欢迎访问作物学报，今天是
作物学报

作物学报 ›› 2026, Vol. 52 ›› Issue (4): 1006-1021.doi: 10.3724/SP.J.1006.2026.53063

所属专题： 玉米：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米根尖及整根响应缺铁的转录组比较研究

杨亚莉(), 徐明睿, 马越飞, 海艺蕊, 刘凯栋, 刘万茂, 孙颖*()   

  1. 宁夏大学农学院, 宁夏银川 750021
  • 收稿日期:2025-08-16 接受日期:2026-01-22 出版日期:2026-04-12 网络出版日期:2026-02-05
  • 通讯作者: *孙颖, E-mail: sunying@nxu.edu.cn
  • 作者简介:E-mail: 18394471072@163.com
  • 基金资助:
    宁夏粮食作物种质创制与生长调控科技创新团队项目(2022BSB03109);国家自然科学基金项目(32260442);宁夏大学研究生创新项目(CXXM2024-09)

Comparative transcriptome analysis of maize root tips and whole roots in response to iron deficiency

Yang Ya-Li(), Xu Ming-Rui, Ma Yue-Fei, Hai Yi-Rui, Liu Kai-Dong, Liu Wan-Mao, Sun Ying*()   

  1. College of Agriculture, Ningxia University, Yinchuan 750021, Ningxia, China
  • Received:2025-08-16 Accepted:2026-01-22 Published:2026-04-12 Published online:2026-02-05
  • Contact: *E-mail: sunying@nxu.edu.cn
  • Supported by:
    Ningxia Grain Crop Germplasm Creation and Growth Regulation Science and Technology Innovation Team Project(2022BSB03109);National Natural Science Foundation of China(32260442);Ningxia University Postgraduate Innovation Program(CXXM2024-09)

RichHTML

11

PDF

160

摘要：

根系响应缺铁的转录组研究多基于整个地下部根系或整根混合样本, 这会导致根尖强烈的转录变化信号被平均化。为解析玉米初生根根尖及整根响应缺铁胁迫的空间转录组特征, 揭示铁吸收与稳态调控的空间分工, 本研究以玉米自交系B73为材料, 设置对照组(25 μmol L-1 Fe-EDTA)与缺铁组(0 μmol L-1 Fe-EDTA) 2个处理, 对三叶期幼苗初生根的根尖(0~2 cm)和整根进行RNA-seq转录组测序。结合差异表达基因(differentially expressed genes, DEGs)分析、多层次基因功能富集分析、加权共表达网络分析(weighted gene co-expression network analysis, WGCNA)及qRT-PCR验证, 系统比较根尖和整根响应缺铁的分子机制。结果显示, 根尖鉴定到4206个DEGs (上调2450个), 显著多于整根(325个, 上调84个), 表明根尖是铁响应的核心区域。功能富集分析表明, 根尖主要激活核糖体组装和TCA循环等代谢通路; 而整根则显著富集于木质素合成、抗氧化防御等过程。多个次生代谢通路, 如苯丙烷生物合成、各种植物次生代谢物的生物合成、类黄酮生物合成, 在初生根不同部位显著富集, 提示次生代谢物通过多途径参与铁稳态调控。铁载体生物合成基因负责麦根酸(mugineic acids, MAs)类植物铁载体(phytosiderophore, PS)合成与铁螯合, 其在根尖中被特异性诱导。多数关键转运蛋白基因在整根中显著表达, 其中, 天然抗性相关巨噬蛋白2 (natural resistance-associated macrophage protein 2)编码基因NRAMP2和黄色条纹蛋白12 (yellow stripe-like protein12)编码基因YSL12在整根中表达上调, 促进了玉米体内铁运输。此外, 多个调控铁稳态的bHLH家族转录因子也在整根显著表达, 暗示其可能在协调铁吸收与再分配过程中发挥作用。本研究阐明了初生根铁响应的空间转录特征, 揭示了玉米初生根通过根尖主导铁载体合成, 整根负责铁转运与应对缺铁的系统防御, 并发现了具有空间特异性的基因与通路, 为理解玉米根系对缺铁胁迫的分子机制提供见解。

关键词: 玉米, 初生根, 缺铁胁迫, 转录组分析, 空间特异性

Abstract:

