{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T15:56:44Z","timestamp":1792598204692,"version":"4.5.0"},"reference-count":47,"publisher":"American Chemical Society (ACS)","issue":"41","license":[{"start":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T00:00:00Z","timestamp":1791158400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100023921","name":"Structural Genomics Consortium","doi-asserted-by":"publisher","award":["1097737"],"award-info":[{"award-number":["1097737"]}],"id":[{"id":"10.13039\/501100023921","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100032455","name":"Male Contraceptive Initiative","doi-asserted-by":"publisher","award":["Agility Grant"],"award-info":[{"award-number":["Agility Grant"]}],"id":[{"id":"10.13039\/100032455","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100032455","name":"Male Contraceptive Initiative","doi-asserted-by":"publisher","award":["Postdoctoral Fellowship"],"award-info":[{"award-number":["Postdoctoral Fellowship"]}],"id":[{"id":"10.13039\/100032455","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100032455","name":"Male Contraceptive Initiative","doi-asserted-by":"publisher","award":["Predoctoral Fellowship"],"award-info":[{"award-number":["Predoctoral Fellowship"]}],"id":[{"id":"10.13039\/100032455","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005972","name":"Deutsche Krebshilfe","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100005972","id-type":"DOI","asserted-by":"publisher"}]},{"name":"German Cancer Research Center (DKTK)"},{"name":"National Institutes of Health (NIH)","award":["P50 HD106793"],"award-info":[{"award-number":["P50 HD106793"]}]},{"name":"National Institutes of Health (NIH)","award":["R35 GM127030"],"award-info":[{"award-number":["R35 GM127030"]}]},{"name":"National Institutes of Health (NIH)","award":["R61\/R33 HD099720"],"award-info":[{"award-number":["R61\/R33 HD099720"]}]},{"name":"National Institutes of Health (NIH)","award":["S10 OD0288697"],"award-info":[{"award-number":["S10 OD0288697"]}]},{"name":"National Institutes of Health (NIH)","award":["S10 OD038355"],"award-info":[{"award-number":["S10 OD038355"]}]},{"name":"National Institutes of Health (NIH)","award":["T32 GM136631"],"award-info":[{"award-number":["T32 GM136631"]}]},{"name":"Bundesministerium f??r Bildung und Forschung","award":["BMBF 03ZU1109FA"],"award-info":[{"award-number":["BMBF 03ZU1109FA"]}]},{"name":"Bundesministerium f??r Bildung und Forschung","award":["BMFTR 01GR2501A"],"award-info":[{"award-number":["BMFTR 01GR2501A"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,21]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Homeodomain-interacting protein kinase 4 (HIPK4) is a dual-specificity kinase that is predominantly expressed in differentiating spermatids, required for sperm development, and a promising target for nonhormonal male contraception. Genetic and functional studies have established an essential role for HIPK4 in spermiogenesis, where it acts at least in part through regulation of the F-actin-scaffolded acroplaxome during spermatid head shaping. The direct molecular targets of HIPK4 and their downstream effectors remain poorly defined, and small-molecule probes would be versatile tools for further investigating HIPK4 functions. Synthetic HIPK4 ligands could also be valuable leads for the development of nonhormonal male contraceptives. Here, we report the discovery of a cyanoquinoline-based series of HIPK4 inhibitors with nanomolar potency. Our lead compounds are selective for HIPK4, both within the HIPK family and across the broader kinome, establishing this scaffold as a useful starting point for probe and lead development. Unexpectedly, we found that a subset of these cyanoquinolines also perturbs HIPK4 proteostasis in a cell-type-specific manner. In spermatids, these compounds induce the formation of detergent-insoluble HIPK4 aggregates and promote interactions between this kinase and the autophagy receptor Tax1-binding protein 1 (TAX1BP1), potentially reinforcing a native mechanism of HIPK4 regulation. Together, our findings establish cyanoquinoline ligands as a new chemotype for probing HIPK4 biology and advancing male contraceptive discovery.