不同时期高温胁迫对夏玉米物质生产性能及籽粒产量的影响

doi:10.3864/j.issn.0578-1752.2022.19.003

摘要/Abstract

摘要：

【目的】黄淮海夏玉米区高温胁迫频发、重发、持续期延长显著影响籽粒产量。本文以不同耐热型玉米品种为材料,探明大喇叭口期（V12）和开花期（VT）高温胁迫对两类品种叶片光合特性、碳同化物积累、分配和籽粒产量的影响。【方法】本研究以耐热型品种郑单958（ZD958）和热敏感型品种先玉335（XY335）为材料,以同时期适宜温度处理（昼32℃12 h/夜22℃12 h）为对照,使用自动控温控湿的高温棚模拟田间自然增温效果,设置V12期、VT期高温胁迫处理（昼38℃12 h/夜28℃12 h）,比较高温胁迫后夏玉米叶面积指数（LAI）、碳代谢酶活性、光合速率、碳同化物积累和分配的动态变化特征,明确夏玉米物质生产性能及籽粒产量对高温胁迫的响应机制。【结果】高温胁迫后,两品种的LAI、碳代谢酶活性、净光合速率和干物质积累量均显著降低,ZD958和XY335的LAI、RuBP羧化酶活性、PEP羧化酶活性、净光合速率和干物质积累量比其对照分别降低了2.98%—4.21%、40.38%—54.46%、16.88%—30.60%、18.14%—25.49%、12.83%—19.38%和3.80%—5.07%、56.56%—76.16%、26.33%—33.66%、22.37%—34.62%、22.07%—26.72%,VT期高温胁迫的降幅大于V12期。高温胁迫后,夏玉米叶片蒸腾速率显著升高,但叶片水分利用效率显著下降。高温下两品种的¹³C同化量均显著降低,V12期高温胁迫后,ZD958和XY335的¹³C同化量分别降低了18.48%和22.82%,籽粒中¹³C同化量占比降低。高温胁迫显著降低穗粒数,千粒重虽有小幅提高,但籽粒产量显著降低。与适宜温度相比,V12期高温胁迫后ZD958穗粒数和产量分别降低了62.53%和45.87%;VT期高温胁迫后穗粒数和产量分别降低了70.53%和66.89%;V12期高温胁迫后XY335穗粒数和产量分别降低了70.50%和62.87%;VT期高温胁迫后分别降低了85.41%和80.61%;VT期高温胁迫降幅大于V12期高温胁迫,XY335的降幅大于ZD958。【结论】高温胁迫降低了夏玉米叶面积指数、叶片RuBP和PEP羧化酶活性,显著降低叶片光合速率和干物质生产性能。高温下穗粒数显著减少,抑制了碳同化物从叶片和茎秆向籽粒的转运,最终导致籽粒产量降低。VT期高温胁迫效应大于V12期,热敏感型品种XY335的降幅显著大于耐热型品种ZD958。

关键词: 夏玉米, 高温胁迫, 碳代谢酶活性, 碳同化物积累与分配, 产量

Abstract:

