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Prelims.qxd 12/19/09 4:25 PM Page i

FOX AND CAMERON’S

Food Science,
Nutrition & Health
This page intentionally left blank
Prelims.qxd 12/19/09 4:25 PM Page iii

FOX AND CAMERON’S

Food Science,
Nutrition & Health
7th edition

Michael E. J. Lean
MA, MB, BChir, MD, FRCP
Professor of Human Nutrition at the University of Glasgow
Consultant Physician at Glasgow Royal Infirmary
Visiting Research Fellow at the Centre for Human Nutrition,
University of Colorado Health Sciences Center, USA

Hodder Arnold
A MEMBER OF THE HODDER HEADLINE GROUP
CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742
© 2006 by Taylor & Francis Group, LLC
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No claim to original U.S. Government works

Version Date: 20121026

International Standard Book Number-13: 978-1-4441-1337-2 (eBook - PDF)

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Prelims.qxd 12/19/09 4:25 PM Page v

Contents
Preface to the 7th edition and Acknowledgements vii
Preface to the 6th edition ix
Acronyms/terms used xi

Chapter 1 Food and its functions 1

Chapter 2 Enzymes and digestion 11

Chapter 3 Food, eating, health and disease 23

Chapter 4 Nutrient dietary requirements and reference values 34

Chapter 5 Obesity in the twenty-first century 39

Chapter 6 Fats, oils and lipids 45

Chapter 7 Milk and dairy products 76

Chapter 8 Carbohydrates 88

Chapter 9 Carbohydrate-rich foods 106

Chapter 10 Amino acids and proteins 132

Chapter 11 Protein-rich foods 152

Prelims.qxd 12/19/09 4:25 PM Page vi

vi Contents

Chapter 12 Water and beverages 168

Chapter 13 Mineral elements 186

Chapter 14 Vitamins and other bioactive food constituents 198

Chapter 15 Fruits, nuts and vegetables 219

Chapter 16 Methods of cooking 235

Chapter 17 Diet and health 249

Chapter 18 Food spoilage and preservation 270

Chapter 19 Toxins, food-borne infections and food hygiene 286

Chapter 20 Food contaminants – adulterants and additives 297

Index 311
Prelims.qxd 12/19/09 4:25 PM Page vii

Preface to the 7th edition

Fox and Cameron has rightly become the leading and new emphasis. Reflecting the change in editor-
textbook in the field of Food Sciences and Nutrition, ship and the changing demands of the food industry
for use both in colleges and in schools, as well as a ref- to meet the needs of consumers, this new edition
erence source for the food and catering companies. includes a great deal more on the impact of foods
The style created by Fox and Cameron was both on health – both through nutrition and through food-
accessible and authoritative, and not without humour borne infections. Referencing is much reduced in
in places. this 7th edition. Rather than referring back to older
For this revised 7th edition, a great debt is still publications, students and their teachers should be
owed to B. A. Fox and A. G. Cameron (Brian Fox accessing the latest scientific regulatory and policy
sadly died in 2002). I have tried to retain their documents on the internet. The impact on the food
human touch and the balance between presenting industry of new, consumer-centred agencies such as
up-to-date scientific issues in the context of foods the Food Standards Agency and the Joint Health
we all know and love (or loathe, in some cases). Claims Initiative, are felt throughout this new
I have been helped enormously by the hard work edition, and students will need to monitor their
of Dr Wendy Wrieden, Centre for Public Health websites for new information.
Nutrition Research, University of Dundee, UK, and The times are certainly changing for the food
we have both relied on help from our secretaries industry. With continued globalization, and 90 per
and academic colleagues in the process of revision. cent of all food regulations in UK now being derived
Some of the book is left unchanged from the lat- from Europe, students and workers in the food indus-
est editions. This is in part a testimony to the huge try need to keep on their toes. This book is designed
amount of material marshalled by Fox and Cameron. to help.
However, all the chapters have been brought up-to- Professor Mike E J Lean
date to incorporate new evidence, new technologies 2006

Acknowledgements

‘The Water Song’ appears on p. 168.

Words and Music by Robin Williamson
© 1968 (renewed) Warner-Tamerlane Publishing Corporation
Administered by Warner/Chappell Music Ltd, London W6 8BS
Reproduced by permission
This page intentionally left blank
Prelims.qxd 12/19/09 4:25 PM Page ix

Preface to the 6th edition

It is universally accepted that an adequate and well- retains the basic structure of previous editions. The
balanced diet is essential for the enjoyment of good first three chapters describe in outline the nature
health but the scientific principles on which this and functions of food, and the changes it undergoes
belief is based are by no means widely understood. in the body when it is eaten, together with a pre-
This book, which bridges the gap between science liminary exploration of the relationship between
and practice in nutrition, is an attempt to make dietary factors and good health. A new chapter fol-
good this deficiency. It is largely concerned with what lows in which the relatively new terms known as
food is, how much of it we should eat and what dietary reference values, which are used to express
happens to it – and to us – when we eat it. It pro- dietary requirements, are introduced. These have
vides a comprehensive and up-to-date survey of the taken the place of the terms used previously, such as
body of knowledge which has come to be known as recommended daily amount, and they are employed
food science, and examines the nutritional signifi- consistently throughout the remainder of the book.
cance of this knowledge. This is a more difficult task The main body of the text is given over to an account
than might at first be supposed because of the many of the nature of nutrients and the foods which con-
scientific disciplines involved, the complex nature tain them and to a description of what happens to
of many apparently simple foodstuffs and the intri- food when it is grown, stored, processed, preserved,
cate biochemical reactions that occur when food is cooked and eaten. A chapter dealing with the rela-
digested, absorbed and converted into the stuff of tionship between diet and health discusses the way
which we are made. in which dietary habits can be modified to promote
We have sought to make the book as self-contained good health and combat disease.
as possible but a study of food science and nutri- Subsequent chapters look at the related topics of
tion, even at the fairly elementary level at which food spoilage and preservation, food poisoning and
this book is pitched, is of necessity far-ranging and food hygiene. The final chapter, which is concerned
inevitably involves an assumption of some previous with the important topic of food contaminants and
knowledge of basic science. Nevertheless, we hope additives, seeks to present a balanced and compre-
that the text will be intelligible to all those who hensive account of what has become a controversial
might wish to read it, and with this in mind, we subject. Although there may be risks involved in
have kept the use of structural formulae to a mini- using some additives, considerable benefits may also
mum. Unfortunately, it has not been possible to be obtained and an attempt is made to balance
avoid using the combinations of initial letters which these two aspects of the use of additives in food.
serve as abbreviations for frequently used groups We have not attempted to provide references to the
of words with particular meanings; these have, in original sources of the information given in this book;
effect, become recognized as part of the specialized such references would have greatly increased the
vocabulary of food science. To assist the reader in length and cost of the book and would, perhaps, have
finding a way through this veritable thicket of alpha- been used by relatively few readers. A short reading
betic abbreviations a Glossary has been provided list is given at the end of many chapters which will
which we hope will prove helpful. assist readers requiring more detailed information.
We have been encouraged by the consistent popu- Details of a number of books on food science and
larity of the book over a long period to conclude nutrition which are of more general interest are given
that it is fulfilling a need and meeting the objectives in a General Reading List at the end of the book.
we set ourselves when it was first written. Conse- Great care has been taken to ensure that the infor-
quently, we have not thought it necessary to make mation contained in this book is as accurate and
radical changes in preparing this sixth edition and it up-to-date as possible but it is almost inevitable that
Prelims.qxd 12/19/09 4:25 PM Page x

x Preface to the 6th edition

errors and ambiguities will occur in a book which the general public who are concerned about the rela-
attempts to summarize a large and expanding field tionship between dietary habits and good health,
of knowledge. We should be grateful if any such and who wisely take an interest in ‘healthy eating’.
inaccuracies, for which we alone are responsible, We hope that all who read the book will enjoy
can be brought to our notice. doing so and that the information it contains will
We trust that the book will continue to be of inter- assist them in the pursuit of good health and the
est and value to all who are concerned with food avoidance of those life-threatening diseases which are
science and nutrition. It is principally intended for now known to have a nutritional component.
students of Food Science and Technology, Catering
and Nutrition and for A-level and BTEC students Brian A Fox
in schools and colleges. We know that it has also Allan G Cameron
been found to be of interest by many members of 1994
Prelims.qxd 12/19/09 4:25 PM Page xi

