Food Microbiology and Food Safety
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1 Food Microbiology and Food Safety Series Editor Michael P. Doyle, Georgia, USA Editorial Board Francis F. Busta, Director, National Center for Food Protection and Defense, University of Minnesota, Minneapolis, MN, USA Bruce R. Cords, Vice President, Environment, Food Safety & Public Health, Ecolab Inc., St. Paul, MN, USA Catherine W. Donnelly, Professor of Nutrition and Food Science, University of Vermont, Burlington, VT, USA Paul A. Hall, President, AIV Microbiology and Food Safety Consultants, LLC, Hawthorn Woods, IL, USA Ailsa D. Hocking, Chief Research Scientist, CSIRO Food Science Australia, North Ryde, Australia Thomas J. Montville, Professor of Food Microbiology, Rutgers University, New Brunswick, NJ, USA R. Bruce Tompkin, Formerly Vice President-Product Safety, ConAgra Refrigerated Prepared Foods, Downers Grove, IL, USA
2 Food Microbiology and Food Safety publishes valuable, practical, and timely resources for professionals and researchers working on microbiological topics associated with foods as well as food safety issues and problems. The series is co-published with the IAFP. The International Association for Food Protection is a non-profit association of food safety professionals. Dedicated to the life-long educational needs of its Members, IAFP provides Members with an information network through its two scientific journals (Food Protection Trends and Journal of Food Protection), its educational Annual Meeting, international meetings and symposia along with international interaction between food safety professionals. More information about this series at
3 Tanja Ćirković Veličković Marija Gavrović-Jankulović Food Allergens Biochemistry and Molecular Nutrition 1 3
4 Tanja Ćirković Veličković Department of Biochemistry Faculty of Chemistry University of Belgrade Belgrade Serbia Marija Gavrović-Jankulović Department of Biochemistry Faculty of Chemistry University of Belgrade Belgrade Serbia ISBN ISBN (ebook) DOI / Springer New York Heidelberg Dordrecht London Library of Congress Control Number: Springer Science+Business Media New York 2014 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright Clearance Center. Violations are liable to prosecution under the respective Copyright Law. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. While the advice and information in this book are believed to be true and accurate at the date of publication, neither the authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that may be made. The publisher makes no warranty, express or implied, with respect to the material contained herein. Printed on acid-free paper Springer is part of Springer Science+Business Media (
5 Dedicated to Sava, Aleksandar, and Nikola v
6 Preface Food allergy is an abnormal immunological reaction to food proteins, which causes an adverse clinical reaction. It can occur as a consequence of conformational crossreactivity to respiratory allergens (oral allergy syndrome), or following sensitization via gastrointestinal tract. Almost all known food allergens that sensitize via gastrointestinal system belong to the prolamin and cupin protein superfamilies of allergens. Those are mainly characterized by resistance to heat and digestion. The discrepancy between the vast numbers of proteins we encounter and the limited number of proteins that actually become allergens, have led scientists to investigate what unique features make proteins destined to be allergens. The information gained from these studies has led to an allergy assessment strategy that characterizes the potential allergenicity of biotechnology products prior to their commercialization. Beside allergen structure, new data are emerging on the influence of various other factors on the allergen survival and uptake by the gut immune system and presentation by the antigen-presenting cells, some of them related to the biophysical and immunomodulatory properties of the food matrix, and other to the way we process food. Nevertheless, the built of knowledge on all those various and complex interactions between food components and gut immune system will help us to better understand food allergies and to manufacture safer food. The most efficient treatment of food allergy is allergen avoidance. Thus, labelling of food for the allergen content is required by European food production regulative. Clear guidelines have been established in the EU regarding the allergenic food labelling according to EC Directive 2000/13/EC and amendments: Directive 2000/1/EC Annex IIIa and the Directive 2007/68/EC. The former included a list of 12 potentially allergenic food (cereals containing gluten, crustaceous, eggs, fish, peanuts, soybeans, milk, nuts, celery, mustard, sesame, and sulphur dioxide) that must always be declared on the label of the foodstuffs present in pre-packaged food traded inside EU. The latter included two more food (lupine and molluscs). This monograph covers the topics of biochemistry, digestibility, and uptake in the gastrointestinal tract of the most important food allergens causing IgE-mediated food allergy that is believed to be responsible for most immediate-type, foodinduced hypersensitive reactions. Currently available tests and strategies for food allergens identification and quantification in food matrices are reviewed in the vii
7 viii Preface monograph, as well as links between food processing, food matrix, and immunomodulatory components of food that can influence food allergy development and onset of allergic reactions. The authors of the book are professors of Biochemistry at the University of Belgrade, Faculty of Chemistry and also lead researchers of the Center of Excellence for Molecular Food Sciences at the Faculty of Chemistry. The monograph is also intended for students of the courses of Food Biochemistry and Nutrition, as well as Molecular Allergology of the postgraduate studies at the University of Belgrade, Faculty of Chemistry. We also hope that the monograph will be valuable to all those who are involved in elucidation of allergenic structures which contribute to elicitation of food allergy and food scientists involved in design of safer and more functional food products.
