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Plant Anatomy . ~ PLANT ANATOMY A. FAHN Professor of Botany The Hebrew University, Jerusalem, Israel Translated from the Hebrew by SYBIL BROmO·ALTMAN PERGAMON PRESS .~. ~~~OOK~. OXFORD' NEW YORK £6'\ f'>,-- ~--!.?Y (.;)-.~ ,/ t- ....- """" v~ ~ - ( '\ TORONTO· SYDNEY· BRAUNSCH 00 A r- {) ".1 . ~. CONTENTS PREFACE vii GENERAL STRUCTURE OF HIGHER PLANTS ....••. 2 THE CELL ••......•..•••...•.•••.....•.....•. 7 3 MERISTEMS . , . 44 Mature Tissues 4 PARENCHYMA ............................. 73 5 COLLENCHYMA ....... ..................... 80 6 SCLERENCHYMA ..........•....•.............. 85 7 XYLEM ••.•.••.• ,.,.. • • . • . •. • . •. •. .• 102 I I . 8 PHLOEM •.•••••••....••••••.••.•.•.•••.•...•.••. 118 ! 9 LATICIFERS .•...•..•.....•.•.....••.•.••..••••. 130 10 EPIDERMIS...................................... 137 I • Primary Vegetative Body Of the Plant I I THE STEM ................................... 162 12 THE LEAF ................................... 193 13 THE ROOT .......................... 240 Secondary Body of the Plant 14 VASCULAR CAMBIUM •.••..•.••.•...••••.....•• 273 15 SECONDARY XYLEM .•••.....•.••...•.•........ 286 16 SECONDARY PHLOEM •.••..•••••.•.•..••..••... 325 17 PERIDERM ••..•.•••..•.•....••..•..•••.- •••..• 331 18 ANOMALOUS SECONDARY GROWTH .............. 349 Pergamon Press Ltd., Headington Hill Hall, Oxford 011 Press Inc., Maxwell House, Fairview Park, Elmsford, New York 10523 n1 pe(ga Pergamon of Canada Ltd., 207 Queen's Quay West, Toronto 1 55 (Amt.) Pty. Ltd. 19a Bounda:y Street, RushcutterslBay, N.S w. 2011 [flo" pre Australia pen:t.!l Vieweg & Sohn GmbH, Burgplatz I, Braunschweig Copyright © J 967 A. FAHN First English Edition 1967 Reprinted with corrections 1969 r-- Reprinted 1972 -___--, 1 I r~!~iral Ph,l! lti~., Cri)"H " CAL I CUl-67JOll ". / N D. .s;- r-:t I D~te Original Hebrew book _published by Hakkibutz Hameuhad Publishing House Ltd. Library of Congress Catalog Card No. 66--24822 Printed in Great Britain by A. Wheaton & Co., Exeter 08 011943 3 PREFACE PLANT anatomy is a basic science and as such is of great importance to stu­ dents of all the plant sciences. Without a thorough knowledge of this field the physiological processes carried out within the plant and the phylogene­ tic relationships between the various plant groups cannot be fully under­ stood. The detailed study of the elements and tissues of which the plant is constructed enables a better understanding of adaptation to special functions as well as of the adaptation of entire plants to different environ­ mental conditions. Without a thorough knowledge of the anatomical and histological structure of plants the results of physiological and ecological oxperiments, for instance, may be incorrectly interpreted. Also, to-day no ~onclusive opinions on evolutionary trends or taxonomic relationships can be suggested on the traditional basis of the study of external morphological characteristics alone; it is now necessary to support such work by the use of the many and varied anatomical and histological characters, which can be observed only from microscopic, and even submicf9scopic, investigation. Anatomy, which draws the attention of the student to the form, varia­ bility and structure of the Itissues comprising the plant body, can be said to develop an aesthetic sense. In addition to this, the awareness of the regularity, and repetition, at different levels, of the structural patterns, as well as of the amazing correlation of structure and function, serves to make anatomy a rewarding field of research. A large section of this book deals with the vegetative plant body. The first introductory chapter briefly presents the general structure of the higher plant. This is followed by tbe descriptions of the different types of cells and tissues that are present in the Tracheophyta. Later chapters de­ scribe how the vegetative plant body, both primary and secondary, is con­ structed of these various tissues. The last section deals with the structure of the flower, fruit and seed. In the chapter on the flower I have covered pollination, fertilization and embryo development. In my opinion this is necessary in order that a full and balanced picture of the development and structure of plants and their tissues can be obtained. An effort has been made, when dealing with the structure of the ele­ ments, tissues and organs of the plant body, to employ the following ap­ proaches-ontogenetic, phylogenetic, physiological and ecological. Atten­ tion has also been paid, wherever possible, to such characteristics that are of importance to agriculture and industry. In view of modern research in plant anatomy and biology as a whole, which ha~ hTf"Illnl"d-""" 1: ....1. .. ~- -- vi Contents Reproductive Organs I 19 THE FLOWER .........•....................•. 