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The Biochemistry of Methylotrophs
C. Anthony (1982) Academic Press, London  

This textbook is no longer in print.
It contains 350 pages, 60 pages of Tables, 60 pages of Figures, a 40 page index and about 1000 references [every methylotroph reference up to 1981]. Copyright exclusive to author.   Contents are summarised below.

I have produced a CD of the book with all pages in both jpg and pdf formats which can be searched in Acrobat Reader.
A copy of the CD is available [£25 or $50].
Email: C.Anthony@soton.ac.uk

  For complete book click on picture
Dedicated to K. G. Wiles, L. J. Zatman and J. R. Quayle, FRS


Foreword by Professor J.R. Quayle FRS

There are two kinds of organism which stand out as the biosynthetic virtuosi
of the living world—the autotrophs and the methylotrophs. The autotrophs
are organisms which can synthesize all their cell constituents from carbon
dioxide as sole carbon source; methylotrophs can perform such total biosynthesis
from reduced carbon compounds containing one or more carbon atoms but
containing no carbon-carbon bonds. Autotrophs have long been the subject of
intense study, if for no other reason than that they include the green
plants as well as microorganisms. Although the first methylotroph, Bacillus
methanicus, was discovered 76 years ago growing aerobically on methane, the
next 40 to 50 years saw the discovery of only a few other methylotrophs
growing on reduced C-1 compounds such as methane, methanol, methylamine
and formate. They were regarded as somewhat exotic organisms and received
scant biochemical attention; it was often speculated that they might merely
couple the energy of oxidation of the C-1 substrate to the reduction of carbon
dioxide and thus in effect be chemosynthetic autotrophs. If this had turned out
to be the case, the subject of methylotrophy would have remained but a minor
variant on a familiar theme. However, it was recognised in the 1950s that
although some methylotrophs were indeed autotrophs in thin disguise, many
other methylotrophic organisms assimilated C-1 units at levels more reduced
than carbon dioxide by unknown assimilatory processes. As a result an
explosive development of this whole field then took place. Biochemists espied
virgin areas of intermediary metabolism and energy transduction, the range of
methylotrophic microorganisms was greatly extended by development of new
enrichment procedures and the new organisms were ripe for genetic analysis
and manipulation. At the same time biotechnologists realized that methane
and methanol offered entirely new industrial possibilities as cheap and
readily available fermentation feedstocks, thus providing an additional
stimulus for research and development.

A wealth of publications resulted from all this activity and for 20 years this
burgeoning literature has been reviewed at frequent intervals by various
investigators. With many of the major biochemical problems now solved, and
the remaining ones clearly defined, it has been clear for some time that there
was an urgent need for a definitive and wide-ranging textbook on the subject.
The present book has been written to meet such a demand and it is hoped that
it will serve as a standard reference book for all aficionados of this field,
whether they be newcomers or veterans. The newcomers can be assured that
it has been written by a biochemist who, since 1960, has himself been a
highly active investigator and an authority on methylotrophy; the veterans
will not need any such prefatory assurance from me.

