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So far there is a lack of knowledge with respect to the role of the acidic microenvironment in the invasive process. This work aimed to study the phenotypic and genetic response of PDAC cells to acidic stress along the different stages of selection. To this end, we subjected the cells to short- and long-term acidic pressure and recovery to pHe 7.4. This treatment aimed at mimicking PDAC edges and consequent cancer cell escape from the tumor. The impact of acidosis was assessed for cell morphology, proliferation, adhesion, migration, invasion, and epithelial\u2013mesenchymal transition (EMT) via functional in vitro assays and RNA sequencing. Our results indicate that short acidic treatment limits growth, adhesion, invasion, and viability of PDAC cells. As the acid treatment progresses, it selects cancer cells with enhanced migration and invasion abilities induced by EMT, potentiating their metastatic potential when re-exposed to pHe 7.4. The RNA-seq analysis of PANC-1 cells exposed to short-term acidosis and pHe-selected recovered to pHe 7.4 revealed distinct transcriptome rewiring. We describe an enrichment of genes relevant to proliferation, migration, EMT, and invasion in acid-selected cells. Our work clearly demonstrates that upon acidosis stress, PDAC cells acquire more invasive cell phenotypes by promoting EMT and thus paving the way for more aggressive cell phenotypes.<\/jats:p>","DOI":"10.3390\/cancers15092572","type":"journal-article","created":{"date-parts":[[2023,5,3]],"date-time":"2023-05-03T05:55:09Z","timestamp":1683093309000},"page":"2572","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Acidic Growth Conditions Promote Epithelial-to-Mesenchymal Transition to Select More Aggressive PDAC Cell Phenotypes In Vitro"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0923-1481","authenticated-orcid":false,"given":"Madelaine Magal\u00ec","family":"Audero","sequence":"first","affiliation":[{"name":"U1003\u2014PHYCELL\u2014Laboratoire de Physiologie Cellulaire, Inserm, University of Lille, Villeneuve d\u2019Ascq, 59000 Lille, France"},{"name":"Laboratory of 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Germany"}]},{"given":"Maya","family":"Yassine","sequence":"additional","affiliation":[{"name":"U1003\u2014PHYCELL\u2014Laboratoire de Physiologie Cellulaire, Inserm, University of Lille, Villeneuve d\u2019Ascq, 59000 Lille, France"}]},{"given":"Giorgia","family":"Chinig\u00f2","sequence":"additional","affiliation":[{"name":"Laboratory of Cellular and Molecular Angiogenesis, Department of Life Sciences and Systems Biology, University of Turin, 10123 Turin, Italy"}]},{"given":"Antoine","family":"Folcher","sequence":"additional","affiliation":[{"name":"U1003\u2014PHYCELL\u2014Laboratoire de Physiologie Cellulaire, Inserm, University of Lille, Villeneuve d\u2019Ascq, 59000 Lille, France"}]},{"given":"Valerio","family":"Farfariello","sequence":"additional","affiliation":[{"name":"U1003\u2014PHYCELL\u2014Laboratoire de Physiologie Cellulaire, Inserm, University of Lille, Villeneuve d\u2019Ascq, 59000 Lille, France"}]},{"given":"Samuele","family":"Amadori","sequence":"additional","affiliation":[{"name":"Laboratory of Cellular and Molecular Angiogenesis, Department of Life Sciences and Systems Biology, University of Turin, 10123 Turin, Italy"}]},{"given":"Chiara","family":"Vaghi","sequence":"additional","affiliation":[{"name":"Laboratory of Cellular and Molecular Angiogenesis, Department of Life Sciences and Systems Biology, University of Turin, 10123 Turin, Italy"}]},{"given":"Albrecht","family":"Schwab","sequence":"additional","affiliation":[{"name":"Institute of Physiology II, University of M\u00fcnster, 48149 M\u00fcnster, Germany"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9757-5908","authenticated-orcid":false,"given":"Stephan J.","family":"Reshkin","sequence":"additional","affiliation":[{"name":"Department of Biosciences, Biotechnologies and Environment, University of Bari, 70126 Bari, 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Gastroenterol. Hepatol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"4846","DOI":"10.3748\/wjg.v24.i43.4846","article-title":"Pancreatic