Mycorrhizal Mediation of Soil : Fertility, Structure, and Carbon Storage

Mycorrhizal Mediation of Soil: Fertility, Structure, and Carbon Storage offers a better understanding of mycorrhizal mediation that will help inform earth system models and subsequently improve the accuracy of global carbon model predictions. Mycorrhizas transport tremendous quantities of plant-derived carbon below ground and are increasingly recognized for their importance in the creation, structure, and function of soils. Different global carbon models vary widely in their predictions of the dynamics of the terrestrial carbon pool, ranging from a large sink to a large source. This edited book presents a unique synthesis of the influence of environmental change on mycorrhizas across a wide range of ecosystems, as well as a clear examination of new discoveries and challenges for the future, to inform land management practices that preserve or increase below ground carbon storage. Synthesizes the abundance of research on the influence of environmental change on mycorrhizas across a wide range of ecosystems from a variety of leading international researchers Focuses on the specific role of mycorrhizal fungi in soil processes, with an emphasis on soil development and carbon storage, including coverage of cutting-edge methods and perspectives Includes a chapter in each section on future avenues for further study 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 B......Page 516 E......Page 517 H......Page 518 M......Page 519 N......Page 521 S......Page 522 Z......Page 524 Back Cover......Page 526 __Mycorrhizal Mediation of Soil: Fertility, Structure, and Carbon Storage__This edited book presents a unique synthesis of the influence of environmental change on mycorrhizas across a wide range of ecosystems, as well as a clear examination of new discoveries and challenges for the future, to inform land management practices that preserve or increase below ground carbon storage.