水稻对镉吸收-迁移-分配的多层次互作网络及基因调控研究进展

doi:10.11924/j.issn.1000-6850.casb2025-0675

中国农学通报 ›› 2026, Vol. 42 ›› Issue (15): 65-76.doi: 10.11924/j.issn.1000-6850.casb2025-0675

水稻对镉吸收-迁移-分配的多层次互作网络及基因调控研究进展

郑兴鑫¹(), 李祖然², 段雨默¹, 陈玲³, 曹力³, 祖艳群¹()

¹ 云南农业大学资源与环境学院，昆明 650201
² 云南农业大学园林园艺学院，昆明 650201
³ 云南农业大学动物科学学院，昆明 650201

收稿日期:2025-08-11 修回日期:2025-12-22 出版日期:2026-08-13 发布日期:2026-08-13
通讯作者:
祖艳群，女，1966年出生，湖南沅江人，教授，博士生导师，博士，研究方向：土壤污染评价及其生态修复、重金属污染土壤的农业安全利用。通信地址：650201 云南昆明盘龙区沣源路，云南农业大学，Tel：0871-65227942，E-mail：649332092@qq.com。
作者简介:
郑兴鑫，女，2000年出生，云南云县人，硕士研究生，研究方向：重金属、农药污染过程及机理。通信地址：650201 云南昆明盘龙区沣源路，云南农业大学，Tel：86-17787267406，E-mail：XingxinZ@foxmail.com。
基金资助:
国家自然科学基金“云南野生稻镉吸收累积的根部响应特征及其机理”(42267062)

Research Advances on Multi-level Interaction Networks and Gene Regulation of Cadmium Absorption, Translocation, and Distribution in Rice (Oryza sativa L.)

ZHENG Xingxin¹(), LI Zuran², DUAN Yumo¹, CHEN Ling³, CAO Li³, ZU Yanqun¹()

¹ College of Resources and Environment, Yunnan Agricultural University, Kunming 650201
² College of Landscape and Horticulture, Yunnan Agricultural University, Kunming 650201
³ College of Animal Science, Yunnan Agricultural University, Kunming 650201

Received:2025-08-11 Revised:2025-12-22 Published:2026-08-13 Online:2026-08-13

摘要/Abstract

摘要：

为探究水稻对镉吸收-迁移-分配的多层次互作网络及基因调控机制，以期为培育低镉累积水稻品种奠定理论基础。本文系统梳理了水稻根系吸收、木质部装载、韧皮部再分配及籽粒累积的镉迁移过程，并整合了低累积相关的遗传调控网络与分子育种技术，并重点解析了转运蛋白、转录因子及非编码RNA在重金属的吸收、迁移、分配及籽粒积累中的作用，同时，探讨了基因过表达与基因编辑技术在降低水稻籽粒镉累积方面的应用潜力。目前对水稻镉吸收-迁移-分配的多层次互作网络及基因调控研究中，仍面临诸多挑战，包括互作网路复杂与调控层级不清、基因-环境互作与田间应用困难、多种重金属协同胁迫对于复杂田间环境下的分子机制尚待阐明。建议可针对关键转录因子或非编码RNA的调控元件进行精准编辑或设计，融合时空特异性基因编辑、合成生物学、多组学整合分析等前沿技术，深入解析并精准调控镉吸收-转运-解毒网络，为定向培育出兼具低镉累积特性与高产稳产优势的新种质提供依据。

关键词: 水稻, 镉, 低累积, 响应机制, 遗传调控

Abstract:

The multi-level interactive networks and genetic regulatory mechanisms of cadmium absorption, translocation, and distribution in rice were discussed in order to provide a theoretical basis for cultivating low-cadmium-accumulating rice. The processes of root uptake, xylem loading, phloem redistribution, and grain accumulation of cadmium in rice were summarized, as well as the genetic regulatory networks for low accumulation and molecular breeding technologies. And the roles of transport proteins, transcription factors, and non-coding RNAs in the absorption, translocation, distribution, and grain accumulation of heavy metals were analyzed as key points. Additionally, the application potential of gene overexpression and gene editing technologies in reducing cadmium accumulation in rice grains were discussed. Current research on the multi-level interaction networks and genetic regulation of cadmium absorption, transport, and distribution in rice still faces challenges, such as the complexity of interaction networks, unclear regulatory hierarchies, difficulties in gene-environment interactions and field applications, and insufficient understanding of the molecular mechanisms underlying co-stress from multiple heavy metals in complex field environments. It is recommended to precisely edit or design of regulatory elements targeting key transcription factors or non-coding RNAs, and integrate spatiotemporal-specific gene editing, synthetic biology, and multi-omics analyses to deeply elucidate and precisely regulate the cadmium uptake-transport-detoxification network. This will provide a basis for the targeted development of new germplasms with both low cadmium accumulation and high, stable yields.

