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Probiotics JOIN US ON THE INTERNET VIA WWW, GOPHER, FTP OR EMAil: WWW: GOPHER: FTP: EMAIL: http://www.thomson.com gopher.thomson.com ftp.thomson.com findit@kiosk.thomson.com A service of ICDP® Probiotics Applications and practical aspects Edited by R Fuller Freelance Consultant in Cut Microecology Reading, UK SPRINGER-SCIENCE+BUSINESS MEDIA, B.v http://avaxhome.ws/blogs/ChrisRedfield First edition © 1997 Springer Science+Business Media Dordrecht Originally published by Chapman & Hali in 1997 Softcover reprint ofthe hardcover lst edition 1997 Typeset in 10/12pt Palatino by Acom Bookwork, Salisbury, Wiltshire ISBN 978-94-010-6476-7 ISBN 978-94-011-5860-2 (eBook) DOI 10.1007/978-94-011-5860-2 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 licences 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 London 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 omission that may be made A catalogue record for this book is available from the British Library § Printed on acid-free text paper, manufactured in accordance with ANSI/NISO Z39.48-1992 (Permanence of Paper) Contents Introduction R Fuller 1.1 Development of commercial preparations 1.2 Factors affecting the response 1.3 Future developments 1.4 Conclusions 1.5 References Probiotics and intestinal infections G.R Gibson, J.M Saavedra, S MacFarlane and G T MacFarlane 2.1 Introduction 2.2 Human colonic microbiota and homeostasis 2.3 Infections of the intestinal tract 2.4 Attachment 2.5 Use of probiotics against intestinal infections 2.6 Probiotics and viral infections of the gastrointestinal tract 2.7 Modulation of the host response to infection 2.8 Effects of probiotics in infants 2.9 Prebiotics and synbiotics 2.10 References Antibiotic-associated diarrhoea: treatments by living organisms given by the oral route (probiotics) G eorthier 3.1 Introduction 3.2 Antibiotic-associated diarrhoea and pseudomembranous colitis 3.3 Treatments by living organisms 3.4 Search for new oral treatments by probiotics 3.5 Acknowledgements 3.6 References Lactose maldigestion P Marteau, T Vesa and J.e Rambaud 4.1 Introduction 1 8 10 10 11 18 19 22 24 27 27 29 31 40 40 40 48 59 60 60 65 65 vi Contents 4.2 Lactose metabolism 4.3 Methods to study lactose digestion 4.4 Digestion and tolerance of fermented dairy products in lactose-intolerant subjects 4.5 Mechanisms for the better tolerance and digestion of lactose from fermented dairy products 4.6 Clinical applications of probiotics or fermented milks in the field of lactose mal digestion or intolerance 4.7 Comments and conclusions 4.8 References 65 71 73 79 83 84 86 Antimutagenic and antitumour activities of lactic acid bacteria A Hosono, H Kitazawa and T Yamaguchi 5.1 Introduction 5.2 Antimutagenic activity of lactic acid bacteria 5.3 Antitumour activity of lactic acid bacteria 5.4 References 89 90 105 126 Stimulation of immunity by probiotics 133 G Famularo, S Moretti, S Marcellini and C De Simone 6.1 6.2 6.3 6.4 6.5 6.6 6.7 6.8 6.9 6.10 6.11 6.12 6.13 6.14 6.15 6.16 6.17 Introduction Microflora, probiotics and immunity Endogenous microflora Morphology of gut-associated lymphoid tissue Intra-epithelial lymphocyte compartment Lamina propria lymphocyte compartment Antigen presentation in the gut Effector response in the gut Modulation of gut functions by immune networks Modulation of cytokine production by probiotics Modulation of gut-associated lymphoid tissue by probiotics Modulation of macrophage functions by probiotics Modulation of resistance towards Salmonella typhimurium by probiotics Modulation of autoimmunity by probiotics Role of probiotics in the treatment of human infections Pathogenic potential of probiotics References Probiotics in cattle J.T Huber 7.1 Introduction 7.2 Baby calves 7.3 Beef cattle 89 133 134 135 137 138 139 139 141 143 144 148 149 150 151 153 155 156 162 162 163 165 Contents vii 7.4 7.5 7.6 7.7 7.8 Lactating cows Market penetration of probiotics How probiotics work in cattle? Conclusions References Intervention strategies: the use of probiotics and competitive exclusion microfloras against contamination with pathogens in pigs and poultry 169 177 177 180 181 186 R.W.A.W Mulder, R Havenaar and J.H.J Huis in 't Veld 8.1 8.2 8.3 8.4 8.5 8.6 8.7 Index Introduction Intestinal flora and gut metabolism Microflora and colonization resistance Use of probiotics Use of competitive exclusion microfloras in poultry Summary References 186 189 190 191 202 204 205 209 Contributors R Fuller, Russet House, 59 Ryeish Green, Three Mile Cross, Reading, RG71ES, UK G.R Gibson, Institute of Food Research, Early Gate, Whiteknights Road, Reading, RG6 2EF, UK J.M Saavedra, Johns Hopkins Hospital, 6000 North Wolfe Street, Brady 320, Baltimore MD 2128-2631, USA S Macfarlane and G.T Macfarlane, Medical Research Council, Dunn Clinical Nutrition Centre, Cambridge, UK G eorthier, Institut National de la Recherche Agronomique, Unite d'Ecologie et de Physiologie du System Digestif, 78352 Jouy-en-Josas Cedex, France P Marteau, T Vesa and J.