{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T13:00:16Z","timestamp":1791378016156,"version":"4.3.2"},"reference-count":91,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"name":"Secretar\u00eda de Ciencia, Humanidades, Tecnolog\u00eda e Innovaci\u00f3n","award":["CF-2023-I-1545"],"award-info":[{"award-number":["CF-2023-I-1545"]}],"id":[{"id":"https:\/\/ror.org\/059ex5q34","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100006087","name":"Direcci\u00f3n General de Asuntos del Personal Acad\u00e9mico, Universidad Nacional Aut\u00f3noma de M\u00e9xico","doi-asserted-by":"publisher","award":["IN208921"],"award-info":[{"award-number":["IN208921"]}],"id":[{"id":"10.13039\/501100006087","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100006087","name":"Direcci\u00f3n General de Asuntos del Personal Acad\u00e9mico, Universidad Nacional Aut\u00f3noma de M\u00e9xico","doi-asserted-by":"publisher","award":["IN218424"],"award-info":[{"award-number":["IN218424"]}],"id":[{"id":"10.13039\/501100006087","id-type":"DOI","asserted-by":"publisher"}]},{"name":"SECIHTI","award":["CVU: 1033319"],"award-info":[{"award-number":["CVU: 1033319"]}],"id":[{"id":"https:\/\/ror.org\/059ex5q34","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Programa de Maestr\u00eda y Doctorado en Ciencias Bioqu\u00edmicas"},{"DOI":"10.13039\/501100005739","name":"Universidad Nacional Aut\u00f3noma de M\u00e9xico","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100005739","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,7]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Mitochondrial biogenesis requires the coordinated synthesis, targeting, and import of nuclear-encoded mitochondrial precursor proteins. Although ribosome-associated chaperones support co-translational protein folding, their genetic contributions to mitochondrial protein import and cellular homeostasis remain incompletely defined. Here, we investigate the roles of the nascent polypeptide\u2013associated complex (NAC) and the ribosome-associated Hsp70 system Ssb1\/2 in Saccharomyces cerevisiae. We show that NAC and Ssb1\/2 have distinct yet partially overlapping functions in the handling of mitochondrial precursor proteins. Loss of NAC activates the mitochondrial retrograde pathway and enhances growth on ethanol as a nonfermentable carbon source without compromising respiratory competence, indicating metabolic adaptation rather than overt mitochondrial dysfunction. In contrast, Ssb1\/2 deficiency disrupts cytosolic proteostasis, sensitizes cells to TORC1 inhibition, and impairs autophagy and mitophagy. Using a tobacco etch virus protease\u2013based import reporter, we show that Ssb1\/2 promotes efficient co-translational distribution of precursor proteins, whereas NAC limits the accumulation of misfolded proteins at the mitochondrial surface. Biochemical analyses further reveal that Ssb1\/2 supports the association of translating cytosolic ribosomes with the mitochondrial outer membrane, while NAC loss partially restores this interaction in the absence of Ssb1\/2. Together, these findings establish NAC and Ssb1\/2 as key components of an integrated network linking co-translational targeting, mitochondrial signaling, and cellular homeostasis.<\/jats:p>","DOI":"10.1093\/genetics\/iyag200","type":"journal-article","created":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T08:46:15Z","timestamp":1785573975000},"source":"Crossref","is-referenced-by-count":0,"title":["Ribosome-associated chaperones integrate co-translational mitochondrial protein import, stress signaling, and proteostasis in\n                    <i>Saccharomyces cerevisiae<\/i>"],"prefix":"10.1093","volume":"234","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-4815-6163","authenticated-orcid":false,"given":"J Ernesto","family":"Bravo-Ar\u00e9valo","sequence":"first","affiliation":[{"name":"Departamento de Gen\u00e9tica Molecular, Instituto de Fisiolog\u00eda Celular, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Coyoac\u00e1n, Cd. Mx. 04510 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ariann E","family":"Mendoza-Mart\u00ednez","sequence":"additional","affiliation":[{"name":"Departamento de Gen\u00e9tica Molecular, Instituto de Fisiolog\u00eda Celular, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Coyoac\u00e1n, Cd. Mx. 