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3"ULLORUM DISEASE STUDEES
RN TURKEYS

Thesis for file Degree of M. S.
MICHIGAN STATE COLLEGE
Ruth Aiexandra Carmen

1945

This is to certify that the

thesis entitled

Pullorun Disease Studies in Turkeys

presented by

Ruth Alexandra Corpron

has been accepted towards fulfilment

of the requirements for

___u._s.__ degree ”Way
8:
Public Health

WWW

Major plkofessor

DaterDchbel‘ 14. l_9_4_5

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-...— ‘H-.-._. ,A U _

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-‘~.-_o-o- n.. ..

uhr'm‘wnn.9v._.n ,u .

 

 

PULIOEJM DISEASE SNDIES IN 'NRKEYS

WILOBUM DISEASE SNDIES m 'IURKEYS

by

Ruth Alexandra Oorpron

A THESIS

Submitted to the School of Graduate Studies
of Michigan State College in partial
fulfillment of the requirements
for the degree of

W W scrum

Department of Bacteriolog and Public Health

lens

V‘. - ‘
l.".f‘_'bl$

 

AMOWLEIBEMENT

I wish to thank Dr. H. J. Stafseth for his gaidance and
help in this work. I also thank Dr. James Bivine and Dr. G.
W. Darby for their help in the handling and. autOpsy of the
turkeys. I acknowledge. too. the helpful suggestions and‘con-
structive criticisms of Dr. W. L. Mallmann in the preparation
of this paper.

I wish to acknowledge the financial aid given by Swift
and company of Chicago and thank the Bureau: of Animal Indus-
try and the Columbus Vaccine company for supplying the anti-

gens used for the whole blood tests.

I.

II.

III.
IV.

TABLE OF CONTEHTS

Introduction

Review of Literature.......................
Experimental

A. Procedures and Results.................

3. Bacteriological Findings...............
conclusions.....................................

References.....o..................o.............

62
71
73

a1-

INERODJO‘MON

l'or many years pullorum disease has caused serious losses
among the poultry of this country and in Dirope. Early investi-
gators in Southern Europe had noticed severe typhoid-like epidem-
ics among chickens, but the causative agmt was not discovered
mainly due to the lack of bacteriological knowledge at the time
(1). '

In 1899 Rettger (2) observed a peculiar epidemic in a brood.
of 17 hen-hatched and mothered chicks. Within ten days of pur-
chase. 1% of the chicks showed severe illness. resulting in the
death of 11 of the affected birds. All were suffering from whit-
ish diarrhea with pasting of the vent. He isolated a gran-negative
organism in pure culture from the livers of the dead birds. Elbe
following summer he observed a disease in chickens which was sympa-
tomatically the same as that he had seen previously. hiring the
next six or seven years he examined numerous chicks that were des-
cribed In their owners as having "white diarrhea." All of these
birds showed the same eysptoms and typical organisms. In 1909 he
suggested that the name, 'bacillary white diarrhea." be given to
the disease and that the causative agent be known as Bacterium
pu_llomm.

In 1925 the nuns of the organism was chnged to Salmonella
mllorum according to Bergey's classification. In 1928 the com-

mon name was changed to 'pullorum disease.ll
In 1916 Rettger, Kirkpatrick and Jones (3) reported the use

of a tube agglutination test for the diagosis of pullorum dis-

ease based on the method used by Vidal for diaglosis of typhoid
fever in lmmans.

In 1931 Goburn and Stafseth of this college (it) reported a
highly concentrated. stained antigen, based on Huddlescn's rapid
method for diagiosis of Bang's disease, for use as a rapid whole
blood test for the diagiosis of pullorum disease in chickens.
About the same time, workers in the Bureau of Animal Industry re-
ported a similar whole blood antigen (5). The whole blood test
is now very widely used in pullorum eradication programs for chick-
ens.

Pullorum disease was unknown or unrecogiized in turkeys un-
til poultrymen began hatching poults in incubators that had been
used for chickms. Hewitt (6) in 1928 was the first to report
the isolation of g; pullorum from baby turkqs that had been
hatched in a chicken incubator. Brunett (7) in 1930 pointed out
the similarity of diseased ovaries in chickens and turkeys. also
the isolation of Mllerum from the heart blood of a four-day—
old poult and the atrophied testicles of a mature turkey.

it first, pullorum disease in turkeys did not appear to be
a menace. but it has been increasing in prevalence over wide
areas in the last few years. Most losses occur in the young
poult. but survivors of the disease may become chronic carriers.
Hinehaw et a1. (8) isolated S. pullorum from eggs and ovaries of
turkeys that had had no contact whatsoever with chickens or chick-
on eggs. thus establishing the fact that the cycle is the same in

both.
Kinshaw et a1. (9). Dickinson (10), Bushnell (ll). Pomroy (12)
and others have cited the merits of the tube agglutination test

and have Inade comparisons with the tube test, whole blood and
serum plate tests in an endeavor to find the ideal method for
the diamosis and eradication of pullorum disease in turkeys.

Kinshaw reported late in 1939 (13) that the whole blood
test was only about half as effective as the standard tube
test. He pointed out the large number of positive reactors
that had titers of only 1/25 or 1/50. According to Schaeffer.
Hall, MacDonald and Binyea (11$) the whole blood test was equiva-
lent to the 1/50 dilution of the tube test for pullorum disease
of chickens. so it is possible that a number of the birds. nega-
tive to the whole blood method. were reactors in the 1/25 dilu-
tion of the tube test.

In our experimental work, we studied the relative value of
the tube agglutination test in comparison to the rapid whole
blood method. we also observed how soon after infection the
two tests would detect agglutinins and how long the birds would
remain reactors. Domroy and Fenstermacher (12) reported that. in
ezqaerimentally-infected nirkeys. the maxim titer ins reached
within a period of two weeks and that at the end of two months
the birds may become negative to the tube agglutination tests.
unless the organisms become established in the visceral crane.
They also reported variation of titer from month to month during
the infection. Titer variations of these birds were observed

over a period of eight months and are shown later in individual
charts.

-lt.

EXPERIMENTAL

Antigen preparation and gsneral technigue used.

i'he solid medium was prepared according to the suggestions
set forth by the Livestock Sanitary Association in 1932 (15).
{the formula is as follows:

Dry granular agar.........20 are.
Bacto-peptone.............10 gms.
Beef extract...............l+ as.
Distilled water.........lOOO m1.
This is adjusted to a pH of 6.s-7.2. In preparing broth, the
agar is omitted from the above formula.

The sterile medium was placed in Iolle flasks and allowed
to harden. The flasks were inverted and the water of condensa-
tion removed by aspiration and then incubated at 37° 0. for 1&8
hours previous to use.

Procedure.

1. The stock culture was streaked lightly on an agar plate to
obtain isolated colonies of Salmonella millerum and at the same
time three agar slants were seeded. {these were incubated at 37°
0. for 2“ hours.

2. me colonies on the plate were checked for smoothness. If
all colonies were smoth, the plate was discarded and one of the
tubes set aside for stock culture.

3. fhe growth was removed from the remaining am slants with
a small amount of 0.85% saline. This suspension was then used
to seed the flasks of broth which were incubated at 37° 6. for

21% hours. Smears were nude of the broth suspension and stained

by Gram's method to check for contaminants. If satisfactory.
the xolle flasks were seeded from the saline suspension and in.
cubated at 37° C. for #8 hours.

1+. The growth on the Xolle flasks was removed with a minimum
amount of phenolized saline (0.5% phenol and 0.85% Hacl). Care
was taken that the washing solution did not remain in contact
with the agar any longer than was necessary to remove the growth,
as agar has a tendency to smsitize the antigen. Washings from
each flask were kept separate until each yield was checked for
purity by microscopic examination. Satisfactory samples were
pooled and filtered through a thin layer of cotton. This con-
stituted the concentrated stock antigen. from which the diluted
antigen was made throughout the emeriments. an antigen was
stored in a refrigerator while not in use.

5. In making the diluted antigen. enougi phenolised saline

(0. 25% phenol and 0. 85% Neal) was added to a volume of the con-
centrate to give a turbidity of McFarland #1. Sufficient 0.1%
Each was added to bring the final pH to 8.2 by electrometric ti-
tration.

6. Each lot of antigen was tested with known positive and known
negative serum.

At the 36th Annual Meeting of the Livestock Sanitary Asso-
ciation (13) it was agreed that a serum dilution of 1/25 be used
as a ”finding dilution' in the tube agglutinatim test for pullorum
disease. In routine testing the serum dilutions are seldom beyond
the 1/50 dilution. HOVCVOI‘. with these experimentally-infected
birds, serum dilutions of 1/12.800 were made until it became evi-

-6—

dart that the agglutination response was diminishing.

