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ANIMAL NUTRITION
 At Pearson, we have a simple mission: to help people
make more of their lives through learning.

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Every day our work helps learning flourish, and

wherever learning flourishes, so do people.

To learn more, please visit us at www.pearson.com/uk

ANIMAL NUTRITION
EIGHTH EDITION

The late P McDonald

Formerly Reader in Agricultural Biochemistry, University of
Edinburgh, and Head of the Department of Agricultural
Biochemistry, Edinburgh School of Agriculture

The late R A Edwards

Formerly Head of the Department of Animal Nutrition,
Edinburgh School of Agriculture

J F D Greenhalgh
Emeritus Professor of Animal Production and Health,
University of Aberdeen

C A Morgan
Formerly animal nutritionist, the Scottish Agricultural College

L A Sinclair
Harper Adams University

R G Wilkinson
Harper Adams University

Harlow, England • London • New York • Boston • San Francisco • Toronto • Sydney
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PEARSON EDUCATION LIMITED
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_____________________________
First published by Oliver & Boyd 1966 (print)
Second edition published 1973 (print)
Third edition published 1981 (print)
Fourth edition published 1988 (print)
Fifth edition published 1995 (print)
Sixth edition published 2002 (print)
Seventh edition published 2011 (print)
Eighth edition published 2022 (print and electronic)
© Pearson Education Limited 2002, 2011 (print)
© Pearson Education Limited 2022 (print and electronic)
© P McDonald, R A Edwards, J F D Greenhalgh, C A Morgan, L A Sinclair and R G Wilkinson
The rights of P McDonald, R A Edwards, J F D Greenhalgh, C A Morgan, L A Sinclair and R G Wilkinson to be identified as authors of
this work have been asserted by them in accordance with the Copyright, Designs and Patents Act 1988.
The print publication is protected by copyright. Prior to any prohibited reproduction, storage in a retrieval system, distribution or
transmission in any form or by any means, electronic, mechanical, recording or otherwise, permission should be obtained from the
publisher or, where applicable, a licence permitting restricted copying in the United Kingdom should be obtained from the Copy-
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The ePublication is protected by copyright and must not be copied, reproduced, transferred, distributed, leased, licensed or publicly
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tions under which it was purchased, or as strictly permitted by applicable copyright law. Any unauthorised distribution or use of this
text may be a direct infringement of the authors’ and the publisher’s rights and those responsible may be liable in law accordingly.
All trademarks used herein are the property of their respective owners. The use of any trademark in this text does not vest in the
author or publisher any trademark ownership rights in such trademarks, nor does the use of such trademarks imply any affiliation
with or endorsement of this book by such owners.
Pearson Education is not responsible for the content of third-party internet sites.
ISBN: 978-1-292-25166-0 (print)
978-1-292-25168-4 (PDF)
978-1-292-25167-7 (ePub)
British Library Cataloguing-in-Publication Data
A catalogue record for the print edition is available from the British Library
Library of Congress Cataloging-in-Publication Data
Names: McDonald, Peter, 1926-2018, author. | Edwards, R. A., author. |
Greenhalgh, J. F. D., author. | Morgan, C. A. (Animal nutritionist),
author. | Sinclair, L. A. (Liam A.), author. | Wilkinson, R. G. (Robert
G.), author.
Title: Animal nutrition / P. McDonald, Formerly Reader in Agricultural
Biochemistry, University of Edinburgh, and Head of the Department of
Agricultural Biochemistry, Edinburgh School of Agriculture, R.A.
Edwards, Formerly Head of the Department of Animal Nutrition, Edinburgh
School of Agriculture, J.F.D. Greenhalgh, Emeritus Professor of Animal
Production and Health, University of Aberdeen, C.A. Morgan, Scottish
Agricultural College, L.A. Sinclair, Harper Adams University College,
R.G. Wilkinson, Harper Adams University College.
Description: Eighth edition. | Harlow, England ; New York : Pearson, 2022.
| Includes bibliographical references and index.
Identifiers: LCCN 2021044367 (print) | LCCN 2021044368 (ebook) | ISBN
9781292251660 (paperback) | ISBN 9781292251684 (ebook) | ISBN
9781292251677 (epub)
Subjects: LCSH: Animal nutrition. | Feeds.
Classification: LCC SF95 .M38 2022 (print) | LCC SF95 (ebook) | DDC
636.08/52--dc23
LC record available at https://lccn.loc.gov/2021044367
LC ebook record available at https://lccn.loc.gov/2021044368
10 9 8 7 6 5 4 3 2 1
26 25 24 23 22
Front cover image: Photos by R A Kearton/Moment/Getty Images
Cover design by Kelly Miller
Print edition typeset in 9.5/12pt Rotation LT Std by Straive
Printed by Ashford Colour Press Ltd, Gosport
NOTE THAT ANY PAGE CROSS REFERENCES REFER TO THE PRINT EDITION
 Contents

Preface to the eighth edition xii

Part 1
THE COMPONENTS OF FOODS 1

1 The animal and its food 3

1.1 Water 4
1.2 Dry matter and its components 5
1.3 Analysis and composition of foods 5
Summary 14
Questions 14
Further reading 14

2 Carbohydrates 16
2.1 Classification of carbohydrates 16
2.2 Monosaccharides 18
2.3 Monosaccharide derivatives 20
2.4 Oligosaccharides 23
2.5 Polysaccharides 26
2.6 Lignin 30
Summary 30
Questions 31
Further reading 31

3 Lipids 32
3.1 Classification of lipids 32
3.2 Fats 33
3.3 Glycolipids 43
3.4 Phospholipids 45
3.5 Waxes 47
3.6 Steroids 48
3.7 Terpenes 51
Summary 51
Questions 52
Further reading 52

4 Proteins, nucleic acids and other nitrogenous compounds 53

4.1 Proteins 53
4.2 Amino acids 53
4.3 Peptides 59
4.4 Structure of proteins 60

v
Contents

4.5 Properties of proteins 61

4.6 Classification of proteins 62
4.7 Nucleic acids 63
4.8 Other important nitrogenous compounds 66
4.9 Nitrates 68
4.10 Alkaloids 68
Summary 68
Questions 69
Further reading 69

5 Vitamins 70
5.1 Introduction 70
5.2 Fat-soluble vitamins 74
5.3 The vitamin B complex 87
5.4 Vitamin C 100
5.5 Hypervitaminosis 101
5.6 Vitamins and gene expression 101
Summary 102
Questions 103
Further reading 103

6 Minerals 105
6.1 Functions of minerals 105
6.2 Natural and supplementary sources of minerals 109
6.3 Acid–base balance 112
6.4 Major elements 115
6.5 Trace elements 124
6.6 Other elements 139
Summary 139
Questions 140
Further reading 140

Part 2
THE DIGESTION AND METABOLISM OF NUTRIENTS 141

7 Enzymes 143
7.1 Classification of enzymes 144
7.2 Nature of enzymes 147
7.3 Mechanism of enzyme action 149
7.4 Specific nature of enzymes 151
7.5 Factors affecting enzyme activity 153
7.6 Nomenclature of enzymes 157
Summary 158
Questions 158
Further reading 158

8 Digestion 159
8.1 Digestion in monogastric mammals and fowl 159
8.2 Microbial digestion in ruminants and other herbivores 175

vi
 Contents

8.3 Alternative sites of microbial digestion 191

8.4 Nutrient digestion and the environment 193
Summary 197
Questions 199
Further reading 199
Historical reference 200

9 Metabolism 201
9.1 Energy metabolism 203
9.2 Protein synthesis 224
9.3 Fat synthesis 232
9.4 Carbohydrate synthesis 238
9.5 Control of metabolism 244
Summary 245
Questions 246
Further reading 246

Part 3
QUANTIFYING THE NUTRIENT CONTENT OF FOODS:
DIGESTIBILITY, ENERGY AND PROTEIN SUPPLY 247

10 Evaluation of feeds: digestibility 249

10.1 Measurement of digestibility 250
10.2 Validity of digestibility coefficients 255
10.3 Digestibility in different sections of the digestive tract 256
10.4 Factors affecting digestibility 259
10.5 Measurement of mineral availability 263
Summary 264
Questions 264
Further reading 265

11 Evaluation of foods: energy content of foods

and energy partition 266
11.1 Energy demand 266
11.2 Energy supply and partition 267
11.3 Animal calorimetry: methods of measuring heat production
and energy retention 274
11.4 Utilisation of metabolisable energy 283
Summary 290
Questions 291
Further reading 292

12 Evaluation of foods: systems for expressing energy

supply and requirements 293
12.1 Energy systems and energy models 294
12.2 Energy systems for ruminants 295
12.3 Energy systems for pigs and poultry 305
12.4 Energy systems for horses 307
12.5 Energy systems for dogs and cats 309

vii
Contents

12.6 Predicting the energy value of foods 310

Summary 312
Questions 313
Further reading 314
Historical references 314

13 Evaluation of foods: protein 315

13.1 Crude protein (CP) 315
13.2 Digestible crude protein (DCP) 317
13.3 Determination of endogenous nitrogen 318
13.4 Measures of protein quality for monogastric animals 320
13.5 Measures of food protein used in practice in the feeding
of pigs and poultry 327
13.6 Measures of food protein used in practice in the feeding
of horses 329
13.7 Measures of food protein quality for dogs and cats 331
13.8 Measures of protein quality for ruminant animals 331
13.9 The UK metabolisable protein system 344
13.10 The UK Feed into Milk (FiM) protein system for dairy cows 346
Summary 351
Questions 352
Further reading 353

Part 4
THE NUTRIENT REQUIREMENTS OF ANIMALS 355

14 Feeding standards for maintenance and growth 357

14.1 Nutrient requirements for maintenance 359
14.2 Nutrient requirements for growth 377
14.3 Nutrient requirements for wool production 391
14.4 Mineral and vitamin requirements for maintenance and growth 393
14.5 Nutritional control of growth 395
Summary 399
Questions 400
Further reading 400
Historical reference 401

15 Feeding standards for reproduction 402

15.1 Nutrition and the initiation of reproductive ability 403
15.2 Plane of nutrition, fertility and fecundity 405
15.3 Egg production in poultry 410
15.4 Nutrition and the growth of the foetus 414
Summary 423
Questions 423
Further reading 424

16 Lactation 426
16.1 Sources of the milk constituents 427
16.2 Nutrient requirements of the lactating dairy cow 431
16.3 Nutrient requirements of the lactating ewe 454

viii
 Contents

16.4 Nutrient requirements of the lactating dairy goat 460

16.5 Nutrient requirements of the lactating sow 464
16.6 Nutrient requirements of the lactating mare 468
16.7 Nutrient requirements of the lactating dog and cat 472
Summary 477
Questions 478
Further reading 479

17 Voluntary intake of food 481

17.1 Food intake in pigs and poultry 482
17.2 Food intake in ruminants 488
17.3 Food intake in horses 493
17.4 Food intake in dogs and cats 494
17.5 Prediction of food intake 495
Summary 497
Questions 497
Further reading 498

Part 5
THE NUTRITIONAL CHARACTERISTICS OF FOODS 499

18 Grass and forage crops 501

18.1 Pastures and grazing animals 501
18.2 Grasses 502
18.3 Legumes 512
18.4 Other forages 515
Summary 518
Questions 519
Further reading 519

