Botany 960:¬Ý Seminar-Plant Physiology¬Ý¬Ý ¬Ý

"Epigenetic mechanisms of gene regulation"

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¬ÝClass meets: Mon 3:30pm-4:30pm in Rm 348 Birge Hall

¬ÝInstructor: Chris Day, 608 262 9749, cday2@wisc.edu

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Class Format:

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1) ¬ÝPresentation:¬Ý At the start of class present the paper for no more than 30 min.

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2)¬Ý Discussion after presentation:¬Ý Whole class will talk about the paper (about 20 min), those auditing are welcome to join in the discussion.

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3)¬Ý Introduction for the following weeks paper (see below).

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Important note:¬Ý One week BEFORE your presentation.¬Ý Present a short introduction of the paper, no more than 5 min.¬Ý Also type out a list of discussion points for people to think about while they read the paper (print enough copies for everyone).

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Week4,¬Ý¬Ý¬Ý week5,¬Ý Week6,¬Ý Week7,¬Ý¬Ý Week8,¬Ý Week9,¬Ý¬Ý Week10, Week11,¬Ý¬Ý Week12,¬Ý Week13,¬Ý¬Ý Week14,¬Ý Week15,¬Ý¬Ý¬Ý Recommended reading

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Week 1:¬Ý Labor day

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Week 2-3:¬Ý Introduction to epigenetics.

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Week 4 (23rd Sept):¬Ý

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Park YD, Papp I, Moscone EA, Iglesias VA, Vaucheret H, Matzke AJ, Matzke MA.

Gene silencing mediated by promoter homology occurs at the level of transcription and results in meiotically heritable alterations in methylation and gene activity.

Plant J. 1996 Feb;9(2):183-94.

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Supplemental material based on class discussion

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Week 5 (30th Sept):¬Ý¬Ý¬Ý

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Wassenegger M, Heimes S, Riedel L, Sanger HL.

RNA-directed de novo methylation of genomic sequences in plants.

Cell. 1994 Feb 11;76(3):567-76.

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¬ÝSupplemental material based on class discussion

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Week 6 (7th Oct):¬Ý¬Ý¬Ý

Questions for week six

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Metzlaff M, O'Dell M, Cluster PD, Flavell RB.

RNA-mediated RNA degradation and chalcone synthase A silencing in petunia.

Cell. 1997 Mar 21;88(6):845-54.

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Week 7 (14th Oct):¬Ý

Questions for week seven

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Mette MF, Aufsatz W, van der Winden J, Matzke MA, Matzke AJ.

Transcriptional silencing and promoter methylation triggered by double-stranded RNA.

EMBO J. 2000 Oct 2;19(19):5194-201.

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¬ÝSupplemental material based on class discussion

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Week 8 (21st Oct):¬Ý¬Ý¬Ý

Questions for week eight

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Llave C, Kasschau KD, Rector MA, Carrington JC.

Endogenous and Silencing-Associated Small RNAs in Plants.

Plant Cell. 2002 Jul;14(7):1605-19.

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Week 9 (28th Oct):¬Ý

Questions for week nine

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Kakutani T, Jeddeloh JA, Flowers SK, Munakata K, Richards EJ.

Developmental abnormalities and epimutations associated with DNA hypomethylation mutations.

Proc Natl Acad Sci U S A. 1996 Oct 29;93(22):12406-11.

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Jeddeloh JA, Stokes TL, Richards EJ.

Maintenance of genomic methylation requires a SWI2/SNF2-like protein.

Nat Genet. 1999 May;22(1):94-7.

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Week 10 (4th Nov):¬Ý

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Volpe TA, Kidner C, Hall IM, Teng G, Grewal SIS and Martienssen RA.

Regulation of Heterochromatic Silencing and Histone H3 Lysine-9 Methylation by RNAi

Science (2002) 297, 1833-1837

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Allshire R

RNAi and Heterochromatin; a Hushed-Up affair

Science (2002) 297, 1818-1819

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Week 11 (11th Nov):¬Ý¬Ý¬Ý

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Jacobsen SE et al.

Ectopic hypermethylation of flower specific genes in Arabidopsis. ¬Ý

Current Biology 2000, 10:179-186.

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Week 12 (18th Nov):¬Ý¬Ý

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Postponed to Dec 2nd.

