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Tulasi Satyanarayana
Sunil Kumar Deshmukh
Mukund V. Deshpande Editors
Advancing
Frontiers in
Mycology &
Mycotechnology
Basic and Applied Aspects of Fungi
Advancing Frontiers in Mycology
& Mycotechnology
Tulasi Satyanarayana
Sunil Kumar Deshmukh
Mukund V. Deshpande
Editors
Advancing Frontiers
in Mycology
& Mycotechnology
Basic and Applied Aspects of Fungi
Editors
Tulasi Satyanarayana Sunil Kumar Deshmukh
Division of Biological Science and Biotech & Management of
Engineering Bioresources Div
Netaji Subhas University of Technology The Energy and Resources Institute
New Delhi, Delhi, India New Delhi, Delhi, India
Mukund V. Deshpande
Division of Biological Sciences
CSIR-National Chemical Laboratory
Pune, Maharashtra, India
This Springer imprint is published by the registered company Springer Nature Singapore Pte Ltd.
The registered company address is: 152 Beach Road, #21-01/04 Gateway East, Singapore 189721,
Singapore
Foreword
The term ‘mycology’ dates back to the eighteenth century, and in its formative
years, the study of fungi was guided by reputed botanists like C. H. Persoon,
E.M. Fries and A. deBary. During the nineteenth and twentieth centuries, phenom-
enal progress in classical mycology took place, including descriptive morphology,
identification, classification and taxonomy of fungi primarily, due to the efforts of
botanists, despite the fact that fungi are not phylogenetically related to plants. The
reason for this essentially stems from the devastating fungal diseases in plants, nota-
bly the crop plants, causing serious problems in agriculture. There was an urgent
need to understand the aetiology of the destructive fungal diseases and devise suit-
able control measures to save the crops. As an example may be mentioned the stud-
ies in India by Prof. M.J. Narasimhan (1930) who collected and studied diverse
heterothallic strains of the destructive Phytophthora1 infection on areca nut palms
and devising fungicidal treatment schedules for controlling the disease in the rain-
forest areas of Karnataka (Mysore) state.
During the early years, traditional mycological knowledge facilitated the (a)
development of various oriental fermented foods, such as miso, tempeh and sake,
based on strains of fungi such as Aspergillus oryzae, Rhizopus oligosporus and oth-
ers (Hesseltine 1983), (b) differentiation of edible and poisonous mushrooms and
(c) understanding the aetiology of ergot poisoning that caused serious convulsions
due to consumption of the ergot sclerotia contaminating the grain which is con-
sumed as food.
During the twentieth century, several milestone advances in biological sciences
based on mycological studies can be recognized. As examples, we may mention the
following:
(a) The Nobel Prize-winning (1958) studies on Neurospora by G.W. Beadle and
E.L. Tatum which resulted in their discovery of how genes act by regulating
definite chemical events that in turn affected the development in organisms.
Based on ingenious selection of mutants deficient in the synthesis of single
growth factors, Beadle and Tatum concluded that mutations to genes affected
1
Recently, the genera Phytophthora and Pythium have been placed in Kingdom Chromista or
Kingdom Straminipila, distinct from Kingdom Fungi (Ho 2018).
v
vi Foreword
the enzymes of organisms. It is noteworthy that these studies provided the early
link between genetics and the upcoming new scientific field of molecular
biology.
(b) The discovery of truly anaerobic chytrids present in the rumen of herbivores
and playing a significant role in the digestion of the cellulosic feed is a mile-
stone discovery of the twentieth century. Classified under Neocallimastigaceae,
these chytrids with chitinous cell walls are truly anaerobes lacking mitochon-
dria, cytochromes and other biochemical features of the oxidative phosphoryla-
tion pathway. The group possesses organelles (hydrogenosomes) for a major
part of anaerobic energy metabolism (Nagpal et al. 2009).
Taxonomy and classification of fungi have been central to the mycological stud-
ies, and this has led to a well-established database for understanding fungal biodi-
versity. Classical taxonomy relied upon the morphological features of the spore
forms. With the advent of biochemical and molecular data becoming available, sev-
eral conceptual changes in fungal taxonomy have arisen. While giving an opportu-
nity to gain newer knowledge, these developments also pose several challenges and
opinion divides in their interpretation and implementation.
Classical taxonomy classified all the zoospore-forming fungi under Phycomycetes
considering their monophyletic origin from algae. Recent advances in biochemical
knowledge have established that the Chytridiomycetes2 with chitinous cell walls,
despite being zoospore-forming, are distinct from the rest of the zoospore-forming
fungi with β-glucan in their cell walls, and these are grouped together under the
Oomycota (as opposed to the Eomycota under which the Zygomycetes, Ascomycetes
and Basidiomycetes are classified). The term Phycomycetes as originally adapted is
no longer valid and, hence, abandoned from present-day mycological literature.
DNA sequencing and bar-coding of fungal species are major developments
which offer challenges to accepted morphology-based taxonomy. The nuclear ribo-
somal internal transcribed spacer (ITS) is widely used as a DNA bar-coding marker
to characterise the diversity and composition of fungal communities. Since the
1990s, the ITS region has been extensively used in molecular methods as well as
ecological studies on fungi.
Estimates of fungal biodiversity have undergone sharp changes following envi-
ronmental DNA sequencing of fungal ITS reads. Currently, there are around 81,000
accepted fungal names out of a total of up to 5.1 million species (Blackwell 2011).
Environmental DNA sequencing has accumulated more than one million fungal ITS
reads. Whether ‘sequence-based voucher-less fungi’ (in the absence of actual speci-
mens) should be given a formal nomenclature is widely debated, and the opinions
of expert mycologists are sharply divided on this point with reference to the estima-
tion of biodiversity of fungi in the natural environment. Several informative reviews
on DNA bar-coding as well as sequence-based fungal nomenclature are published
and may be consulted for a more complete understanding of the problem and its
2
Now chytrids are grouped under Chytridiomycota (N. P. Money, in The Fungi (Third Edition),
2016)
Foreword vii
perspectives (Shenoy et al. 2007; Begerow et al. 2010; Das and Deb 2015; Hibbett
et al. 2016; Hongsanen et al. 2018).
Conventionally, taxonomic principles related to fungi are debated and decisions
taken under the authority of the International Code of Botanical Nomenclature
(ICBN) held once in 6 years during the International Botanical Congress. Recently,
the ICBN has been renamed as International Code of Nomenclature for algae, fungi
and plants. In the Melbourne ICBN held in 2011, a momentous decision was taken
to abolish Article 59 which permitted the use of dual nomenclature for pleomorphic
fungi. In the morphology-based taxonomy, anamorphs (asexual forms) and teleo-
morphs (sexual forms) of fungi discovered and described independently could
receive different but valid names as ruled by Article 59 of the ICBN. In the present
era of phylogenetic molecular analysis, co-ordination of anamorphic and teleomor-
phic elements of a fungus can, in principle, be unequivocally established. Molecular
mycologists are strongly in favour of the unification of the fungal nomenclature and
achieve the ‘one fungus-one name’ ideal. However, there are several practical hur-
dles, and several specialist committees are in serious debates to elucidate the practi-
cal feasibility of implementing the same (Gams 2014).
Simultaneous with the focused attention on fungi as troublesome pathogens of
plants, animals and humans, beneficial attributes of fungi as sources of valuable
metabolites were also realized. Taka-Diastase from A. oryzae and citric acid from
Aspergillus niger are examples of fungal-based processes of manufacture in the late
nineteenth and early twentieth centuries. With the discovery of penicillin and the
dawn of the antibiotic era, the decades of the twentieth century became an era for
screening diverse microorganisms including fungi for discovering novel bioactive
molecules and producing them by fermentation. Several industrially useful enzymes,
hormones, vitamins and growth factors of fungal origin spearheaded the fermenta-
tion technology development. Fungi and their metabolites were also employed in
the biotransformation of organic molecules (e.g. steroids).
The beneficial attributes of fungi far outweighed their earlier negative image, and
this led to a very healthy and positive attitude towards fungi which deserved to be
studied for deriving beneficial products from their metabolism. The term ‘biodiver-
sity’ acquired a newer dimension in terms of its relevance to applied microbiologi-
cal and biotechnological research. An immense focus on the physiological,
biochemical and molecular aspects of mycodiversity became imminent for biopro-
cess development based on novel fungal strains.
Heterologous gene expression and production of mammalian proteins in fungal
hosts have been successfully carried out for chymosin (cheese manufacture enzyme)
in A. niger var. awamori and insulin in Komagataella (Pichia) pastoris.
Spectacular advances in both mycology and mycotechnology during the twenti-
eth century have added new dimensions to mycological research, necessitating
newer approaches to achieve success in this field at present. A different mindset and
training have become necessary in order to carry out meaningful research projects
viii Foreword
in the twenty-first century. Some of the salient features of this new preparation may
be defined as follows:
The endophytic fungus has imbibed the genes from the medicinal plant through
constant association. In the laboratory, culturing it on routine sugar-rich media
while facilitating satisfactory growth, there is no selection pressure for the growing
fungus to conserve these imbibed genes, and over several subcultures on routine
media may even be lost. An alternate suggestion which may be worth considering is
to plan the culture isolation of the endophyte on a relatively simple nutrient medium
fortified with some of the key chemical intermediate components involved in the
biosynthesis of the specific plant metabolite. It is possible on such a medium, the
metabolic genes for the plant metabolite are conserved better, and these may be
subjected to mutation and gene amplification through molecular techniques, leading
to strains which will be able to produce the metabolite at commercially viable lev-
els. Presently, the levels of production observed are very low and insignificant, and
much more studies are warranted before practical and viable technologies for
endophyte-based plant metabolites would become a reality.
