不同滴灌模式对东北春播玉米籽粒淀粉积累及淀粉相关酶活性的影响

doi:10.3864/j.issn.0578-1752.2022.07.006

摘要/Abstract

摘要：

【目的】探明不同滴灌模式对春玉米籽粒淀粉积累及淀粉合成相关酶活性的影响,为提高春玉米产量提供理论指导。【方法】以农华101为试验材料,采用裂区试验,主处理为滴灌模式,设膜下滴灌和浅埋滴灌2种,副处理为灌水量,设传统灌水量40%（W1：1 440 m³·hm^-2）、传统灌水量50%（W2：1 800 m³·hm^-2）和传统灌水量60%（W3：2 160 m³·hm^-2）3 个水平,于苗期—拔节期、拔节期—大喇叭口期、大喇叭口期—吐丝期、吐丝期—乳熟期、乳熟期—收获按照1﹕2﹕2﹕3﹕2比例滴灌。吐丝后20 d开始,每7 d取玉米籽粒1次,采用酸水解-DNS法测定玉米籽粒中总淀粉含量,采用淀粉合成酶试剂盒测定腺苷二磷酸葡萄糖焦磷酸化酶（AGPase）、结合态淀粉合成酶（GBSS）、可溶性淀粉合成酶（SSS）活性,研究玉米籽粒产量、淀粉积累量特征及AGPase、GBSS、SSS酶活性变化。【结果】2019—2020年春玉米有效穗数和穗粒数两种模式间差异不显著, W1处理浅埋滴灌千粒重和产量低于膜下滴灌,W2处理二者无显著性差异,W3处理显著高于膜下滴灌,其中产量分别高5.7%和8.4%,千粒重分别高11.9%、12.1%。方差分析表明,灌水量、滴灌模式和灌水量与滴灌模式互作对千粒重影响差异达显著、极显著水平,年份、灌水量及灌水量与滴灌模式互作对籽粒产量影响达到显著或极显著水平。W1处理浅埋滴灌籽粒淀粉含量和积累量低于膜下滴灌,W2处理从吐丝后20 d起浅埋滴灌低于膜下滴灌,吐丝后55 d二者差异不显著,W3处理吐丝41 d前低于膜下滴灌,随生育进程差异减小,吐丝后55 d显著高于膜下滴灌。3个灌水量下浅埋滴灌的籽粒淀粉活跃积累期较膜下滴灌时间长,且籽粒淀粉积累达到最大速率的时间较膜下滴灌时间延后。活跃积累期对淀粉最终积累量影响系数最大,其次为达到最大速率的时间,生育后期淀粉积累速率与籽粒淀粉总积累量极显著相关、与产量显著相关。浅埋滴灌下吐丝后20—27 d AGPase、GBSS和SSS 3种酶活性3个灌水量下均低于膜下滴灌,随生育进程推移差异减小,从吐丝后48 d起W3处理高于膜下滴灌,吐丝后55 d W1和W2处理亦高于膜下滴灌。除W1处理下籽粒淀粉积累速率与AGPase焦磷酸化酶无显著性相关外,其他处理籽粒淀粉积累速率与AGPase、GBSS、SSS酶活性呈显著性正相关。【结论】在传统灌水量60%时,浅埋滴灌下春玉米籽粒灌浆后期淀粉合成相关酶（AGPase、GBSS、SSS酶）活性强、淀粉活跃积累期延长,淀粉积累能力增强,千粒重增加,籽粒产量最高。浅埋滴灌优化灌溉可通过提高籽粒灌浆后期淀粉合成酶活性,增强淀粉积累能力,进而增加粒重、提高产量,达到节水增效的目的。

关键词: 滴灌模式, 春玉米, 籽粒, 酶活性, 淀粉积累

Abstract:

