽粒間對(duì)籽粒庫(kù)活性的研究結(jié)論,影響作物籽粒庫(kù)活性的因素可概括為兩個(gè)方面,一為籽粒內(nèi)源激素水平,另一為籽粒中多胺和乙烯水平及其平衡關(guān)系。

圖5 玉米籽粒形成期弱勢(shì)粒乙烯釋放率與SAI活性的關(guān)系Fig.5 Relationship between ethylene release rate and soluble acid invertase(SAI)activity of inferior kernels during the blister stage of maize縮寫(xiě)同圖2。Abbreviations are the same as those given in Fig.2.

研究表明,ABA和IAA水平對(duì)作物籽粒庫(kù)活性有正向調(diào)控作用,而GA3則明顯抑制庫(kù)活性。楊建昌等[5]研究表明,水稻灌漿早期噴施低濃度ABA后,籽粒中蔗糖-淀粉關(guān)鍵酶活性顯著提高,說(shuō)明ABA通過(guò)調(diào)控籽粒中蔗糖-淀粉代謝途徑關(guān)鍵酶活性來(lái)促進(jìn)籽粒灌漿。王艷芳等[24]研究認(rèn)為,玉米籽粒中IAA含量高低順序與淀粉含量高低順序有良好的一致性,說(shuō)明高IAA含量有利于淀粉的合成;孫慶泉等[19]也認(rèn)為灌漿期籽粒中IAA含量越高,越利于籽粒的物質(zhì)充實(shí)。很多研究認(rèn)為,在胚乳細(xì)胞活躍增殖期或籽粒活躍灌漿期,籽粒中較高GA3會(huì)使α-淀粉酶等水解酶活性增強(qiáng),促進(jìn)淀粉的水解,降低玉米的粒重[25-27]。也有研究表明,籽粒中的內(nèi)源激素比例影響其庫(kù)活性,王紀(jì)華等[9]和張鳳路等[11]認(rèn)為,當(dāng)ABA水平偏高,而GA3水平偏低時(shí),籽粒中SAI活性升高,也就是相對(duì)較高ABA/GA3比例可促進(jìn)庫(kù)活性,利于同化物質(zhì)在籽粒庫(kù)中的轉(zhuǎn)化卸載。細(xì)胞分裂素類(lèi)物質(zhì)不參與籽粒庫(kù)活性的調(diào)節(jié),Morris等[17]研究表明,作物籽粒中高水平的細(xì)胞分裂素類(lèi)物質(zhì)Z+ZR與籽粒灌漿始期和中期的灌漿速率和灌漿百分比顯著相關(guān),徐云姬等[14]通過(guò)玉米強(qiáng)勢(shì)粒和弱勢(shì)粒的比較研究認(rèn)為,這種調(diào)節(jié)作用可能是通過(guò)調(diào)控胚乳細(xì)胞的分裂和增加庫(kù)容來(lái)實(shí)現(xiàn)的。從本研究的結(jié)果來(lái)看,各內(nèi)源激素含量在成功發(fā)育弱勢(shì)粒和敗育弱勢(shì)粒間皆無(wú)顯著差異,說(shuō)明在玉米籽粒形成期,內(nèi)源激素未參與SAI活性的調(diào)控;但從相關(guān)分析結(jié)果來(lái)看,各內(nèi)源激素含量與弱勢(shì)粒胚乳細(xì)胞分裂速率和灌漿速率顯著相關(guān)(未發(fā)表數(shù)據(jù)),這與徐云姬等[14]的研究結(jié)果一致,說(shuō)明Z+ZR、IAA、GA3和ABA在籽粒形成期主要影響籽粒庫(kù)容量,而與這一時(shí)期籽粒庫(kù)活性無(wú)關(guān),但在線性灌漿期是否參與調(diào)解其他與庫(kù)活性相關(guān)的酶類(lèi)活性有待于進(jìn)一步研究。SAI是灌漿初期轉(zhuǎn)化蔗糖并使其代謝產(chǎn)物向庫(kù)卸載的酶,是打開(kāi)籽粒庫(kù)使其接納光合物質(zhì)的鑰匙;在糖分卸載到籽粒當(dāng)中后,后續(xù)的籽粒淀粉合成中也有多種酶發(fā)揮作用,它們對(duì)庫(kù)活性的影響也會(huì)影響籽粒的最終建成。楊建昌等[15,29]通過(guò)研究水稻、玉米的灌漿動(dòng)態(tài),對(duì)比弱勢(shì)粒和強(qiáng)勢(shì)粒的多種酶活性發(fā)現(xiàn),在水稻和玉米灌漿活躍期,強(qiáng)勢(shì)粒中的焦磷酸化酶(AGPase)、淀粉合酶(StS)和淀粉分支酶(SBE)活性均顯著高于弱勢(shì)粒,但各種內(nèi)源激素是否參與這些酶活性的調(diào)節(jié)進(jìn)而影響籽粒灌漿充實(shí),值得進(jìn)一步深入探討。

圖6 玉米籽粒形成期弱勢(shì)粒多胺含量與SAI活性的關(guān)系Fig.6 Relationship between polyamine content and soluble acid invertase(SAI)of inferior kernels during the blister stage of maize縮寫(xiě)同圖2。Abbreviations are the same as those given in Fig.2.

