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 FOOD SCIENCE AND TECHNOLOGY SERIES

Handbook of Food Science

and Technology 3

Food Biochemistry and Technology

Edited by
Romain Jeantet, Thomas Croguennec
Pierre Schuck and Gérard Brulé
Handbook of Food Science and Technology 3
 Series Editor
Jack Legrand & Gilles Trystram

Handbook of Food Science

and Technology 3

Food Biochemistry and Technology

Edited by

Romain Jeantet
Thomas Croguennec
Pierre Schuck
Gérard Brulé
First published 2016 in Great Britain and the United States by ISTE Ltd and John Wiley & Sons, Inc.
Translated by Geraldine Brodkorb from “Science des aliments” © Tec & Doc Lavoisier 2006.

Apart from any fair dealing for the purposes of research or private study, or criticism or review, as
permitted under the Copyright, Designs and Patents Act 1988, this publication may only be reproduced,
stored or transmitted, in any form or by any means, with the prior permission in writing of the publishers,
or in the case of reprographic reproduction in accordance with the terms and licenses issued by the
CLA. Enquiries concerning reproduction outside these terms should be sent to the publishers at the
undermentioned address:

ISTE Ltd John Wiley & Sons, Inc.

27-37 St George’s Road 111 River Street
London SW19 4EU Hoboken, NJ 07030
UK USA

www.iste.co.uk www.wiley.com

© ISTE Ltd 2016

The rights of Romain Jeantet, Thomas Croguennec, Pierre Schuck and Gérard Brulé to be identified as
the authors of this work have been asserted by them in accordance with the Copyright, Designs and
Patents Act 1988.

Library of Congress Control Number: 2016936913

British Library Cataloguing-in-Publication Data

A CIP record for this book is available from the British Library
ISBN 978-1-84821-934-2
 Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
Gérard BRULÉ

Part 1. Food from Animal Sources . . . . . . . . . . . . . . . . . . . . . . . 1

Chapter 1. From Milk to Dairy Products . . . . . . . . . . . . . . . . . . . . 3

Thomas CROGUENNEC, Romain JEANTET and Pierre SCHUCK
1.1. The biochemistry and physical chemistry of milk . . . . . . . . . . . . . 3
1.1.1. Milk fat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.1.2. Carbohydrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.1.3. Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1.1.4. Milk minerals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1.2. Biological and physicochemical aspects of milk processing . . . . . . . 17
1.2.1. The stability of fat globules . . . . . . . . . . . . . . . . . . . . . . . . 17
1.2.2. Protein stability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1.3. Dairy product technology . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
1.3.1. Liquid milk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
1.3.2. Fermented milk products . . . . . . . . . . . . . . . . . . . . . . . . . 29
1.3.3. Milk powder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
1.3.4. Cheese . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
1.3.5. Cream and butter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Chapter 2. From Muscle to Meat and Meat Products . . . . . . . . . . . 65

Catherine GUÉRIN
2.1. The biochemistry of muscle (land animals and fish) . . . . . . . . . . . . 65
2.1.1. The structure and composition of meat and fish muscle . . . . . . . 66
2.1.2. Muscle structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
2.1.3. Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
vi Handbook of Food Science and Technology 3

2.1.4. Carbohydrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
2.1.5. Vitamins and minerals . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
2.2. Biological and physicochemical changes in muscle . . . . . . . . . . . . 89
2.2.1. Muscle contraction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
2.2.2. Changes in muscle after death . . . . . . . . . . . . . . . . . . . . . . 91
2.3. Meat and fish processing technology . . . . . . . . . . . . . . . . . . . . . 102
2.3.1. Meat processing technology . . . . . . . . . . . . . . . . . . . . . . . 102
2.3.2. Fish processing technology . . . . . . . . . . . . . . . . . . . . . . . . 109

