MANUAL of PTERIDOLOGY Typical New Zealand Lowland Rainforest (Westland) with Cyathea Medullaris, C
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MANUAL OF PTERIDOLOGY Typical New Zealand lowland rainforest (Westland) with Cyathea medullaris, C. dealbata, Hemithelia Smithii etc., on left: Freycinethia Banksii; Podocarpus dacry dioides in background. - Photo I. L. COLLINS. MA UAL OF T IDOLOGY EDITED BY FR. VERDOORN IN COLLABORATION WITH A. H. G. ALSTON, I. ANDERSSON-KoTTO, L. R. ATKINSON, H. BURGEFF, H. G. DU BUY, C. CHRISTENSEN, W. Dopp, W. M. DOCTERS VAN LEEUWEN, H. GAMS, M. J. F. GREGOR, M. HIRMER, R. E. HOLTTUM, R. KRAUSEL, E. L. NUERN BERGK, J. C. SCHOUTE, J. WALTON, K. WETZEL, S. WILLIAMS, H. WINKLER AND W. ZIMMERMANN FOREWORD BY F. O. BOWER WITH 121 ILLUSTRATIONS • Springer-Science+Business Media, B.V 1938 ISBN 978-94-017-5743-0 ISBN 978-94-017-6111-6 (eBook) DOI 10.1007/978-94-017-6111-6 Copyright 1938 by Springer Science+Business Media Dordrecht Originally published by Martinus Nijhoff, The Hague, Netherlands in 1938. Softcover reprint ofthe hardcover 1st edition 1938 All rights reserved, including the right to translate or to reproduce this book or parts thereot in any torm CONTENTS CHAPTER I. Morphology Page § 1 HISTORICAL INTRODUCTION . 1 § 2 THE COMMON PRINCIPLES OF PTERIDOPHYTE MORPHOLOGY. 3 § 3 THE PROT HALL US . 4 § 4 THE GAMETANGIA 5 § 5 THE STEM .... 6 A. GENERAL PROPERTIES 6 B. FORM OF STEM .. 7 C. ORIGIN OF STEM. 8 D. RAMIFICATION. 9 .1. KINDS OF RAMIFICATION. 9 II. SPATIAL RELATIONS OF BRANCHES. 10 a. Spatial relations between branches of a branch system. 10 b. Spatial relations to nodes of stem . 12 c. Spatial relations to nodes and to other branches at the same time 12 d. Spatial relations to nodes and to leaves at the same time 14 e. Spatial relations to leaves. .. ....... 14 f. Spatial relations uncertain or not fixed . 17 (1 Lycopodium Selago, 2 Lepidodendron, 3 Sigillaria, 4 Com parison of spatial relations of lateral branches in Lycopodiinae) g. Taxonomic value of different modes of branching 22 E. KINDS OF STEM PLASTICS 25 F. NODES AND INTERNODES 25 G. TELOME ... 26 § 6 THE SPORANGIUM. 26 § 7 THE ROOT ....... 27 § 8 THE LEAF (STERILE LEAF AND SPOROPHYLL) 30 A. THE MICROPHYLL . 31 I. Phylogenetic origin 31, II. The sterile microphy1l31, III. The fertile microphyll31, IV. Phyllotaxis 35, V. Kinds of microphyll plastics. 42 B. THE MEGAPHYLL ...................... 43 I. Phylogenetic origin 43, II. The sterile megaphyll 44, III. The fertile megaphyll45, IV. Phyllotaxis 46, V. Kinds of megaphyll plastics 46 § 9 THE STROBILUS. 47 § JG SPECIAL PARTS OF THE EMBRYO (1 The foot, 2 The suspensor) . 48 § 11 ORGANS WHICH HAVE GIVEN RISE TO CONTROVERSIES ABOUT THEIR MOR- PHOLOGICAL NATURE. 49 (1 The Psilotum sporophyll, 2 The Selaginella rhizophore, 3 The stigmari an axis, 4 The stigmarian rootlet, 5 The Isoetes stock base, 5 The Pleuromeia stem base, 6 The Nathorstiana stem base) § 12 ApPENDAGES OF THE OUTER SURFACE (HAIRINESS) 63 CHAPTER II. Anatomy § 1 INTRODUCTION. 65 (A. Historical, B. Scope of the chapter) § 2 ANATOMY OF THE STEM . 67 A. PRIMARY ORGANIZATION 67 I. EPIDERMIS . 67 VIII CONTENTS Page II. CORTEX AND GROUND· TISSUE 68 III. STELE. 