拔节期阶段性干旱对小麦茎蘖成穗与结实的影响

doi:10.3864/j.issn.0578-1752.2020.20.004

摘要/Abstract

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.

Key words: winter wheat, phased drought, main stem and tillers, spike formation and seed setting, grain yield

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.

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 Agricultura Sinica, 2020, 53(20): 4137-4151.

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图/表 9

表1

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"

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
年份 Year	品种 Cultivar	处理 Treatment	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

表2

表3

表4

表5

表6

表7

表8

表9

参考文献 33

[1]	LI T Y, ZHANG X, GAO H X, LI B, WANG H, YAN Q Y, OLLENBURGER M, ZHANG W F. Exploring optimal nitrogen management practices within site-specific ecological and socioeconomic conditions. Journal of Cleaner Production, 2019, doi: 10.1016/j.jclepro.2019.118295. doi: 10.1016/j.jclepro.2020.124063 pmid: 32921931
[2]	YUAN Z, YAN D H, YANG Z Y, YIN J, YUAN Y. Temporal and spatial variability of drought in Huang-Huai-Hai River Basin, China. Theoretical and Applied Climatology, 2015,122:755-769.
[3]	银敏华, 李援农, 周昌明, 谷晓博, 张天乐, 杨丹, 吴国军. 调亏灌水和分蘖干扰对冬小麦生长的补偿效应. 应用生态学报, 2015,26(10):3011-3019.
	YIN M H, LI Y N, ZHOU C M, GU X B, ZHANG T L, YANG D, WU G J. Compensation effects of regulated deficit irrigation and tiller disturbance on winter wheat growth. Journal of Applied Ecology, 2015,26(10):3011-3019. (in Chinese)
[4]	汪顺生, 孟鹏涛, 高传昌, 刘慧, 易嘉成. 不同灌溉方式对冬小麦/夏玉米生长发育及产量的影响. 中国农村水利水电, 2015,57(6):86-90.
	WANG S S, MENG P T, GAO C C, LIU H, YI J C. Effects of different irrigation methods on the growth and yield of winter wheat / summer corn. China Rural Water and Hydropower, 2015,57(6):86-90. (in Chinese)
[5]	李娜娜, 宫永超, 蒲艳艳, 张晓东, 贾文斌, 辛富刚, 裴艳婷, 丁汉凤. 不同穗型冬小麦品种分蘖成穗特性的研究进展. 中国农学通报, 2014,30(2):14-18.
	LI N N, GONG Y C, PU Y Y, ZHANG X D, JIA W B, XIN F G, PEI Y T, DING H F. Recent progress of formation of spike characteristics in different spike cultivars of wheat. Chinese Agricultural Science Bulletin, 2014,30(2):14-18. (in Chinese)
[6]	倪雪峰, 朱倩, 刘涛, 闫向泉, 孟自力, 朱伟. 不同农艺措施对‘商麦156’分蘖成穗及产量的影响. 中国农学通报, 2019,35(16):1-5.
	NI X F, ZHU Q, LIU T, YAN X Q, MENG Z L, ZHU W. Agronomic measures affect tillering and spike formation and yield of ‘shangmai 156’. Chinese Agricultural Science Bulletin, 2019,35(16):1-5.(in Chinese)
[7]	YAN J, YU J, TAO G C, VOS J, BOUMAN B A M, XIE G H, MEINKE H. Yield formation and tillering dynamics of direct-seeded rice in flooded and nonflooded soils in the Huai River Basin of China. Field Crops Research, 2010,116(3):252-259.
[8]	佟汉文, 彭敏, 刘易科, 黄玫挺, 邹娟, 朱展望, 陈冷, 陈宇庆, 高春保. 小麦分蘖成穗规律研究进展. 湖北农业科学, 2017,56(24):4700-4702.
	TONG H W, PENG M, LIU Y K, HUANG M T, ZOU J, ZHU Z W, CHEN L, CHEN Y Q, GAO C B. Research progress in law of spike formation from wheat tillers. Hubei Agricultural Sciences, 2017,56(24):4700-4702. (in Chinese)
[9]	高尔明, 赵全志, 刘华山, 杨青华, 刘万代, 梁静静, 王春丽. 砂姜黑土小麦分蘖成穗及其调控研究. 土壤通报, 2001,32(3):140-142.
