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Seafoods: Chemistry, Processing Technology
and Quality
Seafoods: Chemistry, Processing
Technology and Quality
Edited by
FEREIDOON SHAHIDI
Departments of Biochemistry and Chemistry
Memorial University of Newfoundland
and
J. RICHARD BOTTA
Inspection Branch
Canada Department of Fisheries and Oceans
SPRINGER-SCIENCE+BUSINESS MEDIA, B.Y.

First edition 1994
© 1994 Springer Science+Business Media Dordrecht
Origina11y published by Chapman & Hali in 1994
Softcover reprint of the hardcover 18t edition 1994
Typeset in 1O/12pt Times by Cambrian Typesetters, Frimley, Surrey
ISBN 978-1-4613-5913-5 ISBN 978-1-4615-2181-5 (eBook)

DOI 10.1007/978-1-4615-2181-5
Apart from any fair dealing for the purposes of research or private study, or
criticism or review, as permitted under the UK Copyright Designs and
Patents Act, 1988, this publicat ion may not be reproduced, stored, or
transmitted, in any form or by any means, without the prior permission in
writing of the publishers, or in the case of reprographic reproduction only in
accordance with the terms of the licences issued by the Copyright Licensing
Agency in the UK, or in accordance with the terms of licences issued by the
appropriate Reproduction Rights Organization outside the UK. Enquiries
concern ing reproduction outside the terms stated here shou1d be sent to the
publishers at the Glasgow address printed on this page.
The publisher makes no representation, express or imp1ied, with regard to
the accuracy of the information contained in this book and cannot accept
any legal responsibility or liability for any errors or omissions that may be
made.
A catalogue record for this book is available from the British Library
Library of Congress Catalog Card Number: 94-71014
00 Printed on permanent acid-free text paper, manufactured in accordance

with ANSIINISO Z39.48-1992 (Permanence of Paper).
Preface
Seafoods are important sources of nutrients for humans. Proteins and non-
protein nitrogenous compounds play an important role in the nutritional
value and sensory quality of seafoods. Consumption of fish and marine oils
is also actively encouraged for the prevention and treatment of cardio-
vascular diseases and rheumatoid arthritis. Highly unsaturated long-chain
omega-3 fatty acids are regarded as the active components of marine oils
and seafood lipids. The basic chemical and biochemical properties of
seafood proteins and lipids, in addition to flavour-active components, their
microbiological safety and freshness quality, are important factors to be
considered.
A presentation of the state-of-the-art research results on seafoods with
respect to their chemistry, processing technology and quality in one
volume was made possible by cooperative efforts of an international group
of experts. Following a brief overview, the book is divided into three
sections. In Part 1 (chapters 2 to 8) the chemistry of seafood components
such as proteins, lipids, flavorants (together with their properties and
nutritional significance) is discussed. Part 2 (chapters 9 to 13) describes the
quality of seafoods with respect to their freshness, preservation, micro-
biological safety and sensory attributes. The final section of the book
(chapters 14 to 16) summarizes further processing of raw material,
underutilized species and processing discards for production of value-
added products. The up-to-date information on research and development
presented in this volume, together with its extensive data and references,
provide an invaluable overview of the basic and applied concepts of
seafood research for scientists and technologists. The book may also be
used as a supplementary text for food and flavour chemistry courses for
graduate and advanced undergraduate students.
We are grateful to the department of Fisheries and Oceans, Fisheries
Development Division, Newfoundland Region and the Natural Sciences
and Engineering Research Council of Canada, for financial support
towards the organization of the symposium on seafoods. We wish to extend
our thanks to all authors for their efforts and commendable contributions
which made the publication of the book possible.
Fereidoon Shahidi and J. Richard Botta

Series foreword
The 8th World Congress of Food Science and Technology, held in

Toronto, Canada, in 1991 attracted 1400 delegates representing 76
countries and all five continents. By a special arrangement made by the
organizers, many participants from developing countries were able to
attend. The congress was therefore a most important international
assembly and probably the most representative food science and tech-
nology event in that respect ever held. There were over 400 poster
presentations in the scientific programme and a high degree of excellence
was achieved. As in previous congresses much of the work reported
covered recent research and this will since have been published elsewhere
in the scientific literature.
In addition to presentations by individual researchers, a further major
part of the scientific programme consisted of invited papers, presented as
plenary lectures by some of the leading figures in international food science
and technology. They addressed many of the key food issues of the day
including advances in food science knowledge and its application in food
processing technology. Important aspects of consumer interest and of the
environment in terms of a sustainable food industry were also thoroughly
covered. The role of food science and technology in helping to bring about
progress in the food industries of developing countries was highlighted.
This book is part of a series arising from the congress and including
bibliographical details. The series editors are Professor Marvin Tung of the
Technical University of Nova Scotia, Halifax, Nova Scotia, Canada; and
Dr. Gordon Timbers of Agriculture and Agri-Food Canada, Ottawa,
Ontario, Canada. The book presents some of the most significant ideas
which will carry food science and technology through the nineties and into
the new millennium. It is therefore essential reading for anyone interested
in the subject, including specialists, students, and general readers. IUFoST
is extremely grateful to the organizers from the Canadian Institute of Food
Science and Technology for putting together a first class scientific
programme and we welcome the publication of this book as a permanent
record of the keynote papers presented at the congress.
Dr. D. E. Hood
(President, International Union of Food Science & Technology)
Contributors
R. G. Ackman Canadian Institute of Fisheries Technology, Technical

University of Nova Scotia, Box 1000, Halifax, Nova
Scotia, Canada, B3J 2X4
A. Adams Seafood Products Research Center, US Food and
Drug Administration, 22201 23rd Drive Southeast,
Bothell, Washington, 98041-3012, USA
J. R. Botta Inspection Branch, Canada Department of Fisheries
and Oceans, PO Box 5667, St. John's, Newfoundland,
Canada A1C 5X1
K. Colburn Seafood Products Research Center, US Food and
S.Fuke Laboratory of Food Science, Faculty of Education,
Tokyo Gakugei University, 4-1-1, Nukui-kita,
Koganei-shi, Tokyo, Japan
N. F. Haard Institute of Marine Resources, Department of Food
Science & Technology, University of California,
Davis, CA 95616, USA
w. Hill Seafood Products Research Center, US Food and
H. O. Hultin University of Massachusetts Marine Station, PO Box
7128, Lanesville Station, Gloucester, MA 01930-5828,
USA
J. M. Hungerford US Food and Drug Administration, 22201 23rd
Avenue Southeast, Bothell, Washington, 98041-3012,
USA
J.-M. Kuo Department of Marine Food Science, Fisheries
Science College, National Taiwan Ocean University,
Keelung, Taiwan, Republic of China
C. M. Lee Department of Food Science and Nutrition, University
of Rhode Island, Kingston, RI 02881, USA
V1ll CONTRIBUTORS
R. C. Lindsay Department of Food Science, University of Wisconsin-

Madison, Madison, WI 53706, USA
R. Manger Seafoods Products Research Center, US Food and
B. Sun Pan Department of Marine Food Science, Fisheries
Science College, National Taiwan Ocean University,
Keelung, Taiwan, Republic of China
L. M. Sereda Department of Fisheries and Oceans, Inspection
Branch, Central and Arctic Region, 501 University
Crescent, Winnipeg, Manitoba, Canada R3T 2N6
F. Shahidi Departments of Biochemistry and Chemistry,
Memorial University of Newfoundland, St. lohn's,
Newfoundland, Canada, AlB 3X9
Z. E. Sikorski Department of Food Preservation, Technical
University of Gdansk, 80---952 Gdansk-Wrzeszcz,
Gdansk, Poland
K. E. Spencer Canadian Institute of Fisheries Technology, Depart-
ment of Food Science and Technology, Technical
University of Nova Scotia, PO Box 1000, 1320
Barrington Street, Halifax, Nova Scotia, Canada, B31
2X4
M. A. Tung Canadian Institute of Fisheries Technology, Depart-
ment of Food Science and Technology, Technical
University of Nova Scotia, PO Box 1000, 1360
Barrington Street, Halifax, Nova Scotia, Canada, B31
2X4
M. M. Wekell Seafood Products Research Center, US Food and
R. K. York Department of Fisheries and Oceans, Inspection
Branch, Central and Arctic Region, 501 University
Crescent, Winnipeg, Manitoba, Canada R3T 2N6
Contents
1 The chemistry, processing technology and quality

of seafoods - an overview 1
F. SHAHIDI
Part 1 The chemistry of seafood components
2 Seafood proteins and preparation of protein concentrates 3

F. SHAHIDI
2.1 Introduction 3
2.2 Sarcoplasmic proteins 3
2.3 Myofibrillar proteins 4
2.4 Stroma proteins 5
2.5 Non-protein nitrogenous (NPN) compounds 5
2.6 Surimi 6
2.7 Protein concentrates/hydrolysates 6
References 8
3 Protein hydrolysis in seafoods 10

N. F. HAARD
3.1 Introduction 10
3.1.1 Importance 10
3.1.2 Source of proteinases in seafood II
3.2 Digestive proteinases 14
3.2.1 Gastric proteases 14
3.2.2 Intestinal proteinases 15
3.2.3 Hepatopancreas proteinases 16
3.3 Proteinases contained in the muscle fibre 16
3.3.1 Lysosomal proteinases 16
3.3.2 Alkaline proteinases 20
3.3.3 Neutral proteinases 22
3.4 Proteinases in the extracellular matrix of muscle 24
3.4.1 Collagcnascs 24
3.5 Conclusions 25
Acknowledgements 26
References 26
4 Seafood lipids 34
R. G. ACKMAN
4.1 Introduction: the paradox of seafood in health and nutrition 34

4.2 Seafoods 35
X CONTENTS
4.2.1 Marine fish 35

4.2.2 Freshwater fish 40
4.2.3 Shellfish 41
4.3 Cholesterol in fish and shellfish 43
4.4 Natural variations in quality 45
References 45
5 Oxidation of lipids in seafoods 49

H. O. HULTIN
5.1 Introduction 49
5.2 Cellular catalysts of lipid oxidation 49
5.2.1 Oxygen 50
5.2.2 Transition metals 50
5.2.3 Haem iron 55
5.2.4 Singlet oxygen 56
5.2.5 Enzymes -lipoxygenase and cyclooxygenase 56
5.3 Inhibitors of lipid oxidation 57
5.3.1 Initiation inhibitors 58
5.3.2 Propagation inhibitors 59
5.4 Changes post-mortem and during processing 60
5.4.1 Quantitative changes in pro- and antioxidants in fish muscle
post-mortem 60
5.4.2 Processing induced changes 63
5.5 Approaches to inhibiting lipid oxidation in seafoods 65
5.5.1 Limit oxygen 66
5.5.2 Maintain natural antioxidant systems 66
5.5.3 Minimize increase in pro-oxidants 66
5.5.4 Afford early protection from oxidation 67
5.5.5 Maintain low temperature 67
5.5.6 Protect against NaCI effects 67
5.5.7 Remove unstable lipids 68
5.5.8 Remove dark muscle 68
5.6 Summary and future directions 69
Acknowledgements 71
References 71
6 Flavour of fish 75
R. C. LINDSAY
6.1 Introduction 75
6.2 Very fresh fish flavours 75
6.3 Sea-, brine- or iodine-like flavours 77
6.4 Other characterizing flavours 79
6.5 Deterioration offish flavours 79
References 82
7 Flavour of shellfish and kamaboko flavorants 85

