FAX1, a Novel Membrane Protein Mediating Plastid Fatty Acid Export
Publication Date
February 03, 2015
Journal
PLOS Biology
Authors
Nannan Li, Irene Luise Gügel, Patrick Giavalisco, Viktoria Zeisler, et al
Volume
13
Issue
2
Pages
e1002053
DOI
https://dx.plos.org/10.1371/journal.pbio.1002053
Publisher URL
http://journals.plos.org/plosbiology/article?id=10.1371%2Fjournal.pbio.1002053
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/25646734
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4344464
Europe PMC
http://europepmc.org/abstract/MED/25646734
Web of Science
000352080100003
Scopus
84924024823
Mendeley
http://www.mendeley.com/research/fax1-novel-membrane-protein-mediating-plastid-fatty-acid-export
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Mendeley | Further Information

{"title"=>"FAX1, a Novel Membrane Protein Mediating Plastid Fatty Acid Export", "type"=>"journal", "authors"=>[{"first_name"=>"Nannan", "last_name"=>"Li", "scopus_author_id"=>"56535685800"}, {"first_name"=>"Irene Luise", "last_name"=>"Gügel", "scopus_author_id"=>"35198184000"}, {"first_name"=>"Patrick", "last_name"=>"Giavalisco", "scopus_author_id"=>"6507393408"}, {"first_name"=>"Viktoria", "last_name"=>"Zeisler", "scopus_author_id"=>"55376902100"}, {"first_name"=>"Lukas", "last_name"=>"Schreiber", "scopus_author_id"=>"56107745000"}, {"first_name"=>"Jürgen", "last_name"=>"Soll", "scopus_author_id"=>"7007179709"}, {"first_name"=>"Katrin", "last_name"=>"Philippar", "scopus_author_id"=>"7801505281"}], "year"=>2015, "source"=>"PLoS Biology", "identifiers"=>{"pmid"=>"25646734", "scopus"=>"2-s2.0-84924024823", "issn"=>"15457885", "pui"=>"602601535", "doi"=>"10.1371/journal.pbio.1002053", "isbn"=>"1545-7885 (Electronic)\\r1544-9173 (Linking)", "sgr"=>"84924024823"}, "id"=>"7087ebd1-a01b-32f9-9613-f3620edc1ea2", "abstract"=>"Fatty acid synthesis in plants occurs in plastids, and thus, export for subsequent acyl editing and lipid assembly in the cytosol and endoplasmatic reticulum is required. Yet, the transport mechanism for plastid fatty acids still remains enigmatic. We isolated FAX1 (fatty acid export 1), a novel protein, which inserts into the chloroplast inner envelope by α-helical membrane-spanning domains. Detailed phenotypic and ultrastructural analyses of FAX1 mutants in Arabidopsis thaliana showed that FAX1 function is crucial for biomass production, male fertility and synthesis of fatty acid-derived compounds such as lipids, ketone waxes, or pollen cell wall material. Determination of lipid, fatty acid, and wax contents by mass spectrometry revealed that endoplasmatic reticulum (ER)-derived lipids decreased when FAX1 was missing, but levels of several plastid-produced species increased. FAX1 over-expressing lines showed the opposite behavior, including a pronounced increase of triacyglycerol oils in flowers and leaves. Furthermore, the cuticular layer of stems from fax1 knockout lines was specifically reduced in C29 ketone wax compounds. Differential gene expression in FAX1 mutants as determined by DNA microarray analysis confirmed phenotypes and metabolic imbalances. Since in yeast FAX1 could complement for fatty acid transport, we concluded that FAX1 mediates fatty acid export from plastids. In vertebrates, FAX1 relatives are structurally related, mitochondrial membrane proteins of so-far unknown function. Therefore, this protein family might represent a powerful tool not only to increase lipid/biofuel production in plants but also to explore novel transport systems involved in vertebrate fatty acid and lipid metabolism.", "link"=>"http://www.mendeley.com/research/fax1-novel-membrane-protein-mediating-plastid-fatty-acid-export", "reader_count"=>81, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>17, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>18, "Student > Postgraduate"=>6, "Student > Master"=>14, "Other"=>4, "Student > Bachelor"=>6, "Professor"=>7}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>2, "Researcher"=>17, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>18, "Student > Postgraduate"=>6, "Student > Master"=>14, "Other"=>4, "Student > Bachelor"=>6, "Professor"=>7}, "reader_count_by_subject_area"=>{"Unspecified"=>2, "Engineering"=>1, "Biochemistry, Genetics and Molecular Biology"=>21, "Materials Science"=>1, "Agricultural and Biological Sciences"=>53, "Medicine and Dentistry"=>1, "Chemistry"=>1, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>1}, "Materials Science"=>{"Materials Science"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Chemistry"=>{"Chemistry"=>1}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>53}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>21}, "Unspecified"=>{"Unspecified"=>2}}, "reader_count_by_country"=>{"Canada"=>1, "United States"=>2, "Japan"=>1, "Mexico"=>2, "United Kingdom"=>1, "Slovenia"=>1}, "group_count"=>2}

CrossRef

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/1879522"], "description"=>"<p>Plant biomass of <i>FAX1</i> mutant lines and wild-type plants.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300608, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.t001", "stats"=>{"downloads"=>3, "page_views"=>17, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Plant_biomass_of_FAX1_mutant_lines_and_wild_type_plants_/1300608", "title"=>"Plant biomass of <i>FAX1</i> mutant lines and wild-type plants.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879520"], "description"=>"<p>Triacylglycerol (TAG) oils were determined in tissues of 7-week-old, mature flowering plants (compare <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.g006\" target=\"_blank\">Fig. 6</a>). Data (arbitrary units) are expressed relative to the internal standard (PC 34:0) and normalized to mg fresh weight (FW). For overview, we selected representatives of the most abundant species and those significantly different in <i>FAX1</i> mutants. A complete dataset with details on analysis is given in <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s012\" target=\"_blank\">S1 Table</a>; values in mol % are listed in <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s015\" target=\"_blank\">S4 Table</a>. Levels in FAX1 mutants significantly different to wild type are indicated (Student’s <i>t</i>-test, *: <i>p</i> < 0.05, **: <i>p</i> < 0.01). We show high and low abundant TAGs (left and right graphs, respectively); thus for better resolution of differential patterns, <i>y</i>-axes are scaled differently. (A) TAG levels in caulinary leaves of <i>fax1–1</i>, <i>fax1–2</i> knockout and Col-0, WT2 wild-type lines (yellow and black bars, respectively). Mean values (<i>n</i> = 4–6 ± SD), averaged over both <i>fax1</i> knockouts and both wild types, respectively, are shown. (B) TAG content in caulinary leaves of the <i>FAX1</i> over-expressing line ox#4 (green bars; <i>n</i> = 6–12 ± SD) and Col-0 wild type (black bars, <i>n</i> = 5–10 ± SD). (C) TAG levels in flowers of <i>fax1–1</i>, <i>fax1–2</i> knockout and Col-0, WT2 wild-type lines (yellow and black bars, respectively). Mean values (<i>n</i> = 6 ± SD), averaged over both <i>fax1</i> knockouts and both wild types, respectively, are shown. (D) TAG content (<i>n</i> = 7–12 ± SD, for TAG 58:6: <i>n</i> = 5) in flowers of the <i>FAX1</i> over-expressing line ox#4 and Col-0 wild type (green and black bars, respectively). Please note that in comparison to the dataset for <i>fax1</i> knockouts in (A) and (C), usage of a different mass spectrometer for FAX1ox data in (B) and (D) results in different scaling of the relative values (compare <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.g006\" target=\"_blank\">Fig. 6</a>).