Cancer: A Review of Clinical Diagnosis, Epidemiology, Treatment and Outcomes","volume":"24","author":"McGuigan","year":"2018","journal-title":"WJG"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1186\/s13014-019-1345-6","article-title":"Pancreatic Ductal Adenocarcinoma: Biological Hallmarks, Current Status, and Future Perspectives of Combined Modality Treatment Approaches","volume":"14","author":"Orth","year":"2019","journal-title":"Radiat. Oncol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1600253","DOI":"10.1002\/bies.201600253","article-title":"Alternating PH Landscapes Shape Epithelial Cancer Initiation and Progression: Focus on Pancreatic Cancer","volume":"39","author":"Pedersen","year":"2017","journal-title":"BioEssays"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Carvalho, T.M.A., Di Molfetta, D., Greco, M.R., Koltai, T., Alfarouk, K.O., Reshkin, S.J., and Cardone, R.A. (2021). Tumor Microenvironment Features and Chemoresistance in Pancreatic Ductal Adenocarcinoma: Insights into Targeting Physicochemical Barriers and Metabolism as Therapeutic Approaches. Cancers, 13.","DOI":"10.3390\/cancers13236135"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"105796","DOI":"10.1016\/j.biocel.2020.105796","article-title":"PH Gradient Reversal Fuels Cancer Progression","volume":"125","author":"Zheng","year":"2020","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/nrc3110","article-title":"Dysregulated PH: A Perfect Storm for Cancer Progression","volume":"11","author":"Webb","year":"2011","journal-title":"Nat. Rev. Cancer"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1007\/s10555-021-10005-3","article-title":"What Do Cellular Responses to Acidity Tell Us about Cancer?","volume":"40","author":"Blaszczak","year":"2021","journal-title":"Cancer Metastasis Rev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1113\/jphysiol.2005.088344","article-title":"Migration of Human Melanoma Cells Depends on Extracellular PH and Na+\/H+ Exchange: Extracellular Protons Modulate Cell Migration","volume":"567","author":"Stock","year":"2005","journal-title":"J. Physiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1146\/annurev-physiol-021119-034627","article-title":"The Acidic Tumor Microenvironment as a Driver of Cancer","volume":"82","author":"Boedtkjer","year":"2020","journal-title":"Annu. Rev. Physiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1038\/sj.cdd.4401466","article-title":"Alterations of Intracellular PH Homeostasis in Apoptosis: Origins and Roles","volume":"11","author":"Huc","year":"2004","journal-title":"Cell Death Differ."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1038\/35014006","article-title":"Changes in Intramitochondrial and Cytosolic PH: Early Events That Modulate Caspase Activation during Apoptosis","volume":"2","author":"Matsuyama","year":"2000","journal-title":"Nat. Cell Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3938","DOI":"10.1158\/0008-5472.CAN-11-3881","article-title":"Chronic Autophagy Is a Cellular Adaptation to Tumor Acidic PH Microenvironments","volume":"72","author":"Wojtkowiak","year":"2012","journal-title":"Cancer Res."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1186\/s12935-014-0129-1","article-title":"Acidic Extracellular PH Promotes Epithelial Mesenchymal Transition in Lewis Lung Carcinoma Model","volume":"14","author":"Suzuki","year":"2014","journal-title":"Cancer Cell Int."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1431","DOI":"10.1007\/s00109-020-01959-y","article-title":"The Acidic Tumor Microenvironment Drives a Stem-like Phenotype in Melanoma Cells","volume":"98","author":"Andreucci","year":"2020","journal-title":"J. Mol. Med."