The tube agglutination tests were incubated in a water bath
at 52° 0. for four hours followed by 37° c. overnight. Results
were observed the following morning (by Dr. Stafseth and the
author with the aid of a one-tube fluorescent lamp against a
dark background. Interpretations were as follows: M- I complete
agglutination, 3+ I incomplete agglutination. 2+ - partial ag-
glutination, 1+ - slight agglutination. 1' I suspicious and - I
negative. ' ' ' O
Shale blood antigen for thewrgpéd plate test.

Stained K 1&7 whole blood antigen (B. i. I.) and experimental
Redigsn (Ooluhbus Vaccine Gorpauy) were used for the rapid whole
blood tests. The Redigen antigen was discontinued after eight
series of tests because of the difficulty aperienced in reading
reactions. For all. whole blood tests. an electrically-heated
box which kept the temperature between 90°~95° 1‘. was used. An-
tigens were delivered to the plate with a standard size dropper
supplied with the antigen. Blood was procured and mixed with
the standard size nichrome loop. When the reaction was not in--
stantaneous. the plate was removed and rotated by hand before
final results were recorded.

For these studies 20 mature Mammoth Bronze turkey hens
were secured from a known pullorum-free flock. Q: January it,
1915, they were tested and found negative by the standard tube
agglutination test in dilutions of 1/25. 1/50 and 1/100. At

the same time they were negative to the whole blood test using
I 1&7 (B. i. I.) and experimental Redigen (columbus Vaccine Oom-
pany) antigens.

-7-

On January 5, the birds were separated into three groups.

Seven individuals received 73;" 2 m1. ’of a saline suspension_x..7

} \

of a freshly-isolated. 18-hr. “culture. of S. _pullorum, the amount
depending on the size and general condition of the bird. Seven
received 3. h or 5 m1. of the above culture orally, depending
on the size and condition. For this procedure, a 10 ml. pipette
was used to insert the organisms into the esophagus. After re-
moval of the pipette. the beak was held shut and the throat mas-
saged downward to prevent loss of inoculum. Six birds were left
as controls. All infected birds and controls were then housed
in a common pen which was left uncleaned throughout the experi-
ment to see whether the non-infected birds would become reactors
through pen contact.

The first bleeding was done on the fourth day after the
birds had received the organisms. The dilutions of the serum-
antigen mixtures were 1/25, 1/50. 1/100 and 1/200. The results
are shown in Table 1. It will be noted that, although the tube
test revealed the presence of agglutinins. the some of all birds
showed no agglutination with either of the whole blood antigens.
Bird {2621 was definitely sick and died during the night of the
fifth day. Autopsy findings revealed a septicemia with recovery
of 8. mllog from the heart.

It is unfortunate that the serum dilutions were not carried
beyond 1/800 on the sixth day. as all birds inJected intravenously
showed agglutination in this dilution. and it is not known Just
what the titer might have been. The tube and whole blood tests

showed considerable disageement as presented in Table II. In

-8...

the intravenously injected group. with every bird reacting in
dilutions of 1/800 to the tube test, the I ”7 antigen revealed
83.33 per cent positive and 16.66 per cent suspicious reactions.
while the experimental Redigen showed only 1/3 per cent positive.
1/3 per cent suspicious and 1/3 per cent negative. All birds

in the orally infected group were positive in dilutions of 1/100
and below to the tube test. In this group. I 1&7 revealed but
28.57 per cent positive. 57.114 per cent suspicious and 114.28

per cent negative. The Bedigen showed I42. 85 per cent positive
and 57.1” per cent suspicious reactions. In combining the intran-
venously injected and orally infected birds. I 1‘7 showed 53.8“.
per cent positive, 38.% per cent suspicious and 7.69 per cent
negative. Redigen revealed 38.1l6 per cent positive, 166.15 per
cent suspicious and 15.38 per cent negative.

By the ninth day after infection the shale blood tests were
all positive with one exception which gave a partial reaction
with Redigen. An interesting, but unexpected. difficulty occurred
with the tube agglutination tests on this w. Those. with two
exceptions, showed marked “zones of inhibition“ in the lower di-
lutions where the serum concentration was greatest. In most
cases the maximum agglutination was noted in the 1/100. 1/200.
1/‘400 and even 1/800 dilutions. The serun of some of the birds
had as low as 2+or even lein the 1/25 and 1/50 dilutions. Table
III shows the zone reactions.

Zone phenomenon was still present in the next tests which

were done on the 17th day. Bauer in 1922 and da Gosta crux in
1929 (16) had noted that these inhibition zones were most common

Table I

lour Dye

Gomarison of tube and whole blood tests

 

‘4‘

 

 

 

 

 

 

 

 

 

Turkey 1/25 1/50 1/ 100 1/200 In? Bedigen
2622 h” 2* 1+ D a a

a 2630 2+ 1» 1+ - - -

g $33 ’H lH' 2+ 1? o e.
2631 he M 1+ - o e-

E 2621- u. 3. x -. .. ..

0 $28 '4‘: 1+ : e- - O

'5 2627 3+ 1+ .'. - .. .
2623 1+ 1+ .. - . ..
i2 O o O O D D

3 3+ 2+ 9 O o -

a 2637 - - - - - «-

o 2629 3+ - .. - .. .-
2532 - - - - - -
$35 1" - - - D n
2638 1' - - - - eclet atypical
£36 . C O c- . -

'3 262% :- .. - .- -

.‘3 2626 t - - - -clot atypical

n

8 2‘? 9 ? o n - .

 

‘Bird #2621 was very si&. Died 5th day.
from the heart.

8. pullomn isolated

 

 

Table II

-10-

Six Dye
Comparison of tube and whole blood tests

 

 

 

 

 

 

Turkey 1/25 1/50 1/10011/200 1/ 1/800I xiv: Bedigenl
2622 1w I» I» M 2+ 1+ 2 -
- 2630 I» I» I» I» I» u. . 2
5 2633 a: I» u. I» u. u. . + +
, 2631 he. I» 1H- 1H- 3+ , + +
g 2621‘ , - pied‘duT‘L-ing-ni t of anuary 1. -
v 2628 1H- 1» m 1H- 1H- 3+ + -
5 2627 1» 14+ I» M 3+ 2+ 1' t
2623 I» I» 1» 3+ 1+ 2 .‘t t
2625 ‘H' 3‘" 2+ 1+ 1 e- 3 +
26314 I» I» M 3+ 1+ s + +
'3 2537 1H- 3: 1+ -3 ; - .'. +
0 2629 1H- 1H- 3+ 2+ , 1+ + 1
2632 1H 1H- M a: 2+ 3 it +
2635 'H- ‘H 114- 3!- +' t :t
$38 . o C - C e- e- o
.4 2536 Serun Loo unfilysed ”or use
a 262“ O O o D II C - o
:3 2626 - - - - - - - -
8 26 9 u- e- o o - Q o u
26 c O O o e- - u -

 

 

 

 

 

 

 

 

 

I'ZBird {2621. Salmonella pullorum isolated.

e011-

Table 111 line Days
Comparison of tube and whole blood tests

 

 

 

 

    

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I
M”es§§§§§§§ ”me‘m
hfihfifihfifih
£2622i3+3+3+3+h+h+3+3+t + +
326301+2+3+h+h+h+3+2+3 + +
52633l+1+h+h+u+3+2+1+t +slow+
326313+3+u+h+3+3+3+2+1fi «below:
gizsaammu-rgya: + +3
2627 MMM.1+--- '0' 4"
2623 2+ 3+fl3+ 3+3+b+1+t + .1...
Sesgihfltwflizn : I
326373+3+R+3+3I3+2+t- + +
°2229$336$t’*"“2 * *
2 1+ + +
26§§3+3+u+u+m333211+s +slow+
2638 --' ' .. -
H2636--- II D -
gZGZhH'Tl-F99 -- -
236626--- - - -
825:2: -- -

 

Note “some of inhibition'' in the lower dilutions where serum con-
centration is greatest.

l'Bird #2622 died on 10th day. General infection. Mucous enteritis.
_I.iver swollen. 8. Elena from heart. liver. spleen.

