Blood Volume Determinations
in Chickens1
GEORGE W. NEWELL AND C. S. SHAFFNER
Poultry Department, University of Maryland, College Park
(Received for publication July 22, 1949)
HISTORY
A S a normal constituent of the body
- of all animals, blood is present in
relatively large quantities as compared
with other body fluids and has been the
subject of considerable research and study.
Because of inadequate laboratory equip-
ment and proper techniques for the meas-
urement of the amount of blood in the
body, early investigators dealt almost en-
tirely with the chemical constituents of
the blood. In fact, it was as late as 1628
that Dr. William Harvey (Dukes, 1947)
was able to demonstrate the circulation of
blood in the body.
The methods of determining blood
volume fall into two general classifica-
tions: (1) the indirect methods, in which
either dyes or colorless solutions are in-
jected into the blood stream and the dilu-
tion of the dye or of the blood determined
by subsequent samples, or in which carbon
monoxide is inhaled and the blood volume
determined by colorimetric determination
of the amount of oxygen displaced; (2)
the direct method, in which the subject
must be sacrificed and the total blood
volume determined by combining the
quantity lost by bleeding with the quan-
tity remaining in the body.
A complete review of the studies of
early investigators was made by Erlanger
(1921). Erlanger reports that Haller is re-
1
Scientific Paper No. A247. Contribution No.
2182 of the Maryland Agricultural Experiment Sta-
tion (Department of Poultry Husbandry).
78
puted to be the first to attempt any de-
terminations of blood volume. Haller's
work consisted of weighing the blood lost
by criminals on the execution block. Ac-
cording to Erlanger, Welcher used small
laboratory animals and not only weighed
blood loss but also combined it with the
blood washed from the tissues. Welcher's
value of 1/13 or 7.7 percent of the body
weight being blood was used as standard
for many years.
The determination of blood volume by
indirect methods seems to have ori-
ginated with Valentin according to Er-
langer. The solids content of the blood
was determined before and after the in-
travenous injection of a known amount of
water. Erlanger further states that the
use of carbon monoxide inhalation and
gasometric determination of its concen-
tration was first suggested and tested
by Grehant and Quinquad. Haldane and
Smith (1899-1900), and others, attempted
to improve the accuracy of the technique
although their modifications made the
technique difficult and hard to apply.
Smith, Arnold, and Whipple (1920), in
testing this method against other tech-
niques, found it very reliable for deter-
mining red cell volume.
Keith, Rowntree, and Geraghty (1915)
injected a vital dye (vital red) into the
blood stream and determined colori-
metrically its concentration in subsequent
samples of blood. They found only slight
variations between blood volumes of
normal subjects when considered on a
 BLOOD VOLUME DETERMINATIONS IN CHICKENS
body weight basis. Dawson, Evans, and
Whipple (1920) tested some 60 different
dyes, including vital red, covering the
entire color range and found two blue dyes
(T-1824 and T-1835, alkaline) more satis-
factory than vital red. Their reasons for
suggesting the use of the blue dyes are:
(1) they leave the circulation more
slowly than vital red; (2) the blue color
could be read much more accurately on
the type of colorimeter they used; (3)
the blue color does not obscure hemolysis.
Gibson and Evans (1937), using a spectro-
photometer, concur in the use of Evans
Blue (T-1824).
Gregersen (1944) and Nitshe and
Cohen (1947) simplified the technique by
demonstrating that only one sampling of
the blood subsequent to the injection of
the dye need be made. This they took
exactly 10 minutes after the injection.
Gregersen in his study further simplified
the technique by injecting a standardized
amount of dye from an ampule. In this
way, calculation of plasma volume was
eliminated and was determined by ref-
erence to a table of densities predeter-
mined from various dilutions of the dye in
plasma. Nitshe and Cohen injected a
known amount of dye from a carefully
calibrated syringe and calculated the
plasma volume by use of a constant called
K. The value of K was constant for a
particular stock solution of dye. The con-
stant was calculated in order that samples
of plasma containing dye could be com-
pared spectrophotometrically with un-
dyed samples of plasma.
