基施氮肥及灌浆前期喷施锌肥对小麦籽粒富锌及蛋白组分含量的影响

doi:10.3864/j.issn.0578-1752.2022.10.008

摘要/Abstract

摘要：

【目的】叶面喷锌（Zn）是提高小麦籽粒锌含量进而解决人体缺锌问题的有效农艺措施。探明不同施氮（N）量下叶面喷锌后小麦全粒及面粉中的富锌效果及对蛋白组分含量的影响。【方法】基于长期定位试验,于2018—2020年连续进行了两年裂区田间试验。以基施不同用量氮肥（N₀、N₁₂₀、N₂₄₀,施N量分别为0、120、240 kg∙hm^-2）为主区,副区为灌浆前期喷施锌肥处理（Zn₀、Zn₁,分别为喷H₂O、喷0.4% ZnSO₄·7H₂O）,测定了灌浆前期和成熟期各部位锌含量、叶片等营养器官中锌向籽粒的转移量及分配、籽粒和面粉中蛋白质及其组分含量。【结果】与N₀相比,N₁₂₀和N₂₄₀处理籽粒产量显著提高,增幅达88%—114%,但N₁₂₀和N₂₄₀处理之间并无显著差异。叶面喷锌均能显著提高小麦籽粒和面粉锌含量且籽粒达富锌标准,而不受施氮量的影响,其中,N₁₂₀、N₂₄₀处理小麦籽粒锌含量分别比N₀处理提高0.95和1.12倍。与N₀相比,施用氮肥均提高了小麦灌浆前期叶片等营养器官中氮、锌向籽粒的转移量,但降低了二者的转移比例,其中氮转移比例由60.2%下降至48.6%,锌由55.4%下降至42.3%。无论喷锌与否,氮、锌向籽粒的转移量及成熟期籽粒中氮、锌含量均呈显著线性正相关,且喷锌时氮、锌协同效应更为显著。与灌浆前期相比,成熟期小麦籽粒和面粉中储藏蛋白（醇溶蛋白和谷蛋白）含量显著增加,约占蛋白含量的80%—84%。施氮对籽粒和面粉中醇溶蛋白和谷蛋白含量提升幅度高于清蛋白和球蛋白,且以谷蛋白最大,而喷锌不影响籽粒和面粉中蛋白质及其组分含量,但在Zn₁条件下,施氮对籽粒和面粉中谷蛋白含量的提高幅度高于Zn₀条件下,分别提升37.5%和38.1%。【结论】叶面喷锌能够实现籽粒富锌,但不影响籽粒和面粉中蛋白质及其组分含量,表明籽粒和面粉中存在足够的用于锌储存的蛋白质库。因此在潜在缺锌石灰性土壤上,通过合理施用氮肥结合小麦灌浆前期叶面喷锌,能在保证小麦高产稳产的同时提高籽粒氮、锌营养品质。

关键词: 小麦, 叶面喷锌, 基施氮肥, 锌含量, 蛋白组分

Abstract:

【Objective】 Foliar zinc (Zn) application is an effective agronomic biofortification strategy to realize Zn enrichment of wheat grains and to combat the human Zn malnutrition. The aim of this study was to investigate the effects of spraying Zn with different nitrogen (N) inputs on Zn enrichment and content of protein and protein components in wheat whole grain and flour.【Method】 Based on the long-term positioning experiment, the spraying experiment of Zn in wheat for two consecutive seasons was conducted during 2018-2020. A split plot design was used with soil N rates of 0 (N₀), 120 (N₁₂₀) and 240 ∙hm^-2 (N₂₄₀) as the main plot factor, and the foliar application of distilled water (Zn₀) and 0.4% ZnSO₄·7H₂O (Zn₁) as subplot. The indexes were analyzed for this study, including Zn content of various nutritional organs, Zn mobilization and distribution from leaf and other vegetative organs to grain, and protein and protein component content in grains and flour at the early filling stage and maturation stage. 【Result】Compared with N₀, the grain yield under N₁₂₀ and N₂₄₀treatments was significantly increased by 88%-114%, but there was no significant difference between N₁₂₀ and N₂₄₀. Regardless of the N inputs, the foliar Zn application could significantly increase the Zn concentration in grains and flour and the grain Zn content reached the biofortification standard. Among those treatments, the Zn concentration of wheat grains under N₁₂₀ and N₂₄₀ was increased by 0.95 and 1.12 times than that under N₀, respectively. Compared with N₀, the N inputs increased the translocation of N and Zn transferred from leaf and other vegetative organs to grain at early grain filling stage, but reduced the transfer ratio of N and Zn: N decreased from 60.2% to 48.6% and Zn decreased from 42.3% to 26.5%. A significant positive linear correlation was found between the amount of N and Zn mobilization and the content of N and Zn in grain at maturity, and the synergistic effect of N and Zn was more significant at Zn₁. Compared with the early filling stage, the content of storage protein (gliadin and glutenin) in grains and flour at the mature stage increased significantly, accounting for about 80%-84% of the protein content. The content of gliadin and glutenin in grain and flour was increased by N application more than that of albumin and globulin, and the gluten was the largest. The content of protein and protein components in grain and flour were not affected by spraying Zn. However, in terms of Zn₁, the increase of gluten content in grain and flour was higher than that under the condition of Zn₀with the increase of N dosage, which was 37.5% and 38.1%, respectively.【Conclusion】Foliar Zn application could achieve Zn-rich grains but did not affect the content of protein and protein components in grains and flour, indicating that there was sufficient protein pool for Zn storage in grains and flour. Therefore, a reasonable amount of soil N combined with foliar Zn application could increase the N, Zn content and nutritional quality of the grains by ensuring high and stable yield on potentially Zn-deficient calcareous soils.

