Root Ecology,9783540001850

Root Ecology

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ISBN13: 9783540001850
ISBN10: 3540001859
Format: Hardcover
Pub. Date: 2003-08-01
Publisher(s): Springer Verlag
List Price: $349.99

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Summary

In the course of evolution, a great variety of root systems have learned to overcome the many physical, biochemical and biological problems brought about by soil. This development has made them a fascinating object of scientific study. This volume gives an overview of how roots have adapted to the soil environment and which roles they play in the soil ecosystem. The text describes the form and function of roots, their temporal and spatial distribution, and their turnover rate in various ecosystems. Subsequently, a physiological background is provided for basic functions, such as carbon acquisition, water and solute movement, and for their responses to three major abiotic stresses, i.e. hard soil structure, drought and flooding. The volume concludes with the interactions of roots with other organisms of the complex soil ecosystem, including symbiosis, competition, and the function of roots as a food source.

Table of Contents

1 Constraints on the Form and Function of Root Systems 1(32)
D. ROBINSON, A. HODGE and A. FITTER
1.1 Introduction
1(1)
1.2 Problems Associated with Life in Soil
2(2)
1.2.1 Physical Problems
2(1)
1.2.2 Chemical Reactivity
3(1)
1.2.3 Biological Activity
3(1)
1.2.4 Heterogeneity
4(1)
1.3 Evolutionary Solutions
4(7)
1.3.1 Penetration of Soil Pores
5(1)
1.3.2 Heterotrophy
5(1)
1.3.3 Hierarchical Branching
5(3)
1.3.4 Long-Distance Transport
8(1)
1.3.5 Maintenance Costs
8(1)
1.3.6 Dehydration Risk
9(1)
1.3.7 Compensation for Unpredictable Water and Nutrient Supplies
10(1)
1.3.8 Conflicting Design Requirements
10(1)
1.4 Emergent Properties
11(15)
1.4.1 Topology
11(4)
1.4.2 Size
15(4)
1.4.3 Depth
19(1)
1.4.4 Anchorage
20(1)
1.4.5 Rhizosphere
21(2)
1.4.6 Mycorrhizas
23(1)
1.4.7 Specialised Morphologies
24(1)
1.4.8 Global Scale Processes
25(1)
1.5 Concluding Remarks
26(1)
References
27(6)
2 Distribution of Roots in Soil, and Root Foraging Activity 33(28)
M.J. HUTCHINGS and E.A. JOHN
2.1 Introduction
33(2)
2.2 Plant Rooting Patterns in the Vertical and Horizontal Dimensions
35(5)
2.3 Segregation of Root Systems
40(4)
2.3.1 Segregation of Root Systems in the Vertical Dimension
40(2)
2.3.2 Segregation of Root Systems in the Horizontal Dimension
42(2)
2.4 Foraging by Roots
44(11)
2.4.1 Root Foraging Responses to Spatial Heterogeneity in Availability of Soil-Based Resources
45(4)
2.4.2 Morphological vs. Physiological Plasticity: Responses to Total Resource Supply and to the Spatial and Temporal Patterns of Resource Provision
49(1)
2.4.3 Patterns of Root Placement in Heterogeneous Environments and Their Consequences
50(5)
2.5 Summary and Prospects
55(1)
References
56(5)
3 Turnover of Root Systems 61(30)
W.K. LAUENROTH and R. GILL
3.1 Introduction
61(1)
3.2 Overview of the Structure of Root Systems
62(2)
3.2.1 Conifers and Woody Dicots
63(1)
3.2.2 Herbaceous Dicots
63(1)
3.2.3 Monocots
64(1)
3.3 Methods of Assessing Root Turnover
64(4)
3.3.1 Direct Estimates of Root System Turnover Coefficients Based on 14C Turnover
65(1)
3.3.2 Indirect Estimates of Root System Turnover Coefficients
66(2)
3.3.2.1 Biomass
66(1)
3.3.2.2 Ingrowth Cores
66(1)
3.3.2.3 Nitrogen Balance
66(1)
3.3.2.4 Minirhizotrons
67(1)
3.4 The Growth, Life Span, and Death of Roots
68(4)
3.4.1 Effects at the Individual Root Level