19 Silage 521
19.1 Silage, ensilage and silos 521
19.2 Role of plant enzymes in ensilage 523
19.3 Role of microorganisms in ensilage 523
19.4 Nutrient losses in ensilage 527
19.5 Classification of silages 529
19.6 Nutritive value of silages 534
19.7 Whole crop cereal and legume silages 540
Summary 543
Questions 544
Further reading 544

20 Hay, artificially dried forages, straws and chaff 545

20.1 Hay 545
20.2 Artificially dried forages 550
20.3 Straws and related by-products 551
Summary 556
Questions 556
Further reading 556

ix
Contents

21 Roots, tubers and related by-products 557

21.1 Roots 557
21.2 Tubers 561
Summary 563
Questions 563
Further reading 564

22 Cereal grains and cereal by-products 565

22.1 The nutrient composition of grains 565
22.2 Barley 567
22.3 Maize 573
22.4 Oats 575
22.5 Wheat 577
22.6 Other cereals 578
22.7 Cereal processing 580
Summary 584
Questions 585
Further reading 585

23 Protein concentrates 586

23.1 Oilseed cakes and meals 586
23.2 Oilseed residues of minor importance 598
23.3 Leguminous seeds 599
23.4 Processed animal protein (PAP) 602
23.5 Milk products 607
23.6 Single-cell protein and microalgae 609
23.7 Insect protein 611
23.8 Synthetic amino acids 614
23.9 Non-protein nitrogen compounds as protein sources 615
Summary 618
Questions 619
Further reading 619

24 Food additives 621

24.1 Antibiotics 621
24.2 Probiotics 623
24.3 Oligosaccharides 626
24.4 Enzymes 627
24.5 Organic acids 630
24.6 Spray-dried plasma 631
24.7 Modifiers of rumen fermentation 632
24.8 Plant extracts 633
Summary 636
Questions 636
Further reading 636

x
 Contents

Part 6
ANIMAL PRODUCTS AND HUMAN NUTRITION 639

25 Animal nutrition and the consumers of animal products 641

25.1 Comparative nutrition 641
25.2 The contribution of animal products to human requirements 643
25.3 Objections to the use of animal products 647
25.4 Animal products: past, present and future 652
Summary 653
Questions 653
Further reading 653

Appendix 1: Solutions to numerical questions 655

Appendix 2: Notes on tables 661
Index 697
Publisher’s acknowledgements 731

xi
 Preface to the eighth edition

Recent research in the field of animal science has focused on advances in molecular
biology, particularly in the study of gene expression, epigenetics and gene editing,
and exciting advances have been made. However, knowledge of animal biochemistry
and nutrition is still essential if we are to understand the significance and efficient
application of these new findings to further improve animal production, health and
welfare. The application of research and advice in animal nutrition continues to be at
the centre of efficient animal production. Research in dog and cat nutrition has also
progressed since the last edition and information in this area has been expanded in
this new edition.
We have retained the early chapters on basic food chemistry and animal bio-
chemistry to provide a quick reference to questions pertaining to the discipline of
nutrition chemistry in later parts of the book. We have also taken the opportunity to
introduce nutritional topics related to molecular biology and the environment. Each
chapter now has a set of questions to assist with revision of the chapter topic and the
Appendix tables have been revised where new data are available.
Three significant events have occurred since the last edition. In 2016, the British
Society of Animal Science recognised the 50th anniversary of the publication of the
first edition of Animal Nutrition by awarding framed certificates of congratulation to
the original three authors, Peter McDonald, James Greenhalgh and Alun Edwards.
Then, in 2018, came the sad news that Peter McDonald had died and in 2021 that
Alun Edwards, too, had died. Although Peter and Alun had not been actively involved
in the production of recent editions of the book, they had always shown great interest
in its progress. Fittingly, Peter’s funeral service was conducted by another eminent
animal nutritionist, Rev. Dr Neville Suttle.
The production of this edition was assisted by comments and suggestions received
from reviewers and we welcome comments from readers. As with previous editions,
we are grateful to colleagues for their helpful discussions.

C A Morgan, L A Sinclair, R G Wilkinson and J F D Greenhalgh

2022

xii
PART 1

The components of foods

This part describes the chemistry of foods and the components that supply nutrients to the
animal.

Chapter 1 is concerned with the analysis of foods, from the early chemical analysis developed in
the 1800s to categorise chemical and nutrient groups, through to the sophisticated physical and
chemical methods used today to identify individual molecular components.

Chapters 2, 3 and 4 describe the major components of foods that supply energy and amino acids,
i.e. the carbohydrates and lipids, and the proteins.

Chapters 5 and 6 give details of the nutrients required in smaller amounts, the vitamins and
minerals which, nevertheless, are essential for the normal functions of the body and efficient
animal production.
1 The animal and its food

1.1 Water
1.2 Dry matter and its components
1.3 Analysis and composition of foods

Food is material that, after ingestion by animals, is capable of being digested, absorbed
and utilised. n a more general sense, we use the term food to describe edible mate-
rial. rass and hay, for example, are described as foods, but not all their components
are digestible. Where the term food is used in the general sense, as in this text, those
components capable of being utilised by animals are described as nutrients.
The animals associated with human beings cover the spectrum from herbivores – the
plant eaters (ruminants, horses and small animals such as rabbits and guinea pigs) through
omnivores, which eat all types of food (pigs and poultry) to carnivores, which eat chiefly
meat (cats). Under the control of human beings, these major classes of animal still pertain,
but the range of foods that animals are now offered is far greater than they might normally
consume in the wild (for example, ruminants are given plant by-products of various human
food industries and some dog foods contain appreciable amounts of cereals). Nevertheless,
plants and plant products form the major source of nutrients in animal nutrition.
The diet of farm animals in particular consists of plants and plant products, although
some foods of animal origin, such as fishmeal and milk, are used in limited amounts.
Animals depend upon plants for their existence and consequently a study of animal
nutrition must necessarily begin with the plant itself.
Plants are able to synthesise complex materials from simple substances, such as carbon
dioxide from the air, and water and inorganic elements from the soil. By means of pho-
tosynthesis, energy from sunlight is trapped and used in these synthetic processes. The
greater part of the energy, however, is stored as chemical energy within the plant itself
and it is this energy that is used by the animal for the maintenance of life and synthesis
of its own body tissues. Plants and animals contain similar types of chemical substances,
and we can group these into classes according to constitution, properties and function.
The main components of foods, plants and animals are:
Carbohydrates
Water
Lipids
Proteins
Food Organic Nucleic acids
Organic acids
Dry matter Vitamins
Inorganic Minerals
3
Chapter 1 The animal and its food

1.1 WATER
The water content of the animal body varies with age. The newborn animal contains
750–800 g/kg water, but this falls to about 500 g/kg in the mature fat animal. It is
vital to the life of the organism that the water content of the body be maintained:
an animal will die more rapidly if deprived of water than if deprived of food. Water
functions in the body as a solvent in which nutrients are transported about the body
and in which waste products are excreted. Many of the chemical reactions brought
about by enzymes take place in solution and involve hydrolysis. Because of the high
specific heat of water, large changes in heat production can take place within the
animal with very little alteration in body temperature. Water also has a high latent
heat of evaporation, and its evaporation from the lungs and skin gives it a further
role in the regulation of body temperature.
The animal obtains its water from three sources: drinking water, water present in its
food and metabolic water, this last being formed during metabolism by the oxidation of
hydrogen-containing organic nutrients. The water content of foods is variable and can
range from as little as 60 g/kg in concentrates to over 900 g/kg in some root crops. Because
of this great variation in water content, the composition of foods is often expressed on a
dry matter basis, which allows a more valid comparison of nutrient content. This is illus-
trated in Table 1.1, which lists a few examples of plant and animal products.
The water content of growing plants is related to the stage of growth, being
greater in younger plants than in older plants. In temperate climates, the acquisition
of drinking water is not usually a problem and animals are provided with a continu-
ous supply. There is no evidence that under normal conditions an excess of drinking
water is harmful, and animals normally drink what they require.

Table 1.1 Composition of some plant and animal products expressed on a fresh
basis and a dry matter basis

Water Carbohydrate Lipid Protein Ash

Fresh basis (g/kg)
Turnips 910 70 2 11 7
Grass (young) 800 137 8 35 20
Barley grain 140 730 15 93 22
Groundnuts 60 201 449 268 22
Dairy cow 570 2 206 172 50
Milk 876 47 36 33 8
Muscle 720 6 44 215 15
Egg 667 8 100 118 107
Dry matter basis (g/kg)
Turnips 0 778 22 122 78
Grass (young) 0 685 40 175 100
Barley grain 0 849 17 108 26
Groundnuts 0 214 478 285 23
Dairy cow 0 5 479 400 116
Milk 0 379 290 266 65
Muscle 0 21 157 768 54
Egg 0 24 300 355 321

4
 Analysis and composition of foods

1.2 DRY MATTER AND ITS COMPONENTS

The dry matter (DM) of foods is conveniently divided into organic and inorganic
material, although in living organisms there is no such sharp distinction. Many
organic compounds contain mineral elements as structural components. Proteins, for
example, contain sulphur, and many lipids and carbohydrates contain phosphorus.
It can be seen from Table 1.1 that the main component of the DM of pasture grass
is carbohydrate, and this is true of all plants and many seeds. The oilseeds, such as
groundnuts, are exceptional in containing large amounts of protein and lipid material.
In contrast, the carbohydrate content of the animal body is very low. One of the main
reasons for the difference between plants and animals is that, whereas the cell walls
of plants consist of carbohydrate material, mainly cellulose, the walls of animal cells
are composed almost entirely of lipid and protein. Furthermore, plants store energy
largely in the form of carbohydrates such as starch and fructans, whereas an animal’s
main energy store is in the form of lipid.
The lipid content of the animal body is variable and is related to age, the older ani-
mal containing a much greater proportion than the young animal. The lipid content of
living plants is relatively low – that of pasture grass, for example, being 40–50 g/kg DM.
In both plants and animals, proteins are the major nitrogen-containing compounds.
In plants, in which most of the protein is present as enzymes, the concentration is
high in the young growing plant and falls as the plant matures. In animals, the muscle,
skin, hair, feathers, wool and nails consist mainly of protein.
Like proteins, nucleic acids are also nitrogen-containing compounds and they play
a basic role in the synthesis of proteins in all living organisms. They also carry the
genetic information of the living cell.
The organic acids that occur in plants and animals include citric, malic, fumaric,
succinic and pyruvic acids. Although these are normally present in small quantities,
they nevertheless play an important role as intermediates in the general metabolism
of the cell. Other organic acids occur as fermentation products in the rumen, or in
silage, and these include acetic, propionic, butyric and lactic acids.
Vitamins are present in plants and animals in minute amounts, and many of them
are important as components of enzyme systems. An important difference between
plants and animals is that, whereas the former can synthesise all the vitamins they
require for metabolism, animals cannot, or have very limited powers of synthesis,
and are dependent upon an external supply.
The inorganic matter contains all those elements present in plants and animals
other than carbon, hydrogen, oxygen and nitrogen. Calcium and phosphorus are the
major inorganic components of animals, whereas potassium and silicon are the main
inorganic elements in plants.

