11 Plant Biology 111 111 111 111 11 ii Section K – Lipid metabolism The INSTANT NOTES series Series editor B.D. Hames School of Biochemistry and Molecular Biology, University of Leeds, Leeds, UK Animal Biology Ecology Genetics Microbiology Chemistry for Biologists Immunology Biochemistry 2nd edition Molecular Biology 2nd edition Neuroscience Psychology Developmental Biology Plant Biology Forthcoming titles Bioinformatics The INSTANT NOTES Chemistry series Consulting editor: Howard Stanbury Organic Chemistry Inorganic Chemistry Physical Chemistry Forthcoming titles Analytical Chemistry Medicinal Chemistry 11 Plant Biology 111 A.J. Lack & D.E. Evans School of Biological & Molecular Sciences, Oxford Brookes University, Oxford, UK 111 111 111 11 © BIOS Scientific Publishers Limited, 2001 First published 2001 This edition published in the Taylor & Francis e-Library, 2005. “To purchase your own copy of this or any of Taylor & Francis or Routledge’s collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk.” All rights reserved. No part of this book may be reproduced or transmitted, in any form or by any means, without permission. A CIP catalogue record for this book is available from the British Library. ISBN 0-203-64545-6 Master e-book ISBN ISBN 0-203-68112-6 (Adobe eReader Format) ISBN 1 85996 197 5 (Print Edition) BIOS Scientific Publishers Ltd 9 Newtec Place, Magdalen Road, Oxford OX4 1RE, UK Tel. +44 (0)1865 726286. 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Production Editor: Paul Barlass CONTENTS 11 Abbreviations ix Preface xi Section A – Introduction 1 111 A1 Introduction 1 Section B – Structure 3 B1 The plant cell 3 B2 The cell wall 6 B3 Plastids and mitochondria 9 B4 Membranes 11 B5 Nucleus and genome 15 B6 Cell division 18 111 Section C – Vegetative anatomy 23 C1 Meristems and primary tissues 23 C2 Roots 27 C3 Herbaceous stems and primary growth 31 C4 Woody stems and secondary growth 34 C5 Leaves 37 Section D – Reproductive anatomy 41 D1 The flower 41 D2 Pollen and ovules 46 111 D3 The seed 49 D4 Fruits 52 Section E – Physiology and regulation 55 E1 Arabidopsis and other model plants 55 E2 Methods in experimental plant science 57 Section F – Growth and development 63 F1 Features of growth and development 63 F2 Biochemistry of growth regulation 67 111 F3 Molecular action of hormones and intracellular messengers 76 Section G – Sensing and responding to the environment 83 G1 Phytochrome, photoperiodism and photomorphogenesis 83 G2 Tropisms 87 G3 Nastic responses 91 G4 Abscission 94 G5 Stress avoidance and adaptation 96 Section H – Floral development and reproductive physiology 101 H1 Physiology of floral initiation and development 101 11 H2 Breeding systems 103 vi Contents H3 Self incompatibility 107 H4 Seed development, dormancy and germination 111 Section I – Plants, water and mineral nutrition 115 I1 Plants and water 115 I2 Water retention and stomata 121 I3 Movement of nutrient ions across membranes 124 I4 Uptake of mineral nutrients by plants 128 I5 Functions of mineral nutrients 131 Section J – Metabolism 135 J1 Photosynthetic pigments and the nature of light 135 J2 Major reactions of photosynthesis 138 J3 C3 and C4 plants and CAM 144 J4 Respiration and carbohydrate metabolism 148 J5 Amino acid, lipid, polysaccharide and secondary product metabolism 154 Section K – Plant communities and populations 161 K1 Physical factors and plant distribution 161 K2 Plant communities 167 K3 Ecology of different growth forms 171 K4 Populations 174 K5 Contributions to carbon balance and atmosphere 180 Section L – Reproductive ecology 183 L1 Ecology of flowering and pollination 183 L2 Seed ecology 187 L3 Regeneration and establishment 193 L4 Polymorphisms and population genetics 196 Section M – Interactions between plants and other organisms 201 M1 Mycorrhiza 201 M2 Nitrogen fixation 206 M3 Interactions between plants and animals 209 M4 Fungal pathogens and endophytes 213 M5 Bacteria, mycoplasma, viruses and heterokonts 217 M6 Parasites and saprophytes 219 M7 Carnivorous plants 223 Section N – Human uses of plants 227 N1 Plants as food 227 N2 Plants for construction 232 N3 Plants in medicine 236 N4 Plants for other uses 238 N5 Bioremediation 242 Section O – Plant genetic engineering and biotechnology 245 O1 Plant breeding 245 O2 Plant cell and tissue culture 247 O3 Plant genetic engineering 252 Contents vii 11 Section P – Plant diversity 257 P1 Diversity and life cycles 257 P2 The algae 260 P3 The bryophytes 264 P4 Reproduction in bryophytes 269 Section Q – Spore-bearing vascular plants 273 Q1 Early evolution of vascular plants 273 Q2 Clubmosses and horsetails 278 111 Q3 The ferns 284 Q4 Evolution of the seed 291 Section R – Seed plants 293 R1 Early seed plants 293 R2 Conifers 296 R3 Cycads, gingko and Gnetales 301 R4 Evolution of flowering plants 306 R5 Mechanisms of evolution 314 111 Further