NEWCASTLE DISEASE

A REVIEW
1926-1964

J . E . Lancaster
 ALBERT R. MANN
LIBRARY

New York State Colleges

OF
Agriculture and Home Economics

AT

Cornell University
NEWCASTLE DISEASE
A REVIEW OF SOME OF THE LITERATURE
PUBLISHED BETWEEN 1926 AND 1964

John E. Lancaster

1966
Health of Animals Branch

CANADA DEPARTMENT OF AGRICULTURE

Monograph No. 3
 373333
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1966

ii
 FOREWORD

Newcastle disease has probably received more attention from poultrymen and
research workers than any other respiratory disease of poultry. It is worldwide, and
affects fowl that are reared in thousands by modern intensive methods as well as
individual fowl that seek their own food in rural areas.
Control of the disease is essential if the poultry industry of many countries is to
flourish. In this book Dr. Lancaster reviews progress that has been made to date towards
accomplishing this goal. It is not a book that is likely to be of immediate interest to most
poultrymen, though it should benefit them indirectly. It is a research tool that should
be valuable to anyone studying Newcastle disease or its virus.

K. F. WELLS
Veterinary Director General
Ottawa, 1965

iii
 PREFACE

This review is based on articles I have written over the past several years for
publication in a number of different journals. In bringing all these articles together into
one volume I have expanded on them and also added a number of tables and diagrams.
The material covers many aspects of research on Newcastle disease and its virus, but
I have made no attempt to discuss every report that has been published on these subjects.
I have read most of the reports referred to in the original, and where this has not been
possible I have used abstracts published by the Commonwealth Agricultural Bureaux in
The Veterinary Bulletin.
Mention must be made here of three sources of information on Newcastle disease
which have become available since completion of the main work on this book. In 1963,
an international symposium on "Newcastle Disease Virus — An Evolving Pathogen"
was held in Madison, Wisconsin, U.S.A., and the proceedings, edited by Dr. R. P.
Hanson of the Study Center for Newcastle Disease at the University of Wisconsin, have
now been published. Dr. Hanson has also compiled a subject bibliography on Newcastle
disease for the period 1926 to 1962. This bibliography is in manuscript form. Dr. V. N.
Syurin of the Soviet Union has published a book in Russian (Psevdochuma ptits
[Newcastle Disease of Poultry] Moscow, 1963) in which he makes a detailed study of
Russian and foreign literature on the characteristics of viruses, with special reference to
Newcastle disease virus. Dr. Syurin's book includes a more detailed review of Russian
work on Newcastle disease than is given here.
I hope that the present volume, used in conjunction with these other works, will serve
as a useful reference to the literature on Newcastle disease published between 1926
and 1964.
JOHN E. LANCASTER
Ottawa, 1965

iv
 ACKNOWLEDGMENTS

The publication of this review would not have been possible without the kind help of
many individuals and organizations. Grateful acknowledgment is made to:
The Commonwealth Agricultural Bureaux for permission to reproduce the con
tents of three reviews by the author published in The Veterinary Bulletin (Vol. 33,
pp. 221, 279 and 347, and Vol. 34, p. 57);
Dr. R. A. Bankowski of the University of California at Davis, Dr. G. L. Bannister
of the Animal Diseases Research Institute, Canada Department of Agriculture, Dr.
A. P. Waterson of St. Thomas's Hospital, London, and the Controller of Her
Majesty's Stationery Office, London, for providing photographs; Dr. A. S. Greig
for modifying one of the drawings; and
For permission to reproduce drawings and tabular material, the authors whose
names are given in captions; the publishers of the following Journals: Acta
Veterinaria Hungarica (Figure 19), American Journal of Hygiene (Figure 6),
American Journal of Veterinary Research (Figure 15 and Table 7), Animal Health
Yearbook (Figure 5), Annals of the New York Academy of Sciences (Table 20),
Avian Diseases (Figure 7), Bacteriological Reviews (Figure 1), British Veterinary
Journal (Figure 23), Bulletin de l'Office International des Epizooties (Table 2),
Bulletin of the Interafrican Bureau for Animal Health (Figure 3), Canadian
Journal of Comparative Medicine and Veterinary Science (Table 13), Cornell
Veterinarian (Figure 13), Journal of Comparative Pathology and Therapeutics
(Figures 11 and 18 and Tables 11 and 23), Journal of Experimental Medicine
(Figure 14), Journal of Immunology (Figures 21 and 22), The Veterinary Bulletin
(Tables 14, 15, 16 and 18), The Veterinarian (Table 10), The Veterinary Record
(Figure 17 and Table 12), World's Poultry Science Journal (Table 5); and the
American Veterinary Medical Association (Tables 3 and 22), the Controller of
Her Majesty's Stationery Office, London (Table 1 ) , the Eighth World's Poultry
Congress (Table 4), the Italian Society of Veterinary Science (Table 21), the
National Academy of Sciences, Washington, D.C. (Table 8), and the University
of Wisconsin Press, Madison, Wisconsin (Figure 20) .
The author's sincere thanks also go to personnel of the Canada Department of
Agriculture: to the staff of the Library for obtaining articles and reports for review
purposes; to the Art Section of the Information Division and the bio-Graphic Unit of
the Research Branch for preparing text figures; and, especially, to the Editorial Unit of
the Information Division for suggesting improvements to the original manuscript and
for attending to the many details of publication.

v
 CONTENTS

Page Page
Summaries in English and French .... 1 Asymptomatic infections 33
Definition of Newcastle Disease 13 Transport of live poultry 33
Nomenclature 13 Poultry markets 33
Properties of the virus 13 Laying trials 34
Illegal movement 34

PART I — SPREAD OF THE DISEASE Poultry carcasses and offal 34

Spread by human agency 36
Geographic Distribution 17
Human infection 36
Early reports of diseases Warm-blooded animals 38
resembling Newcastle disease 17
Cold-blooded animals 40
First outbreaks of Newcastle
Inanimate causes 41
disease in England 18
Chicken houses, crates and
Spread through Southeast Asia .. 18
brooders 41
India 18
Wind 42
Philippines 18
Aerosols 43
Malaya 18
Water 44
Haiti and Madagascar 19
Poultry vaccines as a means of
United States 19
spread of Newcastle disease 44
Canada 19
Africa 19
Spread through Europe 19
PART II — DIAGNOSIS
Great Britain 22 General Characteristics of the
Ireland 22 Disease 45
Austria 22 Incubation period 45
World distribution in 1962 22 Breed susceptibility 46
Modes of Spread 23 Genetic differences in resistance 46
Wild birds 25 Sex differences in susceptibility 46
Infections in birds in zoological Age susceptibility 46
gardens 25 Effect of season of year 46
Introduction of virus by Climatic influences 47
imported birds 27 Routes of infection 48
Introduction of virus by Effect of the virus on avian
migratory birds 27 physiology 48
Excretion of virus in faeces 27 The Disease in Chickens 48
Game birds 28 Velogenic form 48
Pigeons and doves 28 Mesogenic form 53
Ducks and geese 29 Lentogenic form 56
Turkeys 29 Asymptomatic form 57
Chicken eggs 29 The Disease in Turkeys 57
Young chicks 30 The Disease in Ducks and Geese 58
Growing and adult chickens 31 The Disease in Game Birds 58
Excretion of virus from Diagnosis by Serological Methods 59
respiratory system 31 Haemagglutination (HA Test) .. 59
Excretion of virus in faeces 31 Haemagglutination-inhibition
Reservoir of virus in aqueous (HI Test) 61
humour 33 Other haemagglutinating agents 63

vii
 Page Page
Haemolysis 63 Control with Hyperimmune Serum
Intradermal inoculation 64 Combined with Virus 88
Fluorescent antibody 64 Control with Antibiotics and Other
Serum electrophoresis 64 Medicinal Agents 88
Serum or virus neutralization Control by Vaccination 89

(SN Test) 64 Antigenic plurality 89

Complement-fixation tests 66 Procedures for evaluating
Precipitation test 66 immunity 89

Diagnosis by Virus Isolation 67 Factors that influence

Distribution of virus in the body . 67 development of immunity 92
Embryonating eggs 67 Passive immunity 92
Preparation of inoculum 67 Age at time of vaccination 93
Route of inoculation 68 Virus titre of the vaccine 93
Temperature of egg incubation 69 Viral interference 93
Blind passages 69 Differences between individual
Embryonic mortality 69 birds 95
Lesions in embryos 69 Effect of vaccination on
Distribution of virus in susceptibility to another
embryos 70 disease 95
De-embryonated eggs 70 Other factors that affect
Tissue cultures 70 immunity 95
Mixed virus infections 72 Types of Vaccines — Administration
Diagnosis by Inoculation 73 and Effectiveness 95
Chickens 73 Live and inactivated vaccines
Pigeons 73 compared 95
Ducks 74 Live vaccines 97
Laboratory mammals 74 Lentogenic strains 97

Diagnosis by Challenge Exposure .... 74

Bl strain 98

Identification of Newcastle Disease

Strain F 100
LaSota strain 102
Virus 74
Other lentogenic strains 102
Differential Diagnosis 75
Mesogenic strains 102
Viral diseases 75
Komarov (or Haifa) strain 102
Bacterial diseases 78
Mukteswar strain 103
Nutritional deficiencies 78
Strains originating in the
Toxic drugs and plants 78
United States 104
Avian respiratory diseases 78
Hertfordshire (or Herts)
strain 105
PART III — CONTROL MEASURES Other mesogenic strains 106
Extract of tobacco mosaic virus 106
Control by Slaughter 80
Tissue culture vaccines 106
Effectiveness in various countries 80
Inactivated vaccines 106
Action of Chemicals on Newcastle
Inactivation by beta-
Disease Virus 84
propiolactone 106
Sterilization of Atmospheres Inactivation by formalin 110
Contaminated with Newcastle Inactivation by crystal violet .... 113
Disease Virus 87 Inactivation by other chemicals 113
Control with Hyperimmune Serum .. 87 Inactivation by heat 113

viii
 Page Page
Inactivation by ultraviolet Preparation of vaccines from
irradiation 113 embryonating eggs 125
Inactivation by ultrasonic Egg transmitted diseases 125
treatment 114 Influenceof parental immunity
Vaccines inactivated by different on preparationof vaccines .... 125
methods compared 114 Storage of virus material 127
"Incomplete" Newcastle disease Use of extra-embryonic fluids
vaccine 114 with or without suspensions
Combined vaccines 114 of embryonic tissues 127
Preparation of fresh and
lyophilized vaccine 129
PART IV — VIRUS PROPAGATION Preparation of inactivated vaccine 129
AND VACCINE PRODUCTION Preparation of Newcastle disease
Virus Propagation Methods 116 vaccines from virus propagated
Propagation in Eggs 116 in tissue culture 130
Propagation in avian hosts 117 Storage and Transportation of
Propagation in mammalian hosts 118 Vaccines 130
Propagation in tissue cultures .. 119 Testing and Standardization of
Propagation in yeast cells 124 Vaccines 133
Preparation of Vaccines 125 REFERENCES 135

ILLUSTRATIONS
Figure Page Figure Page
1. Scheme of the structure of 10. The velogenic form of Newcastle
Newcastle disease virus 14 disease — lesions in intestine .... 51
2. Electron micrograph of 1 1. The velogenic form of Newcastle
Newcastle disease virus 15 disease — distribution of lesions 52
3. Newcastle disease in Africa
12. The mesogenic form of
south of the Sahara 20 — nervous
Newcastle disease
4. Infection density of Newcastle 54
symptoms
disease and fowl plague in
13. A comparison of the results of
Europe, 1940-1955 21
HI and SN tests 65
5. Incidence of Newcastle disease
14. Tissue distribution of Newcastle
in Europe in 1962 24
disease virus after intramuscular
6. Suggested scheme for the
vaccination of 1 0-week-old
pathogenesis of Newcastle
chickens 68
disease virus 32
15. Selective infection by Newcastle
7. Mortality due to vaccination
disease virus 71
with a wing-web vaccine 47
8. The velogenic form of Newcastle 16. Newcastle disease in the pigeon . 73

disease — symptoms of paralysis 49 17. Number of outbreaks of

9. The velogenic form of Newcastle Newcastle disease in Great
disease — lesions in Britain and Lancashire,
proventriculus 50 1946-1961 82

ix
Figure Page Figure Page
18. Immune response following non-immune embryos 127
with a BPL vaccine..
vaccination 108 22. Amount of NDV (CGI 79) in
19. Newcastle disease virus in avian allantoic fluid, blood and
tissue culture 121 embryo tissues following
20. Cytopathogenicity of Newcastle allantoic sac inoculation of
disease virus 123 non-immune embryos 128
21. Amount of NDV (Bl strain) in 23. Viability of Komarov strain
allantoic fluid following Newcastle disease lyophilized
allantoic sac inoculation of vaccine 132

TABLES
Page Page
1. Outbreaks of fowl pest in Great 14. Duration of immunity following
Britain, 1954-1961 22 vaccination of chicks up to five
2. Incidence and control of weeks of age with Bl vaccine
Newcastle disease, 1962 23 virus 99
3. Wild birds susceptible 15. Duration of immunity following
experimentally to Newcastle vaccination of chicks up to five
disease virus and methods of weeks of age with Strain F
evaluation 26 vaccine virus 101
4. Probable origin of the first 542 16. Duration of immunity following
outbreaks of Newcastle disease
initial vaccination of chickens
in England and Wales during
with beta-propiolactone
1947 34 inactivated vaccine 107
5. Newcastle disease virus isolated
17. Comparison of effect of
from poultry carcasses imported
Newcastle disease on egg
into England in 1949 35
production in unvaccinated and
6. Reports of duration of viability
BPL-vaccinated flocks 109
of Newcastle disease virus in a
1 8. Duration of immunity following
variety of tissues 36
initial vaccination of chickens
7. Duration of viability of
Newcastle 42
with formalin-inactivated virus .. 1ll
disease virus
8. Effect of temperature on 19. Tissues used for in vitro
infectivity of Newcastle disease propagation of Newcastle disease
virus 43 virus 1 19

9. Characteristics of classical fowl 20. Diseases of birds transmitted

plague and typical Newcastle through eggs 126
disease compared 76
21. Survival of Newcastle disease
10. Avian respiratory diseases —
Strain F virus at different
some characteristics 79
temperatures 131
1 1. Effect of disinfectants on
Newcastle disease virus 85
22. Summary of test requirements
12. Action of formalin on Newcastle for live and modified live virus
disease virus Newcastle disease vaccines 133
86
13. Effect of intranasal Newcastle 23. Summation of 32 potency tests
disease vaccination on chicks on one batch of inactivated
from immune parents 91 Newcastle disease vaccine 134

x
 SUMMARY SOMMAIRE
Maladie de Newcastle
Pseudo-peste aviaire

Spread of the Disease Propagation de la Maladie

Acute diseases of poultry that resemble Depuis le début du dix-huitième siècle,
Newcastle disease have been recorded on enregistre des maladies aiguës de la
since early in the eighteenth century, but volaille ressemblant à la maladie de New
it is generally accepted that Newcastle castle. Cependant ce n'est qu'en 1926 sur
disease, as now recognized, first appeared l'île de Java, en Indonésie, que s'est mani
in epidemic form on the island of Java in festée la première épidémie de maladie de
Indonesia in 1926. From this origin, the Newcastle telle qu'on la connaît aujour
disease spread with a rapidity not previous d'hui. De ce foyer, la maladie s'est répan
ly recorded among diseases of poultry. due avec une rapidité inconnue auparavant
Much of the early global spread was dans les annales des maladies aviaires.
associated with coastal ports or towns, and La maladie tire son nom de la ville por
the disease takes its name from the coastal tuaire de Newcastle-on-Tyne, en Angle
town of Newcastle-on-Tyne in England. terre. Elle a été propagée surtout en raison
One of the exceptions to this general pat de la manutention des volailles dans les
tern was the appearance of the disease in ports et les villes côtières, mais il y a eu,
1927 at the small town of Ranikhet in the aussi, des manifestations isolées, par
Kumaon Hills of India. Another event of exemple dans la petite ville de Raniket,
considerable importance in the history of dans les collines de Kumaon, aux Indes en
the disease was the appearance in Cali 1927. Autre événement important dans
fornia, around 1935, of a mild respiratory l'histoire de la maladie: une affection res
disease termed "avian pneumoencephal- piratoire peu sérieuse, appelée pneumoen-
itis". Several years elapsed before this céphalite aviaire se produisit en Californie,
disease was recognized as a form of New vers 1935; il s'est écoulé plusieurs années
castle disease. avant qu'elle soit reconnue comme une
Many of the early outbreaks of New forme de maladie de Newcastle.
castle disease were associated with deaths Au début, les manifestations de la maladie
of free-flying birds in the vicinity. How de Newcastle ont souvent été marquées
ever, it was not until about 1 950 that the par la mort d'oiseaux volant en liberté
virus was recovered with any regularity dans les envirous. Toutefois, ce n'est qu'en
from free-flying birds such as pigeons 1950 qu'on a constaté la présence régu
(Columba livid), starlings (Sturnus vul lière du virus chez des pigeons (Columba
garis), sparrows (Passer domesticus), livid), étourneaux (Sturnus vulgaris),
pheasants (Phasianidae) and partridges moineaux (Passer domesticus). faisans
(Perdix sp.). There is little evidence of (Phasianidae) et perdix (Perdix sp.). Il
Newcastle disease existing in an endemic existe peu de traces de la maladie de New
form in wild bird populations. Neverthe castle sous forme endémique chez les
less, wild birds have played a part in the oiseaux sauvages; néanmoins, les oiseaux
dissemination of the disease both from one ont joué un rôle dans la dissémination de
country to another and within a country. la maladie tant d'un pays à l'autre qu'à
Chickens are more susceptible to the l'intérieur de pays déterminés.
clinical disease than other domestic Les poulets sont plus susceptibles à la
poultry. However, even in chickens a per maladie que les autres volailles domesti
manent carrier state is uncommon. Tur- ques. Toutefois, il est rare de trouver un

1
 keys are susceptible to the disease, but porteur permanent de virus même chez les
outbreaks in this species often remain poulets. Les dindons aussi sont pré
subclinical. Because of this, apparently- disposés à la maladie mais les cas restent
healthy turkeys have contributed to the souvent non cliniques; c'est pourquoi des
spread of Newcastle disease between coun dindons apparemment en bonne santé ont
tries and also within a country. contribué à répandre la maladie de New

Spread of the disease between individual castle d'un pays à l'autre, aussi bien qu'à
birds and between premises can occur in a l'intérieur d'un même pays.
variety of ways. Movement of domestic La propagation de la maladie entre
poultry has been considered the most im oiseaux et d'un poulailler à l'autre peut se
portant single cause of spread. Although faire de différentes façons; le transport de
the virus has been recovered from the con volailles domestiques a été considéré le
tents of eggs laid by an infected flock, at facteur le plus important. Quoique le virus
the present time egg transmission of the ait été recouvré à l'intérieur d'oeufs pro
virus is seldom reported. However, this venant d'un troupeau oùlamaladieexistait,
form of transmission remains potentially on signale de nos jours très peu de cas de
very dangerous; particularly as modern transmission du virus par les oeufs. Toute
developments in the poultry industry have fois, cette forme de transmission reste
meant that large numbers of hatching toujours un danger, surtout avec les prati
eggs, newly hatched chicks and growing ques modernes de l'industrie telles la dis
chickens are frequently distributed from tribution fréquente, sur de vastes distances,
one central place. de grandes quantités d'oeufs d'incubation,
The virus has survived outside the host de poussins et de poulets en croissance.
for variable, and often long, periods. As a Le virus a parfois vécu hors de son hôte
result, it is generally considered that the pour des périodes de temps variables et
survival of the virus in the environment souvent prolongées; on considère générale
plays an important part in perpetuating ment que sa survivance dans le milieu joue
and spreading the disease. The virus is un rôle important dans la perpétuation et
frequently recovered from the faeces of la propagation de la maladie. On trouve
birds suffering from a natural outbreak or souvent le virus dans les matières fécales
from poultry vaccinated with certain live d'oiseaux infectés naturellement ou de
vaccines. Following excretion, the virus is volailles ayant été vaccinées avec des
transmitted readily by air-borne particles. vaccins vivants. Des excrétions, le virus se
This air-borne spread by means of atmos transmet facilement dans l'air ambiant;
pheric air currents occurs between in La propagation peut alors se produire
dividual chickens in a pen, and between entre les poulets d'un même parquet ou de
different pens and premises. différents parquets ainsi qu'entre des
Other sources of infection related to poulaillers éloignés.
those already mentioned include poultry D'autres sources d'infection compren
markets, and the movement of poultry nent les marchés à volailles de même que
carcasses and poultry offal. Virulent New le déplacement des volailles abattues et des
castle disease virus in poultry vaccines has déchets d'abattage. Des virus actifs de la
also introduced the infection to previously maladie de Newcastle contenus dans des
disease-free areas. vaccins à volailles ont aussi été des agents
Newcastle disease virus can produce d'infection dans des régions auparavant
natural infections in humans. Human in exemptes de la maladie.
fection generally causes a conjunctivitis, Le virus de la maladie de Newcastle est
and sometimes an influenza-like illness. transmissible aux humains. Généralement,
There is no evidence to date of the exis- il cause une conjonctivite et parfois une
tence of the carrier state in man. indisposition ressemblant à l'influenza. Il
Only a few reports have indicated n'a pas encore été prouvé que l'homme
natural Newcastle disease infection in wild puisse être porteur de ce virus.
and domestic mammals. However, many Quelques rapports seulement ont dé
species of laboratory mammals are sus montré que des mammifères sauvages ou

ceptible to the virus by inoculation. Labor domestiques auraient été naturellement

atteints de la maladie de Newcastle. Toute
atory studies have involved bats, cats,
fois, par inoculation, on a réussi à repro
hamsters, mice, monkeys and other species.
duire la maladie chez plusieurs espèces de
There is little evidence
as yet to indicate
mammifères utilisés aux laboratoires. Des
that mammals play a part in the spread of
expériences ont été faites sur des chauves-
the disease.
souris, des chats, des hamsters, des souris,
des singes et autres espèces. Il n'a pas
encore été prouvé que les mammifères
Diagnosis of the Disease
jouent un rôle dans la propagation de la
From a review of the literature publish maladie.
ed during the period 1926-64, it would
appear that Newcastle disease virus has Diagnostic de la Maladie
been able to change in virulence and adapt D'après les documents publiés au cours
to its host. When first identified in 1926 de la période 1927-1964, il semblerait que
the disease was extremely virulent and le virus de la maladie de Newcastle ait fini
caused nearly 100 per cent mortality. The par modifié sa virulence et par s'adapter

form of the disease recognized about ten à son hôte. Lorsqu'elle a été identifiée pour
years later, called avian pneumoencephal- la première fois en 1926, cette maladie
itis, was less virulent. After another interval était extrêmement violente et causait un
taux de mortalité de près de 100 p. 100. La
of ten years, around 1944, asymptomatic
forme de cette maladie reconnue à peu
outbreaks were reported.
près dix ans plus tard sous le nom de
The disease is now considered to have
pneumoencéphalite, était moins virulente.
four main forms: the velogenic (virulent),
Après un autre intervalle de dix ans, vers
the mesogenic (less virulent), the lento-
1944, on signalait des infectations asymp-
genic (mild) and the asymptomatic. The tomatiques.
average incubation period is four to five On considère maintenant que la maladie
days, although considerable variation has se présente sous quatre formes principales:
been reported. vélogénique (virulente), mésogénique
(moins virulente), lentogénique (bénigne)
et asymptomatique. La moyenne de la
Chickens période d'incubation est de quatre à cinq
Velogenic Form jours avec des variations considérables.

This form of Newcastle disease may be

Poulets
peracute, but more typically it causes
Forme vélogénique
marked depression, increased rate of res
Cette forme de la maladie de Newcastle
piration, progressive weakness, diarrhoea,
peut être suraiguë, mais dans ses manifes
and death within a few days. Mortality is
tations les plus typiques elle cause une
usually over 90 per cent. Survivors gener
dépression marquée, une accentuation du
ally exhibit neurotropic involvement. rythme de la respiration, une faiblesse pro
Lesions are mainly haemorrhagic and gressive, de la diarrhée et, après quelques
inflammatory. The haemorrhages are jours, la mort. Le taux de mortalité dépasse

3
usually petechial and are commonly found habituellement 90 p. 100. Les survivants
in the mucosa and submucosa of the pro- souffrent de complications nerveuses.
ventriculus, gizzard and intestinal tract. Les lésions sont surtout hémorragiques
Microscopically, the lesions are essentially et inflammatoires. Des pétéchies sont habi

necrotizing in character. tuellement observées dans la muqueuse et

la sous-muqueuse du proventricule, du
gésier et du conduit intestinal. Sous le
Mesogenic Form
microscope, les lésions présentent les
This form is common in many parts of
caractéristiques essentielles de la nécrose.
the world. In a susceptible flock, the dis
ease appears suddenly and spreads rapid Forme mésogénique
ly. Respiratory distress, sharp drop in egg Cette forme se trouve communément
production and diarrhoea are common dans plusieurs parties du monde. La
symptoms. Mortality varies considerably maladie apparaît subitement et se propage
but is generally lower than in the velogenic rapidement dans les troupeaux prédis

form. Symptoms of paralysis are common posés. Une respiration difficile, de la

diarrhée et une forte réduction de la ponte
in survivors.
en sont les symptômes communs. Le taux
Not only does egg production decrease,
de mortalité varie mais il est généralement
egg quality is also affected: marked moins élevé que dans le cas de la forme
changes occur both in the egg shell and in vélogénique. Les survivants présentent
the albumen. souvent des symptômes de paralysie.
The lesions of haemorrhage and inflam Il se produit une diminution de la ponte
mation vary considerably between individ et des modifications notables tant dans la
ual birds and between outbreaks. Similarly, coquille que dans l'albumen.
Les lésions hémorragiques et inflam
there is marked variation in the organs
matoires varient considérablement d'un
and tissues involved.
individu à l'autre et d'une épidémie à
l'autre. On constate des variantes marquées
Lentogenic Form
entre les organes et les tissus touchés.
Impairment of appetite, mild respiratory
symptoms and sudden
Forme lentogénique
drop in the egg
Cette forme se signale souvent par une
production of laying flocks are common
perte d'appétit,des troubles bénins de la
features of the lentogenic form. In adult
respiration et une diminution soudaine de
fowls, mortality is negligible, and complete
la ponte. La mortalité est négligeable chez
recovery usually occurs within one to eight
les poules adultes qui prennent générale
weeks. huit semaines pour
ment d'une à se
There are usually neither haemorrhagic rétablir.
nor visceral lesions. Microscopic examin Habituellement, dans cette forme il n'y
ation has shown lymphoid infiltration of a pas de lésions hémorragiques ou vis
the respiratory and nervous systems. cérales. L'examen microscopique fait voir
des infiltrations lymphoïdes dans les sys
tèmes respiratoire et nerveux.
Asymptomatic Form
This form is often diagnosed only by Forme asymptomatique
chance. In the absence of clinical signs, Ce n'est souvent que par hasard que l'on
diagnosis is based on serological findings. réussit à diagnostiquer cette forme. En
Serological results have indicated that the l'absence de signes cliniques, le diagnostic
asymptomatic form of Newcastle disease est basé sur des recherches sérologiques:

4
may spread within a flock or may show les résultats obtenus indiquent que la
little or no evidence of spread. forme asymptomatique de la maladie de
Newcastle peut se propager à tout le
Ducks, Geese and Turkeys troupeau ou se limiter à quelques sujets.

Ducks, geese and turkeys are more re

Canards, Oies et Dindons
sistant to Newcastle disease than chickens.
It is not usually possible to distinguish the Les canards, les oies et les dindons sont
four forms of the disease that occur in plus résistants que les poulets; il est habi

chickens. Apart from this general differ tuellement impossible de distinguer chez
eux les quatre formes de la maladie qui
ence, the disease in turkeys resembles that
s'attaquent aux poulets. Hormis cette dif
in chickens. Ducks and geese are more
férence, la maladie de Newcastle des din
resistant than turkeys, usually undergoing
dons ressemble à celle des poulets. Les
symptomless infection and giving negative
canards et les oies y sont encore plus
post mortem findings.
résistants que les dindons; sans présenter
de symptômes, ils sont souvent porteurs de
Diagnosis by Serological virus et l'autopsie peut même donner des

Methods résultats négatifs.

A number of serological methods have

been used in the diagnosis of Newcastle
Diagnostic par les' Méthodes
disease. Of these, the haemagglutination- Sérologiques
inhibition (HI) test has been most widely On utilise un certain nombre de
adopted. This test is based on the property méthodes sérologiques pour diagnostiquer
of Newcastle disease virus to agglutinate la maladie de Newcastle. Parmi celles-ci
the red blood cells (haemagglutination) of l'épreuve d'inhibition-hémagglutination
birds and several species of mammals. The (IH) est la plus acceptée. Cette épreuve
agglutination est basée sur la propriété que possède le
can be inhibited by several
virus d'agglutiner les globules rouges du
substances, chief of which is Newcastle
sang des volailles et de plusieurs espèces de
disease antibody.
mammifères. Plusieurs substances dont la
Two procedures for conducting the HI principale est un anticorps du virus de la
test have been described: alpha and beta.
maladie de Newcastle peuvent avoir un
In the alpha procedure, the Newcastle dis
effet inhibitoire sur l'agglutination.
ease virus is diluted serially and mixed with
On connaît deux procédés pour faire
equal volumes of serum. In the beta pro l'épreuve IH. Dans la méthode alpha, on
cedure, the antibody (serum) is diluted dilue le virus en une série de diverses con
serially and mixed with a constant amount centrations que l'on mélange avec une
of antigen (virus). quantité égale de sérum. La méthode bêta,
The serum neutralization (SN), or virus au contraire, consiste à diluer l'anticorps
neutralization test, involving either grow (sérum) en une série de diverses concentra
ing chickens or chicken embryos, has been tions pour les mélanger avec une quantité

used to identify both Newcastle disease invariable d'antigène (virus).

L'épreuve de neutralisation du sérum
virus and antibody.
(SN) ou de neutralisation du virus faite sur
Other serological methods include hae
des poulets en croissance ou sur des em
molysis inhibition, an intradermal test, the
bryons de poussins est utilisée pour identi
use of fluorescent antibody, complement fier le virus aussi bien que l'anticorps.
fixation, and double diffusion plate tests. Autres méthodes sérologiques: hémolyse-

5
Diagnosis by Virus Isolation inhibition, épreuve intradermique, utilisa
tion d'anticorps fluorescents, fixation du
Serological methods are usually suffi
complément et épreuve de double diffusion
ciently accurate to confirm field evidence.
sur plaque.
However, new foci of infection should be
confirmed by recovery and identification
of the virus. Embryonating chicken eggs Diagnostic par l'Isolement du
are commonly used for virus isolations. Virus
Susceptible chickens, pigeons, laboratory Les méthodes sérologiques suffisent
mammals and tissue culture methods have habituellement à confirmer le diagnostic
been found both efficient and economical. posé d'après les symptômes cliniques.
Toutefois de nouveaux foyers d'infection
Differential Diagnosis devraient être vérifiés en obtenant et en
identifiant le virus. On utilise souvent des
The great variety of symptoms and
embryons de poussins pour isoler le virus.
lesions associated with Newcastle disease
Des poulets, des pigeons, des mammifères
has made differential diagnosis difficult. de laboratoire susceptibles à la maladie, de
Among the infectious diseases that need même que les cultures de tissus ont été
to be differentiated from Newcastle disease trouvés à la fois efficaces et économiques.
are fowl plague, avian encephalomyelitis,
infectious bronchitis, infectious laryngo-
tracheitis, avian duck
Diagnostic Différentiel
leucosis complex,
plague, Mycoplasma gallisepticum, avian La grande variété de symptômes et de

pasteurellosis and ornithosis (psittacosis). lésions qui accompagnent la maladie de

Non-infectious diseases caused by ribo Newcastle rendent difficile un diagnostic

flavin deficiency, vitamin E deficiency,

différentiel. Parmi les maladies infectieu
ses qui lui ressemblent et dont il faut la
toxic drugs and toxic plant seeds may also
distinguer, on compte: la peste aviaire
need to be differentiated.
(fowl plague), l'encéphalomyélite aviaire,
la bronchite infectieuse, la laryngotrachéite
Control Measures infectieuse, le complexe leucémique, la

Slaughter Measures and peste du canard, la mycoplasmose {Myco

plasma gallisepticum), la pasteurellose
International Trade
aviaire et l'ornithose (psittacose). On peut
Slaughter measures have been applied to aussi avoir à différencier la maladie de
eradicate Newcastle disease from a num Newcastle de maladies non infectieuses
ber of countries. In South Africa, England, causées par des carences de riboflavine ou
Wales, Sweden and Australia, where the de vitamine E, par des drogues ou par des

velogenic form of the disease has occurred, graines de plantes toxiques.

eradication by slaughter measures has
been successful. However, where countries
Mesures de Répression
have been invaded by the mesogenic and
lentogenic forms, as in England, Wales, Abattage Obligatoire et
Canada, The Netherlands and Western Commerce International
Germany, the application of slaughter On a eu recours à l'abattage obligatoire
policies has failed to eradicate the disease. pour l'éradication de la maladie de New
The world distribution of Newcastle dis castle en certains pays. Un tel programme
ease and the control policies established by a réussi en Afrique du Sud, en Angleterre,

ft
different countries have an appreciable au Pays de Galles, en Suède et en Aus
influence on the international movement tralie où la forme vélogénique de la
of live poultry, hatching eggs and poultry maladie sévissait. Toutefois, l'abattage
carcasses. In many countries in which obligatoire n'a pas réussi à réprimer les
Newcastle disease has a variable distribu formes mésogénique et lentogénique de la
maladie, dans les pays suivants: Angle
tion, the importation of poultry and poul
terre, Pays de Galles, Canada, Pays-Bas,
try products is very strictly controlled; and
Allemagne de l'Ouest.
in countries such as Australia and New
Zealand,
La répartition mondiale de la maladie
where Newcastle disease does not
de Newcastle et les mesures de répression
exist at present, the importation of live
établies par différents pays ont une in
and dead poultry and hatching eggs is
fluence appréciable sur le mouvement
completely prohibited. To reduce the ex
international des volailles vivantes et
tent of these restrictions, it has been abattues ainsi que des oeufs d'incubation.
suggested that control measures be applied Plusieurs pays où la présence de la maladie
on the basis of "infected region of a de Newcastle varie beaucoup, régissent
country" instead of "infected
country." rigoureusement l'importation de la volaille
Implementation of the concept of "infect et de ses produits. Certains pays indemnes
ed region of a country" might require that de la maladie, comme la Nouvelle-Zélande
Newcastle disease be a notifiable disease et l'Australie, ont complètement prohibé
throughout the world. It was notifiable in les importations de volailles vivantes ou
84 countries in 1962. abattues et d'oeufs
d'incubation. Pour
réduire la portée de ces restrictions, on a
Compulsory notification of outbreaks of
suggéré que des mesures de répression
Newcastle disease is difficult to enforce,
soient prises sur une base de «région in
partly because the disease is often not
fectée dans un pays» plutôt que celle d'un
recognized in its lentogenic and asympto
«pays infecté». Il pourrait résulter de
matic forms and is not, therefore, reported
l'acceptationdu concept d'une «région in
to the authorities. The existence of asymp fectée dans un pays» que la maladie de
tomatic outbreaks, and the use of live virus Newcastle devienne une maladie con
vaccines which can spread the infection to tagieuse «nommée» et à déclaration obli
susceptible poultry, add to the problems gatoire dans tous les pays du monde. Elle
faced by national disease control agencies. l'était dans 84 pays en 1962.
Despite these obvious difficulties, regions L'obligation de déclarer les manifesta
of several countries have been freed from tions de la maladie de Newcastle n'est pas
Newcastle disease for extended periods facile à mettre en application parce que la
through control by slaughter. maladie est difficile à reconnaître dans ses
formes lentogénique et asymptomatique.
De ce fait, on néglige d'en informer les
The Action of Chemicals on autorités. L'existence de formes asympto-
Newcastle Disease Virus matiques et l'emploi des vaccins à virus
vivants qui peuvent propager la maladie
A number of different methods have
parmi les volailles prédisposées, compli
been used to evaluate the viricidal effect
quent les problèmes pour les organismes
of disinfectants on Newcastle disease virus.
nationaux chargés de sa répression. Toute
Some of the findings are tabulated in this
fois, certaines régions dans plusieurs pays
review, though direct comparisons be ont été exemptes de la maladie de New
tween different studies are difficult to castle pour de longues périodes grâce à la
make. répression par l'abattage.

7
Less Common Methods of Action des Produits Chimiques
Control sur le Virus de la Maladie
Hyperimmune serum, serum-virus and de Newcastle
medicinal treatments have been used with Des méthodes variées ont été utilisées
varying degrees of success, and experi pour estimer l'effet des désinfectants contre
mental results so far obtained have been le virus de la maladie de Newcastle. La
inconclusive. présente publication contient des résumés
de ces données; cependant, les comparai
sons directes entre diverses études sont
Control by Vaccination difficiles à établir.
Vaccination has been the most widely
adopted method of controlling Newcastle
Méthodes plus rares de répression
disease. In 1961, over 82 per cent of the
countries reporting the disease were using
L'inoculation de sérum hyperimmun,
l'immunisation mixte (serum-virus) et les
vaccination as their main control proce
traitements médicaux ont été utilisés avec
dure. The following immunological aspects
plus ou moins de succès, et les résultats
are discussed in this review: antigenic
des expériences n'ont pas été concluants.
plurality; procedures for evaluating im
munity; and factors that influence the
development of immunity,
including pas Répression par la Vaccination
sive immunity, age at time of vaccination, La vaccination a été la méthode la plus
vaccine virus titres, viral interference, dif universellement adoptée. En 1961, plus de
ferences between individual birds and 82 p. 100 des pays qui signalaient la pré
susceptibility to another disease. sence de la maladie utilisaient la vaccina

Vaccines are of two main types: live and tion comme principal moyen de répression.
Les aspects immunologiques qui suivent
inactivated. It is convenient to divide live
sont étudiés dans la présente publication:
vaccines into lentogenic and mesogenic
pluralité des antigènes; procédés pour
virus strains. Among the most commonly
estimer le degré d'immunité et les facteurs
used lentogenic strains are the Bl, F and
qui influent sur le développement de l'im
LaSota; among the mesogenic: the Kom- munité: immunité passive, âge au moment
arov, Mukteswar, Roakin and MK107. de la vaccination, titrage des vaccins,
Another type of live vaccine has been interférence virale, différences individuel
developed by the attenuation of virulent les et prédisposition à une autre maladie.
strains of virus and propagation in mam Il existe deux sortes de vaccins: des

malian tissue cell cultures. A number of vaccins vivants et des vaccins tués. Les
authors have described the clinical effects vaccins vivants sont préparés avec des

of vaccination of im
and the duration virus de lignées lentogéniques et mésogéni-
ques. Parmi les lignées lentogéniques le
munity with each of these strains. In this
plus communément employées, on compte
review their findings are compared and
lesBl, F et LaSota, parmi les mésogéni-
discussed.
ques, les Komarov, Mukteswar, Roakin et
A wide variety of agents has been used MK107. Un autre type de vaccin vivant se
to produce inactivated vaccines. These prépare en atténuant certaines lignées de
agents include beta-propiolactone, forma virus et en les propageant dans des cultures
lin, crystal violet, and ultraviolet irradi de cellules provenant de tissus de mammi
ation. A considerable amount of data has fères. Certains auteurs ont décrit les effets
been accumulated on the duration of im cliniques de la vaccination et la durée de
munity produced by inactivated vaccines. l'immunité que procure chacune de ces
lignées; on trouvera dans la présente pub
It is important, also, to consider com
lication des détails et des comparaisons se
bined vaccines administered as a single
rapportant à leurs conclusions.
inoculum, and the administration of two
Pour produire des vaccins tués on utilise
vaccines simultaneously but by different
une grande variété d'agents: bêta-propio-
routes.
lactone, formaline, cristal violet et irradia
The choice of the most suitable vaccine tions par les rayons ultra-violets. On a
to meet any particular set of conditions is accumulé une quantité considérable de
very complex. A comparison of the dura données sur la durée de l'immunité que
tion of immunity engendered by Bl, F, confèrent les vaccins tués.
formalin-inactivated and BPL-inactivated Il est intéressant d'étudier l'inoculation
strains, indicates that, to date, BPL-inacti de vaccins combinés en une seule opéra
vated vaccines have resulted in very satis tion ainsi que l'administration simultanée
de deux vaccins par voies différentes.
factory duration of immunity. There is
little doubt that a killed vaccine of high Il est assez difficile de faire le choix du
antigenicity would be preferable to live vaccin qui convienne à toute une série de

vaccines and would form an essential

conditions données. Une comparaison
quant à la durée de l'immunité que con
part of any eradication scheme. Another
fèrent le Bl, le F, le vaccin tué à la forma-
feature that has become apparent is the
line ou par le BPL a indiqué que jusqu'a
need to conduct one or more revaccina
présent le vaccin tué par le BPL donne des
tions if from one day of age
protection
résultats très satisfaisants. Il y a peu de
through to the end of the laying year is to doute qu'un vaccin tué à antigénicité
be achieved. élevée, serait préférable aux vaccins
As measures for the control of other vivants et entrerait dans les cadres de tout
avian virus diseases advance, there will be programme d'éradication. Il est évident
qu'on doit revacciner une fois ou plus afin
an increasing demand for live Newcastle
d'assurer une protection depuis la nais
disease vaccines prepared from tissue cul
sance jusqu'à la fin de l'année de ponte.
tures, preferably of non-avian origin. One
Au fur et à mesure que progresseront
modified Newcastle disease virus developed
les méthodes de répression des autres
in tissue culture is distinctive because
maladies virales des volailles, il se produira
this virus has shown little or no tendency
un accroissement de la demande de vaccin
to spread and the immunity it engenders vivant contre la maladie de Newcastle,
is of considerable duration. préparé de cultures sur tissus, préférable-
In the control of Newcastle disease by ment d'origine non aviaire. A plusieurs
vaccination, a major issue is whether the points de vue, le virus modifié de la
maladie de Newcastle développé sur des
poultry industry will accept the advantages
cultures de tissus se distingue des autres
to be gained by using vaccines which
parce qu'il ne montre à peu près aucune
have to be administered to each chicken
tendance à se propager. De plus, l'im
individually. For many countries, this
munité qu'il confère dure longtemps.
would entail abandoning mass vaccination
L'une des questions les plus importantes
methods. It is doubtful if this change qui se posent dans la répression de la
would be made voluntarily. Government maladie de Newcastle par la vaccination
assistance or legislation might be neces est celle de savoir si l'industrie avicole
sary. Nevertheless, there is now evidence acceptera les avantages résultant de l'utili

9
that, in general, vaccines requiring in sation de vaccins qu'il faut administrer à

dividual administration give better im chaque oiseau individuellement, ce qui

munity. The development of procedures nécessiterait pour plusieurs pays, l'aban
for the mass administration of inactivated don de la vaccination en masse. Il est
douteux qu'un tel changement soit adopté
or non-spreading tissue culture vaccines
volontairement; une aide ou une législa
could lead to great changes in the global
tion gouvernementale pourrait être néces
picture of Newcastle disease.
saire. Néanmoins, il semble évident main
The world history of Newcastle disease tenant qu'en général les vaccins qu'il faut
has shown that, once the disease has be administrer individuellement assurent une
come established in a country or region, meilleure immunité. La découverte de

it tends to become endemic. Where this méthodes pour l'administration collective

has happened, little progress has been de vaccins tués et non infectieux, prove
made towards eventual eradication. Con nant de cultures faites sur tissus, pourrait
trol measures have tended to be of an entraîner de grands changements dans le
problème global de la maladie.
emergency nature and to involve the use
of live virus and mass vaccination. L'histoire mondiale de la maladie de
Newcastle a démontré que lorsque la
The costs of vaccination are usually maladie s'est établie dans un pays ou une
borne by the poultry industry. Vaccination région, elle a tendance à devenir endémi
of the United States broiler crop in 1956 que. Là où la chose s'est produite, on a
cost more than four million dollars. In réalisé peu de progrès pour l'éradication
Africa, the cost of vaccination has gener éventuelle de cette maladie. Les mesures
ally been considered out of proportion to de répression ont toujours eu un caractère
the economic value of the stock and the d'urgence et comprenaient l'utilisation de

demand for vaccine has been poor. virus vivants et la vaccination collective.

