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Fungal Biology
Ajar Nath Yadav · Shashank Mishra

Divjot Kour · Neelam Yadav
Anil Kumar Editors
Agriculturally
Important Fungi
for Sustainable
Agriculture
Volume 2: Functional Annotation
for Crop Protection
Fungal Biology
Series Editors
Vijai Kumar Gupta
AgroBioSciences (AgBS) and Chemical & Biochemical Sciences (CBS)
Department Mohammed VI Polytechnic University (UM6P)
Benguerir, Morocco
Maria G. Tuohy
School of Natural Sciences
National University of Ireland Galway
Galway, Ireland
About the Series
Fungal biology has an integral role to play in the development of the biotechnology
and biomedical sectors. It has become a subject of increasing importance as new
fungi and their associated biomolecules are identified. The interaction between
fungi and their environment is central to many natural processes that occur in the
biosphere. The hosts and habitats of these eukaryotic microorganisms are very
diverse; fungi are present in every ecosystem on Earth. The fungal kingdom is
equally diverse, consisting of seven different known phyla. Yet detailed knowledge
is limited to relatively few species. The relationship between fungi and humans has
been characterized by the juxtaposed viewpoints of fungi as infectious agents of
much dread and their exploitation as highly versatile systems for a range of
economically important biotechnological applications. Understanding the biology
of different fungi in diverse ecosystems as well as their interactions with living and
non-living is essential to underpin effective and innovative technological
developments. This series will provide a detailed compendium of methods and
information used to investigate different aspects of mycology, including fungal
biology and biochemistry, genetics, phylogenetics, genomics, proteomics, molecular
enzymology, and biotechnological applications in a manner that reflects the many
recent developments of relevance to researchers and scientists investigating the
Kingdom Fungi. Rapid screening techniques based on screening specific regions in
the DNA of fungi have been used in species comparison and identification, and are
now being extended across fungal phyla. The majorities of fungi are multicellular
eukaryotic systems and therefore may be excellent model systems by which to
answer fundamental biological questions. A greater understanding of the cell
biology of these versatile eukaryotes will underpin efforts to engineer certain fungal
species to provide novel cell factories for production of proteins for pharmaceutical
applications. Renewed interest in all aspects of the biology and biotechnology of
fungi may also enable the development of “one pot” microbial cell factories to meet
consumer energy needs in the 21st century. To realize this potential and to truly
understand the diversity and biology of these eukaryotes, continued development of
scientific tools and techniques is essential. As a professional reference, this series
will be very helpful to all people who work with fungi and should be useful both to
academic institutions and research teams, as well as to teachers, and graduate and
postgraduate students with its information on the continuous developments in
fungal biology with the publication of each volume.
More information about this series at http://www.springer.com/series/11224

Ajar Nath Yadav • Shashank Mishra
Divjot Kour • Neelam Yadav • Anil Kumar
Editors
Agriculturally Important
Fungi for Sustainable
Agriculture
Volume 2: Functional Annotation for Crop
Protection
Editors
Ajar Nath Yadav Shashank Mishra
Department of Biotechnology, Dr. Khem Biotechnology
Singh Gill Akal College of Agriculture QCQA Laboratory, Biotech Park
Eternal University Lucknow, Uttar Pradesh, India
Sirmour, Himachal Pradesh, India
Neelam Yadav
Divjot Kour Gopi Nath PG College
Department of Biotechnology, Dr. Khem Veer Bahadur Singh Purvanchal University
Singh Gill Akal College of Agriculture Ghazipur, Uttar Pradesh, India
Eternal University
Baru Sahib, Sirmour, India
Anil Kumar
Rani Lakshmi Bai Central Agricultural
University
Jhansi, Uttar Pradesh, India
ISSN 2198-7777 ISSN 2198-7785 (electronic)

Fungal Biology
ISBN 978-3-030-48473-6 ISBN 978-3-030-48474-3 (eBook)
https://doi.org/10.1007/978-3-030-48474-3
© Springer Nature Switzerland AG 2020

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms or in any other physical way, and transmission or information
storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology
now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Preface
Sustainable agriculture is the only solution in providing food security to feed the
gigantic population, with resource limitation being a foremost challenge for the
global community. Sustainable agriculture requires limited use of harmful chemical
fertilizers and pesticides. Among diverse groups of microbes, soil and plant fungal
communities play an important role in plant growth, development, and soil health.
The beneficial fungal communities help to promote plant growth directly or indi-
rectly via different plant growth-promoting mechanisms viz: releasing plant growth
regulators; solubilization of phosphorus, potassium and zinc; biological nitrogen
fixation or by producing siderophores, ammonia, HCN and other secondary metab-
olites. The plant growth promoting fungal communities with multifunctional PGP
attributes could be used as biofertilizers and biocontrol agents replacing chemical
fertilizers and pesticides in the environmental as eco-friendly agents for sustainable
agriculture and environment. Fungal communities possess a huge sink of capability
by which they act as bioprotectants and biostimulants as well as for mitigation of
different abiotic stress in plants. The utilization of beneficial soil and plant fungal
resources will surely support sustainable agriculture.
The present book on “Agriculturally Important Fungi for Sustainable Agriculture,
Volume 2: Functional Annotation for Crop Protection” covers soil- and plant-asso-
ciated fungal communities and their role in plant growth promotion, and crop pro-
ductivity for sustainable agriculture. This book will be immensely useful to the
biological sciences, especially to microbiologists, microbial biotechnologists, bio-
chemists, researchers, and scientists dealing with fungal biotechnology. We have the
honour that the leading scientists who have extensive, in-depth experience and
expertise in plant-microbe interaction and fungal biotechnology took the time and
v
vi Preface
made efforts to contribute these outstanding chapters. Each chapter is written by

internationally recognized researchers and scientists so that the readers are given an
up-to-date and detailed account of our knowledge of fungal biotechnology and its
innumerable agricultural applications.
Sirmour, Himachal Pradesh, India Ajar Nath Yadav

Lucknow, Uttar Pradesh, India Shashank Mishra
Baru Sahib, Sirmour, India Divjot Kour
Ghazipur, Uttar Pradesh, India Neelam Yadav
Jhansi, Uttar Pradesh, India Anil Kumar
Acknowledgements
All the authors are sincerely acknowledged for contributing up-to-date information
on agriculturally important fungi, their biodiversity and biotechnological applica-
tions for sustainable agriculture and environments. The editors are thankful to all
the authors for their valuable contributions.
All editors would like to thank their families who were very patient and support-
ive during this journey. Our sincere thanks to the whole Springer team who was
directly or indirectly involved in the compilation of this book. We are grateful to the
many people who helped to bring this book to light. Editors wish to thank Mr. Eric
Stannard, Senior Editor, Botany, Springer; Dr. Vijai Kumar Gupta, and Prof Maria
G. Tuohy, Series editor, Fungal Biology Springer; Ms. Saveetha Balasundaram,
Project Coordinator, Springer for generous assistance, constant support, and
patience in initializing the volume.
The editor Dr. Ajar Nath Yadav is grateful to his Ph.D. research scholars Tanvir
Kaur, Rubee Devi, Divjot Kour, Kusam Lata Rana and colleagues for their support,
love, and motivation in all his efforts during this project.
We are very sure that this book will be great interest to the scientists, graduates,
undergraduates, and postdocs who are investigating fungal biology and
biotechnology.
vii
Contents
1 Agriculturally Important Fungi for Crop Protection�� 1

Pavidharshini Selvasekaran and Ramalingam Chidambaram
2 Role of Fungi in Adaptation of Agricultural Crops
to Abiotic Stresses�� 55
Fatma A. Abo Nouh and Ahmed M. Abdel-Azeem
3 Arbuscular Mycorrhizae Associations and Role
in Mitigation of Drought Stress in Plants�� 81
Deepika Goyal, Sushma Mishra, and Prem Kumar Dantu
4 Soil Salinity and Its Alleviation Using Plant
Growth–Promoting Fungi �� 101
Prem Chandra, Enespa, and Ranjan Singh
5 Fungal Secondary Metabolites and Bioactive
Compounds for Plant Defense�� 149
Shomaila Sikandar, Asfand Yar Saqib, and Imran Afzal
6 Fungal Endophytes: A Source for Biological Control Agents�� 181
Hiran Kanti Santra and Debdulal Banerjee
7 Aspergillus Mycotoxins: Potential as Biocontrol Agents�� 217
Fatma A. Abo Nouh, Sara A. Gezaf, and Ahmed M. Abdel-Azeem
8 Trichoderma: Biocontrol Agents for Promoting
Plant Growth and Soil Health�� 239
Hesham A. El Enshasy, Kugan Kumar Ambehabati,
Ashraf F. El Baz, Santosh Ramchuran, R. Z. Sayyed,
Divina Amalin, Daniel J. Dailin, and Siti Zulaiha Hanapi
9 Piriformospora indica: Endophytic Fungus
for Salt Stress Tolerance and Disease Resistance�� 261
Fatma A. Abo Nouh, Hebatallah H. Abo Nahas,
and Ahmed M. Abdel-Azeem
ix
x Contents
10 Fungi: A Bio-resource for the Control of Plant

Parasitic Nematodes�� 285
K. Kiran Kumar
11 Global Scenario of Advance Fungal Research
in Crop Protection�� 313
Vanita Yadav, Murthy M. V. S. N. Lekkala, Challa Surekha,
and Nageswara Rao Reddy Neelapu
12 Functional Annotation of Agriculturally
Important Fungi for Crop Protection: Current
Research and Future Challenges�� 347
Ajar Nath Yadav, Divjot Kour, Tanvir Kaur, Rubee Devi,
and Neelam Yadav
Index�� 357
Contributors
Ahmed M. Abdel-Azeem Systematic Mycology Laboratory, Botany Department,

Faculty of Science, University of Suez Canal, Ismailia, Egypt
Hebatallah H. Abo Nahas Zoology Department, Faculty of Science, Suez Canal
University, Ismailia, Egypt
Fatma A. Abo Nouh Systematic Mycology Laboratory, Botany Department,
Faculty of Science, University of Suez Canal, Ismailia, Egypt
Imran Afzal Department of Biology, Lahore Garrison University, Lahore, Pakistan
Divina Amalin Biological Control Research Unit, Center for Natural Sciences and
Environmental Research, De La Salle University, Manila, Philippines
Kugan Kumar Ambehabati Institute of Bioproduct Development (IBD),
Universiti Teknologi Malaysia (UTM), Johor Bahru, Malaysia
Debdulal Banerjee Microbiology and Microbial Biotechnology, Department of
Botany and Forestry, Vidyasagar University, Midnapore, West Bengal, India
Prem Chandra Department of Environmental Microbiology, School for
Environmental Sciences, Babasaheb Bhimrao Ambedkar University (a central
university), Lucknow, Uttar Pradesh, India
Ramalingam Chidambaram Food Science and Technology, School of Biosciences
and Technology, Vellore Institute of Technology, Vellore, Tamil Nadu, India
Daniel J. Dailin Institute of Bioproduct Development (IBD), Universiti Teknologi
Malaysia (UTM), Johor Bahru, Malaysia
Prem Kumar Dantu Department of Botany, Dayalbagh Educational Institute,
Deemed University, Agra, Uttar Pradesh, India
xi
xii Contributors
Rubee Devi Department of Biotechnology, Eternal University, Sirmour, Himachal