Key words: wheat, foliar Zn application, basal nitrogen fertilization, Zn content, protein component

吴天琪,李雅菲,师江澜,宁鹏,田霄鸿. 基施氮肥及灌浆前期喷施锌肥对小麦籽粒富锌及蛋白组分含量的影响[J]. 中国农业科学, 2022, 55(10): 1971-1986.

WU TianQi,LI YaFei,SHI JiangLan,NING Peng,TIAN XiaoHong. Effects of Basal Nitrogen and Foliar Zinc Application at the Early Filling Stage on Zinc Enrichment and Protein Components Content in Wheat Grain[J]. Scientia Agricultura Sinica, 2022, 55(10): 1971-1986.

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.0578-1752.2022.10.008

https://www.chinaagrisci.com/CN/Y2022/V55/I10/1971

图/表 9

图1

图2

表1

图4

表2

图5

图6

图7

图3

参考文献 62

[1]	GIBSON R S. Zinc deficiency and human health: etiology, health consequences, and future solutions. Plant and Soil, 2012, 361(1/2): 291-299. doi: 10.1007/s11104-012-1209-4. doi: 10.1007/s11104-012-1209-4
[2]	WHO: Vitamin and mineral nutrition information system,Geneva. World Health Organization, 2016, http://www.who.int.
[3]	HOTZ C, BROWN K. Assessment of the risk of deficiency in populations and options for its control. Food and Nutrition Bulletin, 2004, 25: 194-204. doi: 10.1177/156482650402500213
[4]	STEIN A J. Global impacts of human mineral malnutrition. Plant and Soil, 2010, 335(1/2): 133-154. doi: 10.1007/s11104-009-0228-2. doi: 10.1007/s11104-009-0228-2
[5]	DOKOOHAKI H, GHEYSARI M, MEHNATKESH A, AYOUBI S. Applying the CSM-CERES-Wheat model for rainfed wheat with specified soil characteristic in undulating area in Iran. Archives of Agronomy and Soil Science, 2015, 61(9): 1231-1245. doi: 10.1080/03650340.2014.984696. doi: 10.1080/03650340.2014.984696
[6]	HUANG T M, HUANG Q N, SHE X, MA X L, HUANG M, CAO H B, HE G, LIU J S, LIANG D L, MALHI S S, WANG Z H. Grain zinc concentration and its relation to soil nutrient availability in different wheat cropping regions of China. Soil and Tillage Research, 2019, 191: 57-65. doi: 10.1016/j.still.2019.03.019. doi: 10.1016/j.still.2019.03.019
[7]	CAKMAK I, KUTMAN U B. Agronomic biofortification of cereals with zinc: a review. European Journal of Soil Science, 2018, 69(1): 172-180. doi: 10.1111/ejss.12437. doi: 10.1111/ejss.12437
[8]	ZOU C, DU Y, RASHID A, RAM H, SAVASLI E, PIETERSE P J, ORTIZ-MONASTERIO I, YAZICI A, KAUR C, MAHMOOD K, SINGH S, LE ROUX M R, KUANG W, ONDER O, KALAYCI M, CAKMAK I. Simultaneous biofortification of wheat with zinc, iodine, selenium, and iron through foliar treatment of a micronutrient cocktail in six countries. Nitric Oxide, 2019, 67(29): 8096-8106. doi: 10.1021/acs.jafc.9b01829. doi: 10.1021/acs.jafc.9b01829
[9]	GRAHAM R D, WELCH R M, BOUIS H E. Addressing micronutrient malnutrition through enhancing the nutritional quality of staple foods. Principles, perspectives and knowledge gaps. 2001, 70: 77-142. doi: 10.1016/S0065-2113(01)70004-1. doi: 10.1016/S0065-2113(01)70004-1
[10]	BOUIS H E, WELCH R M. Biofortification-A sustainable agricultural strategy for reducing micronutrient malnutrition in the global south. Crop Science, 2010, 50: S-20. doi: 10.2135/cropsci2009.09.0531. doi: 10.2135/cropsci2009.09.0531
[11]	CAKMAK I. Enrichment of cereal grains with zinc: Agronomic or genetic biofortification? Plant and Soil, 2008, 302(1/2): 1-17. doi: 10.1007/s11104-007-9466-3. doi: 10.1007/s11104-007-9466-3