Chapter 3. From Eggs to Egg Products . . . . . . . . . . . . . . . . . . . . 115

Marc ANTON, Valérie LECHEVALIER and Françoise NAU
3.1. Chicken egg – raw material in the egg industry . . . . . . . . . . . . . . 117
3.1.1. Structure and composition . . . . . . . . . . . . . . . . . . . . . . . . . 117
3.1.2. Biochemical and physicochemical properties
of the protein and lipid fractions of egg. . . . . . . . . . . . . . . . . . . . . 120
3.2. Physicochemical properties of the different egg fractions . . . . . . . . 125
3.2.1. Interfacial properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
3.2.2. Gelling properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
3.3. The egg industry: technology and products . . . . . . . . . . . . . . . . . 136
3.3.1. Decontamination of shells . . . . . . . . . . . . . . . . . . . . . . . . . 138
3.3.2. Breaking and separation of the egg white and yolk . . . . . . . . . . 138
3.3.3. Primary processing of egg products – decontamination
and stabilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
3.3.4. Secondary processing of egg products . . . . . . . . . . . . . . . . . 142
3.3.5. Egg extracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Part 2. Food from Plant Sources . . . . . . . . . . . . . . . . . . . . . . . . . 145

Chapter 4. From Wheat to Bread and Pasta . . . . . . . . . . . . . . . . . 147

Hubert CHIRON and Philippe ROUSSEL
4.1. Biochemistry and physical chemistry of wheat . . . . . . . . . . . . . . . 150
4.1.1. Overall composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
4.1.2. Structure and properties of the constituents . . . . . . . . . . . . . . 154
4.2. Biological and physicochemical factors
of wheat processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
4.2.1. Development of texture . . . . . . . . . . . . . . . . . . . . . . . . . . 164
4.2.2. Development of color and flavor . . . . . . . . . . . . . . . . . . . . . 170
4.3. The technology of milling, bread making and pasta making . . . . . . . 172
4.3.1. Processing of wheat into flour and semolina . . . . . . . . . . . . . . 172
4.3.2. Bread making . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
4.3.3. Pasta making. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
 Contents vii

Chapter 5. From Barley to Beer . . . . . . . . . . . . . . . . . . . . . . . . . 205

Romain JEANTET and Ludivine PERROCHEAU
5.1. Biochemistry and structure of barley and malt . . . . . . . . . . . . . . . 205
5.1.1. Morphology of barley grain . . . . . . . . . . . . . . . . . . . . . . . . 206
5.1.2. Biochemical composition of barley . . . . . . . . . . . . . . . . . . . 207
5.1.3. Composition and structure of starch and protein . . . . . . . . . . . 208
5.1.4. Effect of malting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
5.2. Biological and physicochemical factors of processing . . . . . . . . . . 213
5.2.1. Enzymatic degradation of starch and protein . . . . . . . . . . . . . . 214
5.2.2. Fermentability of the wort . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.3. Brewing technology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.3.1. Stages of malting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.3.2. Stages of beer production . . . . . . . . . . . . . . . . . . . . . . . . . 224

Chapter 6. From Fruit to Fruit Juice

and Fermented Products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Alain BARON, Mohammad TURK and Jean-Michel Le QUÉRÉ
6.1. Fruit development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
6.1.1. Stages of development . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
6.1.2. Fruit ripening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
6.2. Biochemistry of fruit juice . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
6.2.1. Pectins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
6.2.2. Pectinolytic enzymes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
6.2.3. Bitter and astringent compounds . . . . . . . . . . . . . . . . . . . . . 245
6.3. Fruit juice processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
6.3.1. Preparation of fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
6.3.2. Pre-treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
6.3.3. Pressing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
6.3.4. Treatment of fruit juice . . . . . . . . . . . . . . . . . . . . . . . . . . 253
6.3.5. Pasteurization, high-pressure treatment,
pulsed electric fields and concentration . . . . . . . . . . . . . . . . . . . . . 262
6.4. Cider . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
6.4.1. French cider . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
6.4.2. Fermentation process . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
6.4.3. Action of microorganisms . . . . . . . . . . . . . . . . . . . . . . . . . 267
6.4.4. Fermentation and post-fermentation . . . . . . . . . . . . . . . . . . . 271

Chapter 7. From Grape to Wine . . . . . . . . . . . . . . . . . . . . . . . . . 275

Thomas CROGUENNEC
7.1. Raw materials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
7.1.1. Grape variety . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
7.1.2. Composition of grapes . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
viii Handbook of Food Science and Technology 3

7.2. Winemaking techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280

7.2.1. State of the harvest and adjustments . . . . . . . . . . . . . . . . . . . 281
7.2.2. Physicochemical processes involved
in winemaking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
7.2.3. Biological processes involved in
winemaking: fermentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285
7.3. Stabilization and maturation of wine . . . . . . . . . . . . . . . . . . . . . 289
7.3.1. Biological stabilization . . . . . . . . . . . . . . . . . . . . . . . . . . 289
7.3.2. Physicochemical stabilization . . . . . . . . . . . . . . . . . . . . . . 290
7.3.3. Maturation of wine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291
7.4. Specific technology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292
7.4.1. Sparkling wines (traditional method) . . . . . . . . . . . . . . . . . . 292
7.4.2. Sweet wines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293

