岳鵬莉,王晨陽(yáng),2,盧紅芳,劉衛(wèi)星,馬 耕,王 強(qiáng),胡陽(yáng)陽(yáng)

(1.河南農(nóng)業(yè)大學(xué)農(nóng)學(xué)院，河南鄭州 450002； 2.國(guó)家小麥工程技術(shù)研究中心，河南鄭州 450002)

灌漿期高溫干旱脅迫對(duì)小麥籽粒淀粉積累的影響

岳鵬莉1,王晨陽(yáng)1,2,盧紅芳1,劉衛(wèi)星1,馬 耕1,王 強(qiáng)1,胡陽(yáng)陽(yáng)1

(1.河南農(nóng)業(yè)大學(xué)農(nóng)學(xué)院，河南鄭州 450002； 2.國(guó)家小麥工程技術(shù)研究中心，河南鄭州 450002)

為探討花后高溫和干旱逆境脅迫對(duì)小麥淀粉組分的影響，采用盆栽和人工氣候室相結(jié)合的方式研究了灌漿期短暫高溫、干旱及其復(fù)合脅迫對(duì)兩個(gè)不同品質(zhì)類型小麥品種籽?？偟矸?、直鏈淀粉和支鏈淀粉含量的影響。結(jié)果表明，小麥籽粒淀粉的積累量符合Logistic方程，但逆境脅迫會(huì)縮短籽粒淀粉積累持續(xù)時(shí)間，降低其積累速率，使其最終淀粉積累量減少。與對(duì)照相比，高溫、干旱及其復(fù)合脅迫顯著降低兩個(gè)小麥品種籽粒的支鏈淀粉和總淀粉含量，直鏈淀粉含量受影響較小，淀粉的直/支比增加。高溫、干旱及其復(fù)合脅迫下，洛麥24的千粒重分別較對(duì)照下降42.03%、21.95%和50.01%，產(chǎn)量分別較對(duì)照下降57.88%、40.75%和61.08%；鄭麥366的千粒重分別較對(duì)照下降34.39%、7.64%和43.16%，產(chǎn)量分別較對(duì)照下降41.58%、30.97%和48.49%。綜上所述，高溫、干旱脅迫抑制支鏈淀粉積累是造成產(chǎn)量下降的重要原因；復(fù)合脅迫對(duì)小麥籽粒淀粉積累量的影響大于干旱或高溫單獨(dú)脅迫；高溫脅迫的影響大于干旱；洛麥24較鄭麥366對(duì)高溫和干旱脅迫更敏感。

小麥；高溫；干旱；淀粉；產(chǎn)量

淀粉是小麥籽粒的重要組成部分，占整個(gè)籽粒干重的 65%～75%[1]。小麥籽粒中的淀粉按葡萄糖分子的鏈接方式分為直鏈和支鏈淀粉兩部分。研究表明，淀粉總含量及其組分對(duì)小麥加工品質(zhì)與面食評(píng)分均有明顯影響[2-4]。在小麥生育過(guò)程中，開(kāi)花至成熟是小麥籽粒與品質(zhì)形成的關(guān)鍵階段，該期間遭受高溫或干旱脅迫會(huì)降低小麥產(chǎn)量，并導(dǎo)致籽粒品質(zhì)變劣。在實(shí)際生產(chǎn)上，我國(guó)北方冬小麥生育后期常遭遇極端的短期高溫或干旱，而且高溫與干旱常相伴疊發(fā)，導(dǎo)致小麥產(chǎn)量下降，并影響面粉加工與食用品質(zhì)[5-7]。研究證明，小麥籽粒灌漿期遭受極端高溫時(shí)，籽粒淀粉合成酶活性下降，積累時(shí)間縮短，淀粉組分改變[8]，而且不同品種對(duì)高溫脅迫的響應(yīng)存在差異[9]。另有研究表明，小麥灌漿期遭受干旱，籽?？偟矸酆亢椭ф湹矸酆匡@著降低，淀粉直/支比改變，導(dǎo)致品質(zhì)變化[10]。戴廷波等[11]研究表明，在高溫與干旱雙重脅迫下，高溫對(duì)小麥的影響大于干旱，且二者存在互作效應(yīng)。

目前，圍繞小麥生育后期單一逆境危害(如高溫或干旱)協(xié)迫對(duì)小麥產(chǎn)量、產(chǎn)量構(gòu)成因素、面粉及加工品質(zhì)的影響已有不少研究報(bào)道[12-13]，但有關(guān)多因素復(fù)合危害的研究相對(duì)較少。由于在我國(guó)北方麥區(qū)小麥生育后期，短期極端高溫天氣與干旱常同時(shí)發(fā)生，關(guān)于二者復(fù)合脅迫對(duì)小麥產(chǎn)量和品質(zhì)影響的研究相對(duì)較少。本試驗(yàn)采用盆栽結(jié)合人工氣候室進(jìn)行小麥生育后期逆境脅迫的模擬，研究小麥灌漿期高溫、干旱及其復(fù)合脅迫對(duì)小麥籽粒淀粉積累量和產(chǎn)量的影響，以期為小麥抗逆調(diào)優(yōu)栽培提供理論依據(jù)。

