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Histological Investigations of the Secondary Phloem of Gymnosperms

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Histological Investigations of the Secondary Phloem of Gymnosperms 582.42/.47:58l.824.2 634.0.174.2:634.0.168 MEDEDELINGEN LANDBOUWHOGESCHOOL WAGENINGEN . NEDERLAND . 67-7 (1967) HISTOLOGICAL INVESTIGATIONS OF THE SECONDARY PHLOEM OF GYMNOSPERMS R. W. DEN OUTER Department of General Botany, Agricultural University, Wageningen, The Netherlands (Received27-11-1967 ) H. VEENMAN &ZONE N N.V. - WAGENINGEN - 1967 Mededelingen Landbouwhogeschool Wageningen 67-7(1967 ) (Communications Agricultural University) is also published as a thesis CONTENTS 1. INTRODUCTION 1 2. MATERIAL AND METHODS 3 2.1. Slides 3 2.2. Drawings 5 2.3. Terms used 5 3. THE DIFFERENT CELL TYPES IN THE SECONDARY PHLOEM 9 3.1. Thecambiu m cells 9 3.1.1. Fusiform cambium initials 9 3.1.2. Ray initials 9 3.2. Sievecell s 10 3.3. Parenchymatouselement s 12 3.3.1. Phloem parenchyma 12 3.3.2. Ray parenchyma cells 13 3.4. Sclereidsan d stonecell s 14 3.5. Albuminouscell s 15 3.6. Phloem fibres 16 3.7. Periderm 17 4. DISTRIBUTION AND ARRANGEMENT OF CELL TYPES IN THE AXIAL SYSTEM OF THE SECONDARY PHLOEM 18 4.1. Sievecells 18 4.1.1. Sievecell sarrange d in broad bands 18 4.1.2. Sievecell sarrange d into tangential bands 19 4.1.3. Sievecell sarrange d into tangential layerso f onecel l wide 19 4.2. The parenchyma cells 19 4.2.1. Scattered amongsiev ecell s 19 4.2.2. In tangential bands 19 4.2.3. In layerso fon ecel lwid e 20 4.3. The albuminouscell s 20 4.4. The phloem fibres 21 4.4.1. Scattered among parenchyma cells 21 4.4.2. Arranged intangentia l layers, 1 cellwid e 21 4.4.3. Arranged intangentia l bands,2- 3cell swid e 21 4.5. Growth rings 21 5. THE PHLOEM RAYS AND THE WOOD RAYS 23 5.1. The phloem rays 23 5.1.1. Structure 23 5.1.2. Development 23 5.1.3. Phylogenetic stages 24 5.2. The wood rays 26 5.2.1. Structure 26 5.2.2. Development 26 6. THE TYPES OF THE SECONDARY PHLOEM, BASED ON ARRANGEMENTS OF CELL TYPES 28 6.1. Secondary phloem consisting of broad bands of sieve cells (Pseudotsuga taxifoliatype ) 28 6.1.1. Pit-contact between sievecell san dphloe m raysvi a thealbuminou s cells ... 28 6.1.1.1. PseudotsugataxifoIiaBMTT 28 6.1.1.2. Pinussylvestris L 31 6.1.1.3. Larix decidua MILL 32 6.1.1.4. Abiesconcolor HOOPES 33 6.1.2. Pit-contact between sieve cells and all the elements of the phloem ray(Tsuga canadensissubtype ) 34 6.1.2.1. Tsugacanadensis CARR 34 6.1.2.2. CedruslibaniA.RlCH 34 6.1.2.3. Piceaspec 35 6.2. Secondary phloem consisting of bands of sieve cells and bands of paren­ chymacell s(Ginkgo biloba type ) 36 6.2.1. Ginkgo bilobaL 36 6.2.2. Podocarpusneri/oliusT>.T>ON 37 6.2.3. Taxusbaccata L 38 6.3. Secondary phloem consisting of a regular sequence of alternating cell types (Chamaecyparispisifera type ) 39 6.3.1. Chamaecyparispisifera ENDL. var.p/umos a OTTO 40 6.3.2. Thujaplicata LAMB 41 6.3.3. ThujopsisdolabrataSiEü.slZvcc 42 6.3.4. Taxodiumdistichum A . RICH 42 6.3.5. Cryptomeriajaponica D . DON 43 7. THE PHYLOGENETIC STAGES OF THE BAST TYPES 45 7.1. Comparison of thestructur e of thedifferen t secondary phloem types .... 