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1007\/s10585-019-09990-1","article-title":"Adaptation to Chronic Acidic Extracellular PH Elicits a Sustained Increase in Lung Cancer Cell Invasion and Metastasis","volume":"37","author":"Sutoo","year":"2020","journal-title":"Clin. Exp. Metastasis"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1739","DOI":"10.1080\/15384101.2016.1231284","article-title":"Phenotypic Changes of Acid-Adapted Cancer Cells Push Them toward Aggressiveness in Their Evolution in the Tumor Microenvironment","volume":"16","author":"Damaghi","year":"2017","journal-title":"Cell Cycle"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"111984","DOI":"10.1016\/j.yexcr.2020.111984","article-title":"LAMC2 Modulates the Acidity of Microenvironments to Promote Invasion and Migration of Pancreatic Cancer Cells via Regulating AKT-Dependent NHE1 Activity","volume":"391","author":"Wang","year":"2020","journal-title":"Exp. Cell Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2395","DOI":"10.1016\/j.bbadis.2018.04.019","article-title":"Acidic PHe Regulates Cytoskeletal Dynamics through Conformational Integrin \u03921 Activation and Promotes Membrane Protrusion","volume":"1864","author":"Li","year":"2018","journal-title":"Biochim. Biophys. Acta (BBA) Mol. Basis Dis."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.bbrc.2020.02.018","article-title":"Invasive Phenotype Induced by Low Extracellular PH Requires Mitochondria Dependent Metabolic Flexibility","volume":"525","author":"Shin","year":"2020","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1524","DOI":"10.1158\/0008-5472.CAN-12-2796","article-title":"Acidity Generated by the Tumor Microenvironment Drives Local Invasion","volume":"73","author":"Estrella","year":"2013","journal-title":"Cancer Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1038\/s41467-019-14262-3","article-title":"TGF\u03b22-Induced Formation of Lipid Droplets Supports Acidosis-Driven EMT and the Metastatic Spreading of Cancer Cells","volume":"11","author":"Corbet","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1111\/iwj.13248","article-title":"Acidic PH Derived from Cancer Cells as a Double-edged Knife Modulates Wound Healing through DNA Repair Genes and Autophagy","volume":"17","author":"Tavakol","year":"2020","journal-title":"Int. Wound J."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"6858","DOI":"10.1038\/s41598-019-43262-y","article-title":"Acidic Stress Triggers Sodium-Coupled Bicarbonate Transport and Promotes Survival in A375 Human Melanoma Cells","volume":"9","author":"Yang","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"6699","DOI":"10.1158\/0008-5472.CAN-06-0983","article-title":"Acidic Extracellular PH Promotes Experimental Metastasis of Human Melanoma Cells in Athymic Nude Mice","volume":"66","author":"Rofstad","year":"2006","journal-title":"Cancer Res."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1016\/j.neo.2019.03.004","article-title":"Extracellular Acidosis Modulates the Expression of Epithelial-Mesenchymal Transition (EMT) Markers and Adhesion of Epithelial and Tumor Cells","volume":"21","author":"Riemann","year":"2019","journal-title":"Neoplasia"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1007\/s12307-016-0187-0","article-title":"Investigating Effects of Acidic PH on Proliferation, Invasion and Drug-Induced Apoptosis in Lymphoblastic Leukemia","volume":"9","author":"Bohloli","year":"2016","journal-title":"Cancer Microenviron."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"30664","DOI":"10.1074\/jbc.M112.339127","article-title":"Autophagy Is a Protective Mechanism for Human Melanoma Cells under Acidic Stress","volume":"287","author":"Marino","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1002\/jcb.22313","article-title":"An Acidic Extracellular PH Disrupts Adherens Junctions in HepG2 Cells by Src Kinases-Dependent Modification of E-Cadherin","volume":"108","author":"Chen","year":"2009","journal-title":"J. Cell. Biochem."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"37360","DOI":"10.1038\/srep37360","article-title":"Acidic PH Promotes Intervertebral Disc Degeneration: Acid-Sensing Ion Channel -3 as a Potential Therapeutic Target","volume":"6","author":"Gilbert","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2228","DOI":"10.1016\/j.celrep.2017.02.006","article-title":"Extracellular Acidic PH Activates the Sterol Regulatory Element-Binding Protein 2 to Promote Tumor Progression","volume":"18","author":"Kondo","year":"2017","journal-title":"Cell Rep."