【Objective】The aim of this study was to explore the effects of different drip irrigation modes on starch accumulation and starch synthesis-related enzyme activities of spring maize, so as to provide a physiological basis for the understanding of the yield formation process in maize under the water-saving conditions. 【Method】 Using maize variety Nonghua 101 as the experimental material, a sub-plot design was selected for this experiment, and two irrigation modes were chosen as the main plot, including mulched drip irrigation (MDI) and shallow drip irrigation (SBDI), and the irrigation amount as the sub-plot included three irrigation levels (W1: 1 440 m³·hm^-2; W2: 1 800 m³·hm^-2; W3: 2 160 m³·hm^-2, which equaled to the amount of 40%, 50% and 60% of the traditional irrigation, respectively). The amount and time of drip irrigation was conducted at the ratio of 1﹕2﹕2﹕3﹕2 for seedling stage to jointing stage, jointing stage to big bell mouth, big bell mouth to silking stage, silking stage to milk stage, and milk stage to harvest stage, respectively. The corn kernels were taken at every 7 days from 20 days after silking. The content of total starch in maize kernel was determined by acid hydrolysis DNS method. The starch synthase activity assay kit was used to determine the activity of adenosine diphosphate glucose pyro phosphorylase (ADPase), bound starch synthase (GBSS), and soluble starch synthase (SSS). On basis of that, the maize grain yield, characteristics of starch accumulation, and the changes enzyme activity of ADPase, GBSS and SSS were investigaged. 【Result】There was no significant difference between effective panicle number and grain number per panicle of the two irrigation modes, and the 1000-grain weight and grain yield in shallow drip irrigation was lower than that of mulched drip irrigation under the treatment of W1, but no significant difference were found under the treatment of W2. The grain yield and 1000-grain weight in shallow drip irrigation were 5.7% and 8.4% higher in grain yield, and 11.9% and 12.1% higher in 1000-grain weight than that in mulched drip irrigation under treatment W3 in 2019 and 2020, respectively. The results of variance analysis showed that irrigation amount, drip irrigation mode and the interaction between irrigation amount and drip irrigation mode had a significant effects (P<0.01 or P<0.001) on 1000 grain weight, and the interaction between years, irrigation amount and drip irrigation mode had a significant effects (P<0.01 or P<0.001) on grain yield. Considering the content and accumulation of starch in grain, the shallow drip irrigation were both lower than that of mulched drip irrigation of W1, and W2 was lower than mulched drip irrigation in 20 days after silking, while no significant difference in 55 days after silking; W3 was lower than mulched drip irrigation in 41 days after silking, while higher than in 55 days after silking. Under the three irrigation amounts, the active accumulation period of starch in grain under shallow drip irrigation was longer than that of mulched drip irrigation, and the time for grain starch accumulation to reach the maximum rate was later than that of shallow drip irrigation. Active accumulation period of starch had the highest influence coefficient on the final accumulation of starch, then was the time of maximum rate. The starch accumulation rate in the late growth stage was highly correlated with the total starch accumulation in grains (P<0.001) and the yield (P<0.01). The activity of ADPase, GBSS and SSS in shallow drip irrigation were lower that of mulched drip irrigation in 20-27 d after silking, while the difference was reducing along with the maize growth; The three enzyme activity were higher under W3 than that of mulched drip irrigation at 48 days after silking, and also under W1 and W2 at 55 days after silking. Except no relationship of accumulation rate of grain starch and activity of ADPase under W1, the accumulation rate of grain starch showed a significantly positive relation with the activities of ADPase, GBSS and SSS under other treatments. 【Conclusion】With the 60% of the traditional irrigation amount in shallow drip irrigation, it showed a higher activity of ADPase, GBSS and SSS at the late filling stage and a longer active starch accumulation period, also enhanced the ability of starch accumulation and increased 1000-grain weight, and possessed the highest maize grain yield. The shallow drip irrigation could increase the activity of the maize starch synthases, enhance the ability of starch accumulation, then increased grain weight and yield, and finally reached the purpose of water-saving and efficiency-improvement.

Key words: drip irrigation modes, spring maize, grain, enzyme activity, starch accumulation

张家桦,杨恒山,张玉芹,李从锋,张瑞富,邰继承,周阳晨. 不同滴灌模式对东北春播玉米籽粒淀粉积累及淀粉相关酶活性的影响[J]. 中国农业科学, 2022, 55(7): 1332-1345.

ZHANG JiaHua,YANG HengShan,ZHANG YuQin,LI CongFeng,ZHANG RuiFu,TAI JiCheng,ZHOU YangChen. Effects of Different Drip Irrigation Modes on Starch Accumulation and Activities of Starch Synthesis-Related Enzyme of Spring Maize Grain in Northeast China[J]. Scientia Agricultura Sinica, 2022, 55(7): 1332-1345.