</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300606, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g007", "stats"=>{"downloads"=>1, "page_views"=>18, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Plastid_FAX1_impacts_TAG_storage_lipid_homeostasis_/1300606", "title"=>"Plastid FAX1 impacts TAG storage lipid homeostasis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879517"], "description"=>"<p>Stem tissue (1 cm at the bottom of the second internode) of the primary inflorescence stalk of 5-week-old <i>fax1–1</i> knockout, Col-0 wild-type, and <i>FAX1</i> over-expressor line ox#2 (left, middle, and right panels, respectively). (A) Light microscopic pictures of stem epidermal cells (bar = 10μm). (B) Transmission electron microscopic pictures of cell walls from stem epidermal cells (bar = 500 nm). (C) Close-ups of cell wall / cuticular layer boundary from cells in (B) (TEM, bar = 200 nm). cut: cuticular layer; cw: cell wall; cyt: cytosol. (D) C29 ketone wax coverage in μg per cm<sup>2</sup> of stem surface from <i>FAX1</i> mutant and wild-type lines (<i>n</i> = 3–7 ± SD; <i>n</i> = 12 for Col-0). For each replicate, three to four stem sections between internode 2–4 of 7-week-old, mature flowering plants were pooled. Asterisks indicate highly significant different contents (**: <i>p</i> < 0.001, Student’s <i>t</i>-test) when compared with Col-0 (for numerical values, see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s001\" target=\"_blank\">S1A Data</a>).</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300603, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g005", "stats"=>{"downloads"=>6, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_FAX1_affects_cell_wall_size_and_cuticular_wax_composition_/1300603", "title"=>"FAX1 affects cell wall size and cuticular wax composition.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879498"], "description"=>"<p>(A) 30-day-old plants of <i>FAX1</i> mutants and wild-type lines. <i>fax1–1</i>, <i>fax1–2</i>: homozygous knockout lines for <i>FAX1</i>; Col-0, WT2: wild-type <i>FAX1</i> alleles, WT2 represents <i>FAX1</i> wild-type progeny, segregated from heterozygous <i>fax1–2</i>; Co#7, Co#54: <i>fax1–2</i> knockout complementation lines; ox#2, ox#4: <i>FAX1</i> over-expressing lines in Col-0 background. (B) 7-week-old flowering plants of <i>FAX1</i> mutants and wild type as specified in (A). (Inset) Comparison of primary inflorescence stalks (bottom parts of 2<sup>nd</sup> internode) from <i>fax1–2</i>, Col-0 and ox#2 plants. (C) Siliques produced by <i>FAX1</i> mutants and wild-type lines as depicted in (B).</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300584, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g003", "stats"=>{"downloads"=>4, "page_views"=>30, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mutation_of_FAX1_in_Arabidopsis_affects_plant_growth_/1300584", "title"=>"Mutation of <i>FAX1</i> in <i>Arabidopsis</i> affects plant growth.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879524"], "description"=>"<p>Impact of <i>FAX1</i> mutation on cellular FA/lipid homeostasis.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300610, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.t003", "stats"=>{"downloads"=>4, "page_views"=>24, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Impact_of_FAX1_mutation_on_cellular_FA_lipid_homeostasis_/1300610", "title"=>"Impact of <i>FAX1</i> mutation on cellular FA/lipid homeostasis.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879523"], "description"=>"<p>Segregation analysis of <i>fax1</i> knockout lines.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300609, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.t002", "stats"=>{"downloads"=>5, "page_views"=>22, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Segregation_analysis_of_fax1_knockout_lines_/1300609", "title"=>"Segregation analysis of <i>fax1</i> knockout lines.