Physiological parameters in broiler lines divergently selected

for the incidence of ascites

S. Druyan,*1,2 D. Shinder,† A. Shlosberg,‡ A. Cahaner,* and S. Yahav†

*The Hebrew University, Faculty of Agricultural, Food and Environmental Quality Sciences, PO Box 12, Rehovot
76100, Israel; †Institute of Animal Science, Agricultural Research Organization, The Volcani Center, PO Box 6,
Bet Dagan 50250, Israel; and ‡Kimron Veterinary Institute, PO Box 12, Bet Dagan 50250, Israel

ABSTRACT Ascites syndrome (AS) is manifested in
flocks of contemporary broilers that are allowed to fully
manifest their genetic potential for rapid growth. After
successful selection, a pair of divergent lines was estab-
lished, AS-susceptible (AS-S) and AS-resistant (AS-R).
These lines facilitate comparisons between genetically
resistant and susceptible healthy young broilers when
reared under standard brooding conditions (SBC). The
aim of the present study was to look for predictive in-
dicators for AS susceptibility by comparing relevant
physiological parameters in the AS-S and AS-R lines
under SBC and after exposure to extreme ascites-in-
ducing conditions (AIC). In this design, a trait differing
significantly between the 2 lines under SBC is expected
to be a reliable indicator for selection against AS sus-
ceptibility in breeding stocks when reared under non-
inducing conditions. Males from the AS-S and AS-R
lines were reared together under SBC to 19 d of age,
then under the AIC protocol. Cumulative incidence of
AS mortality was 93.2% in the AS-S line and only 9%
Key words: ascites, broiler, selected line, heart rate, blood parameter
in the AS-R line, confirming the genetic divergence be-
tween the lines. Exposure to AIC enhanced the imbal-
ance between oxygen demands and supply in the AS-S
birds and induced differences in blood parameter level
between the 2 lines. The AS-S birds exhibited elevated
hematocrit and red blood cell counts and a decline in
oxygen saturation in the arterial blood. No difference in
hemoglobin concentration was found, but calculation of
hemoglobin content per 1,000 red blood cells revealed a
significant reduction in hemoglobin content in the AS-S
birds. Under SBC, there were no significant differences
between the lines for hematocrit, red blood cell count,
hemoglobin concentration, hemoglobin count per 1,000
red cells, and blood oxygen saturation. However, heart
rate during the first week of life was significantly higher
in the AS-S birds than in the AS-R birds on d 1 and 7,
suggesting that high heart rate may potentially serve
as an early criterion for selection against AS suscepti-
bility.
2009 Poultry Science 88:1984–1990
doi:10.3382/ps.2009-00116

INTRODUCTION Since the early 1990s, AS has been a major concern

The general pathogenesis of the metabolic disorder
ascites syndrome (AS) has been well documented (Max-
well et al., 1992; Julian, 1993; Shlosberg et al., 1998;
Currie, 1999; Wideman, 2001; Balog et al., 2003). It is
characterized by a cascade of events that begins when
the oxygen demands for maintenance and growth are
not fully supplied by the cardiovascular system of those
broilers that eventually develop AS (Julian, 2000).
©2009 Poultry Science Association Inc.
Received March 8, 2009.
Accepted May 11, 2009.
1
Current address: Institute of Animal Science, Agricultural Re-
search Organization, The Volcani Center, PO Box 6, Bet Dagan 50250,
Israel.
2
Corresponding author: druyan@agri.huji.ac.il
for the broiler industry worldwide. Ascites syndrome is
observed in flocks of contemporary commercial broilers
that are allowed to fully manifest their genetic poten-
tial for rapid growth (Druyan et al., 2008). Mortality
due to AS may reach 25% in flocks reared under natu-
rally cold conditions (Balog et al., 2003) and up to
50% under applied acute cold exposure (Druyan et al.,
2007b). However, even 1% of AS mortality causes sig-
nificant economic losses to the broiler industry, mainly
because it occurs toward the end of the growing period
(Maxwell and Robertson, 1997).
Successful divergent selection for AS was applied by
Druyan et al. (2007a), whereby 2 divergent lines were
established, AS-susceptible (AS-S) and AS-resistant
(AS-R), with >95% and <5% AS mortality, respec-
tively, when reared together under the same high-chal-
lenge protocol of ascites-inducing conditions (AIC;

