Photosynthesis during leaf development

394 Pages·1985·21.211 MB·English

by  Z. à esták, J. Čatský (auth.), Zdenĕk à estăk (eds.)

Photosynthesis during leaf development
Tasks for vegetation science 11
Series Editors
HELMUT LIETH HAROLD A. MOONEY
University of OsnabrUck, F.R.G. Stanford University, Stanford CA, U.S.A.
Photosynthesis during
leaf development
edited by
ZDENEK SESTAK
Contributors:
Zdenlk Sestdk, Jin Catskj,
Ingrid Tichd, DanuIe Hoddiiovd, Jaromir Kutik,
Jarmi/a So/drovd, Jana Pospisi/ovd,
Jan Zima, Mi/os/av Kase
1985 DR W. JUNK PUBLISHERS
a member of the KLUWER ACADEMIC PUBLISHERS GROUP
DORDRECHT / BOSTON / LANCASTER
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ry, Mongolia, Northern Korea, Poland, Rumania, U.S.S.R., Vietnam, and Yugoslavia:
Academia, Publishing House of the Czechoslovak Academy of Sciences, P.O.B. 896,
11229 Prague 1, Czechoslovakia
for all other countries: Kluwer Academic Publishers Group, Distribution Center,
P.O. Box 322, 3300 AH Dordrecht, The Netherlands
Library of Congress Cataloging in Publication Data
Main entry under tide:
Photosynthesis during leaf development.
(Tasks for vegetation science; 11)
Includes bibliographical references and indexes.
1. Photosynthesis. 2. Leaves-Development.
I. Sestak, Zdenek, 1932 - • II. Series.
QK882.P554 1984 581.1'3342 83-14992
ISBN-13: 978-94-010-8941-8 e-ISBN-13: 978-94-009-5530-1
DOl: 10.1007/978-94-009-5530-1
Scientific Editor: Prof. RNDr. PhMr. Miroslav Penka, DrSc.
Scientific Adviser: RNDr. Bohdan Slavik, DrSc.
Joint edition published by
Dr W. Junk Publishers,
P.O. Box 163,3300 AD Dordrecht, The Netherlands
and Academia, Publishing House of the Czechoslovak Academy of Sciences,
P.O.B. 896, 11229 Prague 1, Czechoslovakia
Copyright
© 1985 by Academia, Prague
Softcover reprint of the hardcover 1s t edition 1985
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, mechanical, photocopying, re
cording, or otherwise, without the prior written permission of the publishers.
CONTENTS
o Leaf Development - Terms and Photosynthetic Aspects 11
(Z. Sestak and J. Gatsky)
0.1 Phases of Leaf Development and Insertion Profile 11
0.2 Common Imperfections of Leaf Ontogenetic Studies 13
0.3 Specialized Monographs and the Scope of this Book 14
1 Ontogeny of Leaf Morphology and Anatomy 16
(Ingrid Ticha)
1.1 Leaf as a Photosynthetic Structure • • • • • • • 16
1.2 Leaf Growth, Leaf Area and Size, Leaf Thickness 16
1.2.1 Leaf Area 16
1.2.2 Leaf Size and Shape • • 20
1.2.3 Leaf Thickness . • • • 21
1.2.4 Leaf Plastochron Index 21
1.3 Leaf Epidermis • . • • • • • 22
1.3.1 Number of Epidermal Cells, their Sizes, and Thickness of Epidermis. 22
1.3.2 Cuticle, Waxes, Trichomes 23
1.3.3 Stomata. • • • • • • • 24
1.3.3.1 Stomata Density • 24
1.3.3.2 Stomata Size • . . 34
1.3.3.3 Absolute Number of Stomata per Leaf • 35
1.3.3.4 Stomatal Index . • • • . • • • 35
1.4 Leaf Mesophyll • • • • . • • . • • . • • • 36
1.4.1 Leaf Anatomy of C3, C4, and CAM Plants 36
1.4.2 Mesophyll Thickness and Tissue Volumes 38
1.4.3 Intercellular Air Spaces and Internal Leaf Surface • 40
1.4.4 Mesophyll Cells • • . . • • 41
1.4.5 Chloroplasts • • • • • • • • 42
1.4.5.1 Chloroplast Numbers 43
1.4.5.2 Chloroplast Sizes. 48
1.4.6 Vascular System 49
1.5 Zalenskii Law. 49
1.6 Conclusions 50
5
2 Chloroplast Development 51
(J. Kulik)
2.1 Chloroplast Structure • • • • • • • • • 51
2.2 Ontogeny of Chloroplast Structure 55
2.2.1 Outlines of Chloroplast Development 55
2.2.2 Non-ontogenetic Changes in Chloroplast Structure • 57
2.2.3 Chloroplast Ultrastructure during Leaf Development 58
