Weed Anatomy PDF
Preview Weed Anatomy
Weed Anatomy Weed Anatomy Hansjoerg Kraehmer Bayer CropScience AG, Industriepark Höchst, H872 65926 Frankfurt am Main Germany Peter Baur Clariant Industrial & Consumer Specialties Industriepark Höchst 65926 Frankfurt am Main Germany A John Wiley & Sons, Ltd., Publication This edition first published 2013 © 2013 by John Wiley & Sons, Ltd. Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific, Technical and Medical business with Blackwell Publishing. Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 2121 State Avenue, Ames, Iowa 50014-8300, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the authors to be identified as the authors of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Kraehmer, Hansjoerg, author. Weed anatomy / Hansjoerg Kraehmer, Bayer Crop Science, Aktiengesellschaft, Frankfurt am Main, Germany, Peter Baur, Head of Competence Center Crop Protection, Global Application Development, Clariant Industrial & Consumer Specialities, Industriepark Höchst, Frankfurt am Main, Germany. Includes bibliographical references and index. ISBN 978-0-470-65986-1 (hardback) – ISBN 978-1-118-50334-8 (epub) – ISBN 978-1-118-50343-0 (epdf) 1. Weeds–Anatomy. I. Baur, Peter, 1965– author. II. Title. SB611.K73 2013 632’.5–dc23 2012040912 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover image: Capitate glandular trichomes of Solanum nigrum L. (blue); SEM, computer-enhanced colours (see figure 7.1 for more details). Cover design by Meaden Creative Set in 10.5/13 pt Classical Garamond BT by Toppan Best-set Premedia Limited 1 2013 Contents CONTENTS Section I Chapter 25 Root Histology 97 Cells and Tissues Chapter 26 Root Tip 99 1 Chapter 27 Xylem Patterns 102 Introduction 3 Chapter 28 Endodermis and Pericycle 109 Chapter 1 Tissues 5 Chapter 29 Rhizodermis, Exodermis and Cortex 114 Chapter 2 Parenchyma 8 Chapter 30 Root Genetics 116 Chapter 3 Collenchyma 10 Chapter 31 Primary and Secondary Growth 117 Chapter 4 Sclerenchyma, a Typical Contributor to Chapter 32 Anomalous Secondary Growth 126 Weediness 12 Section V Section II Complex Tissues and Organs 129 Meristematic, Secretory, Storage and Chapter 33 Leaf 131 Boundary Structures 17 Chapter 34 Flower 142 Chapter 5 Meristems 19 Chapter 35 Androecium 154 Chapter 6 Secretory Structures 24 Chapter 36 Gynoecium 159 Chapter 7 External Secretory Structures 26 Chapter 37 Genetics of Flower Formation 169 Chapter 8 Internal Secretory Structures 30 Chapter 38 Fruit 170 Chapter 9 Stored Compounds 35 Chapter 39 Carpels, Pericarp and Various Fruit Chapter 10 Epidermis 37 Forms 173 Chapter 11 Stomata 40 Chapter 40 Genetics of Fruit Development 184 Chapter 12 Non-glandular Trichomes and Papillae 42 Chapter 41 Seed 185 Chapter 42 Genetics of Seed Development 190 Chapter 43 Secondary Reproduction Section III Characteristics 191 Vascular Elements and Pith 47 Chapter 44 Flower Modifications in Weeds 192 Chapter 45 Seedling and Embryo 196 Chapter 13 Vascular Bundles 49 Chapter 14 Xylem 52 Section VI Chapter 15 Pits 58 Chapter 16 Phloem 60 Vegetative Propagation 203 Chapter 17 Pith 64 Chapter 46 Vegetative Weed Reproduction 205 Section IV Chapter 47 Rhizomes 206 Chapter 48 Tubers and Corms 211 Stem, Root and Growth 67 Chapter 49 Stolons and Runners 217 Chapter 50 Roots with Adventitious Buds 219 Chapter 18 Stem 69 Chapter 51 Bulbs 222 Chapter 19 Dicot Stem – Cortex 71 Chapter 20 Dicot Stem – Patterns of Vascular Tissues 74 Chapter 21 Vascular Bundles and Leaf Traces in Dicots 78 Section VII Chapter 22 Monocot Stem 82 Weediness 225 Chapter 23 Horsetail and Bracken Stem 88 Chapter 24 Root Morphology 92 Chapter 52 Indicators of Weediness 227 v vi Contents Section VIII Monograph 20: Galium aparine L. 370 Monograph 21: Impatiens glandulifera Royle 375 Short Monographs 233 Monograph 22: Lolium multiflorum Lam., Lolium Chapter 53 Introduction to