Mycorrhizal mediation of soil: fertility, structure, and carbon storage

Front Cover......Page 1
MYCORRHIZAL MEDIATION OF SOIL......Page 2
MYCORRHIZAL MEDIATION OF SOIL: FERTILITY, STRUCTURE, AND CARBON STORAGE......Page 4
Copyright......Page 5
Contents......Page 6
List of Contributors......Page 12
 COVER PHOTOS......Page 14
 LIST OF SCIENTIFIC REVIEWERS......Page 15
 1.1 SUCCESSFUL COEXISTENCE OF PLANTS AND FUNGI......Page 16
 1.2 MYCORRHIZAL RESEARCH: PAST, PRESENT, AND FUTURE......Page 17
 References......Page 20
I - MYCORRHIZAL MEDIATION OF SOIL DEVELOPMENT......Page 22
  2.1 THE IMPORTANCE OF RECIPROCAL EFFECTS OF PLANT–MYCORRHIZA–SOIL INTERACTIONS IN THE EVOLUTION AND ASSEMBLY OF TERRESTRIAL ECOSYSTEMS......Page 24
  2.2 PLANTS AND MYCORRHIZAS AS AGENTS OF PEDOGENESIS: COUPLING PLANT PHOTOSYNTHATE ENERGY TO THE ACTIONS OF FUNGAL MYCELIAL NETWORKS......Page 26
  2.3 EVOLUTIONARY ORIGINS OF PLANTS AND MYCORRHIZAS......Page 29
  2.4 COEVOLUTION OF PLANTS, MYCORRHIZAS, AND PHOTOSYNTHATE-DRIVEN WEATHERING AND PEDOGENESIS......Page 36
  2.5 FEEDBACK BETWEEN PLANT-DRIVEN PEDOGENESIS, GLOBAL BIOGEOCHEMICAL CYCLES, AND THE EVOLUTION OF PLANTS AND MYCORRHIZAL FUNCTIONING......Page 40
  2.6 CONCLUSIONS......Page 41
  Acknowledgments......Page 42
  References......Page 43
  3.1 INTRODUCTION......Page 50
  3.2 MECHANISMS OF MINERAL WEATHERING......Page 51
  3.3 FUNGAL WEATHERING IN THE LABORATORY......Page 52
  3.4 FROM LABORATORY TO FIELD......Page 54
   3.4.2 Isotope Tracers......Page 55
   3.4.3 Mineral Incubations......Page 56
   3.4.4 Modeling......Page 57
  References......Page 58
  4.1 INTRODUCTION......Page 62
  4.2 MYCORRHIZAL INTERACTIONS WITH CLIMATE......Page 63
   4.2.1 Environmental Predictors of Mycorrhizas......Page 64
   4.2.2 Distribution of Soil Orders and Mycorrhizas Corresponds to Climate......Page 65
   4.2.3 Climatic and Mycorrhizal Mediation of Decomposition......Page 68
   4.3.1 Soil Phosphorus Dynamics......Page 70
   4.3.2 Physicochemical Properties of Soil Parent Material......Page 71
   4.3.3 Mycorrhizas as a Weathering Agent......Page 73
   4.4.1 Topography Influences on Physical Conditions and Processes......Page 74
   4.4.2 Topography Influences Disturbance Regimes......Page 75
  References......Page 76
  5.1 SUCCESSION......Page 82
  5.2 SUCCESSION IN MYCORRHIZAL FUNGAL COMMUNITIES......Page 84
   5.3.1 The Changing Soil Abiotic Environment during Primary Succession......Page 85
   5.3.2 The Changing Soil Abiotic Environment during Secondary Succession......Page 86
     5.3.2.1.1 SOIL NUTRIENTS: NITROGEN AND PHOSPHORUS......Page 87
     5.3.2.1.3 PH......Page 88
   5.4.2 Do Changing Plant Communities Drive Fungal Communities?......Page 89
   5.4.3 Plant Host Specificity As a Driver of Changes in Fungal Communities......Page 90
   5.5.1 Fungal Community Assembly......Page 91
    5.5.1.1 Dispersal Limitations......Page 92
    5.6.1.1 Nitrogen and the Interacting Drivers Hypothesis......Page 93