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Lee, S., and Shanti, A. (2021). Effect of Exogenous PH on Cell Growth of Breast Cancer Cells. Int. J. Mol. Sci., 22.","DOI":"10.21203\/rs.3.rs-260226\/v1"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1186\/s13046-017-0599-9","article-title":"The Acid-Sensing Ion Channel, ASIC2, Promotes Invasion and Metastasis of Colorectal Cancer under Acidosis by Activating the Calcineurin\/NFAT1 Axis","volume":"36","author":"Zhou","year":"2017","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1038\/s41419-018-1242-1","article-title":"Acidosis Enhances the Self-Renewal and Mitochondrial Respiration of Stem Cell-like Glioma Cells through CYP24A1-Mediated Reduction of Vitamin D","volume":"10","author":"Hu","year":"2019","journal-title":"Cell Death Dis."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1186\/2049-3002-1-23","article-title":"Acidosis Induces Reprogramming of Cellular Metabolism to Mitigate Oxidative Stress","volume":"1","author":"LaMonte","year":"2013","journal-title":"Cancer Metab."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"3088","DOI":"10.1080\/15384101.2015.1078032","article-title":"Extracellular Acidity Strengthens Mesenchymal Stem Cells to Promote Melanoma Progression","volume":"14","author":"Peppicelli","year":"2015","journal-title":"Cell Cycle"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"42369","DOI":"10.1038\/srep42369","article-title":"Extracellular Protonation Modulates Cell-Cell Interaction Mechanics and Tissue Invasion in Human Melanoma Cells","volume":"7","author":"Koch","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1158\/0008-5472.CAN-11-2076","article-title":"Functional Interaction between Responses to Lactic Acidosis and Hypoxia Regulates Genomic Transcriptional Outputs","volume":"72","author":"Tang","year":"2012","journal-title":"Cancer Res."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1007\/s10585-008-9145-7","article-title":"Acid Treatment of Melanoma Cells Selects for Invasive Phenotypes","volume":"25","author":"Moellering","year":"2008","journal-title":"Clin. Exp. Metastasis"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1038\/s41416-020-01173-0","article-title":"Tumour Acidosis Evaluated in Vivo by MRI-CEST PH Imaging Reveals Breast Cancer Metastatic Potential","volume":"124","author":"Anemone","year":"2021","journal-title":"Br. J. Cancer"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Yao, J., Czaplinska, D., Ialchina, R., Schnipper, J., Liu, B., Sandelin, A., and Pedersen, S.F. (2020). Cancer Cell Acid Adaptation Gene Expression Response Is Correlated to Tumor-Specific Tissue Expression Profiles and Patient Survival. Cancers, 12.","DOI":"10.3390\/cancers12082183"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1080\/15384101.2019.1578145","article-title":"Chromatin Accessibility Changes Are Associated with Enhanced Growth and Liver Metastasis Capacity of Acid-Adapted Colorectal Cancer Cells","volume":"18","author":"Zhou","year":"2019","journal-title":"Cell Cycle"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Schnipper, J., Kouba, S., Hague, F., Girault, A., Telliez, M.-S., Gu\u00e9nin, S., Ahidouch, A., Pedersen, S.F., and Ouadid-Ahidouch, H. (2022). Acid Adaptation Promotes TRPC1 Plasma Membrane Localization Leading to Pancreatic Ductal Adenocarcinoma Cell Proliferation and Migration through Ca2+ Entry and Interaction with PI3K\/CaM. Cancers, 14.","DOI":"10.3390\/cancers14194946"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1210","DOI":"10.1002\/ijc.34367","article-title":"Crosstalk between Tumor Acidosis, P53 and Extracellular Matrix Regulates Pancreatic Cancer Aggressiveness","volume":"152","author":"Czaplinska","year":"2023","journal-title":"Int. J. Cancer"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1113\/jphysiol.2005.092957","article-title":"Functional Role of Na+\u2013HCO3\u2212 Cotransport in Migration of Transformed Renal Epithelial Cells: Na+\u2013HCO3\u2212 Cotransport and Cell Migration","volume":"568","author":"Schwab","year":"2005","journal-title":"J. Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"3903","DOI":"10.1096\/fj.09-149518","article-title":"NHE1 Promotes Invadopodial ECM Proteolysis through Acidification of the Peri-invadopodial Space","volume":"24","author":"Busco","year":"2010","journal-title":"FASEB J."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: Ultrafast Universal RNA-Seq Aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Li, B., and Dewey, C.N. (2011). RSEM: Accurate Transcript Quantification from RNA-Seq Data with or without a Reference Genome. BMC Bioinform., 12.","DOI":"10.1186\/1471-2105-12-323"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated Estimation of Fold Change and Dispersion for RNA-Seq Data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/S0166-4328(01)00297-2","article-title":"Controlling the False Discovery Rate in Behavior Genetics Research","volume":"125","author":"Benjamini","year":"2001","journal-title":"Behav. Brain Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"W305","DOI":"10.1093\/nar\/gkp427","article-title":"ToppGene Suite for Gene List Enrichment Analysis and Candidate Gene Prioritization","volume":"37","author":"Chen","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"15545","DOI":"10.1073\/pnas.0506580102","article-title":"Gene Set Enrichment Analysis: A Knowledge-Based Approach for Interpreting Genome-Wide Expression Profiles","volume":"102","author":"Subramanian","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1097\/MPA.0b013e3181c15963","article-title":"Phenotype and Genotype of Pancreatic Cancer Cell Lines","volume":"39","author":"Deer","year":"2010","journal-title":"Pancreas"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"22182","DOI":"10.1039\/C8RA02095G","article-title":"Manipulating Extracellular Tumour PH: An Effective Target for Cancer Therapy","volume":"8","author":"Hao","year":"2018","journal-title":"RSC Adv."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2435","DOI":"10.1242\/jcs.018044","article-title":"Paxillin Comes of Age","volume":"121","author":"Deakin","year":"2008","journal-title":"J. Cell Sci."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"10938","DOI":"10.1074\/jbc.M411313200","article-title":"Acidic Extracellular PH Induces Matrix Metalloproteinase-9 Expression in Mouse Metastatic Melanoma Cells through the Phospholipase D-Mitogen-Activated Protein Kinase Signaling","volume":"280","author":"Kato","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1083\/jcb.201103045","article-title":"Cortactin Phosphorylation Regulates Cell Invasion through a PH-Dependent Pathway","volume":"195","author":"Magalhaes","year":"2011","journal-title":"J. Cell Biol."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Lucien, F., Brochu-Gaudreau, K., Arsenault, D., Harper, K., and Dubois, C.M. (2011). Hypoxia-Induced Invadopodia Formation Involves Activation of NHE-1 by the P90 Ribosomal S6 Kinase (P90RSK). PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0028851"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"940","DOI":"10.3892\/or.2013.2923","article-title":"Protease Activity at Invadopodial Focal Digestive Areas Is Dependent on NHE1-Driven Acidic PHe","volume":"31","author":"Greco","year":"2014","journal-title":"Oncol. Rep."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Debreova, M., Csaderova, L., Burikova, M., Lukacikova, L., Kajanova, I., Sedlakova, O., Kery, M., Kopacek, J., Zatovicova, M., and Bizik, J. (2019). CAIX Regulates Invadopodia Formation through Both a PH-Dependent Mechanism and Interplay with Actin Regulatory Proteins. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20112745"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Yamada, T., Park, C.S., Burns, A., Nakada, D., and Lacorazza, H.D. (2012). The Cytosolic Protein G0S2 Maintains Quiescence in Hematopoietic Stem Cells. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0038280"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1038\/s41389-021-00341-y","article-title":"Guanylate-Binding Proteins Induce Apoptosis of Leukemia Cells by Regulating MCL-1 and BAK","volume":"10","author":"Luo","year":"2021","journal-title":"Oncogenesis"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1093\/carcin\/bgs310","article-title":"GBP-1 Acts as a Tumor Suppressor in Colorectal Cancer Cells","volume":"34","author":"Lipnik","year":"2013","journal-title":"Carcinogenesis"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1016\/j.bbrc.2017.11.050","article-title":"GBP3 Promotes Glioma Cell Proliferation via SQSTM1\/P62-ERK1\/2 Axis","volume":"495","author":"Xu","year":"2018","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"10228","DOI":"10.1096\/fj.201902899R","article-title":"Peroxidasin Is Essential for Endothelial Cell Survival and Growth Signaling by Sulfilimine Crosslink-dependent Matrix Assembly","volume":"34","author":"Lee","year":"2020","journal-title":"FASEB J."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Kurihara-Shimomura, M., Sasahira, T., Shimomura, H., and Kirita, T. (2020). Peroxidan Plays a Tumor-Promoting Role in Oral Squamous Cell Carcinoma. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21155416"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"102090","DOI":"10.1016\/j.redox.2021.102090","article-title":"Peroxidasin Protein Expression and Enzymatic Activity in Metastatic Melanoma Cell Lines Are Associated with Invasive Potential","volume":"46","author":"Kienzl","year":"2021","journal-title":"Redox Biol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1606","DOI":"10.1038\/sj.onc.1209956","article-title":"Androgen Regulation of Soluble Guanylyl Cyclasea1 Mediates Prostate Cancer Cell Proliferation","volume":"26","author":"Cai","year":"2007","journal-title":"Oncogene"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1007\/s10585-012-9464-6","article-title":"Insidious Role of Nitric Oxide in Migration\/Invasion of Colon Cancer Cells by Upregulating MMP-2\/9 via Activation of CGMP-PKG-ERK Signaling Pathways","volume":"29","author":"Babykutty","year":"2012","journal-title":"Clin. Exp. Metastasis"},{"key":"ref_70","first-page":"1626","article-title":"CGMP-Dependent Protein Kinase I\u03b2 Regulates Breast Cancer Cell Migration and Invasion via a Novel Interaction with the Actin\/Myosin-Associated Protein Caldesmon","volume":"126","author":"Schwappacher","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"14797","DOI":"10.1038\/s41598-019-51420-5","article-title":"Soluble Guanylyl Cyclase A1 Subunit Is a Key Mediator of Proliferation, Survival, and Migration in ECC-1 and HeLa Cell Lines","volume":"9","author":"Ronchetti","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Zhou, J., Gao, S., Hsieh, C.-L., Malla, M., and Shemshedini, L. (2017). Peptide B Targets Soluble Guanylyl Cyclase A1 and Kills Prostate Cancer Cells. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0184088"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1016\/j.devcel.2016.04.011","article-title":"PRICKLE1 Contributes to Cancer Cell Dissemination through Its Interaction with MTORC2","volume":"37","author":"Daulat","year":"2016","journal-title":"Dev. Cell"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1186\/s12967-021-02873-8","article-title":"PRICKLE1, a Wnt\/PCP Signaling Component, Is Overexpressed and Associated with Inferior Prognosis in Acute Myeloid Leukemia","volume":"19","author":"Jiang","year":"2021","journal-title":"J. Transl. Med."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"8515","DOI":"10.1080\/21655979.2021.1985858","article-title":"Interferon Alpha-Inducible Protein 27 (IFI27) Is a Prognostic Marker for Pancreatic Cancer Based on Comprehensive Bioinformatics Analysis","volume":"12","author":"Huang","year":"2021","journal-title":"Bioengineered"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"3136","DOI":"10.1038\/s41388-021-01763-z","article-title":"Regulator of Calcineurin 1 Gene Isoform 4 in Pancreatic Ductal Adenocarcinoma Regulates the Progression of Tumor Cells","volume":"40","author":"Lao","year":"2021","journal-title":"Oncogene"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1093\/carcin\/bgn274","article-title":"Bone Morphogenetic Proteins Induce Pancreatic Cancer Cell Invasiveness through a Smad1-Dependent Mechanism That Involves Matrix Metalloproteinase-2","volume":"30","author":"Gordon","year":"2009","journal-title":"Carcinogenesis"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"14","DOI":"10.3892\/ol.2021.13132","article-title":"Interleukin-32 Regulates Downstream Molecules and Promotes the Invasion of Pancreatic Cancer Cells","volume":"23","author":"Takagi","year":"2021","journal-title":"Oncol. Lett."