Dynamic Energy and Mass Budgets in Biological Systems

9,06

MB

444

stron

4871

ID

Faculty of Earthand Life Sciences, Vrije Universiteit Amsterdam

2005

rok

Contents

Preface to the second edition xi

Preface xiii

Book organization xvi

Acknowledgements xviii

1 Energetics and models 1

1.1 Energy and mass fluxes . 1

1.1.1 Hope for generality . 1

1.1.2 Historical setting 3

1.1.3 Energetics 5

1.1.4 Population energetics 6

1.2 The art of modelling 7

1.2.1 Strategies 7

1.2.2 Systems . 11

1.2.3 Physical dimensions . 12

1.2.4 Statistics and support . 14

1.3 Summary 17

2 Basic concepts 19

2.1 Individuals: the basic level of organization 19

2.1.1 Input/ output relationships . 19

2.1.2 State variables . 20

2.2 Body shape: surface area/ volume relationships . 23

2.2.1 Isomorphism . 25

2.2.2 Changing shapes 26

2.3 Body size and composition . 30

2.3.1 Homeostasis . 30

2.3.2 Weights . 31

2.3.3 Masses 33

2.3.4 Biomass composition 34

2.3.5 Energy 35

2.3.6 Storage materials 37

2.4 Concentrations, amounts and fluxes 40

vi Contents

2.4.1 Enzyme kinetics . 41

2.4.2 Synthesizing Units . 43

2.4.3 Production of generalized compounds . 48

2.4.4 Handshaking protocols . 48

2.5 Metabolic modes 51

2.6 Temperature . 53

2.7 Life-stages 59

2.8 Summary 63

3 Energy acquisition and use 65

3.1 Feeding . 66

3.1.1 Feeding methods 66

3.1.2 Feeding and movement costs 71

3.1.3 Functional response . 73

3.1.4 Diet . 76

3.1.5 Food deposits and claims 78

3.2 Digestion 79

3.2.1 Smoothing and satiation 79

3.2.2 Gut residence time . 81

3.3 Assimilation . 81

3.4 Reserve dynamics 82

3.5 The -rule for allocation 87

3.6 Maintenance . 89

3.6.1 Volume-related maintenance costs . 90

3.6.2 Surface-area-related maintenance costs 91

3.7 Growth . 94

3.7.1 Embryonic growth . 96

3.7.2 Growth for non-isomorphs . 108

3.8 Development 111

3.9 Propagation . 113

3.9.1 Reproduction 114

3.9.2 Division . 118

3.10 Summary of the basic deb model . 120

4 Uptake and use of essential compounds 125

4.1 Chemical compounds and transformations 125

4.2 Powers 129

4.3 Mass balance 130

4.3.1 Partitioning of mass fluxes . 131

4.3.2 State versus flux 133

4.3.3 Mass investment in neonates 133

4.3.4 Composition of reserves and structural mass . 134

4.4 Respiration . 135

4.4.1 Respiration Quotient 137

4.4.2 Heat increment of feeding . 138

4.4.3 Aging as a consequence of respiration . 139

4.5 Nitrogen balance 145

4.5.1 Urination Quotient . 146

4.5.2 Ammonia excretion . 147

4.6 Products . 147

4.7 Fermentation 148

4.8 Water balance 151

4.8.1 Doubly labelled water . 152

4.8.2 Plant–water relationships . 152

4.9 Energy balance . 153

4.9.1 Dissipating heat . 153

4.9.2 Indirect calorimetry . 155

4.9.3 Thermodynamic constraints 155

4.10 Summary 156

5 Multivariate DEB models 159

5.1 Several substrates 160

5.1.1 Substitutable substrates 160

Sequential processing 160

Parallel processing . 162

5.1.2 Supplementary substrates . 164

5.1.3 Photosynthesis sensu lato . 164

Pigment systems 165

Carbon fixation . 166

Photorespiration 166

Nitrogen incorporation . 167

5.1.4 Calcification . 167

5.2 Several reserves . 168

5.2.1 Growth . 168

5.2.2 Reserve dynamics 170

5.2.3 Simultaneous nutrient limitation 171

5.2.4 Non-limiting reserves can dam up . 172

5.2.5 Oxygen flux . 174

5.2.6 Ammonia–nitrate interactions . 175

5.3 Several structural masses 177

5.3.1 Organ size and function 179

5.3.2 Roots and shoots 179

5.4 Summary 184

6 Uptake and effects of non-essential compounds 187

6.1 One-compartment kinetics . 189

6.2 Partition coefficient . 191

6.2.1 Kinetics as a function of partition . 191

6.2.2 Kinetics as a function of ionization 193

6.3 Energetics affects toxicokinetics 195

6.3.1 Dilution by growth . 195

6.3.2 Changes in lipid content 196

6.3.3 Bioconcentration coefficient 199

6.3.4 Metabolic transformations . 201

6.4 Toxicants affect energetics . 202

6.4.1 No effects 204

6.4.2 Effects on survival . 205

6.4.3 Effects on growth and reproduction 209

6.4.4 Receptor-mediated effects . 213

6.4.5 Mutagenic effects 214

6.4.6 Effects of mixtures . 217

6.4.7 Population consequences of effects . 217

6.5 Summary 219

7 Case studies 221

7.1 Changing feeding conditions 221

7.1.1 Scatter structure of weight data 221

7.1.2 Step up/down in food availability . 223

7.1.3 Mild starvation . 223

7.1.4 Food intake reconstruction . 223

7.1.5 Prolonged starvation 227

7.1.6 Shrinking 230

7.1.7 Dormancy 231

7.1.8 Emergency reproduction 232

7.1.9 Geographical size variations 232

7.2 Diffusion limitation . 235

7.2.1 Homogeneous mantle 235

7.2.2 Mantle with barrier . 238