1 材料與方法

1.1 試驗(yàn)材料與設(shè)計(jì)

試驗(yàn)選用中筋小麥洛麥24(蛋白質(zhì)含量14%，濕面筋含量30%)和強(qiáng)筋小麥鄭麥366(蛋白質(zhì)含量16%，濕面筋含量33%)為供試材料，于2014-2015年度以盆栽方式在河南農(nóng)業(yè)大學(xué)科教試驗(yàn)園區(qū)進(jìn)行，采用兩因素(品種和脅迫)裂區(qū)設(shè)計(jì)，其中，品種為主區(qū)，脅迫為副區(qū)，設(shè)干旱、高溫和復(fù)合脅迫，以大田生長(zhǎng)為對(duì)照。每處理10盆。

試驗(yàn)用盆高為27 cm，盆口直徑為24 cm。試驗(yàn)用土為耕層土壤，其有機(jī)質(zhì)含量為18 g·kg-1，全氮和堿解氮含量分別為1 g·kg-1和58 mg·kg-1，速效磷含量為68 mg·kg-1，速效鉀含量為205 mg·kg-1，pH值為8，田間持水量為26%，每盆土重10 kg。于2014年10月18日播種，播種前每盆施純氮1.15 g、P2O51.35 g、K2O 1.15 g作為基肥，三葉期每盆定苗12株；拔節(jié)期結(jié)合澆水每盆追施尿素2.5 g。脅迫處理(花后10 d)以前，各盆栽在大田條件下生長(zhǎng)，定期澆水保持土壤相對(duì)含水量在75%左右。

脅迫處理期間，11:00-16:00對(duì)照的平均溫度為29.32 ℃。干旱處理(DS)于高溫處理前7 d進(jìn)行遮雨控制水分，每天隨機(jī)選取10 盆采用稱重法測(cè)定土壤相對(duì)含水量，并于高溫處理時(shí)達(dá)到55%的輕度干旱水平，高溫處理結(jié)束后恢復(fù)正常供水。于花后10 d進(jìn)行高溫處理(HT)，將長(zhǎng)勢(shì)均勻一致的供試盆栽移到人工氣候室，38 ℃高溫處理4 d，每天處理5 h(11:00-16:00)，采用稱重法控制土壤相對(duì)含水量在75%左右。高溫+干旱復(fù)合脅迫(HT+DS)為干旱處理同時(shí)進(jìn)行高溫脅迫處理。處理結(jié)束后將全部材料轉(zhuǎn)移至大田，生長(zhǎng)至成熟。于開(kāi)花期選取同一天開(kāi)花、長(zhǎng)勢(shì)均勻一致的麥穗進(jìn)行標(biāo)記。

1.2 測(cè)定項(xiàng)目與方法

于高溫、干旱處理前和結(jié)束后，各處理隨機(jī)取10個(gè)已標(biāo)記穗子，之后每4 d取樣一次直至成熟。穗子取回立即剝?nèi)⌒←溩蚜＃?05 ℃殺青20 min， 60 ℃烘至恒重。烘干籽粒用高速萬(wàn)能粉碎機(jī)(天津泰斯特儀器有限公司)磨粉，用于直鏈和支鏈淀粉含量的測(cè)定[14]，總淀粉含量為兩者之和。以花后天數(shù)(t)作為自變量，每次測(cè)得的淀粉含量作為依變量(Y)，用Logistic方程Y=K/(1+eA+Bt) 來(lái)擬合小麥籽粒總淀粉、直鏈淀粉和支鏈淀粉的積累過(guò)程[15]，其中，K為理論最大淀粉積累量，A、B為其參數(shù)，通過(guò)對(duì)方程一階求導(dǎo)，可得到淀粉積累速率方程Vt=KeA+Bt/(1+eA+Bt)。以此推導(dǎo)出淀粉積累特征參數(shù)：淀粉積累起始勢(shì)C0=K/(1+eA)(反映淀粉積累潛勢(shì))，積累活躍生長(zhǎng)期D=[ln(1/9)-A]/B(達(dá)到總積累量的90%所經(jīng)歷的時(shí)間)，積累速率達(dá)到最大時(shí)的日期Tmax=-A/B，最大積累速率Rmax=-K×B/4，平均積累速率Rmean=K/D。

成熟期每處理選取3盆計(jì)產(chǎn)、考種。

1.3 數(shù)據(jù)處理

數(shù)據(jù)整理及作圖采用Microsoft Excel進(jìn)行，用 SPSS 19.0 對(duì)數(shù)據(jù)進(jìn)行方差分析。

