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Biofertilizers
Scrivener Publishing
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Beverly, MA 01915-6106
Publishers at Scrivener
Martin Scrivener (martin@scrivenerpublishing.com)
Phillip Carmical (pcarmical@scrivenerpublishing.com)
Biofertilizers
Study and Impact
Edited by
Inamuddin,
Mohd Imran Ahamed,
Rajender Boddula,
and
Mashallah Rezakazemi
This edition first published 2021 by John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA
and Scrivener Publishing LLC, 100 Cummings Center, Suite 541J, Beverly, MA 01915, USA
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Library of Congress Cataloging-in-Publication Data
ISBN 9781119724674
Cover image: Wikimedia Commons

Cover design by Russell Richardson
Set in size of 11pt and Minion Pro by Manila Typesetting Company, Makati, Philippines
Printed in the USA
10 9 8 7 6 5 4 3 2 1
Contents
Preface xxi
1 Biofertilizer Utilization in Forestry 1
Wendy Ying Ying Liu and Ranjetta Poobathy
1.1 Introduction 2
1.2 Mechanisms of Actions of Biofertilizers 3
1.2.1 Facilitation of N Acquisition 3
1.2.1.1 Mutualistic N2 Fixation 4
1.2.1.2 Non-Symbiotic N2 Fixation 5
1.2.2 Facilitation of P Acquisition 5
1.2.2.1 Phosphate Solubilizing Microorganisms 6
1.2.2.2 Mycorrhizas 7
1.2.3 Potassium Solubilization 8
1.2.4 Production of Siderophores 9
1.2.5 Modulation of Phytohormones 10
1.2.6 Phytoprotection 12
1.3 Factors Influencing the Outcome of Forestry-Related
Biofertilizer Applications 13
1.4 Applications of Biofertilizers in Forestry 16
1.5 Conclusion and Future Prospects 18
References 20
2 Impact of Biofertilizers on Horticultural Crops 39
Clement Kiing Fook Wong and Chui-Yao Teh
2.1 Introduction 40
2.2 Microbial Strains Used in Biofertilizers 41
2.3 Impact of Biofertilizer Application on Horticultural Crops 41
2.3.1 Increased Yield and Quality of Crops 41
2.3.1.1 Vegetable Crops 44
2.3.1.2 Fruit Crops 46
2.3.1.3 Ornamental Plants 48
v
vi Contents
2.3.2 Enhanced Nutritional Content of Produce 49

2.3.2.1 Mineral-Biofortified Crops 49
2.3.2.2 Enhanced Secondary Metabolites 50
2.3.2.3 Improved Vitamin Content 51
2.3.3 Improved Tolerance Against Biotic Stress 52
2.3.3.1 Fungal and Bacterial Pathogens 52
2.3.3.2 Viral Pathogens 56
2.3.3.3 Insect Pests 58
2.3.3.4 Nematodes 61
2.3.3.5 Weeds 64
2.3.4 Improved Tolerance Against Abiotic Stress 65
2.3.4.1 Drought 66
2.3.4.2 Salinity 68
2.3.4.3 Heavy Metal 70
2.3.4.4 Cold Stress 71
2.3.4.5 Heat Stress 73
2.3.5 Improved Vegetative Propagation Efficiency 73
2.3.5.1 Propagation by Cuttings 73
2.3.5.2 Grafting 74
2.4 Future Perspectives and Challenges Ahead 75
2.5 Conclusion 79
References 79
3 N2 Fixation in Biofertilizers 105
Rekha Sharma, Sapna Nehra and Dinesh Kumar
3.1 Introduction 106
3.2 Biofertilizers 108
3.2.1 Origin 108
3.3 Biofertilizer: Transporter Constituents 108
3.4 Mechanism of Actions of Biofertilizers 109
3.5 Biochemistry of Manufacture of Biofertilizer 109
3.6 Benefits of Biofertilizer Over Biochemical Fertilizers 110
3.7 Variances Among Organic and Biofertilizer 111
3.8 Types of Biofertilizers 111
3.9 Microorganisms Utilized to Make Biofertilizer 111
3.10 Microorganism in Nitrogen Fixation 113
3.10.1 Biofertilizers: Symbiotic N-Fixers 113
3.10.2 Biofertilizers: Free Living N-Fixers 114
3.10.3 Biofertilizers: Associative Symbiotic
N-Fixers 114
Contents vii
3.11 Phosphorus Solubilizing Microbes 115

3.12 Conclusion and Future Prospect 115
Acknowledgments 116
Abbreviations 116
References 117
4 Organic Farming by Biofertilizers 121
Anuradha and Jagvir Singh
4.2.1 Benefits of Biofertilizers 126
4.2.2 Method of Biofertilizer Application 126
4.2.2.1 Seed Treatment 126
4.2.2.2 Seedling Treatment 127
4.2.2.3 Setts and Tuta Treatment 127
4.2.2.4 Soil Treatment 127
4.2.3 Precautions During Application of Biofertilizers 127
4.3 Classification of Biofertilizers 128
4.3.1 Nitrogen Fixer Bacteria 128
4.3.1.1 Commercial Applications 129
4.3.2 Cyanobacteria as Biofertilizers 130
4.3.2.1 Commercial Applications 130
4.3.2.2 Factors Affecting Cyanobacteria
Biofertilizer 131
4.3.3 Mycorrhiza as Biofertilizers 131
4.3.3.1 Ectotrophic Mycorrhiza 132
4.3.3.2 Endotrophic Mycorrhiza 132
4.3.3.3 Changes in Mineral Compounds 133
4.3.3.4 Manure Value and Its Importance 133
4.3.4 Azolla as Biofertilizer 134
4.3.5 Vermicompost 135
4.3.5.1 Method of Vermicompost 135
4.4 Organic Farming 136
4.4.1 Objectives of Organic Farming 136
4.4.2 Benefits of Organic Farming 136
4.4.3 Benefit for Environment 137
4.4.4 Methods of Organic Farming 137
4.4.5 Techniques for Organic Farming 137
4.4.5.1 Crop Diversity 138
4.4.5.2 Soil Management 138
viii Contents
4.4.5.3 Weed Management 138

4.5 Traditional Agriculture vs. Organic and Inorganic
Farming 139
4.5.1 Problems Created by Traditional Farming 139
4.6 Reasons for Doing Organic Farming 140
4.6.1 To Save Soil Health 140
4.6.2 To Preserve Nutrients 141
4.6.3 To Reduce the Cost of Agriculture 141
4.6.4 To Prevent Hazardous Elements in Animal Products 141
4.6.5 To Protect the Environment 141
4.6.6 Natural and Good Taste 142
4.7 Advantage of Organic Farming 142
4.7.1 Good Nutrition 142
4.7.2 Good Health 142
4.7.3 Freedom From Poison 142
4.7.4 Less Money 143
4.7.5 Great Taste 143
4.7.6 Environmental Safety 143
4.8 Disadvantages of Organic Farming 143
4.8.1 Lack of Information 143
4.8.2 Lack of Outline 143
4.8.3 Making More Money in the Beginning 144
4.9 Conclusion 144
Acknowledgement 144
References 144
5 Phosphorus Solubilizing Microorganisms 151
Rafig Gurbanov, Berkay Kalkanci, Hazel Karadag
and Gizem Samgane
5.1 Phosphorus Pollution 152
5.2 Phosphate Solubilization 153
5.3 Microbial Mechanisms of Phosphate Solubilization 155
5.3.1 Organic Phosphate Solubilization 156
5.3.2 Inorganic Phosphate Solubilization 156
5.4 Phosphate-Solubilizing Bacteria 158
5.5 Phosphate-Solubilizing Fungi 160
5.5.1 Phosphate-Solubilizing Fungi as Plant Growth
Promoters 162
5.5.2 The Methods of using Phosphate-Solubilizing
Fungi in Agriculture 164
Contents ix
5.6 Bacteria-Fungi Consortium for Phosphate Solubilization 165

5.7 Conclusions 167
References 167
6 Exophytical and Endophytical Interactions of Plants
and Microbial Activities 183
A. Mbotho, D. Selikane, J.S. Sefadi and M.J. Mochane
6.2 Beneficial Interactions 185
6.2.1 Arbuscular Mycorrhizal Fungi 186
6.2.2 Plant Growth-Promoting Microorganisms 189
6.2.3 Rhizobia 193
6.2.4 Endophytes 194
6.3 Pathogenic (Harmful) Interactions 194
6.3.1 Oomycetes 195
6.3.2 Fungi 198
6.3.3 Bacteria 199
6.3.4 Viruses 200
6.4 Conclusion 203
References 204
7 Biofertilizer Formulations 211
Sana Saif, Zeeshan Abid, Muhammad Faheem Ashiq,
Muhammad Altaf and Raja Shahid Ashraf
List of Abbreviations 212
7.1.1 Evolution of Biofertilizers 212
7.1.2 Biofertilizers: A Sustainable Approach 213
7.2 Biofertilizer Formulations 215
7.2.1 Selection of Strain 215
7.2.1.1 Microbial Strains 215
7.3 Types of Formulations 227
7.3.1 Carrier-Based/Powder Formulations 230
7.3.1.1 Selection of Carrier Material 230
7.3.1.2 Sterilization of Carrier 235
7.3.2 Granular Formulations 236
7.3.3 Liquid Formulations 236
7.3.3.1 Inoculant Preparation 237
7.3.3.2 Common Additives 238
7.3.4 Cell Immobilization 239
x Contents
7.3.4.1 Polymer Entrapped Formulations 239

7.3.4.2 Advantages and Constrains 243
7.3.5 Fluid Bed-Dried Formulation 243
7.3.6 Mycorrhizal Formulations 244
7.4 Stickers 246
7.5 Additives 246
7.6 Packaging 246
7.7 Conclusion 247
References 247
8 Scoping the Use of Transgenic Microorganisms
as Potential Biofertilizers for Sustainable Agriculture
and Environmental Safety 257
Vasavi Rama Karri and Nirmala Nalluri
8.2 Role of Nitrogen in Plant Growth and Development 260
8.2.1 Microorganisms Involved in Nitrogen Fixation 260
8.3 Importance of Phosphorus 261
8.3.1 Microbes Involved in Phosphate Solubilization 262
8.3.2 Reducing the pH of Soil 262
8.3.3 Mineralization 263
8.3.4 Chelation 263
8.3.5 Promotion of Plant Growth by PSMs 263
8.3.6 Approach of Using PSMs as Biofertilizer
and the Future Perspective 264
8.4 Significance of Potassium (K) 265
8.4.1 Microorganisms Involved in Potassium
Hydrolyzation 265
8.4.2 Effect of KSB on Plant Growth and Yield 266
8.4.3 Abilities and Objections of K Solubilizing Bacteria 266
8.5 Biofertilizers Used in Agriculture 267
8.5.1 Mycorrhiza 268
8.5.2 Plant Growth-Promoting Rhizobacteria (PGPR) 268
8.6 Role of Biotechnology in Agricultural Sector 268
8.6.1 Development of Potent Microbial Strains Through
Genetic Engineering Approach to Produce Efficient
Biofertilizers 269
8.6.2 Genetically Altered Transgenic Azotobacter
vinelandii as an Effective Diazotrophs Biofertilizer 270
8.6.3 Phytostimuators and Biofertilizers 271
8.6.4 Azospirillum 272
Contents xi
8.6.5 Generation of Genetically Modified Transgenic

Azospirillum Strains With Enhanced Levels
of Phytoharmone Secretion 274
8.6.6 Development of Rhizobium Strains With Increased
Competitiveness by Genetic Modification 275
8.6.7 Effect of GM Rhizobial strains on Arbuscular
Mycorrhizal (AM) Fungi 278
8.6.8 Release of Genetically Manipulated Rhizobium
for Field Trails 279
8.7 Conclusion 280
Acknowledgements 281
References 281
9 Biofertilizer Utilization in Agricultural Sector 293
Osikemekha Anthony Anani, Charles Oluwaseun Adetunji,
Osayomwanbo Osarenotor and Inamuddin
9.2 Application of Biofertilizer as Bioaugmentation Agent
for Bioremediation of Heavily Polluted Soil 295
9.3 Advantages of Biofertilizer in Comparison
With Synthetic Fertilizer 296
9.4 Specific Examples of a Biofertilizer for Crop
Improvement in Agricultural Sector 298
9.5 Management of Biotic and Abiotic Stress 301
9.6 Combinatory Effect of Biofertilizer With Other
Substance and Their Effect on Crops 303
9.7 Conclusion and Recommendation to Knowledge 305
References 306
10 Azospirillum: A Salient Source for Sustainable Agriculture 309
Rimjim Gogoi, Sukanya Baruah and Jiban Saikia
10.1.1 The Genus Azospirillum 311
10.1.2 Properties of Azospirillum spp. 312
10.1.2.1 Chemotaxis 312
10.1.2.2 Aerotaxis 313
10.1.2.3 Formation of Cysts and Aggregates
or Flocs 313
10.1.2.4 Survivability in Rhizosphere and Bulk Soil 314
10.1.2.5 Competition With Other Soil
Microorganisms 316
xii Contents
10.1.2.6 Association With Plant Roots 316

10.2 Azospirillum and Induction of Stimulatory Effects
for Promoting Plant Growth 318
10.3 Applications in Various Fields 320
10.4 Current Status 324
10.5 Challenges in Large-Scale Commercial Applications
of Azospirillum Inoculants 327
10.6 Programs Employed for Enhanced Applications
of Azospirillum Inoculants 328
References 330
11 Actinomycetes: Implications and Prospects
in Sustainable Agriculture 335
V. Shanthi
11.2 Role in Maintaining Soil Fertility 338
11.2.1 Nitrogen Fixation 338
11.2.2 Phosphate Solubilization 340
11.2.3 Potassium Solubilization 342
11.3 Role in Maintaining Soil Ecology 342
11.4 Role as Biocontrol Agents 345
11.4.1 Production of Antibiotics 346
11.4.2 Production of Siderophores 348
11.4.3 Production of Hydrogen Cyanide 349
11.4.4 Production of Lytic Enzymes 349
11.5 Role as Plant Stress Busters 351
11.5.1 Resistance From Heavy Metal Toxicity 352
11.5.2 Resistance Against Drought/Water Deficit 354
11.5.3 Resistance Toward Salinity 355
11.6 Conclusion 355
11.7 Future Perspectives 356
References 357
12 Influence of Growth Pattern of Cyanobacterial Species
on Biofertilizer Production 371
Jasti Tejaswi, Kaligotla Venkata Subrahmanya Anirudh,
Lalitha Rishika Majeti, Viswanatha Chaitanya Kolluru
and Rajesh K. Srivastava
12.2 Habit and Habitat of Cyanobacteria 373
Contents xiii
12.3 Morphology and Mode of Reproduction 373

