{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T15:41:06Z","timestamp":1786722066945,"version":"build-2736575974"},"reference-count":86,"publisher":"American Chemical Society (ACS)","issue":"8","license":[{"start":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T00:00:00Z","timestamp":1785196800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T00:00:00Z","timestamp":1785196800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T00:00:00Z","timestamp":1785196800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100004410","name":"T\u00c3\u00bcrkiye Bilimsel ve Teknolojik Arastirma Kurumu","doi-asserted-by":"publisher","award":["221Z294"],"award-info":[{"award-number":["221Z294"]}],"id":[{"id":"10.13039\/501100004410","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,8,14]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Zuotin-related factor 1 (ZRF1) is an essential chromatin-associated regulator with established roles in transcription, differentiation, and cancer; however, its contribution to paracrine signaling within the tumor microenvironment (TME) remains unexplored. Here, we identify ZRF1 as a regulator of nutrient-sensitive tumor-derived paracrine signaling in triple-negative breast cancer (TNBC). Across multiple breast cancer subtypes, ZRF1 depletion altered the expression, secretion, and stress-associated protein patterns of the inflammatory mediator S100A9. This regulation was influenced by metabolic status, where nutrient deprivation selectively destabilized specific S100A9 proteoforms through proteasome-sensitive mechanisms. Bioinformatic analyses revealed a strong association between S100A9 expression and neutrophil infiltration in breast tumors, prompting functional investigation of ZRF1-mediated tumor\u2013neutrophil communication using a multilayered experimental framework. Conditioned medium from ZRF1-deficient tumor cells induced a nonbinary neutrophil activation spectrum rather than a fixed polarization state. This phenotypic remodeling was characterized by altered inflammatory signaling and cytokine (IL-8) dynamics in both differentiated HL-60 neutrophil-like cells and primary human neutrophils. Functionally, ZRF1-dependent paracrine cues regulated neutrophil transendothelial migration over time. Reciprocal coculture platforms and real-time xCELLigence assays further demonstrated that ZRF1 loss sensitizes tumor cells to neutrophil-derived signals, enhancing invasion and driving state-dependent architectural remodeling in 3D tumor spheroids. Collectively, these findings support a model in which a nutrient-sensitive ZRF1-dependent secretory network involving S100A9 contributes to human neutrophil plasticity under metabolic stress. Consequently, this study highlights S100A9 as one component of a broader ZRF1-regulated secretory network and identifies nutrient-sensitive tumor\u2013immune communication as a potential therapeutic vulnerability in aggressive breast cancer.<\/jats:p>","DOI":"10.1021\/acsptsci.6c00020","type":"journal-article","created":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T13:14:36Z","timestamp":1785849276000},"page":"2198-2224","source":"Crossref","is-referenced-by-count":0,"title":["A Nutrient-Sensitive\nZRF1-Dependent Secretory Network\nInvolving S100A9 Contributes to Neutrophil Plasticity in Triple-Negative\nBreast Cancer"],"prefix":"10.1021","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5153-9602","authenticated-orcid":true,"given":"Aysegul","family":"Kaymak Ozdemir","sequence":"first","affiliation":[{"name":"Ege University , , ,","place":["Izmir, Tu\u0308rkiye, 35040"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mahinur","family":"Basci","sequence":"additional","affiliation":[{"name":"Ege University , , ,","place":["Izmir, Tu\u0308rkiye, 35100"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Minenur","family":"Kalyoncu","sequence":"additional","affiliation":[{"name":"Dokuz Eylu\u0308l University Health Campus , , ,","place":["Izmir, Tu\u0308rkiye, 35340"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ayse","family":"Caner","sequence":"additional","affiliation":[{"name":"Ege University , , Bornova, ,","place":["Izmir, Tu\u0308rkiye, 35100"]},{"name":"Ege University , , Bornova, ,","place":["Izmir, Tu\u0308rkiye, 