2.2.3.1 Leaves of Herbs and Deciduous Woody Plants 58
2.2.3.2 Leaves of Evergreen Woody Plants • • • • • . 66
2.2.3.3 Chloroplast Ontogeny in Relation to Ca, C4, or CAM Pathway 70
2.2.4 Chloroplast Substructure during Leaf Development • • • • • • • • • 71
2.2.5 Environmental or Experimental Modifications of the Chloroplast Ontogeny • 72
2.3 Conclusions ••••••••••••••••••••••••••••••• 74
3 Chlorophylls and Carotenoids during Leaf Ontogeny 76
(Z. Sestak)
3.1 The Importance of Chlorophyll and Carotenoids • 76
3.2 Biosynthetic Processes • • • • • • • • 77
3.2.1 Chlorophyll Biosynthesis. • • • • • • • • 77
3.2.1.1 Precursors of Chlorophyll • • • • • 77
3.2.1.2 Enzymes of Chlorophyll Biosynthesis. 80
3.2.1.3 Chlorophyll Biosynthesis in Various Parts of the Leaf Blade 80
3.2.1.4 Chlorophyll Biosynthesis and Chloroplast Ultrastructure. 81
3.2.2 Carotenoid Biosynthesis • • • • • 84
3.3 Pigment Contents during Leaf Ontogeny • 84
3.3.1 Chlorophyll Contents • • • • • • • 84
3.3.1.1 Herbs and Deciduous Trees. 84
3.3.1.2 Evergreen Plants. • • • • • 88
3.3.2 Carotenoid Contents • • • • • • • 89
3.3.2.1 Herbs and Deciduous Trees • 89
3.3.2.2 Evergreen Plants. • • • 91
3.4 Leaf Insertion and Pigment Contents • 91
3.4.1 Chlorophyll and Leaf Insertion 91
3.4.2 Carotenoids and Leaf Insertion • 96
3.5 Pigment Distribution in Leaf Blade • 97
3.5.1 Chlorophylls. • • • 97
3.5.2 Carotenoids • • • • 98
3.6 State of Pigments in Vivo • 99
3.6.1 Chlorophylls in Vivo 99
3.6.2 Carotenoids in Vivo. • 105
3.7 Leaf Age Effects Observed in Detached Leaves. 105
3.8 Conclusion • • • • • • • • • • • • • • • • 106
4 Leaf Optical Properties 107
(DanuIe Hoddiiovd)
4.1 Introduction • 107
4.2 Photoreceptors and Mechanisms • 110
4.2.1 Absorption • • • • • • • 110
6
4.2.2 Transmission • 112
4.2.3 Reflection • . 113
4.3 Leaf as an Optical System • 114
4.3.1 Scattering and Optical Path of Radiation Inside a Leaf. 114
4.3.2 Dorsiventrality of a Leaf • • • • • • 116
4.3.3 Attenuation of Radiation Inside a Leaf • 118
4.4 Variability in Leaf Optical Properties. . . . 120
4.4.1 Leaf Ontogeny. • • • • • • • • • • 120
4.4.2 Insertion Level and Sequential Age of Leaves • 125
4.5 Conclusions ••••••••••••••••• 126
5 Changes in Electron Transport Chain Composition, and Activities of Photo
systems and Photophosphorylation during Leaf Ontogeny . . . . . . . . . . . 128
(Z. Sestdk)
5.1 Principles of Photosynthetic Electron Transport Chain and Photophosphorylation. 128
5.2 Composition of Thylakoids: Components of Electron Transport Chain 130
5.2.1 Plastoquinones . 132
5.2.2 Cytochromes 132
5.2.3 Ferredoxin 133
5.3 Activities of Photosystems 133
5.3.1 Photosystem 2 . . • 134
5.3.2 Photosystem 1 • • . 138
5.3.3 Ratio of Activities of Photosystems 1 and 2 • 140
5.4 Photophosphorylation . . . • . . • . • 142
5.5 General Course of Photochemical Activities • • 144
6 Carbon Fixation Pathways, their Enzymes and Products during Leaf Ontogeny 145
(J. Zima and Z. Sestak)
6.1 Carbon Fixation Pathways • • • • • . 145
6.2 Enzyme Activities during Leaf Ontogeny 146
6.2.1 RuBPC and PEPC in Ca Plants. • 146
6.2.2 RuBPC and PEPC in C4 Plants . 149
6.2.3 RuBPC and PEPC in CAM Plants . 150
6.2.4 Other Enzymes of Carbon Metabolism in Ca Plants 151
6.2.5 Other Enzymes of Carbon Metabolism in C4 and CAM Plants 153
6.3 Changes in Carbon Fixation Pathways and Primary Photosynthates • 153
6.4 Secondary Photosynthates 155
6.5 Conclusion • • • • • . • • • . • • • • • • • • • • . . • . • 156
7 Gas Exchange and Dry Matter Accumulation during Leaf Development 157