Key words: rice, cadmium, low accumulation, response mechanisms, genetic regulation

中图分类号:

S859.84

郑兴鑫, 李祖然, 段雨默, 陈玲, 曹力, 祖艳群. 水稻对镉吸收-迁移-分配的多层次互作网络及基因调控研究进展[J]. 中国农学通报, 2026, 42(15): 65-76.

ZHENG Xingxin, LI Zuran, DUAN Yumo, CHEN Ling, CAO Li, ZU Yanqun. Research Advances on Multi-level Interaction Networks and Gene Regulation of Cadmium Absorption, Translocation, and Distribution in Rice (Oryza sativa L.)[J]. Chinese Agricultural Science Bulletin, 2026, 42(15): 65-76.

图/表 7

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基因符号	表达部位	亚细胞定位	功能	参考文献
OsNRAMP1	根、叶片	质膜	促进镉、锰吸收，调控镉毒性	[22]
OsNRAMP2	根、叶片	液泡膜	介导镉运输及籽粒累积	[23]
OsNRAMP4	根系	细胞膜	改变镉的细胞分布和籽粒镉含量	[21,24]
OsNRAMP5	根系	质膜	介导镉和锰的被动吸收，敲除后显著降低根系镉积累	[25-26]
OsHMA2	茎节	质膜	外排镉，减少向籽粒的转运	[27]
OsHMA3	根系、液泡	液泡膜	液泡区隔镉，减少镉装载和累积	[28-30]
OsLCT1/2	韧皮部	质膜	介导Cd由韧皮部向籽粒分配	[30-31]
OsLCT3	根系	质膜	正调控籽粒Cd积累；敲除显著降低茎叶和糙米镉含量	[32]
OsZIP1	根系	质膜	外排镉、锌和铜，减少根系镉积累	[33-34]
OsZIP5	全株	质膜	介导镉吸收	[35]
OsZIP6/9	根、茎	质膜	调控镉吸收和转运	[35-36]
OsSPR1	根系	质膜	突变体影响镉吸收	[37]
OsABCC1	液泡	液泡膜	螯合镉-植物螯合肽复合物至液泡，减少籽粒镉累积	[38]
OsABCC9	液泡	液泡膜	液泡隔离镉，增强镉耐受性	[30,39]
OsABCG36	根系	质膜	敲除使水稻对镉敏感，增加根部镉积累	[30,40]
OsABCG43	叶片、根系	质膜	调控水稻中的镉累积	[21]
OsABCG48	根系	质膜	外排镉，降低根系对镉的吸收	[41]
OsCCX2	茎节	质膜	调控镉木质部装载	[42]
OsCd1	根、茎	质膜	介导根系对镉的吸收及籽粒积累	[43-44]
CF1	茎节、籽粒	细胞膜	减少籽粒镉含量	[45]
OsMTP11	叶片	反式高尔基体、液泡膜	将镉区室化到液泡，减少镉向籽粒转运	[46]
OsLCD	根、籽粒	细胞质、细胞核	调控籽粒中镉的累积，敲除后籽粒镉含量降低	[30,47]
OsNTP1	根系	叶绿体	介导镉向地上部运输	[48]
OsWRKY15	根系、叶片	细胞核	调控镉胁迫响应基因表达，增强镉耐性	[3]
OsWRKY49	根系	细胞核	调控OsHMA3表达，增强镉耐性	[45,49]
OsVIT1/2	根系	液泡膜	将镉隔离至液泡，降低地上部镉累积	[50]
miR535	全株	非编码RNA	抑制镉转运	[51]
miR166/390	根系	非编码RNA	调控镉累积和镉耐性	[52-53]