Numerous reports of the blood volume
of normal humans are to be found in the
literature. These coupled with reports of
studies on pathologic conditions and sub-
jects under trauma and shock give a fairly
complete picture of the blood volume of
humans. Reports on many of the labora-
tory animals, such as horses, goats, dogs,
79
and rats are also found in the literature of
the past 20 years. However, very few re-
ports of any work of this nature on chick-
ens can be found.
Welcker (1903) in the course of his
studies on the blood volume of a great
number of species of animals, studied
some species of fowls. He reported that
the value for blood weight as a percent of
body weight for the Gallus Bankiva
Domesticus was 3.68. This was based on a
study of only one individual. His values
for the blood of two geese were 5.71 and
6.75 percent of the body weight. He
studied the quantity of blood in several of
the species of wild birds and reported
values varying from 3.43 to 9.35 percent
of the body weight for small and large
birds, respectively.
Latimer (1923) reports on a study by
Zaitschek who weighed the blood lost by
131 chickens and found a loss of 3.8 per-
cent of the body weight when the jugular
vein was severed. The close agreement of
this value with the values reported by
Welcker leads to the conclusion that in
the early studies of Welcker only blood
loss was recorded.
Magnan (1912) studied the blood vol-
ume of several species of wild fowl, using
the original Welcker technique. He deter-
mined the weight of blood lost when the
carotid artery was cut and to this added
the blood obtained by flushing the cir-
culatory system with a 0.7 percent saline
solution. He obtained values for the total
weight of blood ranging from 39 to 86
grams per kilogram of body weight. The
larger values were not for heavier birds
but rather for birds whose natural habitat
was near the sea and which were also good
divers.
Pappenheimer, Goettsch, and Jungherr
(1939) used a modification of the dye tech-
nique for the determination of the blood
volume in chicks ranging from 1-50 days
80 GEORGE W. NEWELL AND C. S. SHAFFNER
of age. Their results indicate a close rela-
tionship between body growth and blood
volume. They conclude that the blood
volume in chickens of this age is approxi-
mately 9.0 percent of the body weight.
Their results also show that the average
cell volume percentage of their chicks not
separated as to sex is 33.0 ±4.65 percent.
PROCEDURE
In this study the blood volume was
determined for a group of New Hamp-
shire chickens ranging in age from six
weeks to adult stock. The young chicks
were all battery raised. The hens were
maintained in batteries and were all in
laying condition. The adult male birds
were from the laying pens on the Univer-
sity poultry farm.
All blood volume determinations were
made by the dye method, using Evans
Blue (T-1824) dye. The technique fol-
lowed was that of Nitsche and Cohen
(1947), modified as described later. The
left wing vein was used for the with-
drawal of the undyed sample and the in-
jection of approximately 0.3 mg. of dye
per kilogram of body weight. The right
wing vein was used for the withdrawal of
the sample of blood containing the dye
after 10 minutes had elapsed for the mix-
ing of the dye in the blood stream.
Most of the methods of blood volume
determinations require the withdrawal of
five to ten ml. of blood for the hematocrit
and plasma volume determinations. Be-
cause of the small size of the chicken, and
because of the necessity of having at least
seven ml. of plasma in the absorption cell
of the spectrophotometer, it was decided
to modify the technique slightly. This
modification consisted of the withdrawal
of only three ml. of blood. One ml. was
used for the hematocrit and two ml. was
diluted with eight ml. of 0.9 percent
saline. The dyed and undyed samples of
blood were both processed in the same
manner to insure that the difference in
optical density would be only due to the
amount of dye present.
Using this sample of dyed blood, the
percentage of red and white blood cells
was determined. This was accomplished
by using Wintrobe hematocrit tubes.
