Cytosolic iron-sulfur cluster assembly (Homo sapiens)

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<?xml version="1.0" encoding="UTF-8"?> <Pathway xmlns="http://pathvisio.org/GPML/2013a" Name="Cytosolic iron-sulfur cluster assembly" Organism="Homo sapiens"> <Comment Source="WikiPathways-description">Iron-sulfur clusters containing 4 atoms of iron and 4 atoms of sulfur (4Fe-4S clusters) are assembled in the cytosol on a heterotetrameric scaffold composed of NUBP2 and NUBP1 subunits (reviewed in Lill et al. 2012, Rouault et al. 2012, Sharma et al. 2010, Lill and Muhlenhoff 2006). The sources of iron and sulfur are uncertain but the process requires a sulfur-containing compound exported from mitochondria via ABCB7 (ABC7). Newly synthesized 4Fe-4S are transferred to apoproteins such as XPD and POLD1 via the CIA targeting complex, composed of NARFL, CIAO1, FAM96B, and MMS19. View original pathway at:[http://www.reactome.org/PathwayBrowser/#DIAGRAM=2564830 Reactome].</Comment> <Comment Source="Reactome-Converter">Pathway is converted from Reactome ID: 2564830</Comment> <Comment Source="Reactome-version">Reactome version: 63</Comment> <Comment Source="Reactome Author">Reactome Author: Lill, R</Comment> <BiopaxRef>cf3</BiopaxRef> <BiopaxRef>a5c</BiopaxRef> <BiopaxRef>fc7</BiopaxRef> <BiopaxRef>bcf</BiopaxRef> <Attribute Key="reactome_id" Value="2564830" /> <Graphics BoardWidth="932.0" BoardHeight="1495.0" /> <DataNode TextLabel="BRIP1 " GraphId="a0572" Type="Protein" GroupRef="e130d"> <Attribute Key="cellular_location" Value="cytosol" /> <Attribute Key="complex_id" Value="R-HSA-2564793" /> <Attribute Key="copies_num" Value="1" /> <Graphics CenterX="100.0" CenterY="815.0" Width="100.0" Height="20.0" ZOrder="32768" FontSize="10" Valign="Middle" /> <Xref Database="Uniprot-TrEMBL" ID="Q9BX63" /> </DataNode> <DataNode TextLabel="NARFL:CIAO1:FAM96B:MMS19" 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Height="20.0" ZOrder="32768" FontSize="10" Valign="Middle" /> <Xref Database="Uniprot-TrEMBL" ID="P18074" /> </DataNode> <DataNode TextLabel="ABC7 dimer" GraphId="feb86" Type="Complex"> <Attribute Key="cellular_location" Value="mitochondrial inner membrane" /> <Graphics CenterX="398.5" CenterY="146.0" Width="105.0" Height="22.0" ZOrder="32768" FontSize="10" Valign="Middle" ShapeType="Hexagon" Color="a52a2a" /> <Xref Database="Reactome" ID="R-HSA-1369041" /> </DataNode> <Interaction GraphId="a22db"> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="532.0" Y="441.0" GraphRef="dbef6" RelX="-0.005847953216374269" RelY="0.9310344827586207" /> <Point X="530.0" Y="534.0" GraphRef="cbe12" RelX="0.0" RelY="0.0" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564828" /> </Interaction> <Interaction GraphId="a43b0"> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="209.0" Y="359.0" GraphRef="a0cd3" RelX="0.0" RelY="0.5099999999999909" /> <Point X="212.0" Y="400.0" GraphRef="aa74c" RelX="0.0" RelY="-1.0" ArrowHead="Arrow" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564824" /> </Interaction> <Interaction GraphId="a6ee5"> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="216.0" Y="270.0" GraphRef="ee357" RelX="-1.0" RelY="0.0" /> <Point X="209.0" Y="308.0" GraphRef="a9fc6" RelX="0.0" RelY="0.0" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564824" /> </Interaction> <Interaction GraphId="ab824"> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="530.0" Y="643.0" GraphRef="b7a72" RelX="0.0" RelY="1.0900000000000318" /> <Point X="605.0" Y="197.0" GraphRef="d041e" RelX="-1.0" RelY="0.0" ArrowHead="Arrow" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564828" /> </Interaction> <Interaction GraphId="abf96"> <Comment Source="Reactome">A complex of four proteins termed CIA targeting complex is involved in the delivery of the NUBP1-NUBP2-bound Fe/S cluster to specific target apoproteins. NARFL (IOP1, Nar1 in yeast) is similar in sequence to iron-only hydrogenases and binds two [4Fe-4S] clusters. These cofactors are essential for biogenesis but their molecular function is unknown. The WD40 domain protein CIAO1 (Cia1 in yeast) forms a propeller-like structure which may serve as a docking site for the other members of the CIA targeting complex. The role of FAM96B (YHR122W, CIA2 in yeast) and MMS19 is still unclear, but they show some specificity for the delivery of Fe/S clusters into certain target apoproteins. They directly interact with numerous Fe/S target proteins including many proteins involved in nuclear DNA integrity and maintenance such as DNA polymerases (POLD1), helicases of the DNA repair pathway (XPD or FANCJ) or of telomere maintenance (RTEL1), and primases. 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The precise role of electrons in the assembly process is unclear but in the absence of the electron chain the stable [4Fe-4S] cluster of NUBP1 is not assembled. CIAPIN1 binds two Fe/S clusters of the [2Fe-2S] and the [4Fe-4S] type. The [2Fe-2S] cluster is reduced by NDOR1-NADPH.</Comment> <BiopaxRef>ced</BiopaxRef> <BiopaxRef>a9a</BiopaxRef> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="209.0" Y="308.0" /> <Point X="209.0" Y="359.0" /> <Anchor Position="0.0" Shape="None" GraphId="a9fc6" /> <Anchor Position="0.99" Shape="None" GraphId="a0cd3" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564824" /> </Interaction> <Interaction GraphId="f2943"> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="425.0" Y="508.0" GraphRef="d4245" RelX="1.0" RelY="-0.03225806451612903" /> <Point X="530.0" Y="534.0" GraphRef="cbe12" RelX="0.0" RelY="0.0" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564828" /> </Interaction> <Interaction GraphId="f4311"> <Comment Source="Reactome">A [4Fe-4S] cluster is assembled on a scaffold composed of the P-loop NTPases NUBP2 (CFD1 in yeast) and NUBP1 (Nbp35 in yeast) in a nucleotide-dependent fashion. The two proteins form a heterotetramer which transiently binds the [4Fe-4S] cluster in a bridged form between two subunits of the complex. The Fe/S cluster is bound to two highly conserved Cys residues in the C-termini of each of these proteins. NUBP1 contains an additional, stably associated [4Fe-4S] cluster at its N-terminus which is essential for function.&lt;br&gt;Mitochondria play a crucial role in cytosolic and nuclear Fe/S protein biogenesis. They export via a mitochondrial ABC transporter (yeast ATM1, human ABCB7) a still unknown, sulfur-containing compound which is essential for Fe/S cluster assembly in the cytosol.&lt;br&gt;The general cytosolic iron donor, the multi-domain monothiol glutaredoxin (human GRX3 or PICOT, yeast Grx3-Grx4) plays a crucial role in cytosolic-nuclear Fe/S protein biogenesis. The precise molecular function of the glutaredoxin is still unclear. </Comment> <BiopaxRef>bdf</BiopaxRef> <Graphics ConnectorType="Segmented" ZOrder="12288" LineThickness="1.0"> <Point X="627.0" Y="290.0" /> <Point X="531.0" Y="410.0" /> <Anchor Position="0.0" Shape="None" GraphId="ae8f2" /> <Anchor Position="0.99" Shape="None" GraphId="c609f" /> <Anchor Position="0.5" Shape="None" GraphId="b8ed6" /> <Anchor Position="0.5" Shape="None" GraphId="baa73" /> </Graphics> <Xref Database="Reactome" ID="R-HSA-2564826" /> </Interaction> <Label TextLabel="mitochondrial matrix" GraphId="a8c3d" GroupRef="c29f2"> <Graphics CenterX="292.5" CenterY="114.5" Width="139.0" Height="33.0" ZOrder="28672" FillColor="ffffff" FontSize="10" Valign="Middle" /> </Label> <Label TextLabel="cytosol" GraphId="ad30e" GroupRef="f09ec"> <Graphics CenterX="63.5" CenterY="694.5" Width="59.0" Height="33.0" ZOrder="28672" FillColor="ffffff" FontSize="10" Valign="Middle" /> </Label> <Label TextLabel="mitochondrial 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