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.0578-1752.2022.07.006

https://www.chinaagrisci.com/CN/Y2022/V55/I7/1332

图/表 12

表1

表2

表3

图1

图2

表4

表5

图3

图4

图5

图6

图7

参考文献 52

[1]	张玉芹, 杨恒山, 高聚林, 张瑞富, 王志刚, 徐寿军, 范秀艳, 毕文波. 超高产春玉米的根系特征. 作物学报, 2011, 37(4):735-743. doi: 10.3724/SP.J.1006.2011.00735
	ZHANG Y Q, YANG H S, GAO J L, ZHANG R F, WANG Z G, XU S J, FAN X Y, BI W B. Root characteristics of super high-yield spring maize. Acta Agronomica Sinica, 2011, 37(4):735-743. (in Chinese) doi: 10.3724/SP.J.1006.2011.00735
[2]	白晓慧. 通辽市2005年—2010年地下水位变化分析. 内蒙古科技与经济, 2016(16):65-66.
	BAI X H. Analysis of groundwater level changes in Tongliao city from 2005 to 2010. Inner Mongolia Science Technology & Economy, 2016(16):65-66. (in Chinese)
[3]	曹玉军, 魏雯雯, 徐国安, 王晓慧, 王洪君, 刘春光, 边少锋, 刘慧涛, 王永军. 半干旱区不同地膜覆盖滴灌对土壤水、温变化及玉米生长的影响. 玉米科学, 2013, 21(1):107-113.
	CAO Y J, WEI W W, XU G A, WANG X H, WANG H J, LIU C G, BIAN S F, LIU H T, WANG Y J. Effects of differed films on soil water, temperature and corn growth characteristics under drip- irrigation conditions in semi-arid region. Journal of Maize Sciences, 2013, 21(1):107-113. (in Chinese)
[4]	杜利, 李援农, 陈朋朋, 王凯瑜, 李昱鹏. 不同残膜量对土壤环境及玉米生长发育的影响. 节水灌溉, 2018(7):4-9, 14.
	DU L, LI Y N, CHEN P P, WANG K Y, LI Y P. Effects of different residual film on the growth and soil environment of maize. Water Saving Irrigation, 2018(7):4-9, 14. (in Chinese)
[5]	林涛, 汤秋香, 郝卫平, 吴凤全, 雷蕾, 严昌荣, 何文清, 梅旭荣. 地膜残留量对棉田土壤水分分布及棉花根系构型的影响. 农业工程学报, 2019, 35(19):117-125.
	LIN T, TANG Q X, HAO W P, WU F Q, LEI L, YAN C R, HE W Q, MEI X R. Effects of plastic film residue rate on root zone water environment and root distribution of cotton under drip irrigation condition. Transactions of the Chinese Society of Agricultural Engineering, 2019, 35(19):117-125. (in Chinese)
[6]	解红娥, 李永山, 杨淑巧, 王娇娟, 吴秀峰, 武宗信. 农田残膜对土壤环境及作物生长发育的影响研究. 农业环境科学学报, 2007, 26(S1):153-156.
	XIE H E, LI Y S, YANG S Q, WANG J J, WU X F, WU Z X. Influence of residual plastic film on soil structure, crop growth and development in fields. Journal of Agro-Environment Science, 2007, 26(S1):153-156. (in Chinese)
[7]	杨恒山, 薛新伟, 张瑞富, 李金琴, 王宇飞, 邰继承, 刘晶. 灌溉方式对西辽河平原玉米产量及水分利用效率的影响. 农业工程学报, 2019, 35(21):69-77.
	YANG H S, XUE X W, ZHANG R F, LI J Q, WANG Y F, TAI J C, LIU J. Effects of irrigation methods on yield and water use efficiency of maize in the West Liaohe Plain. Transactions of the Chinese Society of Agricultural Engineering, 2019, 35(21):69-77. (in Chinese)
[8]	梅园雪, 冯玉涛, 冯天骄, 汪伟, 孙宝忠. 玉米浅埋滴灌节水种植模式产量与效益分析. 玉米科学, 2018, 26(1):98-102.
	MEI Y X, FENG Y T, FENG T J, WANG W, SUN B Z. Brief discussion on the efficient water-saving planting mode of shallow buried drip irrigation. Journal of Maize Sciences, 2018, 26(1):98-102. (in Chinese)
[9]	KAUR A, SINGH N, EZEKIEL R, GURAYAL H S. Physicochemical, thermal and pasting properties of starches separated from different potato cultivars grown at different locations. Food Chemistry, 2007, 101(2):643-651. doi: 10.1016/j.foodchem.2006.01.054
[10]	李志伟, 钟雨越, 吴权明, 王文斌, 高杰, 刘香香, 康慧敏, 郭东伟, 薛吉全. 高直链玉米淀粉的理化特性研究. 西北农林科技大学学报(自然科学版), 2014, 42(7):53-60.
	LI Z W, ZHONG Y Y, WU Q M, WANG W B, GAO J, LIU X X, KANG H M, GUO D W, XUE J Q. Physicochemical properties of high-amylose maize starch. Journal of Northwest A & F University (Natural Science Edition), 2014, 42(7):53-60. (in Chinese)
[11]	XIAO Q L, WANG Y Y, LI H, ZHANG C X, WEI B, WANG Y B, HUNAG H H, LI Y P, YU G W, LIU H M, ZHANG J J, LIU Y H, HU Y F, HUANG Y B. Transcription factor ZmNAC126 plays an important role in transcriptional regulation of maize starch synthesis- related genes. The Crop Journal, 2021, 9(1):192-203. doi: 10.1016/j.cj.2020.04.014
[12]	胡文河, 谷岩, 岳杨, 常莹, 吴春胜. 不同株型玉米籽粒淀粉积累及相关酶活性的研究. 西北农林科技大学学报(自然科学版), 2012, 40(3):104-110.
	HU W H, GU Y, YUE Y, CHANG Y, WU C S. Analysis of starch accumulation and key enzymes activities for starch synthesis in different genotype maize. Journal of Northwest A&F University (Natural Science Edition), 2012, 40(3):104-110. (in Chinese)
[13]	陈江, 王燕, 黄斌全, 胡玉峰, 刘应红, 顾勇, 李炀平, 张军杰, 刘汉梅, 黄玉碧. 不同类型玉米籽粒淀粉积累、相关酶活及基因表达差异分析. 核农学报, 2012, 26(2):217-230.
