李萍^,1, 尚云秋¹, 林祥¹, 刘帅康¹, 王森¹, 胡鑫慧¹, 王东^,1,21山东农业大学/作物生物学国家重点实验室,山东泰安 271018
²淄博禾丰种业科技股份有限公司, 山东临淄 255000

Effects of Drought Stress During Jointing Stage on Spike Formation and Seed Setting of Main Stem and Tillers of Winter Wheat

LI Ping^,1, SHANG YunQiu¹, LIN Xiang¹, LIU ShuaiKang¹, WANG Sen¹, HU XinHui¹, WANG Dong^{,1,2 1}Shandong Agricultural University/State Key Laboratory of Crop Biology, Tai’an 271018, Shandong
²Zibo HeFeng Seed Technology co., ltd., Linzi, Shandong, 255000, China

通讯作者: 王东,E-mail: wangd@sdau.edu.cn

责任编辑: 杨鑫浩
收稿日期:2020-01-2接受日期:2020-03-17网络出版日期:2020-10-16

基金资助:

山东省重大科技创新工程项目.2019JZZY010716 东省泰山产业领军人才项目 家公益性行业(农业)科研专项.201503130

Received:2020-01-2Accepted:2020-03-17Online:2020-10-16
作者简介 About authors
李萍,E-mail: sdauliping@163.com。









摘要
【目的】针对黄淮海地区自然降水季节分布不均、阶段性干旱频发导致小麦产量和水分利用效率低的问题,探索拔节期阶段性干旱对冬小麦主茎和分蘖成穗与结实的影响,可为该地区冬小麦节水栽培提供理论和技术支持。【方法】于2017—2019年冬小麦生长季,在室外遮雨条件下进行盆栽试验。以小麦品种山农29和衡0628为试验材料,在拔节后0—10 d期间设置5个水分处理:充分供水（CK,保持土壤相对含水量75%—80%,土壤有效含水量为42.2—46.7 mm）;拔节后0—5 d轻度干旱（T1,保持土壤相对含水量为65%—70%,土壤有效含水量为33.4—37.8 mm）、重度干旱（T2,保持土壤相对含水量为45%—50%,土壤有效含水量为15.6—20.1 mm）;拔节后0—10 d轻度干旱（T3,保持土壤相对含水量为65%—70%,土壤有效含水量为33.4—37.83 mm）、重度干旱（T4,保持土壤相对含水量为45%—50%,土壤有效含水量为15.6—20.1mm）,测定了茎蘖幼穗发育进程及茎蘖成穗和结实性状等指标。【结果】在拔节后0—10 d期间不同程度干旱对小麦主茎成穗无明显影响,但是随着干旱时间的延长和干旱程度的加大,低位蘖（Ⅲ和Ⅰp）成穗率迅速下降,而高位蘖（Ⅱp和Ⅰ1）成穗率呈先升高后下降趋势。拔节后0—5 d轻度或重度干旱,高位蘖成穗率均较高,单位面积成穗数与CK无显著差异;拔节后0—10 d轻度干旱,高位蘖成穗率虽与CK相近,但由于低位蘖（Ⅲ、Ⅰp）成穗率下降幅度较大,导致单位面积成穗数显著降低,山农29和衡0628单位面积穗数下降幅度分别为4.94%—5.06%和6.77%—8.33%;拔节后0—10 d重度干旱,Ⅱ蘖以上分蘖成穗率均下降,山农29和衡0628单位面积成穗数下降幅度分别为10.97%—11.52%和15.00%—15.55%。拔节后0—5 d轻度干旱,2个品种主茎和各蘖位分蘖的结实性、单穗产量和单位面积产量均与CK无显著差异。拔节后0—5 d重度干旱,2个品种各中位蘖的结实小穗数和穗粒数均显著减少,主茎和高位蘖受影响不明显;山农29各茎蘖单粒重不受影响而单穗产量显著降低;衡0628各茎蘖单粒重和单穗产量显著降低;山农29和衡0628单位面积籽粒产量均显著降低,分别比CK减少5.14%—5.46%和5.45%—6.24%。拔节后0—10 d轻度和重度干旱,2个品种茎蘖的总小穗数、结实小穗数、穗粒数、单粒重、单穗产量和单位面积籽粒产量均显著降低,且以中位蘖下降幅度较大;重度干旱处理各茎蘖的穗粒数和单穗产量及单位面积籽粒产量显著低于轻度干旱处理。山农29和衡0628单位面积籽粒产量在T3处理下分别比CK减少12.87%—13.30%和15.52%—16.59%;在T4处理下分别比CK减少23.18%—25.92%和26.05%—31.22%。【结论】拔节后短时间轻度干旱（拔节后0—5 d保持土壤相对含水量65%—70%,土壤有效水含量33.4—37.8 mm）对小麦成穗和结实无显著影响;干旱时间过长、程度过大则会大幅度降低低位蘖（Ⅲ和Ⅰp）成穗率、总小穗数、结实小穗数、穗粒数、单粒重和单穗产量,导致单位面积籽粒产量显著下降。在拔节后5 d干旱或拔节后10 d轻度干旱条件下,高位蘖（Ⅱp和Ⅰ1）成穗率有所提高,在一定程度上可弥补干旱造成的损失,这可能与低位分蘖受旱后成穗率降低,群体变小,动摇分蘖分配的营养增多、生存空间增大有关,为生产中通过合理措施调控,实现小麦稳产提供了理论依据。山农29对拔节期阶段性干旱的抗性高于衡0628。
关键词： 冬小麦;阶段性干旱;主茎与分蘖;成穗与结实;产量

Abstract
【Objective】To address the problem of low wheat yield and water efficiency caused by irregular distribution of natural precipitation season and the frequent rate of staged drought in the Huang-Huai-Hai Plain of China, this paper provided theoretical as well as technical provisions for water-saving cultivation of winter wheat in this area by exploring the effects of water insufficiency on the main stem and tiller spikes formation and panicles traits during jointing stage.【Method】A 2-year pot experiment under external rain conditions was carried out from 2017 to 2019 in winter wheat growth season, and the two winter wheat varieties of Shannong 29 and Heng 0628 were used as experimental materials. The total five water treatments were set up during 0-10 d after jointing, including full irrigation treatment during whole growing season as control (CK, maintaining soil relative water content of 75%-80%, the effective soil water content of 42.2-46.7 mm), 0-5 d light drought stress after jointing (T1, maintaining soil relative water content of 65%-70%, soil availability water content of 33.4-37.8 mm), 0-5 d severe drought (T2, maintaining soil relative water content of 45%-50%, available soil water capacity is 15.6-20.1 mm), 0-10 d light drought stress after jointing (T3, maintaining soil relative water content is 65%-70%, soil availability water content of 33.4-37.8 mm), and 0-10 d severe drought stress (T4, maintaining the soil relative water content 45%-50%, availability water content of 15.6-20.1mm).【Result】The results revealed that different degrees of drought stress had no significant effect on the main stem of wheat during 0-10 d after jointing, the effective spike rate of low tiller (Ⅲ, Ⅰp ) decreased rapidly with the increasing the drought level and extension of drought time, while the effective spike rate of high tiller (Ⅱp and Ⅰ1) increased first and then decreased. Light or severe drought at 0-5 d after jointing stage, Ⅱp and Ⅰ1 were higher in effective spike rate, and there was no significant difference in spike number per unit area from CK. Although the effective spike rate of Ⅰ1 and Ⅱp was similar to that of CK in 0-10 d light drought after jointing stage, the percentage of spikes per unit area decreased significantly due to the decrease of Ⅲ, Ⅰp and Ⅳ. The number of spike per unit area of Shannong 29 and Heng 0628 was decreased by 4.94%-5.06% and 6.77%-8.33%, respectively. For 0-10 d light drought after jointing, the effective rate of tiller of Ⅱ and more was decreased of severe drought. The number of spikes per unit area of Shannong 29 and Heng 0628 was decreased by 10.97%-11.52% and 15.00%-15.55%, respectively. The treatment of light drought at 0-5 d after jointing stage, the panicle characteristics, single stem grain yield and grain yield per unit area of the two cultivars were not significantly different from CK. The treatment of severe drought from 0 to 5 days after jointing, the fertile spikelets number and the grain number in the two cultivars were significantly reduced, and the main stem and later occurring tillers were not affected. The single grain weight of Shannong 29 was not affected, but the yield of single stem grain yield was significantly reduced, the single grain weight and single stem grain yield of Heng 0628 was significantly reduced, the grain yield per unit area of Shannong 29 and Heng 0628 decreased significantly, which was 5.14%-5.46% and 5.45%-6.24% lower than CK, respectively. For the light and severe drought from 0 to 10 days after jointing, the spikelets number, the fertile spikelets number, grain number, single grain weight, single stem grain yield and grain yield per unit area of the two cultivars were significantly reduced; The median tillers had substantial reduction; The number of grains per ear and the yield of single-stem grain and the grain yield per unit area of severely drought-treated stems were significantly lower than those of light drought treatment. Under the T3 condition, the grain yield per unit area of Shannong 29 and Heng 0628 was decreased by 12.87%-13.30% and15.52%-16.59%, respectively. Under the T4 condition, the grain yield per unit area of Shannong 29 and Heng 0628 decreased by 23.18%-25.92% and 26.05%-31.22%, respectively.【Conclusion】The above results indicated that short-term light drought after jointing (0-5 days after jointing, maintaining soil relative water content of 65%-70% and soil effective water content of 33.4-37.8 mm) had no significant effects on wheat earing and fruiting; The percent of tillers to panicle, the spikelets number, the fertile spikelets number, grain number, single grain weight and single stem grain yield of the low tillers (Ⅲ, Ⅰp) would be greatly reduced, and the grain yield per unit area was significantly reduced increased drought stress and time. However, the effective spike rate of high tillers (Ⅱp and Ⅰ1) in two degrees of 0-5 d after jointing and light drought training of 0-10 d after jointing was increased to some extent, which could make up for the loss caused by drought, which might be related to the decrease of the effective spike rate of low tillers, the decrease of population size, and the increase of nutrition and living space for high tillers. Shannong 29 had the stronger potential to adapt drought stress at jointing stage than Heng 0628.
Keywords：winter wheat;phased drought;main stem and tillers;spike formation and seed setting;grain yield


