小麦籽粒灌浆与脱水特性

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朱冬梅, 王慧, 刘大同, 高德荣, 吕国锋, 王君婵, 高致富, 陆成彬^,江苏里下河地区农业科学研究所/农业部长江中下游小麦生物学与遗传育种重点实验室,江苏扬州 225007

Characteristics of Grain Filling and Dehydration in Wheat

ZHU DongMei, WANG Hui, LIU DaTong, GAO DeRong, Lü GuoFeng, WANG JunChan, GAO ZhiFu, LU ChengBin^,Lixiahe Institute of Agricultural Sciences/Key Laboratory of Wheat Biology and Genetic Breeding in the Middle and Lower Yangtze River, Ministry of Agriculture, Yangzhou 225007, Jiangsu

通讯作者: 陆成彬,E-mail：lcb@wheat.org.cn

责任编辑: 杨鑫浩
收稿日期:2019-08-29接受日期:2019-10-8网络出版日期:2019-12-01

基金资助:

农业部现代农业产业技术体系建设专项.CARS-3-2-11
国家自然科学基金.31700163
扬州市科技计划项目.YZ2017100
扬州市科技计划项目.YZ2018041

Received:2019-08-29Accepted:2019-10-8Online:2019-12-01
作者简介 About authors
朱冬梅,E-mail：zdm@wheat.org.cn

摘要
【目的】研究小麦品种籽粒灌浆与脱水特性,为培育灌浆快、脱水快的少（免）晾晒小麦品种提供选择方法和理论依据。【方法】2015—2016年以长江中下游地区7个主推小麦品种为试验材料,采用Logistic方程拟合、多重比较及相关分析等方法,测定灌浆与脱水指标,生理成熟期及收获期籽粒含水率等。【结果】籽粒灌浆呈“S”型“慢-快-慢”的增长趋势,但不同品种最大灌浆速率、平均灌浆速率及灌浆持续时间差异显著,扬麦11、扬麦158、扬麦16最大灌浆速率及平均灌浆速率较大,花后30 d籽粒干重均达35 g以上,灌浆持续期较短;扬麦15灌浆速率仅次于上述3个品种,但灌浆持续期最长;宁麦13、扬麦20、扬麦22灌浆速率较小。最大灌浆速率、平均灌浆速率以及渐增期、快增期和缓增期的灌浆速率均与千粒重极显著正相关,3个灌浆时期灌浆速率R2>R1>R3,花后30 d灌浆基本完成。籽粒灌浆完成后进入脱水阶段,生理成熟期和收获期籽粒含水率、生理成熟后籽粒脱水速率品种间差异显著,扬麦11、扬麦158、扬麦16生理成熟后籽粒脱水速率较高,扬麦15最低。收获期籽粒含水率与生理成熟期籽粒含水率、籽粒平均脱水速率、生理成熟后2 d籽粒脱水速率显著或极显著相关。【结论】扬麦11、扬麦158、扬麦16灌浆速率大,灌浆完成早,籽粒脱水快。花后30 d粒重>35 g可作为育种材料灌浆快慢的选择指标,生理成熟期后籽粒平均脱水速率可作为衡量小麦品种脱水快慢的选择指标。
关键词： 小麦;扬麦;籽粒灌浆;籽粒脱水

Abstract

【Objective】The characteristic of grain filling and dehydration in wheat was studied, which provided a selection method and a theoretical basis for breeding wheat variety with fast filling and dehydration, and its grain with safety moisture content without drying process at harvesting date. 【Method】 In 2015 and 2016, 7 main wheat varieties in the middle and lower of the Yangtze River Valley were used as tested materials. The grain filling traits, dehydration rate and grain moisture content at physiological maturity and harvest date were measured to explore their profiles by Logistic Growth Equation (LGE) fitting analysis and multiple comparison and correlation coefficient method. 【Result】 The results indicated that grain-filling of 7 varieties fitted LGE best, with an S-like breakthrough curve of slow-fast-slow trend, but there were significant differences in the maximum grain-filling rate, average grain-filling rate and grain-filling duration in different varieties. The maximum grain-filling rates and average grain-filling rates of Yangmai 11, Yangmai 158 and Yangmai 16 were higher, whose dry grain weights at 30 d after anthesis were more than 35 g and the grain-filling durations were shorter; The grain filling rate of Yangmai 15 was the fourth fastest, but the grain-filling duration was the longest among the 7 wheat genotypes; Ningmai 13, Yangmai 20 and Yangmai 22 had the smaller filling rates, respectively. The maximum grain-filling rate, average grain-filling rate, R1, R2 and R3 were significantly positively correlated with 1000-grain weight. The rate of grain filling was R2>R1>R3 in the three filling stages. The filling grain of wheat was basically finished at 30 d after anthesis. After the grain filling stage, it started the dehydration and drying stage. There were significant differences among the 7 genotypes in the grain moisture content at physiological maturity and harvest date, as well as the grain dehydration rate. The grain dehydration rates of Yangmai 11, Yangmai 158 and Yangmai 16 were higher than the others, whereas Yangmai15 was the lowest. The grain moisture content at harvest date was significantly (P<0.01, 0.05) correlated with the grain moisture content at physiological maturity date, the average dehydration rate after physiological maturity, and the grain dehydration rate at 2 d after physiological maturity.【Conclusion】In Yangmai 11, Yangmai 158 and Yangmai 16, the grain-filling was faster and completed earlier, and the grain dehydrated more quickly. The grain weight >35 g at 30 d after anthesis could be used as the selection parameter of the grain-filling rate. The average grain dehydration rate after physiological maturity could be a selection parameter that evaluates the dehydration property of wheat.

Keywords：wheat;Yangmai;grain filling;grain dehydration

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本文引用格式
朱冬梅, 王慧, 刘大同, 高德荣, 吕国锋, 王君婵, 高致富, 陆成彬. 小麦籽粒灌浆与脱水特性[J]. 中国农业科学, 2019, 52(23): 4251-4261 doi:10.3864/j.issn.0578-1752.2019.23.006
ZHU DongMei, WANG Hui, LIU DaTong, GAO DeRong, Lü GuoFeng, WANG JunChan, GAO ZhiFu, LU ChengBin. Characteristics of Grain Filling and Dehydration in Wheat[J]. Scientia Acricultura Sinica, 2019, 52(23): 4251-4261 doi:10.3864/j.issn.0578-1752.2019.23.006

0 引言

【研究意义】全球气候变暖、极端天气频发和灌浆期高温导致小麦减产严重^[1],而且高温导致灌浆加快,时间缩短,粒重降低和品质下降^[2,3],不同品种对高温的反应存在差异^[4]。近年来长江中下游稻麦轮作区由于机插秧和水稻直播面积扩大,致使水稻收获期进一步推迟,影响小麦适期播种,推迟小麦生育进程,增加灌浆期遭遇高温的风险^[5]。蜡熟末期,小麦籽粒的物质积累量最高,达生理成熟,进入脱水阶段,直接关系到小麦收获时期的早迟,以及籽粒能否快速达到安全水分入仓储存。因此,研究小麦籽粒灌浆与脱水特性,筛选培育籽粒灌浆快、脱水快的品种有助于解决当前形势下生产面临的问题,实现小麦规模化生产的少（免）晾晒需求。【前人研究进展】小麦籽粒的灌浆特性已有大量研究,灌浆过程呈“慢-快-慢”的“S”型曲线趋势,且品种间差异较大^[6,7];小麦灌浆速率遗传力较高,快增期灌浆速率狭义遗传力为0.93^[8],灌浆速率与品种的遗传因素相关^[9,10],主要受基因的加性和显性效应控制^[11]。而灌浆持续期与粒重关系的研究结果不尽相同,KAMALUDDIN等^[12]认为小麦灌浆期持续时间的狭义遗传力为0.48;也有研究认为灌浆持续期与粒重无显著相关关系,而平均灌浆速度和最大灌浆速度与粒重极显著正相关,在育种中应选择前、中期籽粒灌浆速度较快的品系^[13],粒重主要是由快增期持续时间和灌浆速度决定的^[14]。关于籽粒灌浆完成后脱水特性的研究多集中在玉米上,收获期玉米籽粒含水率主要取决于生理成熟前后籽粒含水率和脱水速率,具有可遗传性,且品种间差异显著^[15,16,17],生理成熟后籽粒脱水速率受籽粒形状、果皮物理结构、苞叶包裹度、果穗大小及穗轴脱水速率等影响^{[18,19,20,21,22]},与玉米籽粒脱水速率相关的QTL不断被研究发现^[23,24,25]。关于小麦籽粒脱水特性的研究报道相对较少,朱冬梅等^[26]研究认为不同品种在生理成熟期籽粒含水率差异不大,但此后的脱水速率存在差异。何贤芳等^[27]认为蜡熟至收获期小麦籽粒含水量呈“慢-快-慢”的下降趋势,不同小麦品种间籽粒脱水速率差异极显著。【本研究切入点】扬麦16、宁麦13是长江中下游地区近10年来种植面积最大的两个高产小麦品种,生产中发现宁麦13千粒重低且年度间变幅较大;而扬麦16千粒重高且稳定,收获前植株转色好、籽粒水分低,可少（免）晒入库,其灌浆与脱水特性需加强研究和探明。【拟解决的关键问题】本研究为培育适合当前生产形势需求的灌浆快、脱水快的小麦品种提供理论方法,解决小麦生育后期干热风和高温逼熟引起粒重下降、产量降低的问题,确保高产稳产;解决小麦规模化生产难以烘干晾晒的问题,实现丰产丰收。