Both the hematocrit tubes and the dilute
sample of dyed blood were centrifuged for
30 minutes at approximately 3000 rpm.
The percentages of red and white cells
were added together and recorded as cor-
puscular volume.
Nitshe and Cohen (1947) stated that
the light absorption by the dyed as well
as the undyed sample was best determined
using a 620 millimicron filter in the spec-
trophotometer. The spectrophotometer
available for use in this study did not have
a 620 but did have a 610 millimicron
filter. Examination of the light absorption
curve for Evans Blue, as presented by
Gibson and Evans (1937), showed that
the effect of this 10 millimicron difference
in wave length is less than 0.01 percent of
optical density.
After both dyed and undyed samples
have been centrifuged and the optical
density of the dyed sample as compared
to the undyed sample has been deter-
mined, the plasma volume (PV) can be
determined by using the following for-
mula:
in which M = No. of mg. of dye in-
jected, K = Constant, whose value is
determined by dividing the density
of different concentrations of dye in
plasma by the concentrations used, and
D = Density reading of the dyed sample
from the spectrophotometer.
Because two cc. of blood, a part of
which is corpuscles, has been added to
 BLOOD VOLUME DETERMINATIONS IN CHICKENS
eight cc. of saline solution, thus diluting
the dye, this PV will be approximately
five times too large. The exact amount by
which it is too large (Plasma Volume Cor-
rection Factor, PVCF) can be deter-
mined by the following formula:
%PVX2
PCVF =
8+(%PVX2)'
in which % PV is the percent plasma
volume from the hematocrit reading.
The value thus obtained is multiplied
by the PV of the previous formula in
order to determine the True Plasma
Volume (TPV):TPV = PVXPVCF.
The Total Blood Volume (TBV) is
calculated from the formula:
TPV
T V B = — — X100,
in which P is the corrected volume of
plasma per 100 ml. of blood calculated
as follows:
P=100
r Volume of cells
[.Volume of cells and plasma xioo
Thus, the total blood volume is re-
ported in ml. per animal, in which form it
may be related to body weight, surface
area, or any other measure desired.
RESULTS AND DISCUSSION
Before proceeding with the use of the
technique on a large number of chickens,
it was decided to test it by determining
the volume of measured quantities of
blood contained in beakers. Further
tests for accuracy were made by deter-
mining the total blood volume of several
birds and six or seven days later redeter-
mining the blood volume of these same
birds.
The results of the in vitro tests are
shown in Table 1. The samples of blood
were measured in a graduate cylinder
81
and poured into a beaker. The same syr-
inge was used for adding the Evans Blue
to the sample as was used for the later
injections into the chickens. The values
reported are not only the result of dupli-
cate samples but also from different
samples containing the same amount of
blood.
TABLE 1.—Blood volume determinations by the dye
method on measured samples of blood in vitro
No. of Actual Average
calculated
samples volume cc. volume cc.
50 49.1±2.7
60 59.4+3.6
70 70.4±1.2
100 99.5±1.7
It will be noted from Table 1 that the
blood volume as determined by the spec-
trophotometer was very close to the meas-
ured volume. There is some variation
present in the readings, a part of which
may be due to the quantity of blood
clinging to the sides of the graduate
cylinder. The coefficient of correlation
between measured and calculated blood
volumes was 0.99. This high correlation
seemed ample justification for using the
technique as described.
For the in vivo tests, 20 birds (8 males
and 12 females) varying in weight from
900 to 3600 grams were used. The blood
volume was determined twice on these
birds with six or seven days elapsing be-
tween determinations. This lapse of time
was allowed in order that all of the dye
would be lost from the blood stream, al-
though Gibson and Evans (1937) showed
that repeated as well as continuous deter-
minations can be made using the dye
method. Also it was desired that the
chickens would completely recover from
the withdrawal of blood in the previous
determination. The result of these two
determinations shows a coefficient of corre-
82 GEORGE W. NEWELL AND C. S. SHAFENER
lation of 0.93, indicating a close agree-
ment between the two values obtained.
