Mouse&Models&for&Studies&of&Redox&Systems&
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1 Mouse&Models&for&Studies&of&Redox&Systems& Thursday, 5 June 2014 Course: Redox Regulation, Oxidative Stress, and Selenoproteins Karolinska Institutet, Stockholm, 2-6 June 2014 Ed Schmidt, Professor Department of Microbiology & Immunology Montana State University, USA
2 Mouse&Models&for&Studies&of&Redox&Systems:&Outline!!! I. Mouse!Technologies!!A.!Muta3ons!types!!B.!Cre/loxP!technology!!working!with!condi3onal!alleles!!! II.!!!Redox!System!Interac3ons!!C!Complementa3on!between!Trx,!GSH,!and!other!systems!in!hepatocytes:!using!mouse!models! to!unveil!an!obscure!source!of!reduc3on!power!
3 Mouse Models of Redox Biology Among!the!most!important!tools!for!understanding!the!roles!of!genes!and!proteins!in! mammalian!systems!are!mouse!models!! One!of!the!most!valuable!resources!for!designing,!using,!or!evalua3ng!any!mouse! model,!gene,!or!phenotype!is!jackson!lab s!mgi!database!!
4 Jax MGI home-page
5 Allelic&quality& WildCtype!( + ).!Unmutated.! Null!( C )!No!ac3vity.!! Hypomorph.!Has!par3al!or!reduced!ac3vity.! GainCofCfunc3on.!New!ac3vity.!Might!be!inten3onal!or!accidental.! KnockCout.!Most!but!not!all!knockCouts!are!simple!nulls.!Evaluate!allelic!design! carefully.!!look!for!controls!and!valida3on.!exci3ng!new!technologies,!such!as! CRISPER!or!TALENs!bring!increases!in!efficiency,!but!also!limita3ons!and/or! restric3ons!concerning!what!allelic!quality!is!achieved.! KnockCin.!Simple!knockCins!disrupt!the!targeted!gene.!Fusion!knockCins!(e.g.,!most! genectraps )!!might!retain!none,!some,!or!most!ac3vi3es!of!the!endogenous!locus,! but!must!be!cri3cally!validated.!irescdriven!second!cistrons!o]en!have!very!li^le! impact!on!the!endogeous!ac3vity.! Condi3onal!( floxed ).!Change!quality,!for!example!from!func3onally!wildCtype!to! func3onally!null,!under!some!condi3on.!most!commonly,!this!is!driven!by!cre! recombinase.!
6 Cre/loxP! floxed! loxp$ Cre! loxp$ +! Cre! +!
7
8 Using&Mouse&Models&to&Unveil&an&Obscure&Source&of&Reduc=on&Power:& & An&ElectronAHunt& & & &
9 Living!cells!of!aerobic!species!maintain!a!reducing!intracellular!milieu! although!inhabi3ng!an!oxidizing!environment! ox! ox! red! ox! red! ox! red! ox! ox! red! ox! red! red! ox! ox! red! red! ox! ox!
10 Where!does!intracellular!reduc3on!power!come!from?! Cysteine! 2 Cys3ne! 2! 2 GSH!System! Trx!System! Thiol! Biology! Protein!disulfide!reduc3on! Peroxidases! Ribonucleo3de!reductase! Methionine!sulfoxide!reduc3on! Cys3ne!reduc3on!!
11 Where!does!intracellular!reduc3on!power!come!from?! Food! GK! e G6PDH! 6PGDH! Glucose! C! G6P! ec! 6PGluconate! ec! R5P! ATP! ADP! NADP +! NADPH! NADP +! NADPH! GSH!System! Trx!System! Protein!disulfide!reduc3on! Peroxidases! Ribonucleo3de!reductase! Methionine!sulfoxide!reduc3on! Cys3ne!reduc3on!!
12 Glutathione! redox!recycling! Reduced!Glutathione! (GSH)! Various! 2 Oxidized!Glutathione! (GSSG)! 2! Gsr! NADP +! 2 NADPH! Glucose! Food! Pentose!Phosphate!Pathway!