1984
 ASCITES-RESISTANT VERSUS ASCITES-SUSCEPTIBLE BROILER LINES
Druyan et al., 2007b). Research with such lines per-
mits monitoring of differences between AS-R and AS-S
healthy young broilers reared under standard brooding
conditions (SBC).
Several well-known parameters have been used to
predict or monitor AS. Hematocrit was found to be
one of the most reliable indicators for the detection of
AS (Julian and Mirsalimi, 1992; Maxwell et al., 1992;
Lubritz and McPherson, 1994; Shlosberg et al., 1996;
Wideman et al., 1998). Despite the high hematocrit
level, broilers with cold-induced AS exhibited signifi-
cantly lower levels of oxygen saturation of the arterial
blood (SaO2) than their healthy counterparts (Julian
and Mirsalimi, 1992; Druyan et al., 2007b). Broilers
developing AS were characterized by lower heart rate
(Kirby et al., 1997; Olkowski et al., 1997, 2005; Wide-
man et al., 1998) and higher right ventricle weight to
total ventricle weight ratio (RV:TV; Wideman and
Kirby, 1995a,b; Druyan et al., 2007b). However, in all
these studies, differences were found only between broil-
ers that had already developed some degree of AS when
compared with their healthy counterparts. Thus, these
differences were probably secondary to the manifesta-
tions of AS and hence can serve as diagnostic tools but
not for early prediction.
To serve as a predictor of broiler susceptibility to AS,
a potential indicator should differ significantly between
AS-S vs. AS-R individuals when they are reared to-
gether under standard conditions at an early age, when
all birds are healthy. Such an indicator is essential for
indirect phenotypic selection against AS susceptibility.
Scheele et al. (2003, 2005) found that high partial pres-
sure of CO2 tension in venous blood measured at d 11
was a reliable predictor for ascites susceptibility of the
same birds as observed at 5 wk of age. However, the
heritable nature of this association has not been tested
yet. The aim of the present study was to look for pre-
dicting indicators for AS susceptibility by comparing
candidate physiological parameters monitored in the
genetically divergent AS-S and AS-R lines, before and
after exposure to AIC.
MATERIALS AND METHODS
Genetic Lines
Chicks were produced from the divergently selected
AS-S and AS-R lines (Druyan et al., 2007a), in 3 repli-
cated trials, by mating 7 S3 sires to 35 non-sib S3 dams
in the AS-S line and 8 S3 sires to 40 non-sib S3 dams
in the AS-R line. The numbers of chicks in the 3 trials
were 59, 63, and 82 in the AS-S line and 60, 54, and 42
in the AS-R line, respectively.
Experimental High-Challenge AIC Protocol
In each replicated trial, the chicks from both lines
were brooded together under SBC on a concrete floor
1985
covered with wood shavings. At 19 d of age (d 19),
the chicks were placed in individual cages with cool air
(18 to 20°C) blown over them by fans at about 2 m/s.
Growth rate of these chicks was enhanced by using an
accelerated feeding program, comprising prestarter feed
from d 0 to 4 (instead of d 0 to 10), starter feed from
d 5 to 14 (instead of d 11 to 21), grower feed from d
15 to 24 (instead of d 22 to 33), and a high-energy pel-
leted finisher feed from d 25 to 44. The contents of CP
(%) and energy (cal/kg of ME) in these 4 diets were as
follows: 22 and 3,100, 20.5 and 3,125, 19.5 and 3,150,
and 18.3 and 3,225, respectively. Feed and water were
provided ad libitum. Growth rate was further enhanced
by exposure to 23 h/d of light from hatch to the end
of each trial, on d 44, as described by Druyan et al.
(2007b).
Diagnosis of AS
Throughout the phase of AIC, all dead birds were
necropsied and examined to determine the cause of
death. Birds with abdominal ascetic fluid or hydroperi-
cardium were diagnosed as having died due to AS and
were recorded as being AS-S. The few birds that died
from other causes were excluded from the data analy-
ses. At the end of each trial, all surviving birds were
killed by cervical dislocation and necropsied. Those
with abdominal ascetic fluid or hydropericardium were
recorded as being AS-S and all other birds were noted
as being AS-R.
Measurements
Growth and Blood Parameters. Individual BW was
measured once a week and on d 19 when the chicks
were exposed to AIC.
Blood for hematocrit measurement was drawn into
heparinized microcapillary tubes and centrifuged in a
microliter centrifuge (Hettich, Tuttlingen, Germany)
for 7 min. Hemoglobin concentration was determined
colorimetrically as described by Rice (1967) using Sig-
ma diagnostic kit no. 525 (Sigma Chemical Co., St.
Louis, MO) according to the instructions of the manu-
facturer. Blood cells were counted with a Cell Coulter
Counter Model ZM (Coulter Electronics, Luton, UK)
after dilution of whole blood samples (10 µL) to a con-
centration of 5 × 10−4. Hemoglobin concentration (g/
dL) was divided by several red blood cells (1 × 1010/
mL), to determine the hemoglobin quantity per 1,000
red blood cells (ng/1,000 cells).
Heart Rate and SaO2. Percentage saturation of he-
moglobin with O2 in arterial blood and heart rate were
measured using a portable veterinary oximeter (8600V,
Nonin Medical Inc., Plymouth, MN) and a sensor
(2000T, Nonin Medical Inc.). The sensor was placed
over the cephalic vein of the right wing of lightly re-
strained birds held in lateral recumbency; SaO2 and
heart rate were determined simultaneously, usually
1986 Druyan et al.

