ࡱ> f--1-2-3-4-5-6-7-8-9-:-;-<-=->-?-@-A-B-C-D-E-F-G-H-I-J-K-L-M-N-O-P-Q-R-S-T-U-V-W-X-Y-Z-[-\-]-^-_-`-a-b-c-d-e-f-g-h-i-j-k-l-m-n-o-p-q-r-s-t-u-v-w-x-y-z-{-|-}-~----------------i..T//L00J11y27 r bjbjUU QZ7|7|=lJJJJޅޅޅ'''X(/< 7 DDDDyB,zHzt t t t t t t $Z z ^9t ޅXzixXzXz9t .JJDD$ 6...XzZJ"\DLޅDMn .Xzt . .C lbr#ޅyR D7 05/Р',DYI yR T H GJ 2ؗ 8 ؗ yR .JJJJ UVOD Glikozilacija je proces enzimskog pripajanja oligosaharidnih struktura proteinima ili lipidima pri emu nastaju glikokonjugati  glikoproteini, proteoglikani, glikozilfosfatidil inozitol sidra ili glikolipidi. Prisutnost glikanskih struktura na glikokonjugatima utje e na njihovu veli inu, stabilnost, naboj i antigena svojstva. Zbog svoje iznimne strukturne raznolikosti glikani predstavljaju idealan medij za pohranu bioloakih informacija. Molekule koje mogu "protuma iti" te informacije su lektini, fizioloaki receptori glikana. Nalazimo ih u mikroorganizmima, biljkama i ~ivotinjama. Prepoznavanje ugljikohidrata lektinom temelji se na specifi nim interakcijama domene koja prepoznaje ugljikohidrate (engl. carbohydrate recognition domain, CRD) i ugljikohidrata. Lektinska vezna mjesta su niskog afiniteta, ali su brojna i visoke specifi nosti. Lektini su prvi put otkriveni u biljaka krajem proalog stoljea, te su, nedugo nakon otkria, klasificirani u skupine ovisno o ugljikohidratnim strukturama koje prepoznaju. Razvojem molekularne biologije stvoren je temelj za novi na in klasifikacije lektina, ovisno o njihovoj strukturnoj homologiji i evolucijskoj povezanosti. Biljni su lektini podijeljeni prema biljnim vrstama iz kojih su izolirani  lektini mahunarki i lektini ~itarica. }ivotinjski su lektini na osnovi aminokiselinskog slijeda dijela molekule koji prepoznaje ugljikohidrate podijeljeni u etiri skupine: tip C, tip S, tip P te tip I lektina (Tablica 1.1). Tablica 1.1 Obitelji ~ivotinjskih lektina LektiniDosad poznati broj lanovaOvisnost o Ca2+Specifi nosttip C<20+razli itatip S>14-(-Gal, poli-LacNActip P2+/-Man-6-Ptip I>5-razli ita Brojni se bioloaki procesi, kao ato su oplodnja, embrionalni razvoj, diferencijacija, adhezija i imuni odgovor temelje na visokospecifi nim interakcijama oligosaharidnih struktura glikokonjugata i lektina. 1.1 GALEKTINI Galektini su lektini koje karakterizira specifi no prepoznavanje -galaktozidnih jedinica glikokonjugata te veliki broj o uvanih aminokiselinskih ostataka unutar domene odgovorne za prepoznavanje ugljikohidrata  ADDIN EN.CITE Fred Brewer200210201222327315722-32002Sep 19Binding and cross-linking properties of galectins255-62Departments of Molecular Pharmacology, Microbiology and Immunology, Albert Einstein College of Medicine, 1300 Morris Park Ave., Bronx, New York, NY 10461, USA. brewer@aecom.yu.eduFred Brewer, C.Biochim Biophys ActaAnimalsApoptosisBinding SitesCarbohydrate SequenceCrystallography, X-RayCytoskeletal Proteins/chemistryGalectin 1GalectinsGlycoproteins/chemistryHemagglutinins/*chemistry/physiologyHumanLectins/chemistryMembrane Microdomains/chemistryModels, MolecularMolecular Sequence DataOligosaccharides/chemistry*Plant Lectins*Signal Transduction*Soybean ProteinsStructure-Activity RelationshipT-Lymphocytes/chemistry</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12223273</URL></MDL></Cite></EndNote>[1]. Do danas je poznato 14 razli itih galektina sisavaca, a mnogi su galektini izolirani i iz drugih kraljeanjaka, bezkraljeanjaka (kao ato su nematode i insekti) pa ak i iz protista (spu~vi i gljiva) ADDIN EN.CITE <EndNote><Cite><Author>Cooper</Author><Year>2002</Year><RecNum>103</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>1222327115722-32002Sep 19Galectinomics: finding themes in complexity209-31Departments of Psychiatry and Anatomy, University of California, San Francisco, LPPI, Box F-0984, 401 Parnassus Ave., San Francisco, CA 94143-0984, USA. cooper@cgl.ucsf.eduCooper, D. N.Biochim Biophys ActaAmino Acid SequenceAmphibiaAnimalsBirdsCell Cycle Proteins/geneticsComparative StudyFishesGalectin 4Galectins*Gene Expression RegulationHemagglutinins/chemistry/*geneticsHumanInsectsInvertebratesLectins/geneticsMolecular Sequence DataSequence AlignmentTandem Repeat SequencesViruseshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12223271[2]. Kako se prvotno smatralo da je za punu aktivnost ovih lektina potrebna prisutnost reducirajuih agensa (tiola), njihov je stari naziv bio lektini tipa S. Kasnije je utvreno da to ne vrijedi za sve lanove ove obitelji te su nazvani galektinima  ADDIN EN.CITE <EndNote><Cite><Author>Barondes</Author><Year>1994</Year><RecNum>105081247047641994Feb 25Galectins: a family of animal beta-galactoside-binding lectins597-8Barondes, S. H.Castronovo, V.Cooper, D. N.Cummings, R. D.Drickamer, K.Feizi, T.Gitt, M. A.Hirabayashi, J.Hughes, C.Kasai, K.et al.,CellAnimals*Antigens, DifferentiationBase SequenceGalactosides/chemistryGalectin 1Galectin 2Galectin 3Galectin 4*HemagglutininsHuman*LectinsMolecular Sequence Data</KEYWORD><KEYWORD>Terminology</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8124704</URL></MDL></Cite></EndNote>[3]. Svi galektini sadr~e barem jednu domenu CRD izgraenu od oko 130 aminokiselina, od kojih su mnoge visokoo uvane. Prema strukturnim obilje~jima podijeljeni su u tri skupine: proto-tip, kimera-tip i tandem-tip (Slika 1.1).  Slika 1.1 Strukture galektina. Proto-tip sadr~i jednu domenu CRD, kimera-tip sadr~i jednu domenu CRD povezanu s domenom bogatom prolinom i glicinom, a tandem-tip sadr~i dvije CRD domene. Veina je galektina smjeatena u citosolu; meutim, neki od njih nalaze se i u jezgri i u izvanstani nom prostoru, premda ne sadr~e sljedove karakteristi ne za upuivanje proteina izvan stanice niti transmembransku domenu. Njihova to na uloga nije joa u potpunosti razjaanjena, ali je poznato da mnogi galektini sudjeluju u odr~avanju homeostaze modulirajui stani ni rast, kontrolirajui stani ni ciklus te inducirajui ili inibirajui apoptozu  ADDIN EN.CITE <EndNote><Cite><Author>Hsu</Author><Year>2004</Year><RecNum>1</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>14758074</ACCESSION_NUMBER><VOLUME>19</VOLUME><NUMBER>7-9</NUMBER>2004Regulation of cellular homeostasis by galectins507-15Department of Dermatology, University of California-Davis, School of Medicine, Sacramento, CA 95817, USA. dkhsu@ucdavis.eduHsu, D. K.Liu, F. T.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758074</EndNote>[4]. GALEKTIN-3 1.1.1.1 Strukturna obilje~ja galektina-3 Prije no ato usvojeno danaanje nazivlje, galektin-3 bio je poznat i pod imenom CBP30/35  ADDIN EN.CITE <EndNote><Cite><Author>Roff</Author><Year>1983</Year><RecNum>106</RecNum><MDL>06193115258171983Sep 10Endogenous lectins from cultured cells. Isolation and characterization of carbohydrate-binding proteins from 3T3 fibroblasts10657-63Roff, C. F.Wang, J. L.J Biol ChemAnimals*AsialoglycoproteinsCarrier Proteins/*isolation & purificationCells, CulturedChromatography, AffinityEdetic Acid/pharmacologyElectrophoresis, Polyacrylamide GelFibroblasts/analysisIsoelectric PointLectins/*isolation & purificationMiceMolecular Weight*Protozoan ProteinsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.alpha-Fetoproteins/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=6193115[5], Mac-2  ADDIN EN.CITE Ho19821070617342612831982MarMac-2, a novel 32,000 Mr mouse macrophage subpopulation-specific antigen defined by monoclonal antibodies1221-8Ho, M. K.Springer, T. A.J ImmunolAnimals*Antibodies, Monoclonal/biosynthesisAntigen-Antibody Reactions*Antigens, SurfaceAscitic Fluid/immunologyClone Cells/immunologyConcanavalin A/pharmacologyEpitopesHematopoietic Stem Cells/immunologyLymphoid Tissue/immunologyMacrophages/classification/*immunologyMaleMiceMice, Inbred C57BLMolecular WeightPeptones/pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Thioglycolates/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=6173426[6], L-29  ADDIN EN.CITE Cerra198510802411725260191985Sep 5Three soluble rat beta-galactoside-binding lectins10474-7Cerra, R. F.Gitt, M. A.Barondes, S. H.J Biol ChemAgingAmino Acids/analysisAnimals*AsialoglycoproteinsCarbohydrates/pharmacologyChromatography, AffinityElectrophoresis, Polyacrylamide GelGalactosides/*metabolismGalectinsGlycosides/*metabolismHemagglutinins/*isolation & purification/metabolismLung/growth & development/*metabolismMacromolecular SystemsMolecular WeightRatsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.alpha-Fetoproteinshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=2411725[7], L-34  ADDIN EN.CITE Raz19871090367837517311987NovCloning and expression of cDNA for two endogenous UV-2237 fibrosarcoma lectin genes109-16Department of Cell Biology, Weizmann Institute of Science, Rehovot, Israel.Raz, A.Avivi, A.Pazerini, G.Carmi, P.Exp Cell ResAnimalsCloning, MolecularDNA/geneticsGene Expression RegulationGenes, StructuralLectins/*geneticsMiceMolecular WeightSarcoma, Experimental/*geneticsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tumor Cells, Cultured/*physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=3678375[8], LBP  ADDIN EN.CITE Woo199011002332426265131990May 5The major non-integrin laminin binding protein of macrophages is identical to carbohydrate binding protein 35 (Mac-2)7097-9Laboratory of Cancer Biology, New England Deaconess Hospital, Boston, Massachusetts 02115.Woo, H. J.Shaw, L. M.Messier, J. M.Mercurio, A. M.J Biol ChemAmino Acid SequenceAnimalsAntigens, Differentiation/genetics/*isolation & purification/metabolismBasement Membrane/immunologyCarbohydrates/isolation & purificationChromatography, AffinityChromatography, High Pressure LiquidFemaleGalectin 3Laminin/*metabolismMacrophages/*immunologyMembrane Glycoproteins/genetics/*isolation & purification/metabolismMiceMice, Inbred C57BLMolecular Sequence DataMolecular WeightSequence Homology, Nucleic AcidSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=2332426</URL></MDL></Cite></EndNote>[9] i BP  ADDIN EN.CITE <EndNote><Cite><Author>Liu</Author><Year>1984</Year><RecNum>111</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>6206059259171984Sep 10Synthesis of surface immunoglobulin E receptor in Xenopus oocytes by translation of mRNA from rat basophilic leukemia cells10649-52Liu, F. T.Orida, N.J Biol ChemAnimalsBasophils/immunologyCell Membrane/immunologyFemaleImmunoglobulin E/*metabolismLeukemia, Experimental/*immunologyOocytes/*metabolismPoly A/geneticsRNA/geneticsRNA, Messenger/*geneticsRatsReceptors, IgEReceptors, Immunologic/*geneticsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.*Translation, GeneticXenopushttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=6206059[10]. Galektin-3 je monomer relativne molekulske mase 26 200 do 30 300, ovisno o vrsti iz koje je izoliran, premda se pribli~na molekulska masa, odreena denaturirajuom SDS-poliakrilamidnom elektroforezom, ini vea (29-35 kDa). Humani je galektin-3 izgraen od 249 aminokiselina, miaji od 264, a atakorski od 262 aminokiselina, koje tvore tri strukturno razli ite domene: malu N-terminalnu domenu, domenu koja sadr~i ponavljajue elemente i C-terminalnu domenu koja je odgovorna za lektinsku aktivnost galektina-3. Mala N-terminalna domena izgraena je od 12 aminokiselina i vjerojatno je odgovorna za izlu ivanje galektina-3 iz stanice, obzirom da delecija prvih 11 aminokiselina spre ava njegovu sekreciju  ADDIN EN.CITE <EndNote><Cite><Author>Gong</Author><Year>1999</Year><RecNum>2</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>1062681859241999Dec 15The NH2 terminus of galectin-3 governs cellular compartmentalization and functions in cancer cells6239-45Metastasis Research Program, Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Gong, H. C.Honjo, Y.Nangia-Makker, P.Hogan, V.Mazurak, N.Bresalier, R. S.Raz, A.Cancer ResAntigens, Differentiation/biosynthesis/genetics/*physiologyBinding SitesBiological Transport*Cell CompartmentationCell Division/physiologyCell Transformation, NeoplasticDNA, ComplementaryGalectin 3Gene DeletionHemagglutinationHumanMutagenesis, Site-DirectedNeoplasms/metabolismPeptide Fragments/physiologyPhosphorylationProtein Kinases/metabolismSerine/metabolismSupport, U.S. Gov't, P.H.S.TransfectionTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&amp;list_uids=10626818</URL></MDL></Cite></EndNote>[11]. Galektin-3 mo~e biti fosforiliran na polo~aju Ser6 i Ser12, s time da je viae od 90% fosfata prisutno na Ser6, a manje od 10% na Ser12  ADDIN EN.CITE <EndNote><Cite><Author>Huflejt</Author>1993308253806268351993Dec 15L-29, a soluble lactose-binding lectin, is phosphorylated on serine 6 and serine 12 in vivo and by casein kinase I26712-8Department of Psychiatry, University of California, San Francisco 94143.Huflejt, M. E.Turck, C. W.Lindstedt, R.Barondes, S. H.Leffler, H.J Biol ChemAmino Acid SequenceAnimalsAntigens, Differentiation/*metabolismCell LineDogsGalectin 3HybridomasLectins/*metabolismMiceMolecular Sequence DataPhosphorylationProtein Kinases/*metabolismRabbitsRatsSerine/*metabolismSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8253806[12]. Nefosforilirani oblik galektina-3 (pI 8,7) prisutan je isklju ivo u jezgri, a fosforilirani (pI 8,2) u jezgri i citoplazmi  ADDIN EN.CITE <EndNote><Cite><Author>Tsay</Author><Year>1999</Year><RecNum>4</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10527616</ACCESSION_NUMBER>25221999Nov 1Export of galectin-3 from nuclei of digitonin-permeabilized mouse 3T3 fibroblasts250-61Department of Biochemistry, Michigan State University, East Lansing, Michigan, 48824, USA.Tsay, Y. G.Lin, N. Y.Voss, P. G.Patterson, R. J.Wang, J. L.Exp Cell Res3T3 CellsAnimalsAntigens, Differentiation/*metabolismBiological TransportCell Membrane PermeabilityCell Nucleus/*metabolismDigitoninFibroblasts/*metabolismGalectin 3Indicators and ReagentsLectinsMiceSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10527616[13]. ini se da je fosforilacija Ser6 nu~an preduvjet za antiapoptozno djelovanje galektina-3  ADDIN EN.CITE <EndNote><Cite><Author>Yoshii</Author><Year>2002</Year><RecNum>5</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11724777</ACCESSION_NUMBER>27792002Mar 1Galectin-3 phosphorylation is required for its anti-apoptotic function and cell cycle arrest6852-7Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Wayne State University, 110 E. Warren Ave., Detroit, MI 48201, USA.Yoshii, T.Fukumori, T.Honjo, Y.Inohara, H.Kim, H. R.Raz, A.J Biol Chem*AnoikisAntigens, Differentiation/*metabolism*ApoptosisBinding SitesBlotting, Western*Cell CycleCell LineCell SurvivalCisplatin/pharmacologyCyclin A/metabolismCyclins/metabolismDNA/metabolismDNA, Complementary/metabolismDown-RegulationGalectin 3HumanLigandsMutagenesis, Site-DirectedMutationPhosphorylationPrecipitin TestsProtein BindingProtein Kinases/metabolismProtein Processing, Post-TranslationalProtein Structure, TertiaryProto-Oncogene Proteins c-bcl-2/metabolismSerine/chemistrySignal TransductionSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Time FactorsTransfectionTumor Cells, CulturedUp-Regulationhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11724777Takenaka2004601512185824102004MayNuclear export of phosphorylated galectin-3 regulates its antiapoptotic activity in response to chemotherapeutic drugs4395-406Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Takenaka, Y.Fukumori, T.Yoshii, T.Oka, N.Inohara, H.Kim, H. R.Bresalier, R. S.Raz, A.Mol Cell BiolActive Transport, Cell Nucleus/drug effectsAmino Acid SubstitutionAnimalsAntineoplastic Agents/*pharmacologyApoptosis/*drug effects/*physiologyCell Line, TumorCisplatin/pharmacologyFatty Acids, Unsaturated/pharmacologyGalectin 3/chemistry/genetics/*metabolismHela CellsHumanMAP Kinase Signaling SystemMiceModels, BiologicalMutagenesis, Site-DirectedNIH 3T3 CellsPhosphorylationRecombinant Proteins/chemistry/genetics/metabolismSerine/chemistrySupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Transfectionhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15121858[14, 15]. Fosforilacijom se meutim smanjuje afinitet vezanja glikoproteinskih liganada na galektin-3  ADDIN EN.CITE <EndNote><Cite><Author>Hadari</Author><Year>2000</Year><RecNum>7</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10852818</ACCESSION_NUMBER><VOLUME>113 ( Pt 13)2000JulGalectin-8 binding to integrins inhibits cell adhesion and induces apoptosis2385-97Department of Molecular Cell Biology, The Weizmann Institute of Science, Rehovot 76100, Israel. lizick@weizmann. weizmann.ac.ilHadari, Y. R.Arbel-Goren, R.Levy, Y.Amsterdam, A.Alon, R.Zakut, R.Zick, Y.J Cell SciApoptosis/*physiologyBinding Sites/*physiologyCarbohydrates/metabolismCell Adhesion/*physiologyExtracellular Matrix/metabolism*GalectinsGrowth Inhibitors/metabolismIntegrin alpha3beta1Integrins/*metabolismLectins/*metabolismProtein Structure, TertiaryRecombinant Proteins/metabolismTransfectionTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10852818Mazurek20008010961987275462000Nov 17Phosphorylation of the beta-galactoside-binding protein galectin-3 modulates binding to its ligands36311-5Gastrointestinal Cancer Research Laboratory, Henry Ford Health Sciences Center, Detroit, Michigan 48202, USA.Mazurek, N.Conklin, J.Byrd, J. C.Raz, A.Bresalier, R. S.J Biol ChemAntigens, Differentiation/analysis/*metabolismAsialoglycoproteins/*metabolismColonic Neoplasms/chemistry/metabolismGalactosides/*metabolismGalectin 3HumanIsoelectric FocusingLaminin/*metabolismLigandsMucins/*metabolismNeoplasm Proteins/metabolismPhosphoprotein Phosphatase/metabolismPhosphoproteins/analysis/*metabolismPhosphorylationProtein BindingProtein Kinases/metabolismRecombinant Proteins/metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10961987Openo2000901069444325522000Mar 15Galectin-3 expression and subcellular localization in senescent human fibroblasts278-90Department of Biochemistry, Michigan State University, East Lansing, Michigan, 48824, USA.Openo, K. P.Kadrofske, M. M.Patterson, R. J.Wang, J. L.Exp Cell ResAntigens, Differentiation/*metabolismBiological Transport*Cell AgingCells, CulturedFibroblasts/*metabolism/pathologyFluorescent Antibody TechniqueGalectin 3HumanSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10694443[16-18]. Galektin-3 (izoliran iz stani nih ekstrakata) acetiliran je na svom N-terminalnom kraju  ADDIN EN.CITE <EndNote><Cite><Author>Herrmann</Author><Year>1993</Year><RecNum>162</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8253805</ACCESSION_NUMBER><VOLUME>268</VOLUME>351993Dec 15Primary structure of the soluble lactose binding lectin L-29 from rat and dog and interaction of its non-collagenous proline-, glycine-, tyrosine-rich sequence with bacterial and tissue collagenase26704-11Department of Psychiatry, Langley Porter Psychiatric Institute, University of California, San Francisco 94143.Herrmann, J.Turck, C. W.Atchison, R. E.Huflejt, M. E.Poulter, L.Gitt, M. A.Burlingame, A. L.Barondes, S. H.Leffler, H.J Biol ChemAmino Acid SequenceAnimalsAntigens, Differentiation/*chemistry/metabolismBase SequenceCells, CulturedDNA, ComplementaryDogsGalectin 3Glycine/metabolismHumanLectins/*chemistry/metabolismMicrobial Collagenase/*metabolismMolecular Sequence DataProline/metabolismRatsSequence Homology, Amino AcidSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Tyrosine/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8253805[19]. Na malu N-terminalnu domenu nastavlja se domena izgraena od 8-13 ponavljajuih motiva, strukture nalik kolagenu. Nazvana je domena bogata glicinom i prolinom, s obzirom na to da je slijed aminokiselina u motivu: -Pro-Gly-Ala-Tyr-Pro-Gly-X-X-X- (gdje X predstavlja bilo koju aminokiselinu). Taj visokoo uvani slijed aminokiselina je ~25% istovjetan slijedu nekih heterogenih nuklearnih ribonukleoproteinskih (hnRNP) kompleksa. ini se da ova domena sudjeluje u stvaranju dimera ili visokoureenih oligomera  ADDIN EN.CITE <EndNote><Cite><Author>Ahmad</Author><Year>200410014672941279122004Mar 19Galectin-3 precipitates as a pentamer with synthetic multivalent carbohydrates and forms heterogeneous cross-linked complexes10841-7Department of Molecular Pharmacology, Albert Einstein College of Medicine, Bronx, New York 10461, USA.Ahmad, N.Gabius, H. J.Andre, S.Kaltner, H.Sabesan, S.Roy, R.Liu, B.Macaluso, F.Brewer, C. F.J Biol ChemCarbohydrates/*metabolismGalectin 3/chemistry/*metabolism/ultrastructureMicroscopy, ElectronPrecipitationStructure-Activity RelationshipSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14672941[20] koji omoguavaju multivalentne interakcije neophodne za bioloaku aktivnost galektina-3. Ujedno je supstrat za djelovanje metaloproteinaza  2 i  9 (MMP). Kidanjem veze Ala62-Tyr63 djelovanjem MMP nastaje fragment galektina-3 veli ine 22 kDa s o uvanom lektinskom aktivnoau, ali smanjenom mogunoau stvaranja homoagregata. Domena koja prepoznaje ugljikohidrate obuhvaa ~130 aminokiselina C-terminalnog dijela galektina-3. Sadr~i hidrofilna i hidrofobna podru ja, te je stoga globularne prirode. Graena je od 12 -lanaca organiziranih u dvije -nabrane plo e  ADDIN EN.CITE <EndNote><Cite><Author>Rini</Author><Year>1999</Year><RecNum>11</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10508773</ACCESSION_NUMBER><VOLUME>9</VOLUME><NUMBER>5</NUMBER>1999OctNew animal lectin structures578-84Department of Molecular and Medical Genetics, University of Toronto, Toronto, Ontario, M5S 1A8, Canada. james.rini@utoronto.caRini, J. M.Lobsanov, Y. D.Curr Opin Struct BiolAnimalsAntigens, Differentiation/chemistryGalectin 3*GalectinsHumanLectins/*chemistryMembrane Glycoproteins/chemistryModels, MolecularProtein ConformationProtein Structure, SecondaryReceptor, IGF Type 2/chemistryReceptors, Immunologic/chemistryhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10508773</EndNote>[21] (Slika 1.2).  Slika 1.2 Struktura galektina-3. Domena koja prepoznaje ugljikohidrate (prikazana u kompleksu s laktozom) izgraena je od 12 -lanaca (zeleno) organiziranih u dvije -nabrane plo e. Hipotetska konformacija dijela N-terminalne domene prikazana je plavom bojom  ADDIN EN.CITE Hughes2001120115223968372001JulGalectins as modulators of cell adhesion667-76National Institute for Medical Research, Mill Hill, London NW7 1AA, UK. chughes@nimr.mrc.ac.ukHughes, R. C.BiochimieAnimalsCell Adhesion/*physiologyCell Adhesion Molecules/*genetics/physiologyCell CommunicationExtracellular Matrix/physiologyGalectinsGlycoproteins/physiologyHemagglutinins/chemistry/classification/*physiologyHumanMiceMice, KnockoutModels, BiologicalModels, Molecularhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11522396[22]. C-terminalna domena djelomi no je (<30%) homologna BH1 domeni proteina antiapoptoznog faktora Bcl-2. ini se da je slijed NWGR (Asn-Trp-Gly-Arg), prisutan u CRD galektina-3 kao i u BH1 domeni Bcl-2, odgovoran za antiapoptozno djelovanje oba proteina. Domena CRD vjerojatno ima ulogu i prilikom stvaranja oligomera galektina-3  ADDIN EN.CITE Yang1998130952173037121998Mar 24Role of the carboxyl-terminal lectin domain in self-association of galectin-34086-92Division of Allergy, La Jolla Institute for Allergy and Immunology, San Diego, California 92121, USA.Yang, R. Y.Hill, P. N.Hsu, D. K.Liu, F. T.BiochemistryAmino Acid Substitution/geneticsAntigens, Differentiation/genetics/metabolism/*physiologyBinding, CompetitiveCross-Linking ReagentsGalectin 3Hela CellsHumanImmunoglobulin E/metabolismJurkat CellsLactose/pharmacologyLectins/genetics/metabolism/*physiologyLeucine/geneticsMutagenesis, Site-DirectedPeptide Fragments/metabolismProtein Binding/drug effectsProtein Structure, TertiarySepharose/metabolismSupport, U.S. Gov't, P.H.S.Tryptophan/geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9521730Kuklinski199814094535787021998FebHomophilic binding properties of galectin-3: involvement of the carbohydrate recognition domain814-23Department of Biochemistry, Institute for Animal Anatomy and Physiology, University of Bonn, Germany.Kuklinski, S.Probstmeier, R.J NeurochemAntigens, Differentiation/*chemistry/isolation & purification/*metabolismBinding SitesGalectin 3GlycineIodine RadioisotopesKineticsLaminin/metabolismLectins/chemistry/metabolismMembrane Glycoproteins/chemistry/metabolismProlineRadioligand AssayRecombinant Proteins/chemistry/isolation & purification/metabolismSupport, Non-U.S. Gov'tTyrosinehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9453578[23, 24]. 1.1.1.2 LGALS3 gen za galektin-3 Galektin-3 je kodiran genom LGALS3 (engl. lectin, galactoside-binding, soluble), smjetenom kod ljudi na kromosomu 14, lokusu q21-q22  ADDIN EN.CITE Kadrofske1998150943957734911998Jan 1The human LGALS3 (galectin-3) gene: determination of the gene structure and functional characterization of the promoter7-20Department of Biochemistry, Michigan State University, East Lansing 48824, USA.Kadrofske, M. M.Openo, K. P.Wang, J. L.Arch Biochem BiophysAntigens, Differentiation/*geneticsBase SequenceGalectin 3*Genome, HumanHumanLectins/geneticsMolecular Sequence DataPromoter Regions (Genetics)/*geneticsSequence AlignmentSequence AnalysisSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9439577[25]. LGALS3 je izgraen od pribli~no 17 kb, koje su organizirane u aest eksona i pet introna. Postoje dva mjesta po etka transkripcije, nazvana +1a i +1b, koja se nalaze 52, odnosno 50 nukleotida (nt) uzvodno od spoja ekson I ( intron 1. Po etno mjesto translacije nalazi se u eksonu II. Domena bogata prolinom i glicinom kodirana je unutar eksona III, dok je slijed koji kodira C-terminalnu domenu (CRD) smjeaten u potpunosti unutar eksona V. Genski fragment koji obuhvaa slijed od  836 nt do +141 nt (relativno od +1a) ima sna~nu promotorsku aktivnost. U promotorskoj je regiji smjeateno viae  GC box motiva za vezanje posvuda prisutnog transkripcijskog faktora Sp1, ato je osobina promotora konstitutivno eksprimiranih (engl. housekeeping) gena. No kako ekspresija LGALS3 raste prilikom odgovora stanica na stimulaciju serumom, LGALS3 se smatra genom ranog odgovora (engl. immediate early gene). Regija koja odgovara na aktivaciju serumom smjeatena je izmeu  513 i  339 nt i izmeu  339 i  229 nt; dodatna aktivacijska regija smjeatena je izmeu  105 i  15 nt. U promotorskoj regiji identificirani su sljedovi za vezanje mnogih transkripcijskih faktora, primjerice Sp1, AP-1, NF-(B i CREB (engl. cAMP-response element binding factor). Sazrijevanjem pre-mRNA u miajim fibroblastima 3T3 nastaju dva oblika mRNA, koji se razlikuju u 27 bp neprevedenog slijeda eksona I, a u protein se prevode jednakom u estaloau  ADDIN EN.CITE <EndNote><Cite><Author>Cherayil</Author><Year>1989</Year><RecNum>16</RecNum><MDL><REFERENCE_TYPE>0258493117061989Dec 1The Mac-2 antigen is a galactose-specific lectin that binds IgE1959-72Molecular Immunology Laboratory, Cancer Center of the Massachusetts General Hospital, Boston, Massachusetts.Cherayil, B. J.Weiner, S. J.Pillai, S.J Exp MedAnimalsAntigens, Differentiation/*genetics/physiologyBase SequenceCarrier Proteins/*geneticsCloning, MolecularDNA/analysisGalactose/*metabolismGalectin 3GalectinsHemagglutinins/*genetics/physiologyImmunoglobulin E/*metabolismMacrophages/*immunologyMiceMice, Inbred BALB CMice, Inbred DBAMolecular Sequence DataSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=2584931Raz1989170252506949131989Jul 1Identification of the metastasis-associated, galactoside-binding lectin as a chimeric gene product with homology to an IgE-binding protein3489-93Cancer Metastasis Program, Michigan Cancer Foundation, Detroit, Michigan 48201.Raz, A.Pazerini, G.Carmi, P.Cancer ResAmino Acid SequenceAnimalsAntigens, Differentiation, B-Lymphocyte/geneticsBase SequenceBlotting, WesternCloning, MolecularCollagen/geneticsDNA/geneticsGalectinsHemagglutinins/*geneticsLectins/*geneticsMiceMolecular Sequence DataMolecular WeightMultigene FamilyNeoplasm MetastasisNucleic Acid ConformationRNA, Messenger/ultrastructureRatsReceptors, Fc/geneticsReceptors, IgESarcoma, Experimental/*geneticsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&amp;list_uids=2525069</URL></MDL></Cite></EndNote>[26, 27]. Zna enje alternativnog sazrijevanja do danas nije objaanjeno  ADDIN EN.CITE <EndNote><Cite><Author>Gorski</Author><Year>2002</Year><RecNum>18</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>11886849277212002May 24New alternatively spliced form of galectin-3, a member of the beta-galactoside-binding animal lectin family, contains a predicted transmembrane-spanning domain and a leucine zipper motif18840-8Division of Molecular Biology and Biochemistry, School of Biological Sciences, and Department of Oral Biology, Dental School, University of Missouri-Kansas City, Kansas City, Missouri 64110, USA. GorskiJ@umkc.eduGorski, J. P.Liu, F. T.Artigues, A.Castagna, L. F.Osdoby, P.J Biol Chem*Alternative SplicingAmino Acid SequenceAnimalsAntigens, Differentiation/chemistry/genetics/*metabolismBase SequenceCells, CulturedCloning, MolecularDNA, ComplementaryGalactosides/*metabolismGalectin 3HumanIntestines/metabolism*Leucine ZippersMembrane Proteins/*metabolismMolecular Sequence DataOsteoclasts/metabolismRNA, Messenger/genetics/metabolismSequence Homology, Amino AcidSequence Homology, Nucleic AcidSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11886849[28], premda je utvreno da na taj na in mogu nastati forme galektina-3 (veli ine 36 kDa) koje sadr~e transmembransku domenu  ADDIN EN.CITE <EndNote><Cite><Author>Gorski</Author><Year>2002</Year><RecNum>18</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11886849277212002May 24New alternatively spliced form of galectin-3, a member of the beta-galactoside-binding animal lectin family, contains a predicted transmembrane-spanning domain and a leucine zipper motif18840-8Division of Molecular Biology and Biochemistry, School of Biological Sciences, and Department of Oral Biology, Dental School, University of Missouri-Kansas City, Kansas City, Missouri 64110, USA. GorskiJ@umkc.eduGorski, J. P.Liu, F. T.Artigues, A.Castagna, L. F.Osdoby, P.J Biol Chem*Alternative SplicingAmino Acid SequenceAnimalsAntigens, Differentiation/chemistry/genetics/*metabolismBase SequenceCells, CulturedCloning, MolecularDNA, ComplementaryGalactosides/*metabolismGalectin 3HumanIntestines/metabolism*Leucine ZippersMembrane Proteins/*metabolismMolecular Sequence DataOsteoclasts/metabolismRNA, Messenger/genetics/metabolismSequence Homology, Amino AcidSequence Homology, Nucleic AcidSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11886849[28]. LGALS3 je neobi no graen-unutar njega se nalazi joa jedan gen, galig. Naime, drugi intron unutar LGALS3 sadr~i interni promotor, ijom se aktivacijom stvaraju dva transkripta. Oni zbog pomaknutog okvira itanja kodiraju proteine u potpunosti razli ite od galektina-3  ADDIN EN.CITE Guittaut20011901116012327642001Jan 26Identification of an internal gene to the human Galectin-3 gene with two different overlapping reading frames that do not encode Galectin-32652-7Centre de Biophysique Moleculaire (affiliated with the University of Orleans), CNRS UPR4301, Rue Charles Sadron, 45071 Orleans Cedex 02, France.Guittaut, M.Charpentier, S.Normand, T.Dubois, M.Raimond, J.Legrand, A.J Biol ChemAmino Acid SequenceAntigens, Differentiation/*geneticsBase SequenceBlood Proteins/*geneticsCell CompartmentationCell NucleusCytosolGalectin 3HumanMitochondriaMolecular Sequence Data*Open Reading FramesSupport, Non-U.S. Gov'tTissue DistributionTranscription, GeneticTranslation, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11160123[29]. (-836) TCCAGGCCAGCAGATTTGATGTCTGGTGAGGGCCTGCTTTCTGGTTCACAGAGGGAGCCTTCTGGCTGT GTCTTCACAAGGTGGAAGTGGCAAGGGGACTCTCTCCGGCCTCTTTTATTAAGGCACCAATCTCATTCAC CCTATGACCTAATCACTTCCCAAGGCCTCACTTCCTAATACATCACGTGAGGGTTAGGATTTCAACATATG AACTTTGGCGGGATATAAACATTCAGACTATAGCACCCTGACAGTAAAATGAGATAATAATAACTTATCTC TTTCTTCCAACAAAAAGATAAGGTGAAGTTAAAAGGAGGGTATATATATATATATAATGTGAATTTCCTGTG TAAAATGTGTTAAAGAGTTGTCTGATTATTGCTTTATAAGGGAATTGCTTTGAGACTAGGCCTATTGATCTA GAATAAGTAGTCAATTTGTAGTCAGTTCCCTAGGGAATAGACATTGAAAAGATTTTTGGTTTTGTATTCTAC AAATAAAGCAACCTATTAATTGAATTCCTCTCAGCGAATTCTTCACTCAGGTGATTCTGGAGAGGGCGGG GGCAGACGCGGCCGCAGCCCAGGTCCCGGGAGCGCCACGGAACCTAACGGTGGCAGCGGAGGTCGC GCCCCTCGTGCCCGCGCTCTCCCCGTCGGGAGCTTCCTGGTCGCCCCTGCGGCGGCGGCTCGGGGTC TCAGGCCGGCGCGGGGCTCGCCCAGCCTGGTCCGGGGAGAGGACTGGCTGGCAGGGGCGCCGCCCC GCCTCGGGAGAGGCGGGCCGGGCGGGGCTGGGAGTATTTGAGGCTCGGAGCCACCGCCCCGCCGGC  GCCCGCAGCACCTCCTCGCCAGCAGCAGCCGTCCGGAGCCAGCCAACGAGCGGTGAGCTGCGCGGGG CGCGGGGGACGCGGCTCCGGCCGGGCAGGGGAGAGGGCGCCCGGGCGCTGCTTGGGGCGCGGTCC CGGAGAGGGT Slika 1.3 Slijed nukleotida promotorske regije ljudskog LGALS3. Prikazani slijed prote~e se od  836 nt uzvodno i +141 nt nizvodno od mjesta po etka transkripcije (+1a). Mjesta vezanja transkripcijskih faktora obilje~ena su bojama. U sekvencama  AP-1/SIE box i  AP-1/bHLH box niz DNA za vezanje transkripcijskog faktora AP-1 podvu en je jednostrukom linijom, dok je niz za vezanje SIE, odnosno bHLH podvu en dvostrukom linijom. ( ADDIN EN.CITE <EndNote><Cite><Author>Kadrofske</Author><Year>1998</Year><RecNum>15</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>9439577</ACCESSION_NUMBER>34911998Jan 1The human LGALS3 (galectin-3) gene: determination of the gene structure and functional characterization of the promoter7-20Department of Biochemistry, Michigan State University, East Lansing 48824, USA.Kadrofske, M. M.Openo, K. P.Wang, J. L.Arch Biochem BiophysAntigens, Differentiation/*geneticsBase SequenceGalectin 3*Genome, HumanHumanLectins/geneticsMolecular Sequence DataPromoter Regions (Genetics)/*geneticsSequence AlignmentSequence AnalysisSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9439577</URL></MDL></Cite></EndNote>[25]). Ekspresija LGALS3 se mijenja pod utjecajem razli itih stimulansa. Ovisno o tipu stimulansa, aktiviraju se ili inhibiraju pojedini transkripcijski faktori koji se mogu vezati na promotorske regije LGALS3  ADDIN EN.CITE Dumic20001970108784451032000Expression of galectin-3 in cells exposed to stress-roles of jun and NF-kappaB149-58Department of Biochemistry and Molecular Biology, Faculty of Pharmacy and Biochemistry, University of Zagreb, Ante Kovavica 1, 10000 Zagreb, Croatia.Dumic, J.Lauc, G.Flogel, M.Cell Physiol BiochemAlkylating Agents/pharmacologyAntigens, Differentiation/*geneticsGalectin 3Gene Expression RegulationHeat-Shock ResponseHumanNF-kappa B/*metabolismProto-Oncogene Proteins c-jun/*metabolismSupport, Non-U.S. Gov'tTrypsin/pharmacologyTumor Cells, CulturedUltraviolet Rayshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10878445Liu2004163014961764380Pt 12004May 15Nucling mediates apoptosis by inhibiting expression of galectin-3 through interference with nuclear factor kappaB signalling31-41The Institute for Enzyme Research, The University of Tokushima, 3-18-15 Kuramoto-Cho, Tokushima, 770-8503, Japan.Liu, L.Sakai, T.Sano, N.Fukui, K.Biochem Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14961764[30, 31]. O signalnim putevima kojima se regulira ekspresija LGALS3 zna se vrlo malo. Nedavna su istra~ivanja pokazala da u regulaciji ekspresije galektina-3 u makrofagima va~nu ulogu ima signalni put protein kinaza aktiviranih mitogenima (Ras/MEKK1/MKK1) kojim se aktivira transkripcijski faktor AP-1  ADDIN EN.CITE <EndNote>Kim2003164012788225164112003Jun 17Galectin-3 expression in macrophages is signaled by Ras/MAP kinase pathway and up-regulated by modified lipoproteins13-23Department of Pathology and Microbiology, University of South Carolina-School of Medicine, Columbia, SC 29208, USA.Kim, K.Mayer, E. P.Nachtigal, M.Biochim Biophys ActaAnimalsCell DifferentiationCells, CulturedComparative StudyEnzyme Inhibitors/pharmacologyGalectin 3/*metabolismGene Expression Regulation/*drug effectsGenes, jun/drug effectsMAP Kinase Signaling System/*physiologyMacrophages, Peritoneal/cytology/*metabolismMiceMice, Inbred C3HMitogen-Activated Protein Kinase Kinases/antagonists & inhibitorsProtein Kinase C/antagonists & inhibitorsProtein-Serine-Threonine Kinases/antagonists & inhibitorsProto-Oncogene Proteins c-jun/metabolismRabbits*Signal TransductionSupport, Non-U.S. Gov'tTetradecanoylphorbol Acetate/pharmacologyUp-Regulationras Proteins/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12788225[32]. 1.1.1.3 Stani na raspodjela galektina-3 Galektin-3 eksprimiraju razli ite stani ne vrste, primjerice fagocitni leukociti (monociti, makrofagi, dendritske stanice, mastociti i neutrofili) epitelne stanice, fibroblasti i aktivirani limfociti T, kao i velik broj tumorskih stanica  ADDIN EN.CITE Rabinovich20023501222327515722-32002Sep 19Role of galectins in inflammatory and immunomodulatory processes274-84Division de Immunogenetica, Hospital de Clinicas "Jose de San Martin", Facultad de Medicina, Universidad de Buenos Aires, Cordoba 2351, 3er Piso, CP 1120, Buenos Aires, Argentina. gabyrabi@ciudad.com.arRabinovich, G. A.Rubinstein, N.Toscano, M. A.Biochim Biophys ActaAdjuvants, Immunologic/pharmacology/*physiologyAnimalsApoptosisAutoimmune Diseases/etiology/therapyCell Division/drug effectsComparative StudyGalectinsHemagglutinins/immunology/pharmacology/*physiologyHumanHypersensitivity/etiology/therapyImmunityInflammation/etiology/therapyLymph Nodes/drug effectsSpleen/drug effectsSupport, Non-U.S. Gov'tT-Lymphocytes/*drug effectsThymus Gland/drug effectshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12223275</URL></MDL></Cite></EndNote>[33]. Prisutnost galektina-3 utvrena je u gotovo svim stani nim odjeljcima, ovisno o tipu stanice. U jezgri se galektin-3 nalazi u fosforiliranom i nefosforiliranom obliku  ADDIN EN.CITE Tsay1999401052761625221999Nov 1Export of galectin-3 from nuclei of digitonin-permeabilized mouse 3T3 fibroblasts250-61Department of Biochemistry, Michigan State University, East Lansing, Michigan, 48824, USA.Tsay, Y. G.Lin, N. Y.Voss, P. G.Patterson, R. J.Wang, J. L.Exp Cell Res3T3 CellsAnimalsAntigens, Differentiation/*metabolismBiological TransportCell Membrane PermeabilityCell Nucleus/*metabolismDigitoninFibroblasts/*metabolismGalectin 3Indicators and ReagentsLectinsMiceSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10527616Davidson2002220120700751252002MayShuttling of galectin-3 between the nucleus and cytoplasm329-37Cell and Molecular Biology Program, Michigan State University, East Lansing 48824 USA.Davidson, P. J.Davis, M. J.Patterson, R. J.Ripoche, M. A.Poirier, F.Wang, J. L.Glycobiology3T3 CellsAnimalsAntibodies, Monoclonal/immunologyBlotting, WesternCell Nucleus/*metabolismCytoplasm/*metabolismGalectin 3/immunology/*metabolismHumanMiceMicroscopy, FluorescenceProtein TransportSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12070075[13, 34]. U sklopu ribonukleoproteinskog kompleksa, neovisno o svojoj lektinskoj aktivnosti (premda pomou domene CRD) ve~e jednolan anu DNA ili RNA te djeluje kao imbenik sazrijevanja pre-mRNA  ADDIN EN.CITE <EndNote><Cite><Author>Wang</Author><Year>2004</Year>16501523825116731-22004Jul 6Nucleocytoplasmic lectins75-93Department of Biochemistry and Molecular Biology, Michigan State University, 501 Biochemistry Building, East Lansing, MI 48824, USA. wangj@msu.eduWang, J. L.Gray, R. M.Haudek, K. C.Patterson, R. J.Biochim Biophys ActaAnimalsAnnexins/metabolismCell Nucleus/*metabolismCytoplasm/*metabolismGalectins/metabolismHumanLectins/*metabolismLigandsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov&apos;t, P.H.S.