{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T21:21:13Z","timestamp":1792185673900,"version":"4.4.0"},"reference-count":90,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2022,4,14]],"date-time":"2022-04-14T00:00:00Z","timestamp":1649894400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>During recent decades, zebrafish (Danio rerio) have become one of the most important model organisms in which to study different physiological and biological phenomena. The research field of carbonic anhydrases (CAs) and carbonic anhydrase related proteins (CARPs) is not an exception to this. The best-known function of CAs is the regulation of acid\u2013base balance. However, studies performed with zebrafish, among others, have revealed important roles for these proteins in many other physiological processes, some of which had not yet been predicted in the light of previous studies and suggestions. Examples include roles in zebrafish pigmentation as well as motor coordination. Disruption of the function of these proteins may generate lethal outcomes. In this review, we summarize the current knowledge of CA-related studies performed in zebrafish from 1993\u20132021 that was obtained from PubMed search.<\/jats:p>","DOI":"10.3390\/ijms23084342","type":"journal-article","created":{"date-parts":[[2022,4,14]],"date-time":"2022-04-14T21:44:06Z","timestamp":1649972646000},"page":"4342","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":26,"title":["Roles of Carbonic Anhydrases and Carbonic Anhydrase Related Proteins in Zebrafish"],"prefix":"10.3390","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6938-7835","authenticated-orcid":false,"given":"Ashok","family":"Aspatwar","sequence":"first","affiliation":[{"name":"Faculty of Medicine and Health Technology, Tampere University, 33520 Tampere, Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leo","family":"Syrj\u00e4nen","sequence":"additional","affiliation":[{"name":"Faculty of Medicine and Health Technology, Tampere University, 33520 Tampere, Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7323-8536","authenticated-orcid":false,"given":"Seppo","family":"Parkkila","sequence":"additional","affiliation":[{"name":"Faculty of Medicine and Health Technology, Tampere University, 33520 Tampere, Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,4,14]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1146\/annurev.bi.64.070195.002111","article-title":"Human carbonic anhydrases and carbonic anhydrase deficiencies","volume":"64","author":"Sly","year":"1995","journal-title":"Annu. Rev. Biochem."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1016\/j.bbapap.2009.08.026","article-title":"The gamma class of carbonic anhydrases","volume":"1804","author":"Ferry","year":"2010","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1038\/435042a","article-title":"Biochemistry: A cadmium enzyme from a marine diatom","volume":"435","author":"Lane","year":"2005","journal-title":"Nature"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2094","DOI":"10.1038\/s41396-019-0426-8","article-title":"A new widespread subclass of carbonic anhydrase in marine phytoplankton","volume":"13","author":"Jensen","year":"2019","journal-title":"ISME J."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Aspatwar, A., Tolvanen, M.E.E., Barker, H., Syrj\u00e4nen, L., Valanne, S., Purmonen, S., Waheed, A., Sly, W.S., and Parkkila, S. (2022). Carbonic Anhydrases in Metazoan Model Organisms: Molecules, Mechanisms, and Physiology. Physiol. Rev.","DOI":"10.1152\/physrev.00018.2021"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1038\/nrd2467","article-title":"Carbonic anhydrases: Novel therapeutic applications for inhibitors and activators","volume":"7","author":"Supuran","year":"2008","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Syrj\u00e4nen, L., Tolvanen, M., Hilvo, M., Olatubosun, A., Innocenti, A., Scozzafava, A., Leppiniemi, J., Niederhauser, B., Hyt\u00f6nen, V.P., and Gorr, T.A. (2010). Characterization of the first beta-class carbonic anhydrase from an arthropod (Drosophila melanogaster) and phylogenetic analysis of beta-class carbonic anhydrases in invertebrates. BMC Biochem., 11.","DOI":"10.1186\/1471-2091-11-28"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1186\/1756-3305-7-403","article-title":"Beta carbonic anhydrases: Novel targets for pesticides and anti-parasitic agents in agriculture and livestock husbandry","volume":"7","author":"Barker","year":"2014","journal-title":"Parasit. Vectors"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1186\/1756-3305-7-38","article-title":"Bioinformatic analysis of beta carbonic anhydrase sequences from protozoans and metazoans","volume":"7","author":"Barker","year":"2014","journal-title":"Parasit. Vectors"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"505","DOI":"10.3109\/14756366.2014.944178","article-title":"Drosophila melanogaster: A model organism for controlling Dipteran vectors and pests","volume":"30","author":"Barker","year":"2015","journal-title":"J. Enzyme Inhib. Med. Chem."