1.3 ANALYSIS AND COMPOSITION OF FOODS

Originally, the most extensive information about the composition of foods was based
on a system of analysis described as the proximate analysis of foods, which was
devised in the 19th century by two German scientists, Henneberg and Stohmann.
More recently, new analytical techniques have been introduced, and the information
about food composition is rapidly expanding (see the ‘Modern analytical methods’

5
Chapter 1 The animal and its food

section). However, the system of proximate analysis still forms the basis for the
statutory declaration of the composition of foods in Europe.

Proximate analysis of foods

This system of analysis divides the food into six fractions: moisture, ash, crude pro-
tein, ether extract, crude fibre and nitrogen-free extractives.
The moisture content is determined as the loss in weight that results from drying
a known weight of food to constant weight at 100 °C. This method is satisfactory
for most foods, but with a few (such as silage), significant losses of volatile material
(short-chain fatty acids and alcohols) may take place. Therefore, for silages, the mois-
ture content can be determined directly by distilling the water from the sample under
toluene. The distillate is measured and corrected for the presence of fermentation
acids and alcohols. An estimate of the ‘corrected’ dry matter (g/kg) can be calculated
from the oven dry matter (ODM g/kg) using a relationship such as that published by
the Agriculture and Food Research Council:
Corrected DM = 0.99 ODM + 18.9
The ash content is determined by ignition of a known weight of the food at 550 °C
until all carbon has been removed. The residue is the ash and is taken to represent the
inorganic constituents of the food. The major component of ash is silica but ash may,
however, contain material of organic origin such as sulphur and phosphorus from
proteins, and some loss of volatile material in the form of sodium, chloride, potas-
sium, phosphorus and sulphur will take place during ignition. The ash content is thus
not truly representative of the inorganic material in the food, either qualitatively or
quantitatively. Animals do not have a requirement for ash per se but require the indi-
vidual mineral elements that it contains and are determined by methods such as atomic
absorption spectrometry or inductively coupled plasma spectroscopy (see p. 11).
The crude protein (CP) content is calculated from the nitrogen content of the food,
determined by a modification of a technique originally devised by Kjeldahl over
100 years ago. In this method, the food is digested with sulphuric acid, which con-
verts all nitrogen present, except that in the form of nitrate and nitrite, to ammonium
sulphate. The ammonia is liberated by adding sodium hydroxide to the digest, dis-
tilled off and collected in standard acid, and the quantity collected is determined by
titration or by an automated colorimetric method. It is assumed that the nitrogen is
derived from protein containing 16 per cent nitrogen, and by multiplying the nitro-
gen figure by 6.25 (i.e. 100/16) an approximate protein value is obtained. This is not
‘true protein’, since the method determines nitrogen from sources other than protein,
such as free amino acids, amines and nucleic acids, and the fraction is therefore des-
ignated ‘crude protein’. Also, the nitrogen content of the protein varies according to
its amino acid composition and 16 per cent is an average value.
The ether extract (EE) fraction is determined by subjecting the food to a con-
tinuous extraction with petroleum ether for a defined period. The residue, after
evaporation of the solvent, is the ether extract. As well as lipids it contains organic
acids, alcohol and pigments. This procedure is referred to as method A. In the current
official method, the extraction with ether is preceded by hydrolysis of the sample
with hydrochloric acid and the resultant residue is the acid ether extract (method B).

6
 Analysis and composition of foods

The carbohydrate of the food is contained in two fractions, the crude fibre (CF)
and the nitrogen-free extractives (NFE). The former is determined by subjecting the
residual food from ether extraction to successive treatments with boiling acid and
alkali of defined concentration; the organic residue is the crude fibre.
When the sum of the amounts of moisture, ash, crude protein, ether extract and
crude fibre (expressed in g/kg) is subtracted from 1,000, the difference is designated
the nitrogen-free extractives. The nitrogen-free extractives fraction is a heteroge-
neous mixture of all those components not determined in the other fractions. The
crude fibre fraction contains cellulose, lignin (an indigestible component of plant
fibre) and hemicelluloses, but not necessarily the whole amounts of these that are
present in the food: a variable proportion of the cell wall material, depending upon
the species and stage of growth of the plant material, is dissolved during the crude
fibre extraction and thus is contained in the nitrogen-free extractives. This leads to
an underestimation of the fibre and an overestimation of the starch and sugars. Thus,
the nitrogen-free extractive fraction includes starch, sugars, fructans, pectins, organic
acids and pigments, in addition to those components mentioned above.

Modern analytical methods

In recent years the proximate analysis procedure has been severely criticised by
many nutritionists as being archaic and imprecise, and in the majority of laboratories
it has been partially replaced by other analytical procedures. Most criticism has been
focused on the crude fibre, ash and nitrogen-free extractives fractions for the reasons
described above. The newer methods have been developed to characterise foods in
terms of the methods used to express nutrient requirements. In this way, an attempt
is made to use the analytical techniques to quantify the potential supply of nutrients
from the food. For example, for ruminants, analytical methods are being developed
that describe the supply of nutrients for the rumen microbes and the host digestive
enzyme system (Fig. 1.1).

Starch and sugars

Inadequacies in the nitrogen-free extractives fraction have been addressed by the
development of methods to quantify the non-structural carbohydrates, which are
mainly starches and sugars. Sugars can be determined colorimetrically after combi-
nation with a reagent such as anthrone. The official EC method involves extraction
of sugars with dilute ethanol; the solution is then clarified and the ethanol removed.
The sugars are then quantified before and after inversion (giving reducing and total
sugars, respectively) by the Luff-Schoorl method. Starch is determined by dilute
acid hydrolysis of the sample followed by polarimetric determination of the released
sugars. This gives a figure for total sugars (i.e. those originating from the hydrolysed
starch plus the simple sugars in the food). Sugars per se are determined by extracting
the sample with ethanol, acidifying the filtrate and taking a second polarimeter read-
ing. The starch content is calculated from the difference between the two readings
multiplied by a known factor for the starch source. Starch can also be determined
enzymically. For example, in cereals starch is converted to glucose using a-amylase
followed by amyloglucosidase and then the glucose is measured using the glucose
oxidase-peroxidase reagent.

7
Chapter 1 The animal and its food

Food measurements Rumen parameters Absorbed nutrients

Total N
NF N solubility
UDN DUP AA
N degradability
ADIN ERDN Microbial MP
amino
acids
WSC and pectins
Fermentable
Starch – Rate
PFF energy
Extent
Cell walls – Rate VFA
Extent Microbial
fatty acids
ME

VFA LCFA
Non-fermentable
NFF Lactate VFA
energy
Lipid Glucose

Fig. 1.1 Proposed model for characterisation of foods for ruminants.

AA = amino acids, AD N = acid detergent insoluble nitrogen, DUP = digestible undegradable
protein, ERDN = effective rumen degradable nitrogen, CFA = long-chain fatty acids,
ME = metabolisable energy, MP = metabolisable protein, N = nitrogen, NF = nitrogen fraction,
NFF = non-fermentable fraction, PFF = potentially fermentable fraction, UDN = undegradable
nitrogen, FA = volatile fatty acids, WSC = water-soluble carbohydrates.
From Agricultural and Food Research Council 1998 Technical Committee on Responses to Nutrients,
report no. 11, Wallingford, CAB .

Fibre
Alternative procedures for fibre have been developed by Van Soest (Table 1.2).
The neutral-detergent fibre (NDF), which is the residue after extraction with boil-
ing neutral solutions of sodium lauryl sulphate and ethylenediamine tetraacetic acid
(EDTA), consists mainly of lignin, cellulose and hemicellulose and can be regarded
as a measure of the plant cell wall material. The analytical method for determining
NDF was originally devised for forages, but it can also be used for starch-containing
foods provided that an amylase treatment is included in the procedure. By analogy
with the nitrogen-free extractives fraction discussed above, the term non-structural
carbohydrate (NSC) is sometimes used for the fraction obtained by subtracting the
sum of the amounts (g/kg) of CP, EE, ash and NDF from 1,000.
The acid-detergent fibre (ADF) is the residue after refluxing with 0.5 M sulphuric
acid and cetyltrimethyl-ammonium bromide, and represents the crude lignin and
cellulose fractions of plant material but also includes silica.
The determination of ADF is particularly useful for forages as there is a good
statistical correlation between it and the extent to which the food is digested (digest-
ibility). In the UK, the ADF method has been modified slightly by altering the dura-
tion of boiling and acid strength. The term modified acid-detergent fibre (MADF) is
used to describe this determination.

8
 Analysis and composition of foods

Table 1.2 Classification of forage fractions using the detergent methods of

an Soest

Fraction Components
Cell contents (soluble in neutral detergent) Lipids
Sugars, organic acids and
water-soluble matter
Pectin, starch
Non-protein nitrogen
Soluble protein
Cell wall constituents (fibre insoluble
in neutral detergent)
Soluble in acid detergent Hemicelluloses
Fibre-bound protein
Acid-detergent fibre Cellulose
Lignin
Lignified nitrogen
Silica
After an Soest P J 1967 Journal of Animal Science 26: 119.

The acid-detergent lignin determination involves the preparation of acid-detergent

fibre as the preparatory step. The ADF is treated with 72 per cent sulphuric acid,
which dissolves cellulose. Ashing the residue determines crude lignin, including cutin.
The Van Soest methods of fibre analysis and other standard techniques are used
for fractionation of food carbohydrates for ruminants in the Cornell net carbohydrate
and protein system (CNCPS) to derive the following fractions:
1. Total carbohydrate = 100 - (crude protein + fat + ash)
2. Non-structural carbohydrate (NSC) = 100 - (crude protein + fat +
(NDF - NDF protein) + ash)
3. Sugar as a proportion of NSC
4. Starch, pectins, b-glucans, volatile fatty acids = NSC - sugar
5. Lignin
The carbohydrates are then classified according to their degradation rate by rumen
microbes: fraction A – fast (comprising the sugars); fraction B1 – intermediate (starch,
pectins, b-glucans); fraction B2 – slow (available cell wall material represented by
lignin-free NDF); and fraction C – indigestible (unavailable cell wall in the form of
lignin).
In monogastric, and particularly human, nutrition the term dietary fibre is often
used and attention has been focused on its importance in relation to health. Dietary
fibre (DF) was defined as lignin plus those polysaccharides that cannot be digested
by monogastric endogenous enzymes. Initially, epidemiological studies linked a lack
of DF to constipation, gut and bowel disorders, cardiovascular disease and type 2
diabetes; however, the causes of such diseases are multifactorial and in some cases