68(2)
3.4.1.1 Water and Nutrients
68(1)
3.4.1.2 Soil Temperature
69(1)
3.4.1.3 Root Diameter
69(1)
3.4.1.4 Root Symbionts
70(1)
3.4.1.5 Herbivory
70(1)
3.4.2 Effects at the Whole-Plant Level
70(2)
3.4.2.1 Elevated CO2
71(1)
3.4 2.2 Pathogens and Herbivores
72(1)
3.5 Field Estimates of Root Turnover and Net Primary Production
72(6)
3.5.1 Forests
73(2)
3.5.1.1 Temperate
73(1)
3.5.1.2 Boreal
74(1)
3.5.1.3 Tropical
75(1)
3.5.2 Grasslands
75(2)
3.5.2.1 Temperate
75(1)
3.5.2.2 High Latitude
76(1)
3.5.2.3 Tropical
76(1)
3.5.3 Shrublands
77(17)
3.5.3.1 Temperate
77(1)
3.5.3.2 High Latitude
77(1)
3.5.3.3 Tropical
78(1)
3.6 Relationship of Root Turnover to Environmental Factors
78(4)
3.7 Summary and Prospects
82(1)
References
83(8)
4 The Control of Carbon Acquisition by and Growth of Roots 91(34)
J.F. FARRAR and D.L. JONES
4.1 Introduction
91(1)
4.2 Production of Carbohydrate in Source Leaves
92(1)
4.3 Import of Carbohydrates by Roots: the Phloem Path
93(1)
4.4 Import of Carbohydrates by Roots: Phloem Unloading and Short-Distance Transport
94(4)
4.4.1 Fibrous Roots
95(1)
4.4.2 Storage Roots
96(1)
4.4.3 Is There Feedback Control of Import?
96(1)
4.4.4 Are There Plant Growth Substances That Control Import?
97(1)
4.4.5 Are There Genes That Control Import?
98(1)
4.5 Carbon Fluxes Within Roots and Their Role in Growth an Import
98(5)
4.5.1 Fluxes That Increase C Content
99(1)
4.5.2 Fluxes That Cause Loss of C
99(1)
4.5.3 Turnover and Metabolism Within Roots
100(3)
4.5.3.1 Localisation and Compartmentation
100(1)
4.5.3.2 Size of Pools Relative to Fluxes
101(1)
4.5.3.3 Flux to Structure (Including Maintenance)
101(1)
4.5.3.4 Localisation of Metabolism to Different Cell Types
102(1)
4.6 Exudation
103(7)
4.6.1 How Large Is the Root Exudation C Flux?
103(1)
4.6.2 What Are the Dominant Exudate Components?
104(2)
4.6.3 Localisation of Root Exudation
106(1)
4.6.4 Mechanistic Basis of Root Exudation
106(3)
4.6.4.1 Root Exudation Regulated by C Influx
107(1)
4.6.4.2 Root Exudation Regulated by C Efflux
108(1)
4.6.5 Exudation: Conclusion
109(1)
4.7 Integration of Fluxes
110(5)
4.7.1 Shared Control of Carbon Flux
110(1)
4.7.2 Additional Evidence for Shared Control of Import into Roots
111(1)
4.7.3 Mechanisms Underlying Shared Control of Carbon Flux
112(1)
4.7.4 What is Root 'Demand'?
113(1)
4.7.5 The Remarkable Consequences of Darkening
114(1)
4.8 Allocation of C and Dry Weights to Roots Relative to Shoots
115(3)
4.8.1 The Conservation of Shoot/Root Ratio
115(1)
4.8.2 The Case of Phosphate
116(1)
4.8.3 Functional Equilibrium
117(1)
4.9 Summary and Prospects
118(1)
References
119(6)
5 Hydraulic Properties of Roots 125(26)
M.T. TYRES
5.1 Introduction
126(1)
5.2 Root Structure and Possible Pathways of Water Movement
127(3)
5.3 Driving Forces and the 'Composite Membrane'
130(1)
5.4 Methods of Measuring Hydraulic Conductances
131(6)
5.4.1 Root Chamber Methods
132(1)
5.4.2 Nobel Method
132(2)
5.4.3 Root Pressure Probe Method
134(2)
5.4.4 The High Pressure Flowmeter Method
136(1)
5.5 Distribution of Hydraulic Resistances in Roots
137(4)
5.5.1 Axial Water Flow - Poiseuille's Law
137(1)
5.5.2 Radial Water Flow and Role of Endodermis and Exodermis
138(1)
5.5.3 Experiments to Locate Major Barriers to Water and Solute Flow
138(3)
5.6 Models of Solute and Water Flux in Roots (Possible Reinterpretation of Ideas)
141(5)
5.7 The Problem of Scaling for Root or Plant Size
146(2)
5.8 Summary and Prospects
148(1)
References
149(2)
6 Root Growth and Function in Relation to Soil Structure, Composition, and Strength 151(22)