Monographs 235 perenne L. and Lolium rigidum Chapter 54 Weed Anatomy Monographs 237 Gaudin 386 Monograph 1: Abutilon theophrasti Medicus 237 Monograph 23: Myosotis arvensis (L.) Hill and Monograph 2: Alopecurus myosuroides Huds. and Myosotis palustris (L.) Nath. 394 Alopecurus japonicus Steudel 241 Monograph 24: Paspalum dilatatum Poir. 400 Monograph 3: Amaranthus retroflexus L. and Monograph 25: Phalaris minor Retz. and Phalaris Amarantus palmeri S. Wats. 248 paradoxa L. 410 Monograph 4: Ambrosia artemisiifolia L. 259 Monograph 26: Poa annua L. 416 Monograph 5: Apera spica-venti (L.) P. Beauv. 266 Monograph 27: Polygonum amphibium L., Polygonum Monograph 6: Avena fatua L. and Avena sterilis L. 268 aviculare L. and Polygonum Monograph 7: Bidens pilosa L. and lapathifolium L. 422 Bidens tripartita L. 272 Monograph 28: Rottboellia cochinchinensis (Lour.) W. Monograph 8: Bromus secalinus L., Bromus sterilis L. Clayton 429 and Bromus tectorum L. 276 Monograph 29: Setaria faberi Herrm., Setaria glauca Monograph 9: Calystegia sepium (L.) R. Br. and (L.) Beauv. and Setaria viridis (L.) Convolvulus arvensis L. 279 Beauv. 434 Monograph 10: Chenopodium album L. 282 Monograph 30: Sorghum halepense (L.) Pers. 441 Monograph 11: Cirsium arvense (L.) Scop. 293 Monograph 31: Urochloa platyphylla (Munroe ex C. Monograph 12: Conyza canadensis (L.) Cronq. and Wright) R. D. Webster 450 Conyza bonariensis (L.) Cronq. 298 Monograph 32: Xanthium strumarium L. and Xanthium Monograph 13: Cynodon dactylon (L.) Pers. 304 spinosum L. 454 Monograph 14: Cyperus esculentus L., Cyperus iria L., Cyperus rotundus L. and Cyperus Section IX serotinus Rottb. 311 Monograph 15: Digitaria sanguinalis L. 325 Methods for the Preparation Monograph 16: Echinochloa colonum (L.) Link and of Sections 463 Echinochloa crus-galli (L.) Beauv. 333 Monograph 17: Eleusine indica (L.) Link 342 Chapter 55 Tissue Preparation and Staining Monograph 18: Elytrigia repens (L.) Nevski 348 Procedures 465 Monograph 19: Fallopia convolvulus (L.) Löve, Fallopia japonica (Houtt.) Ronse Decraene, References 472 Fallopia sachalinensis (F. Schmidt) Ronse Decraene and Fallopia Index 482 baldschuanica (Regel) Holub 357 SECTION I Cells and Tissues No function without structure | Introduction, 3 1 | Tissues, 5 2 | Parenchyma, 8 3 | Collenchyma, 10 4 | Sclerenchyma, a Typical Contributor to Weediness, 12 Introduction Weeds are plants interfering with the interests by plant population scientists (e.g. Braun-Blanquet of people. They are regarded as problems espe- 1964; Dierschke 1994). Anatomy and morphology cially by farmers, by industrial vegetation managers, are signs and the result of adaptation. In consequence, in ornamental production, in forestry or on sites we try to link ecological characteristics with the form where vegetation can cause safety risks, for example of adaptation. Weediness is the ability of unwanted on railway tracks or along motorways. Some weeds plants to survive in an environment managed by impose a health risk for people by causing allergic humans. Modern weed control tools have reached reactions, such as Ambrosia artemisiifolia L. Others a high degree of efficiency. Evolutionary principles, are toxic to animals, such as Senecio jacobaea L. however, allow weeds to escape our efforts to control which kills horses from time to time when eaten. As them. Resistance, physiological and morphological part of our ability to approach problems systemati- adaptation seem to happen much faster than we had cally, humans try to classify and to prioritise. A famous previously realised. Within 6000 years of agriculture, book cited by weed scientists is The World’s Worst a restricted number of weed species have been selected Weeds (Holm et al. 1977). The title categorises weeds by humans. The flora has changed to some extent in as bad. Weeds are, however, parts of ecosystems