Compulsory vaccination has sometimes Le coût de la vaccination est habituelle

ment supporté par l'industrie avicole. Aux
been adopted in spite of serious difficulties.
Etats-Unis, la vaccination des poulets à
To overcome the difficulties, the cost of
griller produits en 1956 a coûté plus de
the vaccine has been subsidized.
quatre millions de dollars. En Afrique, on
A review of the literature indicates trouve que les frais de vaccination sont
clearly that no one vaccine has proved généralement hors de proportion avec la
ideal for all situations and all geographical valeur économique des sujets et la de
areas. As a result, there would appear to mande pour le vaccin a été faible.
be a need for more government-sponsored La vaccination obligatoire a parfois été

experiments to determine the vaccines and adoptée en dépit de difficultés sérieuses.

methods of use best suited to individual Le coût du vaccin a alors été subventionné
countries. It is axiomatic that these studies pour triompher des difficultés.

need to be supported by adequate labor L'examen des documents indique qu'il

n'existe aucun vaccin idéal pour tous les
atory facilities.
cas et dans toutes les régions géographi
In any discussion on immunization
ques. Aussi semble-t-il que les gouverne
against Newcastle disease, it is necessary devraient études,
ments poursuivre des
to establish whether the protection sought afin de déterminer quels sont les vaccins
is against mortality and paralysis; or et les méthodes d'emploi les mieux adaptés
whether the greater need is to protect à chaque pays. Ces études exigeront des

against the adverse effects of the disease laboratoires convenablement outillés.

on egg production and egg quality. Dans tout travail sur l'immunisation

10
 Virus Propagation and contre la maladie de Newcastle, il est

Vaccine Preparation nécessaire d'établir si la protection est

recherchée contre la mortalité et la paraly
Newcastle disease virus is classified as
est de
sie ou si le besoin le plus important
a Myxovirus and has been given the name protéger contre les effets de la maladie sur
Myxovirus multiforme. The properties of la production et la qualité des oeufs.
the virus and the practical aspects of
vaccine production have been studied
extensively. Different strains of the virus Propagation du Virus et
have been identified on the basis of patho Préparation du Vaccin
genicity and chemical properties. Electron Le virus de la maladie de Newcastle est
microscope studies of the virus have also classé parmi les Myxovirus et on lui a

been made, and the arrangement of its donné le nom de Myxovirus multiforme.
structural components suggested. On a beaucoup étudié les propriétés de ce
The virus has proved adaptable to a virus et les aspects pratiques de la pro
duction du vaccin. Différentes lignées du
number of different living tissue cells of
virus ont été identifiées en tenant compte
avian and mammalian hosts and embryos.
de leur pathogénie et de leurs propriétés
The ease with which Newcastle disease
chimiques. Des études du virus faites au
virus can be propagated is utilized in the
microscope électronique ont permis de
large-scale preparation and use of a variety suggérer la disposition structurale de ses
of vaccines. constituants.
At the present time, the majority of On a trouvé que le virus peut s'adapter
Newcastle disease vaccines is prepared à un certain nombre de tissus cellulaires
from virus propagated in embryonating vivants d'oiseux, de mammifères et d'em

chicken eggs. The main disadvantage to bryons. Sa facilité de multiplication est

using eggs is the risk of their containing a utilisée pour la préparation sur une vaste

variety of infectious agents. To overcome échelle et l'utilisation de vaccins variés.

Presque tous les vaccins contre la
this problem, vaccine strains of virus have
maladie de Newcastle sont préparés de
been propagated in tissue culture. The vast
virus propagés dans des embryons d'oeufs
majority of live vaccines is now dispensed
de poule, mais avec ce procédé on court le
as a lyophilized product and this has
risque que les oeufs contiennent des agents
greatly facilitated transportation and
d'infection. Pour éliminer ce danger, on
storage. propage des lignées de virus pour vaccin
A number of procedures for testing and sur des cultures de tissus.
standardizing Newcastle disease vaccines On a mis au point certaines méthodes
have been devised. However, at the present de vérification et de normalisation des
time, tests and criteria for the safety and vaccins. Toutefois, à l'heure actuelle, les
potency of vaccines are not uniform méthodes de vérification et les normes de
sécurité et d'efficacité ne sont pas identi
throughout the world.
ques dans le monde entier. La grande
majorité des vaccins vivants sont main
tenant distribués comme produits lyophi
lisés.

11
 DEFINITION OF NEWCASTLE DISEASE

Nomenclature castle disease virus into two distinct groups

— both groups shared common antigens
Newcastle disease takes its name from
and had similar reproduction patterns.
the town of Newcastle-on-Tyne in the
Other tests that have been used for strain
county of Northumberland, England,
identification include the intracerebral
where an acute disease occurred in a flock
inoculation of mice (Nitzschke and Sch-
of poultry in the spring of 1926 (Doyle,
mittdiel, 1960; Upton et al., 1953a, 1955)
1927). In the following 15 years, the dis
and the adsorption of haemagglutinating
ease was given more than 20 synonyms.
activity by suspensions of chick embryo
Many of these are listed in the publications
brain cells (Piraino and Hanson, 1960).
of Reis and Nobrega (1956) and Beaudette
Piraino and Hanson (1960) have suggest
(1943). The following are among the
ed that the pathogenicity of different
names that have been most commonly
strains of virus for chickens is related
used: Ranikhet disease (Edwards, 1928),
partly to the avidity of the virus for brain
pseudo-fowl (Picard, 1928; Hutra et
pest
cells. The resistance of haemagglutinin to
al., 1938), pseudo-poultry plague (Johns
heating to 56°C has also been used to dis
tone, 1931), Doyle's disease (Haddow,
tinguish between different strains (Nitz
1938), avian pneumoencephalitis (Beach,
schke and Schmittdiel, 1963). Electron
1943), respiratory nervous disorder (Bang,
microscope examination of a number of
1946) and avian or fowl pest (gefliigelpest)
different strains of virus has revealed no
(Farinas, 1930; Kuppuswamy, 1935).
appreciable morphological differences be
Doyle (1935) thought the name New
tween strains (MacPherson, 1956a; Muss-
castle disease unsuitable;nevertheless he
gay and Weibel, 1962; Reagan et al.,
felt that the terms "pseudo-plague" and
1948b, 1950b).
"avian pseudo-pest" should be avoided be
cause of possible confusion with another The general problem of determining the
disease generally known as fowl plague. particle size of Newcastle disease virus has
The name fowl pest was used in the been investigated by Elford et al. (1948).
Fowl Pest Order of 1936, made under the In normal preparations, the average par
Diseases of Animals Act of the United ticle size is 1 92 microns. However, Reagan
Kingdom of Great Britain and Northern et al. (1948) have reported virus particles
Ireland. In British legal context fowl pest between 1 00 and 1 25 microns in diameter.
still applies to the two separate diseases, According to Cunha et al. (1947) the
Newcastle disease and fowl plague. approximate size of the head piece is in
the range of 80 to 120 microns (Burnet
and Ferry, 1934).
Properties of the Virus Schafer et al. (1949) and Reagan et al.
The physical of Newcastle
properties (1950b, 1956) examined particles of New
disease virus have been reviewed by Elford castle disease virus with an electron micro
et al. (1948) and a number of properties scope and reported that in aqueous solu
have been used to type different strains of tion they appeared to be rounded; whereas
the virus (Acocella, 1955; Anon., 1959; in saline solution they appeared thread-like
Hanson et al., 1949, 1951; Hanson and in shape. This change in shape was con
Brandly, 1955; Kaschula, 1952a; Upton sidered an by Angulo (1951),
artifact
et al., 1953b). By means of pathogenicity Elford (1948) and Schafer et al.
et al.
and serological studies, MacPherson and (1949). Bang (1948), on the other hand,
Swain (1 956) divided eight strains of New considered the filamentous particles to be

13
 SCHEME OF THE STRUCTURE OF NEWCASTLE DISEASE VIRUS

(Modified from SchciUr, 1963)

150 1 (ENVELOPE)

Figure 1.

individual virus particles. This latter view gested that the virus particle is essentially
was based on the fact that the characteris- sperm-shaped. The filamentous forms
tic filamentous particles were not seen in which occur when the virus in saline
is
other virus preparations; that the particles solution revert to the spherical form when
were agglutinated by specific antisera; and replaced in water. These changes have not
that they produced infection in embryos, been related to any loss of infectivity
Similarly, Cunha et al (1947) have sug- (Bang, 1948, 1949). The conversion to

14
Figure 2. — Electron micrograph of Newcastle disease virus (x 150,000). (Courtesy of Dr. A. P.
Waterson, St. Thomas's Hospital, London.)

the filamentous forms can be prevented by This virus density apparently differs ac
partial inactivation with formalin (Bang, cording to the cell type used for propaga
1947). tion. Thus, the infectivity peak densities
All strains of Newcastle disease virus of virus in avian cell culture have been
studied by McCollum and Brandly (1955b) lower than the corresponding figures for
possessed mucinase. It has been suggested virus of mammalian cell origin.
by Mierzejewski (1962) that the catalytic The ribonucleic acid (RNA) fraction of
activity of aldolase in the breakdown of Newcastle disease virus has been found
fructose diphosphate varies between dif non-infective for chick embryos, and there
ferent pathogenic strains of the virus. has been no proof that RNA is responsible
Using three neutralization and sero for infectivity or agglutinins (Benedict et
logical tests, Doll et al. (1956) demon al., 1960). However, the use of lipids
strated that Newcastle disease virus is apparently has maintained the infectivity
serologically and immunologically distinct of the RNA from an Indian strain of virus
from the viruses of mumps, and of human (Dhar et al, 1963).
and swine influenza. Based on morphology, The inactivation of Newcastle disease
there appears to be no resemblance be virus has been examined by radiation and
tween Newcastle disease virus and the Wilson and Pollard (1958) have estimated
viruses of influenza (Cunha et al, 1947). that the total radius of the virus is at least
In caesium chloride density gradients, 510a.
Newcastle disease virus appears to contain According to Cunha et al. (1947), the
infectious with a wide range of
particles virus has an internal differentiation that
densities (Stenback and Durand, 1963). resembles the analogous morphological

15
structure of living cells. Negative contrast particular (Valentine and Isaacs, 1957)
electron microscopy before and after treat has been suggested (Figures 1 and 2).
ment with ether has indicated that the The properdin system can inactivate
is indis Newcastle disease virus (Wedgwood et al.,
internal helical ribonucleo-protein
1956). This inhibition requires all the
tinguishable between various strains of
known constituents of the properdin sys
Newcastle disease virus (Waterson and
tem (Karzon and Bussell, 1960). As a
Cruickshank, 1963). The significance of
result, a theory of resistance to infection
the virus components has been reviewed
with Newcastle disease virus which is not
by Schafer (1963); and an explanation of part of the classical antigen-antibody im
the principles underlying their structure mune concept has been formulated (Kar
and of the arrangement of the RNA in zon, 1956).

16
 PART I: Spread of the Disease
GEOGRAPHIC DISTRIBUTION

Early Reports of Diseases became infected with Newcastle disease

Resembling Newcastle Disease during the 1949 poultry epizootic.

Between 1909 and 1915, diseases now
A disease resembling Newcastle disease considered to be atypical forms of fowl
is believed to have been carried to the plague were reported from several regions
African continent in the mid-nineteenth of the world (Jacotof, 1950). These dis
century by small ships trading with Asiatic eases appear to have had characteristics
ports (Da Camara and Valadao, 1950; in common with Newcastle disease and
Castro Amaro, 1 964) . A disease suggestive were called "peste atypique," or "pseudo-
of Newcastle disease was also observed pest," or were given names with a geo
by Petenyi near Budapest in 1833 (Man- graphical qualification such as "Egyptian
ninger, 1949). pest" and "Madras pest."
At the end of the century there was In March 1 926, Newcastle disease broke
probably an extensive epizootic of New out in the Dutch East Indies near the towns
castle disease in northwest Scotland. Mac- of Batavia, Cheribon and Soerabaia, on the
Pherson (1956b) reached this conclusion island of Java, where it exhibited an ability
after reading a Gaelic poem written in to spread quite unlike that of any known
1898 and doing research in the Hebrides. disease of poultry. The disease was first
He has drawn attention to the fact that in reported by Kraneveld (1926). Later
1 899 and 1900 the Scottish Congested Dis Picard (1928) gave details of the first
tricts Board distributed hatching eggs and reports received from the Civil Veterinary
fowls to the same areas in which outbreaks Department of the Dutch East Indies and
of Newcastle disease occurred between confirmed that in March 1 926 the Veterin
1949 and 1951. Field data and experi ary Institute at Buitenzorg, Java, was
ments with living cormorants (Phalacro- aware of the existence of a serious poultry
corax carbo) led MacPherson to believe disease near the town of Batavia (now
that the cormorant was the source of in Jakarta). The Civil Veterinary Depart
fection in Scotland in 1897, and again in ment reported that the disease had caused
the 1949-51 epizootic. He has suggested enormous losses in a number of districts on
that cormorants and other sea birds are the island and had also been identified in
true reservoirs of
infection and not remote areas throughout the archipelago.
mechanical carriers. He has also suggested In several native villages (kampongs) not
that there may be a long-established a single fowl was left. It was impossible to
biological adaptation to Newcastle disease ascertain the origin of the disease. For
virus in the cormorant, and that the domes Doyle (1948) also, the origin of the out
tic fowl may be only a secondary host. breaks in the Dutch East Indies remained
However, Blaxland (1951) has rejected a mystery, but he thought it unlikely that
this explanation and has maintained that the disease had existed as a latent infection
the cormorants around the Scottish coasts among the native fowls of the islands

1"
because these fowls proved highly sus the disease continued to spread, reaching
ceptible. Japan (Nakamura et al., 1933), southern
India (Kylasamaier, 1931) and Australia
First Outbreaks of Newcastle (Johnstone, 1931) by the end of 1930.
Disease in England Thereafter spread was increasingly rapid
(Dobson, 1952). It has been noted by
The outbreak in a flock of chickens in
MacPherson (1956b) that many of the
Newcastle-on-Tyne in England in 1926
initial outbreaks occurred at seaports or
(Doyle, 1927), which was responsible for
were associated with the movement of
the name Newcastle disease, was thought
ships.
to have been introducedwith infected food
from a ship (Anon., 1962). It is interesting
to note, however, that the original flock India
involved at Newcastle had also been fed a In India, Newcastle disease was first re
ration supplemented with offal collected cognized at the town of Ranikhet in the
from the town. There were two other out United Provinces in July 1927. Almost at
breaks diagnosed in England in 1926: one the same time, a similar disease appeared
in Somerset, and one involving a number in Madras State. As in the Dutch East
of farms in Staffordshire. The latter was Indies, the disease spread rapidly. By the
thought to have resulted from the sale of end of 1927 it had spread throughout the
infected birds in a local market. The United Provinces; by the spring of 1928
mortality in all the 1926 outbreaks in Eng it had reached the Punjab and Bombay.
land was 98 to 100 per cent and, as there Soon afterwards the disease was reported
were few survivors, the disease quickly from all parts of India (Haddow, 1941).
died out.
These English outbreaks were regarded
Philippines
by Doyle (1948) as chance offshoots of
the main stream of infection that origin By comparison with the rapid spread of
ated in the Dutch East Indies. However, Newcastle disease in some countries, rate

the epidemiology of the early outbreaks of spread was slow in the Philippines. This
in England (Dobson, 1952) was very dif was due, perhaps, to the enforcement of
ferent from that experienced in the Dutch quarantine regulations and the movement
East Indies where the disease spread of poultry inward, toward the original
throughout the whole territory within a
focus of infection, rather than outward,

period of six months (Picard, 1928). toward the provinces (Coronel, 1939).
An account of the spread of Newcastle
Spread Through Southeast Asia disease in the Philippines during the ten-
year period 1927-37 has been given by
In the autumn of 1926, Newcastle
Coronel (1939). The disease spread in
disease was reported in Korea (Konno et
all directions from the original focus in
al, 1929; Ochi and Hashimoto, 1929).
the city of Manila. It usually made its
During 1927, outbreaks occurred at
first appearance in a province either in the
Ranikhet in the Kumaon Hills of India capital of the province or in a large town
(Edwards, 1928; Cooper, 1930), at Col on a rail or road route, but it rarely ap
ombo in Ceylon (Sturgess, 1928; Craw
peared in remote districts (barrios).
ford, 1931) and at Manila in the Philip
pine Islands (Farinas, 1930). Thus, in
1926 and 1927 disease was
Malaya
Newcastle
identified of Southeast
in five countries In the province of Wellesley in Malaya,
Asia. From these known foci of infection poultry diseases, including Newcastle

IS
disease, were spread mainly through the Newcastle disease in Canada are given on
activitiesof poultry dealers. The baskets page 84.
in which poultry were carried permitted
the droppings and discharges from in Africa
fected birds to fall on roads and paths
In Africa south of the Sahara, New
between villages (Kuppuswamy, 1935).
castle disease was recognized at the ports
of Mombasa in 1935, Durban in 1944,
Haiti and Madagascar
Cape Town in 1949, and Leopoldville in
As in the Philippines, in both Haiti 1948 (Vandemaele, 1961), as shown in
(Bush, 1954) and Madagascar (Buck, Figure 3. These cases were probably
1947) Newcastle disease first spread caused by fresh introductions of infection
along main traffic routes to market areas. (Scott et al, 1 956) , although evidence was
found that the disease had existed in
United States enzootic form along the coasts of Kenya

In the United States, the appearance and Tanganyika for some time.

circa 1935 of a mild respiratory-nervous A survey conducted in the Congo has

disorder of chickens in California, term shown that the disease there has been con

ed "avian pneumoencephalitis" fined mainly to small flocks owned by

(Beach,
1942), was not identified as Newcastle
Africans (Depoux and Chambron, 1960).
disease until However, in the Union of South Africa
nine years had elapsed
al, outbreaks have involved a number of large
(Beach, 1944, 1946; Brandly et 1944;
poultry flocks (Anon., 1950b). The main
Stover, 1942). The disease reached the
east coast of the United States by 1944 epizootic pathways in Africa have been
closely related to the main rail and road
(Cunha et al, 1947). By August 1946 it
had been diagnosed with certainly in 17 trade routes.

states,including a number on or near the

Atlantic coast (Brandly et al., 1946e; Spread Through Europe
Bruner et al, 1947; Morgan, 1946; The incidence of Newcastle disease and
Stubbs, 1946). fowl plague in Europe during the period
1940-55 has been summarized by Eckert
Canada
(1957), as illustrated in Figure 4. The
Following the identification and subse incidence of Newcastle disease in the same
quent eradication of Newcastle disease countries in 1962 is shown in Figure 5.
from the province of Alberta in 1950, a In many countries, Newcastle disease
serological survey was made of 1 per cent has shown a marked tendency to variation
of the total poultry population of that in pathogenicity. This tendency is for
province. Only one suspicious flock was mesogenic and lentogenic strains to replace
identified. The natural isolation of farm the initial velogenic form of the disease.
flocks in Alberta was probably responsible Such epizootiological changes have been
in part for the successful eradication of the recognized in western Germany (Fritzsche,
disease (Ballantyne and Bigland, 1951). 1963), France (Lissot, 1956), Yugoslavia
During 1952/53 an extensive survey of (Jaksic and Stefanovic, 1957), and Great
Newcastle disease virus antibody was Britain (Anon., 1962b). In some countries
made in Canada. Some 2,130 flocks in epizootiological changes have been in
eight provinces were sampled, and the fluenced by control policies. Further men
results of serological tests showed 12 per tion of this topic is made under the head
cent to have been (Crawley,
infected ing "General Characteristics of the Dis
1954a). Further details of the history of ease" on page 45.

19
 NEWCASTLE DISEASE IN AFRICA SOUTH OF THE SAHARA

Dates of First Outbreaks and Direction of Spread

(Redrawn from Vandemaele 1961)

Figure 3.
-n
N0I1D3JNI A1ISN3Q dO 3H1SADM3N 3SA3SIO aNV HMOd
3novid ni 3doan3 omana 3hi aoiaad 6i of , ss6i

H3D3 JD»A J»d OOO'l I**

uMDJpay) mOJf *Je>Ic3 (ZS61

prrrj o*o ■ o*i

iz
 TABLE — Outbreaks of Fowl Pest in Great Britain —
1

Financial Years 1954/55 to 1960/61 (Anon., 1962b)

Financial No. of No. of Outbreaks

year adult broiler
hens chickens
kept produced Primary' Secondary2 Total

(million) (million)

1 954-55 52 20 162 388 550

1955-56 54 40 154 757 911
1 956-57 58 47 214 822 1,036
1957-58 59 56 236 742 978
1 958-59 63 82 245 537 782
1959-60 66 108 391 2,333 2,724
1 960-61 63 142 342 1,529 1,871

1 A primary outbreak is one which has no established connection with a previous outbreak.
2 A secondary outbreak is one where the infection is known to have spread from another
outbreak.

Great Britain ment of chickens and turkeys by three

poultry dealers was responsible for the
The spread of Newcastle disease in
spread of the disease to 12 premises; and
Great Britain has been well documented
two additional outbreaks occurred on
(Anon., 1962b; Asplin et al, 1949; Cal
adjacent farms.
ender, 1958; Gordon, 1961; Reid, 1955,
1961). In 1950, the tracing of birds moved
from a large poultry show in England dis Austria
closed 255 outbreaks. In 1955, movement In Austria, Newcastle disease first
of infected growing stock from one appeared in 1942 (Grausgruber, 1963).
hatchery resulted in 108 secondary out During the period from 1946 to 1961
breaks. In the same year, illegal movement incidence of the disease in most areas
of infected birds resulted in 187 outbreaks reached a peak between 1947 and 1951
being confirmed in various counties. Out and declined thereafter. In the Burgenland
breaks in broiler chickens in 1960 con area of the country, extensive outbreaks
tributed to an involving 375
epidemic occurred in 1954, 1955 and 1957.
premises. The rapidity of spread of New
castle disease (fowl pest) in Great Britain
World Distribution in 1962
and the difficulty of tracing sources of
infection is illustrated in Table 1. In 1962, 103 countries reported New
castle disease (Table 2). A number of
Ireland countries, including Jordan, Mauritius,
The origin of outbreaks of Newcastle The Netherlands, Pakistan, Romania and
disease which occurred in 1950 in south Venezuela, reported a decrease in its in
east Ireland has not been determined with cidence. On the other hand, Argentina
certainty, though it is possible that the (Anon., 1961a; FAO-WHO-OIE, 1962),
infection originated in an adjacent colony Denmark (Marthedal et al., 1963) and
of sea birds (Anon., 1951-52). Fourteen Luxemburg (Vittoz, 1962), which had
outbreaks were confirmed and a sequence previously been free from the disease, re
of infection was established. The move ported it for the first time. Iceland, Nor-

22
way, Sweden, Finland, the Republic of among the countries free from Newcastle
Ireland, Australia and New Zealand were disease in 1962 (Vittoz, 1963).

TABLE 2 — Incidence and Control of Newcastle Disease, 1962

(Lancaster, 1964b)

Countries reporting !§ o Countries conducting

Geographical area g 2 rag „ mo .52 |

l|
o ^..o

|1
■s!
of h
11
ll
08
ci
1=
§p fl «{
i§

North Africa 5 3 1 — 3 3 1
Central Africa 24 7 8 4 6 3 7
South Africa 22 4 9 9 16 6 7
South America 24 12 7 2 13 6 11
North America
and West
Europe 24 4 9 9 17 10 a
East Europe
and Asia 21 6 12 1 19 15 4
South Asia 23 12 9 1 8 13 9
Oceania 6 — — 4 2 — —

Totals 149 48 (53)' 55 30(26) 84 (81) 56 (46) 45 (43)

1 The figures in parentheses are for 1961 (Lancaster, 1962a).

MODES OF SPREAD

Newcastle disease has been known to The source of an appreciable number

spread in a great many different ways. The of outbreaks has not been determined
known sources of infection are discussed (Gordon etal, 1948; Levine, 1952; Mans-
below but it is conceivable that other joer, 1961); and up to 1941 it had not
modes of spread exist and that they been possible to trace the origin of any
account for outbreaks of the disease whose initial outbreak in a country (Haddow,
origin it has not so far been possible to 1941). It is particularly hard to trace the
trace (Levine, 1952). origin of the disease with certainty when

23
 INCIDENCE OF NEWCASTLE DISEASE IN EUROPE IN 1962

(Compiled from Animal Health Yearbook FAO-WHO-OIE, 1962)

No Evidence

Low Sporadic Incidence

Seasonal Occurrence

Widespread Throughout the Country

Confined to Certain Regions

No Data

N. IRELAND

d
IRELAND

GREAT BRITAIN
itoccurs in a mild form (Levine, 1962a). Cabassi, 1960); and European house
In discussing the dissemination of New martin (Delichon urbica) (Winmill and
castle disease, Brandly (1950) has com Weddell,1961).
mented on the low incidence of the carrier Crows have shown typical nervous
state in this disease as compared with symptoms of Newcastle disease in India
human influenza in which, according to (Haddow, 1941); and they have been
Burnet (1945), one carrier individual in found dead or dying in the vicinity of out
10,000 may suffice to initiate an epidemic. breaks involving chickens in Indonesia
The world history of Newcastle disease (Picard, 1928), Ceylon (Crawford, 1931)
has shown that, in general, once the disease and India (Cooper, 1930; Sahai, 1937a).
has become established in a country or The virus has not, however, been isolated
region, it has tended to become endemic from some crows (Crawford, 1931) and
(Doyle, 1948; Lancaster, 1963a). In areas jackdaws (Keymer 1958, 1961) found
with a dense poultry population, the har dead or dying in the vicinity of other
bouring of the virus in a few survivors, the outbreaks.
resistance of the virus, and the existence of The role of the sparrow (Passer domes-
an appreciable number of potential hosts ticus) is difficult to assess. Gustafson and
are enough to account for the continuing Moses (1953a) and Hartwigk and Nitsch
emergence of new cases (Asplin, 1961 ). (1957) found that fowls developed New
castle disease after being placed in contact
with infected sparrows. Contrary results
Wild Birds have been reported by Maglione (1956).
Newcastle disease virus has a wide range Popovic (1951 ), and also Placidi and San-
of susceptible avian hosts. This has been tucci (1953a), found the sparrow was not
demonstrated in both naturally occurring susceptible to Newcastle disease virus; but
and experimentally produced infections this finding has not been supported by the
(Beach, 1946; Levine, 1952). It is import observations of Pomeroy and Fenster-
ant, therefore, that the part played by wild macher (1948) and Kaschula (1950).
birds be considered whenever the origin or Inconclusive field evidence has been re
means of spread of outbreaks is investi ported by Kee (1928). In recent years it
gated (Schoop et. al, 1955). has been unusual to find crows and spar
Newcastle disease virus has been re rows dying in the vicinity of outbreaks of
covered from the following species of wild Newcastle disease (Parnaik and Dixit,
birds: starling (Sturnus vulgaris) (Gilles 1953).
pie et al., 1950); gannet (Sula bassana) Other natural field infections have in
(Wilson, 1950); sparrow {Passer domes- volved parrots in Africa (Malbrant, 1942;
ticus) (Gustafson and Moses, 1953a; Scott et al., 1956) and jackdaws in Eng
Keymer, 1961; Monda et al, 1960); land (Keymer, 1961).
shag (Phalacrocorax aristotelis) (Blaxland, Many different species of free-flying
1951); grey parrot (Psittacus erithacus) birds have been infected artificially with
(Scott and Winmill, 1960); jackdaw Newcastle disease virus (Table 3).
(Corvus monedula) (Keymer, 1961); koel
(Eudynamis scolopaceus) (Shah and John
Infections in Birds in
son, 1959); great horned owl (Bubo
virginianus) (Ingalls et al., 1951); osprey
Zoological Gardens
(Pandion haliaetus) (Zuydam, 1952b); In an outbreak of Newcastle disease in
parakeet(Palaeornis) (Zuydam, 1952b); a zoological garden in Surabaja, Indonesia,
swan (Cygninae) (Vrtiak, 1958); parrot deaths occurred among paradise birds,
(Amazona ochrocephala) (Cavrini and pheasants, lyre birds and rice birds

25
 TABLE 3—Wild Birds Susceptible Experimentally to
Newcastle Disease Virus and Methods of Evaluation
(Modified from Gustafson and Moses, 1953b)

Name
Methods of Author
Scientific Vernacular evaluation

Bubulcus ibis Stilt bird V Placidi and Santucci (1953a)

Lophortyx Californica California quail Beach (1942)
Colinus virginianus Bobwhite quail
Perdix perdix Hungarian partridge Hl
Fenstermacher ef al. (1946)
Alectoris graeca Chukar partridge
Phasianus colchicus Ring-neck pheasant |
Gennaeus nycthemerus Silver pheasant Beach (1942)
Francolineus capensis Cape pheasant Kaschula (1950)
Columba livia Pigeon Beach (1942)
Fenstermacher et al. (1946)
V Hanson and Sinha (1952)
HI Placidi and Santucci (1953a)
Geopelia striata Japanese dove. Collier and Dinger (1950)
Stone pigeon. Mansjoer (1961)
Striped ground Kraneveld and Mansjoer (1950b)
dove Martini and Kurjana (1950)
Columba palumbus Ringed pigeon
Placidi and Santucci (1953a)
Streptopelia turtur Forest turtledove
Streptopelia Ringed dove
risoria Kaschula (1950)
Streptopelia Laughing dove D, V
senegalensis
Streptopelia chinensis Spotted dove S, D Martini and Kurjana (1950)
tigrina
Zenaidura macroura Mourning dove S, D Jezierski (1950)
Streptopelia (hybrids) Dove S, D Hanson and Sinha (1952)
Munia punctulata Prit
misoria
Munia maja zaperena Bondol S in some ;
Munia ferriginosa Bondol D and serial
Collier and Dinger (1950)
Uroloncha leucogastra Prit passages in
leucogastroides last 2 species
Padda oryzivora Glatik
Ploceus manyar Manyar
Passer melanurus Cape sparrow D, V Kaschula (1950)
Passer domesticus English sparrow S, V Gustafson and Moses (1 953a)
Mansjoer (1961)
Corvus monedula Jackdaw V.HI Baczynski (1960a)
Keymer (1961)
Corvus brachyrhynchos American crow D, HI, V Karstad etal. (1959)

S — symptoms
D —death
HI — haemagglutination inhibition activity of serum
V — virus isolation or virus transmission

26
(Mansjoer, 1961). A similar outbreak in breaks of Newcastle disease have been
Morocco resulted in deaths among chick associated with imported parakeets (Psit-
ens, pigeons and pheasants; whereas some tacidae) (Anon., 1958; Zuydam, 1952b).
predatory species and species belonging to
the orders Passeriformes and Psittaci- Introduction of Virus by
formes did not succumb (Placidi and Migratory Birds
Santucci, 1954).
Migratory birds probably introduced
In one outbreak in a zoological garden
Newcastle disease virus to Cyprus (Crow-
in Germany (Kloppel, 1963; Schoop etal.,
ther, 1952). On the sea coast of Britain,
1955), the disease appeared in specimens
shags, gannets and cormorants have been
of little owl (Athene noctura), raven
considered important agents in the spread
(Bucorvus), eagle (Haliaetus albicilla)
of Newcastle disease to poultry (Blaxland,
and kingfisher (Dacelo gigas) . Among the
1951; MacPherson, 1956b; Reid, 1955;
owls the disease was peracute, but in the
Wilson, 1950); and inland, jackdaws and
other species the disease was more chronic
starlings may have played a similar role
and caused cerebral symptoms. Although
(Keymer, 1961; Locke, 1960).
a number of different species were exposed,
it was noted that the Gallinaceous birds
Excretion of Virus in Faeces
were not affected (Kloppel 1963; Schoop
et al, 1955). A second outbreak occurred The excretion of Newcastle disease virus
in the same zoological garden some 18 in the faeces of several species of birds led
months after the first (Kauker and Siegert, Gillespie et al. (1950), Gustafson and
1957) and involved ostriches (Struthio Moses (1953a, 1953b), Karstad et al.
camelus), a vulture (Pseudogyps Africanus (1959), Seetharaman (1951b) and Spata-
Salvad) and a toucan (Ramphastos dico- lin and Karstad (1959) to believe that
lorus). Newcastle disease has also been poultry farms may become infected
diagnosed in Germany in a king penguin through the faeces of birds. A similar
(A ptenodytes patachonica ) ( Krauss et al., opinion has been expressed by Callender
1963). (1958), and also by Hanson and Sinha
In Italy, the infection was introduced to (1952), although they did not have de
a zoological garden by newly imported finite evidence to support this view. Kas-
parrots (Amazona ochrocephala) and then chula ( 1 950) has suggested that the faeces
spread to a peacock, a jungle fowl, a of wild birds may have played a part in
macaw (Ara severa) and a guinea-fowl the spread of Newcastle disease in South
(Aery Ilium volturinum) (Cavrini and Africa.
Cabassi, 1960). Dissemination by crows and hawks was
In the Bombay Zoological Gardens, likely in India (Sahai, 1937b; Seethara
India, Newcastle disease appeared in a man, 1951b) and in Pakistan
(Khan and
number of Indian partridges that had re Huq, 1963). Haddow (1941 ) has reported
cently been obtained from a dealer (Par- that in India, fowls could be infected with
naik and Dixit, 1953) and within four days virus from crows. However, Gustafson
all 35 birds died. and Moses (1952) failed to demonstrate
transmission of Newcastle disease from
Introduction of Virus by infected sparrows to chickens by close
Imported Birds association; even though the sparrow was
There is danger of spreading Newcastle naturally susceptible to the virus. Further
disease when birds are transported by air more, attempts to transmit a Malayan
from one country to another. For example, strain of Newcastle disease virus to a
in both Scotland and The Netherlands out number of species of Malayan birds have

27
given inconclusive results (Orr and John, The shipment of pheasants, partridges
1946). and other game birds has introduced New
castle disease from Calcutta to the Nether
Game Birds lands (Jansen et al., 1949; Jansen and
Game birds, and pheasants in particu Kunst, 1952; Zuydam, 1949), from the
lar, have been associated with outbreaks mainland of the United States to Hawaii
of Newcastle disease in poultry in Britain (Adler et al., 1951), from Hong Kong to
(Gillespie, 1952), France (Moine, 1950), California (Anon., 1950a), and from Spain
Canada (Moynihan et al., 1951), Italy to the United States (Thompson, 1955). In
(Brandly et al., 1946a), Germany some of these incidents, birds were dead on
(Wagener, 1948), Czechoslovakia (Jera- arrival and this facilitated early diagnosis.
bek, 1961) and The Netherlands (Jansen In one instance, imported partridges (Per
and Kunst, 1952). It has not always been dix perdix) died while being held in a
clear whether the game birds were infected quarantine station (Thompson, 1955).
by poultry or introduced infection to Experimental with Newcastle
infections
poultry. In Germany, Wagener ( 1948 ) disease virus in several species of game
thought that pheasants (Phasianidae) prob birds have been reported by Pomeroy and
ably infected domestic poultry. A similar Fenstermacher (1948) and by the authors
view was taken by Jerabek (1961) in listed in Table 3.
Czechoslovakia. However, Reid (1961)
believed that, in Britain, pheasants did not Pigeons and Doves
constitute a substantial reservoir of infec The part played by domestic pigeons
tion. (Columba livia) in the spread of Newcastle
Outbreaks of Newcastle disease in disease has been examined by a number of
pheasants have also occurred in which authors. Both laboratory and field obser
there has been little or no evidence of vations have indicated that pigeons can
association with domestic poultry (Levine spread the disease.
et al., 1947; Liebengood, 1949; Locke, Kaschula (1952c) reported that pigeons
1960; Skoda and Zuffa, 1956b; Torlone, could excrete virus in the faeces for several
1954; Vrtiak, 1958; Weidenmuller and days after being dosed with virulent virus.
Osthoff, 1953; Zuydam, 1950a), and in However, instances have been recorded
some outbreaks mortality among the where infected pigeons have not spread the
pheasants has been high (Levine et al., disease to chickens during cohabitation
1947; Uzieblo, 1961; Weidenmuller and (Popovic, 1951) unless the pigeons had
Osthoff, 1954). Furthermore, there have been infected intranasally (Schyns and
been indications of Newcastle disease anti Florent, 1951). It has also been observed
bodies in sera from pheasants raised on that pigeons and doves in contact with in
game farms (Andrews, 1963; Andrews fected chickens or contaminated premises
etal, 1963). have remained clinically healthy (Adler et
Natural infections have been reported al, 1951; Crawford, 1931; Kee, 1928; Orr
in guinea-fowl (Numididae) (Crawford, and John, 1946). Reuss (1961a) has con
1931; Farinas, 1930; Kretzer, 1930; cluded that pigeons under natural condi
Moine, 1950); in partridges (Perdix sp.) tions do not spread the virus from infected
(Mantovani and Ceretto, 1953; Parnaik poultry flocks.
and Dixit, 1953; Torlone, 1954; Vrtiak, Reuss ( 1 96 1b) and Walker et al. ( 1 954a)
1958); and in peacocks (Pavocristatus) found that pigeons infected per os excreted
(Jansen and Kunst, 1952). The partridge Newcastle disease virus for at least three
has been considered a likely carrier of the days in sufficient quantity to infect baby
virus (Placidi and Santucci, 1953a). chicks placed in contact with them.

28
Pigeons placed in direct contact with New nected with the spread of infection (Crow-
castle disease-infected chickens developed ther, 1952).
a sub-clinical infection. These pigeons, in
turn, transmitted the disease to young
Turkeys
chicks. Similar results with pigeons have
been reported by Reuss (1961a); and with Turkeys play a more important part
spotted doves (Streptopelia chinesis) by than ducks or geese in the spread of New
Kraneveld and Mansjoer (1950b). castle disease. Early reports of naturally
Naturally occurring outbreaks have occurring outbreaks in turkeys have been
been observed both in flocks of pigeons reviewed by Beaudette (1943, 1949a) and
(Hanson and Sinha, 1952; Marastoni and Fenstermacher et al. (1946). Recent re
Sidoli, 1959; Picard, 1928) and in in ports include those of Buck (1947), Gale
dividual pigeons (Iyer, 1939; Placidi and Gordon
et al. ( 1 96 1 ) , Gray
et al. ( 1948 ) ,

Santucci, 1953a; Vrtiak, 1958; Zoletto, et al.(1954), Levine et al. (1947) and
1958). Recovery of the virus from wild Walker (1948).
pigeons has been reported by Hanson and Apparently-healthy turkeys from infect
Sinha (1952) and by Vrtiak (1958). ed areas have been considered responsible
for spreading Newcastle disease in Great
Britain (Asplin et al., 1949; Gordon et al.,
Ducks and Geese
1948) and in the United States (Fenster
Cases of naturally occurring Newcastle macher et al., 1946). Asplin (1949) has
disease in ducks and geese have been re suggested that some turkeys may remain
ported (Beaudette, 1943; Bush, 1954; carriers. Turkeys imported by air are
Moine, 1 950) but these species have been believed to have transmitted a disease,
regarded as being more resistant to infec thought to be Newcastle disease, from
tion than fowls (Albiston and Gorrie, Italy to Algeria (d'Arces, 1949; Donatien
1942; Asplin, 1947; Beaudette, 1943; and Gayot, 1946). Furthermore, one in
Berthelon and Tournut, 1949; Iyer, 1945; fected turkey that was kept in isolation for
Kaschula et al, 1946). 12 months yielded virulent Newcastle dis
Artificially infected ducks and geese ease from the intestinal contents
virus
have been found to excrete virus from 3 when it was killed (Winmill and Haig,
or 4 days (Asplin, 1947) to 15 days (Tek- 1961).
linskaer al., 1956).
Ducks and geese may play a part in the
dissemination of Newcastle disease virus.
Chicken Eggs
This has been indicated by the isolation of Newcastle disease vaccine virus has been
virus from 48 out of a sample of 265 duck recovered from the surface of the shells of
tissues (Vrtiak, 1958); the recovery of eggs laid by vaccinated birds for four days
virus from the intestinal contents of ducks after vaccination (Zargar and Pomeroy,
six months after infection (Winmill and 1950b). Virus was also recovered from
Haig, 1961); and the observation that an the intestinal contents of these birds and
outbreak of Newcastle disease followed contamination may have come from this
the feeding of uncooked goose viscera source.
(Heller, 1957). Similar conclusions have Examination of tissues taken from 283
been reported from Pakistan by Khan and birds in 105 Newcastle disease-positive
Huq (1963), and from Denmark by flocks has shown 20 out of 47 ovarian
Marthedal et al. ( 1963 ) . However, in an yolks and 3 out of 10 oviduct egg yolks
outbreak in Cyprus there was no field to yield the virus (Beaudette et al., 1948a).
evidence that ducks and geese were con In addition, inflammatory changes in the

29
ovary associated with Newcastle disease (1951a) found that the majority of
infection have been reported (Biswal and embryos they inoculated with Newcastle
Morrill, 1954). disease virus died by the ninth day of
A number of authors have recovered incubation. Virus was recovered from
Newcastle disease virus, (either vaccine these embryos. No virus was recovered
or field strains) from the contents of eggs from embryos dying on the nineteenth
laid two to ten days after vaccination of day, from embryos failing to hatch, or
hens (Bivins et al, 1950; Hitchner et al, from those chicks which did hatch (Doll
1950; Prier et al, 1950; Van Waveren, et al, 1950e). Similar findings, but with
1955; Zargar and Pomeroy, 1950b). Three eggs laid during natural outbreaks, were
isolations were made from the yolks of 85 obtained by Hofstad (1949b) who con
eggs gathered at the beginning of an out cluded that embryos that survived the first
break in a laying flock (Thompson and 1 0 days of incubation were apparently free
Osteen, 1948). Eggs laid during the period from Newcastle disease virus. Embryos
when egg production was severely de with
inoculated less pathogenic strains of
pressed yielded chicks from which virus Newcastle disease virus have been known
was isolatedfour days after hatching to hatch (Asplin, 1952; Rao and Agarwal,
(DeLay, 1947; Mansjoer, 1961); virus was 1960).
also isolated from eggs laid during the
recovery period. Jungherr and Terrell Young Chicks
(1946) and Mansjoer (1961) demon Newcastle disease virus on the surface
strated the presence of virus in eggs laid of the egg, the breaking of infected eggs,
two months after an active outbreak. and the hatching of virus-infected chicks,
Field evidence of Newcastle disease are all means whereby a hatchery can be
virus being passed through the egg to the come contaminated. Newly hatched chicks
chick has been reported (Sinkovics, 1957a; from an infected hatchery have also spread
Walker and Powell, 1950). From the Newcastle disease (Callender, 1958; Jung
information available to him, Beaudette herr and Terrell, 1946). Furthermore, re
(1948a) concluded that virus — either ports have emphasized the dangers in
vaccine or field strains — is deposited in herent in the proximity of a commercial
eggs duringonly a relatively short period. hatchery to a poultry dressing plant (Jung
Furthermore, Placidi and Santucci (1953c) herr and Terrell, 1946) ; to a poultry rear
found no evidence of egg transmission of ing farm (Reid, 1955, 1961); or to a
the virus. Nevertheless, Newcastle disease brooder room (Schmittle and Mansfield,
virus is considered to have been present in 1950) . In some outbreaks in young chicks,
dried eggs (Alegren, 1951) and in com it has been impossible to determine
mercial eggs (Tiefenbacher and Woernle, whether infection occurred in the hatchery
1957) imported from a foreign country. or during transit to the farm (Jungherr,
No experimental evidence of permanent 1948; Olson et al,
1950; Walker, 1948).
egg transmission was obtained by Asplin Pomeroy and Fenstermacher ( 1 948 ) sus
(1952), Bivins et al (1950), Hofstad pected infection of chicks whilst in transit.
(1949b) or Walker and Powell (1950); Infected day-old chicks are thought to
and Beaudette (1948b) considered it safe have introduced the disease to Venezuela
to incubate eggs laid three weeks after (Divo, 1950) and Hawaii (Adler et al,
vaccination. Although some embryos in 1951) .
fected with virulent virus have hatched Older chicks have played an important
(DeLay, 1947), they usually die during part in the spread of the disease in the
incubation. United States (Schmittle and Mansfield,
Doll et al (1950e) and Hitchner et al 1950; Fenstermacher et al, 1946;

30
Pomeroy and Fenstermacher, 1948). Out (Kraneveld and Nasoetion, 1938). In pre
of 148 outbreaks in young chicks reported viously vaccinated chickens, excretion of
by Byerly (1948), 39 (approximately 26 virulent or field virus has been found to
per cent) involved the movement of occur for varying periods: 7 days (Krane
started chicks. veld and Mansjoer, 1950a); 9 days
(Asplin, 1952); 2 weeks (Woernle and
Brunner, 1957); 3 weeks (Adler et al,
Growing and Adult Chickens 1951 ) ; and 5 weeks (Van Waveren, 1955;
The following discussion of the exist Zuydam, 1951b, 1952a). The passage of
ence of Newcastle disease virus among virulent virus through the intestinal tract
growing and adult chickens is based on of immunized fowls in this way apparently
studies involving artificial exposure or does not cause any detectable change in
vaccination. its pathogenicity (Dinter and Bakos, 1953)
or virulence (Zebrowski, 1956).
From studies of this type, involving
Excretion of Virus from either live or inactivated vaccines, several
Respiratory System authors have concluded that mild or in-
The spread of Newcastle disease virus apparent infection with virulent virus is

to susceptible chickens placed in contact possible in vaccinated flocks (Adler et al,,

occurs most readily when the virus is
administered to the donor group by the
respiratory route. This indicates that trans
mission is mainly by air-borne particles
(Andrewes and Allison, 1961 ; Kohn, 1955;
Miller and Miller, 1950; Robin, 1962;
Schyns and Florent, 1951).
In immune chickens, re-infection with
virulent virus appears to be confined to the
respiratory tract and lasts from 4 or 5
days (Doll et al, 1 950c; White et al, 1 954)
to approximately 30 days (Zuydam,
1951b). In non-immune chicks, virus has
been recovered from the respiratory tract
from 6 days (Miller and Miller, 1950;
White et al, 1954) to 21 days (Walker
and McKercher, 1954b) after infection.
There is as yet no evidence that the
respiratory system acts as a permanent
reservoir of the virus, although Beach
1951; Asplin, 1952; Doll et al, 1951b;
Hofstad, 1953a; Zuydam, 1952a). Schmidt
and Bindrich (1956) estimated that in
some clinically immune chickens appreci
able virus multiplication occurred in the
organs which excreted virus. The excre
tion of virus in the presence of circulating
antibody has suggested local multiplication
of the virus in the intestinal wall, un
hindered by the circulating antibody
(Kohn, 1959).
Although they did not determine the
exact means of transmission, Beaudette
etal. (1949) and Zuydam (1953) believed
chickens vaccinated with a mesogenic
strain could be a source of infection by
contact for about three weeks following
vaccination.
When chicks have been vaccinated with
mesogenic strains, the virus has been

( 1942, 1 943 ) has claimed that adult birds, recovered from faeces for 15 days, 19 days
which recovered from the disease, carried (Zuydam, 1951b) and 21 days (Van
virus in the lungs for three months. Waveren, 1955). Hitchner and Johnson
(1948) found that the lentogenic vaccine
strain Bl did not spread from vaccinated
Excretion of Virus in Faeces chicks to other chicks in a separate com
During the early outbreaks of Newcastle partment of the same brooder. Although
disease in Indonesia, the virus was found this Bl strain was excreted in the faeces
in the faeces of chickens suffering from for 7 days (Kohn and Ebert, 1960) to 14
naturally occurring cases of the disease days (Van Waveren, 1955), direct physi

31
 SUGGESTED SCHEME FOR THE PATHOGENESIS OF
NEWCASTLE DISEASE VIRUS
(Redrawn from Kohn, 1959)

AIR

FOOD AND WATER

MUCUS IN
THROAT
FAECES AN D NOSE
I
INGESTED INHALED

INTESTINAL UPPER RESP.

