POST-TRANSCRIPTIONAL CONTROL OF GENE EXPRESSION IN PLANTS Post-Transcriptional Control of Gene Expression in Plants Edited by Witold Filipowicz and Thomas Hahn Reprinted from Plant Molecular Biology, Vol. 32(1, 2), 1996 KLUWER ACADEMIC PUBLISHERS DORDRECHT/BOSTON/LONDON Library of Congress Cataloging-in-Publication Data ISBN-13: 978-94-01 0-6644-0 e-ISBN-13: 978-94-009-0353-1 DOl: 10.1007/978-94-009-0353-1 Published by Kluwer Academic Publishers, P.O. Box 17,3300 AH Dordrecht, The Netherlands Kluwer Academic Publishers incorporates the publishing programmes of D. Reidel, Martinus Nijhoff, Dr. W. Junk and MTP Press. Sold and distributed in the U.S.A. and Canada by Kluwer Academic Publishers, 101 Philip Drive, Norwell, MA 02061, U.S.A. In all other countries, sold and distributed by Kluwer Academic Publishers Group, P.O. Box 322, 3300 AH Dordrecht, The Netherlands. Printed on acid-free paper All Rights Reserved © 1996 Kluwer Academic Publishers Softcover reprint of the hardcover 1s t edition 1996 No part of the material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner. CONTENTS Preface i-ii Dedication iii List of Contributors iv-viii RNA processing and stability Splicing of precursors to messenger RNA in higher plants: mechanism, regulation and sub-nuclear organisation of the spliceosomal machinery G.G. Simpson, W. Filipowicz 1-41 Plant mRNA 3/-end formation H.M. Rothnie 43-61 Control of mRNA stability In higher plants M.L. Abler, P.J. Green 63-78 RNA as a target and an initiator of post-transcriptional gene silencing in transgenic plants D.C. Baulcombe 79-88 RNA structure and regulation of gene expression P. Klatt, D. Riesner, G. Steger 89-106 Trans/ation The plant translational apparatus K.S. Browning 107-144 Translational control of cellular and viral mRNAs D.R Gallie 145-158 Translation in plant: rules and exceptions J. Futterer, T. Hohn 159-189 Fate of translation products Molecular chaperones and protein folding in plants RS. Boston, P. Viitanen, E. Vierling 191-222 Transport of proteins in eukaryotic cells: more questions ahead M. Bar-Peled, D.C. Bassham, N.V. Raikhel 223-249 Plasmodesma I cell-to-cell transport of proteins and nucleic acids L.A. Mezitt, w.J. Lucas 251-273 Proteolysis in plants: mechanisms and functions RD. Vierstra 275-302 Organelles Regulation of gene expression in plant mitochondria S. Binder, A. Marchfelder, A. Brennicke 303-314 Regulation of gene expression in chloroplasts of higher plants M. Sugita, M. Sugiura 315-326 Post-transcriptional regulation of chloroplast gene expression in Chlamydomonas reinhardtii J.-D. Rochaix 327-341 RNA editing in plant mitochondria and chloroplasts A.M. Maier, P. Zeitz, H. Kossel, G. Bannard, J.M. Gualberto, J.M. Grienenberger 343-365 Special topicS Gene expression from viral RNA genomes I.G. Maia, K. Seran, A.-L. Haenni, F. Bernardi 367-391 Optimizing expression of transgenes with an emphasis on post-transcriptional events M.G. Koziel, N.B. Carazzi, N. Desai 393-405 Subject index 407-414 Plasma Cells in I/ membrane"I ~cells.in ~ 3t~ ~ ~' DomalO-2 2222222 ........ -----. :.... ~ Cell Wall For details see Mezitt and Lucas, pp. 251-273 Plant Molecular Biology 32: i-ii, 1996. Preface A recent volume in this series (Signals and Signal Transduction Pathways in Plants (K. Palme, ed.) Plant Mol. BioI. 26, 2137-1679) described the relay races by which signals are transported in plants from the sites of stimuli to the gene expression machinery of the cell. Part of this machinery, the transcription apparatus, has been well studied in the last two decades, and many important mechanisms controlling gene expression at the transcriptional level have been elucidated. However, control of gene expression is by no means complete, once the RNA has been produced. Important regulatory devices determine the maturation and usage of mRNA and the fate of its translation product. Posttranscriptional regulation is especially important for generating a fast response to environmental and intracellular signals. This book opens with chapters