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本文引用格式
李萍, 尚云秋, 林祥, 刘帅康, 王森, 胡鑫慧, 王东. 拔节期阶段性干旱对小麦茎蘖成穗与结实的影响[J]. 中国农业科学, 2020, 53(20): 4137-4151 doi:10.3864/j.issn.0578-1752.2020.20.004
LI Ping, SHANG YunQiu, LIN Xiang, LIU ShuaiKang, WANG Sen, HU XinHui, WANG Dong. Effects of Drought Stress During Jointing Stage on Spike Formation and Seed Setting of Main Stem and Tillers of Winter Wheat[J]. Scientia Acricultura Sinica, 2020, 53(20): 4137-4151 doi:10.3864/j.issn.0578-1752.2020.20.004

0 引言

【研究意义】黄淮海地区是中国小麦主产区,其小麦生产对保障国家粮食安全具有重要作用。然而受全球气候变化的影响,该地区干旱频繁发生,特别是近年来随着气候变暖趋势的加剧,冬春两季干旱发生频率和危害程度逐渐加强,对小麦分蘖成穗和结实产生诸多不利影响,成为该地区小麦生产的重要限制因素^[1,2]。拔节期是小麦分蘖两极分化和穗花分化的重要时期。探索拔节期不同水分条件对小麦茎蘖成穗和结实的调节作用及其生理基础,对春季麦田合理灌溉技术的研发具有重要参考价值和生产指导作用。【前人研究进展】分蘖是小麦等禾本科作物的重要农艺性状。小麦在生长进程中发生无效分蘖,造成一定的“生长冗余”^[3]。在资源受限的情况下,这种生长冗余会对小麦生产造成不利的影响^[4]。国内外****对分蘖成穗特性做了大量研究^[5,6,7]。越来越多的****认为,小麦产量的进一步增加主要依赖于控制无效分蘖的数量,增加动摇分蘖的成穗率,使茎蘖成穗率和群体质量提高^[8,9,10]。小麦分蘖消亡和开花结实受多基因加性效应的控制,且易受幼穗分化过程中栽培因素和环境的影响^[11,12]。小麦拔节期不同程度干旱均影响已发生分蘖的生长发育,最终影响分蘖的成穗与结实^[13,14,15]。将灌溉时间由主茎基部第一节间伸长期（拔节期）推迟到第四节间伸长期（旗叶抽出期）,即拔节期适度干旱虽然显著降低成穗数,但增加了穗粒数、而且延缓了开花后叶片衰老、显著提高叶片净光合速率、籽粒灌浆速率、千粒重、籽粒产量和水分利用效率^[16]。然而拔节期过度干旱则会降低小麦有效穗数和穗粒数,严重干旱条件下小麦千粒重、单株和单位面积籽粒产量亦显著降低^[17,18]。小麦主茎与不同蘖位分蘖间穗花分化存在明显的时间差,主茎穗花分化明显早于分蘖,低位蘖又早于高位蘖,这种时间差造成不同茎蘖的生长条件和营养条件不同,最终导致不同茎蘖经济产量存在明显的差异^[19]。【本研究切入点】前人关于干旱胁迫时段或胁迫程度对小麦成穗和结实影响的研究,多采用主茎或随机采集的茎蘖,而关于主茎和不同蘖位分蘖对拔节期干旱响应差异的研究鲜有报道。【拟解决的关键问题】本试验在盆栽条件下,设置拔节后不同干旱程度及干旱持续时间处理,探索拔节期短时间干旱对主茎和不同蘖位分蘖成穗与结实特性、单茎生产力及单位面积籽粒产量的影响,以期为黄淮海地区冬小麦节水栽培提供理论依据和技术支持。

1 材料与方法

1.1 试验设计

试验于2017—2019年小麦生长季,在山东省泰安市岱岳区道朗镇试验基地遮雨棚内进行。试验点位于116°54′E,36°12′N,属于温带大陆性气候,年平均气温13.0—13.6℃。选用冬小麦品种山农29（具有10 500 kg·hm^-2高产潜力）和衡0628（具有9 000 kg·hm^-2高产潜力）为试验材料。采用盆栽方式,试验用土取自本地高产田0—20 cm耕层土,土壤类型为粉壤土。2017—2018年度试验土壤养分含量为有机质15.5 g·kg^-1,全氮0.8 g·kg^-1,碱解氮85 mg·kg^-1,有效磷33.3 mg·kg^-1,速效钾126 mg·kg^-1;2018—2019年度试验土壤养分含量为有机质15.7 g·kg^-1,全氮0.8 g·kg^-1,碱解氮88 mg·kg^-1,有效磷33.0 mg·kg^-1,速效钾124 mg·kg^-1。土壤过5 mm筛后,称取10 kg装入高26 cm、盆口直径30 cm、盆底直径20 cm的棕色聚乙烯塑料盆内,统一压实后测定土壤最大持水量,两年度分别为28.5%和27.8%。每盆施用N、P₂O₅和K₂O含量均为15%的三元复合肥4.5 g,平铺于距盆口10 cm深处,其上覆土10 cm。播种时挑选饱满、无损、发育良好、大小一致的种子,播种深度为3 cm,每盆播9穴,每穴播2粒,三叶一心期定苗至1株/穴（180 株/m²）。拔节期追施尿素（N含量为46%）1.5 g,追肥时用水将尿素溶解后均匀浇灌于盆内。

试验采用随机区组设计,共5个处理,每个处理种植60盆。设置拔节后0—5 d轻度干旱（T1,保持土壤相对含水量为65%—70%,土壤有效含水量为33.4—37.8 mm）、拔节后0—5 d重度干旱（T2,保持土壤相对含水量为45%—50%,土壤有效含水量为15.6—20.1 mm）、拔节后0—10 d轻度干旱（T3,保持土壤相对含水量为65%—70%,土壤有效含水量为33.4—37.8 mm）、拔节后0—10 d重度干旱（T4,保持土壤相对含水量为45%—50%,土壤有效含水量为15.6—20.1 mm）4个干旱处理,以全生育期充分供水处理为对照（CK,保持土壤相对含水量为75%—80%,土壤有效含水量为42.2—46.7 mm）。拔节前5 d开始控水,达到目标含水量时进行相应时间的处理,干旱处理结束后再将土壤含水量恢复至对照水平。采用称重法控制土壤水分,每2 d称重一次,以相邻2次重量的差值确定补灌水量,以保证各盆内土壤有效含水量和土壤相对含水量控制在设定的范围内。其他管理措施尽量保持一致^[19,20]。