    5.6.1.2 Interacting Drivers Hypothesis Linking Plant and Fungal Communities......Page 94
    5.6.1.3 Shifts in Fungal Communities Depend on Scale......Page 95
     5.6.1.3.2 SPATIAL SCALE: FROM ROOT TIPS TO CONTINENTAL SCALES......Page 96
   5.6.2 Outstanding Questions and Conclusions......Page 97
   References......Page 98
II - MYCORRHIZAL MEDIATION OF SOIL FERTILITY......Page 106
  6.1 INTRODUCTION......Page 108
   6.2.1 Contributions of Mycorrhizal Fungi to Soil Biological Fertility......Page 110
   6.2.2 Contributions of Mycorrhizal Fungi to Soil Chemical Fertility......Page 111
  6.3 SOIL FERTILITY INFLUENCES MYCORRHIZAL FUNGI......Page 112
   6.3.1 Mycorrhizal Function in Agricultural Ecosystems......Page 113
   6.3.2 Mycorrhizal Function in Forest Ecosystems......Page 114
  6.4 PRINCIPLES FOR MANAGEMENT OF MYCORRHIZAL FUNGI FOR SOIL FERTILITY......Page 115
  References......Page 116
  7.1 INTRODUCTION......Page 122
  7.2 NITROGEN NUTRITION WITHIN ARBUSCULAR MYCORRHIZAS......Page 124
  7.3 PHOSPHATE TRANSPORT IN ARBUSCULAR MYCORRHIZAL SYMBIOSIS......Page 127
  7.4 SULFUR METABOLISM AND ARBUSCULAR MYCORRHIZAL SYMBIOSIS......Page 130
  7.5 FROM ROOT TO SHOOT AND BACK: EVIDENCE FOR A SYSTEMIC SIGNALING AND GENE REGULATION IN MYCORRHIZAL PLANTS......Page 132
  Acknowledgments......Page 136
  References......Page 137
   8.1.1 Nitrogen Availability......Page 144
   8.1.2 Phosphorus Availability......Page 145
  8.3 INORGANIC PHOSPHORUS AND NITROGEN ACQUISITION BY ARBUSCULAR MYCORRHIZAL FUNGI......Page 146
  8.4 INORGANIC PHOSPHORUS AND NITROGEN ACQUISITION BY ECTOMYCORRHIZAL FUNGI......Page 148
  8.5 ARBUSCULAR MYCORRHIZAL FUNGI AND ORGANIC NUTRIENT FORMS......Page 150
  8.6 ECTOMYCORRHIZAL FUNGI AND ORGANIC NUTRIENT FORMS......Page 153
   8.6.1 Evidence From Genome Analysis......Page 154
   8.6.2 Evidence From Field and Microcosm Studies......Page 155
   8.6.3 Ectomycorrhizal Fungi and Organic Phosphorus Sources......Page 156
  8.7 CONCLUSIONS......Page 158
  References......Page 159
  9.1 INTRODUCTION......Page 164
   9.1.1 Mycorrhizas and Net Primary Productivity......Page 166
   9.1.2 Mycorrhizas and Plant–Soil Feedback......Page 167
  9.2 MYCORRHIZAS AND SAPROTROPHS......Page 168
  9.3 MYCORRHIZAS AND HERBIVORES......Page 171
  9.4 MYCORRHIZAS AND FUNGIVORES......Page 172
  9.5 MYCORRHIZAS AND BACTERIVORES......Page 175
  9.6 MYCORRHIZAS AND HIGHER TROPHIC LEVELS......Page 176
  9.7 THE WAY FORWARD......Page 177
  References......Page 178
  10.2 AGRICULTURE IN THE PAST......Page 190
   10.3.2 Cultural Systems......Page 192
   10.3.3 Changes in Soils and Plants......Page 193
  10.4 AGRICULTURE IN THE FUTURE......Page 194
   10.4.1 Inoculants......Page 195
   10.4.2 Bioactive Molecules......Page 196
  References......Page 197
  11.1 INTRODUCTION......Page 202
   11.2.1 Rotation Age: Frequency of Harvesting......Page 203
   11.2.2 Large Openings: Clearcuts......Page 204
   11.2.3 Aggregated Retention......Page 206
   11.2.5 Refuge Plants......Page 207