describing the processing of mRNAs, i.e. splicing (Simpson and Filipowicz) and polyadenylation (Rothnie). Usually, neither the splicing nor the polyadenylation signals of animals and fungi are recognized efficiently and correctly in plants, indicating that they are not strictly conserved among the eukaryotic kingdoms. In contrast to animal introns, the processing of plant introns depends on the presence of AU- and U-rich clusters. The known plant polyadenylation sites do not include a defined element downstream of the AATAAA motif as found in animals, the AAUAAA motif is not strictly conserved, and upstream elements are important. More and more examples of alternative mRNA splicing and polyadenylation sites are being identified in plants. Degradation of RNA might be as important as RNA production for the well-being of cells. Some mRNAs encoding regulatory proteins must have a high turnover to avoid undesirable overproduction. Short-lived mRNAs have been found to contain cis elements that label the RNA for degradation (Abler and Green). An especially interesting mechanism of mRNA disposal occurs during posttranscriptional silencing (Baulcombe). In this case, RNA that had apparently been produced originally at high excess is degraded specifically. The mechanism of posttranscriptional silencing is basically unknown, difficult to explain in mechanistic terms and might involve events described by chaos theory. Protein synthesis in eukaryotic cells is much more complex than originally thought. More and more factors governing pretranslational processes (initiation complex formation and scanning), the initiation itself, and elongation are becoming known (Browning). Sequences both on the leader and the trailer of mRNAs play important roles in the efficiency and regulation of translation (Gallie). Furthermore, not all eukaryotic mRNAs are strictly monocistronic. Plant viral RNAs, in particular, are frequently provided with complex leaders containing short open reading frames which control translation, and viral RNAs are often polycistronic (Fiitterer and Hohn). In general, strategies of gene expression from viral genomes are unique and frequently also very complex (Maia et al.). For all the properties and functions of viral and cellular mRNAs, their secondary and tertiary structure is very important (Klaff et al.). An increasing number of RNA-binding proteins which specifically recognize different structural elements in RNA are being characterized. The correct folding of many proteins has been found to depend on the action of chaperonins (Boston et al.). While some proteins remain in the cytoplams, others are transported to various subcellular localizations or are secreted (Bar-Peled et al.). A unique property of plant cells is that some proteins, and also nucleic acids, can be actively transported between cells using gates in the cell wall, the plasmodesmata. This novel macromolecular trafficking pathway may playa role in the programming of plant development (Mezitt and Lucas). Like mRNA, turnover, that of proteins is also regulated. Protein degradation in plants involves many proteolytic pathways operating in different cellular compartments (Vierstra). 11 Due to their probably origin from prokaryotes, plant organelle gene expression uses mechanisms more related to those of prokaryotes than those used by the cell for nuclear encoded genes. Regulation of organelle gene expression occurs mainly at the posttranscriptionallevel, i.e. RNA cis and trans splicing, processing ofpolycistronic transcripts (Rochaix; Sugita and Sugiura; Binder et al.) and finally, RNA editing (Maier et al.). The latter process occurs both in mitochondria and chloroplasts but its biochemistry remains largely unknown. From the many examples of posttranscriptional regulation described in this book it is obvious that optimizing gene expression in transgenic plants cannot stop with the design of optimal promoters but must also consider the stability and translatability of the transcripts as well as certain structural properties