9
Chapter 1 The animal and its food

it is not just DF per se that has the beneficial effects but other aspects of the diet
also (e.g. antioxidants). Nevertheless, DF is a major component related to health in
human beings and it has equally important effects in animals.
The definition of DF has proved difficult, with definitions ranging through physi-
ological/botanical (derived from cell walls of plants, which are poorly digested),
chemical/botanical (non-starch polysaccharides (NSP) of plant cell walls), chemical
(NSP and lignin) and nutritional/physiological (NSP not digested in the small intes-
tine). The common features of DF definitions are carbohydrates (polysaccharides,
oligosaccharides and lignin) resistant to digestion in the small intestine but that may
be fermented in the large intestine and promote beneficial physiological effects. By
virtue of its definition, DF is difficult to determine in the laboratory. The NSP in most
foods, along with lignin, are considered to represent the major components of cell
walls. Methods for measurement of NSP fall into two categories (with slight varia-
tions in the second category, depending on the research laboratory):
■ Enzymic–gravimetric methods, which measure a variety of components and give
no details of polysaccharide type. In the method of the Association of Official
Analytical Chemists for total dietary fibre, samples are gelatinised by heating
and treated with enzymes to remove starch and proteins. The total dietary fibre
is precipitated with ethanol and the residue is dried and weighed.
■ Enzymic–chromatographic methods, which identify the individual carbohydrates
in the dietary NSP. The Englyst method can be used to determine total, soluble
and insoluble dietary fibre. Measurement of NSP by this method involves removal
of starch with the enzymes pullulanase and a-amylase. After precipitation with
ethanol, the NSP residue is then hydrolysed with 12 M sulphuric acid. The individ-
ual monomeric neutral sugar constituents are determined by gas–liquid chroma-
tography (see later) with separate determination of uronic acids. Alternatively, the
total sugars are determined colorimetrically after reaction with dinitrosalicylate
solution. Total NSP and insoluble NSP are determined directly by analysis of
separate subsamples and the soluble NSP are calculated by difference. The major
constituents of NSP are rhamnose, arabinose, xylose, glucose, galactose, mannose
and glucuronic and galacturonic acids. Cellulose is the major source of glucose,
and hemicellulose provides xylose, mannans and galactose. The degradation of
pectins releases arabinose, galactose and uronic acids. Following the adoption
of methods to determine NSP, it became apparent that non-digestible oligosac-
charides and resistant starch also contributed to DF based on their physiological
behaviour. In recognition of this, enzymic procedures have been developed to
determine these components.
In recent years, attention has focused on the importance of both the soluble and
insoluble forms of fibrous material in the human diet. Water-soluble NSP is known
to lower serum cholesterol, and insoluble NSP increases faecal bulk and speeds up
the rate of colonic transit. This last effect is thought to be beneficial in preventing a
number of diseases, including cancer of the bowel.
The NSP of foods may be degraded in the gut of pigs by microbial fermentation,
yielding volatile fatty acids, which are absorbed and contribute to the energy supply.
A further benefit relates to the volatile fatty acid butyric acid, which is reported to be
an important source of energy for the growth of cells in the epithelium of the colon;
thus, the presence of this acid will promote development of the cells and enhance
absorption. The extent of degradation depends on the conformation of the polymers

10
 Analysis and composition of foods

and their structural association with non-carbohydrate components, such as lignin.

In addition, the physical properties of the NSP, such as water-holding capacity and
ion exchange properties, can influence the extent of fermentation. The gel-forming
NSPs, such as b-glucans. influence the viscosity of digesta and thereby reduce the
absorption of other nutrients from the small intestine and depress digestibility and
adversely affect faecal consistency in pigs and poultry (see the section on enzymes
in Chapter 24). On a positive note, the water-holding properties lead to beneficial
effects on the behaviour of pregnant sows by increasing time spent eating and rest-
ing, owing to increased gut fill, and by reducing inappropriate behaviour, such as
bar chewing.

Minerals
A simple ash determination provides very little information about the exact mineral
make-up of the food and, when this is required, analytical techniques involving
spectroscopy are generally used. In atomic absorption spectroscopy, an acid solu-
tion of the sample is heated in a flame and the vaporised atoms absorb energy,
which brings about transitions from the ground state to higher energy levels. The
source of energy for this transition is a cathode lamp, containing the element to
be determined, which emits radiation at a characteristic wavelength. The radiation
absorbed by the atoms in the flame is proportional to the concentration of the
element in the food sample.
Flame emission spectroscopy measures the radiation from solutions of the sample
heated in air/acetylene or oxygen/acetylene flames. Each element emits radiation
at specific wavelengths and there are published tables of flame emission spectra.
Atomic absorption and flame emission spectrometry are being replaced by induc-
tively coupled plasma emission spectroscopy, as this has a greater sensitivity for the
relatively inert elements and can be used to determine several elements simultane-
ously or sequentially. Energy from the inductively coupled plasma source is absorbed
by argon ions and elements to form a conducting gaseous mixture at temperatures
up to 10,000 °C. The electromagnetic radiation emitted from atoms and ions within
the plasma is then measured. Alternatively, the ions can be separated and detected
using a mass spectrometer.
Just as with other nutrients, a measure of the concentration of the element alone
is not sufficient to describe its usefulness to the animal. Attempts have been made to
assess the availability of minerals using chemical methods, such as solubility in water
or dilute acids, but these have had little success. At present, animal experiments are
the only reliable way to measure mineral availability (see Chapter 10).

Amino acids, fatty acids and sugars

As an alternative to the standard Kjeldahl method for the determination of nitrogen
(crude protein) described above, the Dumas method is also now used. In this method
the sample is combusted in pure oxygen; the products are carbon dioxide, water,
oxides of nitrogen and nitrogen. The carbon dioxide and water are absorbed on
columns and the oxides of nitrogen are converted to nitrogen with a column packed
with copper; the resulting total nitrogen is determined in a thermal conductivity
detector. This method, although expensive in equipment, is rapid and does not rely
on hazardous chemicals.
Knowledge of the crude protein content of a food is not a sufficient measure of its
usefulness for non-ruminants. The amino acid composition of the protein is required

11
Chapter 1 The animal and its food

in order to assess how a food can meet the essential amino acid requirements (see
Chapter 4). Similarly, the total ether extract content does not give sufficient infor-
mation on this fraction since it is important to know its fatty acid composition. In
non-ruminants, this has large effects on the composition of body fat and, if soft fat
is to be avoided, the level of unsaturated fatty acids in the diet must be controlled.
In ruminants, a high proportion of unsaturates will depress fibre digestion in the
rumen. When detailed information on the amino acid composition of protein, the
fatty acid composition of fat or the individual sugars in NSP is required, then tech-
niques involving chromatographic separation can be used. In gas–liquid chromato-
graphy, the stationary phase is a liquid held in a porous solid, usually a resin, and the
mobile phase is a gas. Volatile substances partition between the liquid and the vapour
and can be effectively isolated. This form of chromatography is, however, usually a
slow process; in order to speed up the separation procedure, high-performance liquid
chromatography has been developed. In this technique, pressure is used to force a
solution, containing the compounds to be separated, rapidly through the resin held in
a strong metal column. In addition to speeding up the process, high resolution is also
obtained. Gas–liquid chromatography and high-performance liquid chromatography
can also be used for the determination of certain vitamins (e.g. A, E, B6, K), but the
measurement of available vitamins requires biological methods.
An example of the application of high-performance liquid chromatography is seen
with food proteins, which are hydrolysed with acid and the released amino acids are
then determined using one of the following methods:
■ Ion-exchange chromatography – by which the amino acids are separated on the
column, and then mixed with a derivatisation agent, which reacts to give a com-
plex that is detected by a spectrophotometer or fluorimeter.
■ Reverse-phase chromatography – in which the amino acids react with the reagent
to form fluorescent or ultraviolet-absorbing derivatives, which are then separated
using a more polar mobile phase (e.g. acetate buffer with a gradient of acetonitrile)
and a less polar stationary phase (e.g. octadecyl-bonded silica). The availability of
amino acids to the animal can be estimated by chemical methods. For example, for
lysine there are colorimetric methods that depend on the formation of compounds
between lysine and dyes (see Chapter 13).

Measurement of protein in foods for ruminants

The new methods of expressing the protein requirements of ruminants (see
Chapter 13) require more information than just the crude protein (nitrogen) content
of the food. The unavailable nitrogen is measured as acid detergent insoluble nitro-
gen. Information on the rate of degradation in the rumen of the available nitrogen
is also required and this can be estimated by biological methods. In the Cornell
net carbohydrate and protein system, the neutral and acid detergent extractions of
Van Soest, described previously, are used in combination with extraction with a
borate–phosphate buffer and trichloracetic acid solution to derive several protein
fractions. These fractions describe the components that are degraded in the rumen
or digested in the small intestine (see Chapter 13).

Spectroscopy
It is now common for laboratories to use near-infrared reflectance spectroscopy
(NIRS) to estimate the composition of foods. The basis of this methodology lies

12
 Analysis and composition of foods

in the absorption of energy by hydrogen-containing functional groups in organic

compounds present in the food (C–H, O–H, N–H and S–H). The reflected energy
from the sample provides information on its composition but, unlike normal spec-
troscopy, is not related directly to concentration since the sample is heterogenous.
Therefore, empirical relationships are derived by calibrating the reflected spec-
trum with samples of known composition, as determined by standard methods. In
practice, energy in the wavelength range 1,100–2,500 nm is directed on to a cell
containing the dried milled sample, and the diffuse reflected energy is measured
across the spectrum. The spectral data are then related to the known chemical com-
position of the standard samples by multiple linear regression. The relationships are
then validated with a second set of samples of known composition. Once satisfac-
tory relationships have been derived, they can be applied to the spectra of sam-
ples of unknown composition. The technique has been extended to the analysis of
fresh silage samples, eliminating the need to dry and mill the sample. NIRS has the
advantages that it is rapid with minimal sample preparation, it gives instantaneous
results and is non-destructive of the sample, it allows simultaneous measurement of
several parameters with high precision, and it allows a high throughput of samples
at low cost per sample. Nevertheless, it should be recognised that, as the technique
is dependent on chemical or physical calibrations, it cannot be more accurate than
the chemically determined values used to derive the calibration. NIRS is particularly
useful in the context of compound food manufacture, where rapid analysis of raw
materials and finished product is required for efficient mixing and quality control
standards. With forages, particularly grass and cereal silages, NIRS is now routinely
used to determine not only chemical composition but also a range of food character-
istics, including those that are the resultant of a number of nutrient concentrations
such as digestibility, metabolisable energy and nitrogen degradability in the rumen
and potential silage intake (see Chapters 12, 13 and 17). Hand-held NIRS devices
are now available for on-farm analysis of forages. Mineral elements do not have
absorptions in the NIR spectral region and cannot be determined directly by NIRS.
However, minerals can be measured when they are combined in organic complexes
or chelates, or indirectly by their effects on hydrogen bonds. Accuracy of estimation
depends on there being a close degree of correlation between the mineral and the
organic compounds.
Nuclear magnetic resonance spectroscopy is a complex technique that is used to
determine the constituents of foods. This method makes use of the fact that some
compounds contain certain atomic nuclei that can be identified from a nuclear mag-
netic resonance spectrum, which measures variations in frequency of electromagnetic
radiation absorbed. It provides more specific and detailed information of the con-
formational structure of compounds than, for example, NIRS but is more costly and
requires more time and skill on the part of the operator. For these reasons, it is more
suited to research work and for cases in which the results from simpler spectroscopy
techniques require further investigation. Nuclear magnetic resonance spectroscopy
has been useful in the investigation of the soluble and structural components of
forages.