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1042\/BJ20051102","article-title":"Characterization of CA XV, a new GPI-anchored form of carbonic anhydrase","volume":"392","author":"Hilvo","year":"2005","journal-title":"Biochem. J."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1503","DOI":"10.1016\/j.bmc.2012.08.060","article-title":"Analysis of evolution of carbonic anhydrases IV and XV reveals a rich history of gene duplications and a new group of isozymes","volume":"21","author":"Tolvanen","year":"2013","journal-title":"Bioorg. Med. Chem."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Aspatwar, A., Tolvanen, M.E., and Parkkila, S. (2010). Phylogeny and expression of carbonic anhydrase-related proteins. BMC Mol. Biol., 11.","DOI":"10.1186\/1471-2199-11-25"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"498","DOI":"10.1038\/nature12111","article-title":"The zebrafish reference genome sequence and its relationship to the human genome","volume":"496","author":"Howe","year":"2013","journal-title":"Nature"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1002\/aja.1002030302","article-title":"Stages of embryonic development of the zebrafish","volume":"203","author":"Kimmel","year":"1995","journal-title":"Dev. Dyn."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1038\/79951","article-title":"Effective targeted gene \u2018knockdown\u2019 in zebrafish","volume":"26","author":"Nasevicius","year":"2000","journal-title":"Nat. Genet."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2700","DOI":"10.1101\/gr.1725103","article-title":"Efficient target-selected mutagenesis in zebrafish","volume":"13","author":"Wienholds","year":"2003","journal-title":"Genome Res."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"702","DOI":"10.1038\/nbt1409","article-title":"Heritable targeted gene disruption in zebrafish using designed zinc-finger nucleases","volume":"26","author":"Doyon","year":"2008","journal-title":"Nat. Biotechnol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1038\/nbt.2501","article-title":"Efficient genome editing in zebrafish using a CRISPR-Cas system","volume":"31","author":"Hwang","year":"2013","journal-title":"Nat. Biotechnol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"4827","DOI":"10.1242\/dev.115584","article-title":"Precise and efficient genome editing in zebrafish using the CRISPR\/Cas9 system","volume":"141","author":"Irion","year":"2014","journal-title":"Development"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1038\/nbt.1939","article-title":"Heritable gene targeting in zebrafish using customized TALENs","volume":"29","author":"Huang","year":"2011","journal-title":"Nat. Biotechnol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"4327","DOI":"10.1242\/dev.068080","article-title":"Rapid BAC selection for tol2-mediated transgenesis in zebrafish","volume":"138","author":"Bussmann","year":"2011","journal-title":"Development"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1007\/PL00006163","article-title":"Isolation and characterization of a carbonic anhydrase homologue from the zebrafish (Danio rerio)","volume":"44","author":"Peterson","year":"1997","journal-title":"J. Mol. Evol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.gep.2018.04.007","article-title":"Identification and expression of carbonic anhydrase 2, myosin regulatory light chain 2 and selenium-binding protein 1 in zebrafish Danio rerio: Implication for age-related biomarkers","volume":"29","author":"Feng","year":"2018","journal-title":"Gene Expr. Patterns"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"C1250","DOI":"10.1152\/ajpcell.00021.2008","article-title":"Carbonic anhydrase 2-like a and 15a are involved in acid-base regulation and Na+ uptake in zebrafish H+-ATPase-rich cells","volume":"294","author":"Lin","year":"2008","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1002\/cne.20578","article-title":"How can teleostean inner ear hair cells