圖7 玉米籽粒形成期弱勢(shì)粒多胺含量與乙烯釋放率的關(guān)系Fig.7 Relationship between polyamine content and ethylene release rate of inferior kernels during the blister stage of maize縮寫(xiě)同圖2。Abbreviations are the same as those given in Fig.2.

Yang等[5]對(duì)水稻弱勢(shì)粒的研究認(rèn)為,籽粒中多胺和乙烯的平衡關(guān)系影響糖分卸載關(guān)鍵酶的活性。多胺和乙烯有共同的合成前體 S-腺苷甲硫胺酸(SAM),SAM既可以在ACC合成酶作用下轉(zhuǎn)化為乙烯形成的前體物質(zhì)ACC,又可以在S-腺苷甲硫胺酸脫羧酶(SAMDC)及亞精胺合成酶和精胺合成酶的作用下,經(jīng)丙胺基轉(zhuǎn)移反應(yīng)依次形成亞精胺和精胺[29]。Feng等[21]研究認(rèn)為,乙烯和多胺的這種平衡關(guān)系可能受同化物質(zhì)供應(yīng)量的影響,糖分供應(yīng)充足時(shí),乙烯合成受到抑制,而促進(jìn)多胺的合成,進(jìn)而促進(jìn)了籽粒灌漿,但高水平多胺是通過(guò)什么影響籽粒灌漿則并不清楚。本研究則回答了這一問(wèn)題,試驗(yàn)結(jié)果表明,成功發(fā)育弱勢(shì)粒(IcP處理)和發(fā)育不良弱勢(shì)粒(CP處理)的乙烯釋放速率和多胺含量都有顯著差異,籽粒中多胺含量整體表現(xiàn)為IcP>CP,而乙烯釋放速率恰恰相反;相關(guān)分析表明,籽粒中SAI活性與多胺含量顯著正相關(guān),而與乙烯釋放率呈顯著負(fù)相關(guān),且多胺含量與乙烯釋放率也呈顯著負(fù)相關(guān),說(shuō)明乙烯生成會(huì)抑制多胺的合成,從而降低SAI的活性。具體而言,CP處理較高的乙烯含量間接抑制了多胺的合成,使SAI活性降低,進(jìn)而阻礙可溶性糖卸載,造成弱勢(shì)粒敗育或灌漿受限;而IcP處理較低的乙烯含量間接促進(jìn)了多胺的合成,較高的多胺含量使SAI活性提高,而較高的SAI活性則促進(jìn)了可溶性糖的卸載,從而使弱勢(shì)粒成功發(fā)育并灌漿充實(shí)。

實(shí)際上,本研究采用馮漢宇等[20]的方法創(chuàng)造的兩種不同發(fā)育狀態(tài)的弱勢(shì)粒,其根本原因仍是不同的物質(zhì)供應(yīng)水平最終導(dǎo)致了弱勢(shì)粒發(fā)育成功與否,因此將本研究結(jié)果與Feng等[21]和梁紅偉等[13]的結(jié)論綜合分析,可以基本明確不同物質(zhì)供應(yīng)水平通過(guò)影響乙烯、多胺的平衡,進(jìn)而影響SAI活性并最終導(dǎo)致弱勢(shì)粒發(fā)育成功與否的生理途徑(圖8)。在同化物質(zhì)供應(yīng)充足條件下,乙烯合成受到抑制,使更多的SAM在多胺酶等轉(zhuǎn)化酶作用下合成更多的多胺[28],從而提高了SAI活性,促進(jìn)籽粒內(nèi)還原糖的卸載,使籽粒成功建成并灌漿充實(shí);在同化物質(zhì)供應(yīng)不足的條件下,乙烯大量合成,而抑制了多胺合成,多胺含量降低進(jìn)一步影響了SAI活性,還原糖卸載受阻,造成籽粒灌漿受限或最終敗育。