	CHEN J, WANG Y, HUANG B Q, HU Y F, LIU Y H, GU Y, LI Y P, ZHANG J J, LIU H M, HUANG Y B. Analysis of starch accumulation, corresponding enzyme activity and gene expression among different types of corns. Journal of Nuclear Agricultural Sciences, 2012, 26(2):217-230. (in Chinese)
[14]	KEELING P L, BANISADR R, BARONE L, WASSERMAN B P, SINGLETARY G W. Effect of temperature on enzymes in the pathway of starch biosynthesis in developing wheat and maize grain. Functional Plant Biology, 1994, 21(6):807-827. doi: 10.1071/PP9940807
[15]	杨毅, 李昱, 康建宏, 刘萍. 花后高温胁迫对春小麦籽粒淀粉合成的影响. 麦类作物学报, 2015, 35(11):1535-1541.
	YANG Y, LI Y, KANG J H, LIU P. Effect of heat stress after anthesis on starch synthesis in spring wheat. Journal of Triticeae Crops, 2015, 35(11):1535-1541. (in Chinese)
[16]	杜晓东, 赵宏伟, 王敬国, 刘化龙, 杨亮, 许晶, 宋谨同. 氮肥运筹对寒地粳稻淀粉合成关键酶活性及淀粉积累的影响. 作物学报, 2012, 38(1):159-167. doi: 10.3724/SP.J.1006.2012.00159
	DU X D, ZHAO H W, WANG J G, LIU H L, YANG L, XU J, SONG J T. Changes in starch accumulation and activity of enzymes associated with starch synthesis under different nitrogen applications in japonica rice in cold region. Acta Agronomica Sinica, 2012, 38(1):159-167. (in Chinese) doi: 10.3724/SP.J.1006.2012.00159
[17]	WALLWORK M A B, LOGUE S J, MACLEOD L C, JENNER C F. Effect of high temperature during grain filling on starch synthesis in the developing barley grain. Australian Journal of Plant Physiology, 1998, 25(3):173-181.
[18]	赵辉, 戴廷波, 荆奇, 姜东, 曹卫星, 陆玮, 田孝威. 灌浆期高温对两种品质类型小麦品种籽粒淀粉合成关键酶活性的影响. 作物学报, 2006, 32(3):423-429.
	ZHAO H, DAI T B, JING Q, JIANG D, CAO W X, LU W, TIAN X W. Effects of high temperature during grain filling on key enzymes involved in starch synthesisin two wheat cultivars with different quality types. Acta Agronomica Sinica, 2006, 32(3):423-429. (in Chinese)
[19]	王月福, 于振文, 李尚霞, 余松烈. 小麦籽粒灌浆过程中有关淀粉合成酶的活性及其效应. 作物学报, 2003, 29(1):75-81. doi: 10.3724/SP.J.1095.2011.00054
	WANG Y F, YU Z W, LI S X, YU S L. Activity of enzymes related to starch synthesis and their effect during the filling of winter wheat. Acta Agronomica Sinica, 2003, 29(1):75-81. (in Chinese) doi: 10.3724/SP.J.1095.2011.00054
[20]	吕艳梅, 谭伟平, 肖层林, 范美蓉, 廖育林. 高温对优质水稻籽粒淀粉形成及淀粉合成相关酶活性的影响. 华北农学报, 2014, 29(1):135-139.
	LÜ Y M, TAN W P, XIAO C L, FAN M R, LIAO Y L. Effect of high temperature on starch formation of grain and activities of enzymes related to starch synthesis of quality rice varieties. Acta Agriculture Boreali-Sinica, 2014, 29(1):135-139. (in Chinese)
[21]	成臣, 曾勇军, 程慧煌, 谭雪明, 商庆银, 曾研华, 石庆华. 齐穗至乳熟期不同温度对水稻南粳9108籽粒激素含量、淀粉积累及其合成关键酶活性的影响. 中国水稻科学, 2019, 33(1):57-67.
	CHENG C, ZENG Y J, CHENG H H, TAN X M, SHANG Q Y, ZENG Y H, SHI Q H. Effects of different temperature from full heading to milking on grain filling stage on grain hormones concentrations, activities of enzymes involved in starch synthesis and accumulation in rice Nanjing 9108. Chinese Journal of Rice Science, 2019, 33(1):57-67. (in Chinese)
[22]	AHMADI A, BAKER D A. The effect of water stress on the activities of key regulatory enzymes of the sucrose to starch pathway in wheat. Plant Growth Regulation, 2001, 35(1):81-91. doi: 10.1023/A:1013827600528
[23]	石慧清, 龚月桦, 张东武. 花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响. 植物生态学报, 2011, 35(7):769-778. doi: 10.3724/SP.J.1258.2011.00769
	SHI H Q, GONG Y H, ZHANG D W. Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis. Chinese Journal of Plant Ecology, 2011, 35(7):769-778. (in Chinese) doi: 10.3724/SP.J.1258.2011.00769
[24]	闫素辉, 尹燕枰, 李文阳, 梁太波, 李勇, 邬云海, 王平, 耿庆辉, 戴忠民, 王振林. 灌浆期高温对小麦籽粒淀粉的积累、粒度分布及相关酶活性的影响. 作物学报, 2008, 34(6):1092-1096. doi: 10.3724/SP.J.1006.2008.01092
	YAN S H, YIN Y P, LI W Y, LIANG T B, LI Y, WU Y H, WANG P, GENG Q H, DAI Z M, WANG Z L. Effect of high temperature during grain filling on starch accumulation, starch granule distribution, and activities of related enzymes in wheat grains. Acta Agronomica Sinica, 2008, 34(6):1092-1096. (in Chinese) doi: 10.3724/SP.J.1006.2008.01092
[25]	汪敏, 王邵宇, 吴佳佳, 许开放, 汪涛, 何启方, 邢肖丽, 姚文政, 张文静. 花后阴雨对小麦籽粒淀粉合成和干物质积累的影响. 中国生态农业学报(中英文), 2020, 28(1):76-85.
