{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T16:44:04Z","timestamp":1793897044358,"version":"4.8.0"},"reference-count":120,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2023,2,3]],"date-time":"2023-02-03T00:00:00Z","timestamp":1675382400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004564","name":"Ministarstvo Prosvete, Nauke i Tehnolo\u0161kog Razvoja","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004564","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>Essential micronutrients belonging to the transition metals, such as Fe and Cu, are indispensable for plant growth and stress tolerance; however, when present in excess, they can become potentially dangerous producers of reactive oxygen species. Therefore, their homeostases must be strictly regulated. Both microelement deficiencies and elevated concentrations of heavy metals in the soil are global problems that reduce the nutritional value of crops and seriously affect human health. Silicon, a beneficial element known for its protective properties, has been reported to alleviate the symptoms of Cu toxicity and Fe deficiency stress in plants; however, we are still far from a comprehensive understanding of the underlying molecular mechanisms. Although Si-mediated mitigation of these stresses has been clearly demonstrated for some species, the effects of Si vary depending on plant species, growing conditions and experimental design. In this review, the proposed mechanistic models explaining the effect of Si are summarized and discussed. Iron and copper compete for the common metal transporters and share the same transport routes, hence, inadequate concentration of one element leads to disturbances of another. Silicon is reported to beneficially influence not only the distribution of the element supplied below or above the optimal concentration, but also the distribution of other microelements, as well as their molar ratios. The influence of Si on Cu immobilization and retention in the root, as well as Si-induced Fe remobilization from the source to the sink organs are of vital importance. The changes in cellular Cu and Fe localization are considered to play a crucial role in restoring homeostasis of these microelements. Silicon has been shown to stimulate the accumulation of metal chelators involved in both the mobilization of deficient elements and scavenging excess heavy metals. Research into the mechanisms of the ameliorative effects of Si is valuable for reducing mineral stress in plants and improving the nutritional value of crops. This review aims to provide a thorough and critical overview of the current state of knowledge in this field and to discuss discrepancies in the observed effects of Si and different views on its mode of action.<\/jats:p>","DOI":"10.3389\/fpls.2023.1039053","type":"journal-article","created":{"date-parts":[[2023,2,3]],"date-time":"2023-02-03T00:36:27Z","timestamp":1675384587000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":17,"title":["Silicon in action: Between iron scarcity and excess copper"],"prefix":"10.3389","volume":"14","author":[{"given":"Dragana","family":"Nikoli\u0107","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dragana","family":"Bosni\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jelena","family":"Samard\u017ei\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2023,2,3]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1023\/A:1016093317898","article-title":"Organic acids and fe deficiency: a review","volume":"241","author":"Abad\u00eda","year":"2002","journal-title":"Plant Soil"},{"key":"B2","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1016\/j.plaphy.2011.01.026","article-title":"Towards a knowledge-based correction of iron chlorosis","volume":"49","author":"Abad\u00eda","year":"2011","journal-title":"Plant Physiol. Biochem."},{"key":"B3","doi-asserted-by":"publisher","first-page":"767","DOI":"10.1007\/s00425-008-0869-z","article-title":"Contribution of plastocyanin isoforms to photosynthesis and copper homeostasis in arabidopsis thaliana grown at different copper regimes","volume":"229","author":"Abdel-Ghany","year":"2009","journal-title":"Planta"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1233","DOI":"10.1105\/tpc.104.030452","article-title":"Two p-type ATPases are required for copper delivery in arabidopsis thaliana chloroplasts","volume":"17","author":"Abdel-Ghany","year":"2005","journal-title":"Plant Cell"},{"key":"B5","doi-asserted-by":"publisher","first-page":"15932","DOI":"10.1074\/jbc.M801406200","article-title":"MicroRNA-mediated systemic down-regulation of copper protein expression in response to low copper availability in arabidopsis","volume":"283","author":"Abdel-Ghany","year":"2008","journal-title":"J. Biol. Chem."