-12-

in instances where a very light suspension of bacteria was'used
for the antigen. Vith these, flocculation or agglutination usup
ally occurred.in a.tube toward.the middle of the series. In the
light of these observations. attempts were made to eliminate
these inhibition zones in the lower dilutions by running parallel
tests with standard antigen and.with antigens of varying concenp
tration. The serum remaining from.this date was run with antigen
three times as concentrated as standard tube antigen. The parap
llel tests covered a period of 21 days and four bleedings. The
results of these investigations are presented.in Tables 17a.and
IVb through VIIa.and VIIb. In most instances the reaction be-
came slightly more pronounced in the lower dilutions. but results
were not consistent enough to warrant changing the strength of
the antigen.

I Vidal and Sicard in 1897 and Witerberg in 1899 (17) found
that the agglutinins were precipitated mainly in the euglobulin
fraction of the serum. The presence of large amounts of euglo-
bulin. probably indicates the presence of antibodies. It is known
that the serum of new-born or very young animals is markedly de—
ficient in globulin content and especially the euglobulin frac-
tion. This deficiency is compensated for in the case of some
animals by the transfer of globulin from the mother through the
colostrum. The capacity for the young to produce its own euglo-
bulin comes a bit later and with it the appearance of antibodies
in the serum. In reference to poultry; Stafseth and Derby state
(18), “Evidently the tissues must undergp a maturation process

in order to become efficient manufacturers of antibodies. Mature

Seventeen my:

.13.

Standard mt igsn

Comparison of tube and whole blood tests

Table IYe

 

m +++++..+.+
e
R

 

7-
.14.. +++++..+++++++.......

 

..+++++++.

 

.3.......

 

 

09$)” ht rim? .m. . .. T... .t

 

can; assess. .?....s.s
can; sass». missus...
8i saute. assays?
8i asses. harness.
anagrams». assumes.
sufl amass. assumes. . . . . . ..

 

 

 

 

 

 

 

 

a; tress, tenures, a. 3:3.
snsassa. arrears ssn.ns.
q sense newness, unssnm
m sssss sssssss ssssss
evoseeeuasu dose «838

 

 

 

 

 

 

compare with 17b.

~11;-

 

 

 

 

 

 

Table Nb 3:: Antigen 17 We
0 °§§§
Mwsa§§3§§msa
unrnunnuuu
“2630144- lulu- era-+1....
32633MKMMEM3+1+1+3
,2631M3+MM3+2+2+3-¢
326283:2+1H1H-1I+13+2+i+t-n
32627 3+Mlv+h+1s3+2+1+1+4
252++'+M2+2+1+1+..
26233g+ag+3+2+1+---
26$}+3+3:2++-3~-.-~-
§2637 MM 2+O'.'-ee-
Isletslu3+3++t--
2632Mh+h+I3+2+-----
2635" is- sed'
26381--
”2636:-..
8262+...
“262613-molLysedL
32539: - -
26M2+++

 

 

 

 

 

 

 

 

 

 

 

 

-15-

Table Va 2h Days
comparison of tube and whole blood tests

Standard antigen

 

 

 

 

 

  
 

 

 

 

1
8::
c> C) 23 In
M°yna§§ss§s§ah7mnmg
I?“ I? ':? 2? I? 2? t? I? 32' I?
£26303+3+u+3+3+2+1+r-.++ 1
£26331I+1HII+EI3+3+1+11-+ + +
26319?!” 3+2+t'-3- «- - + + +
£2628 3+ 3+ M 3+ 3+ 1+ 1+ 1 .'. - 4. + +
32627u+3+u+3+3.2.1.:--.+ +}
26232+3+at3+3+2+1+1--+. .‘
25259? 1H3+3+1+r-‘-++ +
263n3+3+h+h+2+1+r---++ 1
32637 2+ 2+ 2+ 1+ 3. -- -- - - - ES lif- 4.
26292+2+3+2+1+--.---+ + +
2632 2+ 2+ 2+ 1+ 2 - - - - - + 1 3..+
2535 2+ 2+ 2+ 3+ 2+ 2+: - - .. 1 + 3,
26382+1+— - - -
fi2636: - 4- D II c-
826%---- - -
326261--- .. ..
8263907. 1?- - -
126%].1‘- - o - -

 

 

 

 

 

 

 

 

 

 

 

 

 

 

compare with Vb.

-16-

 

 

 

 

 

Table Yb 3: Antigen 214 Day.
0 O 0 §
Mevwa§§§§§§§3
h E 3 2 2 3 2 > h h
326303:lH-1H~1H-2+ 2+ 1+:t - -
@2633 mu+n+u+3+1+---
2631mu+u+u+3+3+1+:--
52628 a: ‘80 ndt mm uni
.3262? Mutiny 3+ 2+; .. -
2623Mh+h+h+3+2+1+1--
262 I” 1H "4’ 3+ 2+ 1+ 11' O o a
26 h+ h+ 3+ 2+ 1' - ,. 2 - -
E2637 3+ 3+ 3+ 1+ 1% - -. - - -
22293+3+h+3+2+1+----
2 2 + 2+ 1+ - - - - -
2635 3+ 3: a: 3+ 2+ 1+ 3 - - -
2638 t‘ - - -_
M.2636 : .. - -
8 262k 4 - - -
22626 5-!- - -
826 - - - -
2 .. - - -

 

 

 

 

 

 

 

 

 

 

 

 

{cable VIa 31 my:
Comparison of tube and whole blood tests

Standard antigen

 

 

 

 

 

 

 

 

.n 8°°8§3§ 3.7m
N 3 H 8 3 no H 9* u: an}
\ \ \ \ \ \ \ \ \
d‘ LII 1!! d 1:" 1:1 A El J-l_
" 3+ 21+ 13+ M 2+ 1+ 1 - . + +
g 1+ 3: 1H- 3+1 3+ 2+ 2+ 1+ - + t
, 3+ 2+ 3+ 2+ 2+ :t - .. + + g
a t 1 1+ 1+ 2+ 1+ 0' -' O + +'
3 3+ 3+ 11+ 11+ 3+ 2+ :- - - + +
3+3+ 3+ 3+ 2+ 2+ 1+* - + +
3+ 3+ 3+ 2+ 4'21“ t - - - + +
3+ 2+ 2+ 1+ t a - o a + +
a 1» 3+ 2+ 1+ 1+ - - .. - w+ .
0 3+ 3+ 3+ 2+ 2+ 1+ 1+ 3 .. +. +
3+ 3+ V? 3+ 2+ 1: - - .. +' +
3+ 3+ 3+ 2+ 1+ - - - + 1‘1
t 1’ - - -
g t - - -
2+ 5% I - I
t 1' .. . -
t. a - .. ..

 

 

 

 

 

 

 

 

 

 

 

 

 

 

-18..

 

 

 

 

 

 

Table V11: 14:: Antigen 31 Rye
Nike ‘3 O§ 8 0 §
3' a; 2 2 § § 2 .. 23 :2
\\\\\\\\\\
d fid—Ld—AAW‘
826301++h+3+3+2+t - - .. ..
§2633 1w do noT mfn ent
+2631 3+ 3+ 2+ 2+ 1+; - - .. ..
£26283+ 3+ 3+ 3+ 2+1 - .. - -
32627h+h+mn+2+1+t---
2623 114 3+ 2+ 2+ 1+ 1' - - .. -
2:223:31 - - .. .. .. .. ..
32637312+ 1+1 - - .. .. .. ..
02629h+h+ 3+ 2+ 2+- .. .. .. ..
2632M3+2+1+t-----
2635 3+3+1+- .7- - - .. - ..
2638 -‘ -. -
2636 - - -
1426211- - -
29- - c-
82530- - -

 

 

 

 

 

 

 

 

 

 

 

 

Table VIIa

-19-

38W:

Comparison of tube and whole blood tute

Standard antigen

 

 

 

 

   

 

 

 

 

o o x j
“”2§2§§2§§§ W““
1i H g! JFL 131 ('1 fi
326302+3+u+h+3+2+1+x. + +
3 2633 1+ 2+ 3+ h+ h+ 2+ 1+ 3' . + +
g 2631 0' 1+ 1+ 1+ 3' or -' é. a + +
'5 2628 t 3' 2+ 1+ 1+ 3- -- 2 . + +
,3 2627' 1+ 1+ ll+ 3+ 2+ 2+ 1+ 3 .- + +
26232+3+3++3+2+1+1+t- ++ +
262 3+ 3+ 3+ 3+ 1+ - «- . + + +
26 3+ 3+ 3+ 2+ 1+ 1. a. 4 . ¥' ¥ ¥
22637 34' 2+ : an‘ or -' u- a a; an as 1-
02629 3+ 3+ h+ 3+ 2+ t .. - a + + +
26321+2+3+2+1+:,__ +* +
2535 3+ 3+ 3+ 2+2+1+.. .. .. 1 1 +
2638 2"- -‘ ' . .L .L .1
H2636 2+ 2+ 3 u- c- .-
8262h a - - .. .. ..
22626 1dr- ,; - - -
8 $39 w: an i- - -
’Iawh
from 1+ 2+ h+ 3+ h+ 3+ 2+ 1+ : + 1
2627

