富锌水稻研究综述：从遗传育种到营养强化的多维度探索

doi:10.11924/j.issn.1000-6850.casb2025-0428

摘要/Abstract

摘要：

锌是人体必需的微量元素，全球约20亿人受锌缺乏导致的隐性饥饿困扰，水稻作为主粮的锌营养强化是解决该问题的高效途径。本文系统综述了富锌水稻在遗传育种、生理机制与栽培技术等领域的研究进展。归纳了分子标记辅助选择、CRISPR/Cas9基因编辑、航天诱变等育种技术在高锌种质创制中的应用；分析了锌吸收转运蛋白（如OsZIP、OsHMA2）功能及尼克酰胺(NA)介导的韧皮部锌运输机制；总结了锌肥基施结合叶面喷施、磷锌配施、稻渔种养等栽培措施对水稻籽粒锌富集的调控效果。指出OsCKX4过表达株系糙米锌含量达58 mg/kg，OsNAS过表达水稻株系NA合成量提升2倍，锌分配效率显著提高；分析表明，栽培技术协同应用可使水稻籽粒锌含量提升至42 mg/kg，单产突破12750 kg/hm²，实现“高产-高锌”协同提升。研究提出未来应整合多组学技术与智慧农业手段，推动富锌水稻产业化应用，为全球锌营养改善提供解决方案。

关键词: 富锌水稻, 分子育种, 生理机制, 栽培技术, 锌转运蛋白, 生物强化, 营养健康

Abstract:

Zinc is an essential trace element for us. About 2 billion people in the world are suffering from hidden hunger caused by zinc deficiency. Zinc nutrition enhancement of rice is an efficient way to solve this problem. This review systematically synthesized advances in zinc-enriched rice research across genetic breeding, physiological mechanisms, and cultivation techniques. It summarized the application of molecular marker-assisted selection, CRISPR/Cas9 gene editing, and space mutagenesis in developing high-zinc varieties; analyzed the functions of zinc transport proteins (e.g., OsZIP, OsHMA2) and nicotianamine (NA)-mediated zinc phloem transport; and reviewed the effects of cultivation practices such as basal and foliar zinc fertilization, phosphorus-zinc co-application, and rice-fish co-culture on grain zinc enrichment. The review pointed out that OsCKX4 overexpression lines achieved 58 mg/kg zinc in brown rice, while OsNAS overexpression doubled NA synthesis and significantly improved zinc allocation efficiency. It suggested that integrated cultivation techniques elevated grain zinc content to 42 mg/kg, with a yield of 12750 kg/hm², achieving a synergistic ‘high yield-high zinc’ outcome. The review proposed that future efforts should integrate multi-omics and smart agriculture technologies to promote the industrialization of zinc-enriched rice and provide solutions for global zinc nutrition improvement.

Key words: zinc-enriched rice, molecular breeding, physiological mechanism, cultivation technique, zinc transporter, biofortification, nutritional health

吕展轩, 刘冠明. 富锌水稻研究综述：从遗传育种到营养强化的多维度探索[J]. 中国农学通报, 2025, 41(33): 1-5.

LYU Zhanxuan, LIU Guanming. A Review on Zinc-enriched Rice Research: A Multidimensional Exploration from Genetic Breeding to Nutritional Enhancement[J]. Chinese Agricultural Science Bulletin, 2025, 41(33): 1-5.