},{"key":"B6","doi-asserted-by":"publisher","first-page":"1779","DOI":"10.3390\/plants9121779","article-title":"The regulatory role of silicon in mitigating plant nutritional stresses","volume":"9","author":"Ali","year":"2020","journal-title":"Plants"},{"key":"B7","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1007\/s11738-016-2279-3","article-title":"Physiological and biochemical mechanisms of silicon-induced copper stress tolerance in cotton (Gossypium hirsutum l.)","volume":"38","author":"Ali","year":"2016","journal-title":"Acta Physiol. Plant"},{"key":"B8","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/j.plantsci.2006.03.005","article-title":"Phenolics metabolism and lignin synthesis in root suspension cultures of panax ginseng in response to copper stress","volume":"171","author":"Ali","year":"2006","journal-title":"Plant Sci."},{"key":"B9","doi-asserted-by":"publisher","first-page":"1331","DOI":"10.1093\/jexbot\/53.372.1331","article-title":"Role of superoxide dismutases (SODs) in controlling oxidative stress in plants","volume":"53","author":"Alscher","year":"2002","journal-title":"J. Exp. Bot."},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2015.00192","article-title":"Jacks of metal\/metalloid chelation trade in plants\u2013an overview","volume":"6","author":"Anjum","year":"2015","journal-title":"Front. Plant Sci."},{"key":"B11","doi-asserted-by":"publisher","first-page":"12081","DOI":"10.1073\/pnas.0804175105","article-title":"The leaf ionome as a multivariable system to detect a plant\u2019s physiological status","volume":"105","author":"Baxter","year":"2008","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B12","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-020-61718-4","article-title":"Silicon reduces the iron uptake in rice and induces iron homeostasis related genes","volume":"10","author":"Becker","year":"2020","journal-title":"Sci. Rep."},{"key":"B13","doi-asserted-by":"publisher","first-page":"738","DOI":"10.1105\/tpc.111.090431","article-title":"Transcriptome sequencing identifies SPL7-regulated copper acquisition genes FRO4\/FRO5 and the copper dependence of iron homeostasis in arabidopsis","volume":"24","author":"Bernal","year":"2012","journal-title":"Plant Cell"},{"key":"B14","doi-asserted-by":"publisher","first-page":"596","DOI":"10.1104\/pp.78.3.596","article-title":"Free space iron pools in roots: Generation and mobilization","volume":"78","author":"Bienfait","year":"1985","journal-title":"Plant Physiol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"364","DOI":"10.1016\/j.jplph.2018.10.017","article-title":"Silicon ameliorates iron deficiency of cucumber in a pH-dependent manner","volume":"231","author":"Bityutskii","year":"2018","journal-title":"J. Plant Physiol."},{"key":"B16","first-page":"45","article-title":"Vliyanie kremniya na proyavlenie khlorosa rasteniy v usloviyakh defizita zhelesa i marganza","volume":"2","author":"Bityutskii","year":"2010","journal-title":"Agrokhimiia"},{"key":"B17","doi-asserted-by":"publisher","first-page":"554","DOI":"10.3390\/plants8120554","article-title":"Silicon and iron differently alleviate copper toxicity in cucumber leaves","volume":"8","author":"Bosni\u0107","year":"2019","journal-title":"Plants"},{"key":"B18","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1007\/s11104-019-04151-5","article-title":"Silicon alleviates copper (Cu) toxicity in cucumber by increased Cu-binding capacity","volume":"441","author":"Bosni\u0107","year":"2019","journal-title":"Plant Soil"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1679","DOI":"10.1111\/pce.14000","article-title":"FIT and bHLH ib transcription factors modulate iron and copper crosstalk in arabidopsis","volume":"44","author":"Cai","year":"2021","journal-title":"Plant Cell Environ."},{"key":"B20","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1016\/j.plaphy.2018.01.033","article-title":"Silicon induced fe deficiency affects fe, Mn, Cu and zn distribution in rice (Oryza sativa l.) growth in calcareous conditions","volume":"125","author":"Carrasco-Gil","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"B21","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/j.plaphy.2021.03.045","article-title":"Methods for metal chelation in plant homeostasis: Review","volume":"163","author":"Carrillo","year":"2021","journal-title":"Plant Phys. Biochem."},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.01065","article-title":"Interplay between silicon and iron signaling pathways to regulate silicon transporter Lsi1 expression in rice","volume":"11","author":"Chaiwong","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B23","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.envpol.2013.12.024","article-title":"Evidence of sulfur-bound reduced copper in bamboo exposed to high silicon and copper concentrations","volume":"187","author":"Collin","year":"2014","journal-title":"Environ. pollut."