69 a. Rhyniaceae 70 b. Asteroxylon 70 c. Psilotaceae 70 d. Lycopodium 71 e. Selaginella . 72 j. Lepidophytes 73 g. Isoetes. 73 h. Equisetum. 74 i. Calamites . 74 j. Sphenophyllum 74 k. Comparison of stelar structures in microphyllous Pteridophytes 74 (1 Reduction of central part of xylem, 2 Arrangement of proto· and metaxylem, 3 Influence of leaf.traces, 4 Stem xylem and le~~.~race xylem,S Polystely) 1. Flltclnae. • . 78 (1 Reduction of central part of xylem, 2 Arrangement of proto. and metaxylem, 3 Influence of leaf· traces, 4 Stem xylem and leaf. trace xylem) IV. ORIGIN OF PRIMARY ORGANIZATION. • . 85 B. SECONDARY TISSUE FORMATION . 86 (I. Secondary cork tissue formation, II. Secondary vascular tissue) § 3 ANATOMY OF THE ROOT . • . 93 A. PRIMARY ORGANIZATION .. 93 (I. Epidermis, II. Cortex, III. Stele, IV, Origin of primary organization) B. SECONDARY TISSUE FORMATION . 96 (I. Secondary cork tissue formation, II. Secondary vascular tissue) § 4 ANATOMY OF THE LEAF . ... 98 A. THE MICROPHYLL . 98 (1. Epidermis, II. Mesophyll, III. Stele, IV. Secondary tissue formation) B. THE MEGAPHYLL ....................... 100 (I. Epidermis, II. Mesophyll, III. Stele, IV. Secondary tissue formation) CHAPTER III. Experimental Morphology § INTRODUCTION AND HISTORICAL ACCOUNT . • . 105 THE GAMETOPHYTE § 2 FACTORS INFLUENCING GERMINATION. •. 106 § 3 FACTORS INFLUENCING THE DEVELOPMENT AND FORM OF THE PROT HALL US. 107 § 4 REGENERATION OF THE PROTHALLUS 114 § 5 FERTILISATION . • . • . 115 THE SPOROPHYTE § 6 EMBRYOLOGY. 115 § 7 FACTORS UNDERLYING VARIATIONS IN EXTERNAL FORM 116 § 8 CORRELATION OF PARTS . 119 § 9 FACTORS INFLUENCING THE VASCULAR SYSTEM. 119 § 10 ORGANS OF INDETERMINATE NATURE. 123 § 11 REGENERATION. 126 § 12 FACTORS UNDERLYING THE PRODUCTION OF SPORANGIA 130 § 13 EXPERIMENTS BEARING ON THE MORPHOLOGY OF SPOROPHYLLS AND spo· RANGIOPHORES . 131 § 14 DEVIATIONS FROM THE NORMAL LIFE·CYCLE . 132 CONTENTS IX EVOLUTIONARY PROBLEMS Page § 15 THE MECHANISM OF EVOLUTION 136 § 16 THE COURSE OF EVOLUTION 137 § 17 CONCLUSION • • • • . • • • 139 CHAPTER IV. Associations with Fungi and other lower plants § 1 INTRODUCTION • • • • • • • • 141 § 2 THE FOSSIL RECORD • • • • •• 142 § 3 DISEASES OF THE GAMETOPHYTE . 142 § 4 DISEASES OF THE SPOROPHYTE 143 Filicinae ... 143 Equisetinae . 154 Lycopodiinae. 155 § 5 SYMBIOSIS AND EPIPHYTISM . 156 CHAPTER V. Mycorhiza § 1 EINLEITUNG • . • . 159 § 2 THAMNISCOPHAGE MYCORHIZA . 161 a. Primitive Formen 163 b. Marattiaceenmycorhiza . 164 c. Die hochentwickelte Form der Ophioglossaceen 167 § 3 DIE TOLYPOTHAMNISCOPHAGE MYCORHIZA DER PSILOTALES • 174 § 4 DIE PTYOPHAGE MYCORHIZA DER L YCOPODIEN DES PHLEGMARIA-TYPUS 178 § 5 DIE CHYLOPHAGE MYCORHIZA DER LYCOPODIEN. 181 a. Typus des Lycopodium complanatum . 181 b. Typus des L. clavatum und L. annotinum 184 c. Typus des L. inundatum und L. cernuum 188 § 6 RikKBLICK. • • . • • • • • • • 190 CHAPTER VI. Zoocecidia ZOOCECIDIA. 192 LITERATURE 195 CHAPTER VII. Cytology HISTORICAL SKETCH 196 Bibliography. 204 2 THE FILICINAE • • 205 The Gametophyte . 205 Sperma togenesis 206 Oogenesis and Fertilization 212 The Sporophyte. 215 § 3 THE LYCOPODIINAE 220 The Gametophyte . 220 Spermatogenesis 220 The Sporophyte. 222 § 4 EgUISETINAE. • • 226 The Gametophyte . 