04510 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Teresa","family":"Ballado","sequence":"additional","affiliation":[{"name":"Departamento de Gen\u00e9tica Molecular, Instituto de Fisiolog\u00eda Celular, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Coyoac\u00e1n, Cd. Mx. 04510 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ericka","family":"Moreno-M\u00e9ndez","sequence":"additional","affiliation":[{"name":"Centro de Investigaci\u00f3n Sobre el Envejecimiento, Cinvestav , Tlalpan, Cd. Mx. 14330 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9236-2804","authenticated-orcid":false,"given":"Alexander","family":"DeLuna","sequence":"additional","affiliation":[{"name":"Centro de Investigaci\u00f3n Sobre el Envejecimiento, Cinvestav , Tlalpan, Cd. Mx. 14330 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1815-1977","authenticated-orcid":false,"given":"Soledad","family":"Funes","sequence":"additional","affiliation":[{"name":"Departamento de Gen\u00e9tica Molecular, Instituto de Fisiolog\u00eda Celular, Universidad Nacional Aut\u00f3noma de M\u00e9xico , Coyoac\u00e1n, Cd. Mx. 04510 ,","place":["M\u00e9xico"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,8,1]]},"reference":[{"key":"2026100708030396100_iyag200-B1","doi-asserted-by":"publisher","first-page":"519","DOI":"10.3390\/cells12040519","article-title":"Interplays of AMPK and TOR in autophagy regulation in yeast","volume":"12","author":"Alao","year":"2023","journal-title":"Cells"},{"key":"2026100708030396100_iyag200-B2","doi-asserted-by":"publisher","first-page":"7120","DOI":"10.1021\/acs.biochem.5b00801","article-title":"Specific binding of tetratricopeptide repeat proteins to heat shock protein 70 (Hsp70) and heat shock protein 90 (Hsp90) is regulated by affinity and phosphorylation","volume":"54","author":"Assimon","year":"2015","journal-title":"Biochemistry"},{"key":"2026100708030396100_iyag200-B3","doi-asserted-by":"publisher","first-page":"101984","DOI":"10.1016\/j.jbc.2022.101984","article-title":"Positively charged amino acids at the N terminus of select mitochondrial proteins mediate early recognition by import proteins \u03b1\u03b2'-NAC and Sam37","volume":"298","author":"Avenda\u00f1o-Monsalve","year":"2022","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B4","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1515\/hsz-2020-0110","article-title":"From cytosol to mitochondria: the beginning of a protein journey","volume":"401","author":"Avenda\u00f1o-Monsalve","year":"2020","journal-title":"Biol Chem."},{"key":"2026100708030396100_iyag200-B5","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/s00294-019-01035-0","article-title":"Cell organelles and yeast longevity: an intertwined regulation","volume":"66","author":"Banerjee","year":"2020","journal-title":"Curr Genet."},{"key":"2026100708030396100_iyag200-B6","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1091\/mbc.E19-06-0329","article-title":"Msp1 cooperates with the proteasome for extraction of arrested mitochondrial import intermediates","volume":"31","author":"Basch","year":"2020","journal-title":"Mol Biol Cell."},{"key":"2026100708030396100_iyag200-B7","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1007\/978-1-59745-261-8_7","article-title":"Synthesis and sorting of mitochondrial translation products","volume":"457","author":"Bauerschmitt","year":"2008","journal-title":"Methods Mol Biol"},{"key":"2026100708030396100_iyag200-B8","doi-asserted-by":"publisher","first-page":"5528","DOI":"10.3390\/ijms16035528","article-title":"Mechanisms by which different functional states of mitochondria define yeast longevity","volume":"16","author":"Beach","year":"2015","journal-title":"Int J Mol Sci."},{"key":"2026100708030396100_iyag200-B9","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1126\/science.1212642","article-title":"The structure of the eukaryotic ribosome at 3.0 \u00c5 resolution","volume":"334","author":"Ben-Shem","year":"2011","journal-title":"Science"},{"key":"2026100708030396100_iyag200-B10","doi-asserted-by":"publisher","DOI":"10.15252\/embj.2022113240","article-title":"The ribosome-associated chaperone Zuo1 controls translation upon TORC1 inhibition","volume":"42","author":"Black","year":"2023","journal-title":"EMBO J."