-c Rambaud, Hopital Saint-Lazare, 107 bis, rue du fg Saint-Denis, 75475 Paris Cedex, France A Hosono, Shinshu University, Minamiminouwa, Nagano, 399-45, Japan H Kitazawa and T Yamaguchi, Department of Animal Science, Faculty of Agriculture, Tohoku University, Sendai-Aobaku, 981, Japan G Famularo, S Moretti, S Marcellini and C De Simone, Universita de L' Aquila, Cathedra di Malattie Infettive, 671000 L'Aquila, Italy J.T Huber, Department of Animal Sciences, University of Arizona, 205 Shantz Building, Tucson, Arizona 85721, USA R.W.A.W Mulder, DLO Institute of Animal Sciences and Health, Wageningen, The Netherlands R Havenaar, TNO Nutrition and Food Research Institute, Zeist, The Netherlands J.H.J Huis in 't Veld, Department of the Science of Food of Animal Origin, University of Utrecht, The Netherlands CHAPTER Introduction R Fuller 1.1 DEVELOPMENT OF COMMERCIAL PREPARATIONS The history of the probiotic effect has been well documented many times previously (see e.g Bibel, 1982; Fuller, 1992) The consumption of fermented milks dates from pre-biblical times but the probiotic concept was born at the end of the last century with the work of Metchnikoff at the Pasteur Institute in Paris In the century that has elapsed since Metchnikoff's work, the probiotic concept has been accepted by scientists and consumers throughout the world Attempts to refine the practice from the use of traditional soured milks to preparations containing specific microorganisms have occupied the thoughts and endeavours of scientists in many different countries But, in spite of the large amount of effort expended in attempting to explain and define the effect, it has to be admitted that little is known of the way in which probiotics operate There are likely to be several different mechanisms because it seems highly improbable that a mode of action that explains resistance to microbial infection will also hold true for improved milk production or alleviation of lactose malabsorption The lack of fundamental knowledge about the mechanism of the probiotic effect has not deterred the development of a great many probiotic preparations destined for treatment of various conditions in man and animals There are currently over 20 products on the market in the UK The dearth of basic information about the probiotic effect has meant that much of the development has been empirical and not always based on sound scientific principles One factor that has been used in the selection of probiotic cultures has been the ability to adhere to gut epithelial cells of the animal to which the probiotic is being fed R Fuller (ed.), Probiotics © Springer Science+Business Media Dordrecht 1997 Introduction There was a good correlation found between in vitro and in vivo results for adhesion of different strains of bifidobacteria (Crociani et al., 1995) Adhesion is now generally accepted as an important colonization factor and since establishment in the intestine is an essential prerequisite of effective probiotic activity, it is to be applauded as the first step in a rational approach to the selection of micro-organisms for inclusion in probiotic preparations However, it should be appreciated that attachment is not an essential attribute of a successful probiotic organism Rapid growth can achieve the same end In some cases such as fungi (Saccharomyces cerevisiae and Aspergillus oryzae) the effect is gained without either attachment or rapid growth; mere survival is adequate Under these conditions continuous administration is required for the maximum realization of the probiotic effect The numerous probiotic products on the market claim to have many different effects including improved resistance to infectious disease, antitumour activity, increased growth rate and feed conversion in farm animals, improved milk production by cows and increased egg production by poultry The range of micro-organisms contained in the probiotic preparations is wide, comprising bacteria, moulds and yeasts By far the most frequently used of these three groups are the bacteria with lactic acid bacteria (lactobacilli, streptococci, enterococci and bifidobacteria) the predominant genera The emphasis on the lactic acid bacteria stems