AbstractCadmium (Cd) contamination poses significant risks to agricultural productivity and human health, particularly through its accumulation in staple crops such as bread wheat (Triticum aestivum L.). This study evaluated Cd accumulation and tolerance among six bread wheat cultivars exposed to six Cd concentrations (0, 2.5, 5, 10, 15, 20, and 25 mg kg−1 soil).

摘要镉（Cd）污染对农业生产力和人类健康构成重大风险，特别是通过其在面包小麦（Triticum aestivum L.）等主要作物中的积累。本研究评估了暴露于六种Cd浓度（0、2.5、5、10、15、20和25 mg kg-1土壤）的六种面包小麦品种中Cd的积累和耐受性。

Phenotypic assessments and quantitative real-time PCR (qRT-PCR) were conducted to analyze the expression patterns of TaNRAMP and TaZIP genes in various tissues and developmental stages of wheat, which play crucial roles in Cd uptake and transport. Results demonstrated significant variability in Cd accumulation.

。结果表明，Cd积累存在显着差异。

The Barat cultivar exhibited the lowest accumulation in grain (ranging from 0.21 to 8.8 mg kg−1) and the highest tolerance. In contrast, Kavir and Pishtaz displayed elevated Cd levels in both grain and straw, while Parsi accumulated more Cd in straw at lower concentrations (56.9 mg kg−1 in Cd concentration of 10 mg kg−1 soil).

Barat品种在谷物中的积累最低（范围为0.21至8.8 mg kg-1），耐受性最高。相比之下，Kavir和Pishtaz在谷物和秸秆中均显示出较高的Cd水平，而Parsi在较低浓度（10 mg kg-1土壤中Cd浓度为56.9 mg kg-1）的秸秆中积累了更多的Cd。

The gene expression analysis revealed that most cultivars showed increased expression of TaNRAMP genes, particularly TaNRAMP2 in Cd concentration of 10 mg kg−1 soil, which facilitates Cd uptake from the soil, and TaZIP genes, such as TaZIP4 and TaZIP7, involved in transporting Cd within the plant. Notably, the expression of TaZIP1 was significantly lower in cultivars with high Cd accumulation, suggesting a potential regulatory mechanism for Cd tolerance.

基因表达分析表明，大多数品种表现出TaNRAMP基因的表达增加，特别是在10 mg kg-1土壤的Cd浓度下，TaNRAMP2的表达增加，这有助于从土壤中吸收Cd，而TaZIP基因，如TaZIP4和TaZIP7，参与了植物内Cd的运输。值得注意的是，在具有高Cd积累的品种中，TaZIP1的表达显着降低，表明Cd耐受性的潜在调节机制。

Furthermore, cultivars exhibiting higher Cd levels correlated with increased expression of stress-responsive genes, indicating a broader response to Cd stress. These findings highlight Barat’s potential for bread-making applications due to its low Cd accumulation, while Morvarid and Pishtaz which show reduced Cd content in the stra.

此外，表现出较高Cd水平的品种与应激反应基因表达的增加相关，表明对Cd胁迫的反应更广泛。这些发现突出了Barat由于其低Cd积累而在面包制作中的应用潜力，而Morvarid和Pishtaz则显示stra中Cd含量降低。

IntroductionWheat, a major cereal crop for billions of people, belongs to the Poaceae family, comprising various species, with T. aestivum L. (bread wheat) and Triticum turgidum var. durum (pasta wheat) being the most widely cultivated and recognized1. Bread wheat is an essential agricultural product, with approximately 35% of the global food supply relying on it.

引言小麦是数十亿人的主要谷物作物，属于禾本科，包括各种物种，其中小麦（面包小麦）和硬粒小麦（面食小麦）是种植最广泛和公认的1。面包小麦是一种重要的农产品，约占全球粮食供应的35%。

It serves as a significant source of carbohydrates, proteins, minerals, and vitamins for human consumption2,3,4. Furthermore, its versatility in food processing has resulted in the production of an array of products such as bread, pasta, and cereals, establishing wheat as a dietary staple for numerous populations5.The quality and safety of wheat crops are subject to a multitude of influencing factors, among which the presence of heavy metals in the soil emerges as a critical concern.

它是人类消耗的碳水化合物，蛋白质，矿物质和维生素的重要来源2,3,4。此外，它在食品加工方面的多功能性导致了面包，面食和谷物等一系列产品的生产，使小麦成为众多人群的主食5。小麦作物的质量和安全受到多种影响因素的影响，其中土壤中重金属的存在成为一个关键问题。

Heavy metals, including Cd, lead, and arsenic, have the potential to accumulate in wheat plants when present in excessive concentrations within the soil6. These metals are absorbed by the plant’s root system and may subsequently translocate to various plant tissues, including the grain, which constitutes a primary component of the human diet.