Recent years have witnessed increased emphasis on the beneficial attributes of
fungi, recognizing their role in the natural ecosystems and as the source of several
metabolites of value for humankind. Rambold et al. (2013) in an article, suggesting
that mycology should be recognized as a major field in biology, stated ‘Given the
ecological and economic relevance of fungi...Mycology is insufficiently recognized
as a major field of science’. In the theme for the International Mycological Congress
held at Puerto Rico in July 2018, mycological discoveries for a better world and
how fungi contributed to the health of society and of ecosystem were given empha-
sis. Clearly, one can perceive the changing face of mycology from ‘fungi as destruc-
tive agents to be despised to fungi as true benefactors of humanity to be adored’.
In order to harness the full potential of fungi for mankind’s benefit, mycologists
must identify themselves as part of multidisciplinary teams exploring fungi for
novel metabolites. They should broaden their horizons of interest and acquire
knowledge in physiology, biochemistry and molecular biology of fungi. Familiarity
with natural product chemistry and bioprocess engineering for mould metabolites
including mould morphogenesis in fermentors related to growth conditions for
maximum yields are desirable.
Multidisciplinary collaboration to achieve success in technology is a must, and
the status of mycology in the twenty-first century is meaningfully reflected in the
following quote from Kreger (2003):
We all realise that most scientists in the future will be part of multi-disciplinary research
teams…the shift is causing a way scientists need to train … although they must still be an
expert in their specialty, they must also become conversant with techniques that once
seemed beyond their domain … they also need to recognize where their knowledge ends and
where they should seek the help from others….
x Foreword
It gives me pleasure in writing the foreword for this book, Advancing Frontiers
in Mycology and Mycotechnology: Basic and Applied Aspects of Fungi. The invited
chapters included in this book cover most of the aspects of fungal biology and
applied aspects of fungi. I wish to congratulate the editors and all the contributors
for their efforts in bringing out this book. I sincerely hope and wish that the book
will be useful for students, scholars, scientists and teachers of biology, microbiol-
ogy and biotechnology.
References
Begerow D, Nilsson H, Unterseher M, Maier W (2010) Current state and perspectives of fungal
DNA bar coding and rapid identification procedures. Appl Microbiol Biotechnol 87:99–108
Blackwell M (2011) The fungi: 1, 2, 3 … 5.1 million species ? Am J Bot 98: 426–438
Das S, Deb B (2015) DNA bar coding of fungi using ribosomal ITS marker for genetic diversity
analysis: a review. Int J Pure Appl Biosci 3:160–167
Gams W (2014) A new nomenclature for fungi. Mycol Iranica 1:1–5
Hesseltine CW (1983) Microbiology of oriental fermented foods. Annu Rev Microbiol 37: 575–601
Hibbett D, Abarenkov K, Kõljalg U, Öpik M, Vhai B, Cole J, Wang Q, Crous P, Robert V, Helgason
T, Herr JR, Kirk P, Lueschow S, O’Donnell K, Nilsson RH, Oono R, Schoch C, Smyth C,
Walker DM, Porras-Alfaro A, Taylor JW, Geiser DM (2016) Sequence based classification and
identification of fungi. Mycologia 108:1049–1068
Ho HH (2018) The taxonomy and biology of Phytophthora and Pythium. J Bacteriol Mycol Open
Access 6(1):40–45. https://doi.org/10.15406/jbmoa.2018.06.00174
Hongsanen S, Jeewon R, Purahong W, Xie N, Liu J-K, Jayawardana RS, Ekanayaka AH,
Dissanayake A, Raspe O, Hyde KD, Stadler M, Pers D (2018) Can we use environmental DNA
as holotypes? Fungal Divers 92:1–30.
Kreeger C (2003) The learning curve. Nat Biotechnol 21:951–952
Nagpal R, Puniya AK, Griffith GW, Goel G, Puniya M, Sehgal JP, Singh K (2009) Anaerobic
rumen fungi: potential and applications. Chapter 17, 375–393. In: Khachatourians GG, Arora
DK, Rajendran TP, Srivastava AK (eds), Agriculturally important micro-organisms, volume 1,
Academic World International
Narasimhan MJ (1930) Studies in the genus Phytophthora X Heterothallic strains of Phytophthora.
Phytopathology 20:201–214
Rambold G, Stadler M, Begerow D (2013) Mycology should be recognised as a field in biology at
an eye level with other major disciplines – a memorandum. Mycol Prog 12:455–463
Shenoy BD, Jeewon R, Hyde KD (2007) Impact of DNA sequence-data on the taxonomy of ana-
morphic fungi. Fungal Divers 26:1–54
Srinivasan MC (2004) Selective isolation of fungi, Chapter 2. In: Practical mycology for industrial
biotechnologists. Tata McGraw-Hill Publishing co. Ltd., New Delhi, p 26–46
Foreword xi
Srinivasan MC (2008) Unconventional techniques to explore fungal diversity for industrial enzyme
technology. In: Sridhar et al (ed) Novel techniques and ideas in mycology. Fungal Diversity
Research Series 26:261–270
Srinivasan MC (2014) Role of mycology and mycologists in an era of industrial biotechnology.
Kavaka 43:3–5
Stierle A, Strobel G, Stierle D (1993) Taxol and taxane production by Taxomyces andreanae, an
endophytic fungus of Pacific yew. Science 260:214–216
Preface
Fungi are members of the group of eukaryotic organisms, which include microbes
like yeasts, moulds and mushrooms. They are classified as kingdom Fungi and more
closely related to animals than to plants, therefore, placed with the animals in the
monophyletic group. Analyses using molecular phylogenetics support a monophy-
letic origin of fungi. They are heterotrophs, as they acquire their food by absorbing
dissolved molecules, typically by secreting digestive enzymes into their environ-
ment. The origin of fungi can be traced to single-celled marine ancestors in the
Mesoproterozoic era more than one billion years ago. Fungi have since then con-
quered not only land but also almost every potential habitat and substrate. Although
all fungi are heterotrophs, the fungal kingdom comprises a wide range of life strate-
gies ranging from saprotrophy through mutualism to parasitism. The fungal king-
dom encompasses an enormous diversity of taxa with varied ecologies, life cycle
strategies and morphologies ranging from unicellular aquatic chytrids and yeasts to
large mushrooms. However, very little is known about the true diversity of kingdom
Fungi, which had been estimated at five million species. Fungi are immensely
diverse, with 144,000 species named and classified so far at a current rate of around
2,000 per year. Of these, over 8,000 species are known to be detrimental to plants,
and at least 300 can be pathogenic to human beings.
Our fascination for the fungal kingdom is a natural and ancient one based on the
following: (i) the roles of fungi in the production of a variety of foods and beverages
and even as a source of food themselves; (ii) their global ecological impact, espe-
cially as the cause of devastating infections of humans and other animals and of
plants, including many crops grown around the world; and (iii) their roles as funda-
mental model systems in genetics and biological research.
Since the pioneering eighteenth- and nineteenth-century taxonomical works of
Carl Linnaeus, Christiaan Hendrik Persoon and Elias Magnus Fries, fungi have
been classified based on their morphological, physiological and biochemical char-
acteristics. Advances in molecular genetics have opened the way for DNA analysis
to be incorporated into taxonomy, which has sometimes challenged the historical
groupings based on morphology and other traits. Phylogenetic studies published in
the last decade have helped to reshape the classification within kingdom Fungi,
which has been divided into seven phyla (Microsporidia, Chytridiomycota,
Blastocladiomycota, Neocallimastigomycota, Glomeromycota, Ascomycota and
Basidiomycota).
xiii
xiv Preface
Along with bacteria, fungi are the major decomposers in most terrestrial and
some aquatic ecosystems; therefore, they play an essential role in nutrient cycling,
especially as saprobes and symbionts, degrading organic matter to inorganic mole-
cules, which can then re-enter anabolic metabolic pathways in plants and other
organisms. Certain fungi, in particular white-rot fungi, can degrade insecticides,
herbicides, pentachlorophenol, creosote, coal tars and heavy fuels and turn them
into carbon dioxide, water and basic elements. Fungi have been shown to mineralize
uranium oxides, suggesting that they may have application in the bioremediation of
radioactively polluted sites.
The human use of fungi for food preparation and other purposes is extensive and
has a long history. Mushroom farming and mushroom gathering are large industries
in several countries. The global commercial mushroom market was ~US$35 billion
in 2015 and is anticipated to grow to as much as ~US$60 billion by 2021.
Ethnomycology is the study of the historical uses and sociological impact of fungi.
Fungi have the capacity to produce an enormous range of natural products with
antimicrobial or other biological activities; therefore, many species have long been
used or are being developed for industrial production of antibiotics, vitamins and
anticancer and cholesterol-lowering drugs. Very recently, methods have been devel-
oped for genetic engineering of fungi that enable metabolic engineering of fungal
species. For example, genetic modification of yeast species, which are easy to grow
at fast rates in large fermentation vessels, has opened up ways of pharmaceutical
production that are potentially more efficient than production by the original source
organisms.