8 Acknowledgment The authors acknowledge help of the colleagues who read and critically commented on the final version of the manuscript, especially Dr. Dragana Stanić-Vučinić, Dr. Katarina Smiljanić, Dr. Milica Grozdanović, and Dr. Milica Popović. We would also like to thank our families for their sustained support and love. In Belgrade, In Belgrade, Tanja Ćirković Veličković Marija Gavrović-Jankulović ix
9 Contents 1 Food Allergy and Gastrointestinal Tract Food Allergy Basic Facts IgE-Mediated Food Allergy Epidemiology Diagnosis and Management of Food Allergy Development of Sensitization to Food Allergens Oral Allergy Syndrome Mucosal Immune System Oral Tolerance Antigen-Presenting Cells of the Mucosal Immune System Microbiota and Food Allergy Diet-Related Factors that Modulate Immune Response to Dietary Antigens Adjuvants Used in Animal Models of Food Allergy References Intestinal Permeability and Transport of Food Antigens Intestinal Permeability of Antigens Paracellular Transport Pathways Transcellular Transport Pathways Antigen Transcytosis Through Immune Complexes Transport of Antigens via CX3CR1hi Cells Transport of Antigens via Goblet Cells Role of Mucin Protein Degradation in the Gastrointestinal Tract Antigen Presentation in the Gastrointestinal System: Role of APCs of the Gut Intestinal Permeability in Food-Allergic Individuals Intestinal Permeability of Modified Food Allergens References xi
10 xii Contents 3 Biochemistry and Molecular Biology of Food Allergens Characterization of Food Allergens Plant Food Allergen Families The Prolamin Family The Cupin Family The Profilin and Bet v 1 Family Animal Food Allergens Parvalbumin and Fish Allergens Tropomyosins and Other Shellfish Allergens Milk Allergens Egg Allergens References Methods for Allergen Identification and Quantification in Food Matrices Principles of Protein Detection and Quantification Assays Antibody-Based Assays: Immunoassay and ELISA MS-Based Approaches Matrix Effect and Food Processing in Detection of Food Allergens DNA-Based Methods for Allergen Detection References Food Allergens Digestibility Classification of Allergens Based on Digestion Stability: Complete and Incomplete Food Allergens Protein Digestion in the Gastrointestinal Tract Classification of Food Allergens Based on Stability to Digestion Protein Conformational Stability, Digestibility, and Immune Response Digestion Stability of Common Plant Food Allergens Stability of Animal Allergens to Digestion Correlation Between Results from In Vitro Digestion Tests and Allergenicity Impact of Food Processing on Digestibility and Allergenicity of Food Allergens Food Processing and Food Allergenicity Methods for Assessing Allergenicity of Modified Food Allergens Modifications of Food Allergens Enzymatic Modifications of Allergens Thermal Treatments: Boiling, Heating, and Roasting Hydrolysis and Pasteurization of Food Allergens Modern Nonthermal Food Processing Methods: High Hydrostatic Pressure, Gamma Irradiation, and Sonication
11 Contents xiii 5.4 Food Matrix and Protein Digestion Noncovalent Interactions of Food Matrix Components and Food Allergens References Microbiota and Allergic Disease The Intestinal Microbiota Development of Microbiota upon Birth The Intestinal Mucosa Microbiota, the Intestinal Immune System, and Allergy Probiotics and Prebiotics Probiotics in Prevention of Allergic Disease References Phytochemicals and Hypersensitivity Disorders Diet and Hypersensitivity Disorders Dietary Phytochemicals Flavonoids Mechanisms of Polyphenol Interference with Hypersensitivity Disorders Effects of Polyphenols on Antigen Presentation and Maturation of Antigen-Presenting Cells Effects of Polyphenols on Release of Mediators of Allergic Inflammation Effects of Polyphenols on Immune System Regulatory Mechanisms, IgE and Cytokine Production ROS Signaling and Polyphenols Interactions of Polyphenols with Dietary Proteins Polyphenols and Intestinal Barrier Function References Predicting Potential Allergenicity of New Proteins Introduced by Biotechnology Genetically Modified Food Homology and Structural Similarity: Bioinformatics of Food Allergens Allergen Databases Testing Food Allergens Digestion Animal Models of Food Allergy References Index
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