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        <current_status>
            <date>2026-04-01</date>
            <code>REL</code>
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                            <category>citation</category>
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                            <category>em_admin</category>
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                    <metadata>
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                            <category>citation</category>
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                            <category>em_admin</category>
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        <sites>
            <deposition>PDBe</deposition>
            <last_processing>PDBe</last_processing>
        </sites>
        <key_dates>
            <deposition>2025-06-17</deposition>
            <header_release>2026-03-18</header_release>
            <map_release>2026-03-18</map_release>
            <update>2026-04-01</update>
        </key_dates>
        <grant_support>
            <grant_reference>
                <funding_body>German Research Foundation (DFG)</funding_body>
                <code>406260942</code>
                <country>Germany</country>
            </grant_reference>
            <grant_reference>
                <funding_body>German Research Foundation (DFG)</funding_body>
                <code>442219341</code>
                <country>Germany</country>
            </grant_reference>
        </grant_support>
        <title>13S+Beta1+Beta5 proteasome precursor complex</title>
        <authors_list>
            <author>Mark E</author>
            <author>Ramos PC</author>
            <author>Nunes MM</author>
            <author>Dohmen RJ</author>
            <author>Wendler P</author>
        </authors_list>
        <keywords>proteasome biogenesis, Ump1, Pba1-2, cryo-EM, HYDROLASE</keywords>
    </admin>
    <crossreferences>
        <citation_list>
            <primary_citation>
                <journal_citation published="true">
                    <author ORCID="0000-0002-2628-5230" order="1">Mark E</author>
                    <author order="2">Ramos PC</author>
                    <author order="3">Nunes MM</author>
                    <author order="4">Matias AC</author>
                    <author ORCID="0000-0002-5756-6780" order="5">Jurgen Dohmen R</author>
                    <author ORCID="0000-0002-8285-2199" order="6">Wendler P</author>
                    <title>Structural transitions in the stepwise assembly of proteasome core particles.</title>
                    <journal_abbreviation>Nat Commun</journal_abbreviation>
                    <country>UK</country>
                    <volume>17</volume>
                    <year>2026</year>
                    <external_references type="PUBMED">41876489</external_references>
                    <external_references type="DOI">doi:10.1038/s41467-026-70525-w</external_references>
                    <external_references type="ISSN">2041-1723</external_references>
                </journal_citation>
            </primary_citation>
        </citation_list>
        <pdb_list>
            <pdb_reference>
                <pdb_id>9rm1</pdb_id>
                <relationship>
                    <in_frame>FULLOVERLAP</in_frame>
                </relationship>
            </pdb_reference>
        </pdb_list>
        <other_db_list>
            <db_reference>
                <db_name>EMDB</db_name>
                <accession_id>EMD-54048</accession_id>
                <content_type>associated EM volume</content_type>
                <details>13S+Beta1+Beta5 proteasome precursor complex</details>