13
Chapter 1 The animal and its food

SUMMARY
1. Water is an important component of 6. Fibrous constituents can be determined
animal foods. t contributes to the water by application of detergent solutions and
requirements of animals and dilutes the weighing the residue, or by the use of
nutrient content of foods. Water content enzymes followed by weighing or gas–liquid
varies widely between foods. chromatography.
2. The constituents of dry matter comprise 7. ndividual mineral elements are measured
carbohydrates (sugars, starches, fibres), by atomic absorption spectroscopy, flame
nitrogen-containing compounds (proteins, photometry or inductively coupled plasma
amino acids, non-protein nitrogen emission spectroscopy.
compounds), lipids (fatty acids, glycerides),
minerals and vitamins. 8. as–liquid chromatography is used to
determine individual amino acids, fatty acids
3. Analytical techniques have been developed and certain vitamins.
from simple chemical/gravimetric
determinations. 9. Near-infrared reflectance spectroscopy is used
routinely to determine food characteristics
4. Modern analytical techniques attempt to
and to predict nutritive value. Nuclear
measure nutrients in foods in terms of the
magnetic resonance spectroscopy is a research
nutrient requirements of the animal.
technique for determining the chemical
5. Starch is determined by polarimetry. structure of food components.

QUESTIONS
1.1 What are the major components of food dry matter? (p. 3)
1.2 What are the three major components of the organic matter of foods? (p. 5)
1.3 Grass has a dry matter content of 200 g/kg and 137 g carbohydrate/kg. What is
the concentration of carbohydrate in the dry matter?
1.4 Barley grain has a dry matter content of 860 g/kg and 108 g protein/kg DM.
What is the concentration of protein in the fresh matter?
1.5 What are the six fractions quantified by the proximate system of analysis of
foods? (p. 6)
1.6 What is the usually assumed nitrogen content of crude protein?
1.7 What do the following abbreviations mean: NDF, ADF, WSC, NSP, NSC?
(p. 8)
1.8 What is near infrared reflectance spectroscopy and how is it used to determine
the composition of a food? (p. 12)

FURTHER READING
Association of Official Analytical Chemists International 2019 Official Methods of Analysis,
21st edn, Arlington, VA, AOAC International.
Chalupa W and Sniffen C J 1994 Carbohydrate, protein and amino acid nutrition of
lactating dairy cattle. In: Garnsworthy P C and Cole D J A (eds) Recent Advances in
Animal Nutrition, Loughborough, Nottingham University Press, 265–75.

14
 Further reading

Champ M, Langkilde A-M, Brouns F, Kettlitz B and Le Bail Collet Y 2003 Advances in
dietary fibre characterization. 1. Definition of dietary fibre, physiological relevance,
health benefits and analytical aspects. Nutrition Research Reviews 16: 71–82.
European Commission 2009 Commission Regulation (EC) No.152/2009 Methods of
sampling and analysis for the official control of feed.
Givens D I, De Boever J L and Deaville E R 1997 The principles, practices and some future
applications of near infrared spectroscopy for predicting the nutritive value of foods for
animals and humans. Nutrition Research Reviews 10: 83–114.
Reeves J B 2000 Use of near infrared reflectance spectroscopy. In: D’Mello J P F (ed.) Farm
Animal Metabolism and Nutrition, Wallingford, CAB International, 185–207.
Van Soest P J 1994 Nutritional Ecology of the Ruminant, 2nd edn, Ithaca, NY, Comstock.
Van Soest P J, Robertson J B and Lewis B A 1991 Methods for dietary fiber, neutral
detergent fiber and non-starch polysaccharides in relation to animal nutrition. Journal of
Dairy Science 74: 3,583–3,597.

15
 2 Carbohydrates

2.1 Classification of carbohydrates

2.2 Monosaccharides
2.3 Monosaccharide derivatives
2.4 Oligosaccharides
2.5 Polysaccharides
2.6 Lignin

In general, carbohydrates are neutral chemical compounds containing the elements

carbon, hydrogen and oxygen and have the empirical formula (CH2O)n, where n is 3 or
more. However, some compounds with general properties of the carbohydrates also
contain phosphorus, nitrogen or sulphur; and others, e.g. deoxyribose (C5H10O4), do not
have hydrogen and oxygen in the same ratio as that found in water. The carbohydrate
group contains polyhydroxy aldehydes, ketones, alcohols and acids, their simple
derivatives, and any compound that may be hydrolysed to these.

2.1 CLASSIFICATION OF CARBOHYDRATES

The carbohydrates may be classified as shown in Fig. 2.1. The simplest sugars are
the monosaccharides, which are divided into subgroups – trioses (C3H6O3), tetroses
(C4H8O4), pentoses (C5H10O5), hexoses (C6H12O6) and heptoses (C7H14O7) –
depending on the number of carbon atoms present in the molecule. The trioses
and tetroses occur as intermediates in the metabolism of other carbohydrates and
their importance will be considered in Chapter 9. Monosaccharides may be linked
together, with the elimination of one molecule of water at each linkage, to produce
di-, tri-, tetra- or polysaccharides, containing, respectively, two, three, four or larger
numbers of monosaccharide units.
The term sugar is generally restricted to those carbohydrates containing fewer
than ten monosaccharide residues, while the name oligosaccharides (from the Greek
oligos, a few) is frequently used to include all sugars other than the monosaccharides.
Polysaccharides, also called glycans, are polymers of monosaccharide units. They
are classified into two groups, the homoglycans, which contain only a single type
of monosaccharide unit, and the heteroglycans, which on hydrolysis yield mixtures

16
 Classification of carbohydrates

Glyceraldehyde
Trioses (C3H6O3)
Dihydroxyacetone

Tetroses (C4H8O4) Erythrose

Arabinose
Xylose
Monosaccharides Pentoses (C5H10O5) Xylulose
Ribose
Ribulose

Glucose
Hexoses (C6H12O6) Galactose
Mannose
Fructose
Sugars
Heptoses (C7H14O7) Sedoheptulose

Sucrose
Disaccharides Lactose
Maltose
Cellobiose

Oligosaccharides Raffinose
Trisaccharides
Kestose

Tetrasaccharides Stachyose

Arabinans
Xylans
Starch
Dextrins
Glucans Glycogen
Cellulose
Homoglycans Callose

Fructans Inulin
Galactans Levan
Polysaccharides Mannans
Glucosamines

Pectic substances
Hemicelluloses
Heteroglycans Exudate gums
Non-sugars Acidic mucilages
Hyaluronic acid
Chondroitin

Glycolipids
Complex
carbohydrates
Glycoproteins

Fig. 2.1 Classification of carbohydrates.

17
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to be ignorant of this, and he prepared accordingly. A more
thoroughly girded man for the world’s encounter could hardly be
conceived than he at this time. He took no wine nor strong drink; he
lived simply of the simple, on anchorite’s fare, with more than
anchorite resolution, with the temptations of the world always before
him; he clothed plainly, and made the best of everything; he kept no
company, and found every amusement in his science books, his
experiments, in his business, and in simple exercise.
To fill up time till the money patients should come, he became one
of the visitors of the out-patients of Charing Cross Hospital; and to
many a poor representative of the great half-starved, extended a skill
which would have been a blessing to a duke. The Librarian of the
College of Surgeons’ Library knew him as a quiet man, who read
closely, and was not too proud to ask for a translation when an
original bothered him. All who knew him said he was a quiet man,
very reserved and peculiar—a clever man at bottom perchance, but
not easy to be understood and very peculiar.
The connection with the “Westminster Medical” led to Mr. Snow’s
first attempts at authorship. On October the 16th, 1841, he read at
the Society a paper on “Asphyxia and on the Resuscitation of new-
born Children.” The paper in full will be found in the London
Medical Gazette for November the 5th of the same year. The paper is
remarkable for the soundness of its reasonings, and the advanced
knowledge which it displays. The object of the paper was to introduce
to the Society a double air-pump, for supporting artificial
respiration, invented by a Mr. Read, of Regent Circus. The
instrument was so devised that by one action of the piston, the air in
the lungs could be drawn into one of the cylinders, and by the reverse
action, the said air could be driven away, and the lungs supplied with
a stream of pure air from the second cylinder. There was also
advanced, in the concluding part of the communication, a sentence
or two on the cause of the first inspiration, which is well worthy of
note. The cause of the first inspiration, he explained, is probably the
same as the second or the last, viz., a sensation or impression arising
from a want of oxygen in the system. So long as the placenta
performs its functions, the fœtus is perfectly at ease, and feels no
need of respiration; but whenever this communication between the
child and its mother is interrupted, at least in the later months of
pregnancy, the child makes convulsive efforts at respiration similar
to those made by a drowning animal.
On December the 18th, 1841, Mr. Snow was again before the
“Westminster Medical” with a very ingenious instrument which he
had invented for performing the operation of paracentesis of the
thorax. The description of the instrument will be found in the
Medical Gazette of January 28th, 1842.
In the Medical Gazette for November 11th, 1842, Mr. Snow
published a note on a new mode for securing the removal of the
placenta in cases of retention with hæmorrhage; and in the same
journal for March 3rd, 1843, he communicated an essay on the
circulation in the capillary vessels. The essay was selected and
rearranged from papers read before the “Westminster Medical” on
January 21 and February the 4th. We have in this essay an admirable
sketch of the capillary circulation. He advanced, on this occasion, the
idea that the force of the heart is not alone sufficient to carry on the
circulation, but that there is a force generated in the capillary system
which assists the motion. He explained also the great importance of
the cutaneous exhalation, and reasoned that in febrile states,
accompanied with hot skin, the transpiration from the skin is in
reality greater than is normal, and that the good effect of poultices
and similar applications to inflamed skin is due to their influence in
checking the transpiration from the affected part.
But what of practice during all this work at the purer science of
medicine? The story to be told is an old one. Practice did not come, at
least not from the wealthy. He had plenty of practice in so far as
seeing patients was concerned certainly, for he was encumbered with
four sick clubs; and his club practice, together with the out-patient
work at the Charing Cross Hospital, kept the bell ringing all day, and
not unfrequently enlivened the night with the clamorous music. But
the patients with the fees in their hands kept at a respectful distance.
Why? The answer gives another old story—because the practitioner
at 54, Frith Street, Soho, was an earnest man, with not the least
element of quackery in all his composition, with a retiring manner
and a solid scepticism in relation to that routine malpractice which
the people love. I have heard many reasons alleged for the want of
success which attended Mr. Snow’s first labours as a claimant on the
public confidence. These reasons have all had one reading, in that
they refer to every cause but the true one. The true cause was, that a
young man having no personal introduction to the bedsides of
dowagers of the pillmania dynasty, sought to establish his fame on
the basis of a sound and rational medicine—because impressed with
the knowledge of the external origin of disease, he went in for the
removal of external causes, and studied nature in preference to the
Pharmacopœia.
Pushing on in the higher branches of his profession, and aiming
always at the best, the degree of the University of London became a
temptation, and Mr. became Dr. Snow on the 23rd of November,
1843, by passing the M.B. examination. He was enrolled in the
second division on this occasion. On the 20th of December in the
following year, he passed the M.D. examination, and came out in the
first division of candidates.
The harass of London life by this time commenced to tell on Dr.
Snow. He had suffered a few years previously from threatened
symptoms of phthisis pulmonalis, but took plenty of fresh air, and
recovered. He again became slowly unhinged for work, and in the
summer of 1845, was attacked with acute and alarming symptoms of
renal disorder. His friend and neighbour, Mr. Peter Marshall, then of
Greek Street, now of Bedford Square, gave him his able assistance,
and the advice of Dr. Prout, and, I believe, of Dr. Bright, was
obtained. He was induced by their general opinion to change his
mode of living, and even to take wine in small quantities. In the
autumn of 1845, he paid a visit to his friend and old colleague, Mr.
Joshua Parsons, at Beckington, with whom he stayed a fortnight,
enjoying himself very much. The friends resumed their old
controversies, and the Doctor admitted that he had been obliged to
relinquish his vegetable diet in favour of a mixed regimen. He
improved greatly, says Mr. Parsons, during his stay; but it was
obvious that London life and hard study had hold of him. From
Beckington he went to the Isle of Wight, but soon returned to
London and to his work. A little after this, he was elected Lecturer on
Forensic Medicine at the Aldersgate School of Medicine, and held the
appointment till the establishment dissolved in 1849. I have often
heard from him, in his quiet droll way, many laughable stories in
relation to his duties in the forensic chair. When he left off teaching,
he found that, in addition to the labour implied and the cost of
experiments, he had to pay, with the rest of his colleagues, a ransom
for his release.