	GAO E M, ZHAO Q Z, LIU H S, YANG Q H, LIU W D, LIANG J J, WANG C L. Study on tillering and earing of wheat in Shajiang black soil and its regulation. Chinese Journal of Soil Science, 2001,32(3):140-142. (in Chinese)
[10]	王晓宇, 冯伟, 郭天财, 康国章, 王晨阳. 两种穗型小麦品种分蘖衰亡进程中茎蘖碳氮代谢的差异. 西北农业学报, 2010,19(11):38-42,57.
	WANG X Y, FENG W, GUO T C, KANG G Z, WANG C Y. Difference of carbon and nitrogen metabolism in leaves between main cand tillers during tiller senescence of two spike-type winter wheat. Acta Agricuiturae Boreali-Occidentalis Sinica, 2010,19(11):38-42, 57. (in Chinese)
[11]	赵黎明, 李明, 郑殿峰, 顾春梅, 那永光, 解保胜. 灌溉方式对寒地水稻产量及籽粒灌浆的影响. 中国农业科学, 2015,48(22):4493-4506.
	ZHAO L M, LI M, ZHENG D F, GU C M, NA Y G, XIE B S. Effects of irrigation methods on rice yield and grain filling in cold regions. Scientia Agricultura Sinica, 2015,48(22):4493-4506. (in Chinese)
[12]	PENG C R, XIE J S, QIU C F, QIAN Y F, GUAN X J, PAN X H. Study and application of three high and one ensuring cultivation mode of double cropping rice. Agricultural Science & Technology, 2012,13(7):1425-1430.
[13]	崔亚坤, 王妮妮, 田中伟, 戴廷波, 陈艳萍, 袁建华, 分蘖和拔节期干旱对小麦植株氮素积累转运的影响. 麦类作物学报, 2019,39(3):322-328.
	CUI Y K, WANG N N, TIAN Z W, DAI T B, CHEN Y P, YUAN J H. Effect of water deficit during tillering and jointing stages on nitrogen accumulation and translocaton in water wheat. Journal of Triticeae Crops, 2019,39(3):323-328. (in Chinese)
[14]	张继波, 薛晓萍, 李楠, 李鸿怡, 张磊, 宋计平. 干旱胁迫对冬小麦水分关键时期的生理特性和物质生产的影响. 沙漠与绿洲气象, 2019,13(3):124-130.
	ZHANG J B, XUE X P, LI N, LI H Y, ZHANG L, SONG J P. Effect of drought stress on physiological characteristics and dry matter production of winter wheat during water critical period. Desert and Oasis Meteorogy, 2019,13(3):124-130. (in Chinese)
[15]	李彦彬, 朱亚南, 李道西, 高阳. 阶段干旱及复水对小麦生长发育、光合和产量的影响. 灌溉排水学报, 2018,37(8):76-82.
	LI Y B, ZHU Y N, LI D X, GAO Y. Different stage drought and rehydration on wheat growth and development, photosynthesis and yield. Journal of Irrigation and Drainage, 2018,38(8):76-82. (in Chinese)
[16]	胡洋山, 汤颖子, 李治, 晏本菊, 任正隆, 任天恒. 小麦分蘖成穗数相关分子标记在重组自交系(RIL)群体中的有效性验证及实用性评价. 麦类作物学报, 2018,38(1):8-15.
	HU Y S, TANG Y Z, LI Z, YAN B J, REN Z L, REN T H. Verification and practicability evaluation of molecular markers related to tillering number of wheat in recombinant inbred line (RIL) population. Journal of Triticeae Crops, 2018,38(1):8-15. (in Chinese)
[17]	金欣欣, 姚艳荣, 贾秀领, 姚海坡, 申海平, 崔永增, 李谦. 基因型和环境对小麦产量、品质和氮素效率的影响. 作物学报, 2019,45(4):155-164.
	JIN X X, YAO Y R, JIA X L, YAO H P, SHEN H P, CUI Y Z, LI Q. Effects of genotypes and environment on wheat yield, quality and nitrogen efficiency. Acta Agronomica Sinica, 2019,45(4):155-164. (in Chinese)
[18]	杨文平, 单长卷, 胡喜巧, 李杰. 土壤干旱对冬小麦拔节期叶片碳代谢的影响. 河南农业科学, 2008(9):22-24, 28.
	YANG W P, SHAN C J, HU X Q, LI J. Effect of soil drought on carbon metabolism of winter wheat during jointing stage.Henan Agricultural Science, 2008(9):22-24, 28. (in Chinese)
[19]	XU H C, CAI T, WANG Z L, HE M R. Physiological basis for the differences of productive capacity among tillers in winter wheat. Journal of Intergrative Agriculture, 2015,14(10):1958-1970.
[20]	余松烈主编. 山东小麦. 北京: 农业出版社, 1990,80.