            </db_reference>
        </other_db_list>
    </crossreferences>
    <sample>
        <name>13S proteasome precursor complex</name>
        <supramolecule_list>
            <complex_supramolecule supramolecule_id="1">
                <name>13S proteasome precursor complex</name>
                <parent>0</parent>
                <macromolecule_list>
                    <macromolecule>
                        <macromolecule_id>1</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>2</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>3</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>4</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>5</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>6</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>7</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>8</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>9</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>10</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>11</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>12</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>13</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>14</macromolecule_id>
                    </macromolecule>
                    <macromolecule>
                        <macromolecule_id>15</macromolecule_id>
                    </macromolecule>
                </macromolecule_list>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
            </complex_supramolecule>
        </supramolecule_list>
        <macromolecule_list>
            <protein_or_peptide macromolecule_id="1">
                <name>Proteasome subunit alpha type-1</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.02803383</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MSGAAAASAAGYDRHITIFSPEGRLYQVEYAFKATNQTNINSLAVRGKDCTVVISQKKVPDKLLDPTTVSYIFCISRTIG
MVVNGPIPDARNAALRAKAEAAEFRYKYGYDMPCDVLAKRMANLSQIYTQRAYMRPLGVILTFVSVDEELGPSIYKTDPA
GYYVGYKATATGPKQQEITTNLENHFKKSKIDHINEESWEKVVEFAITHMIDALGTEFSKNDLEVGVATKDKFFTLSAEN
IEERLVAIAEQD</string>
                    <external_references type="UNIPROTKB">P21243</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="2">
                <name>Proteasome subunit alpha type-4</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.028478111</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MSGYDRALSIFSPDGHIFQVEYALEAVKRGTCAVGVKGKNCVVLGCERRSTLKLQDTRITPSKVSKIDSHVVLSFSGLNA
DSRILIEKARVEAQSHRLTLEDPVTVEYLTRYVAGVQQRYTQSGGVRPFGVSTLIAGFDPRDDEPKLYQTEPSGIYSSWS
AQTIGRNSKTVREFLEKNYDRKEPPATVEECVKLTVRSLLEVVQTGAKNIEITVVKPDSDIVALSSEEINQYVTQIEQEK
QEQQEQDKKKKSNH</string>
                    <external_references type="UNIPROTKB">P40303</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="3">
                <name>Proteasome subunit alpha type-5</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.028649085999999997</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MFLTRSEYDRGVSTFSPEGRLFQVEYSLEAIKLGSTAIGIATKEGVVLGVEKRATSPLLESDSIEKIVEIDRHIGCAMSG
LTADARSMIEHARTAAVTHNLYYDEDINVESLTQSVCDLALRFGEGASGEERLMSRPFGVALLIAGHDADDGYQLFHAEP
SGTFYRYNAKAIGSGSEGAQAELLNEWHSSLTLKEAELLVLKILKQVMEEKLDENNAQLSCITKQDGFKIYDNEKTAELI
KELKEKEAAESPEEADVEMS</string>
                    <external_references type="UNIPROTKB">P32379</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="4">
                <name>Proteasome subunit alpha type-6</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.025634</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MFRNNYDGDTVTFSPTGRLFQVEYALEAIKQGSVTVGLRSNTHAVLVALKRNADELSSYQKKIIKCDEHMGLSLAGLAPD
ARVLSNYLRQQCNYSSLVFNRKLAVERAGHLLCDKAQKNTQSYGGRPYGVGLLIIGYDKSGAHLLEFQPSGNVTELYGTA
IGARSQGAKTYLERTLDTFIKIDGNPDELIKAGVEAISQSLRDESLTVDNLSIAIVGKDTPFTIYDGEAVAKYI</string>