35100"]},{"name":"Ege University , , Bornova, ,","place":["Izmir, Tu\u0308rkiye, 35100"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Serif","family":"Senturk","sequence":"additional","affiliation":[{"name":"Dokuz Eylu\u0308l University Health Campus , , ,","place":["Izmir, Tu\u0308rkiye, 35340"]},{"name":"Dokuz Eylul University , , ,","place":["Izmir, Tu\u0308rkiye, 35340"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,7,28]]},"reference":[{"issue":"10074","key":"2026081411083401700_cit1","doi-asserted-by":"publisher","first-page":"1134","DOI":"10.1016\/S0140-6736(16)31891-8","article-title":"Breast Cancer","volume":"389","author":"Harbeck","year":"2017","journal-title":"Lancet"},{"issue":"3","key":"2026081411083401700_cit2","doi-asserted-by":"publisher","first-page":"209","DOI":"10.3322\/caac.21660","article-title":"Global Cancer\nStatistics\n2020: GLOBOCAN Estimates of Incidence and Mortality Worldwide for\n36 Cancers in 185 Countries","volume":"71","author":"Sung","year":"2021","journal-title":"CA: Cancer J. Clin."},{"issue":"6797","key":"2026081411083401700_cit3","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1038\/35021093","article-title":"Molecular\nPortraits of Human Breast Tumours","volume":"406","author":"Perou","year":"2000","journal-title":"Nature"},{"issue":"5","key":"2026081411083401700_cit4","doi-asserted-by":"publisher","DOI":"10.1186\/bcr2635","article-title":"Phenotypic\nand Molecular\nCharacterization of the Claudin-Low Intrinsic Subtype of Breast Cancer","volume":"12","author":"Prat","year":"2010","journal-title":"Breast Cancer Res"},{"issue":"5","key":"2026081411083401700_cit5","doi-asserted-by":"publisher","first-page":"660","DOI":"10.1038\/s41588-022-01041-y","article-title":"Breast Tumor Microenvironment Structures\nAre Associated with Genomic Features and Clinical Outcome","volume":"54","author":"Danenberg","year":"2022","journal-title":"Nat. Genet."},{"issue":"8","key":"2026081411083401700_cit6","doi-asserted-by":"publisher","first-page":"554","DOI":"10.1038\/s41568-024-00714-6","article-title":"Towards Targeting the\nBreast Cancer Immune Microenvironment","volume":"24","author":"Harris","year":"2024","journal-title":"Nat.\nRev. Cancer"},{"issue":"4","key":"2026081411083401700_cit7","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1182\/blood-2010-01-259028","article-title":"In Vivo Labeling\nwith 2H2O Reveals a Human Neutrophil Lifespan of 5.4 Days","volume":"116","author":"Pillay","year":"2010","journal-title":"Blood"},{"issue":"3","key":"2026081411083401700_cit8","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1016\/j.ccr.2009.06.017","article-title":"Polarization\nof Tumor-Associated\nNeutrophil Phenotype by TGF-Beta: \u201cN1\u201d versus \u201cN2\u201d\nTAN","volume":"16","author":"Fridlender","year":"2009","journal-title":"Cancer Cell"},{"key":"2026081411083401700_cit9","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.00532","article-title":"An Attempt to Polarize Human Neutrophils\nToward N1 and N2 Phenotypes in Vitro","volume":"11","author":"Ohms","year":"2020","journal-title":"Front.\nImmunol."},{"issue":"6679","key":"2026081411083401700_cit10","doi-asserted-by":"publisher","DOI":"10.1126\/science.adf6493","article-title":"Deterministic\nReprogramming of Neutrophils within Tumors","volume":"383","author":"Ng","year":"2024","journal-title":"Science"},{"issue":"12","key":"2026081411083401700_cit11","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.1016\/j.ccell.2022.10.008","article-title":"High-Resolution Single-Cell Atlas\nReveals Diversity and Plasticity of Tissue-Resident Neutrophils in\nNon-Small Cell Lung Cancer","volume":"40","author":"Salcher","year":"2022","journal-title":"Cancer Cell"},{"issue":"7938","key":"2026081411083401700_cit12","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1038\/s41586-022-05400-x","article-title":"Liver Tumour\nImmune Microenvironment Subtypes and Neutrophil Heterogeneity","volume":"612","author":"Xue","year":"2022","journal-title":"Nature"},{"issue":"2","key":"2026081411083401700_cit13","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1038\/nrc3893","article-title":"S100 Proteins\nin Cancer","volume":"15","author":"Bresnick","year":"2015","journal-title":"Nat. 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Cancer Res."