</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=15238251</URL></MDL></Cite></EndNote>[35]. ini se da galektin-3 mo~e ak modulirati transkripcijsku aktivnost pojedinih transkripcijskih faktora  ADDIN EN.CITE Paron20032301261506930232003Mar 14Nuclear localization of Galectin-3 in transformed thyroid cells: a role in transcriptional regulation545-53Dipartimento di Scienze e Tecnologie Biomediche, P.le Kolbe 1, Universita degli Studi di Udine, 33100 Udine, Italy.Paron, I.Scaloni, A.Pines, A.Bachi, A.Liu, F. T.Puppin, C.Pandolfi, M.Ledda, L.Di Loreto, C.Damante, G.Tell, G.Biochem Biophys Res CommunAnimalsBlotting, WesternCell DifferentiationCell DivisionCell LineCell Line, TransformedCell Nucleus/*metabolismCytoplasm/metabolismElectrophoresis, Gel, Two-DimensionalGalectin 3/*metabolism/*physiology*Gene Expression Regulation, NeoplasticGlutathione Transferase/metabolismHela CellsHumanImmunohistochemistryNuclear Proteins/metabolismOligonucleotides/pharmacologyPlasmids/metabolismProtein BindingProtein Structure, TertiaryRatsRecombinant Proteins/metabolismSpectrometry, Mass, Matrix-Assisted Laser Desorption-IonizationSpectrum Analysis, MassSupport, Non-U.S. Gov'tThyroid Gland/*cytologyTranscription Factors/metabolism*Transcription, GeneticTransfectionTumor Cells, CulturedUp-Regulationhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12615069[36]. Galektin-3 nalazimo i u mitohondrijima  ADDIN EN.CITE Yu200224011839755277182002May 3Galectin-3 translocates to the perinuclear membranes and inhibits cytochrome c release from the mitochondria. A role for synexin in galectin-3 translocation15819-27Department of Pathology, Wayne State University School of Medicine and Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Yu, F.Finley, R. L., Jr.Raz, A.Kim, H. R.J Biol ChemAnnexin VII/*metabolismAntigens, Differentiation/genetics/*metabolismApoptosis/drug effects/physiologyBreast Neoplasms/pathologyCaspases/metabolismCytochrome c Group/*metabolismElectron Transport Complex IV/metabolismEnzyme ActivationEpithelial Cells/metabolismFemaleGalectin 3HumanIntracellular Membranes/metabolismMembrane Glycoproteins/metabolismMicroscopy, ConfocalMitochondria/*metabolismNuclear Envelope/*metabolismProtein TransportRecombinant Fusion Proteins/metabolismStaurosporine/pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11839755[37], u fagosomima makrofaga i egzosomima dendritskih stanica  ADDIN EN.CITE Garin20012501114992915212001Jan 8The phagosome proteome: insight into phagosome functions165-80Laboratoire de Chimie des proteines, Commissariat a l'Energie Atomique, 38054 Grenoble, France.Garin, J.Diez, R.Kieffer, S.Dermine, J. F.Duclos, S.Gagnon, E.Sadoul, R.Rondeau, C.Desjardins, M.J Cell BiolAnimalsBlotting, Western/methodsCell LineElectrophoresis, Gel, Two-DimensionalFluorescent Antibody TechniqueGTP Phosphohydrolases/metabolismHydrolases/metabolismMembrane FusionMembrane Proteins/analysisMicePhagosomes/chemistry/*physiologyProteins/analysisProteome/analysis/*physiologySpectrum Analysis, Mass/methodsSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11149929Thery200126011390481166122001Jun 15Proteomic analysis of dendritic cell-derived exosomes: a secreted subcellular compartment distinct from apoptotic vesicles7309-18Institut National de la Sante et de la Recherche Medical, Unite 520, Institut Curie, Paris, France. clotilde.thery@curie.frThery, C.Boussac, M.Veron, P.Ricciardi-Castagnoli, P.Raposo, G.Garin, J.Amigorena, S.J ImmunolAnimals*Apoptosis/immunologyCell LineCell Membrane/chemistry/secretion/ultrastructureComparative StudyCytoplasmic Vesicles/chemistry/*secretion/ultrastructureCytosol/chemistry/secretion/ultrastructureDendritic Cells/chemistry/*secretion/ultrastructureEndocytosis/immunologyMicePeptide MappingProteome/analysis/*secretion/ultrastructureSubcellular Fractions/chemistry/secretion/ultrastructureSupport, Non-U.S. Gov'tTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=11390481</URL></MDL></Cite></EndNote>[38, 39]. U veini stanica galektin-3 nalazi se mahom u citosolu, isklju ivo u fosforiliranom obliku. Mo~e se u razli itoj mjeri izlu ivati u izvanstani ni prostor, te se potom vezati na izvanstani ne glikoproteine kao i na membranske receptore okolnih stanica ili stanice iz koje je izlu en. Po vezanju liganada, galektin-3 mo~e oligomerizirati. 1.1.1.4 Sekrecija galektina-3 Kapacitet sekrecije galektina-3 pojedinih tipova stanica je jako razli it. Primjerice, mirujui miaji makrofagi WEHI-3 izlu uju izrazito malo (ako uope) galektina-3, stanice BHK  ADDIN EN.CITE <EndNote><Cite><Author>Sato</Author><Year>1994</Year><RecNum>27</RecNum>080205582411994JanControl of Mac-2 surface expression on murine macrophage cell lines216-21National Institute for Medical Research, London, UK.Sato, S.Hughes, R. C.Eur J ImmunolAnimalsAntibodies, MonoclonalAntigens, Differentiation/*biosynthesis/isolation & purificationBlotting, WesternCell LineElectrophoresis, Polyacrylamide GelFlow CytometryGalectin 3LectinsMacrophages/*cytology/*immunologyMicePrecipitin Testshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8020558[40] i MDCK  ADDIN EN.CITE Sato1993280831977420711993JulSecretion of the baby hamster kidney 30-kDa galactose-binding lectin from polarized and nonpolarized cells: a pathway independent of the endoplasmic reticulum-Golgi complex8-18National Institute for Medical Research, London, UK.Sato, S.Burdett, I.Hughes, R. C.Exp Cell ResAmino Acid SequenceAnimalsBinding SitesCalcimycin/pharmacologyCell Membrane/chemistry/physiologyCell PolarityCells, Cultured/drug effects/secretionExocytosisGalactose/metabolismGalectinsHamstersHemagglutinins/chemistry/isolation & purification/*metabolismKidney/cytology/drug effects/*secretionLaminin/metabolismMethylamines/pharmacologyMiceMolecular Sequence DataSequence Homology, Amino Acidhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8319774[41] 10-15%, makrofagi J774.2 30-45%  ADDIN EN.CITE Sato199427080205582411994JanControl of Mac-2 surface expression on murine macrophage cell lines216-21National Institute for Medical Research, London, UK.Sato, S.Hughes, R. C.Eur J ImmunolAnimalsAntibodies, MonoclonalAntigens, Differentiation/*biosynthesis/isolation & purificationBlotting, WesternCell LineElectrophoresis, Polyacrylamide GelFlow CytometryGalectin 3LectinsMacrophages/*cytology/*immunologyMicePrecipitin Testshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8020558[40], a stanice CHO-K1 gotovo 100% ukupne koli ine galektina-3  ADDIN EN.CITE <EndNote><Cite><Author>Lindstedt</Author><Year>1993</Year><RecNum>29</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8505302</ACCESSION_NUMBER><VOLUME>268</VOLUME><NUMBER>16</NUMBER><YEAR>1993</YEAR>Jun 5Apical secretion of a cytosolic protein by Madin-Darby canine kidney cells. Evidence for polarized release of an endogenous lectin by a nonclassical secretory pathway11750-7Department of Psychiatry, University of California San Francisco 94143.Lindstedt, R.Apodaca, G.Barondes, S. H.Mostov, K. E.Leffler, H.J Biol ChemAnimalsBrefeldin ACalcimycin/pharmacologyCell LineChromatography, AffinityCyclopentanes/pharmacologyCysteine/metabolismCytosol/metabolismDogsElectrophoresis, Polyacrylamide GelImmunohistochemistryKidneyKineticsL-Lactate Dehydrogenase/metabolismLectins/biosynthesis/isolation & purification/*metabolismMethionine/metabolismMethylamines/pharmacologyMolecular WeightMonensin/pharmacologyNocodazole/pharmacologySulfur RadioisotopesSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.TemperatureVerapamil/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8505302[42]. Sekrecija galektina-3 ovisna je i o razvojnom stupnju i aktiviranosti stanice. Mirujui makrofagi izlu uju malo, a makrofagi potaknuti tioglikolatom ak 30-40% galektina-3. Galektin-3, kao ni drugi galektini, ne sadr~i signalne sljedove za upuivanje u endoplazmatski retikulum niti transmembransku domenu. Istra~ivanja temeljena na primjeni inhibitora razli itih sekretornih puteva pokazala su da se galektin-3 ne izlu uje klasi nim sekretornim putem kroz endoplazmatski retikulum i Golgijev aparat (Tablica 1.2). Tablica 1.2 Utjecaj razli itih tvari na sekreciju galektina-3  ADDIN EN.CITE Hughes199930010580137147311999Dec 6Secretion of the galectin family of mammalian carbohydrate-binding proteins172-85National Institute for Medical Research, Mill Hill, London, UK. c-hughes@nimr.mrc.ac.ukHughes, R. C.Biochim Biophys ActaATP-Binding Cassette Transporters/metabolismAnimalsAntigens, Differentiation/metabolismBinding SitesBiological TransportCarrier Proteins/*metabolismCell AdhesionCell LineCell Membrane/*metabolismCell PolarityCytosol/metabolismEpithelial Cells/metabolismExtracellular Matrix/metabolismGalectin 3GalectinsHemagglutinins/*metabolismHumanPeriplasm/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10580137[43]. CiljTretmanRelativna sekrecija*BHKMDCKtoplinski ok42C++put ER-Golgibrefeldin ANPNPmonezin++18-20CNP-unos Ca2+A23187++endo/egzocitozametilamin-NPnedostatak seruma-citoskeletcitokalazin BNPkolhicinNPnokodazol-mdr sustavverapamilNPciklosporin+* = izra~eno u odnosu na kontrolu pri 37C NP = nema promjene, + = poveana sekrecija, - = smanjena sekrecija mdr sustav = sustav kojim se ostvaruje otpornost na razli ite lijekove (engl. multi drug resistant system) Zasad joa nije u potpunosti poznat na in sekrecije galektina-3, ali postoje naznake da se on izlu uje procesom ektocitoze  nakon nakupljanja galektina-3 uz stani nu membranu, stvaraju se pupovi membrane unutar kojih se nalazi galektin-3. Oni se odjeljuju od membrane kao izvanstani ne vezikule iz kojih se, nakon naruaavanja njihove strukture, oslobaa galektin-3  ADDIN EN.CITE <EndNote><Cite><Author>Hughes</Author><Year>1999</Year><RecNum>30</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10580137</ACCESSION_NUMBER><VOLUME>1473</VOLUME><NUMBER>1</NUMBER><YEAR>1999Dec 6Secretion of the galectin family of mammalian carbohydrate-binding proteins172-85National Institute for Medical Research, Mill Hill, London, UK. c-hughes@nimr.mrc.ac.ukHughes, R. C.Biochim Biophys ActaATP-Binding Cassette Transporters/metabolismAnimalsAntigens, Differentiation/metabolismBinding SitesBiological TransportCarrier Proteins/*metabolismCell AdhesionCell LineCell Membrane/*metabolismCell PolarityCytosol/metabolismEpithelial Cells/metabolismExtracellular Matrix/metabolismGalectin 3GalectinsHemagglutinins/*metabolismHuman<KEYWORD>Periplasm/metabolism</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10580137</URL></MDL></Cite></EndNote>[43]. Sekrecija se galektina-3 u nekim stanicama mo~e ostvariti i putem vezikula koje se stapaju sa stani nom membranom  ADDIN EN.CITE <EndNote><Cite><Author>Craig</Author><Year>1995</Year><RecNum>31</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>7668406</ACCESSION_NUMBER><VOLUME>242</VOLUME>21995JunImmunoelectron microscopic localization of galectin-3, an IgE binding protein, in human mast cells and basophils211-9Department of Anatomy, Virginia Commonwealth University, Richmond 23298, USA.Craig, S. S.Krishnaswamy, P.Irani, A. M.Kepley, C. L.Liu, F. T.Schwartz, L. B.Anat RecAdultAnimalsAntigens, Differentiation/*metabolismBasophils/*immunology/*ultrastructureCell Degranulation/immunologyCytoplasmic Granules/immunologyDogsGalectin 3HamstersHumanImmunoglobulin E/*metabolismInfantMast Cells/*immunology/*ultrastructureMicroscopy, ImmunoelectronProtein BindingRatsSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7668406Thery200126011390481166122001Jun 15Proteomic analysis of dendritic cell-derived exosomes: a secreted subcellular compartment distinct from apoptotic vesicles7309-18Institut National de la Sante et de la Recherche Medical, Unite 520, Institut Curie, Paris, France. clotilde.thery@curie.frThery, C.Boussac, M.Veron, P.Ricciardi-Castagnoli, P.Raposo, G.Garin, J.Amigorena, S.J ImmunolAnimals*Apoptosis/immunologyCell LineCell Membrane/chemistry/secretion/ultrastructureComparative StudyCytoplasmic Vesicles/chemistry/*secretion/ultrastructureCytosol/chemistry/secretion/ultrastructureDendritic Cells/chemistry/*secretion/ultrastructureEndocytosis/immunologyMicePeptide MappingProteome/analysis/*secretion/ultrastructureSubcellular Fractions/chemistry/secretion/ultrastructureSupport, Non-U.S. Gov'tTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11390481[39, 44] ili izravnom translokacijom posredovanom transmembranskim prijenosnicima  ADDIN EN.CITE Sato1993280831977420711993JulSecretion of the baby hamster kidney 30-kDa galactose-binding lectin from polarized and nonpolarized cells: a pathway independent of the endoplasmic reticulum-Golgi complex8-18National Institute for Medical Research, London, UK.Sato, S.Burdett, I.Hughes, R. C.Exp Cell ResAmino Acid SequenceAnimalsBinding SitesCalcimycin/pharmacologyCell Membrane/chemistry/physiologyCell PolarityCells, Cultured/drug effects/secretionExocytosisGalactose/metabolismGalectinsHamstersHemagglutinins/chemistry/isolation & purification/*metabolismKidney/cytology/drug effects/*secretionLaminin/metabolismMethylamines/pharmacologyMiceMolecular Sequence DataSequence Homology, Amino Acidhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8319774Lindstedt19932908505302268161993Jun 5Apical secretion of a cytosolic protein by Madin-Darby canine kidney cells. Evidence for polarized release of an endogenous lectin by a nonclassical secretory pathway11750-7Department of Psychiatry, University of California San Francisco 94143.Lindstedt, R.Apodaca, G.Barondes, S. H.Mostov, K. E.Leffler, H.J Biol ChemAnimalsBrefeldin ACalcimycin/pharmacologyCell LineChromatography, AffinityCyclopentanes/pharmacologyCysteine/metabolismCytosol/metabolismDogsElectrophoresis, Polyacrylamide GelImmunohistochemistryKidneyKineticsL-Lactate Dehydrogenase/metabolismLectins/biosynthesis/isolation & purification/*metabolismMethionine/metabolismMethylamines/pharmacologyMolecular WeightMonensin/pharmacologyNocodazole/pharmacologySulfur RadioisotopesSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.TemperatureVerapamil/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8505302[41, 42]. 1.1.1.5 Specifi nost galektina-3 prema ugljikohidratnim strukturama Galektin-3, kao i ostali galektini, prepoznaje i ve~e galaktozu. Interakcije izmeu domene koja prepoznaje ugljikohidrate i galaktoze su slabe, s konstantom disocijacije mikromolarnog reda veli ina. Galektin-3 pokazuje meutim puno vei afinitet vezanja za laktozu i N-acetillaktozamin, kao i za mnoge druge oligosaharide koji sadr~e galaktozu (Tablica 1.3). Tablica 1.3 Relativni afiniteti vezanja galektina-3 na glikoligande  ADDIN EN.CITE <EndNote><Cite>Ochieng200433014758076197-92004Extracellular functions of galectin-3527-35Department of Biochemistry, Meharry Medical College, Nashville, TN 37208, USA. jochieng@mmc.eduOchieng, J.Furtak, V.Lukyanov, P.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758076[45]. LigandRelativan afinitet*Lac1,0**1,0***LacNAc6,62-(Fuc-Lac3,03-(Gal-LacNAc23,33,6 LacNAc2-Lac29,13-(GalNAc-(2-(Fuc)-Gal16,73-(GalNAc-(2-(Fuc)-Gal17,33-NAc-LacNAc11,63-NBzl-LacNAc97,63-Nsuc-LacNAc105,03-N(4-MeO-F4-Blz)-LacNAc311,0* izmjereno pri 25-26C ** izmjereno mikrokalorimetrijom  ADDIN EN.CITE Sorme20025301192139632-32002Mar 1Low micromolar inhibitors of galectin-3 based on 3'-derivatization of N-acetyllactosamine183-9Section MIG (Microbiology, Immunology, Glycobiology), Dept. of Laboratory Medicine, Lund University, Solvegatan 23, 22362 Lund, Sweden.Sorme, P.Qian, Y.Nyholm, P. G.Leffler, H.Nilsson, U. J.ChembiochemAmino Sugars/chemical synthesis/*pharmacologyAntigens, Differentiation/*immunologyBinding SitesDrug DesignEnzyme-Linked Immunosorbent AssayGalectin 3Support, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11921396[46] *** izmjereno postupkom ELISA  ADDIN EN.CITE Barboni1999340106197091671999JulKinetic measurements of binding of galectin 3 to a laminin substratum365-73National Institute for Medical Research, Mill Hill, London, UK.Barboni, E. A.Bawumia, S.Hughes, R. C.Glycoconj JAmino Acid SequenceAnimalsAntigens, Differentiation/*chemistry/geneticsCattleGalectin 3HamstersHumanKineticsLaminin/*chemistryMembrane Glycoproteins/*chemistryMiceMolecular Sequence DataMutationProtein BindingProtein Structure, TertiarySupport, Non-U.S. Gov'tSurface Plasmon Resonancehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10619709</URL></MDL></Cite></EndNote>[47] Premda je galektin-3 monomer koji sadr~i samo jedno vezno mjesto za ugljikohidrate, mo~e se pozitivno kooperativno vezati na svoje glikoproteinske ligande. Kineti ka mjerenja su pokazala da se vezanje odvija u dva koraka: brzo vezanje malog afiniteta prati sporo vezanje visokog afiniteta  ADDIN EN.CITE <EndNote><Cite><Author>Barboni</Author><Year>1999</Year><RecNum>34</RecNum><MDL><REFERENCE_TYPE>0106197091671999JulKinetic measurements of binding of galectin 3 to a laminin substratum365-73National Institute for Medical Research, Mill Hill, London, UK.Barboni, E. A.Bawumia, S.Hughes, R. C.Glycoconj JAmino Acid SequenceAnimalsAntigens, Differentiation/*chemistry/geneticsCattleGalectin 3HamstersHumanKineticsLaminin/*chemistryMembrane Glycoproteins/*chemistryMiceMolecular Sequence DataMutationProtein BindingProtein Structure, TertiarySupport, Non-U.S. Gov'tSurface Plasmon Resonancehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10619709[47]. Nedavna su istra~ivanja pokazala da se visoki afinitet posti~e stvaranjem multivalentnog lektinskog kompleksa, putem interakcija izmeu N-terminalnih dijelova molekula galektina-3. Veina do danas poznatih liganada galektina-3 su membranski ili izvanstani ni glikoproteini na koje se galektin-3 ve~e nakon ato se izlu i iz stanice. Valja meutim naglasiti da s unutarstani nim ligandima (osim s citokeratinom i CBP70) galektin-3 ostvaruje interakcije neovisne o lektinskoj aktivnosti. 1.1.1.6 Uloge galektina-3 To na uloga galektina-3 nije poznata, no promjene njegove ekspresije, stani nog smjeataja i sekrecije uo ene su tijekom embrionalnog razvoja, karcinogeneze, metastaziranja tumora, imunim reakcijama, odnosno procesima koji se temelje na meudjelovanju stanica, stanica i izvanstani nog matriksa te prijenosu signala  ADDIN EN.CITE <EndNote><Cite><Author>Ochieng</Author><Year>2004</Year><RecNum>33</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>14758076</ACCESSION_NUMBER><VOLUME>19</VOLUME>7-92004Extracellular functions of galectin-3527-35Department of Biochemistry, Meharry Medical College, Nashville, TN 37208, USA. jochieng@mmc.eduOchieng, J.Furtak, V.Lukyanov, P.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758076Rabinovich2002200120723712362002JunGalectins and their ligands: amplifiers, silencers or tuners of the inflammatory response?313-20Division of Immunogenetics, Hospital de Clinicas Jose de San Martin, School of Medicine, University of Buenos Aires, Cordoba 2351, 3er Piso (1120), Buenos Aires, Argentina.Rabinovich, G. A.Baum, L. G.Tinari, N.Paganelli, R.Natoli, C.Liu, F. T.Iacobelli, S.Trends ImmunolAmino Sugars/metabolismAnimalsApoptosis/physiologyAutoimmune Diseases/immunologyCattleCell CommunicationCell SurvivalChagas Disease/immunologyExtracellular Matrix/metabolismGalectinsHemagglutinins/classification/*physiologyHomeostasisHumanInfection/immunologyInflammation/*metabolismLigandsMammals/immunology/metabolismMiceMice, Inbred DBAModels, MolecularNeoplasms/immunology/pathologyPolysaccharides/metabolismProtein ConformationProtein Isoforms/physiologyT-Lymphocyte Subsets/immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12072371Dumic2000200010817217553-42000Mar-AprTransfer to in vitro conditions influences expression and intracellular distribution of galectin-3 in murine peritoneal macrophages261-6Department of Biochemistry and Molecular Biology, Faculty of Pharmacy and Biochemistry, University of Zagreb, Croatia.Dumic, J.Lauc, G.Hadzija, M.Flogel, M.Z Naturforsch [C]AnimalsAntigens, Differentiation/*metabolismCells, CulturedGalectin 3Macrophages, Peritoneal/*metabolismMaleMiceMice, Inbred BALB CRatsSubcellular Fractions/metabolismSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10817217Dumic2000198010938585332000MayGalectin-3 decreases in mice exposed to immobilization stress241-6Department of Biochemistry and Molecular Biology, Faculty of Pharmacy and Biochemistry, University of Zagreb, Ante Kovacica 1, 10000 Zagreb, Croatia.Dumic, J.Barisic, K.Flogel, M.Lauc, G.StressAging/*physiologyAnimalsAntigens, Differentiation/*metabolismCell LineGalectin 3Glioblastoma/metabolism/pathologyHeat-Shock Response/physiologyImmunoblottingLiver/metabolismMacrophages, Peritoneal/metabolismMiceMice, Inbred C3HMice, Inbred StrainsRestraint, PhysicalSpleen/metabolismStress/*metabolismSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10938585[45, 48-50]. Galektin-3 sudjeluje u regulaciji rasta stanica, inhibiciji apoptoze i procesiranju pre-mRNA  ADDIN EN.CITE Liu20023601222327415722-32002Sep 19Intracellular functions of galectins263-73Department of Dermatology, University of California, Davis, School of Medicine, 4860 Y Street, Suite 3400, Sacramento, CA 95817, USA.Liu, F. T.Patterson, R. J.Wang, J. L.Biochim Biophys ActaAnimalsAntigens, Differentiation/chemistryApoptosisCell CycleCell Cycle Proteins/chemistryCytosol/*chemistryGalectin 1Galectin 3GalectinsHemagglutinins/*chemistry/*physiologyHumanLectins/chemistryOrganelles/*chemistryRNA SplicingSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12223274[51]. Ligandi galektina-3 te funkcije posredovane njegovim vezanjem prikazani su u tablici 1.4. Tablica 1.4 Ligandi galektina-3. LigandFunkcija posredovana vezanjem galektina-3*Bcl-2 antiapoptozno djelovanje  ADDIN EN.CITE Akahani1997370939374857231997Dec 1Galectin-3: a novel antiapoptotic molecule with a functional BH1 (NWGR) domain of Bcl-2 family5272-6Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Akahani, S.Nangia-Makker, P.Inohara, H.Kim, H. R.Raz, A.Cancer ResAmino Acid SequenceAntigens, Differentiation/*biosynthesis*Apoptosis/drug effectsBreast NeoplasmsCell SurvivalCisplatin/pharmacologyConserved SequenceDNA FragmentationFemaleGalectin 3HumanKineticsMembrane Glycoproteins/biosynthesisMolecular Sequence DataPoly(ADP-ribose) Polymerases/metabolismProto-Oncogene Proteins/biosynthesis/chemistryProto-Oncogene Proteins c-bcl-2/*biosynthesis/chemistrySequence AlignmentSequence Homology, Amino AcidSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9393748[52]*Gemin4 procesiranje pre-mRNA  ADDIN EN.CITE Park20013801152282929172001Sep 1Association of galectin-1 and galectin-3 with Gemin4 in complexes containing the SMN protein3595-602Department of Microbiology and Molecular Genetics, Michigan State University, East Lansing, MI 48824, USA.Park, J. W.Voss, P. G.Grabski, S.Wang, J. L.Patterson, R. 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C.Glycoconj JAmino Acid SequenceAnimalsAntigens, CD/metabolismAntigens, CD98Antigens, Differentiation/*metabolismCarrier Proteins/metabolismChromatography, AffinityFemaleGalectin 3Macrophage-1 Antigen/metabolismMacrophages/*chemistry/metabolismMacrophages, Peritoneal/chemistry/metabolismMembrane Glycoproteins/*chemistry/immunology/*metabolismMiceMice, Inbred BALB CPrecipitin TestsRatsSequence AnalysisSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9111144[73, 74]CD66a/CD66b (humani neutrofili)poti e aktivaciju NADH oksidaze i adheziju stanica na izvanstani ni matriks  ADDIN EN.CITE <EndNote><Cite><Author>Feuk-Lagerstedt</Author><Year>1999</Year><RecNum>75</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>10553088163101999Nov 15Identification of CD66a and CD66b as the major galectin-3 receptor candidates in human neutrophils5592-8The Phagocyte Research Laboratory, Department of Medical Microbiology and Immunology, University of Goteborg, Sweden.Feuk-Lagerstedt, E.Jordan, E. 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C.FEBS LettAnimalsAntigens, CD/physiologyAntigens, CD98Antigens, Differentiation/biosynthesis/drug effects/*physiologyCalcium/*metabolismCalcium Channel Blockers/pharmacologyCarrier Proteins/physiologyCell LineFlow CytometryGalectin 3GlycosylationHamstersHumanJurkat CellsKineticsLymphocyte ActivationMembrane Glycoproteins/biosynthesis/physiologyRecombinant Proteins/biosynthesisSupport, Non-U.S. Gov'tT-LymphocytesTetradecanoylphorbol Acetatehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8898087</URL></MDL></Cite></EndNote>[76]CD4+/CD8+ (limfociti T)inhibira apoptozu i modulira adheziju stanica na izvanstani ni matriks  ADDIN EN.CITE <EndNote><Cite><Author>Hughes</Author><Year>2001</Year><RecNum>12</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>115223968372001JulGalectins as modulators of cell adhesion667-76National Institute for Medical Research, Mill Hill, London NW7 1AA, UK. chughes@nimr.mrc.ac.ukHughes, R. 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T.BiochemistryAnimalsAntigens, Differentiation/*metabolism/pharmacologyBinding SitesCells, CulturedCross-Linking ReagentsGalectin 3Lectins/*metabolismLeukemia, Basophilic, AcuteMast Cells/*drug effects/immunologyMembrane Glycoproteins/*metabolismPeritoneal Cavity/cytologyRatsReceptors, IgE/*metabolismSerotonin/secretionSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8347574[81, 82]Lamp-1, Lamp-2 (tumorske stanice)meustani na adhezija i adhezija stanica na izvanstani ni matriks  ADDIN EN.CITE <EndNote><Cite><Author>Ohannesian</Author><Year>1995</Year><RecNum>68</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>7743523</ACCESSION_NUMBER><VOLUME>55</VOLUME>101995May 15Carcinoembryonic antigen and other glycoconjugates act as ligands for galectin-3 in human colon carcinoma cells2191-9Department of Tumor Biology, University of Texas M. D. 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K.Masinick, S.Garrett, M.Hazlett, L. D.Infect ImmunAntigens, Differentiation/immunology/*metabolismBlotting, WesternCells, CulturedCornea/*chemistry/immunologyEpithelium/chemistry/immunologyEye Proteins/immunology/*metabolismGalectin 3HumanLipopolysaccharides/immunology/isolation & purification/*metabolismPseudomonas aeruginosa/immunology/*metabolismSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9199445[84]* ozna ava unutarstani ne ligande 1.1.1.7 Galektin-3 u imunom odgovoru U imunim su procesima uloge galektin-3, bilo ovisne ili neovisne o njegovoj lektinskoj aktivnosti, brojne. Izmeu ostalog, on modulira apoptozu, stani nu adheziju i kemotaksiju imunih stanica. Jak je proinflamatorni signal  ADDIN EN.CITE Rabinovich2002200120723712362002JunGalectins and their ligands: amplifiers, silencers or tuners of the inflammatory response?313-20Division of Immunogenetics, Hospital de Clinicas Jose de San Martin, School of Medicine, University of Buenos Aires, Cordoba 2351, 3er Piso (1120), Buenos Aires, Argentina.Rabinovich, G. A.Baum, L. G.Tinari, N.Paganelli, R.Natoli, C.Liu, F. T.Iacobelli, S.Trends ImmunolAmino Sugars/metabolismAnimalsApoptosis/physiologyAutoimmune Diseases/immunologyCattleCell CommunicationCell SurvivalChagas Disease/immunologyExtracellular Matrix/metabolismGalectinsHemagglutinins/classification/*physiologyHomeostasisHumanInfection/immunologyInflammation/*metabolismLigandsMammals/immunology/metabolismMiceMice, Inbred DBAModels, MolecularNeoplasms/immunology/pathologyPolysaccharides/metabolismProtein ConformationProtein Isoforms/physiologyT-Lymphocyte Subsets/immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12072371Almkvist200442014758082197-92004Galectins as inflammatory mediators575-81The Phagocyte Research Laboratory, Department of Rheumatology and Inflammation Research, Goteborg University, Sweden.Almkvist, J.Karlsson, A.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758082[48, 85], te aktivira mastocite i bazofile, a modulira i aktivaciju limfocita T. Pregled uloge galektina-3 u imunom odgovoru prikazan je u tablici 1.5. Tablica 1.5 Funkcije galektina-3 u imunom odgovoru. FunkcijaMeustani na adhezija ili interakcije stanice i patogenaSudjeluje u adheziji neutrofila na laminin  ADDIN EN.CITE <EndNote><Cite><Author>Kuwabara</Author><Year>1996</Year><RecNum>57</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8621934156101996May 15Galectin-3 promotes adhesion of human neutrophils to laminin3939-44Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, CA 92037, USA.Kuwabara, I.Liu, F. T.J ImmunolAmino Acid Sequence/drug effectsAntibodies, Monoclonal/pharmacologyAntigens, Differentiation/chemistry/*pharmacologyBinding, Competitive/immunologyCalcium/metabolismCarbohydrate Conformation/drug effectsCell Adhesion/drug effects/immunologyFibronectins/metabolismGalectin 3HumanIntegrins/immunologyLaminin/chemistry/*drug effects/*metabolismMagnesium/metabolismMembrane Proteins/chemistry/pharmacologyNeutrophils/chemistry/*drug effects/*metabolismSupport, U.S. Gov't, P.H.S.Temperaturehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8621934[86] Posreduje adheziji neutrofila na vaskularne endotelne stanice  ADDIN EN.CITE Sato20024401182351416842002Feb 15Role of galectin-3 as an adhesion molecule for neutrophil extravasation during streptococcal pneumonia1813-22Glycobiology Laboratory, Research Center for Infectious Disease, Laval University Medical Center, Centre Hospitalier Universitaire de Quebec, Quebec, Canada. 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T.J ImmunolAnimalsAntigens, Differentiation/administration & dosage/metabolism/*physiologyCalcium/metabolismCell Migration InhibitionCell Movement/immunologyCells, CulturedChemotactic Factors/administration & dosage/metabolism/*physiologyChemotaxis, Leukocyte/immunologyComparative StudyDiffusion Chambers, CultureDose-Response Relationship, ImmunologicGalectin 3HumanInjections, IntradermalIntracellular Fluid/metabolismKineticsMacrophages/*immunologyMacrophages, Alveolar/immunologyMiceMice, Inbred BALB CMonocyte Chemoattractant Protein-1/physiologyMonocytes/*immunology/metabolism/pathologyPertussis ToxinProtein Structure, TertiaryReceptors, Chemokine/physiologySignal Transduction/immunologySkin/immunology/pathologySupport, U.S. Gov't, P.H.S.Virulence Factors, Bordetella/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10925302Dong199676088980873952-31996Oct 21Galectin-3 stimulates uptake of extracellular Ca2+ in human Jurkat T-cells165-9National Institute for Medical Research, London, UK.Dong, S.Hughes, R. 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R., Jr.Am J PatholAnimalsAntigens, Differentiation/*physiologyCalcimycin/pharmacologyCell DifferentiationCells, CulturedGalactosides/*metabolismGalectin 3HumanImmunohistochemistryInterferon Type II/pharmacologyLectins/metabolismLipopolysaccharides/pharmacologyMacrophages/cytology/*metabolismMicroscopy, ElectronMonocytes/cytology/drug effects/*metabolism/microbiologySuperoxides/metabolismSupport, U.S. Gov't, P.H.S.Toxoplasma/physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7573347Yamaoka1995800789722815471995Apr 1A human lectin, galectin-3 (epsilon bp/Mac-2), stimulates superoxide production by neutrophils3479-87Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, CA 92037.Yamaoka, A.Kuwabara, I.Frigeri, L. G.Liu, F. T.J ImmunolAntigens, Differentiation/*physiologyCarbohydrates/physiologyChromatography, AffinityElectrophoresis, Polyacrylamide GelGalectin 3HumanImmunoblottingIn VitroLectins/*physiologyNeutrophils/*metabolismProtein Binding/physiologyStructure-Activity RelationshipSuperoxides/*metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7897228Karlsson199893095584029191998May 1Galectin-3 activates the NADPH-oxidase in exudated but not peripheral blood neutrophils3430-8The Phagocyte Research Laboratory, the Department of Medical Microbiology and Immunology, University of Goteborg, Guldhedsgatan 10, S-413 46 Goteborg, Sweden.Karlsson, A.Follin, P.Leffler, H.Dahlgren, C.BloodAlkaline Phosphatase/metabolismAntigens, Differentiation/*physiologyCells, CulturedEnzyme ActivationExudates and Transudates/cytologyGalectin 3HumanMacrophage-1 Antigen/metabolismNADPH Oxidase/*metabolismNeutrophil ActivationNeutrophils/*enzymologyReceptors, Cell Surface/metabolismReceptors, Complement 3b/metabolismRespiratory Burst/drug effectsSignal TransductionSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9558402[79, 89, 90] Poveava stvaranje IL-1 u monocitima  ADDIN EN.CITE <EndNote><Cite><Author>Jeng</Author><Year>1994</Year><RecNum>46</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>7890309</ACCESSION_NUMBER><VOLUME>42</VOLUME><NUMBER>3</NUMBER><YEAR>1994</YEAR>OctAn endogenous lectin, galectin-3 (epsilon BP/Mac-2), potentiates IL-1 production by human monocytes113-6Department of Medical Research, Taichung Veterans General Hospital, Taiwan, ROC.Jeng, K. C.Frigeri, L. G.Liu, F. T.Immunol LettAntigens, Differentiation/metabolism/*physiologyGalectin 3HumanIn VitroInterleukin-1/*biosynthesisLectins/*physiologyMonocytes/*immunology/metabolismProtein Binding/physiologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7890309[91] Zapo inje degranulaciju mastocita  ADDIN EN.CITE <EndNote><Cite><Author>Frigeri</Author><Year>1993</Year><RecNum>83</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8347574</ACCESSION_NUMBER><VOLUME>32</VOLUME><NUMBER>30</NUMBER><YEAR>1993</YEAR>Aug 3Epsilon BP, a beta-galactoside-binding animal lectin, recognizes IgE receptor (Fc epsilon RI) and activates mast cells7644-9Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, California 92037.Frigeri, L. G.Zuberi, R. I.Liu, F. T.BiochemistryAnimalsAntigens, Differentiation/*metabolism/pharmacologyBinding SitesCells, CulturedCross-Linking ReagentsGalectin 3Lectins/*metabolismLeukemia, Basophilic, AcuteMast Cells/*drug effects/immunologyMembrane Glycoproteins/*metabolismPeritoneal Cavity/cytologyRatsReceptors, IgE/*metabolismSerotonin/secretionSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8347574[82] Smanjuje ekspresiju IL-5 u leukocitima  ADDIN EN.CITE Cortegano1998960964724716111998Jul 1Galectin-3 down-regulates IL-5 gene expression on different cell types385-9Immunology Department, Fundacion Jimenez Diaz, Madrid, Spain.Cortegano, I.del Pozo, V.Cardaba, B.de Andres, B.Gallardo, S.del Amo, A.Arrieta, I.Jurado, A.Palomino, P.Liu, F. T.Lahoz, C.J ImmunolAntigens, Differentiation/*pharmacologyCell LineDown-Regulation/genetics/*immunologyEosinophils/drug effects/immunology/*metabolismGalectin 3Gene Expression Regulation/drug effects/*immunologyHumanHypersensitivity/immunologyInterleukin-5/antagonists & inhibitors/biosynthesis/*geneticsLectins/*pharmacologyLeukocytes, Mononuclear/drug effects/immunology/*metabolismSupport, Non-U.S. Gov'tT-Lymphocytes/drug effects/immunology/*metabolismTranscription, Genetic/drug effects/immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9647247Cortegano2000790107045221032000MarInteraction between galectin-3 and FcgammaRII induces down-regulation of IL-5 gene: implication of the promoter sequence IL-5REIII237-42Immunology Department, Fundacion Jimenez Diaz, 28040 Madrid, SpainCortegano, I.Pozo, V.Cardaba, B.Arrieta, I.Gallardo, S.Rojo, M.Aceituno, E.Takai, T.Verbeek, S.Palomino, P.Liu, F. T.Lahoz, C.GlycobiologyAnimalsAntigens, Differentiation/metabolism/*physiologyDown-RegulationFlow CytometryGalectin 3*Gene Expression RegulationHumanIn VitroInterleukin-5/*genetics/metabolismMiceMice, Inbred C57BLOligonucleotides/metabolism*Promoter Regions (Genetics)RNA, Messenger/biosynthesis/metabolismReceptors, IgG/metabolism/*physiologyRecombinant Fusion Proteins/metabolismSupport, Non-U.S. Gov'tTranscription Factors/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10704522[78, 92] Kemoatraktant je za monocite i makrofage  ADDIN EN.CITE Sano20008901092530216542000Aug 15Human galectin-3 is a novel chemoattractant for monocytes and macrophages2156-64Division of Allergy, La Jolla Institute for Allergy and Immunology, San Diego, CA 92121, USA.Sano, H.Hsu, D. K.Yu, L.Apgar, J. R.Kuwabara, I.Yamanaka, T.Hirashima, M.Liu, F. T.J ImmunolAnimalsAntigens, Differentiation/administration & dosage/metabolism/*physiologyCalcium/metabolismCell Migration InhibitionCell Movement/immunologyCells, CulturedChemotactic Factors/administration & dosage/metabolism/*physiologyChemotaxis, Leukocyte/immunologyComparative StudyDiffusion Chambers, CultureDose-Response Relationship, ImmunologicGalectin 3HumanInjections, IntradermalIntracellular Fluid/metabolismKineticsMacrophages/*immunologyMacrophages, Alveolar/immunologyMiceMice, Inbred BALB CMonocyte Chemoattractant Protein-1/physiologyMonocytes/*immunology/metabolism/pathologyPertussis ToxinProtein Structure, TertiaryReceptors, Chemokine/physiologySignal Transduction/immunologySkin/immunology/pathologySupport, U.S. Gov't, P.H.S.Virulence Factors, Bordetella/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10925302[88] Kemoatraktant je za endotelne stanice  ADDIN EN.CITE Nangia-Makker20009901070240715632000MarGalectin-3 induces endothelial cell morphogenesis and angiogenesis899-909Department of Tumor Progression, Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Nangia-Makker, P.Honjo, Y.Sarvis, R.Akahani, S.Hogan, V.Pienta, K. J.Raz, A.Am J PatholAnimalsAntigens, Differentiation/metabolism/*pharmacologyCapillaries/cytologyChemotaxis/*drug effectsCulture Media, Conditioned/pharmacologyDose-Response Relationship, DrugEndothelium, Vascular/*cytology/drug effects/metabolismFemaleGalectin 3HumanImmunoenzyme TechniquesMembrane Glycoproteins/metabolism/*pharmacologyMiceMice, NudeNeovascularization, Pathologic/*chemically induced/pathologyRecombinant Proteins/metabolism/pharmacologySkin Neoplasms/*blood supply/pathologySupport, U.S. Gov't, P.H.S.TransfectionTumor Cells, CulturedUmbilical Veins/cytologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10702407[93] Poti e stvaranje kapilara i inducira angiogenezu  ADDIN EN.CITE <EndNote><Cite><Author>Nangia-Makker</Author><Year>2000</Year><RecNum>99</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10702407</ACCESSION_NUMBER><VOLUME>156</VOLUME><NUMBER>32000MarGalectin-3 induces endothelial cell morphogenesis and angiogenesis899-909Department of Tumor Progression, Karmanos Cancer Institute, Detroit, Michigan 48201, USA.Nangia-Makker, P.Honjo, Y.Sarvis, R.Akahani, S.Hogan, V.Pienta, K. J.Raz, A.Am J PatholAnimalsAntigens, Differentiation/metabolism/*pharmacologyCapillaries/cytologyChemotaxis/*drug effectsCulture Media, Conditioned/pharmacologyDose-Response Relationship, DrugEndothelium, Vascular/*cytology/drug effects/metabolismFemaleGalectin 3HumanImmunoenzyme TechniquesMembrane Glycoproteins/metabolism/*pharmacologyMiceMice, NudeNeovascularization, Pathologic/*chemically induced/pathologyRecombinant Proteins/metabolism/pharmacologySkin Neoplasms/*blood supply/pathologySupport, U.S. Gov't, P.H.S.TransfectionTumor Cells, CulturedUmbilical Veins/cytologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10702407[93] Inducira regrutaciju leukocita in vivo  ADDIN EN.CITE Sato200443014758083197-92004Seeing strangers or announcing "danger": galectin-3 in two models of innate immunity583-91Glycobiology Laboratory, Research Centre for Infectious Diseases Laval University, Medical Centre, Centre Hospitalier Universitaire de Quebec, Ste-Foy, Quebec, Canada. Sachiko.Sato@crchul.ulaval.caSato, S.Nieminen, J.