参考文献 52

[1]	SANDSTEAD H H. Zinc deficiency: a public health problem[J]. American journal of diseases of children, 1991, 40(145):853-859.
[2]	郝元峰, 张勇, 何中虎. 作物锌生物强化研究进展[J]. 生命科学, 2015, 27(8):1047-1054.
[3]	ZHANG G M, ZHENG T Q, CHEN Z, et al. Joint exploration of favorable haplotypes for mineral concentrations in milled grains of rice (Oryza sativa L.)[J]. Frontiers in plant science, 2018(9):447.
[4]	刘铮. 我国土壤中锌含量的分布规律[J]. 中国农业科学, 1994, 27(1):30-37.
[5]	JOHNSON H, WILLIAMS M. Food fortification: a review of its role in preventing micronutrient deficiencies[J]. Nutrition reviews, 2012, 70(10):573-583.
[6]	ZHAO F J, SHEWRY P R. Recent developments in modifying crops and agronomic practice to improve human health[J]. Food policy,2011,36:S94-S101.
[7]	DESCALSOTA G I L, SWAMY B P M, ZAW H, et al. Genome-wide association mapping in a rice MAGIC plus population detects QTLs and genes useful for biofortification[J]. Frontiers in plant science, 2018,9:1347.
[8]	CHEN J, MIAO Z, KONG D, et al. Application of CRISPR/Cas9 technology in rice germplasm innovation and genetic improvement[J]. Genes, 2024, 15(11):1492. doi: 10.3390/genes15111492 URL
[9]	冯绪猛, 郭九信, 王玉雯, 等. 锌肥品种与施用方法对水稻产量和锌含量的影响[J]. 植物营养与肥料学报, 2016, 22(5):1329-1338.
[10]	HUANG S, WANG P, YAMAJI N, et al. Plant nutrition for human nutrition: hints from rice research and future perspectives[J]. Molecular plant, 2020, 13(6):825-835. doi: S1674-2052(20)30142-8 pmid: 32434072
[11]	沈芸, 肖鹏, 包劲松. 水稻营养成分遗传育种研究进展[J]. 核农学报, 2008, 22(4):455-460.
[12]	SUMAN K, NEERAJA C N, MADHUBABU P, et al. Identification of promising RILs for high grain zinc through genotype × environment analysis and stable grain zinc QTL using SSRs and SNPs in rice (Oryza sativa L.)[J]. Frontiers in plant science, 2021,12:587482.
[13]	刘琦, 王张民, 潘斐, 等. 大田条件下水稻锌营养强化方法探究及效果评估[J]. 土壤, 2019, 51(1):32-38.
[14]	王俊, 郭颖, 吴蕊, 等. 不同种植年限和施肥量对日光温室土壤锌累积的影响[J]. 农业环境科学学报, 2009, 28(1):89-94.
[15]	杨丹丹, 吕莹, 董阳均, 等. 水稻云恢1973恢复基因关联分子标记筛选及应用[J]. 西南大学学报(自然科学版), 2024,46(9):55-64.
[16]	HALL J L, WILLIAMS L E. Transition metal transporters in plants[J]. Journal of experimental botany, 2003,54:2601-2613.
[17]	SHAO Z, DING C, LIU H, et al. A zinc metalloproteinase controls rice grain zinc content and weight[J]. Plant communications, 2025, 10(2):101295.
[18]	GAO S, XIAO Y, XU F, et al. Cytokinin-dependent regulatory module underlies the maintenance of zinc nutrition in rice[J]. New phytologist, 2019, 224(1):202-215. doi: 10.1111/nph.15962 pmid: 31131881
[19]	YANG M, LI Y, LIU Z, et al. A high activity zinc transporter OsZIP9 mediates zinc uptake in rice[J]. Plant journal, 2020, 103(5):1695-1709. doi: 10.1111/tpj.v103.5 URL
[20]	李雨彤. 水稻锌铁转运蛋白家族基因OsZIP9的功能研究[D]. 武汉: 华中农业大学, 2021.
[21]	TAN L, QU M, ZHU Y, et al. ZINC TRANSPORTER5 and ZINC TRANSPORTER9 function synergistically in zinc/cadmium uptake[J]. Plant physiology, 2020, 183(3):1235-1249. doi: 10.1104/pp.19.01569 pmid: 32341004
[22]	张琳琳, 韩娟英, 刘振, 等. 迷你型高锌含量水稻的选育及其特征特性[J]. 中国稻米, 2011, 17(6):66-68. doi: 10.3969/j.issn.1006-8082.2011.06.020
[23]	CAKMAK I. Enrichment of cereal grains with zinc: agronomic or genetic biofortification?[J]. Plant and soil, 2008, 302(1/2):1-17. doi: 10.1007/s11104-007-9466-3 URL
[24]	LILAY G H, HUMBERTO C P, GUEDES J G, et al. Rice F-bZIP transcription factors regulate the zinc deficiency response[J]. Journal of experimental botany, 2020, 71(12):3664-3677. doi: 10.1093/jxb/eraa115 pmid: 32133499
[25]	陈志强, 郭涛, 刘永柱, 等. 水稻航天育种研究进展与展望[J]. 华南农业大学学报, 2009, 30(1):1-5.
[26]	LEE S, JEONG H J, KIM S A, et al. OsZIP5 is a plasma membrane zinc transporter in rice[J]. Plant molecular biology, 2010, 73(4-5):507-517. doi: 10.1007/s11103-010-9637-0 pmid: 20419467