of the protein products themselves (Koziel et al.). We would like to dedicate this volume to the memory of Hans Kossel, our colleague and friend, and also coauthor of one of the chapters (Meier et al.) of this book, who passed away suddenly in December 1995. For many years Hans Kossel was a Professor of Molecular Biology and Genetics at the University of Freiburg, Ger many. His research focused on various aspects of the chemistry and biology of nucleic acids. His more recent work on the structure and function of chloroplastgenomes culminated in the discovery of RNA editing in these organelles. Witold Filipowicz and Thomas Hohn Guest Editors iii Hans Kossel (1934-1995) Plant Molecular Biology 32: iv-viii, 1996. iv List of Contributors RNA processing and stability Splicing of precursors to messenger RNA in higher plants: mechanism, regulation and sub-nuclear organ PJ. Green isation of the spliceosomal machinery Plant Research Laboratory Michigan State University G.G. Simpson East Lansing, MI 48824-1312 Friedrich Miescher Institute USA P.O. Box 2543 Phone: 1-517-353-4838 CH-4002 Basel Fax: 1-517-353-9168 Switzerland E-mail: [email protected] Phone: 41-61-697-4128 Fax: 41-61-697-3976 RNA as a target and an initiator of post-transcriptional E-mail: [email protected] gene silencing in transgenic plants W. Filipowicz D.C. Baulcombe Friedrich Miescher Institute The Sainsbury Laboratory P.O. Box 2543 Norwich Research Park CH-4002 Basel Colney Switzerland Norwich NR4 7UH Phone: 41-61-697-6993 UK Fax: 41-61-697-3976 Phone: 44-1603-452571 E-mail: filipowi@fmLch Fax: 44-1603-250024 E-mail: [email protected] Plant mRNA 3' -end formation RNA structure and regulation of gene expression H.M. Rothnie Friedrich Miescher Institute P. Klaff P.O. Box 2543 Heinrich-Heine-Universitat Dusseldorf CH-4002 Basel Institut fUr Physikalische Biologie Switzerland Universitatsstrasse 1 Phone: 41-61-697-6684 D-40225 Dusseldorf 1 Fax: 41-61-697-3976 Germany E-mail: [email protected] Phone: 49-211-8115153 Fax: 49-211-8115167 Control of mRNA stability in higher plants D. Riesner M.L. Abler Heinrich-Heine-Universitat Dusseldorf Plant Research Laboratory Institut fur Physikalische Biologie Michigan State University Universitatsstrasse 1 East Lansing, MI 48824-1312 D-40225 Dusseldorf 1 USA Germany Phone: 1-517-353-4838 Phone: 49-211-8115153 Fax: 1-517-353-9168 Fax: 49-211-8115167 v G. Steger Fate of translation products Heinrich -Heine-U niversitat Dusseldorf Institut fUr Physikalische Biologie Molecular chaperones and protein folding in plants Universitatsstrasse 1 D-40225 Dusseldorf 1 R.S. Boston Germany Department of Botany Phone: 49-211-8114840 North Carolina State University Fax: 49-211-8115167 Raleigh, NC 27695 USA Translation P. Viitanen The plant translational apparatus Dupond de Nemours Co. Central Research and Development K.S. Browning Bldg,402 Department of Chemistry Wilmington, DE 19880-0402 University of Texas USA Austin, TX 78712-1096 Phone: 1-302-695-7032 USA Fax: 1-302-695-4296 Phone: 1-512-471-3973 or 4562 Fax: 1-512-471-8696 E. Vierling E-mail: [email protected] Department of Biochemistry University of Arizona Translation control of cellular and viral mRNAs Tucson, AZ 85721 USA D.R. Gallie Phone: 1-520-621-1601 or 3977 Department of Biochemistry Fax: 1-520-621-3709 University of California E-mail: [email protected] Riverside, CA 92521 USA Transport of proteins in eukaryotic cells: more Phone: 1-909-787-7298 questions ahead Fax: 1-909-787-3590 M. Bar-Peled Translation in plant-rules and exceptions Plant Research Laboratory Michigan State University J. Futterer East Lansing, MI 48824-1312 Institute of Plant Sciences USA ETZ Phone: 517-353-3518 Universitatsstrasse 2 Fax: 517-353-9168 CH-8092 Zurich Switzerland D.C. Bassham Fax: 41-1-6321044 Plant Research Laboratory Michigan State University T. Hohn East Lansing, MI 48824-1312 Friedrich Miescher Institute USA P.O. Box 2543 Phone: 517-353-3518 CH-4002 Basel Fax: 517-353-9168 Switzerland Phone: 41-61-6977266 Fax: 41-61-6973976 E-mail: [email protected]