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Part 1
hysiological Implications of Oscillatory Processes in Plants ....
I Rhythmic Leaf Movements: Physiological
and Molecular Aspects . . . .. ..... ... 3
NAVA MORAN
A bstract . ........... .......... ......... .
.1 Introduction . . ....... ..... ...... ...... 3
1.1.1 Historical Perspective .. ...... ....... 3
1.1.2 The Types of Leaf Movements............. ..... 4
1.2 The Mechanism of Leaf Movement: the Osmotic Motor........ ... 7
1.2.1 Volume Changes ......... ......... .. . ... 7
1.2.2 The Ionic Basis for the Osmotic Motor ...... ..... ..... 8
1,2.3 Plasma Membrane Transporters . .... 10
1.2.4 Tonoplast Transporters -...... . .. . . . . . . . 16
13 Mechanisms of Regulation .. ...... .
.3.1 Regulation by Protein Modification - Phosphorylation . ..... 17
1.3.2 The Perception of ight . .......... 21
13.3 Intermediate Steps . . . .. . .. .23
13.4 Regulation by Other Effectors .. . . ... . . ... ... ..... .28
1A Unanswered Questions . .. ........ .. 30
1.4.1 Acute, Fast Signalling . . ..... ... .. . 31
1.4.2 The Clock Input and Output...... ... . 31
References ... .. ...... . . ..... ... . - 32
2 The Pollen Tube Oscillator: Integrating Biophysics and Biochemistry
into Cellular Growth and Morphogenesis ........................ ... 39
NUNO MORENO, RENATO COLAQO AND Joss A. FEni
Abstract . . ....... ..... ............... ...... . -39
2.1 Finding Stability in Instability . . . . ... 39
2.2 W hy Pollen Tubes? .... .. ... ......... . . . .. . 42
2.3 Growth Oscillations: Trembling with Anticipation? ........ . ... 42
2.4 Under Pressure ... ..... ... 45
2.5 Another Brick in the Cel l Wal . .... . . . ... 46
.6 Cytosolic Approaches to Oscillations: the Ions Within .......... ... 47
2.7 On the Outside: Ions and Fluxes ........ 51
2.8 Actin Cytoskeleton: Pushing it to the Limit .. ..... 54
2.9 Membrane Trafficking and Signalling on the Road .................55
2.10 Conclusions . . . . . . . . . . . . .. .... . 57
References . .. ... ............. . . . 58
3 Ultradian Growth Oscillations in Organs:
Physiological Signal or Noise? . .. .... .. . .. .... ..,.. ... 63
TOBIAS I. BASKIN
Abstract .. - ..... 63
3.1 Introduction ....... .. .......... . ...... -63
3.1.1 Oscillations as Window into Growth .... .. ........ . 63
3.1.2 Growth Versus Movement .... ....... 65
3.2 Circumnutation: Growing Around in Circles? ...... .... ....... 65
33 In Search of ltradian Growth Oscillations .............68
3.4 The Power of Bending in Plants ........... 70
3.5 Conclusion and Perspectives . .............. 73
References . ................. . ... . ... . 73
SNutationin Plants . .... ......... .... ... . 7
SERGIO MUGNAI, ELISA AZZARELLO, ELISA MASI, CAMILLA PANDOLFI
AND STEFANO MANCUSO
Abstract . . . ... ... . ................ 77
4.1 Introduction ............. .... ... ..... ... ...... 77
4.2 Theories and Models for Circumnutation ... ..... ....... . 81
4.2.1 'Internal Oscillator' Model . . ....... ...... ... 83
4.2.2 'Gravitropic Overshoot' Model .... ..... 84
4.2.3 The 'Mediating' Model ......... ....... -85
4.3 Root Circumnutation . . . . ... ... .. . .... . ..... ...... . . 86
References ... . . . ................ 88
Part 2
Stomata Oscillations ....... .... ........ ............. ... ........ 91
5 Oscillations in Plant Transpiration ... . ......... .........93
ANDERS JOHNSSON
Abstract .... ... ..... . 93
