Constitutive Overexpression of the OsNAS Gene Family Reveals Single-Gene Strategies for Effective Iron- and Zinc-Biofortification of Rice Endosperm
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{"title"=>"Constitutive overexpression of the OsNAS gene family reveals single-gene strategies for effective iron- and zinc-biofortification of rice endosperm", "type"=>"journal", "authors"=>[{"first_name"=>"Alexander A.T.", "last_name"=>"Johnson", "scopus_author_id"=>"7410014257"}, {"first_name"=>"Bianca", "last_name"=>"Kyriacou", "scopus_author_id"=>"51261265100"}, {"first_name"=>"Damien L.", "last_name"=>"Callahan", "scopus_author_id"=>"10440617500"}, {"first_name"=>"Lorraine", "last_name"=>"Carruthers", "scopus_author_id"=>"51260960700"}, {"first_name"=>"James", "last_name"=>"Stangoulis", "scopus_author_id"=>"6602971687"}, {"first_name"=>"Enzo", "last_name"=>"Lombi", "scopus_author_id"=>"7003734476"}, {"first_name"=>"Mark", "last_name"=>"Tester", "scopus_author_id"=>"7003273484"}], "year"=>2011, "source"=>"PLoS ONE", "identifiers"=>{"doi"=>"10.1371/journal.pone.0024476", "sgr"=>"80052449864", "issn"=>"19326203", "pui"=>"362504691", "isbn"=>"1932-6203", "pmid"=>"21915334", "scopus"=>"2-s2.0-80052449864"}, "id"=>"8c1d9f03-5312-34d2-ac76-2dfb94750829", "abstract"=>"BACKGROUND: Rice is the primary source of food for billions of people in developing countries, yet the commonly consumed polished grain contains insufficient levels of the key micronutrients iron (Fe), zinc (Zn) and Vitamin A to meet daily dietary requirements. Experts estimate that a rice-based diet should contain 14.5 µg g(-1) Fe in endosperm, the main constituent of polished grain, but breeding programs have failed to achieve even half of that value. Transgenic efforts to increase the Fe concentration of rice endosperm include expression of ferritin genes, nicotianamine synthase genes (NAS) or ferritin in conjunction with NAS genes, with results ranging from two-fold increases via single-gene approaches to six-fold increases via multi-gene approaches, yet no approach has reported 14.5 µg g(-1) Fe in endosperm.\\n\\nMETHODOLOGY/PRINCIPAL FINDINGS: Three populations of rice were generated to constitutively overexpress OsNAS1, OsNAS2 or OsNAS3, respectively. Nicotianamine, Fe and Zn concentrations were significantly increased in unpolished grain of all three of the overexpression populations, relative to controls, with the highest concentrations in the OsNAS2 and OsNAS3 overexpression populations. Selected lines from each population had at least 10 µg g(-1) Fe in polished grain and two OsNAS2 overexpression lines had 14 and 19 µg g(-1) Fe in polished grain, representing up to four-fold increases in Fe concentration. Two-fold increases of Zn concentration were also observed in the OsNAS2 population. Synchrotron X-ray fluorescence spectroscopy demonstrated that OsNAS2 overexpression leads to significant enrichment of Fe and Zn in phosphorus-free regions of rice endosperm.\\n\\nCONCLUSIONS: The OsNAS genes, particularly OsNAS2, show enormous potential for Fe and Zn biofortification of rice endosperm. The results demonstrate that rice cultivars overexpressing single rice OsNAS genes could provide a sustainable and genetically simple solution to Fe and Zn deficiency disorders affecting billions of people throughout the world.", "link"=>"http://www.mendeley.com/research/constitutive-overexpression-osnas-gene-family-reveals-singlegene-strategies-effective-iron-zincbiofo", "reader_count"=>132, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>43, "Student > Doctoral Student"=>6, "Student > Ph. D. Student"=>30, "Student > Postgraduate"=>5, "Student > Master"=>21, "Other"=>3, "Student > Bachelor"=>13, "Lecturer"=>2, "Professor"=>5}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>43, "Student > Doctoral Student"=>6, "Student > Ph. D. Student"=>30, "Student > Postgraduate"=>5, "Student > Master"=>21, "Other"=>3, "Student > Bachelor"=>13, "Lecturer"=>2, "Professor"=>5}, "reader_count_by_subject_area"=>{"Unspecified"=>7, "Agricultural and Biological Sciences"=>95, "Chemical Engineering"=>1, "Chemistry"=>2, "Earth and Planetary Sciences"=>1, "Economics, Econometrics and Finance"=>1, "Engineering"=>1, "Environmental Science"=>6, "Biochemistry, Genetics and Molecular