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"6572","DOI":"10.1038\/s41388-020-01452-3","article-title":"Oncogenic Function of TRIM2 in Pancreatic Cancer by Activating ROS-Related NRF2\/ITGB7\/FAK Axis","volume":"39","author":"Sun","year":"2020","journal-title":"Oncogene"},{"key":"ref_80","first-page":"1558","article-title":"AREG Mediates the Epithelial-mesenchymal Transition in Pancreatic Cancer Cells via the EGFR\/ERK\/NF-\u03baB Signalling Pathway","volume":"43","author":"Wang","year":"2020","journal-title":"Oncol. Rep."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"e21719","DOI":"10.1096\/fj.202100070R","article-title":"A Novel Role of ADGRF1 (GPR110) in Promoting Cellular Quiescence and Chemoresistance in Human Epidermal Growth Factor Receptor 2-positive Breast Cancer","volume":"35","author":"Abdulkareem","year":"2021","journal-title":"FASEB J."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"12586","DOI":"10.1038\/ncomms12586","article-title":"TNIK Inhibition Abrogates Colorectal Cancer Stemness","volume":"7","author":"Masuda","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1007\/s00109-016-1458-4","article-title":"Ankyrin G Expression Is Associated with Androgen Receptor Stability, Invasiveness, and Lethal Outcome in Prostate Cancer Patients","volume":"94","author":"Wang","year":"2016","journal-title":"J. Mol. Med."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"894786","DOI":"10.3389\/fonc.2022.894786","article-title":"Genetic Alterations in Papillary Thyroid Carcinoma with Hashimoto\u2019s Thyroiditis: ANK3, an Indolent Maintainer of Papillary Thyroid Carcinoma","volume":"12","author":"Zeng","year":"2022","journal-title":"Front. Oncol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1016\/j.jgg.2019.11.010","article-title":"DbEMT 2.0: An Updated Database for Epithelial-Mesenchymal Transition Genes with Experimentally Verified Information and Precalculated Regulation Information for Cancer Metastasis","volume":"46","author":"Zhao","year":"2019","journal-title":"J. Genet. Genom."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1186\/s13046-022-02329-x","article-title":"Identification of Distinct Slow Mode of Reversible Adaptation of Pancreatic Ductal Adenocarcinoma to the Prolonged Acidic PH Microenvironment","volume":"41","author":"Wu","year":"2022","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"768579","DOI":"10.3389\/fcell.2022.768579","article-title":"Impact of Extracellular PH on Apoptotic and Non-Apoptotic TRAIL-Induced Signaling in Pancreatic Ductal Adenocarcinoma Cells","volume":"10","author":"Hagelund","year":"2022","journal-title":"Front. Cell Dev. Biol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1952","DOI":"10.1158\/0008-5472.CAN-18-1604","article-title":"Acidification of Tumor at Stromal Boundaries Drives Transcriptome Alterations Associated with Aggressive Phenotypes","volume":"79","author":"Rohani","year":"2019","journal-title":"Cancer Res."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1038\/nrc.2017.77","article-title":"Tumour Acidosis: From the Passenger to the Driver\u2019s Seat","volume":"17","author":"Corbet","year":"2017","journal-title":"Nat. Rev. Cancer"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1007\/s10555-018-09778-x","article-title":"What Is PH Regulation, and Why Do Cancer Cells Need It?","volume":"38","author":"Swietach","year":"2019","journal-title":"Cancer Metastasis Rev."},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"Riemann, A., Schneider, B., Ihling, A., Nowak, M., Sauvant, C., Thews, O., and Gekle, M. (2011). Acidic Environment Leads to ROS-Induced MAPK Signaling in Cancer Cells. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0022445"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1002\/(SICI)1097-4547(19980215)51:4<431::AID-JNR3>3.0.CO;2-D","article-title":"Mechanisms of PHi Regulation Studied in Individual Neurons Cultured from Mouse Cerebral Cortex","volume":"51","author":"Pedersen","year":"1998","journal-title":"J. Neurosci. Res."