MUCOSA MUCOSA

a
EtN FECTION

-L=s{
BLOOD
H
[VISCERA SPLEEN LUNGS

Figure 6.

cal contact was necessary for spread to by wire partitions has been reported ( Ban
susceptible chicks (Bankowski et al., 1957; kowski, 1957; Bankowski et al, 1958a;
Marek, 1960; Markham et al., 1951). Bankowski and Corstvet, 1960). Non-
Contact transmission ceased two weeks vaccinated chicks experimentally infected
after the re-vaccination of chicks with the with field virus excreted virus in the faeces
Bl virus(Hitchner and White, 1956). on the 11th day (Jungherr, 1948); while
Baldelli (1956) concluded that, after oral adults ceased to spread virus by contact
administration, Strain F virus (Asplin, after 34 days (Walker and McKercher,
1952) multiplied first in the lungs, and 1954b).
nine to ten days later was eliminated in Experimental and field data of the type
the faeces. reviewed above suggest that the permanent
Complete absence of spread of a modi carrier state in Newcastle disease is rare
fied live Newcastle disease vaccine ad in chickens (Brandly, 1950; Crowther,
ministered intranasally or intramuscularly 1952; Hofstad, 1951; Jungherr, 1948;
from the vaccinates to non-vaccinated Konev, 1953; Levine, 1952; Nitzschke,
chickens within the same pen or separated 1954; Placidi and Santucci, 1955; Piatt,

32

J
1948). Even if this is true for chickens, did not spread to susceptible chickens and
there may be a more lasting carrier state which did not cause parental antibodies
in turkeys. in young chicks has also been reported
A suggested for the patho
scheme (Bankowski, 1961c). This unusual type
genesis of Newcastle disease virus follow of infection was identified by serological
ing respiratory and alimentary infection methods.
has been given by Kohn (1955,
1959) The possibility that hens can carry non
who believed that multiplication of the pathogenic Newcastle disease virus and
virus in the intestinal tract would occur excrete this virus in their eggs has been
after respiratory infection (Figure 6). The discussed by Sinkovics (1957a). Similar
upper respiratory tract has been consider latent Newcastle disease virus infections
ed the site of election for the early multi have been found in cultures of tissue cells
plication of the virus (Maestrone and inoculatedwith a strain of virulent virus
Coffin, 1961). Kohler (1960) reported (Mason and Kaufman, 1961a).
that Newcastle disease virus was ingested Further mention of asymptomatic infec
by leucocytes, but that the virus did not tions is made on page 57.
appear to multiply within these leucocytes.

Transport of Live Poultry

Reservoir of Virus in Aqueous The movement of domestic poultry has
Humour been a very important factor in the spread
of Newcastle disease. Spread by this means
Clark et al. (1955) found that the
has been reviewed by Beaudette (1943,
aqueous humour of the chicken eye was a
1949a, 1950, 1951b), Brandly et al.
source of virus for 10 days after intra
(1946a) and Brandly ( 1950, 1959). More
muscular inoculation. They reported later
recent reports include those of Anon.
(1957) that they believed the aqueous (1961a), Crowther (1952), Scott et al.
humour was a reservoir for Newcastle
( 1956 ) ,Reid (1961), Vandemaele ( 196 1 ) ,
disease virus in a large percentage of
and Winmill and Haig (1961).
chickens in California. However, more
Of 2,489 outbreaks of Newcastle dis
recent findings have indicated that this
ease in Britain, 56 per cent were a result
material does not constitute a permanent
of traffic in live poultry. Field evidence
reservoir of the virus (Baldwin, 1962;
suggested that in these outbreaks some
Dardiri et al, 1959; Fritzsche, 1961;
recovered birds acted as carriers (Asplin
Pannu and Bankowski, 1962).
etal, 1949).
Levine (1952) considered the actively
Asymptomatic Infections infected bird to be the most important
source of infection. Similarly Haddow
Asymptomatic infections with New
( 1941 ) , in discussing the original outbreak
castle disease virus have been reported in
at Ranikhet, India, suggested that the
which spread of virus could be detected.
infection might have been due to the
For example, chickens hatched two or
importation of poultry from abroad four
three months after an outbreak of clinical
months previously.
Newcastle disease had subsided proved to
be serologically positive. Although no
clinical evidence of disease had been Poultry Markets
observed in the young stock, it was con The significance of poultry markets in
cluded that inapparent spread by carriers the spread of Newcastle disease has been
had occurred (Jungherr and Terrell, 1946). recorded in all geographical regions since
Asymptomatic Newcastle disease which the disease was first recognized (Doyle,

33
1927; Albiston and Gorrie, 1942; Farinas, Illegal Movement
1930; Lissot and Moessel, 1949; Seethara-
Finally, the illegal movement of live
man, 1951b). In Australia, almost every
birds from infected premises or areas has
outbreak has been associated with markets
resulted in new foci of infection (Jansen
(Johnstone, 1931). An analysis of 602
and Kunst, 1952; Reid, 1955, 1961).
outbreaks in England and Wales showed
158 to be primary cases; and from these
primary outbreaks a further 444 outbreaks
resulted — mainly from local spread, direct
Poultry Carcasses and Offal
sales, dealers' transactions and sales Poultry carcasses and offal have been as
through markets (Reid, 1 955). Other out great a source of Newcastle disease infec
breaks of Newcastle disease have been tion as live poultry and they have often
traced to a poultry packing station (Key- carried the disease from one country to
mer, 1961) and to the purchase of fowls another. In Switzerland, where offal from
from ships in dock (Lucam, 1949b). chickens of foreign origin has been con
sidered a highly important source of in
fection, a diagnostic method was develop
Laying Trials
ed to detect infected imports of chicken at
Newcastle disease outbreaks associated the frontier (Hess, 1963). From every
with the movement of live poultry have imported consignment, 30 frozen heads
been reported at laying trials (Beach, per 5 tons of product were examined in
1949a, b; Piatt, 1948). In some trials, birds the laboratory. For the examination,
were identified which had recovered from material from the spinal cord was injected
Newcastle disease but which were, pre into young susceptible chicks. During the
sumably, not excreting virus (Asplin et al., period 1947-62, samples from 21,300 tons
1949; Reid, 1955; Jungherr and Terrell, of imported chicken were examined in
1946). When these birds were traced, a Switzerland (Hess, 1963). Similarly in
subacute form of Newcastle disease was Austria, the importation of refrigerated
discovered in a hatchery (Anon., 1962b). poultry carcasses has been considered the
Furthermore, the tracing of birds moved most important source of initial infection
from a large poultry show in Britain dis (Grausgruber, 1963). In Germany, the
closed 255 outbreaks of Newcastle disease laboratory examination of imported poul
(Reid, 1955). try carcasses has yielded a strain of New-

TABLE 4 — Probable Origin of the First 542 Outbreaks of

Newcastle Disease occurring in England and Wales during 1947
(Gordon et al., 1948)

Origin Per cent

Traffic in live poultry Dealers sales, auction markets and pet shops 42
Feeding infected material Unboiled swill, poulterers waste and table poultry 33
trimmings
Local spread Contiguous premises and mixing and/or straying 28
of poultry
Mechanical spread Visits and handling of birds by dealers and others, 7
infective clothing, crates, etc.
Obscure — 10

34
TABLE 5 — Newcastle Disease Virus Isolated from Poultry Carcasses
Imported into England in 1949 (Dobson, 1952)

Per cent carcasses

infected

Young fowls 69
Hens 66
Old cocks 80
Geese 6.9
Ducks 11
Turkeys 24

castle disease virus pathogenic for chicks resistant fowls; of processing in an infect
(Hartwigk and Gothe, 1958). ed environment; or of surface contamina
Poultry offal has been an important tion from a smaller number of infected
mode of spread not only between countries fowls.
but also within a country. Doyle (1927) Topacio and Coronel (1939) found that
mentioned that the flock involved in the the crop contents and the organs of dead
original 1926 outbreak in England had birds remained infective for seven days
been fed offal collected from the seaport after death. In frozen carcasses, the virus
town of Newcastle. The origin of the 1933 has been recovered from the brain (Hess,
outbreak in England was not determined 1963; Tienfenbacher and Woernle, 1957)
(Dobson, 1939). However, in the 1947 and from spinal cord material (Hess,
outbreaks in England there was evidence 1963). The spread of Newcastle disease
that poultry had had access to uncooked virus from stored edible carcasses to a pen
offal from uneviscerated poultry carcasses of pullets has been described by Gordon
imported 13 days previously from Hun and Asplin (1947).
gary, Poland and other countries where Frozen poultry carcasses are also
Newcastle disease was known to exist thought to have introduced Newcastle
(Andrews, 1948; Gordon and Asplin, disease to Germany (Koser, 1942;
1947; Gordon et al., 1948; Reid, 1961). Wagener, 1948), Sweden (Alegren, 1951;
In 33 per cent of the first 542 outbreaks, Bakos and Nordberg, 1949; Isaksson et al.,
there was a history of poultry having had 1948) and Switzerland (Hess, 1951).
access to offal from imported carcasses Samples taken from chicken carcasses im
(Table 4). Less than 5 per cent of later ported into Switzerland resulted in the
outbreaks were associated with the feeding isolation of Newcastle disease virus from
of poultry offal, whereas outbreaks origi 350 consignments representing a total of
nating from stock movement increased to 1,900 tons of product (Hess, 1963). Hess
over 70 per cent (Gordon et al., 1948). (1963) believed that the absence of New
In a 1949 survey of samples of skin castle disease in Switzerland was largely
taken from frozen carcasses imported into the result of testing samples of imported
England from two countries, Newcastle poultry carcasses.
disease virus was recovered from several Under experimental conditions, the
species of poultry, as shown in Table 5 virus has remained viable in the bone
(Dobson, 1952; Dobson and Simmins, marrow of fowls for 300 days or longer,
1951; Reid, 1961). Asplin (1952) con depending on the storage temperature
sidered that these findings could be the (Table 6). These findings explain how the
result of subclinical infection in partially holding of live poultry at killing plants

35
 TABLE 6 — Reports of Duration of Viability of
Newcastle Disease Virus in a Variety of Tissues

Author Material Storage Duration

temperature viability
(days)

Asplin, 1 949 Bone marrow 34-35"F 134-196

Bone marrow and skin — 4°F Over 300
Skin 34-35'F 98-1 60
Beach, 1943a Lungs Live chickens 60-90
Dalling, 1960 Poultry carcasses Frozen Over 730
Doyle, 1 933 Bone marrow and muscle Chilled 180
Doyle, 1 927 Kidney of infected fowl Approx. 2°C Over 1 00
Iyer, 1 940 Liver and spleen of infected 17'C 21-28
fowl
Hess, 1951 Poultry carcasses Frozen 180
Hartwigk and Gothe, 1958 Infected poultry carcasses — 20°C Over 836
Hartwigk and Gothe, 1958 Infected poultry carcasses Buried in earth 121

prior to slaughter has facilitated spread reports on the isolation of Newcastle

(Reid, 1961). disease virus from a case of conjunctivitis
Although, in general, studies of virus in
man was written by Ingalls and
survival in carcasses have been concerned Mahoney (1949).
with the more virulent strains of New Steele (1959) has pointed out that in
castle disease virus, results reported by respect of Newcastle disease infections, the
Foster and Thompson ( 1957) indicate that milder the strain of virus the broader the
the less virulent strains would probably host spectrum. Thus, the velogenic strains
survive just as long. seldom affect man, whereas the lentogenic
Bl strain has caused severe conjunctivitis
Spread by Human Agency and malaise in man (Dardiri et al, 1962).
Furthermore, re-examination of 1 7 strains
Outbreaks of Newcastle disease in
of atypical influenza virus from human
chicken flocks have been associated with
cases has shown these to be neutralized by
human movement from infected premises
Newcastle disease serum and to have other
(Lancaster, unpublished data). The move
properties resembling Newcastle disease
ment of seasonal land workers from Italy
virus (Solov'ev etal., 1963).
was probably responsible for introducing
The disease in man may be an influenza
the disease to Germany (Wagener, 1948).
like illness
accompanied by headache
Also, Hudson (1937a) and Callender
(Howitt et al, 1948; Negri et el, 1953).
(1958) have reported that infection can
On the other hand, Newcastle disease virus
easily be conveyed from sick to healthy
has been isolated from a human patient
stock by handling.
whose illness was almost inapparent
(Quinn et al, 1952). Although they did
Human Infection
not isolate the virus, Howitt et al. (1948)
Reviews of reported cases of Newcastle have suggested that Newcastle disease
disease in man have been published by virus may be responsible for some atypical
Blood (1950), Evans (1955a), Hanson central nervous system infections in man.
and Brandly (1958), Sinkovics (1957a) The more common symptom is a unilateral
and Thompson ( 1 950) . One of the earliest conjunctivitis (Gustafson and Moses,

}6
1951; Jacotot et al, 1950). In one in haemagglutination at a dilution of 1 : 10 or
stance, conjunctivitis due to Newcastle higher (Collier, 1951). In assessing the
disease virus developed in a laboratory significance of such findings, it must be
worker who had been infected in similar emphasized that there is evidence indicat
circumstances four and a half years earlier ing that mumps virus and Newcastle
(Jacotot et al, 1955). Other cases of con disease virus have a common antigen or
junctivitis, with or without enlargement antigens. Thus, neutralizing and antihae-
of the adjacent lymph nodes, have been magglutinating antibodies against New
reported by Anderson (1946), Burnet castle disease virus were shown to be
(1943), Freymann and Bang (1949), Ku- present in half a group of patients con
jumgiev (1948), Lippman (1952), valescing from mumps (Jungherr et al,
Reagan et el. (1956), Schoop (1954) and 1949). In contrast, other reports have
Slonim and Stranakova (1952). indicated that cross reactions with mumps
In many cases of human infection, an and Newcastle disease viruses cannot be
immune response has been observed (Bang due to common antigens (Wenner et al,
and Foard, 1956b; Blood, 1950; Hunter 1952a, b).
et al., 1951; Jordan and Feller,
1950). A heat stable factor, with complement-
Also, inclusion bodies have been seen in fixing as well as antihaemagglutinating
the cytoplasm of epithelial cells from the activity against Newcastle disease virus,
conjunctiva, though their significance has increased in titre in the sera of a number
not been fully established (Hunter et al., of patients convalescing from mumps
1951; Keeny and Hunter, 1 950; Orlandella, (Jordon and Feller, 1950). Similarly, sera
1955). In some instances, human infec from mumps patients have shown New
tionhas followed the accidental inges castle disease virus antibody when tested
tion of Newcastle disease virus (Divo and by virus neutralization (Kilham et al,
Lugo, 1952). 1949). In addition, a viricidal factor for
During the acute phase of infection, the Newcastle disease virus and a component
virus has been recovered from eye wash haemagglutination by that virus
inhibiting
ings (Jacotot et al., 1950), blood (Negri have been identified in the normal sera of
et al, 1953; Reagan et al, 1956), urine man and some other mammals (Howitt,
(Dardiri et al, 1962), saliva (Divo and 1950; Wenner et al,
1952a, b). Therefore,
Lugo, 1952) and respiratory tissues. In it has been concluded that the serologi
one instance, an outbreak of Newcastle cal diagnosis of Newcastle disease in man
disease conjunctivitis involved 40 persons must be made with caution in the absence
employed in the evisceration of poultry of virus isolation (Evans, 1956; Jordan
(Nelson al,
1952). Nevertheless, there
et and Feller, 1950; Jungherr et al, 1949;
is no evidence of the existence of the car Kilham era/., 1949).
rier state in man. Neither is there any evi It has been emphasized that in many
dence of transfer from one person to an human infections with Newcastle disease
other (Yatom, 1946), although this virus, the antibody response has been low
possibility has been discussed (Hanson and (Anderson, 1946; Freymann and Bang
Brandly, 1958; Mitchell, 1953). 1949; Slonim and Stranakova, 1952).
A survey of 1,363 samples of human Further, Newcastle disease virus may
serum revealed 13 to contain haemag- share antigenic components with other in
glutination-inhibiting and complement- fectious agents (Evans, 1951). Thus, it
fixing antibodies against Newcastle disease has been shown that serum from human
virus (Scatozza, 1957). A similar survey patients convalescing from infectious
showed that 50 per cent of human sera mononucleosis will agglutinate human red
samples inhibited Newcastle disease virus blood cells modified with Newcastle

37
disease virus (Florman, 1949). This Ferrets and Guinea- Pigs
capacity to agglutinate Newcastle disease Although ferrets have been found sus
virus-treated cells is unrelated to the anti ceptible to Newcastle disease virus,
body of infectious mononucleosis (Evans Reagan et al. (1950c) have reported that
and Curnen, 1948). The red cell sensitiz the virus could not be adapted to this
ing property of Newcastle disease virus is species. In some transmission experiments,
distinct from the virus haemagglutinin guinea-pigshave proved to be insuscep
(Evans, 1955b). tible (Wenner and Lash, 1949).

Hamsters
Warm-Blooded Animals
A number of workers have reported the
Several different species of mammals susceptibility of hamsters to Newcastle
have been used in laboratory studies disease virus. Thus, Reagan et al. ( 1 947b,
(Verge, 1948). The results of these studies 1948a, 1950c) passaged a virulent strain
indicate that the ability of Newcastle of virus for 300 passages in the brains
disease virus to multiply in mammalian of Syrian hamsters. The virus produced
hosts is limited (Hanson et al, 1951a; symptoms of irritability, paralysis and
Reagan 1952b). In general, New
et al., rapid death (Reagan et al., 1947a). The
castle disease does not appear to be spon amount of virus required to produce in
taneously transmissible to mammals under fection in hamsters varied with the route
natural conditions 1948). Up to
(Verge, of inoculation. Intracerebral inoculation
now, man has been the only mammalian was found a successful route (Prudovsky
species in which a number of cases of et al., 1961), and 10"10 ml. of virulent
Newcastle disease virus infection have oc virus killed hamsters within 4 to 16 days
curred naturally (Blood, 1950). Also, when administered by this route (Mitroiu
there is no evidence as yet to indicate that and Vior, 1960). Following the intracere
mammals play a part in the spread of the bral inoculation of attenuated Newcastle
disease. However, according to Brandly disease virus, an active immunity against
(1959), the studies described below in challenge infection was produced (Mitroiu
dicate the potential range of infectivity of and Vior, 1960); this was also demon
the virus. Other references to the propaga strated by virus neutralization tests (Rea
tion of Newcastle disease virus in mam gan etal., 1947a).
malian tissue are given on page 1 18.

Horses
Bats Horses have been inoculated with New
The cave bat (Myotus lucifugus) has castle disease virus to produce hyperim

been found susceptible to intracerebral mune serum (Lulic, 1955). This subject
is discussed further on pages 87 and 88.
inoculations with a number of different
strains of Newcastle disease virus (Reagan
et al., 1950a). The virus has also main Mice
tained its pathogenicity for chicken em The successful adaptation of the virus
bryos following intracerebral passage in to unweaned mice following intracerebral
bats (Reagan et el., 1951a). The same inoculation has been reported (Kilham,
species of bat was susceptible to the Cali 1950; Sinkovics, 1960). This mouse-adap
fornia strain 11914 of virus administered ted virus did not propagate in the brain
by intranasal instillation, and typical symp of adult mice and there was evidence of
toms of irritability, paralysis and death re decreased pathogenicity for chicks (Sin
sulted (Reagan and Brueckner, 1951a). kovics, 1960).
Monkeys fox and a dog on the first and fifth day
The intracerebral inoculation of Rhesus after feeding on carcasses of fowls that
monkeys with Newcastle disease virus has died of Newcastle disease (Polci and Sil-
generally resultedin a meningoencepha vagni, 1954). Baldelli (1955) has re
litis (Prudovsky et al., 1961; Rodot, 1953; ported finding that young puppies were
Wenner and Lash, 1949). Intramuscular susceptible to intracerebral inoculation
injection has produced little or no clinical with a strain of Newcastle disease virus. In
effect (Reagan et al., 1951b). Sublethal contrast, other reports have indicated the
infections in monkeys have resulted in the relative insusceptibility of dogs and cats
appearance of haemagglutination-inhibit- (Bonaduce, 1954a; Mitev et al, 1958;
ing antibodies et al., Orlandella, 1954; Rahneberg, 1960; Rea
(Wenner 1952a).
gan and Brueckner, 1952).

Shrews and Rabbits

Hedgehogs
Shrews (Marina brevicauda) (Reagan
The finding, in Morocco, that a North
and Brueckner, 1951b) and rabbits (Wil
African hedgehog (Aethechinus algirus)
cox et al., 1958) have also been used in
became infected after eating another that
laboratory studies.
had died from the disease has suggested
that this animal might act as a reservoir
Cats
of the virus in Morocco and also, perhaps,
The susceptibility of the domestic cat
other countries (Placidi, 1954a).
to Newcastle disease virus has been re
ported by Feldberg and Luttrell (1958); Pigs
and Bolin (1948) isolated Newcastle In studying the part played by the
disease virus from the faeces of a cat
domestic pig in the spread of Newcastle
five months after it was fed infected disease, Asplin (1949) found that, follow
chicks. Furthermore, Nakamura and inoculation,
ing intramuscular virus was
Iwasa (1942) believed that a laboratory excreted of one pig for a
in the urine
cat died of a natural infection caused by maximum of 48 hours. The experimental
Newcastle disease virus. Subsequent inoculation of Newcastle disease virus into
experiments on the infection of cats per os
young pigs has resulted in death 3 to 11
showed that these animals are susceptible
days later (Hofstad, 1950; Placidi,
to the virus: Luttrell and Bang (1958)
1954b), or the development of nervous
and Tokuda (1956) reported that young
symptoms (Bindrich, 1957) and paralysis
and adult cats could be infected with New
(Buck et al., 1954a). Newcastle disease
castle disease virus administered by differ
virus passaged intracerebrally in young
ent routes, and Verge (1948) also con
pigs for eight passages has resulted in the
cluded that cats were naturally susceptible virus becoming for the pig via
pathogenic
to the virus. However, these results were
the nasal route. Fourth to seventh passage
not confirmed by Reagan et al. (1954d)
virus has produced neither disease nor im
who found that kittens were not suscep
munity in chickens (Buck et al., 1954b).
tible to infection with a strain of virus ad
The absence of Newcastle disease haemag-
ministered by the oral or nasal routes,
glutination-inhibition antibodies in 168
but were susceptible to intracerebral in
pigs would suggest lack of contact with the
oculation (Reagan et al., 1954a).
virus. The subcutaneous injection of an
avirulent Newcastle disease virus into a
Dogs and Foxes young pig resulted in the development of a
Newcastle disease virus has been re positive but low Newcastle disease HI titre
covered from the urine and faeces of a (Dyml, 1958).

39
 Rats
The oral dosing or feeding of the com
mon brown rat with Newcastle disease
virus or with chickens dead from the
disease has resulted in the virus being shed
in the excreta for 24 hours (Asplin,
1949), 48 hours (Zuydam, 1951a), 72
hours (Walker et al, 1954c) and 5 days
(Baczynski, 1959). Newcastle disease
virus was not isolated from faeces and
tissues of 12 rats taken from a Newcastle
disease-infected farm (Zuydam,
1951a).
From these studies it was concluded that,
althoughthe rat is generally only a me
chanical carrier, it could be responsible
for causing the disease in chickens
(Placidi andSantucci, 1953c).
Cattle
Two cases of naturally occurring infec
tions with Newcastle disease virus have
been reported in calves (Ozawa and Chow,
1958; Yates et al, 1952). In both instances
the virus was isolated from the lungs of
dying calves which had respiratory symp
toms. Although calves have been artifi
cially infected with Newcastle disease
virus (Helmboldt et al, 1955), there is
no confirming evidence that cattle play
any part in the spread of the disease.
Several studies have been made on the
propagation of Newcastle disease virus
in the lactating mammary gland of cows
and goats (Mitchell et al, 1953a, 1953b,
1954, 1956, 1958; Easterday et al,
1959). The virus has persisted in the
mammary gland for varying periods up
to approximately two weeks and, in most
cases, it has stimulated the production of
neutralizing antibody. The mammary
gland has appeared to be the principal
site of antibody production (Mitchell et
al, 1956).
Cold- Blooded Animals
Reptiles and Fish
Qureshi (1957) found no evidence of
a carrier state following the intracerebral
40
injection of Newcastle disease virus into
goldfish (Carassius auratus), garden
snakes (Thamnophis sirtalis) or baby
turtles (Graptemya geographicalesuerr) .
However, Placidi (1956a) has reported
that the virus of Newcastle disease could
be recovered for several months following
the intracerebral inoculation of the tor
toise (Testudo graeca and Clemmys le-
prosa) and three species of snakes {Natrix
natrix, Natrix viperina and Malpolon
monspessulana) . The virus has also been
propagated in the green turtle (Pseudemys
elegans) (Reagan et al, 1953).
The intracerebral inoculation of New
castle disease virus into the dog fish (Scyl-
lium canicula) has been followed by re
covery of the virus during the subsequent
three weeks. Lesions of meningitis have
been found in the brain, but without any
clinical symptom (Atanasiu and Atanasiu,
1955).
Invertebrates
Insects. Studies on invertebrates, and
on ectoparasites in particular, indicate
that they play a similar role to mammals
in the spread of Newcastle disease. Hof-
stad (1949a) showed that the northern
feather mite (Liponyssus sylviarutn) har
boured the virus after feeding on infected
birds during the viraemic stage. However,
the bite of known infected mites failed
to transmit Newcastle disease to suscep
tible birds. A
similar absence of natural
transmission has been reported for the tick
(Argas persicus) (Komarov, 1940; See-
tharaman, 1951b). Lissot and Moessel
(1949) were of the opinion that ectopara
sites (genera Menopon and Lipeurus)
played no role in the spread of Newcastle
disease. However, Bolin (1948) has re
ported the isolation of Newcastle disease
virus from common chicken lice collected
from hens 35 days after infection with the
virus. In the early outbreaks in the Dutch
East Indies, flies and mosquitoes apparent
ly played no part in spreading the disease
(Picard, 1934).
 Worms. In earthworms (genus Helo- ers or pens two to seven weeks after the
drilus), Newcastle disease virus has been removal of infected chickens (Dobson,
found to survive approximately four days 1939; Jungherr, 1948; Jungherr and Ter
(Baczynski 1960b; Boyd and Hanson, rell, 1946). Levine et al. (1950) con
1958), or 18 days if the worms were kept cluded that premises with birds recovered
at 21°C (Boyd and Hanson, 1958). Al from Newcastle disease do not harbour
though virus fed to planaria (Planaria sufficient virus to infect susceptible birds
maculata) has been recovered after nine one month after the flock recovers from
days there has been no evidence of virus the respiratory symptoms. In poultry
multiplication (Boyd and Hanson, 1958). houses previously occupied by infected
The part played by the large intestinal stock, Newcastle disease virus has sur
round worm (Ascaridia galli) in the vived no more than 7 days during sum
transmission of Newcastle disease virus mer, 14 days during spring and 30 days
was examined by Stefanski and Zebrowski during winter (Fortner, 1952). Schyns
(1958) who found that the virus could and Florent (1951) and Hudson (1937a)
be recovered from these nematodes re also found the virus in uncleaned pens to
moved from chickens dying from New be inactive after 6 to 14 days. In contrast,
castle disease. The virus was not recovered Michalov and Vrtiak (1963) found that
from the eggs of the worms, and there virulent Newcastle disease virus survived
fore A. galli was not considered a true 53 days in hen house litter. Using strains
vector of Newcastle disease. of vaccine viruses, Dardiri et al. (1957)
Snails. The finding that Newcastle failed to recover the virus after 24 hours
disease virus remained alive for several from sterilized dried droppings, shavings,
weeks in the body of the Indonesian giant feathers and feed samples which had
snail (Achatina fulica) has suggested that been placed in a brooder house. Hitchner
the snail might spread the virus (Mans- (1950) also reported failure of the Bl
joer, 1961). strain of vaccine virus to survive in a hen
house after removal of the vaccinated
Inanimate Causes chickens.
Infective secretions and excretions of Under experimental conditions, New
Newcastle disease virus from the digestive
castle disease virus on small amounts of
and respiratory tracts, and, to a lesser
chick down in stoppered vials has re
extent, from the reproductive tract of
mained viable in a hen house for 255 days
poultry, are the chief causes of contamina
(Olesiuk, 1951). At 37°C, Newcastle
tion of buildings and equipment. Various
disease virus has remained viable in the
chemical and physical conditions influence
contents of an egg after 94 to 126 days
the viability of the virus outside the host.
(Olesiuk, 1951) or on pieces of egg shell
Therefore, much of the data referred to
for 1 day (Asplin, 1949) or 7 days
below relates only to the particular experi
(Olesiuk, 1951). The virus has remained
mental and field conditions The
involved.
survival of Newcastle disease virus in the viable on non-sterile burlap for 20 days

environment plays an important part in (Jungherr, 1948); and on sterile burlap

perpetuation and spread from 55 days (Jungherr, 1948) to 108
its (Hanson
and Brandly, 1958). days or longer (Olesiuk, 1951), depend
ing on the storage temperature. It has
Chicken Houses, Crates and been shown that Newcastle disease virus
Brooders will survive for at least 24 hours on pieces
Day-old chicks have become infected of egg shell when exposed to incubator
when placed in uncleaned battery brood conditions (Asplin, 1949; Beamer et al.,

41
 TABLE 7 — Duration of Viability of Newcastle Disease Virus
(Modified from Olesiuk, 1951) Duration of study: 538 days

Test material Incubator Normal room Hen house Refrigerator Deep freeze
37°C 20° to 30"C — 11'to 36'C 3° to 6'C — 26X
(days) (days) (days) (days) (days)

Burlap 12 13 108 123 538

Filter paper 19 37 129 157 538
Cloth 13 44 145 193 538
Broth 30 152 228 538 538
Saline 26 79 228 451 538
Eggshells 7 44 228 258 538
Mash 54 61 172 258 538
Chicken Faeces 41 83 172 538 538
Down 58 163 255 538 538
Soil 14 66 172 235 538
Eggs 110 216 255 538 —
A.A.F' 33 80 199 258 538

1 Undiluted amnioallantoic fluid

1949). The duration of viability on other of spread of the infection to a second

is given in Table 7.
test materials farm on which 72 per cent of 18,000
Asplin (1961) considered that survival chickens subsequently died (Jungherr and
times obtained in the laboratory were Markham, 1962). Farinas (1930) was
probably longer than would be expected able to infect chickens, geese, turkeys and
from field experience. Under farm condi ducks by using boxes that had previously
tions, the presence of decomposing contained native chickens bought in a
organic matter, exposure to light, and public market.
variations in humidity probably shorten
the survival time. Nevertheless, Olesiuk
Wind
(1951) considered it likely that the virus
would survive in the natural environment In areas where the poultry population
from one season to the next. is dense, Newcastle disease virus may be
The survival of Newcastle disease virus carried from one location to another by
in the soil depends on several factors, but the wind (Anon, 1962b; Callender, 1958;
has been found to be from 8 to 22 days Moynihan et al., 1951). This wind-borne
(Boyd and Hanson, 1958). Survival and spread has been particularly noticeable
spread outside the host is also facilitated in countries where large numbers of
by the high resistance of the virus to direct chickens are housed in buildings ventilated
sunlight (Iyer, 1943; Skinner and Bradish, by exhaust fans (Reid, 1961). Spread
1954) ; to a wide range of pH values, from between contiguous premises has also
pH 2 to 11 (Moses et al., 1947; and to been recognized (Jungherr and Terrell,
effects of temperature (Brandly, 1959). 1946; Moynihan et al, 1951; Reid,
Temperature is an important factor and 1961 ), although the exact means involved
its effect is summarizedin Table 8. was not ascertained.
Empty poultry crates returned from a Experimental data supporting the field
farm on which an outbreak of Newcastle evidence have been obtained. DeLay et al.
disease had occurred were the likely means (1948), DeOme et al. (1948) and Mans-
42
TABLE 8 — Effect of Temperature on Infectivity of Newcastle Disease
Virus (Modified from Anon., 1959)

Temperature
Probable stability in time Authority1 and record

60 140 0-5 minutes Hanson — Roakin strain, 5 min. (A)

56 132 30-180 minutes Hanson — GB strain, 180 min. (A)
Roakin strain, 30 min. (A)
B1 strain, 1 5 min. (A)
45 113 12-90 hours Hanson — GB, Roakin and B1 strain, persisted
1 2 hours but not 96 hours. (A)
37 98 6-14 days Hanson — GB, Roakin and B1 strains, persisted
14 days. (A)
Jungherr — Conn, strain, 6, 24 and 72 hours. (C)
Olesiuk — Mass. strain, 10 days. (C)
Torlone — Strain F — titre reduced after 24
hours. (A)
25 77 30-1 00 days Prier — NDV strain, 73-95 days. (A)
Olesiuk — Mass. strain, 9-16 days. (C)
Torlone — Strain F — titre reduced after 48
hours. (A)
Lancaster — Strain F, 30 days. (B)
41 8-1 6 months Jungherr — Conn, strain, 203 days. (C)
(on filter paper 34F)
Prier — NDV strain, 80-1 1 9 days. (A)
Olesiuk — Mass. strain, 123 days. (C)
Reising — B1 strain, titre reduced in 6 months.
0 32 9 months Lancaster — Strain F. (D) ^
— 20 —4 1-10 years Hanson — NDV strains, 4-6 years. (B)
Reising — B1 strain, high titre retained 1 year.
(B)
Lancaster — Strain F, 1 year. (D)

A — Allantoic fluid in sealed glass vials.

B — Allantoic fluid in stopper glass tubes.
C — Allantoic fluid dried on paper or cloth.
D — Allantoic fluid lyophilized.
1 Hanson, 1 955 ; Jungherr,
1 948 ; Lancaster, 1 957a ; Olesiuk, 1 951 ; Prier and Alberts, 1 950 ;
Reising and Hitchner, 1954; Torlone, 1955.

joer (1961) demonstrated the presence Sinha, 1952; Sinha et al, 1957). Sinha
of Newcastle disease virus in air and dust et al. (1952, 1954) recovered Newcastle
samples collected from a poultry house disease virus from the air in the vicinity
containing diseased chickens; and Idnani of infected chickens on the third to sixth
and Seetharaman (1947) used susceptible day after exposure. Under experimental
chickens kept a short distance from an conditions, it was shown that with each
infected bird. minute of aerosol exposure, chickens
would be expected to inhale five particles
Aerosols of Newcastle disease virus (Sinha et al.,
Artificially- and naturally-produced 1954) . Such low concentrations of virus in
aerosols transmit the disease readily the air may be readily infective (Nadel
(Andrewes and Allison, 1961; Hanson and etal, 1957).

41
Water Poultry Vaccines as a Means
of Spread of Newcastle
The spread of Newcastle disease by Disease
contaminated water introduced by the
irrigation of meadows was considered Failure to inactivate a virulent New
likely by Koser (1942). Similarly, castle disease virus completely when pre
Grausgruber (1963) considered that paring a killed vaccine has led to some
rivers contaminated by infected carcasses serious results in the field (Mitchell et al.,
represented a means of dissemination. 1952; Placidi, 1956b; Surin, 1959).
Kraneveld and Mansjoer (1950b) found Henderson (1952) considered that the
that water contaminated with faeces from absence of active virus in an inactivated
infected birds remained infective for at vaccine could never be proved absolutely.
least five days. Idnani and Seetharaman Furthermore, instances have occurred
(1947) also showed that chickens could where commercially manufactured New
be infected by drinking contaminated castle disease, fowl pox and laryngotra-
water. In the Philippines, an important cheitis vaccines have been found to be
source of infection was thought to be the contaminated with a virulent Newcastle
common drinking ponds (Farinas, 1930). disease virus (Zargar andPomeroy,
Thus, Kee (1928) found that healthy 1950a; Brandly, 1959; Hofstad, 1954a;
chickens confined in separate pens near Marek, 1961). Furthermore, a poultry
the diseased birds, but with separate drink vaccine contaminated with Newcastle
ing troughs, did not contract Newcastle disease virus is known to have resulted
disease. in the introduction of the disease to a
It has been shown experimentally that previously free area (Anon., 1964).
Newcastle disease virus will survive 1 1 to The difficulty of detecting a pathogenic
19 days in lake water, depending on aera Newcastle disease virus in a live wing-web
tion, pH, presence of salts and
the Newcastle disease vaccine has been em
organic matter (Boyd and Hanson, 1958) phasized (Rosenwald et al., 1959). Also,
and undetermined factors (Winterfield certain strains of mesogenic vaccine virus
and Seadale, 1956a). In contrast, it has (for example, MK107) have spread from
been possible to recover Newcastle disease inoculated birds to contact birds in other
vaccine virus placed in drinking water in cages of the same unit. There was no
a brooder house for only 24 to 36 hours clinical effect, but an active immunity re
(Dardiri et al., 1957; Luginbuhl et al., sulted (Rosenwald et al., 1959). Under
1955; Marek, 1960; Winterfield and these conditions, the route of spread was
Seadale, 1956a). considered to be aerogenic.

44
 PART Diagnosis

II:
GENERAL CHARACTERISTICS OF THE DISEASE
The causal agent of Newcastle disease the main pathological lesions were in the
was first identified by Doyle (1927) alimentary (Capobianco, 1949). In
tract
who showed that the virus was distinct The Netherlands, the initial severe form of
from the virus that caused fowl the disease has been followed by an in
plague. This finding was based on six crease in the number of mild or subclinical
main factors: period of incubation, symp outbreaks (Hoekstra, 1961). Similarly in
toms, post mortem lesions, blood infec- western Germany, the character of New
tivity, disease infectivity and results of castle disease has changed and the appear
cross-immunity tests. Later, following an ance of lentogenic and mesogenic strains
interchange of virus strains between of the virus has complicated the clinical
workers in Java, the Philippines, India, and pathological picture (Fritzsche, 1963).
Japan and England, cross-immunity and In contrast, the epizootic of Newcastle
serum-virus neutralization tests showed disease which occurred in Kenya in 1955
that six virus strains (designated Java, was equally as virulent as the epizootic
Philippine, Ranikhet, Japan, Korean and which occurred 20 years previously (Scott
Newcastle) were immunologically indis era/., 1956).
tinguishable (Doyle, 1935). The disease now considered to have
is

The marked difference between the four main forms: the velogenic (virulent),
clinical features of the classical or original the mesogenic (less virulent), the
form of Newcastle disease and the disease lentogenic (mild) and the asympto
termed "avian pneumoencephalitis" matic. This classification, though arbitrary,
Beach, 943 delayed for seven years the useful for descriptive purposes and for
is
1
(

recognition of the latter

form of New
as grouping published findings (Lancaster,
a

castle disease (Beach, 1944). The occur usually possible to distinguish

It

1963b).
is

rence of all degrees of severity, from out these forms in fowls but not in other
breaks with 100 per cent mortality to domestic poultry such as ducks, geese and
asymptomatic outbreaks (Bankowski, turkeys which are more resistant to the
1961c), or outbreaks with no characteris virus.
tic symptom or lesion (Beach, 1946), has
also had an important bearing on diag Incubation Period
nosis and diagnostic methods. After artificial infection, the incubation
Another feature that has affected diag period has varied from to 25 days
1

nosis has been the recognition of chang (Cooper, 1930; Doyle, 1927; Farinas,
a

ing clinical picture (Brandly, 1953). In 1930; Guha and Chatterjee, 1950; Jung-
Yugoslavia, for example, the incidence of herr et. al, 1946). The average period has
the nervous form of the disease has in been to days (Beach, 1943, Gomez,
4

creased (Jaksic and Stefanovic, 1957). In 1930; Picard, 1928; Reagan et al., 1954c;
Italy, initial outbreaks generally involved Sahai, 1937a, b), and occasionally day
a

the upper respiratory tract; whereas later or so more (Hudson, 1937a, Jungherr
b;

45
and Minard, 1944), or less (Thompson Sex Differences in Susceptibility
and Osteen, 1952), depending on the
Both sexes are equally susceptible (Fen-
amount of virus inoculated (Albiston and
stermacher etal, 1947; Rodier, 1928).
Gorrie, 1942; Farinas, 1930). With aerosol
infection, the incubation period has been
shorter in warm environments and longer
Age Susceptibility
in cold environments (Sinha et al., 1957).
After contact exposure, the incubation In outbreaks of the less virulent form of
period has varied from 1 to 15 days or the disease, susceptibility decreased as

longer (Beach, 1943; Brandly, 1959; Jung- birds became mature (Brandly et al.,
herr et al., 1946; Khan and Huq, 1963; 1946a, d; Brandly, 1953; Dobson, 1939).
Sahai, 1937a) with the average being 4 to Similar increasing resistance with age has

9 days (Doyle, 1927; Hudson, 1937a, b). been shown towards mesogenic vaccine
strains of virus (Haddow and Idnani, 1946;
Nilakantan et al., 1960a). However, in
Breed Susceptibility natural outbreaks of the velogenic form
There have been reports of no difference of the disease, all ages have been suscep
in susceptibility between improved breeds tible (Rao and Agarwal, 1960; Rodier,
and native breeds in West Africa (Hamil 1928; Sturgess, 1931).
ton, 1951; Hill etal., 1953); India (Cooper, Congenital immunity is an important
1931; Guha and Chatterjee, 1950); Ceylon factor in the resistance of birds to New
(Sturgess, 1931 ); Indonesia
(Doyle, 1948; castle disease and, when present, accounts
(Khan and Huq,
Ressang, 1961); Pakistan for the relative insusceptibility of young
1963); and the Sudan (Karrar and Mus chicks to virulent virus (Brandly et al.,
tafa, 1964). However, among improved 1946c; Doll et al., 1951a) or to a lento-
breeds, Leghorns have been found more genic vaccine virus (Lancaster et al., 1 960).
susceptible Island Reds and
than Rhode
other heavy breeds (Albiston and Gorrie,
1942; Kee, 1928; Piatt, 1948). Effect of Season of Year
In India, outbreaks of Newcastle disease
have been more common during the rainy
Genetic Differences in Resistance season, and the disease has generally sub
In an early report on this topic, Iyer and sided during the very hot weather (Sahai,
Dobson (1941a) stated that the offspring 1937a). Similarly, in the Sudan, the dis
of Newcastle-resistant hens survived chal ease has tended to appear at the end of the
lenge with virulent Newcastle disease virus rainy season (Karrar and Mustafa, 1964).
during the first four weeks after hatching. However, season of the year has not,
It seems likely that this apparent resistance apparently, influenced the virulence of the
was the result of passive transmitted anti disease in the Philippines (Farinas, 1930)
body (see pages 91 to 93). or in Pakistan (Khan and Huq, 1963). In
More recently, Cole and Hutt (1961) Maryland, in the United States, cases con
found that two strains (K and C) of White firmed by virus isolation have shown a
Leghorns showed significant differences in seasonal distribution, being more frequent
susceptibility to a live wingweb vaccine during the winter months (Jungherr and
(Figure 7). Similar genetic variation in Terrell, 1947).
resistance has been reported by Francis Inclement weather may modify the
and Kish (1955) and Godfrey (1952). clinical course of the disease (Brandly,
This variation may account for some of 1953). However, severe epidemics in
the differences in clinical symptoms. Greece and Indonesia during the hot

46
 MORTALITY DUE TO VACCINATION WITH A WING-WEB
VACCINE
Percent
7-

5 -

3 ~

1 -

Days after Vaccination

(Redrawn from Cole and Hott, 1961)

Figure 7.

summer have been reported (Brandly, nervous signs at warm temperatures, and
1953; Mansjoer, 1961). more respiratory signs at cold temper
atures. However, after a study of the
epidemiology of Newcastle disease in
Climatic Influences Europe, Eckert (1957) concluded that
Cold, adverse weather increases the there was no direct relationship between
severity of clinical symptoms (Brandly, the incidence of fowl pest (Newcastle dis
1959); and method of housing has also ease and fowl plague) and seasonal tem
influenced mortality (Francis and Kish, perature or precipitation. Under experi
1955). Under controlled environmental mental conditions, Baetjer et al. (1960)
conditions, Sinha et al. (1957) found more showed that in young chicks the inhalation

47
of warm dry air appeared to inhibit the lone and Squibb, 1962). No pronounced
ciliary mechanism and this allowed a change in packed cell volume, sedimenta
greater spread of virus in the respiratory tion rate, or buffy coat values has been
tract. observed in chickens artificially infected
with Newcastle disease virus (Bierer et al.,

Routes of Infection 1963).