Transcriptomic studies investigating root responses to iron (Fe) deficiency have typically used entire underground root systems or mixed whole-root samples, which can dilute the strong transcriptional signals originating from root tips. To uncover the spatially resolved transcriptomic features of Fe deficiency responses in maize primary roots and elucidate the spatial division of labor in Fe uptake and homeostasis, we employed the maize inbred line B73. Seedlings at the three-leaf stage were grown under control (25 μmol L-1 Fe-EDTA) or Fe-deficient (0 μmol L-1 Fe-EDTA) conditions. RNA sequencing (RNA-seq) was performed on root tips (0-2 cm) and whole primary roots. An integrated analysis—including differentially expressed genes (DEGs), multi-level gene function enrichment, weighted gene co-expression network analysis (WGCNA), and qRT-PCR validation—was used to systematically compare the molecular mechanisms of Fe deficiency responses between root tips and whole roots. A total of 4206 DEGs (2450 upregulated) were identified in root tips, substantially more than the 325 DEGs (84 upregulated) found in whole roots, highlighting root tips as the key region for Fe sensing and response. Functional enrichment analysis revealed that root tips primarily activated metabolic pathways such as ribosome assembly and the TCA cycle, while whole roots were significantly enriched in processes including lignin biosynthesis and antioxidant defense. Several secondary metabolite biosynthesis pathways—including phenylpropanoid biosynthesis, various plant secondary metabolite biosynthesis, and flavonoid biosynthesis—were enriched in distinct root regions, suggesting diverse roles of secondary metabolites in Fe homeostasis. Genes involved in siderophore biosynthesis were specifically induced in root tips, supporting the synthesis of mugineic acids (MAs), the main phytosiderophores (PS) in grasses, and subsequent Fe chelation. Key transporter genes, such as natural resistance-associated macrophage protein 2 (NRAMP2) and yellow stripe-like protein 12 (YSL12), were predominantly expressed and upregulated in whole roots, facilitating Fe translocation within the plant. Additionally, several bHLH family transcription factors, known regulators of Fe homeostasis, were highly expressed in whole roots, indicating their potential role in coordinating Fe uptake and redistribution. This study delineates the spatially partitioned transcriptional landscape of Fe deficiency responses in primary roots, revealing a strategy in which root tips dominate PS biosynthesis, while whole roots coordinate Fe transport and systemic defense. The identification of spatially specific genes and pathways provides new insights into the molecular mechanisms underlying maize root adaptation to Fe deficiency stress.

Key words: Zea mays L., primary root, iron deficiency stress, transcriptomic analysis, spatial specificity

表1

实时荧光定量PCR引物序列"

基因名称
Gene name		 基因ID
Gene ID		 正向引物
Forward primer (5′-3′)		 反向引物
Reverse primer (5′-3′)
ZmACT1 Zm00001d010159 ACCTCTTCGAGCAGAATAGTTG CCTTGACCATTCCAGTGCCA
ZmGAPDH2 Zm00001d035156 CCTACCTCTCCAGCATCCTCC TTCCGATCTTGATCTTGCCCAT
ZmDMAS1 Zm00001d028360 TCTTCGTCACGTCCAAGGTCT GTATTCCATCTGGAGATTGCTGAG
ZmENA1 Zm00001d052532 TCATCCCATTCGCTGTGCTC AGCAATGGCGAGGCAGAATA
ZmNRAMP5 Zm00001d019327 ATCCTGGCAACTTGGAGACG GAGGCCAATGAGGATCACCC
ZmYS1 Zm00001d017429 GTGCTACCAACAGAGTGGCA ATGATGACCGCTGAGACGAC
ZmYSL12 Zm00001d025887 TGGACTGCAGGGTTACAGGGTT GCGAATGAGCACCTTCACGA
ZmTOM1 Zm00001d041111 TACGCTGGTTTTCTTGGTGC GCTTCCTTCCGACACGATCA
ZmFIT Zm00001d004007 TGTTTCCGGGTGGAGACCT CCTAATTGTCCCCTCCCCTTG
ZmIRO2 Zm00001d011847 AGCCTCCATCTTCAGAGAAGTCA CATGCGCCAGCACACTCC
ZmPYE Zm00001d029339 GACTGTGCGCCTTCCTAAGT CCTCCTCTCTACCTTCCTGGAT

图1

缺铁胁迫对玉米幼苗表型、生长及叶绿素含量的影响 A: 玉米幼苗在正常供铁(CK)和缺铁处理(-Fe)下的表型; B: 正常供铁(CK)和缺铁处理(-Fe)对玉米地上部鲜重、地上部干重、根系鲜重和根系干重的影响; C: 正常供铁(CK)和缺铁处理(-Fe)对玉米叶绿素含量的影响。CK: 正常供铁处理; -Fe: 缺铁处理; ns表示无显著差异; ****表示差异极显著。"

表2

玉米初生根不同样本转录组测序数据质量分析"