B. SUN PAN and J.-M. KUO
7.1 Introduction 85
7.2 Volatile compounds of shellfish 85
7.2.1 Alcohols 85
CONTENTS XI
7.2.2 Aldehydes 87
7.2.3 Ketones 88
7.2.4 Furans and other oxygen-containing cyclic compounds 89
7.2.5 Pyrazines and other nitrogen-containing compounds 89
7.2.6 Sulphur-containing compounds 94
7.2.7 Hydrocarbons 98
7.2.8 Phenols 98
7.2.9 Esters 99
7.3 Formation pathway of shellfish volatiles 100
7.3.1 Alcoholsandcarbonyls 100
7.3.2 Alkylpyrazines formation via chemical reactions of carbonyl
compounds 103
7.3.3 Sulphur-containing compounds 103
7.4 Role of lipoxygenase and 5,8, II-tetradecatrien-2-one in shellfish flavour 105
7.4. I Occurrence of lipoxygenase in shrimp 105
7.4.2 Effects of lipoxygenase on shrimp flavour formation 106
7.5 Volatile components of kamaboko f1avorants 108
Acknowlegement 110
References 110
8 Taste-active components of seafoods with special reference to

umami substances 115
S. FUKE
8.1 What are umami and umami substances? 115

8.1.1 Introduction 115
8.2 Synergistic taste effect 116
8.2.1 MSG with 5 '-ribonucleotide (IMP or GMP) 116
8.2.2 a-Amino dicarboxylates and L-homocysteinate with IMP 117
8.2.3 Synergistic taste effect among three compounds (tertiary synergism) 117
8.2.4 Sulphur-containing compounds as flavour enhancers 119
8.2.5 Methyl xanthines and IMP 121
8.3 Seafood taste and umami substances 121
8.3.1 Extractive components 122
8.3.2 Omission test 125
8.4 Taste-active components in seafoods 125
8.4.1 Abalone 125
8.4.2 Sea-urchin 126
8.4.3 Snow crab 126
8.4.4 Scallop 127
8.4.5 Short-necked clam 127
8.4.6 Dried skipjack 127
8.4.7 Salted salmon eggs 128
8.5 Taste specificity of seafoods 128
8.6 Taste-active components in ripened or fermented seafoods 132
8.6.1 Squid muscle 132
8.6.2 Sea-urchin 133
8.6.3 Fish sauces 133
8.6.4 Mullet roe 134
8.7 Changes in extractive components by heating 134
8.8 The roles of components other than extractive components 135
Acknowledgement 136
References 136
XII CONTENTS
Part 2 Quality of seafoods
9 Freshness quality of seafoods: a review 140

1. R. BOTTA
9.1 Introduction 140

9.2 Appearance 141
9.3 Texture 146
9.4 Odour and flavour 153
9.5 Disadvantages of destructive and/or slow analyses 160
9.6 Alternative methods 160
9.7 Conclusion 161
References 162
10 Preservation of seafood quality 168

Z. E. SIKORSKI and B. SUN PAN

10.2 Factors affecting the loss of quality in fresh fish 169
10.2.1 Biological variations 169
10.2.2 Harvesting conditions 171
10.2.3 Post-harvest handling 172
10.3 Chilling of fresh fish 173
10.3.1 Factors affecting the shelf-life 173
10.3.2 Chilling at subzero temperatures 175
10.3.3 Use of chemical preservatives 175
10.3.4 Application of modified atmospheres 176
10.3.5 Radiation preservation 177
10.3.6 Packing of fresh fish 178
10.4 Freezing and frozen storage 179
10.4.1 Factors affecting the quality of frozen products 179
10.4.2 The effect of freezing rate and storage temperature 179
10.4.3 The effect of protein changes 180
10.4.4 The effect of dehydration and lipid changes 182
10.5 Drying of seafoods 183
10.5.1 Role in preservation of seafoods 183
10.5.2 The quality of products 184
10.5.3 The shelf-life of dried seafoods 184
10.6 Smoking 185
10.6.1 The preserving action 185
10.6.2 The sensory quality of smoked seafoods 186
10.6.3 The risk factors 186
10.7 Other methods of preservation 187
10.8 Conclusions 188
References 189
11 Microbiological quality of seafoods: viruses, bacteria and

parasites 196
M. M. WEKELL, R. MANGER, K. COLBURN,
A. ADAMS and W. HILL

11.1.1 Seafood-borne illness 196
11.2 Viruses in seafoods 198
CONTENTS Xlii
11.2.1 Hepatitis A 199

11.2.2 Norwalk virus 200
11.3 Bacteria in seafoods 201
11.3.1 Potential bacterial pathogens in seafoods 202
11.4 Applications of molecular biological techniques in (sea)food
microbiology 205
11.4.1 Gene probes 205
11.4.2 PCR: promise and problems 206
11.5 Parasites in seafoods 208
11.5.1 Anisakids 209
11.5.2 Tapeworms 210
11.5.3 Trematodes 211
11.5.4 Control of parasites in seafoods 212
11.6 Conclusion 212
References 214
12 Microbiological quality of seafoods: marine toxins 220

M. M. WEKELL and 1. M. HUNGERFORD

12.2 Paralytic shellfish poisoning 221
12.3 Amnesic shellfish poisoning 222
12.3.1 Assay methods 223
12.4 Diarrhetic shellfish poisoning 224
12.5 Ciguatera 225
12.6 Scombroid poisoning 227
12.7 Summary 228
References 228
13 Sensory assessment of quality in fish and seafoods 233

R. K. YORK and L. M. SEREDA

13.2 Developments in assessment of fish by expert assessors - grading 237
13.3 Developments in descriptive analysis 240
13.4 Developments in discriminative testing and applied research 243
13.5 Developments in consumer testing 247
13.6 Developments in sensory assessment of environmentally-induced
off-flavours in fish 250
13.7 Conclusion 253
References 253
Appendix 257
Part 3 Further processing of raw material
14 Surimi processing from lean fish 263

C. M. LEE

14.2 Surimi manufacturing from lean fish 264
14.2.1 Conventional and modified surimi manufacturing processes 265
xiv CONTENTS
14.2.2 Factors affecting the surimi quality and yield 267

14.2.3 Factors affecting the frozen storability of surimi 270
14.3 Manufacturing of surimi-based products 275
14.3.1 Factors affecting the surimi sol and gel properties 275
14.3.2 Relationship of sol viscosity to gel properties 276
14.3.3 Comminution time and temperature 276
14.3.4 Gel setting 277
14.3.5 Mechanisms of gel-strengthening effect of various ingredients 279
14 Suggested future work 284
14.4.1 Surimi manufacturing 284
14.4.2 Surimi-based products 285
References 285
15 Surimi processing from fatty fish 288

K. E. SPENCER and M. A. TUNG

15.2 Relevant literature 289
15.2.1 Thesurimiprocess 289
15.2.2 Surimi quality 292
15.2.3 Technological developments 294
15.2.4 Other research findings 297
15.2.5 Surimi processing at Cln 299
15.3 Experimental 299
15.3.1 Process development for Atlantic species 299
15.3.2 Analytical methods 301
15.3.3 Mackerel surimi processing trials 302
15.3.4 Herringsurimi processing trials 304
15.4 Results and discussion 305
15.4.1 Mackerel surimi trials 305
15.4.2 Herring surimi trials 309
15.5 Summary 316
15.6 Conclusions 316
Acknowledgements 317
References 317
16 Seafood processing by-products 320

F. SHAHIDI

16.2 Composition of processing discards 321
16.2.1 Fish processing discards 321
16.2.2 Shellfish processing discards 322
16.3 Discard components and their value-added utilization 323
16.3.1 Proteins 323
16.3.2 Chitinous materials 324
16.3.3 Carotenoid pigments and their use in aquaculture feed 328
16.4 Other value-added components of seafood processing discards 331
16.4.1 Fish enzymes 331
16.4.2 Antifreeze proteins 332
16.4.3 Seafood lipids 332
Acknowledgement 333
References 333
Index 335
1 The chemistry, processing technology and quality
of seafoods - an overview
F. SHAHIDI
The world's annual catch of fish and marine invertebrates is approximately

100 million metric tons, from which only 20% is processed for food use.
Approximately 30% of the latter amount is consumed and the rest is
generally discarded as waste. Many species of fish and invertebrates are
rarely used as food since they possess undesirable flavour or dark colour
and may be small in size with a large proportion of bones and skin. Such
raw materials are generally reduced to meal and oil, silage or protein
concentrates/hydrolysates.
Seafoods have a variable content of protein generally ranging from 12 to
26%. Contrary to animal meat, where ageing brings about a desirable
flavour and texture in the product, seafoods tend to deteriorate rapidly
during the post-mortem period. Therefore, preservation of fresh quality of
seafoods is quite important.
Quality deterioration of seafoods is due to post-mortem biochemical
changes in proteins and non-protein nitrogenous compounds. Enzymatic
and non-enzymatic oxidation of lipids may also play an important role in
modifying the fresh quality of seafoods. Interaction of lipid oxidation
products with proteins may result in cross-linking of protein chains and
toughening as well as discoloration of muscle tissues. Rapid loss of fresh
quality of seafoods may also be due to a variety of changes caused by
hydrolysis, polymerization, deamination, decarboxylation or oxidative
processes. The rate of quality deterioration of seafoods is therefore
dependent on the species being considered, its compositional character-
istics, packaging, storage and processing conditions.
The flavour of seafoods depends primarily on the species being
considered. The content of non-volatile taste-active components, mainly
non-protein nitrogenous compounds and lipids, also affects the aroma
quality of such products.
Presence of highly unsaturated fatty acids (HUFA) in seafoods has been
attributed to their possible health benefits in human nutrition. Long-chain
polyunsaturated omega-3 fatty acids of seafoods have been implicated as
being responsible for reducing the incidence of cardiovascular diseases in
humans. However, HUFA are highly prone to autoxidation as well as
enzymatic breakdown. Therefore, their quality preservation is essential.
F. Shahidi et al. (eds.), Seafoods: Chemistry, Processing Technology and Quality

© Springer Science+Business Media Dordrecht 1994
2 SEAFOODS
There are many unutilized and underutilized species from aquatic origin
which possess a well-balanced amino acid composition and protein quality.
However, due to their small size, high fat content, strong flavours and high
proportion of bones in the carcass they are not used as such in food
preparations. In addition, processing discards from the seafood industry
may provide the necessary raw material for isolation of value-added
components. Recovery of seafood proteins from filleting of fish or from
underutilized fatty fish species by mechanical deboning followed by
aqueous washing affords a functional protein preparation. Such protein
preparations, after addition of a combination of salt, polyphosphates,
sorbitol and possibly sucrose, known as cryoprotectants, are known as
surimi and may be used for preparation of a wide variety of seafood
analogues. Bland taste of such raw materials allows inclusion of flavour
extracts and formulation of a large number of products. Furthermore,
other valuable components such as enzymes, antifreeze proteins, chitin,
carotenoid pigments and flavorants may be isolated from processing
discards of aquatic species.
Despite all beneficial health effects of seafoods, it is imperative to assure
that the fresh quality of seafoods is preserved. Both subjective and
objective methods of analysis are used for this purpose. In addition,
microbiological activity during the post-mortem period must be controlled
in order to avoid health risk from the consumption of seafoods.
Furthermore, the raw material must be scrutinized from possible presence
of toxins, particularly in the cultured species. Finally, depletion of natural
seafood resources in certain areas may provide further opportunities for
the growth of aquaculture industries.
One challenge in the development of aquaculture industries and fish
farming is feed production. A major cost of this industry is availability of
suitable raw material for production of formulated feed. Fortunately,
processing of seafoods and availability of underutilized species provide the
necessary raw material which is required for production of novel feed
products.
2 Seafood proteins and preparation of protein
concentrates
F. SHAHIDI
2.1 Introduction
Seafoods namely fish, crustaceans and molluscs are important sources of

protein in human nutrition. Approximately 11-27% of seafoods consist of
crude proteins. However, contribution of non-protein nitrogenous (NPN)
compounds to this value may reach up to a quarter of this amount. Free
amino acids, peptides, amines, amine oxides, guanidines, quaternary
ammonium compounds, polyamines, nucleotides and their breakdown
products, urea and nucleic acids contribute to the overall content of NPN
in seafoods.
Seafood proteins, like those of all other muscle foods, may be classified
as sacroplasmic, myofibrillar and stroma-type. The sacroplasmic proteins,
mainly albumins, account for approximately 30% of the total muscle
proteins. A large proportion of sarcoplasmic proteins may be composed of
haemoproteins. Seal meat contains up to 10% myoglobin and haemoglobin
and as a result has a very dark colour. The myofibrillar proteins in muscle
are myosin, actin, actomyosin and troponin and these account for 40-60%
of the total crude protein content of fish. The rest of the muscle proteins,
classified as stroma, are mainly collagenous matters.
2.2 Sarcoplasmic proteins
Sarcoplasmic proteins, referred to as 'myogen', are soluble in the muscle

sarcoplasm. They include a large number of proteins such as myoglobin,
enzymes and other albumins. Sarcoplasmic proteins from comminuted fish
may be extracted into water or neutral salt solutions with an ionic strength
of less than 0.15 Debye (Scopes, 1970). Sarcoplasmic proteins may be used
to fingerprint fish species using electrophoretic and isoelectric focusing
techniques (Toom et at., 1982).
The content of sarcoplasmic proteins is generally higher in pelagic fish
species as compared with demersal fish. Dark muscles of some species
contain less sarcoplasmic proteins than their white muscle counterpart
(Suzuki, 1981). However, presence of larger amounts of myoglobin,