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879547", "https://ndownloader.figshare.com/files/1879548", "https://ndownloader.figshare.com/files/1879549", "https://ndownloader.figshare.com/files/1879550", "https://ndownloader.figshare.com/files/1879551", "https://ndownloader.figshare.com/files/1879552", "https://ndownloader.figshare.com/files/1879553", "https://ndownloader.figshare.com/files/1879554", "https://ndownloader.figshare.com/files/1879555", "https://ndownloader.figshare.com/files/1879556", "https://ndownloader.figshare.com/files/1879557", "https://ndownloader.figshare.com/files/1879558", "https://ndownloader.figshare.com/files/1879559", "https://ndownloader.figshare.com/files/1879560", "https://ndownloader.figshare.com/files/1879561", "https://ndownloader.figshare.com/files/1879562", "https://ndownloader.figshare.com/files/1879563", "https://ndownloader.figshare.com/files/1879564", "https://ndownloader.figshare.com/files/1879565", "https://ndownloader.figshare.com/files/1879567"], "description"=>"<div><p>Fatty acid synthesis in plants occurs in plastids, and thus, export for subsequent acyl editing and lipid assembly in the cytosol and endoplasmatic reticulum is required. Yet, the transport mechanism for plastid fatty acids still remains enigmatic. We isolated FAX1 (<i>fatty acid export 1</i>), a novel protein, which inserts into the chloroplast inner envelope by α-helical membrane-spanning domains. Detailed phenotypic and ultrastructural analyses of FAX1 mutants in <i>Arabidopsis thaliana</i> showed that FAX1 function is crucial for biomass production, male fertility and synthesis of fatty acid-derived compounds such as lipids, ketone waxes, or pollen cell wall material. Determination of lipid, fatty acid, and wax contents by mass spectrometry revealed that endoplasmatic reticulum (ER)-derived lipids decreased when FAX1 was missing, but levels of several plastid-produced species increased. FAX1 over-expressing lines showed the opposite behavior, including a pronounced increase of triacyglycerol oils in flowers and leaves. Furthermore, the cuticular layer of stems from <i>fax1</i> knockout lines was specifically reduced in C29 ketone wax compounds. Differential gene expression in FAX1 mutants as determined by DNA microarray analysis confirmed phenotypes and metabolic imbalances. Since in yeast FAX1 could complement for fatty acid transport, we concluded that FAX1 mediates fatty acid export from plastids. In vertebrates, FAX1 relatives are structurally related, mitochondrial membrane proteins of so-far unknown function. Therefore, this protein family might represent a powerful tool not only to increase lipid/biofuel production in plants but also to explore novel transport systems involved in vertebrate fatty acid and lipid metabolism.</p></div>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300624, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>["https://dx.doi.org/10.1371/journal.pbio.1002053.s001", "https://dx.doi.org/10.1371/journal.pbio.1002053.s002", "https://dx.doi.org/10.1371/journal.pbio.1002053.s003", "https://dx.doi.org/10.1371/journal.pbio.1002053.s004", "https://dx.doi.org/10.1371/journal.pbio.1002053.s005", "https://dx.doi.org/10.1371/journal.pbio.1002053.s006", "https://dx.doi.org/10.1371/journal.pbio.1002053.s007", "https://dx.doi.org/10.1371/journal.pbio.1002053.s008", "https://dx.doi.org/10.1371/journal.pbio.1002053.s009", "https://dx.doi.org/10.1371/journal.pbio.1002053.s010", "https://dx.doi.org/10.1371/journal.pbio.1002053.s011", "https://dx.doi.org/10.1371/journal.pbio.1002053.s012", "https://dx.doi.org/10.1371/journal.pbio.1002053.s013", "https://dx.doi.org/10.1371/journal.pbio.1002053.s014", "https://dx.doi.org/10.1371/journal.pbio.1002053.s015", "https://dx.doi.org/10.1371/journal.pbio.1002053.s016", "https://dx.doi.org/10.1371/journal.pbio.1002053.s017", "https://dx.doi.org/10.1371/journal.pbio.1002053.s018", "https://dx.doi.org/10.1371/journal.pbio.1002053.s019", "https://dx.doi.org/10.1371/journal.pbio.1002053.s020"], "stats"=>{"downloads"=>93, "page_views"=>51, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_FAX1_a_Novel_Membrane_Protein_Mediating_Plastid_Fatty_Acid_Export_/1300624", "title"=>"FAX1, a Novel Membrane Protein Mediating Plastid Fatty Acid Export", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879519"], "description"=>"<p>Free fatty acid (FA) and polar lipid species were determined in leaf tissue of 7-week-old, mature flowering plants. Data (arbitrary