	WANG M, WANG S Y, WU J J, XU K F, WANG T, HE Q F, XING X L, YAO W Z, ZHANG W J. Effects of shading and waterlogging following anthesis on starch synthesis and dry matter accumulation in wheat grain. Chinese Journal of Eco-Agriculture, 2020, 28(1):76-85. (in Chinese)
[26]	李双, 司转运, 申孝军, 高阳, 段爱旺. 水氮供应对灌浆期冬小麦籽粒淀粉合成相关酶活性及产量的影响. 麦类作物学报, 2018, 38(4):460-468.
	LI S, SI Z Y, SHEN X J, GAO Y, DUAN A W. Effect of different water and nitrogen levels on starch synthesis enzyme activity in wheat grains during grain filling stage and wheat yield. Journal of Triticeae Crops, 2018, 38(4):460-468. (in Chinese)
[27]	张智猛, 戴良香, 胡昌浩, 董树亭, 王空军, 宁堂原. 玉米灌浆期水分差异供应对籽粒淀粉积累及其酶活性的影响. 植物生态学报, 2005, 29(4):636-643. doi: 10.17521/cjpe.2005.0085
	ZHANG Z M, DAI L X, HU C H, DONG S T, WANG K J, NING T Y. Effects of different water treatments on starch accumulation and related enzyme activity in grain of maize during grain-filling period. Chinese Journal of Plant Ecology, 2005, 29(4):636-643. (in Chinese) doi: 10.17521/cjpe.2005.0085
[28]	薛万来, 牛文全, 张俊. 膜下滴灌土壤水盐运移研究进展. 滴灌排水学报, 2013, 32(4):114-118.
	XUE W L, NIU W Q, ZHANG J. Review of study on soil water and salt transfer under mulched drip irrigation. Journal of Irrigation and Drainage, 2013, 32(4):114-118. (in Chinese)
[29]	何照范. 粮油籽粒品质及其分析技术. 北京: 中国农业出版社, 1985.
	HE Z F. Analysis Technique for Grain Quality of Cereals and Oils. Beijing: China Agriculture Press, 1985. (in Chinese)
[30]	张吉旺, 董树亭, 王空军, 胡昌浩, 刘鹏. 大田遮阴对夏玉米淀粉合成关键酶活性的影响. 作物学报, 2008, 34(8):1470-1474. doi: 10.3724/SP.J.1006.2008.01470
	ZHANG J W, DONG S J, WANG K J, HU C H, LIU P. Effects of shading in field on key enzymes involved in starch synthesis of summer maize. Acta Agronomica Sinica, 2008, 34(8):1470-1474. (in Chinese) doi: 10.3724/SP.J.1006.2008.01470
[31]	胡阳阳, 卢红芳, 刘卫星, 康娟, 马耕, 李莎莎, 褚莹莹, 王晨阳. 灌浆期高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性及淀粉积累的影响. 作物学报, 2018, 44(4):591-600. doi: 10.3724/SP.J.1006.2018.00591
	HU Y Y, LU H F, LIU W X, KANG J, MA G, LI S S, ZHU Y Y, WANG C Y. Effects of high temperature and water deficiency during grain filling on activities of key starch synthesis enzymes and starch accumulation in wheat. Acta Agronomica Sinica, 2018, 44(4):591-600. (in Chinese) doi: 10.3724/SP.J.1006.2018.00591
[32]	RANE J, NAGARAJAN S. High temperature index-for field evaluation of heat tolerance in wheat varieties. Agricultural Systems, 2004, 79(2):243-255. doi: 10.1016/S0308-521X(03)00075-1
[33]	ZHANG H Y, DONG S T, GAO R Q, QUAN L Y. Comparison of starch synthesis and related enzyme activities in developing grains among different types of maize. Journal of plant physiology and molecular biology, 2007, 33(1):25-32.
[34]	李莎莎, 马耕, 刘卫星, 康娟, 陈雨露, 胡阳阳, 张盼盼, 王晨阳. 大田长期水氮处理对土壤氮素及小麦籽粒淀粉糊化特性的影响. 作物学报, 2018, 44(7):1067-1076. doi: 10.3724/SP.J.1006.2018.01067
	LI S S, MA G, LIU W X, KANG J, CHEN Y L, HU Y Y, ZHANG P P, WANG C Y. Effects of long-term irrigation and nitrogen regimes on soil nitrogen content and paste property of wheat grain. Acta Agronomica Sinica, 2018, 44(7):1067-1076. (in Chinese) doi: 10.3724/SP.J.1006.2018.01067
[35]	王龙飞, 杨倩, 李广浩, 陆卫平, 陆大雷. 吐丝后不同阶段干旱胁迫对糯玉米籽粒产量和淀粉品质的影响. 玉米科学, 2021, 29(1):69-76.
	WANG L F, YANG Q, LI G H, LU W P, LU D L. Effect of drought stress at different post-silking stages on grain yield and starch quality of waxy maize. Journal of Maize Sciences, 2021, 29(1):69-76. (in Chinese)
[36]	杜社妮, 白岗栓. 玉米地膜覆盖的土壤环境效应. 干旱地区农业研究, 2007, 25(5):56-59.
	DU S N, BAI G S. Studies on effects of plastic film mulching on soil environment of maize field. Agricultural Research in the Arid Areas, 2007, 25(5):56-59. (in Chinese)
[37]	孙仕军, 杨金鑫, 万博, 谷健, 刘泳圻, 赵旺, 尹光华. 不同滴灌方式对辽西半干旱区春玉米生长及产量的影响. 沈阳农业大学学报, 2021, 52(1):32-39.
	SUN S J, YANG J X, WAN B, GU J, LIU Y Q, ZHAO W, YIN G H. Effects of different drip irrigation methods on the growth and yield of spring maize in semi-arid areas of western Liaoning. Journal of Shenyang Agricultural University, 2021, 52(1):32-39. (in Chinese)
[38]	JENNER C F, UGALDE T D, ASPINALL D. The physiology of starch and protein deposition in the endosperm of wheat. Australian Journal of Plant Physiology, 1991, 18:211-226.