4 SEAFOODS
haemoglobin and cytochrome C in dark muscles may reverse this trend.

Furthermore, fish species from cold-water oceans contain certain character-
istic proteins referred to as antifreeze proteins and glycoproteins which
belong to the sarcoplasmic group.
Sarcoplasmic enzymes are responsible for quality deterioration of fish
after death. These include glycolytic and hydrolytic enzymes. Various
proteinases found in the sarcoplasmic fraction may further catalyse the
degradation of nitrogenous compounds in the muscle tissues. The activity
of these enzymes depends on the fish species, type of muscle tissue as well
as seasonal and environmental factors. The handling and post-harvest
condition of fish also determines the activity of these enzymes.
2.3 Myofibrillar proteins
The largest amount of myofibrillar proteins present in muscles of aquatic

species are myosin, actin, tropomyosin and troponins C, I and T (Suzuki,
1981). These proteins are extracted from comminuted fish with neutral salt
solutions of ionic strength ranging from 0.3 to 1.0 Debye. Myofibrillar
proteins undergo changes during the rigor mortis, resolution of rigor
mortis and long-term frozen storage. The texture of fish products and the
gel-forming ability of fish minces and surimi may also be affected by these
changes.
Myosin is the most abundant myofibrillar fraction of fish muscles and
contributes 50 to 60% to its total amount. The myosin molecule consists of
two heavy chains (200 and 240 kD) associated non-covalently with two
pairs of light chains (16 to 28 kD). In contrast to most proteins of
mammalian origin, the loss of ATPase activity occurs at a faster rate in fish
muscle (Arai et al., 1973; Kimura et al., 1980). The ATPase of squid
myosin is activated by Ca 2+ as well as Mg 2 + with two pH activity maxima
(Tsuchiya et al., 1978).
Paramyosin is a protein found in invertebrates and is present in
quantities ranging from 0.1 to 10 times that of myosin (Offer, 1987).
Paramyosin molecules constitute a core of the thick myofibrils of
invertebrate muscles which is covered by a layer of myosin. Paramyosin is
known to maintain the tension in the muscle tissues.
Actin, the second most abundant myofibrillar protein, constitutes
approximately 20% of the total content of these proteins in fish muscles.
G-actin is the monomeric form of the molecule and in the presence of
neutral salts polymerizes to F-actin. When minced fish is extracted with
neutral salt solutions, actin is co-extracted with myosin in the form of
actomyosin. Similar to myosin, actin shows Ca 2 + and Mg 2 + activated
ATPase characteristics.
Other constituents of myofibrillar proteins include tropomyosin and
SEAFOOD PROTEINS AND PROTEIN CONCENTRATES 5
troponins which account for 10% of the total amount. Several isoforms of
tropomyosin have been described for different sources of muscle foods.
Their number depends on the species from which the muscle is originated,
however, sequence of the homogenous isoforms from different species are
nearly identical. Similarly troponins exist in several isoforms (Nishita and
Ojima, 1990).
2.4 Stroma proteins
The residue after extraction of sarcoplasmic and myofibrillar proteins is

known as stroma. It is composed of collagen and elastin from the
connective tissues. Stroma is soluble in dilute solutions of HCl or NaOH
and contributes up to 10% of the crude muscle proteins.
The collagen content in muscle depends on the species, feeding regime
and state of maturity of the fish. In general, fish muscles contain
approximately 0.2 to 2.2% collagen (Sato et ai., 1986). Although a higher
content of collagen contributes to the toughness of muscles, in fish no such
problems are encountered. However, some species of squid may develop a
tough and rubbery texture upon heat processing.
2.5 Non-protein nitrogenous (NPN) compounds
The content of NPN compounds in seafood depends primarily on the

species, state of freshness, habitat and other effects. Futhermore, content
of NPN compounds in seafoods is also dependent on the fish species; they
account for up to 55% of the total nitrogen in shark muscle tissues. The
dark muscles of fish generally contain a higher amount of NPN compounds
than the light muscles.
Table 2.1 Distribution (%) of non-protein nitrogenous (NPN) compounds in some seafoods"
Free Creatine!
Species amino acids Peptides Nucleotides creatinine Betaines Others h
Mackerel 25 5 10 35 25
Shark 5 5 5 10 75'
Shrimp 65 5 5 5 20
Crab 75 5 15 5
Squid 50 5 15 10 20d
Clam 35 5 35 15 10
"Adapted from Finne (1992).

b Ammonia, urea, trimethylamine oxide and amides.
'Primarily urea.
dYncludes 10% octopine.
6 SEAFOODS
Approximately 85% of NPN compounds in muscle tissues of marine fish

and shellfish is composed of free amino acids, amines, amine oxides,
guanidines, nucleotides and their breakdown products, urea and quaternary
ammonium salts (Ikeda, 1979). The contribution of NPN compounds to
flavour of seafoods is of paramount importance (Table 2.1).
2.6 Surimi
Surimi is mechanically deboned fish flesh which has been washed with
water or very dilute salt solutions at 5-lOo C and to which cryoprotactants
have been added (Lee, 1984). Removal of water-soluble proteins including
enzymes, haemoproteins and other NPN compounds results in product
yields of 5~O%. The content of collagenous matters as well as
myofibrillar proteins is higher in surimi as compared with fish flesh.
However, the amino acid composition of the resultant surimi may be
similar to that of the starting material as was reported for seal surimi
(Synowiecki et al., 1992). Similar results were obtained for thermostable
water dispersions of myofibrillar proteins from mackerel (Venugopal and
Shahidi, 1994).
2.7 Protein concentrates/hydrolysates
After the death of fish digestive enzymes and acids produced in the body,
as a result of glycolysis, quickly attack the muscle tissues of some fish
species. For example, intense activity of enzymes present in the gastro-
intestinal tract of capelin (Mallotus villasus) results in the so-called 'belly
burst'. Incidences of belly burst are greater during the intense feeding
period of these fish. Activation of endogenous enzymes in fish species by
addition of acids such as formic acid has been in practice in fish silage
production. However, exogenous enzymes may be added to fish species
such as male and spent capelin in order to hasten the liquification process,
to produce high quality products and to allow their full utilization as foods.
In addition to enzymatic modification of seafood proteins in the
preparation of concentrates, two other techniques for concentrate prepara-
tion from seafoods may be employed. These are physico-chemical
separation/extraction which includes extraction with organic solvents such
as ethanol and isopropanol as well as aqueous extraction methods.
Modification of proteins via acetylation or succinylation is also possible.
Each of the two techniques given above suffers from certain drawbacks.
For example, the highly-desired functional properties of chemically-
modified proteins prepared by acylation do not compensate for the loss of
nutritional value of the product. The loss of functional properties in solvent
extraction procedures suffers from operational hazards such as fire and

requires use of expensive equipment.
One of the many potential uses of enzymes for modification and
improvement of protein is via controlled hydrolysis. A broad range of
products of intermediate and high cost may be produced. Preparation of
products from both lean and fatty fish species is possible. Lean fish
generally afford the so-called 'type A' concentrate which contains at least
67.5% crude protein, on a dry basis, and not more than 0.75% lipid.
Products of this type are generally colourless, tasteless and odourless.
Concentrates from fatty fish provide the 'type B' product with a lipid
content of up to 10% with a distinct fishy flavour. Both products may be
used as food components; however, type B may be more suitable for
populations accustomed to fish-containing diets (Ockerman and Hansen,
1988).
Protein concentrates with a bland taste and an ivory colour were
prepared from capelin (Mallotus villosus) using the commercially available
Alcalase enzyme (Novo Nordisk A/S, Bagsvaerd, Denmark). AIcalase is
an endo-protease prepared by submerged fermentation of a selected strain
of Bacillus licheninformis and can hydrolyse most peptide bonds in a
protein. Figure 2.1 shows a typical flowsheet for preparation of capelin
protein hydrolysate. The products so obtained, depending on their degree
of dehydration, may be in the liquid, cream/paste or powder forms. At a
degree of hydrolysis of less than 20, reached within 1 to 2.5 h at 45-65°C
(Table 2.2), high quality protein products with a bland taste and an ivory
Table 2.2 Progress of Alcalasc-assisted hydrolysis of capelin over a 2,5 h period a
Degree of hydrolysis ('Yo) at °C

Hydrolysis time
(min) 45 55 65
10 0.78 9.45 10.98

20 9.90 13.20 14.28
30 12.16 15.56 15.93
40 14.00 17.37 17.03
50 15.55 18.62 17.21
60 16,68 19.73 18.26
70 17,95 20.90 18.67
80 19.00 21.40 18.95
90 20,08 22.08 19.22
100 20.92 22.65 19.36
110 21.77 23.20 19.50
120 22.48 23.76 19.55
130 23.20 24.18 19,60
140 23.90 24.59 19.63
150 24.32 25.01 19.63
"Mean values of three determinations using an Alcalase concentration of 48.7 Anson units/kg
protein,
8 SEAFOODS
Water
DILUTION
NaOH Solution I
I
I
pH ADJUSTMENf
AlcalaselNaOH
HYDROLYSIS
Hel solution
ENZYME INACTIVATION
DECOLORIZATIONI
FILTRATION
NaOH Solut;on I I L.--o Solids
I NEUTRALIZATION I
DEHYDRATION
~
C'APELIN PROTEIN CONCENTRATE
Figure 2.1 Flowsheet for preparation of capelin protein concentrate from male and spent
capelin.
colour were obtained. Amino acid composition of the hydrolysate

prepared from capelin was similar to that of the starting material (Table
2.3).
References
Arai, K., Kawamura, K. and Hayashi, C. (1973) The relative thermostabilities of the
actomyosin-ATPase from the dorsal muscles of the various fish species. Bull. Jpn. Soc. Sci.
Fish, 39, 1077-85.
Finne, G. (1992) Non-protein nitrogen compounds in fish and shellfish. In Advances in
Table 2.3 Typical amino acid composition of proteins and protein hydrolysates from capelin
(Mallotus villosus)"
Amino acid Capelin Capelin hydrolysate
Alanine 5.57 6.00