2017—2018年度,试验于2017年10月16日播种,于2018年3月21日开始控水,使土壤含水量逐渐下降,T1和T2处理于3月26日至3月31日正式实施干旱处理,3月31日干旱处理结束后恢复至CK水平;T3和T4于3月26日至4月5日正式实施干旱处理,4月5日干旱处理结束后恢复至CK水平;收获时间为2018年6月12日。2018—2019年度,试验于2018年10月8日播种,于2019年3月29日开始控水,使土壤含水量逐渐下降,T1和T2处理于4月2日至4月7日正式实施干旱处理,4月7日干旱处理结束后恢复至CK水平;T3和T4于4月2日至4月12日正式实施干旱处理,4月12日干旱处理结束后恢复至CK水平。

1.2 测定项目与方法

1.2.1 标记主茎和不同蘖位分蘖 自小麦第一个分蘖出现开始,用不同颜色的回形针标记主茎和新出现的分蘖。用O代表主茎,用Ⅰ、Ⅱ、Ⅲ、Ⅳ分别代表由主茎的第1、2、3、4叶的叶腋中长出的分蘖,用Ⅰp和Ⅰ1分别代表由分蘖Ⅰ上长出的第一和第二个分蘖,用Ⅱp代表由分蘖Ⅱ上长出的第一个分蘖^[20]。

1.2.2 茎蘖成穗和结实性状的调查 成熟期按蘖位分样,每处理取6盆（6次重复）,调查主茎及不同蘖位分蘖的成穗和结实情况,计算每盆有效穗数、每穗总小穗数、每穗结实小穗数、每穗穗粒数、单粒重、单穗产量。每处理取10盆,全部收获脱粒后晾晒至含水量达12.5%左右时称重,计算单位面积籽粒产量。

成穗率=每盆主茎或各蘖位分蘖成穗数/对应每盆主茎或各蘖位分蘖最高发生数×100%。

1.2.3 茎蘖幼穗发育进程的调查 拔节期0 d开始,每隔5 d观察一次,共观察3次。每次取有代表性的植株3—5株在体视显微镜下解剖,观察其幼穗发育状态,确定其幼穗发育阶段。小麦幼穗发育时期（表1）参照WADDINGTION的划分方法^[21],发育阶段用W表示。

Table 1
表1
表1小麦幼穗发育时期
Table 1Spike developmental score of wheat

幼穗发育时期及特征 Spike developmental stage and its characteristic	发育阶段 Developmental score
生长锥伸长期 Transition apex	1.5
单棱期 Early double ridge stage	2
二棱期 Double ridge stage	2.5
颖片原基分化期 Glume primordium present	3
外稃原基分化期 Lemma primordium present	3.25
小花原基分化期 Floret primordium present	3.5
雄蕊原基分化期 Stamen primordium present	4
雌蕊原基分化期 Pistii primordium present	4.25
心皮原基分化期 Carpel primordium present	4.5
三心皮包围胚珠期 Carpel extending round three sides of ovule	5
花柱管闭合,子房仅顶部保持开放状态 Stylar canal closing; ovarian cavity enclosed on all sides but still open above	5.5
花柱管开口,柱头突起形成 Stylar canal remaining as a narrow opening; two short round style primordia present	6
花柱开始伸长 Style begin elongation	6.5
柱头上分化出柱头分枝 Stigmatic branches just differentiating as swollen cell on styles	7
柱头分枝和子房壁上羽毛伸长 Stigmatic branches and hairs on ovary wall elongation	8
柱头分枝和子房壁上羽毛继续伸长,柱头分枝缠绕无絮 Stigmatic branches and hairs on ovary wall continue to elongate; stigmatic branches from a tangled mass	8.5
花柱和柱头分枝直立,羽状柱头分化 Styles and stigmatic branches erect; stigmatic hairs differentiating	9
花柱和柱头分枝向外展开,羽状柱头发育完全 Styles and stigmatic branches spreading outwards. Stigmatic hairs well developed	9.5
花柱向外弯曲,柱头分枝伸展,花粉落在羽状柱头上 Styles curved outwards and stigmatic branches spread wide; pollen grains an well-developed stigmatic hairs	10

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1.3 数据分析

采用Microsoft Excel 2013和DPS软件进行数据处理,采用LSD最小显著差异比较法进行差异显著性检验。

2 结果

2.1 主茎及不同蘖位分蘖幼穗发育进程对拔节期阶段性干旱的响应

衡0628茎蘖幼穗发育进程与山农29相近,部分分蘖稍有滞后;分蘖幼穗发育进程滞后于主茎;随蘖位的升高,主茎与分蘖幼穗发育进程差异逐渐拉大（表2）。2017—2018年度,拔节后0—5 d干旱,主茎穗分化进程处于W5—W6阶段,Ⅰ和Ⅱ多处于W4.25—W5.5阶段或W4—W4.5阶段,Ⅲ、Ⅰp和Ⅳ多处于W4—W5或W4—W4.25阶段,高位蘖（Ⅱp和Ⅰ1）多处于W3.5—W4.25或W3—W4阶段;拔节后10 d,O和Ⅰ蘖穗分化进入W7期,Ⅱ、Ⅲ和Ⅰp蘖多处于W6.5期,IV和Ⅱp多处于W6期,I1蘖多处于W5.5期。2018—2019年度,拔节后干旱处理期间,主茎和各蘖位分蘖穗分化所处的阶段与前一年度相近,趋势基本一致。与充分供水相比,轻度干旱对茎蘖穗分化发育进程无显著影响,重度干旱条件下,低位蘖（Ⅲ、Ⅰp）和高位蘖（Ⅳ）幼穗发育进程有所延缓。

Table 2
表2
表2拔节期阶段性干旱对小麦主茎及不同蘖位分蘖幼穗所处的发育进程阶段的影响
Table 2Effects of staged drought at jointing on development process of young spike of main stem and tillers at different positions of wheat

年份 Year	品种 Cultivar	处理 Treatment	拔节0 d 0 days after jointing stage								拔节5 d 5 days after jointing stage								拔节10 d 10 days after jointing stage
			O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1
2017- 2018	山农29 Shannong29	CK	5	4.5	4.25	4	4	4	3.5	3.5	6	5.5	5	5	5	4.25	4.25	4	7	7	7	6.5	6.5	6	6	5.5
		T1	5	4.5	4.25	4	4	4	3.5	3.5	6	5.5	5.5	5	5	4.5	4.25	4	7	7	7	6.5	6.5	6	6	5.5
		T2	5	4.5	4.25	4	4	4	3.5	3.5	6	5	5	5	4.5	4.5	4.25	4	7	7	6.5	6	6	6	6	5.5
		T3	5	4.5	4.25	4	4	4	3.5	3.5	6	5.5	5.5	5	5	4.5	4.25	4	7	7	7	6.5	6.5	6.5	6	6
		T4	5	4.5	4.25	4	4	4	3.25	3.25	6	5	5	5	4.5	4.5	4.25	4	7	7	6	6	6	6	5.5	5
	衡0628 Heng0628	CK	5	4.25	4	4	4	4	3.25	3.25	5.5	5.5	4.5	4.5	4.25	4.25	4	4	7	7	6.5	6.5	6.5	6.5	6	5.5
		T1	5	4.25	4	4	4	4	3.25	3.25	5.5	5.5	4.5	4.25	4.25	4.25	4	4	7	7	6.5	6.5	6.5	6.5	6.5	5.5
		T2	5	4	4	4	4	4	3.25	3.25	5.5	5	4.5	4.25	4.25	4	4	4	7	7	6.5	6.5	6.5	6	6	5.5
		T3	5	4.25	4	4	4	4	3.25	3.25	5.5	5.5	4.5	4.25	4.25	4.25	4	4	7	7	6.5	6.5	6.5	6.5	6	5.5
		T4	5	4.25	4	4	4	4	3.25	3.25	5.5	5	4.5	4.25	4.25	4	4	4	7	7	6.5	6	6	5.5	5.5	5
2018- 2019	山农29 Shannong29	CK	4.5	4.25	4	4	4	3.5	3.5	3.25	6	5.5	5	5	5	4.5	4.25	4.25	7	7	6.5	6.5	6.5	6	5.5	5
		T1	4.5	4.25	4	4	4	3.5	3.5	3.25	6	5.5	5.5	5	5	4.5	4.25	4.25	7	7	6.5	6.5	6.5	6	5.5	5
		T2	4.5	4.25	4	4	4	3.5	3.5	3.25	6	5	5	5	4.5	4.5	4.25	4	7	7	6.5	6	6	6	5.5	5
		T3	4.5	4.25	4	4	4	3.5	3.5	3.25	6	5.5	5.5	5	5	4.5	4.25	4.25	7	7	6.5	6.5	6.5	6.5	5.5	5
		T4	4.5	4.25	4	4	3.5	3.5	3.25	3	6	5	5	5	4.5	4.5	4.25	4	7	7	6.5	6	6	5.5	5.5	5
	衡0628 Heng0628	CK	4.25	4.25	4	4	4	4	3.25	3.25	6	5.5	4.5	4.5	4.25	4.25	4	4	7	6.5	6.5	6.5	6.5	6	5	5
		T1	4.25	4.25	4	4	4	4	3.25	3.25	6	5.5	4.5	4.25	4.25	4.25	4	4	7	6.5	6.5	6.5	6.5	6	5.5	5
		T2	4.25	4.25	4	4	4	4	3.25	3.25	6	5	4.5	4.25	4	4	4	3.5	7	7	6.5	6.5	6.5	6	5	5
		T3	4.25	4.25	4.25	4	4	4	3.25	3.25	6	5.5	4.5	4.25	4.25	4.25	4	4	7	7	6.5	6.5	6.5	6.5	5	5
		T4	4.25	4.25	4.25	4	4	4	3.25	3.25	6	5	4.5	4.25	4	4	4	3.5	7	7	6	6	6	5	5	5