},{"key":"2026100708030396100_iyag200-B11","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1038\/s41556-019-0294-5","article-title":"Mitochondrial protein-induced stress triggers a global adaptive transcriptional programme","volume":"21","author":"Boos","year":"2019","journal-title":"Nat Cell Biol."},{"key":"2026100708030396100_iyag200-B12","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.tcb.2019.12.003","article-title":"How the mitoprotein-induced stress response safeguards the cytosol: a unified view","volume":"30","author":"Boos","year":"2020","journal-title":"Trends Cell Biol."},{"key":"2026100708030396100_iyag200-B13","doi-asserted-by":"publisher","first-page":"2503","DOI":"10.1091\/mbc.E04-11-0968","article-title":"Homeostatic adjustment and metabolic remodeling in glucose-limited yeast cultures","volume":"16","author":"Brauer","year":"2005","journal-title":"Mol Biol Cell."},{"key":"2026100708030396100_iyag200-B14","doi-asserted-by":"publisher","first-page":"5026","DOI":"10.1111\/febs.70109","article-title":"Tracing the evolutionary pathway: on the origin of mitochondria and eukaryogenesis","volume":"292","author":"Bravo-Ar\u00e9valo","year":"2025","journal-title":"FEBS J."},{"key":"2026100708030396100_iyag200-B15","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1534\/genetics.111.135731","article-title":"Nutritional control of growth and development in yeast","volume":"192","author":"Broach","year":"2012","journal-title":"Genetics"},{"key":"2026100708030396100_iyag200-B16","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/S1097-2765(04)00179-0","article-title":"Mitochondrial signaling: the retrograde response","volume":"14","author":"Butow","year":"2004","journal-title":"Mol Cell."},{"key":"2026100708030396100_iyag200-B17","doi-asserted-by":"publisher","first-page":"628","DOI":"10.1016\/j.cell.2009.08.005","article-title":"Importing mitochondrial proteins: machineries and mechanisms","volume":"138","author":"Chacinska","year":"2009","journal-title":"Cell"},{"key":"2026100708030396100_iyag200-B18","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.202407110","article-title":"Cytoplasmic ribosomes on mitochondria alter the local membrane environment for protein import","volume":"224","author":"Chang","year":"2025","journal-title":"J Cell Biol."},{"key":"2026100708030396100_iyag200-B19","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1007\/s10930-019-09847-2","article-title":"The principles of protein targeting and transport across cell membranes","volume":"38","author":"Chen","year":"2019","journal-title":"Protein J."},{"key":"2026100708030396100_iyag200-B20","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/B978-0-12-407672-3.00001-0","article-title":"Yeast petites and small colony variants: for everything there is a season","volume":"85","author":"Day","year":"2013","journal-title":"Adv Appl Microbiol."},{"key":"2026100708030396100_iyag200-B21","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001100","article-title":"Defining the specificity of cotranslationally acting chaperones by systematic analysis of MRNAs associated with ribosome-nascent chain complexes","volume":"9","author":"del Alamo","year":"2011","journal-title":"PLoS Biol."},{"key":"2026100708030396100_iyag200-B22","doi-asserted-by":"publisher","first-page":"3125","DOI":"10.1042\/BCJ20190584","article-title":"Quality control of protein import into mitochondria","volume":"478","author":"den Brave","year":"2021","journal-title":"Biochem J."},{"key":"2026100708030396100_iyag200-B23","doi-asserted-by":"publisher","first-page":"a033977","DOI":"10.1101\/cshperspect.a033977","article-title":"Chaperone interactions at the ribosome","volume":"11","author":"Deuerling","year":"2019","journal-title":"Cold Spring Harb Perspect Biol."},{"key":"2026100708030396100_iyag200-B24","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1016\/j.cell.2017.06.038","article-title":"Profiling Ssb-nascent chain interactions reveals principles of Hsp70-assisted folding","volume":"170","author":"D\u00f6ring","year":"2017","journal-title":"Cell"},{"key":"2026100708030396100_iyag200-B25","doi-asserted-by":"publisher","first-page":"3289","DOI":"10.1091\/mbc.10.10.3289","article-title":"Nascent polypeptide-associated complex stimulates protein import into yeast mitochondria","volume":"10","author":"F\u00fcnfschilling","year":"1999","journal-title":"Mol Biol Cell."