12.4 Role of a Fertilizer in Plant Growth 375
12.4.1 Synthetic Fertilizers 376
12.4.2 Organic Fertilizers 377
12.4.3 Biofertilizer 377
12.5 Cyanobacteria as Biofertilizer 379
12.6 Production of Cyanobacteria 381
12.7 Methods for In Vitro Culture of Cyanobacteria 382
12.7.1 Macro- and Microelements 382
12.7.2 Temperature 383
12.7.3 Light and Cell Density 383
12.7.4 Media 383
12.8 Methods for Gene Transfer into Cyanobacteria 384
12.8.1 DNA-Mediated Transformation 385
12.8.2 Electroporation 385
12.8.3 Conjugation 386
12.8.4 Biolistic Method 386
12.10 Abbreviations 387
References 388
13 Biofertilizers Application in Agriculture: A Viable Option
to Chemical Fertilizers 393
Rajesh K. Srivastava
13.2 Chemical Fertilizer 397
13.2.1 Customized Fertilizers 400
13.2.2 Fortified Fertilizer 400
13.3.1 Biocompost 403
13.3.2 Trichocard 404
13.3.3 Trichocard Production 405
13.3.4 Azotobacter 405
13.3.5 Phosphorus 406
13.3.6 Vermicompost 406
13.4 Conclusion 408
13.5 Abbreviations 408
References 408
xiv Contents
14 Quality Control of Biofertilizers 413

Swati Agarwal, Sonu Kumari and Suphiya Khan
14.2 Biofertilizer Requirement and Supply 414
14.3 Process of Biofertilizer Quality Control 416
14.4 Requirement of Quality Control 417
14.5 Standards for Biofertilizers Quality Control 419
14.6 Methods for Quality Testing 421
14.6.1 Microbiological Methods 422
14.6.2 Serological Methods 422
14.6.3 Molecular Methods 423
14.7 Conclusion 423
Acknowledgement 423
References 424
15 Biofertilizers: Characteristic Features and Applications 429
Tanushree Chakraborty and Nasim Akhtar
15.3 Characteristic Features and Applications of Biofertilizers 431
15.3.1 Cyanobacteria Biofertilizer 431
15.3.2 Actinomycetes 435
15.3.3 Rhizobium leguminosarum bv. Trifolii 436
15.3.4 Arbuscular Mycorrhizal Fungi (AMF) 436
15.3.5 Bacillus thuringiensis 437
15.3.6 Microalgae 438
15.4 Phosphate Solubilizing Bacteria (PSB) and Fungus (PSF) 438
15.4.1 Azotobacter 439
15.4.2 Azospirillum 440
15.4.3 Paenibacillus 440
15.4.4 Phyllosphere Associated Methylobacterium 441
15.4.5 MO Plus Biofertilizer 441
15.5 Effect of Biofertilizer on Various Plants
(Experimental Design) 442
15.5.1 Azotobacter spp. (AZT) and Azospirillum
spp. (AZP) on Eucalyptus grandis 442
15.5.2 Bradyrhizobium Strains and Streptomyces
griseoﬂavus on Some Leguminous, Cereal,
and Vegetable Crops 443
15.5.3 Rhizobium and Rhizobacteria on Trifolium repens 444
Contents xv
15.5.4 Arbuscular Mycorrhizal and Phosphate

Solubilizing Fungi on Coffee Plants 445
15.5.5 Glutamicibacter halophytocola KLBMP 5180
on Tomato Seedlings 446
15.6 Screening of Microbes for Biofertilizer 447
15.6.1 Screening for Phosphate Solubilization 447
15.6.2 Screening for Potassium Solubilizing 447
15.6.3 Screening for Nitrogen-Fixing 448
15.6.4 Screening for Zinc Solubilization 448
15.6.5 Screening for Ammonia Production 448
15.6.6 Screening for Hydrogen Cyanide (HCN)
Production 448
15.6.7 Screening for Siderophores 448
15.6.8 Screening for Auxin Production 449
15.6.9 Screening for Gibberellic Acid Production 449
15.6.10 Screening for Production of Chitinase 449
15.7 Limitations of Biofertilizers 449
15.8 Success of Biofertilizer 450
15.9 Debottlenecking 453
15.10 Optimization of Biofertilizer 456
15.10.1 Optimization of Phosphate Solubilization 456
15.11 Concomitant of Biofertilizer 458
15.12 New Approach 458
References 460
16 Fabrication Approaches for Biofertilizers 491
Andrew N. Amenaghawon, Chinedu L. Anyalewechi
and Heri Septya Kusuma
16.3.1 Nitrogen-Fixing Biofertilizers 493
16.3.1.1 Rhizobium 494
16.3.1.2 Azospirillum 494
16.3.1.3 Azotobacter 495
16.3.2 Phosphorus-Solubilizing Biofertilizers 495
16.3.3 Phosphate-Mobilizing Biofertilizer (Mycorrhizae) 496
16.3.4 Potassium Biofertilizer 497
16.3.5 Growth-Promoting Biofertilizers 497
16.3.6 Blue-Green Algae (Cyanobacteria) 498
xvi Contents
16.4 Preparation Approaches for Biofertilizers 499

16.4.1 Inoculant Formulation 499
16.4.2 Carriers for Biofertilizer Preparation 500
16.4.2.1 Sterilized Carriers 500
16.4.3 Carrier Form 501
16.5 Methods of Biofertilizer Formulation 501
16.5.1 Solid-Based Carrier Bioformulation 501
16.5.1.1 Peat Formulations 502
16.5.2 Liquid Inoculants Formulation 503
16.5.3 Polymer-Based Formulation 504
16.5.3.1 Alginate Formulations 504
16.5.4 Fluidized Bed Dried Formulation 504
16.5.5 Particles From Gas Saturated Solutions
(PGSS) Method 505
16.5.6 Bionanoformulations 505
16.6 Application Modes for Biofertilizers 506
16.6.1 Seed Treatment 506
16.6.2 Seedling Root Dipping 506
16.6.3 Soil Application 507
16.7 Factors Affecting the Preparation of Biofertilizers 507
16.8 Beneficial Effects of Biofertilizers 508
16.9 Challenges and Limitations of Biofertilizers 509
16.10 Future Prospects 509
16.11 Conclusion 510
References 511
17 Biofertilizers From Waste 517
Rafaela Basso Sartori, Ihana Aguiar Severo,
Álisson Santos de Oliveira, Paola Lasta, Leila Queiroz Zepka
and Eduardo Jacob-Lopes
17.2 Waste Sources 519
17.3 Technologies for Waste Treatment 521
17.3.1 Conventional Technologies 521
17.3.2 Emerging Technologies 522
17.3.2.1 Nutrients Recovery From Wastes
by Microalgae 523
17.3.2.2 Overall Process Operations 526
17.4 Main Applications of Microalgae Biofertilizers 528
17.4.1 Fertility and Soil Quality 528
17.4.1.1 Nitrogen Fixation 528
Contents xvii
17.4.1.2Carbon Sequestration 529

17.4.1.3Soil Organic Matter, Improvement,
and Recovery 530
17.4.2 Promotion of Plant Growth, Disease,
and Pest Control 531
17.4.2.1 Plant Colonization and Hormone
Production 531
17.4.2.2 Disease and Pest Control 532
17.5 Conclusion and Recommendations 532
References 533
18 Biofertilizers Industry Profiles in Market 541
Kashish Gupta
18.1 Biofertilizers and Biofertilizer Technology 541
18.1.1 Benefits of Different Biofertilizers 542
18.2 Limitations in Usage of Biofertilizers 543
18.3 Biofertilizer Market Segments 544
18.4 Biofertilizers Market Drivers in India 546
18.5 Present Scenario of Biofertilizer Market 547
18.6 Key Players of Biofertilizers in Indian Market 549
18.7 Problems in Promotion of Biofertilizer 550
18.8 Popular Marketed Biofertilizers in Indian Market 553
18.9 Recent Trends in Biofertilizer: Liquid Biofertilizer 554
18.9.1 Specialties of Liquid Biofertilizer 554
18.10 Conclusion and Future Scope 555
References 556
19 Case Study on Biofertilizer Utilization in African Continents 561
Osikemekha Anthony Anani and Charles Oluwaseun Adetunji
19.2 Specific Examples of Biofertilizer for Crop Improvement,
Environmental Bioremediation, and Their Advantages
and Challenges in Africa 563
19.3 Conclusion and Future Recommendations 570
References 570
20 Biofertilizers: Prospects and Challenges for Future 575
Tanushree Chakraborty and Nasim Akhtar
20.2 Definition 579
20.2.1 Helper Bacteria 579
20.2.2 The Point of Difference 580
xviii Contents
20.3 Advances in Biofertilizer 580

20.4 Preparation of Biofertilizer 581
20.5 The Carrier Materials 581
20.6 Production System of Biofertilizer 582
20.7 Mechanism of Growth-Promoting Activity
of Biofertilizers 583
20.8 Advantages and Limitations 584
20.9 Future Aspects 584
20.10 Conclusion 585
References 586
21 Biofertilizers: Past, Present, and Future 591
Mukta Sharma and Manoj Sharma
21.2 Biofertilizer: A Brief History 593
21.3 Biofertilizer Classification 594
21.4 Different Paradigms of Biofertilizers 596
21.4.1 Impregnation of Fertilizers and Fertilizer Use
Efficiency 596
21.4.2 Inoculants of Mixtures of Microorganisms 597
21.4.3 Different Formulations of Inoculants 597
21.4.4 Inoculant Carrier 598
21.4.5 Biofertilizer Carriers and Liquid Formulations 599
21.4.6 Controlled Release Techniques: Encapsulation,
Lyophilization, and Drying 600
21.5 Biofertilizers: Current Status 601
21.6 Biofertilizers: Future Paradigm 601
21.7 Conclusion 602
References 603
22 Algal Biofertilizer 607
Muhammad Mudassir Iqbal, Gulzar Muhammad,
Muhammad Shahbaz Aslam, Muhammad Ajaz Hussain,
Zahid Shafiq and Haseeba Razzaq
22.2 Algae and Algal Biofertilizers 609
22.2.1 Algae is a Polyphyletic Functional Group 609
22.2.2 Multifaceted Role of Algal Biofertilizer
in Sustainable Cultivation 610
22.2.3 Biostimulants From Algae 612
22.3 Techniques of Application of Algal Biofertilizer 613
Contents xix
22.3.1 Algal Extracts as Biofertilizer 613

22.3.2 Addition of Algal Strains and Algal
Biofertilizer to Soil 619
22.4 Cultivation of Algae and Production of Algal Biofertilizer 625
22.5 Conclusion 630
References 630
Index 637
Preface
Great attention has been paid to reduce the use of conventional chemical
fertilizers harming living beings through food chain supplements from the
soil environment. Therefore, it is necessary to develop alternative sustain-
able fertilizers to enhance soil sustainability and agriculture productivity.
Biofertilizers are the substance that contains microorganisms (bacteria,
algae, and fungi) living or latent cells that can enrich the soil quality with
nitrogen, phosphorous, potassium, organic matter, etc. They area cost-
effective, biodegradable, and renewable source of plant nutrients/
supplements to improve the soil-health properties. Biofertilizers emerge
as an attractive alternative to chemical fertilizers and as a promising cost-
effective technology for eco-friendly agriculture and a sustainable environ-
ment that holds microorganisms which enhance the soil nutrients’ solu-
bility leading a raise in its fertility and stimulate crop growth and healthy
food safety.
This book provides in-depth knowledge about history and fundamen-
tals to advances biofertilizers, including latest reviews, challenges, and
future perspectives. It covers fabrication approaches and various types of
biofertilizers and their applications in agriculture, environment, forestry,
and industrial sectors. Also, organic farming, quality control, quality assur-
ance, food safety, and case studies of biofertilizers are briefly discussed.
Biofertilizers’ physical properties, affecting factors, impact, and industry
profiles in the market are well addressed. This book is an essential guide for
farmers, agrochemists, environmental engineers, scientists, students, and
faculty who would like to understand the science behind the sustainable
fertilizers, soil chemistry, and agroecology, etc.
Chapter 1 focuses on the various action mechanisms observed in micro-
organisms, those that drive effective biofertilizer functions for forestry-
related utilization. Besides, the chapter discusses the factors influencing
the success of forestry-related biofertilizer applications as well as the cur-
rent use and prospects of biofertilizers in the forestry sector.
xxi
xxii Preface
Chapter 2 highlights the impact of applying biofertilizers on horticul-

tural crops including the possible mechanisms, leading to improved crop
growth and stress tolerance. Possible challenges of biofertilizer application
and recommended solutions to these problems are also discussed to ensure
the efficient use of biofertilizers in the horticulture industry.
Chapter 3 discusses various microorganisms which as act as biofertiliz-
ers and also the nitrogen-fixing bacteria including different symbiotic and
asymbiotic nitrogen-fixing microbes and other substitutes for easy making
of biofertilizers. The major focus is given to innovative methods, for exam-
ple, growing of microorganisms, accumulation of microorganisms, and
conveniences for distribution, applying, and framing of microorganisms
for moving from greenhouse and laboratory to field test.
Chapter 4 highlights the usefulness of organic manure in bio-
farming. Various types of agrochemicals have spoiled our life, environ-
ment, and ecosystem. This chapter provides a detailed discussion about
how organic manure can save the life of our earth and how it is better than
agrochemicals.
Chapter 5 reviews the scientific literature on environmental phospho-
rus pollution and mechanisms of phosphate solubilization through intact
bacteria and fungi or their enzymes. Moreover, inoculation methodolo-
gies, factors affecting the inoculum efficiency, and applications of single or
multiple species as promising biofertilizer components for sustainable and
ecological farming practices are summarized.
Chapter 6 reviews plant-microbe associations occurring both exo- and
endophytically on different plant species. The beneficial and pathogenic
outcomes of these interactions are discussed, highlighting the microorgan-
isms and the plants involved. Furthermore, the importance of research of
these interrelationships is considered concerning use in agriculture for the
development of agricultural agents.
Chapter 7 discusses the different formulation technologies of bio-
fertilizers used to mitigate the harmful effects of chemical fertilizers.
The complete formulation process is discussed, highlighting the signifi-
cance of each step, i.e., types of selected microbes, choice of suitable car-
rier, and addition of sticking materials while unifying the biofertilizer
formulation.
Chapter 8 describes the scope of exploiting efficient transgenic micro-
organisms produced by genetic engineering strategy as potential bio-
fertilizers to enhance the yield of crops through the sustainable farming
approach. Furthermore, environment-friendly benefits of utilizing various
types of microorganisms alternative to chemical fertilizers in improving
soil fertility of agricultural lands are also emphasized.
Preface xxiii
Chapter 9 intends to provide detailed information on the application