},{"key":"B24","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1111\/nph.15343","article-title":"The controversies of silicon\u2019s role in plant biology","volume":"221","author":"Coskun","year":"2019","journal-title":"New Phytol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1146\/annurev.arplant.54.031902.135018","article-title":"Iron transport and signaling in plants","volume":"54","author":"Curie","year":"2003","journal-title":"Annu. Rev. Plant Biol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/aob\/mcn207","article-title":"Metal movement within the plant: contribution of nicotianamine and yellow stripe 1-like transporters","volume":"103","author":"Curie","year":"2009","journal-title":"Ann. Bot."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1259","DOI":"10.1093\/jxb\/eru004","article-title":"Stress-induced electrolyte leakage: the role of k+-permeable channels and involvement in programmed cell death and metabolic adjustment","volume":"65","author":"Demidchik","year":"2014","journal-title":"J. Exp. Bot."},{"key":"B28","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1016\/S0176-1617(89)80171-3","article-title":"Copper-induced damage to the permeability barrier in roots of silene cucubalus","volume":"135","author":"De Vos","year":"1989","journal-title":"J. Plant Physiol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1111\/j.1365-313X.2004.02128.x","article-title":"Arabidopsis yellow stripe-Like2 (YSL2): A metal-regulated gene encoding a plasma membrane transporter of nicotianamine-metal complexes","volume":"39","author":"DiDonato","year":"2004","journal-title":"Plant J."},{"key":"B30","doi-asserted-by":"publisher","first-page":"1085","DOI":"10.1105\/tpc.7.7.1085","article-title":"Stress-induced phenylpropanoid metabolism","volume":"7","author":"Dixon","year":"1995","journal-title":"Plant Cell"},{"key":"B31","doi-asserted-by":"publisher","first-page":"2411","DOI":"10.1093\/jxb\/err359","article-title":"Silicon ameliorates manganese toxicity in cucumber by decreasing hydroxyl radical accumulation in the leaf apoplast","volume":"63","author":"Dragi\u0161i\u0107 Maksimovi\u0107","year":"2012","journal-title":"J. Exp. Bot."},{"key":"B32","doi-asserted-by":"publisher","first-page":"4732","DOI":"10.3390\/su12114732","article-title":"Silicon alleviates copper toxicity in flax plants by up-regulating antioxidant defense and secondary metabolites and decreasing oxidative damage","volume":"12","author":"El-Beltagi","year":"2020","journal-title":"Sustainability"},{"key":"B33","doi-asserted-by":"publisher","first-page":"2001","DOI":"10.1093\/jxb\/erq392","article-title":"Silicon enhances suberization and lignification in roots of rice (Oryza sativa)","volume":"62","author":"Fleck","year":"2011","journal-title":"J. Exp. Bot."},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0138555","article-title":"Silicon promotes exodermal casparian band formation in Si-accumulating and Si-excluding species by forming phenol complexes","volume":"10","author":"Fleck","year":"2015","journal-title":"PLoS One"},{"key":"B35","doi-asserted-by":"publisher","first-page":"864","DOI":"10.1080\/01904167.2019.1589508","article-title":"Silicon alleviates long-term copper toxicity and influences gene expression in nicotiana tabacum","volume":"42","author":"Flora","year":"2019","journal-title":"J. Plant Nutr."},{"key":"B36","doi-asserted-by":"publisher","first-page":"329","DOI":"10.21273\/JASHS.136.5.329","article-title":"Silicon differentially influences copper toxicity response in silicon-accumulator and non-accumulator species","volume":"136","author":"Frantz","year":"2011","journal-title":"J. Am. Soc Hortic. Sci."},{"key":"B37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2013\/436517","article-title":"Lignin: characterization of a multifaceted crop component","volume":"2013","author":"Frei","year":"2013","journal-title":"Sci. World J."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.00006","article-title":"The transcriptional control of iron homeostasis in plants: a tale of bHLH transcription factors","volume":"10","author":"Gao","year":"2019","journal-title":"Front. Plant Sci."},{"key":"B39","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1016\/j.plaphy.2021.08.011","article-title":"Silicon and nitric oxide interplay alleviates copper induced toxicity in mung bean seedlings","volume":"167","author":"Gaur","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2015.00716","article-title":"Local and systemic signaling of iron status and its interactions with homeostasis of other essential elements","volume":"6","author":"Gayomba","year":"2015","journal-title":"Front. Plant Sci."