Chapter 8. From Fruit and Vegetables to

Fresh-Cut Products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
Florence CHARLES and Patrick VAROQUAUX
8.1. Respiratory activity of plants . . . . . . . . . . . . . . . . . . . . . . . . . 298
8.1.1. Measurement and modeling of
respiratory activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
8.1.2. Control of respiratory activity . . . . . . . . . . . . . . . . . . . . . . 301
8.2. Enzymatic browning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
8.2.1. Mechanism and evaluation . . . . . . . . . . . . . . . . . . . . . . . . 302
8.2.2. Prevention of enzymatic browning. . . . . . . . . . . . . . . . . . . . 303
8.3. Unit operations in the production of fresh-cut
products: main scientific and technical challenges . . . . . . . . . . . .. . . 304
8.3.1. Raw materials: selection of varieties and
cultivation methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
8.3.2. Raw material quality control: grading . . . . . . . . . . . . . . . . . . 307
8.3.3. Trimming and mixing . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
8.3.4. Cutting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
8.3.5. Washing and disinfection . . . . . . . . . . . . . . . . . . . . . . . . . 309
8.3.6. Draining and drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
8.3.7. Weighing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
8.3.8. Bagging. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
8.4. Modified atmosphere packaging . . . . . . . . . . . . . . . . . . . . . . . 314
8.4.1. Diffusion of gases through packaging . . . . . . . . . . . . . . . . . . 315
8.4.2. Change in gas content in modified
atmosphere packaging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317
8.5. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319
 Contents ix

Part 3. Food Ingredients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321

Chapter 9. Functional Properties of Ingredients . . . . . . . . . . . . . . 323

Gérard BRULÉ and Thomas CROGUENNEC
9.1. Interactions with water: hydration and
thickening properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
9.1.1. Types of interaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
9.1.2. Influence of hydrophilic components on
water availability and mobility . . . . . . . . . . . . . . . . . . . . . . . . . . 325
9.1.3. Influence of hydration on the solubilization,
structure and mobility of compounds . . . . . . . . . . . . . . . . . . . . . . 325
9.1.4. Effect of the hydration of components
on rheological properties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
9.2. Intermolecular interactions: texture properties . . . . . . . . . .. . . . . 326
9.2.1. Aggregation/gelation by destabilization of
macromolecules or particles . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
9.2.2. Aggregation/gelation by covalent cross-linking . . . . . . . . . . . . 327
9.2.3. Sol–gel transitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
9.2.4. Influence of denaturation kinetics and
molecular interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
9.3. Interfacial properties: foaming and emulsification . . . . . . . . . . . . . 330
9.3.1. Interfacial tension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
9.3.2. Surfactants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332
9.3.3. Emulsification and foaming . . . . . . . . . . . . . . . . . . . . . . . . 332

Chapter 10. Separation Techniques . . . . . . . . . . . . . . . . . . . . . . 335

Thomas CROGUENNEC and Valérie LECHEVALIER
10.1. Proteins and peptides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
10.1.1. Milk proteins and peptides . . . . . . . . . . . . . . . . . . . . . . . . 335
10.1.2. Extraction of lysozyme from egg white . . . . . . . . . . . . . . . . 346
10.1.3. Extraction of gelatin . . . . . . . . . . . . . . . . . . . . . . . . . . . 348
10.1.4. Plant proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
10.2. Carbohydrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351
10.2.1. Sucrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351
10.2.2. Lactose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364
10.2.3. Polysaccharides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
10.3. Lipids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
10.3.1. Production of vegetable oils . . . . . . . . . . . . . . . . . . . . . . . 379
10.3.2. Lipid modification . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
10.4. Pigments and flavorings. . . . . . . . . . . . . . . . . . . . . . . . . . . . 391
10.4.1. Types of pigments and flavorings . . . . . . . . . . . . . . . . . . . 391
x Handbook of Food Science and Technology 3

10.4.2. Extraction/concentration of
colorings and flavors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397
10.4.3. Formulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400

Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403

List of Authors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419
 Introduction

The processing into food of raw materials from hunting, gathering,

fishing and subsequently arable and livestock farming has always had two
objectives: to preserve nutrients in order to defer the time and place of
consumption, and develop products with a wide variety of textures and
flavors to satisfy the sensory needs of consumers. The development of arable
and livestock farming has facilitated an improved control of supply, even
though the provision of agricultural products has long remained very
irregular due to climatic or health risks and the seasonality of certain
products. Furthermore, the importance of stabilization and/or processing has
significantly increased with the rural exodus, which has led to a distancing of
production from consumption areas.