	YU S L. Wheat in Shandong. Beijing: Agricultural Publishing House, 1990,80. (in Chinese)
[21]	ARIEL F, ROXANA S, GUSTAVO A S. Folet development and grain setting differences between modern durum wheats under contrasting nitrogen availability. Journal of Experimental Botany, 2013,64(1):169-184. doi: 10.1093/jxb/ers320 pmid: 23162124
[22]	张伟杨, 钱希旸, 李银银, 徐云姬, 王志琴, 杨建昌. 土壤干旱对小麦生理性状和产量的影响. 麦类作物学报, 2016,36(4):491-500.
	ZHANG W Y, QIAN X Y, LI Y Y, XU Y J, WANG Z Q, YANG J C. Effects of soil drought on physiological characteristics and yield of wheat. Journal of Triticeae Crops, 2016,36(4):491-500. (in Chinese)
[23]	DAVIDSON D J, CHEVALIER P M. Preanthesis tiller mortality in spring wheat. Crop Science, 1990,30(4):832-836.
[24]	VAHAMIDIS P, KARAMANOS A J, ECONOMOU G. Grain number determination in durum wheat as affected by drought stress: An analysis at spike and spikelet level. Annals of Applied Biology, 2019,174(2), 190-208.
[25]	黄德明, 俞仲林, 路季梅, 江华山. 宁麦3号在高产栽培条件下分蘖幼穗的发育特征. 南京农学院学报, 1983,28(2):9-21.
	HUANG D M, YU Z L, LU J M, JIANG H S. Developmental characteristics of tillering young ears of Ningmai 3 under high-yield cultivation conditions. Journal of Nanjing Agriculture University, 1983,28(2):8-21. (in Chinese)
[26]	惠建, 袁汉民. 宁冬11号小麦茎蘖成穗规律研究. 农业科学研究, 2012,33(1):31-35.
	HUI J, YUAN H M. Study on tillers growing into spikes of Ningdong 11 in Ningxia. Journal of Agricultural Sciences, 2012,33(1):31-35. (in Chinese)
[27]	高翔, 宁锟, 宋哲民. 小麦高产品种幼穗分化发育特征的研究. 西北农业学报, 1995,4(3):1-7.
	GAO X, NING K, SONG Z M. Study on characteristics of yong spike differentiation of high-yield wheat. Acta Agriculturae Boreali- Occidentalis Sinica, 1995,4(3):1-7. (in Chinese)
[28]	唐永金. 绵阳11号小麦品种分蘖成穗规律的初步研究. 耕作与栽培, 1987,7(5):43-45.
	TANG Y J. Preliminary study on tillering and spike formation of wheat variety mianyang. Tillage and Cultivation, 1987,7(5):43-45. (in Chinese)
[29]	GONZÁLEN F G, TERRIL I I, FALCÓN M O. Spike fertility and duration of stem elongation as promising traits to improve potential grain number (and yield): Variation in modern Argentinean wheats. Crop Science, 2011,51(4):1693-1702.
[30]	PRIETO P, OCHAGAVÍA H, SAVIN R, GRIFFITHS S, SLAFER G A. Physiological determinants of fertile floret survival in wheat as affected by earliness per se genes under field conditions. European Journal of Agronomy, 2018,99:206-213.
[31]	ZHANG W Y, CHEN Y J, WANG Z Q, YANG J C. Polyamines and ethylene in rice young panicles in response to soil-drying during panicle differentiation. Plant Growth Regulation, 2017,82:491-503.
[32]	SAINIH S, ASPINALLl D. Effect of water deficit on sporogenesis in wheat (Triticum aestivum L.). Annals of Botany, 1981,48(5):623-633.
[33]	VAHAMIDIS P, KARAMANOS A J, ECONOMOU G. Grain number determination in durum wheat as affected by drought stress: An analysis at spike and spikelet level. Annals of Applied Biology, 2019,174(2):190-208.

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

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

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

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

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