of biofertilizer as a sustainable biotechnological tool that could lead to an
increase in agricultural production. Detailed information is provided on
the modes of action of these biofertilizers while specific examples of cases
where biofertilizer has been utilized for improving an increase in agricul-
tural production are also discussed.
Chapter 10 discusses the various characteristic properties, utility, and
challenges of free-living nitrogen-fixing bacteria of the genus Azospirillum
as a commercial inoculant, aimed to enhance the nitrogen availability in
the soil for sustainable agriculture. Mechanistic routes aiding in nitrogen
fixation by the bacteria are comprehensively elaborated.
Chapter 11 discusses the beneficial role of actinomycetes in the field of
sustainable agriculture. Its unique ability to mitigate soil ecosystem and
to promote plant growth by producing important agro-active substances
is highlighted. Also, the role of actinomycetes to serve as a potential and
efficient source of biofertilizer is discussed.
Chapter 12 discusses the influence of growth conditions and other
parameters including morphological patterns of cyanobacterial species,
found in biofertilizer quality and nutrients richness.
Chapter 13 provides the positive influences of biofertilizers’ applica-
tion on agricultural sector via improving the productivity and yields of the
crop. Further, it discusses the role of biofertilizers’ production and promo-
tion, with a viable option to chemical fertilizers that have minimized the
productivity of the crop with negative impacts on soil, water bodies, or
environment components.
Chapter 14 provides detail informationabout the necessity of qual-
ity control of biofertilizers. Quality supervision is crucial and should be
achieved constantly to manage the microbial products in support of the
clients. The rules applied for calculating the quality are restricted to the
concentration and viability of the microbes.
Chapter 15 focuses on the significance of biofertilizers with their micro-
bial composition having an edge over chemical fertilizers. The emphasis is
on recent advances in preparation, mechanism, growth promotion, car-
rier materials, and production system of biofertilizers. The chapter also
discusses the prospects of biofertilizers along with its advantages and
limitations.
Chapter 16 provides an in-depth understanding of biofertilizers and
the various approaches available for their preparation. The chapter ends
with some prospects and recommendations needed for further improve-
ments in the development, preparation, and application of biofertilizers to
achieve green, cleaner, and sustainable food production.
xxiv Preface
Chapter 17 discusses the use of biofertilizers based on waste recycling as

a potential substitute for chemical fertilizers. Initially, an overview of bio-
fertilizers is presented. Furthermore, the main sources of organic waste are
discussed, as well as the appropriate treatment processes. Finally, emerging
technologies and the main applications of biofertilizers are presented.
Chapter 18 clearly describes the know-how of biofertilizer technology,
its segments, and a brief description of the types of biofertilizer available in
the market. The highlight of the chapter is the recent trend in biofertilizer
technology: liquid biofertilizer, its merits over conventional fertilizers, and
future potentials.
Chapter 19 discusses the current situation on the application of biofer-
tilizer in Africa and their mechanism of action. Different types of biofer-
tilizer that have been introduced are also highlighted. Moreover, specific
examples are cited where biofertilizer has led to an increase in agricultural
production.
Chapter 20 focuses on various types of biofertilizers, their properties,
significance, preparation, production, uses, and outcome. It also focuses
on experimental designing on screening and selection of microbes and
their optimization as biofertilizers. The chapter also deals with the suc-
cess of biofertilizers, their limitations, and new approaches to overcome
constraints.
Chapter 21 discusses the use of microorganisms in the form of biolog-
ical fertilizers. It also discusses the characteristics and features of different
formulations in which the biofertilizers are used extensively. The chapter
highlights the development and recent advances in biological fertilizer and
its performance.
Chapter 22 covers the biodiversity of algae and biochemical constitu-
ents of algal biofertilizers with the effects on plant growth and yield. The
state-of-the-art techniques for the mass cultivation of algae are also part of
the discussion. This chapter also focuses on novel strategies for the mass
production of algal biofertilizers.
1
Biofertilizer Utilization in Forestry
Wendy Ying Ying Liu* and Ranjetta Poobathy
School of Biological Sciences, Quest International University, Perak, Malaysia
Abstract
The forest biomes provide crucial ecosystem services which include acting as
carbon sinks, providing habitats for biodiversity, preventing soil erosion, mitigat-
ing climate change, and producing important resources such as timber, fuel, and
bioproducts. However, due to various human activities, the productivity of for-
ests has greatly reduced over time. In order to manage nutrient deficiencies and
phytopathogens, chemical products are utilized in forest nurseries, plantation for-
ests, and natural stands. However, this often leads to nutrient losses via leaching,
gaseous losses, and other detrimental effects. Exploitation of biofertilizers in the
bid to reduce reliance on chemicals could promote the growth and development
of tree species and ultimately increase forest productivity in a more sustainable
manner. Most biofertilizer utilizations are focused in the agriculture and horticul-
ture sectors with less emphasis in forestry, with the exception of mycorrhizae. It
is imperative to recognize the various mechanisms of action of biofertilizers (e.g.,
facilitation of nutrient uptake, phytohormone modulation, and phytoprotection)
to fully exploit the potential of biofertilizer in promoting the ecosystem services of
forest biomes. Hence, this chapter explores the mechanisms of action of effective
microorganisms in biofertilizers, factors influencing the effectiveness of biofertil-
izer application, and applications of biofertilizers in the forestry sector.
Keywords: Biofertilizer, forestry, plant growth promoting microorganisms,

biological nitrogen fixation, mycorrhizae, phytohormones, biocontrol,
sustainability
*Corresponding author: wendy.liu@qiup.edu.my
Inamuddin, Mohd Imran Ahamed, Rajender Boddula, and Mashallah Rezakazemi (eds.)
Biofertilizers: Study and Impact, (1–38) © 2021 Scrivener Publishing LLC
1
2 Biofertilizers
1.1 Introduction
The forest biomes are globally important as they cover 30% of Earth’s ter-
restrial surface with more than three trillion trees at an estimated size of
four billion hectares [1–3]. The forest ecosystem services include, but are
not limited to, acting as carbon sinks, providing biodiversity, protecting
soil quality, and producing a wide array of resources such as wood, tim-
ber, biomass, and coal [4, 5]. The forest ecosystems are widely distributed
worldwide, in boreal, temperate, and tropical regions [6]. For some forest
ecosystems, their distributions are highly correlated to land use and soil
characteristics, whereby nutrient-poor soils are allocated for forests while
high fertility soils are utilized for agriculture crops and grasslands [6]. The
productivity of forests has been on a decline due to accelerated growth of
human and livestock populations, forest fires and undiscerning exploita-
tion of forest products [7]. These human activities have led to continuous
soil erosion causing the forest soils to experience deficiencies of essential
nutrients [8].
Chemical fertilizer usage in forestry, from forest nurseries to plantations
and natural stands, ranges from zero to very minimal in comparison to
its usage in the agriculture sector [9]. In the field, fertilizers are typically
applied only once, or at maximum, a few times over a rotation that takes 25
to 30 years [10–12]. The application of chemical fertilizer, when required,
allows management of nutrient deficiencies in marginal soils, increased
productivity of planted forests and/or maintenance of sustainable soil
nutrients after successive rotations [10, 11]. However, the use of chemical
fertilizers has its drawbacks. Firstly, excessive applications in the field could
lead to nutrient losses via leaching, gaseous losses, and reduced beneficial
forest microbiomes [13]. Also, it is arduous to fertilize forest soils due to the
remoteness, low accessibility, and low fertility of many forests [6]. Hence, it
is more viable to exploit the potential of biofertilizers to reduce the reliance
on chemical fertilizers to promote the growth and development of tree spe-
cies and ultimately promote sustainable forest productivity.
Biofertilizers are products containing beneficial microorganisms, mainly
bacteria and fungi, which can inhabit the rhizosphere and/or plant inte-
riors and subsequently promote plant growth via application onto seeds,
plants, and/or soil [14, 15]. Such microorganisms, also known as plant
growth promoting microorganisms (PGPMs), can directly increase plant
growth by facilitating nutrient acquisition in plants and modulating phyto-
hormones while indirectly promote plant growth by decreasing inhibitory
effects of phytopathogens while making the rhizosphere more favorable for
Biofertilizer Utilization in Forestry 3
plants and other beneficial microorganisms [16–18]. The utilization of bio-

fertilizer in agriculture and horticulture is more extensive and widespread
while the utilization of biofertilizers in forestry is still under investigation
or restricted to forest nurseries, with most studies focusing only on mycor-
rhizae [19, 20]. It is imperative to explore other beneficial microorganisms
and their mechanisms of actions to fully utilize the potential of biofertilizer
in promoting growth of tree species, be it in forest nurseries, plantations,
and/or natural stands, in order to fully harness the ecosystem services of
forest biomes.
1.2 Mechanisms of Actions of Biofertilizers

Biofertilizers could improve plant health and growth, soil nutrient, and
fertility status in forest biomes [21]. Nonetheless, the efficacy of the bio-
fertilizers could be greatly affected by various external factors including
soil characteristics, tree species, and native microbiome composition [22].
Hence, it is crucial to understand the mechanisms of actions of PGPMs
[inclusive of the commonly used term, plant growth promoting rhizobac-
teria (PGPR)] utilized in biofertilizer so as to employ them under appropri-
ate circumstances. In reality, many PGPMs are likely to employ more than a
single mechanism, either simultaneously or at different times under differ-
ent conditions [14, 17]. The mechanisms of action of PGPMs comprise of
aiding nutrient uptake [e.g., nitrogen (N), phosphorus (P), potassium (K),
iron (Fe), zinc (Zn), and sulfur (S), modulation of phytohormones (e.g.,
abscisic acid, cytokinin, ethylene, indole acetic acid (IAA), and gibberellin
(GA)] and biocontrol ability to confer phytoprotection [17, 23–25].
1.2.1 Facilitation of N Acquisition

Nitrogen is vital for plant growth and development as it is a major com-
ponent of nucleic acids, membrane lipid, amino acids, proteins, chloro-
phylls, and enzymes [18, 26]. However, N is a key limiting nutrient in forest
biomes, especially in forest ecosystems that are developed on marginal soils
[27–29]. Despite N abundance in the atmosphere as N2, most plants are
incapable of utilizing it and are highly dependent on soil N bioavailability,
in organic and inorganic N forms, to sustain their growth. However, N2 can
be converted into inorganic forms, such as ammonium and nitrate, by the
aid of N2-fixing microorganisms via a highly energy intensive biological N2
fixation (BNF) process catalyzed by nitrogenase [23, 30]. Nitrogen-fixing
4 Biofertilizers
microorganisms in forest biomes can be categorized into two categories:

(i) symbiotic N2 fixers (e.g., rhizobia in association with leguminous plants
and Frankia spp. in association with actinorhizal plants) and (ii) non-
symbiotic N2 fixers (in free-living, associative or endophytic forms) [17,
23]. Nevertheless, symbiotic N2 fixation contributes to majority of the
amount of fixed N required by plants in comparison to non-symbiotic
N2 fixation [17]. Most literature have referred to symbiotic N2 fixation as
symbioses with the development of root nodules while others have also
included associative N2 fixation as symbiotic as it fits the definition of sym-
biosis which is beneficial association between two different organisms [31].
Nonetheless, the former definition will be used in this review.
1.2.1.1 Mutualistic N2 Fixation

Studies have shown that the legume-rhizobia interaction is a main contrib-
utor to the amount of fixed N2 in many forest biomes [32]. Many, albeit not
all, tree legume species are able to fix atmospheric N2 in association with
rhizobia, a group of gram-negative bacteria that are capable of forming N2
nodules on mainly leguminous roots, under N-deficient conditions [33].
Rhizobia can penetrate leguminous root tissues either via hair infection,
crack entry or epidermal entry [34]. Most legumes employ the root hair
infection strategy where flavonoids secreted by the plants will signal for
rhizobia to secrete signal molecules (Nod factors) that binds to root hair
cell receptors and then results in the curling of root hair, bacterial entry into
the cells of the root hair, division of cortical cells, followed by the forma-
tion and ramification of an infection thread in developing nodule primor-
dia [35, 36]. Subsequently, the bacteria will differentiate morphologically
to form bacteroids and are eventually removed from the infection thread to
form symbiosomes via the synthesis of nitrogenase [35–37]. Ammonium
produced in the nodules will be transported to plant cells where it will
be assimilated into amino acids for the plant’s use via the glutamine syn-
thetase (GS)/glutamate synthase (GOGAT) pathway [38]. While rhizobia
provide N source for plant growth, the legumes provide protection and
photosynthates for the rhizobia as a carbon source [39].
Another common N2-fixation symbiosis in forest biomes is the interac-
tion between actinorhizal plants and the actinomycete Frankia via nodule
formation [40, 41]. Many actinorhizal trees and shrubs can form mutual-
isms with mycorrhizae and tripartite symbiosis (actinorhizal plant-Frankia-
mycorrhiza), thus providing them an edge to grow in soils with poor nutrient
availability [42, 43]. The strategy employed by Frankia in infecting their host
plants will depend on the host and Frankia species involved. Nonetheless,
they are similar to those employed by rhizobia, which are hair infection,
crack entry, or epidermal entry. For actinorhizal plants that belong in the
order Fagales, their intracellular infection progress through via root hairs
while for those in the Rosales, the intracellular entry of Frankia is through
the roots [43]. Nodules produced by actinorhizal plants have indeterminate
growth and are able to fix a substantial amount of N2 that is equivalent to
those produced by legumes [43, 44].
1.2.1.2 Non-Symbiotic N2 Fixation

Many free-living heterotrophic diazotrophs in the rhizosphere, such
as Azotobacter and Klebsiella, are also capable of fixing atmospheric N2,
albeit without direct interaction with any other organisms [45–47]. Hence,
these bacteria have to source for their own energy to carry out the highly
energy-intensive BNF process. They can usually oxidize organic molecules
released through decomposition or the aid of other organisms while some
have chemolithotrophic abilities in utilizing inorganic compounds instead
in order to obtain their energy sources [48]. It has been reported that the
free-living BNF rates are strongly correlated to soil organic matter contents,
with higher rates in woody residues and surface organic layers compared
to mineral soil [48, 49]. Also, as oxygen inhibits nitrogenase activities,
these bacteria will have to be able to act as anaerobes or microaerophiles
during the N2 fixation [50, 51]. Thus, with these restrictions, free-living
N2 fixation, in comparison with symbiotic N2 fixation, will not be able to
contribute greatly to BNF in most of the ecosystems [48, 49]. However, it
was also reported that free-living N2 fixers fix substantial amounts of N in
rain forests [52, 53]. In addition, studies have shown that leguminous trees
in mature tropical forests do not fix as much N due to N deposition [54].
Meanwhile, other bacteria, such as Azospirillum and Herbaspirillum,
can form endophytic and/or associate mutualisms with various varieties
of plants [17, 55]. This manner of N2 fixation is similar to symbiotic N2
fixation without the formation of specialized structures whereby these
diazotrophs receive reduced carbon and other nutrients from the plants
while they provide fixed nitrogen for the plants to use [56, 57].
1.2.2 Facilitation of P Acquisition

Despite N being generally recognized as the most limiting nutrient in
many forest ecosystems, P is also another major limiting nutrient [58,
59]. Phosphorus is vital for plants as a major component of energy source
(ATP) to perform various metabolic processes such as signal transduction,
6 Biofertilizers
macromolecule synthesis, respiration, and photosynthesis [30, 60]. It has

been widely reported that high N deposition that is not matched by equiv-
alent increase in the P inputs could cause nutritional imbalances which
eventually reduce forest growth and productivity [61–63]. Demand for
P has been shown to increase as growth promotion takes place following
the additions of N as plant P concentrations and N:P ratios decrease [64,
65]. Also, there is evidence suggesting that P limitation has detrimentally
affected the rate of N2 fixation in tropical forests that mainly consist of
non-leguminous tree species [66, 67].
Phosphorus can be found abundantly in most soil (as organic and inor-
ganic forms) but the amount of accessible P for plants to utilize is low,
as >90% of soil P are insoluble and precipitated (e.g., phosphotriesters
and aluminium phosphate) [30, 68]. Plants are only able to access solu-
ble P sources such as monobasic (H2PO4) and dibasic HPO2− ( 4 )
ions [30,
140]. Although chemical fertilizer provides sufficient soluble inorganic P,
most of them are immobilized quickly after its application, thus render-
ing them unavailable to plants [17]. Hence, a sustainable alternative would
be to employ microorganisms to aid in P solubilization and mineraliza-
tion. These microorganisms can be divided in two major groups, which
are phosphate solubilizing microorganisms (PSMs) and mycorrhizal fungi.
1.2.2.1 Phosphate Solubilizing Microorganisms