},{"key":"B41","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/j.plaphy.2013.06.007","article-title":"Effect of silicon addition on soybean (Glycine max) and cucumber (Cucumis sativus) plants grown under iron deficiency","volume":"70","author":"Gonzalo","year":"2013","journal-title":"Plant Physiol. Biochem."},{"key":"B42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/jexbot\/53.366.1","article-title":"Cellular mechanisms for heavy metal detoxification and tolerance","volume":"53","author":"Hall","year":"2002","journal-title":"J. Exp. Bot."},{"key":"B43","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1104\/pp.106.077073","article-title":"Reactive species and antioxidants. redox biology is a fundamental theme of aerobic life","volume":"141","author":"Halliwell","year":"2006","journal-title":"Plant Physiol."},{"key":"B44","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1042\/bj2190001","article-title":"Oxygen toxicity, oxygen radicals, transition metals and disease","volume":"219","author":"Halliwell","year":"1984","journal-title":"Biochem. J."},{"key":"B45","doi-asserted-by":"publisher","first-page":"2136","DOI":"10.1093\/jxb\/eraa483","article-title":"Coordinated homeostasis of essential mineral nutrients: A focus on iron","volume":"72","author":"Hanikenne","year":"2021","journal-title":"J. Exp. Bot."},{"key":"B46","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1007\/s007750050297","article-title":"Copper chaperones: function, structure and copper-binding properties","volume":"4","author":"Harrison","year":"1999","journal-title":"JBIC J. Biol. Inorg. Chem."},{"key":"B47","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1111\/j.1469-8137.2007.02051.x","article-title":"Transporters of ligands for essential metal ions in plants","volume":"174","author":"Haydon","year":"2007","journal-title":"New Phytol."},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.580552","article-title":"Root silicon addition induces fe deficiency in cucumber plants, but facilitates their recovery after fe resupply. A comparison with Si foliar sprays","volume":"11","author":"Hern\u00e1ndez-Apaolaza","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2017.00671","article-title":"An ABC transporter is involved in the silicon-induced formation of casparian bands in the exodermis of rice","volume":"8","author":"Hinrichs","year":"2017","journal-title":"Front. Plant Sci."},{"key":"B50","volume-title":"The chemistry of silica : Solubility, polymerization, colloid and surface properties, and biochemistry","author":"Iler","year":"1979"},{"key":"B51","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1093\/jxb\/ern286","article-title":"Nicotianamine and histidine\/proline are, respectively, the most important copper chelators in xylem sap of brassica carinata under conditions of copper deficiency and excess","volume":"60","author":"Irtelli","year":"2008","journal-title":"J. Exp. Bot."},{"key":"B52","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1080\/00380768.1990.10416911","article-title":"Copper binding by the root cell walls of Italian ryegrass and red clover","volume":"36","author":"Iwasaki","year":"1990","journal-title":"Soil Sci. Plant Nutr."},{"key":"B53","volume-title":"Trace elements in soils and plants","author":"Kabata-Pendias","year":"2001"},{"key":"B54","doi-asserted-by":"publisher","first-page":"1824","DOI":"10.1039\/C8MT00287H","article-title":"New aspects of iron\u2013copper crosstalk uncovered by transcriptomic characterization of col-0 and the copper uptake mutant spl7 in arabidopsis thaliana","volume":"10","author":"Kastoori Ramamurthy","year":"2018","journal-title":"Metallomics"},{"key":"B55","doi-asserted-by":"publisher","first-page":"847","DOI":"10.1007\/s00425-014-2220-1","article-title":"Effect of silicon on wheat seedlings (Triticum turgidum l.) grown in hydroponics and exposed to 0 to 30 \u00b5M Cu","volume":"241","author":"Keller","year":"2015","journal-title":"Planta"},{"key":"B56","doi-asserted-by":"publisher","first-page":"112510","DOI":"10.1016\/j.ecoenv.2021.112510","article-title":"Effects of silicon on heavy metal uptake at the soil-plant interphase: A review","volume":"222","author":"Khan","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"B57","doi-asserted-by":"publisher","first-page":"2263","DOI":"10.1111\/pce.13192","article-title":"Changes in iron availability in arabidopsis are rapidly sensed in the leaf vasculature and impaired sensing leads to opposite transcriptional programs in leaves and roots","volume":"41","author":"Khan","year":"2018","journal-title":"Plant Cell Environ."