Biology"=>12, "Medicine and Dentistry"=>2, "Pharmacology, Toxicology and Pharmaceutical Science"=>1, "Sports and Recreations"=>1, "Physics and Astronomy"=>1, "Social Sciences"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>2}, "Social Sciences"=>{"Social Sciences"=>1}, "Sports and Recreations"=>{"Sports and Recreations"=>1}, "Physics and Astronomy"=>{"Physics and Astronomy"=>1}, "Unspecified"=>{"Unspecified"=>7}, "Environmental Science"=>{"Environmental Science"=>6}, "Pharmacology, Toxicology and Pharmaceutical Science"=>{"Pharmacology, Toxicology and Pharmaceutical Science"=>1}, "Chemical Engineering"=>{"Chemical Engineering"=>1}, "Engineering"=>{"Engineering"=>1}, "Chemistry"=>{"Chemistry"=>2}, "Earth and Planetary Sciences"=>{"Earth and Planetary Sciences"=>1}, "Economics, Econometrics and Finance"=>{"Economics, Econometrics and Finance"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>95}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>12}}, "reader_count_by_country"=>{"Netherlands"=>1, "United States"=>3, "Philippines"=>2, "Japan"=>1, "Brazil"=>1, "Italy"=>1, "Malaysia"=>1, "France"=>1, "India"=>2, "Indonesia"=>1}, "group_count"=>2}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/372901"], "description"=>"<div><h3>Background</h3><p>Rice is the primary source of food for billions of people in developing countries, yet the commonly consumed polished grain contains insufficient levels of the key micronutrients iron (Fe), zinc (Zn) and Vitamin A to meet daily dietary requirements. Experts estimate that a rice-based diet should contain 14.5 µg g<sup>−1</sup> Fe in endosperm, the main constituent of polished grain, but breeding programs have failed to achieve even half of that value. Transgenic efforts to increase the Fe concentration of rice endosperm include expression of ferritin genes, nicotianamine synthase genes (NAS) or ferritin in conjunction with NAS genes, with results ranging from two-fold increases via single-gene approaches to six-fold increases via multi-gene approaches, yet no approach has reported 14.5 µg g<sup>−1</sup> Fe in endosperm.</p> <h3>Methodology/Principal Findings</h3><p>Three populations of rice were generated to constitutively overexpress <em>OsNAS1</em>, <em>OsNAS2</em> or <em>OsNAS3</em>, respectively. Nicotianamine, Fe and Zn concentrations were significantly increased in unpolished grain of all three of the overexpression populations, relative to controls, with the highest concentrations in the <em>OsNAS2</em> and <em>OsNAS3</em> overexpression populations. Selected lines from each population had at least 10 µg g<sup>−1</sup> Fe in polished grain and two <em>OsNAS2</em> overexpression lines had 14 and 19 µg g<sup>−1</sup> Fe in polished grain, representing up to four-fold increases in Fe concentration. Two-fold increases of Zn concentration were also observed in the <em>OsNAS2</em> population. Synchrotron X-ray fluorescence spectroscopy demonstrated that <em>OsNAS2</em> overexpression leads to significant enrichment of Fe and Zn in phosphorus-free regions of rice endosperm.</p> <h3>Conclusions</h3><p>The <em>OsNAS</em> genes, particularly <em>OsNAS2</em>, show enormous potential for Fe and Zn biofortification of rice endosperm. The results demonstrate that rice cultivars overexpressing single rice <em>OsNAS</em> genes could provide a sustainable and genetically simple solution to Fe and Zn deficiency disorders affecting billions of people throughout the world.</p> </div>", "links"=>[], "tags"=>["constitutive", "overexpression", "reveals", "single-gene", "strategies", "iron-", "zinc-biofortification", "endosperm"], "article_id"=>133652, "categories"=>["Biotechnology", "Ecology", "Cell Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Constitutive_Overexpression_of_the_OsNAS_Gene_Family_Reveals_Single_Gene_Strategies_for_Effective_Iron_and_Zinc_Biofortification_of_Rice_Endosperm/133652", "title"=>"Constitutive Overexpression of the <em>OsNAS</em> Gene Family Reveals Single-Gene Strategies for Effective Iron- and Zinc-Biofortification of Rice Endosperm", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2011-09-06 01:00:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/739090"], "description"=>"<p>RB, right border; 2 × 35S, dual CaMV 35S promoter; <i>OsNAS</i>, coding sequence of <i>OsNAS1</i> (999 bp), <i>OsNAS2</i> (981 bp) or <i>OsNAS3</i> (1032 bp); nos T, nopaline synthase terminator; 35S, CaMV 35S promoter; nptII, <i>neomycin phosphotransferase II</i>; LB, left border.</p>", "links"=>[], "tags"=>["t-dnas", "constitutive", "overexpression"], "article_id"=>409462, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Schematic_representation_of_the_T_DNAs_used_for_constitutive_overexpression_of_the_three_OsNAS_genes_/409462", "title"=>"Schematic representation of the T-DNAs used for constitutive overexpression of the three <i>OsNAS</i> genes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:37:42"}