Acronyms/terms used

AFD accelerated freeze-drying MRL maximum residue level

ADI acceptable daily intake MAP modified-atmosphere packaging
ACP acid calcium phosphate MAS modified-atmosphere storage
ASP acid sodium pyrophosphate MSG monosodium glutamate
ADP adenosine diphosphate MUFA monounsaturated fatty acid
AMP adenosine monophosphate NHDC neohesperidine dihydrochalcone
ATP adenosine triphosphate NPU net protein utilization
ADH antidiuretic hormone PSD Pesticides Safety Directorate
BV biological value PKU phenylketonuria
BSE bovine spongiform encephalopathy PUFA polyunsaturated fatty acid
COMA Committee on Medical Aspects of POMC pro-opio-melanocortin
Food Policy PEM protein-energy malnutrition
CAS controlled-atmosphere storage PLP pyridoxal-5-phosphate
CHD coronary heart disease RDA recommended daily amount
DEFRA Department for Environment, Food RDI recommended daily intakes
and Rural Affairs RNI reference nutrient intake
DRV dietary reference value SI safe intake
DHA docosahexanoic acid SRSV small round structured virus
ECG electrocardiogram SIADH syndrome of inappropriate antidiuretic
EFAs essential fatty acids hormone
EAR estimated average requirement TVP textured vegetable protein
GMOs genetically modified organisms TDT thermal death time
GDL Glucono-delta-lactone TDF total dietary fiber
HDL high-density lipoprotein UHT ultra-high temperature
HTST high-temperature short-time VLCDs very low calorie diets
JHCI Joint Health Claims Initiative VLDL very low density lipoproteins
LDL low-density lipoprotein cholesterol WHO World Health Organization
LRNI lower reference nutrient intake
This page intentionally left blank
Chap-01.qxd 13-02-2006 12:55 Page 1

1
Food and its functions

Food and foods 1 The use of energy by the body 7

Types of nutrient 3 Energy requirements 8
Food as a source of energy 3 Further reading 10

The basic function of food is to keep us alive and Determining the dietary needs of individuals, or
healthy, to grow and to reproduce. In this book we of populations is the realm of nutritional science.
shall consider how food does this, although we shall The problems involved in determining what foods
also need to think about many other related mat- in what combinations best meet the dietary needs
ters. Indeed, we cannot answer such a fundamental of different countries, or individuals, of the compos-
‘How’ question without first finding out the answer itional merits of various new foods, of how to store
to some simpler ‘What’ questions, such as what is and preserve food with minimum nutritional loss are
food, what happens to it when it is stored, processed, the provinces of food science. However, in order to
preserved, cooked, eaten and digested. The answers use this information it must be applied – food must be
to such questions can only be found out by experi- grown, stored, processed, preserved and transported
ment, and many different sciences play a part in on a large scale – and this is the province of agricul-
helping to provide the answers. In recent years the ture and food technology.
study of food has been accepted as a distinct disci- It becomes clear that promoting health through
pline of its own and given the name food science. diet requires input from human nutritional science,
Food science exists partly to pursue academic food science and food technology.
knowledge (the insatiable curiosity of mankind finds In the following pages we shall study the relation
in food a universal theme of shared interest). As between food science, nutrition and health, but first
Samuel Johnson phrased it, ‘I look upon it that it is important to understand just what we mean by
he who does not mind his belly will hardly mind the term food.
anything else’ (Boswell, The Life of Samuel Johnson,
Vol. 3, Ch. 9). Its interest lies also in the fact that
our knowledge of the subject is growing, leading to FOOD AND FOODS
an unfolding of new perspectives about what is sig-
nificant, while new techniques are being developed The definition of what is, and what is not, a food is
leading to new methods in food processing and of surprisingly taxing, not least because of the need to
analysing nutrients, additives and possible contam- provide control and regulation over the things, which,
inants in food. Food science also promotes the ful- for different purposes, people swallow.
filment of a basic human need for a diet that will Not everything people eat is food. Children, preg-
sustain life and health. To be effective, food science nant women and sometimes others sometimes eat
must be applied, in the manufacture and preparation non-food materials such as coal. This behaviour is
of food and this is the province of food technology. known as ‘pica’. Few people would class coal as food,
Chap-01.qxd 13-02-2006 12:55 Page 2

2 Food and its functions

on the grounds that it is not absorbed and does not from industrial pollution. Heavy metals, radioactive
contribute in any way to biochemical or physio- isotopes, and microbial contamination all have
logical function in the body. But where does that potential negative health effects.
leave chewing gum? Chewing gum is produced by A final and even more tricky consideration is that
‘food companies’ with the intention that it should be foods contain a variety of compounds which can be
taken by mouth, recognizing that it will occasionally absorbed and which have important biological effects.
be swallowed. Although inert, and non-absorbable, A good example is caffeine. It is present in several
chewing gum deserves (and receives) the same regu- common foods (tea, coffee, chocolate). It is also sold,
lation as other less ambiguous foods. and prescribed, as a drug. In doses similar to the
According to the Food and Agriculture Organiza- amount in a cup of coffee it stimulates brain, heart
tion, ‘Food means any substance, whether processed, and lung function, with several beneficial (and some
semi-processed or raw, which is intended for human potentially harmful) consequences. So, it is perfectly
consumption, and includes drink, chewing gum possible for foods to have potent ‘pharmacological’
and any substance which has been used in the actions. Caffeine, depending on how it is packaged,
manufacture, preparation or treatment of ‘food’ but may be subject to either food law or drug law. Fish
does not include cosmetics or tobacco or substances oils are also currently marketed as drugs while obvi-
used only as drugs’ (ftp://ftp.fao.org/codex/manual/ ously being available as part of fish. The distinction
Manual12ce.pdf, p. 41). between a ‘food product’ and a ‘drug/medicinal prod-
It is important to recognize that a food is almost uct’ is a fine one with scope for interpretation. The
always a complex substance, rather than a pure com- European Union (EU) has proposed a definition for
pound of single, uniform composition. Food is char- a medicinal product as ‘Any substance or combin-
acterized by more than just its chemical composition. ation of substances presented for treating or pre-
There are exceptions of course. Table sugar is indis- venting disease in human beings. Any substance or
putably ‘a food’, although it is a pure industrial prep- combination of substances which may be adminis-
aration of (CH2O)6. However, for most foods there tered to human beings with a view to making a
is a characteristic physical form, containing a variety medical diagnosis or to restoring, correcting or modi-
of nutrients with a range of properties. Some grow fying physiological functions in human beings is
that way, others are created by food technology and likewise considered a medicinal product’ (European
manufacturers. Some are consumed as grown or pro- Parliament, 2001).
duced, others form ingredients of still more com- For some new products, it can be a close decision
plex foods, dishes or meals. In the twenty-first whether to market it through food outlets as a ‘food
century, we probably have to extend our concept of product’ with permitted health claims to be made, or
what constitutes a food, at least as perceived by whether to have it registered and sold on prescrip-
consumers, to include its packaging and the imagery tion or over the counter in pharmacies as a drug,
and descriptive or promotional text included, just as which permits medical claims. The technology
we would include the skin of a banana, or the outer required is identical, although greater controls over
leaves of an onion, as part of ‘the food’. We make content variability may be required for a drug. The
judgements about food quality from all these compon- main difference lies in the differences between what
ents, and packaging, natural or man-made, can influ- constitutes a medical claim (drug) and a health
ence food composition even if it is not consumed. claim (food product). Again, there is a grey area for
Foods contain nutrients – components that con- interpretation, and the ultimate practical distinc-
tribute to, and in some cases uniquely provide for, tion between a food or a drug lies in the enormous
biochemical and physiological functions in the amount of highly expensive research required by
body. Foods may also include non-absorbed compon- drug licensing authorities. For a drug, advertising is
ents which may influence bowel health and func- restricted, and the market much more limited, but
tion. Some phenolic compounds, such as tannins vastly greater prices can be charged and drug
and classes of non-starch polysaccharides (e.g. cellu- patents ensure no direct competition for several
lose) probably fall into this category. Food may also years. There is currently great interest in protecting
include contaminants from unusual soil types, or consumers against food companies which make
Chap-01.qxd 13-02-2006 12:55 Page 3