Approximately one-half of the birds
showed a decrease in blood volume and
the rest of them showed a slight increase
in blood volume at the time of the second
determination.
Total Blood Volume—Females—The
blood volume of 113 normal New Hamp-
BL00D
VOLUME
CC.
150
125
Dotted lines Indicate limits
of standard error of the
estimate of Y
0 300 600 900
FIG. 1. The relationship between total blood volume and body weight of normal New Hampshire females.
shire females was determined. Inspection
of a scatter diagram of blood volume
plotted against body weight indicated
that a straight line could not be used to
express the data correctly. It was found by
calculation and plotting the values, that a
second degree parabola type of curve best
fitted the scatter diagram. The equation
for this type of curve is Y = a.-}-bX-\-cX2
(Snedecor 1946). The regression curve
showing the relation between the values
for blood volume and body weight of the
females is shown in Figure 1. The values
for the constants used in calculating the
points on this line are: a =—6.2; b
= 0.11973; c=-0.00002. The standard
error for the estimate of Y is 14.3 c c ,
the limits of which are indicated on Fig-
ure 1 by the broken lines.
The blood volume of females increases
1200 1500 1800 21flO 21i00 2700 3000
BODY WEIGHT IN GR11B
in almost straight line relationship to
body weight between 400 and 1200 grams,
increasing from 44 cc. to 108 cc. Above
1200 grams the relationship levels off and
additional increments of weight are ac-
companied by smaller increments of blood
than at the lower weights.
Individual variation is smaller at
weights below 1200 grams than at heavier
weights. This is undoubtedly caused by the
type of tissue, involved in the weight in-
 BLOOD VOLUME DETERMINATIONS IN CHICKENS
crease. Adipose tissue is low in blood con-
tent and any weight increase due to fat
would not be accompanied by as great an
increase in blood volume as though the
increase were due to general body growth.
Thus, the individual variation in age at
which fat is deposited, the variation in
fat deposition, and the increase in fat
content of the body at heavier weight
BL00D
V0LWE
CC.
360
300
2U0
ISO
1 - a + bX + cX'
a = 1.58
b = 0.10138
c =-0.0000025
60
boo 800 i6o6
BODY VIEIGHT IN GRAMS
FIG. 2. The relationship between total blood volume and body weight of normal New Hampshire males.
would all tend to increase individual vari-
ation in blood volume as well as decrease
the amount of blood volume per unit of
body weight.
The curve calculated from the values
for the blood volume of females indicates
that at weights above 2990 grams the
blood volume would decrease. Thus it
seems obvious that this curve is not ac-
curate for weights above this point. What
is probably true is that in females of
heavier weights the increase in blood
83
volume is so small in relation to body
weight that the formula must be recalcu-
lated and the curve redrawn. Although
this curve seems to hold for the birds
studied, it is not known at this time
whether it will hold true for breeds
which mature at different weights than
New Hampshires.
Total Blood Volume—Males—The
Doited lines Indicate limits
of standard error of the
estimate of T
2000 2u6o SSST "Tste- Toro
blood volume of 110 New Hampshire
males was determined and inspection of a
scatter diagram of the blood volume
plotted against body weight led to the
conclusion that the same formula as used
for the females should be used to express
the relationship. Figure 2 shows the re-
gression line calculated by this formula
and illustrates the relationship between
blood volume and body weight of males.
The values for the constants are: a = 1.98;
6 = 0.10138; c= -0.0000025. The stand-
84 GEORGE W. NEWELL AND C. S. SHAFFNER
ard error for the estimate of Y is 23.5, the
limits of which are shown by the broken
lines in Figure 2.
