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  • Plant Biochemistry

    Hans-Walter Heldt

    in cooperation with Fiona Heldt

    An update and translation of the German third edition

    AMSTERDAM BOSTON HEIDELBERG LONDONNEW YORK OXFORD PARIS SAN DIEGO

    SAN FRANCISCO SINGAPORE SYDNEY TOKYO

  • Acquisition Editor David CellaProject Manager Justin PalmeiroEditorial Coordinator Kelly SonnackMarketing Manager Linda BeattieCover Design Cate BarrComposition SNP Best-set Typesetter Ltd., Hong KongPrinter Courier

    Elsevier Academic Press200 Wheeler Road, Burlington, MA 01803, USA525 B Street, Suite 1900, San Diego, California 92101-4495, USA

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    ISBN: 0-12-088391-033

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    Printed in the United States of America04 05 06 07 08 09 9 8 7 6 5 4 3 2 1

  • Dedicated to my teacher Martin Klingenberg

  • Preface

    This textbook is written for students and is the product of more than threedecades of my teaching experience. It intends to give a broad but conciseoverview of the various aspects of plant biochemistry, including molecularbiology. I have attached importance to an easily understood description ofthe principles of metabolism but also have restricted the content in such a way that a student is not distracted by unnecessary details. In view ofthe importance of plant biotechnology, industrial applications of plant biochemistry have been pointed out wherever appropriate. Thus, specialattention has been given to the generation and utilization of transgenicplants.

    Since there are many excellent textbooks on general biochemistry, I havedeliberately omitted dealing with elements such as the structure and func-tion of amino acids, carbohydrates, and nucleotides; the function of nucleicacids as carriers of genetic information; and the structure and function ofproteins and the basis of enzyme catalysis. I have dealt with topics of generalbiochemistry only when it seemed necessary for enhancing understanding ofthe problem in hand. Thus, this book is, in the end, a compromise betweena general textbook and a specialized textbook.

    This book is a translation of the third German edition. Compared to theprevious English edition, it has been fully revised to take into account theprogress in the field. In particular, the chapter about phytohormones hasundergone extensive changes. As a year has passed since the third Germanedition was completed, the text has been thoroughly updated, taking intoaccount the rapid development of the discipline in 2003.

    I am very grateful to the many colleagues who, through discussions andthe reprints they have sent me on their special fields, have given me the infor-mation that made it possible for me to write the various editions of thisbook. It was especially helpful for me that the colleagues in the followinglist went to great trouble to read critically one or more chapters, to draw myattention to mistakes, and to suggest improvements for which I am particu-larly grateful. My special thanks go to my coworker and wife, Fiona Heldt.Without her intensive support, it would not have been possible for me towrite this book or to prepare the English version.

  • I have tried to eradicate as many mistakes as possible, probably not withcomplete success. I am therefore grateful for any suggestions and comments.

    Hans-Walter HeldtGttingen, January 2004

    Thanks to the following colleagues for their suggestions after looking throughone or more chapters of the various editions of this book:

    Prof. Jan Anderson, Canberra, AustraliaProf. John Andrews, Canberra, AustraliaProf. Tom ap Rees, Cambridge, Great BritainProf. Kozi Asada, Uji, Kyoto, JapanDr. Tony Ashton, Canberra, AustraliaProf. Murray Badger, Canberra, AustraliaProf. Peter Bger, Konstanz, GermanyDr. Sieglinde Borchert, Gttingen, GermanyProf. Peter Brandt, Berlin, GermanyProf. Axel Brennicke, Ulm, GermanyProf. David Day, Perth, AustraliaProf. Karl-Josef Dietz, Bielefeld, GermanyDr. Wolfgang Drge-Laser, Gttingen, GermanyProf. Gerry Edwards, Pullman, Washington, USAProf. Walter Eschrich, Gttingen, GermanyProf. Ivo Feuner, Gttingen, GermanyProf. Ulf-Ingo Flgge, Cologne, GermanyProf. Margrit Frentzen, Aachen, GermanyProf. Wolf Frommer, Tbingen, GermanyDr. R.T. Furbank, Canberra, AustraliaProf. Christine Gatz, Gttingen, GermanyProf. Curtis V. Givan, Durham NH, USAProf. Gowindjee, Urbana, Ill. USAProf. Jan Eiler Graebe, Gttingen, GermanyProf. Peter Grber, Stuttgart, GermanyProf. Dietrich Gradmann, Gttingen, GermanyProf. Erwin Grill, Munich, GermanyDr. Bernhard Grimm, Gatersleben, GermanyProf. Wolfgang Haehnel, Freiburg, GermanyDr. Iris Hanning, Munich, GermanyDr. Marshall D. Hatch, Canberra, AustraliaProf. Ulrich Heber, Wrzburg, Germany