2 結(jié)果與分析

2.1 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)成熟期籽粒淀粉含量的影響

由表1可見(jiàn)，干旱、高溫及其復(fù)合脅迫均顯著降低了小麥籽粒支鏈淀粉和總淀粉含量，對(duì)直鏈淀粉含量的影響相對(duì)較小，使淀粉直/支比顯著增加(鄭麥366干旱處理除外)。干旱、高溫及其復(fù)合脅迫下，洛麥24直鏈淀粉含量分別較對(duì)照下降4.57%、11.95%和14.77%，后兩者與對(duì)照的差異顯著，支鏈淀粉含量分別較對(duì)照下降19.50%、30.72%和37.35%，差異均顯著，總淀粉含量分別較對(duì)照下降15.51%、25.70%和31.31%，差異均顯著；鄭麥366直鏈淀粉含量分別較對(duì)照下降3.01%、7.53%和10.92%，后者與對(duì)照的差異顯著，支鏈淀粉含量分別較對(duì)照下降17.18%、28.93%和30.13%，差異均顯著，總淀粉含量分別較對(duì)照下降13.68%、23.65%和25.39%，差異均顯著。結(jié)果表明，高溫脅迫對(duì)2個(gè)小麥品種淀粉含量的影響明顯大于干旱脅迫，且復(fù)合脅迫具有顯著的疊加效應(yīng)；干旱、高溫脅迫對(duì)中筋品種洛麥24淀粉含量的影響較大。

表1 灌漿期干旱、高溫及其復(fù)合脅迫下小麥籽粒的淀粉含量

Table 1 Effect of heat and drought stress and their combination on starch in wheat grains during grain filling stage

品種Cultivar處理Treatment直鏈淀粉Amylose支鏈淀粉Amylopectin總淀粉Totalstarch直/支Amylose/Amylopectin洛麥24 Luomai24CK17.74a48.61a66.35a0.37cDS16.93ab39.13b56.06b0.43bHT15.62b33.68c49.30c0.46bHT+DS15.12b30.45d45.58d0.50a鄭麥366 Zhengmai366CK17.12a47.26a64.37a0.36bDS16.60ab43.28b59.88b0.38bHT15.83ab37.14c52.97c0.42aHT+DS15.25b36.51c51.76c0.43a

CK:對(duì)照；DS：干旱；HT：高溫。同列數(shù)據(jù)后不同字母表示相同品種不同處理間差異顯著(P<0.05)。下同。

CK：Control;DS:Drought stress;HT:Hight temperature.Different letters following values at the same column mean significant difference among treatments for the same cultivar(P<0.05).The same below.

2.2 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)籽粒直鏈淀粉積累動(dòng)態(tài)的影響

由圖1和表2可見(jiàn)，小麥籽粒直鏈淀粉含量在花后10～26 d急劇增加，26 d后增加趨緩，于成熟期達(dá)最大值，其積累過(guò)程符合Logistic方程Y=K(1+eA+Bt)(R2均大于0.95)。干旱、高溫及其復(fù)合脅迫下，洛麥24的K值與對(duì)照相比分別減少3.96%、10.73%和11.54%，Rmean分別減少5.26%、10.78%和16.53%；鄭麥366的K值分別較對(duì)照減少2.93%、6.27%和9.39%，Rmean分別較對(duì)照減少7.45%、12.06%和15.90%。高溫及復(fù)合脅迫下淀粉積累提前結(jié)束，灌漿期縮短4 d?？梢?jiàn)，高溫脅迫對(duì)小麥籽粒淀粉積累的影響大于干旱，高溫、干旱復(fù)合脅迫具有明顯的疊加效應(yīng)；2個(gè)品種相比，洛麥24受干旱、高溫脅迫影響較大。

表2 灌漿期干旱、高溫及其復(fù)合脅迫下小麥籽粒直鏈淀粉積累的特征參數(shù)

Table 2 Characteristic parameters of amylose accumulation in wheat grains under heat and drought stress and their combination

品種Cultivar處理TreatmentR2K/%ABTmax/dRmax/(%·d-1)Rmean/(%·d-1)C0D/d洛麥24CK0.992917.914.061-0.25116.21.1261.2830.303414.0Luomai24DS0.986617.203.933-0.24116.31.0351.2160.330214.1HT0.984615.993.997-0.24716.20.9891.1450.288314.0HT+DS0.985315.843.805-0.22417.00.8871.0710.345014.8鄭麥366CK0.992817.314.414-0.26716.51.1551.2080.207114.3Zhengmai366DS0.990716.804.369-0.25417.21.0651.1180.210015.0HT0.970716.224.538-0.26017.51.0531.0620.171615.3HT+DS0.963615.684.449-0.25217.60.9891.0160.181215.4

K：生長(zhǎng)終值量；A、B：參數(shù)；C0：積累起始勢(shì)；D：積累活躍生長(zhǎng)期；Tmax：積累速率最大期；Rmax：最大積累速率；Rmean：平均積累速率。表3、表4同。

K：Final amount of growth；A、B：Parameter；C0：Initial potential of accumulation；D：Active growth phase of accumulation；Tmax：Date of maximum accumulation；Rmax：Maximum accumulation rate；Rmean：Average accumulation rate.The same as in table 3 and table 4.