[12]	ZOU C Q, ZHANG Y Q, RASHID A, RAM H, SAVASLI E, ARISOY R Z, ORTIZ-MONASTERIO I, SIMUNJI S, WANG Z H, SOHU V, HASSAN M, KAYA Y, ONDER O, LUNGU O, MUJAHID M Y, JOSHI A K, ZELENSKIY Y, ZHANG F S, CAKMAK I. Biofortification of wheat with zinc through zinc fertilization in seven countries. Plant and Soil, 2012, 361(1/2): 119-130. doi: 10.1007/s11104-012-1369-2. doi: 10.1007/s11104-012-1369-2
[13]	李宏云, 王少霞, 李萌, 田霄鸿, 赵爱青, 国春慧. 不同水氮管理下锌与氮磷肥配合喷施对冬小麦锌营养品质的影响. 中国农业科学, 2014, 47(20): 4016-4026. doi: 10.3864/j.issn.0578-1752.2014.20.010. doi: 10.3864/j.issn.0578-1752.2014.20.010
	LI H Y, WANG S X, LI M, TIAN X H, ZHAO A Q, GUO C H. Effects of combined foliar Zn application with N or P under different water and nitrogen managements on Zn nutritional quality of winter wheat. Scientia Agricultura Sinica, 2014, 47(20): 4016-4026. doi: 10.3864/j.issn.0578-1752.2014.20.010. (in Chinese) doi: 10.3864/j.issn.0578-1752.2014.20.010
[14]	王少霞, 李萌, 田霄鸿, 陈艳龙, 李硕, 刘珂, 贾舟. 锌与氮磷钾配合喷施对小麦锌累积、分配及转移的影响. 植物营养与肥料学报, 2018, 24(2): 296-305. doi: 10.11674/zwyf.17178. doi: 10.11674/zwyf.17178
	WANG S X, LI M, TIAN X H, CHEN Y L, LI S, LIU K, JIA Z. Effects of combined foliar application of Zn with N, P, or K on Zn accumulation, distribution and translocation in wheat. Plant Nutrition and Fertilizer Science, 2018, 24(2): 296-305. doi: 10.11674/zwyf.17178. (in Chinese) doi: 10.11674/zwyf.17178
[15]	AZIZ M Z, YASEEN M, ABBAS T, NAVEED M, MUSTAFA A, HAMID Y, SAEED Q, XU M G. Foliar application of micronutrients enhances crop stand, yield and the biofortification essential for human health of different wheat cultivars. Journal of Integrative Agriculture, 2019, 18(6): 1369-1378. doi: 10.1016/S2095-3119(18)62095-7. doi: 10.1016/S2095-3119(18)62095-7
[16]	ERENOGLU E B, KUTMAN U B, CEYLAN Y, YILDIZ B, CAKMAK I. Improved nitrogen nutrition enhances root uptake, root-to-shoot translocation and remobilization of zinc ((65) Zn) in wheat. The New Phytologist, 2011, 189(2): 438-448. doi: 10.1111/j.1469-8137.2010.03488.x. doi: 10.1111/j.1469-8137.2010.03488.x.
[17]	CHEN X P, ZHANG Y Q, TONG Y P, XUE Y F, LIU D Y, ZHANG W, DENG Y, MENG Q F, YUE S C, YAN P, CUI Z L, SHI X J, GUO S W, SUN Y X, YE Y L, WANG Z H, JIA L L, MA W Q, HE M R, ZHANG X Y, KOU C L, LI Y T, TAN D S, CAKMAK I, ZHANG F S, ZOU C Q. Harvesting more grain zinc of wheat for human health. Scientific Reports, 2017, 7: 7016. doi: 10.1038/s41598-017-07484-2. doi: 10.1038/s41598-017-07484-2
[18]	XIA H Y, XUE Y F, LIU D Y, KONG W L, XUE Y H, TANG Y Y, LI J, LI D, MEI P P. Rational application of fertilizer nitrogen to soil in combination with foliar Zn spraying improved Zn nutritional quality of wheat grains. Frontiers in Plant Science, 2018, 9: 677. doi: 10.3389/fpls.2018.00677. doi: 10.3389/fpls.2018.00677
[19]	PASCOALINO J A L, THOMPSON J A, WRIGHT G, FRANCO F A, SCHEEREN P L, PAULETTI V, MORAES M F, WHITE P J. Grain zinc concentrations differ among Brazilian wheat genotypes and respond to zinc and nitrogen supply. PLoS One, 2018, 13(7): e0199464. doi: 10.1371/journal.pone.0199464. doi: 10.1371/journal.pone.0199464
[20]	WATERS B M, UAUY C, DUBCOVSKY J, GRUSAK M A. Wheat (Triticum aestivum) NAM proteins regulate the translocation of iron, zinc, and nitrogen compounds from vegetative tissues to grain. Journal of Experimental Botany, 2009, 60(15): 4263-4274. doi: 10.1093/jxb/erp257. doi: 10.1093/jxb/erp257