基因符号	表达部位	亚细胞定位	功能	参考文献
OsNRAMP1	根、叶片	质膜	促进镉、锰吸收，调控镉毒性	[22]
OsNRAMP2	根、叶片	液泡膜	介导镉运输及籽粒累积	[23]
OsNRAMP4	根系	细胞膜	改变镉的细胞分布和籽粒镉含量	[21,24]
OsNRAMP5	根系	质膜	介导镉和锰的被动吸收，敲除后显著降低根系镉积累	[25-26]
OsHMA2	茎节	质膜	外排镉，减少向籽粒的转运	[27]
OsHMA3	根系、液泡	液泡膜	液泡区隔镉，减少镉装载和累积	[28-30]
OsLCT1/2	韧皮部	质膜	介导Cd由韧皮部向籽粒分配	[30-31]
OsLCT3	根系	质膜	正调控籽粒Cd积累；敲除显著降低茎叶和糙米镉含量	[32]
OsZIP1	根系	质膜	外排镉、锌和铜，减少根系镉积累	[33-34]
OsZIP5	全株	质膜	介导镉吸收	[35]
OsZIP6/9	根、茎	质膜	调控镉吸收和转运	[35-36]
OsSPR1	根系	质膜	突变体影响镉吸收	[37]
OsABCC1	液泡	液泡膜	螯合镉-植物螯合肽复合物至液泡，减少籽粒镉累积	[38]
OsABCC9	液泡	液泡膜	液泡隔离镉，增强镉耐受性	[30,39]
OsABCG36	根系	质膜	敲除使水稻对镉敏感，增加根部镉积累	[30,40]
OsABCG43	叶片、根系	质膜	调控水稻中的镉累积	[21]
OsABCG48	根系	质膜	外排镉，降低根系对镉的吸收	[41]
OsCCX2	茎节	质膜	调控镉木质部装载	[42]
OsCd1	根、茎	质膜	介导根系对镉的吸收及籽粒积累	[43-44]
CF1	茎节、籽粒	细胞膜	减少籽粒镉含量	[45]
OsMTP11	叶片	反式高尔基体、液泡膜	将镉区室化到液泡，减少镉向籽粒转运	[46]
OsLCD	根、籽粒	细胞质、细胞核	调控籽粒中镉的累积，敲除后籽粒镉含量降低	[30,47]
OsNTP1	根系	叶绿体	介导镉向地上部运输	[48]
OsWRKY15	根系、叶片	细胞核	调控镉胁迫响应基因表达，增强镉耐性	[3]
OsWRKY49	根系	细胞核	调控OsHMA3表达，增强镉耐性	[45,49]
OsVIT1/2	根系	液泡膜	将镉隔离至液泡，降低地上部镉累积	[50]
miR535	全株	非编码RNA	抑制镉转运	[51]
miR166/390	根系	非编码RNA	调控镉累积和镉耐性	[52-53]

转录因子家族	基因	在植物中的作用	植物	参考文献
WRKY	AtWRKY12	负调控拟南芥的镉耐受性；调控镉螯合和区室化	拟南芥	[84]
	AtWRKY13	正调控拟南芥的镉耐受性；抑制镉吸收，减少整体镉累积	拟南芥	[85]
	CaWRKY41	通过清除ROS增强镉耐受性；正向调控镉吸收	辣椒	[86]
	PyWRKY75	过表达增强镉耐性；增强镉吸收和螯合	杨树	[87]
	GmWRKY142	上调镉耐受Ⅰ类似基因赋予镉抗性；抑制镉吸收	大豆	[76]
HSF	SaHsfA4c	通过清除ROS增强镉耐性；促进镉吸收和累积，调控镉向地上部转运	东南景天	[88]
HSF	HSFIA	上调褪黑素生物合成赋予植株镉耐性；增强镉液泡区隔	番茄	[89]
bHLH	AtbHLH104	正调控重金属吸收和解毒相关基因表达；抑制镉向地上部转运	拟南芥	[90]
MYB	SbMYB15	过表达赋予转基因烟草镉和镍耐受性；抑制镉吸收和向地上部转运	烟草	[91]
MYB	AtMYB4	增强抗氧化防御系统和协调PCS1/MT1C表达调控拟南芥镉耐性；增强镉累积能力，间接调控镉累积和分配	拟南芥	[92]
C2H2	ZAT17	与RNA剪接复合体MAC互作，负调控镉耐受性，影响镉响应基因的选择性剪接；正调控镉吸收，抑制镉转运	拟南芥	[93]
bZIP	SpbZIP60	保护光合器官并降低镉含量；抑制镉吸收和累积	拟南芥	[94]
NAC	ANAC004	增强细胞壁半纤维素对镉的固定，减少根细胞Cd²⁺内流；抑制镉吸收和转运，促进镉液泡区隔	拟南芥	[95]