Although the relationship between
blood volume and body weight in males
is curvilinear, the curve in Figure 2 ap-
pears to be nearly straight, with the blood
volume ranging from 48 cc. to 365 cc.
put
CENT
60
50
1*
30
Uoo 800
FIG. 3. The relationship between corpuscular volume as a percentage of total blood volume
and body weight of normal New Hampshire chickens.
for birds of 300 and 4000 grams respec-
tively. This nearly straight line relation-
ship differs considerably from the rela-
tionship for the females, apparently be-
cause the males do not deposit nearly as
much fat in the body as do the females.
Individual variation in blood volume is
greater in magnitude in males than in fe-
males as indicated by the values for the
standard error of the estimate of Y, which
are 23.5 and 13.4 respectively. However,
when the larger volume of blood in the
males is considered, the coefficient of
variability is approximately the same for
both sexes.
Corpuscular Volume—Females—As
shown in Figure 3, the red and white cells
comprise slightly less than 30 percent of
the total blood of the females. This per-
centage remains almost constant through-
MOM +
1600 2000
BODY WEIGHT IK QRAtG
2l|00 3200 3600 lioob
out the weight range studied and is not
influenced by either weight or age. The
birds varied from young chicks to adult
laying stock. This observation is in ac-
cord with that of Juhn and Domm (1930),
who found very little change in erythro-
cyte count in Brown Leghorn females
from hatching time to 490 days of age.
Although blood comprises a smaller per-
cent of the body weight in the heavier
weights studied, the corpuscular volume
of the blood remains the same and any in-
 BLOOD VOLUME DETERMINATIONS IN CHICKENS 85

crease in blood volume is an increase in cockerels ranging in weight from 1000 to

corpuscular volume as well as in plasma 1300 grams were selected. Each of eight
volume. birds received a 10 mg. pellet of testos-
Corpuscular Volume—Males—The re- terone propionate implanted subcutane-
lationship between corpuscular volume, ously in the neck. Seven males received
as a percentage of total blood volume, implants of 15 mg. pellets of diethylstil-
and body weight in males is shown in bestrol. The remaining seven males were
Figure 3. It will be seen that at weights untreated. Hematocrit readings on the
below 1800 grams the corpuscular volume blood of these birds were taken at the
TABLE 2.—The eject of implanting male and female sex hormones
on corpuscular volume and comb area

Mean corpuscular volume* Mean comb area

No.
Treatment birds Before 5 wks. after Before 5 wks. after
implanting implanting implanting implanting
Testosterone propionate 8 28.6 34.4±0.65** 9.96 26.45**
Untreated 7 28.6 31.2±0.83 9.20 17.53
Diethylstilbestrol 7 28.6 28.0± 1.17** 9.92 5.66**

* cc. red and white blood corpuscles per 100 cc. of blood.
** The differences between these values and that of the untreated are highly significant statistically to the
1 percent level.

of males and females is approximately
the same. Between this weight and 2800
grams the corpuscular volume in males in-
creases until it is almost fifty percent
greater than at the lighter weights.
Above 2800 grams the curve again levels
off and rises only slightly throughout the
rest of the weight range studied. It would
appear from this that a part of the in-
crease in blood volume of males over fe-
males is due to this increase in corpuscular
volume.
The study of Juhn and Domm (1930)
showed that the erythrocyte count in
males increased considerably at the time
of sexual maturity. The increase found in
the present study is no doubt as much
affected by age*;as by weight; however
the exact age of the males was not known
and must be deduced from body weight.
To further investigate the relationship
between androgen level and corpuscular
volume, a later test was conducted using
testosterone propionate and diethylstil-
bestrol implants. Twenty-two young
start of the test and again five weeks after
the pellets were implanted. The means
of these hematocrit readings as well as the
means of the comb areas taken at the
same time are shown in Table 2.