<\/jats:p>","DOI":"10.1021\/jacs.6c10097","type":"journal-article","created":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T09:02:16Z","timestamp":1791190936000},"page":"44182-44207","source":"Crossref","is-referenced-by-count":0,"title":["Small-Molecule Modulators\nof HIPK4 Activity and Proteostasis"],"prefix":"10.1021","volume":"148","author":[{"given":"Zaile","family":"Zhuang","sequence":"first","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7499-3189","authenticated-orcid":true,"given":"Riley K.","family":"Togashi","sequence":"additional","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patrick C.","family":"Kearney","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ian","family":"Pass","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven M.","family":"Swick","sequence":"additional","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]},{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fu-Yue","family":"Zeng","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrey A.","family":"Bobkov","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lynn M.","family":"Fujimoto","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shubhankar","family":"Dutta","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Athina","family":"Zerva","sequence":"additional","affiliation":[{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]},{"name":"Buchmann Institute for Life Sciences , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2075-7938","authenticated-orcid":true,"given":"Nicolai D.","family":"Raig","sequence":"additional","affiliation":[{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]},{"name":"Buchmann Institute for Life Sciences , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Debasmita","family":"Saha","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Atoosa","family":"Emami","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1252-1829","authenticated-orcid":true,"given":"Martin P.","family":"Schwalm","sequence":"additional","affiliation":[{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]},{"name":"Buchmann Institute for Life Sciences , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bradley K.","family":"Moon","sequence":"additional","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samuel T.","family":"Howard","sequence":"additional","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5995-6494","authenticated-orcid":true,"given":"Stefan","family":"Knapp","sequence":"additional","affiliation":[{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]},{"name":"Buchmann Institute for Life Sciences , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7202-9468","authenticated-orcid":true,"given":"Thomas","family":"Hanke","sequence":"additional","affiliation":[{"name":"Goethe University , , ,","place":["Frankfurt Am Main, Germany, 60438"]},{"name":"Buchmann Institute for Life Sciences , , ,","place":["Frankfurt Am Main, Germany, 60438"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas D. Y.","family":"Chung","sequence":"additional","affiliation":[{"name":"Sanford Burnham Prebys Medical Discovery Institute , , ,","place":["La Jolla, California, United States, 92037"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9220-8436","authenticated-orcid":true,"given":"James K.","family":"Chen","sequence":"additional","affiliation":[{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]},{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]},{"name":"Stanford University , , , ,","place":["Stanford, California, United States, 94305"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,10,5]]},"reference":[{"issue":"3","key":"2026102111064022500_cit1","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1111\/j.1728-4465.2014.00393.x","article-title":"Intended and unintended pregnancies\nworldwide in 2012 and recent trends","volume":"45","author":"Sedgh","year":"2014","journal-title":"Stud. Fam.\nPlann."},{"issue":"2","key":"2026102111064022500_cit2","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1056\/NEJMoa1000485","article-title":"Adverse\nevents associated\nwith testosterone administration","volume":"363","author":"Basaria","year":"2010","journal-title":"N. Engl. J.