Food as a source of energy 3

health claims, or even medical claims for their prod- Mineral

Carbohydrates Fats Proteins Water Vitamins
ucts, often based on no direct evidence at all. The elements
Bread Butter Meat Vegetables Drinking Fruit
counterargument runs that since antiquity certain Potatoes Margarine Fish Fruit water Vegetables
foods have been attributed with magical qualities Sugar Cheese Cheese Beverages
Biscuits Olive oil Eggs Fruit
in folklore, and to strip these beliefs from food Jam Lard Milk Vegetables
marketing is to deny a rich and evolving food cul-
ture. The leading agency in the UK and Europe is
the Joint Health Claims Initiative, a partnership Energy Growth Control of body
and repair processes
between the food industry, consumer organization
and enforcement authorities, which was established Figure 1.1 The nutrients: diagram showing their
to develop a Code of Practice for health claims on functions and representative foods in which they
are found
foods. The legal and regulatory issues on foods are
too complex to be addressed in detail in this book,
but a clear and helpful introduction can be found functions in the body and important foods that
at http://www.jhci.co.uk. supply them is shown in Fig. 1.1.
Nutrients are often grouped according to their
chemical composition. For example, although differ-
TYPES OF NUTRIENT ent oils and fats, such as olive oil and palm oil, do not
have identical compositions, they are chemically simi-
The two basic functions of nutrients are to provide lar and mostly use the same biochemical pathways
materials for growth and repair of tissues – that is, for digestion, absorption, transport and metabolism.
to provide and maintain the basic structure of our In the same way, different proteins (and carbohy-
bodies – and to supply the body with the energy drates) are constructed and metabolized according
required to perform external activities as well as to the same general chemical patterns. The three
carrying on its own internal activities. The fact that ‘macronutrients’ (fat, carbohydrate and protein) are
the body is able to sustain life is dependent upon its therefore conveniently grouped together. Vitamins
ability to maintain its own internal processes. This are not classified according to chemical type; at the
means that although we may eat all sorts of differ- time of their discovery in food their chemical nature,
ent foods and our bodies may engage in all sorts of which in most cases is complex, was unknown. They
external activities, and even suffer injury or illness, were grouped together because it was known that
the internal processes of the body should absorb small quantities of them were essential to health
and neutralize the effects of these events and carry and they do not fall into the macronutrient classes.
on with a constant rhythm. This is only possible At first they were identified in terms of their effect
because the components of our bodies are engaged on growth and health and distinguished by letters as
in a ceaseless process of breakdown and renewal vitamins A, B, C and so on. Their chemical compos-
(a theme to which we will return). ition is now known, and it has become apparent that
If the body’s internal processes are to be main- they are not chemically related to each other. They
tained constant despite its ceaseless and varying activ- all have different functions but it is still convenient
ity, and in the face of external pressures, there needs to consider them together.
to be systems in place to regulate very precisely all
the processes needed for life. Thus, nutrients have
a third function, namely that of controlling body FOOD AS A SOURCE OF ENERGY
processes, a function that will be considered in the
next chapter. Energy is required for sustaining all forms of life
Although habits and patterns of eating may vary on Earth. The prime source of the earth’s energy is
from person to person, and diets may be selected the sun, without which there could be no life on
from hundreds of different foods, everyone needs this planet. The sun continually radiates energy, a
the same types of nutrients and in roughly the same tiny fraction of which is intercepted by the Earth
proportions. The relation between nutrients, their and stored in various ways; plants and coal, for
Chap-01.qxd 13-02-2006 12:55 Page 4

4 Food and its functions

example, act as energy storehouses. Living plants into short-term storage as ATP, a high-energy phos-
convert the sun’s energy into chemical energy and phate compound, which can release this energy for
some plants of past ages have been converted, over individual steps in biochemical metabolism. Sucrose
many millions of years, into coal. may be converted into carbon dioxide and water by
Plants, by the process of photosynthesis, convert burning it in air. The chemical reaction is the same
carbon dioxide and water into carbohydrate. Photo- as that represented by the equation above, with
synthesis, which is discussed in detail in Chapter 8 exactly the same quantity of heat being liberated as
(p. 88), can only take place in daylight because solar when the oxidation occurs in the body. The differ-
energy is needed for the process. A complex series of ence in the two reactions concerns the speed at which
chemical changes occurs which can be represented they occur and their efficiency. Oxidation in the body
by the following equation: takes place much more slowly than combustion in
air as it takes place in a series of steps, thus ensuring
daylight a slow, controlled and gradual release of energy to
xCO2  yH2O 
→ C12 (H2O)y  xO2
body tissues. The efficiency of combustion within
Carbohydrate
the body is less than that in air because only about
two-thirds of the energy of the sucrose becomes avail-
The formation of carbohydrate is, therefore, the
able as biological energy, with one-third being ‘lost’ as
method used by plants to trap and store a part of
heat which helps to maintain body temperature.
the sun’s energy. Sugar-beet, which synthesizes carbo-
Both fat and proteins also contribute in similar
hydrate in the form of the sugar sucrose, may be
oxidations (see Chapter 8).
taken as an example:
(ˆCH2ˆ)2n  (O2)3n → 3nCO2  nH2O
daylight
12CO2  11H2O 
→ C12 (H2O)11  12O2 Fat
Sucrose The oxidation of proteins depends on which
amino acids are included. The nitrogen is removed
When sucrose is formed from carbon dioxide and and thus most are oxidized like carbohydrate while
water, energy is absorbed and stored as chemical some are oxidized like fat.
bonds within the sucrose molecule. It can be stored The body is sometimes likened to a slow-
indefinitely in this form. combustion stove and macronutrients are described
Animals, unlike plants, cannot store the sun’s as fuel. It is clear that oxidation in the body is a most
energy directly and so must gain it second-hand by important process for it enables the energy stored in
using plants as food. Carnivorous animals and man carbohydrates, fats and proteins to be liberated and
take this process a stage further and also use other ani- made available for use by the body in a closely
mals as food. In this way, chemical compounds which regulated way.
have been photosynthesized and stored in plants are
eaten by man and animals and the stored energy made
The energy value of food
available. For example, the energy that is stored within
the sucrose molecule when it is synthesized by sugar- Energy is usually measured in units called calories. A
beet is liberated when sucrose reverts to carbon diox- calorie is the amount of heat required to raise the
ide and water. This breakdown of sucrose into simpler temperature of 1 g of water by 1°C. As this is rather
units is brought about in the body by digestion and a small unit, energy derived from food may be
oxidation, but the overall reaction is simply the expressed in units which are 1000-times larger and
reverse of that represented above, namely: known as kilocalories (kcal). A kilocalorie is the
amount of heat required to raise the temperature of
C12(H2O)11  12O2 → 12CO2  11H2O
1 kg of water by 1°C. The common abbreviations for
Sucrose
kilocalorie is kcal but Calorie, or Cal (with a capital
When sucrose is converted into carbon dioxide and C) is also used to indicate kilocalories in foods.
water in this way, the energy stored during synthesis Another internationally recognized unit of
is made available for use by the body, by converting it energy is the joule (J), but like the calorie this is an
Chap-01.qxd 13-02-2006 12:55 Page 5