CK0、DS0、HT0、(HT+DS)0:測(cè)量值; CK1、DS1、HT1、(HT+DS)1:理論值; 下同。

CK0、DS0、HT0、(HT+DS)0:Measurement value; CK1、DS1、HT1、(HT+DS)1:Theoretical value;The same as in Fig.2 and Fig.3.

圖1 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)小麥籽粒直鏈淀粉積累動(dòng)態(tài)的影響

Fig.1 Effect of heat and drought stress and their combination on the content of amylose in wheat grains during grain filling stage

2.3 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)小麥籽粒支鏈淀粉積累動(dòng)態(tài)的影響

從圖2和表3可以看出，花后支鏈淀粉含量不斷上升，花后10～26 d增加較快，其積累過(guò)程符合Logistic方程Y=K(1+eA+Bt)(R2均大于0.95)。干旱、高溫及其復(fù)合脅迫下，洛麥24的K值分別較對(duì)照下降19.53%、32.77%和36.69%，Rmean分別下降15.90%、20.95%和26.63%。而鄭麥366的K值較分別較對(duì)照下降6.48%、19.69%和21.54%，Rmean分別下降9.13%、13.03%和16.88%。說(shuō)明高溫對(duì)籽粒支鏈淀粉積累的影響較干旱大，且高溫干旱復(fù)合脅迫有明顯的疊加效應(yīng)。兩品種相比，中筋品種洛麥24對(duì)逆境脅迫更為敏感。

2.4 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)小麥籽?？偟矸鄯e累動(dòng)態(tài)的影響

由圖3可見(jiàn)，花后10～26 d籽?？偟矸酆垦杆僭黾樱腋魈幚黹g差異逐漸增大，花后26 d后增加趨于緩慢，并于成熟期達(dá)到峰值。Logistic方程Y=K(1+eA+Bt)可較好地?cái)M合總淀粉含量的積累過(guò)程(表4)。由總淀粉積累特征參數(shù)可以看出，對(duì)于洛麥24而言，K值在干旱、高溫及其復(fù)合脅迫下分別較對(duì)照下降15.42%、26.36%和29.70%；Rmean值分別較下降12.21%、16.92%和22.48%；鄭麥366K值在干旱、高溫及其復(fù)合脅迫下分別較對(duì)照下降6.38%、16.35%和18.02%；Rmean值分別下降8.47%、11.85%和15.86%。干旱、高溫脅迫對(duì)總淀粉積累的影響表現(xiàn)為高溫干旱復(fù)合脅迫>高溫>干旱；兩品種相比，洛麥24所受影響較鄭麥366大。

表3 灌漿期干旱、高溫及其復(fù)合脅迫下小麥籽粒支鏈淀粉積累特征參數(shù)

Table 3 Characteristic parameters of amylopectin accumulation in wheat grains under heat and drought stress and their combination

品種Cultivar處理TreatmentR2K/%ABTmax/dRmax/(%·d-1)Rmean/(%·d-1)C0D/d洛麥24CK0.993153.454.074-0.19121.32.5552.7970.893619.1Luomai24DS0.994143.013.611-0.17620.51.8962.3521.131418.3HT0.985335.934.162-0.22618.52.0262.2110.550916.3HT+DS0.950533.843.721-0.19918.71.6842.0520.800216.5鄭麥366CK0.991553.733.982-0.18022.12.4162.6950.984019.9Zhengmai366DS0.983450.254.005-0.17622.72.2152.4490.899220.5HT0.960543.154.093-0.19920.62.1432.3440.708018.4HT+DS0.963742.164.328-0.20621.02.1712.2400.548818.8

圖2 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)小麥籽粒支鏈淀粉積累動(dòng)態(tài)的影響

Table 4 Characteristic parameters of total starch accumulation in wheat grains under heat and drought stress and their combination

品種Cultivar處理TreatmentR2K/%ABTmax/dRmax/(%·d-1)Rmean/(%·d-1)C0D/d洛麥24CK0.997470.283.96-0.2019.73.544.021.313317.5Luomai24DS0.996259.443.64-0.1919.02.853.531.515116.8HT0.988751.754.08-0.2317.72.993.340.860715.5HT+DS0.968349.403.75-0.2118.02.573.121.138015.8鄭麥366CK0.994369.293.97-0.2020.23.403.841.281918.1Zhengmai366DS0.990364.874.02-0.1920.73.153.511.149018.5HT0.972957.964.20-0.2219.33.153.380.856917.1HT+DS0.971256.814.32-0.2219.83.103.230.745617.6