                    <external_references type="UNIPROTKB">P40302</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="5">
                <name>Proteasome subunit beta type-1</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.023573603999999998</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MNGIQVDINRLKKGEVSLGTSIMAVTFKDGVILGADSRTTTGAYIANRVTDKLTRVHDKIWCCRSGSAADTQAIADIVQY
HLELYTSQYGTPSTETAASVFKELCYENKDNLTAGIIVAGYDDKNKGEVYTIPLGGSVHKLPYAIAGSGSTFIYGYCDKN
FRENMSKEETVDFIKHSLSQAIKWDGSSGGVIRMVVLTAAGVERLIFYPDEYEQL</string>
                    <external_references type="UNIPROTKB">P38624</external_references>
                </sequence>
                <ec_number>3.4.25.1</ec_number>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="6">
                <name>Proteasome maturation factor UMP1</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.018603623</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <recombinant_expression database="NCBI">
                    <recombinant_organism ncbi="4932">Saccharomyces cerevisiae</recombinant_organism>
                </recombinant_expression>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>DYKDDDDKHHHHHHMNIVPQDTFKSQVSTDQDKSVLSSAVPSLPDTLRQQEGGAVPLSTQLNDRHPLESTLKNWETTQRQ
RQMEQYRQIFGIAEPMKRTMEMEIVNRTDFNPLSTNGSIHRDILLNKECSIDWEDVYPGTGLQASTMVGDDVHSKIEKQL
GI</string>
                    <external_references type="UNIPROTKB">P38293</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="7">
                <name>Proteasome chaperone 1</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.030718073999999998</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MLFKQWNDLPEPKHLLDLPEISKNLQSLEVCPVPKVEFPQDLDVPQYSTAVITTKIMNPLFPKNLLQLTSIGEIKTTLTV
KSPSLPQSSGKHSWNYDENFPNEVDPDQKNDTADETVYGFSFPIYSFGKTLLFSMEENFISISPIFGNMISRSIISQLAQ
FSPDIIVIGTSDKIASMKVMTENECTLQPPEFITGFIGSVLTQLIVGPSKGLKFKCLVAPSEGPNGFEKLSLSDMGSLVD
LCGQWLGFEPSRYSEECYRLWRCDSAAIGAQSGLYI</string>
                    <external_references type="UNIPROTKB">Q05778</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="8">
                <name>Proteasome assembly chaperone 2</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.030762895</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MSCLVLPLVSVGNIPQLSIDWLLNSQANEWEYLEALDSKYLVEFVGPLDRPEDGSDSLYKDADMKYSSALEVFYNKKRGL
FAIQQRTPLVSVNYLNNFIVEIILPFLSKYNISEICIWDSLYAMEDENGVIVRPQEVYSLGEFYFDDEAELLSNLHLNDQ
ESMVNNWLHFTPTSFQDKISVDQPIFKILFQILNASQRPKALRSIKYCSCLANEGDNSLDSQQFLQWIISQKVIKNAPPI
VKFVRPISWQGAYGMADARDKFVDLYN</string>
                    <external_references type="UNIPROTKB">P36040</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="9">
                <name>Proteasome subunit alpha type-2</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.027191828</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MTDRYSFSLTTFSPSGKLGQIDYALTAVKQGVTSLGIKATNGVVIATEKKSSSPLAMSETLSKVSLLTPDIGAVYSGMGP
DYRVLVDKSRKVAHTSYKRIYGEYPPTKLLVSEVAKIMQEATQSGGVRPFGVSLLIAGHDEFNGFSLYQVDPSGSYFPWK
ATAIGKGSVAAKTFLEKRWNDELELEDAIHIALLTLKESVEGEFNGDTIELAIIGDENPDLLGYTGIPTDKGPRFRKLTS
QEINDRLEAL</string>
                    <external_references type="UNIPROTKB">P23639</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="10">
                <name>Proteasome subunit alpha type-3</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.02874823</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MGSRRYDSRTTIFSPEGRLYQVEYALESISHAGTAIGIMASDGIVLAAERKVTSTLLEQDTSTEKLYKLNDKIAVAVAGL
TADAEILINTARIHAQNYLKTYNEDIPVEILVRRLSDIKQGYTQHGGLRPFGVSFIYAGYDDRYGYQLYTSNPSGNYTGW
KAISVGANTSAAQTLLQMDYKDDMKVDDAIELALKTLSKTTDSSALTYDRLEFATIRKGANDGEVYQKIFKPQEIKDILV
KTGITKKDEDEEADEDMK</string>
                    <external_references type="UNIPROTKB">P23638</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="11">