\nMed."},{"issue":"2","key":"2026102111064022500_cit3","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1007\/s11154-011-9183-3","article-title":"Male hormonal contraception: potential\nrisks and benefits","volume":"12","author":"Ilani","year":"2011","journal-title":"Rev. Endocr. Metab. Disord"},{"issue":"1","key":"2026102111064022500_cit4","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0085805","article-title":"Increased\nrisk of non-fatal myocardial infarction following testosterone therapy\nprescription in men","volume":"9","author":"Finkle","year":"2014","journal-title":"PLoS One"},{"issue":"6","key":"2026102111064022500_cit5","doi-asserted-by":"publisher","first-page":"2492","DOI":"10.1210\/en.2010-0941","article-title":"Oral administration\nof a retinoic Acid receptor antagonist\nreversibly inhibits spermatogenesis in mice","volume":"152","author":"Chung","year":"2011","journal-title":"Endocrinology"},{"issue":"1","key":"2026102111064022500_cit6","doi-asserted-by":"publisher","first-page":"111","DOI":"10.2164\/jandrol.110.010751","article-title":"Suppression of spermatogenesis\nby bisdichloroacetyldiamines is mediated by inhibition of testicular\nretinoic acid biosynthesis","volume":"32","author":"Amory","year":"2011","journal-title":"J. Androl."},{"issue":"4","key":"2026102111064022500_cit7","doi-asserted-by":"publisher","first-page":"673","DOI":"10.1016\/j.cell.2012.06.045","article-title":"Small-molecule inhibition of BRDT for male contraception","volume":"150","author":"Matzuk","year":"2012","journal-title":"Cell"},{"issue":"1","key":"2026102111064022500_cit8","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1038\/s41467-023-36119-6","article-title":"On-demand male contraception\nvia acute inhibition of soluble adenylyl cyclase","volume":"14","author":"Balbach","year":"2023","journal-title":"Nat. Commun."},{"issue":"6698","key":"2026102111064022500_cit9","doi-asserted-by":"publisher","first-page":"885","DOI":"10.1126\/science.adl2688","article-title":"Reversible male contraception by targeted inhibition\nof serine\/threonine kinase 33","volume":"384","author":"Ku","year":"2024","journal-title":"Science"},{"key":"2026102111064022500_cit10","doi-asserted-by":"publisher","first-page":"9","DOI":"10.7554\/eLife.50209","article-title":"HIPK4 is essential for murine spermiogenesis","volume":"9","author":"Crapster","year":"2020","journal-title":"Elife"},{"issue":"9","key":"2026102111064022500_cit11","doi-asserted-by":"publisher","first-page":"102327","DOI":"10.1016\/j.jbc.2022.102327","article-title":"Homeodomain-interacting\nprotein kinase HIPK4 regulates\nphosphorylation of manchette protein RIMBP3 during spermiogenesis","volume":"298","author":"Liu","year":"2022","journal-title":"J. Biol. Chem."},{"issue":"3","key":"2026102111064022500_cit12","doi-asserted-by":"publisher","first-page":"hoag066","DOI":"10.1093\/hropen\/hoag066","article-title":"HIPK4 is a novel gene associated with teratozoospermia and male infertility","volume":"2026","author":"Koser","year":"2026","journal-title":"Hum. Reprod. Open"},{"issue":"29","key":"2026102111064022500_cit13","doi-asserted-by":"publisher","first-page":"5649","DOI":"10.1016\/j.febslet.2007.11.022","article-title":"Novel homeodomain-interacting protein kinase family\nmember, HIPK4, phosphorylates human p53 at serine 9","volume":"581","author":"Arai","year":"2007","journal-title":"FEBS Lett."},{"issue":"40","key":"2026102111064022500_cit14","doi-asserted-by":"publisher","first-page":"25875","DOI":"10.1074\/jbc.273.40.25875","article-title":"Homeodomain-interacting\nprotein kinases, a novel family of co-repressors for homeodomain transcription\nfactors","volume":"273","author":"Kim","year":"1998","journal-title":"J. Biol. Chem."