【Objective】 Frequent, recurrent and prolonged high temperature stress had significant effects on grain yield of summer maize in Huang-Huai-Hai region. In this study, we investigated the effects of high temperature stress at the V12 stage and VT stage on leaf photosynthetic characteristics, carbon assimilate accumulation, distribution and grain yield of maize varieties with different heat tolerance. 【Method】 In this study, heat resistant maize variety Zhengdan 958 (ZD958) and heat sensitive maize variety Xianyu 335 (XY335) were used as materials. The normal temperature treatments (day 32℃12 h /night 22℃12 h) were set as the control at the same time. High temperature greenhouse equipped with automatic temperature and humidity control facilities was used to simulate the effect of natural field high temperature, the high temperature stress treatments (day 38℃12 h/night 28℃12 h) were set at V12 and VT stage, respectively. The dynamic characteristics of leaf area index (LAI), carbon metabolism enzyme activities, photosynthetic rate and carbon assimilate accumulation and allocation were compared after high temperature stress, aimed to determine the response mechanism of dry matter production performance and grain yield to high temperature stress. 【Result】 After high temperature stress, LAI, carbon metabolism enzyme activities, net photosynthetic rate and dry matter accumulation of two cultivars were significantly decreased. LAI, RuBP carboxylase activity, PEP carboxylase activity, net photosynthetic rate and dry matter accumulation of ZD958 and XY335 decreased by 2.98%-4.21%, 40.38%-54.46%, 16.88%-30.60%, 18.14%-25.49%, 12.83%-19.38% and 3.80%-5.07%, 56.56%-76.16%, 26.33%-33.66%, 22.37%-34.62%, 22.07%-26.72%, respectively. The decrease range of high temperature stress in VT stage was larger than that in V12 stage. After high temperature stress, transpiration rate of summer maize leaves increased, while leaf water use efficiency decreased significantly. Under high temperature stress, ¹³C assimilation of ZD958 and XY335 decreased by 18.48% and 22.82%, respectively, and the proportion of ¹³C assimilation in grains decreased. The high temperature stress significantly decreased grain number per spike and grain yield, although 1000 grain weight increased slightly. Compared to the optimum temperature, after V12 high temperature stress, the grain number per spike and yield of ZD958 decreased by 62.53% and 45.87%. After VT high temperature stress, grain number per spike and yield decreased by 70.53% and 66.89%. After V12 high temperature stress, the grain number per spike and yield of XY335 decreased by 70.50% and 62.87%. After VT high temperature stress, grain number per spike and yield decreased by 85.41% and 80.61%. The decrease range of high temperature stress in VT stage was larger than that in V12 stage, and XY335 decreased more than ZD958. 【Conclusion】 The high temperature stress reduced LAI, RuBP carboxylase and PEP carboxylase activities, and significantly reduced photosynthetic rate and dry matter production performance of summer maize. Under high temperature stress, the grain number per spike decreased significantly, which inhibited the transportation of carbohydrate from leaf and stem to grain, resulting in lower grain yield. The effects of high temperature stress on dry matter performance and grain yield of summer maize in VT stage was significantly greater than that in V12 stage. The decrease of heat sensitive variety XY335 was significantly greater than that of heat resistant variety ZD958 in two periods.

Key words: summer maize, high temperature stress, carbon metabolism enzyme activity, carbon assimilate accumulation and distribution, yield

张川,刘栋,王洪章,任昊,赵斌,张吉旺,任佰朝,刘存辉,刘鹏. 不同时期高温胁迫对夏玉米物质生产性能及籽粒产量的影响[J]. 中国农业科学, 2022, 55(19): 3710-3722.

ZHANG Chuan,LIU Dong,WANG HongZhang,REN Hao,ZHAO Bin,ZHANG JiWang,REN BaiZhao,LIU CunHui,LIU Peng. Effects of High Temperature Stress in Different Periods on Dry Matter Production and Grain Yield of Summer Maize[J]. Scientia Agricultura Sinica, 2022, 55(19): 3710-3722.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2022.19.003