Arginine 5.99 5.70
Aspartic Acid + Asparagine 8.86 9.89
Cysteine 1.33 1.34
Glutamic Acid + Glutamine 13.19 13.43
Glycine 5.32 5.14
Histidine 2.43 2.09
Hydroxyproline 0.42 0.46
Isoleucine 4.72 4.25
Leucine 8.15 7.60
Lysine 8.47 8.49
Methionine 3.09 2.05
Phenylalanine 3.80 3.19
Proline 3.70 3.67
Serine 4.18 4.24
Threonine 4.28 4.56
Tryptophan 1.07 0.43
Tyrosine 3.34 2.47
Valine 5.71 5.77
aMean values of three determinations.
Seafood Biochemistry: Composition and Quality, (eds. Flick, G.l., lr. and Martin, R.E.),
Technomic Pub. Co. Inc., Lancaster and Basel, pp. 393-401.
Ikeda, S. (1979) Other organic components and inorganic components. In Advances in Fish
Science and Technology, (ed. Connell, 1.1.), Fishing News Books, Surrey, pp. 111-124.
Kimura, 1., Yoshitomi, B., Kenno, K. and Arai, K. (1980) Preparation of highly purified
myosin from mantle muscle of squid Onmastrephes sloani pacificus. Bull. Hpn. Soc. Sci.
Fish, 46, 885-9.
Lee, CM. (1984) Surimi process technology. Food Techno/., 38 (11), 69-80.
Nishita, K. and Ojima, T. (1990) America Lobster troponin. l. Biochem., 108,677-83.
Ockerman, H.W. and Hansen, c.L. (1988) Seafood by-products. In Animal By-Products
Processing, VCH, Weinheim, Germany, pp. 279-308.
Offer, G. (1987) Myosin filaments. In Fibrous Protein Structure, (eds. Squire, J.M. and
Vibert, P.l.), Academic Press, London, pp. 307-357.
Sato, K., Yoshinaka, R., Sato, M. and Shimizu, Y. (1986) Collagen content in the muscle
of fishes in association with their swimming movement and meat texture. Bull. lpn. Soc.
Sci. Fish, 52, 1595-8.
Scopes, R.K. (1970) Characterization and study of sarcoplasmic proteins. In The Physiology
and Biochemistry of Muscle as Food, Volume 2, (eds. Briskey, E.l., Cassen, R.G. and
Marsh, B.B.), The University of Wisconsin Press, Madison, pp. 471-90.
Suzuki, T. (1981) Fish and Krill Processing Technology, Applied Science Publishers, London,
pp. 10--13.
Synowiecki, 1., Hall, D. and Shahidi, F. (1992) Effect of washing on the amino acid
composition and connective tissue content of seal meat. l. Muscle Foods, 3, 25-31.
Toom, P.M., Ward, C.F. and Weber, 1.R. (1982) Identification of fish species by isoelectric
focusing. In Chemistry and Biochemistry of Marine Food Products, (eds. Martin, R.E.,
Flick, G.F., Hebard, C.E. and Ward, D.R.), AVI Publishing Co., Westport, CN,
pp. 51-65.
Tsuckiya, T., Yamada, N. and Matsumoto, 1.1. (1978) Extraction and purification of squid
myosin. Bull. lpn. Soc. Sci. Fish, 44, 175-7.
Venugopal, V. and Shahidi, F. (1994) Thermostable water dispersions of myofibrillar
proteins of Atlantic mackerel (Scomber scombrus). l. Food Sci., in press.
3 Protein hydrolysis in seafoods
N.F. HAARD
3.1 Introduction
3. J.1 Importance
Proteinases are probably studied more by food scientists and technologists
than any other group of enzymes in fish and shellfish. That these enzymes
receive considerable attention is not surprising since protein hydrolysis can
contribute to either improvement or to loss in seafood quality.
3. J. J.1 Quality improvement. The products of protein hydrolysis by

tissue proteases make a positive contribution to the pleasing aroma, taste,
colour and texture of traditional fermentation products such as fish sauce,
lightly salted fish, dried squid and caviar (Table 3.1). Likewise, endogenous
proteases may contribute to the process of preparing protein hydrolysates
for food or animal feed. Digestive proteases from fish are also of value as a
by-product. In recent years, there has been a growing interest in recovering
digestive proteolytic enzymes from fish processing waste for use as
industrial processing aids (reviewed by Simpson and Haard, 1987; Haard,
1992). Suggested uses of digestive proteases from fish include aids for
cheese making, herring fermentation, fish skinning, oxidative stability of
milk and roe processing as well as medical applications.
3. J. J.2 Quality deterioration. The action of tissue proteinases can also

contribute to product quality deterioration (Table 3.2). Post-harvest
biochemical change caused by endogenous enzymes, including proteinases,
is the primary cause of quality loss in iced fish and also limits the efficacy of
storage strategies, such as modified atmospheres, which target bacterial
spoilage (reviewed by Haard, 1992a). Moreover, proteinases can be
directly responsible for unusual textural defects in seafood, e.g. 'belly
burst', 'gaping', and 'mushiness' of bony fish and 'tail meat' softening of
crustacean species and thus may cause poor initial quality and fillet yields.
Heat activated proteinases also adversely influence seafood quality by
causing excessive softening of fish meat or fish gel products during cooking.
Latent proteinases activated by heating to temperatures in the 50-65°C
range cause very rapid hydrolysis of myosin and other muscle proteins. The

PROTEIN HYDROLYSIS IN SEAFOODS 11
Table 3.1 Importance of endogenous proteases to quality improvement of fishery products
Product Quality improvement Enzyme(s) involved
Fermentations yielding food products

Roe processing Flavour Cathepsin D-like and aminopeptidase
activities result in a large increase in free
amino acids during processing (Chiou et
al., 1989)
Fish sauce Yield and flavour Cathepsins A, C and trypsin-like enzyme
are endogenous proteases that appear to
contribute to the process (Orejana and
Liston, 1982; Rosario and Maldo, 1984;
Raksakulkthai et al., 1986)
Dried squid Texture and flavour Cathepsin C appears to contribute to
quality of traditional product (Haard,
1983)
Matjes herring Texture and flavour Trypsin appears to be a key enzyme
(Simpson and Haard, 1984)
Fish protein hydrolysates

Fish silage Yield Pepsins, gastricsins, cathepsin D, and
aminopeptidases appear to be among key
enzymes involved with the process (Haard
et al., 1985)
Hydrolysates Yield Normally supplemented by added
protease(s), but endogenous enzymes also
appear to be important
Enzyme by-products
Gastric mucosa Unique properties Pepsins and gastricsins have been isolated
and may find use as food and feed process
aids (Haard et al.. 1982, 1983; Haard,
1986)
Pyloric ceca Unique properties Trypsins and chymotrypsins have been
isolated and may find use as food, feed
and medical process aids (Simpson and
Haard, 1984. 1985, 1987)
Hepatopancreas Unique properties Trypsin-like enzymes and cathepsin C have
been isolated and may find application as
food, feed and medical process aids
(Hameed and Haard, 1985)
so-called 'modori' phenomenon in kamaboko-type products and shellfish

analogues is an example of this problem and has been extensively studied.
3.1.2 Source of proteinases in seafood
Identification of the specific proteinase(s) responsible for a particular

change is quite complicated because:
(1) there are many different muscle enzymes which hydrolyse protein;
(2) enzymes from other tissues may contaminate muscle;
12 SEAFOODS
Table 3.2 Importance of endogenous proteases to quality deterioration of fishery products
Raw material Quality deterioration Enzyme(s) involved
Holding or processing raw material

Pelagic fish Belly burst, flesh Trypsin-like and other alkaline proteases
softening diffuse from pyloric ceca and intestine to
cause degradation of myofibrillar and
connective tissue proteins (e.g.
Hjelmeland and Raa, 1980; Martinez and
Gildbcrg, 1988)
Crustacean sp. Abdominal muscle Trypsin-like and collagenase enzymes
softening diffuse from hepatopancreas to cause 'tail
meat' softening in products like shrimp
and krill (Nip et al., 1985; Honjo et al.,
1990)
Chum salmon Whitish and very mushy Cathepsins Band L are elevated in Chum
texture salmon during spawning migration
(Konagaya, 1983)
Cod Gaping, poor fillet yield Collagenase(s) and multicatalytic proteases
may be involved with loss of muscle tissue
integrity during post-harvest storage (Lee
and Pan, 1990; Bracho and Haard, 1990)
Thermoprocessed fish products

Jellied fish Extreme texture Cysteine proteases, etc. formed by
softening myxosporidia contaminating muscle are
very active during slow cooking
(Patashnik et al., 1982; Konagaya, 1984)
Fish Loss of gel strength Heat-stable alkaline proteases and neutral
'Modori' proteases (modori inducing) are active
when the product temperature reaches
6O-70°C (Lin and Lanier, 1980; Kinoshita
et al., 1990)
Scombroid fish 'Honeycomb' Degradation of connective tissue appears
to be associated with muscle alkaline
protease(s) and pyloric ceca collagenase
(Pan et al., 1986)
(3) the physiological state of the animal may influence proteinase activity;
(4) a broad taxonomic array of vertebrate and invertebrate organisms
make up seafood;
(5) enzymes may originate from contaminating microorganisms or para-
sites.
Elucidation of enzymes involved with postharvest changes is often done by
studying characteristics like pH optimum and inhibitor sensitivity of tissue
homogenates or impure enzyme isolates. However, conclusions relating
the specific contribution of an enzyme to a change in post-harvest quality
based on inhibition or activation of impure enzyme isolates may be
misleading. For example, the action of an inhibitor may be indirect,
exerting an influence on the conversion of proenzyme to active enzyme by
a zymogen activating proteinase that is not itself directly involved with the
quality change (Hara et ai., 1988).
Exogenous proteinases, i.e. originating from fish spoilage bacteria (e.g.
Venugopal, 1990), bacterial pathogens of fish (e.g. Norqvist et ai., 1990)
and protozoan parasites (e.g. Patashnik et ai., 1982; Konagaya, 1983a;
Chang-Lee et ai., 1989) are beyond the scope of this review. It should,
however, be cautioned that elevated enzyme levels in infected tissue may
originate in the host via stress response and activation of protein catabolism
by tissue enzymes.
3.1.2.1 Intraspecific factors. Within a species, the amount of endogenous

enzyme present in a tissue can be dramatically influenced by intraspecific
factors such as biological age, diet, exercise, water salinity and hydrostatic
pressure as reviewed elsewhere (Haard, 1990; 1992b). Protein synthesis and
degradation in muscle tissue of living animals is particularly sensitive to
malnutrition and catabolic states caused by disease and stress (Horl et ai. ,
1987; Gerard et ai., 1988; Kopple, 1988; Reddy et ai., 1990). Therefore,
the physiological state of the animal prior to harvest can be responsible for
quality change which occurs during subsequent post-harvest storage or
processing of the meat. Physiological status prior to harvest may influence
enzyme content, as during spawning migration of chum salmon where
cathepsin L is elevated (Konagaya, 1983). It may also exert influence on
potential protein hydrolysis via control of endogenous inhibitors, such as
by regulation of the content of carnosine, anserine or I-methyl-histidine
which, in turn, can modify the action of the calpain inhibitor calpastatin in
carp (Toyohara and Makinodan, 1989). Inhibitors of serine proteases
(Hjelmeland and Raa, 1980; Toyohara et ai., 1983) and cysteine proteases
(Toyohara and Makinodan, 1988; Fujita and Horiuchi, 1990) have also
been identified in fish muscle tissue.
3.1.2.2 Endogenous proteinases. Constitutive proteinases important in

seafood include those present in: (1) muscle cells; (2) the extracellular
matrix and connective tissue surrounding muscle cells; and (3) digestive
and other organs.
The characterization of fish proteinases started with digestive proteases,
initially studied more than 100 years ago (Stirling, 1884). Some digestive
enzymes like salmon pepsin were purified more than 50 years ago (e.g.
Norris and Elam, 1940). In more recent years we have gained better
understanding of multiple molecular forms of digestive enzymes (Squires et
ai., 1986; Simpson and Haard, 1987a) and their comparative biochemistry
(Simpson and Haard, 1987; Gildberg, 1988). The presence of lysosomal
enzymes in fish muscle has been known for almost 30 years (Siebert, 1962;
Tappel et ai., 1963) although new catheptic enzymes continue to be
identified in fish and shellfish (e.g. Hara et ai., 1988a; Veno et ai., 1988;
14 SEAFOODS
Zeef and Dennison, 1988). The high molecular weight, heat stable,
alkaline muscle proteinases were identified in fish muscle nearly 20 years
ago (Iwata et at., 1973) and have been extensively studied in connection
with surimi gel weakening. More recently identified proteinases from
seafood tissues include Ca 2 +-activated neutral proteases (e.g. Jiang et at.,
1990), modori inducing neutral proteinases (e.g. Kinoshita et at., 1990),
multicatalytic proteinases or proteosomes (e.g. Folco et at., 1989), and
metalloproteinases with collagen degrading activity (Bracho and Haard,
1990).
3.2 Digestive proteinases
It has long been recognized that digestive proteases play a role in the
softening of abdominal tissues during post-mortem storage of fish (Almy,
1926). Pepsin and trypsin have been detected in the belly cavity adjacent to
muscle of fresh fish (Siebert and Botkke, 1963). Gildberg (1982) has
suggested that leakage of digestive proteinases into the belly cavity of fish
can activate collagenases present in the connective tissue as well as directly
initiate collagen degradation by digestive collagenase. It is interesting that
certain types of feed in the digestive tract can trigger post-mortem autolysis
in so-called 'feedy fish'. Retained organ tissues can also increase the
alkaline proteinase content of minced fish and surimi (Su et at., 1981).
3.2.1 Gastric proteases