1 材料与方法

1.1 试验材料与设计

试验于2015—2016和2016—2017年度在江苏里下河地区农业科学研究所湾头试验基地进行。供试材料为长江中下游地区主推品种扬麦11、扬麦15、扬麦158、扬麦16、宁麦13、扬麦20和扬麦22。试验分别于2015年11月3日和2016年11月1日播种,采用随机区组设计,2次重复,小区面积13.33 m²,种植密度240万株/hm²。基施复合肥800 kg·hm^-2,N、P和K含量均为15%,分蘖期施尿素50 kg·hm^-2,拔节期施尿素200 kg·hm^-2。开花期用多酮和吡虫啉防治赤霉病、白粉病、蚜虫等。

1.2 试验方法

1.2.1 灌浆速率测定在小麦开花期,每小区非边行选择开花时期、穗型大小一致且无病虫害的单穗300个挂牌标记,花后10 d开始取样,以后每隔5 d在固定时间取样1次,直至收获。每次每小区取10个标记的单穗,剥取籽粒,在105℃下烘30 min杀青,80℃烘至恒重,称重计数。灌浆速率用每天每粒小麦增长重量表示,单位mg/(粒·d)。

1.2.2 生理成熟期确定花后30 d开始,每天固定时间取样,取样、杀青、烘干、称重方法同上,单粒重达最大的日期确定为生理成熟期。

1.2.3 脱水速率测定生理成熟期开始每2 d固定时间取样,每小区取10个标记的单穗,快速剥取籽粒,称鲜重,再杀青,烘干称干重,计算籽粒含水率（%）=（籽粒鲜重-籽粒干重）/籽粒鲜重×100%,脱水速率（%）=（前一次籽粒含水率-后一次籽粒含水率）/两次取样相隔天数^[27]。

1.3 脱水期间天气资料

生理成熟后2016—2017年度天气状况总体好于2015—2016年度,最高温度相对略高,且日温差较大,详见表1。

Table 1
表1
表1生理成熟后天气资料
Table 1Weather data after physiological maturity

天气 Weather	年份 Year	生理成熟后天气Weather after physiological maturity
天气 Weather	年份 Year	0 d	1 d	2 d	3 d	4 d	5 d	6 d
最高/最低温度 Highest/lowest temperature (℃)	2016	30/21	26/20	23/17	21/19	23/18	27/20	28/20
最高/最低温度 Highest/lowest temperature (℃)	2017	28/18	29/19	30/18	30/20	23/17	25/14	31/16
天气状况 Weather	2016	多云转阵雨 Cloudy to shower	阵雨 Shower	中雨转小雨 Moderate rain to light rain	小雨 Light rain	多云 Cloudy	阴 Overcast	阴转大雨 Overcast to heavy rain
天气状况 Weather	2017	晴 Sunny	多云 Cloudy	多云 Cloudy	多云 Cloudy	多云 Cloudy	中雨转阴 Moderate rain to overcast	晴 Sunny

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1.4 数据统计与分析

用Logistic方程W=K/（1+eA+Bt）^[28]对籽粒灌浆进程进行拟合,式中t为开花后时间,W为该时间点相应的籽粒干重,A、B为方程对不同品种所确定的参数,K（mg/粒）为拟合理论最高粒质量,e指自然对数函数的底数。对该方程一阶求导,可得籽粒灌浆速率方程,并可得到以下籽粒灌浆特征参数：籽粒生长起始势C0=K/(1+eA);籽粒最大灌浆速率出现时间T_max(d)=-A/B;最大灌浆速率R_max(mg/(粒·d))=-KB/4;灌浆持续时间T（d）;籽粒平均灌浆速率R_mean(mg/(粒·d))=籽粒增重/灌浆持续时间。灌浆速率曲线具有2个拐点,对灌浆速率方程一阶求导,可得2个拐点在t坐标上的值t1和t2;令t3为达到最高粒质量96%时的时间,可得到灌浆过程的3个阶段,依次是灌浆渐增期（T1）、灌浆快增期（T2）和灌浆缓增期（T3）,各阶段籽粒增加的质量分别为W1、W2和W3,对应的灌浆速率分别为R1、R2和R3。

采用Microsoft Excel 2003进行数据处理、作图,用SPSS19.0软件进行显著性检验和方差分析。

2 结果

2.1 籽粒灌浆特性

2.1.1 花后籽粒干重变化与差异性分析不同年度花后不同时期的籽粒干重有明显差异（表2）。2015年花后不同时期平均粒重均高于2016年。2年花后20—30 d籽粒新增干重接近整个灌浆期的1/2,充实度分别达57.1%—96.1%、51.6%—93.0%,说明该时期是小麦籽粒灌浆的关键时期,影响着最终粒重大小和籽粒饱满度优劣。不同品种间的籽粒干重差异显著,但2年度表现不尽一致。扬麦11、扬麦16 2年度不同时期籽粒干重均较高,显著高于宁麦13、扬麦20和扬麦22;而扬麦158 2年度籽粒干重在花后30 d前均显著高于宁麦13、扬麦20和扬麦22;扬麦15 2年度不同时期籽粒干重表现不完全一致,2015年花后30 d之前显著低于扬麦16,但花后35 d与扬麦16无显著差异;2016年与扬麦158无显著差异,但除花后30 d外,其他时期均显著低于扬麦16;宁麦13、扬麦20和扬麦22 2年度不同时期籽粒干重在7个品种中均较低,且三者差异性不显著。

Table 2
表2
表2花后籽粒干重差异性分析
Table 2Comparison of dry weight of grains after anthesis

年份 Year	品种 Cultivar	千粒重 1000-grain weight (g)
年份 Year	品种 Cultivar	10 d	15 d	20 d	25 d	30 d	35 d
2015	扬麦11 Yangmai 11	7.87ab	16.76a	25.65b	36.45a	40.59b	42.17a
	扬麦15 Yangmai 15	7.46bc	14.20c	22.01d	31.06c	38.90b	42.13a
	扬麦158 Yangmai 158	8.17a	15.74b	24.20c	34.90b	42.48a	42.83a
	扬麦16 Yangmai 16	8.14a	15.62b	27.76a	37.85a	43.24a	44.36a
	宁麦13 Ningmai 13	6.70d	12.86d	21.02de	27.38d	33.68d	34.50c
	扬麦20 Yangmai 20	6.94cd	13.16d	20.28ef	28.65d	35.12cd	36.29c
	扬麦22 Yangmai 22	6.60d	12.39d	19.48f	27.77d	36.13c	38.71b
	平均Average	7.41	14.39	22.92	32.01	38.59	40.14
2016	扬麦11 Yangmai 11	7.14b	13.27a	22.52a	33.37a	39.97a	42.06a
	扬麦15 Yangmai 15	6.55cd	11.12cd	18.74c	27.75c	35.72b	37.41b
	扬麦158 Yangmai 158	6.79bc	11.32bc	19.10c	27.65c	34.67b	35.98bc
	扬麦16 Yangmai 16	7.64a	12.41ab	20.63b	30.16b	35.91b	40.35a
	宁麦13 Ningmai 13	5.90e	10.02de	16.67d	24.88d	30.66c	33.37cd
	扬麦20 Yangmai 20	6.10de	10.35cde	17.14d	23.89d	29.71c	32.29d
	扬麦22 Yangmai 22	5.75e	9.79e	16.73d	24.76d	30.56c	33.51cd
	平均Average	6.55	11.18	18.79	27.50	33.88	36.43