In a severe outbreak of Newcastle dis
Chickens 3 to 6 weeks old have been ease in 6-week-old chicks there was a
most susceptible to infection with New significant reduction in the blood serum
castle disease virus by the intramuscular levels of total protein, vitamin A and total
or respiratory routes, and most resistant to carotenoids, but the ascorbic acid level was
virus administered by the alimentary route raised (Squibb et al., 1955). In artificially
(Kohn, 1955). The upper respiratory tract infected chicks, an increase in riboflavin
seems to be the site for the early multipli supplementation of the ration has reduced
cation of the virus following intranasal mortality 10 to 17 per cent (Squibb, 1963).
infection (Maestrone and Coffin, 1964). By way of contrast, an appreciable in
In reviewing the topic of portals of entry, crease over the normal requirements of a
Beaudette (1943) has emphasized that, number of vitamins has resulted in a 13
under field conditions, per os infection per cent increase in Newcastle disease
may represent entrance of the virus mortality.
through either the digestive or the res Following the administration of an in
piratory tract. Further reference to this activated Newcastle disease vaccine to
subject is made under the heading "Live pullets aged 12 weeks, total cholesterol has
Vaccines" on page 97. been determined in samples of serum and
egg yolk (Schiavo, 1963). In these experi
ments, vaccination had no effect on choles
Effect of the Virus on Avian
terol levels.
Physiology
The rise in body temperature in groups
Infection with Newcastle disease virus of vaccinated chickens, subsequently in
has increased nitrogen retention during the fected with virulent virus administered by
incubation stage and depressed nitrogen the subcutaneous and intravenous routes,
retention during the clinical stage (Sans- has been examined by Mantovani (1949).

THE DISEASE IN CHICKENS

Velogenic Form sion, increased rate of respiration, progres

sive weakness and prostration. There is
Symptoms usually an early rise in temperature (Doyle,
In the velogenic form, Newcastle disease 1935; Guha and Chatterjee, 1950; Hud

appears suddenly and spreads rapidly son, 1937b) which increases on the fifth
through a flock. Peracute cases have been or sixth day to a peak of 4° to 6°C above
reported in which birds were found dead normal. In artificially infected birds,
without having shown any symptom Squibb (1961a) found that temperatures
(Sahai, 1937a; Iyer, 1943; Khan and Huq, which remained higher than 42°C for
1963). In more typical cases, initial dull more than 48 hours, and then dropped
ness is followed rapidly by marked depres- below normal, were associated with acute

4S
Figure 8. — The velogenic form of Newcastle disease — symptoms of paralysis. (Courtesy of
Dr. G. L. Bannister, Animal Diseases Research Institute, Canada.)

neurotropic involvement or death. The Guha and Chatterjee (1950), Hilbrich

markedly subnormal temperature before (1963) and Iyer (1943).
death is a characteristic symptom (Cooper, Distension of the crop with gas or sour
1931; Hudson, 1937b; Jungherr et al, fluid is common. There may also be a dis
1946; Sahai, 1937a). Diarrhoea is com charge of mucus from the mouth or nos
mon in the early stages of the disease. trils (Johnstone, 1933; Kylasamaier, 193!;
Fae;es are usually profuse, watery, green Rodier, 1928), and sometimes from the
ish or yellowish, and occasionally blood eye (Kylasamaier, 1931). However, not
stained (Albiston and Gorrie, 1942; many of these symptoms were noticed in
Farinas, 1930; Rodier, 1928). Rate of the 1932 outbreak in Australia (Albiston
respiration is increased, and usually the and Gorrie, 1942) or in outbreaks report
bird shows a characteristic prolonged gasp ed by Gordon and Asplin (1947) and
ing inspiration with outstretched neck and Fabricant (1950).
head and partially opened beak. In arti There is usually rapid dehydration and
ficially infected fowls, gasping respirations cyanosis of the comb and wattles; and
may be absent (Acevedo, 1933), but there occasionally oedema around the head
may be a sharp (Albiston and
cough (Thompson and Osteen, 1952).
Gorrie, 1942; Hudson, 1937b). Photo Birds that survive the acute phase of
graphs of affected birds which show many the disease often show involvement of the
of the above symptoms appear in the pub central nervous system (Ressang, 196!)
of Doyle (1927), Albiston and
lications by weakness or paralysis of the legs and,
Gorrie (1942), Beach (1943), Brandly occasionally, the wings (Figure 8).
(1959), Fritzche and Gerriets (1962), Other symptoms include clonic spasms.

49
Figure 9. — The velogenic form of Newcastle disease — lesions in proventriculus. (Courtesy of
Dr. G. L. Bannister, Animal Diseases Research Institute, Canada.)

muscular tremors, torticollis, opisthotonos, perature mortality was 100 per cent;
emprosthotonos, and abnormal move whereas at a low environmental tempera
ments such as walking in circles. Opacity ture it was 55 per cent. Using different
of the cornea has also been reported management practices, Francis and Kish
(Abrams, 1961; Hill et al., 1953; Thomp (1955) obtained contrary results: they
son and Osteen, 1952). found that chickens exposed to a virulent
Egg production falls abruptly; and soft Newcastle virus in a warm battery room
or imperfectly shelled eggs may be laid. had a much lower mortality than similar
Effects on the production and quality of groups in colony houses.
eggs are more characteristic of the meso- The duration of illness has been report
genic form of the disease and are described ed as one to two days (Sturgess, 1931),
later under that heading. three to four days (Iyer, 1943; Kylasa-
maier, 1931; Rodier, 1928) and ten days
Mortality (Guha and Chatterjee, 1950). Surviving
paralytic birds seldom fully recover (Iyer,
In the velogenic form, mortality is
1943; Rodier, 1928).
usually over
90 per cent. Occasionally,
pens of older birds have survived field
Macroscopic Lesions
infection (Dobson, 1939).
The influence of environmental tempera Post mortem lesions in several hundred
ture was examined by Sinha et al. (1957) infected birds have been described by
who found that at a moderately high tem Culzoni (1949), Doyle (1927), Guha and

50
Figure 70. — The velogenic form of Newcastle disease — lesions in intestine. (Courtesy of
Dr. G. L. Bannister, Animal Diseases Research Institute, Canada.)

Chatter jee (1950), Iyer (1943), Jungherr there is no single lesion which indicates
et al. ( 1 946 ) , Kaschula ( 196 1 ) and Picard the severity of the clinical disease (Jung
(1928). The post mortem findings des herr et al., 1 946) . In some outbreaks there
cribed in 23 published reports have been has been a general absence of gross patho

reviewed by Beaudette ( 1 943 ) . logical changes (Andrews, 1948; Cooper,

Lesions are usually more definite in 1931; Hill et al, 1953; Sahai, 1937a);
acute cases than in chronic cases; but while in others the presence of a simul

51
 THE YELOGENIC FORM OF NEWCASTLE DISEASE -
DISTRIBUTION OF LESIONS

Diagram of the sites of the 12 lesions

seen regularly in the alimentary tract

(Redrawn from Kaschula 1961)

Figure 11.

taneous tapeworm infestation has appar breaks have been associated with diph
ently inhibited the formation of intestinal theritic lesions of the mouth which began
lesions (Gavey, 1958). as small, circular, yellowish elevations
Lesions are mainly haemorrhagic and (Orr and John, 1946). In other outbreaks
inflammatory. Haemorrhages are usually the cloaca has been highly congested
petechial and are commonly found in the (Kuppuswamy, 1935).
mucosa and submucosa of the proven- Bluish-red, raised and haemorrhagic
triculus (Figure 9), gizzard and intestinal necrotic or diphtheritic lesions associated
tract (Figure 10); on the trachea, epicar- with the lymphoid follicles of the intestine
dium, pleura, mesentery, air sac mem and caeca are common. These lesions may
branes; and in the general musculature. be very small haemorrhages or cover areas
Less frequently, severe haemorrhages and 2 inches long (Sturgess, 1931). The
ulceration in the pharynx may occur necrotic membrane is removed only with
(Hudson, 1937b). In Malaya, initial out- difficulty and leaves an ulcerated area. In

52
less acute cases these ulcerative lesions are Stubbs (1946). Typically, in a susceptible
more common. The development and dis flock, this form appears suddenly and
tribution of lesions in the alimentary tract spreads rapidly. Birds show respiratory
(Figure 11) have been described in detail distress, coughing and gasping (Beach,
by Culzoni (1949), de Kock (1954), 1943). There is a marked drop in appetite.
Guha and Chatterjee (1950), Hecke Egg production falls and may stop on the
(1943-44) and Kaschula ( 1961 ) . ninth day (Beach, 1943; Biswal and
In acute cases, the spleen has been Morrill, 1954). Some birds have greenish
enlarged(Kaschula, 1961; Ressang, 1961; or yellowish diarrhoea (Zuydam, 1950b).
Schoening and Osteen, 1 948 ) . Within two or three weeks, respiratory
symptoms usually subside and nervous
Microscopic Lesions symptoms may then appear (Stubbs,
Results of the examination of over 1 60 1946). The latter are of variable incidence
experimental cases have been reported by (Pomeroy and Fenstermacher, 1948).
Jungherr et al. (1946), and of a larger Involvement of the central nervous system
number of field cases by Culzoni (1949) (Figure 12) is more common in young
and Ressang (1961 ). These authors found chicks than in older flocks (Abrams, 1961 ;
that lesions in the spleen, intestine, liver, Baker and Hays, 1947; Beach, 1943). In
gall bladder and heart were essentially some flocks over 3 months of age, nervous
exudative necrotizing in character. Splenic symptoms have not been recognized
lesions have been more severe when the (Beach, 1946).
incubation period has been short (Fuku- Effect on egg production depends
shima and Shimonura, 1934). In the lungs,
largely on the severity of the disease,
central nervous system and eye, lesions individual susceptibility, the stage of the
have been mainly proliferative and hyper- laying period and whether or not moulting
aemic (de Kock, 1954; Thompson and occurs (Schoening and Osteen, 1948).
Osteen, 1952). In one series of specimens, Thus, Biswal and Morrill (1954) found

the lymphocyte was the cell type pre that in some birds the pause in production
dominantly involved in brain lesions dur was 7 days and in others 22 days. In one

ing the acute phase of infection (Karzon outbreak, egg production was adversely
affected for 12 weeks (Knox, 1950); in
and Bang, 1951a). Extensive oedema has
been seen in the liver (Hill et al., 1953) another, the birds never returned to
and cornea (Thompson and Osteen, 1952). normal production (Piatt, 1948). Accord
In artificially infected fowls, severe bron ing to De Moulin (1951 ), decrease in egg
chial haemorrhage has been seen on production results in part from a degenera
histological examination al., tion of the pituitary gland.
(Obel et

1956). In immune birds exposed to viru

A feature of the mesogenic form is its
lent virus, focal fibrosis in the air sacs effect on egg quality. During the early

and lungs has been described (Jungherr stages of the disease, the shell is often

etal, 1946). discoloured, imperfect or missing; and

many eggs are abnormally shaped. After
recovery, the appearance of the shell
Mesogenic Form usually returns to normal; but changes in
interior quality and reduction in egg
Symptoms
weight may be of a more permanent
A general account of the symptoms and nature (Knox, 1950; Quinn et al, 1956).
lesions associated with the mesogenic form There is often an accumulation of small
of Newcastle disease has been given by air bubbles instead of the true air cell. The
Beaudette (1951b), Morgan (1946) and main features include decrease in albumen

53
quality (Lorenz and Newlon, 1 944; Quinn sist for a long time and which is often
el al, 1956), deterioration in keeping qual characterized by nervous symptoms; (4)
ity, and increase in the percentage of stuck phase of humoral antibody formation and
yolks. Quality and weight of the yolk are immunity which is readily revealed by the
not altered. In some outbreaks, the weight haemagglutination-inhibition test.
and thickness of the shell of eggs laid Mortality in the mesogenic form is con
during the recovery phase have decreased; siderably lower than in typical outbreaks
however, the albumen rarely decreases in of the velogenic form. Mortality may vary
weight (Biswal and Morrill, 1954). from 5 to 50 per cent in mature flocks and
may exceed 50 per cent in young chicks
Course and Mortality (Baker and Hayes, 1947; Fenstermacher
Woernle and Siegmann (1954) and etal, 1947; Morgan, 1946; Schoening and
Woernle (1955) have described four dis Osteen, 1948). In one outbreak, losses
tinct phases in the course of Newcastle were in the ratio of five paralysed birds for
disease: (1) phase of panagglutinin forma each dead bird (Goldhaft and Wernicoff,
tion or the appearance of auto- or hetero- 1948). Under experimental conditions, an
agglutinins (Woernle and Siegmann, increased mortality rate in chicks has been
1954); (2) phase of viraemia; (3) phase of associated with depleted vitamin A re
virus and antibody balance which may per serves (Squibb, 1961b).

Figure 12. — The mesogenic form of Newcastle disease — nervous symptoms. (Crown Copy
right. Reproduced by permission of the Controller of Her Majesty's Stationery
Office, London.)

54
Macroscopic Lesions with caseous exudate result (Gross,
1961a, b).
There is considerable variation in the
Some strains of the virus have dermo-
tropisms (Sinha et al, 1952), post mortem
tropic characteristics. These are indicated
findings (Asplin et al, 1949) and patho
by the development of vesicles on the
genicity of mesogenic strains of the virus.
Also, haemorrhagic
wattles and comb (Brandly, 1959) or by a
and inflammatory
clouding of the eyes caused by inflam
lesions vary greatly between flocks and, to
matory cells in
the aqueous humour
some extent, between geographical areas
(Thompson and Osteen, 1952).
(Jungherr and Markham, 1962; Ressang,
In the mesogenic form of the disease,
1961). Thus, an Italian strain has produced
the spleen is usually small, pale, anaemic,
lesions confined mainly to the digestive
and mottled in appearance. However, in
system; whereas an American strain has
acute cases, enlargement of the spleen has
caused lesions mainly in the respiratory
been observed (Watanabe et al., 1952).
and nervous systems (Mantovani et al,
The kidneys often show inflammation and
1954).
evidence of nephritis.
The average incidence of intestinal
lesions produced by four European strains Microscopic Lesions
of the virus in over 1,000 individual
The more pneumotropic strains of virus
chickens was 63 per cent (Jungherr et al., cause inflammation and cellular infiltra
1946). The percentage distribution of tion of the serous membranes of the
lesions in different organs in natural out thoracic and abdominal cavities. Lung
breaks of Newcastle disease in France was lesions are mainly proliferativeand exuda
as follows: liver 74 per cent, proventriculus tive in character (Brandly, 1959). In some
72 per cent, cloaca 72 per cent, heart 52 specimens, atelectasis has been the only
per cent, various 40 per cent (Lucam, abnormality noted (de Moulin, 1951).
1949b). In contrast, in a study of 800 Secondary inflammatory changes in the
chickens conducted by Jungherr et al. pulmonary and abdominal air sacs are
(1946), five American strains caused gross characterized by oedema, cellular infiltra
intestinallesions in 24 per cent of cases. tion and fibroblastic proliferation in the
Kohler (1953) has reported that 56 per more chronic cases. Mild congestion and
cent of the cases he examined showed no mucous exudation is often present in the
macroscopic lesions post mortem. trachea. Catarrh of the bronchi has also

Haemorrhages are common in the

been reported (Jungherr and Minard,
mucosa of the proventriculus and less
1944; Mochizuke et al, 1952).
Inflammation of the ovary with oedema
common in the intestine. In some out
and stromal vacuolation have been des
breaks, there has been severe catarrhal
cribed by Biswal and Morrill
(1954) and
enteritis (Kawashima et al., 1953); in
by Pasley and Auer (1958). Hypertrophy
others, there has been fluid or mucus in
and hyaline necrotic foci have been present
(Watanabe et al, 1952). The
the trachea
in the spleen of many specimens (Jungherr
more pneumotropic strains of virus often
and Minard, 1944).
cause a cloudiness of the air sacs which In the central nervous system, virulent
may develop a film of yellowish
into
strains have caused extensive hyperaemia
exudate (Beach, 1943). This inflammation
and proliferation of the endothelial cells
of the air sacs shows a marked tendency
(Jungherr and Minard, 1944), and mild
to progress and, in combination with an or severe degenerative changes of the
E. coli infection, lesions of severe peri neurones and ganglia (de Moulin, 1951;
carditis, perihepatitis and airsacculitis Jungherr, 1963; Sullivan, 1958). Lesions

55
of encephalomyelitis observed in the early Two periods of leucopenia have been ob
stages following vaccination, have general served: one
immediately after infection
ly disappeared within five months (Salyi which lasts 24 to 96 hours, and the second
and Hodosy, 1 952). The brains of chickens starting on the 1 8th day which lasts seven
showing nervous symptoms have been days (Machado, 1951). On the other hand,
examined with the object of localizing the Pehl (1959) concluded that the cell picture
main histological changes (Auer, 1952; in bone marrow and circulating blood is
Karzon and Bang, 1951a). Different not sufficiently characteristic to distinguish
lesions appear to be associated with vari Newcastle disease from infections caused
ations in the symptoms observed, with by other agents. Furthermore, he thought
typical American and European strains of that the leucopenia which occurs in New
virus differing in the histological lesions castle disease is not diagnostic.
they produce (Potel, 1950). However, in a
paralytic chicken, Karzon and Bang Lentogenic Form
(1951a) found that brain lesions were not Symptoms
related to the virulence of the infecting This form of the disease is characterized
virus. Kohler (1953), who considered by mild respiratory symptoms and by a
microscopic examination of the central sudden drop in the egg production of lay
nervous system a valuable aid in diagnosis, ing flocks. There is usually depression and
found a non-purulent encephalitis in 88.4 impairment of appetite. In some out
per cent and a neuritis in 72.9 per cent of breaks, respiratory involvement is very
the cases he examined. Similarly, Mits- mild (Asplin, 1952; Asplin et al, 1949)
cherlichet al. (1953) have reported that and is only detected when the birds are
89 per cent of the cases they examined roosting. In other instances, there may be
were diagnosed histologically and 80 per no respiratory symptom (Kutlesa, 1952).
cent serologically. There are usually no nervous symptoms
In the pituitary gland, the virus has (Abrams, 1961), and egg production re
caused a degranulation of acidophile and turns to normal within a few weeks. How
basophile cells (de Moulin, 1951; Pasley ever, in some outbreaks nervous symptoms
and Auer, 1958). In the intestines, lesions have predominated (Salyi et al., 1955).
involving proliferation of the reticulo- The course of the disease varies, but
endothelium have been observed (Zan- usually there is complete within
recovery
gerle, 1952). one to eight weeks, depending on the
The haematological picture in the degree of debility. In young chicks, nasal
mesogenic form of the disease has varied. discharge and abnormal respiratory sounds
In 10 spontaneous cases, Chandrasekharan are common.
and Krishnan (1958) observed a slight rise
in the total red blood cell count and a Mortality
reduction in the white cell count. In the In adult fowls mortality is negligible,
differential count, there was a marked but it may reach 50 per cent in young
reduction in lymphocytes and a significant chicks.
increase in heterophiles. In contrast, Ishii
Macroscopic Lesions
and Kobayashi (1952) recorded a marked
Haemorrhagic and visceral lesions are
temporary leucocytosis during the febrile
absent. A mild tracheitis may be seen in
stage. Weidenmuller (1960) found eosino-
early cases and in young birds.
philia, monocytosis and lymphopenia and
suggested that lymphopenia could be used Microscopic Lesions
as a diagnostic criterion in the early stages A disseminated encephalomyelitis has
when serological tests were still negative. been found in a field outbreak (Kutlesa,

56
1952). In a study of vaccinated chicks, Newcastle disease and some of which did
Auer (1953) found no histological evi not, have already been mentioned on page
dence of degeneration in nerve cells. How 33. Thus, Newcastle disease has been
ever, there was an inflammatory reaction diagnosed serologically in the absence of
in the central nervous system. Vascular clinical signs (Reid, 1961). Also, Minard
engorgement and infiltration with lym and Jungherr (1944) have reported high
phocytes have lasted 14 days after vaccin levels of neutralizing antibody in a flock
ation with no clinical symptom being seen in which there was no evidence of the
during this period (Auer, 1954). disease. This was understood to indicate
Gross (1963) examined microscopic a latent infection. In some cases, spread
lesions in air sac membranes, lungs and of an inapparent form of the disease (Jung
trachea of 3-week-old susceptible chickens herr and Terrell, 1946) has been thought
exposed to an aerosol of the Bl strain of to result from carriers of the virus. In one
virus. In the air sac membranes there was instance, inapparent infection had existed
a progressive increase in connective tissue on a large breeding establishment for two
for about 16 days post vaccination; there years before clinical evidence appeared
after, the walls of the air sacs became (Beach, 1951).
thinner. Lymphoid infiltration was marked Bankowski (1961c) has emphasized that
in the air sac membranes and in the lungs asymptomatic Newcastle disease may
around the parabronchi, but became more often be diagnosed only by chance. He has
localized in the tracheal epithelium. It was pointed out that the haemagglutination-
concluded that the Bl strain had resulted inhibition titre fluctuates considerably and
in lesions similar to those associated with that inapparent outbreaks may be missed
infectious bronchitis virus or Mycoplasma unless sufficient of sera are
numbers
gallisepticum infection. examined. However, such variations in
titre are not confined to asymptomatic
Asymptomatic Form infections: marked variation in HI titre
Reports of asymptomatic infections, has also been observed in the mesogenic
some of which resulted in the spread of form of the disease (Fabricant, 1950).

THE DISEASE IN TURKEYS

Symptoms reported an outbreak in turkeys which
spread slowly, and largely in a subclinical
In the mature turkey, there may be mild
respiratory symptoms. Nervous symptoms form. Respiratory symptoms were absent,
are seldom but partial or complete paralysis of one or
seen
(Fenstermacher et al.,
both legs was common.
1946) but have been reported occasionally
(Walker, 1948). In laying flocks, egg pro
duction has dropped almost to zero. Eggs Course and Mortality
have been soft shelled and deformed and After outbreaks in some breeder flocks,
have had watery albumen (Gale et al., fertility and hatchability have returned to
1961 ) . Fertility and hatchability have been normal (Gale et al, 1961). Death may
reduced. occur in mature birds (Fenstermacher et
In flocks of young poults, respiratory al, 1946); and in some outbreaks the
symptoms, depression and inappetence disease has been peracute with high mor
appear early. Gray et al. (1954) have tality (Gordon et al., 1948; Walker,

57
1948). In young poults, mortality of 60 reported by Gale et al. (1961) who noted
per cent or higher has been observed that in some outbreaks there were lesions
( Fenstermacher et al,, 1946). in the respiratory and central nervous
systems: extensive haemorrhage and capil
Macroscopic Lesions in the respiratory
lary congestion system,
Gross lesions are rare, but some pete- in the
and extensive neuronal degeneration
chiae on the heart and a cloudy appear
cerebrum, brain stem and cerebellum. In
ance of the air sacs have been reported.
other outbreaks, focal gliosis with an

Microscopic Lesions occasional perivascular cuffing has been

The histopathology of experimentally observed in the brain and spinal cord
and naturally infected turkeys has been (Gray et al, 1954).

THE DISEASE IN DUCKS AND GEESE

Although these species are more resist lings have succumbed to the disease, and
ant than turkeys, natural outbreaks among mortality in one flock of ducklings was
ducks and geese have been reported (see 100 per cent (Bush, 1954). In contrast, the
page 29). In Australia, locomotory dis experimental inoculation of mature ducks
turbances, abnormal gait and general with virulent Newcastle disease virus has
depression have been observed (Albiston resulted in no clinical symptom and the
and Gorrie, 1942). Elsewhere, ducks and serum has shown no virus neutralizing
geese have undergone symptomless infec antibody (Iyer, 1945). In a few outbreaks,
tion (Khan and Huq, 1963). In England, some ducks have died (Andrews, 1948).
ducks exposed to infected fowls have Post mortem findings have been generally
usually remained apparently healthy negative (Albiston and Gorrie, 1942;
(Asplin, 1947). In Haiti, ducks and gos Asplin, 1947).

THE DISEASE IN GAME BIRDS

Natural outbreaks with varying mor Lucas and Laroche (1958) studied
tality have been reported in pheasants, Newcastle disease in the partridge and
guinea-fowl and partridges. The effect of have reported that, in the acute form,
the disease on these birds has been re death occurred five days after the appear
viewed by Beaudette ( 1943 ) and Brandly ance of symptoms. Post mortem findings
(1959) (see page 28). In one outbreak in were those of enteritis. In the chronic form
partridges (Perdix perdix), a significant of the disease, most partridges made a slow
lesion consisted of numerous microscopic recovery. A strain of virus isolated from a
ulcers in the intestine which appeared to natural infection in a partridge has been
the unaided eye as very small white foci identical to strains isolated from chickens
(Thompson, 1955). In other outbreaks, (Placidi and Santucci, 1953c).
prominent haemorrhages on the serous A natural infection in guinea-fowl has
coat of the gizzard have been reported resulted in symptoms of encephalitis
(Parnaik and Dixit, 1953). (Placidi and Santucci, 1953c).

58
 DIAGNOSIS BY SEROLOGICAL METHODS

Brandly et al., 1946d). Determination of

Haemagglutination (HA Test)
the end titre in the HA titration of virus,
The haemagglutinating property of using bovine and equine erythrocytes, has
Newcastle disease virus has been described been both time-consuming and difficult
by Burnet (1942) who found it capable (Ozawa and Chow, 1958).
of agglutinating the red blood cells of man, All strains of virus are not equally active
guinea-pig, mouse, fowl, sparrow and frog against mammalian erythrocytes (Mac-
(Hyla). The agglutination of human, pherson and Swain, 1956). Winslow et al.
bovine and avian spermatozoa by New (1950) concluded that the ability to
castle disease virus has been described by agglutinate erythrocytes of certain mam
Chu (1953) as similar to the
being malian species differed between strains of
agglutination of red blood cells. Similarly, the virus. It was thought that this property
the agglutination of leucocytes and blood could be used to identify different strains.
platelets has taken place (Jerushalmy Girotto (1954) found that two Italian
et al, 1963). The agglutination of strains did not agglutinate horse red blood
avian red blood cells by Newcastle cells whereas other strains did.
disease virus can be inhibited by a sub Red cells of different chickens have
stance extracted from human and porcine shown variation in their susceptibility to
lungs (Rice and Stevens, 1957). Similar agglutination by Newcastle disease virus
inhibiting substances have been found in (Bang and Foard, 1952; Brandly et al,
sera from humans and Rhesus monkeys 1947; Cochrane and Gray, 1962). Con
(Wenner et al, 1952b), in the allantoic versely, different strains of virus have
fluid of normal 13-day chicken embryos shown variations in agglutination (Beach,
(Williamson et al, 1955) and in normal 1948), and some strains have been non-
human urine (Tamm and Horsfall, 1950). agglutinators of red blood cells (William
In fowl serum, the removal of a non- son et al,1955). With highly pathogenic
specific inhibitor has resulted in an in strains, a minimum concentration of
crease in the neutralization titre (Traub, approximately 10-6 embryo LD50 of virus
1956). Similarly, the removal of an ether- has been necessary to produce haemag
soluble antigen covering the virus has glutination (Hanson et al, 1947). With a
resulted in an increased haemagglutination lentogenic strain, the haemagglutination
sensitivity and a higher titre in the HI test activity has been destroyed after several
(Larski, 1962). Ether disrupts the virus minutes 56°C (Torlone, 1956). For
at
particles, but the ease of disruption varies many strains of Newcastle disease virus,
from strain to strain (Waterson and the resistance of the haemagglutinin at
Cruickshank, 1963). 56°C has been a relatively constant and
The virus has failed to agglutinate the characteristic feature and it might serve as
cells of monkey (Af. irus), ferret, sheep, a means of distinguishing between strains
horse, rabbit, marsupial mouse (Smin- (Nitzschke and 1963) or
Schmittdiel,
thopsis) and tortoise. However, the virus mutations of strains (Goldman and Han
can agglutinate the erythrocytes of camels son, 1955; Hofstad et al, 1963). Further
(Hashmi and Hasnain, 1954b; Placidi and more, heat has had less effect on the ability
Santucci, 1956) buffalos
(Hashmi and of some strains to agglutinate chicken
Hasnain, 1954a), cattle (Cordier et al, cells than it has on theirinfectivity for
1951; Ozawa and Chow, 1958) and other chick embryos (Liao et al, 1953). How
species of animals (Abdel et al, 1960; ever, no conclusive relationship has been

59
demonstrated between the immunogenicity disease virus, a small number of virus
and the heat sensitivity of the haemag- particles permanently remain on the red
glutinating property of substrains of New blood cells. Such sensitized red blood cells
castle disease virus (Hofstad et al, 1963b). can be lysed and stored indefinitely with
Newcastle disease virus first agglutinates out loss of antigenicity (Geurden and
chicken red blood cells by causing the Devos, 1955).
formation of a lattice-like matrix and then Specific antisera against Newcastle dis
elutes from the cells (McCollum et al, ease virus-treated chicken red blood cells

1957). Other characteristics of haemag- has been prepared (Gardner et al, 1954).
glutination have been reported by Hirst However, Swain (1959) found a serum
(1950). factor which reacted with Newcastle dis
The virus appears to be adsorbed more ease virus-treated red blood cells in the
completely and to elute less rapidly from serum of normal animals of many species.
chicken red blood cells at 4°C than at This serum factor appeared to be distinct
room temperature (Florman, 1947; Sagik from specific antibodies to Newcastle dis
and Levine, 1957), giving a clear HA ease virus. Previously, Burnet and Ander
reaction with certain strains (Andrewes et son (1946) had shown that sera from
al, 1955; Berke and Golem, 1950) but human cases of infectious mononucleosis
not with a vaccine strain (Bang and Foard, would agglutinate human red blood cells

1952). Nevertheless, Bohm and Espig sensitized by the action of Newcastle dis
(1961), who made comparisons at four ease virus.
different temperatures, found that higher Modification of the virus antigen by
end-points were obtained at higher tem potassium periodate (KI04) destroys the
peratures, and they concluded that the elution property of the virus (Bang and
HA test should be carried out at room tem Libert, 1952). As a result, the HA test
perature. However, the haemagglutination can be read at any time up to 12 hours;
inhibitor present in normal allantoic fluids whereas, with unmodified antigens, com
is less active atroom temperature than at plete elution of the virus from the red
4°C (Williamson et al, 1955). blood cells occurs within two hours
It has been shown that Newcastle dis (McCollum et al, 1957). When inactivated
ease virus is adsorbed to red blood cells by heat or formalin, the virus retains its
and to platelets in a similar manner ability to agglutinate red blood cells (Berke
(Jerushalmy et al, 1961). However, elu- and Golem, 1950; Bodon, 1953; Brandly
tion of the virus from platelets is much et al, 1946d; Burnet et al, 1945; Walker,
slower than from red blood cells. 1952). The addition of 50 per cent gly
Bang and Libert (1949) have reported cerine to virus suspended in 0.1 per cent
that chicken red blood cells sensitized by formol saline markedly stabilizes the
Newcastle disease virus were agglutinated haemagglutinating antigen (Cabasso et al,
by sera which did not agglutinate normal 1951).
red cells. Thus, sera taken from chickens A non-infectious haemagglutinin whose
five to seven days after inoculation with particles appear to correspond to the in
virulent Newcastle disease virus aggluti complete forms of the virus has been des
nated sensitized cells but not normal red cribed (Rott et al, 1962). Treatment with
blood cells. This haemagglutination of ether has resulted in separation of the
sensitized red blood cells facilitates rapid haemagglutinating components from an
diagnosis (Geurden and Devos, 1955). In American strain of virus but not from an
studying this type of reaction, Anderson Italian strain (Rott et al, 1961). Dialysis
(1947) concluded that, following treat of allantoic fluid or repeated freezing and
ment of red blood cells with Newcastle thawing has reduced the haemagglutina

60
tion titre (Vrtiak et al, 1959). The liber been utilized in the identification of virus
ation of haemagglutinating units by ether in the aqueous humour of naturally or
treatment has produced a suspension of artificially infected chickens (Clark et al,
spherical particles of about 30 micron 1955, 1957; Topolnik, 1957) but, in
diameter with long filaments (Sokol et al, general, results have not been encouraging
1961 ) . Ultrasonic irradiation of Newcastle for diagnosis (Baldwin, 1962; Scott et al,
disease virus has resulted in loss of virul 1956).
ence but an increase in its haemaggluti Rapid diagnosis using tissue extract in
nating property (Garay and Syent-Ivanyi, an HA test has been studied. Tissue ex
1955). tracts for this purpose have been prepared
Using high-speed centrifugation of sus from lungs (McClurkin et al, 1954), liver
pensions of Newcastle disease virus, and spleen (Monti, 1952). The method
Granoff al (1950) demonstrated the
et may be of value in the early stages of
presence of two haemagglutinating par infection before serum antibodies have
ticles of different sizes. The larger appeared appeared. However, Belic (1962) has re
to be the infectious principle; the smaller ported a high percentage of negative HA
particle appeared to be non-infectious. In reactions and has concluded that such
subsequent studies, Granoff and Henle rapid test procedures are of limited value.
(1954) showed that the larger component Peptic or tryptic digestion of the extract
was readily adsorbed on to red blood cells have been found to increase the specificity
in the cold, whereas only small amounts of of the test (Berke and Golem, 1950). On
the smaller particle combined with the the other hand, the use of fowl red blood
cells. cells sensitized by suspensions of infected
Haemagglutination test techniques have organ material has been a reliable pro
been described (Anon., 1946b;
in detail cedure (Grausgruber, 1958).
Berke and Golem, 1950; Chu, 1960; Cun
ningham, 1960; Fabricant, 1949a; Kaplan,
Haemagglutination-lnhibition
1949). A micro-method of conducting the (HI Test)
test, described by Takatsy (1956), was Burnet (1942) and Lush (1943) showed
based on the use of a 0.025 ml. loop that serum from fowls that had recovered
instead of a pipette, together with a metal from the disease contained antibody cap
dropper also calibrated to deliver 0.025 ml. able of inhibiting the haemagglutination
The incubation time for the test was of red blood cells by the virus. The
reduced if the plastic plates containing the haemagglutination-inhibition (HI) test has
reagents were centrifuged for 10 seconds. provided useful quantitative as well as
Possible sources of error and factors qualitative data.
influencing HA test results have been dis Using antigens and sera produced from
cussed by several authors (von Sprock- nine different strains of Newcastle disease
hoff, 1961; Anon., 1959; Bang and Foard, virus, Siegmann and Woernle (1955)
1952; Burnet et al, 1945; Brandly et al, showed that the homologous serum in
1947; Cunha et al, 1947). hibited haemagglutination in far greater
The standard HA test involves the pre dilutions than heterologous sera. Nilakan-
paration of dilutions of the virus antigen tan et al(1962) compared various pro
in constant volumes of a red blood cell tein fractions of plasma from Newcastle
suspension. To facilitate diagnosis, rapid disease immune chickens and showed that
slide or plate tests have been made in the euglobulin contains the maximum
which only one dilution of the reagents is amount of HI and virus-neutralizing anti
used (Walker, 1952; Zargar and Pomeroy, body — no antibodies were demonstrated
1949). The haemagglutinating property has in the albumen fraction of the plasma.

61
 Two procedures for the HI test have glutination of chicken red blood cells by
been used. In the alpha procedure (Anon., Newcastle disease virus (Ginsberg and
1946b; Fabricant, 1949a) the virus sus Horsfall, 1949). Thus, fresh normal
pension is diluted serially and mixed with human sera have caused the virus to lose
equal volumes of the serum under test. In its capacity to agglutinate red cells (Bang
the beta procedure (Anon., 1956; Chu, et al, 1951 ) ; though this phenomenon has
1960; Gentry, 1957) the serum is diluted been influenced by the unsuitability of the
serially and mixed with a constant amount red blood cells of certain chickens (Ander
of virus dilution containing a known son, 1948).
number of HA units. A comparison of the The test has been found to be of little
alpha and beta procedures has been made value in the acute form of the disease
by Brandly et al. (1947). (Scott et al, 1956; Valadao, 1955) ; and in
Electron microscope studies of the HI early infections it should be supplemented
test were conducted by Reagan and by tests for detecting circulating virus
Brueckner (1953) who have reported (Topolnik and Hallauer, 1950). It has
seeing virus-like particles adhering to the been suggested that the strain of invading
surface of agglutinated red blood cells. virus (Doll et al, 1950d) and the genetic
No virus-like particles were evident in the background of the chicken affect HI titre.
presence of immune serum. Some chickens can apparently overcome
Details of HI test techniques have been the virulent virus without developing high
given by several authors (Anon., 1946b, HI titres (Millen, 1960). Among recovered
1956, 1959; Chu, 1960; Crawley, 1954b; birds, there is considerable variation be
Cunningham, 1960; Doll et al, 1950d, tween the serum titres of individuals
1951d; Fabricant, 1949a; Kaplan, 1949). (Bonaduce, 1950b). In mild forms of the
Takatsy (1956) has described and illus disease, some chickens exposed to infec
trated a micro-method. tion have remained negative to the HI test
Discs of absorbent paper
(serodiscs) (Asplin etal, 1949).
have been used to collect samples of whole Several authors have reported that the
blood for subsequent elution and testing by HI test using serum is more satisfactory
the beta procedure (Andrews, 1963). This than post mortem examination for routine
technique resulted in the identification of diagnosis (Nitzschke and Venske, 1956;
85 per cent of pheasants with a serum titre Puteanus, 1953; Schlegel-Oprecht and
of 200 or higher. Fey, 1953). Osteen and Anderson (1948)
The desirability of using a standardized considered the HI test as accurate as the
HI procedure based on a freeze-dried SN test.
Newcastle disease serum has been pro For diagnostic purposes, the HI test
posed by (1963). In addition,
Lessing has been found satisfactory when applied
recommendations have been made for to organ or tissue extracts, blood clots
standardizing reagents and the method of from dead birds (Mitscherlich and Gur-
reporting results (von Sprockhoff, 1961). turk, 1952; Nitzschke and Venske, 1956;
The HI test depends on the presence of Weidenmuller, 1955; Woernle and Sieg-
circulating antibody at a significant level. mann, 1954) or egg yolk (Bornstein et al,
In Newcastle disease, this level is usually 1952; Schmittle and Millen, 1948; Weiden
reached five to ten days after infection muller, 1955). Extracts of liver, spleen
(Hofstad, 1951; Valadao, or two
1955), and proventriculus have been found suit
days after the first respiratory symptoms able for these purposes (Leitner, 1954).
appear (Fabricant, 1950). However, cer However, Schlegel-Oprecht and Fey
tain mammalian sera contain a labile (1953) found there was no advantage to
component which also inhibits haemag- using organ extracts. In contrast, the HI

62
 titres of egg yolks have been considered a tions in poultry. These agents include
reliable quantitative index of the serum a virus, designated myxovirus strain
titres of
the hen (Bornstein et al, 1952). Yucaipa, which immunologically and
is
Zargar and Pomeroy (1949) have des serologically distinct from Newcastle dis
cribed a rapid HI plate test in which one ease virus (Bankowski and Corstvet, 1961).
loopful of whole blood from the wing vein Another agent capable of agglutinating
was mixed with Newcastle disease antigen chicken red blood cells is Mycoplasma
on a glass plate. Topolnik and Hallauer gallisepticum, the causal agent of chronic
(1950) found this test gave positive results respiratory disease (Fahey and Crawley,
in fowls which had been infected five or 1954; Markham and Wong, 1952).
more days previously. A micro-method
using whole blood has been useful for
Haemolysis
immunological and diagnostic investi
gations (Bamberger and Elek, 1955). The ability of the virus to haemolyse
Arapid plate test using serum instead red blood cells in vitro (Burnet, 1949,
of whole blood has been reported by 1950; Kilham, 1949; McCollum and
Dinter et al. (1948) and Luginbuhl and Brandly, 1955a) has been used by Kahnke
Jungherr ( 1949). The latter authors found (1951) and Nilakantan et al. (1963) in a
that in 93 per cent of cases, this test gave haemolysis inhibition test for the detection
comparable results to the tube dilution of antibodies to Newcastle disease virus.
technique. Walker (1952) compared the These authors showed that a correlation
rapid plate tests, using either whole blood existed between the haemolysis inhibition
or serum, with the HI tube test: in 21 test and the HI test.
birds there was disagreement in one in The serum titre which inhibits haemoly
stance only.A modified agglutination test sis corresponds closely with the HI titre
has been described by Raggi ( 1960) in of the serum (Burnet and Lind, 1950);
which each serum sample was centrifuged whereas haemolysis and elution appear to
at 0°C for 20 minutes before mixing with involve separate reactions
(Sagik and
a formolized antigen on a glass plate. In a Levine, 1957). Under suitable conditions,
recently vaccinated flock, this test showed a single virus particle per red blood cell
agglutinins before the HI titres became can cause haemolysis (Sagik and Levine,
significant. 1957). Furthermore, the ability to lyse
A comparative study of four diagnostic chicken red blood cells depends on the
methods was made by Schoenaers and presence of only one intact haemolysin
Cotteleer (1956). The techniques examined unit per virus particle (Wilson, 1958).
were those described by Monti (1952), Radiation studies on the haemolysin in
Mitscherlich et al. (1954), Geurden and dicate that the haemolysin molecules are
Devos (1955) and Woernel and Sieg- flat plates and that approximately 15 such
mann (1952). It was concluded that the molecules are arranged on the virus sur
haemagglutination test using Newcastle face. Although a virus particle with only
disease-sensitized chicken red blood cells one intact haemolysin unit can lyse
(Bang and Libert, 1949; Geurden and chicken red blood cells, it does so at a
Devos, 1955) was the most reliable. much slower rate than a virus with 15
intact haemolysin units (Wilson, 1958).
Full haemolytic activity in Newcastle
Other Haemagglutinating Agents
disease virus suspensions has not been
In addition to Newcastle disease virus, obtained without pretreatment of the virus
other haemagglutinating agents have been by techniques such as freezing and
recovered from respiratory disease condi thawing, precipitation, extensive dialysis

63
(Burnet and Lind, 1950; Granoff and serum of the fluorescent antibody
and
Henle, 1954; Vrtiak et al, 1959; Wilson, technique for bone marrow cultures have
1962b) or drying (Wilson, 1958). Such been given by Jerushalmy et al. (1963).
treatment of the Bl strain of the virus has Other studies with Newcastle disease
resulted in the ability to agglutinate and fluorescent antibodies have been reported
haemolyse human red blood cells (Liu, by Reda et al. (1964).
1952). It has also been shown that fluoro-
carbon treatment may be used to unmask
Serum Electrophoresis
the Newcastle disease virus haemolysin
(Wilson, 1 962a) . The application of high Using the serum electrophoresis tech
pressure to Newcastle disease virus has nique, Lukacevic et al. (1958) found that
been suggested as a means of releasing the serum samples from vaccinated fowls
haemolysin (Atanasiu et al, 1955). In showed a significant increase in gamma
an examination of 15 strains of New globulins.
castle disease virus, all caused haemolysis
of fowl erythrocytes (Nilakantan et al.,
Serum or Virus Neutralization
1963). The heating of the virus in amnio-
allantoic fluid at 50°C for 30 minutes (SN Test)
abolished its haemolytic activity. The antibodies associated with the HI
and the SN tests do not appear to be a
Intradermal Inoculation single entity; neither is their mechanism
of reaction the same (Beach, 1948;
The intradermal inoculation of a skin-
Brandly, et al, 1947; Hanson et al, 1950;
adapted Newcastle disease virus has pro
Schmittle, 1953). However, the HI and
duced a skin reaction only in susceptible
SN titres of experimentally or naturally
chickens (Yates et al, 1953). Chickens infected fowls have shown a close correla
which had shown respiratory symptoms
tion (Beach, 1948; Brandly et al, 1947),
for at least one day had sufficient immunity
especially during the ascending phase of
to prevent the development of this skin
immunity. Generally, HI titres have ap
reaction.
peared at a diagnostic level earlier than
SN titres (Fabricant, 1949a; Osteen and
Fluorescent Antibody
Anderson, 1948) as shown in Figure 13.
Fluorescent antibody has been prepared However, in a number of vaccinated
from the serum globulin of immune chicks chicks, serum-neutralization titres have
and this antibody detected virus antigen been positive at a time when haemag-
in chicks three hours after infection (Mae- glutination-inhibition was not demon
strone and Coffin, 1961). The antigen was strated (Schmidt, 1959). SN titres have
identified in the larynx and vascular walls generally persisted much longer than HI
of internal organs and it was suggested titres (Hanson et al, 1950), but not always
that, for diagnostic purposes, it would be (Fabricant, 1949a).
adequate if tracheal scrapings and impres Only one antibody molecule is required
sion smears from the brain, spleen and for the inactivation of an infectious part
lungs were submitted. Maestrone and icle (Rubin and Franklin, 1957). The
Coffin (1961) showed also that, at room neutralization indices, as determined in
temperature, the virus survived 10 days; embryonating eggs, have shown that nor
whereas after fixation of the smears in mal chicken serum does not contain more
acetone for 1 0 minutes, the survival period than 11 neutralizing doses per ml. (Cun
was one month. Details of the preparation ningham, 1951; Doll et al, 1950c, d). In
of fluorescent antibody from guinea-pig titration of highly virulent strains, grow-

64
 A COMPARISON OF THE RESULTS OF THE HI AND SN TESTS

S N HI
Titre Titre

sss
I I I I I I I I I I I I I I I I I I I I I I I I
0 7 14 21 28 35

Days

(Redrawn from Fabricant, 1949)

Figure 13.

ing or adult chickens have been used in- and embryo titrations, neutralization of
stead of chick embryos. In both chicken 1 ,000 infective doses of virus has been ob-

65
tained and this has been used as a means found very sensitive and specific for dem
of identifying Newcastle disease (Beach, onstrating Newcastle disease virus anti
1944; Doyle, 1935; Osteen and Anderson, body in chicken serum (Wolfe et al.,
1948). 1949; Boulanger and Rice, 1954; Nitz-
In hyperimmunized chickens, there ap schke, 1956; Wenner et al, 1950).
pears to be a direct relationship between There has been good correlation
a high HI titre and the protective value between the serum titres obtained in the
of the serum for day-old chicks (Bodon complement-fixation test and in the HI
et al., 1952). However, there has been no test (Boulanger and Rice, 1954; Russeff,
correlation between the level of specific 1956a). In addition, the complement-fixa
neutralizing antibody and the known tion test has been used to distinguish

a
disease history (Bankowski, 1961c; Min- velogenic from lentogenic strain of

a
ard and Jungherr, 1944). virus (Galassi and Gramezi, 1959).
Techniques used for neutralization tests
have been described in detail in Anon.
(1946a) ; and by Brandly et al. (1946d)
and Minard and Jungherr (1944). They Precipitation Test (Ouchterlony
have also been discussed by Bang and Double Diffusion Plate
Foard (1956a, b) and by Tyrrell and Technique)
Horsfall (1953).
The application of this technique
De-embryonated eggs were found
(Ouchterlony, 1948) to Newcastle disease
particularly suitable by Greuel (1963b).
has been described by Cunningham
Results have shown that the demonstrable
(1960) and by Woernle and
Brunner
antiviral activity or titre of yolks from
eggs laid by immune hens is generally 10- (1961) . Although the reaction has been
considered to be specific (Woernle and
to 100-fold lower than the serum titres
Brunner, 1961), no precipitin has been
of these hens (Brandly et al, 1946c). On
found in sera from high percentage of
a

this basis, it has been thought that eggs

may be just as satisfactory as blood serum
naturally infected or hyperimmunized
for assessing the Newcastle disease status fowls (Schoop and Wachendorfer, 1960).
Furthermore, the degree of precipitation
of a flock (Brandly et al, 1946c).
has not corresponded to the haemagglu-
tinationtitre, and organ extracts from in
Complement- Fixation Tests
fected fowls have been negative. More
The direct complement-fixation test has satisfactory results have been reported by
been of little value in the diagnosis of Wachendorfer (1961), who demonstrated
Newcastle disease in chickens (Rice, 1961) precipitins by gel diffusion in blood of
as a result of incompatibility between cer naturally infected or hyperimmunized
tain avian antibody-antigen complexes fowls. He also reported that chicks vac
and guinea-pig complement (Brumfield cinated with either adsorbed or "drinking-
et al, 1961). Recently, Rott and Reda water" vaccine showed no precipitin reac
(1963) have reported the identification of tion. Similar results in vaccinated chickens
a soluble antigen, termed "nucleoprotein have been reported by Guillon et al.
antigen," which has inhibited complement. (1963) who found that the re-infection
However, the indirect complement-fixa of vaccinated chicks with field virus
tion test (i.e., inhibition of complement resulted in hyperimmunization which
fixation), or modification of
it,

a has been yielded positive gel diffusion results.