13 NADP+ Txnrd1 TrxR1! Trx1 Txn SH SH e C! e C! Thioredoxin!System! e C! e C! NADPH + H + S-S Txnrd1 Cycle enzyme-substrate, oxidized Trx1 Txn enzyme-product, reduced! Enzymes: RNR:!rNDP!+!2e ribonucleotide C!!!!!!!!dNDP! Prx:!XOOH!+!2e reductase peroxiredoxins C!!!!!!!!XOH!+!H 2 O! (Prx) Reactions: Etc.! rndp + H+ H 2 O 2 + 2H+ e C! e C! Enzymes!and!reac3ons! Msr:!RCS=O!+!2e C!!!!!!!RCSCCH 3!+!H 2 O! \&!!!!!!!!!!!!!!CH 3! Other:!RSCSR!+!2e C!!!!!!!2RSH!! dndp + H 2 O 2H 2 O
14 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! TOXICOLOGICAL SCIENCES 82, (2004) doi: /toxsci/kfh268 Advance Access publication September 1, 2004 Analyses of Glutathione Reductase Hypomorphic Mice Indicate a Genetic Knockout Lynette K. Rogers, 1 Toshiya Tamura, Bryan J. Rogers, Stephen E. Welty, Thomas N. Hansen, and Charles V. Smith Center for Developmental Pharmacology and Toxicology, Children s Research Institute, Columbus, Ohio Conclusion: The healthy phenotype exhibited by the [Gsr-null] mice offers clear evidence that [Gsr] is not necessary for viability. Received July 22, 2004; accepted August 23, 2004
15 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! redox& homeostasis& redox& homeostasis&
16 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! TOXICOLOGICAL SCIENCES 82, (2004) doi: /toxsci/kfh268 Advance Access publication September 1, 2004 Analyses of Glutathione Reductase Hypomorphic Mice Indicate a Genetic Knockout Lynette K. Rogers, 1 Toshiya Tamura, Bryan J. Rogers, Stephen E. Welty, Thomas N. Hansen, and Charles V. Smith Center for Developmental Pharmacology and Toxicology, Children s Research Institute, Columbus, Ohio Conclusion: The healthy phenotype exhibited by the [Gsr-null] mice offers clear evidence that [Gsr] is not necessary for viability. Received July 22, 2004; accepted August 23, 2004 Free Radical Biology & Medicine 43 (2007) Original Contribution Effects of thioredoxin reductase-1 deletion on embryogenesis and transcriptome Alla A. Bondareva a,1, Mario R. Capecchi b, Sonya V. Iverson a, Yan Li a, Nathan I. Lopez c, Olivier Lucas a, Gary F. Merrill c, Justin R. Prigge a, Ashley M. Siders a, Maki Wakamiya d, Stephanie L. Wallin a, Edward E. Schmidt a,e, a VMB, Molecular Biosciences, 960 Technology Blvd., Montana State University, Bozeman, MT 59718, USA b Howard Hughes Medical Institute, Eccles Institute of Human Genetics, University of Utah, Salt Lake City, UT 84112, USA c Biochemistry and Biophysics Department, ALS2011, Oregon State University, Corvallis, OR 97331, USA d Department of Neurology, UTMB, Clay Hall Rm , 301 University, Galveston, TX 77555, USA e Center for Reproductive Biology, Washington State University, Pullman, WA 99164, USA Conclusion: Our!results!suggest!a!role!for![TrxR1]! in!allowing!correct!developmental!pa^erning!and! indicate!that!general&cell&physiological&processes& are¬&precluded&by&disrup=on&of&txnrd1. Received 2 February 2007; revised 4 May 2007; accepted 19 May 2007 Available online 31 May 2007
17 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! redox& homeostasis& redox& homeostasis&
18 Txnrd1*/*$hepatocytes!are!fully!replica3onCcompetent!during!regenera3on! in!adult!txnrd1cond/*;albcre1$mice$ Rollins et al., J. Cell Sci. 2010
19 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! ESSENTIAL redox& homeostasis& redox& homeostasis&
20 Glutathione!Biosynthesis! BSO& Reduced!Glutathione! (GSH)! Saline! BSO! Prigge et al., FRBM. 2012
21 808 J.R. Prigge et al. / Free Radical Biology & Medicine 52 (2012) in which very few hepatocytes remained replicative (Fig. 3R and SupReplica3on!in!hepatocytes!requires!either!a!single!copy!of! plementary Figs. S1C and S1D). Quantitative analysis of the proportion of proliferative hepatocyte nuclei under each condition that could txnrd1!or!glutathione! replication under each condition (Figs. 3M 3R, and Supplementary Fig. S1). Results showed similar levels of hepatocyte replication under all conditions except for the txnrd1 cond/null;albcre1 mice treated with BSO, This!shows!complementa3on!between!the!Trx!and!the!GSH! redox!systems!in!hepatocytes! Prigge et al., FRBM. 2012
22 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! BSO! redox& homeostasis& redox& homeostasis&
23 How!redundant!are!the!GSH!and!the!Trx!systems!in!mammals?! redox& homeostasis& redox& homeostasis&
24 redox& homeostasis& redox& homeostasis&
25 Carb Biology e Food! C! e Glucose! C! ec! Thiol Biology redox& homeostasis& redox& homeostasis&
26 TR/GR-null livers are truly TrxR1-deficient MW! 175! 80! An3CTrxR1! 58! 46! An3CTrx1! Dr. Sofi Eriksson
27 TR/GR-null livers truly lack GSSG-reductase activity GSSG-reductase activity (µmol/min/mg*protein)* 40* 20* 0* Gsr: Txnrd1: +/+ +/+ +/+ -/- -/- -/- +/+ +/- -/- +/+ +/- -/- Dr. Sofi Eriksson
28 Carb Biology e Food! C! e Glucose! C! ec! An electron hunt Glutathione! Thiol Biology redox& homeostasis& redox& homeostasis&
29 TR/GR-null livers have wild-type levels of total GSH+GSSG, but perhaps a slightly higher ratio of GSSG:GSH Dr. Sofi Eriksson
30 Do the TR/GR-null livers really require GSH? BSO& Glutathione!Biosynthesis! C survival (percent)! BSO 0-0! 12! control (n = 6) $ Txnrd1 -/- ;Gsr -/- (n = 12) $ 24! 36! time (hours)! //! //! 120!