圖3 灌漿期干旱、高溫及其復(fù)合脅迫對(duì)小麥籽?？偟矸鄯e累動(dòng)態(tài)的影響

2.5 灌漿期高溫、干旱及其復(fù)合脅迫對(duì)小麥籽粒產(chǎn)量的影響

從表5可見(jiàn)，灌漿期干旱、高溫及其復(fù)合脅迫下，兩個(gè)品種的穗粒重均顯著下降，從而使小麥的千粒重和產(chǎn)量顯著降低。干旱、高溫及其復(fù)合脅迫下，洛麥24千粒重分別較對(duì)照下降21.97%、42.03%和50.09%，產(chǎn)量(每盆)分別下降40.75%、57.88%和61.08%；與對(duì)照相比，鄭麥366千粒重分別下降7.62%、34.39%和43.16%，產(chǎn)量分別下降31.49%、43.71%和50.96%。可見(jiàn)，洛麥24受高溫脅迫的影響大于鄭麥366；不同處理間比較，高溫脅迫對(duì)產(chǎn)量的影響大于干旱脅迫，且復(fù)合脅迫具有一定的疊加效應(yīng)。

表5 灌漿期高溫、干旱及其復(fù)合脅迫對(duì)小麥粒重及產(chǎn)量的影響

Table 5 Effect of heat and drought stress and their combination on 1 000-grain weight and yield

品種Cultivar處理Treatment每盆穗數(shù)Spikesperpot千粒重1000-grainweight/g穗粒重Grainweightperspike/g每盆產(chǎn)量Grainyieldperpot/g洛麥24Luomai24CK39.33a37.45a1.12a48.84aDS36.50a29.23b0.78b28.94bHT37.67a21.71c0.60c20.57cHT+DS38.00a18.69c0.42d19.01c鄭麥366Zhengmai366CK39.25a38.88a1.20a48.47aDS38.00a35.91b1.08b33.46bHT37.33a25.51c0.83c27.49cHT+DS39.25a22.10d0.67d23.95d

3 討 論

作為小麥籽粒的重要成分，淀粉含量與組成對(duì)面粉及面食加工品質(zhì)有很大的影響[16]。在小麥籽粒形成與灌漿過(guò)程中，環(huán)境因素諸如高溫、干旱均會(huì)影響籽?？偟矸酆?、直鏈淀粉含量和支鏈淀粉含量，并最終決定了淀粉性狀的優(yōu)劣。有研究表明，就對(duì)淀粉組分的影響看，逆境脅迫對(duì)支鏈淀粉影響更大[11]。小麥籽粒灌漿階段最適宜的溫度為20～24 ℃，超過(guò)30 ℃的高溫則會(huì)使淀粉積累受抑[17-18]。研究發(fā)現(xiàn)，高溫和干旱降低了花后物質(zhì)的同化量，從而降低了粒重和淀粉含量[19]，且高溫與干旱脅迫對(duì)小麥籽粒淀粉的影響具有互作或疊加效應(yīng)[20]。Logistic方程模擬表明，高溫及干旱脅迫下小麥籽粒淀粉積累時(shí)間縮短、積累速率降低，這是導(dǎo)致直鏈、支鏈淀粉最終積累量降低的主要原因；最大積累速率的出現(xiàn)時(shí)間和積累持續(xù)期的調(diào)節(jié)作用甚微。可見(jiàn)淀粉最終積累量的多寡主要由積累速率的高低和積累持續(xù)期決定。說(shuō)明逆境脅迫對(duì)小麥淀粉最終積累量的影響可能是通過(guò)調(diào)控直鏈、支鏈淀粉積累時(shí)間和積累速率來(lái)實(shí)現(xiàn)的。本研究表明，灌漿期高溫、干旱及復(fù)合脅迫均使小麥籽?？偟矸酆椭ф湹矸酆匡@著下降，但對(duì)直鏈淀粉的影響較小，從而導(dǎo)致直/支比升高。同時(shí)，高溫對(duì)淀粉含量積累的影響大于干旱脅迫，且復(fù)合脅迫影響最大，表現(xiàn)出高溫和干旱的顯著疊加效應(yīng)。