},{"key":"2026100708030396100_iyag200-B26","doi-asserted-by":"publisher","first-page":"1551","DOI":"10.1261\/rna.2621111","article-title":"Localization of MRNAs coding for mitochondrial proteins in the yeast Saccharomyces cerevisiae","volume":"17","author":"Gadir","year":"2011","journal-title":"RNA"},{"key":"2026100708030396100_iyag200-B27","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1126\/science.aaa5335","article-title":"The principle of antagonism ensures protein targeting specificity at the endoplasmic reticulum","volume":"348","author":"Gamerdinger","year":"2015","journal-title":"Science"},{"key":"2026100708030396100_iyag200-B28","doi-asserted-by":"publisher","first-page":"2296","DOI":"10.1073\/pnas.95.5.2296","article-title":"The yeast nascent polypeptide-associated complex initiates protein targeting to mitochondria in vivo","volume":"95","author":"George","year":"1998","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2026100708030396100_iyag200-B29","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/S0014-5793(02)02528-0","article-title":"The nascent polypeptide-associated complex (NAC) promotes interaction of ribosomes with the mitochondrial surface in vivo","volume":"516","author":"George","year":"2002","journal-title":"FEBS Lett."},{"key":"2026100708030396100_iyag200-B30","doi-asserted-by":"publisher","first-page":"1786","DOI":"10.15252\/embr.201744261","article-title":"Visualization of cytosolic ribosomes on the surface of mitochondria by electron cryo-tomography","volume":"18","author":"Gold","year":"2017","journal-title":"EMBO Rep."},{"key":"2026100708030396100_iyag200-B31","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/j.ymeth.2014.11.023","article-title":"Assays for the biochemical and ultrastructural measurement of selective and nonselective types of autophagy in the yeast Saccharomyces cerevisiae","volume":"75","author":"Guimaraes","year":"2015","journal-title":"Methods"},{"key":"2026100708030396100_iyag200-B32","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1016\/j.ymeth.2009.04.003","article-title":"Affinity purification of ribosomes to access the translatome","volume":"48","author":"Halbeisen","year":"2009","journal-title":"Methods"},{"key":"2026100708030396100_iyag200-B33","doi-asserted-by":"publisher","first-page":"13695","DOI":"10.1038\/ncomms13695","article-title":"Multivalent contacts of the Hsp70 Ssb contribute to its architecture on ribosomes and nascent chain interaction","volume":"7","author":"Hanebuth","year":"2016","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B34","doi-asserted-by":"publisher","first-page":"330","DOI":"10.1007\/s10930-019-09819-6","article-title":"Transport of proteins into mitochondria","volume":"38","author":"Hansen","year":"2019","journal-title":"Protein J."},{"key":"2026100708030396100_iyag200-B35","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1000407","article-title":"Computationally driven, quantitative experiments discover genes required for mitochondrial biogenesis","volume":"5","author":"Hess","year":"2009","journal-title":"PLoS Genet."},{"key":"2026100708030396100_iyag200-B36","doi-asserted-by":"publisher","first-page":"jcs241539","DOI":"10.1242\/JCS.241539","article-title":"Retrograde signaling mediates an adaptive survival response to endoplasmic reticulum stress in Saccharomyces Cerevisiae","volume":"133","author":"Hijazi","year":"2020","journal-title":"J Cell Sci."},{"key":"2026100708030396100_iyag200-B37","doi-asserted-by":"publisher","first-page":"481","DOI":"10.1111\/1567-1364.12125","article-title":"Management of the endoplasmic reticulum stress by activation of the heat shock response in yeast","volume":"14","author":"Hou","year":"2014","journal-title":"FEMS Yeast Res."},{"key":"2026100708030396100_iyag200-B38","doi-asserted-by":"publisher","first-page":"686","DOI":"10.1038\/nature02026","article-title":"Global analysis of protein localization in budding yeast","volume":"425","author":"Huh","year":"2003","journal-title":"Nature"},{"key":"2026100708030396100_iyag200-B39","doi-asserted-by":"publisher","first-page":"10471","DOI":"10.1038\/ncomms10471","article-title":"Structures of the E. coli translating ribosome with SRP and its receptor and with the translocon","volume":"7","author":"Jomaa","year":"2016","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B40","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1083\/jcb.65.1.1","article-title":"Cytoplasmic type 80S ribosomes associated with yeast mitochondria. IV. Attachment of ribosomes to the outer membrane of isolated mitochondria","volume":"65","author":"Kellems","year":"1975","journal-title":"J Cell Biol."