干旱胁迫下外源褪黑素对冬小麦小花发育及碳营养代谢的调控

doi:10.3864/j.issn.0578-1752.2024.23.006

摘要/Abstract

摘要：

【目的】明确外源褪黑素对干旱胁迫下冬小麦小花发育及碳营养代谢的调控效应。【方法】于2021—2023年以多穗型品种豫麦49-198和大穗型品种周麦22为试验材料，设置干旱胁迫（D）和正常水分（W）2种水分条件，在小麦小花退化高峰前（拔节后20 d左右）进行叶面喷施100 μmol·L^-1褪黑素（MT）和清水对照（CK）处理，重点研究干旱胁迫下外源MT对冬小麦小花发育数量及形态特征、顶展叶片SPAD值、净光合速率、蔗糖含量及其代谢酶活性和产量构成因子的影响。【结果】干旱胁迫导致小麦小花退化和败育增加，而喷施外源MT可有效减少两品种小花退化和败育，增加可孕小花数，但不能完全抵消干旱胁迫带来的负效应；外源MT对两品种正常水分处理也表现出正向调控效应。喷施外源MT可有效提高两品种干旱胁迫和正常水分处理下顶展叶SPAD值、净光合速率、蔗糖磷酸合成酶（SPS）和蔗糖合成酶（SS）活性及穗部蔗糖含量，且干旱处理的增幅高于正常水分处理；喷施外源MT降低了干旱胁迫下两品种茎和叶器官的蔗糖含量，但正常水分处理下则相反。喷施外源MT显著提高了两品种在D和W两水分条件下的穗粒数，相比于不喷施MT处理，豫麦49-198喷施处理的穗粒数增幅分别为19.12%（D）和6.65%（W）；周麦22喷施处理的增幅分别为21.57%（D）和8.73%（W）；喷施MT对两品种同一水分处理下各自的穗数和千粒重表现出一定调控效应，但未达显著水平。两品种相比，外源MT对大穗型品种周麦22的调控效应整体上高于多穗型品种豫麦49-198。【结论】于小花退化高峰前喷施外源MT可有效提高小麦顶展叶SPAD值、净光合速率及碳代谢相关酶活性，促进光合产物合成、茎和叶营养器官蔗糖向穗器官分配转运，为小花发育提供了充足的营养保障以增加可孕小花数，进而提高穗粒数，且对大穗型品种周麦22的调控效应更明显。研究结果为通过施用外源褪黑素实现干旱胁迫下促粒稳产减灾提供了理论依据和技术支撑。

关键词: 冬小麦, 干旱胁迫, 褪黑素, 小花发育, 穗粒数, 碳代谢

Abstract:

【Objective】 This study aimed to clarify the regulatory effects of exogenous melatonin on floret development and carbon nutrient metabolism in winter wheat under drought stress. 【Method】 Two soil water conditions (drought stress treatment: D, and normal moisture treatment: W) were set up using multi-spike variety Yumai 49-198 and large-spike variety Zhoumai 22 as experimental wheat materials, with foliar spraying 100 μmol·L^-1 exogenous melatonin (MT) and clear water control (CK) before the peak of floret degradation (about 20 days after jointing) in 2021-2023, focusing on the effects of exogenous melatonin on the number and morphological characteristics of floret development, SPAD value, net photosynthetic rate, sucrose content and its metabolic enzyme activities of top spread leaves, and yield component factors of winter wheat under drought stress. 【Result】 The drought stress led to an increase in floret degradation and abortion in wheat, while spraying exogenous melatonin could effectively reduce floret degradation and abortion, and increase number of fertile florets of the two varieties, but could not completely counteract the negative effect of drought stress; exogenous melatonin also showed positive regulatory effect on normal water treatments of the two varieties. Spraying exogenous melatonin could effectively increase SPAD value, net photosynthetic rate, carbon metabolism- related enzyme activities of top spread leaves and spike sucrose content of the two varieties under drought stress and normal water treatment, and the increase range was higher in drought treatment than in normal water treatment; exogenous melatonin decreased sucrose content of stem and leaf organs of both varieties under drought stress, but the opposite was true under normal water treatment. Spraying exogenous melatonin significantly increased grain number per spike of two varieties under two moisture treatments, compared with no-spraying MT treatment, the grain number per spike of Yumai 49-198 with spraying MT treatment increased by 19.12% (D) and 6.65% (W), respectively; the grain number per spike of Zhoumai 22 with spraying MT treatment increased by 21.57% (D) and 8.73% (W), respectively; spraying MT showed some regulation effect on spike number and thousand grain weight of the two varieties under the same water treatment, but did not reach a significant level. Compared the differences between two varieties, the regulatory effect of spraying melatonin was overall higher in the large-spike variety Zhoumai 22 than in the multiple-spike variety Yumai 49-198. 【Conclusion】 Spraying exogenous melatonin before the peak of floret degradation could effectively increase the SPAD value, net photosynthetic rate, and carbon metabolism-related enzyme activities of top spread leaves in wheat, and promote synthesis of photosynthesis products and the distribution and transportation of sucrose from stem and leaf nutrient organs to spike organ, which could provide sufficient nutrient security for the development of florets to increase number of fertile florets, thereby increasing grain number per spike, and the regulating effect on the large-spike variety of Zhoumai 22 was more pronounced. The results of this study provided the theoretical basis and technical support for increasing grain number per spike, stabilizing yield and reducing disaster under drought stress through the application of exogenous melatonin.