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

-20-

 

 

 

 

 

 

arable VIIb 2x Antigen. 38 We
‘ o o 8 §
mmaas §§32 ggfi
2 2 2 2 2 2 2 2 2 2
- 26 + + h+ 2+ 2+ - ~ - u a
825§§§+§+u+u+2++:.-.
£2631 3+ 3+ M M '0" a In - 0- :-
326233+3+1++1++2+3+---
32627 2+ 3+h+2+2++ 3 t - -
262 '0' '0' + + 21’ ‘l’ I- c- - -
2623§+§+&L_+ : - - - .
2631+3+3+2+t»- - - - - ~
12637 3+ "V +' :' do u- u u- - O
32629 3+ 3+ In a: 2+ 1 - - - -
2632 3+ 3+ M 3+ + 3 .. - -
2635 3+ 3+ M 3+ :t- :l: :t .. o ..
2638 t u. c,
22636 2+ :t -
5222: :- .7 -
32639 c; : I
26h0 - a -

 

 

 

 

 

 

 

 

 

 

 

birds seem to be the best producers of antibodies." It seems
then, that like young mammals, the serum of young birds is also
deficient in globulin and for this reason birds under eight
weeks of age often do not show agglutinins in the blood.

Boyd and Bernard (19) have proved that since antibodies
are globulins, we would expect the serum globulins. especially
the euglobulins. to increase greatly during infection or immuni-
zation.

In reference to agglutination, which is the result of the
action of the antibody on the antigen. Gruber believed that the
imnnme serum in some way modified the peripheral layer of the bac-
teria. causing it to become adhesive. and that when these sensi-
tized cells touched each other, they clumped or agglutinated.
(20) Goulter, DeKruif, Northrop and Jones believed that the anti-
body is deposited as a thin film on the bacteria and that only
the homologous antibody has this power. Only minute amounts of
the film are necessary because of the fact that bacterial cells
will still agglutinate when the antibody-containing serum is di-
luted 10.000 or even 20,000 times in some instances. (20) Bordet
supposed that the antibocw, by its presence on the surface of the
cells in the antigen. sensitized them to the electrolyte of the
suspension. thus causing them to agglutinats. (l9) Boyd felt that

the combination of the antibody and antigen took place on the sur-
face of the molecules. and once formed, it is firm. He divided

the phenomenon of agglutination into two stages. the stage of com-
bination of antibody and antigen. which in some cases may be al-
most instantansous. and the second stage. the actual clumping of
the bacterial cells.

-22-

In the usual agglutination tests. the strongest response
is found in the lower dilutions where the antibody-ant igsn is
in greatest concentration. If the serum is diluted far enough,
a point will be reached in which there is no clumping and a
homogenous mixture results. This is due to the fact that there
is no longer mfficient antibody content to cause the bacteria
in the antigen to agglutinate.

Occasionally with some scrums. instead of the above reac-
tion, there is an inhibition of agglutination in the lower di-
lutions where the serum concentration is greatest. Topley and
Boyd refer to this as "zone phenomenon,“ while Gay calls it the
"zone of inhibition.“ Jones md Orcott (19) found that these
inhibition zones were due to a thin film which they considered
to be a type of globulin coating the cell and reducing its co-
hesiveness. {Ehey found in the course of their experiments that
if cells coated with globulin from inhibitory serum were washed.
they would late readily agglutinate when combined with non-
inhibitory serum. Zinsser (20) has explained inhibitory action
found in these zones of greatest concentration by assuming that
the antibody in uniting with the bacteria in the antigen carried
over with it some form of inactive protein which acts as a pro-
tective colloid. thus preventing agglutination. It would seem
from these two authors that inhibition zones may be the result
of modified globulin or protein material which overpowers the
ability of the cells to agglutinate in the tubes of greatest
serum concentration.

The mature turkeys used in these experiments were Just
coming into production at the start of the proJect. Becmse

-23—

the birds were starting production, lipoids in the serum would
increase as the period progressed. Further, the birds were
more heavily infected than they would have been naturally. so
their globulin content was probably increased considerably as
shown by the agglutinin response. Inasmch as the serum: con-o
tinned to show inhibition zones, 8. strongly positive reacting
bird, one that had been positive but had become negative, and
a negative bird were bled. These serum were submitted to Miss
Evelyn Sanders of the Central Brucella Station for electrOphore-
sis determination. The results were not significant, merely
showing a decided increase in globulin, the picture being mch
like that of chicken blood. ale did experience considerable
difficulty with lipoid material in preparing the serum for the
determinations.

In a later experiment with young poults, far too young
to come into production, it was interesting to note that on
only one day after they were experimentally infected with g.
mllorum was there slight zone phenomenon in the 1/25 dilu-
tion alone. Those involved five out of twenty-six birds.

There seems to be a possibility that the inhibition zones
noted in the tube agglutination tests. with the mature experiment-
ally infected turkeys, might be due to an increase in some one
or two of the globulin fractions in the serum, that combined

with, but did not permit. the agglutination of the bacteria sus-
pwded in the antigen.
Further the agglutinin response of the turkeys taken on the

days indicated are presented in Tables VIII through XIII, cover-

~2h-

8th Post Infection Bleeding 1&6 Days

Table VI I I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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53 W9

9th Post Infection Bleeding

Table II

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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~26-

 

 

 

 

 

Table 1 10th Post Infection Bleeding 6h Dye
. 0 § 3
Moree§§§§§a§ s
F F F 3 P } P 3 D 3'
a 2630 +- 2+ lH- ‘H- 3+ 33 a e e- +
g 2633 2+ 2+ 3+ 3+ + - - .. .. +
t» 2531 3 5+ 3' 3' 2' t - - - +
£2628 to +' 3' 2' 2 «g» t . so +
a 527 M I” I"? ‘H' 1' o I- e- n +
2623 24 34 3+ 33. i .. .. - . ‘ i
2625 9 9 2+ 3+ 2+ + s .. .. 5 4.
2631; +- 2+ - -- .- - - - .. . an;
'3 2637 - - er 0' f" .. .'. .. .. i low
2229 2+ 3+ 3+ 2+ 4- .- - a .- j +
32 3+ 2+ 3+ 3+ 3+ 1+ - I so i a.
2635 3+ 1H- 3+ 0' st _. o a Q a ’ '2’! +
2638 - - - : ' .L
2636 - - . ' ..
g 2621; - - .. -
a 2:26 - - - -
2 .. - .. .-
8 26 - - .- -

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Table 11 11th Post Infection Bleeding 90 mys

 

 

 

 

 

 

 

U
Maytag newflw‘
:zfiznnfinwufi
263° 2+ 2‘" + - a so + +
g 2633 4- 2+ 2+ + + 1 -. .. + +
p 2631 3+ 3+ 3+ 3+ 2+ + .. - + +
g 2628 2+ 2+ 3+ + + 1 .. - + +
a 2627 3+ 3+ 3+ 3+ 2+ 2+ + :t + +
2623 1H- 3+ 2+ +. i. .. . e- 4- +81:
2623Mh+3+ 2+4: at a .. +"$
8-0 2637 0" n‘ 0 er is u;- . sp as s!-
O 2629 3+ lH' 3+ 3+ 2+ 4' a u + +
2632 2+ 2+ 3+ 3+ 2+ + .- .. + +
2635 2+ +. -. . -. .. .. .. + .
E 2621: I : I I I
" 2626 - - - .. ..
8 26 9 - .. .. .. ..
26 c- a s) - .-

 

 

 

 

 

 

 

 

 

 

 

Table III 12th Post Infection Bleeding 121 Days

-28-

 

Turkey

1/25

1/50

1/100

1/hoo

1/1600

 

Intravenous

”k: £4- »+

.‘f f? “‘15
“$9'*¥P‘”\f

'f' ”3:" '4’ 1/200

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9" 1r! "§r°+.