    viii Preface

  • Prof. Dieter Heineke, Gttingen, GermanyDr. Klaus-Peter Heise, Gttingen, GermanyProf. E. Heinz, Hamburg, GermanyDr. Frank Hellwig, Gttingen, GermanyDr. Gieselbert Hinz, Gttingen, GermanyProf. Steven C. Huber, Raleigh, USADr. Graham Hudson, Canberra, AustraliaDr. Colin Jenkins, Canberra, AustraliaProf. Wolfgang Junge, Osnabrck, GermanyProf. Werner Kaiser, Wrzburg, GermanyDr. Steven King, Canberra, AustraliaProf. Martin Klingenberg, Munich, GermanyProf. Gotthard H. Krause, Dsseldorf, GermanyDr. Silke Krmer, Osnabrck, GermanyProf. Werner Khlbrandt, Heidelberg, GermanyDr. Toni Kutchan, Munich, GermanyProf. Hartmut Lichtenthaler, Karlsruhe, GermanyDr. Gertrud Lohaus, Gttingen, GermanyProf. Ulrich Lttge, Darmstadt, GermanyDr. John Lunn, Canberra, AustraliaProf. Enrico Martinoia, Neuchatel, SwitzerlandProf. Hartmut Michel, Frankfurt, GermanyProf. K. Mntz, Gatersleben, GermanyProf. Walter Neupert, Munich, GermanyProf. Lutz Nover, Frankfurt, GermanyProf. C. Barry Osmond, Canberra, AustraliaDr. Katharina Pawlowski, Gttingen, GermanyProf. Birgit Piechulla, Rostock, GermanyProf. Andrea Polle, Gttingen GermanyProf. Klaus Raschke, Gttingen, GermanyDr. Gnter Retzlaff, Limburger Hof, GermanyDr. Sigrun Reumann, Gttingen, GermanyDr. Gerhard Ritte Potsdam, GermanyProf. Gerhard Rbbelen, Gttingen, GermanyProf. David Robinson, Heidelberg, GermanyProf. Eberhard Schfer, Freiburg, GermanyProf. Dierk Scheel, Halle, GermanyProf. Hugo Scheer, Munich, GermanyProf. Renate Scheibe, Osnabrck, GermanyProf. Ahlert Schmidt, Hannover, GermanyDr. Karin Schott, Gttingen, GermanyDr. Ulrich Schreiber, Wrzburg, Germany

    Preface ix

  • Dr. Danja Schnemann, Aachen, GermanyProf. Gernot Schultz, Hannover, GermanyProf. Jens D. Schwenn, Bochum, GermanyProf. Jrgen Soll, Kiel, GermanyProf. Martin Steup, Potsdam, GermanyProf. Heinrich Strotmann, Dsseldorf, GermanyProf. Gerhard Thiel, Darmstadt, GermanyProf. Rudolf Tischner, Gttingen, GermanyProf. Achim Trebst, Bochum, GermanyProf. Hanns Weiss, Dsseldorf, GermanyProf. Dietrich Werner, Marburg, GermanyProf. Peter Westhoff, Dsseldorf, Germany

    x Preface

  • Contents

    1 A leaf cell consists of several metabolic compartments 1

    1.1 The cell wall gives the plant cell mechanical stability 4The cell wall consists mainly of carbohydrates and proteins 4Plasmodesmata connect neighboring cells 7

    1.2 Vacuoles have multiple functions 91.3 Plastids have evolved from cyanobacteria 101.4 Mitochondria also result from endosymbionts 151.5 Peroxisomes are the site of reactions in which toxic intermediates

    are formed 161.6 The endoplasmic reticulum and Golgi apparatus form a network

    for the distribution of biosynthesis products 181.7 Functionally intact cell organelles can be isolated from plant

    cells 221.8 Various transport processes facilitate the exchange of metabolites

    between different compartments 241.9 Translocators catalyze the specific transport of substrates and

    products of metabolism 26Translocators have a common basic structure 29Aquaporins make cell membranes permeable for water 31

    1.10 Ion channels have a very high transport capacity 321.11 Porins consist of b-sheet structures 37

    Further reading 40

    2 The use of energy from sunlight by photosynthesis is the basis of life on earth 45

    2.1 How did photosynthesis start? 452.2 Pigments capture energy from sunlight 47

    The energy content of light depends on its wavelength 47Chlorophyll is the main photosynthetic pigment 49

    2.3 Light absorption excites the chlorophyll molecule 52The return of the chlorophyll molecule from the first singlet state to the

    ground state can proceed in different ways 55

  • 2.4 An antenna is required to capture light 56How is the excitation energy of the photons, which have been captured

    in the antennae, transferred to the reaction centers? 58The function of an antenna can be illustrated using the antenna of

    photosystem II as an example 59Phycobilisomes enable cyanobacteria and red algae to carry out

    photosynthesis even in dim light 62Further reading 66

    3 Photosy