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2.2 主茎及不同蘖位分蘖成穗率对拔节期阶段性干旱的响应

山农29茎蘖成穗率较衡0628高;主茎与不同蘖位分蘖间比较,其成穗率2个品种均表现为O≥Ⅰ≥Ⅱ>Ⅲ>Ⅰp>Ⅳ>Ⅱp>Ⅰ1（表3）。拔节期不同程度干旱对2个小麦品种茎蘖成穗率的影响在2个年度表现一致,拔节期阶段性干旱对主茎成穗率的影响较小,各处理主茎成穗率均为100%,但对分蘖成穗率的影响较大,随着干旱程度的加剧和干旱时间的延长,低位蘖（Ⅲ和Ⅰp）和高位蘖（Ⅳ）成穗率迅速下降,高位蘖（Ⅱp和Ⅰ1）成穗率呈先增加后降低的趋势。与CK相比,T1和T2处理低位蘖（Ⅰ和Ⅱ）成穗率降低,高位蘖成穗率有所增加;T3处理的低位蘖成穗率显著降低,高位蘖成穗率保持不变;T4处理除O和Ⅰ蘖外,其余分蘖成穗率均显著下降。品种之间比较,低位蘖成穗率在拔节期各干旱条件下的平均降幅表现为衡0628（下降6.45个百分点）>山农29（下降5.19个百分点）,高位蘖的平均增幅表现为衡0628（增加0.14个百分点）<山农29（增加3.79个百分点）。上述结果说明山农29对拔节期阶段性干旱的抗性高于衡0628,拔节后0—5 d轻度干旱处理可增加高位蘖成穗率,提高单位面积穗数。

Table 3
表3
表3拔节期阶段性干旱对小麦主茎及不同蘖位分蘖的成穗率的影响
Table 3Effects of staged drought at jointing on percentage of ear bearing main stem and tillers at different positions of wheat (%)

年份 Year		品种 Cultivar		处理 Treatment		O		Ⅰ		Ⅱ		Ⅲ		Ⅰp		Ⅳ		Ⅱp		Ⅰ1
2017-2018		山农29 Shannong29		CK		100.00a		100.00a		100.00a		79.37a		68.89a		37.78a		22.22bc		17.78b
				T1		100.00a		100.00a		100.00a		75.56ab		64.44a		40.00a		33.33a		26.67a
				T2		100.00a		100.00a		97.78ab		73.33b		55.56b		35.56b		31.11ab		24.44ab
				T3		100.00a		100.00a		95.56ab		68.89c		53.33b		33.33b		26.67abc		22.22ab
				T4		100.00a		97.78a		93.33b		66.67c		48.89c		26.67c		20.00c		12.70c
		衡0628 Heng0628		CK		100.00a		100.00a		100.00a		75.56a		75.56a		40.00a		20.00a		17.78a
				T1		100.00a		100.00a		97.78a		73.33a		73.33a		42.22a		26.67a		22.22a
				T2		100.00a		100.00a		97.78a		68.89b		57.78b		35.56b		24.44a		22.22a
				T3		100.00a		100.00a		95.24ab		66.67b		53.33bc		33.33bc		22.22a		20.00a
				T4		100.00a		95.56a		91.11b		66.67b		40.00c		31.11c		17.78a		11.11b
2018-2019		山农29 Shannong29		CK		100.00a		100.00a		98.41a		79.37a		52.38a		36.51a		11.11b		4.76b
				T1		100.00a		100.00a		96.83ab		77.78a		55.56a		36.51a		20.63a		11.11a
				T2		100.00a		98.41a		95.24ab		73.33b		50.79b		33.33b		17.46a		7.94ab
				T3		100.00a		98.41a		93.65ab		69.84b		47.62c		31.75b		12.70b		4.76b
				T4		100.00a		96.83a		92.06b		55.56c		46.03c		26.98c		7.94c		1.59c
		衡0628 Heng0628		CK		100.00a		98.41a		96.83a		71.43a		44.44b		39.68a		12.70b		6.35b
				T1		100.00a		98.41a		95.24a		68.25ab		49.21a		39.68a		15.87a		12.70a
				T2		100.00a		95.24a		93.65a		65.08b		44.44b		33.33b		12.70b		9.52b
				T3		100.00a		95.24a		92.06ab		60.32c		41.27c		26.98c		12.70b		7.94b
				T4		100.00a		93.65a		90.48b		57.14d		30.16d		20.63d		3.17c		1.59c

同一年份同一品种同列数据后不同小写字母表示处理间在0.05水平差异显著。下同
Values followed different letters of the same variety and column in the same year indicate significant differences at 0.05 level. The same as below

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2.3 主茎及不同蘖位分蘖可见总小穗数对拔节期阶段性干旱的响应

衡0628茎蘖可见总小穗数高于山农29;随蘖位的升高,可见小穗数呈明显的下降趋势,相邻蘖位之间差异不显著（表4）。拔节期不同程度干旱对2个小麦品种茎蘖可见总小穗数的影响在2个年度表现一致,与CK相比,T1和T2处理茎蘖可见总小穗数和平均单茎可见总小穗数呈降低趋势,且2个处理间差异不显著;T3和T4处理各蘖位分蘖小穗数显著降低,主茎小穗数未受显著影响,单茎平均可见小穗数显著降低,2个处理间差异不显著。山农29的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1的可见总小穗数在T3和T4处理下,平均降低幅度为5.64%、6.63%、6.07%、5.79%、5.57%、6.40%、5.18%和1.93%,单茎平均小穗数降低了5.03%,衡0628的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1的可见总小穗数在T3和T4处理下,平均降低幅度为6.51%、7.27%、7.17%、7.20%、6.76%、6.46%、6.05%和1.88%,单茎平均小穗数降低了5.77%。上述结果说明,与全生育期充分供水相比,拔节后0—5 d轻度和重度干旱对2个小麦品种茎蘖小穗数无显著影响,拔节后0—10 d干旱显著降低各成穗分蘖的小穗数,且以低位蘖（Ⅲ和Ⅰp）和高位蘖（Ⅳ）降低幅度较大。2个品种之间比较,山农29各干旱处理茎蘖小穗数平均降幅（3.66%）小于衡0628（4.44%）。

Table 4
表4
表4拔节期阶段性干旱对小麦主茎及不同蘖位分蘖可见小穗数的影响
Table 4Effects of staged drought at jointing on spikelet number of main stem and tillers at different positions of wheat (spikelets/spike)