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tuft-to-H_

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control
re
m
m
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Table m: 13th Post Infection Bleeding 1% Days

 

 

 

 

 

 

 

’ 0
Turkey tn 8 8 8 § § 8 a I“?
§ a 3 § a n S u
0 26 2+ 3+ 2+ 2+ + - - +
8 263?; 2+ + + 1 - — - +
a 263]- 2+ 2+ lH' ll-t + t - +
:2 2628 3+ 3+ + t .. .. - +
E 2627 2+ 3+ 3+ 3+ 4- .‘. .. +
2623 3+ 2+ +6 -' - - - +
262 3+ 3+ 2+ 2+ + - - +
263 :t. - - - - - - ..
'3 2637 cl” 2 -- .. - - - .,
2629 1H- 1H- 1H- 3+ + - - +
2632 t - - - - - - + ‘s'low
2635 3+ 2+ 1" o - - s- t
2623 .. - 4 .-
2535 c - .. -
g 262M +1 - .. -
fl 2226 - - '- e-
2 9 - - - .-
8 26 - - _ -

 

 

 

 

 

 

 

 

 

 

 

-30-

ing agperiod from the h6th to the lh6th day after infection.
Because zone phenomenon still existed on the 121st and lh6th
days, inactivation of the serum.at‘various temperatures for
short periods of time was tried.to see if this procedure would
eliminate the inhibiting zones in the lower dilutions. Shibley
(20) studied inhibiting zones resulting from heated serum. He
noted that Shiga antiwdysentery semm heated for tan minutes
developed zones when the temperature reached 63° C.. widened
the area of inhibition. and readhed its maximum between 72° C.
to 76° 6. along with a drop intiter. All agglutination ceased
at 76° 0. He also noted.that the time serum was submitted to
heating was important and that serums heated at 39° 0. for four
hours developed.these zones. He found.too that heating the serum
alone at 56° 6. increased.the inhibiting action, while heating
it at the same temperature with.homologous organisms restricted
the zone.

In these experiments with inactivation the serum was heated
for much shorter periods at lower temperatures to approximate
somewhat the method of inactivating serum for flocculation or
complement fixation tests. The series were inactivated at 56°
0. fer 20 minutes. 56° 0. for 1/2 hour. “6° C. for one hour and
60° 0. for one to two seconds and.then cooled. The results are

shown in Tables XIVBZYII.

-31.

Table XIV 121 We

nglutinin reaction after serum inactivation at 56° C. for 1/2 hour

 

 

 

 

O 0
May 3 8 § § § § 53¢ a
D l h 2 2 2 2 \_.
3 2630 + 2+ 2+ - - - - -
g 2233 + - - ; - .. .. ..
2 31 old: clay + - - - - -
3 2628 2+ t - - .. .. .. -
3 2627 I 1 .. 2+ 4+ 2+ 1 t -
2623 2+ 1 - - - .. .. -
2623 2+ + + - .. .. - -
263 .. .. - - .. .. - -
a 2637 a ‘ d - o c u -
O 629 " ID t u- t a 9- .-
2632 I. - t 9 b o a- -
$35 1 u o'- - 6. o as -
2638 - - ..
2636 .. - -
'é' 2621+ - - -
a 2626 - - -
'5‘ 26 - - -
o 26 _ _ _ ‘

 

 

 

 

 

 

 

 

 

 

 

-32..

table 17 1% We
nglutinin reaction after serum inactivation at ”6° 6. for 1 hour

 

 

 

 

 

m 2 8 E? f
M” 2 § 5 s i i s. E
U 2630 '.’ 1H- 2'.’ + u- a a
g 2633 3+ 3: t a- II b s- c-
s 2631 3+ 2+ 3+ 3+ 1' .. .. ..
g 2628 3+ 2+ 1’ :t - .. - -
B3 2627 3+ M 3+ 3+ 2+ . .. -
2623 + 3+. 3+ + - .~. - -
262 2+ 2+ 2+ + + .. .. -
263 .. .. - - .. - .. -
F1263] 1c1ay '- - - . .. - -
3 2629 1H- 1H- Nd- 3+ + - - -
2632 ' t O t - a a u .-
2635 2+ ‘+ at d at on o o
2638 O O .-
2636 - - -
8 26214 - - -
‘5 3226 - .. ..
2632. : : :

 

 

 

 

 

 

 

 

 

 

Idblc XVI 1H6 Days

Aggintination reaction after
serum inacti
at 55° 0. for 20 minutes YatiOn

 

 

 

 

 

 

Turkey '6? 3 8 o
2 2 S g i g g g
.2630 4- 2+ 1'
322;: - ' 1' ~ 3 ' ' '
+ - - -
E2628 2+ 3; 3: f - - .. -
a 2627 t + + + : 2 Z :
2623 2+ 1 2 ' t
26 - .. - ' ' ' ‘-
EE2637 .. - : .- - - ~ -
+¢'§2§3 2; 1 - : Z - - -
26,5 1 : 2:. : - - - 3
2638 - .. -
#2636 - .. -
32621; - - .-
32626 - - -
26 - - -
E26 - - .-

 

 

 

 

 

 

 

 

 

 

«an 2,1
Fflfitolhlv
(E
. . blm‘

 

 

-314-

!‘nble m1 116 We

Agglutinntion reaction after serum inactivation
at 60’ for 1-2 seconds and cooled

 

 

o 2

“W 2 a 3 2 E 2 2 g

H H H H H H H H
O 2630 3+ 3+ 2+ + + .. .. -
g 2633 + :t - - - .. - -
,, 2631 + 2+ 2+ + - - - -
g 2628 2+ 1: - - - - .. .-
TE 2627 t + 1’ - - - c- -

 

I

I
lltl-IH-l
III'III
Illllll
IIIOIOI

 

 

2632 i -
2635 2+ + ..
263s - .. -
H 2636 "' '- "
a 262k - - -
‘5 2626 - - -
26 - - ..
2 .. - -

 

 

 

 

 

 

 

 

 

 

flap. M‘.a§‘vfl * .L

 

~35-

The results of tests done on the 182nd, l9lst and 221st
days are given in Tables XVIII through XX. On these days,
as the titer of the birds was decreasing, it was noted that
the x ’47 whole blood antigen was less satisfactory in detect-
ing the weak: reactors. Of the birds with s. titer of 1/50 or
over in the tube test, the whole blood antigen missed approxi-
mately 35 per cent. On the 221st day when the final tests
were done. there was closer agreement between the standard
and whole blood antigens.

from these comparative studies of the two methods, it
would appear that in order to obtain good results with the
whole blood antigen. the serum agglutinins must be well es~
tablished and the titer 1/50 or over. Some birds that have
recovered from an acute infection and have become chronic
carriers with a titer of l/25 or weak 1/50 may be missed on:-
tirely if the poultryman depended wholly upon the mole blood
method for culling his flocks.

Because these experimentally-infected turkeys showed
zone reactions in the lower dilutions for the entire investi-
gation period of eight months, it seems that perhaps a three-
tube diagnostic test, using 1/25, 1/50 and l/lOO dilutions.
would be of value to eliminate the possibility of low or doubt-

ful reactions in the l/25 dilution being zone reactions.

4a

182 Days

fable XVIII

 

2

Quasa-

 

 

 

 

 

 

 

 

 

lhth Post Infection Bleeding

 

 

 

 

 

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mm\H~a+¥a.w/. .3M...%.ma .¢.lm....

v. 03 8 7 35.479 5 h.

e 33322 22332 3 «my/250%

M «garage Ma. Mara Madman/a fifififififi.
aflonehcuann Hugo Haoaoo

 

 

Table III 15th Post Infection Bleeding 191 Days

 

 

 

 

 

O O O 0
Turkey § g a g g g I“?
H H H H H H
a 2630 23 2 + .. - +
3 2633 - .. - - , 3’
§ 2631 - 1 2+ . 1
g 2628 :t s +, 3 7' 3%“;
a 2627 1' + 2... + _ +
2623 + .‘t - - _ _ veggglow
2625 1 ., , _ _ - t
2631: ; .. . - , _ _
'3 2637 .’. .. - .. - .. .'.
2629 2+ 2+ 4- t _ a + weak
2632 :t s 1: - , _ .
2635 3 3" e C- - - ,,
2638 .. - - _
.. 2636 .. - - 9
32222 " ' " -
3 26 .. .. - _
26 - - - _

 

 

 

 

 

 

 

 

 

 

-33.