[21]	DISTELFELD A, CAKMAK I, PELEG Z, OZTURK L, YAZICI A M, BUDAK H, SARANGA Y, FAHIMA T. Multiple QTL-effects of wheat Gpc-B1 locus on grain protein and micronutrient concentrations. Physiologia Plantarum, 2007, 129(3): 635-643. doi: 10.1111/j.1399-3054.2006.00841.x. doi: 10.1111/j.1399-3054.2006.00841.x.
[22]	CAKMAK I, KALAYCI M, KAYA Y, TORUN A A, AYDIN N, WANG Y, ARISOY Z, ERDEM H, YAZICI A, GOKMEN O, OZTURK L, HORST W J. Biofortification and localization of zinc in wheat grain. Journal of Agricultural and Food Chemistry, 2010, 58(16): 9092-9102. doi: 10.1021/jf101197h. doi: 10.1021/jf101197h
[23]	KUTMAN U B, KUTMAN B Y, CEYLAN Y, OVA E A, CAKMAK I. Contributions of root uptake and remobilization to grain zinc accumulation in wheat depending on post-anthesis zinc availability and nitrogen nutrition. Plant and Soil, 2012, 361(1/2): 177-187. doi: 10.1007/s11104-012-1300-x. doi: 10.1007/s11104-012-1300-x
[24]	VELU G, ORTIZ-MONASTERIO I, CAKMAK I, HAO Y, SINGH R P. Biofortification strategies to increase grain zinc and iron concentrations in wheat. Journal of Cereal Science, 2014, 59(3): 365-372. doi: 10.1016/j.jcs.2013.09.001. doi: 10.1016/j.jcs.2013.09.001
[25]	DIONISIO G, UDDIN M N, VINCZE E. Enrichment and identification of the most abundant zinc binding proteins in developing barley grains by zinc-IMAC capture and nano LC-MS/MS. Proteomes, 2018, 6(1): 3. doi: 10.3390/proteomes6010003. doi: 10.3390/proteomes6010003
[26]	PERSSON D P, DE BANG T C, PEDAS P R, KUTMAN U B, CAKMAK I, ANDERSEN B, FINNIE C, SCHJOERRING J K, HUSTED S. Molecular speciation and tissue compartmentation of zinc in durum wheat grains with contrasting nutritional status. The New Phytologist, 2016, 211(4): 1255-1265. doi: 10.1111/nph.13989. doi: 10.1111/nph.13989
[27]	PECK A W, MCDONALD G K, GRAHAM R D. Zinc nutrition influences the protein composition of flour in bread wheat (Triticum aestivum L.). Journal of Cereal Science, 2008, 47(2): 266-274. doi: 10.1016/j.jcs.2007.04.006. doi: 10.1016/j.jcs.2007.04.006
[28]	董明, 王琪, 周琴, 蔡剑, 王笑, 戴廷波, 姜东. 花后5天喷施锌肥有效提高小麦籽粒营养和加工品质. 植物营养与肥料学报, 2018, 24(1): 63-70. doi: 10.11674/zwyf.17106. doi: 10.11674/zwyf.17106
	DONG M, WANG Q, ZHOU Q, CAI J, WANG X, DAI T B, JIANG D. Efficient promotion of the nutritional and processing quality of wheat grain by Zn forliar spraying at 5 days after anthesis. Plant Nutrition and Fertilizer Science, 2018, 24(1): 63-70. doi: 10.11674/zwyf.17106. (in Chinese) doi: 10.11674/zwyf.17106
[29]	HE L, QY W, RENGEL Z, ZHAO P. Zinc fertilization alters flour protein composition of winter wheat genotypes varying in gluten content. Plant, Soil and Environment, 2016, 61(No.5): 195-200. doi: 10.17221/817/2014-pse. doi: 10.17221/817/2014-pse
[30]	BROADLEY M R, WHITE P J, HAMMOND J P, ZELKO I, LUX A. Zinc in plants. New Phytologist, 2007, 173: 677-702. doi: 10.1111/j.1469-8137.2007.01996. doi: 10.1111/j.1469-8137.2007.01996
[31]	DOOLETTE C L, READ T L, LI C, SCHECKEL K G, DONNER E, KOPITTKE P M, SCHJOERRING J K, LOMBI E. Foliar application of zinc sulphate and zinc EDTA to wheat leaves: differences in mobility, distribution, and speciation. Journal of Experimental Botany, 2018, 69(18): 4469-4481. doi: 10.1093/jxb/ery236. doi: 10.1093/jxb/ery236