45 7.2. Characteristics of the different phylogeneticstage s 46 7.3. Thedifferen t phylogeneticstage s 46 8. DISCUSSION 52 8.1. Axial system 52 8.2. Albuminous cellsan d companion cells ' 53 8.3. Sieveelement s 56 8.3.1. Cytologicspecializatio n 56 8.3.2. Morphologicspecializatio n 57 9. SUMMARY 58 ACKNOWLEDGEMENTS 61 SAMENVATTING 62 REFERENCES 65 PLATES 72 1. INTRODUCTION To gain an insight into the structure of the secondary phloem of Gymno- sperms, primarily the different cell types with regard to their appearance, differentiation and degree of development, have to be described. Special atten­ tion has been paid to the presence or the absence of pits, i.e. the location of connections between adjacent cells; in particular amount, kind and location of these pits have been examined. Through these pits an easy movement ofassim ­ ilatesfro m onecel lt oanother , ispossible . The term 'pit' is used in a broad sense,tha t ist o indicate a cavity or a thin place in the cell wall. In primary walls such pits will also be called 'primary pit-fields'. More specifically, the cavities in parenchyma-cell wallswil lb ecalle d pits, those in sieve-cell walls, between adjacent sieve cells, will be called sieve areas. Consequently the cell types are dependent upon pit-contact between these cells for food-conducting functions. The different arrangements of these cell types are investigated, also the directions in which this movement of assimilates maytak eplace . Many statements havebee n madeabou t food-conduction pathways.Th ewa y in which the main transport of food is directly related to the arrangement and the structure of the cell types and the importance of special combinations of different kinds of cell types, have been pointed out. Thus, apart from the comparative anatomy, some physiological aspects have also been alluded to. All these statements about food-conducting pathways require evidence from experiments with tracers etc. So in this thesis only a cytological basis for phys­ iologicalinvestigation sha sbee ncreated . Further both the interrelationship between the different cell types and the combination of their occurrence have been studied. An attempt has been made to classify the Gymnosperms into a small number of different types, based on thecharacteristi c structure ofthei r secondary phloem.Thoug h the arrangement of the cells in the different Gymnosperms varies widely, each of these species belongs to one of the main types or is to be considered as an intermediate between two of those types. The phloem rays turned out to be most important inclassifyin g thevariou sphloe mtypes ,i ncontras t withth ewoo drays . The different types of secondary xylemar e arranged and classified in viewo f dead elements: tracheids, libriform fibres and vessel members. These are determining for a special wood species; the y are,excep t for thelibrifor m fibres, essentialt oth ewater-conductin g function ofth exylem . Theseelement sdevelop , differentiate and modify in the course of time, according to the changing ecologiesituations . The living part of the wood, the parenchymatous partan d thewoo d raysi n particular, also develop, but they are not essential to the vertical water- conducting systembu tthe y provide for the horizontal conduction of water and assimilates. The wood raysar e not interrelated and they do not interfere with the dead cells,whic h aredeterminin g for a special type ofwoo d species.There - Meded. Landbouwhogeschool Wageningen 67-7(1967) 1 fore the wood raysar e not essential in classifying woody species into several evolutionary stages, if only the wood is considered. This is entirely different with the phloem, which is mainly composed of living cells. It is a close, complex and active