(Ingrid Ticha, J. Gatskj, DanuSe Hodaiiova, Jana PospiIilova, M. KaIe and Z. Sestak)
7.1 Photosynthetic Gas Exchange. . . . . . • • • • . . • . . . 157
7.1.1 C02 Fixation Types and their Response in C02 Exchange 160
7.2 Net Photosynthetic Rate • . . • • . . • • • • • . 161
7.2.1 Net Photosynthetic Rate during Leaf Ontogeny 161
7.2.2 LeafInsertion and Net Photosynthetic Rate. . . 172
7.2.3 Net Photosynthetic Rate on Leaf Blade Area . • 181
7.2.4 Modifications of Developmental Pattern in Net Photosynthetic Rate 182
7
7.2.5 Irradiance . • • . • . • • • • • • • • . • • . • • • • • • • •• 183
7.2.5.1 The Dependence of Net Photosynthetic Rate on Leaf Irradiance 183
7.2.5.2 Adaptation to Growth Irradiance . . . . . . . . . • . 186
7.2.5.3 Energy Conversion in Photosynthesis . . . . . . . • • . 187
7.2.5.4 Photosynthesis, Energy Conversion and Spectral Irradiance 188
7.2.6 Carbon Dioxide Concentration •...••....•.• 189
7.2.6.1 Short Term COg Effects on Net Photosynthetic Rate 190
7.2.6.2 Carbon Dioxide Compensation Concentration 193
7.2.6.3 Long Term C02 Effects (C02 Enrichment) 199
7.2.7 Temperature 200
7.2.8 Water Stress . . • 202
7.2.9 Oxygen Effect .•. 203
7.2.10 Mineral Nutrition 204
7.2.11 True (Gross) Photosynthetic Rate and Leaf Ontogeny. 204
7.3 Respiration Rate • • . • . • . . • • • • • • • • . . • 205
7.3.1 Dark Respiration Rate and Leaf Age. . . . • . . • • 206
7.3.2 Environmental Modifications of Ontogenetic Changes in Respiration 209
7.4 Dry Matter Accumulation . . . . . . . • . . . . 210
7.4.1 Dry Matter Accumulation during Leaf Ontogeny 210
7.4.2 Leaf Insertion and Dry Matter Accumulation • 213
7.4.3 Carbon Budget of the Developing Leaf. 214
7.5 Conclusions 215
8 Conductances for Carbon Dioxide Transfer in the Leaf. . . . . . . . . . . . 217
(J. Catsky, Jarmila Solarova, Jana PospiIilova and Ingrid Ticha)
8.1 Carbon Dioxide Transfer in the Gaseous Phase. . . 218
8.1.1 Conductance of Leaf Boundary Layer. . . 219
8.1.2 Epidermal Conductance and its Components. 221
8.1.3 Stomata and Stomatal Conductance. . . . . 222
8.1.4 Ontogenetic Changes in Maximum Values of Stomatal Diffusive Conductance. 223
8.1.5 Leaf Insertion Level and Stomatal Diffusive Conductance. . . . . . . . . 231
8.1.6 Stomatal Conductance and Leaf Blade Heterogeneity. . . . . . . . . . . . 233
8.1.7 Ontogenetic Changes in Responses of Stomatal Diffusive Conductance to
Environmental and Internal Factors. . . . . . . . . . 233
8.1.7.1 Diurnal Course of Stomatal Conductance. . . . . 234
8.1.7.2 Response of Stomatal Conductance to Irradiance. . 235
8.1.7.3 Response of Stomatal Conductance to Water Stress. 235
8.1.7.4 Response of Stomatal Conductance to Temperature. 239
8.1.7.5 Effects of C02, S02, and Nitrogen Supply. 239
8.1.8 Intercellular Conductance. . . . . . . 240
8.2 Carbon Dioxide Transfer in the Liquid Phase. . . . . 240
8.2.1 The Overall Intracellular Conductance. . . . . . 240
8.2.2 Ontogenetic Changes in the Overall Intracellular Conductance 241
8.2.2.1 Comparison of the Ontogenetic Changes in Intracellular Conductance
of Ca and C4 Plants. . . . . . . . . . . . . . . . . . . . . . 243
8.2.3 Components of Intracellular Conductance . . . . . . . . . . . . . . . . 244
8.2.3.1 Transport and Carboxylation Components of Intracellular Conductance 244
8.2.3.2 Transport Component Calculated from Leaf Anatomy . . 244
8.2.4 Transport and Carboxylation Conductances and Enzyme Activities . . . . . 245
8
8.2.4.1 Carbonic Anhydrase ...................... 245