年份 Year	品种 Cultivar	处理 Treatment	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1	平均单茎可见小穗数 Average visible spikelet number per spike
2017-2018	山农29 Shannong29	CK	18.2a	18.1a	17.5a	17.1a	16.5a	16.3a	16.1a	15.7a	17.3a
		T1	18.2a	17.9a	17.1a	16.9a	16.3a	16.1a	16.1a	15.5a	17.1a
		T2	18.0a	17.7ab	17.0ab	16.6ab	16.0ab	15.9ab	15.8a	15.3a	16.9ab
		T3	17.6a	17.2b	16.6b	16.2b	15.6b	15.4b	15.2b	14.9b	16.5b
		T4	17.3a	17.0b	16.3b	15.9b	15.5b	15.4b	15.0b	14.8b	16.4b
	衡0628 Heng0628	CK	20.0a	19.3a	18.8a	18.2a	17.7a	17.0a	16.8a	16.5a	18.6a
		T1	19.7a	19.0a	18.4a	17.8a	17.3ab	16.5a	16.6a	16.4a	18.2a
		T2	19.4a	18.5ab	18.1ab	17.5ab	17.1ab	16.3ab	16.4a	16.1a	17.9ab
		T3	19.3a	18.3b	17.6b	17.0b	16.5bc	15.9b	15.8b	15.6b	17.6b
		T4	19.1a	17.9b	17.3b	16.7b	16.2c	15.8b	15.5b	15.4b	17.4b
2018-2019	山农29 Shannong29	CK	19.5a	19.1a	17.9a	17.5a	17.2a	16.9a	16.7a	16.2a	18.2a
		T1	19.4a	19.0a	17.8a	17.3a	17.1a	16.8ab	16.5a	16.0a	18.0a
		T2	19.2a	18.8ab	17.4ab	17.1ab	16.7ab	16.4ab	16.3a	15.6a	17.8ab
		T3	18.9a	18.2b	16.8b	16.6b	16.4b	16.1b	15.7a	15.4a	17.3b
		T4	18.7a	17.8b	16.4b	16.3b	16.0b	15.8b	15.5b	15.4b	17.2b
	衡0628 Heng0628	CK	21.2a	20.6a	19.7a	19.5a	19.2a	18.5a	18.1a	17.4a	20.0a
		T1	20.9a	20.5a	19.3a	19.1a	18.9ab	18.2a	17.9a	17.0ab	19.5a
		T2	20.8a	20.1ab	19.0ab	18.9ab	18.5ab	17.9ab	17.4ab	16.8ab	19.4ab
		T3	20.6a	19.4b	18.5b	18.4b	18.1b	17.4b	17.1b	16.5b	18.9b
		T4	20.3a	19.0b	18.0b	17.9b	17.7b	17.1b	16.9b	16.2b	18.8b

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2.4 主茎及不同蘖位分蘖结实小穗数对拔节期阶段性干旱的响应

衡0628茎蘖的结实小穗数高于山农29;随蘖位的升高,结实小穗数呈降低趋势。山农29的Ⅲ和Ⅰp之间差异显著,其余相邻蘖位之间差异不显著;衡0628的O和Ⅰ之间、Ⅰ和Ⅱ之间差异显著,其余相邻蘖位之间差异不显著。拔节期不同程度干旱对2个小麦品种各茎蘖结实小穗数的影响表现一致,其中对主茎的影响较小,对分蘖的影响较大（表5）。与CK相比,T1处理对茎蘖结实小穗数和平均单茎结实小穗数无显著影响;T2处理显著降低除Ⅰ之外的低位蘖结实小穗数,对主茎与高位蘖结实小穗数无显著影响;T3和T4处理显著降低茎蘖结实小穗数和单茎平均结实小穗数,2个处理间无显著差异。山农29的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1结实小穗数在T3和T4处理下,较CK平均降幅为6.47%、7.76%、10.76%、9.36%、9.20%、8.44%、9.70%和9.32%,平均单茎结实小穗数降低7.94%;衡0628的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1结实小穗数在T3和T4处理下,较CK平均降幅为6.66%、9.31%、11.21%、10.56%、11.04%、11.18%、10.82%和10.42%,平均单茎结实小穗数降低8.62%。上述结果说明,与全生育期充分供水相比,拔节后0—5 d轻度干旱对茎蘖的结实小穗数无显著影响,短期重度干旱显著降低中位蘖的结实小穗数;拔节后0—10 d轻度干旱和重度干旱处理均显著降低了主茎和分蘖的结实小穗数,且低位蘖（Ⅲ和Ⅰp）和高位蘖（Ⅳ）较其他分蘖大。品种之间比较,山农29各干旱处理茎蘖结实小穗数平均降幅（5.95%）小于衡0628（6.95%）。

Table 5
表5
表5拔节期阶段性干旱对小麦主茎及不同蘖位分蘖结实小穗数的影响
Table 5Effects of staged drought at jointing on fruiting spikelet number of main stem and tillers at different positions of wheat (spikelets/spike)

年份 Year	品种 Cultivar	处理 Treatment	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1	平均单茎结实小穗数 Average fruiting spikelet number per spike
2017-2018	山农29 Shannong29	CK	17.9a	17.6a	16.7a	15.9a	14.8a	14.1a	13.8a	13.1a	16.3a
		T1	17.7ab	17.4a	16.3ab	15.6ab	14.5ab	13.9ab	13.6a	13.1a	15.9ab
		T2	17.5ab	16.9ab	15.8b	15.1b	14.0b	13.5b	13.3ab	12.6a	15.6ab
		T3	17.1bc	16.5bc	15.5bc	14.8bc	13.9bc	13.3bc	12.7b	12.3b	15.3bc
		T4	16.4c	15.9c	14.5c	14.0c	13.1c	12.6c	12.1c	11.5c	14.7c
	衡0628 Heng0628	CK	18.3a	17.3a	16.3a	15.7a	15.2a	14.5a	14.1a	13.5a	16.3a
		T1	18.0ab	16.8ab	15.9ab	15.2ab	14.7ab	13.9ab	13.8a	13.3ab	15.8ab
		T2	17.7ab	16.3ab	15.4b	14.7b	14.3b	13.7b	13.4ab	12.9ab	15.5ab
		T3	17.4b	16.2bc	15.0bc	14.4bc	13.9bc	13.3bc	12.9bc	12.5b	15.2bc
		T4	16.8b	15.3c	14.2c	13.7c	13.0c	12.7c	12.3c	11.8c	14.6c
2018-2019	山农29 Shannong29	CK	19.2a	18.5a	17.4a	16.6a	15.6a	14.9a	14.6a	13.7a	17.3a
		T1	19.1ab	18.4a	17.2ab	16.4ab	15.4ab	14.8ab	14.3a	13.3a	17.0ab
		T2	18.7ab	17.8ab	16.5b	15.7b	14.7b	14.2b	14.0a	13.1a	16.5ab
		T3	18.3b	17.5bc	15.9bc	15.4bc	14.4bc	13.8bc	13.5bc	12.6bc	16.1bc
		T4	17.6b	16.7c	14.8c	14.7c	13.8c	13.4c	13.0c	12.2c	15.6c
	衡0628 Heng0628	CK	19.2a	18.1a	17.1a	16.5a	16.1a	15.4a	15.0a	14.3a	17.3a
		T1	19.0a	17.9a	16.7ab	16.1ab	15.8ab	14.9ab	14.8a	14.0a	16.8ab
		T2	18.5ab	17.2ab	16.1b	15.6b	15.2b	14.5b	14.3ab	13.6ab	16.5ab
		T3	18.2b	16.8bc	15.4bc	15.1bc	14.8bc	14.0bc	13.7b	13.1bc	16.0bc
		T4	17.3b	15.9c	14.7c	14.4c	14.0c	13.1c	13.0b	12.5c	15.6c

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2.5 主茎及不同蘖位分蘖穗粒数对拔节期阶段性干旱的响应

衡0628主茎穗和分蘖穗的穗粒数多于山农29,主茎穗粒数多于分蘖。拔节期不同程度干旱对2个小麦品种茎蘖穗粒数均有明显的影响,2年结果趋势一致,均表现为随干旱程度增强和干旱时间延长,穗粒数呈逐渐降低的趋势（表6）。与CK相比,T1处理各茎蘖穗粒数无显著变化;T2处理主茎和高位蘖穗粒数无显著变化,低位蘖穗粒数显著降低;T3和T4处理主茎和各成穗分蘖的穗粒数均显著降低。T1与T2处理间无显著差异,T2与T3处理间无显著差异,但均显著大于T4处理。各茎蘖之间比较,2个品种在T2、T3和T4处理下均表现为主茎穗粒数平均降幅（7.02%）<高位蘖平均降幅（8.52%）<低位蘖平均降幅（9.77%）。上述结果表明,与全生育期充分供水相比,拔节后0—5 d轻度干旱对各茎蘖穗粒数无显著影响,但重度干旱减少低位蘖和中位蘖穗粒数;拔节后0—10 d轻度和重度干旱均显著降低主茎和分蘖的穗粒数,且中位蘖降低幅度较大。品种之间比较,各干旱处理下山农29主茎穗和分蘖穗穗粒数平均降幅（6.73%）小于衡0628（7.93%）。