Pradesh, India
Enespa Department Plant Pathology, Sri Mahesh Prasad Degree College,
University of Lucknow, Lucknow, Uttar Pradesh, India
Ashraf F. El Baz Genetic Engineering and Biotechnology Research Institute,
Sadat University, Sadat City, Egypt
Hesham A. El Enshasy Institute of Bioproduct Development (IBD), Universiti
Teknologi Malaysia (UTM), Johor Bahru, Malaysia
Sara A. Gezaf Botany Department, Faculty of Science, Al-Arish University,
North Sinai, Egypt
Deepika Goyal Department of Botany, Dayalbagh Educational Institute, Deemed
University, Agra, Uttar Pradesh, India
Siti Zulaiha Hanapi Institute of Bioproduct Development (IBD), Universiti
Teknologi Malaysia (UTM), Johor Bahru, Malaysia
Tanvir Kaur Department of Biotechnology, Eternal University, Sirmour, Himachal
Pradesh, India
Divjot Kour Department of Biotechnology, Dr. Khem Singh Gill Akal College of
Agriculture, Eternal University, Sirmour, Himachal Pradesh, India
K. Kiran Kumar Department of Nematology, ICAR-Central Citrus Research
Institute, Nagpur, Maharashtra, India
Murthy M. V. S. N. Lekkala Department of Biochemistry and Bioinformatics,
Institute of Science, Gandhi Institute of Technology and Management (GITAM),
(Deemed-to-be University), Visakhapatnam, Andhra Pradesh, India
Sushma Mishra Department of Botany, Dayalbagh Educational Institute, Deemed
University, Agra, Uttar Pradesh, India
Nageswara Rao Reddy Neelapu Department of Biochemistry and Bioinformatics,
Institute of Science, Gandhi Institute of Technology and Management (GITAM),
(Deemed-to-be University), Visakhapatnam, Andhra Pradesh, India
Santosh Ramchuran Bioprocess Development Department, CSIR Biosciences,
Pretoria, South Africa
Hiran Kanti Santra Microbiology and Microbial Biotechnology, Department of
Botany and Forestry, Vidyasagar University, Midnapore, West Bengal, India
Asfand Yar Saqib Department of Biology, Lahore Garrison University, Lahore,
Pakistan
R. Z. Sayyed Department of Microbiology, PSGVP Mandal’s, Arts, Science, and
Commerce College, Shahada, Maharashtra, India
Contributors xiii
Pavidharshini Selvasekaran Food Science and Technology, School of Biosciences

and Technology, Vellore Institute of Technology, Vellore, Tamil Nadu, India
Shomaila Sikandar Department of Biology, Lahore Garrison University, Lahore,
Pakistan
Ranjan Singh Department of Environmental Science, School for Environmental
Sciences, Babasaheb Bhimrao Ambedkar University (A Central University),
Lucknow, Uttar Pradesh, India
Challa Surekha Department of Biochemistry and Bioinformatics, Institute of
Science, Gandhi Institute of Technology and Management (GITAM), (Deemed-
to-be University), Visakhapatnam, Andhra Pradesh, India
Ajar Nath Yadav Department of Biotechnology, Dr. Khem Singh Gill Akal
College of Agriculture, Eternal University, Sirmour, Himachal Pradesh, India
Neelam Yadav Gopi Nath PG College, Veer Bahadur Singh Purvanchal University,
Ghazipur, Uttar Pradesh, India
Vanita Yadav Department of Biochemistry and Bioinformatics, Institute of
Science, Gandhi Institute of Technology and Management (GITAM), (Deemed-
to-be University), Visakhapatnam, Andhra Pradesh, India
About the Editors
loc
Ajar Nath Yadav is an Assistant Professor (Sr. Scale)

in the Department of Biotechnology, Dr. Khem Singh
Gill Akal College of Agriculture, Eternal University,
Baru Sahib, Himachal Pradesh, India. He has 5 years of
teaching and 11 years of research experiences in the
field of Microbial Biotechnology, Microbial Diversity,
and Plant-Microbe-Interactions. Dr. Yadav obtained
doctorate in Microbial Biotechnology, jointly from
IARI, New Delhi and BIT, Mesra, Ranchi, India; M.Sc.
(Biotechnology) from Bundelkhand University and
B.Sc. (CBZ) from the University of Allahabad, India.
Dr. Yadav has 296 publications, with h-index of 39, i10-index of 92, and 4045 cita-
tions (Google Scholar). Dr. Yadav is editor of 11 Springer-Nature, 04 CRC Press
Taylor and Francis and 02 Elsevier books. Dr. Yadav has got 12 Best Paper
Presentation Awards, and 01 Young Scientist Award (NASI-Swarna Jayanti Purskar).
Dr. Yadav received the “Outstanding Teacher Award” in 6th Annual Convocation
2018 by Eternal University, Baru Sahib, Himachal Pradesh. Dr. Yadav has a long-
standing interest in teaching at the UG, PG, and PhD level and is involved in taking
courses in microbiology and microbial biotechnology. Dr. Yadav has guided 01
Ph.D. and 01 M.Sc. Scholar and presently he is guiding 03 scholars for PhD degree.
He has been serving as an editor/editorial board member and reviewer for more than
35 national and international peer-reviewed journals. He has lifetime membership
of Association of Microbiologists in India and Indian Science Congress Council,
India. Please visit https://sites.google.com/site/ajarbiotech/ for more details.
xv
xvi About the Editors
Shashank Mishra is presently working as Scientist

“C”, Biotech Park, Lucknow, Uttar Pradesh, India. He
obtained his doctorate degree in Science “Industrial
Biotechnology” in 2015, from Birla Institute of
Technology, Mesra, Ranchi, India; M.Phil.
(Biotechnology) in 2008 from Alagappa University,
Tamil Nadu; M.Sc. (Botany) in 2005 from Dr.
R.M.L. University, Ayodhya; M.Sc. (Biotechnology) in
2004 from Barkatullah University, Bhopal, and B.Sc.
(Botany and Chemistry) in 2001 from Dr.
R.M.L. University, Ayodhya, India. He has made pio-
neering contributions in the area of Microbial
Biotechnology; Natural product synthesis and
Environmental Microbiology for food, pharmaceutical, and Human Health. In his
credit 21 publications [06 research papers, 02 review articles, 03 books, and 10
book chapters] in different reputed international and national journals and publish-
ers with 82 citation, h-index-05, and i10-index 04 (Google Scholar). He has reported
first time with high concentration of phenolic compounds by optimizing various
parameters and published in peer-reviewed and refereed international journals. He
has published 16 abstracts in different conferences/symposiums/workshops. He has
presented 16 papers presentation [12 poster + 04 oral] in conferences/symposiums
and got 01 best poster presentations Award, Dr. Mishra has contributed in organiz-
ing 07 conference/workshops. He has deposited 03 nucleotide sequences to NCBI
GenBank databases: in public domain. Dr. Mishra and group have isolated and char-
acterized 03 microbes (bacteria and microalgae) from Tulsi and paddy plantation
site and transformed ferulic acid into value-added phenolic compounds, viz., vanil-
lin, vanillic acid, and 4-vinylguaiacol. He has a long-standing interest in teaching at
the UG, PG, and Ph.D. level and is involved in taking courses in industrial biotech-
nology, bioprocess engineering and technology, environmental biotechnology, envi-
ronmental microbiology, industrial microbiology, microbial biotechnology and
techniques in microbiology and biotechnology. He is reviewer in 06 international
journals including BMC Microbiology, Indian Phytopathology, PLOS One,
Scientific Reports and Archive of Phytopathology and Plant Protection, and 3
Biotech. He has lifetime membership of Association of Microbiologists of India
(AMI) and Vigyan Bharti (VIBHA).
Divjot Kour is currently working as Project Assistant

in DEST funded project “Development of Microbial
Consortium as Bioinoculants for Drought and Low
Temperature Growing Crops for Organic Farming in
Himachal Pradesh”. Dr. Kour obtained doctorate degree
in Microbial Biotechnology from Department of
Biotechnology, Dr. Khem Singh Gill Akal College of
Agriculture, Eternal University, Baru Sahib, Himachal
Pradesh, India. She obtained M.Phil. in Microbiology
in 2017 from Shoolini University of Biotechnology and
About the Editors xvii
Management Sciences, Solan, Himachal Pradesh; M.Sc. in Microbiology from

University of Jammu, Jammu and Kashmir in 2015, and B.Sc. from University of
Jammu, Jammu and Kashmir in 2014. She has research experience of 04 years. In
her credit 39 publications (11 research/reviews and 28 book chapters) in different
reputed international, national journals and publishers. She is editor of 02 books in
Springer. She has published 18 abstracts in different conferences/symposiums/
workshops. She has presented 09 papers in national and international conferences/
symposiums and got 05 best paper presentation awards. In her credit more than 700
microbes (bacteria and fungi) isolated from diverse sources. She has the member-
ship of National Academy of Sciences and Agro Environmental Development
Society, India.
Neelam Yadav is currently working on microbial

diversity from diverse sources and their biotechnologi-
cal applications in agriculture and allied sectors. She
obtained her post graduation degree from Veer Bahadur
Singh Purvanchal University, Uttar Pradesh, India. She
has research interest in the area of beneficial microbi-
omes and their biotechnological application in agricul-
ture, medicine, environment, and allied sectors. In her
credit 59 research/review/book chapters publication in
different reputed international, national journals and
publishers. She is editor of 02 books in Elsevier, 03 in
CRC press, Taylor & Francis, and 05 in Springer. She is
the Editor/associate editor/reviewer of different international and national journals
including Plos One, Extremophiles, Annals of Microbiology, Journal of Basic
Microbiology, Advance in Microbiology and Biotechnology. She has lifetime
membership of Association of Microbiologists in India, Indian Science Congress
Council, India, and National Academy of Sciences, India.
Anil Kumar is Director, Education, Rani Lakshmi

Bai Central Agricultural University, Near Pahuj dam
Gwalior Road, Jhansi (UP). He has 30 years of research,
teaching, and administrative experience in various
capacities. He had worked as Professor & Head (2006–
till date), Associate Professor (2003–2006), and
Assistant Professor (1993–1999), Dept. of Molecular
Biology & Genetic Engineering, G. B. Pant University
of Agriculture & Technology, Pantnagar. He has to his
credit more than 200 publications including research
papers, reviews, and chapters contributed to books,
popular articles, books/bulletins/manuals. His research
interests focus on diverse but specific areas of agricul-
tural importance at national and international level
employing the tools and techniques of molecular biology and immunology.
Dr. Kumar has strengthened the area of molecular plant pathology, for combating
xviii About the Editors
the Karnal bunt which is an economically important disease of wheat, he followed

three approaches, viz., Plant disease surveillance through molecular/immunological
diagnostics; Pathogen Indexing Programme through Molecular Pathotyping, and
Characterization of disease resistance and Pathogenesis through Molecular signal-
ing investigating the role of MAP kinases and Cystatin gene families as candidate
genes. It was postulated that stoichiometric balance of cystatin and cysteine prote-
ase might be contributing to disease resistance and susceptibility. Dr. Kumar has
filed several patents on synthesis of nano delivery vesicles for facilitation of uptake
of fat soluble vitamins, nano-curcuminoids for better bio-availability, and nano-iron
pro-booster technology for agronomic bio-fortification. His pioneer research work
has been highlighted by several magazines like NATURE and published in several
international journals of repute with citation index: >2398, h-index 27, and i10
index 82. He has been an outstanding teacher and researcher who is credited with
many awards and recognitions, viz., Dr. Radhakrishnan Best Teacher Award, INSA
Best Teacher Award, Dr. C. Subramaniam Outstanding Teacher Award, Outstanding
Faculty Recognition, Dr. B.B. Singh Outstanding Researcher Award, and also con-
ferred three times “Governor’s Award” for best research in the year 2015, 2017, and
2019 from different organizations including university ICAR, INSA, and DBT.
Chapter 1
Agriculturally Important Fungi for Crop
Protection
Pavidharshini Selvasekaran and Ramalingam Chidambaram
Contents
1.1 Introduction 2
1.2 The Role of Fungi as Biofertilizers 3
1.2.1 Advantages of Fungal Biofertilizers 4
1.3 The Role of Fungi as a Biocontrol Agent 5
1.4 Fungal Endophytes 7
1.4.1 Direct Mechanism 8
1.4.2 Indirect Mechanism 10
1.4.3 The Advantages of Fungal Endophytes Over Chemical Methods 15
1.5 Arbuscular Mycorrhizae and Its Role in Agriculture 15
1.6 Penicillium and Its Role as Plant Growth Promotion 17
1.6.1 The Role of Penicillium Under Abiotic Conditions 18
1.7 Trichoderma for Sustainable Agriculture 20
1.7.1 Mycoparasitism 21
1.7.2 Antibiosis 22
1.7.3 Competition and Competence of Rhizosphere 23
1.7.4 Enzyme Production 23
1.7.5 Enhancement of Host Defense Response 24
1.7.6 Growth Promotion of Host Plants 24
1.7.7 Phosphate Solubilization 25
1.7.8 Drought Tolerance 26
1.8 Piriformospora indica 27
1.8.1 Acquisition of Phosphorus 28
1.8.2 Acquisition of Other Nutrients 29
1.8.3 Plant Growth and Development 29
1.8.4 Role in the Tolerance of Biotic and Abiotic Stress 30
1.9 Notable Research on the Role of Fungi in Agriculture 33
1.10 Conclusion and Future Perspective 34
eferences
R 35
P. Selvasekaran · R. Chidambaram (*)

Food Science and Technology, School of Biosciences and Technology,
Vellore Institute of Technology, Vellore, Tamil Nadu, India
e-mail: cramalingam@vit.ac.in
© Springer Nature Switzerland AG 2020 1