转录因子家族	基因	在植物中的作用	植物	参考文献
WRKY	AtWRKY12	负调控拟南芥的镉耐受性；调控镉螯合和区室化	拟南芥	[84]
	AtWRKY13	正调控拟南芥的镉耐受性；抑制镉吸收，减少整体镉累积	拟南芥	[85]
	CaWRKY41	通过清除ROS增强镉耐受性；正向调控镉吸收	辣椒	[86]
	PyWRKY75	过表达增强镉耐性；增强镉吸收和螯合	杨树	[87]
	GmWRKY142	上调镉耐受Ⅰ类似基因赋予镉抗性；抑制镉吸收	大豆	[76]
HSF	SaHsfA4c	通过清除ROS增强镉耐性；促进镉吸收和累积，调控镉向地上部转运	东南景天	[88]
HSF	HSFIA	上调褪黑素生物合成赋予植株镉耐性；增强镉液泡区隔	番茄	[89]
bHLH	AtbHLH104	正调控重金属吸收和解毒相关基因表达；抑制镉向地上部转运	拟南芥	[90]
MYB	SbMYB15	过表达赋予转基因烟草镉和镍耐受性；抑制镉吸收和向地上部转运	烟草	[91]
MYB	AtMYB4	增强抗氧化防御系统和协调PCS1/MT1C表达调控拟南芥镉耐性；增强镉累积能力，间接调控镉累积和分配	拟南芥	[92]
C2H2	ZAT17	与RNA剪接复合体MAC互作，负调控镉耐受性，影响镉响应基因的选择性剪接；正调控镉吸收，抑制镉转运	拟南芥	[93]
bZIP	SpbZIP60	保护光合器官并降低镉含量；抑制镉吸收和累积	拟南芥	[94]
NAC	ANAC004	增强细胞壁半纤维素对镉的固定，减少根细胞Cd²⁺内流；抑制镉吸收和转运，促进镉液泡区隔	拟南芥	[95]

调控类型	基因/RNA符号	靶基因	作用机制	表型效果	参考文献
miRNA	miRNA535	OsLCT	结合OsLCT1 mRNA 3'UTR诱导降解，过表达降低籽粒镉含量	阻断韧皮部镉转运	[51]
lncRNA	lncRNA-CdR1	OsHMA3	染色质重塑激活OsHMA3启动子，过表达株系提高液泡镉隔离效率	降低籽粒镉含量	[102]
DNA甲基化	—	OsNRAMP5	镉胁迫下启动子区去甲基化，促进OsNRAMP5表达	镉吸收：籼稻>粳稻	[43,103]

水稻对镉吸收-迁移-分配的多层次互作网络及基因调控研究进展

Research Advances on Multi-level Interaction Networks and Gene Regulation of Cadmium Absorption, Translocation, and Distribution in Rice (Oryza sativa L.)

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技术	降镉效率	生长影响	应用潜力	参考文献
OsHMA3过表达	*****	低	高，多作物适用	[29,104]
OsNRAMP5敲除	****	中	中，需基因型适配	[26,103]
OsLCT1敲除	***	低	高，易整合至品种	[47,108]
多基因编辑 (OsNRAMP5+ OsLCT1）	****	中	中，复杂性高	[31]

策略	靶基因	效果	优势/局限	参考文献
敲除吸收基因	OsNRAMP5	降低糙米镉含量	高效，但可能导致锰缺乏	[26,47,54,103]
敲除转运基因	OsLCT1	降低籽粒镉含量	不影响必需元素，易应用	[47,108]
多基因编辑	OsNRAMP5+ OsLCT1	叠加降镉效果	协同作用，可能影响生长	[31,47]
组成型过表达	OsHMA3	根部镉含量增加，降低籽粒镉含量	效果优于敲除OsNRAMP5	[29,61,62,104]
组织特异性表达	OsHMA3	节点特异表达，降低籽粒镉含量	避免组成型表达的生长抑制	[68]
启动子编辑	OsNRAMP5调控区	降低表达但不完全敲除，平衡镉吸收与锰需求	减少副作用，维持产量	[103]

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