Table 6
表6
表6拔节期阶段性干旱对小麦主茎及不同蘖位分蘖穗粒数的影响
Table 6Effects of staged drought at jointing on grain number of main stem and tillers at different positions of wheat (kernels/spike)

年份 Year	品种 Cultivar	处理 Treatment	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1
2017-2018	山农29 Shannong29	CK	41.6a	37.4a	34.8a	32.2a	29.5a	27.7a	25.0a	22.8a
		T1	40.9a	36.8ab	33.9ab	31.5ab	29.2ab	27.4ab	25.7a	23.0a
		T2	40.6ab	35.3bc	32.4bc	30.1bc	27.8b	26.1b	24.0ab	22.0ab
		T3	39.4b	35.0c	31.9c	29.6c	27.6b	26.0b	23.6b	21.5b
		T4	37.3c	31.9d	29.1d	27.3d	25.3c	23.7c	22.2c	20.3c
	衡0628 Heng0628	CK	43.5a	40.8a	37.1a	34.8a	32.4a	29.0a	26.5a	25.6a
		T1	42.4a	39.5ab	36.1ab	33.8ab	31.2ab	27.6ab	26.6a	25.5a
		T2	41.5ab	38.2b	34.5bc	32.3bc	30.4bc	27.1b	25.2a	24.4a
		T3	40.3bc	37.7b	34.0c	31.7c	29.6c	26.4b	24.8b	23.5b
		T4	38.8c	34.4c	31.0d	28.9d	26.9d	24.0c	23.5c	22.2c
2018-2019	山农29 Shannong29	CK	47.7a	43.6a	41.9a	38.9a	34.0a	31.9a	29.2a	27.2a
		T1	47.2a	43.8a	41.3ab	38.2ab	33.8ab	31.3ab	28.9a	26.9a
		T2	46.6ab	41.4b	39.3bc	36.7bc	32.1bc	30.1bc	28.4ab	26.3ab
		T3	45.2b	41.3b	38.0c	35.3c	31.9c	29.2c	27.5b	25.1b
		T4	42.4c	38.7c	35.6d	33.7d	29.4d	27.9d	26.0c	23.7c
	衡0628 Heng0628	CK	49.1a	45.3a	42.4a	39.1a	36.5a	33.1a	31.0a	28.6a
		T1	48.5a	44.6ab	41.3ab	38.3ab	36.7a	32.5ab	31.1a	28.9a
		T2	47.1ab	42.8bc	39.6bc	36.6bc	34.0b	30.9bc	29.4ab	27.5a
		T3	45.6b	42.0c	38.4c	35.8c	33.4b	29.7c	28.5b	26.2b
		T4	43.0c	39.5d	35.6d	32.4d	30.6c	27.6d	27.2c	24.3c

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2.6 拔节期阶段性干旱对主茎及不同蘖位分蘖单粒重的影响

山农29主茎穗和分蘖穗平均单粒重高于衡0628;随蘖位的升高,单粒重呈逐渐降低趋势,相邻蘖位之间差异不显著。拔节期不同程度干旱对2个小麦品种茎蘖单粒重的影响在2个年度表现一致（表7）。与CK相比,T1处理主茎穗和分蘖穗的平均单粒重无显著变化;T2处理下,山农29主茎穗和分蘖穗单粒重无显著变化,而衡0628各蘖位分蘖单粒重显著降低;T3和T4处理主茎和各蘖位分蘖单粒重均显著降低。T2与T3处理间无显著差异,T3与T4处理间无显著差异,T2处理显著大于T4处理。山农29的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1结实小穗数在T3和T4处理下,较CK平均降幅为5.96、6.73%、8.05%、9.13%、8.84%、8.49%、7.78%和7.04%;衡0628的O、Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅱp和Ⅰ1单粒重在T3和T4处理下,较CK平均降幅为6.07%、7.79%、8.94%、9.99%、9.29%、8.90%、8.56%和8.26%。上述结果表明,与全生育期充分供水相比,拔节后0—5 d轻度干旱对2个小麦品种茎蘖单粒重无显著影响;拔节后0—10 d干旱显著降低主茎穗和各蘖位分蘖穗的单粒重,且以Ⅲ、Ⅰp和Ⅳ蘖单粒重下降幅度较大。2个品种之间比较,山农29各干旱处理茎蘖单粒重平均降幅（5.55%）小于衡0628（7.71%）。

Table 7
表7
表7拔节期阶段性干旱对小麦主茎及不同蘖位分蘖单粒重的影响
Table 7Effects of staged drought at jointing on single-kernel weight of main stem and tillers at different positions of wheat (mg/kernel)

年份 Year	品种 Cultivar			处理 Treatment	O		Ⅰ		Ⅱ		Ⅲ		Ⅰp		Ⅳ		Ⅱp		Ⅰ1
2017-2018	山农29 Shannong29			CK	51.94a		50.34a		49.36a		48.17a		46.56a		45.46a		44.70a		44.00a
				T1	50.83ab		49.50ab		48.32a		47.89a		45.34a		45.39a		43.85a		43.09ab
				T2	49.85ab		49.12ab		47.89ab		45.93ab		44.57ab		44.05ab		43.17ab		42.54ab
				T3	49.28b		47.91bc		46.06bc		44.16bc		43.01bc		42.35bc		42.11bc		41.72bc
				T4	48.85b		46.05c		44.78c		43.21c		41.80c		40.92c		40.70c		40.33c
	衡0628 Heng0628			CK	45.27ab		43.70a		42.37a		42.33a		41.57a		40.67a		39.90a		39.30a
				T1	46.43a		45.88a		43.57a		43.52a		42.05a		41.88a		41.51a		39.98a
				T2	43.07b		41.38b		39.82b		39.30b		39.52b		38.59b		37.85b		37.06b
				T3	43.00b		41.23bc		39.05bc		38.75bc		38.46bc		37.85bc		37.72b		36.11b
				T4	41.95b		39.38c		37.82c		37.32c		37.15c		36.42c		35.67b		35.10b
2018-2019	山农29 Shannong29			CK	50.77a		50.01a		48.28a		46.39a		45.60a		44.39a		43.20a		41.25a
				T1	50.37a		48.26a		47.06a		45.44a		44.14a		43.07a		42.91ab		40.29ab
				T2	49.16ab		47.83ab		46.32ab		44.48ab		43.56ab		42.65ab		41.30ab		39.33ab
				T3	48.14bc		47.59bc		45.11bc		43.13bc		42.44bc		41.49bc		40.82b		39.04b
				T4	46.91c		45.65c		43.62c		41.34c		40.80c		39.70c		38.50c		37.42c
	衡0628 Heng0628			CK	46.89ab		46.33a		45.11a		43.12a		41.21a		39.89a		39.72a		38.11a
				T1	48.17a		47.81a		46.88a		44.50a		42.94a		40.74a		40.22a		38.23a
				T2	45.45bc		44.05b		42.20b		40.31b		39.12b		37.71b		37.56b		36.05b
				T3	44.79c		43.42bc		41.53bc		39.67bc		37.79bc		37.04bc		36.97bc		35.30bc
				T4	43.39c		42.00c		40.23c		38.09c		36.78c		35.47c		35.25c		34.61c

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2.7 拔节期阶段性干旱对主茎及不同蘖位分蘖单穗产量的影响