PSMs have been shown to effectively increase the accessibility of available
P to plants via solubilization and mineralization of complex P compounds
[69, 70]. There are a few strategies that can be employed by PSMs to sol-
ubilize insoluble P forms, of which the secretion of organic acids (OA)
with low molecular weight, such as acetic, citric, and gluconic acids, is rec-
ognized as a main way P is solubilized [17, 71, 72]. The OA secreted will
either chelate the mineral ions or lower the cell’s pH level in order to acid-
ify these microorganisms and their environments, resulting in the P-ion
being released from P-mineral via the swap of H+ for Ca2+ instead [73].
The efficacy of the solubilization process is highly reliant on on the type,
concentration, and quality of OAs released into the soil, with the quality
of OA being more crucial [74]. The ability of the PSMs to secrete various
OAs simultaneously may enhance the P solubilization process further [71].
Some studies have also recommended that the transformation of insol-
uble P to soluble forms of P could take place without the production of OA
[75, 76]. This was reported by Altomare et al. [77] whereby Trichoderma
harzianum strain T-22 was able to solubilize P, potentially due to chela-
tion and reduction processes, along with providing biocontrol to the plant
without the acidification process. Besides OA production, inorganic acids

secreted by chemoautotrophs (e.g., nitric and sulfuric acids) have been
shown to solubilize P by converting tricalcium phosphate into mono- and
dibasic phosphates [78, 79].
Phosphate-solubilizing microorganisms can mineralize organic phos-
phorus via the secretion of phosphatases and phytases that catalyze the
hydrolysis of phosphoric esters [72]. Acid phosphates, commonly found
in fungi, have been proposed to be the main mechanism in the organic P
mineralization [80]. Meanwhile, phytase releases P from organic materials
that are stored as phytate [72]. Although plants generally are not capable in
acquiring P directly from phytate, PSMs in the rhizosphere could alleviate
the effects due to the plant’s inability to do so [81]. A single PSM strain
could carry out both phosphate solubilization and mineralization [17].
Examples of bacterial genera capable of solubilizing and mineralizing P
include Azotobacter, Bacillus, Bradyrhizobium, Burkholderia, Enterobacter,
Erwinia, Mezorhizobium, Rhizobium, Microbacterium, Paenibacillus,
Pseudomonas, and many others [82]. Meanwhile, fungal PSMs include
those from the genera Aspergillus, Cladosporium, Penicillium, Rhizoctonia,
Rhizopus, Sclerotium, Trichoderma, and others [82, 83].
1.2.2.2 Mycorrhizas
Mycorrhizal fungi can establish symbiotic relationships, either obligately
or facultatively, with many plants, where these mycobionts are reliant on
their host plants for photosynthates and energy while they contribute vari-
ous benefits to their hosts [84]. This group of microorganisms is one of the
most commonly studied PGPMs used in biofertilizer for forestry purposes
[85]. The fungi can increase surface areas for better access to procure nutri-
ents, in particular insoluble phosphorus sources by extending their myce-
lia from root surfaces into the soil [86]. Besides that mycorrhizal fungi are
also able to improve quality and aeration of soil, reduce susceptibility of the
plants to phytopathogens, and promote tolerance of host plants to environ-
mental stresses [84, 87, 88].
Mycorrhiza can colonize their host plant’s root tissues either via intra-
cellular or extracellular mechanisms. Hence, they can be divided into
two main categories, which are endomycorrhiza and ectomycorrhiza,
respectively. Endomycorrhizas can be further categorized as arbuscular,
arbutoid, ericoid, monotropoid, and orchid mycorrhizaes [88, 89]. The
most common mycorrhizal associations are arbuscular mycorrhizal fungi
(AMF) which are obligate mycobionts (generally, Glomeromycetes) that
can form mutualism with more than 80% of vascular plants [90]. Vesicles
8 Biofertilizers
and arbuscules are specialized structures of AMF that assist in mobilizing

and making P available to their host plants and also obtaining mineral and
water sources from further areas in the soil [3, 85]. However, as AMF are
obligate in nature and cannot be grown as pure cultures in isolation from
their hosts, these make AMF biofertilizer production in large scale a very
difficult task [91].
Although ectomycorrhizal fungi (ECMF) are extensively distributed in
many ecosystems, they are only affiliated with only 3% of the vascular plant
families, particularly woody trees (e.g., birch, beech, myrtle, pine, and wil-
low) [85]. Nonetheless, as they are effective in symbiotic association with
many important woody trees, they are crucial in forestry and ecosystem
management. Species of ECMF (>20,000 species) are mainly members of
the phyla Ascomycota and occasionally Basidiomycota [92, 93]. ECMF
are unable to penetrate the cell walls of their host plants but they form an
intercellular interface, the Hartig network, comprising extremely branched
hyphae to form a lattice between epidermal and cortical root cells to allow
metabolic exchange between both fungi and the root [85]. The thick
hyphal sheath, termed as the mantle, formed by ECMF are connected to
extraradical mycelia that extend into the soil to allow minerals and water
to be mobilized, absorbed and translocated to the hosts, often aiding their
survival during adverse conditions [94, 95].
As of late 1950s, ECMF were investigated and utilized as biofertilizer to
promote plant growth as they are able to assist in plant acquisition of P, N,
water, and other minerals in forestry [96]. The application of these fungi
is more common in nurseries whereby tree seedlings are inoculated in the
nursery before planting in field in order to ensure healthy fungi system.
Successful ECMF association is highly dependent on the suitable selection
of plant-host species [85]. Mycorrhizal helper bacteria (MHB), when used
with ECMF, have been shown to assist in the establishment and effective-
ness of the mycorrhizal symbioses [97] by stimulating mycelial growth,
increase the interaction and surface areas for root-mycorrhizae coloniza-
tion via phytohormone and flavonoid production, and mitigate the detri-
mental effect of environmental stress on mycelia [85, 98, 99].
1.2.3 Potassium Solubilization

Potassium is an integral macronutrient for enzyme activation, osmotic
balance, phloem transport, photosynthesis, and protein synthesis in plants
[100, 101]. The concentration of soluble K is generally minimal in soil as
most K occur in the forms of silicate minerals and insoluble rocks [102].
The soil is the main source of K for plants and its accessibility in soil is
reliant on the dynamics and content of K in the biomes [100]. Besides that,
K limitation may detrimentally affect the soil microbial community result-
ing in inefficient nutrient cycling in the forest biomes [103, 104].
It has been suggested that the K dynamics and distribution in plant tis-
sues, soils, and water in the forest biomes could be strongly affected by
biotic factors [105]. The availability of K could also be further affected by
stressors such as timber harvesting, forest fire, and intensification of land
usage [106]. Many forests of which tree species have been continuously
harvested for fuel, fertilizers, and others may have seen a decrease in sub-
stantial amounts of K available in the ecosystem, which may have decel-
erated the K availability for the growth and development of various tree
species [107, 108]. Also, the addition of other major limiting nutrients (N
and P) through atmospheric deposition, without increasing the amount of
K, may be detrimental to the ecology of the forest biomes [105].
Limitations of K in the soil could be addressed by using potassium-
solubilizing microorganisms (KSMs) which produce OAs (e.g., formic,
malic, oxalic, and tartaric acids) to release K from various insoluble min-
erals which include illite, micas, and orthoclases [100]. The OAs can con-
vert K into soluble forms via direct solubilization of rock K or excretion of
chelated silicon. These acids increase the solubilization of K compounds
via provision of protons and formation of complexes with metal ions, such
as Ca2+, present in the soil [109]. Examples of genera of KSMs include
Arthrobacter, Acidithiobacillus, Bacillus, Burkholderia, Paenibacillus, and
others [110, 111].
1.2.4 Production of Siderophores

Iron is an important trace element for plant growth as it acts as a major
component of redox enzymes that aids in oxygen transport, cellular pro-
liferation, chlorophyll synthesis, and nucleic acid synthesis [112]. Despite
being the fourth most copious element on earth, Fe cannot be readily
assimilated by plants and microorganisms as it occurs predominantly as
Fe3+ in nature and tends to form highly insoluble oxides and hydroxides
under aerobic conditions, particularly in calcareous soils [113]. Though
iron is not a limiting factor in many forest biomes, it has reported to be a
limiting factor in some mangrove forests [114–116].
Under Fe-limiting conditions, microorganisms can acquire Fe via the
secretion of siderophores, which are low-molecular mass iron chelators
(~400–1,500 Dalton) with high affinity for Fe3+ ions [17, 117]. Siderophores
will bind to Fe3+ ions and form complexes that are then returned to the
cytosol where the reduction of Fe3+ takes place, followed by the secretion of
10 Biofertilizers
Fe2+ into the cell through a gating machinery, making Fe accessible to the
microorganism [118]. Siderophores are divided into three major families,
which are catecholates (e.g., enterobactin), carboxylates (e.g., rhizobactin),
and hydroxamates (e.g., ferrioxamine B) based on their functional groups
[119]. Thus far, there are 270 types of siderophores that have been structur-
ally characterized out of more than 500 known siderophores [120].
Generally, plants may employ two strategies to sequester iron: (i) low-
ering of pH levels in the rhizosphere, trailed by reduction of Fe3+ ions by
ferric-chelate reductase, allowing the root cells to successively uptake of
Fe2+; and/or (ii) secretion of phytosiderophores for iron solubilization, bind-
ing, and subsequent transport into root cells [121]. Yet, these strategies are
unable to provide sufficient Fe for the plants under Fe-deficient conditions,
especially in calcareous and alkaline soils [121]. Hence, a better alternative
would be to assimilate Fe from microorganisms. Plants can assimilate Fe
from these bacterial or fungal siderophores by two strategies which are
(i) transfer of their Fe-siderophore complexes to plant roots for Fe reduc-
tion to take place and (ii) direct Fe chelation followed by ligand exchange
reaction with phytosiderophores to release Fe2+ [23, 122].
Although the main function of the siderophores is to scavenge for Fe,
they are also able to form complexes with other metals (e.g., aluminium,
cadmium, copper, and nickel) present in the rhizosphere allowing them
to be utilized by the microorganisms [123]. This, in turn, could help to
ameliorate the abiotic stresses caused onto plants by toxic concentrations
of heavy metals in the soil. Besides improving plant growth directly, side
rophore production could also indirectly increase plant growth by inhibit-
ing the growth of phytopathogens by binding to most Fe3+ in the root area
and reducing Fe availability for them to proliferate [124]. This was pro-
posed as an effective biocontrol mechanism as the siderophores produced
by PGPMs have higher affinity for Fe3+ compared to fungal pathogens, thus
out-competing them for available Fe [17].
1.2.5 Modulation of Phytohormones

Phytohormones are organic substances that, at low concentrations (lesser
than 1 mM), have the ability to boost, impede or modify the developmen-
tal activities of plants, particularly in their response to the environment
[30, 125]. When exposed to environmental stresses (e.g., low pH, drought,
salinity, and high temperature), plants usually try to regulate the levels of
endogenous plant hormones so as to reduce the detrimental effects posed
by these stresses [17, 126]. Certain PGPMs are capable of in vitro phyto-
hormone production and could modulate the levels of phytohormones to
reach equilibrium, thus aiding in regulating plants’ responses to environ-

mental stresses [17, 127]. Common phytohormones that are produced by
PGPMs comprise abscisic acid, cytokinin, IAA, and GA [128, 129].
IAA has various functions such as to promote cell division and differ-
entiation, enhance germination rates and percentages of seeds, initiate
and increase the expansion of roots, regulate plants’ responses to stress,
produce metabolites, and negatively affect photosynthesis process [17,
30, 130]. It has been reported that many PGPMs are capable of releasing
IAA as secondary metabolites in a slow and continuous manner, subse-
quently changing the auxin pool and affecting the plants’ physiological
functions [131, 132]. The biosynthesis of IAA by PGPMs could involve
L-tryptophan–dependent or independent pathways. Tryptophan, an amino
acid that occurs at plant root regions, is the main precursor to synthesize
IAA [133]. In addition, tryptophan can indirectly increase IAA production
by inhibiting the formation of anthranilate, a precursor that inhibits IAA
biosynthesis, via negative feedback regulation on the amount of anthra-
nilate synthase [23, 134]. The indole-3-pyruvic acid pathway is the most
frequent IAA biosysnthesis pathway utilized by PGPMs, followed by the
indole-3-acetamide pathway [30, 131].
GAs are diterpenoid phytohormones that are crucial in various pro-
cesses including stem elongation, leaf expansion, seed germination, floral
induction, breaking of dormancy in seeds and tubers, and augmentation in
size and number of fruits [128, 135]. Generally, GAs are produced endoge-
nously by plants to regulate plant growth and development, with GA1, GA3,
and GA4 being the three most common types of GAs that promote plant
growth and shoot elongation [136]. There are three regulatory processes
that control the synthesis of GA which are biosynthesis, reversible conju-
gation, and catabolism [130, 137].
Ethylene can affect plant growth by stimulating root initiation, hindering
root elongation, accelerating the ripening of fruits, reducing wilting, improv-
ing seed germination, and triggering the production of other phytohor-
mones [30]. When exposed to stress conditions, plants will secrete ethylene
at higher concentrations to overcome the stresses [138, 139]. An essen-
tial precursor required for ethylene production is 1-aminocyclopropane-
1-carboxylic acid (ACC) [17, 139]. The production of ACC can be stimu-
lated by the presence of both endogenous and bacterial IAA [17]. However,
high concentrations of ethylene can cause defoliation and alter cellular
processes that will eventually decrease plant growth [23, 30, 138, 140].
The production of ACC deaminase by PGPMs can reduce the negative
impacts of high concentrations of ethylene on plant growth. The benefi-
cial microorganisms, via ACC deaminase, are able to hydrolyze ACC into
12 Biofertilizers
ɑ-ketobutyrate and NH3 which are then utilized as carbon and nitrogen
sources, thus reducing the ethylene levels [139].
1.2.6 Phytoprotection
Despite being used to a much lesser extent in forestry compared to agricul-
ture, pesticides are mostly applied in forest nurseries and planted forests
as they are the most cost-effective manner to manage insect pests, dis-
eases, and weeds [141]. However, continual application of pesticides can
bring about environmental damage to forest biomes (e.g., contamination
of ground water, reduction in soil fertility, and disruption in ecosystem
dynamics) [30]. Hence, the use of PGPMs as biocontrol agents in forests
could be a more sustainable approach to control pests. The biocontrol activ-
ities include antibiosis nutrient competition, hydrogen cyanide (HCN) and
lytic enzyme production, and induced systemic resistance (ISR) [17, 30].
One of the most effective biocontrol mechanisms by PGPMs is antibio-
sis to suppress the proliferation of phytopathogens (generally fungi). Some
of the identified antimicrobial compounds responsible for phytopathogen
inhibition include 2,4-diacetylphloroglucinol (DAPG), ecomycins, pyrrol-
nitrin, pyoluteorin, oomycin A, amphisin, phenazine, and others [142].
However, overdependence on these antimicrobial compounds could lead
to the development of resistance in phytopathogens [30]. Thus, selection
of strains that are capable of producing more than a single antimicrobial
compound as well as synthesizing HCN is preferred to address this [17].
The synthesis of HCN, a volatile secondary metabolite with low molec-
ular weight, can confer selective advantages on the producer, despite the
lack of a role in their growth and primary metabolism. Production of HCN
could inhibit the synthesis of cytochrome-c oxidase and some metalloen-
zymes in the pathogens, consequently preventing electron transport and
disrupting energy supply to the cells [131, 143]. However, the utilization of
HCN on its own does not render significant biocontrol activity, but instead,
it works synergistically with antimicrobial compounds produced in antibi-
osis [17]. It was also reported that HCN could aid in nutrient acquisition
by chelating metal ions and boosting nutrient availability to the plants
[144]. PGPMs could also exert their biocontrol ability by producing lytic
enzymes, such as β-glucanase, chitinase, dehydrogenase, phosphatises,
and proteases that are capable of directly inhibiting fungal pathogens by
affecting the structural integrity of their cell walls [145]. β-glucanase and
chitinase are able to degrade fungal cell walls that are normally made up of
β-1,4-N-acetyl-glucoseamine and chitin. Besides lytic enzyme production,
PGPMs could also suppress phytopathogen growth via competition as they
could colonize surfaces of the plants quickly and utilize many accessible
nutrients, rendering them unavailable to the pathogens for their growth
[17]. Also, as mentioned in the previous section, siderophore production
by PGPMs could restrict the proliferation of these pathogens in the rhizo-
sphere as well due to the lack of iron.
ISR in plants could be triggered by PGPMs with biocontrol ability by
enhancing their ability to defend against specific detrimental environmen-
tal stimuli, triggering their innate defense mechanisms against subsequent
biotic stresses [18, 30]. This resistance involves jasmonate and ethylene sig-
nalling in plants whereby both hormones can positively influence the host
plant’s defense responses to pathogens [17, 23]. Besides these hormones,
many bacterial components have also been shown to cause ISR, including
cyclic lipopeptides, flagella, lipopolysacharrides, homoserine lactones, and
others [146]. Plants with ISR are reported to respond quicker and more
aggressively to pathogen attacks [17]. As ISR does not target pathogens in
a specific manner, it is highly effective in mitigating various pathogenic
diseases [17, 147].
1.3 Factors Influencing the Outcome