5.1 Introduction ...... . . . . . . ... ...... . . 93
5.2 Models for Rhythmic Water Transpiration ..... ................ 95
5.2.1 Overall Description - "Lumped" Model .................. 95
5.2.2 Overall Description - "Composed" Models 97
5.2.3 Self-Sustained Guard Cell Oscillations -
(CaW),, Oscillations . . . 98
5.2.4 W ater Channels ..... . . . .. ....... ... . - 98
5,2.5 Comments on Modelling Transpiration Rhythms .......... 99
53 Basic Experimental Methods Used ........... ...... 99
5.4 Experimental Findings on Transpiration Oscillations .............. 100
5.4.1 Occurrence of Transpiration Rhythms: Period of Rhythms . 01
5.4.2 Some Environmental Parameters Influencing Oscillations . 101
5.4.3 Singularities of Transpiration Rhythms: Test of Models ....... 104
5.5 Ionic Interference with Transpiration Oscillations ...... .... ... 105
5.6 Patchy Water Transpiration from Leaf Surface .................. 106
5.7 Period Doubling and Bifurcations in Transpiration -
a Way to Chaos? . .. . . . . . . . . 107
5.8 Conclusions ... ... ...- ........ ... . . 109
115
References ... ............... ............ ...........
6 Membrane Transport and Cal Oscillations in Guard Cells l..........,. 115
MICHAEL R. BLATT, CARLOS GARCIA-MATA AND SERGEI SOKOLOVSKi
Abstract ..................... ........... 11
6,1 Introduction . ... . . . ........ . .... . . 15
62 Oscillations and the Membrane Platform .................. 116
6.3 Elements of Guard Cell Ion Transport ....... ... 19
6.4 Cal2 and Voltage . ..... ... ....... .... . 121
6.4.1 The Ca> Theme ........... . .......... 122
6.4.2 [Cafl, Oscillations . .. . . ..... .. . . . 123
6,43 Voltage Oscillations .. . .. . . . . .. . .. 124
6.4.4 Membrane Voltage and the '[Ca ']Cassette' . ........ ... . 125
6.5 Concluding Remarks ....... ...... . 127
References -... 128
7 Calcium Oscillations in Guard Cell Adaptive Responses
to the Environment ......... .. .... ........... 135
MARTIN R. McAINSH
Abstract . . . . ..... .... .. . .... . 135
7.1 Introduction . . . ... .... . .................. .... 135
7.2 Guard Cells and Specificity in Ca2 Signalling ......... .. 137
3 Ca Signatures: Encoding Specificityin Ca Signals ...... .... 138
7.4.1 Guard Cell Ca2> Signatures: Correlative Evidence ..... 140
7.4.2 Guard Cell Ca2' Signatures: Evidence
for a Causal Relationship . . ... ...... . . . .. 146
74.3 Guard Cell Ca Signatures: the Role of Oscillations ......... 147
7.5 The Cal Sensor Priming Model of Guard Cell Ca2 Signalling ........ 148
7.6 Decoding Ca2l Signatures in Plants ............. ... . ..... . 149
7.7 Challenging Prospects .... . . . . . . . . . . . . . . . ...... . 150
References . . . . . ... . .. ... ! 52
8 Circadian Rhythms in Stomata: Physiological
and Molecular Aspects ............. .... ......... ... 157
KATHARINE E. HUBBARD, CARLS T. HOTTA, MICHAEL l. GARDNER,
SOENG JN BAER, NEIL DALCIAU, SUHITA DONTAMALA,
ANTONY N. DODD AND ALEX A.R. EBB
Abstract . .. . . . ........ .. - ...... ........... ..... . 157
8.1 Introduction ......... .. .... .. . . . . . . . . . .. . . . . 157
8.2 Mechanisms of Stomatal Movements ....... ......... ..... 159
8.3 The Circadian Clock .. . .. ... ... ....... 162
8.4 Circadian Regulation of Stomatal Aperture ...... ..... . 164
8.5 Structure of the Guard Cell Clock ... .... . ........... . ... ... 166
8.6 Mechanisms of Circadian Control of Guard Cell Physiology ..... 168