Key words: winter wheat, drought stress, melatonin, floret development, grain number per spike, carbon metabolism

张荣, 刘淋茹, 付凯霞, 武紫君, 宋一凡, 王璐媛, 侯阁阁, 贺利, 冯伟, 段剑钊, 王永华, 郭天财. 干旱胁迫下外源褪黑素对冬小麦小花发育及碳营养代谢的调控[J]. 中国农业科学, 2024, 57(23): 4644-4657.

ZHANG Rong, LIU LinRu, FU KaiXia, WU ZiJun, SONG YiFan, WANG LuYuan, HOU GeGe, HE Li, FENG Wei, DUAN JianZhao, WANG YongHua, GUO TianCai. Regulatory of Exogenous Melatonin on Floret Development and Carbon Nutrient Metabolism in Winter Wheat Under Drought Stress[J]. Scientia Agricultura Sinica, 2024, 57(23): 4644-4657.

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参考文献 57

[1]	茹振钢, 冯素伟, 李淦. 黄淮麦区小麦品种的高产潜力与实现途径. 中国农业科学, 2015, 48(17): 3388-3393. doi: 10.3864/j.issn.0578-1752.2015.17.006.
	RU Z G, FENG S W, LI G. High-yield potential and effective ways of wheat in Yellow & Huai river valley facultative winter wheat region. Scientia Agricultura Sinica, 2015, 48(17): 3388-3393. doi: 10.3864/j.issn.0578-1752.2015.17.006. (in Chinese)
[2]	冯伟森, 张学品, 吴少辉, 高海涛, 张灿军. 不同播期对洛旱7号小麦幼穗分化及产量的影响. 河南农业科学, 2011, 40(10): 32-34.
	FENG W S, ZHANG X P, WU S H, GAO H T, ZHANG C J. Effects of different sowing dates on spike characteristics and yield of Luohan 7. Journal of Henan Agricultural Sciences, 2011, 40(10): 32-34. (in Chinese)
[3]	ZHENG C F, ZHU Y J, ZHU H J, KANG G Z, GUO T C, WANG C Y. Floret development and grain setting characteristics in winter wheat in response to pre-anthesis applications of 6-benzylaminopurine and boron. Field Crops Research, 2014, 169: 70-76.
[4]	FERRANTE A, SAVIN R, SLAFER G A. Floret development of durum wheat in response to nitrogen availability. Journal of Experimental Botany, 2010, 61(15): 4351-4359. doi: 10.1093/jxb/erq236 pmid: 20696655
[5]	孔祥, 张鑫, 黄翠, 骆永丽, 李勇, 金敏. 外源亚精胺对水分亏缺下冬小麦小花发育及结实特性的调控效应. 华北农学报, 2019, 34(S1): 49-58. doi: 10.7668/hbnxb.201751727
	KONG X, ZHANG X, HUANG C, LUO Y L, LI Y, JIN M. Effects of exogenous spermidine on floret development and grain setting characteristics of winter wheat under water deficit. Acta Agriculturae Boreali-Sinica, 2019, 34(S1): 49-58. (in Chinese) doi: 10.7668/hbnxb.201751727
[6]	朱元刚, 肖岩岩, 初金鹏, 张秀, 钤太峰, 马鑫, 代兴龙, 贺明荣. 不同播期冬小麦小花发育特性与同化物代谢的相关性. 植物营养与肥料学报, 2019, 25(3): 370-381.
	ZHU Y G, XIAO Y Y, CHU J P, ZHANG X, QIAN T F, MA X, DAI X L, HE M R. Correlation between floret development characteristics and photosynthate metabolism of winter wheat in different sowing dates. Journal of Plant Nutrition and Fertilizers, 2019, 25(3): 370-381. (in Chinese)
[7]	刘北城, 张艳艳, 戎亚思, 魏永康, 段剑钊, 贺利, 王永华, 郭天财, 刘万代, 冯伟. 干旱胁迫下喷施14-羟基芸苔素甾醇对冬小麦穗花发育及碳氮代谢的调控. 植物营养与肥料学报, 2021, 27(6): 1004-1015.
	LIU B C, ZHANG Y Y, RONG Y S, WEI Y K, DUAN J Z, HE L, WANG Y H, GUO T C, LIU W D, FENG W. Regulation of spraying 14-hydroxylated brassinosteroid on spike and fertile floret development and carbon and nitrogen metabolism of winter wheat under drought stress. Journal of Plant Nutrition and Fertilizers, 2021, 27(6): 1004-1015. (in Chinese)