  • {"files"=>["https://ndownloader.figshare.com/files/739172"], "description"=>"<p>WT, three wild type lines of rice; OE-<i>OsNAS1</i>, 30 independent transgenic lines overexpressing <i>OsNAS1</i>; OE-<i>OsNAS2</i>, 39 independent transgenic lines overexpressing <i>OsNAS2</i>; OE-<i>OsNAS3</i>, 24 independent transgenic lines overexpressing <i>OsNAS3</i>. Unpolished grain was analyzed by ICP-OES to determine Fe and Zn concentrations. The three populations of overexpression lines are sorted in order from lowest to highest Fe concentration in panels A and B.</p>", "links"=>[], "tags"=>["zn", "concentrations", "unpolished", "transgenic"], "article_id"=>409540, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Fe_A_and_Zn_B_concentrations_in_unpolished_grain_of_wild_type_and_transgenic_rice_/409540", "title"=>"Fe (A) and Zn (B) concentrations in unpolished grain of wild type and transgenic rice.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:39:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/739264"], "description"=>"<p>(A) NS, null segregant lines of rice; WT, wild type line of rice; OE, overexpression lines of rice. Three sibling T<sub>1</sub> lines, consisting of two OE lines and one NS line, were obtained from a single-insert T<sub>0</sub> mother line in each of the three <i>OsNAS</i> overexpression populations. The three <i>OsNAS1</i> sibling lines are labeled OE1-1, OE1-2 and NS1; the three <i>OsNAS2</i> sibling lines are labeled OE2-1, OE2-2 and NS2; the three <i>OsNAS3</i> sibling lines are labeled OE3-1, OE3-2 and NS3. Unpolished grain was analyzed by LC-MS to determine nicotianamine concentration (mean ± SE, n = 4). Significant differences from WT were determined by Student's <i>t</i> test and are indicated by asterisks (P<0.05). (B) Statistically significant positive correlations were found between unpolished grain NA concentration and Fe (black shapes; r = 0.9769 and p<0.01) and Zn (gray shapes; r = 0.9288 and P<0.01) concentrations for the ten genotypes described in panel A. The six OE1, OE2 and OE3 sibling lines are represented by triangles, diamonds and squares, respectively. The three NS lines appear just next to the WT line (represented by circles) on the scatter chart.</p>", "links"=>[], "tags"=>["zn", "concentrations", "unpolished", "positively", "correlated", "nicotianamine"], "article_id"=>409627, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Fe_and_Zn_concentrations_in_unpolished_grain_are_positively_correlated_with_nicotianamine_NA_concentration_/409627", "title"=>"Fe and Zn concentrations in unpolished grain are positively correlated with nicotianamine (NA) concentration.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:40:27"}
  • {"files"=>["https://ndownloader.figshare.com/files/739375"], "description"=>"<p>WT grain had 23 µg g<sup>−1</sup> DW Fe and 38 µg g<sup>−1</sup> DW Zn while OE-<i>OsNAS2A</i> grain had 64 µg g<sup>−1</sup> DW Fe and 80 µg g<sup>−1</sup> DW Zn, as determined by ICP-OES. (A) Light microscopy photo of a representative grain section with numbers indicating the location of scutellum (1), embryo (2) and endosperm (3); the green box represents the area used to obtain the line scans in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024476#pone-0024476-g005\" target=\"_blank\">Figures 5</a>–<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0024476#pone-0024476-g006\" target=\"_blank\">6</a>. (B–I) Elemental maps of Fe distribution in WT (B) and OE-<i>OsNAS2A</i> (C) grain; Zn distribution in WT (D) and OE-<i>OsNAS2A</i> (E) grain; Mn distribution in WT (F) and OE-<i>OsNAS2A</i> (G) grain; Cu distribution in WT (H) and OE-<i>OsNAS2A</i> (I) grain. The colour scale represents different elemental concentrations, with black and white corresponding to the lowest and highest concentrations, respectively.</p>", "links"=>[], "tags"=>["elemental", "maps", "wt", "longitudinal"], "article_id"=>409744, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_XRF_elemental_maps_of_WT_and_OE_OsNAS2A_longitudinal_grain_sections_/409744", "title"=>"µ-XRF elemental maps of WT and OE-<i>OsNAS2A</i> longitudinal grain sections.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:42:24"}