山农29主茎穗和分蘖穗的单穗产量高于衡0628;随蘖位的升高,单穗产量呈逐渐降低趋势,相邻蘖位之间差异显著。拔节期不同程度干旱对2个小麦品种主茎穗和分蘖穗的单穗产量的影响2年结果趋势一致（表8）。与CK相比,T1处理主茎穗和分蘖穗的单穗产量无显著变化;T2处理主茎穗和高位蘖穗的单穗产量无显著变化,低位蘖穗的单穗产量显著降低,平均单穗产量显著降低;T3和T4处理的主茎穗和分蘖穗单穗产量均显著降低,T3处理显著大于T4处理,但与T2处理无显著差异。T2、T3和T4处理分别与CK相比,山农29平均单穗产量下降幅度分别为5.14%—5.46%、8.53%—8.55%和13.36%—16.68%,衡0628平均单穗产量下降幅度分别为5.45%—6.24%、9.20%—9.69%和12.72%—19.25%。上述结果说明,与全生育期充分供水相比,拔节后0—5 d轻度干旱对2个小麦品种主茎穗和分蘖穗的单穗产量无显著影响,重度干旱降低低位蘖和中位蘖穗的单穗产量;拔节后0—10 d轻度和重度干旱均显著降低主茎穗和分蘖穗单穗产量,且以Ⅲ、Ⅰp和Ⅳ蘖穗单穗产量下降幅度最大。2个品种比较,山农29各干旱处理主茎穗和分蘖穗单穗产量平均降幅（8.09%）小于衡0628（12.37%）。

Table 8
表8
表8拔节期阶段性干旱对小麦主茎及不同蘖位分蘖单穗产量的影响
Table 8Effects of staged drought at jointing on grain yield of main stem and tillers at different positions of wheat (g/stem)

年份 Year	品种 Cultivar	处理 Treatment	O	Ⅰ	Ⅱ	Ⅲ	Ⅰp	Ⅳ	Ⅱp	Ⅰ1	平均单穗产量 Average yield per spike
2017-2018	山农29 Shannong29	CK	1.75a	1.54a	1.49a	1.21a	1.14a	1.03a	0.91a	0.86a	1.38a
		T1	1.73a	1.51ab	1.42ab	1.16ab	1.10ab	1.00ab	0.94a	0.85a	1.33ab
		T2	1.69ab	1.48bc	1.39bc	1.14bc	1.07bc	0.97bc	0.86ab	0.83ab	1.30bc
		T3	1.63b	1.42c	1.32c	1.11c	1.03c	0.93c	0.83b	0.78b	1.26c
		T4	1.55c	1.34d	1.03d	0.95d	0.91d	0.82d	0.78c	0.75c	1.15d
	衡0628 Heng0628	CK	1.65a	1.44a	1.37a	1.18a	1.05a	0.94a	0.88a	0.76a	1.28a
		T1	1.67a	1.47a	1.38a	1.18a	1.07a	0.96a	0.90a	0.79a	1.30a
		T2	1.59ab	1.33b	1.26b	1.09b	0.97b	0.86b	0.83ab	0.73ab	1.21b
		T3	1.52b	1.28b	1.21b	1.04b	0.92b	0.81b	0.75b	0.66b	1.16b
		T4	1.38c	1.16c	0.92c	0.91c	0.83c	0.74c	0.70c	0.59c	1.03c
2018-2019	山农29 Shannong29	CK	1.97a	1.75a	1.62a	1.37a	1.29a	1.06a	0.93a	0.86a	1.58a
		T1	1.98a	1.72ab	1.59ab	1.36ab	1.29ab	1.05ab	0.91a	0.85a	1.54ab
		T2	1.92ab	1.67bc	1.53bc	1.30bc	1.22bc	1.00bc	0.89ab	0.82ab	1.50bc
		T3	1.85b	1.60c	1.46c	1.23c	1.17c	0.95c	0.85b	0.79b	1.45c
		T4	1.77c	1.54d	1.23d	1.14d	1.04d	0.89d	0.79c	0.76c	1.37d
	衡0628 Heng0628	CK	1.86a	1.69a	1.54a	1.32a	1.17a	0.97a	0.88a	0.80a	1.50a
		T1	1.91a	1.74a	1.55a	1.35a	1.17a	0.99a	0.91a	0.81a	1.50a
		T2	1.80ab	1.60b	1.41b	1.20b	1.08b	0.90b	0.84ab	0.76ab	1.41b
		T3	1.73bc	1.54b	1.34b	1.15b	1.03b	0.86b	0.80b	0.73b	1.35b
		T4	1.67c	1.47c	1.15c	1.04c	0.95c	0.80c	0.72c	0.69c	1.31c

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2.8 拔节期阶段性干旱对单位面积籽粒产量及其构成因素的影响

山农29单位面积产量较衡0628高。拔节期不同程度干旱对2个小麦品种单位面积籽粒产量及其构成因素的影响2个年度表现一致,随干旱程度加剧和干旱时间延长,穗数、穗粒数和千粒重呈逐渐降低的趋势（表9）。与CK相比,T1处理单位面积产量无显著变化,T2、T3和T4处理单位面积产量均显著降低。T2处理与CK相比,山农29穗粒数显著降低,衡0628穗粒数和千粒重均显著降低。T3和T4处理与CK相比,穗数、穗粒数和千粒重均显著降低,以T4处理降低幅度最大。上述结果表明,与充分供水相比,拔节后0—5 d轻度干旱对2个小麦品种单位面积籽粒产量及其构成因素无显著影响,重度干旱显著降低2个小麦品种单位面积籽粒产量;拔节后0—10 d轻度和重度干旱,2个小麦品种穗数、穗粒数、千粒重和单位面积籽粒产量均显著降低。2个品种间比较,山农29各干旱处理单位面积籽粒产量平均降幅（12.52%）小于衡0628（17.62%）。

Table 9
表9
表9拔节期阶段性干旱对单位面积籽粒产量及其构成因素的影响
Table 9Effects of staged drought at jointing on yield and its component factors of wheat

处理 Treatments		2017-2018				2018-2019
		穗数 Spike per pot	穗粒数 Kernel per spike	千粒重 1000-kernel weight (g)	产量 Yield (g/pot)	穗数 Spike per pot	穗粒数 Kernel per spike	千粒重 1000-kernel weight (g)	产量 Yield (g/pot)
山农29 Shannong29	CK	47.34a	34.17a	45.97a	65.50a	43.43ab	40.91a	46.69a	68.48a
	T1	48.60a	33.52ab	44.98a	64.67a	44.86a	40.05ab	46.22a	68.87a
	T2	46.60ab	32.22b	44.05ab	60.79b	42.89ab	38.76b	45.03ab	64.86b
	T3	44.80b	31.87b	42.97b	56.78c	41.29b	38.17b	43.70b	59.67c
	T4	42.20c	29.84c	42.05b	48.52d	38.43c	36.26c	42.85b	52.60d
衡0628 Heng0628	CK	47.60a	36.52a	43.77a	61.61a	42.29a	42.09a	46.51a	63.22a
	T1	48.20a	35.34ab	44.72a	62.46a	43.14a	41.16ab	47.01a	64.63a
	T2	45.80ab	34.42b	41.57b	55.19b	40.86ab	39.72bc	43.76b	57.46b
	T3	44.17b	34.82b	40.27b	51.14c	39.57b	38.92c	42.97bc	53.41c
	T4	40.80c	31.75c	39.35b	42.17d	37.22c	37.02c	41.24c	46.75d

同一品种同列数据后不同小写字母表示处理间在0.05水平差异显著
Values followed different letters in the same variety and column are significantly different at 0.05 probability level among the treatments

新窗口打开|下载CSV

3 讨论

3.1 小麦产量构成三因素对拔节期干旱的响应

小麦拔节期是穗、叶、茎等器官同时并进,分蘖迅速向有效和无效两极分化的时期。有研究表明该时期任何程度的干旱均表现出减产效应,穗数和穗粒数显著降低,轻度干旱对千粒重无显著影响,重度干旱亦会导致千粒重显著降低^[15]。另有研究则表明拔节期轻度干旱条件下,穗数显著降低,穗粒数和千粒显著增加,产量显著增加,重度干旱条件下,穗数、穗粒数和千粒重均显著降低,产量显著降低^[22]。上述研究均说明,拔节期的水分条件不仅影响小麦的群体发育和成穗数,对穗粒数甚至粒重亦有显著影响,也反映出旺长小麦通过拔节期适度干旱可在一定程度上优化群体结构,使穗数维持在适宜的范围内,进而通过提高穗粒数和粒重增加籽粒产量。本研究结果表明,拔节后0—5 d期间轻度干旱对2个品种穗数、穗粒数、千粒重和籽粒产量均无显著影响,而拔节后0—10 d期间轻度或重度干旱均导致2个品种产量构成三因素显著降低。说明小麦拔节期短期的轻度干旱并不会对小麦产量构成带来明显的负面影响。进一步分析发现,在拔节后0—10 d内,不同程度干旱对主茎的成穗率无明显影响,但是随着干旱程度的加大和干旱时间的延长,Ⅲ和Ⅰp蘖成穗率迅速下降,而Ⅱp和Ⅰ1蘖成穗率则呈先升高后下降趋势,说明Ⅲ和Ⅰp蘖对该阶段的干旱胁迫相对敏感。Ⅱp和Ⅰ1蘖成穗率的升高在一定程度上可弥补Ⅲ和Ⅰp蘖成穗率下降对单位面积成穗数的影响,这可能是拔节后0—5 d期间轻度干旱条件下,2个品种单位面积穗数无明显变化的原因。

