Table Of Content40. 44.
I ~'
A selection of Hartig's (1837) drawings illustrating his Vergleichende Untersuchungen iiber die
Organisation des Stammes der einheimischen Waldbiiume: Fig. 23. Xylem element (species not
mentioned); Fig.24. course of vascular bundles in a nodal area (species not mentioned);
Fig. 40. sieve fibers ("Siebfasem") in a peripheral section of Pinus sylvestris; Fig. 41. sieve pores
viewed from the ray side (Pinus sylvestris); Fig. 42. sieve pores in sieve fibers of Acer, Aesculus,
Carpinus; a. sieve area of young sieve fiber of Carpinus; Fig. 43. part of a sieve tube of Acer; a.
sieve-like perforated cross walls in sieve tubes of Acer, Robinia (from pp 157-158; Hartig did
not provide legends)
H.-D. Behnke R. D. Sjolund (Eds.)
Sieve Elements
Comparative Structure, Induction and Development
With 266 Figures
Springer-Verlag Berlin Heidelberg New York
London Paris Tokyo Hong Kong
Professor Dr. H.-D. Behnke
Zellenlehre, Universitat Heidelberg
1m Neuenheimer Feld 230
0-6900 Heidelberg, FRO
Professor Dr. R. D. Sjolund
Department of Botany
University of Iowa
Iowa City, Iowa 52242, USA
Cover: Longitudinal section through the phloem of Cheiranthus cheiri with
mature sieve element in center. (From Behnke 1981, Nordic J Bot 1: 381-400)
ISBN-13: 978-3-642-74447-1 e-ISBN-13: 978-3-642-74445-7
DO I: 10.1007/ 978-3-642-74445-7
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Preface
As part of his Comparative Investigations of the Organization of the Trunk of the
Native Forest Trees (Theodor Hartig 1837, Vergleichende Untersuchungen tiber
die Organisation des Stammes der einheimischen Waldbaume. lahresberichte
tiber die Fortschritte der Forstwissenschaften und forstlichen Naturkunde 1:
125-168) Hartig gives an anatomical description of the "composition and
nature" of the then "completely uninvestigated elementary organs" of what he
called the "sap skin" (Safthaut) of trees, a tissue for which Nageli later (1858)
coined the term phloem. Within the "Safthaut" Hartig describes three cell types
in detail, (1) "Siebfasern", (2) "Siebrohren", and (3) "keulenfOrmige Saftrohren"
(club-shaped sap-tubes). While the description of the latter refers to laticifers in
Euphorbia and resin ducts in Acer and Robinia. "Siebfasern" and "Siebrohren"
comprise the sieve elements.
A literal translation of the more significant parts of the description of these
cell types demonstrates that his "Siebrohren" entirely correspond to what has
later been defined as "sieve tubes" but that his "Siebfasern" are less well
defined and in addition to what we call "sieve cells" also include small sieve
tubes as well as spindle-shaped cells of cambium, phloem parenchyma and
sclerenchyma. Both in his "Siebfasern" and "Siebrohren" Hartig describes sieve
areas (his expression is "lense-shaped cavities") and sieve pores (Siebporen).
from page 157:
"1) Sieve fibers
... We are able to shortly characterize them, if we say that they correspond most
to xylem fibers of the conifers both in form and in position. Comparable to
these, they show on the sides facing the rays lense-shaped cavities in the inter
cellular space ... which are penetrated by a great number of smaller pores.
Fig. 40 shows sieve fibers in a peripheral section from Pinus sylvestris. Fig. 41
shows sieve pores as viewed from the ray. ... Their formation within conifers
and deciduous trees basically is the same. In Alnus. Aesculus. Carpinus they are
distinguished by size of both the sieve fibers and the sieve pores, Fig. 42. In Car
pinus it can be seen that in those sieve fibers which are closest to the wood the
concavity of the cell wall, it is true, is already present, and even more depressed
than at a later time, but no porosity is displayed, instead, however, the surface
of the concavity appears to be occupied with small warts, Fig. 42 a. Are they
perhaps small, in the pores lying vesicles? ..."
from page 158:
"2) Sieve tubes
The sieve tube is to be paralleled to the xylem vessel as is the sieve fiber to the
VI Preface
xylem fiber. They are delicate tubes of considerable diameter which are densely
covered by sieve pores at sides not adjacent to rays, while at sides facing the
rays only roundish cavities, contoured by faint shadows, are shown; a detail by
which they are distinguished from sieve fibers, the pores of which are always on
the ray side. Their nature and their relationship to xylem vessels may best be
seen in the chestnut and in the maple from which I have depicted a part of a
tube, Fig.44." [This obviously is a misprint and should refer to Fig. 43, since
according to Hartig's earlier description Fig. 44 refers to sieve fibers of Acer
showing a "transition to pachydermal cells".] "At places where the sieve pores
are arranged very densely, the tube looks like a netlike spiral vessel with pitted
meshes; however, it can be distinctly recognized that the network is formed by
the close proximity of the sieve pores only. Without any visible segmentation of
the sieve tube into single members, sieve-like perforated cross walls can be seen
within the inner lumen at rather wide distances, Fig. 43 a (Acer, Robinia). The
pores are almost regularly hexagonal and sit so close together that the remain
ing cell wall looks like a network. ..." (See frontispiece for reproduction of
figures)
In the 150 years since the publication of Hartig's description of the phloem, our
knowledge about the structure and the function of sieve elements has increased
greatly. The improved resolution made possible initially by refinements in the
light microscope, and more recently through the development of the electron
microscope, has provided modem investigators a tool with which to extend
Hartig's early and limited observations down to the current level of organelles,
membranes and macromolecules. This book reviews the structure of the con
ducting elements of the major plant groups, from the algae to the flowering
plants, from extant and extinct groups (Chaps. 1 to 7 and 14), revealing both
common features and divergent solutions to the problem of nutrient transport.