66
 DIAGNOSIS BY VIRUS ISOLATION

Serological methods may be sufficiently brain (Mitscherlich et al, 1954). Brain

accurate to confirm field information, but tissue probably has the greatest concentra
new foci or extensions of the disease tion of virus (Mantovani, 1948). How
should be confirmed by recovery and ever, Kohler (1960) observed consider
identification of the virus (Beach, 1948; able amounts of Newcastle disease virus
Guillon et al, 1963; Osteen and Ander in the leucocytes for three days after arti
son, 1948; Puteanus, 1953; Walker, ficial infection. The Hertfordshire vaccine
1948). After making a comparative study virus has been recovered from the central
of four serological methods, Schoenaers nervous system 6 to 14 days after vac
and Cotteleer (1956) concluded that the cination (Salyi and Hodosy, 1952). The
inoculation of the developing chick em virus is not as consistently present in
bryo was the best method. For example, bone marrow as in respiratory and spleen
an Indonesian strain of the virus has been tissue (Baskaya et al, 1952).
propagated in the allantoic sac of develop The rate of multiplication of an aviru-
ing chicken embryos (Martini and Koer- lent Newcastle disease virus in different
jana, 1949). Specimens for virus isolation tissues following intramuscular inoculation
should be from cases in the early or even of susceptible 10-week-old chickens was
prodromal stage of the disease, and pre studied by Karzon and Bang (1951a)
ferably from the younger age groups who found the decline in virus titre was
(Anon., 1946a; Beaudette et al, 1948a; slowest in brain tissue (Figure 14). It
Brandly et al, 1946d). follows, therefore, that samples from
several pooled tissues give a higher re
covery rate than samples from any one
Distribution of Virus in the
tissue (Beaudette et al, 1948a; Pannu and
Body Bankowski, 1962).
Asdell and Hanson ( 1 960) and Hofstad
(1951) showed that the virus, when in Embryonating Eggs
jected into a susceptible fowl, spread
rapidly from the site of inoculation and Although embryonating hens' eggs are
generally used, successful results have
could be detected in almost all tissues
been obtained with embryonating duck
within 48 hours, and in all tissues within
72 hours. Electron micrograph studies of eggs (Collier and Dinger, 1950) or young
red blood cells have shown the maximum ducklings (Komaroff and Goldschmitt,
number of virus-like particles to be pre 1946a).
sent 96 hours after inoculation, at which
time the infected fowls have shown Preparation of Inoculum
symptoms of Newcastle disease (Reagan In some early work, unfiltered tissue
et al, 1954c). Nevertheless, laboratory suspensions were not centrifuged and no
results indicate that virus may be more antibiotic was added (Beach, 1943; Beau
readily from some tissues than
recovered dette and Black, 1946; Walker, 1948).
from others. The following tissues have Suspensions were filtered through Berke-
been found particularly suitable: lung and feld,Mandler and Seitz filters (Brandly
trachea (Hofstad, 1951; Miller and et al,1946d; Iyer and Dobson, 1940;
Miller, 1950), respiratory and spleen Rodier, 1928). Among the antibiotics
(Baskaya et al, 1952; Beaudette et al, used to inhibit bacterial contamination,
1948a; Beaudette and Black, 1946) and the relative values of penicillin alone

67
 TISSUE DISTRIBUTION OF NEWCASTLE DISEASE VIRUS
AFTER INTRAMUSCULAR VACCINATION OF 10-WEEK-OLD
CHICKENS

Log
LDM
of Virus

\
5- Spleen
— ■ Lun9

0 1 3 5 7 9

Days after Inoculation

(Redrawn from Karzon and Bang, 1951)

Figure 14.

(Brandly et al, 1946d; Cordier et al, the filtrate was negative for virus on egg
1950), streptomycin alone (Delaplane, inoculation, whereas the antibiotic-
'1947; Thompson and Osteen, 1948), or a treated material was positive (Beaudette
mixture of penicillin and streptomycin etal, 1949).
(Beaudette et al, 1948b) were reported.
Later studies showed that penicillin
Route of Inoculation
was capable of inactivating the virus, to
an extent that depended on the concentra Inoculation into the allantoic sac has
tion of penicillin and the duration of been commonly used (Beaudette et al,
incubation of the mixture before inocula 1948b; Thompson and Osteen, 1948), and
tion of the embryonating eggs (Kohn, various techniques have been described
1953). In a comparison of filtration and (Anon; 1946a; Beaudette et al, 1952; Mc
antibiotic treatment (penicillin and strep Carthy and Dumbell, 1961). A combina
tomycin), it was found that in 22 samples tion of inoculation of the chorio-allantoic

68
membrane and of the allantoic sac, virus isolated from chickens, ducks and a
through the same hole, has been found pheasant. Generally, all strains killed the
valuable in the primary isolation of embryos within 24 to 48 hours; occasion
viruses (Fabricant, 1957; Gorham, 1957). ally an embryo survived 72 hours. How
This technique has also reduced embryo ever, several factors influence embryonic
mortality due to trauma (Bueno et al., mortality; these include or
the presence
1961). absence of parental antibody (Brandly
et al., 1946c), the virulence of the virus

Temperature of Egg Incubation (Anon., 1959; Beach, 1943; Chu, 1958;

Hitchner et al., 1951a; Minard and Jung-
Sinha (1958) has reported that inocul herr, 1944), the temperature of incuba
ated embryos have died 9 to 15 hours
tion, and the route of inoculation. Death
earlier when incubated at 37°C instead of
of the embryo has occurred somewhat
35°C. Bang (1948) found that good virus
earlier after intravenous than after allan
growth occurred at temperatures from toic inoculation (Hanson et al., 1947).
35° to 41 °C, and von Sprockhoff (1960)
However, Bang (1948) found that neither
recorded no difference in gross lesions and
the temperature of incubation nor the
HA titres between embryos incubated at
route of inoculation of embryos had a
35.5° and 38°C. In contrast, Zuschek et al.
consistent effect on the measurement of
(1959) have reported that in embryonat- virus activity. For the latter purpose, the
ing eggs haemagglutinin activity followed 50 per cent embryo was used.
mortality
a log linear response to temperature.
Either a single particle of virulent New
castle disease virus in the inoculum or suc
Blind Passages cessful attack of a single locus has been
considered sufficient to kill the 10-day-old
Beaudette and Black (1946) found
embryo (Nadel et el., 1957).
that approximately two-thirds of a series
of 239 negative samples were determined
as negative on the first inoculation of em-
Lesions in Embryos
bryonating eggs. Of 26 positive samples,
19 were determined as such on the first
In embryos, the virus has been found
specifically destructive to the cells of
egg passage; the remaining seven samples
rapidly differentiating and proliferating
killed every embryo in
the second
tissue (Williamson et al. , 1 956). The results
passage. Thus, it was concluded that blind
of inoculating chick embryos by the chorio
passages were unnecessary in the diagnosis
of Newcastle
allantoic, amniotic and sub-allantoic
disease (Beaudette, 1948a).
methods have been described by Burnet
In contrast, where the virus content of
the inoculum is extremely low, some (1942).
Macroscopic Lesions. These vary with
degree of virus adaptation to the embry-
the strain of infecting virus. With the more
onating egg may be necessary (Brandly
et al, 1946d; Anon., 1959).
virulent (velogenic) strains, small or large
haemorrhages are common, especially on
the wings (Iyer, 1943) and legs (Iyer and
Embryonic Mortality Dobson, 1 940) , and over the cranium and
Suspensions of trachea, lung and dorsal body surfaces (Jungherr et al.,
spleen have produced death in 72 hours 1946). The yolk sac is usually congested.
in 12-day embryos (Perez and Gonzalez, With the lentogenic strains, haemorr
1951). Using 9- to 10-day embryonating hages are seldom seen; but stunting and
chicken eggs, Vrtiak and Polony (1962) curling of embryos dying on the fourth
examined 32 strains of Newcastle disease and succeeding days have been observed

69

(Hitchner et al., 1951; Anon., 1959).
The curling of embryos into a ball-like
form has been considered pathognomonic
of infectious bronchitis (Fabricant,
1949b).
Microscopic Lesions.
(Burnet, 1942) and in the allan
embryos
toic membrane (Granoff, 1955). The
development of Newcastle disease virus
in the chicken embryo has been measured
by an increase in the concentration of
The chorio proteins in the infected allantoic fluid

allantoic membrane has shown prolifera (Kilbourne and Horsfall, 1949). This
tion of the ectoderm with vacuolation (Iyer increased protein concentration has been
and Dobson, 1940), increased density of attributed to the reaction of the develop
the cytoplasm, of the cells
and necrosis ing embryo. After examination of electron
(Burnet and Ferry, 1934; Kilham et al., micrographs, Mussgay and Weibel (1962)
1951). Other changes have been difficult to concluded that morphologically intact
differentiate from non-specific lesions particles of Newcastle disease virus were
(Jungherr et al, 1946). Thus, in an experi capable of entering cells of the develop
ment conducted by Liu and Bang (1953), ing chicken embryo.
a virulent strain of virus did not produce
significant changes in any organ of em
bryos up to the time of death. 7~ De-Embryonated Eggs
The mesoderm has shown haemorrhages that de-embry-
Greuel (1959) found
and oedema (Burnet, 1936). In addition, be used more rapidly
onated eggs could
defective development of the lens, otocyst
and more efficiently than embryonated
or caudal portion of the neural tube has
eggs. The virus was demonstrated in 37
been described (Williamson et al,
1956). infected fowls
out of 45 experimentally
Other microscopic lesions include vacuola
by the inoculation of brain suspension on
tion and cytoplasmic disintegration of
to the chorio-allantoic membrane of de-
cells in tracheal smears (Burnet, 1942); In contrast, only 29
embryonating eggs.
inclusion bodies in cells of allantoic mem
tissues were positive by the inoculation
brane and embryonic liver smears (Collier
of chicken embryos. De-embryonated eggs
and Dinger, 1950) ; and multiple capillary
permitted the use of a larger inoculum and
haemorrhages in the spinal cord (Jung
were not affected by factors which kill
herr et al, 1946). embryos (Greuel, 1963b). In
chicken
comparative studies with de-embryonated
Distribution of Virus in Embryos eggs in which the chorio-allantoic mem
brane has been either removed or left
Newcastle disease virus has an ability virus
intact, virtually all the inoculated
to spread throughout the developing em-
has been adsorbed to the chorio-allantoic
bryonating 1948). Virus con
egg (Bang,
cells (Nadel and Eisenstark, 1956).
centration and haemagglutinating activity
have been found to vary in different fluids
and tissues of infected chick embryos Tissue Cultures
(Burnet, 1942; Hanson et al, 1947), as
in Figure 15. These differences
illustrated Growth of the virus has occurred in a
have been associated with strain of virus medium consisting of chick embryo tissue
and route of inoculation, but the size of and plasma (Topacio, 1934), and in cul
the inoculum has not affected the final tures of cells from a variety of animal
species (Bankowski et al, 1960;
Brandt,
titre of the virus in the allantoic fluid
(Bang, 1948). 1961; Das and Goldberg, 1961; Fontanelli
Evidence of very active virus multipli et al, 1960; Franklin et al, 1957; Gelen-
cation has been obtained in the viscera of czei and Bordt, 1960; Mason and Kauf-

70
 SELECTIVE INFECTION BY NEWCASTLE DISEASE VIRUS

Haemagglutination
Titre

Allantoic Fluid
Amnionic Membrane
Amnionic Fluid

Hours Post Inoculation

(Redrawn from Hanson ef al, 1947)

Figure 15.

man, 1961b; Rubin et al, 1957; Vrtiak cells in plasma media or roller tubes, re
ef al, 1960). Studies with chick embryo ported by Gey and Bang (1951) and Bang

71
( 1 953a) , have been reviewed by Lynn and bryonating eggs for the titration and isola
Morgan (1954). tion of the virus (Fastier, 1954; Gold
Growth in tissue culture has been used wasser and Kohn, 1957; Matewa, 1960).
as a means of virus identification because In addition, tissue cultures have been a
the cytotoxic effect has been shown to be means of distinguishing a mildly patho
caused by the viral particle (Prince and genic from a more pathogenic strain of
Ginsberg, 1957). Treatment of field virus (Mussgay, 1960). The difference in
strains of Newcastle disease virus with the degree of cytopathic effect between
nitrous acid has resulted in mutations strains has coincided with their patho
characterized by different plaque appear genicity for chickens (Subramanyam and

ances in tissue culture (Granoff, 1961; Pomeroy, 1 960) . Thus a virulent strain of
Thiry, 1963). It is considered that the kill Newcastle disease virus has destroyed
ing of mammalian tumour cells by New fibroblastsin tissue culture more rapidly
castle disease virus is not solely a surface than an avirulent strain (Bang and War
reaction (Prince and Ginsberg, 1957). wick, 1957). From the economic point
Serum neutralization tests have been per of view, Jakubik (1962) considered the
formed using cytopathogenicity or hae- tissue culture method to be 85 per cent
magglutinin formation for determining cheaper and to involve about half the work
endpoints (Crowther, 1963; Goldwasser required for isolation in the chick embryo.
and Kohn, 1957; Levine and Sagik, 1956;
Mason and Kaufman, 1955; Pigoury et
al,1962; Rubin and Franklin, 1957; Seif-
Mixed Virus Infections
fert, 1955; Subramanyam and Pomeroy, Considerable strain variation in the

1960). resistance of Newcastle disease and other

According to Seadale and Winterfield avian viruses to inactivation by physical-
(1956), tissue culture titrations based on chemical means has been found (Quiroz
cytopathogenicity are somewhat more sen and Hanson, 1958). A highly pathogenic
sitive than virus assays in chick embryos. strain of Newcastle disease virus has not
The virus has been titrated in tissue cul been inactivated by hydroxylamine, where
tures using indicators to show colour as an attenuated strain has (Rott and

change resulting from the metabolism of Schafer, 1962). Moreover, it has only
non-infected cells (Durand and Eisen- been possible to resolve certain mixtures
stark,1959). This method has also been of Newcastle disease, laryngotracheitis,
found suitable for titrating the specific infectious bronchitis and fowl pox viruses
immune serum. procedures (Quiroz
by physical-chemical
Titration by the plaque method (Rubin and Hanson, 1958). Brandly et al.
et al., 1957) has given titres approximate (1946d); Hanson (1954) and Thompson
ly 1 log. less than embryonated egg titres (1954) showed that hyperimmune sera

expressed as LD.n units per ml. (Bower, would satisfactorily neutralize the virus
1958). A close correlation has been found of Newcastle disease, fowl plague and
in in vitro titration between the haemag- infectious bronchitis and a chronic res
glutinating and haemadsorbing properties piratory disease agent in dual infections
of the virus (Rossi, 1961a). Tissue cul so that the presence of each virus could
tures have been found comparable to em- be demonstrated in embryonating eggs.

72
 DIAGNOSIS BY INOCULATION

Chickens Pigeons
The use of susceptible chickens, free In susceptible pigeons, symptoms usual
from passive antibody, for the isolation ly appear six days after inoculation
and identification of Newcastle disease (Figure 16) and death occurs a few days
virus has been valuable under certain cir later still (Doyle, 1935). The features of
cumstances (Anon., 1959b; Beach, 1943; the experimental disease in pigeons have
Brandly et al., 1946d; Gordon and Asplin, been described by Dobson (1939), Doyle
1947; Walker, 1948). The use of suscep (1935) and Kuppuswamy (1955). The
tible chickens was regarded by Grausgru- site of inoculation
of the pigeon was shown
ber (1958) as being time-consuming and by Kaschula (1951) to influence the de
expensive. Furthermore, due to the varia velopment of symptoms. Pigeons inocul
tion in the susceptibility of individual ated in the neck showed a more rapid
fowls and differences in the virulence of course and developed paralysis of the neck
field strains of virus (Vrtiak and Polony, quicker than those inoculated in the wings
1962), Beller and Siegmann (1955) con and legs.
cluded that diagnosis by the inoculation Olah and Palatka (1962) made a com
of fowls was unsatisfactory by itself and prehensive study of the pathogenicity of
should be supplemented by serological several strains of Newcastle disease virus
methods. Nevertheless, of
the inoculation by intracerebral inoculation of pigeons.
a healthy fowl has facilitated the isolation An intracerebral pathogenicity index was
of virus (Scott et al., 1956). Karzon and used, giving death a value of 4, illness 2
Bang (1951a) showed that whereas a and no ill effect zero. A virulent virus gave
velogenic and a mesogenic strain increased an index of 3; a strain designated Lederle
at the same rate in the extra-neural tissues, was 2; the Roakin strain was 0.9; and Bl,
the velogenic strain caused an increased F and LaSota each gave an index of zero
growth rate in the brain. (Olah and Palatka, 1963).

Figure 16. — Newcastle disease in the pigeon. (Crown Copyright. Reproduced by permission
of the Controller of Her Malesty's Stationery Office, London.)

73
Ducks Dougherty, 1956; Kilham et al, 1952; Liu
and Bang, 1952; Upton et al, 1955).
When compared with chickens, other
Nevertheless, Luttrell and Bang (1958)
domestic poultry are more resistant to
have reported that kittens and adult cats
artificial infection with the virus. How with
can be infected by several routes
ever, ducks have been used by Schofield
Newcastle disease virus; and the intracere
and Hutsen (1952) in the identification bral inoculation of young hamsters has
of the disease.
been used for the recovery of virus from
a human infection (Reagan et el, 1956).
Laboratory Mammals Strains which have induced severe involve
Laboratory mammals are generally ment of the central nervous system in
resistant to the virus unless inoculated in- young mice have generally been highly
tracerebrally (Brandly et al, 1946d; lethal for chickens (Carlotto, 1954; Upton
Brueckner et al, 1951; Groupe and etal, 1953a).

DIAGNOSIS BY CHALLENGE EXPOSURE

Definitive evidence of a previous New the inoculation of fully susceptible
castle disease infection may be obtained chickens.
The value of the challenge method of
by inoculating suitable live specimens with
diagnosis was demonstrated by Bankow-
a known dose of fully virulent virus
ski (1961c) who found some chickens
(Brandly, 1959; Gordon and Asplin, with a negative HI titre refractory to a
1947). This test should be controlled by challenge dose of virus.

IDENTIFICATION OF NEWCASTLE DISEASE VIRUS

Newcastle disease virus has been given ens (neuropathic index).
the name Myxovirus multiforme (An- — Pathogenicity for day-old chick
drewes et al., 1955; Waterson, 1962). ens by intranasal (IN) and intra
An early name for the virus was Tortor muscular (IM) inoculations.
furens (Barger et al, 1958). Criteria for — Pathogenicity for 3-day-old chick
establishing its identity have been given ens (IN, IM).
by Brandley et al. ( 1946d) , Wilner (1964), — Pathogenicity for 5-week-old
and Andrewes and Worthington (1959). chickens (IN, IM).
Ozawa and Chow (1958) have sum — Disease manifestation.
marized these methods and criteria and — Antigenicity (protective antibody).
have used them to identify a field isolate. — Serum neutralization (homolog
Various criteria for identifying New ous LDS0 neutralized).
castle disease virus have been reported — Haemagglutination with chicken
by Hanson (1956). They have been
et al. and other animal erythrocytes.
summarized by Ozawa and Chow (1958) — Haemagglutination inhibition tests.
as follows: — Heat stability of haemagglutination
— Pathogenicity for embryonating at 56°C.
chicken eggs. — Heat stability of embryo infectivity
— Mean death time for minimum at 56°C.
lethal dose. — Mouse toxicity (neuropathic
— Pathogenicity for day-old chick index).

74
 DIFFERENTIAL DIAGNOSIS

The great variety of symptoms and ways, was a separate entity caused by a
lesions exhibited in Newcastle disease has virus immunologically distinct from the
been outlined in the preceding section. virus of fowl plague. Although this distinc
The occurrence of an asymptomatic form tion between the two viruses was not at
has been discussed on page 57. These first recognized by some authors (Hutra
features of the disease make differential et al., 1938; Leynen, 1935; Manninger,
diagnosis difficult unless recourse to 1932, 1936; Picard, 1934), it was con
laboratory procedures is adopted. New firmed by other workers
(Burnet and
castle disease virus is classified as a mem Ferry, 1934; Lush, 1943; Nakamura et al.,
ber of the myxovirus group. The term 1934; Purchase, 1931; Schafer, 1950).
"avian myxovirus" includes fowl plague Thus, Ivanova et al.
(1963) showed that
virus, virus N, duck influenza virus, tern chickens immunized with vaccine strains
virus, myxovirus Yucaipa and perhaps of Newcastle disease virus were fully sus
others. ceptible to inoculation with three different
In the following pages, the similarities strains of fowl plague virus. Another dif
and differences between Newcastle disease ference between the two viruses was that
and a number of other avian diseases are fowl plague virus agglutinated the ery
summarized. This summary is not by any throcytes of a larger number of species and
means complete, but it may serve as a with a more stable antigen-antibody union
guide. than with Newcastle disease virus (Hall-
auer and Kronauer, 1954; Kunst, 1949).
Viral Diseases In more recent studies of this problem,
Schmidt (1960) has concluded that the
Fowl Plague transformation of Newcastle disease virus
The difficulty of distinguishing between into classical fowl plague virus is a pos
Newcastle disease and fowl pest (now sibility. Another consideration is that
generally called fowl plague) that faced Newcastle disease virus may be identical
early investigators has been discussed by or antigenically to atypical fowl
related
Hutra et al. (1938); Jacotot (1950); plague virus (Beller, 1953; Schafer, 1950).
Jacotot and Vallee (1949); Lesbouyries There is, therefore, constant need to dif
(1941, 1951); and Traub (1942). ferentiate between the two diseases, especi
In Europe and some other areas where ally in countries where both might co
fowl plague (geflugelpest) was recognized exist (Lucam, 1949c; Vittoz, 1938, 1963).
before 1926, the later appearance of the Such differentiation has been made by
classical form of Newcastle disease was using fowl plague and Newcastle disease
often not immediately identified. Thus, in vaccines(Nechvatal, 1950), and by cross
some instances, laboratory results have immunity tests (Kujumgiev, 1950). In
shown that a clinical diagnosis of fowl other instances, attempts to differentiate
plague, or a disease resembling fowl between Newcastle disease and fowl
plague, was made in error (Bakos and plague have been made without isolation
Nordberg, 1949; Berke and Golem, 1949; of the virus (Anon., 1948). In addition
Jacotot and Vallée, 1949). to the serological and virus isolation tests
In the original identification of New so far discussed (pages 59 to 74), the
castle disease virus (Doyle, 1927, 1935), following methods have been used to
it was shown that Newcastle disease, al distinguish between Newcastle disease
though it resembled fowl plague in many and fowl plague:

75
 TABLE 9 — Characteristics of Classical Fowl Plague
and Typical Newcastle Disease Compared

Fowl plague Newcastle disease

Incubation period Average 1 54-214 days. Average 4-6 days.

Duration of disease
Symptoms
Lesions
Infectivity of blood
Virus haemagglutination
Virus haemolysin1
Lesions in inoculated
embryos2
A few hours or longer.
Often none. Malaise, diarrhoea,
oedema of head and append
ages.
May be none. Haemorrhages on
heart and in proventriculus and
intestines. Sero-gelatinous exu
date in lungs, pericardium and
subcutis.
Virus present in high dilutions
of blood.
With erythrocytes of many spe
cies. Slow elution of virus.
Negative.
Cytoplasmic inclusions absent.
Haemorrhages in skeletal mus
cles.
Three days or longer.
Respiratory symptoms, diar
rhoea, nervous disturbances
in survivors.
May be none. Haemorrhages
often involving lymphoid
patches of intestine.
Virus present in lower dilu
tions.
Quicker elution of virus.
Present.
Cytoplasmic inclusions. Pe-
techiae on body surface.

' Andrewes and Worthington

(1959)
2 Burnet and Ferry
(1934)

Pigeons. During the years immediately examined by Demnity and Schneider

following the first identification of New (1950).
castle disease, the virus was differentiated Physical properties. The "Herts" strain
from that of fowl plague principally by of Newcastle disease virus has been found
using pigeons, which are susceptible by to have a pH stability of 5.5 to 7.5 and a
inoculation to Newcastle disease but re mean size by electron microscopy of 150
sistant to fowl plague (Dobson, 1952; ±50 microns; whereas the corresponding
Doyle, 1933; Purchase, 1931). Additional figures for fowl plague virus have been
references are given on page 73. pH 7.0 to 9.0 and 83 ±15 microns (Elford
Mice. Although this animal is seldom etal, 1948).
used, results indicate that mice are gen Clinical symptoms and post mortem
erally more susceptible to the virus of fowl findings. The major differences between
plague than to that of Newcastle disease the typical forms of Newcastle disease
(Kranevald and Nasoetion, 1941; Kunst, and fowl plague (summarized in Table 9)
1949). However, little difference has been have been described by Beaudette ( 1951c),
reported by Andrewes and Worthington Fritzsche and Gerriets (1962), Jezierski
(1959). (1953), Jungherr et al. (1946) and Stubbs
Haemagglutination. The agglutination (1946, 1952).
of rabbit erythrocytes by fowl plague virus
but not by Newcastle disease virus has
Avian Encephalomyelitis
been used as a differential (Abdel
test
Aziz et al, 1960) . The possibility of using The comparative pathology of avian
the HI test with high litre sera to distin encephalomyelitis and Newcastle disease

guish between the two viruses was has been reported by Jungherr and Minard

76
(1944) . In avian encephalomyelitis, the Avian Leucosis Complex
central nervous system has shown exten
The neuralymphomatosis component of
sive perivascular granulomatous foci
the complex has been classified as Marek's
whereas the peripheral nervous system
disease (Biggs, 1962); and the differentia
has shown some myelin degeneration. In
tion of this form of the complex from
this disease, there is no respiratory symp
Newcastle disease has been described by
tom and no inflammation of the respira
Mohr (1953). The main difference has
tory tract. In the developing chicken em
been the presence in Newcastle disease
bryo, avian encephalomyelitis virus has
of lymphocytic infiltrations in the lungs
caused no lesion in visceral organs
and characteristic cellular reactions in the
(Casorso and Jungherr, 1959).
central nervous system.

Infectious Bronchitis Duck Plague

The difference between infectious bron The first outbreak of this disease was
chitis and Newcastle disease has been reported in 1923 in The Netherlands, and
outlined by Beaudette (1951c). Delaplane since then a small number of outbreaks
(1945) , Fabricant (1949b, 1950) and has occurred in that country (Jansen,
Stubbs (1946).
1961). Typical lesions include multiple
Recently, the gel diffusion technique the
petechial haemorrhages throughout
has been applied to the diagnosis of New
body. The causal virus has been shown to
castle disease and infectious bronchitis be immunologically distinct from the
(Guillon et al., 1962). A diagnostic pro viruses of fowl plague and Newcastle dis
cedure utilizing this technique has been
ease (Jansen, 1951; Jansen and Kunst,
based on the detection of an increase in
1949).
precipitin titre between two successive
bleedings (Witter, 1962).
Infectious bronchitis virus, in contra Virus N
distinction to the virus of Newcastle This virus has been isolated from dis
disease, has failed to agglutinate sperma eased fowls and, in some respects, the
tozoa (Chu, 1953). virus has resembled Newcastle disease and
fowl plague viruses. Although no serolog
ical relationship has been found between
Infectious Laryngotracheitis
virus N and Newcastle disease virus, some
Factors that enter into the differential antigenic relationship between virus N and
diagnosis of this disease have been re fowl plague virus has been demonstrated
ported by Beaudette (1951c), Delaplane (Dinter, 1949; Dinter and Bakos, 1950).
(1945), Stubbs (1946), and Woernle and Virus N has been placed in the myxovirus
Brunner (1961). group (Rott and Schafer, 1960).
The use of the agar gel diffusion tech
nique in the diagnosis of infectious laryn
gotracheitis has been reported by Jordan Tern Virus
and Chubb (1962). Precipitin antigens Chickens artificially infected with this
have been demonstrated in the absence virus have shown symptoms after three
of virus recovery by egg inoculation. days and have died a few days later. Al
Under field conditions in Italy, simul though macroscopic changes have been
taneous outbreaks of laryngotracheitis and slight, histological lesions of focal tissue
Newcastle disease have been described necrosis have been extensive in severe
(Anon., 1946c). cases. However, the pathological pattern

77
has been considered to be distinct from have also been demonstrated in artifically
that of Newcastle disease or fowl plague infected turkeys (Gale et al, 1960; Page,
(Urs and Becker, 1963). 1959).
In a natural outbreak in chickens, an

Turkey Meningo- Encephalitis ornithosis agent of relatively low virulence

has been recovered from 2-day-old chick
The first report of this disease was made ens exhibiting nervous symptoms similar
by Komarov and Kalmar (1960) who to those of avian encephalomyelitis (Storz
described a disease of turkeys character etal., 1963).
ized by progressive paralysis of legs and
wings and a severe non-purulent lym
phocytic meningo-encephalitis. The causal
Nutritional Deficiencies
virus was classified by Porterfield (1961) Nutritional deficiencies which can cause
as a member of Group B of the arthropod- symptoms and lesions resembling those of
borne viruses. Newcastle disease include:
— Riboflavin deficiency.
Bacterial Diseases — Vitamin E deficiency (nutritional
encephalomalacia) .
Chronic Respiratory Disease — Fatty acid deficiency (Hopkins et

Typically, the disease caused by Myco al, 1963).

plasma gallisepticum has a long incubation
period of 5 to 21 days and runs a more Toxic Drugs and Plants
prolonged course (Chu, 1958). Mild re
Toxic substances which can cause symp
spiratory infections caused by this or other
toms and lesions similar to those of New
agents are often aggravated by exposure
castle disease include:
to avian respiratory viruses (Bankowski, — Agricultural pesticides (Barden and
1 961b) . A pleuropneumonia-like organism
Paver, 1961; Gaafar and Turk,
with neurotropic characteristics has been
1957).
isolated from the brain of a turkey with — Alkyl organophosphorus com
symptoms of encephalitis (Cordy and
(Davies et al, 1960; Lan
pounds
Adler, 1957.)
caster etal., 1960).
— Nitrofurazone (Brion and Fontaine,
Avian Pasteurellosis 1958; Klimes and Kruza, 1962).
and turkeys, with a disfunc
Chickens
— Toxic plant seeds (Kelly et al., 1961;
tion of the central nervous system and Placidi, 1954c).
symptoms similar to those of Newcastle
disease, have been found infected with a Avian Respiratory Diseases
Pasteurella organism localized in the brain
etal, The differential diagnosis of respiratory
(Fenstermacher 1946).
diseases of poultry (Table 10) has been
summarized by Abrams (1961 ), Caporale
Ornithosis (Psittacosis) (1961), Delaplane (1945), Durrell
In the turkey, a common lesion of orni (1952), Chu (1958), Gordon (1956,
thosis is an inflammation of the air sacs, 1961), Jungherr (1953), McMartin
peritoneum and pericardium. These lesions (1963) and Schyns (1961).

78
 TABLE 10 — Avian Respiratory Diseases — Some Characteristics
(Modified from Gordon, 1963)

Disease Incubation Duration Spread Clinical Pathological

period symptoms features

Mycoplasma 10-21 days months slow Profuse oculona Purulent exudate

gallisepticum sal discharge, sinuses, may
swollen face, extend to lower
difficult respiration respiratory tract
Coli 10-21 days many slow Difficult Pericarditis and
septicaemia weeks respiration, loss of airsacculitis
weight, rejected
at market
Mixed 14-21 days months slow Profuse nasal dis Purulent
infections charge, difficult exudation,
'C.R.D., respiration, gasp pericarditis,
ing, poor growth, airsacculitis
etc.
Infectious 1-3 days brief rapid Sneezing, gasping, None, or some
bronchitis depressed egg mucus in bronchi
production and air sacs
Infectious 4-21 days weeks to variable Gasping, cough Blood or diphther
laryngo- months ing, blood clots itic material in
tracheitis trachea
Fowl pox 7-1 4 days 3-4 slow Lesions on comb, As for symptoms
weeks wattles, face, mouth
Haemophilus 1- 2 days 2-3 rapid Slight oculonasal Pus in sinuses
gallinarum weeks discharge
Aspergillus 2—4 days weeks rapid Gasping Nodules in lungs
fumigatus

79
 PART III: Control Measures
CONTROL BY SLAUGHTER
Effectiveness in Various In many countries in which Newcastle
disease has a variable distribution, the
Countries
importation of poultry and poultry pro
The concern shown by national disease ducts is strictly controlled. In countries
control agencies over the control of New where Newcastle disease does not exist at
castle disease is reflected by the fact that present, there is complete prohibition on
in 1962 the disease was notifiable in 84 the importation of live and dead poultry
countries(Table 2). and hatching eggs. To reduce the extent of
In some countries where slaughter mea these restrictions, it has been suggested
sures have been applied they have been that control measures be applied on the
eminently successful; in others they have basis of "infected region of a country" in
failed to prevent the disease from becom stead of "infected country" ( Vittoz, 1 964).
ing established. The variety of ways in
which the disease can spread, the actual
Southeast Ireland
numbers of individual poultry involved,
and prevailing trade and management In County Kilkenny, southeast Ireland,
practices have appreciably handicapped in January 1950, two outbreaks of New
disease eradication. castle disease were reported on the same
Newcastle disease has, however, been day on different premises (Anon., 1951-
eradicated completely from certain coun 52). Immediately, orders for the restric
tries and, in others, large regions have been tion of movement and for slaughter with
kept free of the disease for extended compensation were invoked under the
periods. In Canada, for instance, although Diseases of Animals Act. During subse
the disease has persisted since 1950 in quent field investigations, special emphasis
some of British Columbia, no
districts was placed on tracing the chain of infec
known outbreaks occurred in several prov tion to and from infected flocks. In addi
inces, including Saskatchewan, Manitoba tion, the activities of poultry dealers were
and those on the Atlantic coast, from 1 957 closely investigated. Although the disease
to 1963. Similarly, in Scotland, some coun was confirmed on only 14 premises, over
ties, including West Lothian, Perth and 1,000 premises were visited and tens of
Midlothian, were free from Newcastle thousands of poultry were examined.
disease from 1951 to 1961 (Anon., 1962b) Within two weeks of the first outbreaks,
although there were outbreaks in other it was apparent that control measures were
parts of the country. In 1962, a number meeting with success; and during the next
of countries, including Czechoslovakia, month the size of the infected area was
Brazil, Bulgaria, Peru and South Africa, progressively reduced. At the end of
reported that Newcastle disease was con March 1950, all restrictions were removed
fined to certain regions (FAO-WHO-OIE, except that restocking of infected premises
1962). was not permitted until four to six weeks

80
after cleaning and disinfection had been carried out under the Fowl Pest Order,
completed. and certain ancillary measures were
In this outbreak in southeast Ireland, adopted to control the movement of poul
71 flocks, involving 1,235 poultry and try, hatching eggs and carcasses. At first,
40,202 day-old chicks, were slaughtered. measures were taken to restrict the im
portation of poultry and hatching eggs and
to ensure the boiling of waste food and
Australia the zoning of imported carcasses. Later,
In Australia, the first outbreak occurred infected area restrictions were imposed
near the town of Wonthaggi in the state and the disinfection of poultry premises
of Victoria in November 1930 (Johnstone, and vehicles was required. The general
1931; Albiston and Gorrie, 1942). The procedure adopted to deal with the out
disease spread from the original infected breaks has been described by Reid ( 1961 ) .
area, and during the following two months Failure to report outbreaks was thought
it was identified on 3 1 other farms within to be handicapping the slaughter policy
a radius of 20 miles of Melbourne. In all, (Andrews, 1948), and by the end of 1947,
72 farms eventually became infected and 2,222 outbreaks had been confirmed
22,284 head of poultry were involved. (Figure 17).
The whole state was not considered free In 1947, both the peracute (velogenic)
from the disease until June 1931 (John and the subacute (mesogenic) forms of
stone, 1933). the disease were recognized. The control
The second outbreaks in Victoria, Aus measures adopted were successful in eradi
tralia, started in October 1932 and lasted cating the peracute type by 1953, but the
until March 1933. Restrictions on move of the disease could not be
less acute forms
ment of poultry, quarantine of infected eradicatedfrom many areas. Scotland re
farms and controlled slaughter were en mained free from the disease from 1952
forced, and the outbreak was quickly to 1957.
stopped. While the outbreak lasted, certain Another series of events which reveal
auction rooms were reserved for the sale a number of epidemiological aspects of the
of poultry from outside the quarantine disease is associated with its eradication
area. Other facilities were designated for from the county of Lancashire (Figure
the slaughter and evisceration of healthy 17). Here, concentrated control efforts
poultry from within the quarantine area were begun in 1956; and by 1958 the
(Johnstone, 1933). epidemic in one area of the county had
been terminated. For over two years there
was no recrudescence of the disease in the
Great Britain
area. This success was achieved only by
The control of Newcastle disease by full and rigorous enforcement of the
slaughter in Great Britain has been de slaughter policy (Ritchie, 1962). Whether
scribed in a number of publications this success in one section of the county
(Andrews, 1948; Anon., 1962b; Asplin of Lancashire would have been possible
et al., 1949; Callender, 1958; Dobson, in the face of widespread outbreaks in
1949; Gordon et al, 1948; Gordon, 1961; other parts of the country is debatable,
Gordon and Asplin, 1947; Reid, 1961). In July 1960, the Departmental Com
An interesting episode from the epi mittee on Fowl Pest was appointed by the
demiological point of view started in British Minister of Agriculture, Fisheries
February 1947 when a small number of and Food. Their report (Anon., 1962b)
outbreaks occurred in the county of was presented to Parliament in March
Somerest. Slaughter of affected flocks was 1962. The Committee concluded that

si,
 NUMBER OF OUTBREAKS OF NEWCASTLE DISEASE IN
GREAT BRITAIN AND LANCASHIRE, 1946-1961

1946 '47 '48 '49 *50 '51 '52 '53 '54 '55 '56 '57 '58 '59 '60 '61

(Modified from Collender, 1958)

Figure 17.

control of Newcastle disease rather than Pest Order, the slaughter of flocks affected
eradication should be the immediate aim. with the less acute forms of the disease
As a result, the voluntary use of killed vac ceased in March 1963. Slaughter was to be
cines was encouraged and, although the used only should the peracute form of the
disease was still reportable under the Fowl disease re-appear.

82
Switzerland of mesogenic and strains of
lentogenic
virus have resulted in the abandonment
Control procedures have been adopted of an eradication policy based on compul
in Switzerland with very satisfactory re
sory slaughter and the payment of com
sults (Hess, 1962, These proce
1963). pensation (Fritzsche, 1963). A very
dures have included :
similar situation occurred in The Nether
— The testing of random samples of im lands in 1950 (Hoekstra, 1961). By way
ported poultry for the presence of of contrast, in Sweden, where the first out
Newcastle disease virus. break occurred in 1947, control measures,
— The prohibition of live imports. have eradicated the disease. The last out
— The prohibition of any form of vac break in Sweden occurred in 1956 (Lind-
cination. gren, 1963).
— The slaughter of all infected flocks
immediately after diagnosis has been
established. Yugoslavia
The testing of samples of imported Compulsory vaccination using a live
poultry for the presence of Newcastle virus vaccine has led to a considerable
disease virus was begun in 1947. This reduction in the number of infected farms
procedure was associated with a progres in Yugoslavia (Fiolic, 1957). To accom
sive decline in the number of outbreaks plish complete eradication of the disease
in the country. Field outbreaks ceased in in that country, strict control measures
1960 (Hess, 1963). have been enforced.
Switzerland is frequently exposed to in
fection through large importations of
slaughtered poultry (18,400 tons in 1961), South Africa
and the fact that the national flock of six
million fowls was free from Newcastle The incidence of Newcastle disease and
disease in 1960 has been attributed largely the control measures adopted in the Re
to the routine examination of samples of public of South Africa have been sum
poultry carcasses (Hemsley, marized by Anon. (1950b) and Kluge
imported
1961). (1964). The first outbreak was diagnosed
in the Durban area in 1944 (Figure 3).
Outbreaks which were identified in 1944,
France 1949, 1950, 1951, 1953, 1954, 1961 and
In France, a slaughter policy, in which 1962 were all of the velogenic type with
the compensation paid did not exceed high mortality and were eliminated by
three-quarters of the commercial value of slaughter and other strict control measures
the slaughtered birds, was hard to enforce (Anon., 1950b). These measures included
(Fontaine, 1963). This was because many the designation of infected areas; the con
farmers failed to notify the authorities of trol of movement of poultry and poultry
the existence of the disease, and because products; quarantine and disinfection.
diagnosis was difficult when a strain of Voluntary vaccination, including the use
virus of low virulence was involved. of live vaccines, was permitted within the
infected area. Vaccination teams worked
under official supervision. In July 1960,
Germany, the Netherlands and
the mild (lentogenic) type of Newcastle
Sweden disease was diagnosed in South Africa for
In western Germany the rapid spread the first time (Kluge, 1964). This form
of Newcastle disease and the appearance of the disease was eliminated by 1961.