在小麥生長(zhǎng)發(fā)育過(guò)程中，溫度和水分是關(guān)鍵的生態(tài)因子，對(duì)籽粒灌漿進(jìn)程及籽粒品質(zhì)和產(chǎn)量形成有著極大影響。土壤水分的虧缺可以加速籽粒灌漿進(jìn)程，縮短灌漿持續(xù)時(shí)間[21]，而在小麥生育后期發(fā)生干旱脅迫則會(huì)引起植株過(guò)早衰老、生育時(shí)期縮短，從而使粒重和產(chǎn)量顯著降低[22-23]。高溫對(duì)小麥產(chǎn)量的影響主要是降低粒重[24-25]。有研究表明，灌漿期高溫降低了小麥葉片的光合能力，使葉光合同化效率下降，從而使籽粒中光合產(chǎn)物的積累量嚴(yán)重降低，并明顯縮短胚乳細(xì)胞分裂期，使小麥花后籽粒淀粉積累時(shí)間縮短，進(jìn)而導(dǎo)致籽粒充實(shí)不良，最終導(dǎo)致粒重顯著降低[12,26-27]。本試驗(yàn)中，高溫、干旱及其復(fù)合脅迫均使兩個(gè)品種的穗粒重降低，從而使千粒重和產(chǎn)量下降。灌漿期高溫和干旱脅迫對(duì)小麥粒重和產(chǎn)量的影響與淀粉積累受抑密切相關(guān)[28-29]。本研究表明，高溫、干旱及其復(fù)合脅迫均使小麥籽粒下降，其中高溫與干旱具有顯著的疊加效應(yīng)，且不同品種間存在較大差異，中筋小麥洛麥24產(chǎn)量受逆境脅迫抑制較大，更明確的生理機(jī)制還有待更深入研究。

[1]DALE E M,HOUSLEY T L.Sucrose synthase activity in developing wheat endosperms differing in maximum weigh [J].PlantPhysiology,1986,82:7.

[2] 姚大年,劉廣田.淀粉理化特性、遺傳規(guī)律及小麥淀粉與品質(zhì)的關(guān)系[J].糧食與飼料工業(yè),1997,20(2):36.

YAO D N,LIU G T.Physiochemical characteristics and genetics laws of starch as well as the relationship of wheat starch to qualities [J].CerealandFoodIndustry,1997,20(2):36.

[3] ODA M,YASUDA Y,OKAZAKI S,etal.A method of flour quality assessment for Japanese noodles [J].CerealChemistry,1980,57(4):253.

[4] TOYOKAWA H,RUBENTHALER G L,POWERS J R,etal.Japanese noodle quality,I.Flour components [J].CerealChemistry,1989,66(4):382.

[5] 田良才,張明儀,李晉川.高溫脅迫-小麥超高產(chǎn)的主要障礙[J].山西農(nóng)業(yè)科學(xué),1995,23(2):14.

TIAN L C,ZHANG M Y,LI J C.High temperature stress-main barrier of high productivity on wheat [J].JournalofShanxiAgriculturalSciences,1995,23(2):14.

[6] 鄒 琦.小麥的高溫傷害與高溫適應(yīng)[J].植物學(xué)報(bào),1987,30(4):387.

ZOU Q.Heat injury and heat acclimation in wheat plants [J].JournalofIntegrativePlantBiology,1987,30(4):387.

[7] 張平平,何中虎,夏先春,等.高溫脅迫對(duì)小麥蛋白質(zhì)和淀粉品質(zhì)影響的研究進(jìn)展[J].麥類作物學(xué)報(bào),2005,25(5):129.

ZHANG P P,HE Z H,XIA X C,etal.Effect of heat stress on wheat protein and starch quality [J].JournalofTriticeaeCrops,2005,25(5):129.

[8] JENNER C F.The physiology of starch and protein deposition in the endosperm of wheat [J].AustralianJournalofPlantPhysiology,1991,18:211.

[9] 閆素輝,尹燕枰,李文陽(yáng),等.灌漿期高溫對(duì)小麥籽粒淀粉的積累、粒度分布及相關(guān)酶活性的影響[J].作物學(xué)報(bào),2008,34(6):1092.

YAN S H,YIN Y P,LI W Y,etal.Effect of high temperature during grain filling on starch accumulation,starch granule distribution,and activities of related enzymes in wheat grains [J].ActaAgronomicaSinica,2008,34(6):1092.

[10] 許振柱,于振文,張永麗.土壤水分對(duì)小麥籽粒淀粉合成和積累特性的影響[J].作物學(xué)報(bào),2003,29(4):595.

XU Z Z,YU Z W,ZHANG Y L,etal.The effects of soil moisture on grain starch synthesis and accumulation of winter wheat [J].ActaAgronomicaSinica,2003,29(4):595.

[11] 戴廷波,趙 輝,荊 奇,等.灌漿期高溫和水分逆境對(duì)冬小麥籽粒蛋白質(zhì)和淀粉含量的影響[J].生態(tài)學(xué)報(bào),2006,26(11):3670.