maintain the proper association with the accreting otolith?","volume":"488","author":"Shiao","year":"2005","journal-title":"J. Comp. Neurol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"59","DOI":"10.3389\/fphys.2013.00059","article-title":"Close Association of Carbonic Anhydrase (CA2a and CA15a), Na+\/H+ Exchanger (Nhe3b), and Ammonia Transporter Rhcg1 in Zebrafish Ionocytes Responsible for Na+ Uptake","volume":"4","author":"Ito","year":"2013","journal-title":"Front. Physiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"R470","DOI":"10.1152\/ajpregu.00200.2006","article-title":"Visualization in zebrafish larvae of Na+ uptake in mitochondria-rich cells whose differentiation is dependent on foxi3a","volume":"292","author":"Esaki","year":"2007","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_29","first-page":"3569","article-title":"Cardiac responses to hypercapnia in larval zebrafish (Danio rerio): The links between CO2 chemoreception, catecholamines and carbonic anhydrase","volume":"217","author":"Miller","year":"2014","journal-title":"J. Exp. Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"3837","DOI":"10.1242\/jeb.034116","article-title":"Carbonic anhydrase expression and CO2 excretion during early development in zebrafish Danio rerio","volume":"212","author":"Gilmour","year":"2009","journal-title":"J. Exp. Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"R55","DOI":"10.1152\/ajpregu.00215.2020","article-title":"Use of a carbonic anhydrase ca17a knockout to investigate mechanisms of ion uptake in zebrafish (Danio rerio)","volume":"320","author":"Zimmer","year":"2021","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Postel, R., and Sonnenberg, A. (2012). Carbonic anhydrase 5 regulates acid-base homeostasis in zebrafish. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0039881"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e4128","DOI":"10.7717\/peerj.4128","article-title":"Identification and characterization of a novel zebrafish (Danio rerio) pentraxin-carbonic anhydrase","volume":"5","author":"Patrikainen","year":"2017","journal-title":"PeerJ"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1093\/hmg\/dds438","article-title":"Abnormal cerebellar development and ataxia in CARP VIII morphant zebrafish","volume":"22","author":"Aspatwar","year":"2013","journal-title":"Hum. Mol. Genet."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"R150","DOI":"10.1152\/ajpregu.90685.2008","article-title":"Hypoxia-inducible carbonic anhydrase IX expression is insufficient to alleviate intracellular metabolic acidosis in the muscle of zebrafish, Danio rerio","volume":"296","author":"Esbaugh","year":"2009","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Aspatwar, A., Tolvanen, M.E., Ojanen, M.J., Barker, H.R., Saralahti, A.K., B\u00e4uerlein, C.A., Ortutay, C., Pan, P., Kuuslahti, M., and Parikka, M. (2015). Inactivation of ca10a and ca10b Genes Leads to Abnormal Embryonic Development and Alters Movement Pattern in Zebrafish. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0134263"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e48333","DOI":"10.15252\/embr.201948333","article-title":"pH-controlled histone acetylation amplifies melanocyte differentiation downstream of MITF","volume":"21","author":"Raja","year":"2020","journal-title":"EMBO Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1096","DOI":"10.1016\/j.cub.2015.02.071","article-title":"Chemokine-Dependent pH Elevation at the Cell Front Sustains Polarity in Directionally Migrating Zebrafish Germ Cells","volume":"25","author":"Tarbashevich","year":"2015","journal-title":"Curr. Biol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"33954","DOI":"10.1038\/srep33954","article-title":"A cytosolic carbonic anhydrase molecular switch occurs in the gills of metamorphic sea lamprey","volume":"6","author":"McCormick","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"3283","DOI":"10.1242\/jeb.033910","article-title":"The involvement of SLC26 anion transporters in chloride uptake in zebrafish (Danio rerio) larvae","volume":"212","author":"Bayaa","year":"2009","journal-title":"J. Exp. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"R84","DOI":"10.1152\/ajpregu.00318.2011","article-title":"Rhcg1 and NHE3b are involved in ammonium-dependent sodium uptake by zebrafish larvae acclimated to low-sodium water","volume":"302","author":"Shih","year":"2012","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1002\/humu.9266","article-title":"Carbonic anhydrase II deficiency syndrome (osteopetrosis with renal tubular acidosis and brain calcification): Novel mutations in CA2 identified by direct sequencing expand the opportunity for genotype-phenotype correlation","volume":"24","author":"Shah","year":"2004","journal-title":"Hum. Mutat."