system; one of its functions is the conduction ofassimilates ,mainl y byth e sievecells ,throughou t the plant body; here the living phloem rays play a prominent part. A classification of the different Gymnosperm woods, based on the structure of the secondary phloem, ishardl ypossibl ewithou t consideringth ephloe mrays . Though it will appear that the phloem rays and the axial system of the phloem show different evolutionary trends, the former are still essential to the classification asmentione d above.Apar t from the rays and the axial system of the secondary phloem, the so-called 'Strasburger albuminous cells' are also important inthi sconnection . Theyar eintimatel y associated withth e sieve cells, whichar econsidere d tob eth emos timportan t elemento fth esecondar yphloem . Meded. Landbouwhogeschool Wageningen 67-7 (1967) 2. MATERIAL AND METHODS 2.1. SLIDES The 9 families of the Gymnosperms and the different genera belonging to these families, are: Cycadaceae Austrotaxus Widdringtonia Encephalartos Pseudotaxus Microcycas Taxus Taxodiaceae Cycas Torreya Athrotaxus Zamia Cryptomeria Macrozamia Cephalotaxaceae Cunninghamia Ceratozamia Cephalotaxus Glyptostrobus Metasequoia Ginkgoaceae Cupressaceae Sciadopitys Ginkgo Actinostrobus Sequoia Austrocedrus Sequoiadendron Araucariaceae Callitris Taiwania Agathis Calocedrus Taxodium Araucaria Neocallitropsis Chamaecyparis Pinaceae Podocarpaceae Cupressus Abies Acmopyle Diselma Cathaya Dacrydium Fitzroya Cedrus Microcachrys Fokienia Keteleeria Microstrobus Juniperus Larix Phyllocladus Libocedrus Picea Podocarpus Papuacedrus Tsuga Saxegothaea Pilgerodendron Pseudolarix Tetraclinis Pseudotsuga Taxaceae Thuja Pinus Amentotaxus Thujopsis Samples of stems of several Gymnosperm wood species were taken: in particular theanatomica l andth ecytologica l features of the secondary phloem are investigated and occasionally those ofth e xylem. These wood species are: Cycadaceae Cycas circinatumL . Ginkgoaceae Ginkgo biloba!^. Araucariaceae Agathis spec. Araucaria araucana K. KOCH Podocarpaceae Podocarpusnerifolius D.DO N Dacrydium elatum WALL. Meded. Landbouwhogeschool Wageningen 67-7(1967) 3 Sciadopitys verticillata SIEB,e t Zucc. Taxaceae Taxus baccataL . Torreya nucifera SIEB,e t Zucc. Cephalotaxus harringtonia R. SMITH Cupressaceae Chamaecyparispisifera ENDL. var.plumosa OTTO Biota spec. Cupressussempervirens L . Juniperus chinensisL . Juniperus communis L. Fitzroya cupressoides JOHN. Libocedrus decurrens TORR. Thujaplicata LAMB. Thujopsisdolabrata SIEB,e t Zucc. Cryptomeriajaponica D. DON Taxodiaceae. Cunninghamia Janceolata HOOK Metasequoia glyptostroboides Hu et CHENG Taxodium distichum A. RICH. Sequoiadendron giganteum BUCHH. Pinaceae Abies concolor HOOPES Tsugacanadensis CARR. Cedruslibani A . RICH. Larix decidua Hopes Pseudolarix kaempferi GORD. Picea spec. Pseudotsuga taxifolia BRITT. Pinussylvestris L. Also some samples ofth e Chlamydospermae wereexamined , viz.: Ephedraceae Ephedra spec. Gnetaceae Gnetumgnemon L. With chisel and hammer pieces of wood and bast arecu t from the different stemsa ta heigh t ofabou t oneyar d above thetrun k base;thes epiece s were taken in different seasons ofth e year. For some cytological reactions itwa s necessary to makesection s immediately after cutting thesamples , while for the normal anatomical studies the samples were killed and fixed first in formalin alcohol or Carnoy and next cut and stained.
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