https://www.chinaagrisci.com/CN/Y2022/V55/I19/3710

图/表 10

表1

图1

图2

图3

图4

图5

图6

图7

表2

表3

参考文献 35

[1]	李少昆, 赵久然, 董树亭, 赵明, 李潮海, 崔彦宏, 刘永红, 高聚林, 薛吉全, 王立春, 王璞, 陆卫平, 王俊河, 杨祁峰, 王子明. 中国玉米栽培研究进展与展望. 中国农业科学, 2017, 50(11): 1941-1959.
	LI S K, ZHAO J R, DONG S T, ZHAO M, LI C H, CUI Y H, LIU Y H, GAO J L, XUE J Q, WANG L C, WANG P, LU W P, WANG J H, YANG Q F, WANG Z M. Advances and prospects of maize cultivation in China. Scientia Agricultura Sinica, 2017, 50(11): 1941-1959. (in Chinese)
[2]	任寒, 刘鹏, 董树亭, 张吉旺, 赵斌. 高温胁迫影响玉米生长发育的生理机制研究进展. 玉米科学, 2019, 27(5): 109-115.
	REN H, LIU P, DONG S T, ZHANG J W, ZHAO B. Research advancements of effect of high temperature stress on growth and development of maize. Journal of Maize Sciences, 2019, 27(5): 109-115. (in Chinese)
[3]	王群, 赵向阳, 刘东尧, 闫振华, 李鸿萍, 董朋飞, 李潮海. 淹水弱光复合胁迫对夏玉米根形态结构、生理特性和产量的影响. 中国农业科学, 2020, 53(17): 3479-3495.
	WANG Q, ZHAO X Y, LIU D Y, YAN Z H, LI H P, DONG P F, LI C H. Root morphological, physiological traits and yield of maize under waterlogging and low light stress. Scientia Agricultura Sinica, 2020, 53(17): 3479-3495. (in Chinese)
[4]	ZHANG Q, YANG Z Q. Impact of extreme heat on corn yield in main summer corn cultivating area of China at present and under future climate change. International Journal of Plant Production, 2019, 13(4): 267-274. doi: 10.1007/s42106-019-00052-w
[5]	JHA U C, BOHRA A, SINGH N P. Heat stress in crop plants: Its nature, impacts and integrated breeding strategies to improve heat tolerance. Plant Breeding, 2014, 133(6): 679-701. doi: 10.1111/pbr.12217
[6]	苌建峰. 不同基因型玉米碳氮代谢差异研究[D]. 郑州: 河南农业大学, 2007.
	CHANG J F. Study on differences of carbon and nitrogen metabolism in different genotype maize[D]. Zhengzhou: Henan Agricultural University, 2007. (in Chinese)
[7]	LEMMA A T. Action and reaction of plants to high temperature: Improving response of wheat to heat stress. Current Journal of Applied Science and Technology, 2021, 40(10): 62-70.
[8]	ZHEN F X, ZHOU J J, MAHMOOD A, WANG W, CHANG X N, LIU B, LIU L L, CAO W X, ZHU Y, TANG L. Quantifying the effects of short term heat stress at booting stage on nonstructural carbohydrates remobilization in rice. The Crop Journal, 2020, 8(2): 194-212. doi: 10.1016/j.cj.2019.07.002
[9]	张保仁, 董树亭, 胡昌浩, 王空军. 高温对玉米籽粒淀粉合成及产量的影响. 作物学报, 2007, 33(1): 38-42.
	ZHANG B R, DONG S T, HU C H, WANG K J. Effect of high air temperature during different growth stage on starch synthesis in grain and yield in maize (Zea mays L.). Acta Agronomica Sinica, 2007, 33(1): 38-42. (in Chinese)
[10]	王海梅. 高温胁迫对河套灌区玉米生理指标及产量构成要素的影响. 干旱气象, 2015, 33(1): 59-62.
	WANG H M. Temperature stress on physiological indexes and yield components of maize in Hetao irrigation district. Journal of Arid Meteorology, 2015, 33(1): 59-62. (in Chinese)
[11]	XU Y F, CHU C C, YAO S G. The impact of high temperature stress on rice: Challenges and solutions. The Crop Journal, 2021, 9(5): 963-976. doi: 10.1016/j.cj.2021.02.011
[12]	赵龙飞, 李潮海, 刘天学, 王秀萍, 僧珊珊. 花期前后高温对不同基因型玉米光合特性及产量和品质的影响. 中国农业科学, 2012, 45(23): 4947-4958.
	ZHAO L F, LI C H, LIU T X, WANG X P, SENG S S. Effect of high temperature during flowering on photosynthetic characteristics and grain yield and quality of different genotypes of maize (Zea mays L.). Scientia Agricultura Sinica, 2012, 45(23): 4947-4958. (in Chinese)
[13]	孙宁宁. 玉米叶、粒对高温胁迫的响应[D]. 郑州: 河南农业大学, 2017.
	SUN N N. Responses of maize (Zea mays L.) leaf and kernel to heat stress[D]. Zhengzhou: Henan Agricultural University, 2017. (in Chinese)
[14]	陈传晓. 不同积温带春玉米碳代谢机理及化学调控效应的研究[D]. 保定: 河北农业大学, 2013.