[39]	OU-LEE T M, SETTER T L. Effect of increased temperature in apical regions of maize ears on starch synthesis enzymes and accumulation of sugar and starch. Plant Physiology, 1985, 79(3):852-855. doi: 10.1104/pp.79.3.852
[40]	李永庚, 于振文, 姜东, 余松烈. 冬小麦旗叶蔗糖和籽粒淀粉合成动态及与其有关的酶活性的研究. 作物学报, 2001, 27(5):658-664.
	LI Y G, YU Z W, JIANG D, YU S L. Studies on the dynamic changes of the synthesis of sucrose in the flag leaf and starch in the grain and related enzymes of high-yielding wheat. Acta Agronomica Sinica, 2001, 27(5):658-664. (in Chinese)
[41]	SINGLETARY G W, BANISADR R, KEELING P L. Influence of gene dosage on carbohydrate synthesis and enzymatic activities in endosperm of starch-deficient mutants of maize. Plant Physiology, 1997, 113:293-304. doi: 10.1104/pp.113.1.293
[42]	徐云姬, 顾道健, 秦昊, 张耗, 王志琴, 杨建昌. 玉米灌浆期果穗不同部位籽粒碳水化合物积累与淀粉合成相关酶活性变化. 作物学报, 2015, 41(2):297-307. doi: 10.3724/SP.J.1006.2015.00297
	XU Y J, GU D J, QIN H, ZHANG H, WANG Z Q, YANG J C. Changes in carbohydrate accumulation and activities of enzymes involved in starch synthesis in maize kernels at different positions on an ear during grain filling. Acta Agronomica Sinica, 2015, 41(2):297-307. (in Chinese) doi: 10.3724/SP.J.1006.2015.00297
[43]	TIAN Z X, QIAN Q, LIU Q Q, YAN M X, LIU X F, YAN C J, LIU G F, GAO Z Y, TANG S Z, ZENG D L, WANG Y H, YU J M, GU M H, LI J Y. Allelic diversities in rice starch biosynthesis lead to a diverse array of rice eating and cooking qualities. Proceedings of the National Academy of Sciences of the United States of America, 2009, 106(51):21760-21765.
[44]	ZHANG H Y, DONG S T, GAO R Q, SUN Q Q. Starch accumulation and enzyme activities associated with starch synthesis in maize kernels. Agricultural Sciences in China, 2007, 6(7):808-815. doi: 10.1016/S1671-2927(07)60116-3
[45]	王晓燕, 董树亭, 高荣岐, 张海艳. 不同类型玉米胚乳细胞增殖动态及其与粒重的关系. 华北农学报, 2006, 21(2):23-26.
	WANG X Y, DONG S T, GAO R Q, ZHANG H Y. Endosperm cell proliferating and its relation to kernel weight in different types of maize. Acta Agriculturae Boreali-Sinica, 2006, 21(2):23-26. (in Chinese)
[46]	张海艳, 董树亭, 高荣岐, 李玉全. 玉米籽粒淀粉积累及相关酶活性分析. 中国农业科学, 2008, 41(7):2174-2181.
	ZHANG H Y, DONG S T, GAO R Q, LI Y Q. Starch accumulation and enzymes activities associated with starch synthesis in maize kernels. Scientia Agricultura Sinica, 2008, 41(7):2174-2181. (in Chinese)
[47]	曹颖妮, 胡卫国, 王根平, 刘录祥, 王成社. 糯性和非糯性小麦灌浆期胚乳直/支链淀粉积累及其相关酶活性研究. 西北植物学报, 2010, 30(10):1995-2001.
	CAO Y N, HU W G, WANG G P, LIU L X, WANG C S. Dynamic changes of starch accumulation and enzymes relating to starch biosynthesis of kernel during grain filling in waxy and non-waxy winter wheat. Acta Botanica Boreali-Occidentalia Sinica, 2010, 30(10):1995-2001. (in Chinese)
[48]	刘霞, 姜春明, 郑泽荣, 周筑南, 贺明荣, 王振林. 藁城8901和山农1391淀粉合成酶活性和淀粉组分积累特征的比较. 中国农业科学, 2005, 38(5):897-903.
	LIU X, JIANG C M, ZHENG Z R, ZHOU Z N, HE M R, WANG Z L. Activities of the enzymes involved in starch synthesis and starch accumulation in grains of wheat cultivars GC8901 and SN1391. Scientia Agricultura Sinica, 2005, 38(5):897-903. (in Chinese)
[49]	王自布, 李卫华, 齐军仓, 银永安, 曹连莆, 王泽民, 侯睿睿, 王亮. 小麦籽粒胚乳淀粉合成酶基因表达及酶活性分析. 核农学报, 2010, 24(6):1117-1123.
	WANG Z B, LI W H, QI J C, YIN Y A, CAO L P, WANG Z M, HOU R R, WANG L. Analysis of gene expression of enzymes involved in starch synthesis and enzymes activity in wheat grain. Journal of Nuclear Agricultural Sciences, 2010, 24(6):1117-1123. (in Chinese)
[50]	黄天琪. 结实期高温与水分胁迫对糯玉米淀粉品质的影响研究[D]. 扬州: 扬州大学, 2018.
	HUANG T Q. Effect of high temperature and water stress during grain filling on starch quality of waxy maize[D]. Yangzhou: Yangzhou University, 2018. (in Chinese)
[51]	李媛媛, 杨恒山, 张瑞富, 范秀艳, 李金琴, 罗方. 灌溉定额对浅埋滴灌春玉米生长与产量的影响. 水土保持通报, 2017, 37(2):345-348.
	LI Y Y, YANG H S, ZHANG R F, FAN X Y, LI J Q, LUO F. Effects of irrigation quota on growth and yield of spring maize under shallow subsurface drip irrigation. Bulletin of Soil and Water Conservation, 2017, 37(2):345-348. (in Chinese)
[52]	马金平, 王福星, 张岱, 王双喜. 覆膜对玉米根系分布特性的影响. 农业与技术, 2018, 38(4):21-22.
	MA J P, WANG F X, ZHANG D, WANG S X. Effect of film mulching on the distribution characteristics of maize root system. Agriculture and Technology, 2018, 38(4):21-22. (in Chinese)