226 Spermatogenesis 227 The Sporophyte. 230 § 5 CONCLUSIONS. • • 232 CHAPTER VIII. Karyologie § 1 GESCHICHTLiCHER UBERBLiCK. LiTERATUR. METHODIK 233 Literatur. 235 x CONTENTS Page § 2 DER RUHEKERN, TERMINOLOGIE, MORPHOLOGIE, PHYSIOLOGIE 238 § 3 DIE TYPISCHE KERNTEIL UNG • . • • . • • • • • • . 245 § 4 DIE REIFETEILUNGEN • • . • . • • . • . • . 252 § 5 DIE NUKLEOLEN IN BEZIEHUNG ZU KERNTEILUNG UND CHROMOSOMEN 256 § 6 DIE CHROMOSOMEN. 258 Individualitat 258 Zahl .. 258 Grosse. 260 Form. 261 Bau .. 263 § 7 KERNVERSCHMELZUNG 264 Geschlechtszellen . 264 Befruchtungsvorgang 265 § 8 ABWEICHUNGEN YOM NORMALEN KERNTEILUNGSMODUS 268 § 9 POL YPLOIDIE UND ANEUPLOIDIE . • . • • • • • . 275 § 10 ABWEICHUNGEN YOM NORMALEN KERNPHASENWECHSEL . 281 § 11 SCHLUSS. • • • • . • • . • . • 283 CHAPTER IX. Genetics § 1 HISTORICAL NOTES 284 § 2 GENETICAL EXPERIMENTS WITH STRUCTURAL CHARACTERS 286 § 3 OBLIGATORY APOSPORY. 292 § 4 POLYPLOID SEGREGATION. 293 § 5 MUTATION ••..••. 293 § 6 CHLOROPLAST CHARACTERS 295 § 7 EFFECTS OF ALTERATION OF GENOME QUANTITY. 296 § 8 THE EFFECTS OF GENETICAL FACTORS IN SPOROPHYTE AND GAMETOPHYTE 299 § 9 GENERAL CONSIDERATIONS • • . • . • • . • . • 301 CHAPTER X. Growth, Tropisms and other Movements INTRODUCTION AND REVIEW OF THE MAIN LITERATURE ...... 303 I. GROWTH OF THE GAMETOPHYTE GROWTH OF PROTHALLIA • • . • . • • • . • • . • 304 Influence of hormones on the development of prothallia 305 Regeneration in prothallia . 306 § 2 DEVELOPMENT OF DORSIVENTRALITY . 306 Tropisms of the Gametophyte. 306 Influence of environmental conditions, especially of light 307 Influence of light. ............ 307 1 The dependence of developmental growth on the wavelength of light . .'. .. ....... 308 Germination of spores. .. ....... 308 2 The dependence of transverse phototropism on the wavelength of light . 309 3 Reversal of dorsiventrality by light . 310 Influence of gravity and other tropisms . 310 § 3 GROWTH OF THE RHIZOIDS OF THE FULL GROWN GAMETOPHYTE 311 Movements of the rhizoids of the full grown Gametophyte. 311 Phototropism of prothallium rhizoids of Equisetum. 311 Phototropism and Geotropism of prothallium rhizoids of Filicinae 311 II. GROWTH OF THE SPOROPHYTE § 4 GROWTH TYPES IN FILICINAE . • . • • • • • 312 § 5 INFLUENCE OF DARKNESS ON GROWTH (ETIOLATION) 316 § 6 PHOTOTROPISM OF LEAVES. • • • • • • • • • • • 316 CO~TENTS XI Page § 7 GEOTROPISM OF FERN LEAVES 319 § 8 HYPONASTY OF FERN LEAVES 321 § 9 DORSIVENTRALITY. • . 322 § 10 HETEROPHYLLY. • . • • 322 Heterophylly and light. 323 11 SHOOT FORMATION BY LEAVES. 324 12 GROWTH OF AEROPHORES, LEAF HAIRS OF SALVINIA 324 III. REGENERATION IN FERN LEAVES § 13 REGENERATION • . • • • . • . • . • • 325 Woundhormones ............. 330 Table of regenerations, arranged in order of place of development 330 1 Regeneration on leaves. 330 2 Regenerations on roots and growing points . 334 3 Regeneration on prothallia. 335 IV. SECONDARY THICKENING § 14 SECONDARY THICKENING. • . • .. 336 V. MOVEMENTS IN FERNS WHICH ARE NOT CAUSED BY GROWTH § 15 DIURNAL MOVEMENTS OF THE LEAVES OF MARSILIA. • . .. 337 § 16 MECHANISM OF THE ANNULUS OF THE POLYPODIACEAE AND RELATED PHE- NOMENA • • • • . • • . • . • . • • . • 337 § 17 HYGROSCOPIC MOVEMENTS OF SPORES OF EQUISETUM 339 § 18 MOVEMENTS