of Forestry-Related Biofertilizer Applications
The rhizosphere, extending a few millimetres from the soil surface, is a
substrate that facilitates interactions between bacteria, fungi, and other
microfauna [148–150]. These interactions affect the chemistry and physi-
ology of the surrounding soil, hence presenting major influences on plant
health and nutrition, especially when the total root length and density of
a plant is taken into consideration [148, 151]. Microbes in the rhizosphere
have been observed to interact within and between species as well as with
the host plant, for instance, in the case of co-phosphorus solubilizing bac-
teria (PSB) with vesicular arbuscular mycorrhizae (AM) or N2 fixers, for
instance, Azotobacter and Azospirillum [152, 153]. Rhizospheric fungi are
known to confer benefits on various plant species through a number of
mechanisms, upon which three functional fungal groups are apparent: bio-
fertilizers, biocontrol agents, and bioremediators [148]. Plant growth can
also be boosted via culture with fungi displaying ACC deaminase activity
[154, 155], which results in longer roots as well as reduction of symptoms
following pathogen-induced stress [155, 156].
Generally, the success of any biofertilizer application in forestry depends
on the microbial strain and inoculum production, including formulations,
14 Biofertilizers
and field experimentation strategies, as observed in the application of

Azospirillum [157]. It was found that plants inoculated with PGPR strains
sourced from the plants’ native rhizosphere fare better in terms of germi-
nation and yield [158, 159]. Important considerations include the work-
ability and versatility of the microbial strains at different temperatures: in
the case of a cold climate-targeted PSB, it should be physiologically active
to solubilize P at freezing temperatures. This trait allowed P availability to
the evergreen pine forests during extreme winter conditions [153]. On the
other hand, functionality at ambient temperatures also renders the psy-
chrotolerant PSB species good candidates as P biofertilizer for all condi-
tions [153].
The host plant species itself influences parameters associated with
microbial inoculation: variations in the species, phenology, alterations in
the rhizosphere as mediated by the host, mycorrhizal dependency, as well
as various unknown host plant characteristics are known to influence AMF
colonization as well as spore densities in the host plant [160–162]. The host
species plays the biggest role in this regard, as observed in the rhizosphere
of C. setosum [162]. Carvalho et al. [163] noted that AMF distribution in
salt marshes relies heavily on the host rather than on environment-related
stresses. The host species may also affect the depth of AMF colonization:
spores were located at 40 cm in the subsoil of the rhizospheres of Taxus
chinensis and Pyrus communis [162], attributed to the deep penetration
of roots of the plants. A study on the Pichavaram mangrove in south east
India revealed that areas displaying Rhizophora growth recorded the high-
est total nitrogen content, especially during the monsoon season when the
terrigenous sediments deposit on the mangrove soil [164, 165].
Soil conditions are also integral in the outcome of a biofertilizer-
testing experiment. Soil, as an environment, could present with difficul-
ties due to its unpredictable nature [159, 166]. Research has indicated
that the effectiveness of a PGPR relies on the soil type: plants cultivated
on less fertile soils benefit the most in terms of plant growth from the
application of PGPR [159, 167]. Frommel et al. (1993) concluded that
low soil pH and rainfall as well as high temperatures and Verticillium
soil infection contributed to the poor root colonization by PGPR [159].
The clay soil in the fringes of the Pichavaram mangrove forest is rich
in calcium, magnesium, nitrogen, and phosphorous, besides producing
hydrogen sulfide, partly due to the presence of various species of fungi
and bacteria that confer a host of benefits to the community around
them, including assisting in nitrogen fixation and phosphate availability
[164]. Another factor to consider is the soil moisture content, which may
affect the success of plant growth post-inoculation [159, 169]. This factor,
however, depends on the PGPR type as high moisture content may reduce
soil oxygen levels [159].
The potency of a biofertilizer is also affected by the area in which a par-
ticular microbe is sourced from. The most important groups of PGPRs,
obtained from the ectorhizospheric zone and among endosymbionts,
involve phosphate-solubilizing species such as Acinetobacter, Serratia,
Alcaligenes, Erwinia, Arthrobacter, Aspergillus, Burkholderia, Azospirillum,
Pseudomonas, Bacillus, Enterobacter, Flavobacterium, Penicillium, and
Rhizobium [80, 170–174]. AMF are found in many ecosystems, including
stress-inducing environments such as acid-saline as well as saline soils
[162]. Research has indicated that the AMF distribution depends on both
host physiological and root morphological characteristics [162]. Mangrove
forests have received much attention in terms of the discovery and effects
of biofertilizers on individual tree species [164]. Mangrove areas are known
for harboring populations of nitrogen-fixing Azotobacter [164, 175], of
which three species, i.e., A. vinelandi, A chroococcum, and A. beijerinckii
were identified as potential biofertilizers in mangrove nurseries and prawn
ponds [164, 176]. The high levels of nitrogen resulting from leaf decompo-
sition and the presence of the listed microbes are known to attract amphi-
pods, crabs, fishes, isopods, prawns, and young oysters [177], leading to the
conception of a “mangrove vegetation trap” for seafood harvesting, based
on the requirements above [164, 178, 179]. Cyanobacteria are also promi-
nently featured in mangrove areas. In the case of the Pichavaram mangrove
area in India, aerial N2-fixing cyanobacteria increase in cell number during
the summer and post-monsoon seasons, growing profusely on the lower
and middle portions of the aerial roots [164, 180], and comprise 15% of the
total phytoplankton population [164, 181]. Phormidium species are known
to be tolerant to salinity and present great potential as biofertilizer as well
as shrimp feed formulations [164, 180, 182, 183].
Better results may also be obtained in field experiments involving bio-
fertilizers if combinations of microbes are used rather than single fungal or
bacterial isolates. It was found that phosphate solubilization capacities in
plants can be improved via the usage of a microbial combination instead of
single isolates. Justifications for this phenomenon include improved growth
conditions for other isolates via the metabolic products of one isolate [174,
184]; contribution of P-solubilizing functions by other isolates when one
particular isolate can no longer dissolve phosphate due to changes in
the culture condition [174]; and the microbial diversity involved in the
process [174, 185]. A biofertilizer enriched with N-providing free-living
diazotrophic bacteria and Cunninghamella elegans, a chitosan-producing
fungus, was found to be effective in restricting nematode movement as
16 Biofertilizers
well as reproduction in Pinus pinaster and P. pinea, and mitigating infec-

tion symptoms such as water and total chlorophyll loss, in the former [21].
Phosphate-solubilizing microbes are also known to exert biocontrol poten-
tial [153, 186, 187], a differential influence on AMF within and outside
of plant roots in terms of colonization [153, 188] as well as other growth
parameters for the host plant [153, 189, 190], indicating its potency on the
target plant species as well as the surrounding microbial community.
However, a major hurdle in any study involving biofertilizers is the suc-
cessful transfer of laboratory-based experiments to the field, which has
been shown as not viable in some studies [153, 191], despite the fulfilment
of all the factors listed above. This phenomenon is attributed to the mis-
match in terms of the interactions between the soil biota and the biofer-
tilizer. Hence, it is important to ensure that the ecological characteristics
of all components of a biofertilizer as well as their interactions with the
microbiota are well-characterized [153]. Hayat et al. [153] found that the
PSM isolates, especially Trichoderma paratroviride PWF-1 and Bacillus
megaterium PWB-4, did not display antagonism to one another and other
P-solubilizing fungi, indicating their capability to be established as bacte-
rial or fungal-bacterial combination in order to enhance soil P availability.
The challenge in the fruition of such experiments may also lie in the role of
climate, which is known to influence the effectiveness of an applied PGPR
and hence the outcome of a field experiment greatly [159, 192].
1.4 Applications of Biofertilizers in Forestry

As mentioned in the previous section, efforts to translate laboratory and
greenhouse forestry-related biofertilizer experiments to the field have
been met with minimal success throughout the years. Success is limited
to tree species found in specific locations throughout the globe. Lucy et al.
[159] as well as Mallik and Williams [193] summarized some examples of
research involving the use of PGPR in various forest tree species. A major-
ity of the research were conducted in the growth chamber and greenhouse;
only a minority were transferred to the field [159]. Field testings, how-
ever, indicated increased biomass, shoot and root growth, nutrient uptake,
foliar nitrogen content, pathogenic biocontrol, and seedling emergence in
various pine, beech, oaks, and spruce species inoculated or co-inoculated
with Arthrobacter sp., Pseudomonas sp., Bacillus sp., Azotobacter sp., and
unidentified bacterial strains or AMF [159, 193–219]. In the case of man-
grove forests, known to be lacking in soluble nutrients, especially phospho-
rus [155, 220–222], seedlings were reported to fare better when inoculated
with Azotobacter and Azospirillum [155, 223], phosphobacteria [224–227],

as well as cyanobacteria [155, 228]. Greater success in the experiments was
observed when the rhizospheres of the locations were inoculated with bio-
fertilizers sourced from areas native to both the tree and microbial species
[158, 159].
Climatic conditions, which may differ from laboratory and greenhouse
conditions, influence the success of any biofertilizer experiments. For
instance, extreme cold environments, such as tundra ecosystems, pres-
ent a challenge in terms of microbial interactions, and as such are domi-
nated by psychrophile or psychrotolerant bacteria [153, 229] that tolerate
the cold environment due to abilities such as producing cold-shock pro-
teins, enzymes, genetic changes from thermal shifts, as well as short unsat-
urated membranous acids [153, 230]. Based on this premise, Hayat and
team [153] isolated PSB located in the Himalayan region to quantify for
the isolates’ ecological characteristics as well as their abilities to convert
inorganic P to available P and interact with microbiota present within the
rhizosphere of Pinus spp. The team discovered that 31% and 69% PSB dis-
played ordinary and distinctive P-solubilizing capacities, respectively. Of
the latter, 48% were categorized as “efficient P-solubilisers” (>400 mg L−1).
Out of the 16 efficient PSB isolated, 11 and 4 were located in the lower
subsurface and surface of the Himalayas, respectively. The highest amount
of P-solubilization was observed in Ochrobactrum anthropic, which was
isolated from the pine rhizospheric subsoil. Four out of eight PSB were
found to be resistant to nematode grazing, while another three and one
PSB resisted grazing up to 96 and 120 hours of co-culture, respectively
[153]. The study stressed the importance of isolating microbes native to
the area where the tree species is found in order to increase the fertilizing
potential of the target microorganism.
In order to increase the success of improving plant growth and devel-
opment or soil remediation efforts, microorganisms targeted as biofertiliz-
ers have to be carefully selected based on their characteristics. AMF were
found to be beneficial in terms of plant developmental activities as well as
increasing tolerance to high pH and salinity levels [162, 231–233]. It was
found that AMF inoculation provided more efficient salt stress protection
for the plants when compared to any concentration of plant-available P in
the soil [162, 234]. Salt-tolerant species of AMF that necessitates further
investigation into their roles in plants include Glomus mosseae [162, 235]
and G. caledonium [162] found predominantly in the alkaline and saline
soils of the Yellow River Delta [162]. On the other hand, Trichoderma spe-
cies, a common fungal species, is known to enrich soil via nutrient solubili-
zation, especially in barren or pathogen-infected substrates, which can lead
18 Biofertilizers
to higher plant yields as well as better plant resistance against drought and
pathogenic stress [155, 186, 236, 237]. Trichoderma releases and solubilizes
bound nutrients in the soil [155, 227]. It was reported that an increase of
70% in biomass as well as an increase in the assayed biochemical param-
eters was observed in Rhizophora apiculata and R. mucronata when the
seedlings were inoculated with mangrove-derived Trichoderma. Again, the
results strongly indicated the potential in using biofertilizers derived from
areas native to the target plant [155].
Microbes are sometimes incorporated with certain compounds or sub-
stances to create biostimulants. A biostimulant is considered as such when
a substance is observed to increase or influence a plant’s quality, nutritional
status, or response to stress, regardless of the nutritional situation [238,
239]. Commercial biostimulant products are currently grouped into five
categories: amino acids, fulvic acids, microbial inoculants, humic acids,
and seaweed extracts [238–240]. The use of a biostimulant, reinforced with
A. flavipes cultured in soybean bran and with various amounts of IAA, was
found to be beneficial in the growth of the eucalyptus hybrid E. grandis
x E. urophylla (clone IPB2) when applied in both solid and liquid forms
[239], in which increased root length and mass were observed, especially
in the case of the former. Other Aspergillus species were reported to pres-
ent with similar effects. It was found that A. ustus promoted shoot and
root growth as well as an increase lateral root and root hair numbers in
Solanum tuberosum and Arabidopsis thaliana [239, 241]. Similar effects
were observed in shoots and roots of Cicer arietinum via inoculation with
both Aspergillus niger and Trichoderma harzianum [239, 242]. The natural
compounds present in biofertilizers add value to plant natural resistance
and general nutritional status [21]. Microbial inoculations may stimulate
adventitious root growth, especially in species experiencing difficulties in
rooting, which leads to an increase nutrient and water absorption, host-
plant biomass, as well as stress tolerance levels [239, 243].
1.5 Conclusion and Future Prospects