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758083[94]Imunoregulacija putem povrainskih reaetki multivalentnih galektina-3 Poveava prag aktivacije limfocita T  ADDIN EN.CITE <EndNote><Cite><Author>Demetriou</Author><Year>2001</Year><RecNum>100</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>1121786440968212001Feb 8Negative regulation of T-cell activation and autoimmunity by Mgat5 N-glycosylation733-9Samuel Lunenfeld Research Institute, Mount Sinai Hospital, Toronto, Ontario, Canada.Demetriou, M.Granovsky, M.Quaggin, S.Dennis, J. W.NatureAnimalsAntigens, Differentiation/metabolism*AutoimmunityGalectin 3GlycosylationHumanHypersensitivity, Delayed*Lymphocyte ActivationMajor Histocompatibility ComplexMiceN-Acetylglucosaminyltransferases/deficiency/*metabolismPolysaccharides/metabolismReceptors, Antigen, T-Cell/metabolismSignal TransductionSupport, Non-U.S. Gov'tT-Lymphocytes/*immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11217864[95] Inhibira proliferaciju stanica kotane sri potaknutu GM-CSF  ADDIN EN.CITE Krugluger19971010924153419711997JunGalectin-3 inhibits granulocyte-macrophage colony-stimulating factor (GM-CSF)-driven rat bone marrow cell proliferation and GM-CSF-induced gene transcription97-109Institute of General and Experimental Pathology, Vienna, Austria.Krugluger, W.Frigeri, L. G.Lucas, T.Schmer, M.Forster, O.Liu, F. T.Boltz-Nitulescu, G.ImmunobiologyAnimalsAntigens, Differentiation/genetics/metabolism/*pharmacologyBone Marrow/drug effects/*metabolism*Bone Marrow CellsCell Division/drug effectsGalectin 3Gene Expression Regulation/*drug effectsGranulocyte-Macrophage Colony-Stimulating Factor/*physiologyPolymerase Chain ReactionProtein Binding/geneticsRatsRats, Inbred LewRecombinant Proteins/metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Transcription, Genetic/*drug effectshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9241534[96] 1.1.1.8 Galektin-3 u autoimunim bolestima Za aktivaciju limfocita T potrebno je nakupljanje barem minimalnog broja receptorima na limfocitima T (TCR, engl. T-cell receptors). Nedostatak enzima 1,6 N-acetilglukozaminiltransferaze V (Mgat5) onemoguava stvaranje oligosaharidnih struktura na TCR na koje se ina e ve~e galektin-3. Uloga je galektina-3 vezanog na TCR ograni avanje lateralne mobilnosti TCR. Ukoliko galektin-3 nije vezan na TCR, olakaava se nakupljanje TCR i samim tim aktivacija limfocita T. Miaevi kojima nedostaje Mgat5 (tj. Mgat5-/- miaevi) imaju autoimunu bolest bubrega, poja ani odgoeni tip hipersenzitivnosti te poveanu podlo~nost eksperimentalno izazvanom autoimunom encefalomijelitisu  ADDIN EN.CITE Dennis2002167012417426157332002Dec 19UDP-N-acetylglucosamine:alpha-6-D-mannoside beta1,6 N-acetylglucosaminyltransferase V (Mgat5) deficient mice414-22Samuel Lunenfeld Research Institute, Mount Sinai Hospital, Toronto, Ontario, Canada. Dennis@mshri.on.caDennis, J. W.Pawling, J.Cheung, P.Partridge, E.Demetriou, M.Biochim Biophys ActaAnimalsCell DivisionMiceMice, KnockoutN-Acetylglucosaminyltransferases/genetics/*physiologyPolysaccharides/metabolismReceptors, Antigen, T-Cell/metabolism</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12417426</URL></MDL></Cite></EndNote>[97]. Spre avanje vezanja galektina-3 na TCR postignuto pre-tretmanom limfocita T s laktozom stvara fenokopiju Mgat5-/- nakupljanja TCR. Ti podaci pokazuju da galektin-3 igra va~nu ulogu u ograni avanju regrutacije kompleksa TCR na mjestu prezentacije antigena te da smanjeno vezanje galektina-3 na TCR poveava podlo~nost autoimunim bolestima, kao ato je multipla skleroza  ADDIN EN.CITE <EndNote><Cite><Author>Demetriou</Author><Year>2001</Year><RecNum>100</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11217864</ACCESSION_NUMBER><VOLUME>40968212001Feb 8Negative regulation of T-cell activation and autoimmunity by Mgat5 N-glycosylation733-9Samuel Lunenfeld Research Institute, Mount Sinai Hospital, Toronto, Ontario, Canada.Demetriou, M.Granovsky, M.Quaggin, S.Dennis, J. W.NatureAnimalsAntigens, Differentiation/metabolism*AutoimmunityGalectin 3GlycosylationHumanHypersensitivity, Delayed*Lymphocyte ActivationMajor Histocompatibility ComplexMiceN-Acetylglucosaminyltransferases/deficiency/*metabolismPolysaccharides/metabolismReceptors, Antigen, T-Cell/metabolismSignal TransductionSupport, Non-U.S. Gov'tT-Lymphocytes/*immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11217864[95]. U pacijenata koji boluju od reumatoidnog artritisa utvrena je poveana koncentracija galektina-3 u serumu i sinovijalnoj tekuini  ADDIN EN.CITE <EndNote><Cite><Author>Ohshima</Author><Year>2003</Year><RecNum>169</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>14558084</ACCESSION_NUMBER>48102003OctGalectin 3 and its binding protein in rheumatoid arthritis2788-95Center of Experimental Rheumatology, University Hospital, Zurich, Switzerland.Ohshima, S.Kuchen, S.Seemayer, C. A.Kyburz, D.Hirt, A.Klinzing, S.Michel, B. A.Gay, R. E.Liu, F. T.Gay, S.Neidhart, M.Arthritis RheumArthritis, Rheumatoid/immunology/metabolism/*physiopathologyBiological MarkersC-Reactive Protein/metabolismCarrier Proteins/*blood/*geneticsCell Adhesion/immunologyExtracellular Matrix Proteins/metabolismFibroblasts/cytology/physiologyGalectin 3/*blood/*geneticsGene ExpressionGlycoproteins/metabolismHumanIn VitroInterleukin-1/metabolismOsteoarthritis/immunology/metabolism/physiopathologyRNA, Messenger/analysisSupport, Non-U.S. Gov'tSynovial Fluid/metabolismSynovial Membrane/cytology/immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=14558084</URL></MDL></Cite></EndNote>[98], koja korelira s poremeenom apoptozom limfocita T  ADDIN EN.CITE <EndNote><Cite><Author>Harjacek</Author><Year>2001500115086002882001AugExpression of galectins-1 and -3 correlates with defective mononuclear cell apoptosis in patients with juvenile idiopathic arthritis1914-22Department of Pathology, New England Medical Center, Boston, Massachusetts, USA. miroslav.harjacek@zg.tel.hrHarjacek, M.Diaz-Cano, S.De Miguel, M.Wolfe, H.Maldonado, C. A.Rabinovich, G. A.J RheumatolAntigens, Differentiation/analysis/*biosynthesisApoptosis/*immunologyArthritis, Juvenile Rheumatoid/*immunology/metabolism/pathologyCell Division/immunologyChildGalectin 1Galectin 3Hemagglutinins/analysis/*biosynthesisHumanImmunohistochemistryKi-67 Antigen/analysisLymphocytes/chemistry/*metabolism/pathologyProto-Oncogene Proteins c-bcl-2/analysis/biosynthesisSupport, Non-U.S. Gov'tSynovial Membrane/pathologySynovitis/immunology/metabolism/pathologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11508600[99]. Autoantitijela na galektin-3 ne postoje u ljudi koji boluju od reumatoidnog artritisa, ali je njihova prisutnost utvrena u serumu ljudi koji boluju od sistemskog lupusa eritematozusa, poliomiozitisa/dermatomiozitisa  ADDIN EN.CITE <EndNote><Cite><Author>Lim</Author><Year>2002</Year><RecNum>170</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>1208377729512002Jul 5Identification of autoantibodies associated with systemic lupus erythematosus119-24Cancer Genomics Division, In2Gen Company, 6th Floor, Cancer Research Center, Seoul National University College of Medicine, 28 Yongon-dong, Chongno-gu, Seoul 110-799, Republic of Korea.Lim, Y.Lee, D. Y.Lee, S.Park, S. Y.Kim, J.Cho, B.Lee, H.Kim, H. Y.Lee, E.Song, Y. W.Jeoung, D. I.Biochem Biophys Res CommunAdultAntigens, Differentiation/immunologyAutoantibodies/blood/genetics/*immunologyAutoantigens/immunologyAutoimmune Diseases/immunologyFemaleGalectin 3Gene LibraryHumanLupus Erythematosus, Systemic/diagnosis/*immunologyMalePoly(ADP-ribose) Polymerases/immunologyRNA, Messenger/biosynthesisRibonucleoprotein, U1 Small Nuclear/immunologySupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12083777[100] te Kronove bolesti  ADDIN EN.CITE Jensen-Jarolim2001510117200072152001SepAnti-Galectin-3 IgG autoantibodies in patients with Crohn's disease characterized by means of phage display peptide libraries348-56Department of Pathophysiology, University of Vienna, Austria. erika.jensen-jarolim@akh-wien.ac.atJensen-Jarolim, E.Neumann, C.Oberhuber, G.Gscheidlinger, R.Neuchrist, C.Reinisch, W.Zuberi, R. I.Penner, E.Liu, F. T.Boltz-Nitulescu, G.J Clin ImmunolAmino Acid SequenceAntibodies, Monoclonal/immunologyAntigens, Differentiation/*immunologyAutoantibodies/*blood/immunologyAutoantigens/*immunologyBinding, CompetitiveCell Extracts/immunologyCell LineCrohn Disease/*blood/immunologyEpitopes, B-Lymphocyte/genetics/immunologyGalectin 3HumanImmunoblottingImmunoglobulin G/blood/immunologyLectins/*immunologyMolecular Sequence DataPeptide LibraryPeptides/chemical synthesis/genetics/immunologySequence Analysis, DNASupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11720007[101]. 1.1.1.9 Galektin-3 u tumorima Prisutnost galektina-3 utvrena je na mnogim vrstama tumorskih stanica, neovisno o tome da li su tumori nastali spontano ili su izazvani virusima, zra enjem ili kemijskim agensima. To nu ulogu galektina-3 u tumorskim stanicama, kao ni u drugim bioloakim sustavima, nije mogue definirati, ali je poznato da galektin-3 sudjeluje u sljedeim procesima bitnim za progresiju i metastaziranje tumora: poti e adheziju stani ne linije tumora dojke Evsa-T na laminin, fibronektin, vitronektin  ADDIN EN.CITE Takenaka200452014758078197-92004Galectin-3 and metastasis543-9Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Wayne State University, Detroit, Michigan 48201, USA.Takenaka, Y.Fukumori, T.Raz, A.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758078[102] poti e adheziju stani ne linije tumora dojke BT549 na laminin, kolagen IV  ADDIN EN.CITE <EndNote><Cite><Author>Warfield</Author><Year>1997</Year><RecNum>171</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>9561029</ACCESSION_NUMBER><VOLUME>1721997Adhesion of human breast carcinoma to extracellular matrix proteins is modulated by galectin-3101-12Department of Biochemistry, Meharry Medical College, Nashville, Tenn. 37208, USA.Warfield, P. R.Makker, P. N.Raz, A.Ochieng, J.Invasion MetastasisAntigens, Differentiation/genetics/*physiologyBreast Neoplasms/*metabolism/pathology/physiopathologyCarcinoma/*metabolism/pathology/physiopathologyCell Adhesion/drug effectsCollagen/*physiologyFibronectins/metabolism/*physiologyGalectin 3HumanIntegrin alpha6beta1Integrins/metabolismLaminin/*physiologyNeoplasm Proteins/genetics/*physiologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.TransfectionTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9561029[103] inhibira adheziju stanica tumora dojke, prostate, melanoma, fibrosarkoma na laminin, kolagen IV, fibronektin  ADDIN EN.CITE Ochieng1998580961829024631998May 29Regulation of cellular adhesion to extracellular matrix proteins by galectin-3788-91Department of Biochemistry, Meharry Medical College, Nashville, Tennessee 37208, USA. ochien10@ccvax.mmc.eduOchieng, J.Leite-Browning, M. L.Warfield, P.Biochem Biophys Res CommunAnimalsAntigens, Differentiation/*pharmacologyCell Adhesion/*drug effectsCollagen/metabolismComparative StudyDose-Response Relationship, DrugExtracellular Matrix Proteins/*metabolismFemaleFibronectins/metabolismGalectin 3HumanIntegrin alpha1beta1Integrins/metabolismLaminin/metabolismMaleMiceProtein BindingSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9618290</URL></MDL></Cite></EndNote>[59] poveava ekspresiju integrina (61 u stani noj liniji BT549  ADDIN EN.CITE <EndNote><Cite><Author>Warfield</Author><Year>1997</Year><RecNum>171095610291721997Adhesion of human breast carcinoma to extracellular matrix proteins is modulated by galectin-3101-12Department of Biochemistry, Meharry Medical College, Nashville, Tenn. 37208, USA.Warfield, P. R.Makker, P. N.Raz, A.Ochieng, J.Invasion MetastasisAntigens, Differentiation/genetics/*physiologyBreast Neoplasms/*metabolism/pathology/physiopathologyCarcinoma/*metabolism/pathology/physiopathologyCell Adhesion/drug effectsCollagen/*physiologyFibronectins/metabolism/*physiologyGalectin 3HumanIntegrin alpha6beta1Integrins/metabolismLaminin/*physiologyNeoplasm Proteins/genetics/*physiologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.TransfectionTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9561029[103] poveava ekspresiju integrina (47 u stani noj liniji Evsa-T  ADDIN EN.CITE <EndNote><Cite><Author>Takenaka</Author><Year>2004</Year><RecNum>52</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>14758078</ACCESSION_NUMBER><VOLUME>197-92004Galectin-3 and metastasis543-9Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Wayne State University, Detroit, Michigan 48201, USA.Takenaka, Y.Fukumori, T.Raz, A.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758078</URL></MDL></Cite></EndNote>[102] smanjuje ekspresiju integrina 1 u stani noj liniji MDA-MB-231  ADDIN EN.CITE <EndNote><Cite><Author>Furtak</Author><Year>2001</Year><RecNum>173</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>1173512328942001Dec 14Galectin-3 mediates the endocytosis of beta-1 integrins by breast carcinoma cells845-50Department of Biochemistry, Meharry Medical College, 1005 D. B. Todd Boulevard, Nashville, Tennessee 37208, USA.Furtak, V.Hatcher, F.Ochieng, J.Biochem Biophys Res CommunAntigens, CD29/*physiologyAntigens, Differentiation/pharmacology/*physiologyBreast Neoplasms/immunology/pathology/*physiopathologyCaveolae/physiologyCell Adhesion/drug effects/physiologyCell Membrane/physiologyCell SizeEndocytosis/physiologyFemaleFlow CytometryFluorescent Antibody Technique, IndirectGalectin 3HumanMicrofilaments/drug effects/physiologySupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11735123[104] titi stanice stani ne linije BT549 od apoptoze  ADDIN EN.CITE <EndNote><Cite><Author>Kim</Author><Year>1999</Year><RecNum>174</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10463621</ACCESSION_NUMBER><VOLUME>59</VOLUME><NUMBER>16</NUMBER><YEAR>1999</YEAR>Aug 15Cell cycle arrest and inhibition of anoikis by galectin-3 in human breast epithelial cells4148-54Department of Pathology, Karmanos Cancer Institute, Wayne State University, School of Medicine, Detroit, Michigan 48201, USA. hrckim@med.wayne.eduKim, H. R.Lin, H. M.Biliran, H.Raz, A.Cancer ResAntigens, Differentiation/*pharmacologyApoptosis/*drug effectsBreast/*pathologyCell Adhesion/drug effectsCell Cycle/*drug effectsCell LineEpithelial Cells/*drug effects/pathologyFemaleGalectin 3HumanSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10463621Yoshii2002501172477727792002Mar 1Galectin-3 phosphorylation is required for its anti-apoptotic function and cell cycle arrest6852-7Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Wayne State University, 110 E. Warren Ave., Detroit, MI 48201, USA.Yoshii, T.Fukumori, T.Honjo, Y.Inohara, H.Kim, H. R.Raz, A.J Biol Chem*AnoikisAntigens, Differentiation/*metabolism*ApoptosisBinding SitesBlotting, Western*Cell CycleCell LineCell SurvivalCisplatin/pharmacologyCyclin A/metabolismCyclins/metabolismDNA/metabolismDNA, Complementary/metabolismDown-RegulationGalectin 3HumanLigandsMutagenesis, Site-DirectedMutationPhosphorylationPrecipitin TestsProtein BindingProtein Kinases/metabolismProtein Processing, Post-TranslationalProtein Structure, TertiaryProto-Oncogene Proteins c-bcl-2/metabolismSerine/chemistrySignal TransductionSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Time FactorsTransfectionTumor Cells, CulturedUp-Regulationhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11724777[14, 105] posreduje homotipskoj agregaciji izoliranih tumorskih stanica  ADDIN EN.CITE Glinsky20004701082512560102000May 15Effects of Thomsen-Friedenreich antigen-specific peptide P-30 on beta-galactoside-mediated homotypic aggregation and adhesion to the endothelium of MDA-MB-435 human breast carcinoma cells2584-8Cancer Research Center, University of Missouri, Columbia 65211, USA.Glinsky, V. V.Huflejt, M. E.Glinsky, G. V.Deutscher, S. L.Quinn, T. P.Cancer ResAmino Acid SequenceAntigens, Neoplasm/*metabolismAntigens, Tumor-Associated, Carbohydrate/*metabolismBreast Neoplasms/*pathologyCell Adhesion/*drug effectsCell Aggregation/drug effectsEndothelium/drug effects/metabolismFemaleGalactosides/*metabolismGalectinsHemagglutinins/metabolismHumanMolecular Sequence DataNeoplasm Metastasis/prevention & controlPeptides/*pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&amp;list_uids=10825125</URL></MDL></Cite></EndNote>[106] ve~e antigen T i posreduje adheziji tumorskih stanica na endotel  ADDIN EN.CITE <EndNote><Cite><Author>Inohara</Author><Year>1994</Year><RecNum>176</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>802458120131994Jun 30Identification of human melanoma cellular and secreted ligands for galectin-31366-75Cancer Metastasis Program, Michigan Cancer Foundation, Detroit 48201.Inohara, H.Raz, A.Biochem Biophys Res CommunAmino Acid Sequence*Antigens, CDAntigens, Differentiation/*metabolismCarbohydrate SequenceCarrier Proteins/metabolismGalectin 3Glycoproteins/chemistry/*metabolismHumanIn VitroLigandsMelanoma/*metabolismMembrane Glycoproteins/metabolismMolecular Sequence DataRecombinant ProteinsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8024581Glinsky20004701082512560102000May 15Effects of Thomsen-Friedenreich antigen-specific peptide P-30 on beta-galactoside-mediated homotypic aggregation and adhesion to the endothelium of MDA-MB-435 human breast carcinoma cells2584-8Cancer Research Center, University of Missouri, Columbia 65211, USA.Glinsky, V. V.Huflejt, M. E.Glinsky, G. V.Deutscher, S. L.Quinn, T. P.Cancer ResAmino Acid SequenceAntigens, Neoplasm/*metabolismAntigens, Tumor-Associated, Carbohydrate/*metabolismBreast Neoplasms/*pathologyCell Adhesion/*drug effectsCell Aggregation/drug effectsEndothelium/drug effects/metabolismFemaleGalactosides/*metabolismGalectinsHemagglutinins/metabolismHumanMolecular Sequence DataNeoplasm Metastasis/prevention & controlPeptides/*pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10825125[106, 107] stimulira neovaskularizaciju in vitro  ADDIN EN.CITE Takenaka200452014758078197-92004Galectin-3 and metastasis543-9Tumor Progression and Metastasis Program, Karmanos Cancer Institute, Wayne State University, Detroit, Michigan 48201, USA.Takenaka, Y.Fukumori, T.Raz, A.Glycoconj Jhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14758078[102] Poveana ekspresija galektina-3 utvrena je u stanicama karcinoma skvamoznih stanica glave i vrata  ADDIN EN.CITE <EndNote><Cite><Author>Gillenwater</Author><Year>1996</Year><RecNum>178</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8864733</ACCESSION_NUMBER>1851996Sep-OctExpression of galectins in head and neck squamous cell carcinoma422-32Department of Tumor Biology, University of Texas M. D. Anderson Cancer Center, Houston 77030, USA.Gillenwater, A.Xu, X. C.el-Naggar, A. K.Clayman, G. L.Lotan, R.Head NeckAntigens, Differentiation/*analysisCell Transformation, NeoplasticGalectin 1Galectin 3Head and Neck Neoplasms/*chemistry/pathologyHemagglutinins/*analysisHumanImmunoblotting/methodsImmunohistochemistry/methodsNeoplasms, Squamous Cell/*chemistry/pathologyPhenotypeSupport, Non-U.S. Gov&apos;t</KEYWORD><KEYWORD>Tumor Cells, Cultured</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8864733</URL></MDL></Cite></EndNote>[108], karcinoma ~eluca  ADDIN EN.CITE Lotan1994179081128835641994Feb 15Expression of a 31-kDa lactoside-binding lectin in normal human gastric mucosa and in primary and metastatic gastric carcinomas474-80Department of Tumor Biology, University of Texas M.D. Anderson Cancer Center, Houston 77030.Lotan, R.Ito, H.Yasui, W.Yokozaki, H.Lotan, D.Tahara, E.Int J CancerAntibodies, MonoclonalGalectin 4Gastric Mucosa/*chemistryHemagglutinins/*analysis/metabolismHumanImmunoblottingImmunoenzyme TechniquesImmunohistochemistryJapanLiver Neoplasms/chemistry/secondaryLung Neoplasms/chemistry/secondaryLymphatic MetastasisNeoplasm MetastasisStomach Neoplasms/*chemistrySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tumor Markers, Biological/*analysis/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8112883</URL></MDL></Cite></EndNote>[109], anaplasti nog velikostani nog limfoma  ADDIN EN.CITE <EndNote><Cite><Author>Konstantinov</Author><Year>1996</Year><RecNum>180</RecNum>0854621314811996JanGalectin-3, a beta-galactoside-binding animal lectin, is a marker of anaplastic large-cell lymphoma25-30Department of Molecular and Experimental Medicine, Scripps Research Institute, La Jolla, CA 92037, USA.Konstantinov, K. N.Robbins, B. A.Liu, F. T.Am J PatholAdultAgedAntigens, Differentiation/*analysisFemaleGalectin 3HumanLymphoma, Large-Cell, Ki-1/*chemistryMaleMiddle AgedSupport, U.S. Gov't, P.H.S.Tumor Markers, Biological/*analysishttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8546213</URL></MDL></Cite></EndNote>[110] te tumora sredianjeg ~iv anog sustava  ADDIN EN.CITE <EndNote><Cite><Author>Bresalier</Author><Year>1997</Year><RecNum>181092643628041997Aug 15Expression of the endogenous galactose-binding protein galectin-3 correlates with the malignant potential of tumors in the central nervous system776-87Cancer Research Laboratory, Henry Ford Health Sciences Center, Detroit, Michigan 48202, USA.Bresalier, R. S.Yan, P. S.Byrd, J. C.Lotan, R.Raz, A.CancerAntigens, Differentiation/*metabolismAstrocytoma/metabolism/pathologyBrain Neoplasms/metabolism/*pathology/secondaryGalactose/metabolismGalectin 3Glioblastoma/metabolism/pathologyHumanImmunohistochemistryNeoplasm Proteins/metabolismProtein BindingSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9264362[111] u odnosu na ekspresiju u zdravom tkivu. Povienu koli inu galektina-3 eksprimiraju i stanice malignih neoplazmi atitnja e u odnosu na zdravo tkivo, ali i u odnosu na benigne adenome atitnja e te se galektin-3 smatra markerom prema kojem je mogue razlikovati maligne od benignih neoplazmi atitnja e  ADDIN EN.CITE Xu19951820767719314731995SepDifferential expression of galectin-1 and galectin-3 in thyroid tumors. Potential diagnostic implications815-22Department of Tumor Biology, University of Texas M.D. Anderson Cancer Center, Houston 77030, USA.Xu, X. C.el-Naggar, A. K.Lotan, R.Am J PatholAdenoma/diagnosis/*metabolismAntigens, Differentiation/*metabolismCarcinoma/diagnosis/*metabolismDiagnosis, DifferentialGalectin 1Galectin 3Goiter/metabolismHemagglutinins/*metabolismHumanImmunoblottingImmunohistochemistryLectins/metabolismReference ValuesSupport, Non-U.S. Gov'tThyroid Gland/metabolismThyroid Neoplasms/*diagnosis/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7677193[112]. Takoer je ustanovljeno da metastatsko tkivo eksprimira vee koli ine galektina-3 od primarnih tumora  ADDIN EN.CITE <EndNote><Cite><Author>Schoeppner</Author><Year>1995</Year><RecNum>183</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>777393275121995Jun 15Expression of an endogenous galactose-binding lectin correlates with neoplastic progression in the colon2818-26Gastrointestinal Cancer Research Laboratory, Henry Ford Health Sciences Center, Detroit, Michigan 48202, USA.Schoeppner, H. L.Raz, A.Ho, S. B.Bresalier, R. S.CancerAdenoma/chemistry/pathologyAntigens, Differentiation/*analysisBlotting, WesternCarcinoma/chemistry/mortality/pathologyCell Transformation, NeoplasticColonic Neoplasms/*chemistry/mortality/*pathologyColorectal Neoplasms/chemistry/pathologyGalectin 3HumanImmunohistochemistryIntestinal Mucosa/chemistryLectins/*metabolismNeoplasm MetastasisNeoplasm StagingSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Survival RateTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7773932</URL></MDL></Cite></EndNote>[113]. Poveanje razine cirkulirajueg galektina-3 povezano je s prisutnoau udaljenih metastaza  ADDIN EN.CITE <EndNote><Cite><Author>Iurisci</Author><Year>2000</Year><RecNum>48</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>10778968642000AprConcentrations of galectin-3 in the sera of normal controls and cancer patients1389-93Department of Oncology and Neurosciences, University G. D'Annunzio Medical School, Chieti, Italy.Iurisci, I.Tinari, N.Natoli, C.Angelucci, D.Cianchetti, E.Iacobelli, S.Clin Cancer ResAntibodies, Monoclonal/metabolismAntigens, Differentiation/*blood/immunologyCarrier Proteins/metabolismCell LineGalectin 3Glycoproteins/metabolismHumanImmunoassayLigandsNeoplasms/*blood/pathologyProtein Binding</KEYWORD><KEYWORD>Support, Non-U.S. Gov&apos;t</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10778968</URL></MDL></Cite></EndNote>[114]. Negativna korelacija izmeu ekspresije galektina-3 i progresije tumora utvrena je meutim u stanicama karcinoma dojke  ADDIN EN.CITE <EndNote><Cite><Author>Castronovo</Author><Year>1996</Year><RecNum>184</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>8691344</ACCESSION_NUMBER>17911996MayDecreased expression of galectin-3 is associated with progression of human breast cancer43-8Laboratory of Pathology, National Cancer Institute, Bethesda, MD 20892, USA.Castronovo, V.Van Den Brule, F. A.Jackers, P.Clausse, N.Liu, F. T.Gillet, C.Sobel, M. E.J PatholAntibody SpecificityAntigens, Differentiation/immunology/*metabolismAntigens, Neoplasm/immunology/*metabolismBlotting, NorthernBlotting, WesternBreast/metabolismBreast Neoplasms/immunology/*metabolism/pathologyDisease Progression*Down-RegulationEpithelium/metabolismFemaleGalectin 3HumanImmunoenzyme TechniquesNeoplasm InvasivenessSupport, Non-U.S. Gov'tTumor Markers, Biological/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8691344[115], jajnika i prostate  ADDIN EN.CITE Pacis20001850108810214422000Jul 1Decreased galectin-3 expression in prostate cancer118-23University of Michigan Comprehensive Cancer Center, Division of Hematology/Oncology, Department of Internal Medicine, St. John's Hospital, Detroit, Michigan, USA.Pacis, R. A.Pilat, M. J.Pienta, K. J.Wojno, K.Raz, A.Hogan, V.Cooper, C. R.ProstateAntigens, Differentiation/analysis/*biosynthesis/geneticsBlotting, WesternCell LineGalectin 3*Gene Expression Regulation, NeoplasticHumanImmunohistochemistryMaleProstate/cytology/*pathologyProstatic Neoplasms/chemistry/genetics/*pathologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10881021[116], do im su podaci za stanice tumora kolona kontradiktorni  ADDIN EN.CITE <EndNote><Cite><Author>Schoeppner</Author><Year>1995</Year><RecNum>183</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>7773932</ACCESSION_NUMBER><VOLUME>75</VOLUME><NUMBER>121995Jun 15Expression of an endogenous galactose-binding lectin correlates with neoplastic progression in the colon2818-26Gastrointestinal Cancer Research Laboratory, Henry Ford Health Sciences Center, Detroit, Michigan 48202, USA.Schoeppner, H. L.Raz, A.Ho, S. B.Bresalier, R. S.CancerAdenoma/chemistry/pathologyAntigens, Differentiation/*analysisBlotting, WesternCarcinoma/chemistry/mortality/pathologyCell Transformation, NeoplasticColonic Neoplasms/*chemistry/mortality/*pathologyColorectal Neoplasms/chemistry/pathologyGalectin 3HumanImmunohistochemistryIntestinal Mucosa/chemistryLectins/*metabolismNeoplasm MetastasisNeoplasm StagingSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Survival RateTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7773932Lee199118601842179126121991DecCarbohydrate-binding protein 35 is the major cell-surface laminin-binding protein in colon carcinoma1498-502Laboratory of Cancer Biology, Deaconess Hospital, Harvard Medical School, Boston, Mass 02115.Lee, E. C.Woo, H. J.Korzelius, C. A.Steele, G. D., Jr.Mercurio, A. M.Arch SurgAntigens, Differentiation/*analysisCarcinoma/*chemistryCarrier Proteins/*analysisChromatography, AffinityColonic Neoplasms/*chemistryFlow CytometryGalectin 3Hemagglutinins/*analysisHumanImmunoblottingMembrane Proteins/*analysisNeoplasm Proteins/*analysisSupport, U.S. Gov't, P.H.S.Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1842179Lotz1993187076827049081993Apr 15Decreased expression of Mac-2 (carbohydrate binding protein 35) and loss of its nuclear localization are associated with the neoplastic progression of colon carcinoma3466-70Laboratory of Cancer Biology, Deaconess Hospital, Harvard Medical School, Boston, MA 02115.Lotz, M. M.Andrews, C. W., Jr.Korzelius, C. A.Lee, E. C.Steele, G. D., Jr.Clarke, A.Mercurio, A. M.Proc Natl Acad Sci U S AAdenocarcinoma/*metabolism/pathologyAmino Acid SequenceAntigens, Differentiation/biosynthesis/genetics/*metabolismBase SequenceBlotting, NorthernCell Nucleus/metabolism/*ultrastructureCell Transformation, NeoplasticCloning, MolecularColon/cytology/*metabolism/pathologyColonic Neoplasms/*metabolism/pathologyColonic Polyps/*metabolism/pathologyDNA/genetics/isolation & purificationDNA, Neoplasm/genetics/isolation & purificationGalectin 3HumanImmunohistochemistryIntestinal Mucosa/cytology/*metabolism/pathologyLectins/biosynthesis/genetics/*metabolismMolecular Sequence DataMolecular WeightOligodeoxyribonucleotidesPolymerase Chain Reaction/methodsRNA/genetics/isolation & purificationRNA, Messenger/genetics/isolation & purification/metabolismRNA, Neoplasm/genetics/isolation & purificationSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7682704[113, 117, 118]. 1.2 UPALA Upala je strogo reguliran fizioloaki proces koji je od klju ne va~nosti za odr~avanje homeostaze. Mogu ga izazvati razli iti signali kao ato su infekcija patogenima, fizi ka ozljeda ili UV zra enje. Simptomi su upale crvenilo, oticanje, vruica, bol i oateena funkcionalnost ato nastaje kao posljedica oateenja tkiva i perfuzije terminalnih kapilara usljed eksudacije plazme u meustani ni prostor zbog poveane permeabilnosti kapilara. Ujedno dolazi i do stimulacije receptora boli. Te reakcije ovise o otpuatanju medijatora kao ato su histamin, serotonin, prostaglandini i kinini. Nakon inicijalne reakcije upalni se produkti prenose s mjesta upale krvlju ili dolazi do okon anja upalne reakcije. Eksudacija plazme esto je praena emigracijom krvnih stanica, primjerice granulocita i monocita u meustani ni prostor kao i proliferacijom makrofaga i fibroblasta. Svi su ti dogaaju va~ni za odr~avanje homeostaze, ali ponekad mogu uzrokovati nastanak upalnih bolesti, primjerice kroni nog reumatoidnog artritisa. Upalni je odgovor praen nakupljanjem limfocita T, plazma stanica, monocita, makrofaga, neutrofila i trombocita, koji, kao i tkivne endotelne stanice i fibroblasti, izlu uju mnoatvo lipida, faktora rasta, citokina i enzima koji uzrokuju lokalno oateenje tkiva, a ponekad i proliferaciju i fibrozu. Leukociti se na mjestu ozljede nakupljaju privu eni citokinima, kemokinima i drugim medijatorima upale. Takoer dolazi do indukcije enzima kao ato su ciklooksigenaza-2 (COX-2) i duaik-oksid-sintaze iji produkti (prostaglandini i duaikov oksid) poti u lokalan protok krvi. Molekule koje poti u stvaranje citokina i drugih proteina upalnog odgovora su mikrobni produkti (primjerice, lipopolisaharid-LPS) i sami citokini (primjerice, IL-1, TNF-, IL-17 i IL-18). Te molekule, ve~ui se na razli ite receptore aktiviraju etiri osnovna unutarstani na signalna puta kojima se kontrolira ekspresija gena uklju enih u upalni odgovor na transkripcijskoj i post-transkripcijskoj razini  ADDIN EN.CITE <EndNote><Cite><Author>Saklatvala</Author>2003112014587285702003Control of the expression of inflammatory response genes95-106Kennedy Institute of Rheumatology Division, Imperial College London, I Aspenlea Rd, London W6 8LH, U.K. j.saklatvala@imperial.ac.ukSaklatvala, J.Dean, J.Clark, A.Biochem Soc SympAnimals*Gene Expression RegulationGlucocorticoids/therapeutic useInflammation/drug therapy/*geneticsInflammation Mediators/metabolismMatrix Metalloproteinases/geneticsRNA Processing, Post-TranscriptionalRNA, Messenger/*genetics/metabolismSignal TransductionSupport, Non-U.S. Gov'tTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14587285Tibbles199911301048720555101999Aug 15The stress-activated protein kinase pathways1230-54Division of Experimental Therapeutics, Ontario Cancer Institute, Toronto, Canada.Tibbles, L. A.Woodgett, J. R.Cell Mol Life SciAnimalsCa(2+)-Calmodulin Dependent Protein Kinase/*metabolismHuman*Mitogen-Activated Protein KinasesModels, ChemicalProtein Kinases/*metabolismSignal TransductionSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10487205[119, 120]. To su: signalni put protein kinaza kojim se aktivira transkripcijski faktor NF-B (engl. nuclear factor kappa B) signalni putevi protein kinaza aktiviranih mitogenima (engl. mitogen-activated protein kinase, MAPK): signalni put kinaze regulirane izvanstani nim signalima (engl. extracellular-signal-regulated kinase, ERK) signalni put c-Jun N-terminalne kinaze (engl. c-Jun-N-terminal kinase, JNK) odnosno protein kinaze aktivirane stresom (engl. stress-activated-protein-kinase, SAPK) signalni put kinaze p38 Svi ti putevi vjerojatno reguliraju proces transkripcije, a signalni put kinaze p38 ima i dodatnu va~nu ulogu u post-transkripcijskoj regulaciji jer utje e na stabilnost mRNA gena koji sudjeluju u upalnom odgovoru  ADDIN EN.CITE <EndNote><Cite><Author>Kracht</Author>20021140124534672032002Nov 7Transcriptional and post-transcriptional control of gene expression in inflammation91-106Institute of Phamacology, Medical School Hannover, Carl-Neuberg Strasse 1, D-30625, Hannover, Germany. Kracht.Michael@MH-Hannover.deKracht, M.Saklatvala, J.CytokineAnimals*Gene Expression RegulationHumanInflammation/*geneticsNF-kappa B/metabolismProtein Kinases/metabolismRNA Stability*Signal Transduction*Transcription, GeneticTranslation, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12453467Dean200411501524000616102004OctThe involvement of AU-rich element-binding proteins in p38 mitogen-activated protein kinase pathway-mediated mRNA stabilisation1113-21Kennedy Institute of Rheumatology Division, Faculty of Medicine, Imperial College London, 1 Aspenlea Rd, Hammersmith, London, W6 8LH, UK. jonathan.dean@imperial.ac.ukDean, J. L.Sully, G.Clark, A. R.Saklatvala, J.Cell Signalhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15240006Clark200313701282923454612003Jul 3Post-transcriptional regulation of gene expression by mitogen-activated protein kinase p3837-44Kennedy Institute of Rheumatology Division, Faculty of Medicine, Imperial College London, 1 Aspenlea Road, Hammersmith, London W6 8LH, UK. andy.clark@imperial.ac.ukClark, A. R.Dean, J. L.Saklatvala, J.FEBS LettAnimalsHumanInflammation/physiopathologyMAP Kinase Signaling SystemMitogen-Activated Protein Kinase Kinases/metabolismMitogen-Activated Protein Kinases/*metabolismModels, Biological*Protein Processing, Post-TranslationalSignal TransductionSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12829234[121-123]. Koji e se signalni putevi aktivirati, u kojoj mjeri i na koji na in, ovisi o tipu izvanstani nog poticaja, kao i vrsti i razvojnom stupnju stanice. Osim toga, valja uzeti u obzir da se signalni putevi isprepliu te stoga jedna molekula mo~e biti uklju ena u regulaciju viae signalnih puteva a razli iti signalni putevi u kona nici mogu regulirati ekspresiju i/ili aktivnost iste molekule. 1.2.1 Signalni putevi protein kinaza aktiviranih mitogenima (MAPK) Tri su osnovne skupine MAPK u stanicama sisavaca koje se aktiviraju u upalnom odgovoru: kinaza regulirana izvanstani nim signalima (ERK), c-Jun N-terminalna kinaza (JNK) odnosno protein kinaza aktivirana stresom (SAPK), te kinaza p38. Osnovna je zna ajka signalnih puteva protein kinaza aktiviranih mitogenima o uvan trokomponentni model aktivacije, a sastoji se od tri kinaze u slijedu koje se meusobno aktiviraju (Slika 1.4).  Slika 1.4 Shema trokomponentnog modela aktivacije MAPK. Kaskada MAPK sastoji se od tri kinaze u slijedu koje se meusobno aktiviraju. Pojedine MAPKKK su esto komponente viae signalnih puteva. MAPKK selektivno fosforiliraju pojedine MAPK, osim MKK4, koja osim JNK mo~e aktivirati i p38. MAPK mogu fosforilirati transkripcijske faktore ili kinaze koje potom fosforiliraju transkripcijske faktore (kratice: MKKK-kinaza kinaze protein kinaze aktivirane mitogenima, MKK- kinaza protein kinaze aktivirane mitogenima, MAPK protein kinaza aktivirana mitogenima)  ADDIN EN.CITE Yang20031160145973843202003Nov 27Transcriptional regulation by the MAP kinase signaling cascades3-21School of Biological Sciences, University of Manchester, 2.205 Stopford Building, Oxford Road, Manchester M13 9PT, UK.Yang, S. H.Sharrocks, A. D.Whitmarsh, A. J.GeneAnimalsDNA-Binding Proteins/genetics/metabolism*Gene Expression RegulationHumanMAP Kinase Signaling System/*physiologyNucleosomes/genetics/metabolismSupport, Non-U.S. Gov'tTranscription Factors/genetics/metabolismTranscription, Genetic/*geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14597384[124]. Prva kinaza u slijedu je kinaza kinaze MAPK (MAPKKK) koja se aktivira ili fosforilacijom kinazom kinaze kinaze MAPK (MAPKKKK) ili interakcijom s malim proteinima iz obitelji Ras ili Rho koji ve~u GTP. MAPKKK fosforilira i aktivira sljedeu kinazu u nizu, kinazu MAPK (MAPKK). MAPK su posljednje kinaze u nizu koje uglavnom fosforiliraju brojne transkripcijske faktore (TF), ali i mnoge druge supstrate, poput protein kinaza, fosfolipaza te proteina citoskeleta. Do danas je u stanicama sisavaca identificirano 14 MAPKKK, 7 MKK i 13 MAPK, a postoji i veliki broj njihovih izoformi nastalih alternativnim procesiranjem  ADDIN EN.CITE Chang200111701124203441068242001Mar 1Mammalian MAP kinase signalling cascades37-40Department of Pharmacology, University of California San Diego, La Jolla 92093-0636, USA.Chang, L.Karin, M.NatureAnimalsEnzyme Activation*MAP Kinase Signaling SystemMammalsMitogen-Activated Protein Kinase Kinases/metabolismMitogen-Activated Protein Kinases/metabolismSubstrate SpecificitySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11242034[125]. Fosforilacija transkripcijskih faktora (TF) protein kinazama aktiviranim mitogenima regulira njihovu aktivnost tako to djeluje na: unutarstani nu lokalizaciju TF  ADDIN EN.CITE <EndNote><Cite><Author>Hood</Author><Year>1999</Year><RecNum>118</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10209150</ACCESSION_NUMBER><VOLUME>11</VOLUME><NUMBER>2</NUMBER><YEAR>1999</YEAR>AprIn or out? Regulating nuclear transport241-7Harvard Medical School and Dana-Farber Cancer Institute, Smith 922, 44 Binney Street, Boston, MA 02115, USA. jhood@student.med.harvard.eduHood, J. K.Silver, P. A.Curr Opin Cell BiolBiological Transport/*physiologyCell CompartmentationCell Cycle/physiologyCell Nucleus/*metabolismEnergy MetabolismEukaryotic Cells/cytology/metabolism/ultrastructureFungal Proteins/physiologyGene Expression RegulationHumanModels, BiologicalNuclear Proteins/metabolismPhosphorylationProtein Processing, Post-Translational/physiologySaccharomyces cerevisiae/cytology/physiologySignal Transduction/physiologyStress/genetics/physiopathologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10209150[126] razinu ekspresije TF  ADDIN EN.CITE Gianni200211901211058821142002Jul 15Phosphorylation by p38MAPK and recruitment of SUG-1 are required for RA-induced RAR gamma degradation and transactivation3760-9Institut de Genetique et de Biologie Moleculaire et Cellulaire, CNRS/INSERM/ULP/College de France, BP 163, 67404 Illkirch cedex, France.Gianni, M.Bauer, A.Garattini, E.Chambon, P.Rochette-Egly, C.Embo JAnimalsCOS CellsCarrier Proteins/*metabolismCysteine Endopeptidases/metabolismLigandsMitogen-Activated Protein Kinases/*metabolismMultienzyme Complexes/metabolismPhosphorylationReceptors, Retinoic Acid/genetics/*metabolismSupport, Non-U.S. Gov'tTrans-Activation (Genetics)/*drug effectsTretinoin/*pharmacologyUbiquitin/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12110588Shen200112001150965521182001SepTranscriptional hyperactivity of human progesterone receptors is coupled to their ligand-dependent down-regulation by mitogen-activated protein kinase-dependent phosphorylation of serine 2946122-31Department of Medicine, The Molecular Biology Program, and The Colorado Cancer Center, University of Colorado Health Sciences Center, Denver, Colorado 80262, USA.Shen, T.Horwitz, K. B.Lange, C. A.Mol Cell BiolDown-RegulationFemaleHumanLigandsMAP Kinase Signaling System/physiologyPhosphorylationProtein-Serine-Threonine Kinases/*physiologyReceptors, Progesterone/*physiologySerine*Signal TransductionSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Trans-Activation (Genetics)Tumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11509655[127, 128] vezanje TF na DNA  ADDIN EN.CITE Stefanovsky2001121011741541852001NovAn immediate response of ribosomal transcription to growth factor stimulation in mammals is mediated by ERK phosphorylation of UBF1063-73Cancer Research Centre and Department of Medical Biology, Laval University, Hotel-Dieu de Quebec, 11 cote du Palais, G1R 2J6, Quebec, Canada.Stefanovsky, V. Y.Pelletier, G.Hannan, R.Gagnon-Kugler, T.Rothblum, L. I.Moss, T.Mol CellAnimalsDNA/metabolismDNA-Binding Proteins/genetics/*metabolismEnzyme ActivationEpidermal Growth Factor/*pharmacologyGenes, ReporterHumanMiceMitogen-Activated Protein Kinases/*metabolismMutationPhosphorylation*Pol1 Transcription Initiation Complex ProteinsProtein Structure, SecondaryRNA Polymerase I/metabolismRNA, Ribosomal/genetics/metabolismRecombinant Fusion Proteins/metabolismRibosomes/*geneticsSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Transcription Factors/genetics/*metabolismTranscription, Genetic/drug effects/*physiologyp42 MAP Kinase/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11741541[129] interakciju TF s regulatornim proteinima  ADDIN EN.CITE Li20031220125141342222003Jan 15MAP kinase phosphorylation-dependent activation of Elk-1 leads to activation of the co-activator p300281-91Department of Cell Biology and Neuroscience, University of California, Riverside, CA 92521, USA.Li, Q. J.Yang, S. H.Maeda, Y.Sladek, F. M.Sharrocks, A. D.Martins-Green, M.Embo JAcetyltransferases/metabolismAnimalsChick EmbryoDNA-Binding Proteins/genetics/metabolismEnzyme Inhibitors/metabolismFibroblasts/cytology/metabolismMitogen-Activated Protein Kinases/genetics/*metabolismModels, BiologicalNuclear Proteins/genetics/*metabolismPhosphorylationPromoter Regions (Genetics)Proto-Oncogene Proteins/genetics/*metabolismRecombinant Fusion Proteins/genetics/metabolismSaccharomyces cerevisiae Proteins/metabolismSupport, Non-U.S. Gov'tThrombin/metabolism*Trans-Activation (Genetics)Trans-Activators/genetics/*metabolismTranscription Factors/genetics/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12514134Zhao20031230126202281122003FebERK-dependent phosphorylation of the transcription initiation factor TIF-IA is required for RNA polymerase I transcription and cell growth405-13Division of Molecular Biology of the Cell II, German Cancer Research Center, D-69120, Heidelberg, Germany.Zhao, J.Yuan, X.Frodin, M.Grummt, I.Mol Cell3T3 CellsAmino Acid SequenceAnimalsCell DivisionCell LineHumanMAP Kinase Signaling SystemMiceMitogen-Activated Protein Kinases/*metabolismMitogens/pharmacologyMolecular Sequence DataPhosphorylationProtein Kinases/metabolismRNA Polymerase I/*metabolismRNA, Ribosomal/biosynthesisRecombinant Proteins/genetics/metabolismSupport, Non-U.S. Gov'tTranscription Factors/genetics/*metabolismTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12620228[130, 131]. Pojednostavljeni prikaz mogue regulacije aktivnosti TF signalnim putem MAPK prikazan je na slici 1.5. Signalnim putevima MAPK reguliraju se aktivnosti velikog broja transkripcijskih faktora, primjerice p53, STAT1 (engl. signal transducer and activator protein 1), STAT3 (engl. signal transducer and activator protein 3), C/EBP (engl. CCAAT-enhancer-binding protein ), Sp-1, CREB (engl. cAMP-response-element binding protein), NFAT (engl. nuclear factor of activated T cells) i AP-1 (engl. activator protein 1). ini se da transkripciju najveeg broja gena ija se ekspresija mijenja u stanju upale, uz transkripcijski faktor NF-B, regulira upravo transkripcijski faktor AP-1  ADDIN EN.CITE <EndNote><Cite><Author>Karin</Author><Year>2000</Year><RecNum>128</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>10837071182000Phosphorylation meets ubiquitination: the control of NF-[kappa]B activity621-63Department of Pharmacology, Laboratory of Gene Regulation and Signal Transduction University of California, San Diego, La Jolla, California 92093-0636, USA. karinoffice@ucsd.eduKarin, M.Ben-Neriah, Y.Annu Rev ImmunolAmino Acid SequenceAnimalsDNA-Binding Proteins/*metabolismDown-RegulationHuman*I-kappa BMolecular Sequence DataNF-kappa B/antagonists & inhibitors/*metabolismPhosphorylationProtein-Serine-Threonine Kinases/metabolismSignal TransductionSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Transcription Factors/*metabolismUbiquitins/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10837071Yang20031160145973843202003Nov 27Transcriptional regulation by the MAP kinase signaling cascades3-21School of Biological Sciences, University of Manchester, 2.205 Stopford Building, Oxford Road, Manchester M13 9PT, UK.Yang, S. H.Sharrocks, A. D.Whitmarsh, A. J.GeneAnimalsDNA-Binding Proteins/genetics/metabolism*Gene Expression RegulationHumanMAP Kinase Signaling System/*physiologyNucleosomes/genetics/metabolismSupport, Non-U.S. Gov'tTranscription Factors/genetics/metabolismTranscription, Genetic/*geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14597384</MDL></Cite></EndNote>[124, 132].  