A.G. BENGOUGH
6.1 Introduction
151(1)
6.2 An Introduction to Soil Structure and Some Ways to Quantify It
152(3)
6.3 Root Growth in Bulk Soil
155(8)
6.3.1 Physical Limitations to Root Growth
155(3)
6.3.2 Effects of Soil Strength on Root Growth and Physiology
158(2)
6.3.2.1 Growth of Root Tips in Hard Soil
158(2)
6.3.2.2 Root Branching in Hard Soil
160(1)
6.3.3 Localised Compression of Soil Around Roots
160(2)
6.3.4 Water and Nutrient Uptake
162(1)
6.4 Root Growth in Macropores
163(3)
6.4.1 Root Elongation and Distribution in Macropores
164(1)
6.4.2 Effect of Root Clumping on Water and Nutrient Uptake
165(1)
6.5 Ecological Consequences of Soil Structure and Strength
166(1)
6.6 Summary and Prospects
167(1)
References
168(5)
7 Adaptation of Roots to Drought 173(20)
W.J. DAVIES and M.A. BACON
7.1 Introduction
173(2)
7.1.1 Soil Drying - a Composite Stress
173(2)
7.1.1.1 Changes in Soil Water Status
173(1)
7.1.1.2 The Pathway of Water Movement
174(1)
7.1.1.3 Other Variables
175(1)
7.2 Growth of Roots in Drying Soil
175(7)
7.2.1 Morphological Adaptations to Drying Soil
175(2)
7.2.2 Physiological Adaptation of Roots to Soil Drying
177(1)
7.2.3 The Biochemical Adaptation of Roots to Drought
178(2)
7.2.4 Regulation of the Morphological, Physiological and Biochemical Responses of Roots to Soil Drying
180(2)
7.2.4.1 A Role for Abscisic Acid?
180(2)
7.2.4.2 A Role for Ethylene?
182(1)
7.3 Perception and Signalling of Soil Drying by Roots
182(7)
7.3.1 Roots as Sensors of Soil Water Status
182(6)
7.3.1.1 Abscisic Acid as a Root Signal
183(3)
7.3.1.2 Ethylene as a Root Signal of the Effects of Soil Drying and Soil Compaction
186(1)
7.3.1.3 Adaptive Significance of Chemically Based Signalling of Soil Drying
186(2)
7.3.2 Signals from the Soil
188(1)
7.4 Summary and Prospects
189(1)
References
190(3)
8 Physiology, Biochemistry and Molecular Biology of Plant Root Systems Subjected to Flooding of the Soil 193(22)
M.B. JACKSON and B. RICARD
8.1 Introduction
193(1)
8.2 Inhibition of Root Growth by Partial Oxygen Shortage
193(1)
8.3 Possible Causes of Severe Growth Inhibition and Cell Death in the Absence of Oxygen
194(4)
8.3.1 ATP Supply and Demand
194(3)
8.3.2 Self-Injury from Products of Anaerobic Metabolism
197(1)
8.4 Hypoxic Acclimation to Anoxia
198(5)
8.4.1 Oxygen Sensing and Signal Transduction
198(1)
8.4.2 Regulation of Gene Expression
198(1)
8.4.3 Selective Gene Expression and Enzyme Synthesis
199(1)
8.4.4 Metabolic Basis of Improved Tolerance to Anoxia
200(2)
8.4.4.1 Sugar Transport and Degradation
201(1)
8.4.4.2 Glycolytic and Fermentative Enzymes
201(1)
8.4.5 Cytoplasmic Acidosis
202(1)
8.4.6 Other Routes to Tolerance
202(1)
8.5 Aerenchyma and Avoidance of Anoxia
203(1)
8.6 Stem Hypertrophy, Adventitious Rooting and Related Phenomena
204(1)
8.7 Signalling by Oxygen-Deficient Roots
204(2)
8.8 Summary and Prospects
206(1)
References
207(8)
9 Root Competition: Towards a Mechanistic Understanding 215(20)
H. DE KROON, L. MOMMER and A. NISHIWAKI
9.1 Introduction
215(1)
9.2 What Traits Confer Belowground Competitive Ability?
216(1)
9.3 Mechanisms of Root-Root Interactions
217(5)
9.3.1 Indirect Effects Through Resource Depletion
217(2)
9.3.2 Direct Chemical Interactions
219(3)
9.4 Root Distributions as a Consequence of Root-Root Interactions
222(3)
9.5 Belowground Competition as a Consequence of Root Distribution Patterns
225(6)
9.5.1 Symmetric Competition for Space
225(1)
9.5.2 Symmetric or Asymmetric Competition for Nutrients
226(1)
9.5.3 The Dynamics of Competition
227(4)