Biofertilizers can act as viable alternatives to chemicals in enhancing plant
growth and are increasingly indispensable in both the agricultural and
horticultural field. Besides presenting an important role in improving crop
productivity, it is evident, even with the lack of data, that biofertilizers
present great potential in the remediation of forests and the environment.
However, at present, there is a lack of research in the forest biofertilizer
field. No field data have ever been collected for deciduous trees, while little
field data were recorded for coniferous species. As such, future research in
this field may present a gold mine in terms of useful data that can be used
to further our understanding of various biotic interactions that occur in
any forest.
A majority of laboratory and greenhouse experiments have not been
successfully transferred to the field. The rhizosphere, complex in nature, is
a major consideration in any studies involving biofertilizer and has caused
issues in studies of its structure and function, especially when it comes to
addressing the molecular mechanisms that drive all biotic interactions and
communications within the substrate. This process, however, is currently
being facilitated by the use of modern analytical tools such as functional
genomics, direct soil DNA extraction and amplification followed by com-
munity fingerprinting, reporter gene technology, confocal microscopy, as
well as stable isotope profiling. Recent studies are also gearing toward elu-
cidating the influence of such interactions on the above ground commu-
nities as a function of an ecosystem-based feedback process. Labeling of
inoculants with lux or gfp genes can also be conducted for easy detection
and enumeration in the field. Cross-disciplinary collaboration between
sciences involving microbes, soil, plants, ecology, physiology, molecular
biology, chemistry, and physics addresses the complexity of the rhizo-
sphere and the interactions between all its inhabitants.
Vast swathes of areas have not been tested for the presence of potentially
potent biofertilizers. As such, efforts should be made to document microbes
or plant performance in unexplored or underexplored areas globally for
this purpose. Traditional and ethnobotanical knowledge from the indige-
nous communities should be researched for this purpose. Phytochemical
and pharmacological studies of medicinally important forest plant spe-
cies may reveal active compounds that are synthesized as a result of the
interactions between the microbial fauna found in the rhizosphere. New
therapeutic agents can then be developed based on the phytochemistry,
pharmacognosy, and pharmacology of the plant species.
Further optimization is required for parameters related to the applica-
tion of biofertilizers for maximum output, as some research have displayed
incomplete eradication of plant pathogens such as nematodes, or simply a
reduction in inocula sizes, after application of the biofertilizer. However,
some studies have indicated the presence of isolates resistant to pathogens,
i.e., nematode grazing, indicating their potential in supplying important
elements under pathogenic attack and hence protect target plants from
pathogens. Traits such as these can be explored to further our under-
standing in the working mechanisms of a biofertilizer. Positive effects on
plant developmental activities can be observed with accurate matching
20 Biofertilizers
of a biofertilizer with the host plant species to the surrounding environ-