[8]	GHIGLIONE H O, GONZALEZ F G, SERRAGO R, MALDONADO S B, CHILCOTT C, CURÁ J A, MIRALLES D J, ZHU T, CASAL J J. Autophagy regulated by day length determines the number of fertile florets in wheat. The Plant Journal, 2008, 55(6): 1010-1024. doi: 10.1111/j.1365-313X.2008.03570.x pmid: 18547393
[9]	GUO X L, ZHANG Z, LI J Y, ZHANG S Q, SUN W, XIAO X C, SUN Z C, XUE X Z, WANG Z M, ZHANG Y H. Phenotypic and transcriptome profiling of spikes reveals the regulation of light regimens on spike growth and fertile floret number in wheat. Plant, Cell & Environment, 2024, 47(5): 1575-1591.
[10]	LIN F F, LI C, XU B, CHEN J, CHEN A H, HASSAN M A, LIU B B, XU H, CHEN X, SUN J Q, LI J C. Late spring cold reduces grain number at various spike positions by regulating spike growth and assimilate distribution in winter wheat. The Crop Journal, 2023, 11(4): 1272-1278.
[11]	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. Effects of alternating drought and watering on growth, photosynthesis and yield of wither wheat. Journal of Irrigation and Drainage, 2018, 37(8): 76-82. (in Chinese)
[12]	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. 中国农业科学, 2020, 53(20): 4137-4151. doi: 10.3864/j.issn.0578-1752.2020.20.004.
	LI P, SHANG Y Q, LIN X, LIU S K, WANG S, HU X H, WANG D. Effects of drought stress during jointing stage on spike formation and seed setting of main stem and tillers of winter wheat. Scientia Agricultura Sinica, 2020, 53(20): 4137-4151. doi: 10.3864/j.issn.0578-1752.2020.20.004. (in Chinese)
[13]	ZHANG Z, HUANG J, GAO Y M, LIU Y, LI J P, ZHOU X N, YAO C S, WANG Z M, SUN Z C, ZHANG Y H. Suppressed ABA signal transduction in the spike promotes sucrose use in the stem and reduces grain number in wheat under water stress. Journal of Experimental Botany, 2020, 71(22): 7241-7256. doi: 10.1093/jxb/eraa380 pmid: 32822501
[14]	王沅, 田正国. 小麦小花发育不同时期土壤干旱对成花与成粒的影响. 作物学报, 1982, 8(4): 229-236.
	WANG Y, TIAN Z G. Influence of soil water stress on grain number in different development stages of wheat. Acta Agronomica Sinica, 1982, 8(4): 229-236. (in Chinese)
[15]	刘德帅, 姚磊, 徐伟荣, 冯美, 姚文孔. 褪黑素参与植物抗逆功能研究进展. 植物学报, 2022, 57(1): 111-126. doi: 10.11983/CBB21146
	LIU D S, YAO L, XU W R, FENG M, YAO W K. Research progress of melatonin in plant stress resistance. Chinese Bulletin of Botany, 2022, 57(1): 111-126. (in Chinese)
[16]	ERDAL S. Melatonin promotes plant growth by maintaining integration and coordination between carbon and nitrogen metabolisms. Plant Cell Reports, 2019, 38(8): 1001-1012. doi: 10.1007/s00299-019-02423-z pmid: 31069499
[17]	HUANG B, CHEN Y E, ZHAO Y Q, DING C B, LIAO J Q, HU C, ZHOU L J, ZHANG Z W, YUAN S, YUAN M. Exogenous melatonin alleviates oxidative damages and protects photosystem II in maize seedlings under drought stress. Frontiers in Plant Science, 2019, 10: 677. doi: 10.3389/fpls.2019.00677 pmid: 31178885