Different small letters in the same column indicate significant differences among the treatments at 0.05 level. The same as below
同列数据后的不同小写字母代表处理间在5%水平差异显著。下同

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开花至花后35 d,粒重呈不断增长趋势,前快后缓,即灌浆前期籽粒增重幅度较快,后期增幅逐步减缓（图1）。不同年度间扬麦11、扬麦158、扬麦16灌浆初期籽粒增重较大,且中期物质充实快,花后25 d粒重达27.65—37.85 g,花后30 d均达35 g以上,2015年扬麦16花后30 d籽粒干重甚至高达43.24 g,接近最终粒重。

2.1.2 花后籽粒灌浆速率分析 2年花后籽粒灌浆速率7个品种均呈单峰曲线（图2）,年度间有差异但峰值均出现在花后20—25 d。2015年扬麦16、宁麦13峰值在花后20 d,其他品种均在花后25 d;2016年不同品种峰值表现一致,均在花后25 d。2015年扬麦16峰值最大,为2.43 mg/(粒·d);2016年扬麦11最大,为2.17 mg/(粒·d);2年宁麦13、扬麦20、扬麦22峰值均较小,低于1.67 mg/(粒·d),较扬麦11、扬麦16分别小0.50、0.76 mg/(粒·d)以上。2年度扬麦11、扬麦16花后25 d前籽粒灌浆速率均较高,千粒重亦较高;扬麦158籽粒灌浆速率年度间有差异,因而其千粒重2年不一致;宁麦13、扬麦20、扬麦22花后籽粒灌浆速率相对较低,导致其千粒重均不高。

图1

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图1花后籽粒干重变化趋势

Fig. 1Growth trend of dry grain weight after anthesis

图2

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图2花后籽粒灌浆速率变化趋势

Fig. 2Change trend of grain filling rates after anthesis

2.1.3 籽粒灌浆特征分析 2年试验籽粒灌浆进程Logistic方程拟合结果表明（表3）,决定系数（R²）范围在0.9939—0.9988,说明该方程可以反映不同小麦品种籽粒灌浆进程。整个籽粒灌浆过程分为3个时期,即渐增期（T1）、快增期（T2）和缓增期（T3）。花后0—15 d籽粒干重增长缓慢,持续时间年度间有差异,2年分别为11.72、12.65 d。花后15—30 d籽粒干重快速增长,是籽粒灌浆充实的关键时期,持续时间年度间差异较小,2年分别为14.79、15.64 d,相差不到1 d;但品种间差异较大,变幅为12.59—18.70 d,相差6.11 d。花后30 d至成熟,籽粒干重缓慢增长至稳定,持续时间年度间差异较小,两年分别为18.40、19.46 d;但品种间差异亦较大,变幅为15.67—23.27 d,相差7.60 d。从灌浆持续时间（T）看,年度间变幅相对较小,2年分别为44.91、47.75 d;而品种间变幅较大,为39.94—56.05 d,相差16.11 d;其中扬麦11、扬麦158、扬麦16灌浆持续期相对较短,其次是宁麦13,而扬麦15、扬麦20、扬麦22灌浆持续期较长,尤其是扬麦15最长,分别为49.03 d、56.05 d,比扬麦16长9.09 d、13.73 d。由此可见,灌浆持续时间及各时期长短在品种间变异较大,即灌浆持续时间的长短受遗传因素的影响远大于环境因素。从灌浆速率看,不同品种最大灌浆速率差异较大,变幅为1.39— 2.42 mg/(粒·d);而年度间差异较小,2年分别为1.70、1.95 mg/(粒·d)。2015年扬麦11、扬麦158、扬麦16最大灌浆速率均达2 mg/(粒·d)以上,且平均灌浆速率较高。3个时期灌浆速率均是扬麦11、扬麦158、扬麦16相对较高,其次是扬麦15,而宁麦13、扬麦20和扬麦22相对较小,均表现为R2>R1>R3,R2是R1和R3的1.94—3.57倍。由此可见,R2阶段是小麦籽粒灌浆的主要阶段。

Table 3
表3
表3不同小麦品种籽粒灌浆特征参数
Table 3Characteristic parameters of wheat cultivars in grain filling stage

年份Year	品种 Cultivar	R_max	T_max	R_mean	T1	T2	T3	R1	R2	R3	T	R²
2015	扬麦11 Yangmai 11	2.18	17.73	1.07	11.10	13.27	16.52	0.84	1.91	0.54	40.89	0.9972
	扬麦15 Yangmai 15	1.90	20.52	0.95	12.35	16.34	20.33	0.81	1.66	0.47	49.03	0.9949
	扬麦158 Yangmai 158	2.05	18.66	1.02	11.40	14.53	18.08	0.84	1.80	0.50	44.01	0.9968
	扬麦16 Yangmai 16	2.42	17.97	1.15	11.67	12.59	15.67	0.84	2.12	0.59	39.94	0.9986
	宁麦13 Ningmai 13	1.62	18.47	0.82	11.02	14.88	18.52	0.70	1.42	0.40	44.43	0.9988
	扬麦20 Yangmai 20	1.70	19.64	0.86	11.78	15.71	19.55	0.73	1.49	0.42	47.04	0.9971
	扬麦22 Yangmai 22	1.75	20.81	0.87	12.72	16.18	20.14	0.71	1.53	0.43	49.03	0.9961
	平均Average	1.95	19.11	0.96	11.72	14.79	18.40	0.78	1.71	0.48	44.91
2016	扬麦11 Yangmai 11	2.15	19.13	1.02	12.49	13.29	16.54	0.74	1.89	0.53	42.32	0.9968
	扬麦15 Yangmai 15	1.74	23.43	0.87	14.09	18.70	23.27	0.74	1.52	0.43	56.05	0.9984
	扬麦158 Yangmai 158	1.76	19.30	0.85	12.27	14.06	17.50	0.65	1.54	0.43	43.83	0.9939
	扬麦16 Yangmai 16	1.79	19.34	0.88	11.87	14.95	18.61	0.72	1.57	0.44	45.43	0.9965
	宁麦13 Ningmai 13	1.54	20.21	0.75	12.65	15.12	18.82	0.59	1.35	0.38	46.59	0.9962
	扬麦20 Yangmai 20	1.39	20.60	0.71	12.06	17.07	21.24	0.63	1.22	0.34	50.37	0.9944
	扬麦22 Yangmai 22	1.54	21.30	0.76	13.16	16.27	20.25	0.61	1.35	0.38	49.68	0.9980
	平均Average	1.70	20.47	0.83	12.65	15.64	19.46	0.67	1.49	0.42	47.75

R_max: Maximum grain-filling rate (mg/(grain·d)); T_max: Days reaching the maximum grain-filling rate (d); R_mean: Mean grain filling rate (mg/(grain·d)); T1: Grain-filling time in grain-filling pyramid period (d); T2: Grain-filling time in grain-filling fast increase period (d); T3: Grain-filling time in grain-filling slowly increase period (d); R1: Grain-filling rate in grain-filling pyramid period (mg/(grain·d)); R2: Grain-filling rate in grain-filling fast increase period (mg/(grain·d)); R3: Grain-filling rate in grain-filling slowly increase period (mg/(grain·d)); T: Grain-filling time (d); R²: Fitting coefficient . The same as below
R_max：最大灌浆速率（mg/(粒·d)）;T_max：籽粒最大灌浆速率出现时间（d）;R_mean：平均灌浆速率（mg/(粒·d)）;T1：灌浆渐增期时间（d）;T2：灌浆快增期时间（d）;T3：灌浆缓增期时间（d）;R1：灌浆渐增期灌浆速率（mg/(粒·d)）;R2：灌浆快增期灌浆速率（mg/(粒·d)）;R3：灌浆缓增期灌浆速率（mg/(粒·d)）;T：灌浆持续时间（d）;R²：拟合系数。下同

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2.1.4 籽粒灌浆特征参数与千粒重的相关性籽粒灌浆特征参数与千粒重的相关分析表明（表4）,最大灌浆速率、平均灌浆速率以及渐增期、快增期和缓增期的灌浆速率均与千粒重极显著正相关;到达最大灌浆速率的时间、快增期和缓增期持续天数、灌浆期持续天数与千粒重呈负相关关系,但均不显著。因此,小麦整个灌浆期灌浆速率是影响粒重的关键因素,而灌浆持续时间长短对粒重影响相对较小。