within 1 min. In the first week of life, a Nonin lingual RESULTS

sensor (2000SL, Nonin Medical Inc.) was used, posi-
tioned on the medial aspect of the thigh.
RV:TV. Hearts were collected from all chicks, those
that died during the trials and those killed at the end
of the trials, and were dissected to record the weight of
the right ventricle and of the left ventricle + septum
(total ventricle). The RV:TV ratio was calculated for
each bird.
Statistical Analysis
In the AS-S line, data of the few healthy individuals
(15 of 205) were excluded from the following analy-
ses. Similarly, data of the few ascitic individuals in the
AS-R line (14 of 155) were excluded.
Data were subjected to ANOVA according to the fol-
lowing mixed model:
Y = μ + Line + Trial + Sire/Line
+ Trial × Line + e,
with line (AS-S vs. AS-R) as the main fixed effect
and trial (1, 2, or 3) and sire as random effects nested
within line. Interaction between trial and line was also
included.
No significant differences between trials and inter-
action between trial and line were found; results are
exhibited for the line least squares means and include
all 3 trials together.
Data of blood measurements on d 18 (last day un-
der SBC) and d 25 (6 d after exposure to AIC) were
compared within each line and were analyzed by t-test
(with unequal variances). All of the statistical analyses
were conducted using the JMP software (SAS Institute,
2005).
Incidence of AS and RV:TV
The cumulative incidence of AS was 93.2% in the
AS-S line and only 9% in the AS-R line (Table 1), con-
firming the almost complete genetic divergence between
the 2 selected lines. Data of the few resistant individuals
(15 of 205) in the AS-S line and of the few susceptible
individuals in the AS-R line (14 of 155) were excluded
from the analyses.
By d 28, nine days after the instigation of the AIC
protocol, only 13 (6.3%) AS-S birds had died from AS
(Table 1). However, during the following week (d 28
to 35 of age), substantial AS mortality was observed
in the AS-S line, reaching 26.8%, whereas only 2 birds
(1.3%) died from AS in the AS-R line. On d 44, the cu-
mulative AS mortality reached 65.9% in the AS-S line
and 7.7% in the AS-R line. On d 44, all surviving birds
were necropsied and visually examined for manifesta-
tions of AS morbidity. Fifty-six birds (27.3%) with AS
manifestations were found in the AS-S line and only 2
(1.3%) in the AS-R line (Table 1).
In the AS-S line mean RV:TV ratio was significantly
higher than in the AS-R line, being 0.361 vs. 0.235,
respectively (Table 1).
BW
Table 2 presents mean BW at several ages in the
AS-S and AS-R lines. The initial number of chicks in
each line is also presented (n); in the AS-S line, these
numbers decreased from d 19 to later ages due to AS
mortality under AIC and remained unchanged in the
AS-R line.
At hatch, chicks from the AS-R line were about 4%
heavier than those from the AS-S line (52.3 vs. 50.2 g).