	CHEN C X. Studies on the mechanism of carbon metabolism of spring maize and chemical regulation effects under different accumulated temperature zones[D]. Baoding: Hebei Agricultural University, 2013. (in Chinese)
[15]	孙胜楠, 王强, 孙晨晨, 刘丰娇, 毕焕改, 艾希珍. 黄瓜幼苗光合作用对高温胁迫的响应与适应. 应用生态学报, 2017, 28(5): 1603-1610. doi: 10.13287/j.1001-9332.201705.009
	SUN S N, WANG Q, SUN C C, LIU F J, BI H G, AI X Z. Response and adaptation of photosynthesis of cucumber seedlings to high temperature stress. Chinese Journal of Applied Ecology, 2017, 28(5): 1603-1610. (in Chinese) doi: 10.13287/j.1001-9332.201705.009
[16]	张英华, 杨佑明, 曹莲, 郝杨凡, 黄菁, 李金鹏, 姚得秀, 王志敏. 灌浆期高温对小麦旗叶与非叶器官光合和抗氧化酶活性的影响. 作物学报, 2015, 41(1): 136-144. doi: 10.3724/SP.J.1006.2015.00136
	ZHANG Y H, YANG Y M, CAO L, HAO Y F, HUANG J, LI J P, YAO D X, WANG Z M. Effect of high temperature on photosynthetic capability and antioxidant enzyme activity of flag leaf and non-leaf organs in wheat. Acta Agronomica Sinica, 2015, 41(1): 136-144. (in Chinese) doi: 10.3724/SP.J.1006.2015.00136
[17]	MAESTRI E, KLUEVA N, PERROTTA C, GULLI M, NGUYEN H T, MARMIROLI N. Molecular genetics of heat tolerance and heat shock proteins in cereals. Plant Molecular Biology, 2002, 48(5): 667-681. doi: 10.1023/A:1014826730024
[18]	MORALES D, RODRIGUEZ P, DELL'AMICO J, NICOLAS E, TORRECILLAS A, SANCHEZ-BLANCO M J. High temperature preconditioning and thermal shock imposition affects water relations, gas exchange and root hydraulic conductivity in tomato. Biologia Plantarum, 2003, 47(2): 203-208.
[19]	贾双杰, 李红伟, 江艳平, 赵国强, 王和洲, 杨慎骄, 杨青华, 郭家萌, 邵瑞鑫. 干旱胁迫对玉米叶片光合特性和穗发育特征的影响. 生态学报, 2020, 40(3): 854-863.
	JIA S J, LI H W, JIANG Y P, ZHAO G Q, WANG H Z, YANG S J, YANG Q H, GUA J M, SHAO R X. Effects of drought on photosynthesis and ear development characteristics of maize. Acta Ecologica Sinica, 2020, 40(3): 854-863. (in Chinese)
[20]	穆心愿, 马智艳, 张兰薰, 付景, 刘天学, 丁勇, 夏来坤, 张凤启, 张君, 齐建双, 赵霞, 唐保军. 不同耐/感玉米品种的叶片光合荧光特性、授粉结实和产量构成因素对花期高温的反应. 中国生态农业学报(中英文), 2022, 30(1): 57-71.
	MU X Y, MA Z Y, ZHANG L X, FU J, LIU T X, DING Y, XIA L K, ZHANG F Q, ZHANG J, QI J S, ZHAO X, TANG B J. Responses of the photosynthetic fluorescence characteristics, pollination and yield compositions of different tolerant/susceptible maize varieties to high temperature during flowering. Chinese Journal of Eco-Agriculture, 2022, 30(1): 57-71. (in Chinese)
[21]	闫振华, 刘东尧, 贾绪存, 杨琴, 陈艺博, 董朋飞, 王群. 花期高温干旱对玉米雄穗发育、生理特性和产量影响. 中国农业科学, 2021, 54(17): 3592-3608.
	YAN Z H, LIU D Y, JIA X C, YANG Q, CHEN Y B, DONG P F, WANG Q. Maize tassel development, physiological traits and yield under heat and drought stress during flowering stage. Scientia Agricultura Sinica, 2021, 54(17): 3592-3608. (in Chinese)
[22]	张翼飞. 施氮对甜菜氮素同化与碳代谢的调控机制研究[D]. 哈尔滨: 东北农业大学, 2013.
	ZHANG Y F. Study on regulation mechanism of nitrogen application on nitrogen assimilation and carbon metabolism in sugar beet (Beta Vulgaris L.)[D]. Harbin: Dongbei Agricultural University, 2013. (in Chinese)
[23]	魏爱丽, 张英华, 黄琴, 王志敏. 小麦不同绿色器官光合速率与碳同化酶活性及其基因型差异研究. 作物学报, 2007, 33(9): 1426-1431.
	WEI A L, ZHANG Y H, HUANG Q, WANG Z M. Dynamic characteristics of photosynthetic rate and carbon assimilation enzyme activities of different green organs in different genotypes of wheat. Acta Agronomica Sinica, 2007, 33(9): 1426-1431. (in Chinese)
[24]	黄振喜, 王永军, 王空军, 李登海, 赵明, 柳京国, 董树亭, 王洪军, 王军海, 杨今胜. 产量15000 kg·hm^-2以上夏玉米灌浆期间的光合特性. 中国农业科学, 2007, 40(9): 1898-1906.
	HUANG Z X, WANG Y J, WANG K J, LI D H, ZHAO M, LIU J G, DONG S T, WANG H J, WANG J H, YANG J S. Photosynthetic characteristics during grain filling stage of summer maize hybrids with high yield potential of 15000 kg·ha^-1. Scientia Agricultura Sinica, 2007, 40(9): 1898-1906. (in Chinese)