   11.2.6 Importance of Mycorrhizal Networks......Page 208
   11.2.7 Coarse Woody Debris......Page 209
   11.3.1 Is a Change in Ectomycorrhizal Fungal Community Immediately After Commercial Harvesting Likely to Affect Forest Resilience?......Page 210
   11.3.2 Timing of Planting......Page 211
   11.3.3 Site Preparation and Broadcast Burning......Page 212
   11.4.1 Inoculation: Is Worth it?......Page 213
   11.4.2 Assisted Migration: Seed Sources......Page 214
  11.5 STAND MANAGEMENT......Page 215
   11.5.3 Prescribed Burning......Page 216
  11.6 CONCLUSIONS......Page 217
  References......Page 218
  12.1 INTRODUCTION......Page 228
  12.2 MECHANISMS OF MYCORRHIZAL NUTRITION AND STOICHIOMETRY......Page 229
  12.3 NUTRIENT UPTAKE AND MYCORRHIZAL FUNGI: THE BASICS......Page 230
   12.3.1 Plant Uptake Model......Page 231
   12.3.3 Nutrient Forms and Availability......Page 232
    12.3.3.1 Throughput......Page 233
    12.3.3.2 Mineralization......Page 234
    12.3.3.3 Weathering......Page 235
  12.4 MYCORRHIZAS AND GLOBAL CHANGE......Page 236
   12.4.2 Temperature and Soil Moisture......Page 237
  12.5 MYCORRHIZAS AND NITROGEN DEPOSITION......Page 239
  12.6 WHAT IS NEEDED? A STOICHIOMETRIC CHALLENGE......Page 241
  References......Page 242
III - MYCORRHIZAL MEDIATION OF SOIL STRUCTURE ANDSOIL-PLANT WATER RELATIONS......Page 248
  13.2 SOIL STRUCTURE......Page 250
  13.3 SOIL SALINITY......Page 251
  13.4 SOIL MOISTURE......Page 253
  References......Page 254
  14.1 INTRODUCTION: SOIL AGGREGATION, ITS COMPONENT PROCESSES, AND SIGNIFICANCE OF SOIL STRUCTURE......Page 256
   14.2.1 Arbuscular Mycorrhizal Fungi......Page 257
   14.2.2 Ectomycorrhizal Fungi......Page 259
  14.3 MECHANISMS OF SOIL AGGREGATION......Page 260
   14.3.1 Biophysical Mechanisms......Page 262
   14.3.2 Biochemical Mechanisms......Page 263
   14.3.3 Biological Interaction Mechanisms......Page 265
  14.4 RELATIVE IMPORTANCE OF MYCORRHIZAS......Page 266
   14.4.1 In Relation to Other Biota......Page 267
   14.4.2 Across Different Settings......Page 268
   14.5.2 Relative Importance and Greater Coverage of Ecosystem Types......Page 269
   References......Page 270
  15.1 INTRODUCTION......Page 278
  15.2 ARBUSCULAR MYCORRHIZAL FUNGI AND SALT STRESS......Page 280
  15.3 SALINITY IN COMBINATION WITH DROUGHT AND WARMING......Page 283
  15.4 STUDIES OF SALINITY RESPONSES OF INDIGENOUS ARBUSCULAR MYCORRHIZAL FUNGI......Page 284
  15.6 SIGNALING, MYCORRHIZAL FUNGI, AND SALINITY STRESS......Page 285
  15.7 TRIPARTITE INTERACTIONS AND SALINITY STRESS......Page 286
  15.9 CONCLUSIONS AND FUTURE PERSPECTIVES......Page 288
  References......Page 289
  16.1 INTRODUCTION......Page 294
  16.2 MYCORRHIZAS, PLANTS, AND DROUGHT......Page 295
   16.2.1 Effects of Drought on Mycorrhizas......Page 296
   16.2.2 Mycorrhizas and Host-Plant Drought Tolerance......Page 298
   16.2.3 Conclusions and Suggested Directions for Future Research......Page 300