},{"key":"2026100708030396100_iyag200-B41","doi-asserted-by":"publisher","first-page":"114473","DOI":"10.1016\/j.celrep.2024.114473","article-title":"ATAD1 prevents clogging of TOM and damage caused by un-imported mitochondrial proteins","volume":"43","author":"Kim","year":"2024","journal-title":"Cell Rep."},{"key":"2026100708030396100_iyag200-B42","doi-asserted-by":"publisher","first-page":"878","DOI":"10.1111\/1462-2920.13617","article-title":"Glycerol metabolism and transport in yeast and fungi: established knowledge and ambiguities","volume":"19","author":"Klein","year":"2017","journal-title":"Environ Microbiol."},{"key":"2026100708030396100_iyag200-B43","doi-asserted-by":"publisher","first-page":"25587","DOI":"10.1074\/jbc.273.40.25587","article-title":"Import into mitochondria, folding and retrograde movement of fumarase in yeast","volume":"273","author":"Knox","year":"1998","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B44","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1083\/jcb.200910074","article-title":"A dual function for chaperones SSB-RAC and the NAC nascent polypeptide-associated Complex on ribosomes","volume":"189","author":"Koplin","year":"2010","journal-title":"J Cell Biol."},{"key":"2026100708030396100_iyag200-B45","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3390\/ijms21197262","article-title":"The mitochondrial outer membrane protein Tom70-mediator in protein traffic, membrane contact sites and innate immunity","volume":"21","author":"Kreimendahl","year":"2020","journal-title":"Int J Mol Sci."},{"key":"2026100708030396100_iyag200-B46","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1016\/j.bbamcr.2006.09.009","article-title":"A central role for the peroxisomal membrane in glyoxylate cycle function","volume":"1763","author":"Kunze","year":"2006","journal-title":"Biochim Biophys Acta."},{"key":"2026100708030396100_iyag200-B47","doi-asserted-by":"publisher","first-page":"680","DOI":"10.1038\/227680a0","article-title":"Cleavage of structural proteins during the assembly of the head of bacteriophage T4","volume":"227","author":"Laemmli","year":"1970","journal-title":"Nature"},{"key":"2026100708030396100_iyag200-B48","doi-asserted-by":"publisher","first-page":"2505","DOI":"10.1091\/mbc.E16-03-0187","article-title":"Adaptation to different types of stress converge on mitochondrial metabolism","volume":"27","author":"Lahtvee","year":"2016","journal-title":"Mol Biol Cell."},{"key":"2026100708030396100_iyag200-B49","doi-asserted-by":"publisher","first-page":"3648","DOI":"10.1016\/j.febslet.2013.10.004","article-title":"Rim15-dependent activation of Hsf1 and Msn2\/4 transcription factors by direct phosphorylation in Saccharomyces cerevisiae","volume":"587","author":"Lee","year":"2013","journal-title":"FEBS Lett."},{"key":"2026100708030396100_iyag200-B50","doi-asserted-by":"publisher","first-page":"5711","DOI":"10.1038\/ncomms6711","article-title":"OM14 is a mitochondrial receptor for cytosolic ribosomes that supports co-translational import into mitochondria","volume":"5","author":"Lesnik","year":"2014","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B51","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1128\/MCB.11.1.38","article-title":"Intramitochondrial functions regulate nonmitochondrial citrate synthase (CIT2) expression in Saccharomyces cerevisiae","volume":"11","author":"Liao","year":"1991","journal-title":"Mol Cell Biol."},{"key":"2026100708030396100_iyag200-B52","doi-asserted-by":"publisher","first-page":"6720","DOI":"10.1128\/mcb.19.10.6720","article-title":"A transcriptional switch in the expression of yeast tricarboxylic acid cycle genes in response to a reduction or loss of respiratory function","volume":"19","author":"Liu","year":"1999","journal-title":"Mol Cell Biol."