The implants of diethylstilbestrol were
effective in reducing comb area but did
not change the corpuscular volume during
the test period. The changes in comb area
and in corpuscular volume of the un-
treated males are those which would
occur normally during this five week
period. However, the comb area of the
males receiving the implants of testoster-
one propionate pellets increased consid-
erably more than normal indicating a
higher androgen level in the blood of these
males. Since the corpuscular volume in-
creased in the males receiving the male
sex hormone and there was no change in
the males receiving the female sex hor-
mone, it is evident that the corpuscular
volume increased as a result of the
higher androgen level in the blood. This
conclusion confirms the hypothesis made
86 GEORGE W. NEWELL AND C. S. SHAITNEB.
earlier on the effect of sexual maturity on
corpuscular volume in males and further
corroborates the findings of Juhn and
Domm on the effect of age on the amount
of erythrocytes in the blood. These results
are also in agreement with those reported
by Taber, Davis, and Domm (1943) and
Domm and Taber (1946).
Blood Volume of Thiouracil and Thy-
roprotein Treated Chickens—There was
available at the time of the blood volume
study a number of chickens undergoing a
test on a diet containing thiouracil and
thyroprotein. This treatment was being
tested by this laboratory as a means of
increasing fat deposition. At the time of
the blood volume determinations these
birds were 13 weeks old and for the past
five weeks had received a ration contain-
ing 0.2 percent thiouracil plus one gram
of thyroprotein per hundredweight. Tur-
ner (1948), using the blood loss when
killed as a measure of blood volume,
found no cumulative effect on blood
volume as a result of feeding thyroprotein
to two-year old Leghorn hens. His results
also showed, however, that there was little
change in the body fat of these hens.
The blood volume of a representative
sample of the chickens on the above diet
was determined, using the technique as
presented in this paper, to determine
what effect this treatment might have on
blood volume. The blood volume of nor-
mal chickens of the same sex and weight
was determined by interpolation from the
curves in Figures 1 and 2. The blood vol-
ume of 12 treated males was found to be
17.5 + 2.2 cc. less than that of normal
males of comparable size. The blood vol-
ume of 11 treated females was found to
be 31.6 + 3.5 cc. less than normal females
of comparable weights. These differences
in blood volume are highly significant
statistically. These results indicate that
this treatment causes a greater propor-
tion of the body to be fatty tissue, which
is not high in blood capacity. This is in
accordance with the general observations
on the effect of feeding thiouracil.
SUMMARY AND CONCLUSIONS
A procedure is presented for determin-
ing the blood volume of chickens. This is
a modification of the earlier techniques in
that only three cc. of blood rather than
five or ten cc. need be drawn from the
bird. One cc. of this blood was used for
the hematocrit determination and the
remaining two cc. were diluted with eight
cc. of 0.9 percent saline solution. By use
of the spectrophotometer the optical
density of the solution can be deter-
mined, and by use of the formulae pre-
sented the plasma volume and total blood
volume can be calculated.
Data are presented in support of this
technique in which the calculated volumes
of in vitro samples are in close agreement
with the measured volumes. Also pre-
sented are the results of repeated blood
volume determinations on the same
chicken, which show close agreement be-
tween the two determinations.
Regression curves are given for the
values of blood volume and body weight
as determined for 113 New Hampshire
females and for 110 males of the same
breed. These curves indicate a curvilinear
relationship between blood volume and
body weight, with the blood volume being
approximately 10 percent of the body
weight in young chickens of either sex.
Above 800 grams however the curves
differ and show that a sexual dimorphism
exists in blood volume. The curve for
females levels off, while the curve for
males continues in almost straight line
positive relationship throughout the
weights encountered in this study.
The volume of red and white blood
cells as a percentage of total blood volume
 BLOOD VOLUME DETERMINATIONS IN CHICKENS
remains almost constant in females
throughout the weight range of this study.
In males, this percentage increases
rapidly at weights above sexual maturity,
attaining a value 50 percent greater than
the value for young chickens.
Birds treated with thiouracil had sig-
nificantly less blood than control birds of
an equal weight indicating that weight
increases which result from this treatment
are due to the deposition of fatty tissue
which is low in blood content.
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