A. N. Yadav et al. (eds.), Agriculturally Important Fungi for Sustainable
Agriculture, Fungal Biology, https://doi.org/10.1007/978-3-030-48474-3_1
2 P. Selvasekaran and R. Chidambaram
1.1 Introduction
Fungi are one of the most important pathogens of crop plants in agriculture and
forestry but they are also used as potential biocontrol agents to prevent and control
plant disease. In recent years, the development of fungi for biocontrol of diseases,
pests, and weeds has received a significant amount of interest among the community
of scientists and researchers (Rangel et al. 2018). Public concerns on the usage of
pesticides in the agricultural field and their effect on the environment are increasing
day by day. Many fungi are being developed and mass produced as a commercially
available biocontrol agents and are used in agriculture to promote the growth of
plant and increase plant defense mechanism (Vega et al. 2009) as well as to control
plant disease (Costa et al. 2012), terrestrial weeds (Monteiro, AC, Machado 2012).
These biocontrol agents can also control and reduce aquatic weeds, plant-parasitic
nematodes (Siddiqui and Mahmood 1996), and insects (Alston et al. 2005; Li
et al. 2010).
Recent study in agricultural system on the role of Arbuscular mycorrhiza sug-
gests that they can improve salinity tolerance and drought tolerance of their host
plant by increasing the nutrient uptake, accumulation of organic solutes, and reduc-
ing the oxidative stress as a result of enhanced activity of catalase, peroxidase,
calmodulin, superoxide dismutase, and ascorbate peroxidase (Abdel Latef and
Chaoxing 2014; Chandrasekaran et al. 2014; Huang et al. 2014). An important role
in stress tolerance is enhanced by symbiotic mycorrhizal fungi in both forest and
agricultural ecosystems by improving nutrient uptake and drought tolerance and
restricting base cation leaching as well as by mitigating the toxic effects of alumi-
num and other heavy metals (Finlay 2008; Finlay et al. 2009; Yadav 2017, 2019b).
The non-mycorrhizal fungi such as Trichoderma are well known for mediating
stress reactions in plants and have the potential to parasitize or antagonize plant
pathogenic fungi by stimulating the defense response and increasing the plant
growth (Druzhinina et al. 2011; Sharma et al. 2019). The other fungi species such as
Piriformospora indica induce disease resistance, tolerate salt stress, and promote
the growth of crops through their capacity of glutathione ascorbate cycle mediated
antioxidation activity (Waller et al. 2005). The enhancement of plant growth utiliz-
ing plant bio-stimulants and to access novel molecules by exploiting the fungal
stress response to be used in agriculture is of current interest (Calvo et al. 2014;
Yadav 2018).
Endophytic microbes specifically endophytic fungi are those that have estab-
lished an equilibrium with their host during the evolution and are recognized as a
potential source of a wide variety of bioactive secondary metabolites (Rana et al.
2019a; Tan and Zou 2001; Sonaimuthu et al. 2010; Gangadevi and Muthumary
2008). Their relationship with the host plant can vary from bordering on the patho-
gen to symbiotic. They adopt a different type of symbiosis such as facultative sap-
robic, exploitative, parasitic, and mutualistic (Clay and Schardl 2002). In most
cases, the plant hosts are benefited by asymptomatic relationship. However in few
cases they may exhibit beneficial or pathogenic effects (Photita et al. 2001; Wei
1 Agriculturally Important Fungi for Crop Protection 3
et al. 2007; Neubert et al. 2006) Few endophytic fungi exhibit mutualistic relation-
ship with single plant species and not with the other plant species whereas some
species have the potential to associate with wide range of plant host as well as asso-
ciation of endophyte with specific host tissue has also been reported (Hardoim et al.
2015; Yadav et al. 2020d). Fungal endophytes are ubiquitous and have been found
in the majority of plant species that have been studied to date (Rana et al. 2019b;
Yadav et al. 2019b). However, the literature available on the association between
few plant host and endophytic fungi is limited and has to be explored further
(Brundrett 2007).
Endophytic fungi by secreting selected secondary metabolites have found to be
associated with the plants to promote growth (Dai et al. 2008), protect from insects
and disease (Wilkinson et al. 2000; Tanaka et al. 2005), and improve stress resis-
tance (Lewis 2004). These secondary metabolites produced by endophytic fungi
also have been identified as the source of antidiabetic, insecticidal, anticancer,
immunosuppressive, and biocontrol compounds. These novel compounds help as
remedies with their valuable biological and chemical characteristics for the health
problems of plants, animals, and humans. The fungal endophytes have been repre-
sented as “chemical synthesizer inside the plant” because of the production of a
variety of these useful chemical compounds (Yadav et al. 2020b, c).
Fungal diversity on earth is contributed significantly by endophytic fungi (Photita
et al. 2001). Their biological distribution and diversity are massive in tropical rain-
forest and temperate zone. The detection of novel compounds from this group of
fungi is extensively studied to investigate their potential to be used in pharmaceuti-
cal, medical, industrial, and agricultural sectors. A noteworthy discovery is the use
of these compounds in agriculture as a biocontrol agent (Mane and Vedamurthy
2018). Fungal endophytes contribute to the fitness of plants by enabling the adap-
tion of plant host to abiotic and biotic stresses. They establish a symbiotic relation-
ship between their hosts and confer disease and pest resistance to host by enhancing
their fitness, increasing the growth to resist stresses to the maximum extent, and
promote the production of secondary metabolites (Gautam and Avasthi 2019;
Rastegari et al. 2020a). Therefore, the use of fungi for the purpose of sustainable
agriculture is of major interest.
1.2 The Role of Fungi as Biofertilizers
Biofertilizers are an important source of essential nutrients for plants and crops.
They are inexpensive and eco-friendly. They play a vital role in improving the nutri-
ent status of soil, increase soil fertility, and, thus, crop productivity. Biofertilizers
are the formulation of living microbes including bacteria, actinomycetes, and fungi
(Kour et al. 2020d). They can be applied directly to plant roots, soil, seed, and seed-
lings. Due to their inherent biological activity of the microorganisms, they help in
mobilization and accessibility of nutrients (Pal et al. 2015). Fungal biofertilizers can
be applied alone or in combination to the natural fields and can be beneficial directly
or indirectly to plant development, yield, and growth through various methods (Rai
et al. 2013; Yadav et al. 2019c). The roots of herbs, trees, xerophytes, epiphytes,
aquatics, shrubs, trees, aquatics, hydrophytes, and terrestrial plants are reported to
develop mycorrhizal associations when grown with the insufficient availability of
essential elements such as zinc, phosphorus, nitrogen, iron, sulfur, boron, and cop-
per (Rastegari et al. 2020b; Singh and Yadav 2020).
Fungal biofertilizers that solubilize phosphate are the biological agents that are
commonly employed for improving the growth and development of plants by
enhancing the phosphorus uptake. The phosphate solubilizing property of fungi
contributes significantly to the soil phosphate availability to plants. Several fungi
have phosphate solubilizing property, the most common are Saccharomycopsis
schoenii, Cryptococcus luteolus, Trichosporon beigelii, Rhodotorula aurantiaca A,
Kluyveromyces walti, Neosartorya fisheri var. fischeri, Candida montana,
Penicillium purpurogenum var. rubrisclerotium, and Zygoascus hellenicus (Birhanu
et al. 2017). The fungi that solubilize phosphate belonging to the genera Fusarium,
Aspergillus, Penicillium sp. are also found in the rhizospheric region of vari-
ous plants.
The fungal genera of Penicillium, Chaetomium, and Aspergillus are of wide-
spread occurrence (Yadav et al. 2018, 2019c). The commonly employed fungi for
the production of biofertilizer are Trichoderma which is predominantly present in
agricultural soils. The rhizosphere inhabited by Trichoderma species can also inter-
act and parasitize with other fungi. These species have long been recognized as they
enhance crop nutrition, nutrient acquisition, and augment plant productivity. The
metabolites produced by these species serve as a fungicide against the fungal patho-
gens that cause disease (Harman et al. 2008; Chang 1986; Vinale et al. 2009). The
utilization of Trichoderma as a culture filtrate and its inoculation in soil enhances
biomass production and plant growth. The application of this fungus as a model
organism is feasible as they are easy to cultivate under laboratory conditions for the
evaluation of beneficial interaction between plants and microbes. They can be used
as a novel tool to enhance the productivity of plants (Varma et al. 2012).
1.2.1 Advantages of Fungal Biofertilizers
• They are renewable sources of nutrients and sustain soil health.

• They increase the yield of grains by 20–40% and replace 40–50% of chemical
fertilizers.
• They enhance plant growth by secreting growth-promoting hormones and also
secrete antibiotic and fungi static substances. They do not have an adverse effect
on soil fertility and plant growth.
• Phosphate solubilizing or phosphate mobilizing fungal biofertilizer converts
insoluble soil phosphate into soluble soil phosphate due to the secretion of vari-
ous organic acids. Under optimum condition, they can mobilize or solubilize
about 40–50 kg phosphorus and results in a 20–30% increase in the crop yield.
• They enhance the uptake of P, Zn, S, and water leading to increased yield and
uniform crop growth. They improve the hardiness of the stock transplant and
enhance resistance to root disease.
• They liberate the substance that promotes growth, vitamins, and maintain soil
fertility.
• They stabilize C:N ratio of soil and decompose the residues of the plant.
• Fungal biofertilizers act as an antagonist and help in the biocontrol of disease by
suppressing the incidence of soil-borne pathogens.
• Fungal biofertilizer plays a significant role in the recycling of plant nutrients.
• Fungal biofertilizers are eco-friendly, non-pollutants as well as cost-effective
(Table 1.1) (Pal et al. 2015)
1.3 The Role of Fungi as a Biocontrol Agent
Currently, the rapidly growing research area with a significant role in increased food
production and plant yield is “biocontrol.” The phenomenon of biocontrol agent
helps to sustain the food crop quality and to reduce the risks that result from the
increased utilization of hazardous chemicals and synthetic pesticides. The major
factor responsible for 10–30% of annual crop productivity loss is plant disease. In
spite of the development of water management practice, an agricultural practice that
has posed to be the effective management of plant disease, new techniques in agron-
omy, the development of disease-resistant varieties, there are still pathogens for
which the synthetic chemicals are broadly used for the management of the disease
(Kour et al. 2019b; Yadav et al. 2020a). The attractive choice of using the biocontrol
agent against different pathogens has emerged as the most common and significant
factor responsible for the death of insects in a large populations (Villa et al. 2017).
Several postharvest diseases are biologically balanced using fungal species that
have antagonist properties. The worldwide scientist is attracted by this area of
research. The components of the biocontrol system are influenced by various factors
such as UV light, temperature, pH, as well as abiotic and biotic stresses (Kumar
et al. 2019a, b; Yadav 2019a). The biocontrol agent preparation is significantly
affected by various abiotic stress factors which lead to modification of functionality.
Therefore, knowledge on the survivability of biocontrol agents under certain envi-
ronmental conditions and the development of the procedure to make the biocontrol
agent resist stress tolerance is required to maintain their commercial exploitation
and effectiveness. The experimental studies on various biocontrol agents on differ-
ent plant species have been studied to understand their reaction under varied envi-
ronmental conditions (Sui et al. 2015) (Table 1.2).
Table 1.1 Fungal biofertilizers for crop protection and sustainable agriculture
6
List of agriculturally important fungal biofertilizer Reference

Zinc solubilizing biofertilizer Martino et al. (2003)
• Ericoid mycorrhiza (oidiodendron maius) Franz et al. (1991)
• Penicillium simplicissimum Martino et al. (2003)
• Saccharomyces spp. Wold and Suzuki (1976)
• Aspergillus niger
Potash solubilizing biofertilizer Lian et al. (2008)
• Aspergillus spp. (A. terreus, A. fumigatus, A. niger) Alves et al. (2010)
• Ectomycorrhizal fungi
Phosphate mobilizing biofertilizer Adholeya et al. (2005)
• Arbuscular mycorrhiza, Glomus spp. (G. viscosum, (G. mosseae/G. cerebriforme/G. manihotis/G. aggregatum, G. da Silva (2006)
intraradices, G. versiforme, G. deserticola, G. radiatum, G. globiferum, G. monosporum, G. microcarpum, G. halonatum) Rai et al. (2013)
• Paraglomus, Geosiphon, Acaulospora spp. (A. delicata, A. foveate, Sclerocystis clavispora, A. scrobiculata) Anderson and Cairney (2007),
• Archaeospora, Scutellospora spp. (S. scutata, S. erythropa, S. calospora) Pal et al. (2015)
• Ectomycorrhiza (Pisolithus tinctorius, Amanita sp., Tuber sp., Lactarius sp., Elaphomyces sp., Pisolithus sp.,
Piriformospora indica, Cenococcum sp., Rhizopogon sp.)
• Entrophospora, Gerdemannia, Gigaspora (Gigaspora rosea)
Phosphorus solubilizing biofertilizer Manoharachary et al. (2005)
• Aspergillus spp. (A. tubingensis/A. niger/A. terreus/A. awamori/A. fumigatus/A. tubingensis/A. melleus) Menon and Mohan (2007)
• Penicillium spp. (P. rubrum/P. expansum/P. citrinum, P. simplicissimum/P. frequentans/P. oxalicum/P. bilaji/P. albidum/P. Burton and Knight (2005)
italicum) Harman et al. (2004)
• Trichoderma spp. (T. virens/T. asperellum, T. viride/T. harzianum) Manoharachary et al. (2005)
• Fusarium spp. (F. moniliforme/F. udam) Of (2009)
• Mucor spp. (M. ramosissimus/M. mucedo/M. hiemalis) Pal et al. (2015)
• Candida spp. (C. scotti, C. krissii)
• Tritirachium spp. (T. album/T. egenum)
Biofertilizer enriching compost Manoharachary et al. (2005)
• Trichoderma spp. (T. virens/T. asperellum, T. viride/T. harzianum) Singh and Singh (2008)
• Aspergillus spp. (A. terreus/A. awamori/A. fumigatus, A. tubingensis/A. niger) Whitelaw (1999)
• Pleurotus spp. (P. ostreatus/P. flabellatus) Harman et al. (2004)
• Fusarium spp. (F. solani/F. oxysporum) Of (2009), Pal et al. (2015)
P. Selvasekaran and R. Chidambaram
• Chaetomium spp. (C. bostrychodes, C. olivaceum)