圖8 乙烯、多胺平衡影響弱勢(shì)粒庫(kù)活性并主導(dǎo)弱勢(shì)粒發(fā)育的模式圖Fig.8 Theoretical framework of effects of polyamines and ethylene contents and their balance on sink activity and inferior kernels setting

4 結(jié)論

玉米籽粒形成期的弱勢(shì)粒庫(kù)活性即SAI活性受其內(nèi)源多胺和乙烯含量及其平衡關(guān)系調(diào)控,多胺促進(jìn)SAI活性,而乙烯抑制其活性。多胺和乙烯平衡受物質(zhì)供應(yīng)狀況影響,當(dāng)同化物質(zhì)供應(yīng)充足時(shí),多胺大量合成抑制乙烯釋放,從而促進(jìn)SAI活性使糖分向籽粒中卸載,使弱勢(shì)粒成功建成;當(dāng)同化物質(zhì)供應(yīng)不足時(shí),多胺合成受限而乙烯大量合成,抑制SAI活性,還原糖卸載受阻,籽粒敗育或灌漿受限。

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Relationship of Sink Activity with Endogenous Hormones and Polyamine Contents in Inferior Kernels of Maize

WANG Zhi-Gang1,**,LIANG Hong-Wei1,2,**,GAO Ju-Lin1,*,YU Xiao-Fang1,SUN Ji-Ying1,SU Zhi-Jun1, HU Shu-Ping1,YU Shao-Bo1,LI Ya-Jian1,WEI Shu-Li1,and YANG Zhe1

1College of Agronomy,Inner Mongolia Agricultural University,Hohhot 010019,China;2Inner Mongolia Academy of Agricultural and Animal Husbandry Sciences,Hohhot 010031,China

Sink activity at blister stage of maize is the primary limiting factor of the abortion and/or filling stagnation of inferior kernels.Clarifying the effect of endogenous hormone and polyamine contents on sink activity of inferior kernels in maize is of great importance for regulating inferior kernels in crowding maize colony.In this present study,two typical commercial maize hybrids Zhengdan 958 and Xianyu 335 were planted in 2014 and 2015.At silking stage,two pollination treatments,incomplete pollination(IcP)and complete pollination(CP),were imposed to each cultivar.The objective of incomplete pollination,in which the pollinated filaments within the basal region of ear was decreased by hand-cutting,was to stimulate the inferior kernel setting in the apical region of ear,where the inferior kernels should be aborted or filling-stagnated in complete pollination.The endogenous hormone and polyamine contents were compared between IcP and CP treatments during blister stage,and the relationship of sink activity with endogenous hormone and/or polyamine contents was analyzed.There was no remarkable effect of hybrids and years on sink activity,endogenous hormone and polyamine of kernels.The soluble acid invertase activity of IcPwas significantly higher than that of CP,by 13.5%on an average and 21.8%at the maximum.There was no significant difference in contents of Z+ZR,IAA,GA3,and ABA of the inferior kernels between IcP and CP treatments during blister stage.The polyamine content of IcP was significantly higher than that of CP,but the ethylene producing rate was just the opposite.SAI activity correlated with polyamine content positively,but with ethylene release rate negatively,moreover,ethylene release rate showed negative correlation with polyamine content.The results suggested that,during the blister stage of maize,the contents of Z+ZR,IAA,GA3,and ABA of inferior kernels have nothing to do with SAI activity.SAI activity of inferior kernels is mainly affected by polyamine and ethylene,which is promoted by polyamine content and inhibited by ethylene release rate.The balance between polyamine and ethylene determines the result of inferior kernel development,which is radically regulated by assimilate supply of maize plant.

Maize;Inferior kernel;Sink activity;Hormone;Polyamine

(

):2016-12-05;Accepted(接受日期):2017-04-20;Published online(網(wǎng)絡(luò)出版日期):2017-05-08.

10.3724/SP.J.1006.2017.01196

本研究由國(guó)家自然科學(xué)基金項(xiàng)目(31301265),國(guó)家重點(diǎn)研發(fā)計(jì)劃項(xiàng)目(2016YFD0300103),內(nèi)蒙古自治區(qū)高等學(xué)校青年科技英才支持計(jì)劃(NJYT-14-B04)和國(guó)家科技支撐計(jì)劃項(xiàng)目(2013BAD07B04)資助。

This study was supported by the National Natural Science Foundation of China(31301265),the State Program of China(2016YFD0300103), the Program for Young Talents of Science and Technology in Universities of Inner Mongolia Autonomous Region(NJYT-14-B04),and the National Key Technology Support Program of China(2013BAD07B04).

*通訊作者(Corresponding author):高聚林,E-mail:nmgaojulin@163.com**同等貢獻(xiàn)(Contributed equally to this work)

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