units) are expressed relative to the internal standard (PC 34:0) and normalized to mg fresh weight (FW). For overview, we depict representatives of the most abundant species and those significantly different in <i>FAX1</i> mutants. A complete dataset with details on analysis is given in <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s012\" target=\"_blank\">S1 Table</a>; values in mol % are listed in <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s013\" target=\"_blank\">S2 Table</a>. Levels in FAX1 mutants significantly different to wild type are indicated by asterisks (Student’s <i>t</i>-test, *: <i>p</i> < 0.05, **: <i>p</i> < 0.01). For a better resolution of differential patterns, <i>y</i>-axes are scaled differently. C<sub>16–18</sub> FAs are exclusively and glycolipids in (A) and (B) are mainly synthesized in plastids (for details, see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#sec009\" target=\"_blank\">Discussion</a>). Please note that the diacylglycerol backbone for the “34”-glycolipids can originate from prokaryotic (from plastids) and eukaryotic (from the ER) phospholipid precursors, respectively. C<sub>20–26</sub> FAs and phospholipids in (C) and (D), as well as precursors for “36”- glycolipids, are only produced outside plastids in the cytosol and/or ER. (A), (C) Free FA and lipid levels in caulinary leaves of <i>fax1–1</i>, <i>fax1–2</i> knockout and Col-0, WT2 wild-type lines (yellow and black bars, respectively) were determined by UPLC-Orbitrap MS [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref023\" target=\"_blank\">23</a>]. Mean values (<i>n</i> = 6 ± SD), averaged over both <i>fax1</i> knockouts and both wild types, respectively, are shown. (B), (D) Free FA and lipid content (<i>n</i> = 6–12 ± SD) in caulinary leaves of the <i>FAX1</i> over-expressing line ox#4 and Col-0 wild type (green and black bars, respectively) was measured by UPLC-qTOF MS [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref024\" target=\"_blank\">24</a>]. Please note that in comparison to the dataset in (A) and (C), usage of a different mass spectrometer results in different scaling of the relative values. DGDG: digalactosyl-diacylglycerol; FA: fatty acid; MGDG: monogalactosyl-diacylglycerol; n.d.: not determined; PC: phosphatidyl-choline; PE: phosphatidyl-ethanolamine; PG: phosphatidyl-glycerol; SQDG: sulphoquinovosyl-diacylglycerol.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300605, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g006", "stats"=>{"downloads"=>1, "page_views"=>36, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Plastid_FAX1_impacts_cellular_FA_lipid_homeostasis_in_leaves_/1300605", "title"=>"Plastid FAX1 impacts cellular FA/lipid homeostasis in leaves.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879492"], "description"=>"<p>(A, B) Structural models and alignment (right) of the mature At-FAX1 / human TMEM14C (A) and of At-FAX6 / human TMEM14A (B) proteins. Membrane-spanning and amphiphilic α-helices of FAX and of TMEM14 are depicted in green/yellow-green and blue/light blue, respectively. Please note that At-FAX1 contains an additional N-terminal stretch that most likely folds into another α-helix (gray). First and last amino acid residues are indicated. (C) In vivo green fluorescent protein (GFP)-targeting. Arabidopsis leaf protoplasts were transiently transformed with constructs for FAX1- and NiCo-GFP (chloroplast IE marker; [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref019\" target=\"_blank\">19</a>]). Images show GFP and chlorophyll fluorescence, as well as an overlay of both (bar = 5 μm). (D) Immunoblot analysis of FAX1 in chloroplast subfractions. Equal protein amounts (5μg) of pea chloroplast outer envelope (OE), inner envelope (IE), stroma (str), thylakoids (thy), as well as 2.5μg protein of Arabidopsis microsomal membranes (mm) and chloroplast envelopes (env) were separated by SDS-PAGE and subjected to immunoblot analysis using antibodies directed against Ps-FAX1 and At-FAX1. Antisera against marker proteins LSU (str), LHCP (thy), NiCo (IE), OEP16.1 (OE), and TPR7 (mm, see [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref020\" target=\"_blank\">20</a>]) were used as controls. For LSU and LHCP less protein (1μg, 0.2μg, respectively) was loaded. Numbers indicate molecular mass of proteins in kDa.