Slika 1.5 Regulacija aktivnosti TF signalnim putem MAPK. MAPK mogu fosforilirati (P) transkripcijske faktore (TF) i regulirati njihov ulazak i izlazak iz jezgre. Jednom fosforilirani, TF esto mijenjaju konformaciju regulirajui na taj na in svoje funkcije, kao ato su vezanje DNA i vezanje koaktivatora i korepresora. Koaktivatori i korepresori su, kao i komponente kromatina, takoer podlo~ni regulaciji putem MAPK. U nekim slu ajevima, fosforilacija transkripcijskih faktora modulira njihovu stabilnost. Kratice: MAPK - protein kinaze aktivirane mitogenima, TF transkripcijski faktori, P fosforilacija, A koaktivatori, R korepresori  ADDIN EN.CITE Yang20031160145973843202003Nov 27Transcriptional regulation by the MAP kinase signaling cascades3-21School of Biological Sciences, University of Manchester, 2.205 Stopford Building, Oxford Road, Manchester M13 9PT, UK.Yang, S. H.Sharrocks, A. D.Whitmarsh, A. J.GeneAnimalsDNA-Binding Proteins/genetics/metabolism*Gene Expression RegulationHumanMAP Kinase Signaling System/*physiologyNucleosomes/genetics/metabolismSupport, Non-U.S. Gov'tTranscription Factors/genetics/metabolismTranscription, Genetic/*geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14597384[124]. 1.2.1.1 Transkripcijski faktor AP-1 Transkripcijski faktor AP-1, kodiran protoonkogenima, homodimer je ili heterodimer lanova obitelji proteina Jun (c-Jun, v-Jun, Jun-B, Jun-D), Fos (c-Fos, Fos-B, Fra-1, Fra-2), faktora aktivacije transkripcije (engl. activating transcription factor; ATF-2, ATF-3/LRF-1, B-ATF, JDP-1, JDP-2) ili Maf (v-Maf, c-Maf, Maf-A/B/F/G/K, Nrl). Svi oni pripadaju klasi basic zipper dimernih proteina koji prepoznaju specifi ne sljedove DNA i ve~u se na njih  ADDIN EN.CITE <EndNote><Cite><Author>De Bosscher</Author><Year>2003</Year><RecNum>133</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>129201522442003AugThe interplay between the glucocorticoid receptor and nuclear factor-kappaB or activator protein-1: molecular mechanisms for gene repression488-522Department of Molecular Biology, Ghent University, K. L. Ledeganckstraat 35, 9000 Gent, Belgium.De Bosscher, K.Vanden Berghe, W.Haegeman, G.Endocr RevGene Expression RegulationGlucocorticoids/immunology/metabolism/pharmacology/physiologyHumanInflammation/immunology/metabolismNF-kappa B/immunology/metabolism/*physiologyReceptors, Glucocorticoid/immunology/metabolism/*physiologySignal Transduction/immunologyTranscription Factor AP-1/immunology/metabolism/*physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12920152[133]. Geni fos i jun su geni ranog odgovora (engl. immediate early gene) ato zna i da njihova transkripcija zapo inje neposredno nakon stimulacije stanice  ADDIN EN.CITE <EndNote><Cite><Author>Wisdom</Author><Year>1999</Year><RecNum>189</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10579922</ACCESSION_NUMBER>25311999Nov 25AP-1: one switch for many signals180-5Department of Biochemistry, Vanderbilt University, Nashville, Tennessee, 37232, USA. ronald.m.wisdom@venderbilt.eduWisdom, R.Exp Cell ResMitogen-Activated Protein Kinases/metabolismMultigene FamilyProto-Oncogene Proteins/metabolism*Signal TransductionSupport, U.S. Gov't, P.H.S.Transcription Factor AP-1/genetics/*metabolismTranscription Factors/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10579922[134]. Proteini Jun mogu stvarati stabilne dimere koji prepoznaju sljedove DNA (5'-TGAG/CTCA-3') poznate pod nazivom TREs (engl. phorbol-12-O-tetradecanoate-13-acetate response elements); proteini Fos ne stvaraju stabilne dimere, ali mogu stvarati heterodimere s proteinima Jun koji su stabilniji od homodimera Jun/Jun. Proteini ATF stvaraju homodimere i heterodimere s proteinima Jun koji prepoznaju sljedove CRE (engl. cAMP responsive elements; 5'-TGACGTTCA-3'). U mnoatvo gena koji u svojoj promotorskoj regiji sadr~e sljedove koje prepoznaju pojedine kombinacije komponenti transkripcijskog faktora AP-1 ubrajamo i same gene c-jun i c-fos. Regulacija aktivnosti AP-1 je vrlo slo~ena i odvija se na nekoliko razina koje uklju uju regulacije: razine transkripcije gena jun i fos te stupnja razgradnje pripadnih mRNA (engl. mRNA turnover) stupnja razgradnje proteina Jun i Fos posttranskripcijskih modifikacija proteina Jun i Fos koje mijenjaju njihov transaktivacijski potencijal interakcija s drugim transkripcijskim faktorima koje mogu naruaiti ili poja ati aktivnost AP-1  ADDIN EN.CITE <EndNote><Cite><Author>Karin</Author><Year>1997</Year><RecNum>190</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>9069263921997AprAP-1 function and regulation240-6Department of Pharmacology, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0636, USA. palford@ucsd.eduKarin, M.Liu, Z.Zandi, E.Curr Opin Cell BiolAnimalsApoptosis/physiologyCa(2+)-Calmodulin Dependent Protein Kinase/metabolismGene Expression RegulationGenes, fosGenes, junHumanTranscription Factor AP-1/*physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9069263Karin199519107622446270281995Jul 14The regulation of AP-1 activity by mitogen-activated protein kinases16483-6Department of Pharmacology, University of California, San Diego, School of Medicine, La Jolla 92093, USA.Karin, M.J Biol ChemAnimalsCa(2+)-Calmodulin Dependent Protein Kinase/*physiologyHumanProtein Processing, Post-TranslationalSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Transcription Factor AP-1/*biosynthesisTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7622446Wisdom199918901057992225311999Nov 25AP-1: one switch for many signals180-5Department of Biochemistry, Vanderbilt University, Nashville, Tennessee, 37232, USA. ronald.m.wisdom@venderbilt.eduWisdom, R.Exp Cell ResMitogen-Activated Protein Kinases/metabolismMultigene FamilyProto-Oncogene Proteins/metabolism*Signal TransductionSupport, U.S. Gov't, P.H.S.Transcription Factor AP-1/genetics/*metabolismTranscription Factors/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&amp;dopt=Citation&amp;list_uids=10579922</URL></MDL></Cite></EndNote>[134-136]. Aktivnost AP-1 inducira se pod utjecajem velikog broja razli itih signala (kao ato su prisutnost faktora rasta, citokina, hormona, virusa, farmakoloakih tvari, teakih metala te pod djelovanjem UV- i - zra enja), koji u razli itoj mjeri aktiviraju pojedine kaskade MAPK (JNK, ERK i p38), a one pak poja avaju aktivnost AP-1 fosforiliranjem odreenih supstrata. Primjerice, transkripcijska aktivnost c-Jun se poja ava uslijed amino-terminalne fosforilacije Ser63 i Ser73 djelovanjem JNK  ADDIN EN.CITE Davis200019201105789710322000Oct 13Signal transduction by the JNK group of MAP kinases239-52Howard Hughes Medical Institute, Department of Biochemistry and Molecular Biology, University of Massachusetts Medical School, Worcester 01605, USA. roger.davis@umassmed.eduDavis, R. J.CellApoptosis*MAP Kinase Signaling SystemMitogen-Activated Protein Kinases/classification/*metabolismMorphogenesisNeoplasms/metabolismProtein-Tyrosine Kinase/metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11057897[137], kao i uz prisutnost fosforiliranog transkripcijskog kofaktora CBP (engl. CREB-binding protein)  ADDIN EN.CITE Bannister19951930854510711121995Dec 21Stimulation of c-Jun activity by CBP: c-Jun residues Ser63/73 are required for CBP induced stimulation in vivo and CBP binding in vitro2509-14Wellcome/CRC Institute, University of Cambridge, UK.Bannister, A. J.Oehler, T.Wilhelm, D.Angel, P.Kouzarides, T.OncogeneBinding SitesHumanPhosphorylationProto-Oncogene Proteins c-jun/*metabolismSupport, Non-U.S. Gov'tTrans-Activation (Genetics)Transcription Factors/metabolism/*pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=8545107[138]. c-Jun poti e stani nu proliferaciju smanjujui ekspresiju tumor-supresor gena i poti ui transkripciju gena koji kodira za ciklin D1, do im JunB djeluje upravo na suprotan na in. Zasada ostaje nerazjaanjeno kako c-Jun i JunB, koji prepoznaju gotovo identi ne sljedove DNA, mogu tako opre no djelovati  ADDIN EN.CITE <EndNote><Cite><Author>Shaulian</Author><Year>2002</Year><RecNum>194</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11988758</ACCESSION_NUMBER><VOLUME>4</VOLUME><NUMBER>5</NUMBER>2002MayAP-1 as a regulator of cell life and deathE131-6Department of Experimental Medicine and Cancer Research, School of Medicine, the Hebrew University, Jerusalem, 91120, Israel. eshaulian@md.huji.ac.ilShaulian, E.Karin, M.Nat Cell BiolAnimalsApoptosis/*physiologyCell Division/*physiologyCell Survival/physiology*Cell Transformation, NeoplasticGene Expression RegulationMAP Kinase Signaling SystemMitogen-Activated Protein Kinases/metabolismTranscription Factor AP-1/genetics/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11988758[139]. 1.2.1.2 Transkripcijski faktor NF-B Transkripcijski faktor NF-B (engl. nuclear factor kappa B) nalazi se u gotovo svim tipovima stanica i sudjeluje u regulaciji ekspresije viae od stotinu gena. NF-B je skupni naziv za inducibilne homo- ili heterodimerne transkripcijske faktore sastavljene od proteina obitelji Rel koji ve~u DNA. U sisavaca obitelj proteina Rel ini pet lanova: Rel (c-Rel), RelA (p65/NF-B3) RelB, NF-B1 (p50 i njegov prekursor p105) te NF-B2 (p52 i njegov prekursor p100). Svi oni sadr~e homolognu domenu Rel (engl. Rel homology domain) veli ine oko 300 aminokiselina, koja je odgovorna za njihovu dimerizaciju i vezanje DNA, a ujedno je i regulatorna domena. Unutar nje se nalazi signalni slijed za smjeatanje u jezgru (engl. nuclear localization sequence, NLS). U nestimuliranim stanicama dimeri NF-B nalaze se u citoplazmi povezani s inhibitornim proteinima klase IB. Obitelj proteina IB ine IB-(, IB-, IB-, IB-, IB- i Bcl-3, premda i p105 i p100 mogu djelovati kao IB. Aktivaciju NF-B mogu izazvati razli iti poticaji (primjerice citokini, bakterijski ili virusni produkti, UV i  zra enje) koji, do danas slabo razjaanjenim signalnim putevima, aktiviraju multiproteinski kompleks kinaza IB (IKK) uzrokujui fosforilaciju N-terminalnog dijela (Ser32 i Ser36) IB i njegovu disocijaciju. Signalni slijed za smjeatanje u jezgru NF-B stoga viae nije zaklonjen te se NF-B upuuje u jezgru, a fosforilirani se IB ubikvitinira i usmjerava do 26S proteasoma gdje se hidroliti ki razgrauje (Slika 1.6). Premda je disocijacija IB i translokacija NF-B u jezgru klju na za aktivaciju tog transkripcijskog faktora, njegova se transkripcijska aktivnost mo~e dodatno poja ati fosforilacijom i acetilacijom p65 podjedinice, kao i vezanjem razli itih kofaktora. ini se da u tom procesu, a mo~da i u samoj aktivaciji IKK, sudjeluju i neke od protein kinaza aktiviranih mitogenima  ADDIN EN.CITE <EndNote><Cite><Author>Karin</Author><Year>2004</Year><RecNum>125</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>14708018312004JanThe IKK NF-kappa B system: a treasure trove for drug development17-26Laboratory of Gene Regulation and Signal Transduction Department of Pharmacology, School of Medicine, University of California, San Diego, 9500 Gilman Drive, La Jolla, California 92093-0636, USA. KarinOffice@ucsd.eduKarin, M.Yamamoto, Y.Wang, Q. M.Nat Rev Drug DiscovAdjuvants, Immunologic/chemistry/pharmacologyAnimalsAnti-Inflammatory Agents/chemistry/pharmacology*Drug Design*Enzyme Inhibitors/chemistry/pharmacologyHumanMolecular StructureNF-kappa B/*antagonists & inhibitors/physiologyProtein-Serine-Threonine Kinases/*antagonists & inhibitors/physiologySignal Transductionhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14708018Rothwarf1999126011865184199951999Oct 26The NF-kappa B activation pathway: a paradigm in information transfer from membrane to nucleusRE1Laboratory of Gene Regulation and Signal Transduction, Department of Pharmacology, University of California, San Diego, CA 92093-0636, USA. drothwar@ucsd.eduRothwarf, D. M.Karin, M.Sci STKEAnimalsCell Membrane/*metabolismCell Nucleus/*metabolismHumanNF-kappa B/chemistry/*physiologySignal Transduction/*physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11865184Ruland20031270127526741932003JunTransducing signals from antigen receptors to nuclear factor kappaB93-100Advanced Medical Discovery Institute, Ontario Cancer Institute and University of Toronto, Toronto, ON, Canada.Ruland, J.Mak, T. W.Immunol RevAnimalsHumanLymphocyte Activation/*immunologyNF-kappa B/*immunologyReceptors, Antigen/*immunologySignal Transduction/*immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12752674[140-142].  Slika 1.6 Shematski prikaz aktivacije NF-B. Razli iti stimulansi, uklju ujui proinflamatorne citokine TNF( i IL-1, joa nerazjaanjenim signalnim putem aktiviraju IKK. IKK fosforilira IB-(, omoguavajui prepoznavanje IB pomou E3RS, komponente SCF tipa E3. Posljedi na poliubikvitinilacija IB-( usmjerava IB-( do 26S proteosoma gdje se on proteoliti ki razgrauje. Signalni slijed za smjeatanje u jezgru NF-B viae nije zaklonjen te se NF-B premjeata u jezgru. NF-B u jezgri poti e transkripciju razli itih gena, uklju ujui i IB-(, ime se ponovno inhibira aktivnost NF-B. NF-B poti e transkripciju gena koji kodiraju za medijatore upale, kao ato su TNF(, IL-1 i kemokini, ato pak dovodi do regrutacije dodatnih stanica na mjesto upale  ADDIN EN.CITE <EndNote><Cite><Author>Karin</Author><Year>2000128010837071182000Phosphorylation meets ubiquitination: the control of NF-[kappa]B activity621-63Department of Pharmacology, Laboratory of Gene Regulation and Signal Transduction University of California, San Diego, La Jolla, California 92093-0636, USA. karinoffice@ucsd.eduKarin, M.Ben-Neriah, Y.Annu Rev ImmunolAmino Acid SequenceAnimalsDNA-Binding Proteins/*metabolismDown-RegulationHuman*I-kappa BMolecular Sequence DataNF-kappa B/antagonists & inhibitors/*metabolismPhosphorylationProtein-Serine-Threonine Kinases/metabolismSignal TransductionSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Transcription Factors/*metabolismUbiquitins/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10837071[132].  Slika 1.7 Shematski prikaz signalnih puteva i mehanizama regulacije gena uklju enih u upalni odgovor. Citokini (IL-1, IL-17, IL-18, TNF) ili mikrobni produkti (LPS) poti u nakupljanje receptora i oligomerizaciju TRAF, ato je klju ni dogaaj za aktivaciju kinaze IB te signalnih puteva ERK, JNK i p38. Te kinaze fosforiliraju ili IB i NF-B ili transaktivacijske domene AP-1, aktivirajui na taj na in NF-B i AP-1. Za jaku transkripciju potrebna je, uz aktivaciju NF-B, aktivacija barem joa jednog signalnog puta MAPK. Ona uzrokuje acetilaciju i fosforilaciju histona, koju prati stvaranje viaeproteinskih kompleksa  poja iva a transkripcije. Unutar tih poja iva a NF-B i AP-1 intereagiraju s DNA, drugim transkripcijskim faktorima, koaktivatorima (npr. CBP/p300) i holoenzimom RNA polimerazom II. Posljedica varijacije u strukturi poja iva a je aktivacija specifi nih gena. Novosintetizirana mRNA je stabilizirana signalnim putem p38-MK-2. Inhibicija klju nih koraka prijenosa signala mo~e se postii primjenom glukokortikoida (G), inhibitora protein-kinaza (UO126, SP600125, SB203580), donora NO (SNAP; S-nitro-N-acetil-DL-penicilamin), salicilata i arsenita (kratice: AP-1-transkripcijski factor AP-1, ARE - elementi bogati slijedom AU, AUF1 - faktor vezanja AU 1, CREB - protein koji ve~e CBP, eIF - faktor inicijacije elongacije, ERK - kinaza regulirana izvanstani nim signalima, IB  inhibitor NF-B, IKK  kinaza IB, JNK  c-Jun N-terminalna kinaza, MAPK  protein kinaze aktivirane mitogenima, MEKK  ERK kinaza aktivirana mitogenima, MKK  kinaza MAPK, PABP  protein koji ve~e poliA-rep, P fosforilacija, TTP tristetraprolin, TAK1 protein kinaza 1 aktivirana faktorom rasta tumora, TNF faktor nekroze tumora, TRAF faktor vezanja receptora TNF)  ADDIN EN.CITE Kracht20021140124534672032002Nov 7Transcriptional and post-transcriptional control of gene expression in inflammation91-106Institute of Phamacology, Medical School Hannover, Carl-Neuberg Strasse 1, D-30625, Hannover, Germany. Kracht.Michael@MH-Hannover.deKracht, M.Saklatvala, J.CytokineAnimals*Gene Expression RegulationHumanInflammation/*geneticsNF-kappa B/metabolismProtein Kinases/metabolismRNA Stability*Signal Transduction*Transcription, GeneticTranslation, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12453467[121]. 1.2.2 Tvari s imunomodulatornim djelovanjem 1.2.2.1 Nesteroidne tvari s imunomodulatornim djelovanjem (NSAIDs) Nesteroidne tvari s imunomodulatornim djelovanjem (engl. non-steroidal antiinflammatory drugs, NSAIDs) su jedna od naj eae rabljenih skupina lijekova. Djeluju kao analgetici, antipiretici, antiinflamatorici te spre avaju agregaciju trombocita. Koriste se u terapiji razli itih bolesti (primjerice, astme, ateroskleroze, artritisa). Njihovo se djelovanje temelji ponajviae na sposobnosti inhibicije ciklooksigenaze (COX) drugim imenom prostaglandin H sintaze, klju nog enzima biosintetskog puta prostaglandina. COX je bifunkcionalan enzim katalizira konverziju arahidonske kiseline u prostaglandin G2 (PG2), a potom u prostaglandin H2 (PGH2). On je prete a bioloaki aktivnih prostaglandina - PGE2, PGI2, PGD2 i TXA2 (tromboksana), molekula koje su medijatori upale - djeluju kao vazodilatatori, a imaju i antiproliferativan u inak na limfocite T i B. COX postoji u obliku dviju izoformi-COX-1 i COX-2. COX-1 je konstitutivno eksprimiran u gotovo svim stanicama organizma i sudjeluje u fizioloakim funkcijama (citoprotekciji stanica ~eluca, agregaciji trombocita). COX-2 je proteinski produkt gena ranog odgovora, koji eksprimiraju stanice uklju ene u upalni odgovor (monociti, makrofagi, sinoviociti) i veinom posreduje sintezu prostanoida uklju enih u patoloake procese, kao ato su akutna i kroni na upala. Veina je negativnih popratnih u inaka NSAIDs (primjerice iritacija gastrointestinalnog sustava i krvarenje, disfunkcija trombocita) rezultat inhibicije COX-1, a antiinflamatorna, antipireti ka i analgeti ka djelovanja rezultat su inhibicije COX-2  ADDIN EN.CITE <EndNote><Cite>Hinz200214901180519330022002FebCyclooxygenase-2--10 years later367-75Department of Experimental and Clinical Pharmacology and Toxicology, Friedrich Alexander University Erlangen-Nurnberg, Erlangen, Germany. hinz@pharmakologie.uni-erlangen.deHinz, B.Brune, K.J Pharmacol Exp TherAnimalsCyclooxygenase Inhibitors/pharmacologyGene Expression Regulation, Enzymologic/geneticsHumanInflammation/enzymology/pathologyIsoenzymes/biosynthesis/genetics/*metabolismProstaglandin-Endoperoxide Synthase/biosynthesis/genetics/*metabolismSubstrate Specificity</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=11805193</URL></MDL></Cite></EndNote>[143]. Do danas nije to no utvreno mogu li NSAIDs inhibirati i samu ekspresiju, a ne samo aktivnost COX  ADDIN EN.CITE Barrios-Rodiles19961500878070822531996Aug 23Nonsteroidal antiinflammatory drugs inhibit cyclooxygenase-2 enzyme activity but not mRNA expression in human macrophages896-900Institute of Parasitology of McGill University, Quebec, Canada.Barrios-Rodiles, M.Keller, K.Belley, A.Chadee, K.Biochem Biophys Res CommunAnti-Inflammatory Agents, Non-Steroidal/*pharmacologyAspirin/pharmacologyCell LineCyclooxygenase Inhibitors/*pharmacologyGene Expression/drug effectsHumanIndomethacin/pharmacologyIsoenzymes/*genetics/*metabolismLipopolysaccharides/pharmacologyMacrophages/*drug effects/*enzymologyNaproxen/pharmacologyNitrobenzenes/pharmacologyProstaglandin-Endoperoxide Synthase/*genetics/*metabolismRNA, Messenger/*genetics/*metabolismSulfonamides/pharmacologySupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=8780708</URL></MDL></Cite></EndNote>[144]. U posljednje se vrijeme pojedini NSAIDs sve viae koriste za prevenciju kardiovaskularnih bolesti i mo~danog udara kao i za suportivno lije enje tumora (npr. tumora kolona)  ADDIN EN.CITE <EndNote><Cite><Author>Hull</Author><Year>2003</Year><RecNum>151</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>12927571</ACCESSION_NUMBER><VOLUME>29</VOLUME>42003AugActivity of the non-steroidal anti-inflammatory drug indomethacin against colorectal cancer309-20Molecular Medicine Unit, University of Leeds, Clinical Sciences Building, St. James's University Hospital, Leeds, LS9 7TF, UK. M.A.Hull@leeds.ac.ukHull, M. A.Gardner, S. H.Hawcroft, G.Cancer Treat RevAnimalsAnti-Inflammatory Agents, Non-Steroidal/*pharmacology/therapeutic useAntineoplastic Agents/*pharmacology/therapeutic useAntineoplastic Combined Chemotherapy Protocols/pharmacologyColorectal Neoplasms/*drug therapy/metabolism/*prevention & controlCyclooxygenase Inhibitors/pharmacologyHumanIn VitroIndomethacin/*pharmacology/therapeutic useReceptors, Cytoplasmic and Nuclear/drug effects/metabolismSupport, Non-U.S. Gov'tTranscription Factors/drug effects/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12927571Kim20021520124858699732002NovThe role of mitogen-activated protein kinases and their relationship with NF-kappaB and PPARgamma in indomethacin-Induced apoptosis of colon cancer cells241-5Department of Internal Medicine and Institute of Gastroenterology, Brain Korea 21 Project for Medical Science, College of Medicine, Yonsei University, Seoul, Korea.Kim, T. I.Jin, S. H.Kang, E. H.Shin, S. K.Choi, K. Y.Kim, W. H.Ann N Y Acad SciApoptosis/*drug effectsColonic Neoplasms/*pathologyHumanIndomethacin/*pharmacologyMAP Kinase Signaling System/drug effects/*physiologyMitogen-Activated Protein Kinases/metabolismNF-kappa B/*metabolismReceptors, Cytoplasmic and Nuclear/*metabolismSupport, Non-U.S. Gov'tTranscription Factors/*metabolismTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12485869[145, 146]. Sve je vie dokaza da antitumorsko, a djelomice i protuupalno djelovanje NSAIDs nije samo rezultat inhibicije COX, ve djelovanja NSAIDs na razli ite signalne puteve i transkripcijske faktore (Slika 1.8). Istra~ivanja na tom podru ju do danas su dala vrijedne, ali esto nepotpune i opre ne rezultate, ali se ini da modulacija signalnih puteva i TF ovisi kako o vrsti i primijenjenoj koncentraciji NSAIDs, tako i o tipu stanice, njezinom razvojnom stadiju i okru~enju u kojem se nalazi  ADDIN EN.CITE <EndNote><Cite><Author>Tegeder</Author><Year>200115401164123315122001OctCyclooxygenase-independent actions of cyclooxygenase inhibitors2057-72Pharmazentrum Frankfurt, Klinikum der Johann Wolfgang Goethe-Universitat, Frankfurt, 60590 Frankfurt am Main, Germany.Tegeder, I.Pfeilschifter, J.Geisslinger, G.Faseb JAnimalsAnti-Inflammatory Agents, Non-Steroidal/pharmacologyCyclooxygenase Inhibitors/*pharmacologyDNA-Binding Proteins/antagonists & inhibitorsGrowth Substances/biosynthesisHeat-Shock Response/drug effectsIsoenzymes/metabolismMAP Kinase Signaling System/drug effectsMiceModels, BiologicalNF-kappa B/antagonists & inhibitorsPregnancy Proteins/biosynthesisProstaglandin-Endoperoxide Synthase/metabolismReceptors, Cytoplasmic and Nuclear/antagonists & inhibitorsRibosomal Protein S6 Kinases/antagonists & inhibitorsSupport, Non-U.S. Gov'tTrans-Activators/antagonists & inhibitorsTranscription Factor AP-1/antagonists & inhibitorshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11641233[147].  Slika 1.8 U inak nesteroidnih tvari s imunomodulatornim djelovanjem na pojedine komponente signalnih puteva. Komponente signalnih puteva inhibirane djelovanjem NSAIDs ozna ene su sivim ovalom, a komponente aktivirane NSAIDs bijelim ovalom. (kratice: Akt/PKB  protein kinaza B, APC - adenomatous polyposis coli tumor supresor gen, Cdk  ciklin-ovisna kinaza, CREB - protein koji ve~e CBP, Erk - kinaza regulirana izvanstani nim signalima, GSK3b  kinaza 3  glikogen-sintaze, HSF  faktor toplinskog aoka, Hsp  proteini toplinskog aoka, IKK - kinaza IB, JNK  c-Jun N-terminalna kinaza, MAPK  protein kinaze aktivirane mitogenima, MAPKAP  protein kinaza aktivirana MAPK, MEK/MKK  kinaza protein kinaze aktivirane mitogenima, MLK  protein kinaza iz razli itih obitelji, MSK  protein kinaza aktivirana mitogenima i stresom, NIK  kinaza koja inducira NF-B, PAK  p21-aktivirana kinaza, PDK  kinaza ovisna o fosfoinozitolu, PI-3K  fosfoinozitol-3-kinaza, PKC  protein kinaza C, pRb  retinoblastoma protein, p70-S6K p70-S6 kinaza, p90RSK ribosomska S6 kinaza, Tcf faktor stanica T  ADDIN EN.CITE Tegeder200115401164123315122001OctCyclooxygenase-independent actions of cyclooxygenase inhibitors2057-72Pharmazentrum Frankfurt, Klinikum der Johann Wolfgang Goethe-Universitat, Frankfurt, 60590 Frankfurt am Main, Germany.Tegeder, I.Pfeilschifter, J.Geisslinger, G.Faseb JAnimalsAnti-Inflammatory Agents, Non-Steroidal/pharmacologyCyclooxygenase Inhibitors/*pharmacologyDNA-Binding Proteins/antagonists & inhibitorsGrowth Substances/biosynthesisHeat-Shock Response/drug effectsIsoenzymes/metabolismMAP Kinase Signaling System/drug effectsMiceModels, BiologicalNF-kappa B/antagonists & inhibitorsPregnancy Proteins/biosynthesisProstaglandin-Endoperoxide Synthase/metabolismReceptors, Cytoplasmic and Nuclear/antagonists & inhibitorsRibosomal Protein S6 Kinases/antagonists & inhibitorsSupport, Non-U.S. Gov'tTrans-Activators/antagonists & inhibitorsTranscription Factor AP-1/antagonists & inhibitorshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11641233[147]. Aspirin Aspirin (acetil-salicilna kiselina, ASA, kemijska formula C9H8O4, Mr= 180,16) ireverzibilno inaktivira COX-1 i COX-2 acetiliranjem hidroksilne skupine serina koji se nalazi u veznom mjestu enzima, onemoguavajui na taj na in vezanje arahidonske kiseline na COX  ADDIN EN.CITE <EndNote><Cite><Author>van der Ouderaa</Author><Year>1980</Year><RecNum>156</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE>677059061980Characterization of prostaglandin H2 synthetase139-44van der Ouderaa, F. J.Buytenhek, M.van Dorp, D. A.Adv Prostaglandin Thromboxane ResAcetylationAmino Acid SequenceAnimalsAspirinHemeHemeproteins/analysisIron/pharmacologyMaleMicrosomes/enzymologyMolecular Weight*Prostaglandin-Endoperoxide Synthase/metabolismProtein BindingSeminal Vesicles/enzymologySheepSpectrophotometryhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=6770590DeWitt19901550210816926591990Mar 25The aspirin and heme-binding sites of ovine and murine prostaglandin endoperoxide synthases5192-8Department of Biochemistry, Michigan State University, East Lansing 48824.DeWitt, D. L.el-Harith, E. A.Kraemer, S. A.Andrews, M. J.Yao, E. F.Armstrong, R. L.Smith, W. L.J Biol ChemAmino Acid SequenceAnimalsAspirin/*metabolismBase SequenceBinding SitesBlotting, WesternComparative StudyDNA/geneticsGenetic VectorsHeme/*metabolismMiceMolecular Sequence DataMutationOligonucleotide ProbesProstaglandin-Endoperoxide Synthase/*genetics/metabolismRestriction MappingSequence Homology, Nucleic AcidSerineSheepSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=2108169[148, 149].  Slika 1.9 Struktura aspirina Djelovanja aspirina koja su neovisna o inhibiciji COX uklju uju: modulaciju signalnog puta MAPK koja su rezultat: inhibicije aktivnosti Erk aktivacije p38 MAPK inhibicije ili u pojedinim slu ajevima aktivacije JNK inhibiciju NF-B koja je rezultat inhibicije fosforilacije i/ili proteosomske razgradnje IB. Aspirin se mo~e vezati na IKK spre avajui na taj na in vezanje ATP, ato inhibira kinaznu aktivnost IKK. Na sli an na in aspirin vjerojatno mo~e utjecati i na druge kinaze. inhibiciju AP-1  ADDIN EN.CITE <EndNote><Cite><Author>Dong</Author><Year>199715709092536272151997Apr 11Inhibition of activator protein 1 activity and neoplastic transformation by aspirin9962-70The Hormel Institute, University of Minnesota, Austin, Minnesota 55912, USA.zgdong@wolf.co.netDong, Z.Huang, C.Brown, R. E.Ma, W. Y.J Biol ChemAnimalsAntineoplastic Agents/*pharmacologyAntineoplastic Agents, Hormonal/pharmacologyAspirin/*pharmacologyCa(2+)-Calmodulin Dependent Protein Kinase/antagonists & inhibitorsCarcinogens/pharmacologyCell AdhesionCell Transformation, Neoplastic/*drug effectsCyclooxygenase Inhibitors/pharmacologyDiethylstilbestrol/pharmacologyGenes, junGenes, rasHydrogen-Ion ConcentrationIndomethacin/pharmacologyMice*Mitogen-Activated Protein KinasesPhosphorylationSodium Salicylate/pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tetradecanoylphorbol Acetate/pharmacologyTranscription Factor AP-1/*antagonists & inhibitorsTumor Cells, Culturedp42 MAP Kinasehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9092536[150] aktivaciju STAT1 koja je rezultat poveanja sposobnosti vezanja DNA  ADDIN EN.CITE <EndNote><Cite><Author>Chen</Author><Year>1999</Year><RecNum>158</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10477259</ACCESSION_NUMBER><VOLUME>342 Pt 31999Sep 15Salicylate-enhanced activation of transcription factors induced by interferon-gamma503-7Department of Molecular Genetics and Biochemistry, University of Pittsburgh School of Medicine, Pittsburgh, PA 15261, USA.Chen, L. C.Kepka-Lenhart, D.Wright, T. M.Morris, S. M., Jr.Biochem JAnimalsAspirin/pharmacologyCells, CulturedDNA-Binding Proteins/*metabolismElectrophoresis, Polyacrylamide GelInterferon Type II/*pharmacologyMicePhosphoproteins/*metabolismRNA, Messenger/metabolismSalicylates/*pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Trans-Activation (Genetics)/*drug effectsTrans-Activators/metabolismTranscription Factors/*geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10477259[151] Indometacin Indometacin (1-(4-klorobenzoil)-5-metoksi-2-metil-1-H-indol-3-octena kiselina, kemijska formula C19H16ClNO4 Mr=357,81) djeluje puno ja e od aspirina, a kako esto izaziva ozbiljne popratne pojave (hematoloaka i jetrena oateenja, poremeaje gastrointestinalnog sustava), rabi se rjee nego aspirin, i to naj eae u lije enju te~ih bolesti praenih jakom boli, kao ato je ankilozantni spondilitis i osteoartroza.  Slika 1.10 Struktura indometacina Indometacin uzrokuje sporu i vremenski ovisnu inhibiciju COX-1 i COX-2, koja je rezultat konformacijskih promjena do kojih dolazi ostvarivanjem ionskih interakcija karboksilne skupine indometacina i arginina polipeptidnog lanca COX. Za razliku od aspirina, indometacin vjerojatno ne inhibira niti NF-B niti AP-1 niti MAPK  ADDIN EN.CITE <EndNote><Cite><Author>Pillinger</Author><Year>1998</Year><RecNum>159</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>9826736</ACCESSION_NUMBER><VOLUME>95</VOLUME><NUMBER>24</NUMBER><YEAR>1998</YEAR>Nov 24Modes of action of aspirin-like drugs: salicylates inhibit erk activation and integrin-dependent neutrophil adhesion14540-5Department of Medicine, Room NB16N1, New York University Medical Center, 550 First Avenue, New York, NY 10016, USA. michael.pillinger@mccm.med.nyu.eduPillinger, M. H.Capodici, C.Rosenthal, P.Kheterpal, N.Hanft, S.Philips, M. R.Weissmann, G.Proc Natl Acad Sci U S AAcetaminophen/pharmacologyAntigens, CD/physiologyAntigens, CD18/physiologyArachidonic Acid/pharmacologyAspirin/*analogs & derivatives/*pharmacologyCa(2+)-Calmodulin Dependent Protein Kinase/*bloodCell Adhesion/drug effects/physiologyCell Aggregation/drug effects/physiologyCell Survival/drug effectsCyclooxygenase Inhibitors/pharmacologyHumanIn VitroMacrophage-1 Antigen/physiology*Mitogen-Activated Protein KinasesN-Formylmethionine Leucyl-Phenylalanine/pharmacologyNeutrophils/cytology/drug effects/*physiologySodium Salicylate/*pharmacologySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tetradecanoylphorbol Acetate/pharmacologyp42 MAP Kinasehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9826736[152]. Ve~e se i aktivira receptor  aktiviran proliferatorom peroksisoma (engl. peroxisome proliferator-activated receptor , PPAR), nuklearni hormonski receptor koji nakon vezanja hormona djeluje kao transkripcijski faktor  ADDIN EN.CITE <EndNote><Cite><Author>Lehmann19971600901358327261997Feb 7Peroxisome proliferator-activated receptors alpha and gamma are activated by indomethacin and other non-steroidal anti-inflammatory drugs3406-10Department of Molecular Endocrinology, Glaxo Wellcome Research and Development, Research Triangle Park, North Carolina 27709, USA.Lehmann, J. M.Lenhard, J. M.Oliver, B. B.Ringold, G. M.Kliewer, S. A.J Biol ChemAdipose Tissue/cytologyAnimalsAnti-Inflammatory Agents, Non-Steroidal/*pharmacologyCell Differentiation/drug effectsHumanIbuprofen/pharmacologyIndomethacin/*pharmacologyLiver/ultrastructureMiceMice, Inbred C3HMicrobodies/*metabolismNuclear Proteins/*metabolismReceptors, Cytoplasmic and Nuclear/*metabolismTranscription Factors/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9013583Shimada20041610147595125581-32004Jan 30PPARgamma mediates NSAIDs-induced upregulation of TFF2 expression in gastric epithelial cells33-8Department of Gastroenterology and Institute for Medical Science, Dokkyo University School of Medicine, Mibu, Tochigi 321-0293, Japan. tshimada@dokkyomed.ac.jpShimada, T.Koitabashi, A.Fujii, Y.Hashimoto, T.Hosaka, K.Tabei, K.Namatame, T.Yoneda, M.Hiraishi, H.Terano, A.FEBS LettAnilides/pharmacologyAnti-Inflammatory Agents, Non-Steroidal/*pharmacologyAspirin/pharmacologyCell Line, TumorDose-Response Relationship, DrugEnzyme Inhibitors/pharmacologyEpithelial Cells/drug effects/*metabolismGene Expression Regulation, NeoplasticGenes, ReporterHumanIndomethacin/pharmacologyPeptides/*metabolismRNA, Messenger/metabolismReceptors, Cytoplasmic and Nuclear/genetics/*metabolismReverse Transcriptase Polymerase Chain ReactionStomach Neoplasms/pathologyTrans-Activation (Genetics)/drug effectsTranscription Factors/genetics/*metabolismTranscription, Genetic/drug effectsUp-Regulation/*drug effectshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14759512[153, 154]. PPAR sudjeluje u diferencijaciji adipocita, ali i u modulaciji imunog odgovora suprimiranjem ekspresije proinflamatornih gena u makrofaga (IL-1, TNF-(, duaik-oksid sintaze). Njegovi fizioloaki ligandi nisu to no poznati, premda postoje odreene naznake da se na njega mogu vezati prostaglandini i nesteroidne tvari s imunomodulatornim djelovanjem. Vezanjem agonista PPARs heterodimerizira s retinoi nim receptorom X te se ve~e na svoja vezna mjesta u promotorskim regijama gena. Osim toga, takav mo~e transrepresijom inaktivirati druge transkripcijske faktore, kao ato je NF-B. 1.2.2.2 Steroidne tvari s imunomodulatornim djelovanjem-glukokortikoidi Glukokortikoidi (GC) su vrlo esto upotrebljavana skupina lijekova, koja se koristi u lije enju alergijskih i autoimunih bolesti, reumatoidnog artritisa, reumatske groznice, brojnih ko~nih bolesti, nekih oblika leukemija, a primjenjuju se i kao imunosupresivi nakon transplantacije organa. U cirkulaciji uzrokuju redistribuciju leukocita te inhibiraju funkciju leukocita i tkivnih makrofaga. Protuupalno i imunosupresivno djelovanje GC temelji se ponajviae na inhibiciji transkripcije citokina i kemokina (IL-1, IL-2, Il-3, IL-4, IL-5, IL-6, IL-8, IL-11, IL-12, IL-16, TNF-(, GM-CSF (engl. granulocyte-macrophage colony stimulating factor) IFN- i eotaksina) i citokinskih receptora  ADDIN EN.CITE <EndNote><Cite><Author>Adcock</Author><Year>2001</Year><RecNum>129</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11488985</ACCESSION_NUMBER><VOLUME>7942001AugCross-talk between pro-inflammatory transcription factors and glucocorticoids376-84Thoracic Medicine, National Heart and Lung Institute, Imperial College School of Science, Technology and Medicine, London, United Kingdom. ian.adcock@ic.ac.ukAdcock, I. M.Caramori, G.Immunol Cell BiolAcetylationAnti-Asthmatic Agents/pharmacology/therapeutic useAnti-Inflammatory Agents/pharmacology/therapeutic useAsthma/drug therapy/immunology/metabolism/*physiopathologyCytokines/metabolismGene Expression Regulation/*physiologyGlucocorticoids/*metabolism/pharmacology/therapeutic useHistones/metabolismHumanInflammation Mediators/physiologyModels, BiologicalReceptors, Glucocorticoid/metabolismSignal Transduction/physiologySteroidsTranscription Factors/genetics/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11488985[155]. Prirodni glukokortikoidi proizvode se u izvornom obliku ili kao acetati, a postoji i velik broj sintetskih glukokortikoida koji se donekle razlikuju od prirodnih hormona. Meusobno se razlikuju po trajanju u inka, intenzitetu djelovanja i lokalnoj aktivnosti. Glavni predstavnici sintetskih glukokortikoida su hidrokortizon i deksametazon. Hidrokortizon Hidrokortizon (drugim imenom kortizol, 17-hidroksikortikosteron, kemijska formula C21H30O5, Mr=362,5) je kristalini ni praaak teako topljiv u vodi. Nakon oralne aplikacije resorbira se u probavnom sustavu. U krvi ga je 95% vezano za 2-globulin, a ostatak od 5% je bioloaki aktivan.  Slika 1.11 Struktura hidrokortizona Deksametazon Deksametazon (9-fluoro-11,17,21-trihidroksi-16(-metilpregna-1,4-dien-3,20-dion, kemijska formula C22H29FO5, Mr=392,47) je bijeli kristalini ni prah teako topljiv u vodi. Ima 30 puta ja e protuupalno djelovanje od hidrokortizona. Za farmakoloaki u inak svih glukokortikoida bitna je prisutnost slobodne hidroksilne skupine na C-11 atomu, ali je takoer bitna i ja ina vezanja za receptore. Svi sintetski glukokortikoidi, derivati hidrokortizona, ja e se ve~u na glukokortikoidne receptore.  