处理
Treatment		 原始读段
Raw reads		 有效读段
Clean reads		 原始碱基数
Raw bases		 高质量碱基数
Clean bases
Root_CK1 47.12 46.80 (99.32%) 7.02 6.98 (99.43%)
Root_CK2 47.86 47.46 (99.16%) 7.12 7.06 (99.16%)
Root_CK3 46.92 46.51 (99.13%) 6.98 6.92 (99.14%)
Root_-Fe1 46.88 46.58 (99.35%) 6.99 6.95 (99.43%)
Root_-Fe2 47.26 46.90 (99.23%) 7.04 6.98 (99.15%)
Root_-Fe3 46.83 46.46 (99.20%) 6.97 6.92 (99.28%)
Tip_CK1 46.22 44.97 (97.30%) 6.77 6.58 (97.19%)
Tip_CK2 48.56 47.26 (97.32%) 7.12 6.93 (97.33%)
Tip_CK3 46.71 45.52 (97.45%) 6.84 6.67 (97.51%)
Tip_-Fe1 41.13 39.99 (97.24%) 6.01 5.85 (97.34%)
Tip_-Fe2 50.13 48.45 (96.65%) 7.28 7.04 (96.70%)
Tip_-Fe3 41.06 40.07 (97.57%) 6.02 5.88 (97.67%)

表3

转录组数据与参考基因组比对结果"

处理
Treatment		 总读段数
Total reads		 单独匹配读段数及占比
Unique-mapped and
proportion of all		 多重匹配读段数及占比
Multiple-mapped and
proportion of all		 总匹配读段数及匹配度
Total-mapped and
proportion of all		 基因数量
Gene
number
Root_CK1 46,801,268 44,404,454 (94.88%) 1,771,120 (3.78%) 46,175,574 (98.66%) 29,044
Root_CK2 47,456,152 45,003,702 (94.83%) 1,812,612 (3.82%) 46,816,314 (98.65%) 29,054
Root_CK3 46,514,244 43,924,332 (94.43%) 1,792,289 (3.85%) 45,716,621 (98.29%) 28,903
Root_-Fe1 46,579,498 44,093,238 (94.66%) 1,767,485 (3.79%) 45,860,723 (98.46%) 28,890
Root_-Fe2 46,897,748 44,180,525 (94.21%) 1,675,511 (3.57%) 45,856,036 (97.78%) 28,897
Root_-Fe3 46,457,514 44,105,061 (94.94%) 1,671,691 (3.60%) 45,776,752 (98.53%) 28,893
Tip_Ck1 44,974,496 42,484,726 (94.46%) 1,891,689 (4.21%) 44,376,415 (98.67%) 27,951
Tip_Ck2 47,263,302 44,275,065 (93.68%) 2,065,236 (4.37%) 46,340,301 (98.05%) 27,990
Tip_Ck3 45,517,552 42,942,088 (94.34%) 1,943,621 (4.27%) 44,885,709 (98.61%) 28,153
Tip_-Fe1 39,993,686 37,974,211 (94.95%) 1,440,835 (3.60%) 39,415,046 (98.55%) 28,144
Tip_-Fe2 48,449,490 45,319,605 (93.54%) 1,844,483 (3.81%) 47,164,088 (97.35%) 28,466
Tip_-Fe3 40,066,072 37,753,681 (94.23%) 1,448,787 (3.62%) 39,202,468 (97.84%) 27,686

图2

整根与根尖缺铁处理后转录水平上的变化 A: 样品主成分分析; B: 差异表达基因韦恩图; C: 差异表达基因柱状图。处理同表2。PC1: 第一主成分; PC2: 第二主成分。"

图3

整根及根尖缺铁处理后差异基因富集分析 A, C: 整根、根尖差异表达基因的GO富集图; B, D: 整根、根尖差异表达基因的KEGG富集图。富集得分越高, 富集程度越高; 点越大, 富集的基因数量越多; 点的颜色越红, 富集越显著。"

图4

关键通路基因集富集分析 A: 整根的基因集富集分析; B: 根尖的基因集富集分析。ES: 富集得分; NES: 归一化富集得分; FDR: 假阳性率。"

图5

加权基因共表达网络分析 A: 聚类树状图9个模块显示在分层聚类树上, 每个分支代表一个模块; B: 基因表达模块与根、根尖铁响应的相关性。红色和蓝色分别表示正相关和负相关, 相关系数和P值显示在每个单元格内。C, D: MEdarkmagenta模块基因的GO与KEGG分析; E, F: MEroyalblue模块基因的GO与KEGG分析。ME: 模块特异性基因。"

图6

铁稳态相关基因表达量热图 处理同表2。"