Table 1. Cumulative mortality and morbidity due to the ascites syndrome (AS) at various ages in
the ascites-susceptible (AS-S) and ascites-resistant (AS-R) lines were selected, all kept together under
ascites-inducing conditions from d 19
AS-S line AS-R line
1 2
Age (d) n (N = 205) % n (N = 155) %

Mortality (cumulative)
  28 13 6.3a 0 0b
  35 55 26.8a 2 1.3b
  44 135 65.9a 12 7.7b
Morbidity3
  44 56 27.3a 2 1.3b
Total incidence of AS 191 93.2a 14 9.0b
RV:TV ratio4 0.361a 0.235b
a,b
Percentages within rows (age) without common superscript differ significantly (χ2 test, P < 0.05).
1
n = number of birds with AS.
2
N = total number of birds in the line.
3
Birds that survived to the end of the trial but were diagnosed with manifestations of AS after being killed.
4
RV:TV = right ventricle weight to total ventricle weight; RV:TV was determined on day of death for chicks
that died during the trial and on d 44 for the surviving chick.
 ASCITES-RESISTANT VERSUS ASCITES-SUSCEPTIBLE BROILER LINES 1987
Table 2. Mean and SE of BW measured during standard brooding conditions (SBC, from day of
hatch to d 19) and under high-challenge ascites-inducing conditions (AIC, from d 19 to the end of the
trial on d 44) in chicks from the ascites-susceptible (AS-S) and ascites-resistant (AS-R) lines
Age (d) AS-S (n = 205) AS-R (n = 155) (AS-S − AS-R), g (AS-S − AS-R)/AS-R, %

SBC: d 0 to 19
 0 50.2 ± 0.7 52.3 ± 0.7 −2.1 −3.9§
 7 147.6 ± 2.3 143.5 ± 2.2 4.2 2.9
  14 352.0 ± 6.3 333.9 ± 6.2 18.0 5.4§
  19 584.7 ± 9.8 553.8 ± 10.1 30.9 5.6*
AIC: d 19 to 44
  21 682.8 ± 11.7 653.1 ± 11.7 29.7 4.5
  28 1,018.1 ± 18.6 1,038.0 ± 18.8 −19.9 −1.9
  35 1,355.0 ± 26.3 1,529.3 ± 25.2 −174.3 −11.4***
  42 1,668.7 ± 39.5 2,008.4 ± 25.6 −339.7 −16.9***

§, *, ***Significant differences at the 0.07, 0.05, and 0.001 level of significance, respectively, between AS-S and
AS-R lines.

However, under SBC, the AS-S chicks exhibited signifi-
cantly greater BW than that of the AS-R chicks on d 14
(352.0 vs. 333.9 g) and on d 19 (584.7 vs. 553.8 g; Table
1). Under AIC, most AS-S birds developed AS and, as
expected, exhibited lower BW gain than their healthier
AS-R counterparts. Consequently, AS-S BW was lower
by 11.4% at d 35 and by 16.9% at d 42 in comparison
to the AS-R line (Table 2).
Heart Rate
During the first week of life, under SBC, the AS-S
birds exhibited a significantly higher mean heart rate
than the AS-R birds with a difference of 7.8% on d 1
and 9.8% on d 7 (Table 3). On d 17, there was a re-
duction in the heart rate of AS-S birds and both lines
exhibited similar rates (419.3 vs. 416.9 beats/min). Af-
ter the exposure to AIC, mean heart rate of both lines
decreased by 40 beats/min (378.6 vs. 374.1 beats/min;
Table 3).
75.4% on d 27, apparently due to the development of
AS in the birds of this line. In the healthy AS-R birds,
mean SaO2 remained at a high level (above 90%) and
was significantly higher than that recorded in the AS-S
broilers (Table 3).
Hematocrit
No difference was found between AS-S and AS-R
birds in mean hematocrit values under SBC up to 18
d of age (Table 4). The exposure to AIC increased the
hematocrit level in the AS-S birds, which exhibited sig-
nificantly higher values on d 25 compared with the level
measured on d 18 under SBC. A less-pronounced in-
crease in hematocrit level was found in the AS-R birds
between d 18 to 25 (Table 4). Subsequently, the AS-S
and AS-R lines differed significantly in mean hemat-
ocrit level during the entire AIC phase (Table 4), with
the difference between the 2 lines increasing from 6%
on d 25 to 29.6% on d 41.