Slika 1.12 Struktura deksametazona Glukokortikoidi (GC) su relativno male lipofilne molekule koje difundiraju kroz lipidni dvosloj stani ne membrane i ve~u se na svoje citoplazmatske receptore. Glukokortikoidni receptori (GR) se u citoplazmi nalaze vezani na aaperone (naj eae Hsp 90 i Hsp 70) i ko-aaperonske molekule (poglavito imunofiline). Po vezanju GC na GR dolazi do hiperfosforilacije, koja poti e zamjenu jednog imunofilina (FKBP-51) drugim (FKBP-52) te vezanje transportnog proteina dineina na novonastali kompleks. On se upuuje u jezgru gdje dolazi do njegove disocijacije i konverzije GR u oblik koji mo~e vezati DNA  ADDIN EN.CITE <EndNote><Cite><Author>Saklatvala</Author><Year>2002130012010562432002Glucocorticoids: do we know how they work?146-50Kennedy Institute of Rheumatology Division, Faculty of Medicine, Imperial College of Science, Technology and Medicine, London, UK. j.saklatvala@ic.ac.ukSaklatvala, J.Arthritis ResAnti-Inflammatory Agents/metabolism/pharmacology/*therapeutic useDexamethasone/pharmacologyEnzyme Inhibitors/pharmacologyGlucocorticoids/metabolism/pharmacology/*therapeutic useHumanInflammation/*drug therapyMitogen-Activated Protein Kinases/antagonists & inhibitorsReceptors, Glucocorticoid/metabolismSignal TransductionSupport, Non-U.S. Gov'tTranscription Factors/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12010562Belgi20021310124641492772002OctTraditional therapies: glucocorticoids, azathioprine, methotrexate, hydroxyurea546-54Dermatopharmacology Unit, Southampton General Hospital, UK.Belgi, G.Friedmann, P. S.Clin Exp DermatolAzathioprine/pharmacology/*therapeutic useDermatologic Agents/pharmacology/*therapeutic useGlucocorticoids/pharmacology/*therapeutic useHumanHydroxyurea/pharmacology/*therapeutic useImmunosuppressive Agents/pharmacology/*therapeutic useMethotrexate/pharmacology/*therapeutic useSkin Diseases/drug therapyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12464149[156, 157]. Geni ija je ekspresija pozitivno regulirana djelovanjem GC u svojoj promotorskoj regiji sadr~e elemente odgovora na GC (engl. GC-response elements, GRE), na koje se aktivirani GR ve~e kao homodimer, a geni ija je ekspresija negativno regulirana djelovanjem GC ili u svojoj promotorskoj regiji sadr~e negativne elemente odgovora na GC (engl. negative GC-response elements, nGRE) ili je njihova transkripcija inhibirana izravnim ili posrednim djelovanjem GR na transkripcijsku aktivnost drugih transkripcijskih faktora (kao ato su NF-B, AP-1, CREB, C/EBP, STAT i p53)  ADDIN EN.CITE <EndNote><Cite><Author>Adcock</Author><Year>2001</Year><RecNum>132</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>11448148</ACCESSION_NUMBER><VOLUME>14</VOLUME><NUMBER>32001Glucocorticoid-regulated transcription factors211-9Thoracic Medicine, National Heart & Lung Institute, Imperial College of Science, Technology & Medicine, Dovehouse Street, London SW3 6LY, UK. ian.adcock@ic.ac.ukAdcock, I. M.Pulm Pharmacol TherAsthma/*drug therapy/*physiopathologyCell Differentiation/immunologyDrug ResistanceGene Expression RegulationGlucocorticoids/*pharmacologyHistone Deacetylases/*metabolismHumanInflammationLeukocytes/immunologyReceptors, Glucocorticoid/*physiologySignal TransductionT-LymphocytesTranscription Factors/*biosynthesis/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11448148Lewis1999134010419850341999AugNew targets for anti-inflammatory drugs489-94Signal Pharmaceuticals Inc., 5555 Oberlin Drive, San Diego, CA 92121, USA. alewis@signalpharm.comLewis, A. J.Manning, A. M.Curr Opin Chem BiolAnimalsAnti-Inflammatory Agents, Non-Steroidal/*pharmacologyCa(2+)-Calmodulin Dependent Protein Kinase/drug effectsCytokines/*drug effectsHumanNF-kappa B/drug effects/metabolismTranscription Factors/*drug effects</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10419850</URL></MDL></Cite></EndNote>[158, 159]. Protuupalno djelovanje GC nije mogue objasniti ekspresijom gena koji su pozitivno regulirani GC (kao ato se geni za tirozin- i alanin-aminotransferazu i fosfoenolpiruvat karboksikinazu), a kako geni koji se aktiviraju u upalnom odgovoru ne sadr~e nGRE, ini se da je za protuupalni u inak GC klju no djelovanje GR na transkripcijsku aktivnost TF  ADDIN EN.CITE Almawi20021350117813767112002JanMolecular mechanisms of glucocorticoid antiproliferative effects: antagonism of transcription factor activity by glucocorticoid receptor9-15Department of Medical Biochemistry, Arabian Gulf University, Manama, Bahrain. wassim@agu.edu.bhAlmawi, W. Y.Melemedjian, O. K.J Leukoc BiolAnimalsCell Division/drug effects/physiologyCell Nucleus/physiologyGlucocorticoids/*pharmacologyHumanImmunity/physiologyReceptors, Glucocorticoid/*physiologySignal Transduction/*physiologyTrans-Activation (Genetics)/physiologyTranscription Factors/physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11781376[160]. Na ini kojima se to ostvaruje su: % Indukcija sinteze IB GC u pojedinim tipovima stanica induciraju transkripciju gena za IB, inhibitora transkripcijskog faktora NF-B  ADDIN EN.CITE <EndNote><Cite><Author>Auphan</Author><Year>1995</Year><RecNum>138</RecNum>0756997627052341995Oct 13Immunosuppression by glucocorticoids: inhibition of NF-kappa B activity through induction of I kappa B synthesis286-90Department of Pharmacology, School of Medicine, University of California, San Diego, La Jolla 92093-0636, USA.Auphan, N.DiDonato, J. A.Rosette, C.Helmberg, A.Karin, M.ScienceAnimalsAnti-Inflammatory Agents/*pharmacologyCell LineCell Nucleus/metabolismCytoplasm/metabolismDNA-Binding Proteins/*biosynthesisDexamethasone/*pharmacologyHumanHybridomas*I-kappa B*ImmunosuppressionInterleukin-2/pharmacologyLymph Nodes/drug effects/metabolismMiceNF-kappa B/*antagonists & inhibitors/metabolismReceptors, Glucocorticoid/metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.T-Lymphocytes/drug effects/*metabolismTetradecanoylphorbol Acetate/pharmacologyTranscription Factor AP-1/metabolismTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7569976[161]. % Model proteinskih interakcija "jednostavan model" GR se mo~e vezati na razli ite transkripcijske faktore (NF-AT, NF-B i AP-1) stvarajui na taj na in kompleks koji se ne mo~e vezati na DNA. Taj je mehanizam djelovanja poznat pod nazivom transrepresija  ADDIN EN.CITE Newton19981390965239825411998May 15Effect of dexamethasone on interleukin-1beta-(IL-1beta)-induced nuclear factor-kappaB (NF-kappaB) and kappaB-dependent transcription in epithelial cells81-9Department of Thoracic Medicine, National Heart and Lung Institute, Imperial College School of Medicine, London, UK. robert.newton@ic.ac.ukNewton, R.Hart, L. A.Stevens, D. A.Bergmann, M.Donnelly, L. E.Adcock, I. M.Barnes, P. J.Eur J BiochemCell LineDNA-Binding Proteins/analysisDexamethasone/*pharmacologyGene Expression Regulation/drug effectsGlucocorticoids/pharmacologyGranulocyte-Macrophage Colony-Stimulating Factor/metabolismHumanInflammation/physiopathologyInterleukin-1/*pharmacologyIsoenzymes/metabolismLung/*physiologyNF-kappa B/*metabolismNitric-Oxide Synthase/metabolismNuclear Proteins/metabolismProstaglandin-Endoperoxide Synthase/metabolismRNA, Messenger/metabolismSupport, Non-U.S. Gov'tTetradecanoylphorbol Acetate/pharmacologyTranscription, Genetic/*drug effects/geneticsTransfection/geneticshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9652398Hart200014201061982416112000JanEffects of inhaled corticosteroid therapy on expression and DNA-binding activity of nuclear factor kappaB in asthma224-31Thoracic Medicine, National Heart and Lung Institute, Imperial College School of Medicine and Royal Brompton Hospital, London, United Kingdom.Hart, L.Lim, S.Adcock, I.Barnes, P. J.Chung, K. F.Am J Respir Crit Care MedAdministration, InhalationAdministration, TopicalAdultAndrostadienes/administration & dosage/*therapeutic useAnti-Inflammatory Agents/administration & dosage/*therapeutic useAsthma/*drug therapy/metabolism/pathologyBlotting, WesternBronchi/metabolism/pathologyBronchoalveolar Lavage Fluid/chemistry/cytologyCell CountCells, CulturedComparative StudyCross-Over StudiesDNA/genetics/*metabolismDexamethasone/pharmacologyDouble-Blind MethodEosinophils/cytology/metabolismEpithelial Cells/metabolism/pathologyFemaleGlucocorticoidsHumanInterleukin-1/metabolismMacrophages, Alveolar/cytology/metabolismMaleMembrane Glycoproteins/drug effects/metabolismNF-kappa B/genetics/*metabolismNerve Tissue Proteins/drug effects/metabolismReceptors, Cell Surface/drug effects/metabolismSupport, Non-U.S. Gov&apos;t</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10619824</URL></MDL></Cite></EndNote>[162, 163]. "mjeaoviti model" GR se mo~e vezati na transkripcijske faktore (NF-B i AP-1) koji su ve vezani na DNA i inhibirati njihovu transkripcijsku aktivnost  ADDIN EN.CITE <EndNote><Cite><Author>Nissen</Author><Year>2000</Year><RecNum>141</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>10995388</ACCESSION_NUMBER>14182000Sep 15The glucocorticoid receptor inhibits NFkappaB by interfering with serine-2 phosphorylation of the RNA polymerase II carboxy-terminal domain2314-29Departments of Cellular and Molecular Pharmacology, and Biochemistry and Biophysics, PIBS Biochemistry and Molecular Biology Program, University of California, San Francisco, San Francisco, California 94143-0450, USA.Nissen, R. M.Yamamoto, K. R.Genes DevCell LineHumanI-kappa B/genetics/metabolismIntercellular Adhesion Molecule-1/genetics/metabolismInterleukin-8/antagonists & inhibitors/genetics/metabolismNF-kappa B/*antagonists & inhibitors/metabolismPhosphorylationPrecipitin TestsPromoter Regions (Genetics)Protein BindingProtein Structure, TertiaryRNA Polymerase II/*metabolismReceptors, Glucocorticoid/*physiologyRepressor Proteins/*physiologyResponse ElementsSerine/*metabolismSupport, U.S. Gov't, P.H.S.Tumor Necrosis Factor/metabolismZinc/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10995388Hart200014201061982416112000JanEffects of inhaled corticosteroid therapy on expression and DNA-binding activity of nuclear factor kappaB in asthma224-31Thoracic Medicine, National Heart and Lung Institute, Imperial College School of Medicine and Royal Brompton Hospital, London, United Kingdom.Hart, L.Lim, S.Adcock, I.Barnes, P. J.Chung, K. F.Am J Respir Crit Care MedAdministration, InhalationAdministration, TopicalAdultAndrostadienes/administration & dosage/*therapeutic useAnti-Inflammatory Agents/administration & dosage/*therapeutic useAsthma/*drug therapy/metabolism/pathologyBlotting, WesternBronchi/metabolism/pathologyBronchoalveolar Lavage Fluid/chemistry/cytologyCell CountCells, CulturedComparative StudyCross-Over StudiesDNA/genetics/*metabolismDexamethasone/pharmacologyDouble-Blind MethodEosinophils/cytology/metabolismEpithelial Cells/metabolism/pathologyFemaleGlucocorticoidsHumanInterleukin-1/metabolismMacrophages, Alveolar/cytology/metabolismMaleMembrane Glycoproteins/drug effects/metabolismNF-kappa B/genetics/*metabolismNerve Tissue Proteins/drug effects/metabolismReceptors, Cell Surface/drug effects/metabolismSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10619824[163, 164]. "model transmodulacije" Ukoliko se GRE nalazi u blizini veznih mjesta za druge transkripcijske faktore (AP-1, NF-AT, ANF-1 ili ANF-2), vezanjem GR na GRE mo~e doi do stvaranja proteinskih interakcija s transkripcijskim faktorima koji su ve vezani na DNA. To pak onemoguava stvaranje funkcionalnog preinicijacijskog kompleksa a samim time i inicijaciju transkripcije  ADDIN EN.CITE <EndNote>Bailey20011430112370621532001MarThe role of AP-1 in glucocorticoid resistance in leukaemia391-7Department of Child Health, University of Newcastle upon Tyne, UK.Bailey, S.Hall, A. G.Pearson, A. D.Redfern, C. P.LeukemiaAdultBase SequenceBlotting, NorthernBlotting, WesternChild, PreschoolDNA Primers*Drug Resistance, NeoplasmFemaleGlucocorticoids/*pharmacologyHumanLeukemia, Lymphocytic, Acute/metabolism/*pathologyLeukemia, Lymphocytic, Chronic/metabolism/*pathologyProto-Oncogene Proteins c-fos/metabolismProto-Oncogene Proteins c-jun/metabolismSupport, Non-U.S. Gov'tTranscription Factor AP-1/*physiologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11237062Pearce199814409804760273461998Nov 13Glucocorticoid receptor transcriptional activity determined by spacing of receptor and nonreceptor DNA sites30081-5Department of Medicine, University of California at San Francisco, San Francisco, California 94143, USA.Pearce, D.Matsui, W.Miner, J. N.Yamamoto, K. R.J Biol ChemAnimalsBinding SitesCattleCell LineDNA/*metabolismDexamethasone/pharmacologyElectrophoresis, Polyacrylamide GelGlucocorticoids/pharmacologyMiceNucleic Acid ConformationProto-Oncogene Proteins c-fos/metabolismProto-Oncogene Proteins c-jun/metabolismReceptors, Glucocorticoid/*metabolismRepressor Proteins/metabolismStructure-Activity RelationshipSupport, U.S. Gov't, P.H.S.*Trans-Activation (Genetics)/drug effectsTranscription Factor AP-1/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9804760Herrlich200114501140234120192001Apr 30Cross-talk between glucocorticoid receptor and AP-12465-75Forschungszentrum Karlsruhe, Institute of Toxicology and Genetics, PO Box 3640, D-76021 Karlsruhe, Germany.Herrlich, P.OncogeneAnimalsAnti-Inflammatory Agents/metabolismDNA/metabolismDNA FootprintingHydrocortisone/physiologyMiceModels, Biological*Receptor Cross-TalkReceptors, Glucocorticoid/*metabolismSignal TransductionTrans-Activation (Genetics)Transcription Factor AP-1/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11402341[165-167]. d) "model kompeticije" Transkripcijska aktivnost GR, AP-1 i NF-B mo~e se potaknuti prisutnoau kofaktora kao ato su CBP, p300 ili histon-acetiltransferaze (engl. histone acetyltransferase, HAT), te stoga kompeticija GR i AP-1 ili NF-B za iste kofaktore mo~e uzrokovati promjenu transkripcijske aktivnosti tih TF  ADDIN EN.CITE McKay20001460109355461482000AugCBP (CREB binding protein) integrates NF-kappaB (nuclear factor-kappaB) and glucocorticoid receptor physical interactions and antagonism1222-34Molecular Endocrinology Group, Laboratory of Signal Transduction, National Institute of Environmental Health Sciences, National Institutes of Health, Research Triangle Park, North Carolina 27709-2233, USA.McKay, L. I.Cidlowski, J. A.Mol EndocrinolAnimalsCell LineDexamethasone/metabolismGlucocorticoids/metabolism/pharmacologyHormone Antagonists/metabolism/pharmacologyMifepristone/metabolism/pharmacologyNF-kappa B/genetics/immunology/*metabolismNuclear Proteins/genetics/*metabolismReceptors, Glucocorticoid/drug effects/genetics/*metabolismRecombinant Proteins/genetics/metabolismTrans-Activation (Genetics)Trans-Activators/genetics/*metabolismTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=10935546</URL></MDL></Cite></EndNote>[168]. % Indukcija gena reguliranog glukokortikoidima (GILZ) GC poti u ekspresiju gena GILZ (engl. GC-induced leucine zipper). GILZ je transkripcijski faktor koji mo~e inhibirati translokaciju NF-B i AP-1 stvarajui interakcije s pojedinim komponentama tih TF. GILZ mo~e takoer inhibirati signalni put kinaze regulirane izvanstani nim signalima (ERK) ve~ui se na jednu od MAPKKK, preciznije-Raf  ADDIN EN.CITE <EndNote><Cite>Ayroldi20011470114681759832001Aug 1Modulation of T-cell activation by the glucocorticoid-induced leucine zipper factor via inhibition of nuclear factor kappaB743-53Department of Clinical and Experimental Medicine, Section of Pharmacology, University of Perugia, Via del Giochetto, 06100 Perugia, Italy.Ayroldi, E.Migliorati, G.Bruscoli, S.Marchetti, C.Zollo, O.Cannarile, L.D'Adamio, F.Riccardi, C.BloodActive Transport, Cell Nucleus/drug effectsApoptosis/drug effectsDexamethasone/pharmacologyGene Expression Regulation/drug effectsGlucocorticoids/chemistry/immunology/*pharmacologyHumanInterleukin-2/metabolismLeucine Zippers/*drug effects/immunologyLymphocyte Activation/*drug effectsNF-kappa B/*antagonists & inhibitors/metabolismReceptors, Antigen, T-Cell/antagonists & inhibitors/immunologyReceptors, Interleukin-2/metabolismSupport, Non-U.S. Gov'tT-Lymphocytes/cytology/drug effects/*immunologyTranscription Factors/immunology/*pharmacologyTumor Cells, Culturedhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11468175[169]. % Indukcija gena za MKP-1 GC induciraju ekspresiju fosfataze MAPK-1 (MKP-1, engl. MAPK phosphatase 1). Defosforilacijom MAPK uzrokovanom djelovanjem MKP-1 inaktiviraju se signalni putevi JNK i p38  ADDIN EN.CITE <EndNote><Cite><Author>Franklin</Author><Year>199714809202001272271997Jul 4Conditional expression of the mitogen-activated protein kinase (MAPK) phosphatase MKP-1 preferentially inhibits p38 MAPK and stress-activated protein kinase in U937 cells16917-23Department of Medicine, Division of Medical Oncology, University of Colorado Health Sciences Center, Denver, Colorado 80262, USA. chris.franklin@uchsc.eduFranklin, C. C.Kraft, A. S.J Biol ChemCa(2+)-Calmodulin Dependent Protein Kinase/*antagonists &inhibitors/metabolismCells, CulturedEnzyme ActivationEnzyme Inhibitors/*pharmacologyHumanImmediate-Early Proteins/*pharmacology*Mitogen-Activated Protein KinasesProtein-Tyrosine-Phosphatase/*pharmacologySubstrate SpecificitySupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.Tetradecanoylphorbol Acetate/pharmacologyTranscription Factor AP-1/metabolismTumor Cells, Culturedp42 MAP Kinasehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=9202001</URL></MDL></Cite></EndNote>[170]. 1.3 CILJ ISTRA}IVANJA Osnovni je cilj ovog rada sustavno ispitati u inke imunomodulatornih tvari na ekspresiju LGALS3 i galektina-3 u nediferenciranim i diferenciranim stanicama monocitno-makrofagne loze. Kako bi se mogao ostvariti postavljeni cilj, potrebno je: izabrati prikladnu stani nu liniju monocitno-makrofagne loze koja u detektabilnoj mjeri eksprimira LGALS3 i galektin-3 optimirati metodu za odreivanje razine ekspresije LGALS3, koja uklju uje izolaciju intaktne RNA, reverznu transkripciju praenu lan anom reakcijom polimerazom te odreivanje koli ine nastalih produkata potvrditi specifi nost umna~anja lan anom reakcijom polimerazom sekvenciranjem produkata te reakcije izlo~iti stanice monocitne stani ne linije 72-satnom djelovanju slabe nesteroidne tvari s imunomodulatornim djelovanjem (aspirina) jake nesteroidne tvari s imunomodulatornim djelovanjem (indometacina) slabe steroidne tvari s imunomodulatornim djelovanjem (hidroksikortizona) jake steroidne tvari s imunomodulatornim djelovanjem (deksametazona) niskih, srednjih i visokih terapeutskih koncentracija. odrediti vijabilnost i metaboli ku aktivnost stanica izlo~enih djelovanju imunomodulatornih tvari, odnosno citotoksi nost primijenjenih koncentracija imunomodulatornih tvari diferencirati stanice monocitne stani ne linije u makrofage izlo~iti diferencirane stanice 72-satnom djelovanju slabe nesteroidne tvari s imunomodulatornim djelovanjem (aspirina) jake nesteroidne tvari s imunomodulatornim djelovanjem (indometacina) slabe steroidne tvari s imunomodulatornim djelovanjem (hidroksikortizona) jake steroidne tvari s imunomodulatornim djelovanjem (deksametazona) srednjih i visokih terapeutskih koncentracija prirediti stani ne homogenate kontrolnih stanica i stanica sakupljenih 1, 3, 5, 24, 48 i 72 sata nakon izlaganja navedenim tvarima odrediti koli inu galektina-3 u stani nim homogenatima imunoblot metodom uz primjenu specifi nih protutijela odrediti koli inu mRNA za galektin-3 u kontrolnim stanicama i stanicama sakupljenim 1, 3, 5, 24, 48 i 72 sata nakon izlaganja navedenim tvarima optimiranom metodom za odreivanje razine ekspresije LGALS3 MATERIJAL I METODE 2.1 MATERIJAL Materijal i kemikalije upotrijebljeni u ovom radu nabavljeni su od: Agilent Technologies (Palo Alto, CA, SAD) Amersham Biosciences (Cardiff, Velika Britanija) Applied Biosystems (Foster City, CA, SAD) Biochrom (Berlin, Njema ka) Fermentas (Vilnius, Litva) Fluka (St. Louis, MO, SAD) Invitrogen (Carlsbad, CA, SAD) Kemika (Zagreb, Hrvatska) Merck (Darmstadt, Njema ka) Millipore (Bedford, MA, SAD) Oncogene Research Products (Darmstadt, Njema ka) Pharmacia (Uppsala, `vedska) Promega (Madison, WI, SAD) Qiagen (Qiagen, Valencia, CA, SAD) Roche (Mannheim, Njema ka) Roth (Karlsruhe, Njema ka) Sigma (St. Louis, MO, SAD) 2.1.1 Standardne kemikalije agaroza Sigma albumin goveeg seruma Roth akrilamid/bisakrilamid mix 37,5:1 (30%) Sigma amfotericin Invitrogen amonijev persulfat Sigma aspirin Sigma bicinkonini na kiselina Sigma CuSO4 x 5H2O Sigma deksametazon Sigma dietil-pirokarbonat Sigma DMSO Sigma ECL( Western Blotting Detection Reagents Amersham Biosciences EDTA Roth etanol (96%) Kemika etidij bromid Sigma fetalni telei serum Invitrogen forbol-12-miristat-13-acetat Sigma GBX Developer/Replanisher Sigma GBX Fixer/Replanisher Sigma glicerol (redest.) Kemika glicin Sigma HCl (36%) Kemika hidrokortizon Sigma indometacin Fluka izopropanol Sigma ledena octena kiselina Kemika -merkaptoetanol Sigma metanol Kemika MuL V reverzna transkriptaza Invitrogen NaCl Roth NaOH Sigma PBS - Seromed(, Dulbeco, pH 7,4 Biochrom penicillin G Sigma Platinum Taq polimeraza Invitrogen plazmocin Invitrogen PMSF Sigma ribonukleazni inhibitor Invitrogen SDS Sigma streptomicin-sulfat Sigma Temed Sigma Template Suppression Reagent Applied Biosystems Tris (tris(hidroksimetil(aminometan) - Trizma baze( Sigma Triton X-100 Merck Tween-20 Roth 2.1.2 Boje bromfenol plavo Sigma tripansko modrilo Sigma ksilen-cijanol Sigma 2.1.3 Standardi obojeni standardi molekulskih masa-proteini Sigma standardi molekulskih masa-DNA Fermentas GeneScan-500 TAMRA standard Applied Biosystems 2.1.4 Specifi ne komercijalne smjese analiti kih reagencija -kompleti ABI PRISM( BigDye( Terminator v3.0 Cycle Sequencing Ready Reaction Kit with AmpliTag(DNA Polymerase Applied Biosystems Bicinchoninic Acid Protein Assay Kit Sigma CellTiter 96(AQueous One Solution Reagent Promega DyeEx( 2.0 Spin Kit Qiagen RNA 6000 Nano LabChipKit Agilent Technologies RNeasy Mini Kit Qiagen QIAquick PCR Purification Kit Qiagen 2.1.5 Protutijela atakorsko monoklonsko protutijelo M3/38 razvijeno naspram miajeg galektina-3 koje hibridoma stanice TIB-166 izlu uju u medij (ATCC, SAD) miaje monoklonsko protutijelo razvijeno naspram kokoajeg aktina (Oncogene Research Products, Darmstadt, Njema ka) kozje poliklonsko protutijelo razvijeno naspram cijele molekule atakorskog imunoglobulina G obilje~eno peroksidazom iz hrena (Sigma, St. Louis, MO, SAD) kozje poliklonsko protutijelo razvijeno naspram cijele molekule miajeg imunoglobulina M obilje~eno peroksidazom iz hrena (Oncogene Research Products, Darmstadt, Njema ka) 2.1.6 Pribor filter papir Munktell( 1F Pharmacia Immobilon-PVDF membrane Millipore Lumi-film Chemiluminescent Detection Film Roche 2.1.7 Otopine i puferi Otopina aspirina aspirin 2 M u DMSO Otopina deksametazona deksametazon 0,5 mM u DMSO Otopina indometacina indometacin 5 mM u DMSO Otopina hidrokortizona hidrokortizon 5 mM u DMSO Otopina etidij bromida etidij bromid 5 x 10-5 % u dest. vodi Otopina PMSF PMSF 100 mM u izopropanolu Pufer za elektroforezu u SDS-PAG, pH 8,3 glicin 192,0 mM Tris HCl 25,0 mM SDS 3,5 mM u dest. vodi Pufer za homogenizaciju stanica, pH 8,0 Tris HCl 5 mM NaCl 15 mM PMSF 2 mM Triton X-100 1% u dest. vodi Pufer za nanoenje uzoraka na SDS-PAG, pH 6,8 Tris HCl 0,1875 M glicerol 20,0% -merkaptoetanol 5,0% SDS 6% bromfenol plavo 0,1% u dest. vodi 6 x pufer za nanoaenje uzoraka DNA, pH 6,8 glicerol 30,0% bromfenol plavo 0,25% ksilen-cijanol 0,25 % u dest. vodi Pufer za prijenos na membranu - anodni pufer I, pH 10,4 Tris HCl 0,3 M metanol 20% u dest. vodi Pufer za prijenos na membranu - anodni pufer II, pH 10,4 Tris HCl 0,025 M metanol 20% u dest. vodi Pufer za prijenos na membranu - katodni pufer, pH 9,4 Tris HCl 0,025 M glicin 0,04 M metanol 20% u dest. vodi TAE-pufer, pH 8,3 Tris acetat 40 mM EDTA 1 mM u dest. vodi TBS, pH 7,5 Tris HCl 50 mM NaCl 150 mM u dest. vodi TBST, pH 7,5 Tween-20 0,1% u TBS, pH 7,5 2.1.8 Gelovi za elektroforezu Gel za sabijanje 5% akrilamid/bisakrilamid mix 37,5:1 (30%) 5,00% Tris HCl, pH 6,8 0,25 M SDS 0,10% amonijev persulfat 0,10% Temed 0,05% u dest. vodi Gel za razdvajanje 12% akrilamid/bisakrilamid mix 37,5:1 (30%) 12,00% Tris HCl, pH 8,8 0,375 M SDS 0,10% amonijev persulfat 0,10% Temed 0,05% u dest. vodi 2.2 METODE 2.2.1 Stani ne kulture U pokusima su koriatene stanice stani ne linije ljudskih monocitnih stanica THP-1 te diferencirane stanice stani ne linije THP-1. Stanice su uzgajane u sterilnim uvjetima u plasti nim bocama/petrijevkama za stani ne kulture u termostatu pri 37(C u struji zraka s 5% CO2, relativne vla~nosti 95%. Kao hranidbeni medij koriaten je RPMI 1640 medij obogaen dodatkom 10% fetalnog teleeg seruma, 100 U/ml penicilina G, 100 g/ml streptomicin-sulfata, 250 ng/ml amfotericina te 2,5 g/ml plazmocina. Gustoa stani ne kulture nediferenciranih stanica THP-1 odr~avana je izmeu 1 x 105 i 8 x 105 stanica / ml kulture. Stanice su brojane uporabom hemocitometra. Diferencijacija stanica THP-1 postignuta je tretiranjem stanica THP-1 forbol-12-miristat-13-acetatom (PMA) na sljedei na in: Alikvoti stani ne kulture THP-1 koji sadr~e 4 x 106 stanica, odnosno 1 x 106 stanica centrifugirani su 10 min pri 600g. Talozi stanica resuspendirani su u 1 ml hranidbenog medija, te razrijeeni dodatkom otopine PMA u DMSO do kona ne koncentracije stanica 5 x 105 stanica/ml, koncentracije PMA 10-7 M, te koncentracije DMSO < 0,2%. Stanice su kultivirane u sterilnim uvjetima u plasti nim Petri zdjelicama za stani ne kulture (4 x 106 stanica u 100 x 20 mm Petri zdjelicama, odnosno 1 x 106 stanica u 60 x 15 mm Petri zdjelicama) tijekom 1, 3, 5, 24 ili 48 sati. Po isteku navedenih vremena odreen je postotak neadheriranih stanica u mediju brojanjem stanica pomou hemocitometra. Medij je potom odliven, a adherirane su stanice isprane hladnim PBS. Blagim struganjem silikonskim struga em podignuto je: ( 4 x 106 stanica iz 100 x 20 mm Petri zdjelica u 2 ml hladnog PBS. Nakon centrifugiranja 10 minuta pri 600g stanice su joa dva puta isprane PBS a potom pohranjene na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. ( 1 x 106 stanica iz 60 x 15 mm Petri zdjelica u 350 (l pufera RLT/1 % (-merkaptoetanol te pohranjeno na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.2 Tretiranje stanica THP-1 aspirinom Za potrebe pokusa alikvoti stani ne kulture koji sadr~e 3 x 107 stanica THP-1 prenijeti su u plasti ne boce za stani ne kulture (175 cm2, Nunclon) te je dodano: medija do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml. Te su stanice smatrane kontrolnim, netretiranim stanicama. medija i DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml te koncentracije DMSO 0,2%. Te su stanice smatrane kontrolnim stanicama tretiranim DMSO kako bi se isklju io mogui utjecaj DMSO. medija i 2 M otopine aspirina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije aspirina 0,1 mM te koncentracije DMSO < 0,2%. medija i 2 M otopine aspirina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije aspirina 0,5 mM te koncentracije DMSO < 0,2%. medija i 2 M otopine aspirina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije aspirina 1 mM te koncentracije DMSO < 0,2%. medija i 2 M otopine aspirina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije aspirina 2 mM te koncentracije DMSO < 0,2%. Iz svake pojedine, na opisani na in prireene kulture 1, 3, 5, 24, 48 i 72 sata nakon po etka tretmana odvajani su alikvoti stani ne suspenzije koji sadr~e 4 x 106 stanica te 1 x 106 stanica. Iz kontrolne, netretirane kulture alikvot stanica odvojen je i neposredno prije po etka pokusa, odnosno u vremenu O. Uzorci su centrifugirani 10 minuta pri 600g te su dobiveni talozi, koji sadr~e 4 x 106 stanica, isprani 3 puta hladnim PBS i pohranjeni na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. Talozima koji sadr~e 1 x 106 stanica dodano je 350 (l pufera RLT/1% (-merkaptoetanol. Pohranjeni su na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.3 Tretiranje stanica THP-1 indometacinom Za potrebe pokusa alikvoti stani ne kulture koji sadr~e 3 x 107 stanica prenijeti su u plasti ne boce za stani ne kulture (175 cm2, Nunclon) te je dodano: medija do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml. Te su stanice smatrane kontrolnim, netretiranim stanicama. medija i DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml te koncentracije DMSO 0,2%. Te su stanice smatrane kontrolnim stanicama tretiranim DMSO kako bi se isklju io mogui utjecaj DMSO. medija i 5 mM otopine indometacina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije indometacina 0,1 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine indometacina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije indometacina 1 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine indometacina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije indometacina 5 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine indometacina u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije indometacina 10 (M te koncentracije DMSO 0,2%. Iz svake pojedine, na opisani na in prireene kulture 1, 3, 5, 24, 48 i 72 sata nakon po etka tretmana odvajani su alikvoti stani ne suspenzije koji sadr~e 4 x 106 stanica te 1 x 106 stanica. Iz kontrolne, netretirane kulture alikvot stanica odvojen je i neposredno prije po etka pokusa, odnosno u vremenu O. Uzorci su centrifugirani 10 minuta pri 600g te su dobiveni talozi, koji sadr~e 4 x 106 stanica, isprani 3 puta hladnim PBS i pohranjeni na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. Talozima koji sadr~e 1 x 106 stanica dodano je 350 (l pufera RLT/1% (-merkaptoetanol. Pohranjeni su na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.4 Tretiranje stanica THP-1 hidrokortizonom Za potrebe pokusa alikvoti stani ne kulture koji sadr~e 3 x 107 stanica prenijeti su u plasti ne boce za stani ne kulture (175 cm2, Nunclon) te je dodano: medija do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml. Te su stanice smatrane kontrolnim, netretiranim stanicama. medija i DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml te koncentracije DMSO 0,2%. Te su stanice smatrane kontrolnim stanicama tretiranim DMSO kako bi se isklju io mogui utjecaj DMSO. medija i 5 mM otopine hidrokortizona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije hidrokortizona 0,1 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine hidrokortizona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije hidrokortizona 1 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine hidrokortizona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije hidrokortizona 5 (M te koncentracije DMSO < 0,2%. medija i 5 mM otopine hidrokortizona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije hidrokortizona 10 (M te koncentracije DMSO 0,2%. Iz svake pojedine, na opisani na in prireene kulture 1, 3, 5, 24, 48 i 72 sata nakon po etka tretmana odvajani su alikvoti stani ne suspenzije koji sadr~e 4 x 106 stanica te 1 x 106 stanica. Iz kontrolne, netretirane kulture alikvot stanica odvojen je i neposredno prije po etka pokusa, odnosno u vremenu O. Uzorci su centrifugirani 10 minuta pri 600g te su dobiveni talozi koji sadr~e 4 x 106 stanica isprani su 3 puta hladnim PBS i pohranjeni na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. Talozima koji sadr~e 1 x 106 stanica dodano je 350 (l pufera RLT/1% (-merkaptoetanol. Pohranjeni su na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.5 Tretiranje stanica THP-1 deksametazonom Za potrebe pokusa alikvoti stani ne kulture koji sadr~e 3 x 107 stanica prenijeti su u plasti ne boce za stani ne kulture (175 cm2, Nunclon) te je dodano: medija do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml. Te su stanice smatrane kontrolnim, netretiranim stanicama. medija i DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml te koncentracije DMSO 0,2%. Te su stanice smatrane kontrolnim stanicama tretiranim DMSO kako bi se isklju io mogui utjecaj DMSO. medija i 0,5 mM otopine deksametazona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije deksametazona 0,1 nM te koncentracije DMSO < 0,2%. medija i 0,5 mM otopine deksametazona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije deksametazona 1 nM te koncentracije DMSO < 0,2%. medija i 0,5 mM otopine deksametazona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije deksametazona 10 nM te koncentracije DMSO < 0,2%. medija i 0,5 mM otopine deksametazona u DMSO do kona nog volumena 150 ml, odnosno gustoe stanica 2 x 105 stanica/ml, koncentracije deksametazona 100 nM te koncentracije DMSO 0,2%. Iz svake pojedine, na opisani na in prireene kulture 1, 3, 5, 24, 48 i 72 sata nakon po etka tretmana odvajani su alikvoti stani ne suspenzije koji sadr~e 4 x 106 stanica te 1 x 106 stanica. Iz kontrolne, netretirane kulture alikvot stanica odvojen je i neposredno prije po etka pokusa, odnosno u vremenu O. Uzorci su centrifugirani 10 minuta pri 600g te su dobiveni talozi, koji sadr~e 4 x 106 stanica, isprani 3 puta hladnim PBS i pohranjeni na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. Talozima koji sadr~e 1 x 106 stanica dodano je 350 (l pufera RLT/1% (-merkaptoetanol. Pohranjeni su na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.6 Tretiranje diferenciranih stanica THP-1 aspirinom, indometacinom, hidrokortizonom i deksametazonom Stanice THP-1 diferencirane su uz forbol-12-miristat-13-acetat (koncentracije 10-7 M) tijekom 48 sati prema postupku opisanom u poglavlju 2.2.1. Medij (koji sadr~i PMA) je zatim odliven, a adheriranim je stanicama potom dodan: medij. Te su stanice smatrane kontrolnim, netretiranim diferenciranim stanicama. medij/0,2% DMSO. Te su stanice smatrane kontrolnim diferenciranim stanicama tretiranim DMSO kako bi se isklju io mogui utjecaj DMSO. 0,5 mM otopina aspirina u mediju, u kojoj je koncentracija DMSO < 0,2%. 2 M otopina aspirina u mediju, u kojoj je koncentracije DMSO < 0,2%. 1 (M otopina indometacina u mediju, u kojoj je koncentracija DMSO < 0,2%. 10 (M otopina indometacina u mediju, u kojoj je koncentracija DMSO 0,2%. 1 (M otopina hidrokortizona u mediju, u kojoj je koncentracija DMSO < 0,2%. 10 (M otopina hidrokortizona u mediju, u kojoj je koncentracija DMSO 0,2%. 1 nM otopina deksametazona u mediju, u kojoj je koncentracija DMSO < 0,2%. 100 nM otopina deksametazona u mediju, u kojoj je koncentracija DMSO 0,2%. Stanice su kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata. Medij je potom odliven, a adherirane su stanice isprane hladnim PBS. Blagim struganjem silikonskim struga em podignuto je: ( 4 x 106 stanica iz 100 x 20 mm Petri zdjelica u 2 ml hladnog PBS. Nakon centrifugiranja 10 minuta pri 600g stanice su joa dva puta isprane PBS a potom pohranjene na  20C do priprave stani nog homogenata prema postupku opisanom u poglavlju 2.2.14. ( 1 x 106 stanica iz 60 x 15 mm Petri zdjelica u 350 (l pufera RLT/1 % (-merkaptoetanol te pohranjeno na  20C do izolacije ukupne RNA prema postupku opisanom u poglavlju 2.2.9. 2.2.7 Odreivanje broja ~ivih stanica bojanjem tripanskim modrilom Postotak ~ivih stanica nakon tretiranja stanica opisanog u poglavljima 2.2.1 do 2.2.6 odreen je po sakupljanju stanica metodom bojanja tripanskim modrilom. }ive stanice tripansko modrilo koje prodire u unutraanjost stanice aktivno izbacuju, do im mrtve stanice nemaju to svojstvo, pa boja slobodno prodire u stanicu i zadr~ava se u njoj. Suspenzija stanica u PBS (20 (l) pomijeaana je s jednakim volumenom 0,2% otopine tripanskog modrila. Brkerova komorica napunjena je prireenim uzorkom te su pod mikroskopom (Opton, poveanje 8 x 40) izbrojane neobojane i obojane stanice. Rezultat je izra~en kao postotak ~ivih stanica. 2.2.8 MTS test MTS test (CellTiter 96(AQueous One Solution Cell Proliferation Assay) je kolorimetrijska metoda ispitivanja proliferacije i citotoksi nosti. CellTiter 96(AQueous One Solution reagens sadr~i tetrazolijevu sol (3-(4,5-dimetiltiazol-2-il)-5-(3-karboksimetoksifenil)-2-(4-sulfofenil)-2H-tetrazolij-bromid (MTS) i elektron-akceptor fenazin-etosulfat (PES). MTS (blijedo-~uti supstrat) se reducira enzimima endoplazmatskog retikuluma pri emu nastaje formazan (tamnosmei krajnji produkt). Kako MTS reduciraju samo ~ive stanice, ovim se kolorimetrijskim testom mjeri redukcijski potencijal stanica odnosno metaboli ka aktivnost stanica. Za potrebe ispitivanja proliferacije stanica THP-1 i citotoksi nog djelovanja primijenjenih tvari u ja~ice mikrotitarske plo e (s 96 ja~ica) nanijeto je 50 (l stani ne kulture THP-1 koncentracije 1 x 106 stanica/ml ili 50 (l hranidbenog medija bez stanica. Potom je u ja~ice nanijeto 50 (l otopine razli itih koncentracija aspirina, indometacina, hidrokortizona, deksametazona i DMSO te je kona na koncentracija spojeva bila jedna od navedenih: 0,1 mM, 0,5 mM, 1 mM i 2 mM aspirin 0,1 (M, 1 (M, 5 (M i 10 (M indometacin 0,1 (M, 1 (M, 5 (M i 10 (M indometacin 0,1 nM, 1 nM, 10 nM i 100 nM deksametazon 0,2 % DMSO Za potrebe ispitivanja proliferacije stanica THP-1 diferenciranih10-7 M otopinom PMA i citotoksi nog djelovanja primijenjenih tvari na njih, u ja~ice mikrotitarske plo e (s 96 ja~ica) nanijeto je po 50 (l stani ne kulture THP-1 stanica koncentracije 1x 106 stanica/ml te 50 (l 2 x 10-7 M otopine PMA. Nakon 48 sati inkubacije medij je odsisan, a na adherirane je stanice dodano po 100 (l otopine razli itih koncentracija aspirina, indometacina, hidrokortizona, deksametazona i DMSO, te je kona na koncentracija spojeva bila jedna od navedenih: 0,5 mM i 2 mM aspirin 1 (M i 10 (M indometacin 1 (M i 10 (M indometacin 1 (M i 100 (M deksametazon 0,2 % DMSO Nakon inkubacije tijekom 24, 48 te 72 sata u termostatu pri 37C u struji zraka s 5%CO2, relativne vla~nosti 95%, u ja~ice je dodano po 20 (l reagensa CellTiter 96(AQueous One Solution te je nakon 2 sata inkubacije pri 37C izmjerena apsorbancija otopina pri 490 nm na ureaju Victor2-1420 Multilabel Counter (PerkinElmer). Rezultati su izra~eni kao omjeri apsorpcijskih jedinica izmjerenih za tretirane stanice (umanjenih za apsorpcijske jedinice medija sa spojevima bez stanica) i apsorpcijskih jedinica izmjerenih za netretirane stanice (umanjenih za apsorpcijske jedinice hranidbenog medija). 2.2.9 Izolacija ukupne RNA Ukupna RNA izolirana je iz stanica u kulturi uporabom kompleta RNeasy Mini Kit. Stanice su lizirane dodatkom 350 (l pufera RLT/1% (-merkaptoetanol, u lizat je dodano 350 (l 70% etanola u vodi te je tako prireena smjesa nanijeta na RNeasy mini kolonu. Nakon centrifugiranja 15 sekundi pri 8000g filtrat je odba en, a kolona je isprana puferom RW1 (700 (l) te dva puta puferom RPE (500 (l) (nakon dodatka svakog pojedinog pufera uslijedilo je centrifugiranje 15 sec pri 8000g i odbacivanje filtrata). Ukupna RNA s kolone je eluirana vodom (u kojoj su uniatene ribonukleaze - 30 (l) uz centrifugiranje 1 min pri 8000g. U otopinu RNA odmah je dodano 1 l otopine ribonukleaznog inhibitora (do kona ne koncentracije 1 U/l) te je otopina RNA pohranjena na -80(C. 2.2.10 Odreivanje koncentracije i kvalitete izolirane RNA Provjera integriteta i istoe izolirane ukupne RNA, kao i odreivanje koncentracije RNA provedena je primjenom tehnologije  laboratorija na ipu na ureaju Agilent 2100 bioanalyzer (Agilent Technologies) uporabom kompleta RNA 6000 Nano LabChipKit. Metoda se temelji na elektrokineti kom i tla nom razdvajanju komponenata smjese nanijete na ip-plo icu ija se unutraanjost sastoji od mre~e tankih kanala ispunjenih gelom, puferom i bojom. Na ip se nanosi po 1 l uzorka (otopine izolirane ukupne RNA) kao i standard (otopina molekula RNA razli itih veli ina i poznatih koncentracija). Razdvajanje i analiza rezultata provodi se na ureaju Agilent 2100 bioanalyzer. Rezultati se prikazuju u obliku virtualnog gela ili elektroferograma. Analizom dobivenih vrpci mogue je odrediti koncentraciju ukupne RNA te ujedno provjeriti kvalitetu (intaktnost RNA) i istou (prisutnost DNA) uzorka (slika 2.1).  Slika 2.1 Analiza ukupne RNA na ureaju Agilent 2100 bioanalyser. Elektroferogrami i virtualni gelovi dobiveni analizom ukupne RNA na ureaju Agilent 2100 bioanalyser. a) visokokvalitetna ukupna RNA b) ukupna RNA kontaminirana s DNA c) raspadnuta ukupna RNA. 2.2.11 Reverzna transkripcija praena lan anom reakcijom polimerazom Uzorci ukupne RNA za koje je postupkom opisanim u poglavlju 2.2.10 utvreno da sadr~e intaktnu RNA bez kontaminacije s DNA reverzno su transkribirani uz uporabu oligo dT ishodnica. Reakcijske smjese za reverznu transkripciju pripravljene su kao ato je navedeno u tablici 2.1. Ukupan volumen reakcijske smjese je bio 20 l. Tablica 2.1. Sastav reakcijskih smjesa za reverznu transkripciju. sastojakvolumenkona na koncentracijaMgCl2 (50 mM)2 l5 mM10 x PCR pufer (500 mM KCl, 100 mM Tris/HCl, pH= 8,3)2 l1 xsmjesa dNTP (25 mM dATP, 25 mM dCTP, 25 mM dTTP, 25 mM dGTP)0,8 l1 mMMuL V reverzna transkriptaza (200 U/l)0,252 l2,5 U/lishodnica oligo dT (100 M)0,5 l2,5 Motopina RNA3,2 l~0,8 g/20 lDEPC-H2011,248 l Reverzna transkripcija odvijala se u ureaju za lan anu reakciju polimerazom (GeneAmp(PCR System 2700, Applied Biosystems) tijekom 60 minuta pri 37(C. Potom je uslijedilo zagrijavanje reakcijske smjese na 95(C tijekom 5 minuta kako bi se razdvojili hibridi RNA-cDNA i naglo hlaenje na +4(C. Kako bi se umno~ili dijelovi mRNA za galektin-3 te -aktin, odvojeno po 2 l uzorka nakon reverzne transkripcije te su u njih dodani reagensi potrebni za lan anu reakciju polimerazom prema tablici 2.2 (ukupan volumen reakcijske smjese je 10 l). Tablica 2.2 Sastav reakcijskih smjesa za lan anu reakciju polimerazom. sastojakvolumenkona na koncentracijaMgCl2 (50 mM)0,1 l1,5 mM10 x PCR pufer (500 mM KCl, 100 mM Tris/HCl, pH= 8,3)0,8 l1 xPlatinum Taq polimeraza (5 U/l)0,1 l0,05 U/lsense ishodnica (50 M)0,1 l0,5 Mantisense ishodnica (50 M)0,1 l0,5 MDEPC-H206,8 l Kao ishodnice rabljene su ili ishodnice za -aktin ili ishodnice za galektin-3 (Tablica 2.3). Sense ishodnice bile su na 5'-kraju obilje~ene fluorescentnom bojom 6-karboksifluoresceinom (6-FAM). Tablica 2.3 Ishodnice za lan anu reakciju polimerazom. ishodnicanukleotidni slijedTm (C)veli ina PCR produktasense - za galektin-35'-TGCTGATAACAATTCTGGGC-3'62,5288 bpantisense - za galektin-35'-TAACCCGATGATTGTACTGC-3'60sense - za -aktin5'-CATGTGCAAGGCCGGCTTCG-3'75226 bpantisense - za -aktin5'-GAAGGTGTGGTGCCAGATTT-3'63,9 Reakcija se odvijala u ureaju za lan anu reakciju polimerazom na sljedee na ine, ovisno o tome jesu li rabljene ishodnice za -aktin ili galektin-3: ishodnice za -aktinishodnice za galektin-31. po etna denaturacija95(C ; 1 minuta 45 sekundi95(C ; 1 minuta 45 sekundi2. a) denaturacija b) sljepljivanje ishodnica c) elongacija95(C ; 1 minuta 63(C ; 1 minuta 14 ciklusa 72(C ; 1 minuta95(C ; 1 minuta 58(C ; 1 minuta 21 ciklus 72(C ; 1 minuta3. zavrna elongacija72(C ; 5 minuta72(C ; 5 minuta Slijedilo je naglo hlaenje na +4(C. 2.2.12 Odreivanje nukleotidnog slijeda PCR produkata Kako bismo provjerili jesu li se po izolaciji RNA i reverznoj transkripciji u lan anoj reakciji polimerazom zaista umno~ili upravo i islju ivo dijelovi mRNA koji kodiraju galektin-3 i -aktin, bilo je potrebno provjeriti nukleotidni slijed PCR produkata. Odreivanje nukleotidnog slijeda sastoji se od nekoliko koraka: pro iaavanje PCR produkata nakon lan ane reakcije polimerazom provjera uspjeanosti pro iaavanja i odreivanje koncentracije PCR produkata elektroforezom u agaroznom gelu reakcija sekvenciranja pro iaavanje produkata sekvenciranja odreivanje nukleotidnog slijeda razdvajanjem produkata sekvenciranja na ureaju ABI Prism 310 Genetic Analyzer (Applied Biosystems) pro iaavanje PCR produkata nakon lan ane reakcije polimerazom Pro iaavanje PCR produkata nakon lan ane reakcije polimerazom provedeno je uporabom kompleta QIAquick PCR Purification Kit. Nakon lan ane reakcije polimerazom opisane u poglavlju 2.2.11 (uz jedinu modifikaciju opisanog postupka  umna~anje dijelova cDNA za -aktin i galektin-3 odvijalo se tijekom 30 ciklusa), 15 l smjese pomijeaano je s 75 l pufera PB i nanijeto na QIAquick kolonu. Nakon centrifugiranja 60 sekundi pri 12000g filtrat je ba en a pranje vezane DNA postignuto je dodatkom 750 l pufera PE i centrifugiranjem 2 x 60 sekundi pri 12000g. QIAquick kolona je premjeatena u novu epruvetu, na nju je dodano 30 l pufera EB, te je nakon inkubacije 1 minutu pri sobnoj temperaturi i centrifugiranja 1 minutu pri 12000g sakupljen eluat koji sadr~i PCR produkte pro iaene od dNTP i ishodnica zaostalih nakon lan ane reakcije polimerazom. 2. provjera uspjeanosti pro iaavanja i odreivanje koncentracije PCR produkata elektroforezom u agaroznom gelu 1,5% agarozni gel prireen je zagrijavanjem agaroze u TAE-puferu do vrenja. Otopina ohlaena do 60C izlivena je u kalup. Uzorcima pro iaenih PCR produkata dodana je 1/5 volumena 6 x pufera za nanoaenje uzoraka DNA. U jednu je ja~icu stavljen i standard DNA (GeneRuler( 100 bp DNA Ladder). Elektroforeza je provedena u TAE-puferu pri jakosti elektri nog polja 5-7 V/cm dok tamnoplava boja (bromfenol plavo) nije dosegla rub gela. Gel je bojan 20 minuta u otopini etidij-bromida (0,5 (g/ml) u vodi. Polo~aj DNA u gelu utvren je na temelju fluorescencije interkaliranog etidijevog bromida pri UV-svjetlu valne duljine 312 nm. Na temelju intenziteta fluorescencije mogue je pribli~no odrediti koncentraciju pro iaenih PCR produkata. Pri tom se intenzitet fluorescencije ~eljene vrpce usporeuje s intenzitetom fluorescencije vrpce pribli~no jednake veli ine koja sadr~i poznatu koli inu DNA. reakcija sekvenciranja Reakcija sekvenciranja pro iaenih PCR produkata provedena je uporabom kompleta ABI PRISM( BigDye( Terminator v3.0 Cycle Sequencing Ready Reaction Kit with AmpliTag(DNA Polymerase. Reakcija se temelji na dideoksi-metodi sekvenciranja DNA. Komplet uz polimerazu, pufer i dNTP sadr~i i dideoksi-NTP obilje~ene etirima razli itim rodaminskim fluorescentnim bojama (ddATP je obilje~en bojom dR6G, ddCTP bojom dROX, ddGTP bojom dR110 a ddTTP bojom dTAMRA). 