Table 3
表4
表4灌浆参数与千粒重的相关系数
Table 3Correlation coefficient between grain filling parameters and 1000-grain weight

年份Year	R_max	T_max	R_mean	T1	T2	T3	R1	R2	R3	T
2015	0.879**	-0.242	0.899**	0.071	-0.47	-0.471	0.949**	0.877**	0.881**	-0.402
2016	0.935**	-0.314	0.955**	-0.083	-0.444	-0.444	0.881**	0.935**	0.935**	-0.409

* and ** indicate significant differences at 0.05 and 0.01 levels. The same as below
*和**分别表示在0.05和0.01水平上差异显著性。下同

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2.2 籽粒脱水特性

2.2.1 生理成熟期和收获期籽粒含水率差异分析在生理成熟期,不同品种籽粒含水率差异显著（表5）。籽粒含水率最高的为扬麦15,比最低的扬麦11高4.27%—7.11%,亦显著高于扬麦16和扬麦158,但与扬麦22差异不显著,2016年与扬麦20差异亦不显著。生理成熟期年度间籽粒平均含水率差异较小,2年分别为40.09%、42.22%。

Table 5
表5
表5生理成熟期和收获期籽粒含水率差异比较
Table 5Comparison of grain moisture content at physiological maturity and harvest date (%)

品种 Cultivar	2015		2016
品种 Cultivar	生理成熟期 Physiological maturity	收获期 Harvest	生理成熟期 Physiological maturity	收获期 Harvest
扬麦11 Yangmai 11	39.50c	13.66e	36.22c	13.16e
扬麦15 Yangmai 15	43.77a	20.05a	43.33a	34.00a
扬麦158 Yangmai 158	42.04b	14.82de	39.73b	15.78d
扬麦16 Yangmai 16	42.36b	15.21cde	39.71b	16.78d
宁麦13 Ningmai13	41.97b	17.38bc	39.06b	19.40c
扬麦20 Yangmai20	42.35b	18.34ab	41.18ab	24.74b
扬麦22 Yangmai22	43.59a	17.06bcd	41.41ab	24.99b
平均 Average	42.22	16.65	40.09	21.26

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在试验收获期,不同品种籽粒含水率亦存在显著性差异（表5）。2年扬麦11籽粒含水率均最低,接近国家小麦入库水分标准13%,其次是扬麦158、扬麦16,2015年三者差异不显著,2016年后二者无显著差异,显著高于扬麦11,但均小于谷物临时储存水分标准18%,可直接收获入库,而不需要立即烘干或晾晒。扬麦15 2年收获期籽粒含水率均最高,且显著高于其他所有品种,分别达到了20.05%、34.00%,为籽粒储存高风险水分含量而不能直接临储,需烘干或晾晒。宁麦13、扬麦20、扬麦22 3个品种2015年收获期籽粒含水率差异不显著,2016年宁麦13显著低于扬麦20和扬麦22,但均未达临储水分标准,需烘干或晾晒。收获期年度间籽粒平均含水率差异较大,两年分别为16.65%、21.26%,可能与不同年份收获期间的天气条件有关。

2.2.2 生理成熟后籽粒脱水速率差异分析由表6可知,生理成熟期至收获期籽粒脱水速率年度间有差异,而不同品种间差异显著,不同脱水时期变化趋势不尽相同。2015年籽粒平均脱水速率较2016年高0.98%·d^-1,但生理成熟后4—6 d的脱水速率明显低于2016年,而此前差异相对较小。说明年度间小麦生育后期尤其是成熟收获期的环境条件对不同品种的籽粒脱水干燥影响较大。扬麦11、扬麦16、扬麦158的籽粒平均脱水速率均较高,尤其是2016年显著高于其他品种,而扬麦15籽粒平均脱水速率最低,分别为1.33%·d^-1和3.39%·d^-1。扬麦11、扬麦16、扬麦158在不同时期亦保持较高的脱水速率,且显著高于扬麦15,其中扬麦16在生理成熟后4 d脱水速率保持较高水平。

Table 6
表6
表6生理成熟后籽粒脱水速率比较
Table 6Comparison of grain dehydration rates after physiological maturity date (%·d^-1)

年份 Year	品种 Cultivar	平均脱水速率 Average dehydration rate	生理成熟后脱水速率 Dehydration rate after physiological maturity
年份 Year	品种 Cultivar	平均脱水速率 Average dehydration rate	0-2 d	2-4 d	4-6 d
2015	扬麦11 Yangmai 11	3.69ab	3.14a	1.86b	4.74a
	扬麦15 Yangmai 15	3.39b	1.08e	1.74b	3.98b
	扬麦158 Yangmai 158	3.89a	2.55b	2.13b	4.84a
	扬麦16 Yangmai 16	3.88a	2.01c	2.75a	5.24a
	宁麦13 Ningmai13	3.51ab	1.09e	2.23ab	4.75a
	扬麦20 Yangmai20	3.43ab	1.36de	1.06c	4.98a
	扬麦22 Yangmai22	3.79ab	1.70cd	1.86b	5.11a
	平均 Average	3.67	1.85	1.95	4.81a
2016	扬麦11 Yangmai 11	3.29a	2.04a	2.77a	7.91b
	扬麦15 Yangmai 15	1.33d	0.09e	1.67b	2.45c
	扬麦158 Yangmai 158	3.42a	0.72c	2.41a	12.85a
	扬麦16 Yangmai 16	3.28a	1.24b	2.60a	10.05b
	宁麦13 Ningmai13	2.81b	0.63c	2.38a	8.88b
	扬麦20 Yangmai20	2.35c	0.63c	1.83b	7.85b
	扬麦22 Yangmai22	2.35c	0.35cd	1.89b	8.15b
	平均 Average	2.69	0.81	2.22	8.31

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生理成熟后不同品种不同时期脱水速率变化趋势表现不完全一致,生理成熟后0—6 d的扬麦15、扬麦16、宁麦13、扬麦22均呈快速升高趋势,而扬麦11、扬麦158和扬麦20在2015年经历了一段脱水放缓过程。说明生理成熟后籽粒脱水速率受环境影响年份间略有差异,但品种间差异更显著,籽粒脱水快的基因型确实存在,而扬麦16的籽粒脱水进程年度间稳定性好、脱水快。

2.2.3 籽粒含水率与脱水速率相关性分析由表7分析,2年度间收获期籽粒含水率与生理成熟期籽粒含水率显著或极显著正相关,与脱水速率均呈负相关关系,其中与籽粒平均脱水速率相关性极显著,与生理成熟后2 d籽粒脱水速率相关性显著或极显著,而与生理成熟后4 d、6 d籽粒脱水速率仅2016年相关性极显著。表明生理成熟后籽粒平均脱水速率快慢直接影响了收获期籽粒含水率的高低,可作为衡量小麦品种脱水快慢的选择指标。

Table 7
表7
表7收获期籽粒含水率与生理成熟期籽粒含水率及脱水速率相关系数
Table 7Correlation coefficient between grain moisture content at harvest and grain moisture content at physiological maturity, grain dehydration rate

年份 Year	生理成熟期籽粒含水率 Grain moisture content during physiological maturity	籽粒平均脱水速率 Grain average dehydration rate	生理成熟后脱水速率 Dehydration rate after physiological maturity
年份 Year		籽粒平均脱水速率 Grain average dehydration rate	2 d	4 d	6 d
2015	0.763*	-0.890**	-0.915**	-0.472	-0.539
2016	0.909**	-0.984**	-0.816*	-0.942**	-0.799**