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300578, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g002", "stats"=>{"downloads"=>2, "page_views"=>19, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_FAX1_a_chloroplast_IE_protein_of_the_Tmemb_14_family_/1300578", "title"=>"FAX1, a chloroplast IE protein of the Tmemb_14 family.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879509"], "description"=>"<p>Pictures of flowers, anthers, and mature pollen of 5-week-old <i>fax1–2</i> knockout and Col-0 wild-type plants (left and right panels, respectively). (A) Development of flower buds and young siliques. (B) Close-up of opened flowers. Arrowheads: non-pollinated stigma in <i>fax1–2</i>; arrows: anthers with released pollen in Col-0. (C) Cross sections of mature, dehisced anthers (light microscopy, bar = 50 μm). Black and white arrowheads in <i>fax1–2</i>: fully opened pollen sacks, and dark material covering endothecium/locule boundary, respectively. en: endothecium cells of anthers. (D), (E), (F) Transmission electron microscopic (TEM) pictures of anther cell/pollen grain intersections (D, bar = 5 μm; E, bar = 1μm) and pollen cell wall (F, bar = 500 nm) at mature tricellular pollen stages. White arrowheads in <i>fax1–2</i>: debris material sticking to pollen grains. en: endothecium cell; e: exine layer with e<sub>b</sub>: bacula, e<sub>t</sub>: tectum structures; e<sub>n</sub>: nexine layer (black arrowheads), i: intine layer; po: cytosol of pollen grain; pl: plastid; try: tryphine pollen coat.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300595, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g004", "stats"=>{"downloads"=>1, "page_views"=>17, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_FAX1_function_is_essential_for_pollen_cell_wall_assembly_/1300595", "title"=>"FAX1 function is essential for pollen cell wall assembly.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879491"], "description"=>"<p>(A) Arabidopsis At-FAX1 (At3g57280, 226 amino acids [aa]) is accessible at the ARAMEMNON database [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref018\" target=\"_blank\">18</a>], the sequence for pea Ps-FAX1 (232 aa) was deposited at National Center for Biotechnology Information (NCBI), GenBank acc. no. KF981436. Predicted chloroplast targeting peptides (ChloroP; <a href=\"http://www.cbs.dtu.dk/services/ChloroP\" target=\"_blank\">http://www.cbs.dtu.dk/services/ChloroP</a>), with 33 aa and 39 aa for At-FAX1 and Ps-FAX1, respectively, are marked with red triangles. The Tmemb_14 domain (Pfam|PF03647) of At-FAX1, including the four conserved hydrophobic domains, is indicated. Identical amino acids (49%) are shaded in black, hydrophobic α-helices (ARAMEMNON) are boxed in green, and peptides used for generation of antisera are indicated by red lines. (B) Members of the FAX/Tmemb_14 family in Arabidopsis. Whereas At-FAX1-At-FAX4 are predicted to be in plastids, At-FAX5-At-FAX7 most likely localize to membranes of the secretory pathway. Hydrophobic α-helices (black squares) and subcellular localization are depicted according to ARAMEMNON. Predicted chloroplast targeting peptides (ChloroP) are marked with red triangles. (C) At-FAX1 and Ps-FAX1 [sequence starting with Tmemb_14 domain, see (A)] in comparison to At-FAX6 (At3g20510) and human proteins of the Tmemb_14 superfamily: TMEM14A (Q9Y6G1) and TMEM14C (Q9POS9). Arabidopsis genome initiative (AGI) codes and InterPro accession numbers in brackets. Whereas At-FAX1 and Ps-FAX1 are slightly more similar to TMEM14C (17% identical, 35% similar aa), At-FAX6 shares 28% identical aa with both proteins TMEM14A and 14C. Predicted hydrophobic α-helices (ARAMEMNON) are boxed in green; α-helices in TMEM14A, 14C [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref014\" target=\"_blank\">14</a>] are boxed in blue.