 

 

 

 

 

 

Table II Final Bleeding 221 Days
' Ln 3‘0 8 3

W'fié §§§§§§§
. 263° 2'.’ + 1' + t - - c- I-
§ 2633 + t - ‘- .. - - - .-
fi 263]. +' '6' t 4' :- O O o u
8 2628 1' 4- + + 1' 1' - .. -
E 2627 1 '6‘ 2+ + t C- o c a
2623 1' 1 - - - - - . -
2625 1 - - - - - - - -

a :23“ c- - In a u- n C o o-
7 - O - - O O - C .

0 26% 2+ 2* + 1. o o C u o-
2632 t 1 - - - - - - -
2635 7' 7- . 4 - - - - .
2638 + 4- 2+ 1» t - - - -
.4 2636 3 1' .. - - .. .. - -
8 2621; 3+ 1» 11+ 3+ 3+ 2+ + + I
1; 2626 2+ '6- I - - - .. - -
3 2639 -- t- .... -. -. -- . . ..'.
26ho 3+ 3+ 3+ 3+ 2+ 1 - - -

 

 

 

 

 

 

 

 

 

 

 

 

Menu of agglutinatione followinggintravenoue inJection
of s. pullorum.

Since all of the turkey: injected intravenouely with g.
pullorum in the previoue experiment showed agglutinine in the
tube agglutination test and were negative to both whole blood
methods on the fourth day following infection, teete were made
to deternine the exact day on tick: the tests would become posi-
tive. Three additional turkey hens were obtained. After being
found negative to the tube agglutination and whole blood teete.
they were each injected with 0.5 ml. of a saline nepeneion of
freehly ieolated. 18-hour culture of g. pullorum. mil: teete
were made ueing the etandard tube and whole nethode until both

teete becue definitely poeitive. Iheee reeulte are preeented

 

 

 

 

in table XXI.
Table 111
Lppearance of agglutinine in ”D
tube and I 1&7 whole blood teete
o o o u
o o o o
mmaazaeefifié:
} h 7-? I? h D .-c > > M
a
O
2:; 98 - - - . .-
u 99 - .. .. ..
33 1w "' - O a
98 o a o a
v .. .. .. ..
£2123 - - - ..
98 :3 no a ee
6 - e- - -
4&2199 - - - -

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

~180-

Tabls XXI continued

 

 

 

 

 

O

m, masmsm

fizfinrzfig
6983...
2+1+u

98 2+l+----.

5399 RIMM3+3+l+l+u

F100---.----.

98 ”4"? 2+1+--0-

gaggmiawryyatn

10° 3+2+1+ocuase

 

 

 

 

 

 

 

 

 

 

 

 

 

Beckwith and Jones (20) detected agglutinins on the third
day following a single injection of mtigen with his experimen-
tal animals. i'hey state that the maxim response occurs in
from six to twelve days and that sgglutinins have been detected
in the blood of rabbits from three to eigxt months following
a single injection of antigen.

i'he ngglutinins apparently begin to appear in the blood
of turkeys on the third day following intravenous injection
of §._pu116nm. as shown in the standard tube agglutination
test, but do not appear with the whole blood antigen until the
fifth day. this is probably why the whole blood tests were
negative on the fourth day in the previous findings.

Ponroy and l‘enstcrnaoher (12) have mentioned the fact
that the titer of diseased turkeys may vary rather consider-
ably during the course of pullorum infection. These experi-

mental birds were emined at intervals over a period of 221

-hl-

days following infection to note agglutinin response, variations
in titer and to compare the titers of the intravenously injected
birds with those that had received the organisms by oral admin-
istration.

The shove authors agree with Beckwith and Jones by demon-
strating that the maximum agglutinin response occurs within
two wedzs. They state, however, that the serum of turkeys
mg become negative at the end of two months unless the organ-
isms become established in the visceral organs. Vith the ex—
ception of three instances when the maximatiter was reached
on the 17th day, all the infected birds showed maxim reaction
on the ninth day. Very shortly after the maximum titer was
reached, there was a decline, after which there were fluctua—
tions in the degree of agglutinin response. The majority of
the birds in the intravenously injected group still had titers
of 1/200 at 221 days. One had fallen to 1/50 and the other to
1/25 on the final day.

The orally infected group with one exception had become
non-reactors to both tube and whole blood tests after 221 days.
As will be seen later in the autopsy findings, this one excep-
tion. Bird 2629, with a final titer of 1/2oo yielded §._pullorum
from the ovary-oviduct, giszard—kidney, duodenum-pancreas. mid-
gut and cecal stock.

alerts are presented for each individual to show the de-
gree of response and fluctuation of titer during artificial in-
fection with S. pullorum. Included also are block-type charts
to show the average titers of the intravenously injected and

orally infected birds for the period.

43%... .31. ev .

 

 

 

 

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-57-

msceptibility of the control gotho oral administration

21: Salmon ella pullorum.

The air control birds had been kept in a common pen with
the infected birds throughout the entire experiment to deter-
mine whether they would become reactors through pen contact.
They had remained. negative to the whole blood tests for 191
days. Ior a period of 21 days very early in the experiment.
they had shown incomplete to partial agglutination in the 1/25
and 1/50 dilutions of the tube tests, but the results were not
consistent as different birds reacted each time. This difference
in reaction was attributed to increased globulins or lipoid
material in the serum. because they were coming into produc-
tion at the time. It was evident that there were some "open
cases" in the infected group as a pen of chickens was infected
with S. pullorum from contaminated droppings of the turkeys.

'i'he Storrs Experiment station in 1912 (2) began experi-
ments to determine whether mature chickens would deve10p g.
pullorum infection from contaminated food and drink. This was
done by examining the eggs of the birds that were exposed to
natural infection. They noted that the first evidence of new
infection appeared at about eight months after. the beginning
of exposure when a hen laid an egg from which the organism was
recovered.

In order to determine whether members of the control group
were susceptible to the infection, on the 1‘95th dav they were
each given orally 2 ml. of an 18-hour. mixed broth culture of

8.43u110rum isolated the week before from two chickens and a

-53-

duck. Tests were made to detect when their blood would become
positive to the tube and whole blood tests. In the previous
experiment. with intravenously infected birds. the tube tests
showed agglutinin response on the third day. while the whole
blood antigen did not reveal the positives until the fifth day.
With this orally infected group,, the birds were all negative

to both tests on the third day. In the original orally infected
group, the agglutinins were Just beginning to appear on the
fourth day in the 1/25 and 1/50 dilutions. while the whole
blood tests remained negative. The sixth day following oral
infection these birds were positive to the tube test in l/lOO
dilutions and beyond, the maJority of the reactions being 1/hoo.
Even with these titers. the whole blood test revealed only 28.57
per cent positives with the K 1&7 antigen. .

It was interesting to note that on the eighth day following
infection. only 66 per cent of the control group showed agglutin-
ins in the tube tests. At this time with the whole blood anti-
gen there were two disagreements. In general. the titersk of
the control birds were much lower than what was noted in the
group infected earlier by the oral route. For comparison Tables
XXII, XXIII and XXIV are given to show the difference in response
in the two groups. Table xxfle shows the response of the con-
trol group on the eighth day following infection, while XXIII
and XXIV show the reactions of the original orally infected birds
on the sixth and ninth days. ’

Even on the 2‘4th day, only one bird in the control group
had a titer beyond 1/hoo in the tube test. Tables m and

DWI are presented to give the differences of the two orally

-59-

Table XIII

nglutinin response of Control grow after
oral infection of 2 cc. of 8. pullorum
8 days after infection

 

 

 

 

 

 

 

 

 

 

 

 

Turkey n 2 § §
\ \ \ \
d _.J __l
26 + + 2+ 1+
26%: 3 g i -
262M 3+ 3+ 3+ 3+
2626 e+r c+7 cldyi -
26 1 _.. _. -.
26 0 3+ h+ u+' u+
Table rXIII I

Original orally infected group

 

   

 

 

 

 

 

 

 

 

 

 

 

 

6 days after infection

Turkey in o o o E

eeasée t

2625 h+ 3+ 2+ 1+ - - - 3
263R h+ h+ h+ 3+ 1+ -' - + .