},{"key":"B58","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1016\/j.jplph.2010.09.009","article-title":"Soluble silicon modulates expression of arabidopsis thaliana genes involved in copper stress","volume":"168","author":"Khandekar","year":"2011","journal-title":"J. Plant Physiol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1186\/1471-2229-14-13","article-title":"Silicon mitigates heavy metal stress by regulating p-type heavy metal ATPases, oryza sativa low silicon genes, and endogenous phytohormones","volume":"14","author":"Kim","year":"2014","journal-title":"BMC Plant Biol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1111\/j.1365-313X.2011.04772.x","article-title":"The rice transcription factor IDEF1 directly binds to iron and other divalent metals for sensing cellular iron status","volume":"69","author":"Kobayashi","year":"2012","journal-title":"Plant J."},{"key":"B61","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1007\/s00299-007-0490-9","article-title":"Dynamics of phenolic acids and lignin accumulation in metal-treated matricaria chamomilla roots","volume":"27","author":"Kov\u00e1\u010dik","year":"2008","journal-title":"Plant Cell Rep."},{"key":"B62","first-page":"215","article-title":"Functions and homeostasis of zinc, copper, and nickel in plants","volume-title":"Molecular biology of metal homeostasis and detoxification: From microbes to man","author":"Kr\u00e4mer","year":"2006"},{"key":"B63","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1007\/s11738-010-0581-z","article-title":"The cell wall in plant cell response to trace metals: polysaccharide remodeling and its role in defense strategy","volume":"33","author":"Krzes\u0142owska","year":"2011","journal-title":"Acta Physiol. Plant"},{"key":"B64","doi-asserted-by":"publisher","first-page":"6697","DOI":"10.1007\/s12633-021-01376-x","article-title":"Silicon (Si) alleviates iron deficiency effects in sea barley (Hordeum marinum) by enhancing iron accumulation and photosystem activities","volume":"14","author":"Ksiaa","year":"2022","journal-title":"Silicon"},{"key":"B65","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1104\/pp.109.139717","article-title":"Complexation and toxicity of copper in higher plants. i. characterization of copper accumulation, speciation, and toxicity in crassula helmsii as a new copper accumulator","volume":"151","author":"K\u00fcpper","year":"2009","journal-title":"Plant Physiol."},{"key":"B66","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1104\/pp.15.01589","article-title":"Zinc finger of arabidopsis thaliana12 (ZAT12) interacts with FER-like iron deficiency-induced transcription factor (FIT) linking iron deficiency and oxidative stress responses","volume":"170","author":"Le","year":"2016","journal-title":"Plant Physiol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1023\/A:1004843505053","article-title":"Copper uptake and translocation in chicory (Cichorium intybus l. cv grasslands puna) and tomato (Lycopersicon esculentum mill. cv rondy) plants grown in NFT system. II. the role of nicotianamine and histidine in xylem sap copper transport","volume":"223","author":"Liao","year":"2000","journal-title":"Plant Soil"},{"key":"B68","doi-asserted-by":"publisher","first-page":"670","DOI":"10.21273\/JASHS.133.5.670","article-title":"Alleviation of copper toxicity in arabidopsis thaliana by silicon addition to hydroponic solutions","volume":"133","author":"Li","year":"2008","journal-title":"J. Am. Soc Hortic. Sci."},{"key":"B69","doi-asserted-by":"publisher","first-page":"163","DOI":"10.4067\/S0718-58392017000200163","article-title":"Application of silicon fertilizer affects nutritional quality of rice","volume":"77","author":"Liu","year":"2017","journal-title":"Chil. J. Agric. Res."},{"key":"B70","volume-title":"Mineral nutrition of higher plants","author":"Marschner","year":"1995"},{"key":"B71","doi-asserted-by":"publisher","first-page":"296","DOI":"10.1016\/j.plaphy.2021.04.012","article-title":"Rice responses to silicon addition at different fe status and growth pH. evaluation of ploidy changes","volume":"163","author":"Mart\u00edn-Esquinas","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"B72","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.jplph.2015.03.001","article-title":"Assessment of the role of silicon in the Cu-tolerance of the C4 grass spartina densiflora","volume":"178","author":"Mateos-Naranjo","year":"2015","journal-title":"J. Plant Physiol."},{"key":"B73","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1017\/S1368980008002401","article-title":"Worldwide prevalence of anaemia, WHO vitamin and mineral nutrition information system 1993-2005","volume":"12","author":"McLean","year":"2009","journal-title":"Public Health Nutr."