SiYSL1 regulates the iron absorption, transportation, and grain accumulation in foxtail millet

doi:10.1016/j.jia.2026.03.053

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SiYSL1 regulates the iron absorption, transportation, and grain accumulation in foxtail millet

Shuo Zhang^{1, 2}, Hui Zhi¹, Sha Tang¹, Guanqing Jia¹, Chanjuan Tang¹, Hailong Wang¹, Mingzhao Luo¹, Jiao Chunhai², Haiya Cai², Yanhao Xu², Xianmin Diao^1#

¹China State Key Laboratory of Crop Gene Resources and Breeding/Key laboratory Grain Crop Genetic Resources Evaluation and Utilization, Ministry of Agriculture and Rural Affairs/Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China

²Hubei Key Laboratory of Food Crop Germplasm and Genetic Improvement, Food Crops Institute, Hubei Academy of Agricultural Sciences, Wuhan 430064, China

Highlights

ž The iron (Fe) chelate transporter gene SiYSL1 was cloned using a yellow stripe leaf mutant of foxtail millet.

ž SiYSL1 is specifically expressed in vascular bundles of roots and leaves, spikelets, expanding ovaries, basal endosperm, and embryo-surrounding tissues, appears to play an essential role in Fe uptake, transport and accumulation in seeds.

ž The overexpression of SiYSL1 in rice and foxtail millet significantly increased the Fe contents in seeds and spikelet, providing new genetic resources for the Fe bio-enhancement of food crops.

Abstract
References

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摘要

缺铁（Fe）是最为普遍的人体微量营养缺乏症之一，其导致的缺铁性贫血是一个全球性健康问题。然而，主要粮食作物籽粒的Fe含量普遍偏低。因此，提高谷物中的Fe含量对于以谷物为主食的人群（尤其是发展中国家人群）的健康至关重要。本研究分离到一个谷子叶片黄条纹突变体Siysl1。该突变体表现出典型的Fe缺乏表型，在施加充足Fe条件下突变表型被恢复。与野生型相比，Siysl1突变体苗期的根部，已及拔节期的茎和叶、穗以及种子中的Fe含量均显著降低，但抽穗期叶片的Fe含量略有升高。本研究利用MutMap+法定位并克隆了目标基因SiYSL1，通过CRISPR/Cas9介导的基因敲除验证了SiYSL1的功能。SiYSL1编码一个Fe-PS（植物铁载体）转运蛋白，其表达在缺铁条件下被强烈诱导。组织化学染色显示，SiYSL1在缺铁条件下的幼苗根系和叶片的维管束中，以及小花、膨大的子房、胚乳基部和胚周围组织中特异性表达。这些结果表明，SiYSL1参与调控Fe的吸收及稳态，并在Fe向籽粒的转运过程中发挥重要作用。在水稻和谷子中过表达SiYSL1可显著提高种子及穗部的Fe含量，表明SiYSL1在作物铁生物强化育种中具有潜在应用价值。对SiYSL1启动子中转录因子结合位点进行预测，结合穗部转录组分析，发掘到一些可能调控SiYSL1表达的转录因子。本研究为粮食作物的铁生物强化育种提供了新的基因资源，并为进一步认知作物种子中铁元素积累的机制提供了支撑。

Abstract

Iron (Fe) deficiency is a globally widespread condition in which the body lacks sufficient Fe to produce hemoglobin. However, major food crops generally have low grain Fe contents. Consequently, enhancing grain Fe concentrations is important for improving the health of populations that rely on grains as staple foods. Here, we isolated a yellow stripe leaf mutant of foxtail millet (Setaria italica), designated yellow stripe-like 1 (ysl1). This mutant exhibited typical Fe deficiency symptoms that were alleviated when grown under Fe-sufficient conditions. Compared with the wild-type, Siysl1 showed lower Fe concentrations in seedling roots, shoots, stems, elongation-stage leaves, panicles, and seeds, but a higher Fe concentration in heading-stage leaves. Using MutMap+, we identified and cloned SiYSL1 and validated its function through CRISPR/Cas9-mediated knockout experiments. SiYSL1 encodes an Fe-phytosiderophore transporter and is highly induced under Fe deficiency conditions. Histochemical staining revealed that SiYSL1 is specifically expressed in vascular bundles of roots and leaves of plants grown under Fe deficiency conditions, and in spikelets, expanding ovaries, basal endosperm, and embryo-surrounding tissues. Thus, SiYSL1 appears to regulate Fe uptake and homeostasis, and plays an essential role in Fe translocation to seeds. The overexpression of SiYSL1 in rice and foxtail millet significantly increased seed Fe contents, suggesting its value in crop breeding. Predicted transcription factor binding sites in the SiYSL1 promoter and a spikelet transcriptome analysis indicated that transcription factors regulate SiYSL1 expression. Our study provides new genetic resources for the Fe bio-enhancement of food crops and insights into the mechanisms responsible for seed Fe accumulation.

Keywords: foxtail millet iron absorption transport

Online: 24 March 2026

Fund:

This research was supported by the Hubei Provincial Natural Science Foundation, China (2024AFB365), the National Key Project of Research and the Development Plan of China (2021YFF1000100), and the National Natural Sciences Foundation of China (32241042), the Funds Supporting the High-Quality Development of the Seed Industry in Hubei Province, China (HBZY2023A001-48), and the China Agricultural Research System (CAR-06-04).

About author: Shuo Zhang, E-mail: zhangshuo0728@126.com; #Correspondence Xianmin Diao, E-mail: diaoxianmin@caas.cn

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Shuo Zhang, Hui Zhi, Sha Tang, Guanqing Jia, Chanjuan Tang, Hailong Wang, Mingzhao Luo, Jiao Chunhai, Haiya Cai, Yanhao Xu, Xianmin Diao. 2026. SiYSL1 regulates the iron absorption, transportation, and grain accumulation in foxtail millet. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2026.03.053

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