Additional information about phloem structure and function is provided in the
coverage of special cases of phloem deVelopment (Chaps. 8 and 12), experi
mental systems that provide new insight in sieve element induction and forma
tion (Chaps. 9 to 11), and of specific phloem contents (Chap. 13).
From these pages it will become clear, however, that, despite 150 years of
progress, many fundamental questions concerning sieve-element structure,
development and function remain to be resolved.
Portions of this book were presented during the XIVth International Botani
cal Congress, Berlin 1987, in a symposium commemorating Theodor Hartig's
first description of sieve elements, 150 years ago.
October, 1989 H.-Dietmar Behnke, Heidelberg
and Richard D. Sjolund, Iowa City, Iowa
Contents
v
Preface.
1 Algae _______________________________________________ __
KLAUS SCHMITZ
1.1 Requirement for Medium-Distance and Long-Distance Transport
in Algae ........... .
1.2 Medium-Distance Transport ..
1.3 Long-Distance Transport . . . . 2
1.4 Conducting Cells of Red Algae. 2
1.5 Conducting Cells in Brown Algae 5
1.5.1 General Remarks ..... 5
1.5.2 Conducting Cells in Dictyotales, Scytosiphonales,
Desmarestiales and Fucales . . . 5
1.5.3 Sieve Elements in Laminariales. . . . . . . . . . . 8
2 Mosses
DANIEL C. SCHEIRER
2.1 Introduction .. 19
2.1.1 Overview 19
2.1.2 Terminology 19
2.2 General Organization of Conducting Tissues in Mosses 20
2.2.1 The Gametophyte .. 20
2.2.2 The Sporophyte ............... 20
2.3 Structure of Sieve Elements . . . . . . . . . . . . . 21
2.3.1 General Features of Moss Sieve Elements . 21
2.3.2 Differentiating and Mature Sieve Elements 21
2.4 Associated Parenchyma . .............. 31
3 Seedless Vascular Plants
RAy F. EVERT
3.1 Introduction . . . . . . . . . . 35
3.2 The Sieve-Element Protoplast 35
3.2.1 Nucleus ........ . 37
3.2.2 Endoplastic Reticulum 40
3.2.3 Plastids and Mitochondria 40
3.2.4 Dictyosomes . . . . . . . . 41
VIII Contents
3.2.5 Microtubules and Microfilaments 42
3.2.6 Plasmalemma and Tonoplast 42
3.2.7 Refractive Spherules. 43
3.3 The Wall . . . . . . . . . . . . . . . 48
3.4 The Sieve Areas . . . . . . . . . . . 51
3.5 Parenchymatous Cells Associated with the Sieve Elements 56
3.6 Longevity of the Sieve Elements 57
3.7 Comments on Terminology . . . . . . . . . . . . . . . . . . 57
4 Conifers ______________________________________________ __
ALEXANDER SCHULZ
4.1 Introduction............... 63
4.2 General Description . . . . . . . . . . . 64
4.2.1 Primary and Secondary Phloem 64
4.2.2 Shape and Size of Sieve Cells . 65
4.3 Development of the Sieve Cell 67
4.3.1 The Nucleus 68
4.3.2 Plastids.... 70
4.3.3 Mitochondria 73
4.3.4 Dictyosomes 75
4.3.5 Endoplasmic Reticulum (ER) and Ribosomes 75
4.3.6 Vacuole and Ground Plasm. 79
4.3.7 Structural Proteins . . . . . . . 80
4.3.8 The Wall . . . . . . . . . . . . 80
4.3.9 Intercellular Communication . 81
4.4 Strasburger Cells. . . . . . . . . . . . 83
5 Cycads and Gnetophytes ______________________________
H.-DIETMAR BEHNKE
5.1 Introduction....................... 89
5.2 Cycads . . . . . . . . . . . . . . . . . . . . . . . . . . 91
5.2.1 Organization and Composition of the Phloem 91
5.2.2 Ultrastructure of the Sieve Elements . . . . . . 91
5.2.3 Parenchymatous Cells Associated with the Sieve Elements 95
5.3 Gnetophytes........................... 95
5.3.1 Organization and Composition of the Phloem ....... 95
5.3.2 Ontogeny and Structure of the Mature Sieve Element . .. 97
5.3.3 Parenchymatous Cells Associated with the Sieve Elements 100
6 Dicotyledons _____________________________________________
RAy F. EVERT
6.1 Introduction............. 103