Table 1.2 Fungal communities as biocontrol agents against plant species

S. No. Fungi Plant species Reference
1. Pichia guilliermondii Glycine max
2. Pichia membranifaciens Vitis vinifera Santos and
Marquina (2004)
3. Candida oleophila Vitis vinifera Porat et al. (2003)
4. Lentinus conatus Arachis hypogaea Lakshmanan et al.
(2008)
5. Penicillium roqueforti; Penicillium viridicatum Allium cepa Khokhar et al.
(2013)
6. Streptomyces lydicus WYEC108 P. sativum Yuan and
Crawford (1995)
7. Trichoderma asperellum Tv5SC, T. harzianum Helianthus annuus, Prasad (2015)
Th4d SC Ricinus communis
8. Trichoderma virens IMI-392430, T. Capsicum annuum Rahman et al.
pseudokoningii IMI-392431, T. harzianum (2012)
IMI-392432, T. harzianum IMI-392433, T.
harzianum IMI-392434
9. Trichoderma koningii, Trichoderma Pisum sativum Nelson et al.
harzianum (1988)
10. Pichia angusta Malus domestica Fiori et al. (2008)
11. Penicillium sp. EU0013 Solanum Alam et al. (2011)
lycopersicum;
Brassica oleracea
12. Leucosporidium scottii At 17 Malus domestica Vero et al. (2013)
13. Penicillium citrinum VFI-51 Sorghum bicolor Meesala and
Subramaniam
(2016)
14. Saccharomycopsis schoenii Citrus X sinensis Hashem et al.
(2012)
15. Penicillium adametzioides Vitis vinifera Ahmed et al.
(2015)
1.4 Fungal Endophytes
The efficient and new biological control agents used for the control of pathogens
and insects and the bioremediation of the environment are fungal endophytes (Guo
et al. 2008). The evidence shows that the fungal endophytes play a significant role
in protecting the host from the disease, supporting plant health and plant physiol-
ogy. Fungal endophytes obtain nutrition, propagation opportunities, protection and
shelter from their host by internal plant tissue colonization. This symbiotic relation-
ship helps the host by promoting the overall health and reducing environmental
sensitivity. The best offer to the control of chemical disease is the use of endophytes
(Yao et al. 2017). To control the plant diseases these biocontrol agents utilize direct
and indirect strategies by promoting the accumulation of bioactive compounds,
increasing tolerance against stress, and enhancing fitness. An in-depth
understanding of the mechanism involved in the association of plants and fungal

endophytes is of major interest to optimize the efficacy as well as for the registration
of products that are used for plant protection (Rana et al. 2019a; Suman et al. 2016;
Yadav et al. 2018). The possible mechanism that is considered to be adopted by the
fungal endophytes against the pathogens may be mycoparasitism, induction, anti-
biosis, direct and indirect inhibition, competition, and improvement of host plant
resistance (Yu et al. 2010; Nisa et al. 2015; Gautam and Avasthi 2019). The aspects
that are involved in the control of disease by these biocontrol agents are dis-
cussed below.
1.4.1 Direct Mechanism
Recent studies have proved the ability of the fungal endophytes to protect the host
plant from various diseases and damages due to the activity of plant pathogenic
microorganisms. The fungal endophytes produce bioactive compounds which have
the potential to inhibit the growth of pathogenic microorganism directly. Some
researchers have proposed a few possible mechanisms by the use of fungal endo-
phytes and their secondary metabolites to suppress the pathogens (Yadav et al.
2017b, c). However, the knowledge on the mechanism between plant, endophyte,
and pathogen is to be explored further. In direct mechanism fungal endophytes
secrete lytic enzymes and produce antibiotics or the pathogens are directly sup-
pressed through antibiosis, mycoparasitism, and competition (Bamisile et al. 2018;
Mejía et al. 2008; Ganley et al. 2008; Gautam and Avasthi 2019).
1.4.1.1 Mycoparasitism
Few endophytes exhibit hyper parasitism distinct from antibiosis and competition.
The plant pathogenic fungi are often attacked by these fungi that act as mycopara-
sites either to inhibit or suppress their growth. One of the most popular examples of
mycoparasitism is Trichoderma. Prior to making any contact, Trichoderma detects
the fungal pathogen and the extracellular exochitinase is produced in lower levels
and discharges the oligomers of cell wall from the target fungus (Harman et al.
2004). This mechanism initiates the Trichoderma to release toxic endochitinase that
can degrade the target plant-fungal pathogen. The hyphae of the plant pathogens
including Rhizoctonia solani is parasitized by the Trichoderma species (Grosch
et al. 2006). A novel endophytic fungus Acremonium strictum isolated from Dactylis
glomerate has been reported (Rivera-Varas et al. 2007) for the activity of mycopara-
sitism against Helminthosporium solani. In the process of mycoparasitism the bind-
ing of chemical compounds between the host fungus and mycoparasite occurs
initially. One such example is binding of lecithin in the cell wall of the host fungi to
the carbohydrate present in the cell wall of Trichoderma. Later the establishment of
contact with the host fungus is inhabited by the hyphae of the mycoparasite. Several
lytic enzymes are involved along with the mechanisms in the cell wall degradation
of the host fungi (Cao et al. 2009). These findings have proven that the pest damage
in the agricultural crops can be limited by the mycoparasitic activity of the endo-
phytic fungi and can prove to be a potential alternative to the use of chemical pesti-
cides (Gautam and Avasthi 2019).
1.4.1.2 Competition
One of the important methods used by the fungal endophytes against the prolifera-
tion and infection of the plant pathogen is competition. This process involves com-
petition between the endophytic fungi and pathogens for space and few common
resources (Mejía et al. 2008). Few endophytic organisms can control plant patho-
gens involving such mechanisms and they could be used as an effective biocontrol
agent against the plant disease. The effect on traditional rice varieties by the most
frequently isolated endophytic fungi on the growth of plant and incidence of blast
disease was evaluated by Atugala and Deshappriya (2015). The study concluded
that a range of mechanisms of antagonistic activity was utilized by the endophytic
fungi including the competition for antibiosis, substrate, and mycoparasitism.
Studies across the world (Atugala and Deshappriya 2015; Mejía et al. 2008;
Richmond 2004) demonstrated the use of endophytic fungi as a management strat-
egy against numerous harmful pathogens (Gautam and Avasthi 2019).
1.4.1.3 Antibiosis
A wide spectrum of secondary metabolites is produced by fungal endophytes with

the ability to reduce the attacks from various pests and insects. These fungi possess
antibacterial, antifungal as well as insecticidal properties that strongly inhibit the
growth of plant pathogens and infectious microorganisms (Gautam et al. 2013). The
single or multiple kinds of antibiotics which has been proven to be effective has
been reported to be produced by single fungus including aromatic compounds,
polypeptides, terpenoids, and alkaloids. Researchers (Mejía et al. 2008; Wei et al.
2007) have been carrying out the examination of antibiotic compounds produced by
endophytic fungi and have proven that the plants inoculated with endophytic fungi
induce the defense mechanism in the host plant. It has been determined that the
cultures of endophytic fungi possess the antibiotic properties against several plant
pathogens after a series of tests involving liquid broths of endophytic fungi. It was
also reported by Atugala and Deshappriya (2015); Kim et al. (2007); Wessels (1999)
that the proteins secreted by the species of endophytic fungi act as a protein related
with pathogenicity to suppress the activity of plant pathogens by degrading the cell
wall. Several studies reported that various chemical compounds are produced with
the antibiotic property against pathogens by the fungal endophytes. The examina-
tion of the interaction between the endophytic fungi and Puccinia recondite f. sp.
tritici was done by Dingle and McGee (2003) and found that the leaf rust disease
caused by this fungus was suppressed by the fungal endophytes. An endophytic

fungus Phomopsis cassiae isolated from Cassia spectabilis (Lu et al. 2000a) secretes
a chemical compound 3,11,12-trihydroxycadalene which was found to be effective
against Cladosporium Cladosporioides and Cladosporium sphaerospermum. The
antifungal activities of 183 endophytic fungi isolated from 15 plant species from 15
locations in Korea against pathogens causing rice sheath blight disease, tomato gray
mold disease, rice blast disease, wheat leaf rust disease, tomato late blight disease
and barley powdery mildew diseases such as Corticium sasakii, Botrytis cinerea,
Magnaporthe grisea, P. recondita, and Blumeria graminis f. sp. hordei, respectively,
were evaluated by Park et al. (2003). In another study examination of woody angio-
sperm Theobroma cacao associated with fungal endophytes against foliar pathogen
sp was done (Arnold et al. 2003) and the study revealed that inoculation of leaf tis-
sues by the assemblage of endophytic fungi that are isolated from asymptomatic
and infected host limited the damage caused by a significant foliar pathogen
(Gautam and Avasthi 2019).
1.4.2 Indirect Mechanism
The association of plants with fungal endophytes has a direct impact on the growth
and development of the host plant. The host performance is affected in terms of
morphology, growth, physiology, and biochemistry. The important reason for the
improvement of host health is due to the production of secondary metabolites by
these fungi along with the establishment of a symbiotic relationship (Kaur et al.
2020; Singh et al. 2020; Yadav et al. 2019a). The association of endophytic myco-
biota with the host plant is believed to be responsible for the fate of pathogen attack
(Wei et al. 2007; Vyas and Bansal 2018). These beneficial microbes can decrease
the disease incidence, plant stress, increase the efficiency of nitrogen fixation,
improve the nutrient uptake, growth rate of the shoot and root, and improve resis-
tance (Liu et al. 2001; Kuldau and Bacon 2008). These properties of fungal endo-
phytes can be used to play an essential role in the field of food safety, agriculture,
and maintenance of environmental equilibrium (Rana et al. 2020). The indirect
mechanism adapted by fungal endophytes in the control of pest and disease is
given below.
1.4.2.1 Production of Metabolites
In the adaption of plants to biotic and abiotic stresses, the plant-fungal symbiotic
association plays an important role by the production of secondary metabolites
(Yadav 2019a). The significant factor in the production of various metabolites is the
association between plants and endophytic fungi. The environmental and eco-
friendly approach to control various plant diseases is the use of endophytic fungi
(Rana et al. 2019a, b). The host plant is benefited by the endophytic fungi as they
Table 1.3 Bioactive compounds produced by fungal endophytes