Several pivotal discoveries in biology were made by researchers using fungi as
model organisms. Indeed, seven Nobel prizes have been awarded to scientists study-
ing yeasts and moulds as model organisms that explain fundamental aspects of cell
biology: from Alexander Fleming, Ernst Chain and Howard Florey in 1945 for the
discovery of penicillin from Penicillium notatum; to George Beadle and Edward
Tatum in 1958 for their ‘one gene-one enzyme’ hypothesis in Neurospora crassa; to
Paul Nurse and Leland Hartwell in 2001 for cell division and cancer in
Schizosaccharomyces pombe and Saccharomyces cerevisiae; to Roger Kornberg in
2006 for eukaryotic gene transcription in S. cerevisiae; to Jack W. Szostak (shared)
in 2009 for chromosome telomeres in S. cerevisiae; to Randy Schekman in 2013
(shared) for machinery regulating vesicle traffic in S. cerevisiae; and to Yoshinori
Ohsumi in 2016 for autophagy in S. cerevisiae. Other important model fungi, which
includes Aspergillus nidulans; Candida albicans, a dimorphic, opportunistic human
pathogen; Magnaporthe grisea, a plant pathogen; and Pichia pastoris, a yeast
widely used for eukaryotic protein production, have more recently emerged, which
address specific biological questions relevant to medicine, plant pathology and
industrial uses.
In fact, a fungal species was the first eukaryotic organism to have its genome
completely sequenced (the model budding yeast S. cerevisiae). Advances in genet-
ics and cell biology have contributed to provide a detailed view of how the genome
contributes to the functions of the cell and of the organism. Together, these advances
in genomics and genetics provide a ‘blueprint’ for how these species operate and
Preface xv
have evolved at a cellular level, and consequently, they offer a wealth of knowledge
about how representative species in the fungal kingdom function and the diversity
that lies within. This diversity spans from the most basic way that a fungal cell is
organized, either as a yeast or as a filamentous hypha, to the myriad ways these spe-
cies interact with their environment, from aquatic basal fungi (Chytridiomycota,
Cryptomycota) to fungi that are associated with plants and were critical for their
emergence from the oceans and colonization of the planet to fungi that are patho-
gens of plants or animals. This diversity also extends to the biological behaviour and
cell biology of fungi, including fungi that can sense light and those that have evolved
to be insensitive to light (blind), the modes of sexual reproduction including hetero-
thallism and homothallism, the loss and retention of RNAi pathways, the replace-
ment of regional centromeres by point centromeres and the retention of flagella in
basal fungi versus their loss in fungal branches that evolved the ability to be aerially
dispersed.
This book entitled Advancing Frontiers in Mycology and Mycotechnology: Basic
and Applied Aspects of Fungi is aimed at reviewing major recent developments in
understanding the biology and potential biotechnological applications of fungi.
Parts I, II, III and IV focus on the basic aspects of fungi; their role in environmental
sustainability; their interactions with humans, plants and animals; and bioprospects,
respectively. Part I includes chapters on the diversity of fungi from different envi-
ronments, growth, morphogenesis and genetics and conservation and taxonomy.
Part II deals with the role of fungi in environmental sustainability (biodegradation
and bioremediation) and in nanobiotechnology. The interactions of fungi with
plants, animals and humans, such as mycorrhizal association, endophytism, human
mycoses and mycotoxins, are covered in Part III. The bioprospecting aspects of
entomo- and myco-pathogens, industrial enzymes and secondary metabolites in
healthcare are dealt with in Part IV.
We wish to thank all the contributors for readily accepting our invitation and
submitting their well-written chapters in their areas of specialization within the
stipulated period. We sincerely hope and wish that this book will be useful to stu-
dents, scholars, teachers and scientists in the broad areas of microbiology, life sci-
ences and biotechnology. Thanks are also due to Springer Nature for publishing the
book.
xvii
xviii Contents
Part IV Bioprospects
20 Bioprospecting of Fungal Entomo- and Myco-Pathogens.................... 497
E. K. Pathan, A. V. Patil, and M. V. Deshpande
21 Fungal Enzymes: Sources and Biotechnological Applications............. 515
Naveen Kango, Uttam Kumar Jana, and Ritumbhara Choukade
22 Fungi in Hypogean Environment: Bioprospection Perspective........... 539
S. R. Joshi and Upashna Chettri
23 Secondary Metabolites of Mushrooms: A Potential Source
for Anticancer Therapeutics with Translational Opportunities.......... 563
Sudeshna Nandi, Rimpa Sikder, and Krishnendu Acharya
24 Modulation of Fungal Metabolome by Biotic Stress............................ 599
Geane Pereira de Oliveira, Bruna de Almeida Martins,
Matheus Thomaz Nogueira Silva Lima,
and Jacqueline Aparecida Takahashi
25 Marine Fungi as a Potential Source of Future Cosmeceuticals........... 627
Shivankar Agrawal, Sunil K. Deshmukh, and Colin J. Barrow
Index.................................................................................................................. 671
About the Editors and Contributors
Editors
Dr. Sunil Kumar Deshmukh received his Ph.D. in Mycology from Dr. H.S. Gour
University, Sagar (M.P.) in 1983. The veteran industrial mycologist spent a substan-
tial part of his career at Hoechst Marion Roussel Limited [now Sanofi India Limited],
Mumbai and Piramal Enterprises Limited, Mumbai in drug discovery. He has to his
credit, 8 patents, 120 publications and 9 books on various aspects of Fungi and natu-
ral products of microbial origin. He is the past president of the Mycological Society
of India. He is a fellow of Mycological Society of India (MSI), the Association of
Biotechnology and Pharmacy and the Society for Applied Biotechnology. He is cur-
rently Fellow at Nano-Biotechnology Centre, TERI, New Delhi, and Adjunct
Associate Professor in Deakin University, Australia, working towards the develop-
ment of natural food colors, antioxidants and biostimulants through nanotechnology
intervention.
xxi
xxii About the Editors and Contributors
Dr. Mukund V. Deshpande obtained his PhD in 1982 in Biochemistry and D.Sc. in
Microbiology of the University of Pune in 1994. His extensive work in the area of
fungal biology, especially, fungal differentiation earned him D.Sc. He has worked
extensively on the use of fungi and fungal products in Biotechnology. Dr. Deshpande
successfully completed more than 35 research projects funded by national and inter-
national funding agencies, such as Indo-Swiss Collaboration in Biotechnology
(ISCB) programme of Department of Biotechnology (DBT), New Delhi and Swiss
Development Cooperation (SDC), Berne, Switzerland on development of mycoin-
secticide, Indo-Belarus programme of DBT on biopesticides, Indo-Mexico pro-
gramme of Department of Science and Technology, New Delhi and CONACYT on
fungal dimorphism. Dr. Deshpande is an elected fellow of the Maharashtra Academy
of Sciences (FMASc, 1994) and the Society for Biocontrol Advancement (FSBA,
2010). He is also a recipient of the Department of Biotechnology Overseas (Short-
term) Associateship (1995), and Commonwealth Science Council Fellowship
(1998). He has to his credit more than 140 research papers, reviews and chapters, 8
patents, 6 books and a number of popular articles. He has his own start-up
Greenvention Biotech located in Urli-Kanchan, Pune for the translational activities
in Agricultural Biotechnology.
Contributors
Abstract
The mushrooms have existed approximately 130 million years ago, i.e., long
before human beings evolved on this planet as per the fossil records. Earlier in
Sanskrit mushroom is known as “Ksuonpa.” In Hindi, mushroom is known as
“Khumbi.” Since long mushrooms are worshiped and also considered as divine.
In nature, mushrooms have not only been a source of food for man and other
animals but also have contributed an important role in the cycling of carbon and
other elements through the breakdown of lignocellulolytic plant residues and
animal dung which serves as the substrates for these saprophytic fungi. The his-
torical records of intentionally cultivated mushrooms estimated that the first
mushroom cultivation was started in 600 AD. In the last 20 years, much progress
has been made in the field of mechanization of mushroom cultivation, i.e.,
manure turners, spawning, filling and casing of trays, mechanical harvesting, and
polythene bag method of cultivation. In India, cultivation of edible mushrooms is
of very recent origin, though methods of cultivation of them were known for
many years. The research on different aspects, viz., production, productivity,
spawn production, strain improvement, post-harvest technology disease, and
pest management, was attempted by several scientists. All mushrooms belong to
the group of fungi, a group very distinct from plants, animals, and bacteria. Most
fungi have plant-like cells but miss the most important features of plants. The
known number species of fungi was about 69,000 till 1990, while it was conser-
vatively estimated that 1.5 million species of fungi actually existed in nature.
V. K. Bhalerao (*)
AICRP on Fruits, Mahatma Phule Krishi Vidyapeeth, Rahuri,
Ahmednagar, Maharashtra, India
A. P. Gaikwad
AICRP on Mushroom, College of Agriculture, Pune, India
C. D. Deokar · K. S. Raghuwanshi
Department of Plant Pathology, Mahatma Phule Krishi Vidyapeeth, Ahmednagar, India
Mushrooms are cultivated throughout the world. About 200 species of more
than 2000 edible fungi are widely adopted for human consumption. Out of these,
only 10–12 species are commercially cultivated since the past few decades in
India due to technical advancement. Mushrooms have the capacity to produce
highest proteins per unit area and time by utilizing vertical space which is hun-
dred times more than the traditional agriculture and animal husbandry. This hi-
tech horticulture venture can reduce the pressure on cultivated land to meet the
food shortages all over the world. More than 100 countries are engaged in mush-
room farming today which is increasing at an annual rate of 6–7% per annum.