10 l reakcijske smjese prireuje se mijeaanjem odgovarajueg volumena pro iaenog PCR produkta (koji sadr~i ~5 g PCR produkta), odgovarajueg volumena otopine sense ishodnice (koli ina ishodnica 1,6 pmol), 4 l smjese Ready Reaction mix i vode. Reakcija sekvenciranja se odvija u ureaju za lan anu reakciju polimerazom na sljedei na in: 1. po etna denaturacija95(C; 4 minute2. a) denaturacija b) sljepljivanje ishodnica c) elongacija95(C; 30 sekundi 50(C; 15 sekundi 25 ciklusa 60(C; 4 minute Slijedi naglo hlaenje na +4(C. 4. pro iaavanje produkata sekvenciranja Nakon reakcije sekvenciranja potrebno je iz reakcijske smjese ukloniti viaak dideoksi-NTP obilje~enih fluorescentnim bojama. U tu je svrhu rabljen komplet DyeEx( 2.0 Spin Kit. 1 l 2,2 % SDS dodan je u reakcijsku smjesu nakon reakcije sekvenciranja (u svrhu boljeg pro iaavanja) te je, nakon inkubacije 5 minuta pri 98C i 10 minuta pri 25C smjesa nanijeta na kolonu DyeEx( 2.0 Spin. Nakon centrifugiranja 3 minute pri 800g eluat (koji sadr~i pro iaene produkte sekvenciranja) je uparen do suha u ureaju "Speed Vac" (Applied Biosystems) tijekom 15 minuta pri sobnoj temperaturi. 5. ODREIVANJE NUKLEOTIDNOG SLIJEDA PCR PRODUKATA Upareni pro iaeni produkti sekvenciranja otopljeni su u 17 l reagensa Template Suppression Reagent (TSR, Applied Biosystems), denaturirani pri 95C tijekom dvije minute, te ohlaeni na ledu. Kapilarna elektroforeza i detekcija fluorescentno obilje~enih fragmenata DNA provedene su ureajem ABI Prism 310 Genetic Analyzer (Applied Biosystems). Elektroforeze su provedene tijekom 120 minuta koriatenjem modula Seq POP 6 (1 ml) E uz vrijeme injektiranja uzorka 30 sekundi, volta~i struje tijekom elektroforeze 12,2 kV te temperaturi polimera POP 6 50C. Za analizu rezultata koriaten je program Sequencing Analysis software. 2.2.13 Odreivanje veli ine i koli ine PCR produkata Nakon ato su lan anom reakcijom polimerazom umno~eni dijelovi mRNA za galektin-3 odnosno -aktin iz odreenog uzorka, po 0,5 l tih smjesa pomijeaano je s 0,5 l GeneScan(-500 TAMRA standarda i 11,5 l deioniziranog formamida. GeneScan(-500 TAMRA standard sadr~i fragmente DNA poznatih veli ina obilje~enih fluorescentom bojom TAMRA (karboksitetrametilrodamin). Kapilarna elektroforeza i detekcija fluore scentno obilje~enih fragmenata DNA obavljena je na ureaju ABI Prism 310 Genetic Analyzer (Applied Biosystems). Elektroforeze su provedene tijekom 24 minute koriatenjem modula GS STR POP 4 (1 ml) A uz vrijeme injektiranja uzorka 3 sekunde, volta~i struje tijekom elektroforeze 15 kV te temperaturi polimera POP 4 60C. Za kvantifikaciju koli ine fragmenata DNA koriaten je program GeneScan 2.1 software kojim su odreene povraine peakova na elektroforetogramu koji odgovaraju fragmentima DNA veli ine 226 bp (nastalih umna~anjem dijela mRNA za -aktin) odnosno 288 bp (nastalih umna~anjem dijela mRNA za galektin-3). Rezultati su izra~avani kao omjer povraina peakova za galektin-3 i -aktin odreenog uzorka u odnosu na omjer povraina peakova za galektin-3 i -aktin kontrolnog uzorka (stanica tretiranih 0,2% otopinom DMSO tijekom odgovarajuih vremena). 2.2.14 Priprava stani nog homogenata Sakupljene i isprane stanice (4 x 106 stanica) resuspendirane su u 150 l pufera za homogenizaciju stanica te sonicirane ultrazvu nim sonikatorom (Soniprep 150, MSE Scientific Instruments) 4 x 15 sec uz amplitudu 5 m i frekvenciju 22 kHz na ledu. Nakon centrifugiranja 10 minuta pri 600 g odvojeni su stani ni homogenati od netopljivih taloga. 2.2.15 Odreivanje koncentracije proteina Odmah nakon homogenizacije, koncentracija proteina u stani nim homogenatima odreena je metodom s bicinkonini nom kiselinom (BCA)  ADDIN EN.CITE Smith19852030384370515011985OctMeasurement of protein using bicinchoninic acid76-85Smith, P. K.Krohn, R. I.Hermanson, G. T.Mallia, A. K.Gartner, F. H.Provenzano, M. D.Fujimoto, E. K.Goeke, N. M.Olson, B. J.Klenk, D. C.Anal BiochemHydrogen-Ion ConcentrationIndicators and ReagentsProteins/*analysis*QuinolinesSolubilitySpectrophotometryhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&amp;list_uids=3843705</URL></MDL></Cite></EndNote>[171]. U tu svrhu po 2 l stani nih homogenata razrijeeno je vodom do kona nog volumena 100 l (razrjeenje 1:50). Uzorci (100 (l) inkubirani su 30 minuta pri 37C s 1 ml otopine dobivene mijeaanjem otopine BCA iz kita s 4%-tnom otopinom CuSO4x5H2O u volumnom omjeru 50:1. Nakon ato su uzorci ohlaeni na sobnu temperaturu, intenzitet obojenja otopine izmjeren je pri valnoj duljini 562 nm na spektrofotometru Uvikon 860 (Kontron Instruments). Koncentracija proteina u uzorcima o itana je iz ba~darnog pravca. Kao standardna otopina proteina rabljena je otopina BSA u dest. vodi (podru je koncentracija 0,01  0,1 mg/ml). Uzorci standardne otopine, kao i otopine uzoraka, prireeni su u duplikatu. 2.2.16 Elektroforeza u SDS-poliakrilamidnom gelu Odgovarajuim volumenima uzoraka stani nih homogenata koji su sadr~avali 300 (g proteina dodano je 100 (l pufera za nanoaenje uzoraka na SDS-PAG, pH 6,8 te destilirane vode do kona nog volumena 300 (l (odnosno kona ne koncentracije proteina 1 mg/ml). Tako prireeni uzorci kuhani su 5 minuta pri 96C. Diskontinuirana denaturirajua gel-elektroforeza proteina provedena je u 5%-tnom sabijajuem i 12%-tnom razdvajajuem poliakrilamidnog gelu, debljine 0,5 mm, prema metodi po Laemmliju  ADDIN EN.CITE Laemmli1970201054320632272591970Aug 15Cleavage of structural proteins during the assembly of the head of bacteriophage T4680-5Laemmli, U. K.NatureAutoradiographyCarbon IsotopesColiphages/*metabolismElectrophoresis, DiscGenes, StructuralGenetics, MicrobialKineticsViral Proteins/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=5432063[172]. U pojedinu je ja~icu nanoaeno po 5 (l uzorka. Uzorka proteina poznatih molekulskih masa nanoaeno je po 3 (l. 2.2.17 Postupak Western-blot Elektroforezom razdvojeni proteini prenijeti su na PVDF Immobilon membranu u ureaju za polusuhi elektroforetski prijenos (LKB, Bromma, Pharmacia) pri gustoi elektri ne struje 0,8 mAcm-2  ADDIN EN.CITE <EndNote><Cite><Author>Towbin</Author><Year>1979</Year><RecNum>202</RecNum><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>388439</ACCESSION_NUMBER><VOLUME>7691979SepElectrophoretic transfer of proteins from polyacrylamide gels to nitrocellulose sheets: procedure and some applications4350-4Towbin, H.Staehelin, T.Gordon, J.Proc Natl Acad Sci U S AAnimalsAntigen-Antibody ReactionsAntigens/analysisCelluloseChickensElectrophoresis, Polyacrylamide Gel/*methodsEscherichia coli/analysisImmunologic TechniquesLiver/analysisRibosomal Proteins/analysis/immunologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=388439[173]. Membrane su potom uzdu~no rezrezane na dva dijela na polo~aju gdje se po prilici nalaze proteini veli ine 35 kDa. Membrane su inkubirane 1 sat u puferu TBST, pH 7,5, u svrhu spre avanja nespecifi nog vezanja proteina na slobodna vezna mjesta na membrani. Potom su dijelovi membrane na kojima se nalaze proteini: a) manji od 35 kDa izlo~ene vezanju atakorskog monoklonskog protutijela M3/38 (1:200) razvijenog naspram miajeg galektina-3 preko noi b) vei od 35 kDa izlo~ene vezanju miajeg monoklonskog protutijela (1:2500) razvijenog naspram kokoajeg aktina preko noi. Membrane su zatim isprane (3 puta po 10 minuta) u puferu TBST, pH 7,5, a zatim izlo~ene vezanju odgovarajueg sekundarnog protutijela tijekom jednog sata, odnosno: a) kozjeg poliklonskog protutijela razvijenog naspram cijele molekule atakorskog imunoglobulina G (1:10000) obilje~enog peroksidazom iz hrena b) kozjeg poliklonskog protutijela razvijenog naspram cijele molekule miajeg imunoglobulina M (1:10000) obilje~enog peroksidazom iz hrena. Po ispiranju membrane 3 puta po 10 minuta u puferu TBST, pH 7,5 membrane su inkubirane s reagensom za poja anu kemiluminiscenciju tijekom jedne minute, a zatim je na njih u mraku postavljen film (Lumi-Film Chemiluminescent Detection Film, Roche), pri emu vrijeme izlaganje filma membrani ovisilo o intenzitetu signala. Film je potom razvijen uporabom razvija a (GBX Developer/Replanisher, Sigma) i fiksativa (GBX Fixer/Replanisher, Sigma). Intenzitet signala izmjeren je denzitometrijskom analizom filma koriatenjem programa Scion Image, Microsoft(. Rezultati denzitometrijske analize vrpci izra~eni su u relativnim denzitometrijskim jedinicama. 2.2.18 Statisti ke metode Srednja vrijednost (x) i standardna devijacija (SD) izra unate su pomou ra unala koriatenjem programa Excel, Microsoft(. Razina zna ajnosti (p) izra unata je koriatenjem Student-t testa, istog programa. Statisti ki zna ajnom promjenom smatrana je promjena za koju je p<0,05. REZULTATI Galektin-3, lektin koji specifi no prepoznaje -galaktozidne strukture i kojeg, izmeu ostalog, eksprimiraju monociti i makrofagi, ima va~nu ulogu u imunim reakcijama i upalnom odgovoru. On modulira stani nu proliferaciju, stani nu adheziju, kemotaksiju, fagocitozu, sintezu upalnih medijatora te apoptozu limfocita T. U radu smo ispitali utjecaj razli itih koncentracija nesteroidnih tvari s imunomodulatornim djelovanjem aspirina i indometacina, te steroidnih tvari s imunomodulatornim djelovanjem hidrokortizona i deksametazona na ekspresiju gena LGALS3 i njegovog proteinskog produkta - galektina-3 u stanicama monocitne stani ne linije THP-1. Razina ekspresije LGALS3 i galektina-3 preena je i tijekom diferencijacije stanica THP-1 koja je bila potaknuta forbol-12-miristat-13-acetatom, spojem pod ijim utjecajem stanice THP-1 poprimaju karakteristike makrofaga. Utjecaj razli itih koncentracija aspirina, indometacina, hidrokortizona i deksametazona na ekspresiju LGALS3 i galektina-3 ispitan je i u diferenciranim stanicama stani ne linije THP-1. U svim je pokusima ispitana i vijabilnost i metaboli ka aktivnost stanica. 3.1 Ekspresija LGALS3 i galektina-3 u stanicama THP-1 tijekom kultiviranja Tvari s imunomodulatornim djelovanjem koriatene u ovom radu (aspirin, indometacin, hidrokortizon i deksametazon) su slabo ili uope nisu topljive u vodenim otopinama. Zbog toga su mati ne otopine tvari prireene u u DMSO, a potom razrijeene u vodenoj otopini (preciznije, hranidbenom mediju) do odgovarajuih koncentracija imunomodulatornih tvari. Maksimalna koncentracija DMSO u hranidbenom mediju iznosila je 0,2%. Prije ispitivanja utjecaja razli itih imunomodulatornih tvari na ekspresiju gena LGALS3 i galektina-3 odreene su: vijabilnost i metaboli ka aktivnost stanica stani ne linije THP-1 tijekom 72-satnog kultiviranja u hranidbenom mediju (medij) kao i tijekom 72-satnog kultiviranja u 0,2% otopini DMSO u hranidbenom mediju (medij/0,2% DMSO) razina ekspresije LGALS3 i galektina-3 tijekom 72-satnog kultiviranja stanica stani ne linije THP-1 u hranidbenom mediju (medij) kao i tijekom 72-satnog kultiviranja stanica stani ne linije THP-1 u 0,2% otopini DMSO u hranidbenom mediju (medij/0,2% DMSO). U cilju odreivanja vijabilnosti i metaboli ke aktivnosti stanica THP-1 stanice su kultivirane tijekom 24, 48 ili 72 sata u mediju ili mediju/0,2% DMSO. Pokusi su ponovljeni 2 puta u kvadriplikatu. Postotak ~ivih stanica odreen je metodom bojanja tripanskim modrilom (opisanom u poglavlju 2.2.7). Rezultati su pokazali da nema razlike u vijabilnosti stanica kultiviranih u mediju/0,2% DMSO u odnosu na stanice kultivirane samo u hranidbenom mediju, te da je u svim slu ajevima postotak ~ivih stanica vei od 98%. Metaboli ka aktivnost stanica uz 0,2% DMSO i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da medij/0,2% DMSO ne djeluje citotoksi no na stanice stani ne linije THP-1 niti nakon 72 sata kultiviranja te je njihova metaboli ka aktivnost nepromijenjena (100(2,8%) u odnosu na metaboli ku aktivnost stanica kultiviranih u hranidbenom mediju (Slika 3.1).  Slika 3.1 Metaboli ka aktivnost stanica THP-1 tijekom 72-satnog kultiviranja u mediju/0,2% DMSO se ne mijenja. a) Slika mikrotitarske plo ice nakon MTS testa stanica THP-1 kultiviranih 24 sata u mediju ili mediju/0,2% DMSO. b) Metaboli ke aktivnosti stanica THP-1 kultiviranih tijekom 24, 48 ili 72 sata u mediju odnosno mediju/0,2% DMSO izmjerene MTS testom izra~ene u odnosu na stanice kultivirane tijekom istog vremena u mediju (rezultati kvadriplikata dva neovisna pokusa izra~eni su kao x (SD). Kako bismo utvrdili relativnu razinu ekspresije gena LGALS3, odnosno relativnu koli inu mRNA za galektin-3, iz stanica THP-1 izolirana je ukupna RNA te je postupkom reverzne transkripcije uz primjenu oligo dT ishodnica prevedena u cDNA. Dijelovi mRNA za galektin-3 i -aktin umno~eni su postupkom lan ane reakcije polimerazom uz primjenu specifi nih ishodnica. Nakon lan ane reakcije polimerazom nukleotidni slijed PCR produkata provjeren je metodom sekvenciranja na ureaju ABI Prism 310 Genetic Analyzer (Applied Biosystems), prema postupku opisanom u poglavlju 2.2.12. Ukratko, nakon lan ane reakcije polimerazom PCR produkti su pro iaeni od viaka ishodnica, a uspjeanost pro iaavanja i pribli~na koncentracija pro iaenih PCR produkata odreena je elektroforezom pro iaenih PCR produkata u 1,5% agaroznom gelu (Slika 3.2).  Slika 3.2 Elektroforeza PCR produkata. Nakon izolacije ukupne RNA iz stanica THP-1, reverzne transkripcije i lan ane reakcije polimerazom, PCR produkti su analizirani elektroforezom u 1,5% agaroznom gelu a) prije pro iaavanja i b) nakon pro iaavanja. 1: standard molekulskih masa (viaekratnik 100 bp) 2: PCR produkt veli ine 226 bp (nastao umna~anjem dijela mRNA za -aktin) 3: PCR produkt veli ine 288 bp (nastao umna~anjem dijela mRNA za galektin-3). PCR produkti (5 g) podvrgnuti su reakciji sekvenciranja. Nakon uklanjanja zaostalih fluorescentno obilje~enih nukleotida i viaka ishodnica, nukleotidni je slijed PCR produkata odreen analizom produkata sekvenciranja razdvojenih kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser (Applied Bioystems)(Slika 3.3). Analiza je provedena koriatenjem programa Sequencing Analysis Software, a rezultati su pokazali da nukleotidni sljedovi PCR produkata nastalih uporabom odabranih ishodnica u potpunosti odgovaraju dijelovima mRNA za galektin-3 odnosno -aktin. PCR produkti su veli ine 226 baznih parova (nastali umna~anjem dijela mRNA za -aktin) odnosno 288 baznih parova (nastali umna~anjem dijela mRNA za galektin-3). Utvreno je takoer da su spomenuti produkti ujedno i jedini produkti lan ane reakcije polimerazom te da su odabrane temperature pri kojima se odvijao proces sljepljivanja ishodnica tijekom lan ane reakcije polimerazom (63C za ishodnice za -aktin, odnosno 58C za ishodnice za galektin-3) odgovarajue jer se pri navedenim temperaturama ishodnice ve~u isklju ivo na u potpunosti komplementarne dijelove DNA.  Slika 3.3 Nukleotidni slijed PCR produkta nastalog umna~anjem dijela mRNA za galektin-3 odreen analizom na ureaju AbiPrism 310 Genetic Analyser. S ciljem utvrivanja relativne razine ekspresije gena LGALS3, odnosno relativne koli ine mRNA za galektin-3 tijekom 72-satnog kultiviranja (uzorci sakupljeni nakon 1, 3, 5, 24, 48 ili 72 sata) stanica stani ne linije THP-1 u mediju ili mediju/0,2% DMSO, iz stanica je izolirana ukupna RNA, ~0,8 g ukupne RNA reverzno je transkribirano uz primjenu oligo dT ishodnica te su potom umno~eni dijelovi mRNA za galektin-3 (tijekom 21 ciklusa PCR uz temperaturu sljepljivanja ishodnica 58C) i -aktin (tijekom 14 ciklusa PCR uz temperaturu sljepljivanja ishodnica 63C) primjenom odabranih specifi nih ishodnica. Lan ana reakcija polimerazom ne dose~e krajnju to ku ("end-point") uz navedene reakcijske uvjete ve je joa uvijek u eksponencijalnoj fazi, ato je uvjet da bi ta metoda bila kvantitativna. Na opisani na in dobivene koncentracije PCR produkata prikladne su za analizu koli ine PCR produkata na ureaju AbiPrism 310 Genetic Analyser. PCR produkti su zdru~eni, razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser (Slika 3.4), a relativan omjer njihovih koli ina izra unat je uporabom programa GeneScan Analysis Software.  Slika 3.4 GeneScan elektroforetogram razdvojenih PCR produkata. Ukupna RNA izolirana je iz stanica THP-1 i reverzno transkribirana uz primjenu oligo dT ishodnica. PCR produkti dobiveni primjenom fluorescentno obilje~enih specifi nih ishodnica za mRNA za -aktin tijekom 14 ciklusa PCR i PCR produkti dobiveni primjenom fluorescentno obilje~enih specifi nih ishodnica za mRNA za galektin-3 tijekom 21 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Os ordinate predstavlja relativan intenzitet fluorescencije, a os aspcise veli inu razdvojenih fragmenata izra~eno u baznim parovima (bp). Crni peakovi odgovararaju standardu DNA GeneScan(-500 TAMRA, a plavi PCR produktu -aktina (226 bp) i galektina-3 (288 bp). Rezultati dvaju neovisnih pokusa napravljenih u duplikatu pokazali su da se tijekom 1, 3, 5, 24, 48 i 72 sata kultiviranja stanica THP-1 u mediju kao i u mediju/0,2% DMSO relativni omjeri PCR produkata nastalih umna~anjem dijela mRNA za galektin-3 i -aktin ne mijenjaju (6,28(0,13), odnosno, da se koli ina mRNA za galektin-3 odr~ava na jednakoj razini (Slika 3.5).  Slika 3.5 U stanicama THP-1 tijekom 72-satnog kultiviranja u mediju ili u mediju/0,2% DMSO koli ina mRNA za galektin-3 se ne mijenja. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u mediju ili u mediju/0,2% DMSO. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina (x (SD). Zeleni stupci  kultiviranje u mediju, bijeli stupci  kultiviranje u mediju/0,2% DMSO. Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 ispitana je tijekom 72-satnog kultiviranja postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena ekspresija konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka. Rezultati su pokazali da se koli ina galektina-3 statisti ki zna ajno ne mijenja (p>0,05) tijekom 1, 3, 5, 24, 48 i 72 sata kultiviranja stanica THP-1 u mediju u odnosu na po etnu koli inu galektina-3 (105(5,5% po etne razine). Razina ekspresije galektina-3 statisti ki se zna ajno ne mijenja (p>0,05) niti kada se stanice kultiviraju tijekom 1, 3, 5, 24, 48 i 72 sata u mediju/0,2% DMSO (103(7% po etne razine)(Slika 3.6).   Slika 3.6 Koli ina galektina-3 u stanicama THP-1 tijekom kultiviranja u mediju ili u mediju/0,2% DMSO se ne mijenja. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju i b) mediju/0,2% DMSO (5 g proteina/ja~ici). c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x (SD). Crni stupac-kontrola, zeleni stupci-kultiviranje u mediju, bijeli stupci  kultiviranje u mediju/0,2% DMSO. Utjecaj aspirina na ekspresiju LGALS3 i galektina-3 u stanicama THP-1 Aspirin utje e na signalne puteve i transkripcijske fakore koji izmeu ostalog, sudjeluju i u regulaciji ekspresije gena LGALS3. Stoga smo u ovom radu ~eljeli ispitati u inak aspirina na ekspresiju LGALS3 i galektina-3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u 0,1 mM, 0,5 mM, 1 mM i 2 mM otopini aspirina u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Postotak ~ivih stanica odreen je metodom bojanja tripanskim modrilom (opisanom u poglavlju 2.2.7). Rezultati su pokazali da je postotak ~ivih stanica u svim ispitivanim slu ajevima e"95%, odnosno, da aspirin u primijenjenim koncentracijama ne utje e na vijabilnost stanica THP-1 tijekom 72 sata kultiviranja. Metaboli ka aktivnost stanica kultiviranih uz aspirin koncentracija 0,1 mM, 0,5 mM, 1 mM i 2 mM i citotoksi nost primijenjenih koncentracija aspirina odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da 0,1 mM, 0,5 mM, 1 mM i 2 mM otopine aspirina statisti ki zna ajno ne mijenjaju metaboli ku aktivnost stanica u odnosu na stanice kultivirane u hranidbenom mediju, odnosno da primjenjene koncentracije aspirina nemaju citotoksi an u inak jer je u svim slu ajevima metaboli ka aktivnost bila e"91% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Nakon izolacije RNA iz tretiranih stanica i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u stanicama THP-1 kultiviranih tijekom istih vremena u mediju/0,2% DMSO (Slika 3.7). Razina ekspresije LGALS3 u stanicama THP-1 kultiviranim uz aspirin koncentracije 0,1 mM ne mijenja se tijekom prva tri sata kultiviranja, ali se nakon pet sati kultiviranja primjeuje smanjenje razine ekspresije LGALS3 u odnosu na kontrolu. Ono se kontinuirano nastavlja daljnjim kultiviranjem tijekom 24 sata kada iznosi 64,2(11,6% od razine u kontrolnim stanicama, nakon 48 sati 42,9(5,8%, da bi 72 sat razina ekspresije LGALS3 iznosila 31,1(1,4% od razine u kontrolnim stanicama. Kultiviranjem stanica THP-1 u 0,5 mM otopini aspirina/medij razina ekspresije LGALS3 se smanjuje gotovo na polovinu razine u kontrolnim stanicama ve nakon prvog sata (59,6(7,6%) kultiviranja. Daljnjim se kultiviranjem smanjenje nastavlja, pa nakon 3 sata razina galektina pada na 52,8(8%, 5 sati na 40,4(4,8%, 24 sata na 27(1,9%, 48 sati na 24,2(4% te nakon 72 sata kultiviranja na 25,6(4,8% kontrolne razine. 1 mM aspirin uzrokuje joa zna ajnije, vremenski-ovisno smanjenje razine ekspresije gena za galektin-3. Razina galektina-3 nakon prvog sata iznosi 52,2(1,7% razine u kontrolnim stanicama, odnosno po etne razine, nakon treeg sata 45,4(5,3%, petog sata 24,7(0,2%, 24 sata 28(3,9%, 48 sata 18,2% a nakon 72 sata kultiviranja samo 15,5(0,5%. Najja i pad razine ekspresije LGALS3 u stanicama THP-1 bio je izazvan najveom koriatenom koncentracija aspirina (2 mM). Nakon 72 sata kultiviranja uz 2 mM aspirin razina LGALS3 iznosi samo 12,3(0,04% kontrolne razine, a dostignuta je postupnim smanjenjem ekspresije - nakon 1 sat izlaganja stanica THP-1 aspirinu koncentracije 2 mM razina galektina-3 iznosila je 45(3,4%, 3 sata 40,2(0,8%, 5 sati 25,1(6,7% 24 sata 23(0,3%, a nakon 48 sati 15,4(0,5% po etne razine.  Slika 3.7 Aspirin inhibira ekspresiju LGALS3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz aspirin koncentracije 0,1 mM, 0,5 mM, 1 mM i 2 mM. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Crni stupci  kontrola, rebrasti stupci - 0,1 mM ASA, tamnozeleni stupci  0,5 mM ASA, svjetlozeleni stupci  1 mM ASA, bijeli stupci  2 mM ASA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz aspirin koncentracija 0,1 mM, 0,5 mM, 1 mM i 2 mM ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka. Aspirin u ni~im koncentracijama (0,1 mM i 0,5 mM) ne utje e na razinu ekspresije galektina-3 tijekom prvih 48 sati kultiviranja. Tek se nakon 72 sata kultiviranja primjeuje pad koli ine galektina-3 na ~70% po etne razine (preciznije, kultiviranjem u mediju/0,1 mM ASA smanjuje se koli ina galektina-3 na 70(4%, a u mediju/0,5 mM ASA na 67(6,2% od kontrolne razine). Kada su stanice kultivirane uz 1 mM aspirin pad koli ine galektina-3 zabilje~en je nakon 48 sati; razina galektina-3 tada je iznosila 79(2,8% po etne razine. Daljnjim kultiviranjem doalo je do joa zna ajnijeg smanjenja razine galektina-3 koja je nakon 72 sata iznosila 66(4% po etne razine. Kultiviranje stanica THP-1 uz aspirin koncentracije 2 mM, pad razine galektina-3 izaziva se joa ranije  ve nakon 24 sata razina galektina-3 iznosi 79(2,8% po etne razine. Zatim nastavlja padati te nakon 48 sati iznosi 71(3%, a nakon 72 sata 59(3,4% po etne razine (Slika 3.8).    Slika 3.8 Utjecaj aspirina na ekspresiju galektina-3 u stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/0,1 mM ASA; b) mediju/0,5 mM ASA; c) mediju/1 mM ASA; d) mediju/2 mM ASA (5 g proteina/ja~ici). e) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x ( SD). Crni stupac  kontrola, rebrasti stupci - 0,1 mM ASA, tamnozeleni stupci  0,5 mM ASA, svjetlozeleni stupci  1 mM ASA, bijeli stupci  2 mM ASA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Utjecaj indometacina na ekspresiju LGALS3 i galektina-3 u stanicama THP-1 Djelovanje indometacina, jake nesteroidne tvari s imunomodulatornim djelovanjem, vrlo je slabo istra~eno. Osim o djelovanju indometacina kao inhibitora ciklooksigenaze o kojem se zna neato viae, o njegovom u inku na signalne puteve koji su, izmeu ostalog, uklju eni i u regulaciju ekspresije galektina-3 postoje tek naznake. Stoga smo u ovom radu ~eljeli ispitati u inak indometacina na ekspresiju LGALS3 i galektina-3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u 0,1 (M, 1 (M, 5 (M i 10 (M otopini indometacina u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Postotak ~ivih stanica odreen je metodom bojanja tripanskim modrilom (opisanom u poglavlju 2.2.7). Rezultati su pokazali da je postotak ~ivih stanica u svim ispitivanim slu ajevima e"97%, odnosno, da indometacin u primijenjenim koncentracijama ne utje e na vijabilnost stanica THP-1 tijekom 72-satnog kultiviranja. Metaboli ka aktivnost stanica u prisutnosti primijenjenih koncentracija indometacina i njihova citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da se metaboli ka aktivnost stanica kultiviranih uz indometacin koncentracija 0,1 (M, 1 (M, 5 (M i 10 (M ne mijenja u odnosu na metaboli ku aktivnost stanica kultiviranih samo u hranidbenom mediju. Indometacin u primijenjenih koncentracijama ne djeluje citotoksi no na stanice THP-1 obzirom na to da je u svim slu ajevima metaboli ka aktivnost bila e"96% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Relativna razina ekspresije LGALS3 odreena je nakon izolacije ukupne RNA, reverzne transkripcije i lan ane reakcije polimerazom na ureaju AbiPrism 310 Genetic Analyser. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni istim postupkom u stanicama THP-1 kultiviranih tijekom istih vremena u mediju/0,2% DMSO (Slika 3.9). Izlaganje stanica THP-1 indometacinu u svim primijenjenim koncentracijama inhibira ekspresiju LGALS3 (Slika 3.9). Indometacin koncentracije 0,1 (M uzrokuje smanjenje koli ine mRNA za galektin-3 na 66,7(4,2% po etne razine ve nakon prvog sata kultiviranja. Daljnjim kultiviranjem inhibitorno djelovanje indometacina se nastavlja, te se koli ina mRNA za galektin-3 trei sat smanjuje na 48,1(8,5%, peti sat na 30,2(4,5, 24 sat na 29,2(2,6% i 48 sat na 22,7(0,8%, da bi nakon 72 sata iznosila tek 8%(0,5% po etne razine. Izlaganjem stanica THP-1 indometacinu koncentracije 1 (M razina ekspresije LGALS3 smanjuje se na polovinu po etne razine tijekom prvog sata (53,7(1,4%). Daljnjim kultiviranjem koli ina mRNA za galektin-3 se nastavlja smanjivati  nakon 3 sata kultiviranja iznosi 40,6(7,7%, nakon 5 sati 38,4(3,9%, 24 sata 22,2(4,5%, 48 sati 14,1(0,5% a nakon 72-satnog kultiviranja 15,6(0,5% po etne razine. Kada se stanice THP-1 izlo~e djelovanju indometacina koncentracije 5 (M, koli ina mRNA za galektin-3 smanji se za ~60% tijekom prva tri sata kultiviranja (nakon prvog sata iznosi 35,2(1,1%, a nakon treeg 45,6(7,4% po etne razine). Duljim izlaganjem stanica THP-1 indometacinu koncentracije 5 (M ekspresija LGALS3 se smanjuje joa desetak posto (5h-28,4(3,3%, 24 h-25,7(0,4%, 48h-20,0(5,3%, 72 h-28,2(9% po etne razine). Od svih primijenjenih koncentracija indometacina, najja e inhibitorno djelovanje na ekspresiju LGALS3 ima 10 (M indometacin. On inhibira ekspresiju LGALS3 za ~70% tijekom prvih pet sati kultiviranja (nakon prvog sata koli ina mRNA za galektin-3 iznosi 34,7(1,8%, nakon treeg 30,6(3,1%, a nakon petog 28,1(3,2% po etne razine). Daljnjim kultiviranjem koli ina mRNA za galektin-3 nastavlja padati i iznosi: 24h-23,2(0,7%, 48h-11,6(2,5% i 72 h-8,9(2,1% po etne razine.  Slika 3.9 Indometacin inhibira ekspresiju LGALS3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz indometacin koncentracije 0,1 (M, 1 (M, 5 (M i 10 (M. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Crni stupci  kontrola, rebrasti stupci- 0,1 (M Ind, tamnozeleni stupci  1 (M Ind, svjetlozeleni stupci  5 (M Ind, bijeli stupci  10 (M Ind. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz indometacin koncentracija 0,1 (M, 1 (M, 5 (M i 10 (M ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka (Slika 3.10). Koli ina galektina-3 tijekom prva 24 sata kultiviranja stanica THP-1 uz indometacin svih primjenjenih koncentracija (0,1 (M, 1 (M, 5 (M i 10 (M) odr~ava se na jednakoj razini. Indometacin koncentracije 0,1 (M nakon 48 sati kultiviranja uzrokuje pad razine galektina-3 na 69,0(5,1%, a nakon 72 sata kultiviranja na 48,5(4,3% po etne razine. Promjena koli ine galektina-3 u stanicama THP-1 izlo~enim djelovanju indometacina koncentracije 1 (M uo ava se tek nakon 72 sata kultiviranja, kada iznosi 61,0(6,1% po etne razine. Izlaganje stanica THP-1 indometacinu koncentracije 5 (M uzrokuje pad koli ine galektina-3 za ~20% nakon 48-satnog kultiviranja (80,5(2,7% po etne razine), odnosno ~30% nakon 72-satnog kultiviranja (72(3,8% po etne razine). Joa manju promjenu ekspresije galektina-3 uzrokuje indometacin koncentracije 10 (M: tek nakon 72 sata kultiviranja ona se mijenja i iznosi 81,2(4,4% po etne razine.    Slika 3.10 Utjecaj indometacina na ekspresiju galektina-3 u stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/0,1 (M Ind; b) mediju/1 (M Ind; c) mediju/5 (M Ind; d) mediju/10 (M Ind (5 g proteina/ja~ici). e) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x ( SD). Crni stupac  kontrola, rebrasti stupci-0,1 (M Ind, tamnozeleni stupci  1 (M Ind, svjetlozeleni stupci  5 (M Ind, bijeli stupci  10 (M Ind. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). 3.4 Utjecaj hidrokortizona na ekspresiju LGALS3 i galektina-3 u stanicama THP-1 Prema literaturi, za hidrokortizon, steroidni imunomodulatorni lijek postoje naznake da utje e na signalne puteve koji su, izmeu ostalog, uklju eni i u regulaciju ekspresije galektina-3. Stoga smo u ovom radu ~eljeli ispitati u inak hidrokortizona na ekspresiju LGALS3 i galektina-3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u 0,1 (M, 1 (M, 5 (M i 10 (M otopini hidrokortizona u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Postotak ~ivih stanica odreen je metodom bojanja tripanskim modrilom (opisanom u poglavlju 2.2.7). Rezultati su pokazali da je postotak ~ivih stanica u svim ispitivanim slu ajevima e"97,5%, odnosno, da hidrokortizon u primijenjenim koncentracijama ne utje e na vijabilnost stanica THP-1 tijekom 72-satnog kultiviranja. Metaboli ka aktivnost stanica i citotoksi nost primijenjenih koncentracija hidrokortizona odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da se metaboli ka aktivnost stanica kultiviranih uz hidrokortizon koncentracija 0,1 (M, 1 (M, 5 (M i 10 (M ne mijenja u odnosu na metaboli ku aktivnost stanica kultiviranih samo u hranidbenom mediju. Hidrokortizon u primijenjenih koncentracijama ne djeluje citotoksi no na stanice THP-1 obzirom da je u svim slu ajevima metaboli ka aktivnost bila e"96,4% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Relativna razina ekspresije LGALS3 odreena je nakon izolacije ukupne RNA, reverzne transkripcije i lan ane reakcije polimerazom na ureaju AbiPrism 310 Genetic Analyser. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni istim postupkom u stanicama THP-1 kultiviranih tijekom istih vremena u mediju/0,2% DMSO (Slika 3.11). Hidrokortizon u svim primijenjenim koncentracijama inhibira ekspresiju LGALS3 u stanicama THP-1, a postotak inhibicije je to vei ato je vrijeme izlo~enosti stanica djelovanju hidrokortizona dulje. Kultiviranjem stanica THP-1 u 0,1 (M otopini hidrokortizona/medij razina ekspresije LGALS3 se smanjuje na polovinu ve nakon prvog sata kultiviranja (50,8(8,6%), a smanjenje se nastavlja i tijekom 3 sata (na 42,4(2,4%), 5 sati (na 30,1(8,9%), 24 sata (na 25,7(7,6%), 48 sati (na 13,8(2%) i 72 sata kultiviranja, kada pada na 7,7(0,9% kontrolne razine. Nakon 1 sat kultiviranja stanica THP-1 uz hidrokortizon koncentracije 1 (M koli ina mRNA za galektin-3 se smanjuje na 44,2(8,8% po etne razine i sljedea etiri sata samo pomalo pada (trei sat iznosi 42,9(2,2%, a peti sat 39,3(2,6%). Jednodnevno kultiviranje uzrokuje pad ekspresije LGALS3 na 26,2(0,2%, dvodnevno na 22,2(1,3% a trodnevno na samo 8,3(0,6% po etne razine. Nakon 72 sata kultiviranja uz hidrokortizon koncentracije 5 (M razina LGALS3 iznosi samo 5,4(1,9% po etne razine, a dostignuta je postupnim smanjenjem ekspresije: nakon 1 sat - na 53,5(6%, 3 sata - na 42(2,5%, 5 sati - na 42,9(17,8% 24 sata - na 17,9(4,8% i nakon 48 sati na 11(3,2% po etne razine. Od svih primijenjenih koncentracija hidrokortizona, najja e inhibitorno djelovanje na ekspresiju LGALS3 ima 10 (M hidrokortizon. On inhibira ekspresiju LGALS3 za ~60% tijekom prvih pet sati kultiviranja (nakon prvog sata koli ina mRNA za galektin-3 iznosi 39(1,5%, nakon treeg 36,6(2,7%, a nakon petog 35,6(6,6% po etne razine). Daljnjim kultiviranjem koli ina mRNA za galektin-3 nastavlja padati i iznosi: 24h-21,5(3,5%, 48h-9,1(2% i 72 h-3,8(0,4% po etne razine.  Slika 3.11 Hidrokortizon inhibira ekspresiju LGALS3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz hidrokortizon koncentracije 0,1 (M, 1 (M, 5 (M i 10 (M. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Crni stupci  kontrola, rebrasti stupci-0,1 (M HC, tamnozeleni stupci  1 (M HC, svjetlozeleni stupci  5 (M HC, bijeli stupci  10 (M HC. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz hidrokortizon koncentracija 0,1 (M, 1 (M, 5 (M i 10 (M ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka (Slika 3.12). Tijekom prva 24 sata kultiviranja stanica THP-1 uz hidrokortizon koncentracija 0,1 (M, 1 (M, 5 (M i 10 (M koli ina galektina-3 u odnosu na kontrolu ostaje nepromijenjena. Nakon 48-satnog kultiviranja razina se ekspresije galektina-3 smanjuje ~40% i iznosi, ovisno o primijenjenim koncentracijama hidrokortizona: 63,3(4,9% (uz 0,1 (M HC), 62,9(3,3% (uz 1 (M HC), 59,9(5,1% (uz 5 (M HC) i 58,8(3,9% po etne razine (uz 10 (M HC). Kultiviranjem tijekom 72 sata koli ina galektina-3 smanjuje se na pribli~no polovinu i iznosi: 53,7(5,2% (uz 0,1 (M HC), 54(4,1% (uz 1 (M HC), 50,3(3,8% (uz 5 (M HC) i 42,3(6,2% po etne razine (uz 10 (M HC).   Slika 3.12 Utjecaj hidrokortizona na ekspresiju galektina-3 u stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/0,1 (M HC; b) mediju/1 (M HC; c) mediju/5 (M HC; d) mediju/10 (M HC (5 g proteina/ja~ici). e) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x ( SD). Crni stupac  kontrola, rebrasti stupci-0,1 (M HC, tamnozeleni stupci  1 (M HC, svjetlozeleni stupci  5 (M HC, bijeli stupci  10 (M HC. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). 3.5 Utjecaj deksametazona na ekspresiju LGALS3 i galektina-3 u stanicama THP-1 Deksametazon, steroidni imunomodulatorni lijek utje e na brojne puteve prijenosa signala u stanici. Jedan od njih je signalni put protein kinaza aktiviranih mitogenima (Ras/MEKK1/MKK1) za koji je pokazano da sudjeluje u regulaciji ekspresije galektina-3. Stoga smo u ovom radu ~eljeli ispitati u inak deksametazona na ekspresiju LGALS3 i galektina-3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u hranidbenom mediju uz 0,1 nM, 1 nM, 10 nM ili 100 nM deksametazon, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Postotak ~ivih stanica odreen je metodom bojanja tripanskim modrilom (opisanom u poglavlju 2.2.7). Rezultati su pokazali da je postotak ~ivih stanica u svim ispitivanim slu ajevima e"95%, odnosno, da deksametazon u primijenjenim koncentracijama ne utje e na vijabilnost stanica THP-1 tijekom 72-satnog kultiviranja. Metaboli ka aktivnost stanica uz primijenjene koncentracije deksametazona i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da se metaboli ka aktivnost stanica kultiviranih uz deksametazon koncentracija 1 nM, 1 nM, 10 nM i 100 nM ne mijenja u odnosu na metaboli ku aktivnost stanica kultiviranih samo u hranidbenom mediju. Deksametazon u primijenjenih koncentracijama ne djeluje citotoksi no na stanice THP-1 obzirom da je u svim slu ajevima metaboli ka aktivnost bila e"95,4% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Relativna razina ekspresije LGALS3 odreena je nakon izolacije ukupne RNA, reverzne transkripcije i lan ane reakcije polimerazom na ureaju AbiPrism 310 Genetic Analyser. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni istim postupkom u stanicama THP-1 kultiviranih tijekom istih vremena u mediju/0,2% DMSO (Slika 3.13). Deksametazon koncentracije 0,1 nM uzrokuje smanjenje razine ekspresije LGALS3 na 33,4(1,4% po etne razine ve nakon prvog sata kultiviranja. Daljnjim kultiviranjem do 24. sata ne dolazi do zna ajnije promjene ekspresije (nakon 3 sata iznosi 30,2(5,3%, nakon 5 sati 38,9(0,8%, a nakon 24 sata kultiviranja 32,3(0,1% po etne razine), da bi nakon 48 sati pala na 22,2(3,5% a nakon 72 sata kultiviranja na 15,4(4,6% po etne razine. Sli an u inak ima i deksametazon koncentracije 1 nM: nakon prvog sata kultiviranja koli ina mRNA za galektin-3 se smanji na 30,7(4,7% po etne razine i odr~ava se takvom i nakon 3 sata (31,9(6,3%), 5 sati (34,5(3,1%) i 24 sata inkubiranja (26,1(10% po etne razine). Nakon 48 sati koli ina mRNA za galektin-3 se ponovno zna ajnije smanjuje (na 14,7(0,2%) da bi nakon 72-satnog kultiviranja iznosila samo 9,9(1,7% po etne razine. Pad koli ine mRNA za galektin-3 za 70% (na 28,7(8,3% po etne razine) unutar prvog sata inkubiranja uzrokuje i deksametazon koncentracije 10 nM. Trosatno kao ni peterosatno kultiviranje ne uzrokuje daljnju promjenu ekspresije LGALS3 (nakon 3 sata iznosi 31,4(1,9%, a nakon 5 sati 32,2(2,6% po etne razine). Daljnji pad razine primjeuje se nakon 24 sata (na 18,8(3,1%), nakon 48 sati (na 12,3(2,4%) i 72 sata kultiviranja (na 9,7(5,2% po etne razine). Deksametazon koncentracije 100 nM mijenja ekspresiju LGALS3 na sli an na in i u sli noj mjeri kao ato je primjeeno i kod deksametazona 100 i 1000 puta manje koncentracije. Nakon ato se koli ina mRNA za galektin-3 smanji za ~70% nakon prvog sata kultiviranja (na 34,9(0,3%), odr~ava se na pribli~no jednakoj razini i nakon 3 sata (28,9(2,9%), 5 sati (35,8(12,7%) i 24 sata kultiviranja (kada iznosi 31,8(9,5% po etne razine). Meutim, nakon 48 sati iznosi 13,8(2,9%, a nakon 72 sata samo 10,8(5,5% po etne razine.  Slika 3.13 Deksametazon inhibira ekspresiju LGALS3. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz deksametazon koncentracije 0,1 nM, 1 nM, 10 nM i 100 nM. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Crni stupci  kontrola, rebrasti stupci-0,1 nM Dex, tamnozeleni stupci  1 nM Dex, svjetlozeleni stupci  10 nM Dex, bijeli stupci  100 nM Dex. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz deksametazon koncentracija 0,1 nM, 1 nM, 10 nM i 100 nM ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka (Slika 3.14). Obrazac promjene ekspresije galektina-3 uzrokovane djelovanjem deksametazona sli an je kod svih primijenjenih koncentracija deksametazona. Nakon prvog sata kultiviranja uo ava se blagi pad razine galektina-3 (za ~15%), no ta promjena nije statisti ki zna ajna. Zatim se daljim kultiviranjem koli ina galektina-3 odr~ava na pribli~no jednakoj razini da bi se nakon 24-satnog kultiviranja ponovno vratila na po etnu razinu. Meutim, nakon 48-satnog kultiviranja razina se ekspresije galektina-3 smanjuje 30-50% i iznosi, ovisno o primijenjenim koncentracijama deksametazona: 71(5,1% (0,1 nM Dex), 62,4(3,8% (1 nM Dex), 59,7(6,7% (10 nM Dex) i 51,1(2,8% (100 nM Dex). Daljnjim kultiviranjem koli ina galektina-3 se nastavlja smanjivati te nakon 72 sata kultiviranja iznosi: 62,3(4,3% (0,1 nM Dex), 51,7(7,4% (1 nM Dex), 38,7(5% (10 nM Dex) i 38,3(4,7% po etne razine (100 nM Dex).    Slika 3.14 Utjecaj deksametazona na ekspresiju galektina-3 u stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/0,1 nM Dex; b) mediju/1 nM Dex; c) mediju/10 nM Dex; d) mediju/100 nM Dex (5 g proteina/ja~ici). e) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x ( SD). Crni stupac  kontrola, rebrasti stupci-0,1 nM Dex, tamnozeleni stupci  1 nM Dex, svjetlozeleni stupci  10 nM Dex, bijeli stupci  100 nM Dex. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). 3.6 Ekspresija LGALS3 i galektina-3 tijekom diferencijacije stanica THP-1 pomou PMA Izlaganje stanica monocitne stani ne linije THP-1 djelovanju forbol-12-miristat-13-acetata uzrokuje diferencijaciju stanica THP-1 pri emu one poprimaju karakteristike makrofaga. Stanice THP-1 rastu u suspenziji, a nakon izlaganja djelovanju PMA adheriraju na podlogu. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24 ili 48 sati u 10-7 M otopini forbol-12-miristat-13-acetata (engl. phorbol-12-myristate-13-acetate, PMA) u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila <10-5 %. Postotak neadheriranih stanica u mediju odreen je brojanjem stanica pomou hemocitometra. Rezultati su pokazali da je ve nakon sat vremena izlaganja stanica THP-1 PMA koncentracije 10-7 M adheriralo 80,2(4,1% stanica. Daljnjim kultiviranjem tijekom 3 i 5 sati postotak adheriranih stanica ostaje pribli~no isti (80,7(2,5%, odnosno 84,1(3,1%) da bi se nakon 24 sata dodatno poveao i iznosio 97,7(2,8%. Nakon 48 sati izlaganja stanica THP-1 PMA koncentracije 10-7 M postotak adheriranih stanica je vei od 99%. Nakon izolacije RNA iz stanica THP-1 tretiranih 10-7 M PMA i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u stanicama THP-1 prije tretmana s forbol-12-miristat-13-acetatom. Tijekom prva tri sata kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M koli ina mRNA za galektin-3 ostaje ista. Nakon jednodnevnog kultiviranja raste 2,4 puta (241(7%) a nakon dvodnevnog gotovo 3 puta (292(5%) u odnosu na po etnu koli inu mRNA.              