处理 Treatment		各生育时期灌水量 Irrigation amount in each growth period (m³·hm^-2)					总灌水量 Total irrigation amount
处理 Treatment		苗期—拔节期 Seedling stage- Jointing stage	拔节期—大喇叭口期 Jointing stage- Flare opening stage	大喇叭口期—吐丝期 Flare opening stage- Silking stage	吐丝期—乳熟期 Silking stage- Milk maturity	乳熟期—收获期 Milk maturity- Harvest period	总灌水量 Total irrigation amount
W1	膜下滴灌MDI	144	288	288	432	288	1440
W1	浅埋滴灌SBDI	144	288	288	432	288	1440
W2	膜下滴灌MDI	180	360	360	540	360	1800
W2	浅埋滴灌SBDI	180	360	360	540	360	1800
W3	膜下滴灌MDI	216	432	432	648	432	2160
W3	浅埋滴灌SBDI	216	432	432	648	432	2160

年份 Year	处理 Irrigation method		有效穗数 Effective spike (×10⁴·hm^-2)	穗粒数 Gain number per spike	千粒重 1000 grain weight (g)	产量 Yield (t·hm^-2)
2019	W1	膜下滴灌MDI	6.6814±0.061b	473.95±4.213ab	374.97±6.538c	11.89±0.139b
	W1	浅埋滴灌SBDI	6.6725±0.061b	472.15±4.197b	356.64±9.596d	11.11±0.207c
	W2	膜下滴灌MDI	6.8719±0.063a	475.66±4.228ab	388.32±10.449bc	12.26±0.544b
	W2	浅埋滴灌SBDI	6.8961±0.063a	472.62±4.201b	378.43±10.183bc	12.04±0.557b
	W3	膜下滴灌MDI	6.9359±0.064a	480.62±4.272a	393.03±10.575b	12.53±0.085b
	W3	浅埋滴灌SBDI	6.9281±0.063a	476.37±4.234ab	446.17±12.005a	13.28±0.361a
2020	W1	膜下滴灌MDI	6.9300±0.064a	479.16±4.259b	375.46±10.103c	11.67±0.316b
	W1	浅埋滴灌SBDI	6.9280±0.064a	475.78±4.229b	356.46±9.591d	11.03±0.295c
	W2	膜下滴灌MDI	6.8410±0.063a	489.81±4.354a	395.15±10.632b	12.37±0.561b
	W2	浅埋滴灌SBDI	6.9120±0.063a	487.94±4.337a	383.88±10.329bc	12.12±0.342b
	W3	膜下滴灌MDI	6.9310±0.064a	493.56±4.387a	397.53±10.696b	12.85±0.510b
	W3	浅埋滴灌SBDI	6.9355±0.064a	490.86±4.363a	452.50±12.176a	14.02±0.351a