The gastric mucosal glands of stomached vertebrates secrete zymogens of
aspartic proteinases including pepsins, gastricsins, and chymosins. Until
about 1980 these enzymes were called carboxyl proteinases. Pepstatin, a
pentapeptide produced by some actinomyces, is a very specific inhibitor of
aspartic proteinases. Relatively little work has been done on chymosin and
gastricsin from aquatic organisms and the criteria for classification of these
enzymes is somewhat equivocal (Shamsuzzaman and Haard, 1984); Squires
et al., 1986a). Unlike pepsins, gastricsins appear to be activated by low
concentrations of NaCl (Sanchez-Chang and Ponce, 1981) and this
property may be important in salted seafood products. Fish pepsin may be
represented by four or more isoenzymes (e.g. Squires et al., 1986) but
normally there are two major isoenzyme forms called pepsin I and II
(Gildberg, 1988). The unique properties of fish pepsins have been
reviewed (Haard et at., 1982; Gildberg, 1988) and include broad, relatively
high pH optima (pH 3.5), low thermostability and high molecular activity
at low temperature for cold water species, and more basic amino acids in
the polypeptide chain than mammalian pepsins. Invertebrates and stomach-
less fish with little or no gastric acid secretion do not appear to contain
pepsins although they may contain the homologous enzyme cathepsin D

(Gildberg, 1988).
3.2.2 Intestinal proteinases

Proteases in the intestine of fish are normally secreted from the pyloric
ceca or pancreas depending on species. These include trypsin, trypsin-like
enzyme, chymotrypsin, collagenase, elastase, carboxyl peptidase and
carboxylesterase (Yoshinaka et al., 1981a; Smith, 1989; Chen et al., 1989).
3.2.2.1 Trypsins. Trypsins are serine proteinases which often occur in

multiple or isoenzyme forms (e.g. Hjelmeland and Raa, 1982; Simpson
and Haard, 1987a; Martinez et al., 1988). In some cases multiple enzyme
forms may be different activation products from one zymogen. Comparative
aspects of fish trypsins have been described (Hjelmeland and Raa, 1982;
Simpson and Haard, 1984a; 1984b; Gildberg, 1982; Asgeirsson and
Bjarnsason, 1989; Kolodzeiskaia and Berevka, 1990) and include relatively
low thermal stability and high molecular activity at low temperature for
cold water species, instability at low pH, and a lower content of basic
amino acid residues in the polypeptide chain than mammalian trypsins.
The criteria for classifying an enzyme as trypsin or trypsin-like are
equivocal, although features used to distinguish them normally include
molecular weight and specificity with synthetic substrates. It is interesting
that fish muscle contains trypsin inhibitors which may play a role in
preventing attack by digestive trypsin in post-mortem fish (Hjelmland and
Raa, 1980).
3.2.2.2 Chymotrypsins. Chymotrypsin and chymotrypsin-like serine

proteinases in fish have been studied less than the trypsins. The zymogen
from catfish pancreas exhibited very slow activation by trypsin indicating
the presence of endogenous inhibitors (Yoshinaka et al., 1981). The
properties of capelin and herring chymotrypsins have also been reported
(Kalac, 1978).
There is evidence that chymotrypsins have unique properties when
compared to the enzyme from mammalian sources. Like trypsins,
chymotrypsins from fish are less stable in acidic media than mammalian
chymotrypsin (Kolodzeiskaia and Pivnenko, 1988). Salmon chymotrypsin
is more stable to formaldehyde inactivation and has a different affinity for
aromatic ligands than bovine chymotrypsin (Kolodzeiskaia and Berevka,
1990). Dogfish chymotrypsin has broader protein hydrolytic specificity and
more efficient incorporation of methionine into plastein than bovine
chymotrypsin (Ramakrishna et al., 1987).
16 SEAFOODS
3.2.2.3 Collagenases. Less is known about the comparative properties of

other intestinal proteinases. Collagenase has been isolated from the pyloric
ceca of yellowtail (Yoshinaka et al., 1977), mackerel (Pan et al., 1986),
and the digestive organs of 19 fish species (Yoshinaka et al., 1978).
Collagenase appears to be most abundant in the fatty deposits between the
ceca rather than within the ceca (Yoshinaka et al., 1978). Evidence
provided by Pan et al. (1986) indicates that seepage of digestive collagenase
to muscle tissue contributes to the 'honey comb' disorder in canned
mackerel. Digestive collagenase differs from tissue metalloproteinase
collagenases in being classified, like trypsin and chymotrypsin, as serine
proteases. Elastases (metalloproteinases) have been isolated from the
pancreas of several species of fish (Yoshinaka et al., 1985).
3.2.3 Hepatopancreas proteinases

Autolysis of muscle protein in crustacean seafood (e.g. shrimp, lobster,
crab, krill) and molluscs (e.g. squid) occurs rapidly after harvest and has
been attributed to digestive proteases (Salem et al., 1970). The major
proteinases identified in these organisms are normally families of serine
proteases classified as trypsin-like (de Haen et al., 1975; Chen et al., 1979;
Galgani et al., 1984; Honjo et al., 1990) or as collagenases (Eisen et al.,
1973; Nip et al., 1985; Sakharov and Litvin, 1990). Recently, a new
cysteine proteinase with papain-like activity was shown to represent 80%
of the proteolytic activity in the lumen of the lobster hepatopancreas
(Laycock et al., 1989). Carboxypeptidases (Gates and Travis, 1973;
Zwilling et aI., 1979), cathepsin C (Hameed and Haard, 1985) and
cathepsin D-like proteases have also been purified from crustacean
(Kimoto et al., 1984; Gildberg, 1988) and molluscan (Lee and Pan, 1990)
hepatopancreas.
3.3 Proteinases contained in the muscle fibre
3.3.1 Lysosomal proteinases

In muscle tissue, catheptic and other hydrolytic enzymes are housed in two
species of a subcellular organelle or lysosome. One species is within the
muscle fibre and the other is in the extracellular matrix, originating from
macrophages and connective tissue cells (Canonico and Bird, 1970).
Lysosomes have been isolated from fish muscle (Bird et aI., 1969; Milanesi
and Bird, 1972; Reddi et al., 1972) although they appear to be more fragile
than mammalian Iysosomes and difficult to isolate intact (Whiting et aI.,
1975; Kim and Haard, 1992). A heterogeneity of enzyme distribution in
the lysosomal population of trout muscle has been reported (Milanesi and
Bird, 1972).
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of self-exaltation was to drive many of the minor leaders of his party from
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on Adams which was hopelessly to cripple, if not completely destroy, his
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Even more serious than his flamboyant egotism was his queer lack of
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[165] Only the most execrable taste can pardon the undignified writing of
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instead of ripening his judgment, warped it.’[167] His was a nature of eternal
youth, and in many respects the indiscretions of boyish exuberance cursed
him to the end.
If these personal weaknesses were to weaken him with the leaders of the
second rank, his unpopularity with the rank and file was to come from his
lack of sympathy for, and understanding of, the American spirit. No one
realized it more than he. In justice it must be said that he honestly tried to
suppress his doubts of America; but in moments of depression he burst forth
with expressions that bear the marks of long incubation. ‘Am I a fool—a
romantic Quixote—or is there a constitutional defect in the American mind?’
he wrote King. ‘Were it not for yourself and a few others I would adopt the
reveries of De Paux, as substantial truths, and could say with him that there
is something in our climate which belittles every animal, human or
brute.’[168] And toward the close of his life he wrote Morris: ‘Every day
proves to me more and more that this American world was not made for me.
You, friend Morris, are a native of this country, but by genius an exotic. You
mistake if you fancy that you are more of a favorite than myself, or that you
are in any sort upon a theatre suited to you.’[169] This touch of the exotic, of
which he himself was painfully conscious, was not lost upon his political
enemies. ‘Thus ignorant of the character of this nation, of Pennsylvania, and
of his own city and State of New York, was Alexander Hamilton,’ wrote
Adams.[170] But it was left for another to ‘discover the real secret of his
confusion as to the American character—he had never known the spirit, or
had the training, of the New England town meeting.[171] A marvelous
genius, he thought in terms of world politics at a time when America was
creating a new spirit and system of her own. It was not to weaken his work
as the creator of credit, but it was to dim his vision as an American leader.
If he possessed traits that made him thoroughly hated by some, he had

other qualities that bound his friends to him with bonds of steel. He
commanded affection because he was himself affectionate. His letters to his
wife were uniformly tender and playful. He was idolized by his children. His
comrades in the army loved him because he not only shared their hardships,
but at times helped them to necessities out of his own all but empty pockets.
He was sensitive to the sufferings of many refugees in Philadelphia and New
York, and he would often direct his wife to send money and delicacies to the
women and children.[172] We have many instances of his generosity, like his
attempt to spare Andre the humiliation of the scaffold, and his letter to Knox
protesting against the execution of British officers in retaliation for the
murder of an American.[173] Among the young French officers he was
idolized because of his merry disposition and the cleverness and brilliancy of
his conversation. While prone to hold aloof from the mass, he was a ‘good
fellow’ among those whom he considered his social equals. In social
assemblies of both sexes he fairly sparkled with boyish enthusiasm.[174] In
stag affairs, where he was immensely popular, we may be sure that he was
nothing of a prude. It is not of record that he often drank to excess, but like
most men of his time he loved his wine, and we have it on the best authority
that he sometimes took a wee bit too much.[175] On these convivial
occasions he could always be prevailed upon to sing his one and only song:
‘We’re going to war, and when we die
We’ll want a man of God near by,
So bring your Bible and follow the drum.’
His one serious weakness was an inordinate fondness for women which was
to involve him in the one serious scandal of his career. It was McHenry who
wrote to Pickering, another friend: ‘Far be it from me to attempt to palliate
his pleasures, the indulgence in which Mr. Hamilton himself publicly
lamented.’[176] It was Otis who wrote of his ‘liquorish flirtation’ with a
married woman at a fashionable dinner party.[177] It was Lodge who, in
touching on his overpowering passions, refers to his ‘relations, which had
an unenviable notoriety.’[178] It is Oliver who says that ‘his private
shortcomings cannot be denied,’[179] and that ‘in private life Hamilton was
not always vigilant.’[180] It is the historian of ‘The Republican Court’ who
records that ‘it is true that Hamilton was something of a roué.’[181] And it
was reserved for a descendant to remind us of the story of the alleged
relations with the celebrated Madame Jumel, who, in old age, made an
unsuccessful attempt to live with Aaron Burr,[182] and of the gossip, which
he discredits, that his relations with his sprightly sister-in-law, Mrs. Church,
were more tender than they should have been.[183] This same descendant,
writing with professional authority, explains these moral delinquencies on
the theory that, like other men of genius and great intelligence, he was
prone to ‘impulsively plunge into the underworld in obedience to some
strange promptings of their lower nature.’[184]
And yet, such are the strange inconsistencies of the temperamental—
nothing could have been more beautiful than his home life. His endearing
traits are evident in the passionate devotion of all who knew the Hamilton
of the hearth. If the ties that bound Angelica Church to him were not more
tender than they should have been, her letters indicate something akin to
love.[185] His wife, who must have suffered tortures over the confessions of
the Reynolds pamphlet, clung to him with a faith born perhaps of an
understanding of how much he must have resisted. If he sometimes broke
his vows, there can be no doubt that the shrine of his heart was at his hearth.
‘Colonel Beckwith tells me that our dear Hamilton writes too much and
takes no exercise, and grows fat,’ wrote Angelica Church to Mrs. Hamilton
from London. ‘I hate both the word and the thing, and I desire you to take
care of his health, and his good looks.’[186] Here we have the suggestion of
another frailty which makes all the more notable the intensity of his
sustained efforts and the magnitude of his achievements—the delicacy of
his health. The first, and possibly the last, medical service rendered by
McHenry on becoming a member of Washington’s military family was to
prescribe for Hamilton and make suggestions as to his diet. Early in the war
he who was never robust contracted a malarial infection from which he
suffered every summer throughout his life.[187] His correspondence is
sprinkled throughout with references to his health.[188] While in no sense an
invalid, the magnitude and multiplicity of his labors despite a chronic
physical disability measure the power of mind over matter and indicate
something of his unyielding will.
XI
In view of the sincere or simulated interest in religion shown by