[25]	方芳, 何序晨, 张志豪, 张勤, 关亚静, 胡晋, 胡伟民. 玉米自交系苗期对高温胁迫的响应机制及其抗逆性. 浙江农业学报, 2019, 31(7): 1045-1056. doi: 10.3969/j.issn.1004-1524.2019.07.03
	FANG F, HE X C, ZHANG Z H, ZHANG Q, GUAN Y J, HU J, HU W M. Response mechanism and stress resistance of maize inbred lines to high temperature stress at seedling stage. Acta Agriculturae Zhejiangensis, 2019, 31(7): 1045-1056. (in Chinese) doi: 10.3969/j.issn.1004-1524.2019.07.03
[26]	曲明南. CO₂升高和短期高温胁迫对玉米幼苗生理生化指标的影响[D]. 沈阳: 沈阳农业大学, 2013.
	QU M N. Effects of elevated CO₂ and short term heat stress on physiological and biochemical variables in maize seedlings[D]. Shenyang: Shenyang Agricultural University, 2013. (in Chinese)
[27]	付景, 孙宁宁, 刘天学, 杨豫龙, 赵霞, 李潮海. 高温胁迫对玉米形态、叶片结构及其产量的影响. 玉米科学, 2019, 27(1): 46-53.
	FU J, SUN N N, LIU T X, YANG Y L, ZHAO X, LI C H. Effect of high temperature stress on morphology, leaf structure and grain yield of maize. Journal of Maize Sciences, 2019, 27(1): 46-53. (in Chinese)
[28]	陈岩, 岳丽杰, 杨勤, 张会玲, 柯国华, 刘永红. 高温热害对玉米生长发育的影响及研究进展. 耕作与栽培, 2019, 39(1): 26-31.
	CHEN Y, YUE L J, YANG Q, ZHANG H L, KE G H, LIU Y H. Effects of high temperature on maize development and its research advances. Tillage and Cultivation, 2019, 39(1): 26-31. (in Chinese)
[29]	FUKAYAMA H, UEGUCHI C, NISHIKAWA K, KATOH N, ISHIKAWA C, MASUMOTO C, HATANAKA T, MISOO S. Overexpression of rubisco activase decreases the photosynthetic CO₂ assimilation rate by reducing rubisco content in rice leaves. Plant & Cell Physiology, 2012, 53(6): 976-986.
[30]	LIAN L, LIN Y L, WEI Y D, HE W, CAI Q H, HUANG W, ZHENG Y M, XU H B, WANG F X, ZHU Y S, LUO X, XIE H A, ZHANG J F. PEPC of sugarcane regulated glutathione S-transferase and altered carbon-nitrogen metabolism under different N source concentrations in Oryza sativa. BMC Plant Biology, 2021, 21(1): 287. doi: 10.1186/s12870-021-03071-w
[31]	毕焕改, 董绪兵, 王美玲, 艾希珍. 钙和水杨酸对亚适温弱光下黄瓜幼苗光合酶活性和基因表达的影响. 园艺学报, 2015, 42(1): 56-64.
	BI H G, DONG X B, WANG M L, AI X Z. Foliar spray calcium and salicylic acid improve the activities and gene expression of photosynthetic enzymes in cucumber seedlings under low light intensity and suboptimal temperature. Acta Horticulturae Sinica, 2015, 42(1): 56-64. (in Chinese)
[32]	罗宏海, 李俊华, 张宏芝, 何在菊, 勾玲, 张旺锋. 源库调节对新疆高产棉花产量形成期光合产物生产与分配的影响. 棉花学报, 2009, 21(5): 371-377.
	LUO H H, LI J H, ZHANG H Z, HE Z J, GOU L, ZHANG W F. Effects of source and sink manipulation on transportation and allocation of leaf photosynthetic products during flowering and boll setting stage in high yield cotton of Xinjiang. Cotton Science, 2009, 21(5): 371-377. (in Chinese)
[33]	于康珂, 刘源, 李亚明, 孙宁宁, 詹静, 尤东玲, 牛丽, 李潮海, 刘天学. 玉米花期耐高温品种的筛选与综合评价. 玉米科学, 2016, 24(2): 62-71.
	YU K K, LIU Y, LI Y M, SUN N N, ZHAN J, YOU D L, NIU L, LI C H, LIU T X. Screening and comprehensive evaluation of heat tolerance of maize hybrids in flowering stage. Journal of Maize Sciences, 2016, 24(2): 62-71. (in Chinese)
[34]	高英波, 张慧, 单晶, 薛艳芳, 钱欣, 代红翠, 刘开昌, 李宗新. 吐丝前高温胁迫对不同耐热型夏玉米产量及穗发育特征的影响. 中国农业科学, 2020, 53(19): 3954-3963.
	GAO Y B, ZHANG H, SHAN J, XUE Y F, QIAN X, DAI H C, LIU K C, LI Z X. Effects of pre-silking high temperature stress on yield and ear development characteristics of different heat-resistant summer maize cultivars. Scientia Agricultura Sinica, 2020, 53(19): 3954-3963. (in Chinese)
[35]	许振柱, 周广胜. 全球变化下植物的碳氮关系及其环境调节研究进展-从分子到生态系统. 植物生态学报, 2007, 31(4): 738-747. doi: 10.17521/cjpe.2007.0094
	XU Z Z, ZHOU G S. Relationship between carbon and nitrogen and environmental regulation in plants under global change-From molecule to ecosystem. Journal of Plant Ecology, 2007, 31(4): 738-747. (in Chinese) doi: 10.17521/cjpe.2007.0094