Food as a source of energy 5

inconveniently small unit with which to express the incompletely oxidized in the body. In addition a cor-
energy value of food so that the kilojoule (kJ), which rection may need to be made for non-starch polysac-
is 1000-times larger than the joule, is usually used. charides (NSP, see p. 96), previously called dietary
Sometimes an even larger unit, the megajoule (MJ), fibre. The magnitude of these energy losses may be
is used. A megajoule is 1000-times larger than a appreciated from Table 1.1. There is some variation
kilojoule. and uncertainty about nutrient availability; the ones
The relationship between these units may be quoted may be taken as being sufficiently reliable for
expressed as follows: most purposes, but in special conditions these may be
different. With malabsorptive diarrhoea availability is
1 kcal  4.19  103 J  4.19 kJ  4.19  103 MJ
obviously low, and varies between nutrients. There
An average adult usually needs about 1800– may, for example, be selective fat malabsorption
2500 kcal/day, which is about 8–10 MJ/day. (steatorrhoea) with liver or pancreatic disease. Mild
In the remainder of this chapter both types of unit malabsorption may cause no symptoms.
are given to illustrate their relation to each other. Although the available energy values given in
Elsewhere, however, kilojoules (and megajoules) Table 1.1 are approximate values which can be used
are used. to calculate the energy value of any given diet, the
In order to compare the energy of different foods it available energy value of any food can be found
is simplest to determine the amount of energy pro- using the average figures, provided that its composi-
duced, calculated as heat, when 1 g of the substance tion in terms of carbohydrates, fats and proteins is
is completely oxidized by igniting it in a small cham- known. The energy value of summer milk, for
ber filled with oxygen under pressure. The result example, can be calculated from its analysis, as
obtained represents the heat of combustion of food shown in Table 1.2. By similar calculations the avail-
which is usually expressed as kcal or kJ/g. If the able energy value of other foods may be estimated;
calorie value of sucrose is expressed in this way, it some average values are given in Table 1.3. This
is found to be 3.95 kcal/g. This means that when 1 g table shows that foods, such as butter and mar-
sucrose is completely oxidized the heat produced garine, which contain a high proportion of fat have
is sufficient to raise the temperature of 1000 g of the highest energy values. Foods rich in carbohydrates,
water by 3.95°C. The average values of the heats of containing a high proportion of sugar (jam) or
combustion of the energy-providing nutrients are starch (bread and potatoes), are less concentrated
shown in Table 1.1. sources of energy. Despite this, such foods supply a
In order to express the energy value of nutrients considerable proportion of the energy in an average
in terms of the energy actually made available to the British diet. Indeed, cereal foods contribute no less
body it is necessary to calculate the available energy than one-third of our total energy intake, which is a
values. Such values are always lower than heats of greater proportion than that supplied by any other
combustion because of losses within the body. A class of foodstuff. In many other countries, starchy
small loss results from incomplete absorption from foods, often in the form of rice, supply an even greater
the intestine; such loss is suffered by carbohydrate, fat proportion of the total energy content of the diet.
and protein, and with protein there is an additional As a general rule, the proportion of fat has increased
loss because protein, unlike carbohydrate and fat, is in ‘advanced’ Western market economies.

Table 1.1 Average energy value of nutrients (per gram)

Heat of combustion Available energy value

Nutrient (kcal) (kJ) (kcal) (kJ)

Carbohydrate 4.1 17 3.75 16

Fat 9.4 39 9.00 37
Protein 5.7 24 4.00 17
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6 Food and its functions

Table 1.2 The energy value of summer milk

Amount in 100 g milk Energy/100 g milk

Nutrient (g) (kcal/g) (kJ/g) (kcal) (kJ)

Carbohydrate 4.1 3.75 16 15.4 65.6

Fat 4.0 9.00 37 36.0 148.0
Protein 3.4 4.00 17 13.6 57.8
Total energy provided 65.0 271.4
by 100 g of summer milk
Reference: Food Standards Agency (2002). McCance and Widdowson’s The Composition
of Foods, 6th summary edition. Cambridge: Royal Society of Chemistry.
Crown copyright material is reproduced with the permission of the Controller of HMSO and
the Queen’s Printer for Scotland.

Table 1.3 Typical energy values of some foods with nutrients as percentage of energy

Per 100 g As percentage of energy

Food (kcal) (kJ) Fat Carbohydrate Protein

Apples 47 199 2.0 94.0 3.5

Tomatoes 17 73 16.0 68.0 16.5
Lettuce 14 59 32.0 45.0 23.0
Cabbage (boiled) 16 67 22.5 51.5 25.0
Dates 270 1151 0.7 94.5 4.8
Bread (white) 219 931 6.5 79.0 14.5
Potatoes (old) 74 306 1.3 88.5 10.0
Pasta (boiled) 159 677 8.5 75.0 16.5
Rice (white) 138 587 8.5 84.0 7.5
Beef (stewing) 203 852 42.5 0 57.5
Haddock (steamed) 89 378 6.0 0 94.0
Chicken (roasted) 177 742 38.0 0 62.0
Eggs (boiled) 147 612 66.0 Tr 34.0
Milk (summer) 65 270 55.4 23.6 21.0
Cheese (cheddar) 416 1725 75.5 0.1 24.4
Yogurt (plain) 79 333 34.2 37.0 28.8
Jam (fruit) 261 1114 0 99.1 0.9
Sugar 394 1680 0 99.9 Tr
Chocolate 520 2177 53.1 41.0 5.9
Butter 744 3059 99.4 0.3 0.3
Margarine 746 3067 99.9 0.1 Tr
(polyunsaturated)
Fat spread (60%) 553 2274 99.0 0.9 0.1
(polyunsaturated)
Sunflower oil 899 3696 100.0 0.0 Tr
Tr, trace.
Reference: Food Standards Agency (2002). McCance and Widdowson’s The Composition of Foods,
6th summary edition. Cambridge: Royal Society of Chemistry.
Crown copyright material is reproduced with the permission of the Controller of HMSO and the
Queen’s Printer for Scotland.
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The use of energy by the body 7

THE USE OF ENERGY BY THE BODY supplied to maintain the powerful pumping action of
the heart, the continual expansion and contraction of
As we have already noted, energy is produced in the the lungs and the temperature of the blood. It is
body from food by a series of precisely controlled needed to maintain the ceaseless chemical activity of
steps. Each step results in the release of a small the millions of body cells and the tone of muscles.
amount of energy which is used in the promotion of Living muscle must constantly be ready to contract in
bodily functions and may finally produce heat. response to stimuli transmitted to it by nerves. Such a
Without attempting at this stage any chemical degree of readiness can only be achieved if energy is
explanation of the nature of the steps in the oxida- continually supplied to keep the muscles in a state of
tion process, it is possible to gain a simple picture of mild tension. Muscle tension decreases during sleep
how energy is used by the body. but does not become zero, so that a certain amount of
Energy is required in the body for basal metab- energy, which ultimately appears as heat, is necessary
olism, which includes all the normal processes of cell to maintain it. The BMR is mainly determined by
respiration, maintenance and repair in the resting body weight and composition, which vary with both
status, for thermogenesis or heat production, for age (older people tend to have less muscle and more
growth and for physical activity. A proportion of body fat, so relatively lower BMR) and sex (women
food energy is also used for diet-induced thermo- tend to have higher proportion of body fat, so rela-
genesis to digest and process (store) the nutrients tively lower BMR).
after each meal. This amount is more for protein Table 1.4 gives average BMR values for men and
and carbohydrate than for fat. women of different ages and body weights. It will be
noted that the BMR of men is greater per kilogram
Basal metabolism of body weight than that of women of the same
weight. This is because, for a given body weight,
The term metabolism refers to the sum of all the women’s bodies contain more fat than those of men
chemical reactions going on in the body, and the and fat contributes little to BMR.
energy needed to sustain the body at complete rest Basal metabolic rate also varies with age. As age
is known as the energy of basal metabolism or the increases, BMR (per unit weight) falls; this decrease
basal metabolic rate (BMR). is rather greater for men than for women. Climate
The body requires a constant supply of energy to also affects BMR, its value being increased by 5–10
maintain its internal processes even when resting. per cent in very cold and very hot climates. We are
Even during sleep, when the body is apparently at most comfortable, and BMR is lowest, between 20
rest, energy is needed to ensure that essential internal and 28°C with light clothing. The thyroid gland
processes continue. For example, energy must be regulates cellular metabolism throughout the body.