图7

差异表达基因qRT-PCR验证 以 ZmACT1 和 ZmGAPDH2 作为双内参基因进行归一化。基因相对表达量通过2-ΔΔCt法计算, 并以整根CK组中该基因的表达水平为基准(设定为1)。处理同表2。ns表示无显著差异; *、**、***、****分别表示在0.05、0.01、0.001、0.0001水平差异显著。"

[1] Briat J F, Dubos C, Gaymard F. Iron nutrition, biomass production, and plant product quality. Trends Plant Sci, 2015, 20: 33-40.
doi: 10.1016/j.tplants.2014.07.005
[2] Thomine S, Vert G. Iron transport in plants: better be safe than sorry. Curr Opin Plant Biol, 2013, 16: 322-327.
doi: 10.1016/j.pbi.2013.01.003 pmid: 23415557
[3] Fernández V, Eichert T, Del Río V, et al. Leaf structural changes associated with iron deficiency chlorosis in field-grown pear and peach: physiological implications. Plant Soil, 2008, 311: 161-172.
doi: 10.1007/s11104-008-9667-4
[4] 朱大洲, 武宁, 张勇, 等. 营养导向型作物新品种选育与审定现状、问题与展望. 作物学报, 2023, 49: 1-11.
doi: 10.3724/SP.J.1006.2023.23018
Zhu D Z, Wu N, Zhang Y, et al. Current status, problems and prospects of nutrition-oriented breeding and registration of new crop varieties. Acta Agron Sin, 2023, 49: 1-11 (in Chinese with English abstract).
doi: 10.3724/SP.J.1006.2023.23018
[5] 韩秀萍, 马彦平, 黄灿灿. 微量元素铁与植物营养和人体健康的关系解析. 磷肥与复肥, 2020, 35(12): 33-35.
Han X P, Ma Y P, Huang C C. Analysis of the relationship between iron, plant nutrition and human health. Phosphat Compound Fert, 2020, 35(12): 33-35 (in Chinese with English abstract).
[6] Zheng S J. Iron homeostasis and iron acquisition in plants: maintenance, functions and consequences. Ann Bot, 2010, 105: 799-800.
doi: 10.1093/aob/mcq082
[7] Colombo C, Palumbo G, He J Z, et al. Review on iron availability in soil: interaction of Fe minerals, plants, and microbes. J Soils Sediments, 2014, 14: 538-548.
doi: 10.1007/s11368-013-0814-z
[8] 张妮娜, 上官周平, 陈娟. 植物应答缺铁胁迫的分子生理机制及其调控. 植物营养与肥料学报, 2018, 24: 1365-1377.
Zhang N N, Shangguan Z P, Chen J. Molecular physiological mechanism and regulation of plant responses to iron deficiency stress. J Plant Nutr Fert, 2018, 24: 1365-1377 (in Chinese with English abstract).
[9] Kobayashi T, Nakanishi H, Nishizawa N K. Recent insights into iron homeostasis and their application in graminaceous crops. Proc Jpn Acad Ser B Phys Biol Sci, 2010, 86: 900-913.
doi: 10.2183/pjab.86.900
[10] Nozoye T, Nagasaka S, Kobayashi T, et al. Phytosiderophore efflux transporters are crucial for iron acquisition in graminaceous plants. J Biol Chem, 2011, 286: 5446-5454.
doi: 10.1074/jbc.M110.180026 pmid: 21156806
[11] Kumar A, Joon R, Singh G, et al. The multifaceted role of YSL proteins: iron transport and emerging functions in plant metal homeostasis. Biochim Biophys Acta BBA Gen Subj, 2025, 1869: 130792.
[12] Bashir K, Inoue H, Nagasaka S, et al. Cloning and characterization of deoxymugineic acid synthase genes from graminaceous plants. J Biol Chem, 2006, 281: 32395-32402.
doi: 10.1074/jbc.M604133200 pmid: 16926158
[13] Zhou X J, Li S Z, Zhao Q Q, et al. Genome-wide identification, classification and expression profiling of nicotianamine synthase (NAS) gene family in maize. BMC Genomics, 2013, 14: 238.
doi: 10.1186/1471-2164-14-238 pmid: 23575343