[32]	HAUG W, LANTZSCH H J. Sensitive method for the rapid determination of phytate in cereals and cereal products. Journal of the Science of Food and Agriculture, 1983, 34(12): 1423-1426. doi: 10.1002/jsfa.2740341217. doi: 10.1002/jsfa.2740341217
[33]	WANG Z M, LIU Q, PAN F, YUAN L X, YIN X B. Effects of increasing rates of zinc fertilization on phytic acid and phytic acid/zinc molar ratio in zinc bio-fortified wheat. Field Crops Research, 2015, 184: 58-64. doi: 10.1016/j.fcr.2015.09.007. doi: 10.1016/j.fcr.2015.09.007
[34]	鲍士旦. 土壤农化分析. 3版. 北京: 中国农业出版社, 2000.
	BAO S D . Soil and Agricultural Chemistry Analysis. Beijing: Chinese Agriculture Press, 2000. (in Chinese)
[35]	MILLER L V, KREBS N F, HAMBIDGE K M. A mathematical model of zinc absorption in humans as a function of dietary zinc and phytate. The Journal of Nutrition, 2007, 137(1): 135-141. doi: 10.1093/jn/137.1.135. doi: 10.1093/jn/137.1.135
[36]	赖学华, 丁建国, 赵晶, 郭蔼光. 不同栽培模式下小麦灌浆期蛋白质周转研究. 西北植物学报, 2005, 25(8): 1574-1578. doi: 10.3321/j.issn:1000-4025.2005.08.014. doi: 10.3321/j.issn:1000-4025.2005.08.014
	LAI X H, DING J G, ZHAO J, GUO A G. Protein turnover of wheat in different planting modes at the kernel-filling stage. Acta Botanica Boreali-Occidentalia Sinica, 2005, 25(8): 1574-1578. doi: 10.3321/j.issn:1000-4025.2005.08.014. (in Chinese) doi: 10.3321/j.issn:1000-4025.2005.08.014
[37]	熊淑萍, 王小纯, 李春明, 马新明, 杜少勇, 张营武, 蔺世召. 冬小麦根系时空分布动态及产量对不同氮源配施的响应. 植物生态学报, 2011, 35(7): 759-768. doi: 10.3724/SP.J.1258.2011.00759. doi: 10.3724/SP.J.1258.2011.00759
	XIONG S P, WANG X C, LI C M, MA X M, DU S Y, ZHANG Y W, LIN S Z. Responses of the spatial-temporal distribution of winter wheat (Triticum aestivum) roots and yield to different ratios of nitrogen sources. Chinese Journal of Plant Ecology, 2011, 35(7): 759-768. doi: 10.3724/SP.J.1258.2011.00759. (in Chinese) doi: 10.3724/SP.J.1258.2011.00759
[38]	KUTMAN U B, YILDIZ B, OZTURK L, CAKMAK I. Biofortification of durum wheat with zinc through soil and foliar applications of nitrogen. Cereal Chemistry Journal, 2010, 87(1): 1-9. doi: 10.1094/cchem-87-1-0001. doi: 10.1094/cchem-87-1-0001
[39]	ANDRESEN E, PEITER E, KÜPPER H. Trace metal metabolism in plants. Journal of Experimental Botany, 2018, 69(5): 909-954. doi: 10.1093/jxb/erx465. doi: 10.1093/jxb/erx465
[40]	XUE Y F, YUE S C, ZHANG Y Q, CUI Z L, CHEN X P, YANG F C, CAKMAK I, MCGRATH S P, ZHANG F S, ZOU C Q. Grain and shoot zinc accumulation in winter wheat affected by nitrogen management. Plant and Soil, 2012, 361(1/2): 153-163. doi: 10.1007/s11104-012-1510-2. doi: 10.1007/s11104-012-1510-2
[41]	杨习文, 宋淼, 李秋杰, 周苏玫, 韩少宇, 陈旭, 徐利利, 贺德先. 氮锌配施对小麦锌转运、分配与累积的影响. 应用生态学报, 2020, 31(1): 148-156. doi: 10.13287/j.1001-9332.202001.027. doi: 10.13287/j.1001-9332.202001.027
	YANG X W, SONG M, LI Q J, ZHOU S M, HAN S Y, CHEN X, XU L L, HE D X. Impacts of combined N and Zn application on Zn translocation, partitioning, and accumulation in Triticum aestivum. Chinese Journal of Applied Ecology, 2020, 31(1): 148-156. doi: 10.13287/j.1001-9332.202001.027. (in Chinese) doi: 10.13287/j.1001-9332.202001.027
[42]	PEARSON J N, JENNER C F, RENGEL Z, GRAHAM R D. Differential transport of Zn, Me and sucrose along the longitudinal axis of developing wheat grains. Physiologia Plantarum, 1996, 97(2): 332-338. doi: 10.1034/j.1399-3054.1996.970217.x. doi: 10.1034/j.1399-3054.1996.970217.x.