Fungal endophyte Bioactive compounds Reference
Cladosporium delicatulum Plumbagin Venkateswarulu
(5-hydroxyl-2-methylnaptalene-1,4- et al. (2018)
dione)
Balansia obtecta Ergobalancine Tintjer and
Rudgers (2006)
Melanconium betulinum 3-hydroxypropionic acid Chomcheon et al.
(2005)
Phomopsis phaseoli 3-hydroxypropionic acid Chomcheon et al.
(2005)
Neotyphodium coenophialum Ergovaline Tintjer and
Rudgers (2006)
Acremonium coenophialum Chitinases Gautam and
Avasthi (2019)
Neotyphodium sp. Ergonovine Miles et al. (1996)
Fusarium redolens Peimisine and Pan et al. (2015)
imperialine-3β-D-glucoside
Aspergillus fumigatus fumiclavine, A, fumiclavine B, Panaccione (2005)
fumiclavine C and Chanoclavine
aldehyde
Colletotrichum gloeosporioides Piperine Chithra et al.
(2014)
Rhinocladiella sp., Helminthosporium Bioactive cytochalasines Isaka et al. (2001)
sp., Chalara sp., Phoma sp.,
Hypoxylon sp., Phomopsis sp.,
Xylaria sp.
Xylaria sp. YX-28 7-amino-4-methylcoumarin Xu et al. (2008)
Penicillium sp. Berkeleydione Davis et al. (2005)
Ampelomyces sp. O-methyl alaternin and altersolanol Miller et al.
A (1998)
Morinia pestalozzioides Moriniafungin Gautam and
Avasthi (2019)
Aspergillus niger Aurasperone A, fonsecinone A, Song et al. (2004)
asperpyrone B, rubrofusarin B
Pestalotiopsis adusta Pestalachlorides Li et al. (2008)
Periconia sp. Periconicin A Kim et al. (2004)
Chaetomium chiversii C5-36-62 Radicicol Turbyville et al.
(2006)
Pestalotiopsis jester Hydroxyjesterone, jesterone Li and Strobel
(2001)
Muscodor albus 1-Butano, 3-methylacetate Strobel and Daisy
(2003)
Pestalotiopsis microspora Ambuic acid Li and Strobel
(2001)
Colletotrichum gloeosporioides Colletotric acid Zou et al. (2000)
Epichloe festucae Cyclonerodiol, IAA, diacetamide, Yue et al. (2000)
IEtOH, indole-3-carboxaldehyde,
methylindole-3-carboxylate
prevent the colonization of pathogenic organisms (Arnold et al. 2003). It has been
reported that fungal endophytes secrete a few chemical compounds that inhibit the
plant pathogenic fungi based on the experimental studies. The endophytes isolated
from the medicinal plants have cytotoxic metabolites, a fungicidal and bactericidal
activity which can produce secondary metabolites in the tissue of the host and kill
the pathogens (Zhang et al. 2006). It has been reported that the endophytic fungi
produce secondary metabolites to protect the host plant from disease and pest
(Sudha et al. 2016). These bioactive compounds serve as mediators for specific
communication and interaction with the host plant. The competition, fitness, and
growth of the host plant are triggered by the endopytic fungi upon production of the
secondary metabolites that inhibit the pathogens. These metabolites are categorized
into flavonoids, quinones, chinones, saponins, xanthones, phenolic acid, alkaloids,
tetralones, benzopyranones, tannins, and several others.
The research on the secondary metabolites and their antipathogenic effects pro-
duced by the endophytic fungi to explore their application as a biocontrol agent in
various fields has been conducted worldwide. It was reported that Fusarium spp. E4
and E5 promoted the content of terpenoids and the growth of Euphorbia pekinensis
(Gautam and Avasthi 2019). Another study reported (Strobel and Daisy 2003) the
production of 25–28 volatile compounds by an endophytic fungi Muscodor albus,
isolated from the branches of Cinnamomum zeylanicum. These volatile compounds
are capable of inhibiting the selected bacteria and fungi. Another endophytic fungus
Muscodor crispans of Ananas ananassoides produces a mixture of volatile com-
pounds containing an antibacterial and antifungal property with inhibitory effect
against pathogenic fungi such as Botrytis cinerea, P. palmivora, Pythium ultimum,
Fusarium culmorum, Alternaria helianthi, Rhizoctonia solani, Verticillium dahliae,
and bacteria including Xanthomonas axonopodis (Yuan et al. 2017). Currently,
researchers are investigating the endophytic fungi production of secondary metabo-
lites which can inhibit the insects (Gautam and Avasthi 2019) (Table 1.3).
1.4.2.2 Growth of Root and Shoot
In the process of plant growth, the plant host and mutualistic organisms producing
compounds that regulate the growth of the plant and antimicrobial substances are
asymptomatically colonized by the endophytic fungi to enhance the competitive-
ness, growth, and fitness of the host in nature (Sudha et al. 2016). These fungi help
the host plant in phosphate solubilization whereas the plant provides food and shel-
ter. These fungi also help to enhance the uptake of phosphorus, nitrogen fixation,
production of siderophores including several other plant hormones such as gibberel-
lins, auxin, ethylene, abscisins, and IAA which are important for the regulation of
plant growth (Boddey et al. 2003; Firáková et al. 2007). Hamayun et al. (2009)
examined the endophyte Cladosporium sphaerospermum isolated from Glycine
max for the production of bioactive molecules that promote plant growth. The study
revealed that Cladosporium sphaerospermum produces GA3, GA4, and GA7
growth hormones that are associated with inducing the plant growth in soybean and
rice. It was reported (Khan et al. 2013) that the endophytic fungi Fusarium
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know this street better than I do. That’s your place opposite, isn’t it?
What’s behind that window over the shop?”
“A room I use for storing herbs in,” replied Mr. Ludgrove. “As it
happens, I have been in there once or twice this evening.”
“You didn’t see anything of what was happening in here, I
suppose?” asked Whyland quickly.
“No, I did not,” replied Mr. Ludgrove. “I was only in the room for
a moment or two, selecting bundles of herbs to take downstairs. I
did happen to notice about eleven o’clock that there was a light in
here, but that was all.”
“What about the windows to the right and left of your place?”
“I believe that they belong to offices occupied only in the day-
time. I do not think it at all likely that anyone could have overlooked
this room from them at night.”
“No,” agreed Whyland. “We’re not likely to be lucky to find an
actual witness. All right, Mr. Ludgrove, thank you. Now, young man,
do you know if your father possessed a hypodermic syringe?”
Ted Copperdock, thus addressed, shook his head. “I don’t think
so, Inspector,” he replied. “But I shouldn’t know one if I saw it.
There’s a cupboard over there by the washing stand where he kept a
lot of bottles of stuff.”
Whyland strode over to the cupboard and opened it. It contained
about a dozen bottles of various sizes, each half full of some patent
medicine or other. But of a hypodermic syringe, or even any caustic
potash, there was no trace.
“If he did it himself he must have thrown the syringe away,”
muttered Whyland. “Waters ought to find it; there isn’t a lot of traffic
along here at this time of night. Well, doctor, I don’t think we need
keep you out of bed any longer. I’ll have the body taken to the
mortuary, and perhaps you’ll ring me up at the station later in the
morning?”
The doctor nodded, picked up his bag, which contained the
broken needle and the sample of the incrustation, and left the
house. When he had gone, Whyland turned once again to Ted.
“Can you suggest any reason why your father should wish to take
his life?” he enquired.
“No, Inspector, I can’t,” replied Ted frankly. “The business is
doing very well, and father was only saying the other day that we’d
got a tidy bit put away in the bank. I keep the books myself, and I
know everything’s all right.”
“I see. No money troubles, in fact. You don’t know of any
disappointment which he may have experienced, or anything like
that?”
A faint smile passed across Ted’s face. “I don’t think he had any
disappointment, Inspector,” he replied. “In fact, I should say it was
rather the other way.”
“What do you mean?” said Whyland sharply.
“Why, he always reckoned that nobody knew, but I fancy that we
all guessed sharp enough. He’s hinted to me once or twice lately
that he wasn’t too old to marry again. And——well, from what her
daughter lets drop, Mrs. Tovey wouldn’t mind. He went round there
pretty often, and she always seemed glad to see him.”
Whyland shot a quick glance at Mr. Ludgrove. It was from him
that he had first learnt of this attachment. Ludgrove nodded almost
imperceptibly, and Whyland turned once more to Ted.
“There was nothing preying on his mind, was there?” he asked.
“Well, I’ve thought sometimes that he fair had the wind up about
this black sailor,” replied Ted reluctantly. “I never knew what to make
of that. He told me one day that he’d met him coming out of the
Cambridge Arms, but I never could quite believe it somehow.”
“As it happens, I share your scepticism,” said Whyland. “Mr.
Ludgrove here was in the street at the time, and saw your father
come out of the Cambridge Arms. There was nobody but himself
and your father in sight, he assures me.”
“That is so,” assented Mr. Ludgrove gravely.
“Well, I’m not surprised,” said Ted. “It’s a funny thing, but these
things always happen when he’d been to the Cambridge Arms of an
evening. It was when he came home from there that he found that
counter the other day?”
“Have you ever seen your father definitely under the influence of
liquor?” asked Whyland.
“Why no, not to say actually squiffy. He’d talk freer than usual,
and imagine all sorts of yarns about things that never happened. I
think he got the black sailor on his brain sometimes. When he first
got the counter he made up his mind that the black sailor was going
to get him. But the last day or two he’s been much more cheerful. Of
course, it’s possible that this evening, when he was alone, it got on
his mind again.”
The conversation was interrupted by the return of Waters, the
detective. “I’ve searched as best I can for that syringe, sir, and I
can’t find it,” he reported. “I’ll have another good look as soon as it
gets light, if you like, sir.”
“Yes, do,” replied Whyland. “Now, you were supposed to be
watching this place all the evening. What time did Mr. Copperdock
come in?”
“Between nine and ten, sir. Two fellows came with him, and the
three stood talking at the door for a minute or two. Then they all
went in, and the door was shut behind them. It was close on eleven
when it opened again, and the two men came out. Mr. Copperdock
came downstairs with them, I saw him just inside the door talking to
them. Then they went away, and the door was shut again. A few
minutes after they had gone I saw a light come on in this room.”
“You can’t see into the room from the opposite pavement, I
suppose?”
“Only a bit of ceiling, sir. I noticed the window was open at the
top, and the curtains not properly drawn, like you see them now, sir.
They’ve never been properly drawn since I’ve been watching the
house. The next thing that happened was that Mr. Copperdock’s son
came along at about a quarter past twelve, and let himself in with a
key.”
Inspector Whyland turned to Ted. “What time did you go out?”
he asked.
“About eight o’clock. Dad was just getting ready to go round to
the Cambridge Arms.”
“Where were you between eight and a quarter past twelve?”
“With Miss Tovey,” replied Ted, readily enough, but with an
awkward blush. “We went to a dance, then we had some supper.
After that I saw her home, and stayed there a few minutes. I walked
home from Lisson Grove, found Dad like this, and ran straight across
to fetch Mr. Ludgrove.”
“Nobody but you and your father had a key to the premises, I
suppose?”
“No. One of us always came down to let the charwoman in in the
morning.”
“You are perfectly certain, Waters, that nobody came to the
house between eleven and a quarter past twelve?” enquired
Whyland.
“Certain, sir. I was in the street outside all the time, and never
took my eyes off the place.”
“Very well. You stay here with the body. I’ll arrange for it to be
taken to the mortuary as soon as I can. As for you, young man, you
had better go to bed and try to get some sleep. We shall want you in
the morning. Mr. Ludgrove, if you’ve nothing better to do, I should
like you to come round the house with me. I want to make certain
that nobody can have broken in.”
Mr. Ludgrove nodded, and the two left the room together.
Whyland examined the sitting-room window. It was shut and
fastened, and bore no traces of violence. Then they went downstairs
and looked over the ground floor, without discovering anything in
any way out of the ordinary.
When they reached the office behind the shop, Whyland closed
the door and sank wearily into a chair. “Well, Mr. Ludgrove, what do
you make of it?” he said.
“I couldn’t help overhearing snatches of your conversation with
the doctor,” replied the herbalist. “I confess that I cannot understand
why, if Mr. Copperdock wished to poison himself with a hypodermic
injection, he should select his back for the purpose, unless he had
some confused idea of a lumbar puncture. Yet, on the other hand,
he is not likely to have let someone else drive a needle into him
without a struggle, and of that there is no trace, so far as I could
see.”
“And how did that person get in?” put in Whyland quickly. “That
is, if both Waters and young Ted are telling the truth. Waters is a
good man, and I haven’t the least reason to suspect him. But it’s
just possible that he was dozing somewhere between eleven and
twelve, and that Ted came home before he said he did. His father
wouldn’t be surprised to see him, and he might have walked up
behind him and jabbed the needle in. Then, when his father found
out what he’d done, he got a bit of caustic potash from somewhere
and clapped it on. I know there are lots of difficulties, but at least it’s
possible. At all events, I can’t think of another alternative to the
suicide theory.”
“The case is extraordinarily puzzling,” said Mr. Ludgrove
sympathetically. “If you feel disposed to discuss it, Inspector, I
suggest that you do so in comfort over at my place. I can make you
a cup of cocoa, or, if you prefer it, I can supply you with something
stronger. I always kept a bottle of whiskey in reserve for poor Mr.
Copperdock.”
“Well, it’s very good of you, Mr. Ludgrove,” replied Whyland
gratefully. “What’s the time? After two? I want to stay about here
until it’s light. I’ll just tell Waters where I’m to be found in case he
wants me. Then I shall be very glad to accept your kind hospitality.”
He left the room and returned after an absence of a couple of
minutes. “I can’t make it out,” he said. “Waters swears he never had
his eyes off the place. Still, it won’t do any harm to make enquiries
into young Ted’s movements and verify his statement. It beats me,
but then everything Copperdock did was a puzzle. His name seemed
to crop up in connection with each of these deaths, somehow. Then
there was that yarn about the black sailor, the counter which he said
he found on his bed, and now his amazing death. Well, I’m ready to
go across if you are, Mr. Ludgrove.”
The two passed through the shop into the road. As they crossed
it, Mr. Ludgrove uttered an exclamation of surprise. “Why, the door
of my shop is open!” he said. “I must have forgotten to shut it in my
haste when Ted Copperdock came over for me.”
“Let’s hope no inquisitive visitor has been in to have a look round
while we’ve been over the way,” replied Whyland.
Mr. Ludgrove smiled. “He will have found very little of value to
reward him if he has,” he said. “No, I’m not afraid of burglars. In any
case, it’s a very old-fashioned lock which anyone could force without
any difficulty.”
They had reached the door by now, and Mr. Ludgrove pushed it
open. “Come along, Inspector, we’ll go into the back room,” he said,
leading the way.
At the door of the inner room he paused, and switched on the
light. At a first glance, the room appeared to be exactly as he had
left it to answer Ted’s urgent summons. Then suddenly he clutched
Whyland’s arm, and pointed straight in front of him with a shaking
finger.
On the mantelpiece, propped conspicuously against the clock so
that it could not fail to attract attention, was a white bone counter,
upon which the figure VII had been carefully traced.
Part II.
The Criminal
Chapter XIII.
Enter Dr. Priestley
That eccentric scientist, Dr. Priestley, sat in his study on the
Monday morning following the death of Mr. Copperdock, busily
engaged in sorting out a mass of untidy-looking papers. Most of
them he tore up and placed in the waste-paper basket by his side; a
few he glanced at and put aside. The April sun lit up the room with a
pale radiance, lending an air of Spring even to this dignified but
rather gloomy house in Westbourne Terrace.
Dr. Priestley was thus engaged when the door opened and his
secretary, Harold Merefield, came into the room. There was an air of
heaviness about both men, the old and the young, as though the
Spring had not yet touched them, and Winter held them still in its
grip. One might have guessed that some absorbing work had
monopolized their energies, leaving them no leisure for anything but
the utmost concentration. And one would have guessed right. For
the last six months Dr. Priestley had been engaged upon the writing
of a book which was to enhance his already brilliant reputation. Its
title was Some Aspects of Modern Thought, and in it Dr. Priestley
had, with his usual incontrovertible logic, shattered the majority of
the pet theories of orthodox science. It was, as the reviews were to
say, a brilliant achievement, all the more entertaining from the vein
of biting sarcasm which ran through it.
When Dr. Priestley settled down to writing a book, he
concentrated his whole attention upon it, to the exclusion of
everything else. He allowed nothing whatever to distract his mind,
even for a few minutes. He lived entirely in his subject, refusing
even to read the newspapers, except certain scientific periodicals
which might happen to contain something relevant to the work he
had in hand. As he expected his secretary to follow his example, it
was hardly to be wondered at that both of them looked jaded and
worn out.
“I took the manuscript to the Post Office myself, sir,” said Harold
Merefield listlessly. “Here is the registration receipt.”
“Excellent, my boy, excellent,” replied the Professor, looking up.
“So the work is finished at last, eh? I have been destroying such
notes as we shall not require again. The rest you can file at your
leisure. Dear me, you look as if you needed a change of occupation.”
He stared at his secretary through his spectacles, as though he
had seen him that morning for the first time for many months. “Yes,
I think we both need a change of occupation,” he continued. “I feel
that I should welcome some enticing problem, mathematical or
human. It is time we stepped from our recent absorption back into
the world. Let me see. What is the date?”
“April 28th, sir,” replied Harold with a smile. He knew well enough
that the Professor would have accepted any other day he chose to
mention.
“Dear me! Then the world is six months older than when we
retired from it. No doubt many interesting problems have arisen in
the interval, but I fear that their solutions lie in other hands than
ours. By the way, when does our friend Inspector Hanslet return
from America?”
Harold turned to one of the big presses which lined the walls of
the room, and took from it a folder marked “Inspector Hanslet.” He
consulted this for a moment, then looked up towards his employer.
“At the end of this month, sir. There is no definite date mentioned. I
dare say he is in London already.”
“Perhaps so,” agreed the Professor. “It does not really matter. My
thoughts turned to him naturally, as to one who has in the past
supplied us with some very satisfactory problems. Well, we must be
patient, my boy. I have no doubt that we shall very soon succeed in
finding some congenial work with which to occupy our minds.”