31 ! An electron hunt GSSG! Glutathione! Thiol Biology redox& homeostasis& redox& homeostasis&
32 ! GSH biosynthesis An electron hunt GSSG! Glutathione! Thiol Biology redox& homeostasis& redox& homeostasis&
33 2! Hypothesis:!Cysteine!is!the!only!source!of! reducing!poten3alfor!the!tr/gr-null!livers!! Reduced!Glutathione! (GSH)! e - e - e - NO&REDUCTIVE&RECYCLING&OF&GSSG& Oxidized!Glutathione! (GSSG)! Various!!redox! homeostasis! 2e -
34 Cysteine Where does intracellular reduction power come from? Cysteine! Cys3ne! 2! 2 GSH!System! Trx!System! NADP +! NADPH! Pentose!Phosphate! Pathway! GSSG! Glucose! Food!
35 Met (e - )! Trans-sulfuration Met (e - )! (e- )! SAM Homocysteine e -! (e - )! SAH Cystathionine (e - )! αkb e -! Cys
36 Met (e - )! Trans-sulfuration Met (e - )! (e- )! SAM Homocysteine e -! (e - )! SAH Cystathionine (e - )! αkb e -! Cys
37 Met (e - )! Trans-sulfuration Met (e - )! (e- )! SAM Homocysteine e -! (e - )! SAH PPGG Cystathionine (e - )! αkb e -! Cys
38 Conversion of cystathionine to Cys is required for survival of TR/GR-null livers PPGG control (n = 6) $ //! Txnrd1 -/- ;Gsr -/- (n = 8) $ ! 12! 24! 36! time (hours)! //! 120!
![Liver Label: [35S]Met M c!](/docs-images/91/107205936/images/39-0.jpg "Serum [35S]Met [35S]cystine -/- -/- +/+")
![Txnrd: X M! Label: [35S]M!](/docs-images/91/107205936/images/39-2.jpg "Gsr: Txnrd: -/-/- [35S]cystine -/- -/+/-")
39 Liver Label: [35S]Met M c! Serum [35S]Met [35S]cystine -/- -/- +/+ +/+ +/+ -/- -/- +/+ +/+ +/+ -/- -/- -/- +/- -/- +/+/+ +/+ -/- +/- -/- -/- Gsr: Txnrd: X M! Label: [35S]M! Gsr: Txnrd: -/-/- [35S]cystine -/- -/+/- +/- -/-/- -/-/- Mw Met Met Liver - 30 * - cystine Origin - *! *- GSSG - Origin *! -7 Serum GSH
40 Met (e - )! (3 ATP) + Ser Cost analysis Food (e - )! Met transsulfuration Cost: 5ATP + Met + Glu + Gly - αkb! Cys αkb! GSHbiosynthesis 2 ATP + Glu + Gly e -! GSH! ½(NADPH) Glucose Pentose Phosphate Pathway Cost: ½ NADPH ½(NADPH) ½(NADP + ) : - 2ATP-equivalents/mol e - Met + Ser + Glu + Gly GSH + αkb : - 5ATP-equivalents/mol e - αkb BCKDC*! ATP! Succinyl CoA! GTP + NADH + QH 2 : + 24ATP-equiv/mol e - GDP, NAD +, Q! *branched-chain α-ketoacid dehydrogenase complex
41 Dietary methionine provides the reducing-potential and sulfur-amino acids that allow survival of TR/GR-null livers in Gsr-null mice
42 Normal Hepatocyte! TR/GR-null Hepatocyte! e -! Food (any) Met Food (any) e -! Met cystine GSSG Glucose PPP e -! Gcl e -! e -! Gsr GSH NADPH e -! energy TrxR1 Trx1 Cys dntps redoxhomeostasis Met SAM 1-C metab protein GSSG e -! synth e -! e -! DNA synth GSSG Glucose PPP NADPH Cys e-! e -! GSH dntps redoxhomeostasis energy e -! protein synth Met DNA synth e -! SAM 1-C metab e -! e -! H 2 O! e -! H 2 O!
43 Can&Exogenous&An=oxidants&Support&Redox&Homeostasis?& Vitamin C Vitamin E Vitamin A Lycopene Green tea Selenium NAC GSH Polyphenols Lipoic acid Uric acid Carrotene Ubiquinol Tannins Methionine
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