[43]	陈娟, 王少霞, 田霄鸿, 陈艳龙, 朱文玲, 李秀双, 刘珂, 杨畅. 锌与农药配合喷施对小麦锌累积分配及转移的影响. 西北农林科技大学学报(自然科学版), 2019, 47(3): 67-76. doi: 10.13207/j.cnki.jnwafu.2019.03.010. doi: 10.13207/j.cnki.jnwafu.2019.03.010
	CHEN J, WANG S X, TIAN X H, CHEN Y L, ZHU W L, LI X S, LIU K, YANG C. Effect of combined foliar application of zinc and pesticides on accumulation, distribution and transfer of zinc in wheat. Journal of Northwest A & F University (Natural Science Edition), 2019, 47(3): 67-76. doi: 10.13207/j.cnki.jnwafu.2019.03.010. (in Chinese) doi: 10.13207/j.cnki.jnwafu.2019.03.010
[44]	WANG X Z, LIU D Y, ZHANG W, WANG C J, CAKMAK I, ZOU C Q. An effective strategy to improve grain zinc concentration of winter wheat, Aphids prevention and farmers’ income. Field Crops Research, 2015, 184: 74-79. doi: 10.1016/j.fcr.2015.08.015. doi: 10.1016/j.fcr.2015.08.015
[45]	WANG S X, SUN N H, YANG S, TIAN X H, LIU Q. The effectiveness of foliar applications of different zinc source and urea to increase grain zinc of wheat grown under reduced soil nitrogen supply. Journal of Plant Nutrition, 2021, 44(5): 644-659. doi: 10.1080/01904167.2020.1849286. doi: 10.1080/01904167.2020.1849286
[46]	LI M, WANG S X, TIAN X H, ZHAO J H, LI H Y, GUO C H, CHEN Y L, ZHAO A Q. Zn distribution and bioavailability in whole grain and grain fractions of winter wheat as affected by applications of soil N and foliar Zn combined with N or P. Journal of Cereal Science, 2015, 61: 26-32. doi: 10.1016/j.jcs.2014.09.009. doi: 10.1016/j.jcs.2014.09.009
[47]	GONZALEZ D, ALMENDROS P, OBRADOR A, ALVAREZ J M. Zinc application in conjunction with urea as a fertilization strategy for improving both nitrogen use efficiency and the zinc biofortification of barley. Journal of the Science of Food and Agriculture, 2019, 99(9): 4445-4451. doi: 10.1002/jsfa.9681. doi: 10.1002/jsfa.9681
[48]	LI M, WANG S X, TIAN X H, HUANG Y P. Improving nutritional quality of wheat grain through foliar zinc combined with macronutrients. Agronomy Journal, 2018, 110(1): 38-46. doi: 10.2134/agronj2017.08.0437. doi: 10.2134/agronj2017.08.0437
[49]	DOOLETTE C L, READ T L, HOWELL N R, CRESSWELL T, LOMBI E. Zinc from foliar-applied nanoparticle fertiliser is translocated to wheat grain: a 65Zn radiolabelled translocation study comparing conventional and novel foliar fertilisers. Science of the Total Environment, 2020, 749: 142369. doi: 10.1016/j.scitotenv.2020.142369. doi: 10.1016/j.scitotenv.2020.142369
[50]	WIESER H, SEILMEIER W. The influence of nitrogen fertilisation on quantities and proportions of different protein types in wheat flour. Journal of the Science of Food and Agriculture, 1998, 76(1): 49-55. doi: 10.1002/(sici)1097-0010(199801)76:1<49:aid-jsfa950>3.0.co;2-2. doi: 10.1002/(sici)1097-0010(199801)76:1<49:aid-jsfa950>3.0.co;2-2
[51]	GOESAERT H, BRIJS K, VERAVERBEKE W S, COURTIN C M, GEBRUERS K, DELCOUR J A. Wheat flour constituents: how they impact bread quality, and how to impact their functionality. Trends in Food Science & Technology, 2005, 16(1/2/3): 12-30. doi: 10.1016/j.tifs.2004.02.011. doi: 10.1016/j.tifs.2004.02.011
[52]	赵鹏, 杨帆, 睢福庆, 王巧燕. 氮锌配施对冬小麦氮利用、产量及籽粒蛋白质含量的影响. 中国农业大学学报, 2013, 18(3): 28-33.