},{"issue":"9","key":"2026102111064022500_cit15","doi-asserted-by":"publisher","first-page":"2527","DOI":"10.1091\/mbc.9.9.2527","article-title":"Activation\nof androgen receptor function by a novel nuclear protein kinase","volume":"9","author":"Moilanen","year":"1998","journal-title":"Mol. Biol. Cell"},{"issue":"7873","key":"2026102111064022500_cit16","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction\nwith AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"2026102111064022500_cit17","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1186\/s12964-014-0082-6","article-title":"Effect of tyrosine\nautophosphorylation on catalytic activity and subcellular localisation\nof homeodomain-interacting protein kinases (HIPK)","volume":"13","author":"van der Laden","year":"2015","journal-title":"Cell Commun. Signal"},{"key":"2026102111064022500_cit18","doi-asserted-by":"publisher","first-page":"11","DOI":"10.2147\/JEP.S110702","article-title":"Comprehensive kinase profile of pacritinib, a nonmyelosuppressive\nJanus kinase 2 inhibitor","volume":"8","author":"Singer","year":"2016","journal-title":"J. Exp. Pharmacol."},{"issue":"11","key":"2026102111064022500_cit19","doi-asserted-by":"publisher","first-page":"1046","DOI":"10.1038\/nbt.1990","article-title":"Comprehensive\nanalysis of kinase inhibitor selectivity","volume":"29","author":"Davis","year":"2011","journal-title":"Nat.\nBiotechnol."},{"key":"2026102111064022500_cit20","unstructured":"Flynn, G. A.; Bajji, A.; Huynh, K.\n          HIPK4 inhibitors and uses thereof\n          US11629143B2\n          2023."},{"issue":"6","key":"2026102111064022500_cit21","doi-asserted-by":"publisher","first-page":"560","DOI":"10.1089\/adt.2009.0222","article-title":"ADP-Glo: A Bioluminescent\nand homogeneous ADP monitoring assay for kinases","volume":"7","author":"Zegzouti","year":"2009","journal-title":"Assay Drug Dev. Technol."},{"issue":"2","key":"2026102111064022500_cit22","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1038\/nchembio.1404","article-title":"A conserved water-mediated\nhydrogen bond network defines\nbosutinib\u2019s kinase selectivity","volume":"10","author":"Levinson","year":"2014","journal-title":"Nat.\nChem. Biol."},{"issue":"13","key":"2026102111064022500_cit23","doi-asserted-by":"publisher","first-page":"3915","DOI":"10.1021\/jm9002928","article-title":"Hybrid compound design\nto overcome the gatekeeper T338M mutation in cSrc","volume":"52","author":"Getlik","year":"2009","journal-title":"J. Med. Chem."},{"issue":"48","key":"2026102111064022500_cit24","doi-asserted-by":"publisher","first-page":"46265","DOI":"10.1074\/jbc.M207135200","article-title":"Structure of the epidermal growth\nfactor receptor kinase domain alone and in complex with a 4-anilinoquinazoline\ninhibitor","volume":"277","author":"Stamos","year":"2002","journal-title":"J. Biol. Chem."},{"issue":"2","key":"2026102111064022500_cit25","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1016\/j.chembiol.2017.10.010","article-title":"Quantitative,\nwide-spectrum\nkinase profiling in live cells for assessing the effect of cellular\nATP on target engagement","volume":"25","author":"Vasta","year":"2018","journal-title":"Cell Chem. Biol."},{"issue":"15","key":"2026102111064022500_cit26","doi-asserted-by":"publisher","first-page":"12534","DOI":"10.1021\/acs.jmedchem.4c00411","article-title":"Back-pocket optimization of 2-aminopyrimidine-based\nmacrocycles leads to potent EPHA2\/GAK kinase inhibitors","volume":"67","author":"Gerninghaus","year":"2024","journal-title":"J. Med. Chem."},{"key":"2026102111064022500_cit27","doi-asserted-by":"publisher","first-page":"118076","DOI":"10.1016\/j.ejmech.2025.118076","article-title":"Development\nof pyrazolo[1,5-a]pyrimidine based\nmacrocyclic kinase inhibitors targeting AAK1","volume":"299","author":"Mensing","year":"2025","journal-title":"Eur. J. Med. Chem."},{"issue":"12","key":"2026102111064022500_cit28","doi-asserted-by":"publisher","first-page":"5214","DOI":"10.1039\/D2CS00193D","article-title":"PROTACs:\npast, present and future","volume":"51","author":"Li","year":"2022","journal-title":"Chem. Soc. Rev."},{"issue":"1","key":"2026102111064022500_cit29","doi-asserted-by":"publisher","first-page":"103387","DOI":"10.1016\/j.drudis.2022.103387","article-title":"Drug delivery\nchallenges and formulation aspects of proteolysis targeting chimera\n(PROTACs)","volume":"28","author":"Saraswat","year":"2023","journal-title":"Drug Discovery Today"},{"issue":"9","key":"2026102111064022500_cit30","doi-asserted-by":"publisher","first-page":"2758","DOI":"10.1021\/acschembio.8b00692","article-title":"Quantitative\nLive-Cell Kinetic Degradation and Mechanistic Profiling of PROTAC\nMode of Action","volume":"13","author":"Riching","year":"2018","journal-title":"ACS Chem. Biol."