Hamilton where political interests were involved, it would be interesting to
know just what he thought and felt. The records here are slight. During his
youth he passed through the period of religious exaltation not uncommon in
the average life. Not only was he attentive to public worship, but he prayed
fervently and with eloquence in the seclusion of his room.[189] About this
time he wrote a hymn, ‘A Soul Entering into Bliss,’ which is said to have
had some literary merit.[190] We hear no more concerning his religious
fervor for many years until he pretended, if he did not feel, an intense
indignation against the revolutionary reaction aimed at the church
establishment in France. He was shocked that ‘equal pains have been taken
to deprave the morals as to extinguish the religion of the country.’[191]
A few years more, and, with the fall of his party, he outlined to Bayard a
‘Plan of Conduct’ for Federalists with a view to its rehabilitation, and
proposed an association to be denominated ‘The Christian Constitutional
Society,’ having for its objects ‘the support of the Christian Religion’ and
‘the support of the Constitution.’[192] This hints strongly of the Old World
idea of the union of Church and State. In Connecticut the clergy had been
the shock troops of Federalism, and it is quite possible that the political
advantage of an alliance between the Church and his party appealed to
Hamilton.
At any rate, he was a member of no church. One of his descendants
assures us that ‘he was a man of earnest, simple faith, quite unemotional in
this respect, so far as display was concerned, but his belief was very
strong.’[193] Strong as it was, it never led him to the altar.
Leaving his idol’s death-bed, Oliver Wolcott wrote his wife that ‘Colonel
H. in late years expressed his conviction of the truths of the Christian
Religion, and his desire to receive the Sacrament—but no one of the clergy
who have yet been consulted will administer it.’[194] At length, life ebbing
away, a bishop consented after being earnestly solicited the second time.
Thus in his dying hour, Hamilton declared: ‘It has for some time past been
the wish of my heart, and it was my intention to take an early opportunity of
uniting myself to the church.’ The natural deduction from the meager
information we have is that his intensive political and professional activities
and consuming ambitions gave him little time to meditate on religion. He
certainly never gave it the consideration of his greatest political opponent
whom his party attacked as an enemy of Christianity. But he used the
Church, whenever possible, to advance his political views—and with effect.
Quite as problematical as his religious feeling was his attitude toward
Washington. It was the policy of the Federalists to capitalize politically the
popularity of the man of Mount Vernon, and they succeeded, as we shall
find, to a marked degree. Even so, some of Hamilton’s most partial
biographers[195] have commented on the absence of any deep affection
between the two, and Dr. Hamilton is not convincing with his observation
that his ancestor signed his letters to Washington, ‘Very affectionately.’[196]
As a matter of fact none of his letters to Washington denote real affection.
This would be more impressive, however, but for the singular absence of
the note of affection in all his political correspondence. But in one of his
letters we find the very opposite of either affection or admiration. This was
his letter to General Schuyler on the occasion of Hamilton’s withdrawal
from Washington’s military family, and it does not speak well for the
reliability of his son’s biography that he deliberately mutilated the letter. It
was in this that he wrote that he had found his chief ‘neither remarkable for
delicacy or good temper’ and complained of his ‘self love.’ Here we have
the confession that ‘for three years past I have felt no friendship for him and
have professed none.’[197]
In his letter to Lear, the secretary, when Washington died he probably
came perilously near to summing up his attitude in a sentence: ‘I have been
much indebted to the kindness of the General, and he was an Aegis very
essential to me.’ And then, the significant postscript: ‘In whose hands are
his papers gone? Our very confidential situation will not permit this to be a
point of indifference to me.’[198]
Such a man was Hamilton, a Colossus, brilliant, fascinating, daring, and

audacious—a constructive statesman of the highest order, a genius of the
first rank, with all the strength and the weaknesses of genius. Such the man
who sat down at the mahogany desk to write the documents that were to
give credit to a nation and a programme to a party.
CHAPTER III
HAMILTON IN THE SADDLE
T HERE was quite enough in the picture of the handsome, penniless

Hamilton, at the age of thirty-two, striding upon the national scene with
the air of a conqueror to undertake the solution of the problem on which
the existence of the young Republic depended, to appeal to the popular
imagination. The mystery and romance of his history, the dash in his
manner, the shimmer of his genius, interested all and fascinated many of his
contemporaries. The audacious gayety with which he faced his task
imparted a feeling of confidence to those who did not know, as many did,
just what was in his mind. He set to work with an enthusiasm that smacked
of inspiration, for it was a task to his taste.
With the startling effect of a magician at his tricks he created the
machinery of his complicated department, selected his assistants with
discrimination, trained them with meticulous care in their duties, outlined
his plans for revenue immediately required, and sat down with joy to the
preparation of his ‘Report on the Public Credit,’ which was to proclaim the
public faith and establish the Nation’s credit.
The mere presence of this youthful figure at the mahogany desk
commanded confidence. Here was a man who was primarily interested in
the rights of property, who believed in the sanctity of contracts and had the
courage of his convictions. Even as he was writing his ‘Report,’ he loomed
large as the man of the hour. His close associates foresaw the nature of his
recommendations. The mercantile and financial interests plumed
themselves upon a triumph. Within a month after his appointment a
contemporary rhymester put in verse the counting-room conception of the
man:
’...young Hamilton’s unshaken soul
The wayward hosts of anarchy control—
And while the Senate with his accents rung
A full conviction followed from his tongue.’[199]
His plans, given in confidence to some, were soon whispered among the
politicians and the merchants of New York, Philadelphia, and Boston, and
the market price of public securities in the cities rose fifty per cent two
months before Congress convened.
It was not until in early January that the ‘Report’ was read in House and
Senate. His wish to present it personally was denied, not by his political
enemies as his partial biographers contend, but by the supporters of his
plan.[200] In the galleries of the House eager speculators were closely
packed. They overflowed and filled the lobbies. Some were drawn by mere
curiosity, some were the original creditors who had waited long for their
reward, but the greater number were speculators, who, in anticipation of
such a recommendation, had bought freely of the skeptical holders at
ridiculously low prices. Not a few of these poured forth into Wall Street at
the conclusion with the exhilarating knowledge that a fortune was within
their grasp.
In the Senate the ‘Report’ was heard in secret and in ‘awful silence,’ for
the elder statesmen met behind doors closed and locked. Most of these
listened with approval, but the rheumatic Maclay, who had been puzzled for
some time with ‘the extraordinary rise in public securities,’ wrote that night
in his journal that Hamilton ‘recommends indiscriminate funding, and, in
the style of the British Minister, has sent down his bill.’ There were some
complaints that ‘a committee of speculators in certificates could not have
formed it more to their advantage.’ In truth, ‘it occasioned many serious
faces,’ and Maclay himself was ‘struck of a heap.’[201] But the prevalent
note was one of jubilation. In New York, enthusiasm in the coffee-houses;
in Boston, ‘great applause’;[202] in other commercial cities, Philadelphia,
Charleston, Baltimore, approbation, with reprobation for objections.[203]
All men of honor sympathized with the purpose of discharging the debt.
The repudiationists were among the ignorant and the vicious. Few at the
moment found fault with the funding system, though some would have
preferred a speedy liquidation through the sale of the public lands. Then—
suddenly—a low murmur of protest, followed by acrimonious attacks.
Thousands of the original creditors had been ‘swindled’ out of their
certificates for a song—were these, who rendered Revolutionary services,
to be taxed to ensure exorbitant profits to the speculators? Why should the
Federal Government assume the debts contracted by the separate States—
debts unevenly distributed? And what was the purpose of the proposal that
the Government should be prohibited from paying more than two per cent
of the principal a year? The indignation of the insurgents, at first a glimmer,
became a flame. The greater part of the certificates were in the hands of the
prosperous who had taken advantage of the necessities of the original
holders—Revolutionary soldiers, small farmers, hard-pressed country
merchants. The funding system would tax all the people to pay to the rich a
hundred cents on the dollar for evidence of debts that had cost them fifteen
and twenty. With the people taxed to pay the interest—it was proposed to
perpetuate the debt. Thus, for generations, perhaps, as many reasoned, the
Government would operate for the enrichment of the few already rich, and
the masses would pay the piper.
Had Hamilton been disposed to frankness, he would have smiled his
acknowledgment of the charge. One of his biographers has conceded that
through this system he hoped to ‘array property on the side of the
Government,’ by giving it a financial interest in the Government, and ‘to
assure to the property of the country a powerful influence upon the
Government.’[204] Having ‘been unable to introduce a class influence into
the Constitution by limiting the suffrage ... with a property qualification,’ he
hoped through his financial system to accomplish his purpose in another
way.[205]
There was nothing diabolical in the plan—coming from one who looked
upon the masses as lawless and unfit for self-government. His obsession
was a strong, stable government—and to sustain it he required the
interested devotion of the propertied class. The astonishing thing is that the
comparatively crude Maclay from the wilds of Pennsylvania and the
leather-lunged James Jackson from sparsely settled Georgia should have
caught the full significance of it all before it dawned on Jefferson and
Madison. The latter thought the ‘Report’ ‘well digested and illustrated,’ and
‘supported by very able reasoning,’ but after a while he, too, was depressed
with the injustice to the original creditors who ‘were most instrumental in
saving their country,’ and concluded there was something ‘radically wrong
in suffering those who rendered a bona fide consideration to lose seven
eighths of their dues, and those who had no particular merit toward their
country to gain seven or eight times as much as they advanced.’[206]
II
Meanwhile, speculation was manifesting itself with incredible audacity

