The Genera of Lactic Acid Bacteria The Lactic Acid Bacteria Volume 2 The Genera of Lactic Acid Bacteria Edited by B.J.B. WOOD Department of Bioscience and Biotechnology University of Strathclyde Glasgow UK and W.H. HOLZAPFEL Federal Research Centre for Nutrition Institute of Hygiene and Toxicology Karlsruhe Germany mJ SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. First edition 1995 © 1995 Springer Science+Business Media Dordrecht Originally published by Chapman & Hali in 1995 Softcover reprint ofthe hardcover lst edition 1995 Typeset in 10/12pt Times by Cambrian Typesetters, Frimley, Surrey Cover photograph is a scanning electron micrograph of Lactococcus lactis subsp. lactis Bu2-60. (Courtesy of Horst Neve, Kiel, Germany.) ISBN 978-1-4613-7666-8 ISBN 978-1-4615-5817-0 (eBook) DOI 10.1007/978-1-4615-5817-0 Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the Iicences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries conceming reproduction outside the terms stated here should be sent to the publishers at the Glasgow address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. A catalogue record for this book is available from the British Library Library of Congress Catalog Card Number: 94-73502 9 Printed on permanent acid-free text paper, manufactured in accordance with ANSIINISO Z39.48-1992 (permanence of Paper). Series preface The Lactic Acid Bacteria is planned as a series in a number of volumes, and the interest shown in it appears to justify a cautious optimism that a series comprising at least five volumes will appear in the fullness of time. This being so, I feel that it is desirable to introduce the series by providing a little of the history of the events which culminated in the decision to produce such a series. I also wish to indicate the boundaries of the group 'The Lactic Acid Bacteria' as I have defined them for the present purposes, and to outline my hopes for future topics in the series. Historical background lowe my interest in the lactic acid bacteria (LAB) to the late Dr Cyril Rainbow, who introduced me to their fascinating world when he offered me a place with him to work for a PhD on the carbohydrate metabolism of some lactic rods isolated from English beer breweries by himself and others, notably Dr Dora Kulka. He was particularly interested in their preference for maltose over glucose as a source of carbohydrate for growth, expressed in most cases as a more rapid growth on the disaccharide; but one isolate would grow only on maltose. Eventually we showed that maltose was being utilised by 'direct fermentation' as the older texts called it, specifically by the phosphorolysis which had first been demonstrated for maltose by Doudoroff and his associates in their work on maltose metabolism by a strain of Neisseria meningitidis. I began work on food fermentations when I came to Strathclyde University, and I soon found myself involved again with the bacteria which I had not touched since completing my doctoral thesis. In 1973 J.G. Carr, C.V. Cutting and G.C. Whiting organised the 4th Long Ashton Symposium Lactic Acid Bacteria in Beverages and Food, and from my participation in that excellent conference arose a friendship with Geoff Carr. The growing importance of these bacteria was subsequently confirmed by the holding, a decade later, of the first of the Wageningen Conferences on the LAB. Discussions about the LAB, and the fact that they were unusual and important enough to have entire conferences devoted to them, with Mr George Olley of Applied Science Publishers Ltd (who had proposed the idea which resulted in the production of Microbiology of Fermented Foods) gave rise to the idea that there might be scope for a book on them. vi THE GENERA OF LACTIC ACID BACTERIA Subsequent discussions with Geoff Carr refined 'a book' into 'a multi­ volume series' and rather slowly Geoff and I began to lay rough plans for such a series, a process greatly helped by our preference for planning sessions in pubs. Sadly, Geoff died before the plans reached fruition, but his thinking contributed much to my development of the ideas which we hatched together, and I think that he would have approved of the final form which the series is taking. I have tried to achieve a multi-national spread among the authors, which would certainly have met with his approval. I have also tried to achieve a direct and clear style of writing through my directions to authors, and my editing, as both he and Cyril Rainbow valued clarity very highly, and deplored the sort of scientific writing which seemed to have been generated under the delusion that proper science should demonstrate its importance by being as obscure as possible. The lactic acid bacteria Most of us think that we know what we mean when we use the expression 'lactic acid bacteria'. We tend to think of them as a reasonably coherent group, and this is convenient for a great many purposes. In fact, however, the LAB remind me of a cloud in the sky, which can look rather solid and well-bounded when viewed from a distance but becomes more and more fuzzy and intangible the closer that one approaches to it. The LAB are all Gram-positive; anaerobic, micro-aerophilic or aero-tolerant; catalase negative; rods or cocci; most importantly they all produce lactic acid as the sole, major or an important product from the energy-yielding fermentation of sugars. It used to be thought that all LAB were non-motile and non­ sporing, although we now have the Sporolactobacilli, and motile organisms are reported which would otherwise fit with the LAB. Listeria species produce lactic acid as a major metabolic product, but I doubt if many people working with the LAB as normally understood, would wish to admit Listeria to membership of the club. The LAB as a group had a 'squeaky clean' image, with such adverse effects as have been reported being, in general, ascribed to unusual circumstances such as abnormal immune status. This despite the Streptococci, as originally defined, including both beneficial and harmful organisms. Yet Listeria is closely related (according to modern classifications) to undoubted LAB, and Streptococcus as now defined seems to consist almost entirely of rather disagreeable specimens, so it seems a little irrational to include one of these genera and exclude the other; I can offer no real defence of my decision to do so despite this evidently logical argument, other than the hoary old one of 'accepted practice'. After much discussion, and with due acknowledgement of the rapidly evolving situation which is leading to the SERIES PREFACE VB establishment of numerous new genera, the genera which receive chapters in Volume 2 will be regarded as the principal ones for the purposes of this series. This whilst acknowledging the right of individual authors to define the LAB in the manner which seems most appropriate to the story which they have to tell, and recognising that there will be well-established additional genera of LAB before the series has reached its conclusion. Future plans Future plans will inevitably be modified in the light of experience gained as the series develops, not least the feedback from readers and the comments and suggestions of reviewers. At present the following volumes are envisaged (not necessarily in the order listed: Genetics of the LAB. Physiology and Biochemistry. Uses. This volume will cover their role in food fermentations, lactic acid production, vitamin assays, etc. H is hoped and believed that together these volumes will form a lasting primary source and, as the computer experts would say, 'benchmark' against which future progress in our understanding of this diverse group of bacteria can be measured. Brian J.B. Wood Preface There is very little to say about this volume which can usefully add to the material in chapter 1. I would, however, like to take the opportunity to thank various people, not least my co-editor Professor Wilhelm Holzapfel. While I recognise the paramount importance of taxonomy as the map by which we find our way around the confusing worlds of biology, I must admit that my grasp of the finer points of this discipline is not always secure. Wilhelm's hand on the tiller was always more confident than mine could ever be, but perhaps even more important was his remarkably extensive network of contacts. I am certain that his standing contributed significantly to our success in persuading busy people to give up their time to write difficult reviews. He also contributed significantly to my hitherto disgracefully limited understanding of the newer methods which are so changing both the methods employed in taxonomical investigations and (as a consequence thereof) our appreciation of the subtle relationships between, and even the evolution of, the kingdom Prokaryota. Preparation of this book began with Elsevier Applied Science, and I was very pleased that it was one of the titles selected by Chapman and Hall when the transfer to them took place.