},{"key":"2026100708030396100_iyag200-B53","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1146\/annurev.genet.40.110405.090613","article-title":"Mitochondrial retrograde signaling","volume":"40","author":"Liu","year":"2006","journal-title":"Annu Rev Genet."},{"key":"2026100708030396100_iyag200-B54","doi-asserted-by":"publisher","first-page":"22715","DOI":"10.1074\/jbc.M504257200","article-title":"Manganese activation of superoxide dismutase 2 in the mitochondria of Saccharomyces cerevisiae","volume":"280","author":"Luk","year":"2005","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B55","doi-asserted-by":"publisher","first-page":"214","DOI":"10.1179\/135100009X12525712409454","article-title":"Buffer modulation of menadione-induced oxidative stress in Saccharomyces cerevisiae","volume":"14","author":"Lushchak","year":"2009","journal-title":"Redox Rep."},{"key":"2026100708030396100_iyag200-B56","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.47791","article-title":"Cytoplasmic protein misfolding titrates Hsp70 to activate nuclear Hsf1","volume":"8","author":"Masser","year":"2019","journal-title":"ELife"},{"key":"2026100708030396100_iyag200-B57","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/j.molcel.2019.07.006","article-title":"Msp1 clears mistargeted proteins by facilitating their transfer from mitochondria to the ER","volume":"76","author":"Matsumoto","year":"2019","journal-title":"Mol Cell."},{"key":"2026100708030396100_iyag200-B58","doi-asserted-by":"publisher","DOI":"10.21769\/BioProtoc.5610","article-title":"Optimized mechanical isolation of mitochondria from Saccharomyces cerevisiae preserving Atg32 for quantitative analysis","volume":"16","author":"Mendoza-Mart\u00ednez","year":"2026","journal-title":"Bio Protoc."},{"key":"2026100708030396100_iyag200-B59","doi-asserted-by":"publisher","first-page":"102","DOI":"10.3389\/fgene.2011.00102","article-title":"Loss of mitochondrial membrane potential triggers the retrograde response extending yeast replicative lifespan","volume":"2","author":"Miceli","year":"2011","journal-title":"Front Genet."},{"key":"2026100708030396100_iyag200-B60","doi-asserted-by":"publisher","first-page":"1714","DOI":"10.3390\/cells11101714","article-title":"Yeast chronological lifespan: longevity regulatory genes and mechanisms","volume":"11","author":"Mirisola","year":"2022","journal-title":"Cells"},{"key":"2026100708030396100_iyag200-B61","doi-asserted-by":"publisher","first-page":"8256","DOI":"10.1038\/ncomms9256","article-title":"TORC1 controls G1-S cell cycle transition in yeast via Mpk1 and the greatwall kinase pathway","volume":"6","author":"Moreno-Torres","year":"2015","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B62","doi-asserted-by":"publisher","first-page":"937","DOI":"10.1038\/s41467-017-00635-z","article-title":"The Hsp70 homolog Ssb affects ribosome biogenesis via the TORC1-Sch9 signaling pathway","volume":"8","author":"Mudholkar","year":"2017","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B63","doi-asserted-by":"publisher","first-page":"4347","DOI":"10.1128\/mcb.21.13.4347-4368.2001","article-title":"Transcriptional profiling shows that Gcn4p is a master regulator of gene expression during amino acid starvation in yeast","volume":"21","author":"Natarajan","year":"2001","journal-title":"Mol Cell Biol."},{"key":"2026100708030396100_iyag200-B64","doi-asserted-by":"publisher","first-page":"1075","DOI":"10.1016\/S0092-8674(05)80012-1","article-title":"SRP samples nascent chains for the presence of signal sequences by interacting with ribosomes at a discrete step during translation elongation","volume":"81","author":"Ogg","year":"1995","journal-title":"Cell"},{"key":"2026100708030396100_iyag200-B65","doi-asserted-by":"publisher","first-page":"928","DOI":"10.1126\/science.125.3254.928","article-title":"Tetrazolium overlay technique for population studies of respiration deficiency in yeast","volume":"125","author":"Ogur","year":"1957","journal-title":"Science"},{"key":"2026100708030396100_iyag200-B66","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0143457","article-title":"Functional dissection of the nascent polypeptide-associated complex in Saccharomyces cerevisiae","volume":"10","author":"Ott","year":"2015","journal-title":"PLoS One"},{"key":"2026100708030396100_iyag200-B67","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1038\/s41580-018-0092-0","article-title":"Mitochondrial proteins: from biogenesis to functional networks","volume":"20","author":"Pfanner","year":"2019","journal-title":"Nat Rev Mol Cell Biol."