and mendacity. The greater part of the securities in the hands of original
creditors were in the hands of soldiers, farmers, and merchants in the
remote interior. To most of these, they had come to mean so much worthless
paper. No telegraph could flash the news into the back country of Georgia
and North Carolina that Congress was about to legislate to par the promises
to pay. Weeks or months would pass before the proceedings in New York
could be known and comprehended by holders of the paper living in the
woods of the Carolinas or on the banks of the Savannah. Poor, and mostly
ignorant, they had no correspondents in the coffee-houses to write them of
the activities at Federal Hall; and even if they had, it required weeks for a
letter to reach them.
But members of Congress knew what to expect—for they were the
actors in the drama; and their friends, the capitalists and merchants of the
cities, knew—for they had been informed. The unscrupulous and
adventurous soldiers of fortune on the scene comprehended the opportunity
at a glance. The day after the ‘Report’ was read, the city buzzed with the
gossip of the speculators. One Senator, making calls in the congressional
circle, found it almost the sole topic of conversation. He heard that Robert
Morris of the Senate, who had been consulted by Hamilton, ‘must be deep
in it, for his partner ... had one contract for $40,000 worth.’ It was
whispered that ‘General Heister had brought over a sum of money for Mr.
Morris for this business.’ Senator Langdon, it was noted, was living with a
Mr. Hazard ‘who is an old and intimate friend of Mr. Morris,’ and he
admitted that he had followed buying certificates for some time past.’ ‘Ah,’
said the visiting Senator, ‘so you are one of the happy few who have been
let in on the secret’—and Mr. Hazard seemed abashed. It was understood
that Representative Fitzsimons of Philadelphia was likewise concerned in
the business.
Four days after the ‘Report’ was read, ‘expresses with very large sums of
money on their way to North Carolina for purposes of speculation in
certificates’ splashed and bumped over the wretched winter roads, the
drivers lashing the straining horses. Two fast-sailing vessels, chartered by a
member of Congress who had been an officer in the war, were ploughing
the waters southward on a similar mission—and this scandalous proceeding
was to be mentioned frequently in the subsequent debates. ‘I really fear,’
wrote Maclay, ‘the members of Congress are deeper in this business than
any others.’[207] Whether they were deeper or not, they were deep enough,
and numerous enough to hold the balance of power in the body that
legislated the certificates to par. These ranged from Robert Morris, the chief
legislative agent of Hamilton in the Senate, to Fisher Ames, who was his
most eloquent defender in the House.[208] In later years Jefferson was to
record in justice to Ames that his speculative activities had been greatly
exaggerated and that he had acted as an agent in the enterprises of his
Boston friends, Gore and Mason.[209]
So thoroughly did this money-madness take possession of the minds of
men that even the puritanic John Quincy Adams was to write his father,
without a homily, that by September of 1790, Christopher Gore, the richest
lawyer in Massachusetts, and one of the strongest Bay State members of
Hamilton’s machine, had ‘made an independent fortune in speculation in
the public funds’; and that other leaders of the bar[210] had ‘successfully
engaged in speculation’ by playing at ‘that hazardous game with moneys
deposited in their hands’ by clients at a distance. They took the chance of
becoming ‘masters of sums to an equal amount before they have been
called upon for payment.’[211] Maclay thought ‘there is no room to doubt
but that a connection is spread over the whole continent on this villainous
business.’[212] Everywhere men with capital—and a hint—were feverishly
pushing their advantage by preying on the ignorance of the poor. Thus,
paper held for years by the private soldiers was coaxed from them for five,
and even as low as two, shillings on the pound by speculators, including
leading members of Congress, who knew that provision for the redemption
of the paper had been made.
In all this, Hamilton had no part and no responsibility beyond having
made indiscreet disclosures of which his friends availed themselves, and
through buying and selling through his agents in New York and
Philadelphia for his brother-in-law.[213] Just how he viewed the scandalous
proceedings in the earlier stages we do not know. They were not without
defense from his supporters. The obsequious John Fenno took notice of the
gossip with a defense of speculation in the ‘Gazette.’ Were not moneyed
men ‘the props of the infant credit of the United States?’[214] The dark
insinuations of the gossips, the criticism of the ‘rabble,’ we may be sure
caused Hamilton no concern. Surveying the field at the beginning of the
battle, he must have been content. He saw the financiers, the commercial
interests of the large centers, including the speculators, enlisted under his
banner. The influential Society of the Cincinnati, composed of
Revolutionary officers, men of means who had been able to hold on to their
paper, gave dignity to his cause. With its compact organization in every
State, and its system of correspondence, it was an engine of tremendous
power. The social and intellectual circles were flying his flag. He looked
upon his work and called it good.
III
With the first discussion in the House, it was apparent that speculation
was to play a conspicuous part in the debates. The speculators packed the
galleries, overflowed into the lobby, causing the complacent Theodore
Sedgwick of Massachusetts, himself a speculator, to insist that the ‘ardent
expectations of the people on this subject want no other demonstration than
the numerous body of citizens assembled within these walls.’ The effect
was different on the pugnacious Jackson of Georgia. ‘Since this Report has
been read,’ he shouted, with a contemptuous glance at the eager gallery,
‘the spirit of speculation ... has arisen and been cherished by people who
had access to information the Report contained, that would have made a
Hastings blush to have been connected with, though long inured to preying
on the vitals of his fellow man. Three vessels, sir, have sailed within a
fortnight from this port freighted for speculation.’[215]
The unctuous Sedgwick was melting suavity. Speculation within
reasonable bounds was not bad, but action should be taken with all possible
speed to stop it; and the troublesome Jackson returned to the attack—this
time on New York City. He wished to God Congress had met in the woods
and out of the neighborhood of a populous town. The gallant veterans,
driven by economic necessity to the wilderness, were being robbed by these
speculators of the pittance a grateful country had bestowed. Since the
assumption of State debts was proposed, why not postpone action until the
various legislatures could express the sentiment of the States? ‘Then these
men may send out other vessels to countermand their former orders; and
perhaps we may yet save the distant inhabitants from being plundered by
these harpies.’[216]
This line of attack had not been anticipated, and Hamilton was not the
man to take anything for granted. His well-groomed figure was seen
moving nervously about the lobbies of Federal Hall, within a few days after
the commencement of the debate. One of his enemies observed that he
‘spent most of his time running from place to place among the
members.’[217] In the evenings he gathered his more influential supporters
about him at his home. At his table he brought his most seductive charms to
bear upon the doubting. Time was all-important and indefinite delay might
be fatal.
With the thunder of Jackson’s ugly charges reverberating through the
streets, taverns, coffee-houses, Hamilton was ‘moving heaven and earth for
his funding system.’ The commercial interests and the members of the
Cincinnati hastened to join the lobby, which began to seek out the wavering
or the doubtful in their lodging-houses. A fashionable minister found his
way to the quarters of Speaker Muhlenberg and Senator Maclay to extol the
policies of the dynamic young Secretary, and ‘argued as if he had been in
the pulpit.’ Time, too, for a redoubling of effort, for there were rumors that
Madison, the strongest man in the House, had been unpleasantly impressed
with the fast-sailing vessels and the expresses jolting over the roads
southward. A bitter attack had appeared in one of the papers which gossip
ascribed to the popular George Clinton.[218]
In the House—still harping were the foes on speculation, when with a
benevolent expression Sedgwick rose with saccharine urbanity to regret the
vice of speculation, and declare himself ‘totally disinterested,’ albeit he was
financially concerned. It was only his distress over speculation that
admonished him to speedy action to minimize the evil. It was really
unfortunate that so much heat had been engendered. After all, were not ‘a
great and respectable body of our citizens creditors of the United States?’ It
would be tragic were these animosities to create ‘factions among the
people.’
‘A danger there?’ bellowed Jackson, the incorrigible infant terrible. ‘Do
not gentlemen think there is some danger on the other side? Will there not
be grounds for uneasiness when the soldier and the meritorious citizen are
called upon to pay the speculator more than ten times the amount they ever
received from him for their securities?’[219]
Meanwhile the fight was spreading from Federal Hall to the newspapers
where congressional courtesy imposed no restrictions on the temper.
Sinister stories were finding their way into print. ‘Several officials in
conjunction with Robert Morris and wealthy contractors “were” at the
bottom of this new arrangement.’ If it succeeded, Robert Morris would
benefit $18,000,000, Jeremiah Wadsworth would profit $9,000,000 and
Governor George Clinton would make $5,000,000.[220]
It was under these conditions, with the speculators packing the galleries,
with the lobbyists, legitimate and illegitimate, buzzing through the
corridors, with the most amazing rumors floating about the streets, that
James Madison, who had remained silent heretofore, rose in a crowded
House to fire the first fun in the Jeffersonian war on the financial policies of
Alexander Hamilton.
IV
Here was a man at whom the Federalist leaders dare not sneer. A
stranger, looking down from the gallery, would have been at a loss to
understand the deference with which members hung upon his words. His
personal appearance was disappointing. The short little man dressed in
sober black, with a bald head, and a little protuberant in front, whose lower
limbs were slight and weak,[221] was surely not meant to ride on the
whirlwind and direct the storm. The impression of physical weakness he
conveyed did belie the fact. In the mild blue eyes there was much to suggest
the meditative philosopher, nothing to hint of the fighter. His voice was so
weak that even in the cozy little chamber he could scarcely be heard.[222]
He spoke in low tones, without gesture or excitement, almost like a man
communing with himself in the seclusion of his closet. And yet he
commanded a hearing vouchsafed to few. It was the triumph of character.
Here, too, was a man with a background second to none in the infant
Republic. An ailing body had obsessed him in youth with the premonition
of an early death, and, feeling the futility of entering on any pursuit, he had
sought consolation in his books. He not only consumed, he assimilated. He
not only read, he thought. Thus he became something more than a learned
man—he developed into a political philosopher ‘worthy to rank with
Montesquieu and Locke.’[223] At the time he rose to propose an amendment
to Hamilton’s plan there was not a man in America who was his peer in the
knowledge of constitutional law or history. Nor was there a man, either,
whose support Hamilton more eagerly coveted. Even the jealous Ames
conceded him to be ‘our first man,’ consoling himself for the concession
with the comment that ‘I think him too much of a book politician and too
timid in his politics,’ and that ‘he speaks decently as to manner and no
more.’[224]
But the ill-natured jealousy of the more ornamental Ames failed to take
account, as most of his colleagues did, of the important practical use to
which he had put his knowledge of the battles he had fought and the
victories he had won. No one in either branch of Congress or at the head of
any of the departments had approached his services in the framing of the
Constitution. It was his genius that conceived the Virginia plan which
became the basis of the agreement. At many critical junctures his speeches
had dissipated the gathering darkness with their light. His pen, unknown to
many at the time, had recorded the story of the Convention. His
contributions to ‘The Federalist’ had been quite as important, if not so
numerous, as those of Hamilton; and the fight he waged in the Virginia
Convention for ratification was quite as Titanic and conclusive as that of
Hamilton in New York, but with this difference—Hamilton was confronted
by Melancthon Smith, while Madison had to cross swords with Patrick
Henry, with the powerful George Mason and the accomplished Pendleton.
He was not an orator of frills and fancies, magnetic and dramatic,
appealing to the passions and emotions, but he was formidable in debate. In
the speeches of none of his contemporaries is found such erudition, more
driving logic, such tact and moderation of statement, or greater nobility of
sentiment, fairness, justice. If they are a bit heavy in their sobriety, the
occasion called for something remote from theatrical frivolity. His grace
was in his reasoning, not his rhetoric—and yet his style would have given
him a foremost place at Saint Stephen’s.
It is not surprising that such a man should not have been a favorite with
the crowd. There was a diffidence in his manner, a formality and precision
in his method, a quiet dignity in his bearing that discouraged familiarity. He
was too absorbed in his work to fit in with the social festivity of his time.
Only at his own table and among his intimates did he appear in the rôle of
‘an incessant humorist’ and ‘keep the table in roars of laughter over his
stories and his whimsical way of telling them.’[225] Even his letters read
like state papers. But there were a few, greater than Ames, who appreciated
him. These were the three most important personages of his time—
Washington, Hamilton, and Jefferson.
Washington consulted him and made use of his pen. Hamilton cultivated
him. Jefferson loved him as a son. His relations with the latter were no less
than beautiful. Through many years they constantly interchanged visits,
corresponded regularly, and traveled together whenever possible. A
strikingly incongruous pair they must have seemed as they plodded along
country roads together, or rode to and from Philadelphia together in
Jefferson’s carriage—the tall, thin, loose-jointed, and powerful master of
Monticello, and the short, frail, bald-headed Madison. But the incongruity
was in their physical appearance only, for they had much in common—a
common sweetness of disposition, a common code of political principles
and morals, a common liberality of views, and a common passion for
knowledge. The older man paid tribute to his protégé’s qualities long after
both had passed from active public life: his ‘habits of self-possession which
placed at ready command the rich resources of his luminous and
discriminating mind’; his language ‘soothing always the feelings of his
adversaries by civilities and softness of expression’; his ‘pure and spotless
virtue which no calumny has ever attempted to sully’—all qualities that
made him a congenial companion for the philosopher who shared them in a
large degree.[226] Observing Jefferson’s happiness at the inauguration of his
successor, a lady who knew them both intimately wrote what all who knew
them felt: ‘I do believe father never loved a son more than he loves Mr.
Madison.’[227] But when Madison rose that cold February day to make his
first attack on Hamilton’s programme, he acted on his own volition and
without consultation with the man who was to be his chief.
The character of Madison’s speech in favor of discrimination between