SaO2
The AS-S and the AS-R lines exhibited similar means
of SaO2 on d 1, 7, and 17 under SBC (Table 3). After
the exposure to the challenging AIC protocol, mean
SaO2 of the AS-S line decreased from 92.7% on d 17 to
Hemoglobin
Hemoglobin concentration in the whole blood tended
to have similar levels in the chicks from the 2 lines when
reared under SBC. Exposure to AIC slightly increased
the hemoglobin concentration in both lines. Despite the

Table 3. Means and SE of heart rate (beats/min) and oxygen saturation of hemoglobin in arterial blood (SaO2) measured during
standard brooding conditions (SBC, from day of hatch to d 19) and under ascites-inducing conditions (AIC, from d 19 to end of trial
on d 44) in chicks from the ascites-susceptible (AS-S) and ascites-resistant (AS-R) lines
          Heart rate (beats/min)           SaO2 (%)

Age (d) AS-S AS-R %diff1 (%) AS-S AS-R %diff (%)

SBC: d 0 to 19
 1 435.2 ± 8.3 403.8 ± 7.7 7.8** 96.0 ± 0.6 96.0 ± 0.5 0
 7 449.8 ± 8.1 409.4 ± 7.5 9.8*** 95.4 ± 0.5 95.8 ± 0.5 −0.4
  17 419.3 ± 5.5 416.9 ± 5.1 0.6 92.7 ± 0.9 94.1 ± 0.9 −1.4
AIC: d 19 to 44
  27 378.6 ± 5.9 374.1 ± 6.6 1.2 75.4 ± 1.1 90.4 ± 1.2 −16.6***
1
%diff = (AS-S − AS-R)/AS-R.
**, ***The difference between the means of the AS-S vs. AS-R lines differ from zero at the 0.01 and 0.001 level of significance, respectively.
1988 Druyan et al.

exposure to AIC and the manifestation of AS, no differ-

Table 4. Means and SE of hemoglobin concentration (g/dL), hematocrit (%), red blood cell count (RBC, units), and hemoglobin per 1,000 erythrocytes (ng/1,000 cells) measured
during standard brooding conditions (SBC, from day of hatch to d 19) and under ascites-inducing conditions (AIC, from d 19 to end of trial on d 44) in chicks from the ascites-

−24.2**
−15.7*
−11.3*
ences in hemoglobin level were found between the AS-S

%diff

1.7
2.6
4.1
(%)
and AS-R lines (Table 4).

Hemoglobin (ng/1,000 cells)

1,029.4 ± 60.9
1,237.4 ± 77.0
1,177.5 ± 77.6

1,120.5 ± 30.2
850.9 ± 25.6
862.2 ± 41.9
Red Blood Cell Count and Hemoglobin

AS-R
per 1,000 Cells
The red blood cell count did not differ among lines
under SBC, although a slightly higher number was
found in the AS-S line (Table 4). Exposure to AIC re-

1,046.8 ± 66.8
1,269.1 ± 84.0
1,225.2 ± 65.2

944.8 ± 32.5
754.7 ± 33.4
653.5 ± 63.7
sulted in an increased red blood cell count in both lines;

AS-S

†
however, this increase was significant only in the AS-S
line. On d 25, six days after the instigation of the AIC
protocol, mean cell count in the AS-S line was 9.4%

†, ††, †††The difference between the means of d 18 vs. d 25 within line differs from zero at the 0.05, 0.01, and 0.001 level of significance, respectively.
higher than that of the AS-R line. This difference be-

*, **, ***The difference between the means of the AS-S vs. AS-R lines differs from zero at the 0.05, 0.01, and 0.001 level of significance, respectively.
tween the lines was maintained on d 32 (Table 4) and

24.9***
%diff
increased to 24.9% by d 41(14.6 vs. 11.7 × 109 in AS-S

(%)

9.4*
9.3*
5.1
7.2
6.3
and AS-R, respectively).
During the SBC period, no significant difference in
the calculation of hemoglobin content respective to red

7.8 ± 0.3
8.0 ± 0.3
9.6 ± 0.5

10.9 ± 0.4
12.8 ± 0.4
11.7 ± 0.5
RBC (×109)

AS-R
blood cell count was found between the lines, although
numerically it was slightly higher in the AS-S line. After
exposure to AIC, the AS-S birds exhibited a significant
reduction of this parameter in comparison to the reduc-

8.2 ± 0.3
8.6 ± 0.3
10.2 ± 0.4

11.8 ± 0.5
13.9 ± 0.5
14.6 ± 0.8
tion found in the AS-R birds (Table 4). Subsequently,

AS-S

†
the AS-S and AS-R line differed significantly in the
quantity of hemoglobin per 1,000 cells (Table 4) during
the AIC phase with 15.7, 11.3, and 24.2% difference
between the 2 lines on d 25, 32, and 41, respectably.