The very high levels of mechanization and automation were achieved in mush-
room farming in developed countries of Europe and America. As per FAO Stat,
the present world production of mushrooms is around 3.5 million tonnes, which
is more than 25 million tonnes (estimated) as per claims of Chinese Association
of Edible Fungi.The mushrooms can be cultivated under varied climatic condi-
tions. Some of the economically important mushrooms cultivated all over the
world under temperate, subtropical, and tropical conditions are Agaricus bispo-
rus, Lentinula edodes, Flammulina velutipes, Agaricus bitorquis, Pleurotus spp.,
Auricularia spp., Agrocybe aegerita, Volvariella spp., Calocybe indica,
Ganoderma lucidum, etc.
Mushrooms are rich source of proteins, carbohydrates, valuable salts, and
vitamins in diet of human being. Mushroom fungus has the ability to secrete a
wide variety of hydrolyzing and oxidizing enzymes which have potential for
biotechnological applications. More than 100 medicinal edible mushrooms have
been identified. However, important medicinal mushrooms are Lentinus edodes
(shiitake mushroom), Ganoderma lucidum (reishi mushroom), Grifola frondosa
(maitake mushroom), Pleurotus ostreatus (oyster mushroom), Agaricus bisporus
(button mushroom), Coriolus versicolor (PSK), Boletus edulis, Tremella fucifor-
mis, Auricularia polytricha, Hericium erinaceus, and Cordyceps sinensis.
Keywords
Mushrooms · Oyster mushroom · Agaricus sp. · Medicinal values · Cultivation ·
Commercial production · Nutritional resource
1.1 Introduction
The mushrooms have existed approximately 130 million years ago, i.e., long before
human beings evolved on this planet as per the fossil records. The origin of the term
“Mushroom” has different views. In Latin, “Fungo” means to flourish. The Greek
term mushroom was derived from the word “Sphonggos” or “Sphoggos” which
means “Sponge” and referred to the sponge-like structure of some species. In
French, the term mushroom was derived from the word “Mousseron” (muceron),
“Mousse,” or “Moss.”
1 The Mystical World of Mushrooms 5
About three and half millennia ago, the Greek hero Perseus founded Mycenae
city after a legendary mushroom. “Mycenae” is derived from Greek word “mycol-
ogy” (i.e., mykes – mushroom + logos – discourse). Earlier in Sanskrit, mushroom
is known as “Ksuonpa.” In Hindi, mushroom is known as “Khumbi.” The other
words for fleshy capped fungi are Chatra, Kukurmutta, Kavaka, Bhumi kavak, and
Bhustrana. Aryans during migration into the Indian subcontinent around in
1500 B.C. carried with them an intoxicating drink “Soma” which they used in their
religious rites. According to Wasson) (1969), the “Soma” in Rig Veda refers to
Amanita muscaria.
Since long mushrooms are worshiped and also considered as divine. There are
lots of superstitions about the mushrooms. According to the ancient Indian, Greek,
and Roman myths, mushrooms sprang from a stroke of lightning. The Indians in
Mexico believed that mushrooms are sacred because they are borne of the sexual
intercourse between a bolt of lightning and the earth. The mushroom-shaped stone
carvings have also been found in Central America and the highlands of Guatemala.
These objects resemble very closely to the fruit body of Amanita muscaria.
In nature, mushrooms have not only been a resource of food for man and other
animals but also have an important role in the cycling of carbon and other elements
through the breakdown of lignocellulolytic plant residues and animal dung which
serves as the substrates for these saprophytic fungi. In this way, mushroom species
as agents of decay helps in keeping the environment clean.
The mushrooms were long appreciated because of their flavor, texture, and
medicinal or tonic attributes. The recognition, that mushrooms are nutritionally a
very good food is much more recent.
The popularity of mushroom is desired from the following highly preferred char-
acteristics of food:
The historical records of intentionally cultivated mushrooms are shown in Table 1.1.
The first artificial cultivation of Auricularia auricula mushroom was estimated
around 600 AD in China on wood logs and the same time other wood rotting mush-
rooms, such as Flammulina velutipes (800 AD) and Lentinula edodes (1000 AD).
In Europe, the first cultivated fungi, the mushroom, was introduced in the seven-
teenth century. Agaricus bisporus was not cultivated until 1600 AD which is the
leading mushrooms. The Pleurotus ostreatus was first cultivated in the USA, and
several other species of Pleurotus were initially cultivated in India.
The first record of the cultivation of mushrooms was during 1638–1715, i.e., the
reign of Louis XIV. Treschow shortly before 1700 suggested a method of growing
A. bisporus in hotbeds under heat. Tourneforte, a French man in 1707, published
description and know-how of growing mushrooms which is similar to the present
methods.
Van Griensven (1988) reviewed the historical events in mushroom cultivation,
whereas a detailed history was elaborated by Shu-ting and Miles Philip. The
cultivation of mushrooms aboveground is originated in Sweden. Bahl (2002) reviewed
that Lundberg described mushroom growing in greenhouses in year 1754 whereas,
the French started growing mushrooms underground in the quarries around Paris on
horse manure in 1800 and Callow reported shelf bed method in a peculiar type of
house for mushroom cultivation in 1831.
By the end of the nineteenth century, French mycologists Matrochot and
Constantin (1894) discovered the cause of mushroom disease La mole (Mycogone
perniciosa Magnus) and started sulfur fumigation. They were able to germinate
spores for obtaining sterile spawn. Miss Fergussen (1902) of Corne described
detailed method for germinating spores.
Duggar (1905), an American, perfected a method of making pure culture spawn
from mushroom tissue. Lambert (1929) introduced and marketed pure spore culture
bottled spawn in the USA. The cultivation of mushroom aboveground is originated
in Sweden. Sinden (1938) patented his grain spawn process. Thomas, Austin, and
Jary worked on pests of mushrooms. Ware, Glasscock, and Bewley in the UK and
Lambert and Beach in the USA worked on diseases and competitive molds of
mushrooms.
In 1945, MGA (Mushroom Growers Association of England and Ireland), MRA
(Mushroom Research Association Ltd.), and MGP (Midlands Group of Publication)
were formed and established. In the same year, an organization called Mushroom
Growers Association of Great Britain and North Ireland was established. This asso-
ciation publishes a mushroom journal, which is supplied to the 50 nations.
In 1946, the first mushroom research station was founded in England with Dr.
R.L. Edwards as its first director. The mushroom research at this station is mainly
devoted to Agaricus bisporus and involves all aspects of this single species. In 1954,
1 The Mystical World of Mushrooms 7
Table 1.1 Historical evidence of commonly cultivated mushrooms (Miles and Chang 1997)
Species Record first cultivated (est.)
Agaricus bisporus 1600 (Atkins 1979)
Agaricus bitorquis 1961
Agrocybe cylindracea 1950
Amanita caesarea 1984
Armillaria mellea 1983
Auricularia auricula 600
Coprinus comatus 1984
Dictyophora duplicata 1982
Flammulina velutipes 800
Ganoderma spp. 1621
Gloestereum incarnatum 1989
Grifola frondosus 1983
Hericium coralloides 1984
Hericium erinaceus 1960
Hohenbuehelia serotina 1982
Hypsizygus marmoreus 1973
Lentinus edodes 1000
Lentinus tigrinus 1988
Lyophyllum ulmarium 1987 (Wang and Zhang 1987)
Morchella spp. 1986
Oudemansiella radicata 1982
Pholiota nameko 1958
Pleurotus citrinopileatus 1981
Pleurotus cystidiosus 1969
Pleurotus ferulae 1958
Pleurotus flabellatus 1962 (Bano and Srinvatava 1962)
Pleurotus Florida 1958
Pleurotus ostreatus 1900
Pleurotus sajor-caju 1974 (Jandaik 1974)
Poria cocos 1232
Sparassis crispa 1985
Tremella fuciformis 1800
Tremella mesenterica 1985
Tricholoma gambosum 1991
Tricholoma lobayense 1990
Tricholoma mongolicum 1991
Volvariella volvacea 1700
the mushroom research station was transferred along with three staff members and
government grants to the new glasshouse crop research station at Littlehampton,
Sussex.