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1242\/jcs.195297","article-title":"Cancer Cell Behaviors Mediated by Dysregulated PH Dynamics at a Glance","volume":"130","author":"White","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"2997","DOI":"10.1038\/s41467-018-05261-x","article-title":"Systems Analysis of Intracellular PH Vulnerabilities for Cancer Therapy","volume":"9","author":"Persi","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Russell, S., Xu, L., Kam, Y., Abrahams, D., Ordway, B., Lopez, A.S., Bui, M.M., Johnson, J., Epstein, T., and Ruiz, E. (2022). Proton Export Upregulates Aerobic Glycolysis. BMC Biol., 20.","DOI":"10.1186\/s12915-022-01340-0"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"464S","DOI":"10.1042\/bst023464s","article-title":"Protein Catabolism in Metabolic Acidosis: Inhibition of Glycolysis by Low PH Suggests a Role for Glucose","volume":"23","author":"Bevington","year":"1995","journal-title":"Biochem. Soc. Trans."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"120163","DOI":"10.1016\/j.lfs.2021.120163","article-title":"The Role of Tumor Acidification in Aggressiveness, Cell Dissemination and Treatment Resistance of Oral Squamous Cell Carcinoma","volume":"288","author":"Nunes","year":"2022","journal-title":"Life Sci."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"136","DOI":"10.4049\/jimmunol.2100475","article-title":"Extracellular PH Controls Chemotaxis of Neutrophil Granulocytes by Regulating Leukotriene B4 Production and Cdc42 Signaling","volume":"209","author":"Oster","year":"2022","journal-title":"J. Immunol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"310616","DOI":"10.1155\/2012\/310616","article-title":"Turnover of Focal Adhesions and Cancer Cell Migration","volume":"2012","author":"Nagano","year":"2012","journal-title":"Int. J. Cell Biol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1038\/nrc1713","article-title":"The Role of Disturbed PH Dynamics and the Na+H+ exchanger in metastasis","volume":"5","author":"Cardone","year":"2005","journal-title":"Nat. Rev. Cancer"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"847","DOI":"10.1016\/j.ejcb.2012.04.004","article-title":"PH Regulators in Invadosomal Functioning: Proton Delivery for Matrix Tasting","volume":"91","author":"Brisson","year":"2012","journal-title":"Eur. J. Cell Biol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"85","DOI":"10.2174\/1574892811308010085","article-title":"Na+-H+ Exchanger, PH Regulation and Cancer","volume":"8","author":"Reshkin","year":"2012","journal-title":"Recent Pat. Anti-Cancer Drug Discov."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"6244","DOI":"10.1038\/onc.2017.219","article-title":"The Interactome of Metabolic Enzyme Carbonic Anhydrase IX Reveals Novel Roles in Tumor Cell Migration and Invadopodia\/MMP14-Mediated Invasion","volume":"36","author":"Swayampakula","year":"2017","journal-title":"Oncogene"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1038\/s41416-018-0216-5","article-title":"Carbonic Anhydrase IX Is a PH-Stat That Sets an Acidic Tumour Extracellular PH in Vivo","volume":"119","author":"Lee","year":"2018","journal-title":"Br. J. Cancer"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1007\/s10555-008-9169-0","article-title":"EMT, the Cytoskeleton, and Cancer Cell Invasion","volume":"28","author":"Yilmaz","year":"2009","journal-title":"Cancer Metastasis Rev."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Ranieri, D., Guttieri, L., Raffa, S., Torrisi, M.R., and Belleudi, F. (2021). Role of FGFR2c and Its PKC\u03b5 Downstream Signaling in the Control of EMT and Autophagy in Pancreatic Ductal Adenocarcinoma Cells. Cancers, 13.","DOI":"10.3390\/cancers13194993"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.3892\/mmr.2013.1604","article-title":"Knockdown of Slug by RNAi Inhibits the Proliferation and Invasion of HCT116 Colorectal Cancer Cells","volume":"8","author":"Qian","year":"2013","journal-title":"Mol. Med. 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