[18]	WANG K X, XING Q F, AHAMMED G J, ZHOU J. Functions and prospects of melatonin in plant growth, yield, and quality. Journal of Experimental Botany, 2022, 73(17): 5928-5946.
[19]	ZOU J N, JIN X J, WANG Y X Z C Y R M C Z M X. Effects of melatonin on photosynthesis and soybean seed growth during grain filling under drought stress. Photosynthetica, 2019, 57(2): 512-520.
[20]	WANG J J, YAN D, LU Z Z, LIU R, HONG Y, WANG Y, PENG Z, YU C X, GAO Y R, LIU Z Y, DUAN L S, LI R Z. Integration of the metabolome and transcriptome reveals diurnal variability in the effects of melatonin on salt tolerance in maize seedlings. Journal of Plant Growth Regulation, 2024, 43(5): 1672-1688.
[21]	WANG M L, ZHANG T, DING F. Exogenous melatonin delays methyl jasmonate-triggered senescence in tomato leaves. Agronomy, 2019, 9(12): 795.
[22]	KE Q B, YE J, WANG B M, REN J H, YIN L N, DENG X P, WANG S W. Melatonin mitigates salt stress in wheat seedlings by modulating polyamine metabolism. Frontiers in Plant Science, 2018, 9: 914. doi: 10.3389/fpls.2018.00914 pmid: 30018628
[23]	崔金梅, 郭天财, 朱云集, 王晨阳, 马新明. 小麦的穗. 北京: 中国农业出版社, 2008.
	CUI J M, GUO T C, ZHU Y J, WANG C Y, MA X M. Spike of Wheat. Beijing: China Agriculture Press, 2008. (in Chinese)
[24]	高俊凤. 植物生理学实验指导. 北京: 高等教育出版社, 2006.
	GAO J F. Experimental Guidance for Plant Physiology. Beijing: Higher Education Press, 2006. (in Chinese)
[25]	中国科学院上海植物生理研究所, 上海市植物生理学会. 现代植物生理学实验指南. 北京: 科学出版社, 1999.
	Shanghai Institute of Plant Physiology, Chinese Academy of Sciences, Shanghai Society for Plant Physiology. Guide to Modern Plant Physiology Experiments. Beijing: Science Press, 1999. (in Chinese)
[26]	王兆龙, 曹卫星, 戴廷波, 周琴. 不同穗型小麦品种小花发育与结实特性研究. 南京农业大学学报, 2000, 23(4): 9-12.
	WANG Z L, CAO W X, DAI T B, ZHOU Q. Characteristics of floret development and grain set in three wheat genotypes of different spike sizes. Journal of Nanjing Agricultural University, 2000, 23(4): 9-12. (in Chinese)
[27]	郑春风, 朱慧杰, 朱云集, 郭天财, 王晨阳. 冬小麦小花发育及结实特性对叶面喷硼的响应. 植物营养与肥料学报, 2016, 22(2): 550-556.
	ZHENG C F, ZHU H J, ZHU Y J, GUO T C, WANG C Y. Responses of floret development and grain setting characteristics of winter wheat to foliar spray boron. Journal of Plant Nutrition and Fertilizers, 2016, 22(2): 550-556. (in Chinese)
[28]	朱云集, 崔金梅, 郭天财, 王晨阳, 王永华. 河南省小麦生产发展中几个关键技术问题的商榷. 河南农业科学, 2011, 40(8): 54-57.
	ZHU Y J, CUI J M, GUO T C, WANG C Y, WANG Y H. Discussion of key technique problems in development of wheat production in Henan Province. Journal of Henan Agricultural Sciences, 2011, 40(8): 54-57. (in Chinese)
[29]	MIRALLES D J, RICHARDS R A, SLAFER G A. Duration of the stem elongation period influences the number of fertile florets in wheat and barley. Functional Plant Biology, 2000, 27(10): 931-940.