In 1959 at Horst in Holland, a fine center on mushroom was opened. In the later
years of the eighteenth century, French growers discovered the usefulness of gyp-
sum powder to prevent greasiness in compost. Bewley and Lambert worked on
Other documents randomly have
different content
treaty of July 22 with the Wyandots, Delawares, Shawanees, and
other tribes, binding them to take up arms against the British, had
then arrived, and this news lessened the interest of both parties in
the Indian question. None of the American negotiators were
prepared to break off negotiations on that point at such a time, and
Clay was so earnest to settle the matter that he took from Gallatin
and Adams the task of writing the necessary acceptance of the
British ultimatum. Gallatin and Clay decided to receive the British
article as according entirely with the American offer of amnesty, and
the note was so written.[44]
With this cordial admission of the British ultimatum the Americans
coupled an intimation that the time had come when an exchange of
general projects for the proposed treaty should be made. More than
two months of discussion had then resulted only in eliminating the
Indians from the dispute, and in agreeing to maintain silence in
regard to the Lakes. Another great difficulty which had been
insuperable was voluntarily removed by President Madison and his
Cabinet, who after long and obstinate resistance at last authorized
the commissioners, by instructions dated June 27, to omit
impressment from the treaty. Considering the frequent positive
declarations of the United States government, besides the rejection
of Monroe’s treaty in 1807 and of Admiral Warren’s and Sir George
Prevost’s armistice of 1812 for want of an explicit concession on that
point, Monroe’s letter of June 27 was only to be excused as an act of
common-sense or of necessity. The President preferred to represent
it as an act of common-sense, warranted by the peace in Europe,
which promised to offer no further occasion for the claim or the
denial of the British right. On the same principle the subject of
blockades was withdrawn from discussion; and these concessions,
balanced by the British withdrawal from the Indian ultimatum and
the Lake armaments, relieved the American commissioners of all
their insuperable difficulties.
The British commissioners were not so easily rescued from their
untenable positions. The American note of October 13, sent as usual
to London, was answered by Bathurst October 18 and 20,[45] in
instructions revealing the true British terms more completely than
had yet been ventured. Bathurst at length came to the cardinal point
of the negotiation. As the American commissioners had said in their
note of August 24, the British government must choose between the
two ordinary bases of treaties of peace,—the state before the war, or
status ante bellum; and the state of possession, or uti possidetis.
Until the middle of October, 1814, the uti possidetis, as a basis of
negotiation, included whatever country might have been occupied by
Sir George Prevost in his September campaign. Bathurst from the
first intended to insist on the state of possession, but had not
thought proper to avow it. His instructions of October 18 and 20
directed the British commissioners to come to the point, and to claim
the basis of uti possidetis from the American negotiators:—
“On their admitting this to be the basis on which they are ready to
negotiate, but not before they have admitted it, you will proceed to
state the mutual accommodations which may be entered into in
conformity with this basis. The British occupy Fort Michillimackinaw,
Fort Niagara, and all the country east of the Penobscot. On the other
hand the forces of the United States occupy Fort Erie and Fort
Amherstburg [Malden]. On the government of the United States
consenting to restore these two forts, Great Britain is ready to restore
the forts of Castine and Machias, retaining Fort Niagara and Fort
Michillimackinaw.”
Thus the British demand, which had till then been intended to
include half of Maine and the whole south bank of the St. Lawrence
River from Plattsburg to Sackett’s Harbor, suddenly fell to a demand
for Moose Island, a right of way across the northern angle of Maine,
Fort Niagara with five miles circuit, and the Island of Mackinaw. The
reason for the new spirit of moderation was not far to seek. On the
afternoon of October 17, while the British Cabinet was still
deliberating on the basis of uti possidetis, news reached London that
the British invasion of northern New York, from which so much had
been expected, had totally failed, and that Prevost’s large army had
precipitately retreated into Canada. The London “Times” of October
19 was frank in its expressions of disappointment:—
“This is a lamentable event to the civilized world.... The subversion
of that system of fraud and malignity which constitutes the whole
policy of the Jeffersonian school ... was an event to which we should
have bent and yet must bend all our energies. The present American
government must be displaced, or it will sooner or later plant its
poisoned dagger in the heart of the parent State.”
The failure of the attempt on Baltimore and Drummond’s bloody
repulse at Fort Erie became known at the same time, and coming
together at a critical moment threw confusion into the Ministry and
their agents in the press and the diplomatic service throughout
Europe. The “Courier” of October 25 declared that “peace with
America is neither practicable nor desirable till we have wiped away
this late disaster;” but the “Morning Chronicle” of October 21–24
openly intimated that the game of war was at an end. October 31,
the Paris correspondent of the London “Times” told of the cheers
that rose from the crowds in the Palais Royal gardens at each recital
of the Plattsburg defeat; and October 21 Goulburn wrote from Ghent
to Bathurst,[46]—
“The news from America is very far from satisfactory. Even our
brilliant success at Baltimore, as it did not terminate in the capture of
the town, will be considered by the Americans as a victory and not as
an escape.... If it were not for the want of fuel in Boston, I should be
quite in despair.”
In truth the blockade was the single advantage held by England;
and even in that advantage the Americans had a share as long as
their cruisers surrounded the British Islands.
Liverpool wrote to Castlereagh, October 21,[47] commenting
severely on Prevost’s failure, and finding consolation only in the
thought that the Americans showed themselves even less patriotic
than he had supposed them to be:—
“The capture and destruction of Washington has not united the
Americans: quite the contrary. We have gained more credit with them
by saving private property than we have lost by the destruction of
their public works and buildings. Madison clings to office, and I am
strongly inclined to think that the best thing for us is that he should
remain there.”
Castlereagh at Vienna found himself unable to make the full
influence of England felt, so long as such mortifying disasters by
land and sea proved her inability to deal with an enemy she
persisted in calling contemptible.
On the American commissioners the news came, October 21, with
the effect of a reprieve from execution. Gallatin was deeply moved;
Adams could not believe the magnitude of the success; but as far as
regarded their joint action, the overthrow of England’s scheme
produced no change. Their tone had always been high, and they
saw no advantage to be gained by altering it. The British
commissioners sent to them, October 21, the substance of the new
instructions, offering the basis of uti possidetis, subject to
modifications for mutual convenience.[48] The Americans by
common consent, October 23, declined to treat on that basis, or on
any other than the mutual restoration of territory.[49] They thought
that the British government was still playing with them, when in
truth Lord Bathurst had yielded the chief part of the original British
demand, and had come to what the whole British empire regarded
as essentials,—the right of way to Quebec, and the exclusion of
American fishermen from British shores and waters.
The American note of October 24, bluntly rejecting the basis of uti
possidetis, created a feeling akin to consternation in the British
Cabinet. At first, ministers assumed that the war must go on, and
deliberated only on the point to be preferred for a rupture. “We still
think it desirable to gain a little more time before the negotiation is
brought to a close,” wrote Liverpool to the Duke of Wellington,[50]
October 28; and on the same day he wrote to Castlereagh at Vienna
to warn him that the American war “will probably now be of some
duration,” and treating of its embarrassments without disguise.[51]
The Czar’s conduct at Vienna had annoyed and alarmed all the great
Powers, and the American war gave him a decisive advantage over
England; but even without the Russian complication, the prospect
for ministers was not cheering.
“Looking to a continuance of the American war, our financial state
is far from satisfactory,” wrote Lord Liverpool; “... the American war
will not cost us less than £10,000,000, in addition to our peace
establishment and other expenses. We must expect, therefore, to
hear it said that the property tax is continued for the purpose of
securing a better frontier for Canada.”
A week passed without bringing encouragement to the British
Cabinet. On the contrary the Ministry learned that a vigorous
prosecution of hostilities would cost much more than ten million
pounds, and when Liverpool next wrote to Castlereagh, November 2,
[52] although he could still see “little prospect for our negotiations at
Ghent ending in peace,” he added that “the continuance of the
American war will entail upon us a prodigious expense, much more
than we had any idea of.” A Cabinet meeting was to be held the next
day, November 3, to review the whole course of policy as to
America.
Throughout the American difficulties, from first to last, the most
striking quality shown by the British government was the want of
intelligence which caused the war, and marked the conduct of both
the war and the negotiations. If the foreign relations of every
government were marked by the same character, politics could be no
more than rivalry in the race to blunder; but in October, 1814,
another quality almost equally striking became evident. The
weakness of British councils was as remarkable as their want of
intelligence. The government of England had exasperated the
Americans to an animosity that could not forget or forgive, and every
dictate of self-interest required that it should carry out its policy to
the end. Even domestic politics in Parliament might have been more
easily managed by drawing public criticism to America, while in no
event could taxes be reduced to satisfy the public demand.[53]
Another year of war was the consistent and natural course for
ministers to prefer.
So the Cabinet evidently thought; but instead of making a
decision, the Cabinet council of November 3 resorted to the
expedient of shifting responsibility upon the Duke of Wellington. The
Duke was then Ambassador at Paris. His life had been threatened by
angry officers of Napoleon, who could not forgive his victories at
Vittoria and Toulouse. For his own security he might be sent to
Canada, and if he went, he should go with full powers to close the
war as he pleased.
The next day, November 4, Liverpool wrote to Wellington,
explaining the wishes of the Cabinet, and inviting him to take the
entire command in Canada, in order to bring the war to an
honorable conclusion.[54] Wellington replied November 9,—and his
words were the more interesting because, after inviting and
receiving so decided an opinion from so high an authority, the
Government could not easily reject it. Wellington began by reviewing
the military situation, and closed by expressing his opinion on the
diplomatic contest:[55]—
“I have already told you and Lord Bathurst that I feel no objection
to going to America, though I don’t promise to myself much success
there. I believe there are troops enough there for the defence of
Canada forever, and even for the accomplishment of any reasonable
offensive plan that could be formed from the Canadian frontier. I am
quite sure that all the American armies of which I have ever read
would not beat out of a field of battle the troops that went from
Bordeaux last summer, if common precautions and care were taken of
them. That which appears to me to be wanting in America is not a
general, or a general officer and troops, but a naval superiority on the
Lakes.”