partly from the fact that there is evidence that the lactic acid bacteria occupy a central role in the gut flora, which enables them to influence the composition of the flora to the benefit of the host This has been elegantly demonstrated by the work of Tannock and his group in New Zealand By developing a population of mice with a lactobacillus-free gut microflora, they have been able to study the way in which the gut lactobacilli can affect the metabolism and growth of the host animal which, in this case, is the mouse Their results are summarized in Table 1.1 The use of lactic acid bacteria was also stimulated by the early work of Metchnikoff who espoused the view that the normal gut microflora Table 1.1 Effect of lactobacilli on mice (from Tannock, 1995) Increased bile salt hydrolase activity More unconjugated bile acids Growth rate unaffected Azoreductase activity reduced P-Glucuronidase activity reduced Enzyme activities associated with duodenal enterocytes unaffected Serum cholesterol concentration unaffected 197 Use of probiotics 4_1 10 Days (prob: day 10) 13 15 17 Fig 8.5 Average log numbers of Enterococcus faecium (cfu g-l ileal chyme) in ileal samples from fistulated piglets (n = 4) in a group feeding the probiotic LBCME (5 x 106 E jaecium/g feed) from day to 10 (e) and a group feeding LBCME in combination with tylosin (50 ppm) from day to 10 (A) in comparison to a control group (0) without probiotic and tylosin (Havenaar, 1993) gastrointestinal fluids, it can be suggested that Ent faecium multiplies in the small intestine by a factor of 50-100, while B toyoi showed no increase or decrease After the probiotic was left out from the feed, the numbers of both bacterial species dropped to the basic levels, which indicates that both probiotic strains did not colonize the small intestine Samples from the mucosa of the small intestine showed that Ent faecium was associated with the mucosa side, in contrast to B toyoi, which could not be isolated from the mucosa, but only from the intestinal contents However, histological slides (light microscopy) did not show adherence of Ent faecium to the brush borders Therefore, the bacteria were probably embedded in the mucus layer If tylosin was added to the feed in combination with the probiotic bacteria, the numbers of Ent faecium did not increase significantly as compared with the basic levels and the numbers in the control group (Fig 8.5) Apparently the growth of Ent faecium in the intestine was inhibited by tylosin The numbers of B toyoi in the ileum were not influenced by tylosin (Fig 8.6) A challenge experiment in piglets showed that the infection with 198 Intervention strategies - 0' -1. -' -' -' -' - - 10 13 15 17 Days (prob: day 10) Fig 8.6 Average log numbers of Bacillus toyoi (cfu g-l ileal chyme) in ileal samples from fistulated piglets (n =4) in a group feeding the probiotic Toyocerin (1 x 105 B toyoi g-l feed) from day to 10 (e) and a group feeding Toyocerin in combination with tylosin (SO ppm) from day to 10 CA) in comparison to a control group (0) without probiotic and tylosin (Havenaar, 1993) enteropathogenic E coli resulted in intestinal disturbances, irrespective of the ingested Ent faecium or B toyoi administered in the feed The increase in the average severity index of the symptoms up to day 915 and the decrease thereafter followed a similar pattern in the infected control group and in both probiotic groups (Fig 8.7) The numbers of haemolytic E coli in the contents (duodenum: 1-6 x 103 ml-1; ileum: 2-8 x 105 ml-1 ) and on the ileum mucosa (5-9 x 104 2) of the small intestine were not significantly different between the challenged control and probiotic groups (Havenaar and Huis in 't Veld, 1993) Field trials with Ent faecium strain 68 (Cernivet-LBC) resulted in reduced numbers of ruiemolytic E coli (0141 K85ab) from herds suffering from coli-enterotoxaemia, but not in a significant effect on the clinical symptoms of the disease (Deprez et al., 1989) The oral administration of Ent faecium to pregnant sows (twice daily) resulted in lower incidences of scouring in the offspring as compared to the control group without probiotic (Maeng, Kim and Shin, 1989) The same strain of Ent faecium also reduced the numbers of pathogenic E coli and the 199 Use of probiotics 12 10 12 15 18 Days after E coli K88 challenge 21 Fig 8.7 Average severity of symptoms (expressed as an index) of (n = 15 per group) after challenge with enteropathogenic Escherichia coli the control group (0), in two groups receiving the probiotic LBC-ME Toyocerin (T) in comparison to a non-E coli-infected control group (0) naar and Huis in 