	ZHAO P, YANG F, SUI F Q, WANG Q Y. Effect of combined application of Zn and N fertilizers on nitrogen use, grain yield and protein content in winter wheat. Journal of China Agricultural University, 2013, 18(3): 28-33. (in Chinese)
[53]	TRIBOI E. Environmentally-induced changes in protein composition in developing grains of wheat are related to changes in total protein content. Journal of Experimental Botany, 2003, 54(388): 1731-1742. doi: 10.1093/jxb/erg183. doi: 10.1093/jxb/erg183
[54]	DIER M, HÜTHER L, SCHULZE W X, ERBS M, KÖHLER P, WEIGEL H J, MANDERSCHEID R, ZÖRB C. Elevated atmospheric CO₂ concentration has limited effect on wheat grain quality regardless of nitrogen supply. Journal of Agricultural and Food Chemistry, 2020, 68(12): 3711-3721. doi: 10.1021/acs.jafc.9b07817. doi: 10.1021/acs.jafc.9b07817
[55]	ROSSMANN A, BUCHNER P, SAVILL G P, POWERS S J, HAWKESFORD M J, MÜHLING K H. Foliar N application at anthesis stimulates gene expression of grain protein fractions and alters protein body distribution in winter wheat (Triticum aestivum L.). Journal of Agricultural and Food Chemistry, 2019, 67(46): 12709-12719. doi: 10.1021/acs.jafc.9b04634. doi: 10.1021/acs.jafc.9b04634
[56]	石玉, 张永丽, 于振文. 施氮量对不同品质类型小麦子粒蛋白质组分含量及加工品质的影响. 植物营养与肥料学报, 2010, 16(1): 33-40.
	SHI Y, ZHANG Y L, YU Z W. Effects of nitrogen fertilization on protein components contents and processing quality of different wheat genotypes. Plant Nutrition and Fertilizer Science, 2010, 16(1): 33-40. (in Chinese)
[57]	MARTRE P, PORTER J R, JAMIESON P D, TRIBOÏ E. Modeling grain nitrogen accumulation and protein composition to understand the sink/source regulations of nitrogen remobilization for wheat. Plant Physiology, 2003, 133(4): 1959-1967. doi: 10.1104/pp.103.030585. doi: 10.1104/pp.103.030585
[58]	GIULIANI M M, GIUZIO L, DE CARO A, FLAGELLA Z. Relationships between nitrogen utilization and grain technological quality in durum wheat: I. nitrogen translocation and nitrogen use efficiency for protein. Agronomy Journal, 2011, 103(5): 1487-1494. doi: 10.2134/agronj2011.0153. doi: 10.2134/agronj2011.0153
[59]	SHEWRY P R, MITCHELL R A C, TOSI P, WAN Y F, UNDERWOOD C, LOVEGROVE A, FREEMAN J, TOOLE G A, MILLS E N C, WARD J L. An integrated study of grain development of wheat (cv. Hereward). Journal of Cereal Science, 2012, 56(1): 21-30. doi: 10.1016/j.jcs.2011.11.007. doi: 10.1016/j.jcs.2011.11.007
[60]	GUPTA R B, MASCI S, LAFIANDRA D, BARIANA H S, MACRITCHIE F. Accumulation of protein subunits and their polymers in developing grains of hexaploid wheats. Journal of Experimental Botany, 1996, 47(9): 1377-1385. doi: 10.1093/jxb/47.9.1377. doi: 10.1093/jxb/47.9.1377
[61]	TAO Z Q, WANG D M, CHANG X H, WANG Y J, YANG Y S, ZHAO G C. Effects of zinc fertilizer and short-term high temperature stress on wheat grain production and wheat flour proteins. Journal of Integrative Agriculture, 2018, 17(9): 1979-1990. doi: 10.1016/S2095-3119(18)61911-2. doi: 10.1016/S2095-3119(18)61911-2
[62]	WANG Y X, SPECHT A, HORST W J. Stable isotope labelling and zinc distribution in grains studied by laser ablation ICP-MS in an ear culture system reveals zinc transport barriers during grain filling in wheat. The New Phytologist, 2011, 189(2): 428-437. doi: 10.1111/j.1469-8137.2010.03489.x. doi: 10.1111/j.1469-8137.2010.03489.x.