These views did not altogether accord with those of Americans,
who could not see that the British generals made use of the Lakes
even when controlling them, but who saw the troops of Wellington
retire from one field of battle after another,—at Plattsburg,
Baltimore, and New Orleans,—while taking more than common
precautions. Wellington’s military comments showed little interest in
American affairs, and evidently he saw nothing to be gained by
going to Canada. His diplomatic ideas betrayed the same bias:—
“In regard to your present negotiations, I confess that I think you
have no right, from the state of the war, to demand any concession of
territory from America.... You have not been able to carry it into the
enemy’s territory, notwithstanding your military success and now
undoubted military superiority, and have not even cleared your own
territory on the point of attack. You cannot on any principle of equality
in negotiation claim a cession of territory excepting in exchange for
other advantages which you have in your power.... Then if this
reasoning be true, why stipulate for the uti possidetis? You can get no
territory; indeed, the state of your military operations, however
creditable, does not entitle you to demand any.”
After such an opinion from the first military authority of England,
the British Ministry had no choice but to abandon its claim for
territory. Wellington’s letter reached London about November 13,
and was duly considered in the Cabinet. Liverpool wrote to
Castlereagh, November 18, that the Ministry had made its decision;
the claim for territory was to be abandoned. For this retreat he
alleged various excuses,—such as the unsatisfactory state of the
negotiations at Vienna, and the alarming condition of France; the
finances, the depression of rents, and the temper of Parliament.[56]
Such reasoning would have counted for nothing in the previous
month of May, but six months wrought a change in public feeling.
The war had lost public favor. Even the colonial and shipping
interests and the navy were weary of it, while the army had little to
expect from it but hard service and no increase of credit. Every
Englishman who came in contact with Americans seemed to suffer.
Broke, the only victor by sea, was a lifelong invalid; and Brock and
Ross, the only victors on land, had paid for their success with their
lives. Incessant disappointment made the war an unpleasant
thought with Englishmen. The burning of Washington was an exploit
of which they could not boast. The rate of marine insurance was a
daily and intolerable annoyance. So rapidly did the war decline in
favor, that in the first half of December it was declared to be
decidedly unpopular by one of the most judicious English liberals,
Francis Horner; although Horner held that the Americans, as the
dispute then stood, were the aggressors.[57] The tone of the press
showed the same popular tendency, for while the “Times” grumbled
loudly over the Canada campaign, the “Morning Chronicle” no longer
concealed its hostility to the war, and ventured to sneer at it, talking
of “the entire defeat and destruction of the last British fleet but one;
for it has become necessary to particularize them now.”[58]
While the Cabinet still waited, the first instalment of Ghent
correspondence to August 20, published in America October 10,
returned to England November 18, and received no flattering
attention. “We cannot compliment our negotiators,” remarked the
“Morning Chronicle;” and the “Times” was still less pleased. “The
British government has been tricked into bringing forward demands
which it had not the power to enforce.... Why treat at all with Mr.
Madison?” In Parliament, November 19, the liberal opposition
attacked the Government for setting up novel pretensions. Ministers
needed no more urging, and Bathurst thenceforward could not be
charged with waste of time.
During this interval of more than three weeks the negotiators at
Ghent were left to follow their own devices. In order to provide the
Americans with occupation, the British commissioners sent them a
note dated October 31 calling for a counter-project, since the basis
of uti possidetis was refused.[59] This note, with all the others since
August 20, was sent by the Americans to Washington on the same
day, October 31; and then Gallatin and Adams began the task of
drafting the formal project of a treaty. Immediately the internal
discords of the commission broke into earnest dispute. A struggle
began between the East and the West over the fisheries and the
Mississippi.
The treaty of 1783 coupled the American right of fishing in British
waters and curing fish on British shores with the British right of
navigating the Mississippi River. For that arrangement the elder
Adams was responsible. The fisheries were a Massachusetts interest.
At Paris in 1783 John Adams, in season and out of season, with his
colleagues and with the British negotiators, insisted, with the
intensity of conviction, that the fishing rights which the New England
people held while subjects of the British crown were theirs by no
grant or treaty, but as a natural right, which could not be
extinguished by war; and that where British subjects had a right to
fish, whether on coasts or shores, in bays, inlets, creeks, or harbors,
Americans had the same right, to be exercised wherever and
whenever they pleased. John Adams’s persistence secured the article
of the definitive treaty, which, without expressly admitting a natural
right, coupled the in-shore fisheries and the navigation of the
Mississippi with the recognition of independence. In 1814 as in 1783
John Adams clung to his trophies, and his son would have waged
indefinite war rather than break his father’s heart by sacrificing what
he had won; but at Ghent the son stood in isolation which the father
in the worst times had never known. Massachusetts left him to
struggle alone for a principle that needed not only argument but
force to make it victorious. Governor Strong did not even write to
him as he did to Pickering, that Massachusetts would give an
equivalent in territory for the fisheries. As far as the State could
influence the result, the fisheries were to be lost by default.
Had Adams encountered only British opposition he might have
overborne it as his father had done; but since 1783 the West had
become a political power, and Louisiana had been brought into the
Union. If the fisheries were recognized as an indefeasible right by
the treaty of 1783, the British liberty of navigating the Mississippi
was another indefeasible right, which must revive with peace. The
Western people naturally objected to such a proposition. Neither
they nor the Canadians could be blamed for unwillingness to impose
a mischievous servitude forever upon their shores, and Clay believed
his popularity to depend on preventing an express recognition of the
British right to navigate the Mississippi. Either Clay or Adams was
sure to refuse signing any treaty which expressly sacrificed the local
interests of either.
In this delicate situation only the authority and skill of Gallatin
saved the treaty. At the outset of the discussion, October 30, Gallatin
quietly took the lead from Adams’s hands, and assumed the
championship of the fisheries by proposing to renew both privileges,
making the one an equivalent for the other. Clay resisted obstinately,
while Gallatin gently and patiently overbore him. When Gallatin’s
proposal was put to the vote November 5, Clay and Russell alone
opposed it,—and the support then given by Russell to Clay was
never forgotten by Adams. Clay still refusing to sign the offer,
Gallatin continued his pressure, until at last, November 10, Clay
consented to insert, not in the project of treaty, but in the note
which accompanied it, a paragraph declaring that the commissioners
were not authorized to bring into discussion any of the rights
hitherto enjoyed in the fisheries; “From their nature, and from the
peculiar character of the treaty of 1783 by which they were
recognized, no further stipulation has been deemed necessary by the
Government of the United States to entitle them to the full
enjoyment of all of them.”
Clay signed the note,[60] though unwillingly; and it was sent,
November 10, with the treaty project, to the British commissioners,
who forwarded it to London, where it arrived at the time when the
British Cabinet had at last decided on peace. Bathurst sent his reply
in due course; and Goulburn’s disgust was great to find that instead
of breaking negotiation on the point of the fisheries as he wished,
[61] he was required once more to give way. “You know that I was
never much inclined to give way to the Americans,” he wrote,
November 25.[62] “I am still less inclined to do so after the
statement of our demands with which the negotiation opened, and
which has in every point of view proved most unfortunate.”
The British reply, dated November 26,[63] took no notice of the
American reservation as to the fisheries, but inserted in the project
the old right of navigating the Mississippi. Both Bathurst and
Goulburn thought that their silence, after the American declaration,
practically conceded the American right to the fisheries, though
Gambier and Dr. Adams thought differently.[64] In either case the
British note of November 26, though satisfactory to Adams, was far
from agreeable to Clay, who was obliged to endanger the peace in
order to save the Mississippi. Adams strongly inclined to take the
British project precisely as it was offered,[65] but Gallatin overruled
him, and Clay would certainly have refused to sign. In discussing the
subject, November 28, Gallatin proposed to accept the article on the
navigation of the Mississippi if the British would add a provision
recognizing the fishing rights. Clay lost his temper, and intimated
something more than willingness to let Massachusetts pay for the
pleasure of peace;[66] but during the whole day of November 28,
and with the same patience November 29, Gallatin continued urging
Clay and restraining Adams, until at last on the third day he brought
the matter to the point he wished.
The result of this long struggle saved not indeed the fisheries, but
the peace. Clay made no further protest when, in conference with
the British commissioners December 1, the Americans offered to
renew both the disputed rights.[67] Their proposal was sent to
London, and was answered by Bathurst December 6, in a letter
offering to set aside for future negotiation the terms under which the
old fishing liberty and the navigation of the Mississippi should be
continued for fair equivalents.[68] The British commissioners
communicated this suggestion in conference December 10, and
threw new dissension among the Americans.
The British offer to reserve both disputed rights for future
negotiation implied that both rights were forfeited, or subject to
forfeit, by war,—an admission which Adams could not make, but
which the other commissioners could not reject. At that point Adams
found himself alone. Even Gallatin admitted that the claim to the
natural right of catching and curing fish on British shores was
untenable, and could never be supported. Adams’s difficulties were
the greater because the question of peace and war was reduced to
two points,—the fisheries and Moose Island,—both interesting to
Massachusetts alone. Yet the Americans were unwilling to yield
without another struggle, and decided still to resist the British claim
as inconsistent with the admitted basis of the status ante bellum.