年份 Year	处理 Treatment	平均光照强度 Average light intensity (μmol·m^-2·s^-1)	平均CO₂浓度 Average CO₂ concentration (μmol·mol^-1)	相对湿度 Relative humidity (%)
2019	CK	1141.6a	420.0a	80.0a
2019	HT	1136.8a	416.3a	78.8a
2020	CK	1127.3a	419.3a	84.5a
2020	HT	1114.5a	411.9a	82.8a

年份 Year	处理时期 Treatments stage	处理 Treatment	¹³C同化量 ¹³C assimilate amount (mg/plant)				所占比例 Proportion (%)
年份 Year	处理时期 Treatments stage	处理 Treatment	总量 Total	茎秆 Stem	叶片 Leaf	籽粒 Grain	茎秆 Stem	叶片 Leaf	籽粒 Grain
2019	V12	ZD-CK	20.02a	4.95a	2.51d	12.56a	24.73	12.54	62.74
		ZD-HT	16.32bc	6.25a	4.18c	5.87b	38.30	25.61	35.97
		XY-CK	19.19ab	4.86a	2.30d	12.03a	25.33	11.99	62.69
		XY-HT	14.81c	6.52a	4.55b	3.73d	44.02	30.72	25.19
	VT	ZD-CK	19.70a	4.86a	2.53d	12.31a	24.67	12.84	62.49
		ZD-HT	15.64c	6.97a	4.13c	4.55c	44.57	26.41	29.09
		XY-CK	19.74a	4.73a	2.37d	12.63a	23.96	12.01	63.98
		XY-HT	13.34c	6.18a	5.05a	2.12e	46.33	37.86	15.89
2020	V12	ZD-CK	20.69a	4.53a	4.07a	12.09b	21.89	19.67	58.43
		ZD-HT	18.33bc	6.70a	4.19a	7.43c	36.55	22.86	40.53
		XY-CK	20.07ab	4.42a	3.86a	11.8b	22.02	19.23	58.79
		XY-HT	14.90de	6.63a	4.12a	4.16e	44.50	27.65	27.92
	VT	ZD-CK	21.87a	4.40a	3.89a	13.57a	20.12	17.79	62.05
		ZD-HT	16.96cd	6.50a	5.38a	5.10d	38.33	31.72	30.07
		XY-CK	21.51a	5.10a	4.10a	12.31b	23.71	19.06	57.23
		XY-HT	13.59e	5.74a	4.57a	3.28f	42.24	33.63	24.14