[14] Zhang X, Xiao K, Li S Z, et al. Genome-wide analysis of the NAAT, DMAS, TOM, and ENA gene families in maize suggests their roles in mediating iron homeostasis. BMC Plant Biol, 2022, 22: 37.
doi: 10.1186/s12870-021-03422-7 pmid: 35039017
[15] Nozoye T, Nakanishi H, Nishizawa N K. Characterizing the crucial components of iron homeostasis in the maize mutants ys1 and ys3. PLoS One, 2013, 8: e62567.
[16] Nozoye T, Nakanishi H, Nishizawa N K. Transcriptomic analyses of maize ys1 and ys3 mutants reveal maize iron homeostasis. Genom Data, 2015, 5: 97-99.
[17] Curie C, Panaviene Z, Loulergue C, et al. Maize yellow stripe1 encodes a membrane protein directly involved in Fe(III) uptake. Nature, 2001, 409: 346-349.
doi: 10.1038/35053080
[18] Kobayashi T, Nozoye T, Nishizawa N K. Iron transport and its regulation in plants. Free Radic Biol Med, 2019, 133: 11-20.
doi: 10.1016/j.freeradbiomed.2018.10.439
[19] Yokosho K, Yamaji N, Ueno D, et al. OsFRDL1 is a citrate transporter required for efficient translocation of iron in rice. Plant Physiol, 2009, 149: 297-305.
doi: 10.1104/pp.108.128132 pmid: 19011004
[20] Gao F, Dubos C. Transcriptional integration of plant responses to iron availability. J Exp Bot, 2021, 72: 2056-2070.
doi: 10.1093/jxb/eraa556 pmid: 33246334
[21] Shen C J, Yue R Q, Sun T, et al. OsARF16, a transcription factor regulating auxin redistribution, is required for iron deficiency response in rice (Oryza sativa L.). Plant Sci, 2015, 231: 148-158.
doi: 10.1016/j.plantsci.2014.12.003
[22] Wu X, Jia Y F, Ma Q, et al. The transcription factor bZIP44 cooperates with MYB10 and MYB72 to regulate the response of Arabidopsis thaliana to iron deficiency stress. New Phytol, 2024, 242: 2586-2603.
doi: 10.1111/nph.v242.6
[23] Liu W, Li Q W, Wang Y, et al. Ethylene response factor AtERF72 negatively regulates Arabidopsis thaliana response to iron deficiency. Biochem Biophys Res Commun, 2017, 491: 862-868.
doi: 10.1016/j.bbrc.2017.04.014
[24] Yan J Y, Li C X, Sun L, et al. A WRKY transcription factor regulates Fe translocation under Fe deficiency. Plant Physiol, 2016, 171: 2017-2027.
doi: 10.1104/pp.16.00252 pmid: 27208259
[25] Sun L, Wei Y Q, Wu K H, et al. Restriction of iron loading into developing seeds by a YABBY transcription factor safeguards successful reproduction in Arabidopsis. Mol Plant, 2021, 14: 1624-1639.
doi: 10.1016/j.molp.2021.06.005 pmid: 34116221
[26] Long T A, Tsukagoshi H, Busch W, et al. The bHLH transcription factor POPEYE regulates response to iron deficiency in Arabidopsis roots. Plant Cell, 2010, 22: 2219-2236.
doi: 10.1105/tpc.110.074096
[27] Xu J Q, Zhu X Y, Yan F, et al. Identification of quantitative trait loci associated with iron deficiency tolerance in maize. Front Plant Sci, 2022, 13: 805247.
doi: 10.3389/fpls.2022.805247
[28] Bauer P, Ling H Q, Guerinot M L. FIT, the FER-like iron deficiency induced transcription factor in Arabidopsis. Plant Physiol Biochem, 2007, 45: 260-261.
doi: 10.1016/j.plaphy.2007.03.006
[29] Li S Z, Ma S, Song Z Z, et al. Regulatory mechanisms of iron homeostasis in maize mediated by ZmFIT. Crop J, 2024, 12: 1426-1436.
doi: 10.1016/j.cj.2024.06.013