2637 h+ a: 1+ -- - e - i

2629 h+ h+ 3+ 2+ 1+ 1+ +

2632 h+ h+ h+ 3+ 2+ - - 1

2635 u+ n+ u+ h+ 3+ 1+ -- '1

Table in? '

Original orally infected group
39 s after infection

 

 

 

 

 

 

 

 

 

 

 

 

 

”kegfigsaee‘a ge-
-4 :=L_ “ LEE; “' “ “ BEE. “
-=L1 rflhvdi—wli-
2623 2+ 3+ 3: 3+ 3+ 3+ h+ 1+ 3 +
:22 .3+ 3+ 3+ 3+ 2+ 1+ 0 e +
3+ 3+ 1+ + + + 2+ 1+ e +
2637 3+ 3+ fi+ 3+ 3+ 3+ 2+ : - +
2629 a: 3+ u+ h+ h+ 3+ 3+ 1+ 3 +
2632 h+ h+ M+ h+ 3+ 3+ 1+ I +
2535 3+ 3+ h+ wh+ h+ 2+ 1+ ' e +

 

-60-

Table XXV

control group - orally infected
2“ days after infection

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

<3 C) c: ‘3 ‘3 23
xx (3 <3 C) {8 r—
Mkevunnuigqgae
P1 Pl r4 rt .4 .4 r4 F! rt 54
2636 t i - :- ee e- e- - :8
262% 3+ N+ h+ 3+ 3+ 2+ + + 1 +
2626 2+ + t - - - - .. +
2239 -. -. -. . _. 1. - ,_ ,~, ,,
2 0 3+ 3+ 3+ 3+ 2+ - - - - +
Table xxv:
Original orally infected.gxoup
2h days after infection
<3 53 c: g;
Mkovna§§ 3§§a§ “'3
Pt '4 r4 rd +4 +4 r! r! F: e+ be
2623 2+ 3+ 3+ 3+ 3+ 2+ + 1 - - +
2625 3+ 3+ h+ h+ 3+ 3+ + t - - +
263’} 3+ 3+ M 1|"? 2+ +‘ t o'- O s- ‘V
2637 2+ 2+ 2+ +- t- - .-. .;. .. . +
2629 2+ 2+ 3+ 2+ +- e ..'. .. . .. +
2632 2+ 2+ 2+ + e - - - - - +
2635 2+ 2+ 2+ 3+ 2+ + 1' a - - +

 

 

 

 

 

-51-

infected groups on the 2’4th day following infection.
In the light of the experiment done at the Storrs station
with chickens, it appears that mature turkeys exposed naturally

to S. pullorum develop the disease after a longer period of

 

time than was allowed for this group. It would have been an
advantage to trap-nest the control hens and examine the eggs
for the organism. By infecting them with S. pullorum cultures
orally, they proved to be susceptible to the disease but to a

lesser extent than the birds in the original group.

-62..
Bacteriological findings.

The turkeys were killed and dressed at the Frozen locker
System and returned to the laboratory for evisceration of or-
gans for bacteriological examination. Gross lesions were noted
and sections of various organs that were apt to harbor _S_.__p_u_l~
1n______rum were taken. is this was an attempt to merely recover
the organism from experimentally infected birds, the organs
were in some cases grouped together for the sake of convenience
and conservation of mediums. The following groupings were
made: (1) ovary and oviduct; (2) liver, gall bladder, spleen;
(3) gizzard and kidney; (‘4) crOp; (5) duodenum and pancreas;
(6) mid-gut; (7) cecal-stock; (8) lungs, heart. air sacs.

Each of the above group of tissues were ground well in
sterile mortars with a small amount of sterile sand and broth.
A loopful of the broth suspension was streaked directly on 8. 8.
plates (Difco) and incubated at'37° G. for ’48 hours. At the
same time a small portion of each tissue was placed in tetrathi-
onate broth for enrichment, and after incubation at 37’ C. for
18 hours. the cultures were streaked ligitly on S. 8. plates
with a bent wire. After incubation at 37° C. the plates were
examined at 21+ and 148 hours and suspicious colonies picked.

For the initial determination of type, they were seeded into
lactose motility agar. (Darby modification of 131ch motility
agar) and into dextrose broth containing 0.5 per cent dextrose
and l per cent Andrade's indicator. After incubation at 37° C.
for 21+ hours, all definite coliform organisms showing acid and

gas in both lactose motility and dextrose broth were discarded.

-63..

Non-motile organisms showing no lactose fermentation. but acid
and gas in dextrose. were set aside for further study and posi-
tive identification of S. pullorum. Non-motile organisms showing
no fermentation of lactose. acid. but no gas. in dextrose were
incubated for another 21% hours. If no gas developed. they were
then retained at room temperature for one month and examined

at four to six day intervals for possible slow lactose fermenta-
tion.

Growth from the suspicious tubes was seeded into dulcitol.
maltose. sucrose and adonitol broth. Tryptone broth was inocun
lated for indol determination. All cultures that were non-
motile with acid and gas in dextrose, but no change in lactose,
dulcitol, maltose, sucrose and adonitol, were considered to be
S. pullorum if they were also non-indol producers. Gram stains
confirmed the typical morphology. Spot agglutination tests were
done by mixing a little growth from the lactose motility tube in
a drop of physiological saline on a microscopic slide. To this
was added a lOOpf‘ul of a known positive serum of high titer. If
agglutination occurred, the presence of S. pullorum was considered
positive.

The results of the post-mortal: examinations and cultures
are shown in Tables XXVII. mm and XXIX. It was surprising
to find that in some instances the only tissue yielding .3; pullorum
was the section of duodenum-pancreas. The question arose as to
which one of these two organs was harboring the organisms. How-
ever. Johnson and Anderson (23) report lesions observed at the
autOpsy of 161 turkeys. They state that "the most consistent

lesion was slight duodenitis with occasional petechia in the duo-

-614-

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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-67.

denal mucosae.” Mallmann in 1928 (21). in mentioning the in-
cidaice of S. pullorum in the intestines of chicks, pointed

out the desirability of culturing the intestines in routine
diagmstic procedures. Posell (22) in 13 of 71 poults recovered

S. pullorum from the intestine, so he believed that culturing

 

the intestine routinely would increase the incidence of m-
_1_g__ru_xg diagnosis in turkeys. Four of the birds yielded the or-
ganism from the midp-gut or cecal stock as well as from the duo-
denum area. Bushnell and Porter (25) state that if the intestine
is not cultured it will lead to an important source of error in
diagiosis.

S. pullorum was isolated several times from the ovary-oviduct
culture. Brunett first isolated the organism from a mature tur-
key in 1929 (7) and since then Hinehaw and others have recovered
the organism from the ovary on several occasions. In fact Hin-
shaw (2“) states, "The lesions are confined principally to the
reproductive organs." In hens he observed non-functioning ovaries
with caseous plugs in the oviduct which yielded S. pullorum in
pure culture.

These mature turkeys did not show S. pullorum in the heart-
lung-air sac culture. Johnson and Anderson (23) reported that
the lungs were usually free from gross lesions with only an oc-
casional bird showing congestion of these organs. Posell (22)
has pointed out that, in young turkeys. the disease is usually
septicemic end that the organism can usually be recovered from
the heart. liver and lungs. Adult birds in the acute stage of
the disease may harbor the organisms in these organs.

It has been cited previously that Pomroy and Fenstermacher

-68-

(12) stated that with experimentally infected turkeys that
the agglutinin response may become negative in two months after
infection unless the organism becomes established.in the viscer-
al organs. It was of interest to note that with these experi-
mental birds, in every instance where the final titer was

l/lOO or over, S. pullorum was isolated from at least one

group of organs. Tw0 birds showing doubtful reactions in 1/25
and 1/50 dilutions in the tube test, yielded the organism. Bird
#2632 (oral) with a doubtful reaction in the 1/25 and 1/50 di-
lutions and slow response to the whole blood antigen, yielded
Sgwpullorum from the liver-spleen, duodenum—pancreas. and an
encysted egg. Bird #2625 had a doubtful reaction in 1/25 but
the organism.was recovered.from the ovaryboviduct. These birds
may have been missed if the agglutination tests alone were taken
into consideration. Bird #2631 had a final titer of 1/50 but
the organism was recovered from the ovary-oviduct, gizZard-
kidney, duodenum-pancreas, mid-gut and cecal stock. This bird
also showed extensive tubercular lesions throughout its organs
which may have lowered its agglutinin response. There were two
birds with. final titers of 1/50. from which the organism was not
recovered.

Only 50 per cent of the control birds, that had been in-
fected orally 2h days previous to autopsy to determine suscep-
tibility. yielded 8. pullorum from the tissues. Two positive
individuals had the organism in the duodenum-pancreas and one
of these still had the organism in the crop. is would be ex-
pected. the bird with the highest titer, yielded the Organism

from the most tissues.

 

iILAJJ. .h Id I

-69..