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3 讨论

本研究表明最大灌浆速率、平均灌浆速率以及渐增期、快增期、缓增期的灌浆速率与千粒重极显著正相关,而灌浆持续天数与千粒重相关不显著,这与高德荣等^[29]研究结果基本一致。蔡庆生等^[30]、罗爱花等^[31]认为灌浆速率与千粒重呈显著正相关关系,灌浆持续天数与千粒重关系不明显。BRDAR等^[32]研究认为千粒重与平均灌浆速率和最高灌浆速率高度正相关,而与灌浆期维持时间负相关,灌浆期延长可以补偿灌浆速率低的缺陷。上述研究的灌浆速率、灌浆持续期与千粒重关系异同可能与小麦生长发育期的光、温、水、肥环境有关。任正隆等^[33]比较了5个品种的灌浆特性,发现花后10 d内品种间灌浆速率无差异,花后15 d始品种间差异显著,不同品种几乎同期达到籽粒最大干重。本研究表明供试品种花后0—15 d内灌浆速率较小,籽粒干重增长缓慢;花后15 d开始籽粒干重快速增长,灌浆速率不断加快,花后20—25 d达最大;花后30 d干重缓慢增长至稳定,灌浆速率大幅度下降。扬麦11、扬麦16灌浆速率大,灌浆完成快,生理成熟早。张平平等^[34]研究亦认为扬麦16峰值灌浆速率高,快速灌浆期稳定且持续时间长,后期灌浆速率低。本研究中扬麦11、扬麦16、扬麦158花后30 d粒重均达35 g以上,而宁麦13、扬麦20、扬麦22此时粒重较低且年度间不稳定,2015年仅30 g左右。因此,在育种上花后30 d粒重可作为灌浆快慢的选择指标,据此筛选籽粒灌浆速率快的品种,尤其是花后30 d籽粒充实饱满的品种,能够有效避开高温逼熟、干热风的影响,再结合“考种看籽粒”的育种观点获得较高产量水平的优良品种。

籽粒灌浆完成后进入脱水阶段,籽粒的脱水速率不仅与环境、遗传、品种形态特征等相关^[35,36],还与种子脱水过程中生理物质有关^[37]。本研究发现不同品种在生理成熟期籽粒含水率及此后籽粒脱水速率均存在显著差异。扬麦11、扬麦16生理成熟后籽粒平均脱水速率快,快速达到谷物临时储存安全水分,成熟收获早于其他品种,可直接入库,不需要烘干或晾晒。扬麦15等品种生理成熟期籽粒含水率高,脱水速率慢,同期收获期籽粒含水率最高,需要烘干晾晒或推迟收获。长江中下游地区大面积生产实践也充分表明扬麦15全生育期较扬麦16长4—5 d,扬麦16表现出灌浆快、成熟早、籽粒含水量低的现象。虽然籽粒干燥脱水受收获期天气状况影响,但其与生理成熟后籽粒平均脱水速率极显著负相关。因此,平均脱水速率可作为衡量小麦品种脱水快慢的选择指标。

在生产上,扬麦16收获前“熟相”快速转变,且麦穗干燥下弯,以及此前种植面积最大的扬麦11表现亦如此,这一现象可能与其完成生理成熟后籽粒快速脱水有关。从遗传角度分析,扬麦11、扬麦16均为扬麦158衍生后代,是否遗传了扬麦158灌浆和脱水的优良基因或存在其他遗传机制,还有待进一步深入研究。

4 结论

花后30 d籽粒灌浆基本完成,扬麦16、扬麦11、扬麦158灌浆速率大,灌浆完成早。育种实践可根据花后30 d粒重选择灌浆快的品系,花后30 d粒重>35 g可作为灌浆快慢的选择指标。

扬麦16、扬麦11、扬麦158生理成熟后籽粒平均脱水速率快,快速达谷物临时储存安全水分,可直接收获入库,少（免）烘干晾晒。生理成熟后籽粒平均脱水速率可作为衡量小麦品种籽粒脱水快慢的选择指标。

参考文献原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

[1]

ASSENG

, FOSTER

, TURNER N

. The impact of temperature variability on wheat yields
Global Change Biology, 2011,17(2):997-1012.

DOI:10.1111/j.1365-2486.2010.02262.x URL [本文引用: 1]

With current annual production at over 600 million tonnes, wheat is the third largest crop in the world behind corn and rice, and an essential source of carbohydrates for millions of people. While wheat is grown over a wide range of environments, it is common in the major wheat-producing countries for grain filling to occur when soil moisture is declining and temperature is increasing. Average global temperatures have increased over the last decades and are predicted to continue rising, along with a greater frequency of extremely hot days. Such events have already been reported for major wheat growing regions in the world. However, the direct impact of past temperature variability and changes in averages and extremes on wheat production has not been quantified. Attributing changes in observed yields over recent decades to a single factor such as temperature is not possible due to the confounding effects of other factors. By using simulation modelling, we were able to separate the impact of temperature from other factors and show that the effect of temperature on wheat production has been underestimated. Surprisingly, observed variations in average growing-season temperatures of +/- 2 degrees C in the main wheat growing regions of Australia can cause reductions in grain production of up to 50%. Most of this can be attributed to increased leaf senescence as a result of temperatures > 34 degrees C. Temperature conditions during grain filling in the major wheat growing regions of the world are similar to the Australian conditions during grain filling. With average temperatures and the frequency of heat events projected to increase world-wide with global warming, yield reductions due to higher temperatures during the important grain-filling stage alone could substantially undermine future global food security. Adaptation strategies need to be considered now to prevent substantial yield losses in wheat from increasing future heat stress.

[2]

刁操铨 . 作物栽培学各论. 北京: 中国农业出版社, 1994: 112.

[本文引用: 1]

DIAO C

. The Theory of Crop Cultivation. Beijing: China Agriculture Press, 1994: 112. (in Chinese)

[本文引用: 1]

[3]

刘萍, 郭文善, 浦汉春, 封超年, 朱新开, 彭永欣 . 灌浆期短暂高温对小麦淀粉形成的影响
作物学报, 2006,32(2):182-188.

URL [本文引用: 1]

Wheat grain quality is affected by the environment. Temperature is a major component in environmental variation and has a marked effect on grain filling in wheat. Short period of heat shock, such as over 30℃ for a few days, happens frequently during grain filling in China and many wheat-growing areas in the world, and has become an important factor limiting wheat quality. Starch is the major storage substance in grains of wheat, comprising 65%–70% of dry weight. Investigating the effects of transient high temperature during grain filling period on starch formation and its context is useful for understanding the mechanism of heat damage and improving wheat quality under high temperature stress.
Four temperature levels, i.e. daily average temperature 25℃, 30℃, 35℃ and 40℃, respectively, were set in a phytotron, and wheat plants during different grain filling stages, including the 15th–17th, 19th–21st, 25th-27th and 33rd-35th day after anthesis (DAA), were treated. The wheat cultivars tested were Yangmai 9 with weak-gluten and Yangmai 12 with medium-gluten. Results showed that grain starch content was highest in the treatment of 30℃ and was lowest in the treatment of 40℃ in two cultivars .Under the same temperature, the effects of heat shock during the 25th–27th DAA were greatest, while that during the 33rd–35th DAA were lowest (Table 1). For the treatments of the 25th–27th DAA, the higher the temperature, the lower the soluble sugar and sucrose contents and sucrose phosphate synthase (SPS) activity in flag leaves. Sucrose synthase (SS), starch branching enzyme (SBE) activities were lowest in 40℃ treatment, and SS and SBE activities reached maximum in 25–30℃ treatments (Fig. 2–4). As for starch morphology, the granules in grains were in ellipsoid shape and combined loosely with protein sheath for 40℃ treatment of Yangmai 9 (PlateⅠ3, 4), but were damaged and in flat ball shape with a crack for that of Yangmai 12 (PlateⅠ7, 8). Starch granules of Yangmai 9 were slightly damaged for 30℃ treatment (PlateⅠ1, 2). At the 1st day after 30℃ heat shock treatment, the cracked granules were observed for Yangmai 12, but disappeared at maturity (PlateⅠ5, 6). While in 40℃ treatment, the crack could still be observed at maturity. It indicated that the heat damage at 30℃ for 3d could be considerably compensated other than at 40℃. It was probable that a kind of protective mechanism for enzymes existed in the endosperm, which had the critical temperature of 30℃. Resistance to high temperature varied with cultivars, and Yangmai 9 was stronger than Yangmai 12.

LIU

, GUO W

, PU H

, FENG C

, ZHU X

, PENG Y

. Effects of transient high temperature during grain filling period on starch formation in wheat (Triticum aestivum L.)
Acta Agronomica Sinica, 2006,32(2):182-188. (in Chinese)

URL [本文引用: 1]

ZHAO

, DAI T

, JING

, JIANG

, CAO W

. Leaf senescence and grain filling affected by post-anthesis high temperatures in two different wheat cultivars
Plant Growth Regulation, 2007,51(2):149-158.