[30] Svistoonoff S, Creff A, Reymond M, et al. Root tip contact with low-phosphate media reprograms plant root architecture. Nat Genet, 2007, 39: 792-796.
doi: 10.1038/ng2041 pmid: 17496893
[31] Ondzighi-Assoume C A, Chakraborty S, Harris J M. Environmental nitrate stimulates abscisic acid accumulation in Arabidopsis root tips by releasing it from inactive stores. Plant Cell, 2016, 28: 729-745.
doi: 10.1105/tpc.15.00946
[32] Viana W G, Scharwies J D, Dinneny J R. Deconstructing the root system of grasses through an exploration of development, anatomy and function. Plant Cell Environ, 2022, 45: 602-619.
doi: 10.1111/pce.v45.3
[33] Ruiz Herrera L F, Shane M W, López-Bucio J. Nutritional regulation of root development. Wires Dev Biol, 2015, 4: 431-443.
doi: 10.1002/wdev.183 pmid: 25760021
[34] Li Y, Wang N, Zhao F T, et al. Changes in the transcriptomic profiles of maize roots in response to iron-deficiency stress. Plant Mol Biol, 2014, 85: 349-363.
doi: 10.1007/s11103-014-0189-6 pmid: 24648157
[35] Zhang Y, Zheng G H, Liu P, et al. Morphological and physiological responses of root tip cells to Fe2+ toxicity in rice. Acta Physiol Plant, 2011, 33: 683-689.
doi: 10.1007/s11738-010-0590-y
[36] 魏琦, 何冠华, 张登峰, 等. 基于抗旱玉米自交系SL001的抗旱优异基因资源挖掘. 作物学报, 2025, 51: 3171-3183.
doi: 10.3724/SP.J.1006.2025.53024
Wei Q, He G H, Zhang D F, et al. Mining of excellent drought-tolerant gene resources based on drought-tolerant maize inbred line SL001. Acta Agron Sin, 2025, 51: 3171-3183 (in Chinese with English abstract).
doi: 10.3724/SP.J.1006.2025.53024
[37] 张锦辉, 肖姿仪, 李旭华, 等. 玉米突变体caspl2b2的耐盐特性评价及转录组分析. 作物学报, 2024, 50: 3144-3154.
doi: 10.3724/SP.J.1006.2024.43017
Zhang J H, Xiao Z Y, Li X H, et al. Evaluation of salt tolerance and transcriptome analysis of maize mutant caspl2b2. Acta Agron Sin, 2024, 50: 3144-3154 (in Chinese with English abstract).
doi: 10.3724/SP.J.1006.2024.43017
[38] Vert G, Grotz N, Dédaldéchamp F, et al. IRT1, an Arabidopsis transporter essential for iron uptake from the soil and for plant growth. Plant Cell, 2002, 14: 1223-1233.
doi: 10.1105/tpc.001388
[39] Inoue H, Takahashi M, Kobayashi T, et al. Identification and localisation of the rice nicotianamine aminotransferase gene OsNAAT1 expression suggests the site of phytosiderophore synthesis in rice. Plant Mol Biol, 2008, 66: 193-203.
doi: 10.1007/s11103-007-9262-8
[40] 乔孟欣, 李素贞, 陈景堂. 玉米铁还原酶基因ZmFRO2的功能分析. 生物技术通报, 2020, 36(11): 9-20.
doi: 10.13560/j.cnki.biotech.bull.1985.2020-0281
Qiao M X, Li S Z, Chen J T. Functional analysis of ferric reductase gene ZmFRO2 in maize. Biotechnol Bull, 2020, 36(11): 9-20 (in Chinese with English abstract).
[41] Ueno D, Yamaji N, Ma J F. Further characterization of ferric-phytosiderophore transporters ZmYS1 and HvYS1 in maize and barley. J Exp Bot, 2009, 60: 3513-3520.
doi: 10.1093/jxb/erp191 pmid: 19549626
[42] Takatsuka H, Umeda M. Hormonal control of cell division and elongation along differentiation trajectories in roots. J Exp Bot, 2014, 65: 2633-2643.
doi: 10.1093/jxb/ert485 pmid: 24474807
[43] Blancaflor E B, Masson P H. Plant gravitropism: unraveling the ups and downs of a complex process. Plant Physiol, 2003, 133: 1677-1690.
