A
Stomata
SECOND EDITION
Colin Willmer
University of Stirling,
UK
and
Mark Fricker
University of Oxford,
UK
CHAPMAN & HALL
London • Glasgow • Weinheim • New York • Tokyo • Melbourne • Madras
Contents
Preface to the second edition
ix
Preface to the first edition
xi
Acknowledgements
1
1.1
1.2
1.3
xiii
Introduction
General considerations about stomata
Historical aspects
The importance of research into stomatal functioning and
behaviour
References
1
1
2
7
8
2
2.1
2.2
2.3
2.4
The distribution of stomata
Location and occurrence of stomata
Stomatal frequency
Stomatal patterning
Heterogeneity in stomatal characteristics and responses
References
12
12
17
24
26
30
3
3.1
3.2
3.3
The structure and development of stomata
Anatomy and morphology of the leaf epidermis
Development and differentiation of stomata
The ultrastructure of guard cells
References
36
36
49
64
82
4
4.1
4.2
4.3
4.4
4.5
5
5.1
5.2
5.3
5.4
.
,
The mechanics of stomatal movements
Introduction
Water relations of guard cells
.
The physico-chemical aspects of stomatal movements
Morphological changes of the plasma membrane and
tonoplast of guard cells during stomatal movements
Physiological and mechanical properties of GCPs
References
92
92
92
96
103
104
106
The theory of gas diffusion through stomata
Introduction
Fick's law and the driving forces for diffusion
The analogy between electrical circuits and leaf resistances
Diffusion through a single pore and multipores
110
110
110
113
115
vi
CONTENTS
5.5
5.6
5.7
6
6.1
6.2
6.3
6.4
6.5
6.6
6.7
6.8
6.9
6.10
6.11
6.12
7
7.1
7.2
7.3
7.4
7.5
7.6
7.7
7.8
7.9
8
8.1
8.2
8.3
8.4
8.5
Cuticular conductance
Boundary layer conductance
Water use efficiency
References
. - -
120
121
123
124
Stomatal responses to environmental factors
126
126
Introduction
Interaction of environmental factors which influence stomatal
behaviour
126
Stomatal responses to light quality and quantity
131
144
Carbon dioxide
Temperature
148
152
Plant water status and atmospheric humidity
161
Mineral nutrition
Circadian rhythms and shorter-term oscillations in stomatal
161
movements
Stomatal activity during the life cycle of a leaf
165
168
Stomatal behaviour in canopies
Gaseous environmental pollutants and stomata
169
Plant pathogens and stomata
172
177
References
The influence of hormones and other naturally occurring
compounds on stomatal behaviour
Introduction
Abscisic acid
Other sesquiterpenoids (xanthoxin, al\-trans farnesol,
vomifoliol, phaseic acid)
Other hormones
Fatty acids
Phytotoxins
Phytoalexins
Other compounds
Artificial control of stomata
References
Ionic relations of stomatal movement and signal
transduction in guard cells
Introduction
The involvement of ions in stomatal movements
Ion fluxes and the driving forces for ion movement
Summary of ion movement events durins stomatal
movements
Signal perception and transduction by guard cells
References
192
192
193
208
209
212
212
215
215
216
218
228
228
228
244
CONTENTS
9
9.1
9.2
9.3
9.4
9.5
9.6
9.7
10
10.1
10.2
10.3
10.4
10.5
10.6
10.7
10.8
10.9
10.10
Index
The metabolism of guard cells
Introduction
The flow of carbon from starch to malate during stomatal
opening
The fate of malate during stomatal closure
The role of chloroplasts in guard cells
Metabolite transport into guard cells and sucrose
metabolism
The energy relations of stomatal movements
Lipid metabolism in guard cells
References
310
310
Recent developments and future research
Introduction
Stomatal distribution and patterning
Cell and tissue differentiation
Mechanical relations of the epidermis
Diffusion of gases and the role of stomata
Environmental effects
Hormone action
Ion transport and signalling
Intermediary metabolism and its regulation
Molecular biology of stomata
References
354
354
354
355
356
356
356
358
360
360
361
364
315
330
335
339
341
344
345
367
VII