},{"key":"B74","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1007\/s11104-008-9642-0","article-title":"Copper phytotoxicity affects root elongation and iron nutrition in durum wheat (Triticum turgidum durum l.)","volume":"310","author":"Michaud","year":"2008","journal-title":"Plant Soil"},{"key":"B75","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1104\/pp.109.144675","article-title":"Complexation and toxicity of copper in higher plants. II. Different mechanisms for copper versus cadmium detoxification in the copper-sensitive cadmium\/zinc hyperaccumulator thlaspi caerulescens (Ganges ecotype)","volume":"151","author":"Mijovilovich","year":"2009","journal-title":"Plant Physiol."},{"key":"B76","doi-asserted-by":"publisher","first-page":"4553","DOI":"10.1021\/cr900112r","article-title":"Iron uptake and transport in plants: the good, the bad, and the ionome","volume":"109","author":"Morrissey","year":"2009","journal-title":"Chem. Rev."},{"key":"B77","doi-asserted-by":"publisher","first-page":"1375","DOI":"10.1104\/pp.121.4.1375","article-title":"Early copper-induced leakage of k+ from arabidopsis seedlings is mediated by ion channels and coupled to citrate efflux","volume":"121","author":"Murphy","year":"1999","journal-title":"Plant Physiol."},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.00416","article-title":"Silicon alleviates iron deficiency in barley by enhancing expression of strategy II genes and metal redistribution","volume":"10","author":"Nikolic","year":"2019","journal-title":"Front. Plant Sci."},{"key":"B79","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1093\/pcp\/pcr188","article-title":"Identification of zn-nicotianamine and fe-2\u2032-deoxymugineic acid in the phloem sap from rice plants (Oryza sativa l.)","volume":"53","author":"Nishiyama","year":"2012","journal-title":"Plant Cell Physiol."},{"key":"B80","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1023\/A:1001009100026","article-title":"Silicon and copper interaction in the growth of spring wheat seedlings","volume":"39","author":"Nowakowski","year":"1997","journal-title":"Biol. Plant"},{"key":"B81","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1039\/c0em00549e","article-title":"Effects of silicon on copper toxicity in Erica andevalensis cabezudo and Rivera: A potential species to remediate contaminated soils","volume":"13","author":"Oliva","year":"2011","journal-title":"J. Environ. Monit."},{"key":"B82","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/0098-8472(94)90035-3","article-title":"Copper-induced changes on growth, metal content and photosynthetic function of alyssum montanum l. plants","volume":"34","author":"Ouzounidou","year":"1994","journal-title":"Environ. Exp. Bot."},{"key":"B83","doi-asserted-by":"publisher","first-page":"2089","DOI":"10.1080\/01904169809365546","article-title":"Amelioration of copper toxicity by iron on spinach physiology","volume":"21","author":"Ouzounidou","year":"1998","journal-title":"J. Plant Nutr."},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.697592","article-title":"Interactions of silicon with essential and beneficial elements in plants","volume":"12","author":"Pavlovic","year":"2021","journal-title":"Front. Plant Sci."},{"key":"B85","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1093\/aob\/mcw105","article-title":"Silicon enhances leaf remobilization of iron in cucumber under limited iron conditions","volume":"118","author":"Pavlovic","year":"2016","journal-title":"Ann. Bot."},{"key":"B86","doi-asserted-by":"publisher","first-page":"1096","DOI":"10.1111\/nph.12213","article-title":"Silicon alleviates iron deficiency in cucumber by promoting mobilization of iron in the root apoplast","volume":"198","author":"Pavlovic","year":"2013","journal-title":"New Phytol."},{"key":"B87","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1093\/jxb\/47.1.41","article-title":"Translocation of copper and other micronutrients in tomato plants (Lycopersicon esculentum mill.): Nicotianamine-stimulated copper transport in the xylem","volume":"47","author":"Pich","year":"1996","journal-title":"J. Exp. Bot."},{"key":"B88","doi-asserted-by":"publisher","first-page":"905","DOI":"10.1007\/s00709-021-01708-z","article-title":"Silicon and nitric oxide synergistically modulate the production of essential oil and rosmarinic acid in salvia officinalis under Cu stress","volume":"259","author":"Pirooz","year":"2022","journal-title":"Protoplasma"},{"key":"B89","doi-asserted-by":"publisher","first-page":"26021","DOI":"10.1074\/jbc.M202547200","article-title":"Biochemical and genetic analyses of yeast and human high affinity copper transporters suggest a conserved mechanism for copper uptake","volume":"277","author":"Puig","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B90","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.stress.2021.100008","article-title":"Iron homeostasis in plants and its crosstalk with copper, zinc, and manganese","volume":"100008","author":"Rai","year":"2021","journal-title":"Plant Stress"},{"key":"B91","doi-asserted-by":"publisher","first-page":"1553","DOI":"10.1002\/rcm.3523","article-title":"Formation of metal-nicotianamine complexes as affected by pH, ligand exchange with citrate and metal exchange. A study by electrospray ionization time-of-flight mass spectrometry","volume":"22","author":"Rell\u00e1n-\u00c1lvarez","year":"2008","journal-title":"Rapid Commun. Mass Spectrom. Int. J. Devoted to Rapid Dissem. Up-to-the-Minute Res. Mass Spectrom."