and history due to climate changes and to agricultural are regarded as elements of biodiversity by many sci- practice. A few species, however, have managed to entists today (Radosevich et al. 2007). For these survive over centuries. We will concentrate on these reasons, we will not use value-driven categories in this apparently best-adapted weeds and we will highlight book. We will also include rare weeds, that is weeds their characteristics. The basis of our categories was that are endangered and protected in some countries defined by working groups of the European Weed (e.g. according to the UK Biodiversity Action Plan). Research Society. One might even ask if these plants can be called weeds Any plant can be a weed by definition (i.e. there as they don’t compete with crops and no longer nega- are no typical structures or features within a plant that tively interfere with the interests of people. Proceed- make it a weed). There are, however, characteristic ings of the Weed Science Society of America (WSSA) features that allow some weed species to survive and formerly classify weeds as either frequent or trouble- proliferate in arable field situations: stolons, rhizomes, some. Frequency can be quantified and measured; this bulbs, tubers, buds on roots, seed morphology and is not so easy with troublesome. We will therefore try reproductive organs. The uptake of herbicides and their to concentrate on frequency and use tools described distribution within a plant depend on leaf structures Weed Anatomy, First Edition. Hansjoerg Kraehmer and Peter Baur. © 2013 John Wiley & Sons, Ltd. Published 2013 by John Wiley & Sons, Ltd. 3 4 Section 1 Cells and Tissues and barriers between parenchyma and vascular our book. They tolerated the collection of specimens bundles. The morphological terms used in this book during private excursions and, from time to time, were derived from classical textbooks written, for quite a mess of plant material at home. example, by K. Esau (1969, 1977), A. Fahn (1990) We are grateful to our employer, Bayer Crop- or Bowes and Mauseth (2008), C.B. Beck (2010), Science, for permission to use screening plant mate- E. Strasburger (1891). Thoughts about specific weed rial and to publish SEM pictures produced for various structures were inspired by W.C. Dickison’s book purposes. Integrative Plant Anatomy. Rich sources of details are Nigel Balmforth and Carys Williams from Wiley- A.R. Metcalfe’s series of Plant Anatomy (Metcalfe Blackwell accompanied us with help and assistance 1960, 1971; Metcalfe and Chalk 1965) and the Ency- early on. The positive statements of a number of peers clopedia of Plant Anatomy. all over the world led our publisher to the final deci- In Section 9, below, you will find literature on sion in favour of this worthwhile enterprise. staining techniques and microscopy. Classical books on plant anatomy start with cells and tissues. They continue with characteristic plant Morphology, Anatomy, tissues, for example parenchyma, collenchyma, scler- enchyma, epidermis, meristems, phloem, xylem and Physiology – Explanation secretory structures. Organs and plant parts are of Terms described thereafter: stem, root, leaves and reproduc- tive organs. We will try to follow this structure with examples from weeds in the first part of this book. The term morphology is derived from the Greek For an introduction into cytology and the character- words ‘morphe’, meaning form or shape, and ‘logos’, istics of protoplasts, we recommend to read the which may be translated here as science. In essence, appropriate chapters in the latest issue of Esau’s Plant plant morphology is the science of form, shape, struc- Anatomy (Evert 2006). Many textbooks have devoted ture and outer appearance of plants. The definition separate chapters to primary and secondary growth. of each of these descriptive terms