</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300577, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g001", "stats"=>{"downloads"=>6, "page_views"=>21, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Plant_FAX_and_human_Tmemb_14_proteins_/1300577", "title"=>"Plant FAX and human Tmemb_14 proteins.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/1879521"], "description"=>"<p>The empty plasmid pDR195 (-) and the mature At-FAX1 cDNA in pDR195 (FAX1) were introduced into <i>faa1/faa4</i> and <i>fat1</i> yeast mutants, respectively. (A) Serial dilutions (OD<sub>600</sub> of 10<sup>–1</sup>, 10<sup>–2</sup>, and 5<sup>–3</sup>) of rapidly growing yeast cells on SD-ura plates (0.1% glucose, 1% tergitol). Control plates (left) in comparison to plates with 3.6 mM α-linolenic acid (C<sub>18:3</sub>; right). (B), (C) Growth of <i>fat1</i> (B) and <i>faa1/faa4</i> (C) cells in liquid SD-ura [see (A)]. The OD<sub>600</sub> was monitored within 29 h incubation at 30°C. White and light gray bars: growth of pDR195 (-) and matFAX1/pDR195 (FAX1) at control conditions. Gray and black bars: growth of (-) and FAX1 in presence of 3.6 mM α-linolenic acid (LIN). Error bars depict SD (<i>n</i> = 4), for numerical values, see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s001\" target=\"_blank\">S1B Data</a>. a (<i>p</i> < 0.05), b (<i>p</i> < 0.005), c (<i>p</i> < 0.0005), * (<i>p</i> < 0.00005) indicate significantly different values (Student’s <i>t</i>-test) compared to (-) cells without and with LIN, respectively. (D) Representative growth curves of <i>fat1</i> cells in liquid SD-ura (2% glucose, 0.5% Brij 58, 0.7% KH<sub>2</sub>PO<sub>4</sub>) in the presence of 10μM cerulenin (CER, inhibitor of FA-biosynthesis) and 100 μM palmitic acid (PAL, C<sub>16:0</sub>) or 100 μM α-linolenic acid (LIN, C<sub>18:3</sub>). Assays were performed according to [<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref026\" target=\"_blank\">26</a>,<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.ref027\" target=\"_blank\">27</a>]. For comparison with stearic (C<sub>18:0</sub>) and oleic acid (C<sub>18:1</sub>), see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s005\" target=\"_blank\">S4 Fig.</a> White and black circles and triangles: growth of pDR195 (-) and matFAX1/pDR195 (FAX1) cells supplemented with PAL and LIN, respectively. Red/blue lines and numbers indicate maximal difference of cell density ratios for (-) versus FAX1 (compare <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1002053#pbio.1002053.s001\" target=\"_blank\">S1B Data</a>).</p>", "links"=>[], "tags"=>["FAX 1 function", "export", "pollen cell wall material", "lipid", "FAX 1 relatives", "er", "endoplasmatic reticulum", "FAX 1 mediates", "novel transport systems", "Export Fatty acid synthesis", "plastid", "mitochondrial membrane proteins", "DNA microarray analysis", "differential gene expression", "FAX 1 mutants", "C 29 ketone wax compounds", "yeast FAX 1", "fax 1 knockout lines", "Novel Membrane Protein Mediating Plastid Fatty", "FAX 1"], "article_id"=>1300607, "categories"=>["Uncategorised"], "users"=>["Nannan Li", "Irene Luise Gügel", "Patrick Giavalisco", "Viktoria Zeisler", "Lukas Schreiber", "Jürgen Soll", "Katrin Philippar"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1002053.g008", "stats"=>{"downloads"=>1, "page_views"=>20, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_FAX1_mediates_FA_transport_in_yeast_/1300607", "title"=>"FAX1 mediates FA-transport in yeast.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-02-03 03:09:25"}

PMC Usage Stats | Further Information

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Relative Metric

{"start_date"=>"2015-01-01T00:00:00Z", "end_date"=>"2015-12-31T00:00:00Z", "subject_areas"=>[]}

F1000Prime | Further Information

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