Table 1.4 Basal metabolic rate (BMR) values

Males Females
Age (years) Weight (kg) BMR (MJ/day) Weight (kg) BMR (MJ/day)

10–17 30 5.0 30 4.6

65 7.6 60 6.3
18–29 60 6.7 45 4.8
80 7.9 70 6.4
30–59 65 6.8 50 5.2
85 7.7 70 5.9
Reference: Department of Health (1991). Dietary Reference Values for food Energy and Nutrients
for the United Kingdom. Report on Health and Social Subjects No. 41. London: HMSO.
Crown copyright material is reproduced with the permission of the Controller of HMSO and the
Queen’s Printer for Scotland.
Chap-01.qxd 13-02-2006 12:55 Page 8

8 Food and its functions

Excess thyroid activity increases BMR. If thyroid requirement of the muscles. It is useful, therefore,
function is defective, BMR declines. Many other to relate the degree of physical activity to the energy
hormones affect metabolic rate including insulin that must be supplied by the diet.
and adrenaline. The problem of equating physical activity with
energy requirement is complicated by the fact that
the body is unable to convert energy that is supplied
Growth
by food completely into mechanical work. The effi-
During the 9 months of pregnancy it is calculated ciency of conversion by the body, considered as a
that some 70 000 kcal (293 MJ) (DOH 1991, p. 30) machine, is of the order of 15–20 per cent. If the
of energy are required to produce the baby, increase higher value is taken it means that 100 units of energy
the size of the placenta and reproductive organs, supplied by food enable the body to perform phys-
allow for the energy (increased BMR) needed for ical work, for example, running, equivalent to 20
the newly formed baby tissues and create additional units. The other 80 units appear as heat and account
stores of fat in the mother which will be used for the fact that heat is lost from the body surface at
during lactation. Once the baby has been born, an increased rate when physical work is done. How-
lactation requires around 600 kcal (2.5 MJ) per day ever, this total energy cost and thus dietary need is
(Garrow et al. 2000). 100 units.
Newly born infants grow at a remarkable rate and Overweight and obese people need more energy
in the first 3 months of life 23 per cent of food for movement, in proportion to their extra weight.
energy is required for growth. This figure falls to
6 per cent by the time the infant is 1 year old and to ENERGY REQUIREMENTS
2 per cent by the fifth year. In general terms it has
been estimated that the formation of 1 g of new The energy requirement of an individual is the same
tissue requires 20 kJ of food energy. as energy expenditure, unless there is some growth
or weight loss. The estimated average requirement for
Physical activity energy by a group of people may be defined using
the concept of physical activity level (PAL). This
Energy is needed to enable the body to perform is the ratio of overall daily energy expenditure to
external work. Physical activity requires a supply of BMR. Daily energy expenditure may be calculated
energy additional to that needed to maintain by adding together energy expenditure on an hour-
muscle tone and other internal processes. The simplest by-hour basis at work, in non-work activities and
physical act, such as standing up, involves the use of while sleeping.
many muscles, and the greater the degree of phys- The values of PAL in different activities are shown
ical activity in daily life the greater is the energy in Table 1.5. Different occupations can be grouped

Table 1.5 Calculated physical activity level (PAL) of adults

Occupational activity
Non-
occupational Light Moderate Moderate/heavy
activity Male Female Male Female Male Female

Non-active 1.4 1.4 1.6 1.5 1.7 1.5

Moderately active 1.5 1.5 1.7 1.6 1.8 1.6
Very active 1.6 1.6 1.8 1.7 1.9 1.7
Reference: Department of Health (1991). Dietary Reference Values for Food Energy and Nutrients
for the United Kingdom. Report on Health and Social Subjects No. 41. London: HMSO. Crown
copyright material is reproduced with the permission of the Controller of HMSO and the Queen’s
Printer for Scotland.
Chap-01.qxd 13-02-2006 12:55 Page 9

Energy requirements 9

to indicate approximate PAL figures for estimated Table 1.6 Estimated average requirements (EAR) for
average 24 hour energy expenditure is as follows: energy (MJ/day)

1 Light. Includes professional and technical work- Physical activity level (PAL)
ers, administrative and managerial, sales repre- Weight (kg) 1.4 1.5 1.6 1.8 2.0
sentatives, clerical workers,
2 Moderate. Includes sales workers, domestic Males 10–18 years
helpers, transport workers, light construction 30 7.0 7.5 8.0 9.0 9.9
workers, e.g. joiners, 45 8.5 9.1 9.7 11.0 12.2
60 10.1 10.8 11.5 12.9 14.4
3 Moderate/heavy. Includes labourers, agricultural
Males 19–29 years
and fishing and forestry workers, heavy construc-
60 9.3 10.0 10.7 12.0 13.4
tion workers, e.g. bricklayers. 70 10.2 11.0 11.7 13.2 14.6
80 11.1 11.9 12.7 14.3 15.9
In general terms, most population groups in the UK Males 30–59 years
may be assumed to have light occupations and non- 65 9.5 10.2 10.8 12.2 13.5
active, non-work activities and, consequently, have a 75 10.2 10.9 11.6 13.1 14.5
PAL value of 1.4. 85 10.8 11.6 12.4 13.9 15.5
The EAR values shown in Table 1.6 are obtained Females 10–18 years
by multiplying the appropriate PAL value by BMR, 30 6.4 6.9 7.3 8.2 9.2
values of which are shown in Table 1.4. 45 7.6 8.1 8.7 9.8 10.8
60 8.8 9.4 10.0 11.3 12.5
For people over 60 years of age a standard PAL
Females 18–29 years
value of 1.5 is used. The EAR values for men
45 6.8 7.2 7.7 8.7 9.7
and women aged over 60 years decrease with 60 8.1 8.7 9.2 10.4 11.5
increasing age, although precise information for the 70 8.9 9.6 10.2 11.5 12.8
energy requirements of the elderly is lacking. It is Females 30–59 years
estimated that for men and women aged over 75 50 7.3 7.9 8.4 9.4 10.5
years the EAR values are 8.8 and 7.6 MJ/day 60 7.8 8.4 8.9 10.0 11.2
respectively. 70 8.3 8.9 9.5 10.7 11.8
For young children and babies EAR values are Reference: Department of Health (1991). Dietary Reference Values for
calculated by interpolating from values calculated food Energy and Nutrients for the United Kingdom. Report on Health
from data for older children. and Social Subjects No. 41. London: HMSO. Crown copyright
material is reproduced with the permission of the Controller of
HMSO and the Queen’s Printer for Scotland.