源 Source	千粒重 1000-grain weight		产量 Yield
源 Source	F	P	F	P
年份 Year (A)	1.285	0.268	4.877	0.037
灌水量 Irrigation volume (B)	91.730	0.000	49.252	0.000
滴灌模式 Drip irrigation mode (C)	5.770	0.024	0.026	0.874
年份×灌水量A×B	0.300	0.744	0.953	0.400
年份×滴灌模式A×C	0.000	0.992	0.229	0.637
灌水量×滴灌模式B×C	44.674	0.000	15.906	0.000
年份×灌水量×滴灌模式A×B×C	0.020	0.980	0.372	0.724

处理 Treatment		籽粒淀粉积累方程 Equation of starch accumulation in grain	活跃积累期 Active accumulation period (d)	最大积累速率 Maximum rate of accumulation (g·d^-1)	平均积累速率 Average rate of accumulation (g·d^-1)	达到最大速率的时间 Time to reach maximum velocity (d)	R²
W1	膜下滴灌MDI	y=0.243659/(1+129.895×e^-0.234907t)^0.657312	18.06	1.23608	0.633045	18.9315	0.995
W1	浅埋滴灌SBDI	y=0.214038/(1+126.942×e^-0.232139t)^0.793003	19.43	1.15071	0.539682	19.8665	0.994
W2	膜下滴灌MDI	y=0.246778/(1+190.999×e^-0.246529t)^0.600757	18.17	1.26721	0.651644	19.2379	0.994
W2	浅埋滴灌SBDI	y=0.250457/(1+64.3241×e^-0.206384t)^0.971553	19.99	1.28076	0.591956	20.0359	0.993
W3	膜下滴灌MDI	y=0.261923/(1+70.8877×e^-0.210563t)^0.751041	18.26	1.26044	0.647571	18.877	0.992
W3	浅埋滴灌SBDI	y=0.313065/(1+281.947×e^-0.244806t)^0.739513	21.25	1.76071	0.784787	21.813	0.998

变量 Variable	直接系数 Direct coefficient	X1	X2	X3	X4	X5	X6
X1	2.6396		2.6112	1.9411	-0.7934	-1.8436	2.3242
X2	2.6254	2.6396		-1.8512	0.8148	1.7085	-2.3939
X3	-2.7222	2.6396	2.6254		-0.2599	-1.8967	1.0063
X4	0.9173	2.6396	2.6254	-2.7222		-1.1674	2.5279
X5	-2.3799	2.6396	2.6254	-2.7222	0.9173		-1.3152
X6	2.5594	2.6396	2.6254	-2.7222	0.9173	-2.3799