处理 Treatment	2018—2019		2019—2020
处理 Treatment	氮转移量 N mobilization (mg/plant)	氮转移比例 N transfer ratio (%)	氮转移量 N mobilization (mg/plant)	氮转移比例 N transfer ratio (%)
N₀Zn₀	4.5±0.5 b	53.6	3.7±0.3 c	58.4
N₁₂₀Zn₀	13.4±2.3 a	39.4	6.4±0.3 b	47.2
N₂₄₀Zn₀	14.7±1.0 a	41.9	9.9±0.9 a	51.0
N₀Zn₁	4.6±0.8 b	44.3	3.3±0.2c	62.6
N₁₂₀Zn₁	11.9±0.7 a	42.5	6.5±0.5 b	57.5
N₂₄₀Zn₁	10.0±0.8 a	14.7	9.3±1.4 a	55.3
变异来源
N	<0.001		<0.001
Zn	0.104		0.601
N×Zn	0.281		0.882

处理 Treatment	2018—2019		2019—2020
处理 Treatment	锌转移量 Zn mobilization (×10^-3 mg/plant)	锌转移比例 Zn transfer ratio (%)	锌转移量 Zn mobilization (×10^-3 mg/plant)	锌转移比例 Zn transfer ratio (%)
N₀Zn₀	3.5±0.1 c	53.6	1.7±0.3 c	41.2
N₁₂₀Zn₀	5.9±0.1 bc	42.9	4.2±0.3 abc	35.1
N₂₄₀Zn₀	4.6±0.4 bc	34.3	3.2±1.3 bc	38.4
N₀Zn₁	3.4±0.4 c	48.9	2.1±0.5 c	39.2
N₁₂₀Zn₁	15.3±0.5 a	27.7	10.1±0.7 a	23.7
N₂₄₀Zn₁	12.8±0.3 ab	18.6	11.3±0.8 ab	14.6
变异来源
N	0.042		0.014
Zn	0.017		0.024
N×Zn	0.226		0.245