},{"key":"2026100708030396100_iyag200-B68","doi-asserted-by":"publisher","first-page":"814","DOI":"10.1111\/febs.14024","article-title":"\u0391\u03b2\u2032-NAC cooperates with Sam37 to mediate early stages of mitochondrial protein import","volume":"284","author":"Ponce-Rojas","year":"2017","journal-title":"FEBS J."},{"key":"2026100708030396100_iyag200-B69","doi-asserted-by":"publisher","first-page":"13004","DOI":"10.1074\/jbc.M414093200","article-title":"Evolution of mitochondrial oxa proteins from bacterial YidC. Inherited and acquired functions of a conserved protein insertion machinery","volume":"280","author":"Preuss","year":"2005","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B70","doi-asserted-by":"publisher","first-page":"3211","DOI":"10.1111\/febs.70356","article-title":"The ribosome-associated complex regulates cytosolic translation upon mitoprotein-induced stress","volume":"293","author":"Qian","year":"2026","journal-title":"FEBS J."},{"key":"2026100708030396100_iyag200-B71","doi-asserted-by":"publisher","first-page":"2253","DOI":"10.1038\/nprot.2006.397","article-title":"Identification of fungal cell wall mutants using susceptibility assays based on calcofluor white and Congo red","volume":"1","author":"Ram","year":"2006","journal-title":"Nat Protoc."},{"key":"2026100708030396100_iyag200-B72","doi-asserted-by":"publisher","first-page":"9750","DOI":"10.1126\/sciadv.adn9750","article-title":"NAC regulates metabolism and cell fate in intestinal stem cells","volume":"11","author":"Ramalho","year":"2025","journal-title":"Sci Adv."},{"key":"2026100708030396100_iyag200-B73","doi-asserted-by":"publisher","first-page":"384","DOI":"10.3390\/biology10050384","article-title":"The unfolded protein response: an overview","volume":"10","author":"Read","year":"2021","journal-title":"Biology (Basel)."},{"key":"2026100708030396100_iyag200-B74","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1111\/j.1365-2958.2009.06659.x","article-title":"Dual localization of fumarase is dependent on the integrity of the glyoxylate shunt","volume":"72","author":"Regev-Rudzki","year":"2009","journal-title":"Mol Microbiol."},{"key":"2026100708030396100_iyag200-B75","doi-asserted-by":"publisher","first-page":"R1177","DOI":"10.1016\/j.cub.2017.09.015","article-title":"The origin and diversification of mitochondria","volume":"27","author":"Roger","year":"2017","journal-title":"Curr Biol."},{"key":"2026100708030396100_iyag200-B76","doi-asserted-by":"publisher","first-page":"2763","DOI":"10.1093\/molbev\/msm211","article-title":"The tetratricopeptide repeats of receptors involved in protein translocation across membranes","volume":"24","author":"Schlegel","year":"2007","journal-title":"Mol Biol Evol."},{"key":"2026100708030396100_iyag200-B77","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1515\/hsz-2019-0444","article-title":"Evolution of mitochondrial protein import\u2014lessons from trypanosomes","volume":"401","author":"Schneider","year":"2020","journal-title":"Biol Chem."},{"key":"2026100708030396100_iyag200-B78","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1007\/s00294-003-0381-8","article-title":"Transcriptional control of nonfermentative metabolism in the yeast Saccharomyces cerevisiae","volume":"43","author":"Schuller","year":"2003","journal-title":"Curr Genet."},{"key":"2026100708030396100_iyag200-B79","doi-asserted-by":"publisher","first-page":"6491","DOI":"10.1038\/s41598-019-43032-w","article-title":"Carbon catabolite repression in yeast is not limited to glucose","volume":"9","author":"Simpson-Lavy","year":"2019","journal-title":"Sci Rep."},{"key":"2026100708030396100_iyag200-B80","doi-asserted-by":"publisher","first-page":"60","DOI":"10.1016\/j.molcel.2016.05.014","article-title":"Defining the essential function of yeast Hsf1 reveals a compact transcriptional program for maintaining eukaryotic proteostasis","volume":"63","author":"Sol\u00eds","year":"2016","journal-title":"Mol Cell."},{"key":"2026100708030396100_iyag200-B81","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1016\/S0076-6879(05)09019-1","article-title":"The Delitto Perfetto approach to in vivo site-directed mutagenesis and chromosome rearrangements with synthetic oligonucleotides in yeast","volume":"409","author":"Storici","year":"2006","journal-title":"Methods Enzymol."