8.6.1 Calcium-Dependent Models for Circadian
Stomatal Movements . . . ..... . 169
8.6.2 Calcium-Independent Models for Circadian
Stomatal Movements ............. . 170
8.7 Circadian Regulation of Sensitivity of Environmental
Signals ('Gating') ..... . .. . . 17
S Conclusions .... . .. . ..... 172
References ....... ........ _.............. ........ ..172
Part 3
Rhythms, Clocks and Development ................... ....... 179
9 How Plants Identify the Season byUsing a Circadian Clock . ............ 181
WOLFGANG ENGELMANN
Abstract , _ _ _ , . ......... .................. _ 181
9.1 Introduction and History . ...... .... 181
9.2 Examples for Photoperiodic Reactions .................... 184
9.3 Binning Hypothesis and Critical Tests ................. .... 185
9.4 The Circadian Clock and its Entrainment to the Day ......... 189
9.5 Seasonal Timing of Flower Induction ........ . 91
References . .......... .......................................194
10 Rhythmic Stem Extension Growth and Leaf Movements
as Markers of Plant Behaviour: the Integral Output from
Endogenous and Environmental Signals ............................ 199
JOHANNES NORMANN, MARCO VERVLUET-SClEEBAUM,
JOLANA T.P. ALBRECHTOVA AND EDGAR WAGNER
A bstract .. . .. .. .. ..... .. .. . .... ..... ...... .. . 199
101 Introduction . . .. . . . . 200
10.1.1 Life is Rhythmic. ....200
10.1.2 Rhythm Research: Metabolic and Genetic
Determination of Rhytmic Behaviou......... 20
10.2 Rhythmicityin Chenopodiun spp .............. ....... 203
10.2.1 Rhythmic Changes in Interorgan Communication
of Growth Responses . ...................... .. 206
10.2.2 Local Hydraulic Signalling: the Shoot Apex in Transition .209
10.2.3 Membrane Potential as the Basis for Hydro-
Electrochemical Signalling, Interorgan Communication
and Metabolic Control .. ..... . ........... . . 212
10.3 Conclusions and Perspectives: Rhythms in Energy Metabolism as
Determinants for Rhythmic Growth and Leaf Movements .. ..... 213
References ... . .. .. . .. .. 215
11 Rhythms and Morphogenesis ............. ............. .......... 219
PETER W. BARLOW AND JACQUELINE LOCK
Abstract ................ . .... ... .. 219
11.11 Introduction . .... ............... . .. . 219
112 Developmental Theories and Their A plication to Rhythmic
Morphogenesis .... .. .. .... ... ..... . ..... .. 220
11.3 Rhythmic Patterns of Cellular Development Within Cell Files ..... 221
114 Organogenetic Rhythms . . . . . . . . . . . . . ..... . 227
114.1 Angiosperm Shoot Apices and Their Phyllotaxies ...... . 228
11.4.2 ThePlastochron -. . . . . . . . . . . . . . . .. 231
11.43 A Petri Net Representation of the Plastochron .......... 232
11.4.4 Rhythms of Cell Determination and the Plastochron ....... 236
11.5 The Cycle of Life .. ........ ... . . ..., . .237
11.6 A Glimpse of Cell Biology and Morphogenetic Rhythms ..... .. 238
References . .. ..... ... . . .. . 240
12 Molecular Aspects of the Arabidopsis Circadian Clock ......... .. 245
TRAcEY ANN CUIN
A bstract . . . . . ...... .. ... ... . ... . . . . . .. . 245
12.1 Introduction . .. ................. .. ..... . .. .... .. . 245
12.1.1 Defining Features of Circadian Rhythms . ......... .... . 246
12,1,2 Overview of the Circadian System in Arabidopsis ....... 246
12.2 Entrainment ~ Inputs to the Clock .... ...... . . . 247
12.2.1 Light ......._ . ... ... . . .. .... 247