't Veld, 1993) piglets K88 in (e) or (Have- severity of the induced disease in gnotobiotic piglets (Underdahl et al., 1982) However, in this study the animals were only associated with Ent faecium before the challenge with enteropathogenic E coli, while the control group was not associated with any micro-organism before challenge Germfree animals have a strongly reduced development of the intestinal tissue and the immune system The daily administration of another strain of Ent faecium (BI0-4R) to piglets resulted in a significant 20-fold reduction in the numbers of coliforrns in faecal samples (Ozawa et al., 1983) Claims are made that probiotics may optimize the microbial ecology in the gastrointestinal tract and consequently promote the general health of the host and/or the resistance against enteropathogens The majority of the commercial probiotics make claims such as growth promotion, without unequivocal scientific data and information about the underlying mechanism(s) Selection criteria can only be developed if they are based on scientific evidence which provides the explanation for the expected probiotic effect Besides mechanisms, appropriate in vitro and animal models must be available for screening and validation, 200 Intervention strategies respectively Finally, sensitive and reproducible (microbiological) methods are necessary for identification of the specific probiotic strain(s) in a complex microflora and to quantify the expected effects Only in this way can the credibility of the probiotic concept be improved 8.4.2 Use in poultry Table 8.1 gives the results of trials with commercially available probiotic products after continuous (0-4 weeks) administration to broilers through the feed or drinking water This period was chosen because colonization of potentially pathogenic micro-organisms occurs during the early days of life of the broilers and if positive effects of this intervention strategy tool are expected, these should become evident during this period The group of Enterobacteriaceae was chosen as an indicator for the effects on Salmonella bacteria From these experiments it is concluded that the administration of probiotic products, at the given dosage during the 4-week trial period, did not influence body weight The initial numbers of lactobacilli and other organisms present in the probiotic products seem not to exert any effect on the level of contamination in the caecal contents Additional pH measurements in several parts of the gastrointestinal tract showed a large variation and no effect related to the administration of probiotic products could be observed These trials led to the conclusion that probiotic compounds used under these conditions have no effect and are no tool in the intervention strategy against Salmonella colonization (see also Carazzoni et al., 1993) The health of birds with a stable gastrointestinal microflora is maintained by using antibiotics in preventive dosages Low dosages of antibiotics have under certain conditions shown to be effective as growth promotors, but these low dosages have negative effects on lactobacilli and other lactic acid-producing bacteria Lactobacilli constitute the largest part of aerobic microflora in the gastrointestinal tract and are very susceptible to antibiotics To decrease the use of antibiotics, which may result in bacterial resistance and in formation of residues in organs and tissues of treated birds, the use of probiotics should be considered There are some differences in the time-span in which action occurs (Antibiotics are active in the short term, whereas the effects of probiotics may last days or weeks.) The name 'probiotics' is confusing and some years ago the suggestion, by a panel of European scientists, was made to call these products Ecological Health Control Products Lactobacillus reuteri could be the sole organism able to exert colonization resistance L reuteri is the only Lactobacillus species common to the gastrointestinal tract of humans and animals L reuteri produces and 201 Use of probiotics Table 8.1 Commercially available probiotics and their effects on (a) bodyweight and (b) Enterobacteriaceae and lactobacilli after a 4-week trial (a) Product Micro-organism Dosa~e (kgC) Average weight (kg) CV ('Yo) Fralac FraSacc Gist Brocades BioPlus Streptococcus faecium SF68 Saccharomyces cerevisiae Unknown yeast Bacillus subtilis Bacillus licheniformis Bacillus toyoi Saccharomyces cerevisiae Lactobacillus, Streptococcus, yeast, enzymes Lactobacillus acidophilus Lactobacillus plantarum Lactobacillus, Streptococcus 0.080 0.400 2.000 Q.400 0.500 0.500 1.000 1.1097 1.1095 1.0577 1.1378 1.1092 1.0698 1.1210 10.1 8.8 13.0 7.3 10.7 7.2 11.8 1.000 1.1202 lOA 0.005 0.025 1.0938 1.2021 13.3 9.6 Toyocerin KemPro LactoSacc BioSavor AllLac (b) Product name Control Fralac FraSacc Gist Brocades BioPlus Toyocerin KemPro LactoSacc BioSavor AllLac Enterobacteriaceae contents after week 9.0 >9.0 >9.0 >9.0 >9.0 >9.0 >9.0 >9.0 >9.0 7.9 7.7 7.9 7.7 7.9 6.9 7.3 8.9 8.9 6.9 Lactobacilli contents after week 9.0 5.9 7.0 6.5 6.8 7.0 8.5 7.8 7.8 6.9 5.0 5.2 5.2 6.1 5.3 [...]