The production of certain foods that still form the basis of our diet today
dates back several centuries or even millennia, as in the case of bread, cheese
and wine for example. These products, particularly those derived from
fermentation, were developed based on empirical observations, with no
knowledge of the raw materials or phenomena involved in their processing.
It was not until the work of Pasteur in the 19th Century that microorganisms
gained a key role in the development and processing of agricultural products.

The agri-food industry has undergone a major change over the past few
decades in order to better meet the quality requirements of consumers; while
traditional food is the result of a series of increasingly understood and
controlled biological and physicochemical phenomena, this is not the case
for a number of new products designed to meet market expectations. These

Introduction written by Gérard BRULÉ.

xii Handbook of Food Science and Technology 3

products are the result of an assembly of various ingredients (Figure I.1), the
control of which is a real challenge for food technologists and engineers.

chemical
Constituents enzymatic – microbial Functional ingredients
Proteins - Lipids engineering
Texture agent
Sugars – Aromas
Sweetener
Coloring
Aroma, coloring
Biological agents
microorganisms - enzymes
Separation Assembly

Raw materials Thermal & mechanical Foods

treatments
maturation - grinding – water
heating - salting - etc.

Figure I.1. New products and assembly technology

I.1. From empiricism to rational technology

The oldest forms of processing (milk to cheese, grain to bread or beer,

grapes to wine, muscle to meat, etc.) were based on biological phenomena
that could occur naturally under specific water content and temperature
conditions, since the biological agents (enzymes or microorganisms)
responsible for processing and the reaction substrates and growth factors
were present in the raw materials and/or immediate environment; it was
enough to simply mature (milk, meat), crush, grind and sometimes hydrate
(fruit, grain) in order for biological reactions to take place. This is why these
types of products were able to develop based solely on the observation of
natural processes.

The knowledge acquired since the end of the 19th Century in the field of
microbiology and the early 20th Century in the field of enzymology has
gradually helped explain the biological phenomena involved in the
development of certain food products. Based on this knowledge, the food
industry has sought to control these processes rather than witness them,
 Introduction xiii

which is how the fermentation and then an enzyme industry arose, producing
and marketing biological agents for each type of processing. The use of
fermentation and in some cases enzymes has become indispensable given the
existing food safety requirements, which include increasingly stringent
hygiene conditions in production and processing, and technological
treatments to eliminate potential pathogenic microorganisms (microfiltration,
heat treatment). This change has led to a reduction in the endogenous
biological potential, which needs to be replenished by adding fermentation
steps and enzymes. Reconstituting microbial ecosystems through the
assembly of exogenous flora requires the identification of the endogenous
flora and their role in the characteristic features of the food; progress in the
field of molecular biology should enable significant progress in this area.

Over the past 40 years, many teams have focused on the study of food
science, which has resulted in a better understanding of the composition of
various raw materials and the biological and physicochemical mechanisms
involved in the development of texture, flavor and aroma; this work has
allowed the food industry to better identify the key technological tools in the
development of quality, and to rely less on empiricism and more on
technology.

I.2. From traditional foods to assembly technology

The quality requirements of consumers are increasingly specific and

segmented. Food must be absolutely safe (no pathogens, toxins, residues or
contaminants), have the closest possible nutritional profile to that
recommended by nutritionists, meet sensory needs, integrate practicality
(ease of storage and use) and convey social values (fair trade, environmental
protection and animal welfare) while remaining at an affordable price. These
market expectations are identified by the marketing services of the food
industry for specific target consumers whose needs depend on several factors
(gender, age, activity, health, metabolic disorders, food trends, etc.). These
services, in consultation with nutrition and health specialists, identify which
nutrients and micronutrients (minerals and vitamins) to assemble and define
the structure, sensory characteristics and practicality of the food based on
consumer research. The path from conception to completion can sometimes
be difficult, since food is a complex and thermodynamically-unstable
system, which can be defined as a continuous, usually aqueous phase, a
three-dimensional protein and/or polysaccharide network and dispersed
xiv Handbook of Food Science and Technology 3

elements (gas, fat globules, solids); the aim of technologists is to stabilize

this system throughout the marketing period while taking into account
mechanical (transport) and thermal (refrigeration, freezing, thawing)
constraints.