!2.2.2 Pathways to the Central Oscillator ...... . ..249
12.2.3 Negative Regulation of Photoentrainment ..... . ...... 253
12.2.4 Temperature Entrainment ...... . ..... ..... ..... . 253
12.3 The Central Oscillator .... . . .. .. . . . .... . 254
12.3.1 The CCAI/LHY-TOC Model for the Arabidopsis
Central Oscillator . ............ ..... . . . . . . 254
12.3.2 Is There more than One Oscillator Within Plants? ......... 256
12.3.3 Regulation of the Circadian Oscillator .......... ...... 257
12.4 Outputs of the Circadian System ........... ............. .258
12.5 Concluding Remarks .... . . . . . . 259
R eferences ................... . .... . ..... .. 259
Part 4
Theoretical Aspects of Rhythmical Plant Behaviour .......... ..... 265
13 Rhythms, Clocks and Deterministic Chaos in Unicellular
Organisms ............ . .,267
DAVID LLOYD
Abstract ..... . . . . .. ....... . .. ...... 267
3.1 Tim e in Biology . .... . ...... . . ................ ... . . 268
13,2 Circadian Rhythms ............... .. - .... ... .......... . 270
13.2.1 Circadian Timekeeping in Unicellular Organisms ..... .... 270
13.2.2 Cyanobacterial Circadian Rhythms ...................... 270
13,3 Ultradian Rhythms: the 40-Min Clock in Yeast .... .......... 271
13.4 Oscillatory Behaviour During the Cell Division Cycles
of Lower Organisms .............. ..... .... ..... 277
35 Ultradian Gating of the Cell Division Cycle .... . ..... 278
13.51 Experimental Systems ............... . .. 278
115.2 The Model ... .... 279
13.5.3 Computer Simulations . ....... ...... 279
13.6 Chaos in Biochemistry and Physiology._............... ... 282
13 7 Functions of Rhythms .. ..... .... .. . ... 284
13.8 Biological Functio ins of Chaotic Performance ..........286
13.9 Evolution of Rhythmic Performance . ...... ... . 286
References .. . ... . 288
14 Modelling Ca2l Oscillations in Plants .................. ..*..... 295
GERALD SCHONKNECT. AND. CLAUDIA BAUER
Abstract . . . .295
14.1 Introduction ........ . .. . ... . _ 296
14.2 Developing a Mathematical Model ......... ..... 297
14.3; Discussion of the Model .... . 304
References ......... , ,.... 309
Considerations, Modelling and Experimental Evidence ................. 313
15.3, Case Study I1 Stomatal Patterns ...... ...... 32
15A Experimental Observations of Complex Rhythms Plants 327
15,5 A Path Towards Systems Biology ........ ...... 330
.References .335
S Modeling Osclations of Mebrane Potential ifference ,34
MARY JANE BEiLBY
Abstract __,341
16.1 Introduction. .... .. ...... ..... _ _341
---:
16.2 Single Tansporter Oscillations ..... ... 34
162,11 Proton Pump and the Background State i Charophytes.... 342
16.2,2 Putative KV Pump and the Background State 4
in .Ventricaria. ventrosa. .... 346
16.3' Two Transporter Interaction , ,.... ...... __.,346
16.3,1 Proton Pump and the Background State in Hypertorict
Regulation in Lamprotha.mni.on spp. ....... .... 346
16.3.2 Interaction of the Proton Purn.p an.d the.Proto
Channel in Chan EPP .. ....... ... 348
16.4 Multiple Transporter Interaction .... 35
<16.4.1 Hypotonic Regulation in Salt-ToJ.erant Charophytes. 350
16&4C2 Repetitive Action .Potentials . Salt-Senqitive
Charophytes it High Salinity ... .. .. 352
.16.5 Conclusions . ...... ... 354
References ...*. . . ........ . . ... .. .. . . 354
Subjecml-ifftlA A pagaHpA ka»nR«»aA« pi [ fBaBa lpll l- Bf -e/ t Inde . .. . .. . .. . . _ _ _ _ 35