DAI T B,ZHAO H,JING Q,etal.Effects of high temperature and water stress during grain filling on grain protein and starch formation in winter wheat [J].ActaEcologicaSinica,2006,26(11):3670.

[12] 王 玨,封超年,郭文善,等.花后高溫脅迫對(duì)小麥籽粒淀粉積累及晶體特性的影響[J].麥類作物學(xué)報(bào),2008,28(2):260.

WANG Y,FENG C N,GUO W S,etal.Effects of high temperature after anthesis on starch traits of grain in wheat [J].JournalofTriticeaeCrops,2008,28(2):260.

[13] 谷艷芳,丁圣彥,高志英,等.干旱脅迫下冬小麥光合產(chǎn)物分配格局及其與產(chǎn)量的關(guān)系[J].生態(tài)學(xué)報(bào),2010,30(5):1167.

GU Y F,DING S Y,GAO Z Y,etal.The pattern of photosynthate partitioning in drought-stressed winter wheat and its relationship with yield [J].ActaEcologicaSinica,2010,30(5):1167.

[14] 何照范.糧油籽粒品質(zhì)及其分析技術(shù)[M].北京:農(nóng)業(yè)出版社,1985:290.

HE Z F.Analysis Technique for Grain Quality in Cereals and Oils [M].Beijing:Agriculture Press,1985:290.

[15] 李世清,邵明安,李紫燕,等.小麥籽粒灌漿特征及影響因素的研究進(jìn)展[J].西北植物學(xué)報(bào),2003,23(11):2031.

LI S Q,SHAO M A,LI Z Y,etal.Review of characteristics of wheat grain fill and factors to influence it [J].ActaBotanicaBoreali-OccidentaliaSinica,2003,23(11):2031.

[16] 杜 朝,楊學(xué)舉,劉桂茹,等.小麥面粉淀粉特性與烘烤品質(zhì)關(guān)系的研究[J].河北農(nóng)業(yè)大學(xué)學(xué)報(bào),2002,25(4):29.

DU C,YANG X J,LIU G R,etal.Studies on the relations between starch properties of wheat flour and baking quality [J].JournalofAgriculturalUniversityofHebei,2002,25(4):29.

[17] 劉 萍,郭文善,浦漢春,等.灌漿期短暫高溫對(duì)小麥淀粉形成的影響[J].作物學(xué)報(bào),2006,32(2):182.

LIU P,GUO W S,PU H C,etal.Effects of transient high temperature during grain filling period on starch formation in wheat [J].ActaAgronomicaSinica,2006,32(2):182.

[18] GUEDIRA M,PAULSEN G M.Accumulation of starch in wheat grain under different shoot/root temperatures during maturation [J].FunctionalPlantBiology,2002,29:495.

[20] NAKAMURA Y,YUKI K,PARK S Y.Carbohydrate metabolism in the developing endosperm of rice grains [J].PlantCellPhysiology,1989,56:833.

[19] 苗建利,王晨陽(yáng),郭天財(cái),等.高溫與干旱互作對(duì)兩種筋力小麥品種籽粒淀粉及其組分含量的影響[J].麥類作物學(xué)報(bào),2008,28(2):254.

MIAO J L,WANG C Y,GUO T C,etal.Drought stresses on content and composition of grain starch in two wheat cultivars with different gluten strength [J].JournalofTriticeaeCrops,2008,28(2):254.

[21] 吳少輝,高海濤,王書(shū)子,等.干旱對(duì)冬小麥粒重形成的影響及灌漿特性分析[J].干旱地區(qū)農(nóng)業(yè)研究,2002,20(2):49.

WU S H,GAO H T,WANG S Z,etal.Analysis on the effect of drought on the grain weight grow and the character of the grain filling of winter wheat [J].AgriculturalResearchintheAridAreas,2002,20(2):49.

[22] 王晨陽(yáng),馬冬云,朱云集,等.冬小麥不同水氮運(yùn)籌對(duì)面條煮制品質(zhì)的影響[J].中國(guó)農(nóng)業(yè)科學(xué),2004,37(2):256.

WANG C Y,MA D Y,ZHU Y J,etal.Effects of different irrigation and nitrogen application regimes in winter wheat on cooking qualities of Chinese noodle [J].ScientiaAgriculturaSinica,2004,37(2):256.

[23] 王晨陽(yáng),郭天財(cái),彭 羽,等.花后灌水對(duì)小麥籽粒品質(zhì)性狀及產(chǎn)量的影響[J].作物學(xué)報(bào),2004,30(10):1031.

WANG C Y,GUO T C,PENG Y,etal.Effects of post-anthesis irrigation on grain quality indices and yield in winter [J].ActaAgronomicaSinica,2004,30(10):1031.

[24] KOPOBHH A H.Effects of low and high temperature during grain filling period on yield in spring wheat [J].OverseasAgronomyTriticeaeCrops,1983,5:35.