ment, which is advantageous in that a natural or an artificially constructed
microbe can be used for this purpose. Future research in this area can espe-
cially benefit from studies involving synergism and environmental per-
sistence in all components of a biofertilizer in boosting plant development,
especially on the creation of an efficient and effective novel biofertilizer
delivery systems catering to the traits of the plant and microbial species
involved. Outcomes from this research can then be used to supplement in
situ reforesting efforts through the use of biofertilizers.
Future biofertilizer production strategies may also capitalize on year-
round management and application strategies that are large-scaled, cheap,
sustainable, and, independent of climate as well as environmental fac-
tors. Further field studies involving commercial biofertilizers and deliv-
ery mechanisms are required to determine their effectiveness. Sustainable
methods of producing biofertilizers can be initiated through the use of
agro-industrial waste. Ultimately, factors such as effective strains, inocu-
lum production, host compatibility, appropriate formulations, commercial
potential, and technology transfer capacities are integral in the establish-
ment of effective biofertilizers for sustainable forestry.
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least for the moment safe in unconsciousness from the screaming, tearing,
grabbing world.
The next morning, then, when Laura came down punctually at nine
o’clock to breakfast—for however late she went to bed her restless vitality,
once it was broad daylight, prevented her being able to stay there, which
made her unpopular in country houses,—she found Charles in the dining-
room, standing with his back to the fire.
‘How much you must love me,’ she remarked sarcastically, being, after a
bad night, a little cross.
‘I don’t love you at all at this moment,’ said Charles.
‘Then is it breakfast you want?’
‘No,’ said Charles.
‘Can it be Sally?’
‘Yes,’ said Charles.
‘Fancy,’ said Laura; and poured herself out some coffee.
‘How is she?’ asked Charles after a pause, ignoring such silliness.
‘Oh, quite well,’ said Laura. ‘She was tired last night.’
‘Tired! I should think so,’ said Charles severely. ‘I’ve come to ask her if
she will let me take her into the country for the day. It’s my intention to get
her away from your crowd for a few hours.’
‘Rescue her, in fact,’ said Laura, munching, her back to him.
‘Exactly,’ said Charles, who was angry.
‘I expect Tom’—Tom was Lord Streatley—‘will be here soon, wanting
to rescue her too,’ remarked Laura, glancing out of the window to where
she could see Charles’s touring car standing, and no chauffeur. ‘He won’t
bring his chauffeur either. Have some?’ she asked, holding up the coffee-
pot.
‘Can’t you be a little beast when you give your mind to it,’ said Charles.
‘Well, you scolded me last night because I had rescued her, and now here
you are——’
Laura broke off, and hastily drank some coffee. She didn’t really want to
quarrel with Charles; she never had yet. In fact, till Sally appeared on the
scene she had never quarrelled with any of her family. Besides in her heart,
though she was cross that morning, not having slept well for the first time
for years because of being worried and conscience-stricken and anxious,
she was glad that Charles should take Sally off her hands. She had so much
to do that day, so many important engagements; and if Sally went with her
everybody would instantly be upset, and if she left her at home she would
be a prey to Streatley. Other people wishing to prey on her could be kept
out by a simple order to the servants, but not her own brother. And
Streatley, when he was infatuated, was a gross creature, and there would be
more trouble and wretchedness for poor Kitty his wife, let alone God
knowing what mightn’t happen to Sally.
If Sally had to be with one or the other of them, Charles was far the
better; but what a very great pity it was, Laura thought as she pretended to
be absorbed in her breakfast, that she hadn’t let her go back the day before
to where she belonged. It wasn’t any sort of fun quarrelling with her dearest
brother Charles, and seeing him look as if he hadn’t slept a wink. Besides,
Sally was going to have a baby. At least, so she had informed Laura during
the night, basing her conviction on the close resemblance between her
behaviour in fainting, and her subsequent behaviour when she came to in
being violently sick, and the behaviour of somebody called Mrs. Ooper,
who had lived next door at Islington, and every spring, for seven years
running, had fainted just like that and then been sick,—and sure as fate,
Sally had told Laura in a feeble murmur, there at Christmas in each of those
seven years had been another little baby.
‘I don’t want no doctor,’ she had whispered, putting out a cold hand and
catching at Laura’s arm when, dismayed at Sally’s sickness just as they had
at last been able to undress her and get her into bed, she was running to the
telephone to call hers up.
‘But, my darling,’ Laura had said, bending over her and smoothing back
the hair from her damp forehead with quick, anxious movements, ‘he’ll
give you something to make you well again.’
‘No, ’e won’t,’ Sally had whispered, looking up at her with a faint, proud
smile, ‘ ’cos I ain’t ill. I know wot’s ’appenin’ all right. It’s a little baby.’
And then she had told Laura, who had to stoop down close to hear, about
Mrs. Ooper.
Well, Laura didn’t know much about babies before they were born, but
she was sure a person who was expecting any ought to be with her husband.
She couldn’t kidnap whole families; she hadn’t bargained for more than one
Luke. And during the few hours that remained of the night, after she had
seen Sally go off to sleep with an expression of beatitude on her face, she
had tossed about in her own bed in a fever of penitence.
When would she learn not to interfere? When would she learn to hang on
to her impulses, and resist sudden temptation? Up to then she had never
even tried to. And a vision of what Sally’s unfortunate young husband must
be feeling, and of course his mother too, who might be tiresome but hadn’t
deserved this, produced the most painful sensations in Laura’s naturally
benevolent heart.
She would make amends,—oh, she would make amends. She would take
Sally to Cambridge herself on Saturday, when she was through with her
London engagements, and find rooms for her, and explain everything to the
young man, and beg his pardon. Perhaps, too, she could tell him a little of
Sally’s fear of his mother, and perhaps she might be able to persuade him
not to let her live with them; for Laura had often noticed, though each time,
being a member of the Labour Party, with shame and regret, that the
persuasions of the daughter of a duke are readily listened to. But she didn’t
want to make amends that day,—she was too busy; and she couldn’t send a
telegram, or anything like that, letting the Lukes know where Sally was,
because it would only bring them about her ears in hordes, and she simply
hadn’t time that day for hordes. Laura’s intentions, that is, were admirable,
but deferred.
‘Isn’t she coming down?’ asked Charles at last, for Laura, with her back
to him pretending to eat her breakfast, had said no more.
‘She’s having breakfast upstairs,’ said Laura.
‘Why didn’t you tell me?’ he asked, annoyed.
‘Because you say I’m a little beast, so I may as well do the thing
thoroughly.’
Charles went across to the bell.
‘No—don’t ring,’ said Laura jumping up. ‘I’ll go and tell her.’ And she
went to the door, but hesitated, and came back to him, and laid her hand on
his arm.
He withdrew his arm.
‘Charles—are you so angry with me?’ she asked.
‘You’ve behaved simply disgracefully,’ he answered in a voice of deep
disgust. ‘You would sacrifice anybody to provide your friends with a new
sensation.’
Laura looked at him. It was true; or had been true. But she wasn’t going
to ever any more, she was going to turn over a new leaf—next day, when
she had finished with all her tiresome and important engagements.
‘You sacrificed that child’—began Charles, passionately indignant when
he thought of the unconscious figure on the floor.
‘Don’t you sacrifice her,’ interrupted Laura. And when Charles stared at
her, too angry for speech, she added hastily, ‘Oh, don’t let’s quarrel, Charles
darling. I’m sure you’ll take the greatest care of her. I’ll go and fetch her.
Drive slowly, won’t you—and bring her back safe. Tomorrow I’m going to
hand her over to her husband.’
§
Now in his heart Charles knew that this was the only right thing to do.
Sally ought never to have been taken away from her husband, and, having
been taken, ought to be returned to him. At once. Not tomorrow, but at
once. He didn’t know the circumstances, except what Laura had hurriedly
told him the night before after supper, about having found her in a train,
dissolved in tears because her father was sending her back to a mother-in-
law who was awful to her, and she had brought her home with her just to
comfort her, just to let her recover; but it was plain that such conduct on
Laura’s part was indefensible. If ever anybody ought to be safe at home it
was Sally. She should be taken there without losing a moment. Disgraceful
of Laura to put it off for another day and night, while she kept her fool
engagements. Having behaved so wickedly, she ought, without losing an
hour, to set things straight again.
Charles felt strongly about Laura’s conduct; yet, though he himself could
have set things straight by simply driving Sally back to the Lukes that
morning, he didn’t do so. That was because he couldn’t. He was in love,
and therefore couldn’t.
There are some things it is impossible to do when you are in love,
thought Charles, who recognised and admitted his condition, and one is to
hand over the beloved to a brute. Luke was a brute. Clearly he was, from
what Sally had said the night before. He was either angry—angry with that
little angel!—or he oh-Sallied. A cold shudder ran down Charles’s spine.
The thought made him feel really sick, for he was a tender-stomached as
well as a tender-hearted young man, and possessed an imagination which
was sometimes too lively for comfort. It wouldn’t be his hand that delivered
her up to a young brute; nor, he suddenly determined, on the butler’s
hurrying out to Laura, who was standing on the steps seeing him and Sally
off, and saying with urgency, ‘Lord Streatley to speak to Mrs. Luke on the
telephone,’ would it be his hand that delivered her up to an old one. At once
on hearing the message he started the car, and was out of the square before
Laura could say anything. There was Sally, tucked up beside him in Laura’s
furs, and looking more beautiful in broad sunshine even than he
remembered her the night before,—a child of light and grace if ever there
was one, thought Charles, a thing of simple sweetness and obedience and
trust; and was he going to bring her back to another evening’s exploitation
by his sister and her precious friends, with that old scoundrel, his elder
brother, all over her?
Never, said Charles to himself; and headed his car for Crippenham.
§
Crippenham was where his father was. What so safe as a refuge for Sally
as his father? He was ninety-three, and he was deaf. A venerable age; a
convenient failing. Convenient indeed in this case, for the Duke, like
Charles, took little pleasure in the speech of the lower classes. Also he was
alone there till Laura should come back to him on the following day,
because nobody was ever invited to Crippenham, which was his yearly rest-
cure, and nobody ever dared even try to disturb its guarded repose.
Charles felt that it was, besides being the only, the very place. Here Sally
could be kept remote and hidden till Laura—not he; he wouldn’t be able to
do such a thing—restored her to where she belonged; here she would be
safe from the advances of Streatley, who couldn’t follow her anywhere his
father was, because the old man had an aversion to the four surviving fruits
of his first marriage, and freely showed it; and here he would have her to
himself for a whole evening, and part at least of the next day.
Also, it would serve Laura right. She would get a fright, and think all
sorts of things had happened when they didn’t come back. Well, thought
Charles, she deserved everything she got. Under the cloak of protecting and
comforting Sally she had been completely selfish and cruel. Charles was
himself astonished at the violence of his feelings towards Laura, with whom
he had always been such friends. He didn’t investigate these feelings,
however; he didn’t investigate any of his other feelings either, not excepting
the one he had when he asked Sally, soon after they had turned the corner
out of the square, if she were warm enough, and she looked up shyly at him,
and smiled as she politely thanked him, for his feelings since the evening
before no longer bore investigation. They were a mixed lot, a strong lot.
And it vexed Charles to know that even as early in the day as this, and not
much after half past nine in the morning, he wished to kiss Sally.
This wasn’t at all the proper spirit of rescue. He drove in silence. He
couldn’t remember having wished to kiss a woman before at half past nine
in the morning, and it annoyed him.
Sally, of course, was silent too. Not for her to speak without being
spoken to, and she sat mildly wondering that she should be going along in a
car at all. Laura had come up to her bedroom and said her brother was there,
wanting to take her out for a little fresh air. Do her good, Laura had said,
though Sally had never known good come of fresh air yet; but, passive as a
parcel, she had let herself be taken. Why, however, she should be going for
a joy-ride with this lord she didn’t know, though she supposed it was as
good a way as another of getting through the intimidating day among the
picks of the basket, and anyhow this way there was only one of them, and
anyhow he wasn’t the big old one with the hairs on his hands.
Queer lot, these picks, thought Sally. Didn’t seem to have anything to do
to keep them at home; seemed to spend their time going somewhere else.
Fidgety. And a vision of her own life as it was going to be once she was
settled in those rooms at Cambridge, getting ready for her little baby, and
cleaning up, and making things cosy for her man, flooded her heart with a
delicious warmth. Laura had promised to help her find the rooms, and take
her to where Mr. Luke would be. Mr. Luke wouldn’t be angry any more
now, thought Sally—he’d be too pleased about the little baby; and Laura
seemed to know exactly where they would find him, and had assured her he
wouldn’t want to have Mrs. Luke living with them. Laura was queer too, in
Sally’s eyes, but good. Indeed Sally, feeling very much the married woman
after what had happened the evening before, feeling motherly already,
feeling exalted by the coming of her baby to a height immensely above
mere spinsterhood, went so far as to say to herself of Laura, with indulgent
affection, ‘Nice kid.’
§
They lunched at Thaxted. It was still only half past twelve, and Charles
had managed to be three hours doing the forty odd miles. There was a
beautiful church at Thaxted in which he could linger with her, for he didn’t
want to get to Crippenham till tea-time, and Crippenham was only about
nine miles beyond Cambridge, off the Ely road between Waterbeach and
Swaffham Prior.
Up to Thaxted, Charles was filled with an embarrassingly strong desire
to appropriate Sally for ever to himself. He hadn’t an idea how to do it, but
that was his wish. She sat there silent, beautiful beyond his dreams—and
how often and how wistfully had he not dreamt of what a woman’s beauty
might be!—pathetic, defenceless in the midst of a rudely jostling, predatory
world, like a child with a priceless pearl in its hand among the poor and
hungry, and he passionately loved her. As the miles increased, so did
Charles’s passion. He looked at her sideways, and each time with a fresh
throb of wonder. He wove dreams about her; he saw visions of magic
casements and perilous seas, and she behind them, protected, guarded,
worshipped by him alone; his soul was filled with poetry; he was lifted
above himself by this Presence, this Manifestation; he thought in terms of
music; the whole of England sang.
But at Thaxted he felt different, and began to think Sally ought to be
with those she belonged to; and by the time it was evening, and he was
meditating alone in the garden at Crippenham, he was quite sure of it.
At Thaxted he ordered the best lunch he could—Sally’s mouth watered
as she listened,—and while it was being got ready he took her into the
church. She was inattentively polite. The brisk movements of a big, close-
cropped man in a cassock, who strode busily about and made what seemed
to Sally a curtsey each time he crossed the middle aisle, appeared to interest
her much more than Charles’s remarks on the clear, pale beauty of the
building. It was rather like taking a dog to look at things. Charles didn’t
consciously think this, but there was an unawareness about Sally when
faced by the beauties of Thaxted Church, and when faced, coming down, by
the beauties of certain bits of the country that singing April morning, which
was very like, Charles subconsciously thought, the unawareness of a dog.
Ah, but how far, far more beautiful she herself was than anything else, he
thought; how exquisite she looked in Laura’s chinchilla wrap, with the
exalted thoughts of the men who had built the church, thoughts frozen into
the delicate greys, and silvers, and rose-colours of that fair wide place, for
her background.
The man in the cassock left off doing whatever he was doing on catching
sight of Sally, and, after looking at her a moment, came up and offered, his
eyes on her face, to show them round the church; a little cluster of
Americans dissolved, and flowed towards her; and a woman dressed like a
nun broke off her prayers, and presently sidled up to where she stood.
Charles removed her.
Thaxted is a quiet place, and he strolled with her through its streets till
their food should be ready. Its streets, quiet to begin with, didn’t stay quiet.
The people of Thaxted, for some reason incomprehensible to Charles,
because no two women could be more unlike, seemed to think Sally was
Mary Pickford. He heard whispers to that effect. Did they then think, too,
that he was the person known, he understood, as Doug?
He removed her a second time.
Perhaps the inn was as good a place as any to wait in. He had, however,
to engage a private room for their lunch, because so many people came in
and wished to lunch too; and it was when Sally had eaten a great deal of
greengage tart and cream—bottled greengages, Charles feared, but she said
she liked them—and drunk a great deal of raspberry syrup which had, he
was sure, never been near real raspberries and couldn’t be very good for
her, and then, while he was having coffee and she tea—he had somehow
stumbled on the fact that she liked tea after meals, and he watched with
concern the strength and number of the cups she drank—it was then that she
began to thaw, and to talk.
Alas, that she should. Alas, that she didn’t remain for ever silent,
wonderful, mysterious, of God.
Once having started thawing, it wasn’t in Sally’s generous nature to stop.
She thawed and thawed, and Charles became more and more afflicted. Lord
Charles—so, the night before, she had learned he was called—was
evidently a chip off the same block as her friend Laura; kind, that is. See
what a lovely dinner he was giving her. Also he had been much more like a
gentleman that day, and less like somebody who wanted to be a husband;
and after the greengage tart she began to warm up, and by the time she had
got to the cups of tea she felt great confidence in Charles.
‘Kind, ain’t you,’ she said with her enchanting smile, when he
suggested, much against his convictions, another pot of tea.
‘Isn’t everybody?’ asked Charles.
‘Does their best,’ said Sally charitably. ‘But it’s up ’ill all the way for
some as I could mention.’
By this time Charles was already feeling chilled. The raspberry syrup
and the cups of strong tea had estranged him. This perfect girl, he thought,
ought to be choice too in her food, ought instinctively to reject things out of
bottles, and have no desire for a second helping of obviously bad pastry.
Still, she was very young. He too, at Eton, had liked bad tuck. After all,
queer as it seemed, she had only got to the age he was at then.
He made excuses for her; and, it appearing to him important that he
should be in possession of more facts about her than those Laura had told
him the evening before, said encouragingly, ‘Do mention them.’
Sally did. She mentioned everybody and everything; and soon he knew
as much about her hasty marriage, hurried on within a fortnight to the first
man who came along, her return from her honeymoon to South Winch, the
determination of her mother-in-law to keep her apart from her husband, her
flight, helped by her father-in-law, back to her father, his rejection of her,
and her intention to rejoin her husband next day at Cambridge whether he
liked it or not, as he could bear.
He couldn’t bear much. It wasn’t only how she said it, but what she said.
Charles, who had at first been afflicted by her language, was now afflicted
by her facts. He shifted uneasily in his chair. He smoked cigarette after
cigarette. His thin brown face was flushed, and he looked distressed. In that
strange, defective, yet all too vivid speech which he so deeply deplored, she
drew for him a picture of what seemed sheer exploitation, culminating in
his own sister’s flinging herself hilariously into the game. This child; this
helpless child, who would obey anybody, go anywhere, do anything she
was told—in Charles’s eyes, as he listened and drew her out, she became
the most pathetic thing on earth. Everybody, it appeared, first grabbed at her
and then wanted to get rid of her. Everybody; himself too. Yes, he too had
grabbed at her, under a mealy-mouthed pretence of helping her, and now he
too wanted—not to get rid of her, that seemed too violent, too brutal a way
of putting it, but to hand her over, to pass her on, to send her back to that
infernal young Luke, who himself was trying to escape from her and leave
her to his mother. And the courage of the child! It was the courage of
ignorance, of course, but still it seemed to Charles a lovely thing, that was
afraid of nothing, of no discomfort, of no hard work, if only she might be
with her husband in their own home. Charles discovered that that was
Sally’s one wish, and that her simple ambition appeared to be to do what
she called work her fingers to the bone on behalf of that odious youth.
‘Mr. Luke,’ said Sally, who was unacquainted with any reason why she
shouldn’t say everything she knew to anyone who wished to hear, ‘Mr.
Luke, ’e thinks ’e can’t afford a ’ome yet for me, and so——’
‘Then he oughtn’t to have married you,’ flashed out Charles, infuriated
by the young brute.
‘Seemed ’e couldn’t ’elp it,’ said Sally. ‘Seemed as if it ’ad to be. ’E
——’
‘Oh yes, yes,’ interrupted Charles impatiently, for he hated hearing
anything about Jocelyn’s emotions. ‘Of course, of course. That was a quite
foolish remark of mine.’
‘Five ’undred pounds a year ’e got,’ went on Sally, ‘and me able to make
sixpence go twice as far as most can. Dunno wot ’e’s talkin’ about.’
And indeed she didn’t know, for she shared Mr. Pinner’s opinion that
five hundred a year was wealth.
‘Fair beats me,’ she added, after a thoughtful pause.
Well, thought Charles, the Moulsford family had behaved badly, and,
under the cloak of sympathy and wishing to help, his and Laura’s conduct
had been most base; but they were certainly going to make up for it now.
By God, yes. Crippenham, which he had at first thought of from sheer
selfishness as the very place to get Sally to himself in, was evidently now
the place of all others from which she could be helped. Quite close to
Cambridge, within easy reach of young Luke, and in it, all-powerful even
now in spite of his age, certainly all-powerful when it came to putting the
fear of God into an undergraduate, or whatever he was, his ancient but still
inflammable father. Naturally at ninety-three the old man consisted
principally of embers; but these embers could still be fanned into a partial
glow by the sight of a good horse or a beautiful woman, and Charles would
only need to show Sally to him to have the old man on her side. Not able to
hear, but able to see: what combination could, in the case of Sally, possibly
be more admirable?
He drove on after lunch, his conscience clear; so clear that before
leaving Thaxted he sent Laura a telegram telling her they were going to
Crippenham, because he no longer wanted her to be made anxious,—for
those only, thought Charles, are angry and wish to make others
uncomfortable who are themselves in the wrong. He was no longer in the
wrong; or, rather, he was no longer thinking with rapture of the wrong he
would like to be in if Sally could be in it with him. Her speech made a gulf
between them which his fastidious soul couldn’t cross. There had to be h’s
before Charles could love with passion. Where there were none, passion
with him collapsed and died. On this occasion it died at the inn at Thaxted
towards the end of lunch; and he was grateful, really, however unpleasant at
the moment its dying was. For what mightn’t have happened if she had
gone on being silent and only saying yes and no, and smiling the divine,
delicious smile that didn’t only play in her dimples but laughed and danced
in her darling eyes? Charles was afraid that in that case he would have been
done for. Talking, she had saved him; and though he still loved her, for no
man could look at Sally and not love her, he loved her differently,—kindly,
gently, with a growingly motherly concern for her welfare. After Thaxted
there was no further trace in his looks and manner of that which had made
Sally suspect him of a wish to be a husband.
But she was surprised when he asked her, as they drove along, whether
she would mind if he took her to his father in the country for the night,
instead of back to what he called noisy London. Laura was in London; why
should she be taken somewhere else, away from her? And to his father too
—to more picks, fresh ones; just as she was beginning to shake down nicely
with the ones she knew. Surely the father of the picks would be the most
frightening of all?
So she said, ‘Pardon?’ and looked so much alarmed that Charles,
smiling, explained that his father was staying at that moment quite near
Cambridge, and it would be convenient for the search for rooms she had
told him Laura had promised to undertake with her next day.
‘He’s quite harmless,’ Charles assured her, for she continued to look
alarmed—if where she was to be taken to next was near Cambridge, it must
also be near Woodles, and suppose her father were to happen to see her?
—‘and he’s all alone there till Laura goes back to him tomorrow. It will
cheer him up to have us. He’s ninety-three.’
Ninety-three? ‘Oh, my,’ said Sally politely. ‘ ’E ain’t ’alf old. Poor old
gentleman,’ she added with compassion, old people having been objects of
special regard and attention in the Pinner circle.
But for the rest of the drive she was silent, for she was trying to thread
her way among her indistinct and entangled thoughts, all of which seemed
confusedly to press upon her notice that she oughtn’t to be where she was at
all, that if she was anywhere it ought to be with her husband, and that with
every hour that passed she was sinking deeper and deeper in wrong-doing.
‘Soon be in right up to the neck,’ she said to herself with resigned
unhappiness; and sincerely wished it were that time tomorrow, and she
safely joined up with Mr. Luke, and finished for good and all with these
soft-spoken but headstrong picks.
XIII
§
While they, along the roads, were drawing every minute nearer, the
unconscious Duke was sitting in his plain study, having his plain tea, which
had been set beside him by his plain parlourmaid. This is not to say that the
parlourmaid was ill-favoured, but only that she wasn’t a footman.
There were no footmen at Crippenham. There was hardly anything there,
except the Duke. For years it had been his conviction that this annual
fortnight of the rest that is obtained by complete contrast prolonged his life.
Something evidently prolonged it, and the Duke was sure it was
Crippenham. There he went every Easter alone with Laura, because it was a
small house, and an ugly house, and a solitary house, and had nothing to
recommend it except that it was the exact opposite of every other
Moulsford possession.
Only Charles could come and go as he pleased; only he could dare break
in without notice on the sacred yearly business of prolonging life. Although
he had had ninety-three years of it, the Duke still wanted more. He liked
being alive, and it pleased him to keep Streatley waiting. Streatley, and the
other three children of his first marriage—absurd, he thought, to have to
refer to those four old things as children—were unpopular with their father.
He had never at any time cared much for them, and had begun to be really
angry with them when he was a lively seventy, and perceived that the
possession of children bordering on a heavy fifty made him seem less
young than he felt himself to be. Now that they were practically seventy
themselves, and old seventies too, and he not looking a day different, he
hoped, from what he had looked thirty years before, he was angrier with
them than ever. He admitted that other people might be old at ninety-three,
but he wasn’t; he was the exception. He didn’t feel old, and he didn’t, he
considered, look old, so what was all this talk of age? The press never
mentioned him without the prefix venerable; people pretended he was deaf,
when he could hear as well as any man if he wasn’t mumbled at; Laura was
continually making him sit out of draughts, just as if he were a damned
invalid; arms were offered him if he wanted to walk a few steps—he
couldn’t appear in the House without some officious member of it, usually
that ass Chepstow, who was eighty if a day himself, ambling across to help;
and every time he had a birthday the newspapers tumbled over each other
with their offensively astonished congratulations. Couldn’t a man be over
ninety without having it perpetually rubbed into him that he was old?
What he loved was his brood of young ones—Laura, Terry, and Charles;
and of this lively trio the dearest to him was Charles. So that, looking up
from his seedcake and seeing his last born coming into the room, not only
entirely unexpectedly but with a young woman, though he was surprised he
wasn’t angry; and when on their coming close to him he perceived the
exceeding fairness of the young woman, his surprise became pleasurable;
very pleasurable; in fact, pleasurable to excess.
He stared up at Sally a moment, not listening to what Charles was
saying, and then struggled to get on to his feet. Younger than his three
young ones ... much, much younger than his three young ones ... youth, ah,
youth ... lovely, lovely youth....
Charles wanted to help him, but was thrust aside. ‘Poor old gentleman,’
said Sally, catching him by the arm as he seemed about to lose his balance
and drop back into the chair.
‘Married?’ asked the Duke, breathing hard after his exertion, and
looking at Charles.
Charles shook his head.
‘ ’Course I’m married,’ said Sally with heat.
‘He means us,’ said Charles.
‘Us?’ repeated Sally, much shocked.
‘You’re going to be, then,’ said the Duke, looking first at her and then at
Charles, his face red with pleasure.
Charles shook his head again, and laughed.
But the Duke didn’t laugh. He stared at him a minute, and then said,
‘Fool.’
‘I got a nusband,’ said Sally indignantly.
‘He can’t hear,’ said Charles. ‘He’s very deaf.’
‘What does she say?’ asked the Duke. ‘Speak clearly, my dear—no,
don’t shout,’ he added; though Sally, far from going to shout, wasn’t even
opening her mouth. Poor old gentleman, she thought, gazing at him in silent
compassion; fancy him still being anybody’s father.
The Duke took her hand in a dry, cold grip.
‘Like shakin’ ’ands with a tombstone,’ thought Sally. And she was filled
with so great a pity for anything so old that she didn’t feel shy of him at all,
and in the coaxing voice of one who is addressing a baby she said, ‘ ’Ave
yer tea while it’s ’ot—do, now.’
Charles looked at her astonished. Nearly everybody was afraid of his
father. She reminded him of the weaned child in Isaiah, who put its hand
fearlessly on the cockatrice’s den.
‘What does she say?’ asked the Duke, gazing at her with delight.
‘This is Mrs. Luke, Father—a friend of Laura’s,’ shouted Charles, ‘and
I’ve brought her——’
‘Write it down, my dear,’ said the Duke, not heeding Charles, and
drawing Sally into a chair next his own and pushing paper and a pencil
towards her with his shaking old hands. ‘Write down what you were saying
to me.’
Charles became anxious. He felt sure Sally couldn’t write anything
down. Nor could she; for if her spoken words were imperfect her written
ones were worse, so that to be given a pencil and paper by the Duke and
told to write might have been embarrassing if she hadn’t, owing to his
extreme age and evident dilapidation, felt he wasn’t, as she said to herself,
all there. Poor old gentleman, she thought, full of pity. What she saw, sitting
heavily in the chair, breathing hard and blinking at her so kindly, was just,
thought Sally, the remains, the left-overs; like, she said to herself, her
images being necessarily domestic, Sunday’s dinner by the time one got to
Friday,—not much good, that is, but had to be put up with. No; there was
nothing frightening about him, poor old gentleman. More like a baby than
anything else.
‘ ’Ave yer tea while it’s ’ot,’ she said again, gently putting the paper and
pencil aside. ‘Do you good,’ she encouraged, ‘a nice ’ot cup of tea will.’
‘He can’t hear, you know,’ said Charles, much relieved by Sally’s
attitude. But with what confidence, he thought, couldn’t a thing so gracious
approach the most churlish, disgruntled of human beings; and his father
wasn’t either churlish or disgruntled,—he only looked as if he were, and
frightened people, and when he saw they were frightened he didn’t like
them, and frightened them more than ever.
The Duke, watching Sally’s every movement with rapt attention, thought
when she put her hand on the teapot to feel if it was still hot that she wanted
tea herself, and bade Charles ring the bell and order more to be brought, and
meanwhile he took the cup she offered him obediently, his eyes on her face.
He hadn’t got as far, being still in too great a condition of amazement at her
beauty, as wondering which of the ancient families of England had
produced this young shoot of perfection, and not being able to hear a word
she said took it for granted that the delicate-ankled—he was of the
practically extinct generation that looks first at a woman’s ankles,—slender-
fingered creature belonged to his own kind. True her hands were red hands;
surprisingly red, he thought, on her presently taking off her gloves, which
she rolled up together into a neat tight ball, compared to the flawless
fairness of her face; but they were the authentic shape of good-breeding,
even if her nails——
The Duke was really surprised when his eyes reached Sally’s nails.
Charles drew a chair close up to his father, and began his explanations.
He was determined the old man should attend, and shouted well into his ear
as he told him that he had motored Laura’s friend, Mrs. Luke, down from
London, where she had been staying with Laura at Goring House, to
Crippenham for the night because it was quieter, and she hadn’t been well
——
‘I’m all right,’ interrupted Sally, who had been listening in an attitude of
polite attention.
‘Oh, my dear child—when you fainted,’ protested Charles in his
ordinary voice, raising a deprecating hand.
‘Speak up,’ said the Duke, impatiently.
‘ ’Course I fainted,’ said Sally, looking pleased.
‘What does she say?’ asked the Duke.
‘Yes—and were unconscious for at least half an hour,’ said Charles.
‘That’s right. And sick,’ said Sally, looking proud.
‘Sick? Were you sick as well? Then see how really ill——’
‘Speak up, speak up,’ said the Duke testily.
But Sally said nothing further, and merely smiled indulgently at Charles.
‘What did she say?’ asked the Duke, not wishing to lose a word that fell
from that enchanting mouth.
‘She said,’ shouted Charles, ‘that she is quite well now.’
‘Of course she is,’ said the Duke, staring at her face and forgetting her
nails. ‘Anyone can see she is as perfectly well as she is perfectly beautiful.’
‘Oh lor,’ thought Sally, ‘now ’e’s goin’ to begin.’
§
That afternoon and evening were a triumph for her if she had known it,
but all she knew was that she was counting the hours to next day, and
Jocelyn, and the settling down at last to her home and her duties. The old
man was her slave. Crippenham and everything in it was laid at her feet,
and the Duke only lamented that it should be to this one of his houses that
she had come, where he couldn’t, he was afraid, make her even decently
comfortable. Positively at Crippenham there was only one bathroom. The
Duke seemed to regard this as a calamity, and Sally listened with mild
wonder to the amount he had to say about it.
‘Fair ’arps on it, don’t ’e, poor old gentleman,’ she remarked to Charles;
and bending over to the Duke’s ear—Charles looked on in astonishment at
the fearless familiarity of the gesture—she tried to convey to him that it
wasn’t Saturday night till the next night, and that by then she’d be in
Cambridge, so there was no need for him to take on.
‘Eh?’ said the Duke. ‘What does she say?’ he asked Charles.
‘She says,’ shouted Charles, ‘that it doesn’t matter.’
How very glad he was that his father was so deaf. Often he had found his
deafness trying, but how glad he was of it now. Not Saturday night....
Charles fell silent. It was then Friday. Could it be that since the previous
Saturday——?
The Duke, however, knew nothing of Sally except what his eyes told
him, and accordingly he was her slave. When she presently went up to
Laura’s room with the housekeeper, who had instructions to place
everything of Lady Laura’s at Mrs. Luke’s disposal—Crippenham had no
spare rooms, only a room each for the Duke, and Charles, and Laura, the
other six or seven bedrooms being left unfurnished and kept locked up—
and Charles, who from long practice could make his father hear better than
anyone except Laura, settled down to telling him as much about Sally as he
thought prudent, the old man listened eagerly, his hand behind his ear,
drinking in every word and asking questions which showed that if he was
really interested in a subject he still could be most shrewd.
He was delighted that Sally should have run away from her mother-in-
law, said it was proof of a fine, thoroughbred spirit, and asked who her
father was.
Charles said his name was Pinner.
The Duke then inquired whether he were one of the Worcestershire
Pinners, and Charles said he didn’t know.
The Duke then rambled off among his capacious memories, and
presently brought back a Pinner who had been at Christchurch with him,
and who had married, he said, one of the Dartmoors, an extremely
handsome woman, fair too, who was probably the girl’s grandmother.
Charles merely bowed his head.
The Duke then asked who the Lukes, apart from this boy-husband at
Ananias, were; for, he said, except the fellow in the Bible, he couldn’t
recollect ever having heard of a Luke before.
Charles said all he knew was that they lived at South Winch.
‘What?’ cried the Duke. ‘Has she married beneath her?’—and was so
really upset that for a time he blinked at Charles in silence. Because he felt
that if only this dear son of his had secured the beautiful young creature he
could have died content; and it seemed to him a double catastrophe that not
only should his boy have missed her, but that she should have been caught
into a misalliance with some obscure family in a suburb.
‘Upon my word, Charles,’ he said, after a dismayed silence, ‘that’s a
pity. A very great pity.’
And rambling off into his memories again, he said it was a good thing
that poor Jack Pinner was dead, for no man had a keener family feeling than
he, and it would have broken his heart to think his grand-daughter had made
a mistake of that kind.
He couldn’t get over it. He had never, in the whole of his long life, seen
anyone to touch this girl for beauty, and that she should, at the very outset
of what ought to have been a career of unparalleled splendour and success,
have dropped out of her proper sphere and become entangled in a suburb
really shocked him. Kings at her feet, all Europe echoing with her name—
this seemed to the Duke such beauty’s proper accompaniment.
‘Tut, tut,’ he said, his hands, clasped on the top of his stick, shaking
more than usual, ‘tut, tut, tut. What was her mother thinking of?’
‘Her mother is dead,’ said Charles.
‘Her father, then. Jack Pinner was no fool. I don’t understand how his
son—where is he, by the way? I heard something about the Worcestershire
estates having been sold after the war——’
Charles said he didn’t know where her father was, because, although
Sally had told him the shop was at Woodles, he had never heard of
Woodles, which indeed is not marked on any map, so that he felt he wasn’t
lying in saying he didn’t know.
The Duke, however, appeared to be seized by a sudden fierce desire to
track down his old friend’s reprehensible son and tell him what he thought
of him, and Charles was dismayed, for no good, he was sure, could come of
tracking down Mr. Pinner. Sally, he knew, was anxious her father shouldn’t
find out her disobedience to his orders, and though of this disobedience
Charles held Laura guilty, not Sally, yet he didn’t suppose Mr. Pinner would
look at it like that, and it was, besides, important, Charles considered, that
his father, who had always had a rooted objection to any woman who
wasn’t well-bred, should go on thinking Sally was a Worcestershire Pinner.
It seemed, then, to Charles a good thing to keep his father and Mr. Pinner
apart, and it was therefore with regret that he listened to the old man asking
Sally the moment he next saw her, which wasn’t till dinner, for she stayed
up in her room till fetched down by the scandalised housekeeper, to whom
it was a new experience that His Grace should be kept waiting even a
minute after the gong had sounded, where her father was.
‘ ’Im?’ said Sally, turning pale but forced by nature and her upbringing
to an obedient truthfulness. ‘ ’E’s at Woodles, ’e is.’ And, ‘Oh my
gracious,’ she added to herself, ‘they ain’t goin’ to tell ’im I’m ’ere?’
‘What does she say?’ the Duke asked Charles.
‘She says,’ shouted Charles, following his father, who was shuffling
along leaning on Sally’s arm, to the dining-room, and shouting with
outward composure but inward regret, ‘that he is at Woodles.’
‘Woodles? Woodles?’ repeated the Duke. ‘Never heard of it. Is it in
Worcestershire?’
Sally shook her head. She didn’t know where Worcestershire was, but
she felt pretty sure Woodles wasn’t in it.
‘I dunno wot it’s in,’ she said. And then, impelled as always to the naked
truth, she added, ‘Close by ’ere, any’ow.’
‘What does she say?’ inquired the Duke, turning again to Charles.
‘She says,’ shouted Charles, obliged to hand on the answer correctly
with Sally listening, but doing so with increased regret, ‘that it isn’t far
from here.’
‘How very lucky,’ said the Duke, ‘and how very odd that I shouldn’t
have known he was so near.’ And he added, when he had been lowered into
his chair at the head of the table by the parlourmaid, who held one arm, and
his servant, who held the other, ‘I’d like to have a talk with that father of
yours, my dear.’
Sally turned paler.
‘Your grandfather was one of my oldest friends,’ continued the Duke,
with difficulty unfolding his table-napkin because of how much his hands
shook.
‘I ain’t got no grandfather,’ said Sally anxiously, who had never heard of
him till that moment.
‘She says,’ began Charles reluctantly—‘You know,’ he muttered quickly
to Sally, for how could he tell the old man what she had said? ‘you have a
grandfather—or had. You must have. Everybody has them.’
‘What? What?’ said the Duke impatiently. ‘Send a message round
tonight, Charles, and say with my compliments that I’d very much like to
see Pinner. Tell him I’m too old to go to him, so perhaps he’ll be obliging
enough to come to me some time tomorrow. You can say his father was at
Oxford with me if you like, and that I’ve only just heard he is in the
neighbourhood. Say his daughter——’
‘Now don’t—now don’t go doin’ a thing like that,’ Sally faintly begged
of Charles.