},{"issue":"15","key":"2026081411083401700_cit18","doi-asserted-by":"publisher","DOI":"10.3390\/cells11152274","article-title":"Multifunctional\nRole of S100 Protein Family in the\nImmune System: An Update","volume":"11","author":"Singh","year":"2022","journal-title":"Cells"},{"issue":"1","key":"2026081411083401700_cit19","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.cell.2012.04.042","article-title":"A CXCL1 Paracrine\nNetwork Links Cancer Chemoresistance and Metastasis","volume":"150","author":"Acharyya","year":"2012","journal-title":"Cell"},{"issue":"4","key":"2026081411083401700_cit20","doi-asserted-by":"publisher","first-page":"243","DOI":"10.2174\/156800908784533445","article-title":"S100A8 and S100A9 Overexpression\nIs Associated with\nPoor Pathological Parameters in Invasive Ductal Carcinoma of the Breast","volume":"8","author":"Arai","year":"2008","journal-title":"Curr. 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Life Sci."},{"issue":"1","key":"2026081411083401700_cit24","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-19722-9","article-title":"Polycomb Group-Mediated\nHistone H2A Monoubiquitination in Epigenome Regulation and Nuclear\nProcesses","volume":"11","author":"Barbour","year":"2020","journal-title":"Nat. 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Are There Implications for the Retinoic Acid\nSyndrome?","volume":"22","author":"Sham","year":"1996","journal-title":"Blood Cells, Mol., Dis."},{"issue":"78","key":"2026081411083401700_cit32","doi-asserted-by":"publisher","DOI":"10.3791\/50677","article-title":"Quantitative\nIn Vitro Assay to Measure\nNeutrophil Adhesion to Activated Primary Human Microvascular Endothelial\nCells under Static Conditions","author":"Wilhelmsen","year":"2013","journal-title":"J. Visualized\nExp."},{"key":"2026081411083401700_cit33","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1016\/j.atherosclerosis.2019.04.210","article-title":"Understanding Mechanobiology in Cultured\nEndothelium: A Review of the Orbital Shaker Method","volume":"285","author":"Warboys","year":"2019","journal-title":"Atherosclerosis"},{"key":"2026081411083401700_cit34","doi-asserted-by":"publisher","first-page":"245","DOI":"10.2144\/000114416","article-title":"A real-time\nassay for neutrophil\nchemotaxis","volume":"60","author":"Cano","year":"2016","journal-title":"Biotechniques"},{"issue":"50","key":"2026081411083401700_cit35","doi-asserted-by":"publisher","DOI":"10.3791\/2792","article-title":"A Real-Time\nElectrical Impedance Based Technique to\nMeasure Invasion of Endothelial Cell Monolayer by Cancer Cells","author":"Rahim","year":"2011","journal-title":"J. Visualized Exp."},{"issue":"6","key":"2026081411083401700_cit36","doi-asserted-by":"publisher","first-page":"1957","DOI":"10.1158\/1078-0432.CCR-18-2544","article-title":"Tumor-Contacted\nNeutrophils\nPromote Metastasis by a CD90-TIMP-1 Juxtacrine\u2013Paracrine Loop","volume":"25","author":"Wang","year":"2019","journal-title":"Clin. Cancer Res."},{"issue":"4","key":"2026081411083401700_cit37","doi-asserted-by":"publisher","first-page":"628","DOI":"10.1038\/nprot.2010.6","article-title":"In Vitro Angiogenesis:\nEndothelial Cell Tube Formation\non Gelled Basement Membrane Extract","volume":"5","author":"Arnaoutova","year":"2010","journal-title":"Nat. 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Rheumatol."},{"key":"2026081411083401700_cit42","doi-asserted-by":"publisher","DOI":"10.1016\/j.lfs.2020.118699","article-title":"Tumor-Associated\nNeutrophils as New Players in Immunosuppressive Process of the Tumor\nMicroenvironment in Breast Cancer","volume":"264","author":"Hajizadeh","year":"2021","journal-title":"Life Sci."},{"issue":"3","key":"2026081411083401700_cit43","doi-asserted-by":"publisher","first-page":"51","DOI":"10.62347\/JQDQ1527","article-title":"Prognostic Impact of Tumor-Associated\nNeutrophils in\nBreast Cancer","volume":"17","author":"Kakumoto","year":"2024","journal-title":"Int. J. Clin. Exp. Pathol."},{"issue":"10","key":"2026081411083401700_cit44","doi-asserted-by":"publisher","first-page":"2689","DOI":"10.22034\/APJCP.2017.18.10.2689","article-title":"Tumor-Associated\nNeutrophils in Breast Cancer Subtypes","volume":"18","author":"Soto-Perez-de-Celis","year":"2017","journal-title":"Asian\nPac. J. Cancer Prev."