The struggle with the British commissioners then became warm. A
long conference, December 12, brought no conclusion. The treaty of
1783 could neither be followed nor ignored, and perplexed the
Englishmen as much as the Americans. During December 13 and
December 14, Adams continued to press his colleagues to assert the
natural right to the fisheries, and to insist on the permanent
character of the treaty of 1783; but Gallatin would not consent to
make that point an ultimatum. All the commissioners except Adams
resigned themselves to the sacrifice of the fisheries; but Gallatin
decided to make one more effort before abandoning the struggle,
and with that object drew up a note rejecting the British stipulation
because it implied the abandonment of a right, but offering either to
be silent as to both the fisheries and the Mississippi, or to admit a
general reference to further negotiation of all subjects in dispute, so
expressed as to imply no abandonment of right.
The note was signed and sent December 14,[69] and the
Americans waited another week for the answer. Successful as they
had been in driving their British antagonists from one position after
another, they were not satisfied. Adams still feared that he might not
be able to sign, and Clay was little better pleased. “He said we
should make a damned bad treaty, and he did not know whether he
would sign it or not.”[70] Whatever Adams thought of the treaty, his
respect for at least two of his colleagues was expressed in terms of
praise rarely used by him. Writing to his wife, September 27,[71]
Adams said: “Mr. Gallatin keeps and increases his influence over us
all. It would have been an irreparable loss if our country had been
deprived of the benefit of his talents in this negotiation.” At the
moment of final suspense he wrote again, December 16:—
“Of the five members of the American mission, the Chevalier
[Bayard] has the most perfect control of his temper, the most
deliberate coolness; and it is the more meritorious because it is real
self-command. His feelings are as quick and his spirits as high as
those of any one among us, but he certainly has them more under
government. I can scarcely express to you how much both he and Mr.
Gallatin have risen in my esteem since we have been here living
together. Gallatin has not quite so constant a supremacy over his own
emotions; yet he seldom yields to an ebullition of temper, and
recovers from it immediately. He has a faculty, when discussion grows
too warm, of turning off its edge by a joke, which I envy him more
than all his other talents; and he has in his character one of the most
extraordinary combinations of stubbornness and of flexibility that I
ever met with in man. His greatest fault I think to be an ingenuity
sometimes trenching upon ingenuousness.”
Gallatin’s opinion of Adams was not so enthusiastic as Adams’s
admiration for him. He thought Adams’s chief fault to be that he
lacked judgment “to a deplorable degree.”[72] Of Clay, whether in his
merits or his faults, only one opinion was possible. Clay’s character
belonged to the simple Southern or Virginia type, somewhat
affected, but not rendered more complex, by Western influence,—
and transparent beyond need of description or criticism.
The extraordinary patience and judgment of Gallatin, aided by the
steady support of Bayard, carried all the American points without
sacrificing either Adams or Clay, and with no quarrel of serious
importance on any side. When Lord Bathurst received the American
note of December 14, he replied December 19, yielding the last
advantage he possessed:[73] “The Prince Regent regrets to find that
there does not appear any prospect of being able to arrive at such
an arrangement with regard to the fisheries as would have the effect
of coming to a full and satisfactory explanation on that subject;” but
since this was the case, the disputed article might be altogether
omitted.
Thus the treaty became simply a cessation of hostilities, leaving
every claim on either side open for future settlement. The formality
of signature was completed December 24, and closed an era of
American history. In substance, the treaty sacrificed much on both
sides for peace. The Americans lost their claims for British
spoliations, and were obliged to admit question of their right to
Eastport and their fisheries in British waters; the British failed to
establish their principles of impressment and blockade, and admitted
question of their right to navigate the Mississippi and trade with the
Indians. Perhaps at the moment the Americans were the chief
losers; but they gained their greatest triumph in referring all their
disputes to be settled by time, the final negotiator, whose decision
they could safely trust.
CHAPTER III.
England received the Treaty of Ghent with feelings of mixed anger
and satisfaction. The “Morning Chronicle” seemed surprised at the
extreme interest which the news excited. As early as November 24,
when ministers made their decision to concede the American terms,
the “Morning Chronicle” announced that “a most extraordinary
sensation was produced yesterday” by news from Ghent, and by
reports that ministers had abandoned their ground. When the treaty
arrived, December 26, the same Whig newspaper, the next morning,
while asserting that ministers had “humbled themselves in the dust
and thereby brought discredit on the country,” heartily approved
what they had done; and added that “the city was in a complete
state of hurricane during the whole of yesterday, but the storm did
not attain its utmost height until toward the evening.... Purchases
were made to the extent of many hundred thousand pounds.” The
importance of the United States to England was made more
apparent by the act of peace than by the pressure of war. “At
Birmingham,” said the “Courier,” “an immense assemblage witnessed
the arrival of the mail, and immediately took the horses out, and
drew the mail to the post-office with the loudest acclamations,”—
acclamations over a treaty universally regarded as discreditable.
The “Times” admitted the general joy, and denied only that it was
universal. If the “Times” in any degree represented public opinion,
the popular satisfaction at the peace was an extraordinary political
symptom, for in its opinion the Government had accepted terms
such as “might have been expected from an indulgent and liberal
conqueror.... We have retired from the combat,” it said, December
30, “with the stripes yet bleeding on our back,—with the recent
defeats at Plattsburg and on Lake Champlain unavenged.” During
several succeeding weeks the “Times” continued its extravagant
complaints, which served only to give the Americans a new idea of
the triumph they had won.
In truth, no one familiar with English opinion during the past ten
years attempted to deny that the government of England must admit
one or the other of two conclusions,—either it had ruinously
mismanaged its American policy before the war, or it had disgraced
itself by the peace. The “Morning Chronicle,” while approving the
treaty, declared that the Tories were on this point at odds with their
own leaders:[74] “Their attachment to the ministers, though strong,
cannot reconcile them to this one step, though surely if they would
look back with an impartial eye on the imbecility and error with
which their idols conducted the war, they must acknowledge their
prudence in putting an end to it. One of them very honestly said,
two days ago, that if they had not put an end to the war, the war
would have put an end to their Ministry.” Whatever doubts existed
about the temper of England before that time, no one doubted after
the peace of Ghent that war with the United States was an
unpopular measure with the British people.
Nevertheless the “Times” and the Tories continued their
complaints until March 9, when two simultaneous pieces of news
silenced criticism of the American treaty. The severe defeat at New
Orleans became known at the moment when Napoleon, having
quitted Elba, began his triumphal return to Paris. These news,
coming in the midst of Corn Riots, silenced further discussion of
American relations, and left ministers free to redeem at Waterloo the
failures they had experienced in America.
In the United States news of peace was slow to arrive. The British
sloop-of-war “Favorite” bore the despatches, and was still at sea
when the month of February began. The commissioners from
Massachusetts and Connecticut, bearing the demands of the
Hartford Convention, started for Washington. Every one was intent
on the situation of New Orleans, where a disaster was feared.
Congress seemed to have abandoned the attempt to provide means
of defence, although it began another effort to create a bank on
Dallas’s plan. A large number of the most intelligent citizens believed
that two announcements would soon be made,—one, that New
Orleans was lost; the other, that the negotiation at Ghent had ended
in rupture. Under this double shock, the collapse of the national
government seemed to its enemies inevitable.
In this moment of suspense, the first news arrived from New
Orleans. To the extreme relief of the Government and the
Republican majority in Congress, they learned, February 4, that the
British invasion was defeated and New Orleans saved. The victory
was welcomed by illuminations, votes of thanks, and rejoicings
greater than had followed the more important success at Plattsburg,
or the more brilliant battles at Niagara; for the success won at New
Orleans relieved the Government from a load of anxiety, and
postponed a crisis supposed to be immediately at hand. Half the
influence of the Hartford Convention was destroyed by it; and the
commissioners, who were starting for the capital, had reason to
expect a reception less favorable by far than they would have met
had the British been announced as masters of Louisiana. Yet the
immediate effect of the news was not to lend new vigor to Congress,
but rather to increase its inertness, and to encourage its dependence
on militia, Treasury notes, and good fortune.
A week afterward, on the afternoon of Saturday, February 11, the
British sloop-of-war “Favorite” sailed up New York harbor, and the
city quickly heard rumors of peace. At eight o’clock that evening the
American special messenger landed, bringing the official documents
intrusted to his care; and when the news could no longer be
doubted, the city burst into an uproar of joy. The messenger was
slow in reaching Washington, where he arrived only on the evening
of Tuesday, February 13, and delivered his despatches to the
Secretary of State.
Had the treaty been less satisfactory than it was, the President
would have hesitated long before advising its rejection, and the
Senate could hardly have gained courage to reject it. In spite of
rumors from London and significant speculations on the London
Exchange, known in America in the middle of January, no one had
seriously counted on a satisfactory peace, as was proved by the
steady depression of government credit and of the prices of
American staples. The reaction after the arrival of the news was
natural, and so violent that few persons stopped to scrutinize the
terms. Contrary to Clay’s forebodings, the treaty, mere armistice
though it seemed to be, was probably the most popular treaty ever
negotiated by the United States. The President sent it to the Senate
February 15; and the next day, without suggestion of amendment,
and apparently without a criticism, unless from Federalists, the
Senate unanimously confirmed it, thirty-five senators uniting in
approval.