},{"key":"B92","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1093\/pcp\/pcp170","article-title":"Identification of a tri-iron (III), tri-citrate complex in the xylem sap of iron-deficient tomato resupplied with iron: new insights into plant iron long-distance transport","volume":"51","author":"Rell\u00e1n-\u00c1lvarez","year":"2010","journal-title":"Plant Cell Physiol."},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.986991","article-title":"Effect of nano-silicon on the regulation of ascorbate-glutathione contents, antioxidant defense system and growth of copper stressed wheat (Triticum aestivum l.) seedlings","volume":"13","author":"Riaz","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B94","doi-asserted-by":"publisher","first-page":"694","DOI":"10.1038\/17800","article-title":"A ferric-chelate reductase for iron uptake from soils","volume":"397","author":"Robinson","year":"1999","journal-title":"Nature"},{"key":"B95","doi-asserted-by":"publisher","first-page":"2173","DOI":"10.1093\/pcp\/pcr149","article-title":"Characterization of flavins in roots of fe-deficient strategy I plants, with a focus on medicago truncatula","volume":"52","author":"Rodr\u00edguez-Celma","year":"2011","journal-title":"Plant Cell Physiol."},{"key":"B96","first-page":"155","article-title":"Mobilization of iron in the rhizosphere of different plant species","author":"R\u00f6mheld","year":"1986","journal-title":"Adv. Plant Nutr."},{"key":"B97","doi-asserted-by":"publisher","first-page":"1","DOI":"10.7831\/ras.3.1","article-title":"Role of iron in plant growth and metabolism","volume":"3","author":"Rout","year":"2015","journal-title":"Rev. Agric. Sci."},{"key":"B98","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1111\/nph.12276","article-title":"Copper speciation and isotopic fractionation in plants: Uptake and translocation mechanisms","volume":"199","author":"Ryan","year":"2013","journal-title":"New Phytol."},{"key":"B99","doi-asserted-by":"publisher","first-page":"15348","DOI":"10.1074\/jbc.M313321200","article-title":"The arabidopsis copper transporter COPT1 functions in root elongation and pollen development","volume":"279","author":"Sancen\u00f3n","year":"2004","journal-title":"J. Biol. Chem."},{"key":"B100","doi-asserted-by":"publisher","first-page":"1072","DOI":"10.1111\/j.1469-8137.2009.02908.x","article-title":"Dissecting iron deficiency-induced proton extrusion in arabidopsis roots","volume":"183","author":"Santi","year":"2009","journal-title":"New Phytol."},{"key":"B101","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1046\/j.1469-8137.1997.00691.x","article-title":"Physiological effects of copper on iron acquisition processes in plantago","volume":"135","author":"Schmidt","year":"1997","journal-title":"New Phytol."},{"key":"B102","doi-asserted-by":"publisher","first-page":"1608","DOI":"10.1073\/pnas.70.5.1608","article-title":"A bound form of silicon in glycosaminoglycans and polyuronides","volume":"70","author":"Schwarz","year":"1973","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B103","doi-asserted-by":"publisher","first-page":"711","DOI":"10.1093\/jxb\/erj073","article-title":"The significance of amino acids and amino acid-derived molecules in plant responses and adaptation to heavy metal stress","volume":"57","author":"Sharma","year":"2006","journal-title":"J. Exp. Bot."},{"key":"B104","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2012\/217037","article-title":"Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions","volume":"2012","author":"Sharma","year":"2012","journal-title":"J. Bot."},{"key":"B105","doi-asserted-by":"publisher","first-page":"1333","DOI":"10.1105\/tpc.011817","article-title":"PAA1, a p-type ATPase of arabidopsis, functions in copper transport in chloroplasts","volume":"15","author":"Shikanai","year":"2003","journal-title":"Plant Cell"},{"key":"B106","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1093\/aob\/mcx060","article-title":"Formation of silica aggregates in sorghum root endodermis is predetermined by cell wall architecture and development","volume":"120","author":"Soukup","year":"2017","journal-title":"Ann. Bot."