DOI:10.1007/s10725-006-9157-8 URL [本文引用: 1]

High temperature is a major factor affecting grain yield and plant senescence in wheat growing regions of central and east China. In this study, two different wheat cultivars, Yangmai 9 with low-grain protein concentration and Xuzhou 26 with high-grain protein concentration, were exposed to different temperature regimes in growth chambers during grain filling. Four day/night temperature regimes of 34°C/22°C, 32°C/24°C, 26°C/14°C, and 24°C/16°C were established to obtain two daily temperatures of 28 and 20°C, and two diurnal day/night temperature differences of 12 and 8°C. Concentration of a lipid peroxidation product malondialdehyde (MDA), activities of the antioxidants superoxide dismutase (SOD) and catalase (CAT), chlorophyll concentration (SPAD) in flag leaves and kernel weight were determined. Results show that activities of SOD and CAT in leaves increased markedly on 14days after anthesis (DAA) for the high-temperature treatment (34°C/22°C) and then declined. As a result, MDA concentration in leaves increased significantly under high temperature (34°C/22°C and 32°C/24°C). Compared with optimum temperature treatment, high temperature reduced the concentration of soluble protein and SPAD values in flag leaves. Grain-filling rate increased slightly initially, but decreased significantly during late grain filling under high temperature. As a result, final grain weight was reduced markedly under high temperature. Decreases in the activities of SOD and CAT and increases in MDA concentration in leaves were more pronounced with a 12°C of day/night temperature difference when under high temperatures. Kernel weight was higher under 12°C of day/night temperature difference under optimum temperatures (24°C/16°C and 26°C/14°C). The responses to high-temperature regimes appeared to differ between the two wheat cultivars with different grain protein concentrations. It is concluded that a larger diurnal temperature difference hastened the senescence of flag leaves under high-temperature conditions, but retarded senescence under optimum temperature treatments of 26°C/14°C and 24°C/16°C.

[5]

高德荣, 张晓, 康建鹏, 别同德, 张伯桥, 张晓祥, 程顺和 . 长江中下游麦区小麦迟播的不利影响及育种对策
麦类作物学报, 2014,34(2):279-283.

[本文引用: 1]

GAO D

, ZHANG

, KANG J

, BIE T

, ZHANG B

, ZHANG X

, CHENG S

. Negative effects of late sowing on wheat production in middle and lower reaches of Yangtze river valley and breeding strategies
Journal of Triticeae Crops, 2014,34(2):279-283. (in Chinese)

[本文引用: 1]

[6]

孙花, 柴守玺, 刘小娥, 常磊 . 不同熟期小麦籽粒灌浆特性的研究
甘肃农业大学学报, 2009,12(6):12-18.

[本文引用: 1]

SUN

, CHAI S

, LIU X

, CHANG

. Studies on grain filling characteristics in different maturity type wheat
Journal of Gansu Agricultural University, 2009,12(6):12-18. (in Chinese)

[本文引用: 1]

[7]

杨丽娟, 董昀, 盛坤, 王映红, 赵宗武 . 超强筋小麦新品种新麦26籽粒灌浆特性研究
河南农业科学, 2011,40(11):35-37.

URL [本文引用: 1]

为了给优质超强筋小麦新品种新麦26的推广利用提供理论依据,于2009-2010年在新乡市农业科学院小麦研究所对大田试验条件下新麦26的籽粒灌浆特性进行了研究。结果表明,小麦千粒重呈"S"型增长。新麦26比新麦18灌浆持续时间短3d,平均灌浆速率(1.40mg/(粒.d))和最高灌浆速率(2.48mg/(粒.d))分别较新麦18高2.2%、4.6%,前中期灌浆速率快、积累量大。新麦26灌浆期间籽粒含水量稳定阶段短,灌浆完成后失水快。因而认为,新麦26具有灌浆时间短、灌浆速率快、粒质量高、落黄好的特点。在推广应用新麦26时,应注意提高单位面积穗数和穗粒数,并适时收获防止籽粒干物质回流。

YANG L

, DONG

, SHENG

, WANG Y

, ZHAO Z

. Grain filling characteristics of new strong gluten wheat cultivar Xinmai 26
Journal of Henan Agricultural Sciences, 2011,40(11):35-37. (in Chinese)

URL [本文引用: 1]

王敏, 姚维传 . 小麦灌浆特性的遗传研究:I. 遗传模型及基因效应
遗传, 1996,18(5):23-26.

URL [本文引用: 1]

利用8个小麦品种双列杂交的F~1|及其亲本, 按Hayman分析法研究了小麦籽粒灌浆特性的遗传规律。结果表明,小麦籽粒灌浆特性符合加性-显性遗传模型。基因作用以加性效应为主,呈部分显性。籽粒灌浆期的显性方向指向增效,籽粒灌浆速率的显性方向指向减效。所有性状均有较高的狭义遗传力h^2|~N|=0.71-0.93,可在杂种早代选择。

WANG

, YAO W

. Inheritance of grain filling duration and rate in wheat: I. Genetic model and gene effect

Hereditas,

1996,18(5):23-26. (in Chinese)

URL [本文引用: 1]

王瑞霞, 张秀英, 伍玲, 王瑞, 海林, 闫长生, 游光霞, 肖世和 . 不同生态环境条件下小麦籽粒灌浆速率及千粒重QTL分析
作物学报, 2008,34(10):1750-1756.

DOI:10.3724/SP.J.1006.2008.01750 URL [本文引用: 1]

The duration and rate of grain filling determine the individual grain size, thousand-grain weight (TGW), and final grain yield. Several reports have focused on the physiological basis of grain filling in wheat (Triticum aestivum L.), but rare on the genetic mechanism and QTL mapping due to its complexity. To identify QTLs related to grain filling, the F_7:8 generation of 142 recombinant inbred lines (RILs) derived from the cross between Yu 8679 (large spike) and Heshangmai (small spike) were planted in four ecological environments in Beijing (2006 and 2007), Hefei (2007), and Chengdu (2007). Three agronomic traits including mean grain filling rate (GFR_mean), maximum grain filing rate (GFR_max), and TGW were evaluated. A genetic map comprising 170 SSR and 2 EST markers (Tx23-24 and Tx37-38) was constructed based on the 142 RILs. According to the genetic map and phe-notypic data, quantitative trait loci were identified for these agronomic traits using the composite interval mapping (CIM) method. A total of 54 QTLs located on chromosomes 1A, 1B, 2A, 2D, 3A, 3B, 3D, 4A, 4D, 5A, 5B, 6D, and 7D for the three traits were identified over four environments. Among them, 17 for GFR_mean, 16 for GFR_max, and 21 for TGW, accounted for variations of GFR_mean, GFR_max, and TGW by 7.17–20.83%, 6.31–15.95%, and 4.36–16.80%, respectively. Ten genomic sections involving chromosomes 1A, 1B, 2A, 3B, 4D, 6D, and 7D with “pleiotropic effects” were detected. These QTLs with pleiotropic effects are useful for understanding the relationship between grain filling and other related grain yield traits at gene level.

WANG R

, ZHANG X

, WU

, WANG

, HAI

, YAN C

, YOU G

, XIAO S

. QTL mapping for grain filling rate and thousand-grain weight in different ecological environments in wheat
Acta Agronomica Sinica, 2008,34(10):1750-1756. (in Chinese)

DOI:10.3724/SP.J.1006.2008.01750 URL [本文引用: 1]

孙进先, 魏秀华, 王国飞, 于新华, 王德高, 张其鲁 . 品种、播期、灌水和施氮量对小麦灌浆速率的影响
山东农业科学, 2010,7:48-50.

URL [本文引用: 1]

采用正交试验设计研究了品种、播期、灌水和施氮量对小麦籽粒平均灌浆速率的影响,并对其灌浆曲线进行拟合.结果显示,品种是影响小麦灌浆速率的主要因素,其次为施氮量,均达显著水平;灌水和播期对平均灌浆速率的影响较小.各因素对小麦灌浆的影响曲线,以三次多项式拟合效果最好.表明,灌浆速率与品种的遗传因素相关,增施适量氮肥可有效提高灌浆速率,而增加灌水量有延长灌浆持续期的趋势,推迟播期则有缩短灌浆持续期的风险.

SUN J

, WEI X

, WANG G

, YU X

, WANG D

, ZHANG Q

. Effects of cultivar, sowing date, irrigation and nitrogen fertilizer rate on wheat grain filling rate
Shandong Agricultural Sciences, 2010,7:48-50. (in Chinese)

URL [本文引用: 1]

MASHIRINGWANI N

, MASHINGAIDZE

, KANGAI

, OLSEN

. Genetic basis of grain filling rate in wheat ( Triticum aestivum L. emend. Thell.)
Euphytica, 1994,76(1/2):33-44.