9.6 Summary and Prospects
231(1)
References
231(4)
10 Root Exudates: an Overview 235(22)
INDERJIT and L.A. WESTON
10.1 Introduction
235(2)
10.2 Examples of Root Exudation
237(1)
10.3 Methods of Measuring Root Exudation
238(3)
10.4 Fate and Movement of Exudates in Soil
241(2)
10.5 Case Study: Root Exudation by Sorghum
243(3)
10.6 Influence on Inorganic Nutrient Availability
246(2)
10.7 Influence on Soil Organisms
248(1)
10.8 Other Roles of Root Exudates
248(2)
10.9 Summary and Prospects
250(1)
References
251(6)
11 Mycorrhizas 257(40)
F.A. SMITH, S.E. SMITH and S. TIMONEN
11.1 Introduction
257(2)
11.2 Classification and Root Structures
259(8)
11.2.1 Arbuscular Mycorrhizas
259(3)
11.2.2 Ectomycorrhizas and Ectendomycorrhizas
262(2)
11.2.3 Mycorrhizas of the Ericales
264(1)
11.2.4 Orchid Mycorrhizas
265(1)
11.2.5 Surprises in Store?
265(2)
11.2.6 Fungus-Plant Interfaces and Interactions
267(1)
11.3 Mycorrhizal Plant Communities and Their Distribution
267(3)
11.4 The Mycorrhizosphere
270(4)
11.4.1 External Hyphae
270(2)
11.4.2 The Soil Environment
272(1)
11.4.3 Bacteria Associated with Mycorrhizal Fungi
272(2)
11.5 Functional Bases of Mycorrhizal Symbioses
274(3)
11.5.1 Transfer of Nutrients and Carbon
274(2)
11.5.1.1 Individual Plants
274(1)
11.5.1.2 Linked Plants
275(1)
11.5.2 Non-nutritional Factors
276(1)
11.6 Diversity in Plant Growth Responses
277(5)
11.6.1 Carbon Costs of Mycorrhizal Symbioses
278(1)
11.6.2 Growth Rates, Nutrient Demand and Mycorrhizal Responsiveness
279(3)
11.7 Plant-Fungal Interactions at the Community Level
282(3)
11.7.1 Plant Density, Competition and Succession
282(3)
11.7.2 The Mycorrhizal Fungal Community
285(1)
11.8 Summary and Prospects
285(2)
References
287(10)
12 Signalling in Rhizobacteria-Plant Interactions 297(34)
L.C. VAN LOON and P.A.H.M. BAKKER
12.1 Introduction
297(1)
12.2 Plant Growth Promotion by Rhizobacteria
298(5)
12.3 Rhizobium-Plant Interactions
303(5)
12.4 Disease Suppression by Rhizobacteria
308(6)
12.4.1 Competition for Substrate
309(1)
12.4.2 Competition for Iron by Siderophores
309(2)
12.4.3 Antibiosis
311(3)
12.4.4 Lytic Activity
314(1)
12.5 Rhizobacteria-Mediated Induced Systemic Resistance
314(6)
12.6 Summary and Prospects
320(1)
References
321(10)
13 Interactions Between Oxygen-Releasing Roots and Microbial Processes in Flooded Soils and Sediments 331(32)
P.L.E. BODELIER
13.1 Introduction
331(3)
13.2 Methodology in Rhizosphere Microbiology
334(1)
13.3 Quantitative and Qualitative Aspects of Root Oxygen Release
335(4)
13.4 Interactions Between Oxygen-Releasing Roots and Aerobic Microbial Processes involved in C- and N-Cycling
339(7)
13.4.1 Heterotrophic Bacteria
339(2)
13.4.2 Methane-Consuming Bacteria
341(3)
13.4.3 Nitrifying Bacteria
344(2)
13.5 Interactions Between Oxygen-Releasing Roots and Anaerobic Microbial Processes Involved in C- and N-Cycling
346(7)
13.5.1 Denitrifying Bacteria
346(2)
13.5.2 Iron- and Sulphate-Reducing Bacteria
348(2)
13.5.3 Methanogenic Bacteria
350(2)
13.5.4 Nitrogen-Fixing Bacteria
352(1)
13.6 Summary and Prospects
353(2)
References
355(8)
14 Root-Animal Interactions 363(24)
J.B. WHITTAKER
14.1 Introduction
363(1)
14.2 The Organisms Involved
364(3)
14.3 Indirect Effects of Aboveground Grazing on Roots
367(3)
14.4 Direct Herbivory on Roots
370(4)
14.5 Interactions Between Above- and Belowground Herbivory
374(2)
14.6 Physiological Responses
376(2)
14.7 Community Responses
378(2)
14.8 Summary and Prospects
380(1)
References
381(6)
Subject Index 387

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