360 20 THE FRUIT .................................. 439 21 THE SEED ..............•.................... 466 GLOSSARY OF TERMS ......................•...... 't~0 AUTHOR INDEX 495 SUBJECT INDEX viii Preface bility in the definitions of the various elements and tissues is stressed throughout the book. i In many cases I have endeavoured to point out problems which as yet constitute serious gaps in our knowledge, and which await further research. The inclusion of a large number of illustrations has enabled the text to be written in a concise form. The great majority of the micrographs are original and have been made from slides in the collection of.the laboratory of Plant Anatomy at the Hebrew University of Jerusalem. The drawings are in p",rt original or have been taken from previous publications of the author, and the rest have been redrawn and adapted from various books and articles. In the case of the latter the original author is cited in the le-' gend and the reference is given at the end of the relevant chapter or Chap­ ter I. In a book of this size and scope it is impossible to deal with all the relevant facts in detail and therefore many references are given in the text. It is hoped that readers will refer to these and other articles, books, etc., in order to broaden their knowledge. For convenience, details of these references are given at the end of each chapter. \ This book was originally written in Hebrew for the use of students study­ ing in Israel. Thus many of the examples cited )Ire of plants growing in this and neighbouring regions. / I express my thanks to all those who helped me in the preparation of tlie original'Hebrew book. I am indebted to·my friend S. Stoler for his critical reading of the manuscript and for his valuable suggestions; to Mrs. Ella Werker for her great help in the preparation of the manuscript and for seeing the book through press; to Mrs. Batya Amir for the careful and accurate execution of most of the drawings; to Y. Shehori for his aid in the· preparation of mast of the photographs appearing in tire book; and to Mrs. Irena Fertig for her assistance in reading the pr09fs. My thanks are also extended to all those who have put at my disposal photographs and.draw­ ings, as well as to my colleagues at the Hebrew University who gave me valuable advice at all times. I especially thank my students, throughout the years, who have encouraged me to write this book. In connection with this revised English edition I am indebted to Dr. C. \ R. Metcalfe and Sir George Taylor who suggested and "encouraged me to have my book translated. I greatly appreciate the criticism and advice that Dr. Metcalfe extended after having read the English manuscript. I thank Dr. F. A. L. Clowes who undertook to edit the English. I gratefully acknow­ ledge the permission so generously granted to me by the Hakkibutz Hame­ uhad Publishing House Limited to translate the original text. I also thank Mrs. Sybil Broido-Altman for undertaking the translation. Once again I thank Mrs. Ella Werker, who assisted in the collation. A.FAHN Jerusalem, April, 1965 CHAPTER I GENERAL STRUCTURE OF HIGHER PLANTS PLANTS that bear seeds are termed spermatophytes. These plants pro­ duce spores (newly formed embryo sacs and pollen grains) and therefore they are sporophytes. These plants develop from a zygote which results 'v CotyJeEion -Hypocotyl -Root 2 3 FIG. 1. Schematic drawings of longitudinal sections ofa dicotyledonous plant at various ages. 1. Embryo. 2, Seedling. 3, Mature plant. (Adapted from Troll, 1948.) from the fertilization of an egg cell by a male gamete. At the start the zygote divides into two cells which themselves undergo further divisions to form the embryo. The embryo usually consists of radicle, hypocotyl, cotyledons and olumule (Pi!!. L no n TJ,p. p".,h .." ......0 ............ " ~~ ____ 1_..J 2 Plant Anatomy and dormant within the seed which develops from the entire ovule. IUs not always clear whether the embryo possesses a radicle ~roper or whether it merely has a root apical meristem. It is difficult to make a definite dis­ tinction between the radicle and hypocotyl, and therefore the axis of the embryo is called the hypocotyl-root axis. With the germination of the seed the embryo renews its growth. The radicle grows and penetrates deeper into the soil. In some species the hypocotyl elongates and so raises the cotyledons above soil level where they become green (epigeal germination). In other species the hypocotyl does not elongate, .or it elongates only very slightly, and the cotyledons remain below soil level where they eventually rot (hypogeal germination). The plumule, which is situated above the junction of the cotyledons to the hypocotyl, elongates and gives rise to the stem and leaves (Fig. 1, nos. 2, 3) .. That part of the
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