is, however, not The most frequent weed species, however, are either easy. Philosophical discussions about the form of annuals, biennials or non-woody perennials. For this plants concentrate today, more or less, on three dif- reason, we will not refer to noxious trees nor bushes. ferent aspects: on the characterisation of organ types, Only higher plants are considered, that is algae or on how plant parts were modified during evolution, mosses are excluded. The second part of our book what role they play and on the relative importance of consists of short monographs of weeds with particular these parts when describing the development of a reference to anatomy. plant (e.g. Classen-Bockhoff 2001). Anatomy is also We thank all our colleagues who have assisted us derived from a Greek word, from anatomos, which with the preparation of this book and have helped means dissected. It deals with the ‘inner’ appearance us in improving the layout. M. Hess, C. Rosinger, of plants or plant parts. Anatomy describes tissues and C. Ueno, M. Hills, J. Koehler and L. Lorentz dis- organs as they can be seen when a plant is cut into cussed our approach with us regularly. B. Rueffer, M. pieces with a razor blade, a microtome knife or when Linder, P. Remmert and S. Engels assisted us with a plant is analysed by means of electromagnetic waves. plant material. S. Teitscheid and Stephanie Giessler Physiology comes from ‘physis’, nature, and explains produced our illustrative SEM pictures and Philipp the function and development of plants or plant parts. Baur contributed by processing SEM picture files. H. You will find all these aspects in our book where Reitzammer checked our manuscript from time to appropriate. The major emphasis, however, is placed time. Our families encouraged us to go ahead with on the anatomical characteristics of weeds. Chapter 1 Tissues No structure without substructure The reproductive development of higher plants starts differentiates into a seedling and characteristic tissues with a fertilised egg cell. This cell and its descendants or functional plant parts (Figure 1.1), which later on divide and form cell clusters. The growing embryo result in an organism consisting of stem, leaf and root. Calyptra (root cap) Radicle (A) (B) Figure 1.1 Germinating seed of Abutilon theophrasti Medik. Weed Anatomy, First Edition. Hansjoerg Kraehmer and Peter Baur. © 2013 John Wiley & Sons, Ltd. Published 2013 by John Wiley & Sons, Ltd. 5 6 Section 1 Cells and Tissues In between, a number of distinct tissues develop ● Vascular tissues called phloem and xylem: they which can be classified according to their appearance transport water and assimilates from one plant and function. The following terms will be used in the part into another. next chapters. ● Meristem: these consist of embryonic cells which divide, form new tissues and are already apparent ● Epidermis with interfacial cuticle: the plants’ in seedlings, as shown in Figure 1.2. outer protective layer of cells; it allows the exchange of water, some ions, CO and O with ● Secretory tissues: many plants excrete all kinds of 2 2 the environment; many agrochemicals can enter secondary products, for example oils, resins, gums, the plant via this layer. mucilages and others; these products are stored, transported or excreted by secretory tissues. ● Parenchyma: ground tissue of cells with thin walls; this kind of tissue often fills spaces Explanations of our figures often describe how between other tissues or between organs. tissues were dissected. Figure 1.3 depicts three major ● Collenchyma and sclerenchyma: tissues that ways of generating sections as you may find them in stabilise the form of stem, leaf and root. most of our examples. Cotyledons Apical meristem Figure 1.2 Embryo of Galium aparine L., longitudinal section.
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