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Week 13 (25th Nov):¬Ý¬Ý

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Nathan Springer will focus on chromatin and transcriptional regulation but will not address PTGS or RNAi except briefly.¬Ý This will cover many epigenetic phenomena not covered in the course (including paramutation, imprinting and PcG mediated silencing).¬Ý

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Then he will introduce the NSF Functional Genomics of Chromatin project that the Kaeppler lab is involved in. Before class have a look at the www.chromdb.org website to become familiar with what is present in plants.¬Ý Nathan will talk in more detail about the methyltransferases, methyl-binding domain proteins and SET domain proteins in regards to their putative function and their evolution.¬Ý This will lead into a brief discussion about the histone code hypothesis.¬Ý ¬ÝIn this regard you should read the two reviews below before coming to class.

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Jenuwein T. and Allis C.D.

Translating the Histone Code

Science (2001) 293; 1074-1080

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Strahl B.D. and Allis C.D.

The language of covalent histone modifications

Nature (2000) 403; 41-45

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Week 14 (2nd Dec):¬Ý

Questions for week fourteen-

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Chris Pires we will discuss epigenetics in the context of polyploidy.¬Ý Specifically, we will discuss allopolyploidy - which involves the union of two genomes from different species.¬Ý Chris is providing you with two papers.¬Ý The first paper is mandatory:

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Thomas C. Osborn, J. Chris Pires, James A. Birchler, Donald L. Auger, Z. Jeffery Chen, Hyeon-Se Lee, Luca Comai, Andreas Madlung, R.W. Doerge, Vincent Colot, Robert A. Martienssen (in press).¬Ý Understanding mechanisms of novel gene expression in polyploids.¬Ý Trends in Genetics.

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This is a preprint of a paper that was accepted on 21 November, 2002 (it was handed out at class, if you need a copy see Chris or someone who attended class on Nov 25th)¬Ý Please do not distribute it to anyone without permission.¬Ý It reviews most of the recent literature on polyploidy pertaining to gene expression with a focus on rapid genetic and epigenetic changes that occur immediately upon polyploidization.¬Ý The authors are members of a polyploid consortium (see http://polyploid.agronomy.wisc.edu/).

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The second paper is optional but gives nice examples of experimental approaches being used to examine polyploidy and epigenetics.

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A Madlung, RW Masuelli, B Watson, SH Reynolds, J Davidson, L Comai.

Remodeling of DNA Methylation and Phenotypic and Transcriptional Changes in Synthetic Arabidopsis Allotetraploids.

Plant Physiology (2002) 129 (June issue): 733-746.-

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The second paragraph of this paper sets the stage: "a consequence of the union of two genomes, abnormal phenotypes have been reported.¬Ý The causes of these phenotypes are largely unknown. McClintock described similar phenomena as ³ÑÍ߀"genomic shock" -³ÑÈÝ‚ which she defined as a preprogrammed response to an unusual challenge resulting in extensive restructuring of the genome.¬Ý This ³Ñ͵Æ"unusual challenge³ÑÈÝ‚" involve epigenetic gene silencing, which results from homologous DNA-DNA or DNA-RNA interactions.¬Ý The hybridization of redundant and diverged homeolgous sets of genes in allopolyploids might trigger widespread gene silencing and changes in chromatin structure and DNA methylation patterns.³Ñ"

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Week 15 (9th Dec):¬Ý¬Ý

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Compile timeline and decide who should win the Nobel prize.¬Ý Timeline will be unraveled based on the model we discussed in class.¬Ý Listen for venue change in class.

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Recommended reviews:

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Vaucheret H, Fagard M.

Transcriptional gene silencing in plants: targets, inducers and regulators.

Trends Genet. 2001 Jan;17(1):29-35.

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Matzke M, Matzke AJ, Kooter JM.

RNA: guiding gene silencing.

Science. 2001 Aug 10;293(5532):1080-3.

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Voinnet O.

RNA silencing: small RNAs as ubiquitous regulators of gene expression.

Curr Opin Plant Biol. 2002 Oct;5(5):444.

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Martienssen RA, Colot V.

DNA methylation and epigenetic inheritance in plants and filamentous fungi.

Science. 2001 Aug 10;293(5532):1070-4.

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Finnegan EJ.

Epialleles - a source of random variation in times of stress.

Curr Opin Plant Biol. 2002 Apr;5(2):101-6.

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Pikaard CS.

Genomic change and gene silencing in polyploids.

Trends Genet. 2001 Dec;17(12):675-7.