He returned to the business of sorting his papers, while Harold
sat down at the table reserved for his use, thankful to be able to do
absolutely nothing for a few minutes. His idea of a change of
occupation was not to plunge at once into some abstruse
mathematical investigation which would involve him in the writing up
of endless notes. If only Hanslet would come back and divert the
Professor’s thoughts into some other channel! But of Hanslet, since
he had departed for New York during the previous year to co-
operate with the American police in running to earth a gang of
international swindlers, nothing had been heard.
Inspector Hanslet was rapidly becoming the foremost figure at
Scotland Yard. He was a man who, without being brilliant, possessed
more than the usual quickness of perception. He could, in his own
phrase, see as far through a brick wall as most people, and to this
attribute he added an agility of mind remarkable in a man whose
training had been of a stereotyped kind. Early in his career he had
become acquainted with Dr. Priestley, and the Professor, to whom a
problem of any kind was as the breath of his body, had since
encouraged him to come to Westbourne Terrace and discuss his
difficulties. To many of these the Professor’s logical mind had
suggested the solution. Since he refused to allow his name to be
mentioned, the credit for his deductions descended upon Hanslet. As
a matter of fact, the authorities knew very well how matters stood,
and Hanslet was always employed upon those cases which promised
to be complicated, since it was an open secret that he could call
upon the advice and assistance of Dr. Priestley.
It was evident that the sudden reaction of having nothing to do,
after his unremitting labours of the past six months, was having an
unfavourable effect upon Dr. Priestley’s temper. He roamed about the
study, pulling out a file from time to time, and finding fault with
Harold because some item did not come immediately to his hand. It
was not until it was time to dress for dinner that he desisted from
this irritating occupation. And even at dinner he was silent and
morose, obviously seeking in vain for some new interest which
should occupy his restless thoughts. But hardly had he and Harold
finished their coffee, which they always had in the study after dinner,
than Mary the parlourmaid opened the door softly. “Inspector
Hanslet to see you, sir,” she announced.
The Professor turned so abruptly in his chair as seriously to
endanger the coffee cup he was holding. “Inspector Hanslet!” he
exclaimed. “Why show him in, of course. Good evening, Inspector, it
was only this morning that Harold and I were speaking of you. I
hope that you enjoyed yourself in America.”
“I did indeed, Professor,” replied Hanslet, shaking hands warmly
with Dr. Priestley, and nodding cheerily to Harold. “Not that I’m not
very glad to be home again; one’s own country’s best, after all. I
landed at Southampton last Wednesday.”
“And now you have come back to tell us of your experiences,”
said the Professor. “I am sure we shall be most interested to hear
them. Did you succeed in your object?”
“Oh, yes, we rounded them up all right,” replied Hanslet. “My
word, Professor, you ought to go over to New York and see the
things the fellows do over there. As far as scientific detection goes,
they’ve got us beat to a frazzle. You’d appreciate their methods. And
they’re a cheery crowd, too. They gave me no end of a good time
while I was over there.”
“Well, sit down, and tell us all about it,” said the Professor,
motioning Hanslet towards a comfortable chair. “You will relieve the
tedium I am feeling at having nothing to do.”
Hanslet sat down, and, as he did so, looked enquiringly at the
Professor. “You say you’ve nothing to do, sir? Well, I’m very glad to
hear that. The truth is that I didn’t come here to tell you my
experiences. As a matter of fact, I meant to take a month’s leave
when I got back, but the Chief asked me to wait a bit and take over
a case which has been puzzling the Yard for several months. And I
wanted to ask your advice, if you would be good enough to listen.”
The Professor rubbed his hands together briskly. “Excellent,
excellent!” he exclaimed. “I told you this morning, Harold, that a
problem was bound to turn up before long. By all means tell me your
difficulties, Inspector. But let me beg of you to keep to facts, and not
to digress into conjecture.”
Hanslet smiled. The Professor’s passion for facts was well-known
to him from past experience. “Well, I expect you know as much
about it as I do,” he began. “Ever since Tovey the greengrocer was
killed last November, there’s been a lot in the papers——”
But the Professor interrupted him. “I should perhaps have
explained, Inspector, that since last October I have scarcely opened
a newspaper. My whole mind has been concentrated upon a task
which is now happily finished. The name of Tovey the greengrocer
is, I regret to say, utterly unfamiliar to me. I should be glad if you
would treat me as one who has only lately reached this world from
the planet Mars, and give me the facts without presuming that I
have any previous knowledge of them.”
“Very well, Professor,” replied Hanslet. “You must have heard of a
series of deaths under peculiar circumstances which have occurred
in Praed Street, not half a mile away from here? Why, I read about
them in New York! They caused a great sensation.”
“I am not concerned with popular sensations,” said the Professor
coldly. “I admit that some rumours of such happenings penetrated
the isolation with which I have endeavoured to surround myself, but
I dismissed them from my mind as likely to introduce a disturbing
factor. I repeat that you had better repeat the facts, as briefly as
possible.”
“Very well, Professor, I will tell you the story exactly as it was told
to me at the Yard,” replied Hanslet. “You will be able to see how
much is fact and how much conjecture. As I was not on the spot
myself, I cannot vouch for the details. Will that do?”
The Professor nodded, and turned to Harold. “Make a note of the
names and dates mentioned by Inspector Hanslet,” he said. “Now,
Inspector, you may proceed.”
Hanslet, whose memory for names and facts was rarely at fault,
recounted as briefly as he could the course of events from the
murder of Mr. Tovey in November, to the finding of Martin’s body in
the cellar of Number 407, in January. The Professor interrupted him
now and then to ask a question, but in the main he allowed him to
tell the story in his own way. When he had finished, and the
Professor had expressed himself satisfied, Hanslet continued.
“The man who’s been in charge of the case is a fellow called
Whyland, keen enough on his job, but a bit lacking in imagination. I
had a chat with him yesterday, and he confessed that he was
completely at the end of his tether. Up till last Saturday evening, he
told me, he was pretty sure that he could lay his hand on the
criminal, but that night something happened which entirely upset his
calculations.”
“What was that?” enquired the Professor, who was listening
intently.
“Why, for one thing, the man whom he suspected of the murders
has been killed,” replied Hanslet. “Not that there was anything
amazing in that, for he seems to have been a trifle unbalanced in
any case, and his death may possibly have been due to suicide. No,
what altogether upset Whyland’s apple-cart was that another man
received a counter, some time after the death of the man whom
Whyland suspected of delivering them.”
“It is remarkable how frequently hypotheses founded upon pure
conjecture are upset by one simple fact,” remarked the Professor
acidly. “Now, what was the name of this man whom Whyland
suspected, and who so inconsiderately spoilt the theory by his
premature death?”
“Samuel Copperdock,” replied Hanslet, turning to Harold, who
wrote the name on his pad.
“Copperdock?” repeated the Professor. “An unusual name, and
yet I seem to have heard it before in some connection. Copperdock,
Copperdock! Let me think——”
“You’ve probably seen the name above his shop, Professor,” said
Hanslet. “He was a tobacconist in Praed Street. Or you may have
seen it some months ago in the paper. He was a witness at the
inquest on Tovey, who was the first man murdered.”
But the Professor shook his head. “No, if my memory serves me,
I heard the name many years ago, in some connection which
escapes me for the moment. However, the point would not appear to
have any importance. I must apologize for interrupting you,
Inspector. You were saying that another man received a counter
after this man Copperdock’s death, but I do not think you mentioned
his name?”
“Ludgrove. Elmer Ludgrove,” said Hanslet. “Rather an interesting
personality, from what Whyland tells me. He keeps a herbalist’s
shop, and is a bit of a character in his way. He’s a man of some
education, between fifty and sixty, a very dignified old boy with a
striking white beard, which I expect is a bit of an asset in his trade.
He doesn’t say much about himself, but does a lot of good in his
own quiet way. All the poorer people in the neighbourhood come to
him if they’re in any sort of trouble, and he freely admits he hears a
good many secrets. Whyland thought he would be a useful chap to
get on the right side of, and often used to drop in to see him. He
says he got more than one valuable hint from him. He was also
pretty certain that this chap Ludgrove shared his suspicions of
Copperdock, but he would never say so outright. You see,
Copperdock was a friend of his.”
The Professor nodded. “I see,” he said. “And it was this Mr.
Ludgrove who received the counter you say?”
“Yes, and, what’s more, Whyland was with him when he found it.
The poor old boy was terribly shaken for the moment, Whyland says,
but after a bit he pretended to treat it as a joke. I’ve seen him since,
and he’s pretty plucky about it, knowing as he does that everybody
who has received one of these infernal numbered counters has died
a sudden death. He says that he is an old man, anyhow, alone in the
world and with only a few more years to live in any case, so that his
death will be no great blow to anybody.”
“A most philosophic attitude,” agreed the Professor. “But to return
to Mr. Copperdock, I should like to hear the circumstances under
which he met his death.”
Hanslet related the events of the previous Saturday night in
considerable detail, up to the time when Whyland and Ludgrove
entered the latter’s sanctum. “There’s not much more to add,” he
continued, “except that the doctor’s suspicions were confirmed as to
the poison. The Home Office people examined the fragment of
broken needle, and I heard this afternoon that they found traces of
a remarkable virulent synthetic alkaloid. You’ll know what that is
better than I do, Professor.”
“Yes, I know,” replied the Professor grimly. “I have reason to. It
was with one of these synthetic alkaloids—there are a number of
them—that Farwell tipped the spines of the hedgehog to which I so
nearly fell a victim.¹ You remember that incident, I dare say?”
“I do, indeed,” said Hanslet warmly. “What’s more, the Home
Office people say that a dose of the stuff would produce almost
immediate paralysis, and death within a few minutes. The
incrustation was potassium carbonate all right, almost certainly the
result of putting caustic potash on the place. But that only makes
the business more puzzling. If Copperdock poisoned himself, how did
he have time to apply the caustic potash before he was paralysed? If
someone else did it, why should they apply the caustic, and how did
they get in and out of the house? Remember, Whyland’s man Waters
had the place under observation all the time.”
“Then you are inclined to favour the theory of suicide?” asked the
Professor.
“On the whole, yes,” replied Hanslet. “Oh, by the way, I forgot to
tell you that soon after daylight Waters found the syringe, with the
other part of the needle still in it, by the side of the road under
Copperdock’s window. There had been a heavy shower of rain about
half past three, and the syringe was covered with mud and filth. The
analysts could not find any traces remaining of the poison, but the
end of the needle proved that it was the one that had been used.
That points to suicide, a murderer wouldn’t chuck away his weapon
like that where anyone could see it.
“Besides, if you come to think of it, suicide fits in best with what
we know. It is a fact that Copperdock’s mind was to some extent
unhinged. He declared that he met the black sailor, when a reliable
witness declares that no such person was about. In fact, the only
person besides Copperdock who seriously claims to have seen this
black sailor is a degenerate youth who is also a convicted pick-
pocket. It is highly probable that the counters were numbered, and
the envelopes containing them typed in Mr. Copperdock’s office.
Whyland assures me that the only link between the victims was
Copperdock, not in any definite form, certainly, but still definite
enough to make the coincidence remarkable. I am inclined to believe
that Copperdock was at the bottom of it all somehow. My difficulty
will be to prove it.”
“You think, I gather, that this Mr. Copperdock suffered from a
peculiar form of homicidal mania, which finally culminated in his
taking his own life?” suggested the Professor. “I admit that such
cases are not unknown, but the theory involves you in many
difficulties. I mention only one of them, the first that occurs to me.
Where did he obtain this synthetic alkaloid? These substances are
not articles of commerce, they are not, so far as I am aware, used in
medicine. They are only produced experimentally in research
laboratories. Farwell had a well-equipped laboratory, as you probably
remember, which accounts for his use of such a poison. But how
could a man in Copperdock’s position procure it?”
Hanslet shrugged his shoulders. “I don’t know, Professor,” he
replied. “I confess that I turn to the theory of Copperdock as the
murderer because it seems to present fewer difficulties than any
other. The whole thing seems to me to involve a mass of
contradictions, whichever way you look at it. It’s for that very reason
I came to see you, Professor. But you must at least admit that
madness in some form must be responsible. What rational motive
could there be for the murder of half a dozen men entirely
unconnected with one another, and whose deaths could be of no
possible benefit to the murderer?”
“I am prepared to admit nothing until I have further examined
the facts,” replied the Professor severely. “Now, Harold, will you read
me your notes upon the first murder? Thank you. I should like all
details relating to Mr. Tovey, please, Inspector.”
It was long past midnight before they reached the end of the
catalogue, and the Professor was satisfied that he knew everything
which Hanslet could tell him.
“You will, of course, let me know if any fresh facts come to light,”
he said, as Hanslet rose to take his leave. “Meanwhile, I will consider
the matter. If I come to any definite conclusions I will let you know.
Good-night, and pray accept my most sincere thanks for presenting
me with a most absorbing problem.”
¹ See The Ellerby Case by John Rhode. ↩︎
Chapter XIV.
The Morlandson Trial
The Professor came down to breakfast next morning looking
even more weary than on the previous day. Harold, looking at him
anxiously, guessed that he had hardly slept at all during the night.
Some absorbing train of thought, whether started by Hanslet’s story
of the previous evening or not, had taken possession of his brain.
But, in spite of his weariness, there was a queer gleam in the
piercing eyes behind the powerful spectacles, which Harold knew
from past experience to be the light of battle.
“I have some work for you to-day, my boy,” he said, as soon as
the meal was over. “I want you to go to the British Museum and look
up the reports of the trials for murder at the Old Bailey during the
first ten years of the present century. Among them you will find the
trial of a doctor for the murder of one of his patients by giving him
an overdose of morphia. I believe that the doctor’s name began with
an M, and I fancy that his patient had a title. More than this I cannot
tell you, my memory, I regret to say, is not what it used to be. I
want you to make a précis of that trial and of the sentence.”
“Very good, sir,” replied Harold, and forthwith started on his
quest. He could not guess the purpose for which the Professor
required this information, but it could obviously have nothing to do
with these intriguing murders in Praed Street which Hanslet had
described. Unless, perhaps, the Professor had seen some parallel
between the methods of the unknown criminal and those of this
vague doctor whose name began with an M. You never could predict
the direction from which the Professor would approach a problem.
All that you could be certain of was that it would be different from
the one you anticipated.
Arrived at the Museum, where he was a frequent visitor on
similar errands, he went carefully through the index to the Law
Reports. It was not until he came to the year 1906, that he met with
anything which corresponded to the data which the Professor had
given him. Then he found a reference to the trial of one Doctor
Morlandson for the murder of Lord Whatley. This must be the case
to which his employer had referred. He turned up the records, and
proceeded to make a careful abstract.
It appeared that Lord Whatley had been a man of middle age,
and of some considerable wealth. Dr. Morlandson was his regular
medical attendant, and in 1905 he had been compelled to warn his
patient that he was suffering from cancer, and that though an
operation might be successful, there was grave doubt that it would
permanently remove the source of the trouble. However, Lord
Whatley consented to undergo the operation. He was removed to a
nursing home, and a specialist was called in. The patient went
through the ordeal satisfactorily, and after a while he returned
home.
But, by the beginning of the following year, the symptoms
reasserted themselves, and Dr. Morlandson informed his patient,
who insisted that he should be told the truth, that nothing more
could be done, and that Lord Whatley had nothing to look forward to
but perhaps a year or two of suffering. The relations between the
two men were rather those of close friendship than of doctor and
patient, and, subsequent to Dr. Morlandson’s pronouncement, they
saw a great deal of one another. Morlandson devoted as much time
as he could spare from his practice to sitting with Lord Whatley, who
was a childless widower and did not encourage the visits of his
friends and relations.
By the end of February it appeared that the disease was
progressing even more rapidly than Dr. Morlandson had anticipated.
He administered frequent injections of morphia, and his patient was
rarely conscious. Morlandson continued to spend the greater part of
his time with him, and in Lord Whatley’s brief intervals of
consciousness his doctor, and the nurses who had been called in,
were the only people he spoke to.
He died early in March, in the presence of Dr. Morlandson and
one of the nurses, without regaining consciousness. A cousin of Lord
Whatley’s, who happened to be his nearest relative, was in the
house, and Morlandson informed him that he would return home
and bring the necessary certificate with him later in the day.
Morlandson, who lived about a mile away, started to walk home.
When he had almost reached his own house, he heard a sound of
confused shouting, and saw a runaway horse attached to a milk-cart,
coming towards him. Without a moment’s hesitation he rushed for
the horse’s head, and had almost succeeded in stopping him, when
he slipped and fell. One of the horse’s hoofs struck him on the head
and he was left unconscious on the road.