[25] 敬海霞,王晨陽(yáng),左學(xué)玲,等.花后高溫脅迫對(duì)小麥籽粒產(chǎn)量和蛋白質(zhì)含量的影響[J].麥類作物學(xué)報(bào),2010,30(3):459.

JING H X,WANG C Y,ZUO X L,etal.Effect of post-anthesis high temperature stress on grain yield and protein content of different wheat cultivars [J].JournalofTriticeaeCrops,2010,30(3):459.

[26]郭文善,施勁松,彭永欣,等.灌漿期高溫對(duì)小麥光合產(chǎn)物運(yùn)轉(zhuǎn)的影響[J].核農(nóng)學(xué)報(bào),1998,12(1):21.

GUO W S,SHI J S,PENG Y X,etal.Effect of high temperature on transportation of assimilate from wheat flag leaf during grain filling stage [J].JournalofNuclearAgriculturalSciences,1998,12(1):21.

[27] DON C,LOOKHART G,NAEEM H,etal.Heat stress and genotype affect the gluten in particles of the gluten in macropolymer-gel fraction [J].JournalofCerealSciences,2005,42:69.

[28] 范雪梅,姜 東,戴廷波,等.花后干旱和漬水對(duì)不同品質(zhì)類型小麥籽粒品質(zhì)形成的影響[J].植物生態(tài)學(xué)報(bào),2004,28(5):680.

FAN X M,JIANG D,DAI T B,etal.Effects of post-anthesis drought and waterlogging on the quality of grain formation in different wheat varieties [J].ChineseJournalofPlantEconogy,2004,28(5):680.

[29] JENNER C F.The physiology of starch and protein deposition in the endosperm of wheat [J].AustralianJournalofPlantPhysiology,1991,18:211.

Effect of Heat and Drought Stress on Starch Accumulation During Grain Filling Stage

YUE Pengli1,WANG Chenyang1,2,LU Hongfang1,LIU Weixing1,MA Geng1,WANG Qiang1,HU Yangyang1

(1.College of Agriculture,Henan Agriculture University,Zhengzhou,Henan 450002,China;2.National Engineering Research Center for Wheat,Zhengzhou,Henan 450002,China)

High temperature(HT),drought stress(DS) and their combination(HT+DS) after anthesis affected the starch concentration and its components as well as the starch characteristics significantly in wheat grains. A pot experiment was performed at the scientific & educational station of Henan agricultural university in 2014-2015 wheat growing season. Luomai 24 and Zhengmai 366,wheat cultivars with different gluten-strengths,were grown in 24 cm pots with 10 kg loam soil. The experiment was arranged with a completely randomized design with three replicates. Plants were initially grown in the field environment and then transferred to a greenhouse for HT treatment. HT treatment began from the 10th day to 13th day after anthesis with a high temperature of 38 ℃ from 11:00 am to 16:00 pm,and after that pots were returned to the field environment. The results showed that the starch accumulation amount in wheat grain depended on the accumulation duration and rate. HT,DS and HT+DS all caused decrease of both starch accumulation duration and accumulation rate,which resulted in the decrease of final starch accumulation quantity. Compared with the CK,the concentration of amylopectin and total starch were all decreased significantly under stress treatments,the amylose was less sensitive to the stress treatments,which led to an increase in the ratio of amylose to amylopectin.Compared with the CK,HT,DS and HT+DS decreased the 1 000-grain weight of Luomai 24 by 42.03%,21.95% and 50.01%,respectively; and decreased grain yield per pot by 57.88%,40.75% and 61.08%,respectively. As for Zhengmai 366,HT,DS and HT+DS decreased 1 000-grain weight by 34.39%,7.64%,43.16%,and declined grain yield per pot by 41.58%,30.97% and 48.49%,respectively. Consequently,the inhibition of starch accumulation was one of the important influence factors for the decline of wheat grain weight,and which affected the wheat grain yield ultimately.HT treatments had a greater effect than DS on the starch quality,Luomai 24 was more sensitive to HT and DS than Zhengmai 366.

Wheat; High temperature; Drought stress; Starch; Grain yield

時(shí)間：2016-11-04

2016-03-30

2016-04-21

國(guó)家科技支撐計(jì)劃項(xiàng)目(2015BAD26B01,2013BAC09B01)；河南省小麥產(chǎn)業(yè)技術(shù)體系崗位專家項(xiàng)目(S2010-01-G07)；河南省教育廳科學(xué)技術(shù)研究重點(diǎn)項(xiàng)目(14A210012)

E-mail: ypl1412@163.com

王晨陽(yáng)(E-mail: xmzxwang@163.com)

S512.1；S311

A

1009-1041(2016)11-1489-08

網(wǎng)絡(luò)出版地址：http://www.cnki.net/kcms/detail/61.1359.S.20161104.0925.022.html