In comparing the two mediums used in isolation of LIB—“l"
w. some differences were noted in the incidence of recovery.
From Table XXX it will be seen that in several instances no
growth was obtained.with direct seeding of 83 agar plates. and
that the corresponding tissue culture after 15~18 hours' enrich-
ment in tetrathionate broth before seeding yielded S.Jullorum.
There were 20 per cent positive cultures recorded that were en-
tirely lost with direct seeding because of lack of growth. Where
the direct method.yielded one or more pmaxive cultures, in prac-
tically every instance the organism‘was recovered.from additional
tissues with the enrichment medium.

One drawbad: with the tetrathionate enrichment broth was
that organisms of the Proteus group were apt to spread over the
sub-culture plates. thus making the isolation of typical m-
£93112 colonies difficult. To avoid this. only extremely light
streaking of the broth culture was used. This gave satisfactory
results and discrete colonies were obtained.

Posell found (22) that the direct method yielded 3. pullorum
from three birds in three lots; while with enrichment wiflh tetra-
thionate broth.prior to plating, the organisms were recovered

from eight birds in three lots.

 

‘70-

Table m
Comparison of SS agar and tetrathionate broth

Artificial infection of 8. pullorum

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

SS agar Tetrathionate broth
Turkey123u567smrke123h567s
a 2630 ng ng ng as a. .- .. 2630 .. + .- .. + .- .. ..
8 2633 ng 11g ng " '- 1' '- ng 2633 e- e- 0- e-o as so v a
S 2631 0- '— ng 11g 4' es s- 263]. 'f e- + t“! ‘I' e- e- .-
32628++ng-—-- 2628+++»+..--
a 2627 ng ng - mg + ng - 2627 + e4 .. es + .. + ..
2623 N C- c- e- we on v. a 2623 I. on as as I. is e- o
2662 + ~ *I' - 00 *- H v- 3662 on 00 an as n In as one
3} ... - .. e:- a 9 H e- 3 e- s- e- a a s- ee -
23; 2637 D s-o v» e- we e- s c- 2637 to e- ro a set s- fit me
o 2629 ng ng ng mg + ng ng mg 2629 + . + .. + + + -
*2632 ng + 11g 21g ng 11g e- . *2632 so a s- s- + e- n- .-
2635 ng ng '4 P! w n - - 2635 '- ng s- ea 9 as as -
2638 mg mg mg - ng ng - 2638 .- .. + .- .. . .. -
H 2636 ng II II 11g .- .- o 2636 es "en en a o a e- u
8 2621!» mg + ng - + + - 21 2621+ ng + e- .. + + .. ..
8 26 ng ng ng - ng e .. 26? .. .. .. c- - .- .. ..
26 ng - ng . .. .. e- 26 O a .. e. .. + .. .. .
‘Gyst 4-

Va}.- No growl“! .

 

-71-

CCNGIUSIONS

The tube method.was found to be more sensitive and more
consistent in reaction than the whole blood method. The tube
method revealed.positive reactors earlier than the stained anti-
gen. and positives were detected by the tube method that were
sometimes not revealed by the whole blood antigen.

Throughout the eight-month investigation period, marked
zones of inhibition were observed.in the lower dilutions uhere
the serum concentration was greatest. These were not eliminated
by varying the concentration of the antiganér inactivation of
the serum at varying times and intervals. Zone phenomenon may
be explained by increased globulins and lipoid material which
may have interfered.with agglutination in the tubes with the
greatest amount of serum. The data.presented.indicate/ that
a three-tube test, \hich would include a 1/100 dilution, might
detect some positive birds that give weak or doubtful reactions
in the 1/25 or 1/50 dilutions.

‘Lfter intravenous injection of S. pullorum, agglutinins
appeared.on the flaird day in the tube test and.not until the
fifth with the whole blood antigen. The I 1‘7 antigen was a
little more than half as effective in detecting birds of lower
titer or early in the course of infection as was the standard
tube test. The peak titers of intravenously injected and orally
infected birds occurred on about the ninth day, after thich the

titer decreased in fluctuating descent. The orally infected

 

 

 

-72..

birds became negative earlier than did those injected intraven-
ously.
Tetrathionate broth enrichmsnt prior to plating onto 83

agar yields more positive cultures than with direct seeding

o f the plates.

1.

2.

3.

I;

7.

8.

10.

11.

12.

REFERENCE 8

Mallmann. W. 1..: Bacterium Pullorum Studies. Mich. Ag. Exp.
Sta. Tech. aul. _6_§_, 1925.

Rettger. L. F. and Plastridge. W. Na Pullorum Disease of
Domestic Fowl. Storrs Ag. Exp. Sta. Bul. 1E. 1932.

Bettger. L. L. Kirkpatrick. W. N. and Jones. R. 1.: Bacil-
lary White Diarrhea in Young Chicks. Storrs Ag. Exp.
Sta. Bul. g; (Dec.). 1915.

coburn. D. R. and Stafseth. H. Jo: A Field Test for Pul-
lorum Disease. Jour. A. V. M. A. lg: p. 2M. 1931.

Schaffer. Jacob 11.. MacDonald. A. F.. Hall. H. J. and
Bunyea. Hubert: A Stained Antigen for the Rapid
Whole Blood Test for Pullorum Disease. Jour. A. V.

M‘ A. 19;; 2369 1931’

Hewitt. 3. 1.: Bacillary White Diarrhea in Baby Turkeys.
Cornell Vet. $8.3 272, 1928.

Brunett. E. 1..: Pullon Disease in the Mature Turkey,
Poultry Sci. 9; 356. 1930.

Hinehaw. W. 2.: Diseases of Turkeys in the United States:
Proc. of 7th World's Poultry Gong. and lxpos.. cleve-
land. 1939.

Hinshaw, W. 3.. McNeil. 3.. Taylor. J. T.: Four Years'
Progress in Eradication of Pullorum Disease from Tur-
key; Flocks. 146th in. U. S. Livestock San. Assoc. Report.
19 2.

Dickinson. E. M.. Rosenwald. A. 8. and Norrill. D. 3.:
Comparison of the Tube and Rapid Serum Agglutination
Tests for the Detection of Pullorum Disease in Tara
keys. Ore. Agric. Exp. Sta. Tech. 311. _6_. 191m.

Enamel-1, In Do: Pal-10m TCSting 1n mteys, Pan-1t. 3C1.
£31; 208. 19145.
2’ L..}.
Pomroy. B. 8.; Fenstermacher. 3.: Salmonella Infections
A in Turkeys. Am. Jour. Vet. Res. 3; 119. 191$}.

Hinehaw. W. 3.. Jones. E. 3.. Barr. J. R. and Neinmeyar.
V. 3.: comparison of the Tube and Whole Blood Tests
£011;o Pullorum Disease in Turkeys. Cornell Vet. O; 30.
19 .

1’4.

15.

16.

17.

18.

19 .
20.

21.

23.
2h.

25.

26.

-714.

Bunysa. Hubert. Hall. V. A. and Dorset. MA A Sinplified
Agglutination Test for Pullorum Disease. Jour. A. V.

M. .1. 15; nos, 1929.

36th Meeting of the U. S. Livestock San. Assoc.. Jour. A.
Vs Me ‘0 2‘ 1‘88. 1932.

Topley. W. W. 3.: Outline of Immunity. 1933.

Nuttal. George E. L: Blood Immunity and Blood Relation-
Ships. 190,4.

Stafseth. H. J. and Darby. C. W.8 Pullorum Disease. Its
Nature and Its Control. Mich. Ag. Exp. Sta. mart.

Bul. _2_1(Feb.). 191:5.
Boyd. V. C.: Fundamentals of Immunology. 1918.

Gay and Associates: Agents of Disease and Host Resistance.
1935. .

Hallmann. W. In: Salmonella Pullorum in the Intestinal
Contents of Baby Chicks. Jour. Inf. 1313.33; 16. 1929.

Posell. J. J.: Intestinal Qiltures in.Detecting Salmonel-
lozis in Young Turkeys. Am. Jour. Vet. Res. 3; 257.
19 2.

Johnson. D. P. and Anderson, G. v.: Pullorum Disease in
Turkeys. Jour. Inf. Dis. iii; 337. 1936.

Kinshaw. U. 3.: Diseases of Turkeys. Calif. Ag. Exp. Sta.
Tech. 3111. 613. 1937.

Bushnell. L. D. and Porter. J. J.: A Study of Kethods for
Isolation of Salmonella Pullorum. Poult. Sci. 311; 212.

191.5.

Rettger. L. 1.. Kirkpatrick. W. 3.. Stoneburn. 1'. 11.:
Bacillary White Diarrhea in Young Chicks. Storrs Ag.
up. Sta. 1m. 7’1; 153. 1912.

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