   16.3.1 Pinyon Pine Mortality and Mycorrhizas......Page 301
   16.3.2 Implications for Other Ecosystems......Page 307
  Acknowledgments......Page 308
  References......Page 309
  17.1 INTRODUCTION......Page 314
  17.2 INFLUENCE OF VEGETATION ON SOIL HYDRAULIC PROPERTIES......Page 315
   17.3.1 Arbuscular Mycorrhizal Fungi......Page 316
   17.3.2 Ectomycorrhizal Fungi......Page 321
  17.4 MYCORRHIZAL FUNGAL ROLE IN HYDRAULIC REDISTRIBUTION AND HYDRAULIC CONNECTIVITY IN THE VADOSE ZONE......Page 322
  17.5 MYCORRHIZAL FUNGAL ROLE IN REDUCING SOIL EROSION......Page 323
  17.6 CONSEQUENCES FOR INDIVIDUAL PLANTS, COMMUNITIES, AND ECOSYSTEMS, AND IMPLICATIONS FOR TERRESTRIAL ECOSYSTEMS RESPONSE TO GLOBAL CHANGE......Page 324
  17.7 KNOWLEDGE GAPS, RESEARCH NEEDS, AND FUTURE RESEARCH DIRECTIONS......Page 325
  References......Page 327
  18.1 INTRODUCTION......Page 334
  18.3 THE ROLE OF MYCORRHIZAS IN WATER UPTAKE......Page 335
  18.4 MYCORRHIZAL NETWORKS AND THEIR ROLE IN HYDRAULIC REDISTRIBUTION AND DROUGHT RESPONSES......Page 338
  18.5 ROOTING DEPTH......Page 342
  18.6 THE ROLE OF DROUGHT IN GLOBAL FOREST DECLINE......Page 343
  18.7 CLIMATE CHANGE PROJECTIONS FOR DROUGHT EFFECTS ON FORESTS AND THE DOMINO EFFECT......Page 344
  18.8 INCORPORATING MYCORRHIZAL NETWORKS IN FOREST MANAGEMENT......Page 345
  18.9 KNOWLEDGE GAPS AND FUTURE RESEARCH DIRECTIONS......Page 346
  18.10 CONCLUSIONS......Page 348
  References......Page 349
IV - MYCORRHIZAL MEDIATION OF ECOSYSTEM CARBON FLUXES AND SOIL CARBON STORAGE......Page 356
  19.1 THE CARBON CYCLE......Page 358
  19.3 POSITION OF MYCORRHIZAL FUNGI WITHIN THE SOIL FOOD WEBS......Page 359
  19.4 MYCORRHIZAL SYMBIOSIS AND THE SOIL C CYCLING......Page 361
  19.5 FUNCTIONAL DIVERSITY IN MYCORRHIZAL SYMBIOSES WITH RESPECT TO C CYCLING......Page 362
   19.5.2 Ectomycorrhiza......Page 363
  19.6 OPEN QUESTIONS, EXPERIMENTAL CHALLENGES......Page 364
  References......Page 366
  20.1 INTRODUCTION......Page 372
  20.2 TWO CONCEPTS OF SAPROTROPHY......Page 374
  20.4 ENZYMATIC EVIDENCE......Page 376
  20.5 CARBON SIGNATURES......Page 378
  20.6 ECTOMYCORRHIZAL FUNGI INVOLVED......Page 379
  20.7 NONENZYMATIC NUTRIENT MINING BY ECTOMYCORRHIZAL FUNGI......Page 380
  20.8 STOICHIOMETRIC CONSIDERATIONS......Page 381
  20.9 MODELING STUDIES......Page 383
  20.10 ARBUSCULAR MYCORRHIZAL FUNGI......Page 384
  20.11 SAPROTROPHIC CAPABILITIES OF ECTOMYCORRHIZAL FUNGI: THE WAY FORWARD......Page 385
  References......Page 386
  21.1 INTRODUCTION......Page 390
   21.2.2 Arbuscular Mycorrhizas......Page 391
   21.2.3 Orchid Mycorrhizas and Life-Cycle–Dependent C Flux......Page 392
   21.2.5 Ectomycorrhizas......Page 394
  21.3 HOW DOES C AVAILABILITY (CO2 AND SHADING) INFLUENCE THE CARBON FLUX BETWEEN PLANT AND MYCORRHIZAL FUNGAL COMMUNITIES?......Page 395
   21.3.1 Abiotic Influences on C Flux Between Plants and Mycorrhizal Fungi......Page 396
   21.3.2 Temporal Dynamics in C Allocation......Page 398
   21.3.3 Community Diversity......Page 399