Yet the treaty was not what the Government had expected in
declaring the war, or such as it had a right to demand. The
Republicans admitted it in private, and the Federalists proclaimed it
in the press. Senator Gore wrote to Governor Strong:[75] “The treaty
must be deemed disgraceful to the Government who made the war
and the peace, and will be so adjudged by all, after the first
effusions of joy at relief have subsided.” Opinions differed widely on
the question where the disgrace belonged,—whether to the
Government who made the war, or to the people who refused to
support it; but no one pretended that the terms of peace, as far as
they were expressed in the treaty, were so good as those repeatedly
offered by England more than two years before. Yet the treaty was
universally welcomed, and not a thought of continued war found
expression.
In New England the peace was received with extravagant delight.
While the government messenger who carried the official news to
Washington made no haste, a special messenger started from New
York at ten o’clock Saturday night, immediately on the landing of the
government messenger, and in thirty-two hours arrived in Boston.
Probably the distance had rarely been travelled in less time, for the
Boston “Centinel” announced the expense to be two hundred and
twenty-five dollars; and such an outlay was seldom made for rapidity
of travel or news. As the messenger passed from town to town he
announced the tidings to the delighted people.[76] Reaching the
“Centinel” office, at Boston, early Monday morning, he delivered his
bulletin, and a few minutes after it was published all the bells were
set ringing; schools and shops were closed, and a general holiday
taken; flags were hoisted, the British with the American; the militia
paraded, and in the evening the city was illuminated. Yet the terms
of peace were wholly unknown, and the people of Massachusetts
had every reason to fear that their interests were sacrificed for the
safety of the Union. Their rejoicing over the peace was as
unreasoning as their hatred of the war.
Only along the Canadian frontier where the farmers had for three
years made large profits by supplying both armies, the peace was
received without rejoicing.[77] South of New York, although less
public delight was expressed, the relief was probably greater than in
New England. Virginia had suffered most, and had felt the blockade
with peculiar severity. A few weeks before the treaty was signed,
Jefferson wrote:[78]—
“By the total annihilation in value of the produce which was to give
me sustenance and independence, I shall be like Tantalus,—up to the
shoulders in water, yet dying with thirst. We can make indeed enough
to eat, drink, and clothe ourselves, but nothing for our salt, iron,
groceries, and taxes which must be paid in money. For what can we
raise for the market? Wheat?—we only give it to our horses, as we
have been doing ever since harvest. Tobacco?—it is not worth the
pipe it is smoked in.”
While all Virginia planters were in this situation February 13, they
awoke February 14 to find flour worth ten dollars a barrel, and
groceries fallen fifty per cent. They were once more rich beyond
their wants.
So violent and sudden a change in values had never been known
in the United States. The New York market saw fortunes disappear
and other fortunes created in the utterance of a single word. All
imported articles dropped to low prices. Sugar which sold Saturday
at twenty-six dollars a hundred-weight, sold Monday at twelve
dollars and a half. Tea sank from two dollars and a quarter to one
dollar a pound; tin fell from eighty to twenty-five dollars a box;
cotton fabrics declined about fifty per cent. On the other hand flour,
cotton, and the other chief staples of American produce rose in the
same proportion. Nominally flour was worth seven and a half dollars
on Saturday, though no large amounts could have been sold; on
Monday the price was ten dollars, and all the wheat in the country
was soon sold at that rate.
Owing to the derangement of currency, these prices expressed no
precise specie value. The effect of the peace on the currency was for
a moment to restore an apparent equilibrium. In New York the
specie premium of twenty-two per cent was imagined for a time to
have vanished. In truth, United States six-per-cents rose in New York
from seventy-six to eighty-eight in paper; Treasury-notes from
ninety-two to ninety-eight. In Philadelphia, on Saturday, six-per-
cents sold at seventy-five; on Monday, at ninety-three. The paper
depreciation remained about twenty per cent in New York, about
twenty-four per cent in Philadelphia, and about thirty per cent in
Baltimore. The true value of six-per-cents was about sixty-eight; of
Treasury notes about seventy-eight, after the announcement of
peace.
As rapidly as possible the blockade was raised, and ships were
hurried to sea with the harvests of three seasons for cargo; but
some weeks still passed before all the operations of war were
closed. The news of peace reached the British squadron below
Mobile in time to prevent further advance on that place; but on the
ocean a long time elapsed before fighting wholly ceased.
Some of the worst disasters as well as the greatest triumphs of
the war occurred after the treaty of peace had been signed. The
battle of New Orleans was followed by the loss of Fort Bowyer. At
about the same time a British force occupied Cumberland Island on
the southern edge of the Georgia coast, and January 13 attacked the
fort at the entrance of the St. Mary’s, and having captured it without
loss, ascended the river the next day to the town of St. Mary’s,
which they seized, together with its merchandise and valuable ships
in the river. Cockburn established his headquarters on Cumberland
Island January 22, and threw the whole State of Georgia into
agitation, while he waited the arrival of a brigade with which an
attack was to be made on Savannah.
The worst disaster of the naval war occurred January 15, when
the frigate “President”—one of the three American forty-fours, under
Stephen Decatur, the favorite ocean hero of the American service—
suffered defeat and capture within fifty miles of Sandy Hook. No
naval battle of the war was more disputed in its merits, although its
occurrence in the darkest moments of national depression was
almost immediately forgotten in the elation of the peace a few days
later.
Secretary Jones retired from the Navy Department Dec. 19, 1814,
yielding the direction to B. W. Crowninshield of Massachusetts, but
leaving a squadron ready for sea at New York under orders for
distant service. The “Peacock” and “Hornet,” commanded by
Warrington and Biddle, were to sail with a store-ship on a long
cruise in Indian waters, where they were expected to ravage British
shipping from the Cape of Good Hope to the China seas. With them
Decatur was to go in the “President,” and at the beginning of the
new year he waited only an opportunity to slip to sea past the
blockading squadron. January 14 a strong westerly wind drove the
British fleet out of sight. The “President” set sail, but in crossing the
bar at night grounded, and continued for an hour or more to strike
heavily, until the tide and strong wind forced her across. Decatur
then ran along the Long Island coast some fifty miles, when he
changed his course to the southeast, hoping that he had evaded the
blockading squadron. This course was precisely that which Captain
Hayes, commanding the squadron, expected;[79] and an hour before
daylight the four British ships, standing to the northward and
eastward, sighted the “President,” standing to the southward and
eastward, not more than two miles on the weather-bow of the
“Majestic,”—the fifty-six-gun razee commanded by Captain Hayes.
The British ships promptly made chase. Captain Hayes’s squadron,
besides the “Majestic,” consisted of the “Endymion,” a fifty-gun
frigate, with the “Pomone” and “Tenedos,” frigates like the
“Guerriere,” “Macedonian,” and “Java,” armed with eighteen-pound
guns. Only from the “Endymion” had Decatur much to fear, for the
“Majestic” was slow and the other ships were weak; but the
“Endymion” was a fast sailer, and especially adapted to meet the
American frigates. The “Endymion,” according to British authority,
was about one hundred and fifty-nine feet in length on the lower
deck, and nearly forty-three feet in extreme breadth; the “President,”
on the same authority, was about one hundred and seventy-three
feet in length, and forty-four feet in breadth. The “Endymion” carried
twenty-six long twenty-four-pounders on the main deck; the
“President” carried thirty. The “Endymion” mounted twenty-two
thirty-two pound carronades on the spar deck; the “President”
mounted twenty. The “Endymion” had also a long brass eighteen-
pounder as a bow-chaser; the “President” a long twenty-four-
pounder as a bow-chaser, and another as a stern-chaser. The
“Endymion” was short-handed after her losses in action with the
“Prince de Neufchatel,” and carried only three hundred and forty-six
men; the “President” carried four hundred and fifty. The “Endymion”
was the weaker ship, probably in the proportion of four to five; but
for her immediate purpose she possessed a decisive advantage in
superior speed, especially in light winds.
At two o’clock in the afternoon, the “Endymion” had gained so
much on the “President” as to begin exchanging shots between the
stern and bow-chasers.[80] Soon after five o’clock, as the wind fell,
the “Endymion” crept up on the “President’s” starboard quarter, and
“commenced close action.”[81] After bearing the enemy’s fire for half
an hour without reply, Decatur was obliged to alter his course and
accept battle, or suffer himself to be crippled.[82] The battle lasted
two hours and a half, until eight o’clock, when firing ceased; but at
half-past nine, according to the “Pomone’s” log, the “Endymion” fired
two guns, which the “President” returned with one.[83] According to
Decatur’s account the “Endymion” lay for half an hour under his
stern, without firing, while the “President” was trying to escape. In
truth the “Endymion” had no need to fire; she was busy bending
new sails, while Decatur’s ship, according to his official report, was
crippled, and in the want of wind could not escape.
In a letter written by Decatur to his wife immediately after the
battle, he gave an account of what followed, as he understood it.[84]
“The ‘Endymion,’” he began, ... “was the leading ship of the enemy.
She got close under my quarters and was cutting my rigging without
my being able to bring a gun to bear upon her. To suffer this was
making my capture certain, and that too without injury to the enemy.
I therefore bore up for the ‘Endymion’ and engaged her for two hours,
when we silenced and beat her off. At this time the rest of the ships
had got within two miles of us. We made all the sail we could from
them, but it was in vain. In three hours the ‘Pomone’ and ‘Tenedos’
were alongside, and the ‘Majestic’ and ‘Endymion’ close to us. All that
was now left for me to do was to receive the fire of the nearest ship
and surrender.”
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