年份 Year	处理时期 Treatment stage	处理 Treatment	干物质积累量 Dry matter accumulate amount (g/plant)			穗粒数 Grains per ear	千粒重 1000-grain weight (g)	产量 Yield (g/plant)	收获指数 HI
年份 Year	处理时期 Treatment stage	处理 Treatment	大喇叭口期 V12	开花期 VT	完熟期 R6	穗粒数 Grains per ear	千粒重 1000-grain weight (g)	产量 Yield (g/plant)	收获指数 HI
2019	V12	ZD-CK	111.63a	143.15c	311.50bc	591.2a	299.12bc	151.93a	0.49a
		ZD-HT	90.00c	120.78e	219.68d	219.5d	306.17a	82.25b	0.37c
		XY-CK	104.28b	157.03b	308.48c	515.2c	298.64bc	153.26a	0.49a
		XY-HT	76.42d	115.89e	211.52d	152.0e	308.10a	57.44c	0.26d
	VT	ZD-CK	—	153.35b	329.54a	562.6ab	293.87c	159.58a	0.47b
		ZD-HT	—	133.68d	221.2d	165.8e	304.90ab	52.83c	0.24e
		XY-CK	—	177.53a	320.09ab	544.3bc	293.70c	151.26a	0.48ab
		XY-HT	—	138.35cd	199.92e	79.4f	301.83ab	29.33d	0.14f
2020	V12	ZD-CK	97.31a	140.18b	311.12b	571.1a	305.195d	173.46a	0.56ab
		ZD-HT	86.80b	116.49c	275.84c	247.1b	311.663bc	99.69d	0.35d
		XY-CK	84.04b	140.34b	284.78c	545.5a	309.672c	161.39bc	0.58a
		XY-HT	64.73c	102.42d	243.80e	188.1cd	318.725a	64.47f	0.26ef
	VT	ZD-CK	—	152.81a	326.09a	544.5a	311.207bc	169.35ab	0.51c
		ZD-HT	—	139.68b	260.05d	194.9c	314.107b	77.92e	0.28e
		XY-CK	—	146.00ab	300.27b	550.4a	309.438c	157.59c	0.53bc
		XY-HT	—	124.94c	212.47f	151.9d	321.509a	52.77g	0.24f