83
Canada try population was dense, control mea
sures failed to eliminate the disease. As a
In Canada, Newcastle disease was first
of On
result, the slaughter policy was discon
recognized in 1 948 in the province
tinued in 1954.
tario. An eradication policy was immedi
The reasons for the termination of the
ately enforced under the Animal Contagi
Canadian slaughter policy may be sum
ous Diseases Act, and during the follow
marized as follows (Lancaster, unpub
ing three years only a small number of
lished data) :
outbreaks were identified in the province.
However, in 1951 and 1952 the number — The exposure of the Canadian poul
of outbreaks increased to a total of 68. In try industry to the continued risk of
the province of British Columbia, New the introduction of the disease in im
castle disease first appeared in February ported live poultry and hatching eggs.
1950 and spread very rapidly. The peak —The difficulty of identifying the mild
of the epizootic occurred in May 1950 form of the disease in the field.
and 133 flocks were involved. In the other — The favourable progress made in the
provinces of Canada, only sporadic and development of Newcastle disease
isolated outbreaks have occurred. vaccines.
A federal policy was carried
slaughter — The characteristics of the Canadian
out in Canada for six years: from Febru poultry industry, especially the wide
ary 1948 to March 1954. This policy was spread movement of live poultry,
supported by two additional measures: hatching eggs and poultry carcasses,
one requiring certification that all im which did not assist disease eradica
ported live poultry and hatching eggs tion procedures based on slaughter.
originate from flocks free of Newcastle
disease; the second requiring the cleaning Although the official slaughter policy
and disinfection of poultry crates after was terminated in Canada in 1954, New
each use (Wells, 1948). In certain prov castle disease has continued to be named
inces, eradication of Newcastle disease in the regulations made under the Animal
was achieved; but in areas where the poul Contagious Diseases Act.

ACTION OF CHEMICALS ON NEWCASTLE

DISEASE VIRUS
A number of different methods have that the virus (usually as diluted crop
been adopted to evaluate the viricidal contents) was inactivated after 30 to 40
activity of disinfectants against Newcastle minutes by 1 and 2 per cent formalin (37
disease virus. Using an emulsion of mouth per cent formaldehyde) and by 1:10,000
exudate from infected chickens as the solution of potassium permanganate. One
source of virus, Doyle (1927) examined procedure used successfully in the field
the effect of a number of disinfectants was to disinfect the daily
environment
(Table 11) and concluded that Newcastle with 2 to 3 per cent creolin solution and
disease virus was more susceptible to the to supply the poultry with drinking water
action of alkali than to that of acid. containing 1 : 2000 potassium permangan
Kohn (1958) showed that when New ate (Farinas, 1930). Similar recommenda
castle disease virus was brought into con tions for field control have been made by
tact with gizzard contents at pH 2.6, its Johnstone (1931) and Haddow (1941).
viability was considerably reduced. Stud Tilley and Anderson (1947) used a
ies reported by Farinas (1930) showed method whereby test solutions were mixed

84
 TABLE 11 — Effect of Disinfectants on Newcastle Disease Virus
(Doyle, 1927)

Minimum strength required to kill the virus after

Disinfectant contact for one hour at room temperature (1 5'C)

Methyl alcohol 1 2
Ethyl alcohol 1 2
Ether 1 5
Acetone 1 2
HCI Not killed by 1 25
NaOH N/50
Antiformin Not killed by 1 100
Formalin Not killed by 1 50
Mercuric chloride Not killed by 1 100
Oil of cloves Not killed by 1 100
Carbolic 1 20
Izal 1 500
Potassium permanganate 1 5,000
Copper sulphate 1 20
Hydrogen peroxide 1 2
Lysol 1 5,000
Cresol 1 1,000

with amnio-allantoic fluid as the source reported. When

similar to those previously
of virus. The mixtures were held at 20°C Asplin (1949) studied the action of for
and, after 5 minutes exposure, aliquots malin on Newcastle disease virus he found
were removed and inoculated into embryo- that a virus suspension remained active
nating eggs. A
similar procedure was after exposure to 2 per cent commercial
adopted by Cunningham (1948), except formalin for one hour at 18.3°C (Table
that a 3-minute reaction time was used. 12). On the other hand, Beamer and Prier
When comparing his results with those of (1950) have reported that 0.5 per cent
Tilley and Anderson (1947), Cunning formalin inactivated the virus in 30 min
ham ( 1 948 ) indicated that 2 per cent utes at room temperature.
sodium hydroxide, 1 per cent liquor cre- A different method of assessing the ef
solis saponatus, and 3 per cent phenol fectiveness of distinfectants has been re
were effective against the virus during a 3 ported by Reuss (1957). He placed hy
to5 minute reaction period. Similarly, podermic needles contaminated with New
Weidenmuller (1951) concluded that 2 castle disease virus-infected allantoic fluid
per cent sodium hydroxide was an effec in the disinfectant and then inserted them
tive disinfectant.On the other hand, into embryonating eggs. Using this meth
Moses (1948) reported that the destruc od, Reuss (1957) found that the virus
tion of Newcastle disease virus by 2 per was inactivated by a 10-minute exposure
cent sodium hydroxide was irregular. to 3 per cent formalin solution.
Asplin (1949) generally used a mini Another method of evaluating disinfec
mum virus-disinfection reaction time of tants was studied by Haussman and Grafe
one hour. This precluded a direct com (1957) who reported that there was a
parison being made with other results close relationship between viricidal action
(Anon., 1959). Nevertheless, Asplin and the action on the Newcastle disease
(1949) concluded that his results were virus haemagglutinin. Accordingly, it has

85
 TABLE 12 — Action of Formalin on Newcastle Disease Virus
(Asplin, 1949)

Concentration Time of Temperature Result

of formalin exposure 'F
per cent

0.2 10 days 34-35 Inactive

0.1 16 days 34-35 Inactive
0.05 50 days 34-35 Inactive
0.025 90 days 34-35 Inactive
2.0 1 hour 65 Active
2.0 12 hours 65 Inactive
1.0 12 hours 65 Inactive
0.25 1 hour 98 Inactive
0.1 1 hour 98 Active
0.1 6 hours 98 Inactive

been suggested that disinfectants might be gation of brooder and incubator rooms as
evaluated on the basis of reduction in well as incubators has been reported by
haemagglutination titre. Schmittle and They
Mansfield (1950).
treated approximately 5,500 cubic feet of

Fumigation with Formalin space with 8 U.S. gallons of 37 per cent

formaldehyde solution. When the tempera
Using five different test materials con ture of the building was raised to 37.8"C
taminated with Newcastle disease virus,
and the humidity to 100 per cent, the
Beamer et al. (1949) showed that, in a
fumigation time was 20 hours. This
forced draft incubator, fumigation with
fumigation apparently inactivated New
35 ml. formalin released by 17 gm. potas
castle disease virus on egg shells placed
sium permanganate per 100 cubic feet of
in various locations in the rooms and
space was sufficient to inactivate the virus
incubators.
during a period of 2 to 3 hours. In thick
films of yolk or allantoic fluid, destruction
Other Disinfectants
of the virus was achieved with 70 ml.
formalin and 34 gm. potassium perman The distinfectant properties of a solution
ganate per 100 cubic feet of space. Beamer of the amino acid di(octyl-amino-ethyl)-
et al. (1949) have emphasized that, to pre glycine has been examined. Reuss (1962)
vent thick deposits from protecting the found a 2 per cent solution of this dis
virus, all contaminated surfaces in the in infectant inactivated Newcastle disease
cubator should be cleaned thoroughly be virus in test tubes in 20 minutes; and a
fore fumigation.Greuel (1963a) has con 5 per cent solution inactivated the virus on
cluded that exposure for 5 to 10 minutes a contaminated hypodermic syringe with
to 30 ml. formalin released by 20 gm. in 15 minutes (Reuss, 1963).
potassium permanganate is sufficient to in In tests where disinfectants of undis
activate a considerable proportion of New closed composition have been used against
castle disease virus adsorbed onto sur Newcastle disease virus in different media,
faces. The use of smaller amounts of for the viricidal action has been reduced in the
malin and potassium permanganate has presence of albumen (Grafe and Hauss-
been recommended by Nobrega (1955). mann, 1956; Haussmann and Grafe,
The value of formalin alone in the fumi 1956). There is much less protein in virus.

86
suspensions in allantoic fluid than in organ Using both in vitro and in ovo methods,
suspensions (Grafe and Haussmann, McLimans et al. (1957) tested a variety
1957). of compounds containing a terminal o -
Certain chemicals have been found to ketoaldehyde or o - hydroxy-aldehyde
have little or no viricidal action against grouping against Newcastle disease virus.
Newcastle disease Thus, neither
virus.
One of the compounds, designated "Keth-
ascorbic acid nor cysteine hydrochloride
oxal," was found to be a potent inactiva
at a concentration of 0.2 mg. per ml. has
ting agent in vitro against the virus. Lyo-
had any effect on the virus (Sinha and
philized virus has been inactivated after
Datta, 1950a). In contrast, a low con
centration of a lecithin-like fraction from 4 hours exposure to ethylene oxide (car-
serum has been capable of inactivating boxide) under a pressure of 1,500 mm.;
Newcastle disease virus at 37°C (Utz, while wet virus has been inactivated in 3
1949). hours (Mathews and Hofstad, 1953).

STERILIZATION OF ATMOSPHERES
CONTAMINATED WITH NEWCASTLE
DISEASE VIRUS
Robin (1962) has reported that, under shown that the disinfectant aerosol was
experimental conditions, an aerosol con not harmful to hatching eggs or chicks.
taining volatile organic acids used for Similarly, in a broiler flock, the use of
periods of 3 minutes each day was suffi an aerosol of triethylene glycol has ap
cient to prevent the air-borne transmission peared to reduce the spread of Newcastle
of virulent Newcastle disease virus to disease (Ellis et al., 1952). However,
susceptible chickens. adequate concentration of the vapour may
The effect on Newcastle disease virus of be difficult to obtain under general farm
an aerosol of a glycol mixture containing conditions. The ultraviolet irradiation of
a quaternary ammonium compound was the air of a poultry house by means of
examined by Walker et al. (1953). First "Sterilamps" did not prevent the spread
a fog of the disinfectant was produced of infection during a natural outbreak of
in a chamber; then a fine virus suspension Newcastle disease among broilers in bat
was blown in. Air samplings taken at inter teries, and the disease spread to laying
vals from 10 to 120 minutes showed no hens housed on other floors in the same
evidence of live virus. Furthermore, it was building (Levine and Hofstad, 1947).

CONTROL WITH HYPERIMMUNE SERUM

Early studies on the prophylactic and production of appreciable quantities of
therapeutic value of hyperimmune serum immune (Farinas, 1930; Haddow,
serum
have been reviewed by Beaudette (1943). 1941). To meet this problem, Mitchell
The majority of these studies involved and Walker (1951b) and Spalatin (1948)
small numbers of fowls under field or hyperimmunized horses and they found
laboratory conditions. In general, results the antiserum had considerable neutraliz
were not encouraging. ing power when tested against virulent
One of the difficulties has been the virus in chickens. Less satisfactory results

87
have been reported by Cooper ( 193 1 ) and ( Zuffa et al., 1956) , or 48 hours previous
Lulic (1955). Moynihan et al. (1954) ly (Nai and Garinei, 1945). Although the
found that 0.5 ml. of antiserum adminis cost of this type of serum has sometimes
tered between 24 hours and 72 hours after been considered prohibitive (Anon..
virus exposure failed to prevent the devel 1943), satisfactory results have followed
opment of Newcastle disease. its use in infected flocks; and mortality has
Immune serum from goats
prepared been considerably reduced (Capaul et al.,
has not been very (Anon.,
satisfactory 1963; Lulic, 1955), or has ceased 6 to 10
1943 ) ; it has delayed death of inoculated days after treatment (Nobili et al., 1960).
chickens for only two to four days Spalatin (1948) and Tanasugarn (1961)
(Placidi et al., 1952). However, a potent have reported that the injection of im
serum has been prepared from goats mune serum has markedly reduced the
(Fomina and Ochkina, 1951) and also anticipated mortality in infected flocks.
from a calf (Coronel, 1939).
Alternative procedures have been to
Hyperimmune serum has been prepared
inject egg yolk (Wills and Luginbuhl,
in fowls and turkeys by injections with
1963) or immune serum and gamma
virulent virus (Majewska and Zebrowski,
globulin simultaneously (Vasington et al.,
1955; Nai and Garinei, 1945; Skoda
and Zuffa, 1956a; Zuffa et al, 1956) and 1960). It has also been shown that hens

has been treated with antibiotics (Nobili passively immunized to Newcastle disease

et al., 1960). Immune serum in amounts virus can transfer a portion of this im
of 0.5 or 1 ml. has protected fowls against munity to their offspring (Grun and

virulent virus administered simultaneously Wogan, 1963).

CONTROL WITH HYPERIMMUNE SERUM

COMBINED WITH VIRUS
This method of immunization has been that the intranasal instillation of immune
generally unsatisfactory (Coronel, 1939; serum simultaneously with attenuated
Iyer, 1943; Seetharaman, 1951b), al- virus resulted in satisfactory immunity,
though Zuffa and Skoda (1959) found

CONTROL WITH ANTIBIOTICS AND OTHER

MEDICINAL AGENTS
The therapeutic effect of antibiotics in hibition of secondary invaders (Reuss,
the controlof Newcastle disease infection 1960; Tanasugarn, 1961).
has been reported. Giovaneli (1962), for
Substances which have shown little or
example, found that symptoms disap
no curative value against the disease in
peared and mortality ceased after injection
clude: xenalamine (Gagliardi and Girotto,
ofa mixture of three antibiotics. Penicillin
1961), sulphapyridine (Anon., 1943),
administered intramuscularly has also
proved of some value (Michalewicz,
vitamin A (Bonaduce, 1950a) and corti

1952). However, the oral administration costeroid (Hababou Sala, 1960). In con
of oxytetracycline has had no influence trast, increased riboflavin supplementation
on the course of experimental Newcastle has reduced mortality 10 and 17 per cent

disease, although it has reduced mortality in groups of artificially infected chickens

in chicks — probably because of the in (Squibb, 1963).

88
 CONTROL BY VACCINATION
In a summary of national control mea ada, 1954; Rao and Agarwal, 1962;
sures in effect during 1961, it was shown Thompson and Osteen, 1952; Vandem
that out of 103 countries reporting the aele, 1961). There are some exceptions:
disease, 85 had adopted vaccination as the in Spain (Blanco, 1949), the United
main control procedure (Table 2). The States (Flowers et al, 1960) and Guate
costsof vaccination are usually borne by mala (Correa, 1963; Correa and Rosales,
the poultry industry. Vaccination of the 1961) vaccines have failed to protect
United States broiler crop in 1956 cost against an indigenous field virus, especi
more than four million dollars (Hanson ally when the field virus has been highly
and Brandly, 1958). In Africa, the cost virulent (Valdes Ornelas, 1964). How
of vaccination has generally been con ever, it is important to differentiate be
sidered out of proportion to the economic tween vaccine breaks and vaccination
value of the stock (Kaschula, 1950; breaks (Jungherr and Markham, 1962).
Vandemaele, 1961) and the demand for The latter involve many factors relating
vaccine has been poor ( Winmill and Haig, to the handling and administration of
1961). vaccine (Davis et al, 1951; Larose and
Compulsory vaccination has sometimes Van Roekel, 1959; Marek; 1957; Tenni-
been (Fiolic, 1957) in spite of
adopted son, 1963).
serious (Fontaine, 1963; Win-
difficulties
mill and Haig, 1961). To overcome the Procedures for Evaluating
difficulties, the cost of the vaccine has
Immunity
been subsidized (Garside, 1962).
When vaccination was first adopted, the
Studies of vaccination for control of
potency of vaccines could be readily
the disease under varying methods of
assessed because the disease frequently
husbandry were begun soon after identifi
caused over 90 per cent mortality. How
cation of Newcastle disease in 1926 and
ever, in certain countries, Newcastle dis
the literature on the subject is now con
ease has since changed to a form in which
siderable 1964a). To review
(Lancaster,
mortality is considerably lower (Fritzsche,
every report on vaccination is beyond the
1963; Jansen and Kunst 1952; Lissot,
scope of this publication; nevertheless, an
1956; Reid, 1961 ; Skoda and Zuffa, 1958),
attempt is made in the following pages to
or the nervous symptoms more numerous
survey various facets of the subject.
(Jaksic and Stefanovic, 1957). In these
countries, reduction of mortality can,
Antigenic Plurality therefore, no longer be used as the sole
Results reported by Gill et al (1959), for the evaluation of vaccines.
criterion
Gualandi (1949), Traub (1944), Upton Moreover, resistance to infection of the
etal. (1953) and Valdes Ornelas (1964) respiratory epithelium, immunity to sys
have led to a questioning of the feasibility temic infection as denoted by clinical
of using any one vaccine in all situations. signs, and resistance to a decrease in egg
However, several strains of vaccine virus production may be independent of each
have been used successfully in widely dif other (Bankowski and Corstvet, 1960,
ferent geographical regions and against 1962b; Markham et al, 1951a, 1957;
different clinical manifestations of the Raggi and Lee, 1962), or show a lack of
disease (FAO-OIE, 1959; Jungherr and correlation (Bankowski et al, 1958b). As
Markham, 1962; Marek, 1957; Mitchell an alternative procedure to determine im
and Walker, 1953; Pagnini, 1950; Ques- munity, an intradermal test has been used

84
(Wasserman and Yates, 1953; Yates et al, antibody and HI antibody do not always
1954). develop or decline at the same rate follow
Probably a still better procedure for ing vaccination (Keeble and Wade, 1963;
determining immunity is to expose vacci Raggi and Lee, 1960; Schmidt, 1959)
nated and non-vaccinated chickens to makes it impossible to regard the immune
susceptible chickens which have been in bodies associated with haemagglutination
fected with a virulent field virus (Bankow- inhibition, virus neutralization and speci
ski and Corstvet, 1962a; Mazzaracchio fic refractivity to infection as a single
and Orfei, 1956; Taylor, 1953). However, entity (Brandly et al, 1947; Dardiri and
in this procedure the environmental tem Yates, 1962; Hanson et al, 1950; Sch-
perature (Francis and Kish, 1955; Sinha mittle, 1953). Thus, Karzon and Bang
et al, 1957), as well as other stress ( 1951 ) showed that the neutralization test
factors (Schultz and Feiling, 1954), can in embryo and the haemagglutination
influence the resulting mortality. Vaccin inhibition test yielded parallel results in
ated birds, when exposed by injection or the measurement of antibody during early
by contact to virulent virus, may show no convalescence. Later in convalescence, the
mortality or sign of infection but may HI titres were lower.
experience a temporary viraemia (Gill et No definite relationship has been found
al, 1959; Hofstad, 1956) or a respiratory between HI and SN titres (Nakamura
infection (Doll et al, 1950c, 1951b; Ban- et al, 1956) and respiratory infection
kowski et al, 1957). (Levine and Fabricant, 1950). However,
The use of the haemagglutination- the SN test has been thought to give a
inhibition (HI) test as a quantitative truer picture of immunity than the HI
measure of immune response has already test (Atanasiu and Gareau, 1951; Hitch
been reviewed on pages 61 to 63. It must ner and Reising, 1954), especially when
be emphasized that the HI response can the observation period extends beyond 50
not be compared directly with the immune weeks (Gill and Stone, 1964), and it
status as measured by challenge with viru could, therefore, be useful to supplement
lent virus (Doll et al, 1950a, b, 1951a; the HI test (Crowther, 1963).
Hamann, 1958; Hitchner and Reising, It follows that the methods and criteria
1953a; Hitchner et al, 1951a; Ileri, 1950, adopted for evaluating the immunity
Levine and Fabricant, 1952; Mark ham engendered by Newcastle disease vaccines
et al, 1954; Mazzaracchio and Orfei, have been variable (Hofstad, 1953b;
1955; Raggi and Lee, 1962; Simmins and Johnson et al, 1954) and this has pre
Baldwin, 1963; Valadao, 1955; Winter- vented a close comparison of all the
field and Seadale, 1957; Winterfield et al, results reported. Bankowski and Corstvet
1957). Schmidt and Schmidt (1955) have ( 1 960) were of the opinion that immunity
reported that 8.5 to 95 per cent of fowls in Newcastle disease consisted of many
with vaccination HI
titres up to 1:16 died measurable as well as unmeasurable
after experimental infection; whereas factors.
fowls with vaccination HI titres of 1:32 In the discussion that follows, more
and over resisted infection. Furthermore, emphasis is placed on the results of ex
Crowther (1963) has suggested that, be posure of vaccinated birds to virulent
cause different strains of Newcastle dis virus and results obtained in the field than
ease virus vary in their ability to stimulate on serological responses. In adopting these
the formation of HI antibody, the HI test criteria, it is recognized that a drop in
should not be used as the sole basis of egg production or the presence of respira
comparison for different vaccine strains. tory symptoms would be a more critical
The fact that serum neutralizing (SN) test of immunity than mere survival
90
 TABLE 13 — Effect of Intranasal Newcastle Disease Vaccination of Chicks from Immune Parents

(Lancaster et al., 1960)

Number of chickens showing the same maximum HI titre1

in
No. Period of 22 days Period of 22 days Period of 23 days
group following 1st vaccination2 following 2nd vaccination3 following 3rd vaccination4

No. No. No.

<
<

1
1
1
1
1
1

:8
:8
:8
:8
:1
:8
:8

>
>

1
1
1
1
6
1
1
1
1
1

:1

tested<1 :16 :32>1 :32 tested :32 :32 tested :16 :32 :62 28 :8

2
-

5
1
1
2
3
3
2
3

102 16 2

13
23

36
2
28
28

12
3 2
222 33 12 25 23

Not vaccinated Not vaccinated

8
1
1
5
8
365 32 11 11 13 -

Expressed as the reciprocal of the highest serum dilution causing complete inhibition of haemagglutination

2
Age at first vaccination days
Age at second vaccination 58 days

1 2 3 4
1
Age at third vaccination 08 days
(Hofstad, 1953b). In this connection, hand, the administration of a virulent
Bankowski (1961a) found that the level strain of virus drinking water has
in
of immunity which protected hens against resulted in satisfactory immunity in chicks
a drop in egg production was related to possessing maternal antibody (Gagliardi
the amount of virus in the vaccine. How and Girotto, 1960). The results given in
ever, during declining immunity, transi Table 13 indicate that satisfactory HI
tory respiratory symptoms and a slight serum titres do not result from the intra
drop in egg production have appeared to nasal vaccination with Bl virus of
be unavoidable when vaccinated birds are parentally-immune chicks at 2 days of age
exposed to virulent field virus (Garside, (Lancaster et al., 1960).
1962; Hitchner and White, 1956). Never In young chicks, the degree of immunity
theless, significantly fewer soft-shelled (Doll et al., 1950b; Richey and Schmittle,
eggs have been produced following the 1962), or of virus multiplication in the
challenge of vaccinated birds than follow oral mucosa (Gagliardi and Irsara, 1958),
ing the challenge of unvaccinated birds has been directly influenced by the amount
(Gill and Stone, 1964). of yolk-transmitted antibody, and the
virulence of the vaccine virus (Gagliardi
and Girotto, 1960). As a result of the use
Factors that Influence of the B 1 vaccine during a five to six year
Development of Immunity period, with resulting passive immunity in
young chicks, the overall duration of im
Passive lmmunity munity has apparently been reduced from
The literature on the transmission of 12 weeks to 6 weeks (Bankowski et al.,

maternal antibodies to the chick has been 1957).

reviewed by Beaudette and Bivins (1953) Usually, the best response in chicks
and Brandly et al (1946b) and it has having maternally immunity
transferred
been discussed by Hanson (1957) and has followed administration of a live
Grun and Wogan (1963). At hatching vaccine by the respiratory or conjunctival
time, chicks from immune dams have routes (Beaudette and Bivins, 1953; Born
shown a much lower serum HI titre than stein et al, 1952; Hitchner, 1950; Hitch
that of the dam. Bornstein et al. (1952) ner and Reising, 1953b; Markham et al.,
found that this titre in the chick rose to a 1951a, b, 1954; White et al, 1953; Zuffa
peak at 3 days of age and progressively and Skoda, 1959). In passively immune
declined thereafter. day-old chicks, intranasal instillation has
Usually, congenital passive immunity resulted in a better immune response than
or the administration of immune serum that obtained by aerosol vaccination ( Rao
has interfered with the development of et al,
1963). Such findings have led to the
active immunity in response to Newcastle conclusion that to delay vaccination be
disease vaccine given intramuscularly cause of maternal immunity is not justi
(Bankowski et al., 1958a; Bornstein et al., fied (Crawley, 1954; Hitchner et al, 1950;
1952; Doll et al., 1950b; Keeble and Markham et al, 1951b; White et al,
Wade, 1963; Keeble et al., 1963; Mark- 1953). However, when an inactivated
ham et al., 1954), subcutaneously (Haig vaccine has been used, better immunity
et al., 1962; Zuffa and Skoda, 1959), has resulted when chicks from immunized
intranasally (Bankowski and Corstvet, hens were first vaccinated at 8 days
1962a; Richey and Schmittle, 1962) or (Keeble et al, 1963) or, preferably, 21
in drinking water (Marek and Raszewska, days of age (Keeble and Wade, 1963).
1959; Winterfield and Seadale, 1956b; The value of passive immunity in pro
Winterfield et al, 1957). On the other tecting young birds has been examined.

92
A number of reports have indicated the of susceptible day-old chicks can result
absence of protective immunity in young in a durable immunity lasting 5 months
chicks from vaccinated flocks (Doll et al, (Asplin, 1952; Lancaster, 1957b) to 12
1951a; Monti, 1954; Olson et al., 1950; months (Hitchner, 1950). It is believed
Reuss and Hilbrich, 1960; Zureck, 1958). that at about 1 to 2 months of age,
Some of these results can be explained by chickens are sufficiently developed to give
a low level of immunity in the parent an optimal response to a vaccine (Ban
flock (Doll et al, 1950b) or by the fact kowski and Rosenwald, 1956; Gupta and
that, in chicks, congenital antibodies give Rao, 1959b; Raggi and Lee, 1962;
little protection to the respiratory tract Schmidt, 1959) or other antigen (Wolfe
against Newcastle disease virus (Levine and Dilks, 1948).
and Fabricant, 1 950) . In contrast, a num When considering the influence of age,
ber of authors have found that chicks it should be mentioned that there is some
from immune hens resist artificial infec evidence that resistance to Newcastle
tion (Mitchell and Walker, 1953; Russeff disease increases progressively as birds
and Miteff, 1957) or develop asympto mature (Baldelli, 1957; Brandly et al,
matic infection (Doll et al, 1951a), de 1946c; Cole and Hutt, 1961; Gill et al,
pending on the virulence of the challenge 1959).
virus.
This resistance due to passive immunity Virus Titre of the Vaccine
lasts for varying periods up to about 4
In addition to the antigenic characteris
weeks after hatching (Anon., 1962a;
tics of the vaccine, the titre and the dose
Brandly etal, 1946b; Christie et al, 1963;
of virus play an important part in the
Clancy etal, 1949; Garinei, 1945; Hitch-
level of immunity produced (Bankowski
ner et al, 1950; Maglione and Dotta,
and Hill, 1954; Brandly et al,- 1946a;
1957; White et al, 1953). However, field
Crawley, 1954; Hitchner and Reising,
studies have shown that in heavily in
1954; Raggi and Lee, 1962, Winterfield
fected areas passive immunity cannot be
and Seadale, 1 957). Thus, under field con
depended for the protection of
upon
chicks under 4 weeks of age (Hitchner
ditions, White-Stevens (1961) considered
that the degree of immunity was directly
etal, 1950; Levine and Fabricant, 1950).
related to the dosage of virus particles or
It must be emphasized that in many of
the amount of antigen (Keeble and Wade,
the reports referred to on pages 95 to 115
it not been 1963). Bankowski and Corstvet (1962b)
has possible to determine
found that a single injection of a vaccine
whether the chicks used were from im
at 10~7 ELD per 0.2 ml. gave nearly the
mune or susceptible parents. This is
same level of immunity as two doses of
particularly true of reports of field results.
vaccine of 10^4,5 ELD administered nine
weeks apart. For a water-administered
Age at Time of Vaccination vaccine, the minimum titre should be
10-7 ELD (Winterfield and Sea-
per ml.
Chicks, even in the absence of passive
dale, 1957). For spray administration of
immunity, do not show the maximum
Bl virus, the titre should be between 10^7
response to antigens (Bankowski et al,
and 10-8 ELD per 50 ml. (Crawley, 1954).
1957; Brandly et al, 1946a; Doll et al,
1950b; Hitchner et al, 1950; Keeble and
Wade, 1963; Mansjoer, 1961; Nakamura
Viral Interference
et al, 1956; Waller and Gardiner, 1952; The different features which comprise
Wasserman and Yates, 1953; Wolfe and the overall phenomenon of viral inter
Dilks, 1948). Nevertheless, vaccination ference and viral infection of the cell have

93
 been discussed in detail
by Goret and in suspended L cells (Cantell et al., 1962).
Provost (1964). When a mesogenic live Newcastle dis
Hanson and Alberts (1959) and Han ease vaccine is administered to birds in
son et al. (1956) showed that when a the early of infection, recovery
stages
virulent field strain of Newcastle disease frequently follows, or further spread of
virus was administered simultaneously with the disease is stopped (Harnach and Polak,
infectious bronchitis virus by the intranasal 1964; Mihalka, 1963). This interference
route, there was interference with the sub or cell block has been associated with the
sequent development of Newcastle disease use of the Mukteswar vaccine (Daubney
infection. Other workers have found that and Mansi, 1948; Generoso and Men-
the interference resulted in poor immunity doza, and has been used in the
1950)
to the infectious bronchitis vaccine control of a number of outbreaks
(Ban- (Karc-
kowski et al, 1955). Raggi et al. (1963) zewski et al., 1955). Interference between
obtained similar results in chick embryos. the Mukteswar vaccine virus and field
An homolagous interference has been virus has been demonstrated to occur
established in chicken cells by the use between 20 and 72 hours after vaccination
of ultraviolet-irradiated Newcastle disease (Gupta and Rao, 1959a; Haddow and
virus. As a result, the cells lost their ability Idnani, 1946; Karczewski et al., 1955;
to adsorb active virus. It has been suggest Nilakantan et al, 1960a; Russeff, 1956).
ed that this type of interference occurred A similar Newcastle disease viral inter
at the cell surface (Baluda, 1957, ference has been observed in tissue cul
1959).
An tures (Durand, 1961).
egg-adaptedstrain of Newcastle
disease virus has also inhibited the growth Protection against virulent virus has
of a strain of the same virus adapted to been demonstrated one to three days after
the brains of new-born mice (Sinkovics, vaccination with the Hertfordshire virus
1957b). (Buzna and Hodosy, 1951; Gualandi,
Morimoto et al. 1952; Schmidt, 1952); after six hours with
(1962) found that the
propagation of Newcastle disease virus in another mesogenic vaccine strain (Jezier-
certain cell ski, 1953) and after two to four days with
cultures was inhibited by
Russian spring-summer encephalitis virus
a formolized vaccine (Brandly et. al,
and by Japanese encephalitis virus. In 1946a).
terference between other heterologous With lentogenic strains of Newcastle
viruses have included: the interference in disease virus, viral interference has not
birds inoculated with a mixture of attenu always been demonstrated. Thus Doll et
ated fowl plague and Newcastle disease al. (1950a), Hitchner and Johnson (1948)
viruses (Daubney and Ishak,
1953); the and Nilakantan et al. (1960a) found
interference between western equine en vaccine strains Bl and F afforded little
cephalomyelitis and Newcastle disease protection when birds were exposed to
viruses when propagated in chick embryo virulent virus soon after vaccination.
fibroblast monolayers (Levine, 1958, Similar results have been observed under
1962b); the interference between mumps field conditions (Lancaster, unpublished
virus and Newcastle disease virus in em- data). In contrast, it has been suggested
bryonated hens' eggs (Sinkovics, 1957a); that the vaccination of young chicks with
the interference between influenza and Strain F virus intranasally (Rao and
Newcastle disease viruses in chick-embryo Agarwal, 1960), or Bl virus intraocularly
cultures (Tyrell, 1955); and the inter (White and Appleton, 1953), results in
ference by ultraviolet-irradiated Newcastle protection due to viral interference 48
disease virus and vesicular stomatitis virus hours later.

94
Differences Between lndividual Birds fourth and eighth day after vaccination.
Thus, vaccination may act as a predis
In any group of vaccinated chickens,
posing factor to diseases of the avian
not every individual develops a satis leucosis et al.,
complex (Consoli 1955).
factory immune response (Asplin, 1952;
Garside, 1962; Lancaster et al., 1960), Other Factors that Affect Immunity
regardless of the kind and quality of the A reduction in the degree and duration
antigen (Bankowski and Corstvet, 1962b; of immunity has been associated with
Markham et al., 1951b). Furthermore, the intercurrent infectious diseases and in
speed of the immune response varies ternal (Blanco, 1949; Brandly,
parasites
between individuals; and approximately 5
1948; Hoekstra, 1961); with poor condi
to 7 per cent of susceptible birds have
tion (Garside, 1962; Marek et al., 1961;
shown an abnormally poor response with
Pomeroy and Brandly, 1953);
(Markham, 1962).
moulting (Schiavo, 1960); and with
Differences between individuals in their
caponization (Gualandi, 1953). The last
immune response to Newcastle disease
mentioned effect was not, however, ob
antigen may be influenced by a deficiency served by Brandly et al. (1946a).
of gamma globulin (Hanson, 1957); by The stress placed on poultry during
the genetic
background of the chicken
transport to and from a market may result
(Millen, 1960); or simply by the varying
in a breakdown of vaccination immunity
ability of different birds to respond to the
(Schultz and Feiling, 1954). Furthermore,
virus antigen (Ram, 1961).
the prolonged feeding of tobacco powder
as a parasiticide appears to inhibit the

Effect of Vaccination on Susceptibility production of Newcastle disease HI anti

bodies following vaccination (Papparella,
to Another Disease
1955). According to Squibb (1963), the
Hanson and Alberts (1959) and Han addition of B complex vitamins above
son et al. (1956) have shown that chickens normal requirements does not appear to
exposed to small quantities of Newcastle influence antibody response, as determined
disease virus may be more susceptible to by HI titres. Similarly, Cho (1963) has
the same virus on re-exposure. This viral reported no evidence that the bursa of
sensitization may play a role in some Fabricius plays a significant role in the
vaccination failures (Hanson, 1957). production of antibodies to Newcastle
Another feature relevent to Newcastle disease virus (Bl strain).
disease vaccinationis the fact that approxi In addition to these factors, a four-fold
mately 25 per cent of a group of vacci increase in Newcastle disease HI antibody
nated fowls have shown an erythroblastosis has been associated with maximum egg
and moderate leucocytosis between the production (Markham et al., 1956a).

TYPES OF VACCINES— ADMINISTRATION

AND EFFECTIVENESS
Live and Inactivated kinds: live and inactivated. The inactivated
vaccines were the first to be studied and
Vaccines Compared the results obtained have been reviewed
by Beaudette (1943, 1948b, 1949a,
Newcastle disease vaccines are of two 1 95 1b ) , Brandly et al. ( 1946b ) , Fabricant

95
(1956) and Thompson (1951). However, double vaccinations is carried out (Fabri
in the early 1930's investigations began cant, 1956). It has been suggested that a
on the preparation and use of live vaccines killed vaccine of high antigenicity would
(Iyer and Dobson, 1940; Topacio, 1934) give better immunity than live vaccines
and this literature has been reviewed by (Levine and Fabricant, 1952). If developed,
Beaudette et al (1950), Brandly (1959), it would probably form an essential
Fritzsche and
Gerriets (1962), Kruger part of any eradication scheme (Osteen
(1961) and Reis and Nobrega (1956). etal, 1961).
It has been found that under some In 1948, a formolized adsorbed vaccine
conditions, and in some geographical was used in Spain (Botija and Loizelier,
areas, live vaccines have certain advan 1948). Although extensive vaccination of
tages over inactivated vaccines and this large flocks was possible, the difficulty of
has resulted in their extensive use (Dall- vaccinating small isolated farm flocks was
ing, 1958; Davis et al, 1951). a major obstacle to eradication. Some 14
The concentrated of live vaccines
use years in 1962, Newcastle disease
later,
has apparently led to the elimination of was reported to be widespread (Figure 5).
Newcastle disease for significant periods Botija and Loizelier (1948) did not expect
of time from Cyprus (Crowther, 1952) the vaccine would eradicate the disease
and parts of Canada (Anon., 1962b). In entirely but they believed that vaccination
other countries it has been thought that in conjunction with sanitary measures
some emergencies fully justified the use should keep it under control.
of live vaccines (Beach, 1946, 1952; In areas of the United States, an in
Osteen et al, 1961; Schoening and activated vaccine has not been used as
Thompson, 1955; Stover, personal com extensively as had been anticipated
munication). Nevertheless, it is well (Beach, 1946). In one field study in the
recognized that live vaccines seldom lead United States, on premises where New
to complete eradication of Newcastle dis castle disease was known to exist, Schoen
ease (Levine, 1962a; Osteen et al., 1961; ing et al. (1949) have reported that the
Schoening and Thompson, 1955). use of formalin-inactivated vaccine did
It has been generally held that in not entirely prevent the disease, but
activated vaccines give a more transient enabled vaccinated birds to withstand a
or less durable immunity than live virus severe infection with relatively small losses
vaccines (Bankowski and Rosenwald, compared with the unvaccinated controls.
1956; Brandly et al, 1946b; Fabricant, The advantages associated with in
1953; Hofstad, 1953b; Osteen et al, 1961; activated vaccines have been summarized
Pomeroy and Brandly, 1953; Thompson, by Beach (1946) and Garside (1962),
1951). Beaudette (1951a) has reported and the disadvantages of live vaccines
that only 60 to 85 per cent of young to have been outlined by Hemsley (1962),
mature chickens develop immunity when Hoekstra (1961) and Osteen et al. (1961).
vaccinated with an inactivated vaccine. Comparative tests involving either lento-
However, inactivated vaccines are now genic or mesogenic live vaccines and
available which, within certain limits, inactivated vaccines have given results in
induce satisfactory immunity (Appleton favour of the live vaccines (Kaschula,
et al, 1963; Levine, 1962a; Woernle, 1950; Levine and Fabricant, 1952; van
1955), especially when prepared from a Waveren and Zuijdam, 1953; Zuydam,
local strain of virus (Woernle and Sieg- 1953). Other studies have shown little
mann, 1954) or one found antigenically difference between live and inactivated
suitable (Hanson et al, 1951b; Koch, vaccines (Miyamoto and Nagashima,
1955) and when a proper regimen of 1957); however Schmidt (1959) found
96
the immune response to an inactivated instillation of the vaccine resulted in

a
adsorbed vaccine to be greater than that more durable response when compared
produced by the Bl strain. with the aerosol method. They emphasized
the need to use special nebulizers with
reconstituted freeze-dried vaccine.
Live Vaccines
The viability of Bl and viruses in

F
In this section live vaccines are con drinking water adversely affected by

is

a
sidered under two main headings: lento- variety of conditions and has been en
genic strains and mesogenic strains. This hanced by the use of organic stabilizing
materials (Marek, 1960; Winterfield and
is,

perhaps, rather an arbitrary division

but based principally on the fact that Seadale, 1956a). The mechanism of the
is
it

lentogenic strains can be administered to immune response following vaccination by

young chicks in which the mesogenic the drinking water method has been dis
strains generally cause severe reaction. cussed by Baldelli (1956), Burnstein and
a

There are also number of other differ Bang (1958), Gagliardi and Irsara (1958),
a

ences between these two main types and by Winterfield et al. (1957).
(Anon., 1959; Hanson and Brandly, In intranasal or inhalation vaccination
1955). The lentogenic strains take longer with Bl virus, the virus may or may not
to kill chick embryos and also appear be confined to the turbinate region where
unable to multiply proliferates (Burnstein and Bang, 1958).
in

the central nervous

it

system of the chicken; whereas the meso However, the initial multiplication of an
genic strains readily multiply in the brain attenuated strain which occurred on the
of chickens of any age introduced by mucosa of the portal of entry has spread
if

the intracerebral route. The intracerebral to the mucosae of other parts of the head
inoculation of virus into day-old chickens (Gagliardi, 1959).
To relate the optimum immune
is,

therefore, sensitive method of dis res

a

tinguishing between these two types of ponse with methods of administering live
Newcastle disease virus (Hanson, 1956). vaccines, comparative studies have been
made of the following routes and methods:
intraocular, intranasal, intratracheal, intra
Lentogenic Strains
muscular, intravenous, wing-web, sub
The various mechanical factors and cutaneous, drinking water, spray and dust.
principles involved in the preparation of The most effective was considered to be
viral suspensions suitable for air-borne the intraocular by Kaschula (1952) and
methods of vaccination have been dis White and Appleton (1953); the intra
cussed by Bankowski and Hill (1954), nasal by Glinski and Szemberowa(1960),
Crawley (1954), Hitchner and Reising Hitchner and Johnson (1948), Johnson
(1953a), and Markham et al. (1955a). (1956), Marek et al. (1961), Markham
Studies using the Bl virus labelled with et al. (1954), Rao and Agarwal (1960);
radio-active phosphorus and administered the intramuscular by Bran et al. (1959,
as spray have shown that virus small 1961); by Gualandi
in

the intravenous
a

amounts can be detected in the abdominal (1949); drinking water by Johnson (1956);
air sacs 30 minutes after vaccination. Wynohradnyk et al. (1958); spray by
It

has been concluded that comparatively Johnson (1956) and Marek et al. (1961);
few virus particles need to reach the and dust by Dardiri et al. (1957) and
respiratory epithelium in order to stimu Johnson (1956). The protection afforded
late measurable response (Johnson et al., by mixing live vaccine virus with the
a

954b. Rao et al. 963 in studies with food was considered slight by Gagliardi
1

,
(
)

Strain virus, showed that intranasal

F

(1953).
97
B1 or Blacksburg Strain 1954; Bran et al, 1959; Hitchner and
Johnson, 1948; Hitchner, 1950; Hitchner
The main characteristics of this strain et al., 1950) to 15 per cent for one or two
and the indications for its use were first weeks (Hoekstra, 1959, 1961; van
reported by Hitchner and Johnson (1948) Waveren, 1955) and 20 to 50 per cent
of Blacksburg, Virginia, in the United for two to four weeks (Doll et al., 1950a).
States. The initial studies showed that all Duration of immunity. Determination
ages from day-old chicks to birds in full of the degree and duration of the im
production could be vaccinated. Within a munity engendered by the Bl virus has
relatively short time this vaccine strain presented difficulties, as already outlined
was being used extensively in the United on page 89. This has hindered the estab
States (Schoening and Thompson, 1955) lishment of uniform vaccination proce
and also in other parts of the world dures and programmes for different
(Anon., 1962a; Surin, 1959). methods of husbandry and degrees of
Clinical effects of vaccination. In young exposure.
chicks, clinical effects produced by the Bl A number of reports on the duration of
strain have depended largely on the route immunity after initial vaccination of
or method of vaccination. Thus, adminis young chicks are summarized in Table 14.
tration by intranasal or conjunctival drop The data in this table are confined to
or in drinking water has usually had little observations based on challenge exposure
or no clinical effect (Hitchner and John made two or more months after vacci
son, 1948; Luginbuhl et al., 1955; Lulic nation.
and Spalatin, 1956; Hutson, 1953; Mark- Results of tests conducted approxi
ham et al., 1951; Miyamoto and Naga- mately one month vaccination of
after
shima, 1957; Raggi and Lee, 1962; Russeff young chicks have been reported by Doll
and Miteff, 1957; Winterfield et al., 1957), et al. (1950a, b, c) , Hitchner and Johnson
but some instances of more marked clini (1948), Hitchner and Reising (1953a),
cal effect have been reported (Doll et al., Hoekstra (1961), Johnson and Gross
1950a). Administration as a dust or (1951, 1952), Jungherr and Markham
aerosol has produced either no adverse (1962), Markham et al. (1950), Miya
reaction (Crawley and Fahey, 1 954; John moto and Nagashima (1957), Pette
son and Gross, 1951; Markham et al., (1961), Rao (1955), Richey and Sch-
1955a); a slight respiratory reaction last mittle (1962), White et al. (1954) and
ing a few (Hitchner and Reising,
days Winterfield et al. (1957).
1952, 1953b; Price et al., 1955; White Relatively fewer reports describe the
et al., 1954) ; or a severe respiratory reac immunity resulting from vaccination at
tion with appreciable mortality (Bankow- 8 weeks of age and older. Vaccination
ski and Hill, 1954), especially if the of such chickens has resulted in satis
vaccinated chicks were exposed to E. coli factory immunity for 12 weeks (Bosgra
(Gross, 1961a) or Mycoplasma (Bankow- and Roerink, 1961), 20 weeks (van
ski, 1961b). The vaccination of young Waveren, 1955), 26 weeks (Bran et al.,
chicks in incubators or in chick boxes has 1959; Miyamoto and Nagashima, 1957),
given variable results (Johnson and Gross, 32 weeks (Raggi and Lee, 1960) and 56
1952), and the spray method has been weeks (Bankowski et al., 1957; Raggi and
considered unsuitable for baby chicks Lee, 1962).
(Hitchner and Reising, 1953b). The problem of revaccination in the
In susceptible pullets, vaccination with presence or absence of residual immunity
B 1 virus has caused a drop in egg produc may be summarized as follows. The intra
tion ranging from negligible (Bankier, nasal B 1 vaccination of passively immune

98
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99
chicks less than 1 week of age results Strain F
in appreciable neutralization of the vaccine The suitability of this lentogenic strain
virus (Lancaster et al, 1960; Brandly, as a vaccine virus was first reported by
1955) . This neutralization is lessened by Asplin (1952) of Weybridge, England.
administering the vaccine via the respira Reports on its use in Europe, Asia, Africa
tory route which is the natural route of and America have also been reviewed
infection (Burnstein and Bang, 1958; (Lancaster, 1962b). Strain F closely
Hitchner et al,
1950). Nevertheless, such resembles the Bl strain in many of its
vaccinated chicks are generally susceptible properties (Anon., 1959). It multiplies
when 1 month of age. Revaccination at slowly in chickens of all ages
(Quesada
this time has given a good anamnestic et al., 1960) and is well suited to their
response and a satisfactory immunity vaccination.
against virulent virus. By way of contrast, Clinical effects of vaccination. In young
the similar vaccination of young chicks chickens and laying hens, Strain F virus
carrying no maternal antibody has re has sometimes produced mild respiratory
sulted in a satisfactory immune response. symptoms (Asplin, 1952; Asplin et al,
Revaccination four or more weeks later, 1949). However, egg production has not
in the presence of antibody, has resulted been seriously affected (Asplin, 1952;
in a poor anamnestic response (Lan Binaghi and Nardelli, 1955; Lancaster,
caster et al, 1960), and such chicks have 1957a; Mitev and Gagov, 1960b). No
been found susceptible to virulent virus adverse effect on chickens of any age was
two months later. The immune status of reported by Ageeva (1962), Anon.
et al.
commercial chicks is seldom known with (1960), Binaghi Nardelli (1955),
and
certainty and this increases the difficulty Borzemska (1962), Mazzaracchio and
of establishing a vaccination programme Orfei (1956), Petek and Gagliardi (1954),
suitable for all situations. For this reason, or Rao and Agarwal ( 1960) , although a
the age at which primary and subsequent number of different routes of vaccination
vaccinations are conducted must be care were used. Contrary findings were report
fully evaluated for each particular set of ed by Russeff and Miteff (1956) who
circumstances. observed a slight transient paralysis of the
When the Bl strain has been used for feet, and by Thorne and MacLeod (1960)
both the initial vaccination and revaccin- who found that their strain of this virus
ations, considerable differences in pro caused nervous and intestinal symptoms in
cedure have given adequate and satis groups of 4-month-old cockerels. The
factory immunity. Thus, chicks initially findings of the last-named authors do not
vaccinated at 1 to 7 days of age have correspond with the evidence at present
been revaccinated at 4 weeks (Hitchner available on the lentogenic characters of
and Reising, 1952; Johnson, 1956) or 19 Strain F virus.
weeks (Hitchner et al., 1951b; Johnson, Duration of immunity. A number of
1956) . Chicks initially vaccinated at about reports of immunity, as
on the duration
2 weeks of age have been revaccinated determined by exposure to virulent virus,
at 4 weeks (Dardiri and Yates, 1962), are summarized in Table 15. Other re
8 weeks (Meyn and Pette, 1961), 12 ports describing immunity tests with
weeks (Hoekstra, 1961) and 20 weeks Strain F virus have been written by
(Hitchner and Reising, 1952). When the Ageeva et al. (1962), Baldelli (1956), Bor
initial vaccination has been delayed until zemska (1962), Gagliardi and Irsara
8 weeks, revaccination has been conducted (1958), Marek (1960), Marek et al. (1961),
at 27 weeks of age (Bankowski et al., Mitev and Gagov (1960b), Petek and
1957) . Gagliardi (1954), Rao and Agarwal

100
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(I960), Winterfield et al (1957) and least four months has been reported (Ober-
Wynohradnyk et al. (1958). feld, 1962).
Studies on revaccination with Strain When administered as an aerosol, the
F virus have been of two main kinds. A LaSota strain has been found effective
procedure which has given satisfactory (Borzemska, 1962).
results has been to revaccinate with Strain
F virus every one to three months (Mitev Other Lentogenic Strains
and Gagov, 1960b; Petek and Gagliardi,
Other lentogenic vaccine strains de
1954). An alternative method has been scribed by (1961) Vega and
Quiroz
to administer Strain F virus intranasally
Pagnini (1956) have proved satisfactory
to day-old chicks and revaccinate with the have
in rather local situations. Reports
mesogenic Mukteswar vaccine virus at immunity and
also indicated that poor
about 6 weeks of age. This method has followed the
post vaccination mortality
proved practical in India (Rao and Agar- of a live
intraocular administration
wal, 1962), Malaya and Hong Kong to day-old chicks
attenuated vaccine
(FAO-OIE, 1959). (Gharib, etal, 1961).