[30]	FLINTHAM J E, BÖRNER A, WORLAND A J, GALE M D. Optimizing wheat grain yield: Effects of Rht (gibberellin-insensitive) dwarfing genes. The Journal of Agricultural Science, 1997, 128(1): 11-25.
[31]	MIRALLES D J, KATZ S D, COLLOCA A, SLAFER G A. Floret development in near isogenic wheat lines differing in plant height. Field Crops Research, 1998, 59(1): 21-30.
[32]	SERRAGO R A, MIRALLES D J, SLAFER G A. Floret fertility in wheat as affected by photoperiod during stem elongation and removal of spikelets at booting. European Journal of Agronomy, 2008, 28(3): 301-308.
[33]	邱泽生, 刘捷平, 黄勤妮, 丁以珊, 张承谦, 王沅. 冬小麦的小花发育与碳氮代谢的关系. 作物学报, 1980, 6(3): 139-146.
	QIU Z S, LIU J P, HUANG Q N, DING Y S, ZHANG C Q, WANG Y. The relation between floret development and carbon nitrogen metabolism in winter wheat. Acta Agronomica Sinica, 1980, 6(3): 139-146. (in Chinese)
[34]	王志敏, 王树安, 苏宝林. 蔗糖供给对小麦离体培养穗小花结实的影响. 北京农业科学, 1996, 14(4): 17-20.
	WANG Z M, WANG S A, SU B L. Effect of sucrose supply on the fruiting of spike florets in wheat culture in vitro. Beijing Agricultural Sciences, 1996, 14(4): 17-20. (in Chinese)
[35]	王志敏, 王树安, 苏宝林. 乙烯对小麦小花发育和结实的影响. 种子, 1996, 15(2): 13-15.
	WANG Z M, WANG S A, SU B L. The effect of ethylene on floret development and grain set in wheat. Seed, 1996, 15(2): 13-15. (in Chinese)
[36]	黄明, 吴金芝, 李友军, 王贺正, 付国占, 陈明灿, 张振旺. 干旱对不同抗旱性小麦旗叶光响应特征和产量的影响. 干旱地区农业研究, 2020, 38(3): 64-73.
	HUANG M, WU J Z, LI Y J, WANG H Z, FU G Z, CHEN M C, ZHANG Z W. Effects of drought on flag-leaf photosynthetic characteristics to light and yield of wheat with different drought resistance. Agricultural Research in the Arid Areas, 2020, 38(3): 64-73. (in Chinese)
[37]	郭瑞盼, 辛泽毓, 王志强, 郭晓阳, 张丽婷, 王俊哲, 林同保. 干旱胁迫对小麦非结构性碳水化合物代谢的影响及其与抗旱性的关系. 华北农学报, 2015, 30(2): 202-211. doi: 10.7668/hbnxb.2015.02.035
	GUO R P, XIN Z Y, WANG Z Q, GUO X Y, ZHANG L T, WANG J Z, LIN T B. Effect of drought stress on non-structure carbohydrate metabolism of wheat and its relationship with drought resistance. Acta Agriculturae Boreali-Sinica, 2015, 30(2): 202-211. (in Chinese)
[38]	胡程达, 杨光仙, 成林. 干旱对冬小麦光合产物积累和分配的影响. 中国农业气象, 2014, 35(3): 243-249.
	HU C D, YANG G X, CHENG L. Effects of drought on distribution and accumulation of photosynthetic matter in winter wheat. Chinese Journal of Agrometeorology, 2014, 35(3): 243-249. (in Chinese)
[39]	ZHU Y G, ZHANG X, XIAO Y Y, CHU J P, DAI Z M. Variation of floret development and grain setting characteristics in winter wheat responses to delayed sowing. Journal of the Science of Food and Agriculture, 2022, 102(11): 4892-4908. doi: 10.1002/jsfa.11853 pmid: 35246843
[40]	张艳艳, 关涵文, 刘淋茹, 贺利, 段剑钊, 王晨阳, 郭天财, 冯伟. 不同水分条件下施磷对冬小麦穗花发育及产量的影响. 作物学报, 2023, 49(10): 2753-2765.
	ZHANG Y Y, GUAN H W, LIU L R, HE L, DUAN J Z, WANG C Y, GUO T C, FENG W. Effects of phosphorus application on spike and fertile floret development and yield of winter wheat under different water treatments. Acta Agronomica Sinica, 2023, 49(10): 2753-2765. (in Chinese) doi: 10.3724/SP.J.1006.2023.21097