DOI:10.1007/BF00024018 URL [本文引用: 1]

[12]

KAMALUDDIN, SINGH R

, ABDIN M

, KHAN M

, ALAM

, KHAN

, PRASAD L

, JOSHI A

. Inheritance of grain filling duration in spring wheat ( Triticum aestivum L. em thell)
Journal of Plant Biology, 2007,50(4):504-507.

DOI:10.1007/BF03030690 URL [本文引用: 1]

To understand the genetic control of grain filling duration (GFD), i.e., the number of days from anthesis to physiological maturity, we studied the F₁, F₂, BC1 and BC₂ generations of six spring wheat crosses from nine varieties/genotypes. Generation mean analysis for gene effects indicated that one or more types of epistasis were significant in all crosses. In each pairing, the F₁ and F₂ means were either intermediate or closer to the mean of the parent having the longer GFD. Our narrow-sense heritability estimate was reasonably high, at 47.67 (based on diallel analysis). This demonstrated that progress could be made from the selection in these crosses for either long or short GFD. The two early varieties that had identical maturity durations differed in their GFD values, indicating that maturity dates are not good criteria when choosing parents for modifying GFD. To utilize favorable additive × additive effects during this selection, we suggest that a single seed descent (SSD) or bulk popula-tion approach be adopted. In comparison, dominance effects would prove quite useful in hybrid wheat breeding programs.

[13]

任明全, 徐向阳 . 不同小麦品种籽粒灌浆特性的研究
华北农学报, 1993,8(3):28-32.

DOI:10.3321/j.issn:1000-7091.1993.03.006 URL [本文引用: 1]

Twelve winter wheat cultivars that were widely planted in South Huang-Huai region were used to study their grain filling characters.Conclusions were obtained as follows: Significantly positive relationships were found between mean grain filling rate, maximum grain filling rate and 100 grain weight, kernel volume, kernel maximum volume.It was grain filling index, not 100 grain weight, that had significant relationship with grain filling duration.Among the cultivars used, Zhcngzhou 853, Shannong 7859 had the biggest mean grain filling rate and maximum grain filling rate, with the shortest grain filling duration.They belonged to the genotype of late flowering.The grain filling durations of Ji 5418 and Xuzhou 21 had the longest grain filling duration, and they belonged to the genotype of early flowering.In addition, the suitable grain filling pattern in South Huang-Hai region was also discussed.

REN M

, XU X

. Studies on the grain filling characters of wheat cultivars
Acta Agriculturae Boreali-Sinica, 1993,8(3):28-32. (in Chinese)

DOI:10.3321/j.issn:1000-7091.1993.03.006 URL [本文引用: 1]

冯素伟, 胡铁柱, 李淦, 董娜, 李笑慧, 茹振钢, 程自华 . 不同小麦品种籽粒灌浆特性分析
麦类作物学报, 2009,29(4):643-646.

DOI:10.7606/j.issn.1009-1041.2009.04.019 URL [本文引用: 1]

为了给小麦高产栽培及新品种选育提供依据，对9个小麦品种的籽粒灌浆过程进行了研究。结果表明，小麦籽粒干重呈“S”型曲线增长，籽粒灌浆期可分为渐增期、快增期、缓增期三个阶段。相关分析表明，粒重主要是由快增期持续时间和灌浆速度决定的，与整个灌浆持续期关系不明显。在小麦灌浆快增期，灌浆速度越快，持续时间越长，干物质积累越多，粒重就越高。不同品种间籽粒灌浆特性存在一定的差异；综合分析表明，襄麦29和百农矮抗58 1籽粒灌浆速度较快，粒重较高。

FENG S

, HU T

, LI

, DONG

, LI X

, RU Z

, CHENG Z

. Analysis on grain filling characteristics of different wheat varieties

Journal of Triticeae Crops,

2009,29(4):643-646. (in Chinese)

DOI:10.7606/j.issn.1009-1041.2009.04.019 URL [本文引用: 1]

PURDY J

, CRANE P

. Inheritance of drying rate in mature corn(Zea mays L.)
Crop Science, 1967,7(4):294-297.

DOI:10.2135/cropsci1967.0011183X000700040003x URL [本文引用: 1]

[16]

张林, 王振华, 金益, 于天江 . 玉米收获期含水量的配合力分析
西南农业学报, 2005,18(5):534-537.

[本文引用: 1]

ZHANG

, WANG Z

, JIN

, YU T

. Combine ability analysis of water content in harvest stage in corn
Southwest China Journal of Agricultural Sciences, 2005,18(5):534-537. (in Chinese)

[本文引用: 1]

[17]

王克如, 李少昆 . 玉米籽粒脱水速率影响因素分析
中国农业科学, 2017,50(11):2027-2035.

DOI:10.3864/j.issn.0578-1752.2017.11.008 URL [本文引用: 1]

Grain moisture content of maize is the key factor that affects the quality of mechanical harvesting, safe storable level, and economic benefits. It has become an important technical and economic problem. At present, the high moisture content of grain in the corn harvest period not only restricts the popularization of corn combine harvesting technology, but also affects the change of maize harvest and production mode, and seriously affects the grain quality of maize. A review of relevant literature both at home and abroad shows that the moisture content of grain in harvesting period is controlled mainly by the dry-down rate of grain before and after physiological maturity, and this trait is heritable, and it has a significant difference among maize hybrids. The difference of grain drying rate is closely correlated with many agronomic traits of maize, such as length and thickness of bract, thickness of cob, shape of kernel, and size of ear. The ecological factors such as air humidity (saturation degree of environmental water deficit), temperature, solar radiation, wind speed, rainfall and so on, have important influences on grain drying rate at the late growth stage of maize. Agronomic measurements such as planting density, row spacing, irrigation and fertilization, also have some influence on drying rate of grain. The optimum period of harvesting grain with combine machine can be predicted by the grain moisture content and the drying rate of grain after physiological maturity. In this paper, it is suggested that, at present, the selection of maize hybrids having characteristics of suitable early maturity, and rapid drying rate of ear at grain filling stage and low moisture content of grain at physiological maturity is the key measure to realize grain mechanical harvest in maize production areas in China. At the same time, due to the combined effects on drying rate of grain by genotype, ecological and meteorological factors and cultivation measures, meanwhile, the corn planting regions are wide, the planting patters are diverse, and the maize variety types used are various in China, further studies on the physiological mechanism of kernel dry-down are needed, and systematic observation characteristics of drying rate of grain should be carried out, that will provide a theoretical basis and technical supports for promotion of grain machinery harvesting technology and improvement of maize grain quality.

WANG K

, LI S

. Analysis of influencing factors on kernel dehydration rate of maize hybrids
Scientia Agricultura Sinica, 2017,50(11):2027-2035.

DOI:10.3864/j.issn.0578-1752.2017.11.008 URL [本文引用: 1]

CRANE P

, MILES S

, NEWMAN J

. Factors associated with varietal differences in rate of field drying in corn
Agronomy Journal, 1959,51(6):318-320.

DOI:10.2134/agronj1959.00021962005100060003x URL [本文引用: 1]

[19]

PURDY J

, CRANE P

. Inheritance of drying rate in “mature” corn ( Zea mays L.)
Crop Science, 1967,7(4):294-297.

DOI:10.2135/cropsci1967.0011183X000700040003x URL [本文引用: 1]

[20]

TROYER A

, AMBROSE W

. Plant characteristics affecting field drying rate of ear corn
Crop Science, 1971,11(4):529-531.

DOI:10.2135/cropsci1971.0011183X001100040019x URL [本文引用: 1]

[21]

郭佳丽, 吕志尧, 吕颖颖, 胡海军, 姚晓云, 贾森, 李凤海, 史振声 . 玉米粒部性状对籽粒脱水速率的影响
玉米科学, 2014,22(4):33-38.

[本文引用: 1]

GUO J

, Lü Z

, Lü Y

, HU H

, YAO X

, JIA

, LI F

, SHI Z

. Effect of kernel characteristics on kernel dehydration rate of maize
Journal of Maize Sciences, 2014,22(4):33-38. (in Chinese)

[本文引用: 1]

[22]

张树光, 冯学民, 高树仁, 孙生林 . 玉米成熟期籽粒含水量与果穗性状的关系
中国农学通报, 1994,10(2):15-17.