Slika 3.15 PMA inducira ekspresiju LGALS3 u stanicama THP-1. Stanice THP-1 kultivirane su tijekom 1, 3, 5, 24 ili 48 sati uz PMA koncentracije 10-7 M. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Crni stupac  kontrola, bijeli stupci  10-7 M PMA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u stanicama stani ne linije THP-1 tijekom 48-satnog kultiviranja uz PMA koncentracije 10-7 M ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24 ili 48 sati. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka. Izlaganje stanica THP-1 forbol-12-miristat-13-acetatu koncentracije 10-7 M tijekom 1, 3 ili 5 sati ne utje e na razinu ekspresije galektina-3. Meutim, koli ina galektina-3 se nakon 24-satnog kultiviranja uz PMA koncentracije 10-7 M poveava gotovo 3 puta (292(9%), a nakon 48-satnog kultiviranja 3,5 puta (346(8%) u odnosu na po etnu razinu galektina-3 (Slika 3.16).    Slika 3.16 PMA inducira ekspresiju galektina-3 u stanicama THP-1. a) Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 kultiviranih tijekom 1, 3, 5, 24 ili 48 sati u mediju/10-7 M PMA (5 g proteina/ja~ici). b) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu (x ( SD). Crni stupac  kontrola, bijeli stupci  10-7 M PMA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). 3.7 Ekspresija LGALS3 i galektina-3 u diferenciranim stanicama THP-1 tijekom kultiviranja S ciljem odreivanja vijabilnosti i metaboli ke aktivnosti diferenciranih stanica THP-1, nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M medij je odliven, te su tako dobivene diferencirane stanice THP-1 dodatno kultivirane tijekom 24, 48 ili 72 sata u mediju ili mediju/0,2% DMSO. Pokusi su ponovljeni 2 puta u kvadriplikatu. Metaboli ka aktivnost stanica uz 0,2% DMSO i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da prisutnost 0,2% DMSO ne djeluje citotoksi no na diferencirane stanice stani ne linije THP-1 niti nakon 72 sata kultiviranja te je njihova metaboli ka aktivnost nepromijenjena (98,4(3,4%) u odnosu na metaboli ku aktivnosti stanica kultiviranih u hranidbenom mediju. Relativna razina ekspresije gena LGALS3, odnosno relativna koli ina mRNA za galektin-3 tijekom kultiviranja (nakon 1, 3, 5, 24, 48 ili 72 sata) diferenciranih stanica stani ne linije THP-1 u mediju ili mediju/0,2% DMSO odreena je nakon izolacije RNA i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom. Koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u stanicama THP-1 nakon 48-satnog kultiviranja uz PMA koncentracije 10-7 M (Slika 3.17). Rezultati su pokazali da se razina ekspresije LGALS3 ne mijenja tijekom 72-satnog kultiviranja diferenciranih stanica THP-1 u mediju, kao niti tijekom 72-satnog kultiviranja diferenciranih stanica THP-1 u mediju/0,2% DMSO u odnosu na po etnu razinu ekspresije LGALS3, odnosno na razinu ekspresije u stanicama THP-1 kultiviranih tijekom 48 sati uz PMA koncentracije 10-7 M.  Slika 3.17 U diferenciranim stanicama THP-1 tijekom 72-satnog kultiviranja u mediju ili u mediju/0,2% DMSO koli ina mRNA za galektin-3 se ne mijenja. Stanice THP-1 diferencirane 48-satnim kultiviranjem uz PMA koncentracije 10-7 M kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata u mediju ili u mediju/0,2% DMSO. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Bijeli stupac  kontrola, rebrasti stupci-kultiviranje u mediju, zeleni stupci  kultiviranje u mediju/0,2% DMSO. Razina ekspresije galektina-3 u diferenciranim stanicama stani ne linije THP-1 ispitana je tijekom 72-satnog kultiviranja postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena ekspresija konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat stanica THP-1 koje su kultivirane u hranidbenom mediju. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka. Rezultati su pokazali da se koli ina galektina-3 ne mijenja tijekom 1, 3, 5, 24, 48 i 72 sata kultiviranja stanica THP-1 u mediju u odnosu na po etnu koli inu galektina-3 u diferenciranim stanicama THP-1 (odnosno, nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M). Razina ekspresije galektina-3 ostaje ista i kada se stanice kultiviraju tijekom tijekom 1, 3, 5, 24, 48 i 72 sata u mediju/0,2% DMSO, te je koli ina galektina-3 i dalje oko 3,5 puta vea od koli ine galektina-3 u nediferenciranim stanicama THP-1 (Slika 3.18).    Slika 3.18 Koli ina galektina-3 ne mijenja se u diferenciranim stanicama THP-1 tijekom kultiviranja u mediju ili u mediju/0,2% DMSO. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1 koje su nakon 48-satnog kultiviranja uz PMA koncentracije 10-7 M kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju i b) mediju/0,2% DMSO (5 g proteina/ja~ici). c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu 2 (x (SD). Crni stupac  kontrola 1 (nediferencirane stanice THP-1), bijeli stupac  kontrola 2 (diferencirane stanice THP-1 nakon 48-satnog kultiviranja uz 10-7 M PMA), rebrasti stupci-kultiviranje u mediju, zeleni stupci  kultiviranje u mediju/0,2% DMSO. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05). 3.8 Utjecaj aspirina na ekspresiju LGALS3 i galektina-3 u diferenciranim stanica THP-1 Nakon 48-satnog kultiviranja stanica THP-1 uz forbol-12-miristat-13-acetat koncentracije 10-7 M, medij koji sadr~i PMA je odliven, te su diferencirane stanice THP-1 kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u 0,5 mM i 2 mM otopini aspirina u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Metaboli ka aktivnost stanica i citotoksi nost primijenjenih koncentracija aspirina odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da 0,5 mM i 2 mM otopina aspirina statisti ki zna ajno ne mijenjaju metaboli ku aktivnost stanica u odnosu na diferencirane stanice kultivirane u hranidbenom mediju. Primjenjene koncentracije aspirina nemaju citotoksi an u inak na diferencirane stanice THP-1 obzirom da je u svim slu ajevima metaboli ka aktivnost bila e"87% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Nakon izolacije RNA iz tretiranih stanica i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u diferenciranim stanicama THP-1 kultiviranim tijekom istih vremena u mediju/0,2% DMSO (Slika 3.19). Kultiviranje diferenciranih stanica THP-1 uz aspirin koncentracije 0,5 mM uzrokuje smanjenje koli ine mRNA za galektin-3 na 65,4(5,2% nakon prvog sata kultiviranja, a nakon treeg sata na 87,1(3,5%. Nakon pet sati kultiviranja koli ina mRNA za galektin-3 pribli~no je jednaka po etnoj koli ini, te se daljnjim kultiviranjem ne mijenja. Aspirin koncentracije 2 mM uzrokuje smanjenje ekspresije LGALS3 na 46,7(7,3% po etne vrijednosti nakon prvog sata kultiviranja. Daljnjim kultiviranjem poveava se koli ina mRNA za galektin-3, ali je nakon tri sata joa uvijek manja od po etne vrijednosti (82,1(4,2%). Peti sat pribli~ava se po etnoj vrijednosti (92,3(4,3; p>0,05), koju dose~e nakon 24-satnog kultiviranja. Uo ava se daljnji porast koli ine mRNA za galektin-3 nakon 48 sati (115(7,2) i 72 sata kultiviranja (112,3(8,1), premda te promjene nisu statisti ki zna ajne.  Slika 3.19 Utjecaj aspirina na ekspresiju LGALS3 u diferenciranim stanicama THP-1. Nakon 48-satnog kultiviranja stanica THP-1 uz PMA (10-7 M), diferencirane stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz aspirin koncentracije 0,5 mM i 2 mM. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Bijeli stupci  kontrola, rebrasti stupci  0,5 mM ASA, zeleni stupci  2 mM ASA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05). Razina ekspresije galektina-3 u diferenciranim stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz aspirin koncentracija 0,5 mM i 2 mM ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani 1, 3, 5, 24, 48 ili 72 sata nakon dodatka medija s aspirinom. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat nediferenciranih stanica THP-1 te stani ni homogenat diferenciranih stanica THP-1, dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7M. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka 2 (diferenciranih stanica THP-1 dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7M) (Slika 3.20). Tijekom 72-satnog kultiviranja diferenciranih stanica THP-1 uz aspirin koncentracije 0,5 mM razina galektina-3 se ne mijenja. U diferenciranim stanicama THP-1 izlo~enim djelovanju aspirina koncentracije 2 mM nakon prvog sata kultiviranja koli ina galektina-3 statisti ki se zna ajno smanji za pribli~no 30% (71,2(5,5% po etne razine. Zatim raste te je nakon 3 sata neato manja od po etne razine (93,3(7%, p>0,05), a nakon 5 sati odgovara po etnoj razini. Nakon 24-satnog kultiviranja po inje se poveavati (111,0(9,0%, p>0,05), nakon 48 sati iznosi 124(5,1%, a nakon 72 sata kultiviranja uz aspirin koncentracije 2 mM iznosi 125(7,2% po etne razine.    Slika 3.20 Utjecaj aspirina na ekspresiju galektina-3 u diferenciranim stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1, koje su nakon 48-satnog kultiviranja uz PMA (10-7 M) kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/0,5 mM ASA; b) mediju/2 mM ASA (5 g proteina/ja~ici); c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu 2 (x (SD). Crni stupac  kontrola 1 (nediferencirane stanice THP-1), bijeli stupac  kontrola 2 (diferencirane stanice THP-1 nakon 48-satnog kultiviranja uz 10-7 M PMA), rebrasti stupci  0,5 mM ASA, zeleni stupci  2 mM ASA. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05). 3.9 Utjecaj indometacina na ekspresiju LGALS3 i galektina-3 u diferenciranim stanica THP-1 Nakon 48-satnog kultiviranja stanica THP-1 uz forbol-12-miristat-13-acetat koncentracije 10-7 M, medij koji sadr~i PMA je odliven te su diferencirane stanice THP-1 kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u 1 M i 10 M otopini indometacina u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Metaboli ka aktivnost stanica uz primijenjene koncentracije indometacina i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da 1 M i 10 M otopine indometacina statisti ki zna ajno ne mijenjaju metaboli ku aktivnost stanica u odnosu na diferencirane stanice kultivirane u hranidbenom mediju. Primjenjene koncentracije indometacina nemaju citotoksi an u inak na diferencirane stanice THP-1 obzirom da je u svim slu ajevima metaboli ka aktivnost bila e"94% metaboli ke aktivnosti stanica kultiviranih u hranidbenom mediju. Nakon izolacije RNA iz tretiranih stanica i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u diferenciranim stanicama THP-1 kultiviranim tijekom istih vremena u mediju/0,2% DMSO (Slika 3.21). Nakon prvog sata kultiviranja diferenciranih stanica THP-1 uz indometacin koncentracije 1 M ekspresija LGALS3 smanjena je na 71,4(3,8%, ali se ve nakon petog sata ponovno uspostavlja razina ekspresije jednaka po etnoj. Ona se daljnjim kultivirajem tijekom 24, 48 i 72 sata ne mijenja. Kultiviranje diferenciranih stanica THP-1 tijekom 1, 3, 5, 24, 48 i 72 sata uz indometacin koncentracije 10 M ne utje e na razinu ekspresije LGALS3. Ona je pribli~no jednaka po etnoj razini tijekom cijelog perioda inkubiranja, premda se nakon prvog sata mo~e uo iti blago smanjenje koli ine mRNA za galektin-3 (88,9(7,3; p>0,05).  Slika 3.21 Utjecaj indometacina na ekspresiju LGALS3 u diferenciranim stanicama THP-1. Nakon 48-satnog kultiviranja stanica THP-1 uz PMA (10-7 M), diferencirane stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz indometacin koncentracije 1 M i 10 M. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Bijeli stupci  kontrola, rebrasti stupci  1 M Ind, zeleni stupci  10 M Ind. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u diferenciranim stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz indometacin koncentracija 1 M i 10 M ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat nediferenciranih stanica THP-1 te stani ni homogenat diferenciranih stanica THP-1, dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka 2 (diferenciranih stanica THP-1 dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7M) (Slika 3.22). Koli ina galektina-3 u diferenciranim stanicama THP-1 kultiviranim uz indometacin koncentracije 1 M tijekom prvih pet sati statisti ki se zna ajno ne mijenja. Zatim se koli ina galektina-3 poveava te iznosi 123,3(4,7% nakon 24 sata kultiviranja te 120,2(5,8% nakon 48 sati kultiviranja, da bi nakon 72-satnog kultiviranja ponovno pala na po etnu vrijednost (95,5(4,4%). Kultiviranje diferenciranih stanica THP-1 uz indometacin koncentracije 10 M tijekom pet sati ne utje e na razinu ekspresije galektina-3. Jednodnevno pak kultiviranje podi~e razinu ekspresije galektina-3 na 122,3(4% po etne razine, dvodnevno na 137,2(5,6%, a nakon trodnevnog kultiviranja koli ina galektina-3 ponovno je jednaka po etnoj koli ini (107,1(3,3%).    Slika 3.22 Utjecaj indometacina na ekspresiju galektina-3 u diferenciranim stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1, koje su nakon 48-satnog kultiviranja uz PMA koncentracije 10-7 M kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/1 M Ind; b) mediju/10 M Ind (5 g proteina/ja~ici); c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu 2 (x ( SD). Crni stupac  kontrola 1 (nediferencirane stanice THP-1), bijeli stupac  kontrola 2 (diferencirane stanice THP-1 nakon 48-satnog kultiviranja uz 10-7 M PMA), rebrasti stupci  1 M Ind, sivi stupci  10 M Ind. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05); dvije zvjezdice ozna avaju statisti ki zna ajnu razliku u odnosu na 48. sat. 3.10 Utjecaj hidrokortizona na ekspresiju LGALS3 i galektina-3 u diferenciranim stanica THP-1 Nakon 48-satnog kultiviranja stanica THP-1 uz forbol-12-miristat-13-acetat koncentracije 10-7 M, medij koji sadr~i PMA je odliven, te su diferencirane stanice THP-1 kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u 1 M i 10 M otopini hidrokortizona u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Metaboli ka aktivnost stanica uz primijenjene koncentracije hidrokortizona i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da hidrokortizon koncentracije 1 M uzrokuje lagano poveanje metaboli ke aktivnosti diferenciranih stanica THP-1 (118,6(4,4% kontrolne vrijednosti). Hidrokortizon koncentracije 10 M uzrokuje poveanje metaboli ke aktivnosti diferenciranih stanica THP-1 na 124,9(9,2% kontrolne vrijednosti. Nakon izolacije RNA iz tretiranih stanica i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u diferenciranim stanicama THP-1 kultiviranim tijekom istih vremena u mediju/0,2% DMSO (Slika 3.23). U diferenciranim stanicama THP-1 razina ekspresije LGALS3 smanjuje se na 46,7(2,3% nakon jednog sata kultiviranja uz hidrokortizon koncentracije 1 M. Sli an u inak uo ava se i nakon tri sata kultiviranja (51,3(7,2%), da bi se nakon petog sata razina ekspresije LGALS3 vratila na po etnu vrijednost (95,1(3,5%). Premda daljnjim kultiviranjem tijekom 24, 48 i 72 sata koli ina mRNA za galektin-3 pokazuje trend blagog rasta te promjene u usporedbi s koli inom u kontrolnim uzorcima nisu statisti i zna ajne (11(5,2%; 108(6,2% odnosno 112(5,2%). Hidrokortizon koncentracije 10 M uzrokuje pad koli ine mRNA za galektin-3 na 55,4(6,2% po etne vrijednosti nakon 1 sata kultiviranja. Sli an se u inak uo ava i nakon 3 sata (52,3(4,8%) i pet sati kultiviranja, kada razina ekspresije LGALS3 iznosi 57,1(3,3% po etne vrijednosti. Potom se koli ina mRNA za galektin-3 poveava te nakon 24 sata iznosi 82,1(2,9% po etne vrijednosti, da bi nakon 48 sati, kao i nakon 72 sata kultiviranja dosegla po etnu razinu.  Slika 3.23 Utjecaj hidrokortizona na ekspresiju LGALS3 u diferenciranim stanicama THP-1. Nakon 48-satnog kultiviranja stanica THP-1 uz PMA (10-7 M), diferencirane stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz hidrokortizon koncentracije 1 M i 10 M. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Bijeli stupci  kontrola, rebrasti stupci  1 M HC, zeleni stupci  10 M HC. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05); dvije zvjezdice ozna avaju statisti ki zna ajnu razliku u odnosu na 24. sat. Razina ekspresije galektina-3 u diferenciranim stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz hidrokortizon koncentracija 1 M i 10 M ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat nediferenciranih stanica THP-1 te stani ni homogenat diferenciranih stanica THP-1, dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka 2 (diferenciranih stanica THP-1 dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7M) (Slika 3.24). Smanjenje koli ine galektina-3 za ~40% uo eno je nakon jednog sata kultiviranja stanica THP-1 uz hidrokortizon koncentracije 1 M. Tada iznosi 58,3(7,2% po etne razine. Daljnje kultiviranje uzrokuje lagani porast koli ine galektina-3  nakon treeg sata koli ina galektina-3 je joa uvijek manja od po etne razine, ali samo za ~20% (79,8(3,1% po etne razine). Do petog sata koli ina galektina-3 zna ajno raste (za ~50%) u odnosu na kontrolu, odnosno za 82% od razine dostignute nakon 3 sata (145,0(4,8% po etne razine). Nakon 24 sata razina galektina-3 iznosi 137,0(6,1%, a nakon 48 sati 138(5,5% po etne vrijednosti. Nakon 72-satnog kultiviranja uz hidrokortizon koncentracije 1 M, diferencirane stanice THP-1 eksprimiraju 121,0(2,9% po etne koli ine galektina-3. Hidrokortizon koncentracije 10 M takoer uzrokuje pad razine ekspresije galektina-3 u diferenciranim stanicama THP-1 tijekom prvih pet sati kultiviranja. Nakon jednog sata ona iznosi 65,0(4,4%, nakon treeg 67,2(3,2%, a nakon petog sata kultiviranja 69,1(5,3% po etne razine. Meutim, nakon 24 sata kultiviranja ponovno se uspostavlja razina ekspresije uo ena kod kontrolnih uzoraka (112,0(6,1%), ato zna i da je u odnosu na 5. sat doalo do statisti ki zna ajnog porasta razine galektina-3 za ~60%. Daljnjim kultiviranjem (48 i 72 sata) razina galektina-3 ostaje nepromijenjena.    Slika 3.24 Utjecaj hidrokortizona na ekspresiju galektina-3 u diferenciranim stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1, koje su nakon 48-satnog kultiviranja uz PMA koncentracije 10-7 M kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/1 M HC; b) mediju/10 M HC (5 g proteina/ja~ici); c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu 2 (x ( SD). Crni stupac  kontrola 1 (nediferencirane stanice THP-1), bijeli stupac  kontrola 2 (diferencirane stanice THP-1 nakon 48-satnog kultiviranja uz 10-7 M PMA), rebrasti stupci  1 M HC, zeleni stupci  10 M HC. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05). dvije zvjezdice ozna avaju statisti ki zna ajnu razliku u odnosu na 5. sat. 3.11 Utjecaj deksametazona na ekspresiju LGALS3 i galektina-3 u diferenciranim stanica THP-1 Nakon 48-satnog kultiviranja stanica THP-1 uz forbol-12-miristat-13-acetat koncentracije 10-7 M, medij koji sadr~i PMA je odliven, te su diferencirane stanice THP-1 kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u 1 nM i 100 nM otopini deksametazona u hranidbenom mediju, u kojem je istovremeno koncentracija DMSO bila d"0,2%. Metaboli ka aktivnost stanica uz primijenjene koncentracije deksametazona i citotoksi nost odreeni su testom MTS (prema postupku opisanom u poglavlju 2.2.8). Rezultati su pokazali da deksametazon koncentracije 1 nM ne djeluje citotoksi no na diferencirane stanice THP-1; ini se da se njihova metaboli ka aktivnost ak lagano poveava (na 111(5,5% kontrolne vrijednosti, p>0,05). Izlaganje diferenciranih stanica THP-1 deksametazonu koncentracije 100 nM poveava njihovu metaboli ku aktivnost na 127(8,3% kontrolne razine. Nakon izolacije RNA iz tretiranih stanica i reverzne transkripcije po emu je slijedila lan ana reakcija polimerazom, koli ina PCR produkta nastalih umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR odreena je analizom na ureaju AbiPrism 310 Genetic Analyser uz primjenu programa GeneScan 2.1 software. Relativna koli ina mRNA za galektin-3 izra~ena je omjerom povraine peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu. Kontrolom su smatrani omjeri povraine peakova PCR produkata galektina-3 i -aktina dobiveni opisanim na inom u diferenciranim stanicama THP-1 kultiviranim tijekom istih vremena u mediju/0,2% DMSO (Slika 3.25). Rezultati su pokazali da tijekom prvog sata kultiviranja diferenciranih stanica THP-1 uz deksametazon koncentracije 1 nM dolazi do smanjenja koli ine mRNA za galektin-3 na 82,3(3,1%, ali se ve nakon tri sata kultiviranja ponovno uspostavlja po etna razina ekspresije LGALS3 (95,4(2,2%). Daljnjim kultiviranjem (tijekom 24, 48, 72 sata) razina ekspresije odr~ava se na pribli~no jednakoj razini. Gotovo identi an u inak na ekspresiju gena za galektin-3 ima i deksametazon koncentracije 100 nM. I on u diferenciranim stanicama THP-1 uzrokuje smanjenje koli ine mRNA za galektin-3 na 83(4,3% nakon jednog sata kultiviranja. Nakon 5 sati kultiviranja razina ekspresije LGALS3 je samo neznatno manja od po etne razine (94(4,8%), koja ostaje nepromijenjena nakon 24, 48 i 72 sata kultiviranja.  Slika 3.25 Utjecaj deksametazona na ekspresiju LGALS3 u diferenciranim stanicama THP-1. Nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M, diferencirane stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz deksametazon koncentracije 1 nM i 100 nM. Nakon izolacije ukupne RNA i reverzne transkripcije, PCR produkti nastali umna~anjem dijela mRNA za galektin-3 tijekom 21 ciklusa PCR i dijela mRNA za -aktin tijekom 14 ciklusa PCR su zdru~eni i razdvojeni kapilarnom elektroforezom na ureaju AbiPrism 310 Genetic Analyser. Rezultati dvaju neovisnih pokusa napravljenih u duplikatu izra unati primjenom programa GeneScan 2.1 software izra~eni su kao omjeri povraina peakova PCR produkata galektina-3 i -aktina u odnosu na kontrolu (x (SD). Bijeli stupci  kontrola, rebrasti stupci  1 nM Dex, zeleni stupci  100 nM Dex. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu (p<0,05). Razina ekspresije galektina-3 u diferenciranim stanicama stani ne linije THP-1 tijekom 72-satnog kultiviranja uz deksametazon koncentracija 1 nM i 100 nM ispitana je postupkom Western blot uz denzitometrijsku analizu vrpci. Uzorci su sakupljani nakon 1, 3, 5, 24, 48 ili 72 sata. U svakom je uzorku istim postupkom odreena koli ina konstitutivno eksprimiranog proteina - -aktina. Rezultati denzitometrijske analize vrpci galektina-3 normirani su prema rezultatima dobivenim za vrpce -aktina (za svaki pojedini uzorak). Kako bismo mogli usporeivati rezultate dobivene za uzorke analizirane na razli itim gelovima/membranama na svaki je gel nanijet isti stani ni homogenat nediferenciranih stanica THP-1 te stani ni homogenat diferenciranih stanica THP-1, dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7 M. Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina pojedinog uzorka u odnosu na relativnu povrainu vrpci galektina-3/-aktina kontrolnog uzorka 2 (diferenciranih stanica THP-1 dobivenih nakon 48-satnog kultiviranja stanica THP-1 uz PMA koncentracije 10-7M) (Slika 3.26). Tijekom prvog sata kultiviranja diferenciranih stanica THP-1 uz deksametazon koncentracije 1 nM te 100 nM ekspresija galektina-3 se ne mijenja. U stanicama kultiviranim uz deksametazon koncentracije 1 nM ve se trei sat kultiviranja ekspresija galektina-3 poveava za >50% (155,2(4,5%), a zatim joa raste te iznosi: 5 sat-172,4(5,8%, 24 sat-173,2(7,1%, 48 sat  170,0(5,3%. Nakon 72-satnog kultiviranja iznosi 154,1(2,8% po etne razine. Sli an u inak na ekspresiju galektina-3 ima i deksametazon koncentracije 100 nM. Premda nakon tri sata kultiviranja nema promjene ekspresije galektina-3, peti sat koli ina galektina-3 iznosi 164,3(7,2% po etne razine te nastavlja rasti, pa 24. sat iznosi 184,5(5,8%, a 48. sat 185,2(4,6%. Nakon 72-satnog kultiviranja uz deksametazon koncentracije 100 nM iznosi 158,4(5,4% po etne koli ine.    Slika 3.26 Utjecaj deksametazona na ekspresiju galektina-3 u diferenciranim stanicama THP-1. Imunoblot analiza ekspresije galektina-3 u homogenatima stanica THP-1, koje su nakon 48-satnog kultiviranja uz PMA koncentracije 10-7 M kultivirane tijekom 1, 3, 5, 24, 48 ili 72 sata u a) mediju/1 nM Dex; b) mediju/100 nM Dex (5 g proteina/ja~ici); c) Rezultati denzitometriranja vrpci iz dva neovisna pokusa napravljena u duplikatu izra~eni su kao relativna povraina vrpci galektina-3/-aktina u odnosu na kontrolu 2 (x ( SD). Crni stupac  kontrola 1 (nediferencirane stanice THP-1), bijeli stupac  kontrola 2 (diferencirane stanice THP-1 nakon 48-satnog kultiviranja uz 10-7 M PMA), rebrasti stupci  1 nM Dex, zeleni stupci  100 nM Dex. Zvjezdica ozna ava statisti ki zna ajnu razliku u odnosu na kontrolu 2 (p<0,05). RASPRAVA Galektin-3 je lektin 29-32 kDa, koji specifi no prepoznaje (-galaktozidne strukture glikoproteina, a ija to na fizioloaka uloga, kao ni mehanizam regulacije, do danas nisu u potpunosti razjaanjeni. Galektin-3 sudjeluje u mnogim bioloakim procesima, izmeu ostalog, ima va~nu ulogu u modulaciji imunih reakcija i upalnom odgovoru. Prisutan je u izvanstani noj tekuini, ima sposobnost aktiviranja brojnih stanica, posreduje u meustani nim interakcijama te interakcijama izmeu stanica i izvanstani nog matriksa, kao i homotipskim interakcijama izmeu stanica  sve to ini ga pogodnom molekulom za regulaciju upalnog odgovora. Osim toga, galektin-3 mo~e inhibirati apoptozu, te stoga ekspresija galektina-3 mo~e produljiti opstanak upalnih stanica, a samim time i upalni odgovor. Rezultati in vitro istra~ivanja pokazali su da galektin-3 djeluje kao jak proinflamatorni signal koji modulira stani nu proliferaciju, stani nu adheziju, kemotaksiju, fagocitozu, aktivaciju mastocita i bazofila, sintezu upalnih medijatora te apoptozu limfocita T. Upalni odgovor je visokoreguliran fizioloaki proces koji je od klju ne va~nosti za odr~avanje homeostaze. Precizna fizioloaka kontrola upale omoguava brzu reakciju organizma na infekciju ili ozljedu bez pogubnih posljedica za organizam uzrokovanih pretjeranom biokemijskom reakcijom. Signalni putevi kojima se regulira upalni proces su kaskade protein kinaza aktiviranih mitogenima kojima se u kona nici moduliraju aktivnosti brojnih transkripcijskih faktora. Na te signalne puteve, kao i izravno na transkripcijske faktore, utje u i tvari s imunomodulatornim djelovanjem  glukokortikoidi i nesteroidne tvari s imunomodulatornim djelovanjem (NSAIDs). Najzna ajniji transkripcijski faktori, klju ni za indukciju gena uklju enih u upalni odgovor te za nastanak i progresiju bolesti uzrokovanih kroni nom aktivacijom imunog sustava (kao ato su primjerice astma i ateroskleroza te autoimune bolesti, primjerice multipla skleroza i reumatoidni artritis) su transkripcijski faktori NF-B i AP-1. Vezna mjesta za NF-B i AP-1 prisutna su i u promotorskoj regiji gena za galektin-3 (LGALS3), pa je vrlo vjerojatno da tvari s imunomodulatornim djelovanjem utje u i na ekspresiju LGALS3 i galektina-3. Zbog este uporabe ovih tvari u terapijske svrhe te va~nosti galektina-3 u fiziologiji stanica monocitno-makrofagne loze, u ovom je radu ispitan utjecaj nesteroidnih (aspirina i indometacina) i steroidnih (hidrokortizona i deksametazona) tvari s imunomodulatornim djelovanjem primijenjenih u razli itim terapijskim rasponima na ekspresiju LGALS3 i galektina-3 u stanicama monocitno-makrofagne loze. Kao prvi korak valjalo je izabrati odgovarajuu stani nu liniju. Prema literaturnim podacima, mnoga su istra~ivanja funkcije, lokalizacije i ekspresije galektina-3 provedena na stanicama promijelocitne stani ne linije HL-60  izmeu ostalog, galektin-3 po prvi je puta izoliran upravo iz tih stanica. Meutim, rezultati naaih ispitivanja (koji nisu prikazani u ovom radu) pokazali su da stanice stani ne linije HL-60 eksprimiraju izrazito malu, jedva detektabilnu koli inu LGALS3 i galektina-3. Premda naizgled za uujue, taj rezultat nije neobjaanjiv. Mnoge stani ne linije uzgajaju se u kulturi ve viae od 10 godina  stani na linija HL-60 ak gotovo 30 godina. Takve stanice esto imaju zna ajno razli ite osobine ne samo u odnosu na izvorno tkivo, ve i u odnosu na stanice iste stani ne linije uzgajane u drugim laboratorijima. Za stanice HL-60 postoje odreeni podaci o promjenama njihovih karakteristika, kao ato su sposobnost proliferacije i diferencijacije (procesi u kojima sudjeluje i galektin-3), tijekom viegodinjih kultiviranja  ADDIN EN.CITE Collins1977204027127227056351977Nov 24Continuous growth and differentiation of human myeloid leukaemic cells in suspension culture347-9Collins, S. J.Gallo, R. C.Gallagher, R. E.NatureAdultAneuploidyCell DifferentiationCell DivisionCells, CulturedCulture MediaFemaleHumanKaryotypingLeukemia, Myeloid/genetics/*pathologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=271272Macfarlane1988205031626846831988MarA phorbol ester tolerant (PET) variant of HL-60 promyelocytes291-302Department of Medicine, Veterans Administration Hospital, Iowa City, Iowa.Macfarlane, D. E.Gailani, D.Vann, K.Br J HaematolCell Differentiation/drug effectsCell LineDrug ToleranceKaryotypingLeukemia, Myelocytic, Acute/*genetics/pathologyMicroscopy, ElectronSupport, Non-U.S. Gov'tSupport, U.S. Gov't, Non-P.H.S.Support, U.S. Gov't, P.H.S.Tetradecanoylphorbol Acetate/*pharmacologyTime FactorsTumor Cells, Cultured/*drug effectsVariation (Genetics)http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=3162684[174, 175]. O igledno je promjena konstitutivne razine galektina-3 jedna od takvih promijenjenih karakteristika, barem u stanicama HL-60 na kojima smo mi proveli istra~ivanja. Kao zadovoljavajua stani na linija pokazala se monocitna stani na linija THP-1. Rezultati imunoblot analize uz primjenu specifi nog protutijela M3/38 (koje prepoznaje domenu koja sadr~i ponavljajue sljedove u N-terminalnom dijelu galektina-3) dobiveni kemifluorescentnom detekcijom pokazali su da je konstitutivna razina ekspresije galektina-3 relativno visoka. Nakon odabira odgovarajue stani ne linije, pristupilo se optimiranju metode za odreivanje razine ekspresije LGALS3, odnosno koli ine mRNA za galektin-3 (u rezultatima je prikazan samo kona an rezultat postupka optimiranja). Metoda uklju uje izolaciju ukupne RNA iz stanica, reverznu transkripciju i lan anu reakciju polimerazom te analizu koli ine PCR produkata na ureaju AbiPrism 310 Genetic Analyser uporabom programa GeneScan Analysis software. Premda izolacija RNA mo~e esto rezultirati raspadnutom RNA, kontaminacijom DNA ili nedovoljnom koli inom RNA, rezultati analiza kvalitete i koli ine izolirane RNA na ureaju Agilent 2100 bioanalyzer pokazali su da niti jedan uzorak izolirane RNA ne sadr~i primjese DNA, ve samo intaktnu RNA, relativno visoke koncentracije (analize su obavljene na svim uzorcima sakupljenim tijekom svih eksperimenata). Lan anom reakcijom polimerazom ~eljelo se umno~iti dijelove mRNA za galektin-3 i dijelove mRNA za konstitutivno eksprimirani -aktin. Prilikom odabira ishodnica, zadovoljeni su uvjeti va~ni za u inkovitost umna~anja (najviae tri uzastopna ponavljanja iste baze, pribli~no jednaka duljina ishodnica (20-21 bp), dopuateni udio GC-parova 50-70%, dopuateni raspon Tm 60-75C, pribli~no jednaka duljina nastalih PCR produkata). Premda smo pokuaali umno~iti fragmente mRNA za oba gena u istoj reakcijskoj smjesi tijekom jedne lan ane reakcije polimerazom (multipleks PCR), kako bi u najveoj moguoj mjeri izbjegli "tube to tube" varijacije, rezultati tih pokuaaja nisu bili zadovoljavajui. Naime, konstitutivna razina ekspresije -aktina je puno vea od konstitutivne razine ekspresije galektina-3. Premda se na ureaju AbiPrism 310 Genetic Analyser mogu analizirati i kvantificirati PCR produkti ija se koli ina razlikuje do 20 puta, u trenutku kada je koli ina PCR produkata za galektin-3 detektabilna (nakon 21 ciklusa PCR), koli ina PCR produkata za -aktin je izrazito velika. Zaato jednostavno ne ponoviti analizu koriatenjem razrijeenih uzoraka i zdru~iti rezultate analize koli ine PCR produkata za galektin-3 i -aktin? Na~alost, takav pristup mogao bi navesti na pogreane zaklju ke. Naime, ustanovili smo da je nakon 21 ciklusa umna~anja dijela mRNA za -aktin lan ana reakcija polimerazom ve dosegnula svoju krajnju to ku, te da koli ina nastalog produkta viae nije proporcionalna po etnoj koli ini mRNA. Stoga smo umna~anje dijelova mRNA za galektin-3 i za -aktin ipak morali provesti u zasebnim epruvetama, tijekom druk ijih reakcijskih uvjeta (14 ciklusa umna~anja za -aktin, 21 ciklus za galektin-3). Jedna od prednosti kvantifikacije PCR produkata na ureaju AbiPrism 310 Genetic Analyser je ato je izrazito osjetljiv, te je mogue kvantificirati vrlo male koli ine PCR produkata (za usporedbu, na agaroznom se gelu jedva mogu detektirati PCR produkti za galektin-3 nastali tijekom 28 ciklusa umna~anja). Stoga je mogue umno~iti dijelove mRNA i prekinuti lan anu reakciju polimerazom joa dok je u svojoj eksponencijalnoj fazi, ato je osnovni uvjet za pouzdanu kvantifikaciju PCR produkata. Joa jedan od uvjeta pouzdane kvantifikacije je jednaka u inkovitost amplifikacije PCR produkata razli itih gena. Provjerena je na taj na in ato su prireena razrjeenja cDNA, te su isti volumeni (a razli ite koli ine cDNA) podvrgnuti lan anim reakcijama polimerazom tijekom 14 ciklusa (za umna~anje gena za -aktin) i 21 ciklusa (za umna~anje gena za galektin-3). Rezultati su pokazali da su omjeri PCR produkta za galektin-3 i -aktin uvijek jednaki, iz ega zaklju ujemo da se PCR produkti umna~aju jednakom u inkovitoau. Nakon odabira odgovarajue stani ne linije i optimiranja metoda analize, ispitana je ekspresija LGALS3 i galektina-3 u nediferenciranim stanicama THP-1 nakon 1, 3, 5, 24, 48 i 72 kultiviranja u kulturi po etne gustoe 2 x 105 st/ml. Obzirom da stanice THP-1 rastu u suspenziji te su kultivirane kao asinhrona stani na kultura, bilo je za o ekivati da se konstitutivna razina ekspresije kako LGALS3, tako i galektina-3 odr~ava na jednakoj razini tijekom 72-satnog kultiviranja. Rezultati su pokazali da je pretpostavka to na. Takoer je utvreno da 72-satno kultiviranje ne utje e niti na vijabilnost niti na metaboli ku aktivnost stanica, bez obzira na gotovo peterostruko poveanje gustoe stani ne linije, ime je ve bila gotovo konfluentna. Stanice monocitne stani ne linije THP-1 diferencirane su tijekom 48-satnog izlaganja forbol-12-miristat-13-acetatu (PMA) koncentracije 10-7 M. Ve nakon prvog sata kultiviranja uz PMA, e"80% stanica THP-1 adherira na podlogu (postotak adheriranih stanica sli an je i nakon 3 i 5 sati kultiviranja), ali se koli ina mRNA za galektin-3 niti galektina-3 ne mijenja zna ajno. Tijekom prvih pet sati kultiviranja uz PMA, premda u velikom postotku ve adherirane na podlogu, stanice THP-1 (promjera 10-15 m) zadr~avaju sferi an oblik. Nakon 24 sata kultiviranja poprimaju ameboidan oblik makrofaga (promjera 30-50 m). No nama zanimljiviji podatak jest da se nakon 24- i 48-satnog kultiviranja uz PMA podosta poveava razina ekspresije LGALS3 (2,4 odnosno 2,9 puta) i galektina-3 (2,9 odnosno 3,5 puta). Koji su mogui mehanizmi regulacije ekspresije LGALS3 i galektina-3 tijekom diferencijacije stanica THP-1? Na osnovi strukture promotorskog dijela gena za galektin-3 otkrivene 1998. godine pretpostavljeno je da transkripcija gena za galektin-3 mo~e biti regulirana transkripcijskim faktorima Sp1, AP-1, NF-B i CREB. Zaista, rezultati naaih prethodnih istra~ivanja  ADDIN EN.CITE <EndNote><Cite><Author>Dumic</Author><Year>2000</Year><RecNum>197</RecNum><MDL><REFERENCE_TYPE>0108784451032000Expression of galectin-3 in cells exposed to stress-roles of jun and NF-kappaB149-58Department of Biochemistry and Molecular Biology, Faculty of Pharmacy and Biochemistry, University of Zagreb, Ante Kovavica 1, 10000 Zagreb, Croatia.Dumic, J.Lauc, G.Flogel, M.Cell Physiol BiochemAlkylating Agents/pharmacologyAntigens, Differentiation/*geneticsGalectin 3Gene Expression RegulationHeat-Shock ResponseHumanNF-kappa B/*metabolismProto-Oncogene Proteins c-jun/*metabolismSupport, Non-U.S. Gov'tTrypsin/pharmacologyTumor Cells, CulturedUltraviolet Rayshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10878445[30] pokazali su da je bazalna razina ekspresije galektina-3 u adherirajuim stanicama A1235 uglavnom regulirana transkripcijskim faktorom AP-1, dok je indukcija njegove ekspresije izazvane UV-zra enjem posredovana transkripcijskim faktorima NF-B i AP-1. Ekspresija LGALS3 mijenja se pod utjecajem razli itih stimulansa, ali prvu naznaku o moguim mehanizmima regulacije te ekspresije pru~ila su nedavna istra~ivanja Kim i sur., na osnovi kojih se mo~e zaklju iti da u regulaciji ekspresije LGALS3 va~nu ulogu ima signalni put Ras-Raf-MKK1/MKK3/p38/AP-1  ADDIN EN.CITE <EndNote><Cite>Kim2003164012788225164112003Jun 17Galectin-3 expression in macrophages is signaled by Ras/MAP kinase pathway and up-regulated by modified lipoproteins13-23Department of Pathology and Microbiology, University of South Carolina-School of Medicine, Columbia, SC 29208, USA.Kim, K.Mayer, E. P.Nachtigal, M.Biochim Biophys ActaAnimalsCell DifferentiationCells, CulturedComparative StudyEnzyme Inhibitors/pharmacologyGalectin 3/*metabolismGene Expression Regulation/*drug effectsGenes, jun/drug effectsMAP Kinase Signaling System/*physiologyMacrophages, Peritoneal/cytology/*metabolismMiceMice, Inbred C3HMitogen-Activated Protein Kinase Kinases/antagonists & inhibitorsProtein Kinase C/antagonists & inhibitorsProtein-Serine-Threonine Kinases/antagonists & inhibitorsProto-Oncogene Proteins c-jun/metabolismRabbits*Signal TransductionSupport, Non-U.S. Gov'tTetradecanoylphorbol Acetate/pharmacologyUp-Regulationras Proteins/*metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12788225[32]. Diferencijacija stanica THP-1 inducirana je forbol-12-miristatom-13-acetatom, esto rabljenom tvari u pokusima kojima se istra~uje stani na diferencijacija, prijenos signala i genska ekspresija. Aktivirajui protein kinazu C, kinazu reguliranu izvanstani nim signalima (ERK) te c-Jun N-terminalnu kinazu (JNK), PMA pokree kaskadu dogaaja koji rezultiraju aktivacijom transkripcijskih faktora kao ato su NF-B i AP-1 te diferencijacijom odreenih stani nih tipova. Vjerojatno je upravo aktivacija NF-B i AP-1 uzrok poveanja ekspresije LGALS3. Osim poveane koli ine mRNA za galektin-3, poveanje koli ine samog galektina-3 mo~e biti posljedica i aktivacije translacije, posebice ako se uzme u obzir da je porast koli ine mRNA (2,9 puta) neato manji od porasta koli ine samog proteina (3,5 puta). Poznato je naime da se faktori inicijacije translacije aktiviraju fosforilacijom koja je potaknuta djelovanjem PMA  ADDIN EN.CITE Morley19972063270942873841831997Dec 1Signalling through either the p38 or ERK mitogen-activated protein (MAP) kinase pathway is obligatory for phorbol ester and T cell receptor complex (TCR-CD3)-stimulated phosphorylation of initiation factor (eIF) 4E in Jurkat T cells327-32Biochemistry Laboratory, School of Biological Sciences, University of Sussex, Brighton, UK. s.j.morley@sussex.ac.ukMorley, S. J.FEBS LettCa(2+)-Calmodulin Dependent Protein Kinase/*metabolismCarcinogens/pharmacologyHumanJurkat Cells*Mitogen-Activated Protein Kinase Kinases*Mitogen-Activated Protein KinasesPeptide Initiation Factors/*metabolismPhosphorylationProtein-Serine-Threonine Kinases/*metabolismProtein-Tyrosine Kinase/*metabolismReceptor-CD3 Complex, Antigen, T-Cell/*metabolism*Signal Transduction/drug effectsSupport, Non-U.S. Gov'tT-Lymphocytes/*metabolismTetradecanoylphorbol Acetate/pharmacologyhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=9428738Miyamoto200020701107206614941-22000Nov 15Signal transduction pathways that contribute to increased protein synthesis during T-cell activation28-42Molecular Hematology Branch, NHLBI, Bethesda, MD 20892-1654, USA. smiyamot@ucdavis.eduMiyamoto, S.Kimball, S. R.Safer, B.Biochim Biophys Acta1-Phosphatidylinositol 3-Kinase/antagonists & inhibitors/metabolismAndrostadienes/pharmacology*Carrier ProteinsCells, CulturedEukaryotic Initiation Factor-2B/metabolismEukaryotic Initiation Factor-4EHumanIonomycin/pharmacologyKinetics*Lymphocyte Activation/drug effectsMAP Kinase Signaling System/drug effectsMitogen-Activated Protein Kinases/antagonists & inhibitors/metabolismModels, BiologicalPeptide Initiation Factors/metabolismPhosphoproteins/metabolismPhosphorylation/drug effectsProtein Kinase C/antagonists & inhibitors/metabolismProteins/biosynthesisProto-Oncogene Proteins/metabolismRibosomal Protein S6 Kinases/metabolism*Signal Transduction/drug effectsSirolimus/pharmacologySupport, U.S. Gov't, P.H.S.T-Lymphocytes/drug effects/enzymology/*immunology/*metabolismTacrolimus/pharmacologyTetradecanoylphorbol Acetate/pharmacology*Translation, Genetic/drug effectshttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11072066[176, 177]. Nakon ato smo raspravili mogue mehanizme kojima se omoguava poveanje ekspresije galektina-3 tijekom pretvorbe monocitnih stanica u makrofagne, valja se zapitati zaato je stanici potrebna vea koli ina galektina-3. Sazrijevanje monocita u makrofage praeno je promjenom morfologije i fiziologije stanica, ato uklju uje poveanu ekspresiju membranskih proteina, unutarstani nih enzima i receptora. Kako monocitne stanice rastu u suspenziji, a makrofagne adherirane na podlogu, prilikom pretvorbe stanice moraju ostvariti interakcije s podlogom. Sudjeluje li mo~da i galektin-3 u ostvarivanju tih interakcija? Obzirom na to da je poznato da galektin-3 sudjeluje u adheziji stanica na izvanstani ni matriks, mogue je da sudjeluje i u