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"488","DOI":"10.1073\/pnas.72.2.488","article-title":"A zinc protein isolated from human parotid saliva","volume":"72","author":"Henkin","year":"1975","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1006\/bbrc.1998.9356","article-title":"Gustin from human parotid saliva is carbonic anhydrase VI","volume":"250","author":"Thatcher","year":"1998","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"3867","DOI":"10.1073\/pnas.78.6.3867","article-title":"Gustin concentration changes relative to salivary zinc and taste in humans","volume":"78","author":"Shatzman","year":"1981","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Melis, M., Atzori, E., Cabras, S., Zonza, A., Cal\u00f2, C., Muroni, P., Nieddu, M., Padiglia, A., Sogos, V., and Tepper, B.J. (2013). The gustin (CA6) gene polymorphism, rs2274333 (A\/G), as a mechanistic link between PROP tasting and fungiform taste papilla density and maintenance. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0074151"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1007\/s11248-010-9441-2","article-title":"Gene expression profiling in the submandibular gland, stomach, and duodenum of CAVI-deficient mice","volume":"20","author":"Pan","year":"2011","journal-title":"Transgenic Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1186\/s12929-014-0082-2","article-title":"The role of carbonic anhydrase VI in bitter taste perception: Evidence from the Car6\u2212\/\u2212 mouse model","volume":"21","author":"Patrikainen","year":"2014","journal-title":"J. Biomed. Sci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1007\/s11248-016-9961-5","article-title":"Altered gene expression in the lower respiratory tract of Car6 (\u2212\/\u2212) mice","volume":"25","author":"Patrikainen","year":"2016","journal-title":"Transgenic Res."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"3264","DOI":"10.2174\/138161210793429823","article-title":"Carbonic anhydrase related protein VIII and its role in neurodegeneration and cancer","volume":"16","author":"Aspatwar","year":"2010","journal-title":"Curr. Pharm. Des."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1007\/978-94-007-7359-2_8","article-title":"Carbonic anhydrase related proteins: Molecular biology and evolution","volume":"75","author":"Aspatwar","year":"2014","journal-title":"Subcell. Biochem."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1111\/j.1460-9568.1990.tb00460.x","article-title":"A Collection of cDNA Clones with Specific Expression Patterns in Mouse Brain","volume":"2","author":"Kato","year":"1990","journal-title":"Eur. J. Neurosci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1006\/mpev.1996.0006","article-title":"Functional diversity, conservation, and convergence in the evolution of the alpha-, beta-, and gamma-carbonic anhydrase gene families","volume":"5","author":"Tashian","year":"1996","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1023\/A:1026433321974","article-title":"Expression of the acatalytic carbonic anhydrase VIII gene, Car8, during mouse embryonic development","volume":"29","author":"Lakkis","year":"1997","journal-title":"Histochem. J."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/S0306-4522(02)00066-0","article-title":"Developmental expression of carbonic anhydrase-related proteins VIII, X, and XI in the human brain","volume":"112","author":"Taniuchi","year":"2002","journal-title":"Neuroscience"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S0169-328X(02)00563-6","article-title":"cDNA cloning and developmental expression of murine carbonic anhydrase-related proteins VIII, X, and XI","volume":"109","author":"Taniuchi","year":"2002","journal-title":"Brain Res. Mol. Brain Res."