},{"issue":"1","key":"2026081411083401700_cit45","doi-asserted-by":"publisher","first-page":"207","DOI":"10.31557\/APJCP.2025.26.1.207","article-title":"The Role of Tumor-Associated Neutrophils in Early Luminal\nHER2-Negative Breast Cancer Progression","volume":"26","author":"Zakurdaev","year":"2025","journal-title":"Asian\nPac. J. Cancer Prev."},{"key":"2026081411083401700_cit46","doi-asserted-by":"publisher","DOI":"10.1016\/j.pharmthera.2020.107692","article-title":"Interleukin-8: A Chemokine\nat the\nIntersection of Cancer Plasticity, Angiogenesis, and Immune Suppression","volume":"219","author":"Fousek","year":"2021","journal-title":"Pharmacol. Ther."},{"key":"2026081411083401700_cit47","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.656908","article-title":"The Role of Tumor Necrosis\nFactor in Manipulating the Immunological Response of Tumor Microenvironment","volume":"12","author":"Laha","year":"2021","journal-title":"Front. Immunol."},{"issue":"7","key":"2026081411083401700_cit48","doi-asserted-by":"publisher","DOI":"10.3390\/cancers12071791","article-title":"Interleukin-1\u03b2 and Cancer","volume":"12","author":"Re\u0301be\u0301","year":"2020","journal-title":"Cancers"},{"key":"2026081411083401700_cit49","doi-asserted-by":"publisher","first-page":"141","DOI":"10.2147\/JIR.S284941","article-title":"Studying Neutrophil Function in Vitro:\nCell Models and Environmental Factors","volume":"14","author":"Blanter","year":"2021","journal-title":"J. Inflamm.\nRes."},{"issue":"1","key":"2026081411083401700_cit50","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00280-013-2321-8","article-title":"Mechanisms\nof Action of Tasquinimod on the Tumour Microenvironment","volume":"73","author":"Raymond","year":"2014","journal-title":"Cancer Chemother. Pharmacol."},{"issue":"21","key":"2026081411083401700_cit51","doi-asserted-by":"publisher","first-page":"4546","DOI":"10.1016\/j.cell.2023.08.043","article-title":"The Local\nMicroenvironment Drives\nActivation of Neutrophils in Human Brain Tumors","volume":"186","author":"Maas","year":"2023","journal-title":"Cell"},{"issue":"1","key":"2026081411083401700_cit52","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.it.2015.11.008","article-title":"Neutrophils in the Tumor Microenvironment","volume":"37","author":"Powell","year":"2016","journal-title":"Trends Immunol."},{"key":"2026081411083401700_cit53","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1038\/nri3399","article-title":"Neutrophil recruitment\nand function in health and inflammation","volume":"13","author":"Kolaczkowska","year":"2013","journal-title":"Nat.\nRev. Immunol."},{"issue":"1","key":"2026081411083401700_cit54","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms7915","article-title":"Extracellular\nMRP8\/14 Is a Regulator of \u03922 Integrin-Dependent Neutrophil Slow\nRolling and Adhesion","volume":"6","author":"Pruenster","year":"2015","journal-title":"Nat. Commun."},{"issue":"6","key":"2026081411083401700_cit55","doi-asserted-by":"publisher","first-page":"3233","DOI":"10.4049\/jimmunol.170.6.3233","article-title":"Proinflammatory\nActivities of S100: Proteins S100A8, S100A9, and S100A8\/A9 Induce\nNeutrophil Chemotaxis and Adhesion","volume":"170","author":"Ryckman","year":"2003","journal-title":"J. Immunol."},{"key":"2026081411083401700_cit56","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1543403","article-title":"Cannabidiol Polarizes Human Neutrophils\ntoward a Cancer-Promoting Phenotype","volume":"16","author":"Khoury","year":"2025","journal-title":"Front.\nImmunol."},{"issue":"20","key":"2026081411083401700_cit57","doi-asserted-by":"publisher","first-page":"2159","DOI":"10.1182\/blood-2018-11-844548","article-title":"Neutrophil\nPlasticity in the Tumor\nMicroenvironment","volume":"133","author":"Giese","year":"2019","journal-title":"Blood"},{"issue":"12","key":"2026081411083401700_cit58","doi-asserted-by":"publisher","first-page":"1431","DOI":"10.20892\/j.issn.2095-3941.2025.0547","article-title":"From Bone Marrow to the Tumor Microenvironment:\nHow Neutrophil Maturation Shapes Cancer Immunity","volume":"22","author":"Shi","year":"2025","journal-title":"Cancer Biol. 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