pmid: 14681531
[44] Giehl R F H, Lima J E, von Wirén N. Localized iron supply triggers lateral root elongation in Arabidopsis by altering the AUX1-mediated auxin distribution. Plant Cell, 2012, 24: 33-49.
doi: 10.1105/tpc.111.092973
[45] Song H, Chen F, Wu X, et al. MNB1 gene is involved in regulating the iron-deficiency stress response in Arabidopsis thaliana. BMC Plant Biol, 2022, 22: 151.
doi: 10.1186/s12870-022-03553-5 pmid: 35346040
[46] Zhang X, Xue C W, Wang R N, et al. Physiological and proteomic dissection of the rice roots in response to iron deficiency and excess. J Proteom, 2022, 267: 104689.
doi: 10.1016/j.jprot.2022.104689
[47] 王哲, 孙亿敬, 李娇爱, 等. 高等植物铁元素的吸收、转位和调控. 上海师范大学学报(自然科学版), 2017, 46: 729-739.
Wang Z, Sun Y J, Li J A, et al. Iron uptake, translocation, and regulation in higher plants. J Shanghai Norm Univ (Nat Sci), 2017, 46: 729-739 (in Chinese with English abstract).
[48] 董晓雨, 郝梦琪, 郭东林. 黄色条纹/黄色条纹样蛋白与植物中的铁转运功能. 中国生物化学与分子生物学报, 2017, 33: 886-892.
doi: 10.13865/j.cnki.cjbmb.2017.09.06
Dong X Y, Hao M Q, Guo D L. YS/YSL protein and its iron transport function in plants. Chin J Biochem Mol Biol, 2017, 33: 886-892 (in Chinese with English abstract).
[49] 李素贞, 陈景堂. 植物锌铁转运相关蛋白家族的研究进展. 生物技术通报, 2013, 29(2): 8-14.
Li S Z, Chen J T. Progresses in studying of protein families involved in Zn/Fe transporting in plants. Biotechnol Bull, 2013, 29(2): 8-14 (in Chinese with English abstract).
[50] Waters B M, Chu H H, Didonato R J, et al. Mutations in Arabidopsis Yellow Stripe-Like1 and Yellow Stripe-Like3 reveal their roles in metal ion homeostasis and loading of metal ions in seeds. Plant Physiol, 2006, 141: 1446-1458.
doi: 10.1104/pp.106.082586 pmid: 16815956
[51] DiDonato R J Jr, Roberts L A, Sanderson T, et al. Arabidopsis Yellow Stripe-Like2 (YSL2): a metal-regulated gene encoding a plasma membrane transporter of nicotianamine-metal complexes. Plant J, 2004, 39: 403-414.
doi: 10.1111/j.1365-313X.2004.02128.x pmid: 15255869
[52] Zheng L Q, Fujii M, Yamaji N, et al. Isolation and characterization of a barley yellow stripe-like gene, HvYSL5. Plant Cell Physiol, 2011, 52: 765-774.
doi: 10.1093/pcp/pcr009
[53] 刘元峰, 李素贞, 郭晋杰, 等. 植物YSL家族基因研究进展. 生物技术通报, 2017, 33(9): 1-9.
doi: 10.13560/j.cnki.biotech.bull.1985.2017-0375
Liu Y F, Li S Z, Guo J J, et al. Research progress on YSL transporters gene family. Biotechnol Bull, 2017, 33(9): 1-9 (in Chinese with English abstract).
[54] Koike S, Inoue H, Mizuno D, et al. OsYSL2 is a rice metal-nicotianamine transporter that is regulated by iron and expressed in the phloem. Plant J, 2004, 39: 415-424.
doi: 10.1111/tpj.2004.39.issue-3
[55] Yuan Y X, Wu H L, Wang N, et al. FIT interacts with AtbHLH38 and AtbHLH39 in regulating iron uptake gene expression for iron homeostasis in Arabidopsis. Cell Res, 2008, 18: 385-397.
[56] Ogo Y, Nakanishi Itai R, Nakanishi H, et al. The rice bHLH protein OsIRO2 is an essential regulator of the genes involved in Fe uptake under Fe-deficient conditions. Plant J, 2007, 51: 366-377.
doi: 10.1111/j.1365-313X.2007.03149.x pmid: 17559517
[57] 申红芸, 熊宏春, 郭笑彤, 等. 植物吸收和转运铁的分子生理机制研究进展. 植物营养与肥料学报, 2011, 17: 1522-1530.