},{"key":"2026100708030396100_iyag200-B82","doi-asserted-by":"publisher","first-page":"8007","DOI":"10.1074\/jbc.M116.761122","article-title":"Two alternative binding mechanisms connect the protein translocation Sec71-Sec72 Complex with heat shock proteins","volume":"292","author":"Tripathi","year":"2017","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B83","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1111\/j.1567-1364.2009.00555.x","article-title":"Transcriptional regulation of nonfermentable carbon utilization in budding yeast","volume":"10","author":"Turcotte","year":"2010","journal-title":"FEMS Yeast Res."},{"key":"2026100708030396100_iyag200-B84","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1128\/mmbr.05018-11","article-title":"Biology of the heat shock response and protein chaperones: budding yeast (Saccharomyces cerevisiae) as a model system","volume":"76","author":"Verghese","year":"2012","journal-title":"Microbiol Mol Biol Rev."},{"key":"2026100708030396100_iyag200-B85","doi-asserted-by":"publisher","first-page":"eaan4146","DOI":"10.1126\/science.aan4146","article-title":"MitoCPR-A surveillance pathway that protects mitochondria in response to protein import stress","volume":"360","author":"Weidberg","year":"2018","journal-title":"Science"},{"key":"2026100708030396100_iyag200-B86","doi-asserted-by":"publisher","first-page":"1049","DOI":"10.1038\/nsmb853","article-title":"SRP meets the ribosome","volume":"11","author":"Wild","year":"2004","journal-title":"Nat Struct Mol Biol."},{"key":"2026100708030396100_iyag200-B87","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1126\/science.1257522","article-title":"Targeting and plasticity of mitochondrial proteins revealed by proximity-specific ribosome profiling","volume":"346","author":"Williams","year":"2014","journal-title":"Science"},{"key":"2026100708030396100_iyag200-B88","doi-asserted-by":"publisher","first-page":"29222","DOI":"10.1074\/jbc.M704201200","article-title":"Translation-coupled translocation of yeast fumarase into mitochondria in vivo","volume":"282","author":"Yogev","year":"2007","journal-title":"J Biol Chem."},{"key":"2026100708030396100_iyag200-B89","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/S0092-8674(02)01250-3","article-title":"Molecular chaperones Hsp90 and Hsp70 deliver preproteins to the mitochondrial import receptor Tom70","volume":"112","author":"Young","year":"2003","journal-title":"Cell"},{"key":"2026100708030396100_iyag200-B90","doi-asserted-by":"publisher","first-page":"1504","DOI":"10.1038\/s41467-020-15313-w","article-title":"The ribosome-associated complex RAC serves in a relay that directs nascent chains to Ssb","volume":"11","author":"Zhang","year":"2020","journal-title":"Nat Commun."},{"key":"2026100708030396100_iyag200-B91","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1007\/s10482-018-1189-3","article-title":"Yeast molecular chaperone gene SSB2 is involved in the endoplasmic reticulum stress response","volume":"112","author":"Zhao","year":"2019","journal-title":"Antonie van Leeuwenhoek"}],"container-title":["GENETICS"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/genetics\/advance-article-pdf\/doi\/10.1093\/genetics\/iyag200\/70430251\/iyag200.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/genetics\/article-pdf\/234\/2\/iyag200\/70430251\/iyag200.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/genetics\/article-pdf\/234\/2\/iyag200\/70430251\/iyag200.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T12:03:10Z","timestamp":1791374590000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/genetics\/article\/doi\/10.1093\/genetics\/iyag200\/8748742"}},"subtitle":[],"editor":[{"given":"M","family":"McMurray","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,8,1]]},"references-count":91,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,8,1]]},"published-print":{"date-parts":[[2026,10,7]]}},"URL":"https:\/\/doi.org\/10.1093\/genetics\/iyag200","relation":{},"ISSN":["1943-2631"],"issn-type":[{"value":"1943-2631","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,10]]},"published":{"date-parts":[[2026,8,1]]},"article-number":"iyag200"}}