The spectators of the accident picked him up, and he was carried
into his own house. A colleague was summoned, and declared that
he was suffering from severe concussion. This diagnosis proved
correct, and Morlandson lay in a state of semi-consciousness for
nearly a week. On his recovery, he found the house in possession of
the police.
Lord Whatley’s cousin, hearing of the accident to Dr. Morlandson,
and learning that he could not possibly attend to his duties for some
time to come, was at a loss for the want of a death certificate. He
therefore sent for another doctor—not the man who was attending
Morlandson—and asked him to sign the certificate. This the doctor
would have done, had not one of the nurses, whom Morlandson had
reprimanded for some breach of duty, made some vague insinuation
that everything was not as it should be. The doctor insisted upon
examining the body, and as a result of this examination he
communicated with the authorities. A post-mortem was held, and
Lord Whatley was proved to have died of an overdose of morphia.
The experts gave it as their opinion that the deceased would not
have died of the disease from which he was suffering for another
year at least. A warrant was immediately issued for Dr. Morlandson’s
arrest.
When Lord Whatley’s will came to be read, it was found that he
had left the sum of ten thousand pounds to Morlandson, conditional
upon his being his medical attendant at the time of his death. This
bequest was contained in a codicil executed early in February.
Morlandson came up for trial at the Old Bailey in July. The
prosecution alleged that the codicil disclosed the motive for the
murder, and submitted that Morlandson, fearing lest Lord Whatley
should change his doctor before he died, had made certain of
securing the legacy by poisoning him. They pointed out that, but for
Morlandson’s accident, he would have been able to certify cancer as
the cause of Lord Whatley’s death, and no suspicion would have
been aroused.
Dr. Morlandson’s counsel put in a very striking defence. In effect,
he pleaded guilty to the act of poisoning, but affirmed that this was
done at Lord Whatley’s express command. He had already suffered
considerably and undergone an ineffectual operation, and refused to
contemplate the further agony to which he was condemned. As soon
as Morlandson had informed him that his case was hopeless, he had
begged him to put a end to his sufferings at once, pointing out that
such a course would cause no grief or inconvenience to anyone.
Morlandson had at first refused, but at last, upon the solemn
assurance of Lord Whatley that he would find some means of
committing suicide unless his wishes were complied with, he
consented to inject morphia in increasing doses. This Lord Whatley
agreed to, and whenever he was conscious Morlandson begged him
to reconsider his determination. Finally, knowing that the disease
was incurable, and that the man he cared for as his friend could only
endure months of suffering under his very eyes, he bade him
farewell and administered the fatal dose. The news of the bequest
came as a complete surprise to him.
Morlandson’s defence raised in an acute form a controversy
which had been going on for many years. Many people held that he
was completely justified in his action, that his offence was purely
technical, and that at the most it merited a short term of
imprisonment. But the jury, in spite of a hint from the judge, found
Morlandson guilty of murder and refused to add a rider
recommending him to mercy. Sentence of death was duly
pronounced, but the Home Secretary, the Court of Criminal Appeal
not being then in existence, ordered a reprieve, and the sentence
was commuted to one of twenty years’ penal servitude. Morlandson’s
wife, to whom he was deeply attached, died before a year of it had
expired.
This was the substance of the notes which Harold Merefield
brought back to Dr. Priestley. The latter read them through carefully,
then gave them back to his secretary. “Yes, I thought that I was not
mistaken,” he said. “The facts of the case come back to me very
clearly now. It made a considerable sensation at the time, owing to
the principle involved. Right or wrong, Morlandson was acting in
accordance with his lights. His evidence, I remember, was given with
an air of passionate conviction. This Lord Whatley was his friend,
and he had saved him from suffering at the expense of twenty years
of his own life. I wonder whether he survived his sentence? It would
be most interesting to learn.”
The Professor relapsed into his favourite attitude of thought, his
eyes fixed upon the ceiling, his hands, with the tips of the fingers
touching, laid upon the table in front of him. He remained like this
for many minutes before he spoke again.
“It would be so interesting that I feel impelled to take steps to
discover the facts,” he said. “After lunch I shall visit the record
department of Scotland Yard. While I am away, you can complete
the filing of those papers I gave you yesterday, relating to the work
which we have just completed.”
Harold received these instructions without any great enthusiasm.
He was not greatly interested in the case of this Dr. Morlandson,
since it had occurred so many years ago and could have no possible
bearing upon any problem of the present day. In his recollection of
this forgotten trial the Professor seemed to be neglecting entirely the
problems presented by the murders in Praed Street. Perhaps he had
decided that they were not worthy of his notice. It was not every
problem submitted to him which appealed to him sufficiently to
induce him to devote his energies to its solution.
He spent the afternoon in the study, working half-heartedly and
awaiting the Professor’s return. But it was not until nearly dinner-
time that his employer came in, and then he could see by his
expression that the result of his search had in some way
disappointed him. Dinner was passed in almost complete silence,
and the two returned once more to the study.
“I have discovered the subsequent history of Dr. Morlandson,”
announced the Professor abruptly, as soon as he had finished his
coffee. “I will recount to you the result of my researches at Scotland
Yard. You can make notes of them, and file them with your précis of
his trial.”
Harold produced pencil and paper, and the Professor proceeded
to give an account of how he had spent the afternoon. After some
delay the authorities at Scotland Yard, who were always anxious to
carry out any of Dr. Priestley’s requests, even though they were
ignorant of the motive behind them, had found the record of
Morlandson’s career after his sentence. He had been sent to
Dartmoor, and had served his time there. He had been released on
licence in 1920, having undergone fourteen years of his sentence.
He had then remained for a short time in London, arranging his
affairs, but had not communicated with anybody but his solicitor, to
whom he had expressed his intention of spending the rest of his life
in the most complete seclusion, and devoting himself to chemical
research, for which he had always had a bent during the period in
which he was in practice.
Before the catastrophe which had overtaken him, Morlandson
had been a tall, spare man, clean-shaven, and with carefully brushed
dark hair. Upon his release he had developed a slight stoop, and
although he was still clean-shaven and smart in his appearance, his
hair had gone nearly white. He told his solicitor that he knew he had
only a few years longer to live, but that he hoped that during that
period his researches would confer some benefit upon suffering
humanity. He proposed to commence them as soon as he could find
a suitable spot for the purpose, where he could live entirely alone.
A few weeks after his release, he found a half-ruined cottage
which answered to his requirements, situated in a peculiarly desolate
part of the Isle of Purbeck, in Dorsetshire. He took up his residence
here, repaired the cottage, and added to it a laboratory, built of
concrete. Under the terms of his licence he was compelled to report
to the police, and they kept an eye upon his movements. They might
have saved themselves the trouble. Once he was established in his
cottage, his furthest excursion was to Corfe Castle, the nearest
town, to obtain supplies. He lived entirely alone, and invariably
walked across the heath to and from his cottage. But, even while
living this hermit existence, he was always carefully dressed and
shaved. He made no attempt to conceal his identity, but called
himself Mr. Morlandson, having dropped the prefix “Doctor.” He had,
of course, been struck off the register, and could not have practised
as a doctor even had he desired to do so.
The local superintendent to whom he reported conceived a liking
for him, and occasionally walked across the heath to visit him. He
invariably found him at work in his laboratory, which was plentifully
stocked with chemicals of various kinds. He would never allow
smoking in the laboratory, for, as he pointed out to the
superintendent, the substances with which he was experimenting
were highly inflammable, and there was consequently grave risk of
fire unless proper precautions were taken.
One night, rather more than a year after Morlandson’s release,
flames were seen from Corfe Castle across the heath in the direction
of his cottage. The superintendent leapt on his bicycle, and dashed
off to the scene. When he arrived, he found the laboratory burning
like a furnace, and quite unapproachable. The flames had caught the
cottage, which was by then past saving, especially as the only
available water supply was from a well fitted with a small bucket.
The superintendent, at considerable risk to himself, managed to
enter the sitting-room of the cottage, but could see no trace of
Morlandson.
By morning the fire had burnt itself out. The cottage had been
completely destroyed, only two or three feet of the outer walls
remaining. The laboratory, being built of concrete, had fared rather
better. The greater part of the walls remained, as did the steel door,
which formed the only entrance. The place had no windows, but had
been lighted from above through sky-lights in the roof. These and
the roof itself had completely disappeared. The iron door was found
to be locked upon the inside.
When it had been broken down, the interior of the laboratory
showed how fierce the fire had been. Every trace of wood had been
consumed, and solid metal fittings had been melted into
unrecognizable shapes. Among the debris on the floor lay a charred
human skeleton, upon one of the fingers of which was a half-melted
gold ring, of which enough remained for the superintendent to
identify it as having been habitually worn by Morlandson. The
remains of the unfortunate man were huddled up by the door, the
key of which was in the lock. It was clear that Morlandson had tried
to make his way out when the fire broke out, but had been
overcome by the fumes of the burning chemicals before he could
achieve his purpose. He had been in the habit of locking the door in
order to secure himself from interruption.
“You have made notes upon this?” asked the Professor. “Good.
File them away. I confess that there are many things about this man
Morlandson which I do not yet understand. I was able to supplement
your account of the trial by an examination of the original records,
which I was allowed to make. These gave me considerable food for
thought. I believe that, through a pure accident, I have stumbled
upon one of the most curious occurrences of modern times. I can,
as yet, only conjecture, and so far my conjectures are wholly
unsupported by fact. Much research will be necessary before these
facts can be established, and it is possible that I may not be spared
for a sufficient time to carry out this research.”
“Not be spared, sir!” exclaimed Harold, startled by the grave tone
of the Professor’s voice. “Why, you have many years before you yet,
I hope.”
“Death comes to us all, sooner, perhaps, than we expect,” replied
Dr. Priestley. “And I feel, this evening, that death may be closer to
me than I have supposed. Ah, do I hear someone in the hall?”
With a nervous movement, entirely foreign to him, Dr. Priestley
rose from his chair and stood facing the door. Harold, with a queer
feeling of expectation, walked towards it and opened it. In the hall
stood Inspector Hanslet, handing his coat and hat to Mary.
“Good evening, Mr. Merefield, I thought I’d look round and see if
the Professor had any information for me,” he said. “May I see him?”
“Yes, come in by all means,” replied Harold, with a sudden sense
of relief. “But I shouldn’t stay too long, if I were you. He’s rather
tired and nervy to-night.”
Hanslet nodded, and Harold led the way into the study. “It’s
Inspector Hanslet, sir,” he said.
The Professor appeared to have entirely recovered his usual
equanimity. “Ah, good evening, Inspector,” he said blandly. “I half
expected that you would be round this evening. I am very glad to
see you.”
“I thought I would come round, on the chance that you had
some hint to give me,” replied Hanslet. “I can’t make head or tail of
the business I told you about last night. The more I think about it,
the more puzzling it seems. It’s the utter lack of motive that makes it
all so inexplicable.”
“I believe, mind, I say only that I believe, that I have discovered
the motive,” said the Professor quietly.
“You have!” exclaimed Hanslet excitedly. Then, seeing the slow
movement of the Professor’s head, he smiled. “I know you won’t tell
me until you are certain,” he continued. “But at least tell me this. Are
there likely to be any more of these mysterious deaths?”
“There will be one more, unless I am able to prevent it,” replied
the Professor.
Chapter XV.
The Bone Counters
Mr. Ludgrove, as Hanslet had said to Dr. Priestley, bore the shock
of the finding of the numbered counter extremely well. He had
refused to make any alteration in usual habits, and it was with the
greatest difficulty that Whyland could persuade him to allow a
constable to sleep in the house at night.
“I can assure you that this mysterious warning does not terrify
me,” he had said. “I am an old man, and death cannot be far off in
any case. I am not sure that I should not prefer a violent end to
some lingering illness which might leave me helpless for months
before it killed me. But, if you think that by keeping a close watch
over me you can gain some clue to the distributor of these counters,
by all means do so.”
He was in this frame of mind when Hanslet came to see him on
the Sunday afternoon. Whyland brought him round and introduced
him, and Mr. Ludgrove welcomed him with his usual courtesy.
“I have heard of you, Inspector Hanslet, and I am indeed proud
to make your acquaintance. Sit down, and make yourself
comfortable.”
“Thank you, Mr. Ludgrove,” Hanslet replied. “I thought you
wouldn’t mind my coming to have a chat with you. Whyland here
has told me all about these queer happenings in this street of yours,
and of the help which you have been to him.”
“I am afraid that I have been of very little help,” said Mr.
Ludgrove with a smile. “Inspector Whyland has been kind enough to
appreciate beyond their value any suggestions I have made.”
“Well, that’s as may be,” replied Hanslet. “Now, Mr. Ludgrove, I
am going to ask for further assistance on your part. You know as
much about these counters as I do. They seem to have been sent,
so far, to six men, all of whom have died shortly after they received
them. Whyland tells me that he has utterly failed to establish any
connection between these men. Except for the fact that Tovey and
Copperdock were close friends, they all seem to be comparative
strangers to one another, and have never been associated in any
common enterprise. You see what I mean, of course?”
“I do, indeed. In fact, Inspector Whyland and I discussed the
point, long ago. It might be possible to imagine a motive for the
murder of a group of men who were inspired by a common motive
or who belonged to some common society. The difficulty is,
assuming that the agency which compassed their deaths was the
same in each case, to imagine a motive for the actions of that
agency.”
“Exactly!” exclaimed Hanslet warmly. “I see that you appreciate
my point as clearly as I do myself. But now we have a fresh line of
investigation. You yourself are added to the list of those who have
received the counter. Can you explain why you should have been
singled out?”
Mr. Ludgrove shook his head. “As you may suppose, the subject
has occupied my thoughts ever since I found the counter,” he
replied. “I am an old man, as I have said before, and for the last
twenty years or more I have led a retired life, retired, I mean, in the
sense that I have taken no part in the affairs of the world. I have
had enemies as well as friends; few men who have reached my
years could say otherwise. But most of the contemporaries of my
youth are dead, and in any case I do not believe that any of the
enemies I may have made would be so vindictive as to seek my life.”
“Let us look at it another way, then,” said Hanslet. “Can you
imagine any way in which you, in common with the six men who
have already died, could have made an unconscious enemy?”
“I cannot,” replied Mr. Ludgrove. “Of those six men, I knew only
two personally, Mr. Copperdock fairly well, and Mr. Colburn slightly.
Both of these I have known only since I came to live in Praed Street,
five years ago. Tovey, I had heard Copperdock speak of. The name
of Richard Pargent, I had seen mentioned in the newspapers. The
other two were complete strangers to me. I cannot imagine how we
could have committed any act in common which would draw down
upon us the vengeance of a single assassin.”
“Then you do not believe that these deaths are the work of a
single assassin, Mr. Ludgrove?” enquired Hanslet with interest.
“Not of a single man, acting upon any rational motive,” replied
Mr. Ludgrove. “Even in the brain of a homicidal maniac there is
usually traceable some dim guiding principle. He either conceives a
hatred for a certain class of person, or he kills indiscriminately,
usually selecting the people nearest to hand. In this case the
selection was anything, but indiscriminate. Mind, I am assuming for
the moment, as apparently you are yourself, that the death of all six
was the direct sequel of their receipt of a numbered counter. If you
adopt the theory that a single man is responsible, you may as well
believe in the existence of the black sailor.”
“I am afraid that we are already committed to him,” said Hanslet
with a smile. “You see, we offered a reward for him, and it would
never do for the police to admit that they had offered a reward for a
ghost. Whyland, what is your honest opinion of this black sailor?”
“Entirely between ourselves and this most comfortable room, I
have never believed in his existence for a moment,” replied Whyland
readily. “But what could I do? That young rip, Wal Snyder, swore to
having seen him, and I couldn’t shake him.”
“Whether young Snyder saw him or not,” remarked the herbalist,
“your reward has made him a very real person to the poorer classes
of this district. One or other of my customers sees him every night,
usually during the hour which immediately follows the closing of the
public houses. And, as a rule, they come here hot-foot to tell me
about it.”
“I’m sorry, Mr. Ludgrove,” laughed Whyland. “I wouldn’t have
done it if I could have helped it. By the way, I suppose that you are
perfectly satisfied that none of these odd customers of yours know
anything about this business?”
“Perfectly,” replied Mr. Ludgrove. “They are a strange lot, I admit,
professing a code of morals which in some respects would shock the
conscience of a savage, and many of them are not above any petty
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Agriculturally Important Fungi for Sustainable