ostvarivanju interakcija stanica i podloge in vitro. Prema naaih prethodnim istra~ivanjima, presaivanjem adherirajuih stanica podignutih tripsinizacijom s podloge dolazi do promjene (preciznije, smanjenja) koli ine galektina-3, kako membranskog, tako i citosolnog, ali se nakon jednodnevnog kultiviranja ponovno uspostavlja konstitutivna razina galektina-3  ADDIN EN.CITE Dumic2000200010817217553-42000Mar-AprTransfer to in vitro conditions influences expression and intracellular distribution of galectin-3 in murine peritoneal macrophages261-6Department of Biochemistry and Molecular Biology, Faculty of Pharmacy and Biochemistry, University of Zagreb, Croatia.Dumic, J.Lauc, G.Hadzija, M.Flogel, M.Z Naturforsch [C]AnimalsAntigens, Differentiation/*metabolismCells, CulturedGalectin 3Macrophages, Peritoneal/*metabolismMaleMiceMice, Inbred BALB CRatsSubcellular Fractions/metabolismSupport, Non-U.S. Gov'thttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=10817217[49]. Rezultati ovog istra~ivanja pokazuju da premda je velika veina stanica THP-1 tijekom prvih pet sati kultiviranja uz PMA adherirala, nije doalo do promjene ekspresije galektina-3. To joa uvijek ne zna i da on ne sudjeluje u procesu adhezije, obzirom na to da monocitne stanice imaju izlu eni galektin-3 vezan na svoje membranske receptore  njegove glikoproteinske ligande. Osim toga, koli ina galektina-3 povea se 3 puta nakon 24-satnog kultiviranja uz PMA, kada je uo eno i trostruko poveanje njihova promjera, a samim time i povraine kojom se ostvaruju interakcije s podlogom. No uklju enost galektina-3 u ostvarivanje interakcija stanica s podlogom valjalo bi dodatno istra~iti. Galektin-3 u monocitima poti e unos Ca2+ te uzrokuje poveanje stvaranja interleukina-1. Djeluje kao kemoatraktant za monocite, ali i za makrofage. No u makrofagima galektin-3 ima i mnoge druge zna ajne uloge. Osim ato izaziva oksidativni prasak u makrofagima, ini se da sudjeluje i u procesu fagocitoze. Veliki kapacitet fagocitoze jedna je od osnovnih karakteristika makrofaga. To je jedan od klju nih procesa u ostvarivanju uroene imunosti ali i u patogenezi autoimunih bolesti. Nedavna saznanja prema kojima je galektin-3 jedna od osnovnih komponenti fagosoma miaje makrofagne stani ne linije i egzosoma dendritske stani ne linije navode na zaklju ak da je taj protein uklju en u endocitozu i prezentaciju antigena. Sano i sur. su utvrdili da je galektin-3 nu~an za u inkovitu fagocitozu IgG-opsoniziranih eritrocita i apoptotskih stanica in vitro i in vivo  ADDIN EN.CITE Sano200321101289720611232003AugCritical role of galectin-3 in phagocytosis by macrophages389-97Division of Allergy, La Jolla Institute for Allergy and Immunology, San Diego, California, USA.Sano, H.Hsu, D. K.Apgar, J. R.Yu, L.Sharma, B. B.Kuwabara, I.Izui, S.Liu, F. T.J Clin InvestAnemia, Hemolytic, Autoimmune/immunology/pathology/physiopathologyAnimalsApoptosisErythrocytes/immunology/physiologyGalectin 3/deficiency/genetics/*physiologyImmunoglobulin G/metabolismIn VitroKupffer Cells/physiologyMacrophages, Peritoneal/*physiologyMiceMice, Inbred C57BLMice, KnockoutOpsonins/metabolismPhagocytosis/*physiologyReceptors, IgG/metabolismSupport, Non-U.S. Gov'tSupport, U.S. Gov't, P.H.S.T-Lymphocytes/cytology/immunology/physiology</KEYWORD></KEYWORDS><URL>http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&amp;db=PubMed&amp;dopt=Citation&amp;list_uids=12897206</URL></MDL></Cite></EndNote>[178]. Obzirom na to da nije uo ena promjena koli ine niti mRNA za galektin-3 niti galektina-3 tijekom 72- satnog kultiviranja diferenciranih stanica THP-1 u mediju, zaklju ujemo da se nakon porasta koli ine galektina-3 prilikom diferencijacije monocita u makrofage, ukoliko se oni ne izlo~e nikakvom dodatnom stimulansu, u njima odr~ava uspostavljena konstitutivna razina ekspresije LGALS3 i galektina-3. Prilikom izbora koncentracija tvari s imunomodulatornim djelovanjem uz prisutnost kojih e se stanice kultivirati, smjernice su bile terapeutske koncentracije tih tvari u krvi ljudi koji se nalaze pod terapijom (premda i te koncentracije jako variraju ovisno o tipu bolesti). Takoer se ~elio pokriti airi raspon terapeutskih koncentracija (odnosno niske, srednje i visoke). Meutim, obzirom da se situacija in vitro u mnogo emu razlikuje od situacije in vivo, prije no ato se pristupilo ispitivanju razine ekspresije LGALS3 i galektina-3 provjerena je citotoksi nost imunomodulatornih tvari odreenih koncentracija te su za daljnje pokuse odabrane samo one koncentracije koje ne djeluju citotoksi no niti na nediferencirane niti na diferencirane stanice THP-1 tijekom 72-satnog kultiviranja. Kada se aspirin koristi kao analgetik, njegove koncentracije u plazmi su oko 0,5 mM, a kada se rabi kao protuupalni lijek do 2,7 mM. Vee koncentracije su toksi ne. Sukladno tome, rezultati naaih pokusa pokazali su da 4 mM aspirin djeluje citotoksi no na stanice THP-1 (rezultati nisu prikazani). Stoga su odabrane koncentracije aspirina bile 0,1 mM, 0,5 mM, 1 mM i 2 mM. Indometacin je koriaten za pokuse u koncentracijama 0,1 M, 1 M, 5 M i 10 M, obzirom da je raspon terapeutskih koncentracija 1,3 M do 8,3 M. U istim je koncentracijama rabljen i hidrokortizon, do im su primijenjene koncentracije deksametazona bile 0,1 nM do 100 nM, obzirom da je ustanovljeno da deksametazon u veim koncentracijama djeluje citotoksi no na stanice THP-1. Aspirin, indometacin, hidrokortizon i deksametazon u odabranim koncentracijama ne utje u zna ajno na cjelokupno metaboli ko stanje stanice koje bi za posljedicu imalo odumiranje stanice. `toviae, u diferenciranim stanicama THP-1 hidrokortizon (10 M) i deksametazon (100 nM) ak blago poveavaju metaboli ku aktivnost stanica. Niti jedna od navedenih tvari nije topljiva u vodi, pa su za potrebe eksperimenata prireene njihove mati ne otopine u DMSO. Po prireivanju radnih koncentracija tih tvari, kona ne koncentracije DMSO u medijima u kojima su stanice THP-1 inkubirane bile su d" 0,2%. Sam DMSO koncentracije 0,2% tijekom 72-satnog kultiviranja nije uzrokovao statisti ki zna ajnu promjenu niti jednog ispitivanog parametra (vijabilnosti, metaboli ke aktivnosti, ekspresije LGALS3 i ekspresije galektina-3). To je u suglasju s naaim o ekivanjima i literaturnim podacima, prema kojima tek 2% DMSO (ato je jedan red veli ine vea koncentracija od one primijenjene u ovom istra~ivanju!) ima citotoksi an u inak na stanice. Zaklju ujemo, takoer, da DMSO navedene koncentracije ne utje e zna ajno na mehanizme regulacije ekspresije LGALS3 i galektina-3. Tvari s imunomodulatornim djelovanjem iji je utjecaj na ekspresiju LGALS3 i galektina-3 u ovom radu ispitan pripadaju dvjema osnovnim skupinama protuupalnih lijekova - nesteroidnim tvarima s imunomodulatornim djelovanjem (NSAIDs) i glukokortikoidima. Oni razli itim mehanizmima, koji doduae nisu u potpunosti razjaanjeni, ostvaruju protuupalne u inke. U nediferenciranim, odnosno monocitnim stanicama THP-1 sve ispitane tvari s imunomodulatornim djelovanjem u svim primijenjenih koncentracijama inhibiraju ekspresiju LGALS3, preciznije, uzrokuju smanjenje koli ine mRNA za galektin-3. Ono se uo ava ve nakon prvog sata kultiviranja, osim uz najni~u koncentraciju aspirina (0,1 M), kod koje se tendencija pada primjeuje nakon petog sata kultiviranja, a statisti ki zna ajnom postaje nakon jednodnevnog kultiviranja. Postotak inhibicije varira od 66% do ak 96% te je u korelaciji je s vremenom izlaganja tvarima s imunomodulatornim djelovanjem. Takoer je u korelaciji s primijenjenom koncentracijom aspirina, dok je u slu aju ostalih tvari ta korelacija ne postoji. Koji su mogui mehanizmi kojima tvari s imunomodulatornim djelovanjem ostvaruju inhibiciju ekspresije gena za galektin-3? Odavno je poznato da aspirin acetiliranjem inhibira aktivnost ciklooksigenaze i vjeruje se da je veina protuupalnih u inaka aspirina posljedica smanjenja aktivnosti tog enzima, klju ne molekule biosintetskog puta prostaglandina. Sa prili nom sigurnoau mo~emo zaklju iti da to nije meutim na in kojim aspirin inhibira ekspresiju LGALS3. No injenica da je puno vea koncentracija aspirina potrebna za utiaavanje upale nego za inhibiciju ciklooksigenaze navela je znanstvenike na istra~ivanja kojima je utvreno da aspirin mo~e modulirati ekspresiju gena i proteina, kao i aktivnost proteina utje ui na signalne puteve protein kinaza aktiviranih mitogenima (MAPK) i/ili na transkripcijske faktore NF-B, AP-1 i STAT1. Stoga je logi no za pretpostaviti da je inhibicija, izravna ili posredna, transkripcijskih faktora NF-B i/ili AP-1, ija vezna mjesta nalazimo i u promotorskoj regiji LGALS3, odgovorna za smanjenje transkripcije tog gena, a samim time i koli ine mRNA. Kao i aspirin, i indometacin ubrajamo u skupinu nesteroidnih tvari s imunomodulatornim djelovanjem. Osim njegovog u inka na aktivnost cikooksigenaze, ostala su njegova djelovanja slabo istra~ena. Utvreno je, meutim, da indometacin ve~e i aktivira PPAR, nuklearni hormonski receptor koji nakon vezanja hormona djeluje kao transkripcijski faktor. No vezna mjesta za PPAR ne nalaze se u promotorskoj regiji LGALS3 (a osim toga, indometacin ne aktivira, ve inhibira ekspresiju LGALS3). Premda su rezultati nekih istra~ivanja naveli na zaklju ak da indometacin ne mo~e inhibirati aktivnost niti transkripcijskog faktora NF-B niti AP-1, prema drugim istra~ivanjima utvreno je da je takva inhibicija ipak mogua, i to inhibicija NF-B procesom transrepresije. Stoga mo~emo zaklju iti da je i opa~eno smanjenje koli ine mRNA za galektin-3 posljedica inhibicije NF-B. Uzimajui u obzir injenicu da do danas nije pokazano da indometacin mo~e inhibirati transkripcijski faktor AP-1 (ato joa uvijek ne zna i da to nije mogue!) te injenicu da je inhibitorni u inak terapeutskih koncentracija indometacina sli an, ako ne ak i sna~niji od aspirina, mo~e se pretpostaviti da je bazalna razina ekspresije LGALS3 u monocitnim stanicama regulirana u veoj mjeri transkripcijskim faktorom NF-B nego AP-1. Jedan od na ina djelovanja steroidnih tvari s imunomodulatornim djelovanjem  glukokortikoida (GC), posredovan je njihovim vezanjem na receptore (GR), nakon ega oni djeluju kao transkripcijski faktori. Ukoliko se u promotorskim regijama gena nalaze vezna mjesta za te transkripcijske faktore, posljedi no e doi do poveanja ekspresije tih gena. Ako se meutim u promotorskoj regiji nalaze negativni elementi odgovora na GC (nGRE), mo~e doi do utiaavanja genske ekspresije. Kako analiza promotorskog dijela gena za galektin-3 nije potvrdila postojanje takvih nGRE, zaklju ujemo da je inhibicija LGALS3 ostvarena na drugi na in. Literaturni podaci pokazuju da glukokortikoidi (hidrokortizon i deksametazon) mogu inhibirati transkripcijske faktore NF-B, AP-1, CREB, C/EBP, STAT i p53 razli itim na inima, kao ato su transrepresija, inihicija razgradnje IB, izravno vezanje na transkripcijske faktore, kompeticija za iste kofaktore i joa neki dodatni mehanizmi. Stoga se ini da su ponovno transkripcijski faktori NF-B i AP-1 odgovorni za smanjenje ekspresije LGALS3 uo eno u monocitnim stanicama izlo~enim glukokortikoidima, ali tek podrobnija istra~ivanja dala bi detaljniji uvid u mehanizme kojima se ta inhibicija ostvaruje. Valja osim toga uzeti u obzir da smanjenje koli ine mRNA mo~e biti uzrokovano ne samo smanjenom transkripcijom doti nog gena, ve i poveanim stupnjem razgradnje te mRNA. Kako to da postoji korelacija izmeu intenziteta inhibicije LGALS3 i primijenjene koncentracije aspirina, ali ne i indometacina, hidrokortizona i deksametazona? Jedno od moguih objaanjenja je da je u slu aju primjene ni~ih koncentracija aspirina broj molekula aspirina joa uvijek puno manji od broja njegovih receptora. To zna i da primjenom veih koncentracija aspirina dolazi do sve veeg zasienja receptora, te posljedi no do poja avanja u inka te tvari. ini se meutim da kada se indometacin, hidrokortizon i deksametazon koriste ve i u najni~im koncentracijama dolazi do zasienja njihovih receptora, te se poveanjem koncentracija tih tvari ne poja ava njihov u inak. Kako se smanjenje koli ine mRNA za galektin-3 uzrokovano izlaganjem monocitnih stanica THP-1 tvarima s imunomodulatornim djelovanjem odra~ava na ekspresiju samog proteina? Niti jedna od ispitivanih tvari s imunomodulatornim djelovanje niti u jednoj od primijenjenih koncentracija nije uzrokovala promjenu koli ine galektina-3 tijekom prvih pet sati kultiviranja. Istovremeno, osim u slu aju najni~e primijenjene koncentracije aspirina (0,1 mM), koli ina mRNA za galektin-3 smanjena je nakon 1, 3 te 5 sati kultiviranja uz sve ispitivane tvari svih koncentracija za 30% do ak 70%. To se mo~e objasniti vremenskim odmakom procesa transkripcije i translacije. Nakon jednodnevnog kultiviranja joa uvijek se ne uo ava pad koli ine galektina-3 (osim uz 2 mM aspirin), premda je u gotovo svim slu ajevima postotak inhibicije ekspresije LGALS3 oko 70%. To je mo~da posljedica poveanja razine translacije i/ili o uvanja postojeih koli ina galektina-3, odnosno smanjenja stupnja njegove razgradnje. No nakon 48- i 72-satnog kultiviranja za o ekivati je da se ve dugotrajno i sna~no smanjenje ekspresije LGALS3 mora odraziti na razinu ekspresije tog proteina. Ona je u veini slu ajeva zaista i smanjena, ali u puno manjoj mjeri nego ekspresija samog LGALS3. Intenzitet smanjenja razine ekspresije galektina-3 u korelaciji je s vremenom izlo~enosti ispitivanim tvarima te s primijenjenom koncentracijom aspirina, hidrokortizona i deksametazona, do im je u slu aju indometacina u negativnoj korelaciji s primijenjenom koncentracijom te tvari. Osim toga, to smanjenje nije u korelaciji s razinom smanjenja koli ine mRNA, s obzirom na to da je u pojedinim slu ajevima ja i pad razine galektina-3 primjeen u stanicama koje imaju viae mRNA za galektin-3. Odr~anje relativno velike koli ine galektina-3 u odnosu na koli inu pripadajue mu mRNA teako je objasniti o uvanoau samog proteina (odnosno njegovom ne-razgradnjom) obzirom da je nakon 48-satnog (a posebice 72-satnog) kultiviranja gustoa stani ne kulture viae nego dvostruko vea. Izlu uju li mo~da stanice THP-1 znatne koli ine galektina-3 u izvanstani ni prostor, koji tek naknadno mogu vezati na sebe, primjerice ukoliko se pod utjecajem tvari s imunomodulatornim djelovanjem povea ekspresija i/ili afinitet njegovih membranskih liganada? Mogue, s obzirom na to da je nedavno ustanovljeno da fetuin, serumski glikoprotein kojeg u obilju ima u fetalnom serumu - jednoj od osnovnih komponenti hranidbenog medija, uzrokuje brzu (unutar 1 minute) sekreciju unutarstani nog galektina-3. To je jedno od razumnih objaanjenja ovih zanimljivih i mo~da neobi nih rezultata. Sve u svemu, o igledno je da je ekspresija galektina-3 u monocitnim stanicama THP-1 izlo~enim djelovanju aspirina, indometacina, hidrokortizona i deksametazona regulirana ne samo na razini transkripcije, ve i translacije, a mo~da i nekim drugim mehanizmima. ini se da je monocitnim stanicama galektin-3 toliko potreban da pronalaze u inkovite na ine odr~anja njegove proteinske razine bez obzira na drasti no smanjenje pripadajue mu mRNA. Kako tvari s imunomodulatornim djelovanjem utje u na ekspresiju LGALS3 i galektina  3 u diferenciranim, odnosno makrofagnim stanicama THP-1? Dok se u monocitnim stanicama opi obrazac promjene koli ine mRNA za galektin-3 mo~e opisati kao sna~na, vremenski ovisna inhibicija transkripcije gena za galektin-3, u makrofagnima je situacija druk ija. Mo~e se uo iti da sve ispitivane tvari u po etku inhibiraju ekspresiju LGALS3 nakon ega dolazi do uspostave konstitutivne razine mRNA. U slu aju aspirina i hidrokortizona intenzitet je te inhibicije u korelaciji s koncentracijom primijenjene tvari, ato kod indometacina i deksametazona nije zamijeeno. No kako to da kultiviranje uz odreenu tvar neke koncentracije ima tako razli it u inak na stanice iste loze? To nije toliko udno ni neobi no. Premda je zapravo rije  o istim stanicama (THP-1), o ito je da su zbog djelovanja PMA promijenile mnoge svoje karakteristike. Jedna od njih je i razina ekspresije gena  podsjetimo se da je ekspresija LGALS3 tri puta vea u diferenciranim nego u nediferenciranim stanicama. To je poveanje vjerojatno rezultat aktivacije transkripcijskih faktora AP-1 i/ili NF-B. Na tako aktivirane transkripcijske faktore tvari s imunomodulatornim djelovanjem u odreenoj koncentraciji imaju o ito puno slabiji u inak. To je vrlo jednostavno objaanjenje razlike u stupnju inhibicije ekspresije LGALS3. Zanimljivo je osim toga, ato postotak inhibicije u nediferenciranim stanicama vremenom raste, a u diferenciranima se vrlo brzo uspostavlja konstitutivna razina ekspresije LGALS3. Primjer razli itog u inka jednake koncentracije odreene tvari tijekom istog vremena na monocitne i makrofagne stanice prikazan je na slici 4.1.  Slika 4.1 Utjecaj hidrokortizona na ekspresiju a) LGALS3 i b) galektina- 3 u nediferenciranim i diferenciranim stanicama THP-1. Nediferencirane i diferencirane stanice THP-1 kultivirane su tijekom 1, 3, 5, 24, 48 ili 72 sata uz hidrokortizon koncentracije 1 M. a) relativne koli ine mRNA za galektin-3 (normirane prema -aktinu) i b) relativne koli ine galektina-3 (normirane prema -aktinu) izra~ene su u odnosu na kontrolu (nediferencirane ili diferencirane stanice kultivirane tijekom istih vremena u mediju)(x (SD).Rebrasti stupci  nediferencirane stanice, zeleni stupci  diferencirane stanice. `to se istovremeno dogaa s galektinom-3? Kultiviranjem makrofagnih stanica THP-1 uz indometacin i deksametazon koli ina galektina-3 u po etku se odr~ava na jednakoj razini, a zatim postupno poveava, sve to 72 sata kada u odnosu na 48 sat blago padne, i to u slu aju indometacina pribli~no na po etnu razinu, a u slu aju deksametazona, na razinu koja je joa uvijek zna ajno vea od po etne. Aspirin i hidrokortizon ovisno o primijenjenim koncentracijama, uzrokuju pad koli ine galektina-3. Nakon pada dolazi do poveanja koli ine galektina-3 tijekom vremena, ime se ponovno uspostavlja konstitutivna razina ekspresije, koja se potom ak i premaauje. Ako se ti podaci usporede s podacima o razini ekspresije mRNA za galektin-3 u istim stanicama, dolazi se do zaklju ka da vee koncentracije primjenjivanih tvari uzrokuju ja e smanjenje koli ine mRNA. Ukoliko je ostvaren dovoljan postotak inhibicije ekspresije LGALS3, to se o ituje i na proteinskoj razini. ini se da je u tom razdoblju koli ina galektina-3 regulirana veinom na razini transkripcije. Prisjetimo se da to nije bio slu aj u monocitnim stanicama, kao i da se u njima puno vei postotak inhibicije ekspresije LGALS3 nije odra~avao na proteinskoj razini. Meutim, uspostavu konstitutivne razine ekspresije LGALS3 prati daljnje poveanje koli ine galektina-3. Poveava li se u tom periodu intenzitet translacije postojeih mRNA? Smanjuje li se razgradnja postojeih proteina uz kontinuiranu sintezu novih? Ve~u li se vee koli ine izlu enog galektina-3 na membranu? To su pitanja na koja tek treba nai odgovor. Sli ni odnosi koli ine mRNA i proteina primijeeni su i u nekim drugim istra~ivanjima, primjerice, Andersson i Sundler  ADDIN EN.CITE Andersson200020801109774812122000DecSignalling to translational activation of tumour necrosis factor-alpha expression in human THP-1 cells1784-7Department of Cell and Molecular Biology, Section for Molecular Pathogenesis, Lund University, S-221 00, Lund, Sweden.Andersson, K.Sundler, R.CytokineAnisomycin/pharmacologyBlotting, NorthernBlotting, WesternButadienes/pharmacologyCalcium/metabolismCell LineCycloheximide/pharmacologyDactinomycin/pharmacologyDose-Response Relationship, DrugEnzyme Inhibitors/pharmacologyFlavonoids/pharmacologyHumanImidazoles/pharmacologyIonophores/pharmacologyMAP Kinase Signaling SystemMitogen-Activated Protein Kinases/metabolismMonocytes/metabolismNitriles/pharmacologyProtein Synthesis Inhibitors/pharmacologyPyridines/pharmacologyRNA, Messenger/metabolism*Signal TransductionSupport, Non-U.S. Gov'tTetradecanoylphorbol Acetate/pharmacologyTime FactorsTranslation, GeneticTumor Necrosis Factor/biosynthesis/*metabolismp42 MAP Kinase/metabolismhttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11097748[179] su pokazali da stanice THP-1 eksprimiraju mRNA za faktor nekroze tumora, ali jedva detektabilnu koli inu samog proteina. Nakon stimulacije stanica s PMA, poveava se razina proteina, ali ne i mRNA. Nekoliko je studija pokazalo loau korelaciju izmeu razine mRNA i stvarne razine ekspresije gena, odnosno koli ine samog proteina. Primjerice, Lorenz i sur.  ADDIN EN.CITE <EndNote><Cite><Author>Lorenz</Author><Year>2003</Year><RecNum>209</RecNum><IDText>991</IDText><MDL><REFERENCE_TYPE>0</REFERENCE_TYPE><ACCESSION_NUMBER>12833524362003JunFrom transcriptome to proteome: differentially expressed proteins identified in synovial tissue of patients suffering from rheumatoid arthritis and osteoarthritis by an initial screen with a panel of 791 antibodies991-1002Institute of Immunology, Rostock, Germany.Lorenz, P.Ruschpler, P.Koczan, D.Stiehl, P.Thiesen, H. J.ProteomicsAntibodies/metabolismArthritis, Rheumatoid/*metabolism/pathologyBlotting, WesternComparative StudyFemaleGene Expression ProfilingHumanMiddle AgedOligonucleotide Array Sequence AnalysisOsteoarthritis/*metabolismProteins/analysis/*metabolismProteome/*metabolismSupport, Non-U.S. Gov'tSynovial Membrane/*metabolism/pathologyTranscription, Genetichttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=12833524[180] su utvrdili da od 58 proteina ija je promjena ekspresije utvrena u pacijenata oboljelih od reumatoidnog artritisa u odnosu na oboljele od osteoartritisa samo kod njih 16 (28%) te su promjene u skladu s promjenama na razini transkripcije. Shipp i sur.  ADDIN EN.CITE Shipp200221068011786909812002JanDiffuse large B-cell lymphoma outcome prediction by gene-expression profiling and supervised machine learning68-74Dana-Farber Cancer Institute, Harvard Medical School, Boston, Massachusetts, USA. margaret_shipp@dfci.harvard.eduShipp, M. A.Ross, K. N.Tamayo, P.Weng, A. P.Kutok, J. L.Aguiar, R. C.Gaasenbeek, M.Angelo, M.Reich, M.Pinkus, G. S.Ray, T. S.Koval, M. A.Last, K. W.Norton, A.Lister, T. A.Mesirov, J.Neuberg, D. S.Lander, E. S.Aster, J. C.Golub, T. R.Nat MedAntineoplastic Combined Chemotherapy Protocols*Artificial IntelligenceCyclophosphamideDoxorubicinGene Expression Profiling/*methodsHumanLymphoma, B-Cell/*diagnosis/drug therapy/mortalityLymphoma, Large-Cell, Diffuse/*diagnosis/drug therapy/mortalityOligonucleotide Array Sequence AnalysisPredictive Value of TestsPrednisoneSupport, Non-U.S. Gov'tTreatment OutcomeVincristinehttp://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11786909[181] su detektirali visoku razinu ekspresije mRNA za galektin-3 u difuznim limfomima velikih limfocita T u odnosu na folikularne limfome, ali ona nije korelirala s razinom ekspresije proteina. Osvrnimo se joa na trenutak na rezultate imunoblot analize stani nih homogenata diferenciranih stanica THP-1 izlo~enih djelovanju hidrokortizona i deksametazona. Uo ava se prisutnost dodatnih vrpci neposredno ispod vrpce galektina-3. Obzirom da na istom blotu te vrpce ne postoje ispod vrpce kontrolnih uzoraka, zaklju ujemo da to nije posljedica sekundarnog obojenja, a vjerojatno niti nespecifi nog vezanja. Odgovaraju li te vrpce mo~da fosforiliranom galektinu-3? Naime, zna se da galektin-3 mo~e biti fosforiliran na polo~aju Ser6 i Ser12, a nedavna istra~ivanja su pokazala da galektin-3 mo~da mo~e biti fosforiliran i na nekom od tirozina. Fosforilacijom se mijenja izoelektri na to ka galektina-3 (pI nefosforiliranog oblika iznosi 8,7, a fosforiliranog 8,2), ato vjerojatno mo~e promijeniti i njegovu pokretljivost tijekom SDS-PAG elektroforeze. Fosforilirani galektin-3 pokazuje smanjenu lektinsku aktivnost, ali je fosforilacija nu~an preduvjet za antiapoptozno djelovanje galektina-3. Da li je poveana metaboli ka aktivnost diferenciranih stanica izlo~enih djelovanju hidrokortizona i deksametazona rezultat fosforilacije i antiapoptoznog djelovanja galektina-3? Na to, kao i na mnoga druga pitanja o funkciji galektina-3 tek treba pronai odgovor. zaklju ci Na temelju provedenih pokusa, dobivenih rezultata i rasprave mogue je zaklju iti sljedee: Aspirin u koncentracijama d" 2 mM, indometacin u koncentracijama d"10 M, hidrokortizon u koncentracijama d" 10 M, deksametazon u koncentracijama d" 100 nM, kao ni DMSO koncentracije 0,2%, ne djeluju citotoksi no niti na nediferencirane niti na diferencirane stanice THP-1. Kultiviranjem nediferenciranih stanica tijekom 72 sata u standardnim uvjetima razina ekspresije LGALS3 kao ni galektina-3 se ne mijenja. DMSO koncentracije 0,2% ne utje e na ekspresije LGALS3 i galektina-3 niti u nediferenciranim niti u diferenciranim stanicama THP-1 tijekom 72-satnog kultiviranja. Diferencijacija stanica THP-1 iz monocitnih u makrofagne stanice poti e ekspresiju LGALS3 i galektina-3. Razina ekspresije LGALS3 i galektina-3 u stanicama stani ne linije THP-1 ovisi o diferencijacijskom stupnju stanica. Nakon diferencijacije stanica THP-1 iz monocitnih u makrofagne, daljnje kultiviranje stanica u standardnim uvjetima tijekom 72 sata ne mijenja uspostavljenu razinu ekspresije LGALS3 i galektina-3. Tvari s imunomodulatornim djelovanjem (aspirin, indometacin, hidrokortizon i deksametazon) mijenjaju ekspresiju LGALS3 i galektina-3 u stanicama stani ne linije THP-1, a promjene ovise o vrsti i koncentraciji primijenjene tvari, vremenu izlo~enosti stanica istima te diferencijacijskom stupnju stanica. U inak svake pojedine tvari s imunomodulatornim djelovanjem u pojedinoj koncentraciji na razinu galektina-3 u kvalitativnom se i kvantitativnom smislu razlikuje u odnosu na njezin u inak na razinu mRNA za galektin-3 u stanicama monocitne kao i u stanicama makrofagne stani ne linije THP-1: sve tvari u primijenjenim koncentracijama u monocitnim stanicama THP-1 inhibiraju ekspresiju LGALS3 i galektina-3. Stupanj inhibicije na razini mRNA ovisi o koncentraciji imunomodulatornih tvari i u korelaciji je s vremenom izlo~enosti stanica njihovom djelovanju. Inhibitorni je u inak na proteinskoj razini u odnosu na inhibitorni u inak na razini mRNA vremenski odgoen. u diferenciranim stanicama aspirin, indometacin, hidrokortizon i deksametazon u po etku djeluju inhibitorno na ekspresiju LGALS3 nakon ega dolazi do uspostave konstitutivne razine mRNA. u inak slabih imunomodulatornih tvari - aspirina i hidrokortizona na razinu galektina-3 u diferenciranim stanicama THP-1 razlikuje se od u inka jakih imunomodulatornih tvari  indometacina i deksametazona; iako dugotrajnijim izlaganjem objema vrstama tvari razina galektina-3 raste, slabe imunomodulatorne tvari u po etku na razinu galektina-3 djeluju inhibitorno. Literatura  ADDIN EN.REFLIST 1. Fred Brewer, C., Binding and cross-linking properties of galectins. Biochim Biophys Acta, 2002. 1572(2-3): 255-62. 2. Cooper, D.N., Galectinomics: finding themes in complexity. Biochim Biophys Acta, 2002. 1572(2-3): 209-31. 3. 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CCAAT-enhancer-binding protein COX-1ciklooksigenaza 1 engl. cyclooxygenase 1COX-2ciklooksigenaza 2 engl. cyclooxygenase 2CRDdomena koja prepoznaje ugljikohidrate engl. carbohydrate recognition domainCREelementi odgovora na cAMP engl. cAMP responsive elementsCREBprotein koji se ve~e na element odgovora na cAMP engl. cAMP-response element binding factorDaDalton (gmol-1)dATPdeoksiadenozin-trifosfat engl. deoxyadenosine-triphosphatedCTPdeoksicitidin-trifosfat engl. deoxycytidine-triphosphateDEPCdietil-pirokarbonat engl. diethyl-pirocarbonateDexdeksametazon engl. dexamethasonedGTPdeoksigvanozin-trifosfat engl. deoxyguanosine-triphosphateDMSOdimetil-sulfoksidDNAdeoksiribonukleinska kiselina engl. deoxyribonucleic aciddNTPdeoksinukleozid-trifosfat engl. deoxynucleoside-triphosphatedTTPdeoksitimidin-trifosfat engl. deoxythymidine-triphosphateEDTAetilen-diamin-tetraoctena kiselina engl. ethylenediaminetetraacetic acidERendoplazmatski retikulumERKkinaza regulirana izvanstani nim signalima engl. extracellular-signal-regulated kinase6-FAM6-karboksifluoresceinFucfukozaGalgalaktozagal-3galektin-3GalNacN-acetilgalaktozaminGCglukokortikoidiGlcNAcN-acetilglukozaminGM-CSFfaktor koji stimulira granulocitne i makrofagne kolonije engl. granulocyte-macrophage colony stimulating factorGRglukokortikoidni receptoriGREelementi odgovora na glukokortikoide engl. GC-response elementsHAThiston-acetiltransferaza engl. histone acetyltransferaseHChidrokortizon, kortizol engl. hydrocortisone, cortisolhnRNPheterogeni nuklearni ribonukleoproteiniHspproteini toplinskog aoka engl. heat-shock proteinsIFNinterferonIKKkinaza IBILinterleukinIndindometacin engl. indomethacinIBinhibitor NF-(BJNKc-Jun N-terminalna kinaza, drugim imenom SAPK engl. c-Jun-N-terminal kinasekbkilobazaLaclaktozaLacNAcN-acetillaktozaminLGALS3gen za galektin-3 engl. lectin, galactoside-binding, solubleLPSlipopolisaharidMan-6-Pmanoza-6-fosfatMAPKprotein kinaza aktivirana mitogenima engl. mitogen-activated protein kinaseMAPKKkinaza protein kinaze aktivirane mitogenima engl. mitogen-activated protein kinase kinaseMAPKKKkinaza kinaze protein kinaze aktivirane mitogenima engl. mitogen-activated protein kinase kinase kinaseMgat51,6 N-acetilglukozaminiltransferaza V engl. 1,6 N-acetylglucosaminyltransferase VMKP-1fosfataza protein kinaze aktivirane mitogenima engl. MAPK phosphatase 1MMPmetaloproteinaza matriksa engl. matrix metalloproteinaseMrrelativna molekulska masamRNAglasni ka ribonukleinska kiselina engl. messenger ribonucleic acidMTStetrazolijeva sol (3-(4,5-dimetiltiazol-2-il)-5-(3-karboksimetoksifenil)-2-(4-sulfofenil)-2H-tetrazolij-bromidNF-(Bnuklearni faktor (B engl. nuclear factor kappa BNFATnuklearni faktor aktiviranih stanica T engl. nuclear factor of activated T cellsnGREnegativni elementi odgovora na glukokortikoide engl. negative GC-response elementsNSAIDsnesteroidne tvari s imunomodulatornim djelovanjem engl. non-steroidal antiinflammatory drugsntnukleotidPAGpoliakrilamidni gel engl. polyacrylamide gelPBSfizioloka otopina (154 mM NaCl) puferirana fosfatnim puferom (Na2HPO4/KH2PO4), pH 7,4 engl. phosphate buffered salinePCRlan ana reakcija polimerazom engl. polymerase chain reactionPMAforbol-12-miristat-13-acetat engl. phorbol-12-myristate-13-acetatePMSFfenil-metil-sulfonil-fluorid engl. phenyl-methyl-sulphonyl-fluoridPPAR receptor  aktiviran proliferatorom peroksisoma engl. peroxisome proliferator-activated receptor PVDFpoliviniliden-difluorid engl. polyvinylidene difluorideRNAribonukleinska kiselina engl. ribonucleic acidSAPKprotein kinaza aktivirana stresom, drugim imenom JNK engl. stress-activated-protein-kinaseSDSnatrijev dodecilsulfat engl. sodium dodecylsulphateSp-1transkripcijski faktor Sp1STATmolekule koje provode signale i aktiviraju transkripciju engl. signal transducer and activator protein 1TAMRAkarboksitetrametilrodaminTBSpuferirana otopina tris(hidroksimetil)aminometana engl. tris(hydroxymethy)aminomethane buffered salineTCRreceptori na limfocitima T engl. T-cell receptorsTemedN, N, N, N-tetrametil-etilen-diamin engl. N, N, N, N-tetramethyl-ethylenediamineTFtranskripcijski faktorTmtemperatura mekanjaTNFfaktor nekroze tumora engl. tumour necrosis factorTREselementi odgovora na forbol-12-O-tetradekanoat-13-acetat engl. phorbol-12-O-tetradecanoate-13-acetate response elementsTristris(hidroksimetil)aminometan engl. tris(hydroxymethy)aminomethaneTween-20polioksietilen sorbitan monolaurat engl. polyoxyethylenesorbitan monolaurate POPIS SLIKA I TABLICA Slika 1.1Strukture galektinaSlika 1.2Struktura galektina-3Slika 1.3Slijed nukleotida promotorske regije ljudskog LGALS3Slika 1.4Shema trokomponentnog modela aktivacije MAPKSlika 1.5Regulacija aktivnosti TF signalnim putem MAPKSlika 1.6Shematski prikaz aktivacije NF-BSlika 1.7Shematski prikaz signalnih puteva i mehanizama regulacije gena uklju enih u upalni odgovorSlika 1.8U inak nesteroidnih tvari s imunomodulatornim djelovanjem na pojedine komponente signalnih putevaSlika 1.9Struktura aspirinaSlika 1.10Struktura indometacinaSlika 1.11Struktura hidrokortizonaSlika 1.12Struktura deksametazonaSlika 2.1Analiza ukupne RNA na ureaju Agilent 2100 bioanalyserSlika 3.1Metaboli ka aktivnost stanica THP-1 tijekom 72-satnog kultiviranja u mediju/0,2% DMSO se ne mijenjaSlika 3.2Elektroforeza PCR produkataSlika 3.3Nukleotidni slijed PCR produkta nastalog umna~anjem dijela mRNA za galektin-3 odreen analizom na ureaju AbiPrism 310 Genetic AnalyserSlika 3.4GeneScan elektroforetogram razdvojenih PCR produkataSlika 3.5U stanicama THP-1 tijekom 72-satnog kultiviranja u mediju ili u mediju/0,2% DMSO koli ina mRNA za galektin-3 se ne mijenjaSlika 3.6Koli ina galektina-3 u stanicama THP-1 tijekom kultiviranja u mediju ili u mediju/0,2% DMSO se ne mijenjaSlika 3.7Aspirin inhibira ekspresiju LGALS3Slika 3.8Utjecaj aspirina na ekspresiju galektina-3 u stanicama THP-1Slika 3.9Indometacin inhibira ekspresiju LGALS3Slika 3.10Utjecaj indometacina na ekspresiju galektina-3 u stanicama THP-1Slika 3.11Hidrokortizon inhibira ekspresiju LGALS3Slika 3.12Utjecaj hidrokortizona na ekspresiju galektina-3 u stanicama THP-1Slika 3.13Deksametazon inhibira ekspresiju LGALS3Slika 3.14Utjecaj deksametazona na ekspresiju galektina-3 u stanicama THP-1Slika 3.15PMA inducira ekspresiju LGALS3 u stanicama THP-1Slika 3.16PMA inducira ekspresiju galektina-3 u stanicama THP-1Slika 3.17U diferenciranim stanicama THP-1 tijekom 72-satnog kultiviranja u mediju ili u mediju/0,2% DMSO koli ina mRNA za galektin-3 se ne mijenjaSlika 3.18Koli ina galektina-3 ne mijenja se u diferenciranim stanicama THP-1 tijekom kultiviranja u mediju ili u mediju/0,2% DMSOSlika 3.19Utjecaj aspirina na ekspresiju LGALS3 u diferenciranim stanicama THP-1Slika 3.20Utjecaj aspirina na ekspresiju galektina-3 u diferenciranim stanicama THP-1Slika 3.21Utjecaj indometacina na ekspresiju LGALS3 u diferenciranim stanicama THP-1Slika 3.22Utjecaj indometacina na ekspresiju galektina-3 u diferenciranim stanicama THP-1Slika 3.23Utjecaj hidrokortizona na ekspresiju LGALS3 u diferenciranim stanicama THP-1Slika 3.24Utjecaj hidrokortizona na ekspresiju galektina-3 u diferenciranim stanicama THP-1Slika 3.25Utjecaj deksametazona na ekspresiju LGALS3 u diferenciranim stanicama THP-1Slika 3.26Utjecaj deksametazona na ekspresiju galektina-3 u diferenciranim stanicama THP-1Slika 4.1Utjecaj hidrokortizona na ekspresiju a) LGALS3 i b) galektina-3 u nediferenciranim i diferenciranim stanicama THP-1Tablica 1.1Obitelji ~ivotinjskih lektinaTablica 1.2Utjecaj razli itih tvari na sekreciju galektina-3Tablica 1.3Relativni afiniteti vezanja galektina-3 na glikoligandeTablica 1.4Ligandi galektina-3Tablica 1.5Funkcije galektina-3 u imunom odgovoruTablica 2.1Sastav reakcijskih smjesa za reverznu transkripcijuTablica 2.2Sastav reakcijskih smjesa za lan anu reakciju polimerazomTablica 2.3Ishodnice za lan anu reakciju polimerazom ~ivotopis Roena sam 5. srpnja 1973. godine u Zagrebu, gdje sam zavraila osnovnu i srednju akolu. Godine 1992. upisala sam studij Molekularne biologije na Prirodoslovno-matemati kom fakultetu Sveu iliata u Zagrebu. Godine 1994. dodijeljena mi je dvogodianja stipendija grada Zagreba za 50 najboljih studenata. Dobitnica sam Rektorove nagrade za godinu 1994. i 1996. Diplomirala sam 1996. godine s izvrsnim uspjehom. Sljedee sam godine upisala poslijediplomski studij Molekularne i stani ne biologije na Prirodoslovno-matemati kom fakultetu. Od 1997. godine radim kao znanstvena novakinja na Zavodu za biokemiju i molekularnu biologiju Farmaceutsko-biokemijskog fakulteta Sveu iliata u Zagrebu. Tri mjeseca (od 18. sije nja 1999. do 7. o~ujka 1999. godine, te od 31. sije nja 2000. do 6. o~ujka 2000. godine) provela sam na studijskom boravku u Department of Pharmacology and Neuroscience, University of Dundee, u Dundeeju, `kotska, Velika Britanija, u sklopu ostvarivanja znanstvene suradnje. Akademski stupanj magistra prirodnih znanosti iz podru ja biologije stekla sam 17. studenog 2000. godine. Sudjelovala sam na brojnim domaim i meunarodnim znanstvenim skupovima i te ajevima. lanica sam Hrvatskog druatva za biokemiju i molekularnu biologiju. Autor sam sljedeih znanstvenih radova citiranih u CC/SCI: Dumi J., Dabeli S. and Flgel M. (2004) Sweet attachment to molecular biology. Period. biol. 106(2): 127-133. Dabeli S., Flgel M., Maravi G. and Lauc G. (2004) Stress Causes Tissue-Specific Changes in the Sialyltransferase Activity. Z. Naturforsch. 59:276-280. Dumi J., Dabeli S. and Flgel M. (2002) Curcumin  a Potent Inhibitor of Galectin-3 Expression. Food Technol. Biotechnol. 40: 281-287. Lauc G., Peter-Katalini J., Dabeli S. and Flgel M. (1999) Purification and MALDI-MS characterization of a stress - associated glycoprotein from sera of professional soldiers. Biol. Chem. 380:443-450. Lauc G., Dabeli S., `upraha S., and Flgel M. (1998) Glycobiology of stress. Ann. N. Y. Acad. Sci. 851:397-403. Lauc G., Dabeli S., Dumi J. and Flgel M. (1998) Stressin and natural killer cell activity in professional soldiers. Ann. N. Y. Acad. Sci., 851:526-530. 1. Uvod PAGE  PAGE 22  2. Materijal i metode PAGE 56 PAGE 83  3. Rezultati 4. Rasprava PAGE 1 PAGE 94  5. Zaklju ci PAGE 97  6. Literatura 7. Popis kratica i simbola PAGE 110  8. Popis slika i tablica PAGE 114  9. }ivotopis PAGE 117  } } c) b) a) vrijeme (sekunde) vrijeme (sekunde) vrijeme (sekunde) (+141) (+1a)(+1b) AP-2 NF- B NF- B AP-4 AP-1 AP-1/bHLH box AP-1/SIE box GC Box 5 GC Box 4 GC Box 3 GC Box 2 GC Box 1 CRE CRE citoplazma jezgra 0 P TF MAPK RSK MSK MAPKAPK2 MAPKAPK3 MAPK MAPKK MAPKKK ERK JNK p38 MKK3 MKK8 MKK7 MKK4 MEK1 MEK2 MAPK ASK1 MLK MEKK Raf CRE CRE CRE Galektini 4 6 8 9 12 Galektin 3 Galektini 1 2 5 7 10 11 13 14 CRD CRD CRD CRD Tandem-tip Kimera-tip Proto-tip gal-3 bp 200 300 400 500 3 2 1 medij/0,2% DMSO medij THP-1 u mediju medij medij/ 0,2% DMSO THP-1 u mediju/ 0,2% DMSO a) b) } P degradacija posredovana proteosomom vezanje koaktivatora i korepresora modifikacija kromatina P vezanje DNA A R TF promjena konformacije TF -aktin ishodnice 500 400 300 200 3 1 bp 2 gal-3 -aktin a) b) 3600 2700 1800 900 0 150 200 250 300 340 350 400 450 490 500 100 75 0 medij medij / 0,2% DMSO a) medij b) medij/0,2% DMSO gal-3 -aktin K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h kontrola medij medij / 0,2% DMSO c) 0 K 0 * kontrola 0,1 mM ASA 0,5 mM ASA 1 mM ASA 2 mM ASA * * * * * * * * * * * * * * * * * * * * a) 0,1 mM ASA b) 0,5 mM ASA gal-3 -aktin K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h c) 1 mM ASA d) 2 mM ASA gal-3 -aktin K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h e) kontrola 0,1 mM ASA 0,5 mM ASA 1 mM ASA 2 mM ASA * * * * K 0.0 * * * kontrola 0,1 M Ind 1 M Ind 5 M Ind 10 M Ind 3 * * * * * * * * * * * * * * * * * * * * * * * * gal-3 -aktin K 1h 3h 5h 24h 48h 72h a) 0,1 M Ind b) 1 M Ind gal-3 -aktin K 1h 3h 5h 24h 48h 72h c) 5 M Ind d) 10 M Ind -aktin gal-3 K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h e) kontrola 0,1 M Ind 1 M Ind 5 M Ind 10 M Ind * * * * * * 0.0 K 0.0 kontrola 0,1 M HC 1 M HC 5 M HC 10 M HC * * * * * * * * * * * * * * * * * * * * * * * * a) 0,1 M HC b) 1 M HC K 1h 3h 5h 24h 48h 72h gal-3 -aktin gal-3 -aktin K 1h 3h 5h 24h 48h 72h c) 5 M HC d) 10 M HC K 1h 3h 5h 24h 48h 72h gal-3 -aktin gal-3 -aktin K 1h 3h 5h 24h 48h 72h 0.0 e) * * * * * * * * K kontrola 0,1 M HC 1 M HC 5 M HC 10 M HC 0.0 * * * * * * * * * * * * * * * * * * * * * * * * kontrola 0,1 nM Dex 1 nM Dex 10 nM Dex 100 nM Dex a) 0,1 nM Dex b) 100 nM Dex K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h gal-3 -aktin c) 10 nM Dex d) 100 nM Dex K 1h 3h 5h 24h 48h 72h gal-3 -aktin K 1h 3h 5h 24h 48h 72h gal-3 -aktin e) 0 * * * * * * * * K kontrola 0,1 nM Dex 1 nM Dex 10 nM Dex 100 nM Dex 0 kontrola 10-7 M PMA * * gal-3 -aktin K 1h 3h 5h 24h 48h a) b) 0 kontrola 10-7 M PMA * * 0 kontrola (10-7 M PMA) medij / 0,2% DMSO medij a) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h b) * 0 medij medij / 0,2% DMSO kontrola 2 (10-7 M PMA) kontrola 1 c) kontrola 2 mM ASA 0,5 mM ASA 0 * * * * a) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h b) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h c) kontrola 1 kontrola 2 (10-7 M PMA) 2 mM ASA 0,5 mM ASA 0 * * * * 0 * kontrola 10 M Ind 1 M Ind a) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h b) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h c) kontrola 1 kontrola 2 (10-7 M PMA) 10 M Ind 1 M Ind 0 * * * * * ** ** * * * * * * 0 kontrola 10 M HC 1 M HC ** ** a) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h b) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h kontrola 1 kontrola 2 (10-7 M PMA) 10 M HC 1 M HC 0 c) * * * * * * * * * * ** ** ** 0 * * kontrola 100 nM Dex 1 nM Dex a) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h b) gal-3 -aktin K1 K2 1h 3h 5h 24h 48h 72h 0 * * * * * * * * * * kontrola 1 kontrola 2 (10-7 M PMA) 100 nM Dex 1 nM Dex c) b) a) diferencirane stanice nediferencirane stanice  EMBED ISISServer  0 2 1 relativne jednice vrijeme 72 h 48 h 24 h 5 h 3 h 1 h 1.2 1.0 0.8 0.6 0.4 0.2 0.0 (% od kontrole) relativna apsorbancija vrijeme 72 h 48 h 24 h 100 80 60 40 20 0 4 5 6 7 1 h 3 h 5 h 24 h 48 h 72 h vrijeme relativne jedinice 0.2 0.4 0.6 0.8 1 1.2 1.4 1 h 3 h 5 h 24 h 48 h 72 h vrijeme relativne jedinice 0.2 0.4 0.6 0.8 1 1.2 1 h 3 h 5 h 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