reading 317 Index 325 111 111 11 ABBREVIATIONS 11 2,4-D 2,4-dichlorophenoxyacetic acid IP3 inositol triphosphate 2,4,5-T 2,4,5-trichlorophenoxyacetic LDP long day plant acid LSD lysergic acid diethylamine ABA abscisic acid mRNA messenger ribonucleic acid ABP auxin-binding protein NAA naphthalene acetic acid 111 ACC 1-aminocyclopropane-1- NADP nicotinamide adenine carboxylic acid dinucleotide phosphate ACS ACC synthase NADPH nicotinamide adenine ADP adenosine diphosphate dinucleotide phosphate AMP adenosine monophosphate (reduced) AS asparagine synthase NE nuclear envelope ATP adenosine triphosphate NPA 1-N-naphthylphthalamic acid bp base pair NR nitrate reductase CAM crassulacean acid metabolism Pa Pascals CaMPK calmodulin-dependent protein PCR polymerase chain reaction 111 kinases PEP phosphoenolpyruvate CFP cyan fluorescent protein PG polygalacturonase CoA coenzyme A PGS plant growth substances DAG diacylglycerol Pi inorganic phosphate DDT 1,1-bis(p-chlorophenyl)-2,2,2- ppm parts per million trichloroethane PIP2 phosphatidyl inositol DNA deoxyribonucleic acid bisphosphate ems ethylmethanesulfonate RAPD random amplified polymorphic ER endoplasmic reticulum DNA FADH flavin adenine dinucleotide RET resonance energy transfer 111 (reduced) RFLP restriction fragment length Fd ferredoxin-dependent polymorphism GA gibberellic acid RNA ribonucleic acid GA Golgi apparatus RNase ribonuclease GARE gibberellic acid response SAM s-adenosyl methionine element SAUR soybean auxin upregulated gene GDH glutamate dehydrogenase SDP short day plant GFP green fluorescent protein SI self-incompatibility GM genetically modified T-DNA transferred DNA GOGAT glutamate synthase TGN trans Golgi network 111 GS glutamine synthase UTP uridine diphosphoglucose GSH glutathione UV ultraviolet GST glutathione-S-transferase VIR virulence region for infection GUS β-glucuronidase YFP yellow fluorescent protein IAA indole-3-acetic acid 11 PREFACE 11 Plant science has always been a fundamental area of biology, but the emphasis in the subject has changed radically in the last two decades with a plethora of new information, much of it deriving from techniques in molecular biology. This has deepened our understanding of plant processes and has illu- minated almost all aspects of plant biology. The ability to analyze genomes and to transfer genes has 111 opened possibilities for plant biotechnology and genetic manipulation undreamed-of in earlier decades. There have been advances in ecological knowledge that, with increased awareness of the richness of biodiversity, have shed new light on the relationships between plants, other organisms and their interdependence. Plant breeders, ecologists and many people outside plant biology have become acutely conscious of the aesthetic and economic value of the resources, so often dwindling, of the plant kingdom. In this book we have covered all these aspects of modern plant biology. We have written it keeping in mind an undergraduate faced with a range of advanced courses, needing an affordable text that gives insight into the whole range of plant science. Its scope and depth are suitable for a first and second year undergraduate student of plant biology; specialism will need an advanced text. We have also aimed it 111 at molecular biologists and biotechnologists needing an accessible route to understanding the basis of the systems on which they work. It is intended to provide the fundamental background required for true understanding. It should aid undergraduates in their learning and give insight for specialists into areas of plant science not their own. As in all Instant Notes books we have provided ‘Key Notes’ at the start of each section. These are intended solely as revision notes, e.g. before an exam, to prompt a reader’s memory after reading the section fully. We have kept technical and jargon terms to the mini- mum needed for understanding; any such term is defined at first mention. We have assumed minimal previous knowledge of biology and hope that the book will prove useful to journalists, environmen- talists and those with a genuine interest in the key issues of plant biology as they seek to be informed about the issues that they deal with. 111 The book is divided into four major sections; structure (Sections B–D); physiology and regulation (Sections E–J); ecology, genetics and wider significance (Sections K–O); and plant diversity (Sections P–R). Throughout we have included insights from modern and advanced techniques and, where pos- sible, explain how understanding is gained. Much of the recent genetic and molecular research has been done on a few key plant species, of which Arabidopsis thaliana (arabidopsis) is pre-eminent. This small short-lived weed of arable land has become the single best-known plant species and is used as a model for other plants, the ‘Drosophila’ of the plant kingdom.