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Biofertilizers: Study and Impact 1st

Polymers Coatings: Technology and Applications 1st

Biodiesel Technology and Applications Inamuddin

Impact 1st Edition Ali Rahim

Biosensors Nanotechnology, 2nd Edition Inamuddin

Biodegradable Materials and Their Applications

Interdisciplinary Team Teaching: A Collaborative Study

Applied Soil Chemistry Inamuddin

Flexible Supercapacitor Nanoarchitectonics Inamuddin

Study and Impact

Wiley Global Headquarters

Limit of Liability/Disclaimer of Warranty

Library of Congress Cataloging-in-Publication Data

Cover image: Wikimedia Commons

Printed in the USA

2.3.2 Enhanced Nutritional Content of Produce 49

3.11 Phosphorus Solubilizing Microbes 115

4.4.5.3 Weed Management 138

5.6 Bacteria-Fungi Consortium for Phosphate Solubilization 165

7.3.4.1 Polymer Entrapped Formulations 239

8.6.5 Generation of Genetically Modified Transgenic

10.1.2.6 Association With Plant Roots 316

12.3 Morphology and Mode of Reproduction 373

14 Quality Control of Biofertilizers 413

15.5.4 Arbuscular Mycorrhizal and Phosphate

16.4 Preparation Approaches for Biofertilizers 499

17.4.1.2Carbon Sequestration 529

20.3 Advances in Biofertilizer 580

22.3.1 Algal Extracts as Biofertilizer 613

Chapter 2 highlights the impact of applying biofertilizers on horticul-

Chapter 9 intends to provide detailed information on the application

Chapter 17 discusses the use of biofertilizers based on waste recycling as

School of Biological Sciences, Quest International University, Perak, Malaysia

Keywords: Biofertilizer, forestry, plant growth promoting microorganisms,

*Corresponding author: wendy.liu@qiup.edu.my

plants and other beneficial microorganisms [16–18]. The utilization of bio-

1.2 Mechanisms of Actions of Biofertilizers

1.2.1 Facilitation of N Acquisition

microorganisms in forest biomes can be categorized into two categories:

1.2.1.1 Mutualistic N2 Fixation

1.2.1.2 Non-Symbiotic N2 Fixation

1.2.2 Facilitation of P Acquisition

macromolecule synthesis, respiration, and photosynthesis [30, 60]. It has

1.2.2.1 Phosphate Solubilizing Microorganisms

without the acidification process. Besides OA production, inorganic acids

and arbuscules are specialized structures of AMF that assist in mobilizing

1.2.3 Potassium Solubilization

1.2.4 Production of Siderophores

1.2.5 Modulation of Phytohormones

reach equilibrium, thus aiding in regulating plants’ responses to environ-

1.3 Factors Influencing the Outcome

and field experimentation strategies, as observed in the application of

well as reproduction in Pinus pinaster and P. pinea, and mitigating infec-

1.4 Applications of Biofertilizers in Forestry

with Azotobacter and Azospirillum [155, 223], phosphobacteria [224–227],

1.5 Conclusion and Future Prospects

of a biofertilizer with the host plant species to the surrounding environ-

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