[27]	REBECCA F, GERARD C, PAUL J, et al. Functional expression of AtHMA4, a P1B-type ATPase of the Zn/Co/Cd/Pb subclass[J]. The plant journal, 2003, 35(2):164-176. doi: 10.1046/j.1365-313X.2003.01790.x URL
[28]	万吉丽. 锌在水稻体内运输、分配及积累的生理机制及基因型差异[D]. 杭州: 浙江大学, 2010.
[29]	徐勤松, 施国新, 杜开和, 等. Zn诱导的菹草叶抗氧化酶活性变化和超微结构损伤[J]. 植物研究, 2001, 21(4):569-573.
[30]	STEPHAN U W, SCHMIDKE I, PICH A. Phloem translocation of Fe, Cu, Mn, and Zn in Ricinus seedlings in relation to the concentrations of nicotianamine, an endogenous chelator of divalent metal ions, in different seedling parts[J]. Plant and soil, 1994, 165(2):181-188. doi: 10.1007/BF00008060 URL
[31]	ZHAN J H, ZOU W L, LI S Y, et al. OsNAC15 regulates tolerance to zinc deficiency and cadmium by binding to OsZIP7 and OsZIP10 in rice[J]. International journal of molecular sciences, 2022, 23(19):11771. doi: 10.3390/ijms231911771 URL
[32]	CHEN W R, FENG Y, CHAO Y. Genomic analysis and expression pattern of OsZIP1, OsZIP3, and OsZIP4 in two rice (Oryza sativa L.) genotypes with different zinc efficiency[J]. Russian journal of plant physiology, 2008, 55(3):400-409. doi: 10.1134/S1021443708030175 URL
[33]	SPEROTTO R A, BOFF T, DUARTE G L, et al. Identification of putative target genes to manipulate Fe and Zn concentrations in rice grains[J]. Journal of plant physiology, 2010, 167(17):1500-1506. doi: 10.1016/j.jplph.2010.05.003 pmid: 20580124
[34]	BARRETO M S C, GOMES F P, DE CARVALHO H W P, et al. Desorption kinetic and sequential extraction of Pb and Zn in a contaminated soil amended with phosphate, lime, biochar, and biosolids[J]. Environmental science and pollution research, 2023, 30(57):120793-120804. doi: 10.1007/s11356-023-30643-0
[35]	WISAWAPIPAT W, CHRISTL I, BOUCHET S, et al. Temporal development of arsenic speciation and extractability in acidified and non-acidified paddy soil amended with silicon-rich fly ash and manganese- or zinc-oxides under flooded and drainage conditions[J]. Chemosphere, 2024,351:141140.
[36]	李沿霖, 陈娟, 楚丽丽, 等. 解锌微生物的分离、筛选及在低锌石灰性土壤的应用[J]. 应用生态学报, 2024, 35(10):2765-2774. doi: 10.13287/j.1001-9332.202410.010
[37]	王孝忠, 田娣, 邹春琴. 锌肥不同施用方式及施用量对我国主要粮食作物增产效果的影响[J]. 植物营养与肥料学报, 2014, 20(4):998-1004.
[38]	刘露. 不同水稻品种中磷锌互作效应及机制的差异研究[D]. 武汉: 华中农业大学, 2020.
[39]	GARCÍA-GÓMEZ C, OBRADOR A, GONZÁLEZ D, et al. Comparative effect of ZnO NPs, ZnO bulk and ZnSO₄ in the antioxidant defences of two plant species growing in two agricultural soils under greenhouse conditions[J]. Science of the total environment, 2017,589:11-24.
[40]	周琼. 稻渔复合系统水土环境营养元素运移研究[D]. 银川: 宁夏大学, 2022.
[41]	HOU Y, JIA R, ZHOU L, et al. Integrated rice-fish farming dynamically altered the metal resistances and microbial-mediated iron, arsenic, and mercury biotransformation in paddy soil[J]. Environmental pollution, 2025,373:126107.
[42]	HASLETT B S, REID R J, RENGEL Z. Zinc mobility in wheat: uptake and distribution of zinc applied to leaves or roots[J]. Annals of botany, 2001, 87(3):379-385. doi: 10.1006/anbo.2000.1349 URL
[43]	周梦, 高尚, 王雪艳, 等. 水稻分蘖特性、产量及其构成因素对基施锌肥的响应[J]. 农业科技通讯, 2022(6):56-61.
[44]	王记安, 谢春甫, 刘长兵, 等. 机械插秧密度对水稻广两优5号生长及产量的影响[J]. 湖北农业科学, 2014, 53(14):3236-3238.
[45]	王芝. 乡村振兴背景下孝昌县乡村旅游可持续发展研究[D]. 乌鲁木齐: 新疆农业大学, 2022.
[46]	GIBSON R S. Zinc nutrition in developing countries[J]. Nutrition research reviews, 1994, 7(5):151-173. doi: 10.1079/NRR19940010 URL
[47]	WELCH R M, GRAHAM R D. Breeding for micronutrients in staple food crops from a human nutrition perspective[J]. Journal of experimental botany, 2004, 55(12):353-364. doi: 10.1093/jxb/erh064 URL
[48]	浙江大学. 负调控PICKs基因在降低植物中植酸含量中的应用[P]. 中国专利,202311046950.3,2023-12-01.
[49]	周钦彩, 朱明, 胡正文. 不同土壤类型水稻测土配方施肥对肥料利用率的影响分析[J]. 南方农业, 2018, 12(36):44-45.
[50]	张标金, 罗林广, 魏益华, 等. 锌镉互作对水稻幼苗镉和矿物质积累的影响[J]. 广东农业科学, 2015, 42(5):1-6.
[51]	赵波. 籽粒富锌铁差异水稻品种锌铁吸收利用特性研究[D]. 合肥: 安徽农业大学, 2023.
[52]	黄雷, 朱子龙, 程茂梓, 等. 孝昌太子米产业的发展概况与未来展望[J]. 中国种业, 2024(12):26-31.