8.2.4.2 Carboxylation Conductance and Ribulose-1,5-bisphosphate Carboxylase 246
8.2.5 Excitation Conductance and Photosynthetic Efficiency . 246
8.2.6 C02 Transport on Individual Cell Structures . 248
8.3 Conclusions • . . . . . . • • . . . . . . . • 249
9 Photorespiration during Leaf Ontogeny 250
(J. Catsky and Ingrid Tichtf)
9.1 Photorespiration and Photosynthesis . . . . . . . . . . 250
9.2 Measures of Photorespiration Rate. . . . . . . . . . . 251
9.3 Ontogenetic Changes in Photorespiration Rate in Cs Plants 252
9.4 Ontogenetic Changes in Light Respiration in C4 Species. 261
9.5 Conclusions .•. . . . • . • • • . • • . . . . . 262
10 Integration of Photosynthetic Characteristics during Leaf Development 263
(Z. Sestak, Ingrid Tichd, J. Catsky, Jarmila Soldrova, Jana PospfSilova and Danuse Hodd
novtf)
10.1 General Considerations . • • • . . . . 263
10.2 Leaf Structure and Photosynthesis . . . 264
10.3 Chloroplast Formation and Degeneration. 268
10.4 Chlorophyll and Photosynthetic Activities 269
10.5 Activities of Photochemical Reactions and the Photosynthetic Rate. 272
10.6 Activities of Photosynthetic Enzymes and Photosynthetic Rate. 273
10.7 Photosynthesis and Conductances for Carbon Dioxide Transfer . . 274
10.8 Photosynthesis and Respiration. . . . . . . . . . . . . . . . 276
10.9 Photosynthesis and Dry Matter Accumulation. The Carbon Balance of the Leaf. 280
10.10 The Ratio between Photosynthetic and Transpiration Rates 281
10.11 Conclusions . • • . • . • • .• ••••... 283
11 References 287
Authors' Index 365
Subject Index 386
Plant Index. . 393
9
o
LEAF DEVELOPMENT - TERMS
AND PHOTOSYNTHETIC ASPECTS
z. Sestak and J. Catskj
Photosynthesis is the primary metabolic process in nature: without photosynthe
sis life on our planet would cease. Green plants together with photosynthetic
bacteria are the only primary producers on Earth: they provide energy and biomass
for animals and men and keep the oxygen stable in the atmosphere.
The most important producers of biomass and oxygen are higher plants (mainly
forming tropical forests) which produce about 70% of the total biomass, i.e.
about 12 X 1013 kg per year (e/. Lieth and Whittaker 1975, Whittaker and Likens
1975), a great deal of it being of immediate significance for man as nutrition or
raw materials. Crops on cultivated land produce about 5% of the total biomass,
i.e. less than 1010 t per year.
Even if we take into account the important contribution of photosynthesis of
ears, stems and other green plant parts to the total plant biomass production, the
major role in most plants is played by the leaves, the total surface of which is in
the biosphere about 65x 107 km2• During its life, however, the leaf morphology,
structure of its tissues and cells, and its composition change, and physiological
and biochemical activities (including those embraced by the concept of photo
synthesis) develop in a manner similar to that of any creature.
0.1 PHASES OF LEAF DEVELOPMENT
AND INSERTION PROFILE
During leaf development, or leaf life span, generally three phases may be distingui
shed: (1) leaf formation related to leaf area increase, (2) the period of leaf maturity
after maximum leaf area is reached, and (3) senescence associated with a decrease
in leaf area.
Leaf formation begins with the origin of leaf primordia at the shoot apex. Cell
division is replaced by cell enlargement as the chief means ofleaf expansion when
leaves have attained about one third or half of maximum leaf area (c/. Avery 1933,
Hannan 1968). The duration of leaf maturity is different in herbs, deciduous
11