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1534\/genetics.105.044487","article-title":"Carbonic anhydrase-related protein VIII deficiency is associated with a distinctive lifelong gait disorder in waddles mice","volume":"171","author":"Jiao","year":"2005","journal-title":"Genetics"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"T\u00fcrkmen, S., Guo, G., Garshasbi, M., Hoffmann, K., Alshalah, A.J., Mischung, C., Kuss, A., Humphrey, N., Mundlos, S., and Robinson, P.N. (2009). CA8 mutations cause a novel syndrome characterized by ataxia and mild mental retardation with predisposition to quadrupedal gait. PLoS Genet., 5.","DOI":"10.1371\/journal.pgen.1000487"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"826","DOI":"10.1002\/ajmg.b.31227","article-title":"Phenotypical spectrum of cerebellar ataxia associated with a novel mutation in the CA8 gene, encoding carbonic anhydrase (CA) VIII","volume":"156","author":"Kaya","year":"2011","journal-title":"Am. J. Med. Genet. B Neuropsychiatr. Genet."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1042\/bj20030110","article-title":"Carbonic anhydrase-related protein is a novel binding protein for inositol 1,4,5-trisphosphate receptor type 1","volume":"372","author":"Hirota","year":"2003","journal-title":"Biochem. J."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"480","DOI":"10.1006\/geno.1996.0223","article-title":"Human MN\/CA9 gene, a novel member of the carbonic anhydrase family: Structure and exon to protein domain relationships","volume":"33","author":"Stanbridge","year":"1996","journal-title":"Genomics"},{"key":"ref_62","first-page":"2877","article-title":"Cloning and characterization of MN, a human tumor-associated protein with a domain homologous to carbonic anhydrase and a putative helix-loop-helix DNA binding segment","volume":"9","author":"Pastorek","year":"1994","journal-title":"Oncogene"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"C402","DOI":"10.1152\/ajpcell.00228.2007","article-title":"Carbonic anhydrases IV and IX: Subcellular localization and functional role in mouse skeletal muscle","volume":"294","author":"Scheibe","year":"2008","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"155","DOI":"10.3748\/wjg.v11.i2.155","article-title":"Carbonic anhydrases in normal gastrointestinal tract and gastrointestinal tumours","volume":"11","author":"Saarnio","year":"2005","journal-title":"World J. Gastroenterol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1053\/gast.1997.v112.pm9024293","article-title":"Carbonic anhydrase IX, MN\/CA IX: Analysis of stomach complementary DNA sequence and expression in human and rat alimentary tracts","volume":"112","author":"Parkkila","year":"1997","journal-title":"Gastroenterology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"R1256","DOI":"10.1152\/ajpregu.00735.2004","article-title":"Expression of membrane-associated carbonic anhydrase isoforms IV, IX, XII, and XIV in the rabbit: Induction of CA IV and IX during maturation","volume":"288","author":"Purkerson","year":"2005","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"e4068","DOI":"10.7717\/peerj.4068","article-title":"An update on anticancer drug development and delivery targeting carbonic anhydrase IX","volume":"5","author":"Aspatwar","year":"2017","journal-title":"PeerJ"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1007\/s10555-007-9064-0","article-title":"Regulation of tumor pH and the role of carbonic anhydrase 9","volume":"26","author":"Swietach","year":"2007","journal-title":"Cancer Metastasis Rev."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"164","DOI":"10.4161\/cc.3.2.618","article-title":"Hypoxia inducible carbonic anhydrase IX, marker of tumour hypoxia, survival pathway and therapy target","volume":"3","author":"Potter","year":"2004","journal-title":"Cell Cycle"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1038\/sj.bjc.6600936","article-title":"Diagnostic, prognostic and therapeutic implications of carbonic anhydrases in cancer","volume":"89","author":"Potter","year":"2003","journal-title":"Br. J. Cancer"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1808","DOI":"10.1054\/bjoc.2000.1111","article-title":"Human tumour-associated cell adhesion protein MN\/CA IX: Identification of M75 epitope and of the region mediating cell adhesion","volume":"82","author":"Pastorek","year":"2000","journal-title":"Br. J. Cancer"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1038\/sj.bjc.6601264","article-title":"Soluble form of carbonic anhydrase IX (CA IX) in the serum and urine of renal carcinoma patients","volume":"89","author":"Hyrsl","year":"2003","journal-title":"Br. J. Cancer"},{"key":"ref_73","first-page":"977","article-title":"Carbonic anhydrase IX (CA IX) mediates tumor cell interactions with microenvironment","volume":"13","author":"Zavadova","year":"2005","journal-title":"Oncol. Rep."