There is no night without its morning. The eventful medical year of

1846 proved the turn of tide season with our struggling Esculapian.
In this year, the news came over from America that operations could
be performed without pain under the influence of sulphuric ether.
The fact was just such an one as would at once attract the earnest
attention of Dr. Snow. It was a physiological, as well as a practical
fact. It was rational in its meaning, and marvellously humane in its
application. The question once before him, was in a scientific sense
his own. His previous experimental studies on respiration and
asphyxia had prepared him for this new inquiry. He lost no time,
therefore, in investigating the new fact; he took it up for its own sake,
however, not from any thought, at the time, of a harvest of gold.
The first inhalations of ether in this country were not so successful
as to astonish all the surgeons, or to recommend etherization as a
common practice. The distrust arose from the manner in which the
agent was administered. Dr. Snow at once detected this
circumstance; and, as he explains in the pages of the work now in the
hands of the reader, remedied the mistake by making an improved
inhaler. He next carried out many experiments on animals and on
himself, and brought the administration to great perfection. One day,
on coming out of one of the hospitals (I am giving the narrative as he
gave it to me), he met Mr. —— (a druggist whom he knew) bustling
along with a large ether apparatus under his arm. “Good morning!”
said Dr. Snow. “Good morning to you, doctor!” said the friend; “but,
don’t detain me, I am giving ether here and there and everywhere,
and am getting quite into an ether practice. Good morning, doctor!”
“Good morning to you!” Rather peculiar! said the doctor to himself;
rather peculiar, certainly! for the man has not the remotest chemical
or physiological idea on the subject. An “ether practice! If he can get
an ether practice, perchance some scraps of the same thing might fall
to a scientific unfortunate.” Consequently, with his improved inhaler,
Dr. Snow lost no time in asking to be allowed to give ether at St.
George’s Hospital. He got permission to give it there to the out-
patients, in cases of tooth-drawing. Dr. Fuller, of Manchester-square,
standing by, was surprised to see with what happy effects ether was
administered when administered properly. A day or two afterwards,
an operation having to be performed, and the surgeon (I believe, Mr.
Cutler) not approving of the ether in the way in which it had
previously acted, Dr. Fuller remarked on the superiority of Dr.
Snow’s mode of administering it; and the result was, that he was
asked to give it on operating days. He did so with great success. He
administered it at University College with the same success. Liston,
then the leading operator, struck with the new man who came before
him in such an able and unaffected way, took him by the hand; and
from that time the ether practice in London came almost exclusively
to him. Science for once put assumption in its right place.
The new field once open, it were impossible but that he should
cultivate it diligently. The Westminster Medical Society was often
favoured with his communications and experiments on etherization;
and in the September of 1847, he embodied, in his first work, the
whole of his experience up to that time. The work was remarkable for
the care with which it was written, the science which it displayed,
and the complete mastery of the subject which it everywhere
conveyed.
What had been a mere accidental discovery, I had almost said a
lucky adventure, was turned by the touch of the master into a
veritable science. The book was readily appreciated by the
profession, and was just beginning to sell, when the discovery of the
application of chloroform threw ether into the shade and the book
with it.
Dr. Snow, though a man of great firmness when once his mind was
made up, was always ready for new inquiry. Chloroform, therefore,
was no sooner brought before the profession by Dr. Simpson, than he
began to institute a series of independent researches, and having
satisfied himself personally as to the effects and greater practicability
of chloroform, he at once commenced its use, and forgot sooner
almost than others all predilections for ether. In 1848, he
commenced a series of experimental papers on narcotic vapours in
the Medical Gazette, and continued them until 1851, when the
Medical Gazette virtually ceased to exist. The papers on narcotics, in
accordance with his other and earlier productions, were stamped
with the evidences of profound and careful research, and still more
careful deduction. I infer that they have been more talked about than
read, for few people seem to be aware of the enlarged and positive
physiological arguments which they contain. Chloroform and ether
are not alone discussed, but all narcotics. Narcotics are not alone
considered, but various of the great functions of life. The records of a
vast number and variety of experiments are here related, and an
amount of information, original in kind, collected, which will always
remain as a memorable record in the history of medical literature.
But the great points in these papers are those in which the author
enters on the physiological action of narcotics. Here appear the
generalizations and insights into the relations of allied phenomena
which mark the man of true power. His greatest deduction on these
matters, and the proofs on which it is based, are to be found in his
observations, where he explains that the action of the volatile
narcotics is that of arresting or limiting those combinations between
the oxygen of the arterial blood and the tissues of the body, which are
essential to sensation, volition, and all the animal functions. He
demonstrated that these substances modify and, in large quantities,
arrest the animal functions in the same way, and by the same power
as that by which they modify and arrest combustion, the slow
oxidation of phosphorus and other kinds of oxidation unconnected
with the living body when they (the narcotics) are mixed with the
atmospheric air.
In his modest way, he often spoke to me, with honest pride, on this
observation. He himself thought it the best observation he had ever
made, and believed that it would not be lost as an historical truth.
Placing a taper, during one of our experiments, in a bottle through
which chloroform vapour was diffused, and watching the declining
flame, he once said, “There, now, is all that occurs in narcotism; but
to submit the candle to the action of the narcotic without
extinguishing it altogether, you must neither expose it to much
vapour at once, nor subject it to the vapour too long; and this is all
you can provide against in subjecting a man to the same influence. I
could illustrate all the meaning of this great practical discovery of
narcotism on a farthing candle, but I fear the experiment would be
thought rather too commonplace.”
The year of the world’s fair in London, 1851, may be considered a
fortunate one for Dr. Snow. His affairs had taken a new turn, and the
tide was fairly in his favour. He had a positive holiday, physical and
mental. The harass of the professional struggle was over, the world
was opening its eyes to his intrinsic merits; old friends flocked
around him, brought to the grand show in town, and all was well. He
did but little this year, except to write a characteristic letter to Lord
Campbell, who was pushing on a bill in the House of Lords, called
the “Prevention of Offences Bill,” in which a clause was introduced to
prevent, by severe punishment, any attempt that might be made by
any person to administer chloroform or other stupifying drug for
unlawful purposes. Dr. Snow, believing that Lord Campbell was
actuated in introducing this clause by the fact of certain trials having
recently occurred for the offence of using chloroform unlawfully, and
being himself convinced that, in two of the cases (the one the case of
a robbery in Thrale-street; the other, of a robbery attempted on
London Bridge), the evidence against the prisoners, of attempting to
produce insensibility by chloroform, was without any reason or
possibility, he opposed the afore named clause in the bill on the
ground that if it became law numerous frivolous and false charges
would be constantly brought up against innocent people, or against
guilty persons, but persons not guilty of the special charge laid
against them, that, namely, of administering a volatile narcotic by
inhalation. Knowing that weakness of human nature which leads a
man, in the presence of all evidence, never to admit intoxication as
possible in his own proper person, Dr. Snow felt that, in any case
where an intoxicated person had been robbed, such person might
allege that he had been made insensible by narcotic vapour. The two
cases specially noticed in his letter admitted readily of such
interpretation, and were clearly not cases in which chloroform had
been administered. Lord Campbell, on the receipt of Dr. Snow’s
letter, referred to it in very complimentary terms in the Lords’, but
intimated that the reasoning of the letter did not alter his
determination. The editor of the Medical Gazette, Dr. Alfred Taylor,
opened fire on Dr. Snow; and for two or three weeks a sharp contest
occurred between the two doctors; but the matter soon rested, each
author retaining his own opinions, and both agreeing to differ.
Dr. Snow’s amiable but firm nature led him often to this
ultimatum. Freedom of expression was a right he always claimed;
but for this reason he extended the same privilege to others. He was
never stirred into provocation by any difference of opinion. It was
enough for him to form carefully his own opinions, and then to hold
to what he had said, so long as he felt, from his internal convictions,
that he was right.
In the year 1848, Dr. Snow, in the midst of his other occupations,
turned his thoughts to the questions of the cause and propagation of
cholera. He argued in his own mind that the poison of cholera must
be a poison acting on the alimentary canal by being brought into
direct contact with the alimentary mucous surface, and not by the
inhalation of any effluvium. In all known diseases, so he reasoned, in
which the blood is poisoned in the first instance, there are developed
certain general symptoms, such as rigors, headache, and quickened
pulse; and these symptoms all precede any local demonstration of
disease. But in cholera this rule is broken; the symptoms are
primarily seated in the alimentary canal, and all the after symptoms
of a general kind are the results of the flux from the canal. His
inference from this was, that the poison of cholera is taken direct
into the canal by the mouth. This view led him to consider the
mediums through which the poison is conveyed, and the nature of
the poison itself. Several circumstances lent their aid in referring him
to water as the chief, though not the only, medium, and to the
excreted matters from the patient already stricken with cholera, as
the poison. He first broached these ideas to Drs. Garrod and Parkes,
early in 1848; but feeling that his data were not sufficiently clear, he
waited for several months, and having in 1849 obtained more
reliable data, he published his views in extenso in a pamphlet
entitled “The Mode of Communication of Cholera”. During
subsequent years, but specially during the great epidemic outbreak
of the disease in London in 1854, intent to follow out his grand idea,
he went systematically to his work. He laboured personally with
untiring zeal. No one but those who knew him intimately can
conceive how he laboured, at what cost, and at what risk. Wherever
cholera was visitant, there was he in the midst. For the time, he laid
aside as much as possible the emoluments of practice; and when
even, by early rising and late taking rest, he found that all that might
be learned was not, from the physical labour implied, within the
grasp of one man, he paid for qualified labour. The result of his
endeavours, in so far as scientific satisfaction is a realization, was
truly realized, in the discovery of the statistical fact, that of 286 fatal
attacks of cholera, in 1854, occurring in the south districts of the
metropolis, where one water company, the Southwark and Vauxhall,
supplied water charged with the London fæcal impurities, and
another company, the Lambeth, supplied a pure water, the
proportion of fatal cases to each 10,000 houses supplied by these
waters, was to the Southwark and Vauxhall Company’s water 71, to
the Lambeth 5.
There was, however, another fact during this epidemic, which
more than the rest drew attention to Dr. Snow’s labours and
deductions. In the latter part of August 1854, a terrific outbreak of
cholera commenced in and about the neighbourhood of Broad-street,
Golden-square. Within two hundred and fifty yards of the spot where
Cambridge-street joins Broad-street, there were upwards of five
hundred fatal attacks of cholera in ten days. To investigate this
fearful epidemic was at once the selfimposed task of Dr. Snow. On
the evening of Thursday, the 7th of September, the vestrymen of St.
James’s were sitting in solemn consultation on the causes of the
visitation. They might well be solemn, for such a panic possibly never
existed in London since the days of the great plague. People fled from
their homes as from instant death, leaving behind them, in their
haste, all the mere matter which before they valued most. While,
then, the vestrymen were in solemn deliberation, they were called to
consider a new suggestion. A stranger had asked, in modest speech,
for a brief hearing. Dr. Snow, the stranger in question, was admitted,
and in few words explained his view of the “head and front of the
offending”. He had fixed his attention on the Broad-street pump as
the source and centre of the calamity. He advised the removal of the
pump-handle as the grand prescription. The vestry was incredulous,
but had the good sense to carry out the advice. The pump-handle was
removed, and the plague was stayed. There arose hereupon much
discussion amongst the learned, much sneering and jeering even; for
the pump-handle removal was a fact too great for the abstruse
science men who wanted to discover the cause of a great natural
phenomenon in some overwhelming scientific problem. But it
matters little. Men with great thoughts in their heads, think of little
things which little men cover with their wide-spread feet. It matters
little, for the plague was stayed; and whoever will now read
dispassionately the report of a committee, afterwards published by
the vestry, and the demonstrative evidence of the Rev. Mr.
Whitehead, will find that the labours and suggestion of Dr. Snow, in
reference to the Broad-street epidemic of cholera, must become each
day better and better appreciated, as time, which never yet told a lie,
tells the tale and points the moral of the event which is here so
imperfectly described. Some who, at first, were amongst those who
held up the labours of our friend to ridicule, or passed them over in
contemptuous silence, have, indeed, since modified their opinions,
and have either tacitly accepted his facts, or have done far worse by
attempting to put them forward as though they were the work of no
single man, or of some one unknown, or as though their connection
with a theory destroyed the originality of the facts themselves. It was
my privilege, during the life of Dr. Snow, to stand on his side. It is
now my duty, in his death, as a biographer who feels that his work
will not be lost, to claim for him not only the entire originality of the
theory of the communication of cholera by the direct introduction of
the excreted cholera poison into the alimentary system; but,
independently of that theory, the entire originality of the discovery of
a connection between impure water supply and choleraic disease.
The whole of his inquiries in regard to cholera were published in
1855, in the second edition of his work on the “Mode of
Communication of Cholera”—a work in the preparation and
publication of which he spent more than £200 in hard cash, and
realized in return scarcely so many shillings.
In 1856, he made a visit to Paris in company with his uncle, Mr.
Empson, who having personally known the present Emperor many
years, had on this occasion special imperial favours shown to him, in
which the nephew participated. During the visit, Dr. Snow lodged a
copy of his work on Cholera at the “Institute”, in competition for the
prize of £1,200 offered for the discovery of a means for preventing or
curing the disease. The decision of the judges has since been
published, but no note seems to have been made of Dr. Snow’s
researches.
The Medical Society of London, reformed under that name in
1849–50, by amalgamation with the Westminster Medical, was at
this time the principal scene of Dr. Snow’s scientific exertions. In
1852, the Society elected him as Orator for the ensuing year; and at
the eightieth anniversary of the Society, held on March the 8th, at
the Thatched House Tavern, he delivered an admirable oration on
“Continuous Molecular Changes, more particularly in their Relation
to Epidemic Diseases.” He made no claim to the orator’s gown; but