6.2 The Sieve-Tube Member Protoplast 103
6.2.1 Nucleus............ 105
Contents IX
6.2.2 Endoplasmic Reticulum. . 106
6.2.3 Plastids and Mitochondria 107
6.2.4 Dictyosomes ........ 109
6.2.5 Microtubules and Microfilaments 110
6.2.6 Plasmalemma and Tonoplast 110
6.2.7 P-Protein 111
6.3 The Wall ......... 119
6.4 The Sieve Plate. . . . . . 121
6.5 The Lateral Sieve Areas . 124
6.6 Parenchymatous Cells Associated with Sieve-Tube Members 125
6.7 Longevity of Sieve-Tube Members . . . . . . . . . . . . . . . 130
7 Monocotyledons
ELEFTHERIOS P. ELEFTHERIOU
7.1 Introduction 139
7.2 Ontogeny .... 139
7.3 The Protoplast . 141
7.3.1 Nucleus. 141
7.3.2 Plastids . 146
7.3.3 Endoplasmic Reticulum and Mitochondria 151
7.3.4 Microtubules and Dictyosomes . 151
7.3.5 P-Protein ......... 154
7.3.6 Vacuoles and Tonoplast . 154
7.4 Cell Wall ............ 155
7.5 Thick-Walled Sieve Elements. 155
7.6 Sieve Plates. . . . . . . . . . . 156
8 Sieve Elements in Internodal and Nodal Anastomoses of the
Monocotyledon Liana Dioscorea _______________
H.-DIETMAR BEHNKE
8.1 Introduction.......................... 161
8.2 The Vascular Construction in the Aerial Stem of Dioscorea 163
8.3 The Specific Composition of Phloem Anastomoses . 168
8.4 Ultrastructure of the Sieve Elements of Anastomoses 170
8.4.1 Connecting Sieve-Tube Members (CST) . . . 170
8.4.2 Anastomosal Sieve-Tube Members (ANST) . . 173
8.4.3 Anastomosal Sieve Elements (ANSE) . . . . . 173
8.5 Parenchymatous Cells Associated with the Sieve Elements of
Anastomoses. . . . . . . . . . . . . . . . . . . . . . . . . 175
8.6 Some Physiological Implications of Nodal Anastomoses ... 177
X Contents
9 Sieve Elements in Plant Tissue Cultures:
Development, Freeze-Fracture, and Isolation
RICHARD D. SJOLUND
9.1 Introduction ............... 179
9.2 Phloem Function in Vitro . . . . . . . . 180
9.3 Phloem Development in Callus Tissues 181
9.4 P-Protein, Callus Phloem and Wounding 181
9.5 Freeze-Fracture Studies Using Callus Sieve Elements 185
9.6 Sieve-Area Pores . . . . . . . . . . . . . . . . . . . . . 185
9.7 The Sieve-Element Reticulum (SER). . . . . . . . . . 187
9.8 Isolation and Partial Purification of Callus Sieve Elements 193
9.9 Antibody Formation Against Callus Sieve Elements .... 195
10 Wound-Sieve Elements ___________________
ALEXANDER SCHULZ
10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . 199
10.2 Tissue Changes During Wound-Phloem Development. 199
10.2.1 Tissue of Origin . . . . . . . . . . . . . . . 199
10.2.2 Developmental Pattern of Wound Phloem 201
10.2.3 Size and Shape of Wound-Sieve Elements. 203
10.3 Cytoplasm of Wound-Sieve Elements 205
10.3.1 Nucleus and Vacuole . . . . . . 205
10.3.2 Sieve-Element Plastids ..... 207
10.3.3 Mitochondria and Dictyosomes 208
10.3.4 Endoplasmic Reticulum and Ribosomes 208
10.3.5 P-Protein and Microtubules. . . . . . . . 209
10.4 Symplastic Connections of Wound-Sieve Elements. 210
10.4.1 Connections Between Sieve Elements . . . . 210
10.4.2 Connections from Sieve Elements to Other Cell Types 214
10.5 Companion Cells . . . . . . . . . . . . . . . . . . . . . . . 215
10.6 Comparison Between Wound-and Bundle-Sieve Elements . 215
11 Sieve Elements of Graft Unions ________________
RAINER KOLLMANN and CHRISTL GLOCKMANN
11.1 Introduction . . . . . . . . . . . . . . . . 219
11.2 Grafting Procedure ..... . . . . . . . 221
11.3 Histology and Cytology of the Graft Union . 221
11.3.1 General Aspects of Development .. 221
11.3.2 Sieve-Element Contact at the Graft Interface 224
11.4 Function of Phloem Connections in Graft Unions . 232
11.5 Questions Concerning the Mechanism of Sieve-Element
Formation in Graft Unions . . . . . . . . . . . . . . . . . ?33