Sheffield 1982 J. R. Quayle

My Preface
In 1960, when I started working with methylotrophs I had half a dozen
reprints (the total available) on these organisms. The growth of the literature
since then has been exponential, and the decision to write this book is based
on the well-established practice of harvesting in the mid-log phase; this is in
order to avoid being completely submerged beneath the methylotrophic
A preface usually defines the limits of the author's intentions; the ground
covered and the likely readership. I shall do likewise; this book covers
everything to do with the biochemistry of methylotrophs and it is written to
suit everyone.
Perhaps I had better attempt to justify this bold claim. My first aim was
to cover all aspects of methylotrophy in the detail required by anyone active
in research in this field, whether they be microbiologists, microbial physiologists,
"metabolic" biochemists or enzymologists. Such an aim, however,
might tend to diminish the usefulness of a book to those using it as an
introduction to any aspect of the subject; this is because more fundamental
aspects can be obscured by detail, and submerged beneath lines of references
to the original sources. I have tried to avoid this in two ways. When possible
I have arranged references in separate tables so as to leave the text clear.
More important, each chapter starts with a descriptive section which I have
tried to make comprehensible to a senior undergraduate. The evidence for
conclusions initially presented is then discussed in separate sections. When
in doubt I have erred, I hope, on the side of too much explanation rather than
too little.
As the subject of the book is biochemistry, the first chapter was to have
been a very brief introductory guide to the methylotrophs themselves.
This intention was abandoned when I realised that the biochemists for whom
it was written could probably cope with any biochemistry they might come
across in the literature, but might find the microbes themselves bewildering
beyond belief. I hope that the first chapter now provides a complete account
of the methylotrophic bacteria in a way that allows a ready evaluation of
which are likely to be biochemically similar and which are probably different.
Chapters 2, 3, 4 and 5 cover the carbon assimilation pathways of methylo.-
trophic bacteria. The pathways, and evidence for them, are first described;
this is followed by summaries of the distribution of the pathways amongst
the methylotrophs; the individual enzymes are then described and regulation
of the assimilation pathways discussed.
Chapters 6 and 7 deal with the enzymes (many of them novel) involved
in the oxidation of methane, methanol and methylated amines.
Chapter 8 is devoted to electron transport and energy transduction; and
Chapter 9 attempts to interrelate all the biochemistry discussed in Chapters
2 to 8 in a discussion of growth yields and bioenergetics in methylotrophs.
Because the biochemistry of oxidation and assimilation of methanol in
yeasts differs in most important respects from that in methylotrophic bacteria,
Chapter 10 is exclusively concerned with metabolism in methylotrophic
Chapter 11 is a self-contained chapter on methanogens and methanogenesis.
Although few methanogens are actually methylotrophs, the recent rapid
advances in our knowledge of the unique biochemistry of these "archaebacteria"
made the temptation to include this complete review irresistible.
Chapter 12 deals with the commercial exploitation of methylotrophs.
The biochemistry of microbes growing on methane or methanol makes them
especially suitable for commercial exploitation; this includes their use as
single cell protein (SCP); their use for the production of carboxylic acids,
amino acids and vitamins; their use as biocatalysts; and their use in electroenzymology and biofuel cells. Although not dealing with detailed practical
aspects of such processes, this chapter aims to provide a foundation for
understanding this rapidly-developing area of biotechnology.
February 1982 C. Anthony

My first acknowledgement must be an expansion of the Dedication of this
book, which is to K. G. Wiles, L. J. Zatman and J. R. Quayle, FRS.
"Willy" Wiles was my biology teacher in the sixth form at Watford Boys'
Grammar School; his enthusiastic eccentric insistence that Life is "interesting",
and worth some effort in understanding, was both exceptional
and captivating. Len Zatman was first my tutor then my Ph.D. supervisor
in the Microbiology Department at Reading University, and in spite of this
he has remained a good friend. Rod Quayle is, of course, the "Godfather" of
methylotrophy, honoured above all men by the international community of
methylophiles. To him I owe a second thanks for generously agreeing to
write a Foreword to "The Biochemistry of Methylotrophs".
I have received a great deal of help in the preparation and writing of this
book, not least from the many scientists who have kindly sent me reprints
and manuscripts prior to publication. Pat Goodwin (nee Dunstan), my first
research student, deserves a special thanks for her encouragement and
valuable discussions in the early stages of preparation, and I must also
thank my present research students and assistants who have probably
suffered from both my absence and from my presence during the last two years
(Matthew Beardmore-Gray, Steve Ford, Steve Froud, Ashley Lawton,
David O'Keeffe and Dudley Page). David O'Keeffe has worked with me for
seven years and to him I owe a special debt of gratitude — both for this, and
for his very generous help in keeping the lab running and for accepting many
extra burdens during the final stages of production of this book.
A final, most important word of thanks I owe my wife, Elizabeth, for her
support and encouragement, without which "The Biochemistry of Methylotrophs"
would not have been written.


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