The progress made in recent years in food science has provided insight
into the key role of various biological components and particularly their
structure, whether native or modified by technological treatments, in the
development of texture, the thermodynamics of dispersed systems and the
role of interfaces. This knowledge allows us to better understand and control
the instability of food using technological processes or functional
ingredients; the industry has a very large range of functional ingredients that
is used to create texture and stabilize complex multiphase systems. It is
therefore possible, by assembly, to create new foods that meet the quality
requirements of the market.
12 Handbook of Food Science and Technology 3

sometimes be glycosylated. Phosphoserines, arranged in clusters, display a

strong affinity for divalent or polyvalent cations, which depending on their
type can make casein insoluble. The sensitivity of casein to calcium increases
with the rate of phosphorylation. κ-casein does not precipitate in the presence
of calcium.

Caseins have few sulfuric amino acids, which limits their nutritional value.
αs2- and κ-caseins each have two cysteines involved in intermolecular
disulphide bonds. While αs2-casein is mostly present as covalent homodimers,
κ-casein forms polymers of up to 15 κ-casein units.

1.1.3.2. Structure of the casein micelle

Casein micelles are spherical particles formed by the aggregation of
different caseins (αS1, αS2, β and κ), some peptide fragments resulting from the
proteolysis of β-casein by plasmin (γ-casein) and salt components, the main
ones being calcium and phosphate. Table 1.4 shows the average composition
of casein micelles.

Caseins Salt components

αS1 33 Calcium 2.9

αS2 11 Magnesium 0.2
β 33 Inorganic 4.3
κ 11 Citrate 0.5
γ 4
Total caseins 92 Total salt components 8.0

Table 1.4. Average composition of casein micelles in % (w/w)

The composition of casein micelles varies slightly depending on their

diameter, which varies between 50 and 600 nm for an average diameter of
about 150 nm. Regardless of the size of the micelle, the proportion of αS1 and
αS2 casein varies marginally, whereas the ratio of β- to κ-caseins increases
with the size of the micelle. Micelle organization, or the arrangement and
distribution of the various micelle components and their types of association,
is still a subject of intense debate. The non-charged regions of caseins form
rigid structures maintained by hydrophobic associations and hydrogen bonds;
colloidal calcium phosphate, in the form of nanoclusters, shields the negative
 From Milk to Dairy Products 13

charges of phosphoserine clusters and allows the association of casein

micelles. κ-casein is structured into inhomogeneous clusters almost
exclusively located on the micelle surface. Without phosphoserine clusters, κ-
casein remains associated to the casein micelle by its hydrophobic N-terminus,
but prevents further micelle growth. Its charged hydrophilic C-terminus
protrudes about 5 – 10 nm into the solvent phase, making the micelle appear
hairy.

Table 1.5 shows some of the properties of casein micelles. Their

composition and physicochemical properties are highly dependent on the
solvent phase.

Property Values

Average diameter (nm) 150

2
Area (cm ) 8 × 10–10
Volume (mL) 2.1 × 10–15
Density (hydrated) 1.0632
-1
Hydration (g H2O g of proteins) 3.7
-1
Voluminosity (mL g of proteins) 4.4
Molecular weight (hydrated) (Da) 1.3 × 109
Molecular weight (dehydrated) (Da) 5 × 108
Water content (%, w/w) 63
Number of caseins per micelle 2 × 104
Number of nanoclusters of calcium phosphate per micelle 3 × 103
Number of micelles per L of milk 1017 – 1019
Mean free distance between micelles (nm) 240
Zeta potential (mV) –13

Table 1.5. Average physicochemical properties of casein

micelles at 20°C and pH 6.7 (modified from [MCM 84])

1.1.3.3. Whey proteins

Whey proteins are defined as the protein fraction that remains soluble at pH
4.6. β-lactoglobulin, α-lactalbumin, bovine serum albumin (BSA),
immunoglobulins and lactoferrin represent more than 90% of all whey
proteins. They are mostly globular proteins with a high sensitivity to heat
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