Key points
Definitions
● Nutritional science
– What people eat and absorb
– What people are
– How these effect what people (can) do
● Food science: what is in food
● Food technology: optimises quality and/or profitability of foods

Chapter summary
Foods provide a variety of nutrients in a matrix form. All foods and most nutrients contribute to the
energy supply, as part of a chain which harnesses energy (calories) from the sun and presents it as a
form which can contribute to metabolism, growth and physical activity. Amongst the macronutrients
which supply energy, the richest source is fat (9 kcal/g) followed by protein (4) and carbohydrate (3.75).
Chap-01.qxd 13-02-2006 12:55 Page 10

10 Food and its functions

FURTHER READING
DEPARTMENT OF HEALTH (1991). Dietary Reference Values for Food Energy and Nutrients for the United
Kingdom. Report on Health and Social Subjects No. 41. London: HMSO.
EUROPEAN PARLIAMENT (2001). Directive 2001/83/EC of the European Parliament on the
Community code relating to medicinal products for human use. Official Journal L311 28.11 2001, p. 71.
FOOD STANDARDS AGENCY (2002). McCance and Widdowson’s The Composition of Foods, 6th sum-
mary edition. Cambridge: Royal Society of Chemistry. ftp://ftp.fao.org/codex/manual/Manual12ce.pdf
GARROW JS, JAMES WPT, RALPH A (2000). Human Nutrition and Dietetics, 10th edn. London:
Churchill Livingstone.
Chap-02.qxd 13-02-2006 12:56 Page 11

2
Enzymes and digestion

Enzymes 12 Digestion 17
Cell metabolism 16 Further reading 22

The human body is composed of some hundred- the many functions of the cell, involves thousands
thousand million cells, each of which is a functional of different reactions. Each of these reactions com-
unit, enclosed within its cell membrane. These cells prises many steps which must be carried out in a
are grouped together in the body to form tissues definite sequence with the result that the chemical
with specialized functions. Thus, some cells comprise operations of a cell are much more complicated,
connective tissue and bind together the various and need much greater integration, than those of
organs of the body, others are concerned with muscu- a chemical factory. They must also be capable of
lar and nervous tissue, while others form the skeletal adaptation to enormous variations in conditions and
framework of bone that contributes strength and fluctuating nutrient supply.
rigidity to the body. Most organs contain several In order to sustain life, the cell’s activities must be
functional cell types. controlled and organized into a self-regulating and
Individual cells are so tiny that their internal struc- self-renewing pattern. But how can such control
ture can only be observed when magnified many be achieved, and how is it that although almost all
hundreds or thousands of times with the help of an human cells are built according to the same basic
electron microscope. Within the cell there are a pattern, they are able to perform a multitude of dif-
number of different types of very much smaller ferent functions? The answer to these questions is
‘subcellular organelles’. These have specialized func- to be found in the existence of a group of crucially
tions, and their size, structure and number vary widely important protein substances called enzymes.
according to demands, or to regulation by local Enzymes control all the chemical changes (that
nutrient supply, and instructions delivered by nerves is, the metabolism), which occur in living cells. They
or hormones to the cell. regulate the building up (anabolic) reactions that
The complex activities needed to sustain life in result in the formation of complex substances such
the human body take place within the body’s cells. as proteins from single building units. They also regu-
The activity of a cell is often likened to that of a late the breaking-down (catabolic) reactions that
chemical factory in which a great variety of raw result in release of energy. Enzymes can activate or
materials are processed and converted into finished catalyse chemical reactions or inhibit them. Enzymes
products. In a single cell many different raw materials often increase the cycling or turnover of substances
are required, though they are largely composed of in cells (i.e. stimulate both synthesis and breakdown
only four elements: carbon, hydrogen, oxygen and of compounds). This allows processes to be more
nitrogen. The processing stage, which is concerned rapidly regulated (in terms of direction and speed
with the conversion of these simple raw materials into of reaction) by other factors. Anabolic and catabolic
the more complex substances required to carry out processes involve very many steps, and each step is
Chap-02.qxd 13-02-2006 12:56 Page 12

12 Enzymes and digestion

controlled by its own enzyme. This control must A feature of all enzymes is that their tertiary struc-
be carefully regulated so that the life of the cell ture includes a site that ‘recognizes’ and can bind
continues smoothly at all times, with the whole temporarily to immobilize its substrate.
metabolic process being kept carefully balanced.
Classification of enzymes
ENZYMES The substance upon which an enzyme acts is called
the substrate, and enzymes are usually named after
How different cells perform different functions can this substance. Thus, the enzyme that acts on (breaks
be explained in terms of the enzymes that are pre- down) urea is called urease and that which acts on
sent. Some 1000 different enzymes have been recog- maltose is called maltase. It is a general rule that
nized in the body, but in any one cell only a selection enzymes are named after the substrate upon which
are present. Even so, most cells contain about 200 they act and given the suffix -ase (from Latin). In
different enzymes, each of which is responsible for common with most general rules, however, there
controlling a particular step. The complement of are notable exceptions, mainly those enzymes which
enzymes present in a cell automatically selects and were named before the rule gained general accept-
controls those reactions which are to proceed. Since ance. Some of these, such as pepsin and trypsin, will
enzymes are proteins, their presence in cells in turn be encountered later.
depends on the DNA of the cell, including the DNA Enzymes may be classified in a number of ways,
sequence which for the synthesis of each enzyme but one of the most useful is to group them accord-
and for its own storage and metabolism. ing to the type of reaction which they control. The
five main groups of enzyme are shown in Table 2.1.
Of these, the first two are the most important in
The chemical nature of enzymes connection with what follows. Hydrolases control
the hydrolysis of the substrate, that is its reaction
One of the most widely known sources of enzymes with water and, as we shall see later in the chapter,
is yeast, which is one of the simplest possible types this type of enzyme is of paramount importance
of living organism. This is how the name enzyme in digestion. Oxidases control the oxidation of the
arose, for it means literally ‘in yeast’. The fermenta- substrate, and this usually takes the form of removal
tion of sugar in grape juice by yeast and the leaven- of hydrogen as indicated in the equation shown
ing effect of yeast in making bread have been used in Table 2.1. There is often more than one enzyme
for many centuries. These are actions of enzymes. which carries out a particular function – very simi-
A long time elapsed between the discovery of the lar ‘iso-enzymes’ may be produced in different
first known enzyme in yeast and the isolation of an tissues – each with its own gene and separate
enzyme in the pure state. For many years it was
believed that enzymes were living organisms. This
idea was only shown to be false at the end of the last Table 2.1 Classification of enzymes
century when the German chemist Buchner extracted
Reaction General
a cell-free liquid from yeast cells that had a similar
Name catalysed equation
enzyme activity to the original living cells. So, although
enzymes are made by living cells, they themselves Hydrolases Hydrolysis AB  H2O → AOH  BH
are not living. Oxidases Oxidation ABH2 → AB  2H
All enzymes are proteins, the structure of which is Isomerases Intramolecular ABC → ACB
described in Chapter 10. The properties of proteins – rearrangement
their ability to change their shape, their sensitivity Transferases Transfer of a AB  C → A  BC
to changes of conditions of temperature and acidity, group
their capacity to oppose changes of acidity that Synthetases Addition of A  B → AB
would upset the smooth working of the cell – make one molecule
them peculiarly suited to control cell metabolism. to another
Chap-02.qxd 13-02-2006 12:56 Page 13