the address was too forcible and first class not to call forth the
enthusiasm of the audience. It was admirably received; and few of us
who were present on that interesting occasion will forget the simple
and genuine earnestness of our beloved associate, as in the twinkling
twilight he carried us along with the smooth current of his thoughts.
He spent nearly twelve months in the preparation of this oration. It
was intended to convey, in the most pleasing manner at his
command, a broad view of his observations on the communication of
certain spreading diseases. He advanced, on this occasion, the idea
that intermittent fever, and perhaps yellow fever, are, like cholera,
carried by their poisons direct into the alimentary system.
Two years after this event, having, meantime, passed the office of
vice-president, the Society elected him to the highest honour it can
confer,—to the presidential chair. He took his place as President, in
his unassuming manner, on March 10th, 1855, delivering a short but
pleasing address. Throughout the year he carried out the duties of his
office with great success. One of his presidential acts was peculiarly
graceful. One evening, while presiding, Dr. Clutterbuck (then the
father, or oldest member of the Society) came into the meeting. The
venerable and distinguished old man, then long past his eightieth
year, had lately been a stranger to the assembly, and was known but
to few of the members. The President, as Dr. Clutterbuck entered the
room, himself rose, and in a way that was irresistible in its simple
courtesy resigned his chair to the veteran Esculapian. “It is near fifty
years,” said Dr. Clutterbuck with emotion, as he took the proffered
seat, “since I last occupied this honourable position.” At the next
anniversary meeting, held on March the 8th, 1856, Dr. Clutterbuck
came to his last meeting, and to see (so the fates willed it) his friend
the President play also his last part in presidential duties. At the
anniversary dinner on that same day, the President reviewed, in
feeling terms, his own career in the professional strife, and expressed
that his success in life had originated in his acquaintance with the
Society over which he then governed by the general will.
In addition to the fellowship of the Medical Society, Dr. Snow
belonged to the Royal Medical and Chirurgical, Pathological, and
Epidemiological societies. He was also a member of the British
Medical Association. The Medical Society, from its old associations,
was, however, that in which he took the most active part. Next to
this, the Epidemiological Society claimed his regard. When Mr.
Tucker first contemplated the formation of the Epidemiological
Society, Dr. Snow was one of the first with whom he held
consultation, and from whom he received that able support which
enabled him to found that excellent institution. From the first of the
Society, Dr. Snow was an active member. He was on many of its
committees; he was a member of council, and a frequent contributor
to its Transactions. He used often to meet with opponents to his
peculiar opinions at the meetings of this Society, but he always
retained friendships.
The position which he took as an epidemiologist was original, and
in opposition to the views of many eminent men who had in the
matters relating to public health considerable influence, scientific
and political. He could not consequently, and did not, expect to go on
his way unopposed. But he did sometimes expect a more deliberate
and considerate attention to his hard wrought labours than he
received or deserved. He used constantly, though no great professor
of Shakespearian lore, to deplore the long admitted fact, that nothing
so inevitably tends to transform an earnest inquiring and
enthusiastic man, into a supercilious, superficial, and cold-hearted
egotist, as translation from the stool of self-reliance and
independence, into the gilded chair of office and brief authority.
It must be admitted that Dr. Snow’s views on the spread of
epidemics were extreme in character; but from the slight which they
too hastily received, they were not, I believe, properly understood. It
has often been said that he encouraged by his arguments the
perpetuation of certain offensive arts and occupations which are
injurious to the public health; and in 1855, several journalists
commented on him severely for this supposed error. But the fact is,
he never presumed that any man could breathe with impunity other
gaseous mixture than oxygen and nitrogen in atmospheric
proportion. He knew too well the effect of inhaling chemical
substances to allow of such supposition to enter his mind. But he
contended, in regard to pure epidemic disorders, distinguished by
specific symptoms, that these have a specific poison, which is
propagated by certain fixed laws, which attains its progression and
increase in and through animal bodies; which is communicated from
one animal body to another, and which is the same in its essence
from first to last. This was his position, and he adhered to it. No
mere emanation arising from evolution of foul smelling gas can, per
se, according to his views, originate a specific disease, such as small-
pox or scarlet-fever; as well expect that the evolution of such gas
should plant a plain with oaks or a garden with crocuses. True,
small-pox may occur over a cesspool as an oak may spring up from a
manure heap; but the small-pox would never appear over the
cesspool in the absence of its specific poison; nor the oak rise from
the manure heap in the absence of the acorn which seeded it.
In 1855, Dr. Snow gave evidence before the select committee on
the “Public Health and Nuisances Removal Bill,” in which evidence
he strove to convey the impressions which are condensed above.
Feeling that he had not been correctly understood, he afterwards
wrote a letter to Sir Benjamin Hall, in which he set forth the whole of
his argument very distinctly and sensibly. He indicated in this letter
that he was no defender of nuisances, but that whereas a bad smell
cannot simply because it is a bad smell give rise to specific disease, so
an offensive business conducted in a place where it ought not be
should be proceeded against by ordinary law as a nuisance, without
using in regard to it the word pestiferous, or otherwise dragging in
and distorting the science of medicine. As time rolls on, it will
probably be elicited that the groundwork of Dr. Snow’s theory is
sound. That if he committed error, it was in adhering too closely to
the abstract fact, and in not allowing sufficient importance to the
favouring influence of impure conditions in the propagation and
distribution of the specific poisons of the specific diseases.
At all events, the view he had maintained originally, he maintained
to the end, and throughout conscientiously; and the aspersions that
the object of his argument was to support his special theory
regarding the communication of cholera, are utterly unfounded. In
the present year, 1858, he read at the Epidemiological Society, and
published in the Medical Times and Gazette, a repetition of his
previous opinions, strengthening them by a statistical record,
showing that the mortality of persons working at so-called offensive
occupations is at certain ages lower, and at certain ages slightly
higher, than in the general population. When the paper was read at
the Epidemiological Society, Mr. Edwin Chadwick made a long series
of objections to the paper, and complained that the argument was
illogical. It was so, doubtless, on Mr. Chadwick’s premises; but on
the premises advanced by Dr. Snow, as to the specific propagation of
specific diseases by specific poisons, physiological problems on
which, from his experimental researches and knowledge, he was far
the best authority, his arguments were perfectly logical, and perfectly
consistent.
In relation to public health, Dr. Snow contributed many other
observations. In the first number of the Sanitary Review, he
communicated a valuable paper, previously read at the
Epidemiological Society, on the “Comparative Mortality of Town and
Rural Districts”; and, previous to his decease, he was busily occupied
in investigating the question of adulteration of bread with alum. He
made several analyses of different specimens of bread, but his papers
merely leave a brief record of the fact, without any comments or
results.
We return for a few moments to some further points connected
with his researches on inhalation. In addition to his experiments
with volatile narcotics, he carried out for a long time a series of
inquiries with other medicinal substances, and administered many
remedies by inhalation at the Brompton Hospital, during a period of
twenty months. In 1851, he recorded the result of this experience at
the Medical Society of London, and explained the modes of
administering various agents. Some, as morphia and stramonium,
were inhaled with the aid of heat; others, as hydrocyanic acid and
conia, were inhaled at the ordinary temperature. The particulars of
these experiments will be found in a short paper in the London
Journal of Medicine for January 1851.
He continued steadily to investigate the effects of various volatile
agents for the production of insensibility, and arrived by frequent
experiment to such a degree of positive knowledge regarding agents
of this class, that the composition and boiling point of any new
chemical body having been supplied, he could predict whether or not
its vapour would produce narcotism by inhalation. Other than the
volatile narcotics referred to in his present essay, he performed a
variety of experiments with carbonic acid, carbonic oxide, cyanogen,
hydrocyanic acid, Dutch liquid, ammonia, nitrogen, amylovinic
ether, puff-ball smoke, allyle, cyanide of ethyle, chloride of amyle, a
carbo-hydrogen from Rangoon tar, a carbo-hydrogen coming over
with amylene, and various combinations of these. His grand search
was for a narcotic vapour which, having the physical properties and
practicability of chloroform, should, in its physiological effects,
resemble ether in not producing, by any accident of administration,
paralysis of the heart. The fact that in almost every fatal case from
chloroform the result had occurred from the action of the narcotic on
the central organ of the circulation, was never absent from his
thoughts. An agent having this effect, however intrinsically valuable,
was not to be put in the hands of every person for administration.
“There would be a great uproar,” he remarked on one occasion, “if a
student were to undertake on the operating table to tie the femoral
artery, and were to open the femoral vein. Yet at some of our
hospitals, the administration of chloroform has been entrusted to the
porter, who would only grin in ignorance, if informed that each time
his services were required, he performed the grand act of suspending
for a time the oxidation of the whole body, and of inducing a
temporary death; and who would tell you, if you asked him the
composition of chloroform, that it was smelling stuff.” He spoke this
from no selfish feeling, but with that kind of regret which an
educated engineer would feel, on referring to the fact of a railway
porter who, knowing nothing of steam, how to put it on, when to take
it off, or why it propelled, had mounted an engine and driven a host
of confiding passengers to their destruction. This is the way in which
he expressed himself, and it would be difficult to show that he was
not correct.
Intent on the discovery of some new anæsthetic, which might be
more safely entrusted to general use, Dr. Snow began, in 1856, to
experiment with amylene. As usual, he went to work cautiously and
with precision. First he ascertained the boiling point of the specimen
supplied to him; then the point of saturation of air with the vapour at
different temperatures; then the effects of inhalation of the vapour
by inferior animals, and the quantity required to be inspired, with
the air breathed, to produce insensibility. These were the usual steps
in all his inquiries of this kind. When he had obtained any substance
which would produce insensibility favourably on animals, he pushed
it, in one or two experiments, to its extreme in animals of different
kinds; and having produced death by the inhalation, both by giving
rapidly a large dose, and by giving a small dose for a long period, he
observed the mode of death, whether it occurred by cessation of the
heart, or by cessation of the respiration primarily. If the agent
seemed to promise favourably from these inquiries, he commenced
to try it on man; and the first man was invariably his own self. His
friends, knowing his unflinching courage in the ardour of his
inquiries, often expostulated with him in regard to the risks he ran. It
was of no avail. He felt the personal trial a duty, and he did it. I do
not believe, as some have supposed, that these personal experiments
had any effect in producing his early death; but it is certain that he
underwent many risks in the performance of his investigations, and
that he held his own life of least consideration when the lives or well-
being of others were under consideration.
There is yet another trait in his character which I cannot but
notice, and which I would respectfully commend to all physiological
inquirers. While he held it as a necessity to use inferior animals for
the purpose of experiment, he never touched living thing with the
physiologist’s finger without having before him some definite object;
and never performed experiment on any animal without providing
with scrupulous care against the infliction of all unnecessary
suffering. The interests of humanity were, according to his rule, best
advanced by the practice of a humanity that was universal.
He paid considerable attention to the subject of local anæsthesia,
and tried numerous methods for attaining to a knowledge of a
perfect local anæsthetic. He performed experiments with freezing
mixtures, with chloroform; and for the production of rapid and
efficient benumbing by cold, he tried, in 1854, the effects of applying
solid carbonic acid to the skin. At one of the meetings of the Medical
Society, he reported at length the results he had arrived at; but he
was never satisfied with them, and soon relinquished the inquiry, in
order to concentrate his energies on the discovery of what he felt
sure must be discovered ultimately,—an anæsthetic which might be
inhaled with absolute safety, and which would destroy common
sensation without destroying consciousness.
To some extent he succeeded in this latter direction, in his
discovery of the physiological effects of amylene; and for some time
he was sanguine as to the great safety of the new agent. But the
deaths which he has so faithfully recorded as occurring in his own
hands from amylene, removed his expectations, and he discontinued
its use as soon as he learned the risks which might follow its
administration.
By his earnest labours Dr. Snow soon acquired a professional
reputation, in relation to his knowledge of the action of anæsthetics,
which spread far and wide; and the people, through the profession,
looked up to him from all ranks, as the guide to whom to entrust
themselves in “Lethe’s walk”. On April the 7th, 1853, he administered
chloroform to Her Majesty at the birth of the Prince Leopold. A note
in his diary records the event. The inhalation lasted fifty-three
minutes. The chloroform was given on a handkerchief, in fifteen
minim doses; and the Queen expressed herself as greatly relieved by
the administration. He had previously been consulted on the
occasion of the birth of Prince Arthur, in 1850, but had not been
called in to render his services. Previous to the birth of Prince
Leopold, he had been honoured with an interview with His Royal
Highness the Prince Albert, and returned much overjoyed with the
Prince’s kindness and great intelligence on the scientific points
which had formed the subject of their conversation. On April 14th,
1857, another note in the diary records the fact of the second
administration of chloroform to Her Majesty, at the birth of the
Princess Beatrice. The chloroform again exerted its beneficent
influence; and Her Majesty once more expressed herself as much
satisfied with the result.
Inquisitive folk often overburthened Snow, after these events, with
a multitude of questions of an unmeaning kind. He answered them
all with goodnatured reserve. “Her Majesty is a model patient,” was
his usual reply: a reply which, he once told me, seemed to answer
every purpose, and was very true. One lady of an inquiring mind, to
whom he was administering chloroform, got very loquacious during
the period of excitement, and declared she would inhale no more of
the vapour unless she were told what the Queen said, word for word,
when she was taking it. Her Majesty, replied the dry doctor, asked no
questions until she had breathed very much longer than you have;
and if you will only go on in loyal imitation, I will tell you everything.
The patient could not but follow the example held out to her. In a few
seconds she forget all about Queen, Lords, and Commons; and when
the time came for a renewal of hostilities, found that her clever
witness had gone home to his dinner, leaving her with the thirst for
knowledge still on her tongue.