0.8
−0.4
0.5

−3.3
0.9
5.5
%diff
(%)
DISCUSSION
Hemoglobin (g/dL)

7.2 ± 0.2
8.9 ± 0.2
8.4 ± 0.3

9.0 ± 0.1
9.9 ± 0.2
10.1 ± 0.3
AS-R

Comparisons between AS-S and AS-R commercial

broilers depend solely on the phenotype [i.e., only in-
dividuals affected by AS (mortality or morbidity) are
identified as AS-S]. Under low-challenge AIC, relatively
7.2 ± 0.2
8.9 ± 0.2
8.4 ± 0.2

8.7 ± 0.1
10.0 ± 0.2
slow-growing AS-S broilers have a correspondingly low- 10.7 ± 0.5
AS-S

er demand for oxygen and hence do not develop AS.

Moreover, in floor-rearing systems, some AS-S broilers
manage to reduce or avoid cold stress by crowding to-
gether and so remain healthy. Misidentification of AS-S
susceptible (AS-S) and ascites-resistant (AS-R) lines

18.2***
29.6***

chicks, as being AS-R, may lead to biased findings re-

6.0**
%diff1
(%)

−1.2
0.8
1.4

garding the true genetic association between the mea-

sured traits and the genetic difference in susceptibility
Hematocrit (%)

or resistance of broilers to AS. Using the fully divergent

24.8 ± 0.3
26.7 ± 0.3
27.6 ± 0.4

29.3 ± 0.3
34.0 ± 0.5
35.8 ± 0.7

AS-S and AS-R lines largely avoids this problem; the

AS-R

††

AS-related genetic status of each individual is known

from its line affiliation, and not from its AS-related
%diff = (AS-S − AS-R)/AS-R.

phenotype.
24.5 ± 0.3
26.9 ± 0.3
28.0 ± 0.3

31.1 ± 0.3
40.1 ± 0.6
46.4 ± 1.1

The design of the present study allowed measurement

AS-S

†††

of AS-related traits in AS-S and AS-R lines during the

first 18 d under SBC and then monitored the fate of
each broiler genotype by moving it to AIC, which ef-
ficaciously induced AS in the susceptible birds. Results
AIC: d 19 to 44
SBC: d 0 to 19

from the 3 trials showed that 93.2% of the AS-S birds

developed AS and 91% of the AS-R birds remained
Age (d)