Enzymes 13

system for regulation. A very common feature of thousands, that of nature’s catalysts is measured in
enzyme function is that activation of an enzyme, millions.
or the expression of the gene, are often inhibited by How do enzymes work? We can best approach this
the product of the reaction catalysed. This prevents by considering first how an ordinary non-catalytic
wastage of energy on unnecessary metabolism, and reaction proceeds. Suppose a reaction involves the
prevents accumulation. conversion of reacting substances represented by A
into products represented by B. The reaction will
not start until A has received a ‘push’ in the form
Catalytic action of enzymes of energy, often supplied in the form of heat. The
reason for this can be appreciated from Fig. 2.1.
Enzymes are organic catalysts; they operate by speed- Before A can react to form B it must surmount
ing up a chemical process while remaining unchanged the energy hump shown by moving along path
at the end of the reaction. In many respects their (i), and this requires an amount of energy DG*
action is similar to that of the more familiar inorganic (uncatalysed). When A has absorbed energy DG*
catalysts such as are often used in manufacturing (uncatalysed), known as the activation energy, it is in
processes. In the manufacture of margarine, vegetable an activated state and can decompose to form B.
oils are converted into solid fats by chemical reaction This process can be likened to that of transferring
with hydrogen. In the absence of a catalyst the con- a ball from one side of a hill to the other. If Fig. 2.1
version of the oils into fats is very slow indeed, but the represents a hill, the problem is that of transferring
addition of small quantities of finely divided nickel a ball from X to Y. If the only path lies over the
produces a remarkable increase in the rate of the reac- summit of the hill, then it is necessary to push the
tion; moreover, the nickel catalyst may be used time ball up the hill – that is, to work on it by supplying
after time, as it is not used up in the process. energy – until it reaches the top. Once there, it will
It is remarkable that only one part of nickel is run down the other side to Y of its own accord. The
needed to catalyse the conversion of several-thousand ball may now be at a lower level than it was at its
parts of oil into fat, but this achievement appears starting point, as is the case in Fig. 2.1.This means that
quite insignificant when compared with the startling an amount of energy DG* has been released, though
catalytic power of enzymes. One of the enzymes con- this could not have been achieved without first
cerned with the breakdown of starch during digestion pushing the ball to the top of the hill, which involved
is amylase, produced by the pancreas. Only one part supplying it with energy DG* (uncatalysed).
of amylase is needed to effect the conversion of four In the cells of the human body, activation energy
million parts of starch into the sugar maltose. Where cannot be supplied in normal ways, such as heat,
the efficiency of man-made catalysts is measured in because this would damage the cells. The function

Energy G*
(uncatalysed)

(i) G*
(catalysed)
(ii) Activation
energy

Energy of
reactants A X G°
Energy of
Y products B

Figure 2.1 Reaction paths: (i) non-catalytic

Reactants Products path; (ii) catalytic path
Chap-02.qxd 13-02-2006 12:56 Page 14

14 Enzymes and digestion

of enzymes is to enable the reaction to proceed at a the time taken to reach the equilibrium position.
much lower activation energy than would other- In the absence of the enzyme the reaction may be so
wise be possible. In terms of Fig. 2.1, we must replace slow that, for all practical purposes, it does not pro-
the reaction path (i) over the summit of the hill by ceed at all. In a cell, thousands of different reactions
one at a lower level such as (ii) involving a lower are possible but the function of the enzymes pre-
activation energy DG* (catalysed). sent is to speed up particular ones, so that some
Enzymes catalyse reactions by replacing a single- reactions proceed rapidly while others proceed at a
step of a high-energy mechanism with a two- or relatively insignificant rate. In this way, cell metab-
multi-stage process, each step of which involves a olism is controlled and directed so that different cells
low activation energy. If the enzyme catalyst is are able to fulfil different functions.
represented as E and the substrate as A we have:

A  E 7 AE Selectivity of enzymes
AE 7 B  E
Overall reaction: A 7 B Enzymes are usually highly selective in which reac-
tion they catalyse. Frequently one enzyme will catal-
Within each cell is its nucleus, which mainly con- yse only one cell reaction. It is this characteristic
tains genetic material (chromosomes) and this is sur- which allows them to preserve order in living cells.
rounded by a watery fluid called cytoplasm, which Thus, although many other reactions may occur in a
contain water-soluble enzymes. The cytoplasm cell, the rate at which they proceed is insignificant
contains a network of membrane-like material, the compared with that of the catalysed reaction.
endoplasmic reticulum, which is studded with small The selective power of enzymes is sometimes com-
dark bodies known as ribosomes. The cytoplasm also pared with the action of a key in a lock: the enzyme is
contains a number of bodies, among which are the the lock and only certain molecules can act as the key
egg-shaped mitochondria and the smaller lysosomes. which exactly fits it. If a reaction between two mol-
These bodies contain enzymes which are lipid- ecules is catalysed by an enzyme the lock can be imag-
soluble within their membranes. Ribosomes are ined as having grooves into which the two molecules
responsible for translating the DNA code in chromo- fit side by side; this leads to a brief union between the
somes by assembling amino acids in the correct order enzyme and the two molecules which are acting as
to make proteins and enzymes. Mitochondria house keys, as is shown in stage 2 of Fig. 2.2. The key mol-
the enzymes for oxidation (combustion) of nutrients ecules are thus brought together and converted into
to generate chemical energy (as ATP). Lysosomes an active state which enables them to react with each
include the enzymes which release enzymes to attack other. After reaction new molecules are formed, but
foreign material and infecting organisms. as shown in stage 3 of the diagram, the enzyme
Enzymically catalysed reactions proceed by way of remains unchanged and can catalyse further reaction.
a temporary enzyme–substrate complex represented In order to achieve a good fit between lock and
by AE. If both stages require little energy the reac- key molecules, an enzyme frequently needs the help
tion path is as represented by Fig. 2.1 (ii) and the of another substance to be effective. Three main
reaction will be a rapid and near spontaneous one types of enzyme promotor have been distinguished,
with the enzyme E being regenerated. Thus, we have namely coenzymes, cofactors (or activators) and pros-
a simple picture of how enzymes act as catalysts. thetic groups (see p. 149). Coenzymes are smaller than
It should be noted that although enzymes speed enzyme molecules and are not proteins. They are
up reactions, and direct substrates towards one spe- not permanently bound to the enzyme but may
cific reaction when others are possible, they cannot become attached to it during enzyme reaction (Fig.
turn impossible reactions into possible ones. Neither 2.2, stage 1) only to be released later (once stage 3,
can they affect the equilibrium position of a reversible Fig. 2.2, has been completed). Coenzymes are closely
reaction; this means that the amount of product in a related to vitamins or are the vitamins themselves.
reaction is the same whether an enzyme is involved One of the functions of the B group of vitamins
or not. The presence of the enzyme merely reduces seems to be to provide the body with suitable
Chap-02.qxd 13-02-2006 12:56 Page 15

Enzymes 15

Enzyme surface

C C C

Stage 1 Stage 2 Stage 3

Lock and key molecules Union and activation of New molecules formed
before union. lock and key molecules leaving enzyme surface
C is the coenzyme or unchanged Figure 2.2 The lock and key
prosthetic group theory of enzyme reaction

starting materials from which to make the coen- other substance (not even a similar sugar). Maltase
zymes that it needs. The exact function of coenzymes and sucrase, however, catalyse the hydrolysis not
is still only partly understood, but they certainly play only of maltose and sucrose but also that of certain
an active and vital part in many reactions involving other similar sugars.
oxidizing enzymes. This is shown by the fact that if a A further example of enzymes which show a
coenzyme needed by an enzyme is absent, the remarkable selectivity are those which catalyse the
enzyme can exert no catalytic effect. This can be used hydrolysis of proteins. The three enzymes, pepsin,
to measure the status of essential nutrients. For exam- trypsin and chymotrypsin, each select certain links
ple, thiamine (vitamin B1) is a coenzyme for the of protein molecules and catalyse hydrolysis only at
enzyme transketolase. Transketolase activity in blood these links (see p. 148).
is a simple laboratory measure of thiamine status.
In some cases it is found that metallic ions, such as
magnesium, or non-metallic ions, such as chloride, are Sensitivity of enzymes
required to increase the activity of enzymes. Such
substances are known as cofactors (or activators). Enzymes are very sensitive to effects of temperature
Prosthetic groups are non-protein groups which and the environment.All enzyme activity is destroyed
are permanently bound to the enzyme. on boiling because, being proteins, enzymes are dena-
Sometimes several molecules, which are similar tured (see p. 141) by high temperatures. At low
to each other, can approximately fit the grooves of temperatures enzyme activity is greatly slowed down
the same lock. In such cases the enzyme does not but as the enzymes are not denatured and, because
distinguish between them and acts as a catalyst to enzyme reactions have a low activation energy,
them all. In most cases, however, enzymes show great enzymes may retain some catalytic activity even at
powers of discrimination as the following examples subzero temperatures. In general, plant enzymes work
show.The three enzymes, maltase, lactase and sucrase, best at about 25°C and those in warm-blooded ani-
are present in the small intestine, and during diges- mals at about 37–40°C. An increase in temperature
tion these enzymes catalyse the hydrolysis of the usually increases the rate of a chemical reaction, but
sugars maltose, lactose and sucrose respectively. in the case of an enzyme reaction it may also lead to
These three enzymes have considerable specificity, inactivation of the enzyme.
and in the case of lactase, complete specificity, since Figure 2.3 shows the effect of temperature on the
it will catalyse the hydrolysis of lactose and of no rate of catalysis. At 37°C, the initial rate of reaction is
Chap-02.qxd 13-02-2006 12:56 Page 16