LaSota Strain
Mesogenic Strains
The LaSota strain of lentogenic virus
Komarov (or Haifa) Strain
differs in some respects from the Bl and
F strains. One difference is a lower mean This strain was isolated in Palestine
death time for chick embryos (Anon., from an outbreak of Newcastle disease
1959). Other differences, as compared associated with 80 per cent mortality.
with Bl and F, include a greater spread Serial intracerebral passage through
ing potential for LaSota (Marek, 1960) ducklings (Komaroff and Goldsmidt,
and often more post vaccination respira 1946a) modified the virus to the extent
tory symptoms (Winterfield et al, 1957). that adult fowls showed no symptom after
When administered in drinking water, the injection and remained healthy when ex
serological response has been higher with posed to virulent virus 18 days later
LaSota than with Bl and F strains. As a (Komarov and Goldsmit, 1946b).
result, it has been suggested that the Clinical effects of vaccination. In grow
LaSota strain be used at 18 weeks of age ing chickens over 4 weeks of age, the
to revaccinate chickens given Bl or F Komarov strain has been well tolerated
strains at an earlier age (Winterfield et al., (Crowther, 1952; Komarov and Gold
1957). smit, 1947), although, in young birds in
When the immunity engendered by poor condition, a few cases of paralysis
water-administered LaSota vaccine has have resulted (Thorne and MacLeod,
been challenged with virulent virus, only 1960).
28 per cent of the chickens have been Vaccination of laying hens has caused
immune 11 weeks after vaccination. a fall in egg production (Golem and
However, two doses of the vaccine at 5 Berke, 1950; Ileri, 1950; Thorne and
and 28 days of age have resulted in 94 per MacLeod, 1960) which has lasted four
cent protection against virulent virus at 15 weeks (Crowther, 1952).
weeks of age (Dardiri et al., 1957). Im Duration of immunity. The Komarov
munity has lasted 42 weeks after the sec strain has been administered intraocular-
ond dose of water-administered vaccine at ly (Castro Amaro, 1964), subcutaneously
8 and 21 weeks of age (Stumpel, 1962). and intramuscularly (Komarov and Gold
Under field conditions, an immunity of at smit, 1947) with similar results. The

102
wing-web stick method has also been ly used in Asia (FAO-OIE, 1959; Memo.,
widely used (Madhusudan, 1957; Van- 1955; Seetharaman, 1951a, b), in parts of
demaele, 1961). Illustrations of the wing- Northern Croatia (Lukacevic, 1955), in
web vaccination method have been given Bosnia (Marusic, 1955) and in Burma
by Bankowski and Rosenwald ( 1956) and (Peatt, 1945).
Martini and Kurjana (1950); and the Clinical effects of vaccination. In young
method has been described by Schoening chicks, the R2B virus has produced a
and Thompson (1955). severe reaction; and (Gupta
mortality
Vaccination of growing or mature birds and Rao, 1959b; Haddow and Idnani,
has resulted in an immunity against 1946) may reach 30 per cent (Rao and
virulent virus
lasting 7 weeks (van Agarwal, 1960). It has caused paralysis
Waveren, 1955), 3 months (Komarov and (Daubney and Mansi, 1948) in about 2
Goldsmit, 1947; van Waveren and Zuij- per cent of birds (Memo., 1955).
dam, 1953), 8 months (da Camara and In chickens over 6 weeks of age, the
Valadao, 1961), 8 to 12 months (Ileri, Mukteswar strain has been well tolerated
1956), 9 months (Komarov et al., 1948a) (Dhanda et al., 1958; Haddow and Idnani,
and 13 months (Karrar and Mustafa, 1946), although mortality varying from
1964; Thorne and MacLeod, 1960). 1.3 per cent (Generoso and Mendoza,
Serial passage of the Komarov strain 1950) to 6 per cent (Haddow and Idnani,
in bovine kidney tissue culture has at 1946) and 16 per cent (van Waveren and
tenuated the neuropathogenicity to the Zuijdam, 1953) has occurred. Nervous
level of the lentogenic strains of New symptoms have also followed vaccination
castle disease virus, without loss of anti (Jaksic and Stefanovic, 1957).
genicity (Huygelen and Peetermans, As with other mesogenic viruses, the
1963). Mukteswar strain causes a marked reduc
In comparative tests based on patho tion in egg production lasting one to three
genicity and antigenicity, the Komarov weeks (Haddow and Idnani, 1946; Ileri,
strain has been preferred to the Mukte- 1956), or as much as six weeks (Memo.,
swar (Komarov and Goldsmit,
strain 1955). During this period, egg production
1947), a formolized vaccine (da Camara may decrease 10 to 16 per cent (Agcanas
and Valadao, 1961), Strain F (Thorne and Rigor, 1 95 1 ) or even 60 per cent
and MacLeod, 1960) and the Roakin (Dixit, 1950).
strain (Ileri, 1950). One of the Mukteswar stocks of vaccine
virus has shown increased pathogenicity to
White Leghorn chickens causing 56 per
Mukteswar Strain
cent mortality, as compared with 4 per
Reviews of the early work on the devel cent mortality in Rhode Island Reds. This
opment of a live vaccine in India have selective pathogenicity has been attributed
been given by Haddow and Idnani ( 1 946) , to repeated passage through White Leg
Iyer (1943), Iyer and Hashmi (1945) horn embryos (Nandi, 1955).
and Seetharaman (1951b). It was found Duration of immunity. Using the sub
that after repeated passage in chick em cutaneous route, Haddow and Idnani
bryos, an attenuated strain, designated (1946) found immunity against virulent
"R..B", was obtained (Haddow and virus to be durable for 9 to 15 months.
Idnani, 1946). From these studies the Similar results have been reported by
vaccine virus, now usually designated the Cakalowa et al. (1955) and Daubney and
Mukteswar strain, was developed (Dhanda Mansi (1948). Immunity lasting three to
et al., 1958; Gupta and Rao, 1959b; four years was noted by Nilakantan et al.
Memo., 1955). This strain has been wide (1960b) and Seetharaman (1951a). In

103
contrast, Bornstein et al. (1949) were of that the number of doses of mesogenic
the opinion that the immunity produced wing-web vaccines used in the United
by the Mukteswar strain was durable for States has declined appreciably since 1953
no more than one year. Significant HI (Anon., 1962b). Nevertheless, mesogenic
titres have been present in 11 to 64 per vaccine strains of American origin have
cent of fowls vaccinated 6 to 12 months been used in a number of countries in
previously (Lukacevic, 1955). Africa (Kaschula, 1950; Vandemaele,
Using the wing-web stick and intramus 1961) and Europe (van Waveren, 1955).
cular methods of vaccination Dhanda et Clinical results of vaccination. In birds
al. (1958), Generoso and Mendoza under 6 weeks of age the Roakin and
(1950) and van Waveren and Zuijdam MK107 vaccines can cause severe mor
(1953) found that immunity lasted 6 to bidity and some mortality (Anon., 1962b;
8 months but declined at 12 months
(van Cole and Hutt, 1961; Van Roekel, 1956;
Waveren, 1955). A similar 8 months' van Waveren, 1955). Thus, 50 per cent of
duration has resulted from a number of week-old chicks have died after vaccina
different vaccination routes (Nilakantan tion (Clancy et al., 1949). A lower but
et al, 1960a), including administration significant vaccination mortality has been
in drinking water (Forsek et al, 1957). related to the genetic characteristics of two
Nilakantan et al. (1960a) have concluded strains of White Leghorns (Cole and Hutt,
that there is little difference antigenically 196 1 ) . Evidence of a similar strain resis
between the Mukteswar and Komarov tance to Newcastle disease virus (Roakin
strains. strain) has been reported by Francis et al.
Using electrophoresis of serum samples, (1964). However, Beaudette et al. (1949)
Lukacevic et al.
(1958) showed that vac and Cordier-Boullangier et al. (1955b)
cination with the Mukteswar virus resulted found that, with the Roakin virus, mortal
in a significant increase in gamma globulin ity was negligible in chickens vaccinated at
which reached a similar level to that at 5 weeks of age. In 4-week-old chicks, the
tained in experimentally infected fowls. Roakin virus has resulted in a significant
decrease in weight gain following vaccina
tion (Francis et al., 1964).
Strains originating in the United States With the MK107 virus, mortality has
also been negligible in 4-week-old chicks
Two mesogenic strains of the virus have (Markham et al., 1949). A differential
been widely used in the United States as characteristic between the Roakin and
live wing-web administered vaccines. One MK107 viruses has been reported by
strain (Roakin) was identified during the Rosenwald (1959) who found that
et al.
screening of 105 strains (Beaudette et al., whereas the Roakin strain consistently
1949). The other strain (MK107) was failed to spread to contact birds, the
subjected to serial passage in chick and MK107 just as consistently spread from
duck embryos following its isolation inoculated birds to contact birds in other
(Clancy et al., 1949; Markham et al., cages in the same unit.
1949). A number of reports have shown that
Reports and reviews relating to the de wing-web are well tolerated by
vaccines
velopment of wing-web vaccines in the birds 4 to 16 weeks of age (Anon., 1962a;
United States include those by Beach Quinn and Thompson, 1952; Thompson
(1949c, 1951), Beaudette (1948b), Van and Osteen, 1952; Van Roekel, 1956),
Roekel et al. (1948) and Van Roekel even when chickens on infected farms are
(1956). vaccinated (Beach, 1949c; Beaudette et
It would appear from published figures al, 1949; Markham et al, 1949). How

104
ever, in some vaccinated flocks, a slight and reviewed by Iyer (1943) and Kran-
systemic reaction, respiratory symptoms eveld (1950). A strain of virus originally
or, more rarely, paresis may be observed isolated from an outbreak of Newcastle
(Beaudette et al, 1949; Van Roekel, disease in Hertfordshire, England, was
1956). These reactions have been most used. After 14 to 33 passages in the chick
severe in winter
(Davis et al,
1950). embryo, the virus did not kill adult fowls;
In susceptible hens, vaccination with and vaccinated chickens survived exposure
wing-web vaccine viruses has reduced egg to virulent virus (Iyer and Dobson,1940).
production and egg quality to almost the Other studies with the Hertfordshire vac
same extent as natural outbreaks of New cine virus have also been made.
castle disease (Beach, 1949c; Beaudette Clinical effects of vaccination. In chicks
et al, 1949; Kaschula, 1950; Van Roekel possessing passive immunity, the Hertford
et al., 1948; Van Roekel, 1956; van shire strain has caused no mortality
Waveren, 1955). (Szakmary and Beke, 1955). However,
Duration of immunity. Wing-web vac chicks under 8 weeks of age have been
cines have produced an adequate and per very susceptible (Iyer and Dobson, 1940;
sistent immunity (Cordier-Boullangier et Schneider, 1954). The vaccine has proved
al, 1955b;Richter, 1953; Van Roekel satisfactory for chickens over 12 weeks
et al., 1948; Van Roekel, 1956; Thompson old (Schneider,1954); although some
and Osteen, 1952), sufficient to protect cases of paralysis have followed vaccina
against a drop in egg production for one tion (Mazzaracchio and Orfei, 1955;
laying season (Bankowski and Rosenwald, Salyi and Hodosy, 1952), and there has
1956). The immunity engendered by the been a severe drop in egg production
Roakin strain has been durable for 12 (Pagnini, 1954). Clinical effects have been
months (van Waveren, 1955; van Waveren less severe when an adsorbed but living

and Zuijdam, 1953). The MK107 strain Hertfordshire vaccine has been used
has resulted in immunity against virulent (Schmidt, 1952).
virus lasting 10 weeks (Markham et al., Duration of immunity. The intramuscu
1954) to 4 months (Clancy et al, 1949). lar or subcutaneous vaccination of
A feature of many wing-web mesogenic chickens over 3 weeks of age and of adults
vaccines has been the ability of the vaccine has resulted in a durable immunity lasting
virus to spread to adjacent flocks (Beach three to five months (Gualandi, 1951;
1949c). Kaschula (1950) was of the opin Mazzaracchio and Orfei, 1954; Szakmary
ion that there was no danger of the Roakin and Beke 1955). Thereafter, during the
strain becoming virulent. Although the succeeding nine months, the immunity has
mesogenic vaccine viruses are usually ad progressively declined (Mazzaracchio and
ministered either by the wing-web or intra Orfei, 1955), although Teklinska (1951b)
muscular routes, Kaschula (1952b) found found vaccinated fowls were protected for
that, for revaccination with the Roakin one year. Some chickens vaccinated with
virus, intraocular instillation was the best the Hertfordshire virus have been immune
method. to experimental infection although HI
titres were considered negative (Hamann,
1958).
Hertfordshire (or Herts)
Gualandi (1950) concluded that chick
Strain
with live Hertfordshire vac
ens vaccinated
Initial studies on the attenuation of a cine did not excrete the virus and were
virulent strain of Newcastle disease virus not a source of infection, a finding that
for use as a live immunizing agent were contrasts with other results (Lancaster,
conducted by Iyer and Dobson (1940) 1963a).

105
 In some areas of Hungary, a strictly authors considered that virus administered
controlled nation-wide compulsory vac by the vent was less likely to be neutralized
cination with the Hertfordshire strain of by circulating antibodies from the initial
virus, together with veterinary and police vaccination.
control to ensure 100 per cent vaccination Further reference to mesogenic strains
coverage, resulted in a marked decrease is made under the heading "Viral Inter
in the incidence of Newcastle disease ference" on page 93.
(Zsigmond and Gyorgy, 1957).

Extract of Tobacco
Other Mesogenic Strains Mosaic Virus
Other mesogenic strains of Newcastle A small-scale experiment by Marxer
disease virus have been developed and et al. ( 1958) indicated that injection of an
examined as immunizing agents. Some extract of tobacco mosaic virus into chick
have been attenuated by passage through ens engendered immunity to Newcastle
mammals (Reagan et al, 1947b; 1948c, disease. Subsequent by San
investigation
d, 1952c) . Others have been attenuated by ger et al (1961) failed to confirm these
passage through duck and chick embryos results.
(Papparella, 1956; Prier, 1951; Collier
and Dinger, 1950); young ducklings
(Pillai, 1949); doves
Tissue Culture Vaccines
striped ground
{Geopelia striata) (Mansjoer, 1961; Mar The use of tissue-culture-propagated
tini and Kurjana, 1950) or tissue cultures vaccine viruses is discussed on page 130.
(Rusev, 1960; Russeff, 1960). Moreover,
some mesogenic strains have been found
suitable for use in Indonesia
Inactivated Vaccines
(Donker-
Voet and Kurjana, 1950; Mansjoer, This section summarizes the work that
1961) and in certain countries of Africa has been done up to now on the inactiva-
(Jezierski, 1953) and of South America tion of viruses by different substances and
(Nobrega, 1955). On the other hand, procedures and the results obtained.
serial passage in chicken embryos has in
creased the virulence of a vaccine strain
Inactivation by
of virus to the point where its use in the
field could not be recommended
Beta-Propiolactone
(Geh-
ring, 1958). The satisfactory inactivation of viruses
Mesogenic or weakly pathogenic vac by beta-propiolactone (BPL) was first re
cine viruses have been used successfully ported by LoGrippo and Hartman (1955),
for revaccination of chickens initially vac and this method was also used by Mack
cinated with an inactivated virus (Adler and Chotisen (1955, 1956) to prepare an
et al, 1951; Brandly et al, 1946c; Dardiri inactivated Newcastle disease vaccine. A
et al, 1961; Kaschula, 1952); and with general review of virus inactivation by
a lentogenic virus (FAO-OIE, 1959; Rao BPL has been given by LoGrippo (1960).
and Agarwal, 1962; van Waveren, 1955). He showed that, with rabies virus, the use
Winterfield and Hitchner (1961) of BPL and ultraviolet irradiation in com
showed that revaccination by the vent bination resulted in increased viricidal ac
route with the live weakly pathogenic tion, with the antigenic component being
strain "N-47" gave a better serological better preserved than when BPL was used
response than did the Roakin strain by the alone. The inactivation of Newcastle dis
wing-web or intramuscular routes. These ease virus at 37°C has been attained by

106
TABLE 16 — Duration of Immunity following Initial Vaccination of
Chickens with Beta-propiolactone Inactivated Vaccine
(Lancaster, 1964a)

Author Concentration Age when Duration of immunity

of B.P.L. vaccinated against virulent virus

Mack and Chotisen, 1 955, 1 956 0.025% adult at least 1 6 days

Winmill and Weddell, 1961 0.025% 4 weeks at least 3 months
Simmins and Baldwin, 1963 0.025% 2 weeks at least 8 months
Haig et al., 1962 0.033% 8 weeks at least 8 months
Sullivan et al., 1 958 0.1% 2 weeks approximately 2!4 months
Gill era/., 1959 0.1% 2 weeks approximately 3 months
Keeble and Wade, 1963 0.1% 3 weeks at least 13 days
Keeble and Wade, 1 963 0.1% 1 0-1 9 days at least 10 weeks
Keeble and Coid, 1 962 0.1% 2 days at least 1 2 weeks
•
Piercy et al., 1 962 3 weeks at least 1 8% weeks
Appleton et al., 1 963 0.2% 2 weeks poor
•
Christie ef al., 1 963 11-25 days at least 6 weeks
m
Akat, 1 962 1-14 days from 2 to 6 months
m
Cooper, 1 963 adult at least 1 3 weeks

*
Details not available to writer of this Review

(a) a concentration of BPL of 1/1,000 to Clinical effects of vaccination. Day-old

1/2,000 for 30 minutes; (b) a concentra chicks without maternal antibody have
tion of 1/5,000 for 60 minutes; or (c) a been vaccinated (Haig et al., 1962) with
concentration of 1/7,000 for more than out subsequent retardation of growth
two hours (Cherby and Valette, 1964). (Keeble and Coid, 1962).
In field trials, a virus inactivated by The vaccination of laying flocks with a
BPL has revealed a potency similar to that commercial BPL-inactivated vaccine has
of a live vaccine (Gill and Stone, 1960). had no significant effect on egg produc
The relative value of formalin and beta- tion; nor any respiratory symptoms
propiolactone as inactivating agents for (Cooper, 1963).
Newcastle disease virus has been dis Duration of Immunity. Results of a
cussed by Piercy et al. (1963) and by number of studies are summarized in
Appleton et al. (1964). The former Table 16.
authors believed that prolonged exposure It was shown by Simmins and Baldwin
of virus to BPL at low temperatures might (1963) and Piercy et al. (1962) that Strain
adversely affect the antigenicity of the F virus (described under the heading
vaccine. Appleton et al. (1964) felt that "Lentogenic Strains") produced as good a
more study with BPL was needed because BPL-inactivated vaccine as the more viru
birds vaccinated with BPL-inactivated lent Herts strain. Thus, should inactivation
virus had shown a poor immunity against of the virus be incomplete, there would be
challenge. The addition of adjuvants to no ill effect among vaccinated birds if
BPL-inactivated Newcastle disease virus Strain F virus were used. The presence of
has been examined by Sullivan et al. active virus residue in a BPL vaccine was
(1958) and Gill (1959).
et al. These considered Appleton et al.
possible by
studies have been extended by several (1963). Pini et al. (1963) found in
other workers, as shown in Table 16, and activated virulent virus to be more im
most recently by Hofstad et al. (1963). munogenic than Strain F virus.

107
 IMMUNE RESPONSE FOLLOWING VACCINATION WITH A
BPL VACCINE
1280

I I I I I I I 1
2 4 6 8 10 12 14 16

Interval after vaccination (weeks)

(Redrawn from Keeble and Wade, 1963)

Figure 18.

Of the results summarized in Table 16 hours; whereas Haig et al. (1962), Mack
the generally poor immune response re and Chotisen (1955), Keeble and Wade
ported by Appleton et al. (1963) should (1963), Simmins and Baldwin (1963),
be noted particularly. Under the condi Winmill and Weddell (1961) used 37°C
tions of their experiment, these authors for two hours; and Gill et al. (1959) and
found that inactivation with formalin Sullivan et al. (1958) used 21°C for two
resulted in a better vaccine than did BPL. hours.
Appleton et al. (1964) have discussed the Vaccination programmes utilizing BPL-
variables involved in the preparation and inactivated vaccines. In some studies (Sul
evaluation of inactive Newcastle disease livan et al., 1958), initial vaccination was
vaccines and have suggested that the bal at 2 weeks of age; revaccination was at
ance between the concentration of BPL 6 weeks of age. More recently initial vac
and amnio-allantoic fluid needs further cination at 2 to 3 weeks of age has been
investigation. Appleton et al. (1963) used followed by revaccination at 12 to 20
an inactivating temperature of 4°C for 48 weeks of age. Such procedures have given

108
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109
a marked secondary response, as shown in To increase the immune response, dif
Figure 18 (Keeble and Wade, 1963), and ferent adjuvants and adsorbents have been
a duration of immunity after the second tested (Brandly et al, 1946b; Gill et al,
vaccination of 17 weeks (Simmins and 1959; Traub, 1943b). The adsorption of
Baldwin, 1963), 20 weeks (Gill et al, a formalin-inactivated virus with alumin
1959) and approximately 52 weeks (Gar- ium hydroxide gel has been shown to
side, 1962). increase the immunizing properties of the
In England and Wales, a modification vaccine (Nakamura et al, 1956).
of the above schemes has been to recom Considerable differences in the immune
mend three vaccinations at intervals before response associated with different adju
the pullets come into lay (Anon., 1963a; vants have been noted
(Generoso and
Smith, 1963), and another to conduct the Agustin, 1947; Jacotot and Vallee, 1959;
initial vaccination when the chicks are 10 Skoda and Zuffa, 1957). The adjuvant
days of age (Hemsley, 1963). However, probably has created favourable condi
Keeble et al. (1963) considered it best, in tions for the steady antigenic response
the case of chicks from well-immunized (Rao, 1955). In the absence of an adju
parent stock, to delay the initial vaccin vant, formolized vaccine has been more
ation until the chicks are about 3 weeks effective when administered intravenously
of age. (Brandly et al, 1946b; Madison, 1947;
Figures published by Simpson (1963) Nakamura et al,
1956). A formolized
show the effect of Newcastle disease on vaccine, from embryonic fluids
prepared
egg production in unvaccinated flocks and only, has been found to be better than one
in flocks vaccinated with a BPL vaccine prepared from fluids, membranes and
(Table 17). embryos (Perez-Rebelo, 1 962). The use of
a minimum concentration of formalin is
Inactivation by Formalin also a factor for consideration (Appleton
Formalin has been used extensively for et al, 1963).
the inactivation of Newcastle disease virus The evaluation of immunity produced
and the overall results obtained have been by formalin-inactivated vaccines has been
reviewed by Hofstad et al (1963). Viru complicated by the use of different routes
lent strains of virus have generally been of inoculation and various strains and
so treated (Surin, 1959), it being con doses of challenge virus (Appleton et al,
sidered that these strains produce better 1963; Hofstad, 1953b, 1956). Although
immunological agents than mild or lento- survival after challenge may not evaluate
genic strains. However, Hanson et al. immunity accurately (Hofstad, 1953b),
(1951b) have shown that this is not neces this criterion has been generally adopted.
sarily the case. Because strain differences Clinical effects of vaccination. Formalin-
in immunogenic potency exist, it has been inactivated vaccines have been adminis
considered advisable to use several strains tered to chicks at 1 to 2 days of age with
in the production of a vaccine (Brandly few, if any, clinical side effects (Rao,
et al.,1946b). Formolized virus has been 1955, 1956; Chute, 1952; Hofstad, 1954b;
administered by the subcutaneous, intra Waller and Gardiner, 1952).
muscular, and intraperitoneal
intravenous Under field conditions and with
routes (Beach, 1945; Brandly et al, 1946b; chickens of varying ages, the vaccine has
Nakamura et al, 1937, 1956). Treatment been considered safe (Atanasiu and
with formalin has been found more effec Gareau, 1951; Haddow and Idnani, 1941;
tive than other methods of inactivation by Jacotot and Vallée, 1959). Egg production
Acevedo and Mendoza (1947), Appleton has not been affected (Placidi and San-
et al. (1963) and Coronel (1947). tucci, 1953c), except where incomplete

110
 TABLE 18 — Duration of Immunity Following Initial Vaccination of
Chickens with Formalin Inactivated Virus (Lancaster, 1964a)

Author Age when % surviving challenge1

vaccinated Months after vaccination

2 3 4 7 9 11 12

Chute, 1952 1 day 0 0 02

Waller and Gardiner, 19533 1 day 72 66 62 48
Rao, 1956 2 days 100 30
Miyamoto and Nagashima, 1957 4 days 100 80
Dardiri era/., 1957 10 days 98
Waller and Gardiner, 19533 10 days 96 84 88 80
Hanson et al., 1951b 3 weeks S
Hofstad, 1 953a4 3 wscks 82
Jacotot and Valine, 1 962 3 weeks 100
Rao, 1 956 6 weeks 100
Mitchell and Walker, 1952 8 weeks 100 100 90
Jezierski, 1 953 young s S
Hofstad, 1 953a4 3 months 100 100 100
Nakamura et al., 1956 4 months 75 66
Mitchell and Walker, 1951a adult 100 87
Mitchell and Walker, 1953 adult 90
Mitchell et al., 1952 adult 90
Miyamoto and Nagashima, 1957 adult 80 60
Traub, 1 944 adult

1 In many reports the end point of the duration of immunity was not determined
2 No immunity compared with non-vaccinated controls
3 Data taken from Fabricant
(1956)
4 Results with smaller challenge dose
S — Immunity considered satisfactory

inactivation or dissociation of the virulent chickens over 1 day of age is sum

virus in the vaccine has resulted in frank marized in Table 18. This summary is
disease (Mitchell et al., 1952; Placidi, based on the use of virulent virus for
1956b). challenging the resulting immunity.
One criticism against the addition of Authors who have described duration of
adjuvants to the vaccines for broilers has immunity tests for chickens under 1
been the persistence of the adjuvant in the month of age include: Appleton et al.
muscle tissue (Schoening and Osteen, (1963); Doll et al. (1951b); Generoso
1948). In addition, certain oils, when used and Agustin (1947); Grillo Torrado and
as adjuvants, have caused some post Smitsaart (1962); Hofstad (1953a); Lucam
vaccinal reaction (Generoso and Agustin, (1949a); Perez-Rebelo (1962); Rao (1955);
1947) of the granuloma type (lacotot and Schneider (1956); Traub (1943a) and van
Vallée, 1962). However, the use of an Waveren (1955).
aluminium hydroxide gel has not caused A number of authors have reported
any problems under field conditions favourable results after the use of formo-
(Eskelund, personal communication). lized vaccine. In some instances the results
Duration of immunity. The immunity were not determined by challenge; in
that follows the initial vaccination of others serological procedures were used.

Ill
These authors include: Acevedo and weeks later has resulted in a strong
Mendoza (1947), Atanasiu and Gareau immunity against virulent virus (Hofstad,
(1951), Balducci et al. (1954), Beach 1956; Miyamoto and Nagashima, 1957;

(1945), Botija and Loizelier (1948), da Miyamoto et al., 1957) lasting six months
Camara and Valadao (1961), Coronel (Miyamoto and Nagashima, 1957) to nine
(1947), Dedie and Starke (1952), Ellis and months (Nakamura et al., 1956). When
Crook (1953), Freudenberg (1950), Had- fowls aged 15 to 18 months have been
dow and Idnani (1941), Kaschula (1950, vaccinated twice at an interval of 12 days,
1952b), Lucam (1949a, b), Mansjoer the immunity has lasted 8 to 12 months,
(1961), Netter and Nguyen-ba-Luong depending on the adjuvant used (Jacotot
(1956), Pagnini (1950), Placidi and San- and Vallée, 1959).
tucci (1953c) and Schjerning-Thiesen In a vaccination programme which re
(1951). duced the mortality on a large farm,
In contrast, a number of authors have formolized vaccine was used at 15 to 17
had poor results after the use of formo- days; 10VS weeks; and 6 to 7 months
lized vaccines. These include: Adler et al. (Ellis and Crook, 1953).
( 195 1), Beach ( 1945), Clancy et al. ( 1949), The value of using a live vaccine for
Iyer (1943), Gualandi (1950), Levine and revaccination after a formolized vaccine
Fabricant (1952), Nakamura et al. (1937), has been examined. Thus, in the initial
Pomeroy (1951), Sturgess (1931) and stages of a vaccination programme,
Winmill and Weddell (1961). formolized vaccine was used for laying
Two methods have been adopted to stock and followed by a live vaccine two
prolong the immunity resulting from or three weeks later (Adler et al, 1951).
initial vaccination with a formolized A similar programme has been suggested
vaccine: revaccination with a formol in by Le Dosseur and Lissot (1949). How
activated vaccine, and revaccination with ever, a much longer interval has been
a live vaccine. recommended by Kaschula (1952b), who
The degree of immunity following two considered that at least 12 weeks, pre
doses of formol-inactivated Newcastle ferably five months, should elapse before
disease vaccine depends to a great extent a live wing-web vaccine is used after
on the interval between doses (Hofstad, initial vaccination with a formolized
1954b; Woernle and Brunner, 1957). A vaccine.
second dose of vaccine 2 weeks after the When Dardiri and Yates (1962) ad
first has not
appreciably improved im ministered formolized vaccine to chickens
munity (Doll etal, 1951b; Hofstad 1953b; at 2 weeks of age, followed by revacci
van Waveren, 1955), whereas a second nation with Bl virus at 4 to 16 weeks of
dose 3 weeks 1956) or 8
(Schneider, age, satisfactory immunity lasted 14
to 12 weeks later has resulted in a sub months after the last vaccination. Hilbrich
stantial degree of immunity (Hofstad, and Reuss (1963) studied a reverse pro
1954b; Miyamoto et al.. 1957; Rao, 1956; cedure in which the initial vaccinations of
Waller and Gardiner, 1952) lasting at least young chicks were with Bl virus, followed
eight months after the last dose of vaccine at 2 to 6 months of age by a formolized
(Hofstad, 1953b). When the initial dose is adsorbed vaccine. The resulting immunity
administered at 3 weeks of age or less, lasted at least 15 months.
revaccination should be delayed for at The comparative efficiency of formalin-
least four to eight weeks (Dardiri et al., inactivated vaccines and live virus vac
1961; Hofstad, 1953a; Rao, 1956). When cines has been studied. The formolized
the first dose is administered at 6 weeks vaccine was considered less effective than
of age or older, revaccination 16 or more the Komarov vaccine by da Camara and

112
Valadao (1961) and less effective than the 1930; Nakamura et al., 1937; Topacio and
Mukteswar vaccine by Zuijdam (1952a). Coronel, 1939), even though adjuvants
Netter and Nguyen-ba-Luong (1956) con were added (Fortner et al., 1959). In con
cluded there was little difference between trast, inactivation with the sodium salt of
the Mukteswar virus and a formalin- 8 - hydroxy - 7 - iodoquinoline - 5 - sulphonic
adsorbed vaccine. In contrast, when com acid has produced a satisfactory vaccine
pared with the lentogenic vaccine viruses, (Kraneveld and Nasoetion, 1948). Also,
Bl and LaSota, a formolized vaccine has an inactivated vaccine involving 50 per
sometimes given better protection on cent glycerin has given good results when
challenge (Dardiri et al., 1957),or there administered intradermally (Teklinska,
has been little difference (Miyamoto and 1951).
Nagashima, 1957). At a concentration of 0.5 g/ml, ure-
thane has been found to inactivate a
strain of Newcastle disease virus, as deter
Inactivation by Crystal Violet
mined by the absence of chick embryo
The use of crystal violet for the in infectivity. Administration of the treated
activationof Newcastle disease virus has virus by the wing-web route has given the
given variable results. Thus, Iyer and same HI titres as those resulting from the
Dobson (1941b), Iyer (1943) and Win- use of a commercial wing-web Newcastle
mill and Weddell ( 1961 ) found the results disease vaccine (Bower and Eisenstark,
with this chemical were irregular and 1954).
unsatisfactory. In contrast, a crystal violet
ethylene glycol vaccine has engendered an
Inactivation by Heat
immunity lasting 12 months (Doyle and
Wright, 1950), especially when a booster Dutcher et al. (1960) observed that
injection was given three or four weeks high temperature for a very short period
after the initial dose (Legenhausen and of time inactivated Newcastle disease virus
Sinkiewicz, 1959); Legenhausen et al., with a minimum destruction of antigenic
1959). Two doses of the vaccine have proteins. A series of three intramuscular
given protection against death 61 weeks vaccinations at intervals of three to seven
after vaccination (Bankowski and Corst- days resulted in a satisfactory immunity
vet, 1960). On the other hand, crystal lasting five to seven months.
violet vaccines have given less protection
when compared with either live wing-web
Inactivation by Ultraviolet
vaccines (Thompson and Osteen, 1952;
van Waveren and 1953;
Irradiation
Zuijdam, van
Waveren, 1955) or a tissue culture vaccine The practical application of ultraviolet
(Bankowski and Corstvet, 1960; Bankow irradiation in the inactivation of viruses
ski et al., 1958b). They have also been for the preparation of vaccines has been
found difficult to prepare in bulk (Mit reviewed by Taylor (1960). Prolonged
chell and Walker, 1951a). irradiation has killed the virus with an
associated loss of antigenic property

Inactivation by Other Chemicals (Sinha and Datta, 1950b). The infective

property of Newcastle disease virus has
A number of reports have described been destroyed more readily by ultraviolet
generally results following
unsatisfactory light than the less sensitive cytotoxic
the inactivation of Newcastle disease virus activity (Prince and Ginsberg, 1957) or
by a variety of chemicals (Coronel, 1939; the less sensitive agglutination property
Bindrich and Schmidt, 1955; Farinas, (Atanasiu and Suoto Patuleia, 1952). The

113
phenomenon of multiplicity reactivation fornia 11914) in embryonating eggs,

of Newcastle disease virus after exposure Nadel and Eisenstark (1955a) found the

to ultraviolet rays has either not been allantoic fluid to be non-infective, yet to

detected (Barry, 1962) or has been very have a positive HI titre. This fluid was
used a vaccine administered intra-
weak (Drake, 1962). Other studies on as
ocularly to day-old days
chicks, and 21
the inactivation of the virus by ultraviolet
irradiation have been reported by Drake later the immunity was challenged with
and Lay (1962). virulent virus. The "incomplete" virus
Oil-emulsified vaccine, utilizing ultra vaccine resulted in a level of immunity
violet irradiated virus, when administered that was the same as that engendered by
at 10 days and again at 42 days of age, has a commercial intraocular vaccine (Nadel
given protection for at least 20 weeks after and Eisenstark, 1955b). Similarly, Rott
the second vaccination (Legenhausen et al. 1963) have reported the
(1962,
and Sinkiewicz, 1959). Such durable im isolationand identification of the incom
munity was not obtained by Brandly et al. plete form and the viral microsome in
embryonating chicken eggs. Particles
(1946b).
designated "incomplete virus" have also
been demonstrated in intracytoplasmic
Inactivation by Ultrasonic
inclusion bodies found in tumor cell cul
Treatment
tures infected with Newcastle disease virus
Ultrasonic wave treatment was em
(Adams and Prince, 1957). Chromato
ployed by Michelsen (1951) and resulted graphic analysis has shown the presence
in no appreciable reduction in the infec of a non-infectious haemagglutinating
tion titre of Newcastle disease virus. component of Newcastle disease virus
(Wilson, 1962b).
VaccinesInactivated by Different
Methods Compared Combined Vaccines
Using three strains and two substrains
of Newcastle disease virus, Hofstad et al. To facilitate mass vaccination proce
dures, studies have been made in which a
(1963) compared the immunogenicity of
live Newcastle disease vaccine has been
Newcastle disease virus preparations,
combined with other virus vaccines and
mostly as allantoic fluid, inactivated by:
administered as a single inoculum (Beau-
(a) gamma irradiation under different
dette, 1949b). Thus, a live lentogenic
conditions; (b) 0.1 per cent formalin at
Newcastle disease vaccine combined with
2° to 4°C; (c) 1:1,000 beta-propiolactone
infectious bronchitis virus vaccine has
for two hours at 37°C. The immunity was
been administered as a spray or dust
challenged with virulent virus five to eight
(Crawley, 1954b; Markham et al, 1955a,
weeks after vaccination. The overall re
b, 1956b, 1957; Wardsworth and Young,
sults showed that beta-propiolactone as an
1955), in drinking water (Hitchner and
inactivating agent was superior to gamma
White, 1956; Hoekstra, 1961; Luginbuhl
irradiation and formalin in preserving the
et al., 1955) or by other means (Jansen
immunogenicity of Newcastle disease
and Richter, 1959).
virus.
The advisability of combining New
castle disease and infectious bronchitis
"Incomplete" Newcastle vaccines has been questioned on the
Disease Vaccine grounds that the combination may prevent
At one stage in the propagation of the the development of satisfactory immunity
virulent Newcastle disease virus (Cali to either or both diseases (Bankowski and

114
Rosenwald, 1956; Bankowski et al, 1955; web (Bengelsdorff and Schneider, 1963a).
Hitchner and White, 1956). On the other Newcastle disease virus has also been
hand, good protection against both dis combined with a fowl plague virus as a
eases can also be conferred by a combined mixed vaccine (Daubney and Ishak, 1953;
vaccine administered by different routes Lucam, 1949d).
(Markham et al, 1956b). To meet certain situations, it has been
Mesogenic strains of Newcastle disease found convenient to administer two
virus have been combined with fowl or vaccines simultaneously, but separately,
pigeon pox vaccines for wing-web or and by different routes. This procedure
intramuscular administration
(Cordier- has been used for Newcastle disease and
Boullangier et al, 1955a; Delpy and Hars, fowl or pigeon pox virus vaccines (Bran
1953; Dhanda et al, 1958; Jansen and et al, 1959; Gupta and Rao, 1959b;
v.d. Vlerk, 1954; Komarov et al, 1948b; Hartwigk and Untermann, 1962; Hutson,
Madhusudan, 1957; Prier, 1951; Suhaci 1953; Jezierski, 1953; Lancaster, 1957b)
et al, 1958; Van Roekel et al, 1948; and for Newcastle disease and infectious
Verge, 1954; Weidenmuller, 1954) or by bronchitis vaccines (Bankowski et al,
the feather follicle route (Richter, 1956). 1957).
Similarly, the lentogenic Bl strain has A triple vaccine, comprising live
been combined with fowl pox virus for viruses of Newcastle disease, infectious
wing-web administration. This has result bronchitis and infectious laryngotracheitis,
ed in good immunity to fowl pox but has been administered by the vent method.
insufficient protection against Newcastle This procedure has resulted in satisfactory
disease. Better immunity has followed two immunity to Newcastle disease (Winter-
injections of the vaccine into each wing field and Hitchner. 1961).