[41]	ZHANG N, SUN Q Q, ZHANG H J, CAO Y Y, WEEDA S, REN S X, GUO Y D. Roles of melatonin in abiotic stress resistance in plants. Journal of Experimental Botany, 2015, 66(3): 647-656. doi: 10.1093/jxb/eru336 pmid: 25124318
[42]	苗含笑, 李东晓, 王久红, 孙亚倩, 刘立, 刘睿, 李红叶. 褪黑素对干旱胁迫下小麦生长发育和产量的影响. 干旱地区农业研究, 2020, 38(5): 161-167, 191.
	MIAO H X, LI D X, WANG J H, SUN Y Q, LIU L, LIU R, LI H Y. Effects of melatonin on the growth and yield of wheat under drought condition. Agricultural Research in the Arid Areas, 2020, 38(5): 161-167, 191. (in Chinese)
[43]	陈丽珊, 周红艳, 林伟伟. 外源褪黑素对盐胁迫下水稻苗期碳氮代谢的影响. 生态学杂志, 2023, 42(7): 1635-1643.
	CHEN L S, ZHOU H Y, LIN W W. Effects of exogenous melatonin on carbon and nitrogen metabolism of rice seedlings under salt stress. Chinese Journal of Ecology, 2023, 42(7): 1635-1643. (in Chinese)
[44]	刘佳奇, 李丽, 杨红红, 孙敏, 冯汉青. 盐胁迫下褪黑素对小麦种子萌发和幼苗生理特性的影响. 麦类作物学报, 2022, 42(7): 857-863.
	LIU J Q, LI L, YANG H H, SUN M, FENG H Q. Effect of melatonin on seed germination and seedling physiological characteristics of wheat under salt stress. Journal of Triticeae Crops, 2022, 42(7): 857-863. (in Chinese)
[45]	SUN L Y, LI X N, WANG Z S, SUN Z W, ZHU X C, LIU S Q, SONG F B, LIU F L, WANG Y J. Cold priming induced tolerance to subsequent low temperature stress is enhanced by melatonin application during recovery in wheat. Molecules, 2018, 23(5): 1091.
[46]	曹亮, 杜昕, 于高波, 金喜军, 张明聪, 任春元, 王孟雪, 张玉先. 外源褪黑素对干旱胁迫下绥农26大豆鼓粒期叶片碳氮代谢调控的途径分析. 作物学报, 2021, 47(9): 1779-1790. doi: 10.3724/SP.J.1006.2021.04151
	CAO L, DU X, YU G B, JIN X J, ZHANG M C, REN C Y, WANG M X, ZHANG Y X. Regulation of carbon and nitrogen metabolism in leaf of soybean cultivar Suinong 26 at seed-filling stage under drought stress by exogenous melatonin. Acta Agronomica Sinica, 2021, 47(9): 1779-1790. (in Chinese) doi: 10.3724/SP.J.1006.2021.04151
[47]	FU Y Y, LI P H, LIANG Y P, SI Z Y, MA S T, GAO Y. Effects of exogenous melatonin on wheat quality under drought stress and rehydration. Plant Growth Regulation, 2024, 103(3): 471-490.
[48]	李春雨, 陈春宇, 毛浩田, 张怀渝, 陈洋尔. 干旱胁迫下外源褪黑素对小麦生长和光系统活性的影响. 麦类作物学报, 2022, 42(7): 846-856.
	LI C Y, CHEN C Y, MAO H T, ZHANG H Y, CHEN Y E. Effect of exogenous melatonin on the growth and photosystem activity of wheat under drought stress. Journal of Triticeae Crops, 2022, 42(7): 846-856. (in Chinese)
[49]	SONG L L, LIU S W, YU H T, YU Z F. Exogenous melatonin ameliorates yellowing of postharvest pak choi (Brassica rapa subsp. chinensis) by modulating chlorophyll catabolism and antioxidant system during storage at 20 ℃. Scientia Horticulturae, 2023, 311: 111808.
[50]	ZAFAR S, AKHTAR M, PERVEEN S, HASNAIN Z, KHALIL A. Attenuating the adverse aspects of water stress on wheat genotypes by foliar spray of melatonin and indole-3-acetic acid. Physiology and Molecular Biology of Plants, 2020, 26(9): 1751-1762. doi: 10.1007/s12298-020-00855-6 pmid: 32943813