},{"key":"B107","doi-asserted-by":"publisher","first-page":"6807","DOI":"10.1093\/jxb\/erz387","article-title":"Formation of root silica aggregates in sorghum is an active process of the endodermis","volume":"71","author":"Soukup","year":"2019","journal-title":"J. Exp. Bot"},{"key":"B108","doi-asserted-by":"publisher","first-page":"945","DOI":"10.1007\/s10534-016-9966-9","article-title":"Binding of transition metals to monosilicic acid in aqueous and xylem (Cucumis sativus l.) solutions: a low-T electron paramagnetic resonance study","volume":"29","author":"Stevic","year":"2016","journal-title":"Biometals"},{"key":"B109","doi-asserted-by":"publisher","first-page":"1263","DOI":"10.1105\/tpc.010256","article-title":"Role of nicotianamine in the intracellular delivery of metals and plant reproductive development","volume":"15","author":"Takahashi","year":"2003","journal-title":"Plant Cell"},{"key":"B110","doi-asserted-by":"publisher","first-page":"1311","DOI":"10.1007\/s42729-020-00214-0","article-title":"Silicon increases leaf chlorophyll content and iron nutritional efficiency and reduces iron deficiency in sorghum plants","volume":"20","author":"Teixeira","year":"2020","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"B111","doi-asserted-by":"publisher","first-page":"1789","DOI":"10.1039\/C4MT00111G","article-title":"Roles for root iron plaque in sequestration and uptake of heavy metals and metalloids in aquatic and wetland plants","volume":"6","author":"Tripathi","year":"2014","journal-title":"Metallomics"},{"key":"B112","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1093\/pcp\/pcn192","article-title":"52Fe translocation in barley as monitored by a positron-emitting tracer imaging system (PETIS): Evidence for the direct translocation of fe from roots to young leaves via phloem","volume":"50","author":"Tsukamoto","year":"2009","journal-title":"Plant Cell Physiol."},{"key":"B113","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1093\/aob\/mcs039","article-title":"Silicon modifies root anatomy, and uptake and subcellular distribution of cadmium in young maize plants","volume":"110","author":"Vacul\u00edk","year":"2012","journal-title":"Ann. Bot."},{"key":"B114","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1111\/j.1365-3040.1990.tb01304.x","article-title":"Effects of metals on enzyme activity in plants","volume":"13","author":"Van Assche","year":"1990","journal-title":"Plant Cell Environ."},{"key":"B115","doi-asserted-by":"publisher","first-page":"1223","DOI":"10.1105\/tpc.001388","article-title":"IRT1, an arabidopsis transporter essential for iron uptake from the soil and for plant growth","volume":"14","author":"Vert","year":"2002","journal-title":"Plant Cell"},{"key":"B116","doi-asserted-by":"publisher","first-page":"100045","DOI":"10.1016\/j.stress.2021.100045","article-title":"Silicon improves photosynthetic activity and induces antioxidant enzyme activity in Tanzania Guinea grass under copper toxicity","volume":"3","author":"Vieira-Filho","year":"2022","journal-title":"Plant Stress"},{"key":"B117","doi-asserted-by":"publisher","first-page":"3762","DOI":"10.1104\/pp.104.045005","article-title":"Apoplastic binding of aluminum is involved in silicon-induced amelioration of aluminum toxicity in maize","volume":"136","author":"Wang","year":"2004","journal-title":"Plant Physiol."},{"key":"B118","doi-asserted-by":"publisher","first-page":"47644","DOI":"10.1074\/jbc.M309338200","article-title":"Expression profiles of arabidopsis thaliana in mineral deficiencies reveal novel transporters involved in metal homeostasis","volume":"278","author":"Wintz","year":"2003","journal-title":"J. Biol. Chem."},{"key":"B119","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1071\/FP08288","article-title":"Copper in plants: acquisition, transport and interactions","volume":"36","author":"Yruela","year":"2009","journal-title":"Funct. Plant Biol."},{"key":"B120","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.plantsci.2017.12.006","article-title":"Ectopic expression of SsPETE2, a plastocyanin from suaeda salsa, improves plant tolerance to oxidative stress","volume":"268","author":"Zhou","year":"2018","journal-title":"Plant Sci."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2023.1039053\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,2,3]],"date-time":"2023-02-03T00:36:37Z","timestamp":1675384597000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2023.1039053\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,2,3]]},"references-count":120,"alternative-id":["10.3389\/fpls.2023.1039053"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2023.1039053","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,2,3]]},"article-number":"1039053"}}