the original holders and the purchasers of securities was not so open to
attack as that of the impulsive and loose-thinking Jackson. He began in a
manner to conciliate his hearers, matching Hamilton in his insistence on the
sanctity of the debt and the necessity for its discharge. The question is—to
whom is the money due? There could be no doubt in the case of the original
holders who had not alienated their securities. The only rival pretensions
were those of the original holders who had assigned and the present holders
of the assignments.
‘The former may appeal to justice,’ he said, ‘because the value of the
money, the service or the property advanced by them has never been really
paid to them. They may appeal to good faith, because the value stipulated
and expected is not satisfied by the steps taken by the Government. The
certificates put in the hands of the creditors, on closing their settlements
with the public, were of less value than was acknowledged to be due; they
may be considered as having been forced on the receivers. They cannot
therefore be adjudged an extinguishment of the debt. They may appeal to
the motives for establishing public credit, for which justice and faith form
the natural foundation. They may appeal to humanity for the sufferings of
the military part of the creditors who never can be forgotten while sympathy
is an American virtue.’
Admitting that the purchaser also had a claim, he proposed a plan
designed, as he thought, to do justice to both—to pay the original holder in
full, and, where there had been an assignment, the assignee to receive the
highest market value and the original holder whatever remained over.[228]
The plan spread consternation. At the Knoxes’ dinner table that night,
where members of Congress and diplomats were gathered, it was almost the
sole topic of conversation. In the coffee-houses, where the speculators
gathered about their mugs, Madison was denounced as a dreamer and an
enemy of public faith. The more cautious regretted the insurmountable
difficulties of the scheme. This was felt by Madison as the one legitimate
argument in opposition, and writing Jefferson three days later he made the
admission with the suggestion that ‘they might be removed by one half the
exertions that will be used to collect and color them.’[229] It was not until
four days later that the Hamiltonian leaders attacked the plan with their
heavy artillery. One by one they rushed to the assault. ‘It is not pretended,’
cried Sedgwick, ‘that any fraud or imposition has been practiced’—which is
precisely what was charged. If the original holders lost, it was their own
fault. It was too bad. He really sympathized with their misfortunes. But
business was business. There was ‘no fraud on the part of the holder,’
echoed Laurance of New York—who knew that the town was humming
with the charge. At any rate, ‘the general opinion of men of property is in
favor of it.’ No public bodies like Chambers of Commerce were against the
Hamilton plan. As for ‘the people’—newspapers and pamphlets could not
be taken as expressive of public opinion. William Smith of Charleston had
heard few advocates of discrimination ‘in society.’ As for the newspapers,
they appeared on both sides. And why so much sympathy with the original
holders?
It was reserved for Ames, whose friend Gore was getting rich on
speculation, to take a stouter stand. Why should not ‘the seller who sold for
a trifle be taxed to pay the purchaser?’ he asked. ‘He certainly ought to fare
as other citizens do. If he has property, then the plea of necessity is
destroyed; if he has none, then his taxes will be a mere trifle.’ And public
opinion against it? Then ‘all the more duty on Government to protect right
when it may happen to be unpopular; that is what Government is framed to
do.’ Away with maudlin sentiment—it was not the function of the State to
‘rob on the highway to exercise charity.’[230]
Meanwhile the commercial organizations of the larger towns were
summoned to the field against discrimination, and they responded—even in
Richmond. ‘It is the natural language of the towns,’ wrote Madison, ‘and
decides nothing.’[231] As the debate proceeded, Wall Street swarmed with
the curious who could not get into the House where the speculators packed
the galleries, and lined up deep behind the railing in the rear of the chamber.
Petitions began to pour in. Passions rose. ‘I do not believe the crowd in the
gallery consists of original holders,’ shouted one speaker with a
contemptuous glance at the covetous group bending over the railing.[232]
Soldiers! ‘Poor soldiers!’ sneered Wadsworth—he who had sent the two
fast-sailing vessels to the South—‘I am tired of hearing about the poor
soldiers. Perhaps soldiers were never better paid in any part of the
country.’[233]
Two days later, Madison returned to the attack in a speech unusually
spirited for him. Only when he had parted with his self-respect ‘could he
admit that America ought to erect the monuments of her gratitude, not to
those who saved her liberties, but to those who had enriched themselves on
her funds.’ It was his last effort. He had spent himself to the utmost. A
spectator entering the House late in the day found him ‘rather jaded.’[234]
He had incurred the hate of the Hamiltonians without having consolidated
all the opposition in favor of his plan.
Three days later—it was Sunday—that extreme democrat Senator
Maclay, who was indifferent to Madison’s plan because opposed to funding
altogether, sat down in his boarding-house and framed a plan of his own
looking to the extinguishment of the debt through the sale of public lands.
Having satisfied himself, he went forth in search of Thomas Scott, his
colleague. But ‘shame to tell it—he a man in years and burdened with
complaints—had lodged out and was not home yet.’ Pity that ‘a good head
should be led astray by the inordinate lust of its concomitant parts.’ At
length the old ‘roué’ was found, and he urged that it be submitted to
Madison at once.
The next day found Maclay indignantly chafing at Madison’s lodging-
house because it was ‘a long time’ before he appeared. As the radical from
Pennsylvania read his plan, it seemed to him that Madison ‘attended to no
one word, being so much absorbed in his own ideas.’ Maclay handed him
the paper, and Madison handed it back without glancing at it. Alas, thought
the radical, ‘his pride seems of the kind that repels all
communications.’[235] It was not an easy task to organize the forces of
Democracy.
The next day Madison’s plan was voted down. It was found long
afterward that of the sixty-four members of the House, twenty-nine were
security-holders.
VI
One thing, however, had been accomplished—the public interest had

been awakened. The tongue of criticism had been loosened. The man in the
street began to hold forth. It was all beyond him—as problems of finance
were beyond Madison himself; but he could understand that a policy had
been adopted that would be advantageous to the rich, profitable to the
speculator, and mean loss to the common soldier. In the commercial centers
of the cities Madison became anathema. Young Adams reported to his
father that in Boston ‘Mr. Madison’s reputation has suffered from his
conduct,’ albeit so respectable a character as Judge Dana had adopted
Madison’s views.[236] The immediate reaction through letters to the papers
was so bitter that Fenno was moved to a homily under the caption, ‘Honor
Your Rulers,’ in which he pointed to such outrageous derelictions as
expressions of doubt concerning the propriety of the proceedings of
Congress.[237] These expressions had gone far beyond a mere questioning
of the wisdom of Congress. ‘A War Worn Soldier’ thought it ‘happy there is
a Madison who fearless of the blood suckers will step forward and boldly
vindicate the rights of the widows and orphans, the original creditors and
the war worn soldier.’[238] Another ‘Real Soldier’ described ‘the poor
emaciated soldier, hungry and naked, in many instances now wandering
from one extreme part of the country to another.... But thank God there lives
a Madison to propose justice....’[239] An uglier and more pointed note was
struck by ‘A Farmer’ in Pennsylvania. ‘Would it not be a good regulation,’
he wrote, ‘to oblige every member of Congress ... to lay his hand on his
heart and to declare that he is no speculator; and that he did not come
forward to claim for himself the price of the blood or the limb or the life of
the poor soldier?’[240] Another wrote to ‘gentlemen who by superior wealth
have monopolized the public securities’ that if honor and public faith called
for the maintenance of the paper at par then, there was more occasion for it
‘when they were in the hands of those poor people to whom they were
justly due, who had implicitly pinned their faith on your sheaves.’[241] ‘An
Old Soldier’ recalled Washington’s pledge to see justice done the common
soldier. ‘Ample means are said to be now about to be provided, not for their
relief, but to enable eight or nine hundred per cent gain on the purchase
money of the speculator.’[242] ‘Ah well,’ wrote ‘A Citizen’ of Boston,
‘Madison, Jackson and others in favor of discrimination in funding the
public debt have probably immortalized their memories.’[243]
Their letters probably reflect the talk among the workers on the wharves,
the pioneers on the fringe of the forests, the gossips of the taverns. Rightly
or wrongly, a spirit of resentment had been aroused—a feeling in the breast
of many that their interests were being subordinated by the Government.
This sentiment was to grow and to increase the trouble of Hamilton in the
next step toward the adoption of his funding system.
VII

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Seafoods Chemistry Processing

Technology and Quality Fereidoon

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Chemistry and technology of emulsion polymerisation

Handbook of Banana Production Postharvest Science

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SPRINGER-SCIENCE+BUSINESS MEDIA, B.Y.

ISBN 978-1-4613-5913-5 ISBN 978-1-4615-2181-5 (eBook)

00 Printed on permanent acid-free text paper, manufactured in accordance

Fereidoon Shahidi and J. Richard Botta

The 8th World Congress of Food Science and Technology, held in

R. G. Ackman Canadian Institute of Fisheries Technology, Technical

R. C. Lindsay Department of Food Science, University of Wisconsin-

1 The chemistry, processing technology and quality

Part 1 The chemistry of seafood components

2 Seafood proteins and preparation of protein concentrates 3

3 Protein hydrolysis in seafoods 10

4.1 Introduction: the paradox of seafood in health and nutrition 34

4.2.1 Marine fish 35

5 Oxidation of lipids in seafoods 49

7 Flavour of shellfish and kamaboko flavorants 85

8 Taste-active components of seafoods with special reference to

8.1 What are umami and umami substances? 115

Part 2 Quality of seafoods

9 Freshness quality of seafoods: a review 140

9.1 Introduction 140

10 Preservation of seafood quality 168

10.1 Introduction 168

11 Microbiological quality of seafoods: viruses, bacteria and

11.1 Introduction 196

11.2.1 Hepatitis A 199

12 Microbiological quality of seafoods: marine toxins 220

12.1 Introduction 220

13 Sensory assessment of quality in fish and seafoods 233

13.1 Introduction 233

Part 3 Further processing of raw material

14 Surimi processing from lean fish 263

14.1 Introduction 263

14.2.2 Factors affecting the surimi quality and yield 267

15 Surimi processing from fatty fish 288

15.1 Introduction 288

16 Seafood processing by-products 320

16.1 Introduction 320

The world's annual catch of fish and marine invertebrates is approximately

F. Shahidi et al. (eds.), Seafoods: Chemistry, Processing Technology and Quality

Seafoods namely fish, crustaceans and molluscs are important sources of

2.2 Sarcoplasmic proteins

Sarcoplasmic proteins, referred to as 'myogen', are soluble in the muscle

F. Shahidi et al. (eds.), Seafoods: Chemistry, Processing Technology and Quality

haemoglobin and cytochrome C in dark muscles may reverse this trend.

2.3 Myofibrillar proteins

The largest amount of myofibrillar proteins present in muscles of aquatic

2.4 Stroma proteins

The residue after extraction of sarcoplasmic and myofibrillar proteins is

2.5 Non-protein nitrogenous (NPN) compounds

The content of NPN compounds in seafood depends primarily on the

"Adapted from Finne (1992).

Approximately 85% of NPN compounds in muscle tissues of marine fish

2.7 Protein concentrates/hydrolysates