[51]	王志敏, 王树安. 小麦穗茎生长与穗粒数的关系. 北京农业科学, 1996, 14(3): 4-7.
	WANG Z M, WANG S A. Relationship between stem growth and number of grains per spike of wheat. Beijing Agricultural Sciences, 1996, 14(3): 4-7. (in Chinese)
[52]	FISCHER R A. Number of kernels in wheat crops and the influence of solar radiation and temperature. The Journal of Agricultural Science, 1985, 105(2): 447-461.
[53]	田中伟, 王妮妮, 李怡香, 崔亚坤, 蔡剑, 姜东, 戴廷波. 分蘖期和拔节期干旱对小麦主茎和分蘖穗粒形成的影响. 麦类作物学报, 2018, 38(6): 734-741.
	TIAN Z W, WANG N N, LI Y X, CUI Y K, CAI J, JIANG D, DAI T B. Effect of water deficit on main stem and tiller grain number formation of wheat during tillering stage and jointing stage. Journal of Triticeae Crops, 2018, 38(6): 734-741. (in Chinese)
[54]	YE J, YANG W J, LI Y L, WANG S W, YIN L N, DENG X P. Seed pre-soaking with melatonin improves wheat yield by delaying leaf senescence and promoting root development. Agronomy-Basel, 2020, 10(1): 84.
[55]	叶君, 于美玲, 曹梦龙, 崔国惠, 王小兵, 吴晓华, 张海斌, 赵轩微, 李元清, 李娟, 路战远, 耿志勇. 水分亏缺条件下褪黑素对春小麦光合特性及产量性状的影响. 北方农业学报, 2021, 49(2): 24-30. doi: 10.12190/j.issn.2096-1197.2021.02.04
	YE J, YU M L, CAO M L, CUI G H, WANG X B, WU X H, ZHANG H B, ZHAO X W, LI Y Q, LI J, LU Z Y, GENG Z Y. Effects of melatonin on photosynthetic characteristics and yield traits of spring wheat under water deficit. Journal of Northern Agriculture, 2021, 49(2): 24-30. (in Chinese) doi: 10.12190/j.issn.2096-1197.2021.02.04
[56]	朱云集, 崔金梅, 王晨阳, 郭天财, 夏国军, 刘万代, 王永华. 小麦不同生育时期施氮对穗花发育和产量的影响. 中国农业科学, 2002, 35(11): 1325-1329.
	ZHU Y J, CUI J M, WANG C Y, GUO T C, XIA G J, LIU W D, WANG Y H. Effects of nitrogen application at different wheat growth stages on floret development and grain yield of winter wheat. Scientia Agricultura Sinica, 2002, 35(11): 1325-1329. (in Chinese)
[57]	郑春风, 任伟, 朱云集, 张玉亭, 孙克刚, 杜君, 张运红, 和爱玲. 冬小麦小花发育及结实特性对叶面喷6-BA的响应. 植物营养与肥料学报, 2017, 23(3): 774-780.
	ZHENG C F, REN W, ZHU Y J, ZHANG Y T, SUN K G, DU J, ZHANG Y H, HE A L. Responses of floret development and grain setting characteristics of winter wheat to foliar spray of 6-benzylaminopurine. Journal of Plant Nutrition and Fertilizer, 2017, 23(3): 774-780. (in Chinese)

年份 Year	品种 Cultivar	处理 Treatment	每盆穗数 Spike number per pot	穗粒数 Grain number per spike	千粒重 1000-grain weight (g)
2021—2022	豫麦49-198 Yumai 49-198	WCK	63.00a	35.50b	46.55c
		WMT	62.75a	38.24a	47.27bc
		DCK	40.50b	23.38d	48.13ab
		DMT	41.00b	28.04c	49.01a
	周麦22 Zhoumai 22	WCK	47.50a	38.61b	48.61b
		WMT	47.00a	42.33a	49.24b
		DCK	34.75b	24.13d	50.93a
		DMT	35.75b	29.61c	51.70a
2022—2023	豫麦49-198 Yumai 49-198	WCK	47.02a	36.00b	45.12b
		WMT	48.35a	38.01a	45.94b
		DCK	35.05b	24.00d	48.65a
		DMT	36.25b	28.40c	49.38a
	周麦22 Zhoumai 22	WCK	39.80a	39.46b	47.00b
		WMT	40.85a	42.55a	47.84b
		DCK	27.22b	25.90d	50.12a
		DMT	28.28b	31.10c	50.80a