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"R1079","DOI":"10.1152\/ajpregu.00626.2004","article-title":"Oxygen-dependent gene expression in fishes","volume":"288","author":"Nikinmaa","year":"2005","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"359","DOI":"10.4161\/cbt.4.4.1646","article-title":"Achieving hypoxia-inducible gene expression in tumors","volume":"4","author":"Marignol","year":"2005","journal-title":"Cancer Biol. Ther."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1016\/S0167-4781(01)00193-2","article-title":"cDNA sequence of human carbonic anhydrase-related protein, CA-RP X: mRNA expressions of CA-RP X and XI in human brain","volume":"1518","author":"Okamoto","year":"2001","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1006\/bbrc.1998.9449","article-title":"Sequence and tissue expression of a novel human carbonic anhydrase-related protein, CARP-2, mapping to chromosome 19q13.3","volume":"253","author":"Bellingham","year":"1998","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.3109\/14756366.2012.727813","article-title":"An update on carbonic anhydrase-related proteins VIII, X and XI","volume":"28","author":"Aspatwar","year":"2013","journal-title":"J. Enzym. Inhib. Med. Chem."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1006\/geno.1999.5938","article-title":"Human carbonic anhydrase XIV (CA14): cDNA cloning, mRNA expression, and mapping to chromosome 1","volume":"61","author":"Nishimori","year":"1999","journal-title":"Genomics"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"8514","DOI":"10.1073\/pnas.0702899104","article-title":"Carbonic anhydrase XIV deficiency produces a functional defect in the retinal light response","volume":"104","author":"Ogilvie","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1111\/j.1755-148X.2008.00505.x","article-title":"Novel MITF targets identified using a two-step DNA microarray strategy","volume":"21","author":"Hoek","year":"2008","journal-title":"Pigment Cell Melanoma Res."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/j.theriogenology.2012.10.016","article-title":"Characterization of the carbonic anhydrases 15b expressed in PGCs during early zebrafish development","volume":"79","author":"Wang","year":"2013","journal-title":"Theriogenology"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1113\/jphysiol.2009.184739","article-title":"Zebrafish (Danio rerio) gill neuroepithelial cells are sensitive chemoreceptors for environmental CO2","volume":"588","author":"Qin","year":"2010","journal-title":"J. Physiol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1038\/nrc1075","article-title":"Tumour-cell invasion and migration: Diversity and escape mechanisms","volume":"3","author":"Friedl","year":"2003","journal-title":"Nat. Rev. Cancer"},{"key":"ref_85","first-page":"1347","article-title":"Emerging targets in cancer management: Role of the CXCL12\/CXCR4 axis","volume":"6","author":"Cojoc","year":"2013","journal-title":"Onco Targets Ther."},{"key":"ref_86","first-page":"1539","article-title":"The CXCR4\/CXCL12 axis in cutaneous malignancies with an emphasis on melanoma","volume":"29","author":"Mitchell","year":"2014","journal-title":"Histol. Histopathol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1038\/nrc2620","article-title":"Transitions between epithelial and mesenchymal states: Acquisition of malignant and stem cell traits","volume":"9","author":"Polyak","year":"2009","journal-title":"Nat. Rev. Cancer"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.gde.2004.12.002","article-title":"Mechanisms of cancer cell invasion","volume":"15","author":"Sahai","year":"2005","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1016\/j.ceb.2010.09.005","article-title":"The nuts and bolts of germ-cell migration","volume":"22","author":"Tarbashevich","year":"2010","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"170","DOI":"10.4161\/cam.5.2.14403","article-title":"Dynamics of the Rho-family small GTPases in actin regulation and motility","volume":"5","author":"Spiering","year":"2011","journal-title":"Cell Adhesion Migration"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/8\/4342\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T22:54:06Z","timestamp":1760136846000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/8\/4342"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,4,14]]},"references-count":90,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2022,4]]}},"alternative-id":["ijms23084342"],"URL":"https:\/\/doi.org\/10.3390\/ijms23084342","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,4,14]]}}}