healthy. These data indicate that comparisons between

  13
  18

  25
  32
  41
1

the 2 lines could provide information about the associa-

 6
 ASCITES-RESISTANT VERSUS ASCITES-SUSCEPTIBLE BROILER LINES
tion between AS susceptibility-resistance and the trait
that was measured.
In this design, a trait differing significantly between
the 2 lines under SBC is expected to be a good in-
dicator for selection against AS susceptibility in com-
mercial flocks. Mean hatch weight was similar in both
lines, but the AS-S chicks exhibited a slightly higher
growth rate and consequently their mean BW on d 19
was significantly (P = 0.032) higher. However, both
lines exhibited a similar relative growth rate to d 19.
The difference in mean BW at d 19 between the lines
could be related to the environmental conditions un-
der which the initial divergent selection (generation S0)
was conducted 4 yr before these trials (Druyan et al.,
2007a). Due to relatively low challenge AIC, AS was
induced only in AS-S broilers with genetically higher
growth rate (which increased the AS challenge) and
not in AS-S broilers with genetically lower growth rate.
With this growth-rate-dependent differential genotype-
to-phenotype expression, the observed incidence of AS
per family was biased downward in slow-growing fami-
lies and upward in fast-growing families. Consequently,
the divergent selection in S0 also influenced differences
in BW (Druyan et al., 2007a, 2008), but not due to a
true genetic association between high growth rate and
AS. Similar divergence in early growth rate and BW
was found between another pair of AS-S and AS-R lines
that were selected divergently under hypobaric condi-
tions (Anthony and Balog, 2003; Balog et al., 2003;
Pavlidis et al., 2007).
In the present study, exposure to extreme AIC en-
hanced the imbalance between oxygen demand and
supply in susceptible broilers and induced differences
in blood parameters between the 2 lines. No signifi-
cant differences in hematocrit level, mean red blood
cell count, hemoglobin concentration, and hemoglobin
count per 1,000 red blood cells were found under SBC.
As reported in numerous previous reports (Julian and
Mirsalimi, 1992; Shlosberg et al., 1996; Olkowski et al.,
1997, 2005; Wideman et al., 1998; Luger et al., 2003),
after the instigation of AIC protocol, the AS-S birds
exhibited an increase in hematocrit levels and red blood
cell count. These changes were also seen in the AS-R
line but with a lower, nonsignificant magnitude. The
high hematocrit level and increase in the red blood cell
count in the AS-S line were concomitant with a decline
in SaO2 of birds from that line. In the present study,
both lines exhibited an increased red blood cell count
that was accompanied by a slight increase in blood he-
moglobin concentration, but further calculations of he-
moglobin content per 1,000 red blood cells revealed a
significant reduction in hemoglobin content in the AS-S
birds compared with the AS-R birds. The differences
were similar in magnitude to the differences between
the lines in mean red blood cell count at the same ages,
due to the lack of difference between the lines in he-
moglobin concentration (Table 4). These results are
in agreement with the previous study of Luger et al.
1989
(2003), who found that plasma volume was similar in
ascitic and healthy broilers, but the hematocrit was
significantly higher in birds with AS, as a result of a
significant increase (by 23%) in the number of erythro-
cytes that were partly immature. The lower SaO2 in the
AS-S broilers may therefore be associated with a sig-
nificant increase in the number of immature red blood
cells and the significant decline in hemoglobin content,
which lowered the total capacity oxygen of the blood.
These changes in birds developing AS are in agree-
ment with numerous previous reports (Julian and Mir-
salimi, 1992; Shlosberg et al., 1996; Olkowski et al.,
1997, 2005; Wideman et al., 1998; Luger et al., 2003;
Druyan et al., 2007b). However, these changes are sec-
ondary manifestations of AS and hence are useful for
diagnosis of birds that are already developing AS, but
are not suitable for selection that needs a predictive
indicator in healthy birds at a young age.
Heart rate was significantly higher in the AS-S birds
than in the AS-R birds on d 1 and 7, with only the
lowest quartile of individual heart rates in the AS-S
line overlapping the range of individual heart rates of
the highest quartile in the AS-R line, and may cause a
false identification of AS-R birds with high heart rate
as AS-S. These results are in agreement with Druyan
et al. (2005), where S3 chicks from the AS-S line had
a significantly higher heart rate on day of hatch com-
pared with S3 chicks from the AS-R line. It was re-
ported (Tazawa et al., 1992) that an increase in heart
rate begins shortly after hatch, reaching a peak close to
4 wk of age. Thereafter, a slow decline in heart rate oc-
curs (Wideman, 1999). In this study, the AS-R line fit-
ted that description well, heart rate on d 1 (404 beats/
min) rose mildly toward d 17 (417 beats/min). How-
ever the AS-S line demonstrated an increase in heart
rate only between d 1 to 7, with a decline thereafter
toward d 17. Mild hypoxia was found to attribute to
an increase in heart rate (Besch and Kadono, 1978;
Faraci, 1986), suggesting that AS-S birds in this study
experienced oxygen shortage already during the time
of hatch. Scheele et al. (2003, 2005) found that higher
mean partial pressure of CO2 venous blood (a marker
for lung ventilation rate) of birds on d 11 coincidences
with higher ascites susceptibility. These results, as well
as the results of the present study, indicate that AS-S
birds are under oxygen shortage at an early age. How-
ever, the present results suggest that as long as the
susceptible birds are under SBC, higher heart rate can
compensate for a mild oxemia and no other physiologi-
cal parameter was affected.
In recent years, breeding companies have been select-
ing against broilers with low SaO2, measured by oxime-
ter in flocks of selection candidates at about 5 wk of age
(Powell et al., 2008). However, due to low heritability
and dominance of high SaO2 (Navarro et al., 2006),
the usefulness of low SaO2 as an effective indicator for
further selection against AS susceptibility is expected
to be limited. The results of the present study suggest
1990 Druyan et al.

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Olkowski, A. A., J. A. Abbott, and H. L. Classen. 2005. Patho-
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