   21.4.2 Why Does C Flow Between Plants and Fungi Need to Be Regulated?......Page 400
   21.4.3 C-for-Nutrient Exchange: Initial Evidence Through to Current Hypotheses......Page 401
  Acknowledgments......Page 403
  References......Page 404
  22.1 MYCORRHIZAS AND HYPHAE-ASSOCIATED MICROBES......Page 410
  22.2 CARBON ALLOCATION FROM MYCORRHIZAL FUNGI TO THE HYPHAE-ASSOCIATED MICROBES IN THE HYPHOSPHERE......Page 412
   22.3.1 Mycorrhiza Helper Bacteria......Page 415
    22.3.3.1 Phosphorus......Page 416
   22.3.4 Induced Protection of Plants Against Pathogens......Page 418
  22.4 DYNAMICS OF THE MYCORRHIZOSPHERE ASSOCIATIONS UNDER FLUCTUATING ENVIRONMENTAL CONDITIONS......Page 419
  22.5 UNRESOLVED QUESTIONS ON TRADING CARBON AND NUTRIENT BETWEEN MYCORRHIZAS AND HYPHAE-ASSOCIATED MICROBES......Page 421
  References......Page 422
  23.1 INTRODUCTION......Page 428
   23.2.2 Ectomycorrhizal Fungi......Page 430
   23.2.3 Ericoid Mycorrhizal Fungi......Page 433
   23.2.4 Biomass Turnover......Page 434
   23.2.5 Biomass Recycling and Decomposition by Mycorrhizal Fungi......Page 436
  23.3 SECRETIONS OF MYCORRHIZAL MYCELIA......Page 437
   23.4.1 Mycorrhizal Fungal Necromass Decomposition......Page 440
   23.4.2 Effects of Mycorrhizal Tissue Quality......Page 441
  23.5 INCORPORATION INTO STABLE CARBON......Page 442
   23.5.1 Sequestration of Carbon Into Secondary Minerals......Page 443
   23.5.2 Sequestration of Carbon Into Soil Organic Matter......Page 444
  23.6 CONCLUSIONS......Page 447
  References......Page 449
   24.1.1 How Different Are Roots and Mycorrhizal Fungi in Relation to Saprotrophs?......Page 456
   24.1.2 Mycorrhizal Effects on Soil Organic Matter......Page 458
  24.2 MYCORRHIZAL FUNGI AS A SOURCE OF C IN SOIL......Page 459
   24.2.1 Provision of Labile C to Microbial Saprotrophs......Page 460
   24.2.2 Importance in Different Mycorrhizal Fungi and Effect on Soil Organic Carbon......Page 463
  24.3 COMPETITION FOR NUTRIENTS AND HABITAT......Page 464
   24.3.1 Organic Nutrient Uptake......Page 465
  24.4 INTERACTIONS AMONG MYCORRHIZAL FUNGI, SOIL FAUNA, AND SOIL ORGANIC CARBON......Page 467
  24.5 CONCLUSION......Page 468
  References......Page 469
  25.1 INTRODUCTION......Page 476
  25.2 BIOCHAR AND MYCORRHIZAS......Page 478
   25.3.1 Soil pH......Page 481
   25.3.4 Soil Temperature......Page 482
   25.3.6 Organic Inhibitors and Signals......Page 483
   25.4.1 Biochar Decreases Nutrient Stress in Plants......Page 484
   25.4.2 Biochar Reduces Disease Severity in Plants......Page 485
  25.5 CONCLUSIONS......Page 486
  References......Page 487
  26.1 INTRODUCTION......Page 494
  26.2 EXISTING MODEL FRAMEWORKS......Page 496
  26.3 CRITICAL MYCORRHIZAL FUNCTIONS FOR TERRESTRIAL BIOSPHERE MODELS......Page 502
  26.4 MYCORRHIZAL FUNGI AS TRAIT INTEGRATORS......Page 506
  26.5 CHALLENGES MOVING FORWARD......Page 508
  References......Page 509
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Back Cover......Page 526