3.2 茎蘖幼穗发育与成穗的关系及其对拔节期干旱的响应

分蘖是否成穗与分蘖自身生长速度、叶龄及幼穗发育程度有关^[23,24]。有研究认为分蘖与主茎的幼穗分化起步差距不能太大,与主茎穗分化起步差距多于两期的分蘖均为无效分蘖,在水分胁迫条件下,与主茎穗分化起步差距超过一期者大多数为无效分蘖^[25]。还有研究表明小花分化期是决定分蘖是否成穗的关键时期,各成穗分蘖进入小花原基分化期的时间相差较小,而无效分蘖进入小花分化期的时间滞后,与成穗分蘖的差距较大^[26]。高翔等^[27]的研究也证明能成穗的Ⅰ、Ⅱ、Ⅲ、Ⅰp、Ⅳ、Ⅰ1蘖与主茎在小花分化期同步,无效分蘖的穗分化在雌雄蕊分化期或更早阶段停止。唐勇金^[28]的研究则表明拔节期幼穗处于小花分化期至雌雄蕊分化期的分蘖均能成穗,主茎出现第6片可见叶时,分蘖叶龄在2.5以上且穗分化处于小花分化期者能成穗;拔节期叶龄在3.0以上且穗分化在小花分化期的二级分蘖少数能成穗。本研究结果表明,主茎O和分蘖I在拔节后0 d时处于W4.25（雌蕊原基分化期）至W5（三心皮包围胚珠期）或W4.25（雌蕊原基分化期）至W4.5（心皮原基分化期）,在拔节10 d后二者仍保持一致,成穗率无显著差异;分蘖穗分化在拔节后0 d时与主茎差距大于0.5期者成穗率显著低于主茎;虽然Ⅱ蘖之后发生的相邻分蘖,穗分化差异较小,但是成穗率彼此间存在显著差异;在拔节期重度干旱条件下,分蘖Ⅲ和Ip的幼穗发育进程迟缓,成穗率迅速下降。

3.3 茎蘖幼穗发育与结实的关系及其对拔节期干旱的响应

二棱期至顶端小穗形成是小穗发育的关键时期,在该阶段,随干旱胁迫程度的增加,不孕小穗率呈升高趋势^[14]。自顶端小穗形成到开花期是小花发育的关键时期,在该阶段,花粉发育早期尤其是小孢子母细胞减数分裂期易受干旱胁迫的影响,从而导致可育小花数显著降低^[29,30,31]。研究表明小麦在花粉减数分裂期遭受干旱胁迫,叶片水势从-0.8 MPa下降到-2.3 MPa,小花退化数量在70%以上,穗粒数显著降低^[32]。与充分灌水处理相比,水分胁迫程度每增加0.2 MPa,每穗粒数减少12.4%—58.7%,且主要集中在穗顶端和基部位置^[33]。本研究在小麦拔节初期设置阶段性干旱处理,结果表明在拔节后0—10 d内,小麦主茎和不同蘖位分蘖幼穗分化正处于颖片原基分化期（W3）至柱头分支在柱头上分化为肿胀的细胞时期（W7）。在此阶段,主茎和不同蘖位分蘖幼穗发育受干旱胁迫影响的程度存在明显差异。拔节后0—5 d,主茎幼穗处于三心皮包围胚珠期（W5）至花柱作为一个狭窄的开口、两个短的圆形花柱原基期（W6）,无论轻度还是重度干旱,穗粒数均无明显变化;分蘖I和II多处于雌蕊原基分化期（W4.25）至花柱管关闭、子房腔仅顶部保持开放状态期（W5.5）或处于雄蕊原基分化期（W4）至心皮原基分化期（W4.5）,在轻度干旱条件下穗粒数无显著变化,重度干旱条件下穗粒数显著降低;分蘖III、Ip和IV多处于雄蕊原基分化期（W4）至三心皮包围胚珠期（W5）或处于雄蕊原基分化期（W4）至雌蕊原基分化期（W4.25）,在轻度干旱条件下穗粒数无显著变化,重度干旱条件下穗粒数大幅度减少;分蘖IIp和I1蘖多处于小花原基分化期（W3.5）至雌蕊原基分化期（W4.25）或处于颖片原基分化期（W3）至雄蕊原基分化期（W4）,无论轻度还是重度干旱,穗粒数均无明显变化。上述结果说明在小麦拔节初期主茎和各蘖位分蘖受干旱胁迫影响的程度与其幼穗发育所处的时期有关,在小麦幼穗分化的雄蕊原基分化期至三心皮包围胚珠期遭受严重干旱会显著降低其结实性,减少穗粒数,而在该阶段遭受轻度干旱,或在颖片原基分化期之前、三心皮包围胚珠期之后遭受短期（5 d）干旱对其结实性影响较小。

4 结论

小麦主茎和不同蘖位分蘖对拔节期干旱的响应存在明显差异,Ⅲ、Ⅰp和Ⅳ蘖成穗率对该阶段干旱相对敏感,下降幅度较大,这可能与干旱期间主茎和不同蘖位分蘖幼穗分化所处的时期不同有关,在小花原基分化期至雌雄蕊原基分化期受胁迫影响最大。在小麦幼穗分化的雄蕊原基分化期至三心皮包围胚珠期遭受阶段性重度干旱（土壤相对含水量为45%—50%,土壤有效含水量为15.6—20.1 mm）对穗粒数有显著影响,而在该阶段遭受轻度干旱（土壤相对含水量65%—70%,土壤有效水含量33.4—37.8 mm）或在颖片原基分化期之前或在三心皮包围胚珠期之后遭受短期（5 d）的重度干旱对穗粒数影响都较小。在群体层面上,拔节后短时间轻度干旱对小麦成穗数和结实特性无显著影响,这与Ⅱp和Ⅰ1蘖成穗率升高在一定程度上弥补了Ⅲ、Ⅰp和Ⅳ蘖成穗率下降对单位面积穗数的影响有关;但干旱时间过长、程度过大会大幅度降低Ⅲ、Ⅰp和Ⅳ蘖成穗率、总小穗数、结实小穗数、穗粒数、单粒重和单穗产量,导致单位面积籽粒产量显著下降。山农29对拔节期阶段性干旱的抗性高于衡0628。

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Wheat yield depends on the number of grains per square metre, which in turn is related to the number of fertile florets at anthesis. The dynamics of floret generation/degeneration were studied in contrasting conditions of nitrogen (N) and water availability of modern, well-adapted, durum wheats in order to understand further the bases for grain number determination. Experiments were carried out during the 2008-2009 and 2009-2010 growing seasons at Lleida (NE Spain). The first experiment involved four cultivars (Claudio, Donduro, Simeto, and Vitron) and two contrasting N availabilities (50 kgN ha(-1) and 250 kgN ha(-1); N50 and N250) while experiment 2 included the two cultivars most contrasting in grain setting responsiveness to N in experiment 1, and two levels of N (N50 and N250), under irrigated (IR) and rainfed (RF) conditions. In addition, a detillering treatment was imposed on both cultivars under the IR+N250 condition. The number of fertile florets at anthesis was increased by ~30% in response to N fertilization (averaging across treatments and spikelet positions). The effect of N and water availability was evident on floret developmental rates from the third floret primordium onwards, as these florets in the central spikelets of all genotypes reached the stage of a fertile floret in N250 while in N50 they did not. In this study, clear differences were found between the cultivars in their responsiveness to N by producing more fertile florets at anthesis (through accelerating developmental rates of floret primordia), by increasing the likelihood of particular grains to be set, or by both traits.

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