                <name>Probable proteasome subunit alpha type-7</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.031575068</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MTSIGTGYDLSNSVFSPDGRNFQVEYAVKAVENGTTSIGIKCNDGVVFAVEKLITSKLLVPQKNVKIQVVDRHIGCVYSG
LIPDGRHLVNRGREEAASFKKLYKTPIPIPAFADRLGQYVQAHTLYNSVRPFGVSTIFGGVDKNGAHLYMLEPSGSYWGY
KGAATGKGRQSAKAELEKLVDHHPEGLSAREAVKQAAKIIYLAHEDNKEKDFELEISWCSLSETNGLHKFVKGDLLQEAI
DFAQKEINGDDDEDEDDSDNVMSSDDENAPVATNANATTDQEGDIHLE</string>
                    <external_references type="UNIPROTKB">P21242</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="12">
                <name>Proteasome subunit beta type-2</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.028299888999999998</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MAGLSFDNYQRNNFLAENSHTQPKATSTGTTIVGVKFNNGVVIAADTRSTQGPIVADKNCAKLHRISPKIWCAGAGTAAD
TEAVTQLIGSNIELHSLYTSREPRVVSALQMLKQHLFKYQGHIGAYLIVAGVDPTGSHLFSIHAHGSTDVGYYLSLGSGS
LAAMAVLESHWKQDLTKEEAIKLASDAIQAGIWNDLGSGSNVDVCVMEIGKDAEYLRNYLTPNVREEKQKSYKFPRGTTA
VLKESIVNICDIQEEQVDITA</string>
                    <external_references type="UNIPROTKB">P25043</external_references>
                </sequence>
                <ec_number>3.4.25.1</ec_number>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="13">
                <name>Proteasome subunit beta type-3</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.022627842</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MSDPSSINGGIVVAMTGKDCVAIACDLRLGSQSLGVSNKFEKIFHYGHVFLGITGLATDVTTLNEMFRYKTNLYKLKEER
AIEPETFTQLVSSSLYERRFGPYFVGPVVAGINSKSGKPFIAGFDLIGCIDEAKDFIVSGTASDQLFGMCESLYEPNLEP
EDLFETISQALLNAADRDALSGWGAVVYIIKKDEVVKRYLKMRQD</string>
                    <external_references type="UNIPROTKB">P25451</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="14">
                <name>Proteasome subunit beta type-4</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.022545675999999997</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MDIILGIRVQDSVILASSKAVTRGISVLKDSDDKTRQLSPHTLMSFAGEAGDTVQFAEYIQANIQLYSIREDYELSPQAV
SSFVRQELAKSIRSRRPYQVNVLIGGYDKKKNKPELYQIDYLGTKVELPYGAHGYSGFYTFSLLDHHYRPDMTTEEGLDL
LKLCVQELEKRMPMDFKGVIVKIVDKDGIRQVDDFQAQ</string>
                    <external_references type="UNIPROTKB">P22141</external_references>
                </sequence>
            </protein_or_peptide>
            <protein_or_peptide macromolecule_id="15">
                <name>Proteasome subunit beta type-5</name>
                <natural_source database="NCBI">
                    <organism ncbi="4932">Saccharomyces cerevisiae</organism>
                </natural_source>
                <molecular_weight>
                    <theoretical units="MDa">0.031670539</theoretical>
                </molecular_weight>
                <number_of_copies>1</number_of_copies>
                <enantiomer>LEVO</enantiomer>
                <sequence>
                    <string>MQAIADSFSVPNRLVKELQYDNEQNLESDFVTGASQFQRLAPSLTVPPIASPQQFLRAHTDDSRNPDCKIKIAHGTTTLA
FRFQGGIIVAVDSRATAGNWVASQTVKKVIEINPFLLGTMAGGAADCQFWETWLGSQCRLHELREKERISVAAASKILSN
LVYQYKGAGLSMGTMICGYTRKEGPTIYYVDSDGTRLKGDIFCVGSGQTFAYGVLDSNYKWDLSVEDALYLGKRSILAAA
HRDAYSGGSVNLYHVTEDGWIYHGNHDVGELFWKVKEEEGSFNNVIG</string>
                    <external_references type="UNIPROTKB">P30656</external_references>
                </sequence>
                <ec_number>3.4.25.1</ec_number>
            </protein_or_peptide>
        </macromolecule_list>
    </sample>
    <structure_determination_list>
        <structure_determination structure_determination_id="1">
            <method>singleParticle</method>
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50 mM Tris-HCl</details>
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