重金属，包括Cd，铅和砷，当土壤中浓度过高时，有可能在小麦植株中积累6。这些金属被植物的根系吸收，随后可能转移到各种植物组织中，包括构成人类饮食主要成分的谷物。

Accumulation of heavy metals in wheat grains raises significant health concerns, as heavy metals are known to exhibit toxicity, posing risks to human health through the consumption of contaminated grains7,8,9,10. Therefore, the execution of efficient agronomic practices, including the adoption of wheat cultivars with low accumulation rates11,12, play a crucial role in safeguarding the nutritional quality and safety of wheat-based products in the food supply chain.As a toxic and unnecessary element in humans and animals, Cd may become a serious menace through transfer from plants and food chai.

小麦籽粒中重金属的积累引起了严重的健康问题，因为已知重金属具有毒性，通过食用受污染的谷物对人类健康构成风险7,8,9,10。因此，实施有效的农艺措施，包括采用低积累率的小麦品种11,12，在保障食品供应链中小麦产品的营养质量和安全方面起着至关重要的作用。作为人类和动物中有毒且不必要的元素，Cd可能通过从植物和食物中转移而成为严重的威胁。

AFss = [Cd] in straws/[Cd] in soil.

AFss=秸秆中的[Cd]/土壤中的[Cd]。

AFgs = [Cd] in grains/[Cd] in soil.

AFgs=谷物中的[镉]/土壤中的[镉]。

TFsg = [Cd] in grains/[Cd] in straws.

TFsg=谷物中的[镉]/稻草中的[镉]。

Furthermore, variations in Cd concentrations were precisely analyzed among different wheat cultivars. The significant differences between the Cd treatments from the same cultivar were analyzed using one-way ANOVA via Duncan’s multiple range test.Gene expression assayBased on preliminary experiments, three wheat cultivars were selected for gene expression analysis due to their demonstrated highest diversity in response to Cd stress, covering the spectrum of sensitivity to Cd: Morvarid (sensitive), Pishtaz (semi-sensitive), and Barat (resistant).

此外，精确分析了不同小麦品种之间Cd浓度的变化。通过Duncan的多范围检验，使用单向ANOVA分析了来自同一品种的Cd处理之间的显着差异。基因表达测定基于初步实验，选择了三个小麦品种进行基因表达分析，因为它们对Cd胁迫的反应表现出最高的多样性，涵盖了对Cd的敏感性范围：Morvarid（敏感），Pishtaz（半敏感）和Barat（抗性）。

This selection ensured a comprehensive understanding of Cd uptake and translocation mechanisms by capturing a wide range of genetic responses. The experimental setup involved cultivating seeds in soil treated with two Cd concentrations, 5 mg kg−1 and 10 mg kg−1, chosen because they induced significant differences among the cultivars.

这种选择通过捕获广泛的遗传反应，确保了对Cd吸收和转运机制的全面了解。实验设置涉及在用两种Cd浓度（5 mg kg-1和10 mg kg-1）处理的土壤中培养种子，因为它们诱导了品种之间的显着差异。

Subsequently, samples of the blade, sheath, and root were collected at three crucial stages of development: tillering (Zadok’s growth stages 20–29), booting (Zadok’s growth stage 45), and flowering (Zadok’s growth stage 58). Then, RNA was extracted in triplicate utilizing the DENAzist column RNA isolation kit, in accordance with the manufacturer’s instructions.

随后，在发育的三个关键阶段收集叶片，鞘和根的样品：分蘖（Zadok的生长阶段20-29），孕穗（Zadok的生长阶段45）和开花（Zadok的生长阶段58）。然后，根据制造商的说明，使用DENAzist柱RNA分离试剂盒一式三份提取RNA。

The concentration and purity of the isolated RNA were evaluated through the utilization of a NanoDrop Spectrophotometer (NanoDrop Technologies) and agarose gel electrophoresis to guarantee the quality of the RNA samples. After verifying the integrity of the RNA, quantification of the RNA samples was carried out, leading to the commencement of cDNA library construction.

通过使用NanoDrop分光光度计（NanoDrop Technologies）和琼脂糖凝胶电泳来评估分离的RNA的浓度和纯度，以保证RNA样品的质量。在验证了RNA的完整性后，对RNA样品进行了定量，从而开始了cDNA文库的构建。

For the cDNA library preparation, aimed at reverse transcription quantitative polymerase chain reaction (RT-qPCR), the initial step involved treating the isolated RNA samples wi.

对于旨在逆转录定量聚合酶链反应（RT-qPCR）的cDNA文库制备，第一步涉及处理分离的RNA样品。

Data availability

数据可用性

The datasets used and/or analysed during the current study available from the corresponding author on reasonable request.