From the literary and general history of Dr. Snow, let us turn for a
few pages to his history personal. I will take the mean of the last
eight years of his life,—the period in which I knew him,—as the
period from which to draw particulars. He was of middle size, and,
some years since, slender; but of late he had become of slightly fuller
build. His long life in comparative student loneliness had made him
reserved in manner to strangers; but with private friends he was
always open, and of sweet companionship. With his increased
popularity he became less reserved to strangers; and within the last
few years he so far threw off restraint as to visit the opera
occasionally. But he moderated every enjoyment, and let nothing
personal stand in the way of his scientific pursuits. He was the
impersonation of order. He had his time and place for everything; he
kept a diary, in which he recorded the particulars of every case in
which he administered chloroform or other anæsthetic, with
comments on the results of the administration, and hints as to
dangers avoided or chanced. He kept a record of all his experiments,
and short notes of observations made by his friends. He rose early,
and retired early to rest,—at eleven o’clock. He seemed, whenever he
was waited on, as though he had nothing in hand, and were always
open to an engagement.
Anything and everything of scientific interest arrested his
attention, and his kindliness of heart was at all times in the
foreground. When I was living at Mortlake, he would run down, on
request, after his day’s duties were over, to a post-mortem, to see a
poor patient, or to take part in an experiment, returning as cheerily
as though he had been to receive the heaviest fee. I name this as but
one example of his kindly nature; there are many who could
corroborate the example in like personal manner.
He laid no claim to eloquence, nor had he that gift. A peculiar
huskiness of voice, indeed, rendered first hearings from him painful;
but this was soon felt less on acquaintance, and the ear once
accustomed to the peculiarity, the mind was quickly interested in the
matter of his discourse, for he always spoke earnestly, clearly, and to
the point. In the societies he spoke very often, and gave expression to
views, on which he had spent great thought, with a generous freedom
which, in so far as the fame of his originality was concerned, had
been better held in reserve. It had been better, that is to say, for him
to have carefully elaborated some of his views in the closet, and
published them fully, than to have sent them forth in the hurry of
debate. Had he lived, he would possibly have collected many stray
labours thus put forward, and have given to them the matured
consideration which they deserved. One of his views, on which he
would have bestowed great attention, refers to the origin of various
morbid growths, as cancer. He believed that these morbid formations
are all of local origin; that they arise in the parts of the body where
they are found, from some perversion of nutrition; and that the
constitutional effects are secondary to and dependent on the local
disorder. He had made many observations on this important subject,
notices of which are to be found scattered, here and there, in the
proceedings of the Medical Society of London, but no connected
record was ever completed.
His private conversation was both instructive and amusing; he was
full of humorous anecdotes, which he told in a quiet, and irresistibly
droll style; and when he laughed, his goodnatured face laughed in
every feature.[1] His anecdotes were never given in set form, but were
elicited by some circumstance or other which might happen to
suggest them. Once, when a friend of ours related at dinner some of
the economical measures of an odd old doctor who was known to
some of the company, he gave us an anecdote, showing how a man
may work too hard for his money. “When I was a very young man,”
he said, “I went for a brief period to assist a gentleman who had a
large parochial practice. I found his surgery in a very disorderly state,
and thinking on my first day with him that I would enhance myself in
his opinion by my industry, I set to work, as soon as his back was
turned, to cleanse the Augean stable. I took off my coat, cleared out
every drawer, relieved the counter of its unnecessary covering,
relabelled the bottles, and got everything as clean as a new pin. When
the doctor returned, he was quite taken by storm with the change,
and commenced to prescribe in his day book. There was a patient
who required a blister, and the worthy doctor, to make dispensing
short, put his hand into a drawer to produce one. To his horror, the
drawer was cleansed. Goodness! cried he, why where are all the
blisters? The blisters, I replied, the blisters in that drawer? I burnt
them all; they were old ones. Nay, my good fellow, was the answer,
that is, the most extravagant act I ever heard of; such proceedings
would ruin a parish doctor. Why, I make all my parochial people
return their blisters when they have done with them. One good
blister is enough for at least half a dozen patients. You must never do
such a thing again, indeed you must not. I did not, for he and I soon
found a good many miles of ground between us, though we never had
any more serious misunderstanding.”
His replies, when under the fire of cross-question, were ready and
commonsense. Once, he observed that in his opinion sulphuric ether
was a safer narcotic than chloroform. Why, then, said a listener, do
you not use ether? I use chloroform, he resumed, for the same reason
that you use phosphorus matches instead of the tinder box. An
occasional risk never stands in the way of ready applicability. On
another occasion, after one of the meetings of the “Medical Society,”
when the subject of a specific cholera cell had been under debate,
some one asked him, as a poser and rather ironically, where he
thought the first cholera cell came from? “Exactly,” he replied, with a
droll face. “But to begin, do you tell me where the first tiger or the
first upas tree came from; nay, tell me where you came from yourself,
and I will then tell you the origin of the first cholera cell, and give you
the full history of the first case; but I want a model before I venture
on the description of ultimate facts.”
As an author, his style was plain, clear, and smoothly elegant. His
argument was always carefully studied and as carefully rendered. He
sent manuscript to the printer which required scarcely a letter of
correction. Both in writing and speaking, he made the expression of
truth his first business. Neither provocation nor temptation could
ever lead him aside from that principle. His readings were select. He
chiefly read scientific works, old and new. He had great relish for
some of the old medical writers—the masters in physic. He had read
Bacon, but agreed with Harvey’s criticism that Bacon wrote science
like a lord chancellor. He had a notion that there had been a history
long previous to any we know of from existing records, in which the
sciences generally had risen to a greater perfection than they are at
this present. His conversance with Sprengel’s History of Medicine
had possibly led him to this opinion. He was fond of general history
also, but studied it little. He never read novels, because the hours
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