Shen H Y, Xiong H C, Guo X T, et al. Progress of molecular and physiological mechanism of iron uptake and translocation in plants. Plant Nutr Fert Sci, 2011, 17: 1522-1530 (in Chinese with English abstract).
[58] Dakora F D, Phillips D A. Root exudates as mediators of mineral acquisition in low-nutrient environments. Plant Soil, 2002, 245: 35-47.
doi: 10.1023/A:1020809400075
[59] Jin C W, You G Y, Zheng S J. The iron deficiency-induced phenolics secretion plays multiple important roles in plant iron acquisition underground. Plant Signal Behav, 2008, 3: 60-61.
doi: 10.4161/psb.3.1.4902 pmid: 19704773
[60] Cesco S, Neumann G, Tomasi N, et al. Release of plant-borne flavonoids into the rhizosphere and their role in plant nutrition. Plant Soil, 2010, 329: 266.
[61] Paffrath V, Tandron Moya Y A, Weber G, et al. A major role of coumarin-dependent ferric iron reduction in strategy I-type iron acquisition in Arabidopsis. Plant Cell, 2024, 36: 642-664.
doi: 10.1093/plcell/koad279
[62] Robe K, Stassen M J J, Watanabe S, et al. Coumarin-facilitated iron transport: an IRT1-independent strategy for iron acquisition in Arabidopsis thaliana. Plant Commun, 2025, 6: 101431.
doi: 10.1016/j.xplc.2025.101431
[63] Ishimaru Y, Kakei Y, Shimo H, et al. A rice phenolic efflux transporter is essential for solubilizing precipitated apoplasmic iron in the plant stele. J Biol Chem, 2011, 286: 24649-24655.
doi: 10.1074/jbc.M111.221168 pmid: 21602276
[1] 刘恩波, 陈静, 李红星, 于宁宁, 任佰朝, 赵斌, 刘鹏, 张吉旺. 遮阴改变源-库平衡和调节碳水化合物代谢进而抑制夏玉米幼穗发育[J]. 作物学报, 2026, 52(6): 1891-1901.
[2] 梁进宇, 尹嘉德, 王红丽, 张国平, 侯慧芝, 董博, 马明生. 基于无人机高光谱和机器学习的旱地饲用玉米叶片氮含量估测[J]. 作物学报, 2026, 52(6): 1788-1801.
[3] 孙淑凤, 许振南, 黄嘉鑫, 翁建峰, 李新海. 玉米MAPK家族全基因组鉴定及其对拟轮枝镰孢菌感染的响应[J]. 作物学报, 2026, 52(5): 1291-1308.
[4] 张宁宁, 滕雨菲, 任娜娜, 魏兴卓, 闫书豪, 樊可心, 王永宏, 陈文康, 张兴华, 朱万超, 徐淑兔, 薛吉全. 201份玉米自交系抗旱表型评价及可塑性分析[J]. 作物学报, 2026, 52(5): 1309-1325.
[5] 杨欣雨, 崔文涛, 迪力尼格尔·阿力木, 汪凯翔, 吴鹏昊, 任姣姣. 玉米穗上叶片数全基因组关联分析和全基因组选择[J]. 作物学报, 2026, 52(5): 1573-1590.
[6] 韩亚鑫, 何冠华, 张小琼, 张登峰, 李永祥, 刘旭洋, 王天宇, 黎裕, 邹华文, 李春辉. 基于RNA-Seq和BSA-Seq联合分析挖掘玉米侧根密度基因资源[J]. 作物学报, 2026, 52(5): 1341-1352.
[7] 张鸿蓉, 王菲儿, 李盼, 仇海龙, 朱静, 赵连豪, 南运有, 何蔚, 樊志龙, 胡发龙, 柴强, 殷文. 减量20%灌水与25%有机肥替代化肥提高青贮玉米产量的光合特性[J]. 作物学报, 2026, 52(5): 1487-1500.
[8] 杨扬, 常诗惠, 田红丽, 易红梅, 王璐, 任洁, 范亚明, 刘亚维, 王凤格, 赵久然. 不同生态区国审玉米品种的遗传多样性分析[J]. 作物学报, 2026, 52(5): 1352-1364.
[9] 蔡宏玮, 于爱忠, 姜科强, 王鹏飞, 王玉珑, 霍建喆, 庞小能, 尹波, 尚永盼. 干旱灌区有机肥替代部分化肥促进甜玉米产量提升的关键机制[J]. 作物学报, 2026, 52(4): 1166-1180.
[10] 田红丽, 杨扬, 范亚明, 易红梅, 郭丹丹, 王凤格, 赵久然. 适于玉米品种鉴定的一套三等位变异SNP新型标记组合[J]. 作物学报, 2026, 52(4): 993-1005.
[11] 张超, 郭欢, 李忠玲, 岳淑宁, 赵娜. 基于BSA-seq技术定位玉米籽粒花青素关联基因[J]. 作物学报, 2026, 52(3): 780-789.
[12] 郭向阳, 涂亮, 王栋, 刘鹏飞, 王安贵, 易强, 任洪, 李刚, 祝云芳, 吴迅, 蒋喻林, 田丰, 陈泽辉. 热带Suwan种质在我国玉米种质改良中的创新与利用[J]. 作物学报, 2026, 52(3): 655-664.
[13] 马亮, 马璐, 张舒钰, 章慧敏, 王仁明, 宋旭东, 张振良, 冒宇翔, 陆虎华, 陈国清, 郝德荣, 周广飞. 玉米苞叶数目转录组分析及候选基因鉴定[J]. 作物学报, 2026, 52(3): 790-801.
[14] 孟成, 王哲. 玉米ZmPFK基因家族全基因组鉴定及响应胁迫表达分析[J]. 作物学报, 2026, 52(3): 764-779.
[15] 李新浩, 邢梦柯, 周梓惠, 李思烨, 任昊, 王洪章, 赖华江. 外源褪黑素通过协调光反应与暗反应增强玉米苗期的耐热性[J]. 作物学报, 2026, 52(3): 839-856.
Viewed
Full text


Abstract

Cited
  Shared   
  Discussed   
No Suggested Reading articles found!
京ICP备15008789号-2