本研究中使用和/或分析的数据集可根据合理要求从通讯作者处获得。

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Mirzaghaderi, G. rtpcr: A Package for Statistical Analysis of Real-Tme PCR Data in R. https://doi.org/10.21203/rs.3.rs-4193266/v1 (2024).Download referencesFundingNo funding was received for conducting this study.Author informationAuthor notesFatemeh Beigi, Mahdiye Mortazavi and Farzaneh Najafi contributed equally to this work.Authors and AffiliationsDepartment of Biotechnology, College of Agriculture, Isfahan University of Technology, Isfahan, IranZinat Abdolmalaki, Aboozar Soorni, Fatemeh Beigi, Mahdiye Mortazavi, Farzaneh Najafi, Rahim Mehrabi & Badraldin Ebrahim Sayed-TabatabaeiDepartment of Soil Science, College of Agriculture, Isfahan University of Technology, Isfahan, IranMehran ShirvaniDepartment of Agronomy and Plant Breeding, College of Agriculture, Isfahan University of Technology, Isfahan, 84156-83111, IranMohammad Mahdi MajidiAuthorsZinat AbdolmalakiView author publicationsYou can also search for this author in.

Mirzaghaderi，G。rtpcr：用于R中真实Tme PCR数据统计分析的软件包。https://doi.org/10.21203/rs.3.rs-4193266/v1（2024年）。下载参考资金进行这项研究没有收到资金。作者信息作者注：Fatemeh Beigi，Mahdiye Mortazavi和Farzaneh Najafi对这项工作做出了同样的贡献。作者和附属机构伊斯法罕理工大学农业学院生物技术系，伊斯法罕，IranZinat Abdolmalaki，Aboozar Soorni，Fatemeh Beigi，Mahdiye Mortazavi，Farzaneh Najafi，Rahim Mehrabi和Badraldin Ebrahim Sayed Tabatabaeid伊斯法罕理工大学农业学院土壤科学系农业，伊斯法罕理工大学，伊斯法罕，84156-83111，IranMohammad Mahdi MajidiAuthorsZinat AbdolmalakiView作者出版物您也可以在中搜索这位作者。

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PubMed Google ScholarMohammad Mahdi MajidiView author publicationsYou can also search for this author in

PubMed谷歌学术评论Mahdi MajidiView作者出版物您也可以在

PubMed Google ScholarContributionsZA, FB, MM, and FN: conceptualization, investigation (responsible for most experimental work), formal analysis, validation (qRT-PCR), visualization, and writing review & editing. AS: conceptualization, formal analysis (responsible for all bioinformatic analysis), validation, visualization, supervision, and writing - original draft as well as review & editing.

PubMed Google ScholarContributionsZA，FB，MM和FN：概念化，调查（负责大多数实验工作），正式分析，验证（qRT-PCR），可视化以及写作评论和编辑。AS：概念化，形式分析（负责所有生物信息学分析），验证，可视化，监督和写作-原稿以及评论和编辑。

RM, BEST, MSh, and MMM: conceptualization, investigation; formal analysis, writing-review and editing. All authors contributed to the article and approved the submitted version.Corresponding authorCorrespondence to.

RM，BEST，MSh和MMM：概念化，调查；形式分析，写作评论和编辑。所有作者都为文章做出了贡献，并批准了提交的版本。对应作者对应。

Aboozar Soorni.Ethics declarations

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Additional informationPublisher’s noteSpringer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.Electronic supplementary materialBelow is the link to the electronic supplementary material.Supplementary Material 1Rights and permissions

。电子补充材料流是指向电子补充材料的链接。补充材料1权利和许可

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Reprints and permissionsAbout this articleCite this articleAbdolmalaki, Z., Soorni, A., Beigi, F. et al. Exploring genotypic variation and gene expression associated to cadmium accumulation in bread wheat.

转载和许可本文引用本文Abdolmalaki，Z.，Soorni，A.，Beigi，F。等人探索与面包小麦中镉积累相关的基因型变异和基因表达。

Sci Rep 14, 26505 (2024). https://doi.org/10.1038/s41598-024-78425-zDownload citationReceived: 30 July 2024Accepted: 30 October 2024Published: 03 November 2024DOI: https://doi.org/10.1038/s41598-024-78425-zShare this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy to clipboard.

《科学报告》1426505（2024）。https://doi.org/10.1038/s41598-024-78425-zDownload引文接收日期：2024年7月30日接受日期：2024年10月30日发布日期：2024年11月3日OI：https://doi.org/10.1038/s41598-024-78425-zShare本文与您共享以下链接的任何人都可以阅读此内容：获取可共享链接对不起，本文目前没有可共享的链接。复制到剪贴板。

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KeywordsBread wheatCultivarsSoilCadmiumGene expression

关键词小麦品种土壤管理基因表达