... 389-95 Fuller, R (19 92) History and development of probiotics, in Probiotics: The Scientific Basis (ed R Fuller) , Chapman & Hall, London, pp 1-8 Gibson, G .R and Roberfroid, M.B (1994) Dietary modulation of the human colonic microbiota: introducing the concept of prebiotics J Nutr ., 12 5, 1401- 12 Hamilton-Miller, J.M.T ., Shap, P and Smith, CT (1996) 'Probiotic' remedies are not what they seem Brit Med J .,. .. 31 2, 55-6 Hughes, V.L and Hillier, S.L (1990) Microbiologic characteristics of Lactobacillus products used for colonisation of the vagina Obst Gynaecol ., 7 5, 24 4-8 Lauland, S (1994) Commercial aspects of formulations, production and marketing of probiotic products, in Human Health: The Contribution of Microorganisms (ed AW Gibson ), Springer, London, pp 159-73 Lin, M.Y ., Savaiano, D and Harlander, S... Bibel, D.J (19 82) Bacterial interference, bacteriotherapy and bacterioprophylaxis, in Bacterial Interference (ed R Aly and H .R Shinefield ), CRC Press, Florida, pp.1- 12 Couwehand, A.c and Conway, P.L (1996) Purification and characterisation of a component produced by Lactobacillus fermentum that inhibits the adhesion of K88 expressing Escherichia coli to porcine ileal mucus J Appl Bacterial ., SO,311-18... Bacteriol ., 7 9, 23 0-6 Pusztai, A, Grant, G ., King, T.P and Clark, E.M.W (1990) Chemical probiosis, in Recent Advances in Animal Nutrition (ed W Haresign and D.J.A Cole ), Butterworth, London, pp 47-60 Sanders, M.E (1993a) Summary of conclusions from a consensus panel of experts on healthy attributes of lactic cultures: significance of fluid milk products containing cultures J Dairy Sci ., 7 6, 1819 -28 Sanders,... 1974; Finegold, Sutter and Mathisen, 1983; Gibson and Macfarlane, 1995) Strictly anaerobic bacteria far outnumber other types of micro-organism, with Gram-negative rods belonging to the Bacteriodes fragilis group predominating Numerically, other Gram-negative organisms such as fusobacteria and enterobacteria constitute a relatively minor proportion of the total faecal flora Gram-positive rods including... large intestine, including age, drug therapy, diet, host physiology, peristalsis, local immunity and microbe-microbe interactions (Freter, 19 92; Salminen, Isolauri and Onnela, 1995) An important role of the normal flora is related to improved colonization resistance and reduction in the metabolic activities of harmful organisms A number of mechanisms exist whereby components of the normal gut microbiota... Dodd and Gasson (1994) and De Vuyst and Vandamme (1994) discuss the properties, producer strains, genetic information and target organisms of bacteriocins associated with these bacteria In addition, bifidobacteria, although strictly speaking not part of the lactic acid group, also secrete antimicrobial agents (Meghrous et al ., 1990; Gibson and Wang, 1994; O'Riordan, Condon and Fitzgerald, 1995) From... bifidobacteria on pathogenic micro-organisms M-A extracts from eight different species of bifidobacteria (Bif infantis NCFB 22 0 5, Bif longum NCFB 22 5 9, Bif pseudolongum NCFB 22 4 4, Bif catenulatum NCFB 22 4 6, Bif bifidum NCFB 22 0 3, Bif breve NCFB 22 5 7, Bif adolescentis NCFB 22 3 0, Bif angulatum NCFB 22 38) directly inhibited growth of a range of pathogenic bacteria, including species belonging to the genera Salmonella,... al ., 1977; Faure et al ., 1984 ), with the presence of lactoferrin also being a contributory factor (Drasar and Roberts, 1990) As a result, attempts have been made to modify infant feed formulae to more closely resemble that of humans, including adjustments to the protein content (Modler, McKellar and Yaguchi, 1990 ), and the use of probiotics An early report by Robinson and Thompson (19 52) showed that... anti-lipopolysaccharide antibody production (Yasui and Ohwaki, 1991) Solis and Lemmonnier (1993) reported that several agents, including bifidobacteria, L acidophiIus, L casei, L heiveticus and yoghurt, when incubated with blood mononuclear cells from healthy adults, produced varying increases in interleukin I-B, tumour necrosis factor alpha and gamma interferon More recently, Kitazawa et al (1994) carried out