  • {"files"=>["https://ndownloader.figshare.com/files/739521"], "description"=>"<p>Line scans begin on the outer margin of grain and continue 135 µm towards the endosperm; data is displayed as average count (mean ± SE, n = 23). Counts are plotted on logarithmic scale in the y-axis to account for the low P counts (blue) relative to Fe (red) and Zn (green). As the rice grain contains a single aleurone layer that is rich in phosphorus (P, primarily in the form of phytic acid) while endosperm contains little P, 1–50 µm was assigned to the aleurone layer, 51–90 µm to the subaleurone layer and 91–135 µm to the endosperm.</p>", "links"=>[], "tags"=>["scans", "fe", "zn", "wt"], "article_id"=>409890, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Line_scans_for_P_Fe_and_Zn_in_WT_A_and_OE_OsNAS2A_B_grain_/409890", "title"=>"Line scans for P, Fe and Zn in WT (A) and OE-<i>OsNAS2A</i> (B) grain.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:44:50"}
  • {"files"=>["https://ndownloader.figshare.com/files/739595"], "description"=>"<p>Line scans begin on the outer margin of grain and continue 135 µm towards the endosperm; average WT counts indicated by the black line and average OE-<i>OsNAS2</i> counts indicated by the gray line (mean, n = 23). The location of the successively higher Fe peaks in aleurone, subaleurone and endosperm regions of OE-<i>OsNAS2A</i> grain is indicated on the figure.</p>", "links"=>[], "tags"=>["scans", "fe", "wt"], "article_id"=>409969, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.g006"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Line_scans_for_Fe_in_WT_and_OE_OsNAS2A_grain_/409969", "title"=>"Line scans for Fe in WT and OE-<i>OsNAS2A</i> grain.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-09-06 02:46:09"}
  • {"files"=>["https://ndownloader.figshare.com/files/739670"], "description"=>"<p>The Fe:Zn signal ratio is presented for each of the three layers.</p>", "links"=>[], "tags"=>["counts", "fe", "zn", "subaleurone", "endosperm", "layers", "wt"], "article_id"=>410030, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.t002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Average_counts_of_Fe_and_Zn_in_aleurone_subaleurone_and_endosperm_layers_of_WT_and_OE_OsNAS2A_grain_/410030", "title"=>"Average counts of Fe and Zn in aleurone, subaleurone and endosperm layers of WT and OE-<i>OsNAS2A</i> grain.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2011-09-06 00:00:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/739708"], "description"=>"<p>Grain samples from WT, one transgenic line overexpressing <i>OsNAS1</i> (OE-<i>OsNAS1S</i>), two independent transgenic lines overexpressing O<i>sNAS2</i> (OE-<i>OsNAS2B</i> and OE-<i>OsNAS2J</i>) and one transgenic line overexpressing <i>OsNAS3</i> (OE-<i>OsNAS3B</i>) were analyzed by ICP-OES. The percentage of Fe and Zn concentration in polished grain, relative to unpolished grain concentration, is presented in the last two columns of the table.</p>", "links"=>[], "tags"=>["fe", "zn", "unpolished", "polished", "wt", "transgenic"], "article_id"=>410073, "categories"=>["Biotechnology", "Ecology", "Plant Biology"], "users"=>["Alexander A. T. Johnson", "Bianca Kyriacou", "Damien L. Callahan", "Lorraine Carruthers", "James Stangoulis", "Enzo Lombi", "Mark Tester"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0024476.t001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Concentrations_of_Fe_and_Zn_in_unpolished_and_polished_grain_of_WT_and_transgenic_rice_/410073", "title"=>"Concentrations of Fe and Zn in unpolished and polished grain of WT and transgenic rice.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2011-09-06 00:01:13"}

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