115
 PART IV: Virus Propagation and
Vaccine Production

VIRUS PROPAGATION METHODS

Propagation in Eggs hours after inoculation; and on the death
of the embryo, the amount of virus in the
Burnet and Ferry (1934) found New tissues approached that in the allantoic
castle disease virus was highly infective fluid. In contrast, the vaccine virus (Bl
for developing chicken embryos : death of
strain) appeared in the tissues of the
embryos occurred 30 to 48 hours after
embryo 24 to 30 hours after allantoic sac
inoculation. The technique used for inocu inoculation, and the titre of the virus in the
lation of the chorio-allantoic membrane tissues was appreciably lower than in the
has been described in detail by Burnet allantoic Furthermore, the vaccine
fluid.
(1936). virus did not kill embryos within four
Early studies on the use of the develop days. One feature of the Bl strain was its
ing embryo in the propagation of viruses ability to cause severe pneumonia with
have been reviewed by Iyer and Dobson cellular infiltration and inflammatory
(1940). Using 9- to 13-day-old embryo exudate in embryos six days after inocu
nating eggs, these authors inoculated a
lation. It appeared that the lungs of 16- to
Berkefeld V filtrate of a 1 per cent saline 1 7-day-old embryos were particularly sus
emulsion of liver and spleen from an in ceptible by the Bl strain.
to destruction
fected fowl. Usually, all embryos were Soon after the development of pneumonia,
dead by the second day. Material for sub the embryos began to die (Liu and Bang,
sequent passages was harvested on the
1953). Other reports have indicated that
second or third day post inoculation. the infectivity of Newcastle disease virus
Several factors relating to the virulence reaches a maximum shortly after the death
and propagation of Newcastle disease virus of inoculated embryos (Atanasiu and
in embryonating eggs were examined by
Gareau, 1952).
Florman (1948), Murchelano and Han
Certain chicken embryos, when exposed
son (1960) and Zebrowski (1957). Under
to large doses of Newcastle disease virus,
certain circumstances, a decline in anti
have shown a state of refractivity to the
genicity was recorded (Beach et al, 1948).
infection (Karrar, 1963). It is possible
Liu and Bang (1953) compared the
that this refractory state is indicative of
growth and pathogenesis in the chicken
viral toxicity (Burnet, 1955; Westwood,
embryo of three strains of virus differing
1961). Although some embryos have died
considerably in virulence. All three strains
within four days, no haemagglutinin has
reached a maximum titre in the allantoic
been detected (Karrar, 1963).
fluid 24 hours following inoculation into
the allantoic sac. However, the virulent A number of substances have tended
virus (Strain CGI 79) appeared in the to protect chick embryos against infection
brain and liver of the embryo some 20 with Newcastle disease virus. These in

116
clude Trypan blue and related compounds in embryonating eggs have already been
(Finkelstein, 1961a) and polylysine (Green referred to on pages 69 and 70, and the
et al., 1953). Furthermore, when a cul results of egg passage on the virulence of
ture of E. coli or its endotoxin has been Newcastle disease virus have been re
inoculated into the allantoic sac of viewed under the heading "Live Vaccines"
embryonating embryos some hours before on page 97.
the subsequent inoculation of Newcastle
disease virus, the results have been either Propagation in Avian Hosts
increased resistance or greater suscepti
Ducks
bility of the embryo to the virus, depend
ing upon the interval between inoculation One of the first reports on the modifica
of E. coli (or its endotoxin) and the virus tion of Newcastle disease virus by passage
(Finkelstein, 1961b). in a bird other than a fowl was made by
When embryos have been inoculated at Komarov and Goldsmit (1946b) who
45 hours incubation, a high percentage succeeded in carrying the virus through
has shown defective development of the 14 serial passages by intracerebral inocu
auditory sac and the embryonic lens lation of 4- to 8-week-old ducks. Thirty
(Blattner and Williamson, 1951), and the serial passages have been reported by
neural tube and visceral arches (William Syurin (1957), and 32 passages by Mitev
son et al., 1953). Eggs inoculated at 48 and Gagov (1958). The virus has also
hours incubation have shown no gross been passaged by intramuscular or sub
defect. cutaneous inoculation of ducklings 1 to
The virus in chorio-allantoic fluid has 10 days old (Seetharaman, 1951b). In
been subjected to purification procedures other studies, a virulent strain of virus has
and chemical analysis (Cunha et al., 1947). been attenuated by serial intracerebral
These studies have indicated that the passage in ducklings loss of anti
without
haemagglutination activity is proportional genicity (Martini and Kurjana, 1950).
to the virus content. Further purification
has resulted in a virus preparation without Pigeons
any haemagglutinating property (Edlinger The susceptibility of pigeons to New
and Vaux-Saint-Cyr, 1955). From the castle disease virus was early established
genetic point of view, both the sire and the by Doyle (1927) and this species has since
dam have apparently contributed to the been used for identifying the disease
variation in rate of viral growth in their (Beaudette, 1943) and for determining
eggs (Retae/a/., 1963). the pathogenicity of different strains (Olah
In embryonating chicken eggs, inter and Palatka, 1963). It has also been
ference between mumps and Newcastle found possible to carry virulent strains of
disease viruses has been demonstrated virus through a number of passages in
(Sinkovics, 1957a). An interferon has pigeons by means of intracerebral inocu
been shown to be present in eggs infected lation of brain suspension (Olah and
with Newcastle disease virus (Levine, Palatka,1963). Following nearly 400 pas
1962b). This interferon has reduced the sages of the virus by the subcutaneous
amount of Western equine encephalo route in pigeons, some attenuation of
myelitis virus produced by cells. virulence to fowls has resulted (Seethara
Newcastle disease virus has been main man, 1951b).
tained by serial passage in duck embryos
with aresulting attenuation for fowls Doves
(Seetharaman, 1951b). After 15 intracerebral passages in a
Macroscopic and microscopic lesions species of dove, the virus became avirulent

117
for fowls but retained its immunogenicity chicks has been reduced without loss of
when inoculated intramuscularly (Mitev antigenicity (Tokuda, 1956). Following
and Gagov, 1958). Also, serial passage the initial inoculation of mouse brain, the
in striped ground doves has resulted in virus begins to multiply and becomes non
considerable attenuation of a virulent infectious; later the infectivity titre of the
strain of virus (Martini and Kurjana, tissues rises, but falls again later as the
1950). process is repeated (Cairns, 1951).
An extensive pneumonia and pulmon
ary consolidation has been produced in
Propagation in Mammalian Hosts
the mouse by Newcastle disease virus
With the exception of man, Newcastle (Hanson et al, 1951a; Ginsberg, 1951).
disease is not spontaneously transmissible This effect has been considered to be due
to mammals under natural conditions to the toxicity of the virus rather than
(Verge, 1948). multiplication of the virus in the mouse
lung (Davenport, 1952). The toxic effects
Hamsters of Newcastle disease virus in the mouse
et al (1948a, 1949) passaged have been modified by viral interference,
Reagan
strain of Newcastle disease virus by a receptor destroying enzyme, or by
a virulent
xerosin (Grope and Dougherty, 1956).
300 times in the brains of Syrian hamsters.
Virus of the 92nd passage was non Following the intravenous injection of
mice, the virus has been cleared from the
pathogenic for chickens, although paraly
blood by the reticulo-endothelial cells of
sis and death followed the intracerebral
the liver (Brunner et al, 1960).
inoculation of hamsters. Hamsters inject
ed with virulent virus by the intranasal,
Guinea-Pigs
intraperitoneal and subcutaneous routes
have also been shown to develop signs of
In some studies, it has been possible to
or pneumonia or both make only a few passages in the brains of
encephalomyelitis
(Prudovsky et al, 1961). The results of guinea-pigs (Tokuda, 1956); whereas
these studies suggest that there is an other studies have indicated that the virus
of susceptible cells in the could be adapted to this species (Syurin,
abundance
nervous but that their 1957; Verge and Placidi, 1956), produc
central system,
capacity to produce virus is relatively low. ing a non-purulent disseminated enceph
alomyelitis (Syurin and Skalinski, 1957).
Fowls inoculated with guinea-pig-adapted
Bats
virus have developed HI antibodies and
Reagan et al. (1952a) found the large
have resisted inoculation with virulent
brown bat (Eptesicus fuscus) susceptible virus for a period of six months (Syurin,
to Newcastle disease virus administered
1957).
by different routes. The virus was re
covered from a number of organs taken Rabbits
at death or from bats surviving 14 days. Young rabbits have been found sus
ceptible to intracerebral inoculation of six
Mice strains of Newcastle disease virus (Reagan
Strains of virus have been successfully et al,
1954b). Symptoms of irritability
passaged in the brains of mice, especially and general paralysis resulted. In contrast,
when a susceptible strain of mice (Reagan the inoculationof virus into the anterior
et al., 1952c) or suckling mice (Tokuda, chamber of the eye has caused no symp
1956) has been used. After 43 passages tom in rabbits, although HI antibodies
in mice, infectivity for 3- to 5-week-old have been demonstrated in the blood

118
 TABLE 19 — Tissues used for In Vitro Propagation of
Newcastle Disease Virus (Bankowski, 1964)

Host Tissues

Chicken embryo Whole undifferentiated

Heart and lung
Kidney
Liver
Heart
Lungs
Fibroblasts
Skeletal muscle
Intestine
Chorio allantoic membrane
Amniotic membrane
Chicken, postembryonic Leukocytes (PE) (P)
Macrophage (PE) (P)
Porcine Kidney (PE) (P)
Lymph node (F) (L)
Bovine Kidney (F) (P)
Skin and muscle (L)
Rabbit Kidney (PE) (P)
Corneal endothelium (PE) (P)
Rat WRC-256-Carcinoma (L)
Mouse L-Fibroblasts (PE) (L)
Ehrlich's ascites tumor (PE) (L)
Guinea pig Kidney
Monkey Kidney
Human Lung (F)(P)
Heart (L)
Uterus (U12)
Maben-epithelial-adeno-carcinoma of lung (PE) (L)
H.ep.-2 (epidermoid carcinoma of larynx) (PE) (L)
K.B. (carcinoma of nasopharynx) (PE) (L)
Hela (carcinoma of cervix) (PE) (L)
Detroit 98 (adult sternal marrow) (PE) (L)

Source and type, if known or applicable :

L — line cell
F —fetal
PE — postembryonic
P — primary

serum (Bonaduce, 1954c). The adminis Monkeys

tration of antibiotics has not affected the Following passage of Newcastle disease
production of HI antibodies in rabbits virus through five different species of
(Bonaduce, 1954b). mammals, the virus has been adapted to
Following alternate passage in chick serial passage in monkeys with an encepha
embryos and newborn rabbits, Newcastle litis usually resulting (Collier et al., 1950).
disease virus has been adapted to serial
intracerebral passage in rabbits (Mitev
and Gagov, 1960a). After 21 passages in Propagation in Tissue Cultures
rabbits, the virus lost its pathogenicity for Goret and Provost (1963) have tabu
chickens but retained some antigenicity. lated the origin of some of the main cell

119
lines used in tissue culture work; and the Propagated in this manner, a virulent
properties of Newcastle disease virus when strain of virus retained its antigenic
propagated in a number of different cell properties. Similarly, the pathogenicity has
cultures have been described by Durand not been markedly affected (Day et al,
and Eisenstark (1962) and reviewed by 1953; Rusev, 1959; Seadale and Winter-
Fontanelli (1960), Sanders et al.
et al. field, 1956). A change in the neuropatho-
(1953) and Bankowski (1964). Bankow- genicity of a strain of Newcastle disease
ski (1964) has listed the cell types found virus has been produced by a method
susceptible to various strains of Newcastle involving the propagation in suspensions
disease virus (Table 19) and they are also of microglial cells obtained from the
summarized in Anon. (1963b). brains of chick embryos (Piraino and
Hanson, 1959). However, in monolayers
Avian Tissue Cells of chick embryo lung epithelium, there
Using minced chicken embryos in has been little evidence of significant
plasma and Tyrode's solution, Topacio amounts of infectious virus within the cell
(1934) reported 31 successive passages of (Rubin etal, 1957).
a strain of Newcastle disease virus. Details Using roller-tube cultures of epithelial
of the cultivation of Newcastle disease and mesenchymal cells from chick
virus in tissue cell cultures have since been embryos, Pereira and (1954)
Gompels
described by Seiffert (1955) and Buthala observed a progressive in the
increase
and Mathews (1957), and reviewed by virus content of the fluid phase which
Csikvary et al. (1961). In addition, the reached a maximum between the fourth
effect of the virus on the metabolic path and eighth day. Some cultures have shown
ways of embryonic chicken fibroblasts has characteristicchanges in the morphology
been examined by Magee and Sagik of infected epithelial cells (Bang, 1953a;
(1959). Hotz and Bang, 1957). In the chorio
No difference has been detected in the allantoic membrane, virulent strains of
infectivity peak densities of Newcastle Newcastle disease virus have rapidly des
disease virus when propagated in embryo troyed the host cell cytoplasm. Less viru
fibroblast monolayers of chicken, duck lent strains have caused hyperplasiaof the
and goose (Stenback and Durand, 1963). membrane with virus sometimes being
Zuschek et al. (1958a) reported that a released from cells without causing cellu
maximum titre of virus, when propagated lar destruction (Bang, 1953b). Certain
in isolated chorio-allantoic membranes cultures of chick embryo cells have yield
suspended in Tyrode's solution, was ob ed about 300 particles of virus per cell
tained at pH 9.5 in the presence of 0.2 (Kohn and Goldwasser, 1957). Electron
gm. CaCl, and 0.4 gm. KC1 per litre. microscopy has revealed Newcastle dis
Later Zuschek et al. (1958b) described ease virus particles in the cytoplasm 30
the effects of various concentrations of minutes after the inoculation of chick
antimetabolites on the growth of New embryo cultures (Mussgay and Weibel,
castle disease virus. They also found that 1962). Thus it has been concluded that
the virus grew best in chorio-allantoic morphologically intact Newcastle disease
membranes incubated at 42°C, although virus particles are capable of entering a
haemagglutinin was not demonstrable cell.
(Zuschek et al, 1959). Virus propagated in chicken embryo
In trypsinized chick embryo tissue cul skin tissue cultures has shown greater viru
ture, Newcastle disease virus has given a lence to young chicks when compared with
maximum infective titre of 10-7-5 per 0.1 the same strain of virus propagated in
ml. in 24 hours (Csikvary et al, 1961). whole embryo tissue culture medium (Day

120
 NEWCASTLE DISEASE VIRUS IN AVIAN TISSUE CULTURE

LD50 HA Titre

Hour

(Redrawn from Csikvary et al, 1961)

Figure 19.

et al., 1953). In chick embryo tissue cul determined in embryonating eggs; and
tures, the infectivity, haemagglutinating afterwards, following a number of 6-hour
and immunizing activities have developed multiplication cycles, the maximum infec
in a parallel manner (Cessi, 1959); al tive titre occurs at 20 hours post inocu
though virus titres have been lower than lation (Csikvary et al., as illus
1961),
those obtained by propagation in chick trated in Figure 19. This peak infective
embryos (Strizhachenko, 1961). Studies titre has remained largely unchanged up
using suspensions of chicken embryo tissue to the 50th hour post inoculation. The HA
cells have also shown that the inoculation titre of the culture has reached a maxi
of a virulent strain of Newcastle disease mum after 38 hours.
virus results first in a drop in virus titre as A comparison of the propagation rates

121
of a virulent strain of Newcastle disease infective virus has not per se interfered
virus and Strain F virus (Asplin, 1952) with the ability of an infected cell to
has shown that the latter virus is liberated divide (Wheelock and Tamm, 1961). The
more slowly from infected chicken embryo virus has been found to spread from cell
cells than the virulent virus (Mussgay, to cell through the intracellular milieu
1960). Another difference noted, when (Wheelock and Tamm, 1961), and pro
cultures of cells from chicken trachea gressive inhibition of mitotic activity in
have been used, is that the cells contained infected cells has been correlated with
a virulent virus for 12 weeks and a lento- continued production of viral antigen.
genic for at least 16 weeks after
virus Cytopathic changes have developed after
infection (Morehouse et al., 1963a). mitotic activity has decreased to low levels
Ultraviolet irradiation of cells has re (Wheelock and Tamm, 1961).
sulted in a higher yield of virus per cell In individual cells of HeLa cultures,
(Rosenberg and Rosenbergova, 1962). non-infectious haemagglutinins have been
Cell cultures from explants of trachea, identified (Bader and Morgan, 1961),
lungs and kidneys of chickens have been and also the synthesis of Newcastle dis
maintained for 4 to 16 months without ease virus ribonucleic acid (Wheelock,
subculture (Morehouse et al., 1963a). 1963). Bankowski (1964) showed that
Adult fowl spleen tissue with added fowl the propagation of the Bl strain in HeLa
serum has been used successfully in the cell cultures resulted in a predominance of
propagation of Newcastle disease virus infection of individual cells (Figure 20).
(Hayashi, 1956; Hayashi et al., 1962). In cell cultures of guinea-pig bone
marrow, formation of new viral antigen
Mammalian Tissue Cells has begun four hours after infection
Some strains of Newcastle disease virus (Jerushalmy et al., 1963).
produce distinct plaques in human cell The virus has been propagated in cul
lines without any previous adaptation of tures of embryonic human lung (Chap-
the virus to the plaque technique. The use ronieve and Pereira, 1955), human car
of a low concentration of agar in the over cinoma cells (Pigoury et al., 1962) and
lay followed by a non-vital staining of the human leucaemic bone marrow cells,
cultures after removal of the agar, has (Henle, 1960). Using fluorescein-conjug-
permitted the identification and titration ated antibody stain, Johnson and Scott
in HeLa cells of strains and mutants of (1964) demonstrated the presence of
Newcastle disease virus (Zavada, 1963). Newcastle disease viral antigen in the
A variant strain produced by continued nucleus and cytoplasm six hours after
passage in HeLa cells has been found non infection of HE p-2 cell coverslip cultures.
pathogenic to fowls, although it has Studies with Newcastle disease virus in
retained its immunizing properties (Hal- mouse L strain cells have shown that the
lauer, 1958). adsorption by an L cell of a single in
Using the fluorescent-cell counting fectious particle was sufficient to result in
technique, Newcastle disease virus pro loss of cellular integrity. In these cultures
duction has been shown to begin in HeLa the cytotoxic effect was accompanied by
cells 4 to 5 hours after infection, and to synthesis of large amounts of non-infec
reach maximum titres in 6 to 7 hours tious haemagglutinin and small amounts
(Wheelock, 1963), or in 24 hours (Tyr of infectious virus (Wilcox, 1959a). In
rell, 1955). In HeLa cell monolayers, the other cultures, cytopathogenicity has
addition of Newcastle disease virus has occurred only when the concentration of
increased the rate of release of potassium virus exceeded one egg infectious
dose per
(Klemperer, 1960); but the production of cell (Mason and Kaufman, 1960).

122
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AllSldAIUf) JO BIUJOJIlBQ )B )SIABQ
lBndiAidul Bjal-l sllao BuiMoqs snoirBA saBejs fo uoijarauaBad daondui Aq u/Brjs
iB awduoy) sBuBro x )0QP
BijAouAs fo B-/ofj sllao daondui Aq uiBrjs XIN31 uoijcJrosdBiuaBH fo uayo/uo dor
poolq sllao 01 BijAouAs auiduoy) sBuBro x )001
 Infection of L cells with small amounts vation (Kumagai et al, 1961); an in
of Newcastle disease virus has resulted in crease in this cytopathic effect has been
cell populations resistant to the virus. produced by the addition of hog cholera
These cultures found to contain
were virus (Matumoto et al, 1961). In tissue
small amounts of virus which persisted cultures of fresh swine kidney cortex,
at extremely low levels for extended Newcastle disease virus has produced cyto
periods (Paucker et al, 1962; Wilcox, pathic changes which first appeared as
1959b). A scheme of the events occurring cytoplasmic granules and vacuoles (Shi-
upon initial infection of L cells by New mizu, etal, 1957).
castle disease virus indicates that a cell The danger of introducing Newcastle
which adsorbs an infectious virus particle disease virus by employing chicken plasma
will produce infectious particles (Rod in any tissue culture system was clearly
riguez and Henle, 1964). demonstrated by Chanock (1955) during
In some mouse embryo tissue cultures, studies on variants of poliomyelitis virus.
Newcastle disease virus has not propa This latter virus was serially propagated in
gated well (Kono, 1962); in others the monkey kidney epithelial cells with the
virus has persisted for 1 1 days (Klapotke, production of granular degeneration. Cells
1952). which escaped the cytopathogenic effect
In calf kidney tissue, Newcastle disease of the Newcastle disease virus were found
virus has been either incompletely cyto- resistant to the cytopathogenic effect of
pathic (Dannacher and Fedida,
1962) or some types of poliomyelitis virus.
has shown a decrease in this effect (Rus-
seff, 1962). Tortoise Tissue Cells
The virus has shown appreciable mul
Newcastle disease virus has undergone
tiplication in cultures of rabbit corneal
ten consecutive passages in monolayers of
endothelial cells (Oh, 1961).
kidney epithelium cells of the tortoise
Other mammalian tissue cultures used
(Testudo graeca) (Schindarow and Todo-
for the propagation of Newcastle disease
rov, 1962). There was a pronounced cyto
virus have included swine tissue (Kumagai
pathic effect and nuclear inclusion bodies
et al, 1958; Bengelsdorff and Schneider,
appeared in the infected cells. The forma
1963b), monkey cells (Andre and Aude-
tion of these inclusion bodies was neutral
baud, 1960), Ehrlich ascites cells (Adams
ized by immune serum from a fowl (Fau-
and Prince, 1957; Flanagan, et al, 1955)
connier, 1963).
and rat carcinoma cells (Bang and War
wick, 1957). The virus has also been pro
pagated in seven other mammalian cell Propagation in Yeast Cells
cultures (Stenback and Durand, 1963). Sarajew (1954) cultivated Newcastle
Cell cultures derived from swine em disease virus in yeast cells in liquid and
bryo lymph nodes have been found to be solid culture media. Maximum multiplica
sensitive to small amounts of virus and to tion of the virus occurred after 24 hours
support the development of high titres of at 16° to 28°C. Repeated subculturing of
virus (Morehouse et al, 1963b). In swine Newcastle disease virus for 25 generations
testicular cell monolayer cultures, New in yeast cells resulted in loss of virulence
castle disease virus has produced cyto- for fowls with no detectable loss of anti
pathic changes after six to eight days culti genicity against virulent virus.

124
 PREPARATION OF VACCINES

Preparation of Vaccines from Cottral (1952), Cox (1952) has com

mented that in the commercial production
Embryonating Eggs
of vaccines, involving millions of eggs, no
contaminating virus agent has been
Egg-Transmitted Diseases
detected by the various types of safety
Factors that lead to the production of tests employed. Nevertheless, egg trasmis-
normal chicken embryos have been the sible agents must be regarded as being

subject of an extensive review by Landauer potentially able to contaminate vaccines.

(1961) and are summarized in Anon. Techniques used to inoculate embryo

(1959, 1963b). Not only the general nating eggs have been discussed on pages
health of the embryo, but also the import 68 and 69. Both the Roakin and Bl

ance of infectious agents in the egg has strains of vaccine virus have produced
been studied. Thus, Burmester et al. gross defects in chick embryos when eggs
(1956) found a commercial live New have been inoculated at less than 72 hours
castle disease vaccine caused a highly signi incubation (Williamson et al., 1953).
When inoculated at ten days incubation,
ficant incidence of visceral lymphomatosis
in inoculated chickens; from field evi the Bl strain has produced stunting and

dence, Gallego Piedrafita curling of embryos dying on the fourth

(1952) sug
gested that a contaminated formolized and succeeding days (Hitchner et al.,
Newcastle disease vaccine was the likely 1951a). This abnormal development may
cause of the appearance of fowl paralysis be due to changes in the flow of fluids
in a number of vaccinated flocks; Hejl and from the embryo into the allantoic cavity.
Faber (1959) found bacterial contamin When eggs inoculated with a virulent
strain of Newcastle disease virus have
ants in 35 of 53 samples of commercially
available Newcastle disease vaccines; Wil been incubated at 32.2°C, the embryonic

liams (1957) identified organisms of survival time has increased from approxi
faecal origin and various fungi in other mately 40 hours to 70 hours (McLimans
vaccines of egg embryo origin; Thompson et al., 1950). When large doses of virulent

(1954) thought that the causal agent of Newcastle disease virus have been inocu
chronic respiratory disease was trans lated into embryonating eggs, a number

mitted in a Newcastle disease vaccine, and of embryos have survived an otherwise

demonstrated that a culture of Newcastle lethal infection (Karrar, 1963). In these
disease virus contaminated with the agent resistant embryos, there has been little or
of chronic respiratory disease could be no multiplication of virus. The exact cause
carried through several embryo passages of this abnormal behaviour has not been
without the contamination being sus ascertained, however viral toxicity has
pected; Kruger (1961) emphasized the been considered likely.
possible transmission of leucosis virus.
Cottral (1952) has summarized diseases The Influence of Parental
of birds transmitted through eggs (Table Immunity on the Preparation of
20), and he has emphasized that, whereas
Vaccines
the different etiological agents in eggs have
been found, it has not been established Antibody passively transmitted from an
in all cases that the agents can cause dis immunized dam has an appreciable in
ease in hatched chicks as a result of egg fluence on the embryonic mortality rate
transmission. In discussing the report of following the inoculation of Newcastle
125
 TABLE 20 — Diseases of Birds Transmitted Through Eggs
(Modified from Cottral, 1952)

Disease Etiological Principal Selected

agent hosts references

Avian Virus-like Chickens Burmester et al..

lymphomatosis agent 1956
Avian Virus Chickens Van Roekel, 1959
encephalomyelitis
Newcastle disease Virus Chickens Lancaster, 1963a
Infectious Virus Turkeys Jerstad et al.,
sinusitis 1949
Psittacosis Virus Parakeets Meyer, 1934
Pullorum disease Salmonella Chickens Rettger and
pullorum Stonebunn, 1 909
Avian Mycobacterium Chickens Feldman, 1959
tuberculosis avium
Fowl typhoid Salmonella Chickens, Beaudette, 1 925
gallinarum turkeys,ducks
Paratyphoid Salmonella Ducks, pigeons. Wilson, 1947
infections sp. chickens, turkeys
Chronic respiratory Mycoplasma Chickens, turkeys Fahey and Crawley,
disease gallisepticum 1954

disease virus.Thus, with the Bl strain, no significant effect upon the final virus
Hitchner et al., (1951) found that em titre. However, for the propagation of Bl
bryos from Newcastle disease-immune vaccine virus, eggs from immunized hens
parents showed a mortality of 33 per have been considered unsatisfactory (Szak-
cent on the sixth day post inoculation; mary and Toth, 1963). Similarly, Topol-
whereas susceptible embryos gave a mort nick and Beganovic ( 195 1 ) have reported
ality of 67 per cent on the fourth day post higher haemagglutination titres in eggs
inoculation. The corresponding mortality laid by non-immune hens than in eggs laid
pattern for Strain F virus was 15 to 17 by immune hens.
per cent for immune embryos six days Using the Bl strain of virus, Reta et al.
post inoculation, and 75 per cent for sus found that both the sires and dams
( 1 963 )
ceptible embryos four days post inocula made statistically significant contributions
tion (Lancaster, unpublished data; Rao to the virus titres in the amnio-allantoic
andAgarwal, 1960). fluid, as determined by the HA test. This
Reports on the preparation of New was interpreted to indicate that variations
castle disease vaccine have indicated that in the rate of viral growth in eggs was in
eggsfrom vaccinated hens could be used fluenced by genetic effects.
(Daubney. 1953; Brandly et al., 1946c). With Strain F virus, no difference has
Furthermore, no difference has been been detected in the HA titres of fluids
found in the viral multiplication in 1 1 -day- from susceptible embryonating eggs com
old embryos from non-immune and im pared with eggs from immune dams (Lan
mune hens (Liu and Bang, 1953). caster, unpublished data). Nevertheless,
With the Bl Hitchner et al.
strain, whenever possible, embryos free from
(1951) reported a slight lag of approxi parental antibody should probably be used
mately two hours in immune embryos, al in the preparation of vaccines (Lancaster,
though the passive antibodies present had 1957a).

126
 AMOUNT OF NDV (Bl STRAIN) IN ALLANTOIC FLUID
AND EMBRYO TISSUES FOLLOWING ALLANTOIC SAC
INOCULATION OF NON-IMMUNE EMBRYOS

Log Dilution
- 9

-7

0/0 8 16 24 32 48 60 72 84 96

Hours after Inoculation

(Redrawn from Liu and Bang, 1953)

Figure 21.

Storage of Virus Material Use of Extra-Embryonic Fluids

The ability of strains of Newcastle dis
with or without Suspensions of
ease virus to survive for long periods faci Embryonic Tissues
litates the repeated production of vaccine Vaccines consisting of chorio-allantoic
from the same initial virus suspension. fluid mixed with some embryonic tissue
The virus in amnio-allantoic fluid has re have been prepared (Rao et al, 1963;
mained viable for over 28 months at 1°C Szakmary and Toth, 1963). However,
(Bonaduce, 1950c). However, lyophilized Gagliardi (1960) has reported that the
virus has been inactivated within 10 virus-containing fluid loses some of its
months when stored at 37°C (Hofstad and infectivity and immunogenicity when
Yoder, 1963). mixed with yolks from immunized hens.

127
 AMOUNT OF NDV (CG179) IN ALLANTOIC FLUID, BLOOD
AND EMBRYO TISSUES FOLLOWING ALLANTOIC SAC
INOCULATION OF NON-IMMUNE EMBRYOS
Log Dilution

-10-

I I I I I I I
0 8 16 24 32 40 4*

Hours after Inoculation

(Redrawn from Liu and Bang, 1953)

igure 22.

This effect may be due to specific antibody (Figure 21). Thus, the lungs of 16- to 17-
a the yolk and yolk should not, therefore, day-old embryos have been found very
ie included in the vaccine. susceptible to the Bl virus (Liu and Bang,
With the Bl strain of vaccine virus, the 1953). Under certain conditions, the
imount of virus in the tissues of the em- maximum yield of Newcastle disease virus
>ryo following allantoic sac inoculation per allantoic cell has been about 1,000
emains about 2.5 to 3.0 logs lower than particles (Horsfall, 1956).
he amount of virus in the allantoic fluid Atanasiu and Gareau (1952) found

28
that the infectivity of Newcastle disease transported in the field for periods up to
virus reached a maximum shortly after the three weeks. In countries or districts
death of inoculated embryos. Thereafter, where adequate freeze-drying equipment
the HA titre fell rapidly if the eggs were is not available, fresh vaccine might, per
kept at 37°C. Contrary results have been haps, be prepared and, with care, used
reported by Chang al.
(1956)
et who satisfactorilyin the field (Lancaster, 1956).
found that both the infectivity and hae- Nevertheless, in India, arranging for field
magglutination titres of the allantoic fluids workers to have a continuous supply of
continued to rise following further incuba the fresh (undried) vaccine and ice for
tion up to 24 hours after death of the em its preservation, was the most difficult part
bryos. A decrease in both titres began 36 of a Newcastle disease vaccination cam
hours after embryonic death (Chang et al, paign (Naidu, 1959).
1960).
The immunizing capacity of an inacti
vated vaccine prepared from egg fluids
Preparation of Inactivated
corresponds to the HA titre of the fluid.
Thus, the HA test has been considered a Vaccine
useful control procedure in the prepara
tion of vaccines from embryonating eggs The action of a number of physical
(Cordier et al, 1950; Hanson et al, 1947; agents on viruses has been reviewed by
Hill, 1957). Nevertheless, the embryo Pollard (1960). It has been considered
LD50 titration is much more sensitive as that true inactivation of a virus particle,
an indicator of the presence of virus than that is, complete destruction of infectivity
is the HA test (Hanson et al, 1947). and capacity of multiplication, must in
In the case of a virulent strain of New clude an irreversible change in its nucleic
castle disease virus (CG 179), the amount acid (Gard, 1960). Thus, the degree of
of virus in the embryonic tissues has close virus inactivation by beta-propiolactone
ly approached the amount in the allantoic cannot be reversed by dilution or pro
fluid (Liu and Bang, 1953), as illustrated longed storage (Lo Grippo, 1960).
in Figure 22. However, Cordier et al. Some of the problems associated with
(1950) have reported that the virus from the inactivation of virulent Newcastle dis
pulped embryos has reached a higher HA ease virus have been mentioned on page
titre than that from the amnio-allantoic 00. One of these problems relates to the
fluid. The influence of the route of inocu presence of active virus residue in an other
lation of Newcastle disease virus on selec wise inactivated product. In this connec
tive infection of the fully susceptible em tion, a report by Cordier et al. (1950)
bryonating egg reported by Hanson et al. has indicated that the adsorption of live
(1947) is shown in Figure 15 (page 71). virus on aluminium hydroxide results in
some loss of virulence of the non-adsorbed
virus.
Preparation of Fresh and Use of infected amniotic-allantoic fluid
Lyophilized Vaccine inactivated with formalin and diluted 1 : 80
in a thin aluminium hydroxide suspension
In particular situations, it has been im has resulted in better immunity than the
possible to prepare of
adequate amounts use of thicker commercial vaccines (Perez-
lyophilized vaccine of either lentogenic or Rebelo, 1962). The use of infected mem
mesogenic vaccine strains (Lancaster, branes and chicken embryos apparently
1957a). Under tropical conditions, fresh increases non-active proteins in the vac
(undried) Strain F vaccine has been cine.

129
Preparation of Vaccines from incubated stationary and tubes rotated in
roller drum. In the case of the stationary
Virus Propagated in Tissue a
cultures, the haemagglutination and the
Culture titres were lower and took
infectivity
longer to reach their maximum than the
To overcome some of the disadvantages corresponding titres attained in the cul
inherent in the use of embryonating eggs tures on the shaking machine.
(Bankowski, 1958), Bl and F vaccine The virulent field strain, designated
strains have been propagated in tissue California 11914, has been propagated in
cultures (Bengelsdorff and Schneider, suspensions of chick embryo cells (Bank
1963b; Brandt, 1961; Cessi, 1959; Gelenc- owski, 1957; Bankowski and Boynton,
zei and Bordt, 1960; Mayr, 1961; More 1948). Serial passage resulted in attenua
house et al., 1963a; Rossi, 1961a, b; tion without loss of antigenicity (Bank
Subramanyam and Pomerov, 1960). This owski, 1958) and this adapted strain has
subject has been reviewed by Fontanelli, been designated "TCND". The virus could
etal. (1960). also be propagated in cultures of HeLa
Russeff (1962) considered that the use cells and in bovine or pig kidney cells
of calf tissue culture eliminated the risk (Bankowski, 1958; Bankowski and Hyde,
of contamination by avian leucosis virus 1957; Provost et al., 1962). Syncytia
in the vaccine. However, a number of in were formed in HeLa cells, as shown in
vestigators have reported poor develop Figure 20 (Bankowski, 1964). One fea
ment of haemagglutinin and some loss of ture of this tissue culture virus was its in
antigenicity of the lentogenic and other ability to spread to susceptible in-contact
strains of viruses when propagated in tis chickens (Bankowski et al., 1958a). One
sue culture. Thus, the virus titres obtained dose of the vaccine administered intra
by propagation in chicken embryo fibro muscularly to susceptible chicks at 5 days
blasts have been lower than those obtained of age immunity lasting
or over induced
by using chicken embryos (Strizhachenko, at least 13 weeks(Bankowski et al., 1950,
1961), although no loss of antigenic pro 1958a). Revaccination at 39 days of age
perties has been noted for the mesogenic following Bl vaccine at 5 days of age
Komarov strain (Huygelen and Peeter- resulted in an immunity against virulent
mans, 1963) or Hertfordshire strain virus lasting 93 weeks (Bankowski et al.,
(Markovits and Toth, 1962). Good im 1963). Two doses of the vaccine gave
munizing activity of tissue culture virus immunity lasting from 33 weeks (Bank
has also been reported by Hallauer and owski, 1957) to 101 weeks after the
Kronauer (1960) and by Rossi (1961b). second vaccination (Bankowski and Cor-
Iwasaki (1954) examined a number stvet, 1960). When compared with the
of factors that influence the propagation Bl strain and a crystal violet vaccine, the
of Newcastle disease virus in chick embryo tissue culture vaccine gave appreciably
tissue cultures and compared three meth greater protection against virulent virus
ods of virus propagation using: culture (Bankowski and Hill, 1954; Bankowski
tubes attached to a shaking machine, tubes etal, (1958b).

STORAGE AND TRANSPORTATION OF VACCINES

Studies have been made on the storage 1960) or without. Strain F virus, as fresh
of fresh (non-lyophilized) virus, either unpreserved allantoic fluid, has been kept
with the addition of preservative (Bella, at a temperature of 4° to 6°C for at least

130
TABLE 21 — Survival of Newcastle Disease Strain F Virus at Different
Temperatures (Modified from Monda and DeBonis, 1959)

Vaccine at 1 5" to 27'C Vaccine at 37°C

Length of HA % chickens Length of HA % chickens

exposure titre surviving1 exposure titre surviving1
(days) (days)

25 80 100 25 40 100
35 80 100 35 40 80
45 80 80 45 40 0
55 40 25 55 20 0
65 40 10 65 20
75 20 0 75 10 —

1 % vaccinated chickens in groups of 5 surviving challenge

six months without loss of immunizing 27°C (Table 21).

property (Lancaster,1957a). Fresh, un The resistance of Strain F virus vac
diluted Mukteswar virus vaccine has re cine to adverse conditions has been re
tained its immunizing property after an ported by Torlone (1956) who found that
exposure of 7 days to an atmospheric tem the virus survived two months at 4°C
perature of 24° to 32°C (Generoso and after being kept initially for 48, hours at
Mendoza, 1950), or for 33 days at a 25°C. Strain F fluid has
virus as allantoic
temperature of 18° to 22°C (Gualandi, also been relatively
stable at temperatures
195 1 ) . Lyophilized Mukteswar vaccine of 25° and 37°C for 48 hours (Torlone,
has remained viable for 30 days at room 1955). Nevertheless, Newcastle disease
temperature 1957). A formolized
(Datta, virus is susceptible to exposure to daylight.
vaccine adsorbed onto aluminium hydrox Thus, unfiltered amniotic fluid infected
ide when frozen in a refrigerator and sub with the virus, when exposed to daylight
sequently thawed, has been found to be for four hours at 4°C has shown a de
antigenic, although there had been some crease in titre of about 3 log units (Skinner
lossof antigenicity (Placidi and Santucci, and Bradish, 1954). Exposure to light
1953b). However, a formolized vaccine therefore could cause serious errors in
with an adjuvant, following storage for infectivity titrations.
600 days at room temperature, has been More comprehensive studies have been
capable of stimulating protective immunity made on live Newcastle disease virus vac
in fowls (Walker and McKercher, 1954a). cines prepared as alyophilized product.
With the Bl virus, a frozen vaccine pre The immunizing property of lyophilized
pared with 20 per cent glycerin has main Strain F virus has been unimpaired after
tained an adequate titre during 12 months storage for one year at -22°C or nine
storage (Reising and Hitchner, 1954). months at 0°C (Lancaster, 1957a), as
Thawed vaccine preparations have gradu shown in Table 8. Similar material, when
ally lost titre when stored at 5°C. stored for two years at 4°C has shown
Using lyophilized Strain F virus in a loss of less than one log unit in virus

amnio-allantoic fluid diluted 1 in 2 in potency as determined by chick embryo

skimmed milk, Monda and DeBonis infectivity tests (Anon., 1961b). Other
(1959) found the immunizing power to be live lyophilized vaccines have been stored
unimpaired after 35 days storage at 15° to for over one year at -10°C without loss

13!
 VIABILITY OF KOMAROV STRAIN NEWCASTLE DISEASE
LYOPHILIZED VACCINE

Log Dilutions
-9

- 1
I I I I I I I i
1 3 5 7 9 11 13 15

Weeks

(Redrawn from Thorite and Mac Laod, 1960)

Figure 23.

of virus concentration (Borzemska et al, The Bl strain, when diluted 1:10,000

1961). Egg titrations have shown that, in tap water, has remained viable at room
with desiccated vaccine of the Komarov temperature for at least 11 hours (Lugin-
strain, the virus titre has not dropped buhl et al, 1955), although the same
greatly during 12 months storage at -15°C strain at 37°C has shown a lack of stability
(Thome and MacLeod, 1960). At room (Fernandez Espinosa et al, 1961 ).
temperature, the same dried vaccine has Studies have been on the
conducted
shown a progressive decline in titre aver purification of the virus.
and concentration
aging about half a log per week over Thus, an increased concentration of New
eight weeks. Thus, Thorne and MacLeod castle disease virus in amnio-allantoic
(1960) concluded that this vaccine could fluid has been obtained by adsorption on
withstand field temperatures up to a week aluminium followed by elution
hydroxide
without the titre falling below the stand with buffered saline (Zinca et al,, 1960)
ard (Figure 23). Kruger (1961) has sug or by thermo-diffusion (Medgyesi, 1958,
gested that the storage time should be 1951). Purification of the virus has been
three to nine months, depending on condi obtained by calcium phosphate chromato
tions. graphy (Reda and Rott, 1962) or by

132
thermodiffusion (Medgyesi, 1951). Chro- tains at least two components, one of
matographic results have indicated that which may be relatively non-infectious
one strain of Newcastle disease virus con- (Wilson, 1962b).

TESTING AND STANDARDIZATION OF VACCINES

Details for the testing of Newcastle dis when the ratio of mesogenic to lentogenic
ease vaccines have been reported by virus was 1 : 1 0,000. This is the same dilu
Gehring and Geiss (1958) and Oshel tion detected by Olah and Palatka ( 1963 ) .
( 1963 ) and are also given in Anon. ( 1 962c In the Federal Republic of Germany,
and 1963b). the manufacturers of the Bl vaccine have
Additional information on the testing of been obliged to test the vaccine according
vaccines has been provided by Olah and to prescribed procedures. The tests for
Palatka (1962, 1963) using the intracere purity, safety and efficacy have been out
bral inoculation of pigeons. These authors lined by Fritzsche (1963) and involve the
showed that a 1:10,000 dilution of viru use of embryonating eggs and day-old
lent Newcastle disease virus could be chicks. In West Germany, Government
detected in a Bl vaccine; whereas virulent testing of Newcastle disease vaccines has
virus alone produced symptoms and death been in operation since 1959 (Eissner,
in pigeons at a dilution of 10-6. Thus, it 1961).
appeared that large quantities of Bl virus The need for careful regulatory control
interfered to a certain extent with the of poultry vaccine manufacture has been
infectivity of virulent virus; a finding simi emphasized by Brandly (1950), Johnson
lar to that of Rosenwald et al. (1959). et al. (1954a) and Levine (1962a). In
Olah and Palatka (1963) concluded that addition, Eissner (1961) has discussed
the pigeon was more reliable than the standardization based on an adsorbed lyo-
day-old chick for testing the safety of B 1 philized standard, and he has suggested
vaccine. However, it must be emphasized that commercial vaccines should contain
that the day-old chick intracerebral patho not less than 40 protecting units per ml.
genicity index has been used to distinguish The practicability of using a single inacti
lentogenic and mesogenic strains of New vated standard for the calibration of both
castle disease virus over a wide range of inactivated and live vaccines has been dis
virus concentrations (Hanson, 1956). In cussed by Stableforth (1961).
addition (Hanson, 1956) has reported the A procedure for examining live virus
detection of a mesogenic virus in a mixture vaccines is summarized in Table 22.

TABLE 22 — Summary of Test Requirements for Live and Modified Live

Virus Newcastle Disease Vaccines (Van Houweling, 1963)
Salmonella Must not be present when examined by culture.
Extraneous bacterial contamination Must not be more than 10 colonies per bird
dose based on plate count.
Extraneous pathogenic viral contamination Must be none based on either bird or embryo
inoculation.
Chick safety Not more than 2 of 25 chicks, 5 days old or
less, may develop severe vaccination reaction.
Virus content There must be 10 s EIDS0/CC. after incubation
of the sample for 7 days at 37°C.
Immunizing capability 8 of 10 vaccinated birds must be completely
protected by one dose ; 8 of 10 controls must
be susceptible to the same challenge dose.

133
TABLE 23 — Summation of Thirty-two Potency Tests on One Batch of
Inactivated Newcastle Disease Vaccine (Coid et al., 1963)

Dilution of vaccine Chicks surviving /Chicks challenged

Place "G" Place "A"

1 :25 161 /223 (72%) 123/156 (79%)
1 : 50 121 /227 (53%) 87/156 (56%)
1 : 100 85/227 (37%) 60/156 (39%)
Controls 18/423 (4%) 10/341 (3%)

The results of 32 potency tests on a quantities of Newcastle disease vaccine are

single batch of BPL-inactivated vaccine traded commercially between different
during a period of 37 weeks have been countries, and more uniform and accep
reported by Coid et al. (1963) who table standards need to be established
showed that doubling the amount of internationally for Newcastle disease vac
antigen increased the percentage of chick cines. Further study and agreement is re
ens protected (Table 23). quired,so that poultry raisers in regions
At the present time, tests and criteria of the world where vaccination against
for the safety and potency of Newcastle Newcastle disease is permitted can be
disease vaccines are not uniform through provided with the most suitable antigens
out the world. However, appreciable for the immunization of their flocks.

134
 REFERENCES

ABDEL AZIZ, A.H., EL-NASSARI, B.B. AND IBRA ALBISTON, H.E. AND GORRIE, C.J.R., 1942.
HIM., k. Haemagglutination activ
1960. Newcastle disease in Victoria. Aust. vet.
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