16 Enzymes and digestion

enzyme trypsin which is present in pancreatic juice

37°C
requires a slightly alkaline medium before it can
catalyse protein hydrolysis. When food passes from
Amount of product formed

the stomach into the small intestine the hydrochloric

acid is completely neutralized and the medium
70°C becomes alkaline. Under these conditions pepsin
becomes inactivated and trypsin carries on the
digestion of proteins.

CELL METABOLISM
Time All activities that occur in cells are controlled by
Figure 2.3 The effect of temperature on the rate of enzymes. However, these enzymes are not evenly
catalysis by an enzyme distributed throughout the cell but are dispersed
among the different parts of the cell so that each
rapid, but after a time the reaction rate slows down part has a distinct role in maintaining the life of
and stops, no further product being formed. This may the cell.
have one of several causes. For example, the reaction The nucleus of a cell contains all the genes (coded
may be complete, all the substrate having reacted, or in DNA), which can produce any of the enzymes
it may be that the products of reaction have made the found in cells. The nucleus also contains a small
environment unfavourable for enzyme activity and number of enzymes that, together, specifically con-
the enzyme has been deactivated. If the temperature trol cell (nuclear) growth and division. The surround-
is raised to 70°C, the initial rate of reaction is ing cytoplasm contains water-soluble enzymes that
increased. This is because the greater energy input control a variety of anabolic and catabolic processes.
increases the energy of substrate and enzyme mol- The mitochondria are important because they are the
ecules, and these molecules gain the activation energy powerhouses of the cell and contain a number of oxi-
needed for reaction to take place more rapidly. dases responsible for the manufacture of high-energy
Although the initial rate of formation of products is materials used for energy production. The important
rapid, it soon stops because the enzyme is rapidly job of protein synthesis in the cell is controlled by
inactivated at the higher temperature. The net result enzymes found in the endoplasmic reticulum. Some
is that less product is formed at the higher tempera- of the enzymes found only in mitochondria are syn-
ture than at the lower. Enzymes catalyse reaction thesized from RNA inside mitochondria, rather than
efficiently in man, the temperature being high enough from DNA in the nucleus. Mitochondria reproduce
to give rapid formation of products but low enough in a parallel but separate way to the rest of the cell.
to avoid inactivation of the enzyme. Finally, one of the most interesting bodies in the cell
Enzyme activity is also dependent upon the acidity is the lysosome, also sometimes called the ‘suicide
or alkalinity (the pH) of the medium in which the bag’. Lysosomes contain a sufficient variety of hydro-
enzyme acts. Most enzymes operate most efficiently lases capable of destroying nearly all the components
in an environment that is nearly neutral, and if the of the cell. Normally these ‘suicide’ enzymes are
medium becomes strongly acid or alkaline the enzyme safely contained within the impermeable membrane
becomes completely inactivated. Some enzymes, that encloses the lysosome, but if the cell becomes
however, can only operate in an acid or alkaline injured or dies, the enzymes are released and the cell
solution. For example, the enzyme pepsin is present destroys itself and also any infecting agent.
in gastric juice and during digestion it catalyses Each cell, and each component of a cell, is sur-
the initial hydrolysis of proteins. It can only act in rounded by a membrane. The membrane allows only
strongly acid conditions such as are produced by the those raw materials needed by the cell to pass through
hydrochloric acid in the stomach. Conversely, the it, all other substances being prevented from
Chap-02.qxd 13-02-2006 12:56 Page 17

Digestion 17

entering. This selection mechanism ensures that only food component, is the proportion that is actually
those substances required for a particular job are absorbed after passing through the two (incomplete)
available, and it also ensures that only those enzymes processes of digestion and absorption. Once in the
required to control this function are allowed through bloodstream, nutrients are distributed to all the cells
the membrane. Most cell membranes are ‘semi- of the body where they sustain the complex processes
permeable’, that is, they allow water, and sometimes of metabolism. So you are not what you eat –
small molecules dissolved in water, to pass freely although what you are reflects what you eat.
but larger molecules can only pass through if there
are specific transport mechanisms.
In later chapters we shall develop the theme of The role of enzymes in digestion
what happens to nutrients in metabolism in rather
more detail, but enough has been said in this The chemical processes involved in digestion are
introductory survey to show that all the activities brought about by enzymes. The chemical breakdown
and functions of the body which constitute life are of food molecules into absorbable components,
entirely dependent upon enzymes. which in the absence of enzymes would be very
slow indeed, is thereby speeded up so that digestion
is completed in a matter of hours. Thus, in the space
of some 3–4 hours a remarkable change in the nature
DIGESTION of the food has occurred. Substances such as starch,
which may contain as many as 150 000 atoms in a
‘You are what you eat.’
single molecule, have been converted into molecules
(An error! – Anon) containing only 24 atoms (simple sugars such as glu-
It is indeed a very odd thing – an extraordinary and cose). The breakdown of protein molecules is almost
remarkable thing – that no matter what we eat, the equally spectacular, as an average protein molecule
structure of the body, both flesh and blood, changes is split up into about 500 amino acid molecules
very little. There is no obvious similarity between during digestion. These two examples perhaps make
the nature of the food we eat and the nature of our clearer the magnitude of the chemical task performed
bodies. Yet within a few hours of being eaten, food by the enzymes of the digestive system.
is transformed into flesh and blood. This transforma- Each stage of digestion involves hydrolysis and is
tion is so complete that it cannot be accomplished catalysed by a hydrolysing enzyme or hydrolase. The
before food has undergone a drastic breaking-down hydrolysis can be represented:
process known as digestion.
AB  H2O  AOH  BH.
Digestion is both physical and chemical: the physi-
cal process involves the breakdown of large food par- The equation shows how water is involved in split-
ticles into smaller ones while the chemical process ting up a molecule AB into two smaller molecules
involves the breakdown of larger molecules into AOH and BH. In some instances (e.g. sucrose), a sin-
smaller ones. Foodstuffs are mainly complicated, gle step involving the breakdown of a molecule into
insoluble substances that must be converted into sim- two parts is sufficient to produce a smaller soluble
pler, soluble, more active ones before they can be used molecule that can be absorbed. In other instances
by the body. Not all nutrients need digesting, how- (e.g. proteins), a very large number of hydrolytic
ever, as there are some such as water and simple sug- steps is required before breakdown is complete.
ars (e.g. glucose) that do not need to be broken down The digestive process involves a fairly small
and many vitamins and mineral salts which must not number of different enzymes which catalyse the
be broken down if they are to be useful. Whether or chemical breakdown of proteins, carbohydrates and
not nutrients need to be broken down by digestion fats. The names of the hydrolases that catalyse the
they cannot be utilized by the body until they have hydrolysis of these different nutrients are shown in
passed into the bloodstream, a process which is Table 2.2. Unfortunately, different authorities use
known as absorption. The availability of a nutrient, or different names for these enzymes. There is an
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