[本文引用: 1]

ZHANG S

, FENG X

, GAO S

, SUN S

. Study on kernel moisture content and ear characters of maize hybrids with different maturity time
Chinese Agricultural Science Bulletin, 1994,10(2):15-17. (in Chinese)

[本文引用: 1]

[23]

SALA R

, ANDRADE F

, CAMADRO E

, CERONO J

. Quantitative trait loci for grain moisture at harvest and field grain drying rate in maize ( Zea mays L.)
Theoretical and Applied Genetics, 2006,112(3):462-471.

DOI:10.1007/s00122-005-0146-5 URL [本文引用: 1]

Hybrids with low grain moisture (GM) at harvest are specially required in mid- to short-season environments. One of the most important factors determining this trait is field grain drying rate (FDR). To produce hybrids with low GM at harvest, inbred lines can be obtained through selection for either GM or FDR. Thus, a single-cross population (181 F _2:3-generation plants) of two divergent inbred lines was evaluated to locate QTL affecting GM at harvest and FDR as a starting point for marker assisted selection (MAS). Moisture measurements were made with a hand-held moisture meter. Detection of QTL was facilitated with interval mapping in one and two dimensions including an interaction term, and a genetic linkage map of 122 SSR loci covering 1,557.8cM. The markers were arranged in ten linkage groups. QTL mapping was made for the mean trait performance of the F _2:3 population across years. Ten QTL and an interaction were associated with GM. These QTL accounted for 54.8 and 65.2% of the phenotypic and genotypic variation, respectively. Eight QTL and two interactions were associated with FDR accounting for 35.7 and 45.2% of the phenotypic and genotypic variation, respectively. Two regions were in common between traits. The interaction between QTL for GM at harvest had practical implications for MAS. We conclude that MAS per se will not be an efficient method for reducing GM at harvest and/or increasing FDR. A selection index including both molecular marker information and phenotypic values, each appropriately weighted, would be the best selection strategy.

[24]

刘显君, 王振华, 王霞, 李庭锋, 张林 . 玉米籽粒生理成熟后自然脱水速率QTL的初步定位
作物学报, 2010,36(1):47-52.

DOI:10.3724/SP.J.1006.2010.00047 URL [本文引用: 1]

以吉846 (脱水快，1.18% d^-1)和掖3189 (脱水慢，0.39% d^-1)为亲本衍生出的232个重组自交系(F₇)为作图群体，构建了具有101个SSR标记位点的玉米遗传连锁图谱，覆盖玉米基因组1 941.7 cM，标记间平均距离为19.22 cM。通过1年2点试验(双城和哈尔滨，2007年)评价了232个重组自交系籽粒生理成熟后的自然脱水速率。采用WinQTL2.5对该性状数量性状位点(QTL)进行了初步定位和遗传效应分析，结果共检测出9个显著影响玉米籽粒生理成熟后自然脱水速率的QTL，分别位于第2、3、4、5和6染色体上，加性增效作用均来源于亲本吉846。其中在第2和6染色体上的2个QTL (qKdr-2-1, qKdr-6-1)在2个环境下均稳定表达，分别位于SSR标记bnlg198-umc1516和phi126-phi077之间，其累积表型贡献率为15.49%。具有较快脱水速率的等位基因均来自吉846。所检测到的QTL将在分子辅助选育具有较快脱水速率的材料中具有较大的应用潜力。

LIU X

, WANG Z

, WANG

, LI T

, ZHANG

. Primary mapping of QTL for dehydration rate of maize kernel after physiological maturing

Acta Agronomica Sinica,

2010,36(1):47-52. (in Chinese)

DOI:10.3724/SP.J.1006.2010.00047 URL [本文引用: 1]

WANG Z

, WANG

, ZHANG

, LIU X

, DI

, LI T

, JIN X

. QTL underlying field grain drying rate after physiological maturity in maize ( Zea mays L.)
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DOI:10.1007/s10681-012-0676-2 URL [本文引用: 1]

Grain moisture in maize at harvest depends on the grain drying rate (GDR) after physiological maturity. The maize plants with high GDR can reduce grain moisture rapidly, which will shorten the drying time after harvest and prevent the grain to be mildew and enhance maize quality. In this study, A total of 280 recombinant inbred lines that were derived from a cross between Ji846 (high drying rate, 1.18 % day(-1)) and Ye3189 (slow drying rate, 0.39 % day(-1)) were used to construct genetic linkage map and identify QTL underlying GDR in different environments. A genetic linkage map was constructed containing 97 SSR and 49 AFLP markers, which covered 2356.8 cM of the maize genome, with an average distance of 16.1 cM. Composite interval mapping identified 14 QTL for GDR after physiological maturity located on chromosomes 2, 3, 5, 6 and 8. The additive effects of QTL were all from Ji846. The range of phenotypic variation explained by the QTL was 5.05-16.28 %. But only two QTL (qKdr-2-1, qKdr-3-6) were identified across both locations. qKdr-2-1 positioned between the markers phi090-umc1560 on chromosome 2 explained 15.59 % of the phenotypic variance, and the other qKdr-3-6 positioned between the markers phi046-bnlg1754 on chromosome 3 explained 10.28 % of the phenotypic variance.

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扬州大学学报: 农业与生命科学版, 2015,36(2):77-78.

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高德荣, 王慧, 刘巧, 朱冬梅, 张晓, 吕国锋, 张晓祥, 江伟, 李曼 . 迟播早熟高产小麦新品种的培育
中国农业科学, 2019,52(14):2379-2390.

DOI:10.3864/j.issn.0578-1752.2019.14.001 URL [本文引用: 1]

【Objective】The objectives of this study were to investigate the growth characteristics during the period from sowing to elongation and from anthesis to maturity in wheat cultivars (varieties), and to explore the possibility of breeding new cultivars with the feature of rapid growth at both early and late stages to achieve high yield and early maturity under late sowing condition in the middle and lower reaches of the Yangtze river. 【Method】 In the seasons of 2012-2013 and 2013-2014, experiments were conducted to compare leaf number, stem and tiller number and the parameters of grain filling among 18 new wheat lines and cultivars for screening the cultivar with rapid development at both early and late growth stages. In the season of 2014-2015, the differences of stem and tiller number, leaf area index (LAI), dry matter accumulation, yield and yield components were analyzed among Yangmai 16 and other 4 cultivars under the conditions of the normal (5th Nov.) and late (19th Nov.) sowing time to clarify the growth mechanics of high-yield about Yangmai 16 with rapid development at both early and late growth stages under late-sowing treatment.【Result】 Fu F101 showed a rapid development at early growth stage with a significantly higher number of leaves and tillers (before winter and during winter period) than Zhen 10216 in the 2012-2013, Ning 09-118 and Zhen 10216 had longer grain filling and rapid development at late growth stage with higher maximum grain filling rate (G_max) and mean grain filling rate (G_mean) thus higher grain weight than other cultivars. The 2013-2014 trial showed similar results with Fu F101 performing rapid development at early stage and Zhen 10216 at late stage. Yangmai 16 produced even more tillers than Fu F101 before winter and similar tillers to Fu F101 during winter period. Meanwhile, both G_mean and G_max of Yangmai 16 were similar to Zhen 10216 with G_max being more than 2.0 mg/(grain·d). LAI, stem and tiller number and dry matter accumulation of Yangmai 16 in winter period were significantly higher than Ningmai 13 and other cultivars. In 2014-2015 trial, the yield of Yangmai 16 was lower than Yangmai 22 and similar to Yangmai 20, Yangmai 23 and Ningmai 13 under normal sowing time. However, under late sowing, the yield of Yangmai 16 was significantly higher than Yangmai 20 and Ningmai 13 and had the lowest yield reduction (5.2%) compared to other cultivars.【Conclusion】Significant differences were found among varieties in the rate of development at early and late growth stages. The rapid development at both early and late stages of Yangmai 16 makes it an excellent variety with not only a reasonable yield under normal sowing date but minimum yield reduction under late sowing. In summary, rapid development at both early and late stages can be used as a key criterion for selecting wheat varieties suitable for late sowing.

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, WANG

, LIU

, ZHU D

, ZHANG

, Lü G

, ZHANG X

, JIANG

, LI

. Breeding of new wheat varieties with early maturity and high yield under late sowing
Scientia Agricultura Sinica, 2019,52(14):2379-2390. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2019.14.001 URL [本文引用: 1]

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CAI Q

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. The relations of dry matter accumulation of grain growth stages to grain weight in wheat
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LUO A

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