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Agriculturally Important Fungi for Sustainable Agriculture

Advances in Plant Microbiome and Sustainable Agriculture

Agriculturally Important Microbes for Sustainable

Recent Advancement in White Biotechnology Through Fungi

Advanced Intelligent Systems for Sustainable Development

Water Stress and Crop Plants A Sustainable Approach 2

Nutrient Dynamics for Sustainable Crop Production Ram

In Silico Approach for Sustainable Agriculture Devendra K.

Ajar Nath Yadav · Shashank Mishra

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made efforts to contribute these outstanding chapters. Each chapter is written by

Sirmour, Himachal Pradesh, India Ajar Nath Yadav

1 Agriculturally Important Fungi for Crop Protection�������������������������� 1

10 Fungi: A Bio-resource for the Control of Plant

Ahmed M. Abdel-Azeem Systematic Mycology Laboratory, Botany Department,

Rubee Devi Department of Biotechnology, Eternal University, Sirmour, Himachal

Pavidharshini Selvasekaran Food Science and Technology, School of Biosciences

Ajar Nath Yadav is an Assistant Professor (Sr. Scale)

Shashank Mishra is presently working as Scientist

Divjot Kour is currently working as Project Assistant

Management Sciences, Solan, Himachal Pradesh; M.Sc. in Microbiology from

Neelam Yadav is currently working on microbial

Anil Kumar is Director, Education, Rani Lakshmi

the Karnal bunt which is an economically important disease of wheat, he followed

Pavidharshini Selvasekaran and Ramalingam Chidambaram

P. Selvasekaran · R. Chidambaram (*)

© Springer Nature Switzerland AG 2020 1

1.2 The Role of Fungi as Biofertilizers

1.2.1 Advantages of Fungal Biofertilizers

• They are renewable sources of nutrients and sustain soil health.

1.3 The Role of Fungi as a Biocontrol Agent

List of agriculturally important fungal biofertilizer Reference

• Chaetomium spp. (C. bostrychodes, C. olivaceum)

Table 1.2 Fungal communities as biocontrol agents against plant species

1.4 Fungal Endophytes

understanding of the mechanism involved in the association of plants and fungal

1.4.1 Direct Mechanism

A wide spectrum of secondary metabolites is produced by fungal endophytes with

caused by this fungus was suppressed by the fungal endophytes. An endophytic

1.4.2 Indirect Mechanism

1.4.2.1 Production of Metabolites

Table 1.3 Bioactive compounds produced by fungal endophytes

1.4.2.2 Growth of Root and Shoot

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Sirmour, Himachal Pradesh, India Ajar Nath Yadav

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10 Fungi: A Bio-resource for the Control of Plant

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1.4 Fungal Endophytes

1.4.1 Direct Mechanism

1.4.2 Indirect Mechanism

1.4.2.1 Production of Metabolites

1.4.2.2 Growth of Root and Shoot