},{"issue":"1","key":"2026102111064022500_cit31","doi-asserted-by":"publisher","first-page":"1294","DOI":"10.1038\/s41598-018-19202-7","article-title":"Structural\nbasis of thalidomide enantiomer binding to cereblon","volume":"8","author":"Mori","year":"2018","journal-title":"Sci. Rep."},{"issue":"17","key":"2026102111064022500_cit32","doi-asserted-by":"publisher","first-page":"20616","DOI":"10.1021\/acs.jmedchem.6c01320","article-title":"Macrocyclization of\nbroad-spectrum kinase inhibitor\nbosutinib leads to potent and selective quinoline-based HIPK4 inhibitor\nAZ137","volume":"69","author":"Zerva","year":"2026","journal-title":"J. Med. Chem."},{"issue":"6","key":"2026102111064022500_cit33","doi-asserted-by":"publisher","first-page":"1714","DOI":"10.1016\/j.cell.2020.10.038","article-title":"Mapping the degradable\nkinome provides a resource for expedited degrader\ndevelopment","volume":"183","author":"Donovan","year":"2020","journal-title":"Cell"},{"issue":"9","key":"2026102111064022500_cit34","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1080\/15548627.2016.1192750","article-title":"Autophagy regulates\nspermatid differentiation via degradation of\nPDLIM1","volume":"12","author":"Shang","year":"2016","journal-title":"Autophagy"},{"issue":"7","key":"2026102111064022500_cit35","doi-asserted-by":"publisher","first-page":"1753","DOI":"10.1080\/15548627.2020.1783822","article-title":"Autophagy\ncore protein ATG5 is required for elongating spermatid\ndevelopment, sperm individualization and normal fertility in male\nmice","volume":"17","author":"Huang","year":"2021","journal-title":"Autophagy"},{"issue":"5","key":"2026102111064022500_cit36","doi-asserted-by":"publisher","first-page":"424","DOI":"10.1111\/j.1440-1681.1996.tb02752.x","article-title":"Structure-activity\nrelationships for 4-anilinoquinazolines as potent inhibitors at the\nATP binding site of the epidermal growth factor receptor in vitro","volume":"23","author":"Denny","year":"1996","journal-title":"Clin. Exp. Pharmacol. Physio.l"},{"issue":"8098","key":"2026102111064022500_cit37","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.1038\/s41586-025-09763-9","article-title":"Inhibitors\nsupercharge kinase turnover through\nnative proteolytic circuits","volume":"649","author":"Scholes","year":"2026","journal-title":"Nature"},{"issue":"9","key":"2026102111064022500_cit38","doi-asserted-by":"publisher","first-page":"1883","DOI":"10.1038\/sj.emboj.7601077","article-title":"Autoregulatory control of the p53\nresponse by caspase-mediated processing of HIPK2","volume":"25","author":"Gresko","year":"2006","journal-title":"Embo J."},{"issue":"1","key":"2026102111064022500_cit39","doi-asserted-by":"publisher","first-page":"5212","DOI":"10.1038\/s41467-021-25572-w","article-title":"Reconstitution defines\nthe roles\nof p62, NBR1 and TAX1BP1 in ubiquitin condensate formation and autophagy\ninitiation","volume":"12","author":"Turco","year":"2021","journal-title":"Nat. Commun."},{"issue":"7","key":"2026102111064022500_cit40","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1038\/cr.2014.70","article-title":"Atg7 is required for\nacrosome biogenesis during spermatogenesis in\nmice","volume":"24","author":"Wang","year":"2014","journal-title":"Cell Res."},{"issue":"3","key":"2026102111064022500_cit41","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1242\/dev.147074","article-title":"Sirt1 regulates acrosome biogenesis by modulating\nautophagic\nflux during spermiogenesis in mice","volume":"144","author":"Liu","year":"2017","journal-title":"Development"},{"issue":"2","key":"2026102111064022500_cit42","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1093\/humupd\/dmab043","article-title":"Autophagy: a multifaceted player\nin the fate of sperm","volume":"28","author":"Wang","year":"2022","journal-title":"Hum. Reprod. 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