Comparison of the broody behavior characteristics
of different breeds of geese
Y. Yao,∗ Y. Z. Yang,∗ T. T. Gu,∗ Z. F. Cao,∗ W. M. Zhao,∗ H. R. Qin,† Q. Xu,∗,1 and G. H. Chen∗,1
∗
College of Animal Science and Technology, Yangzhou University, Yangzhou, Jiangsu Province 225009, China;
and † Jiangsu Animal Husbandry & Veterinary College, Taizhou, Jiangsu Province 225300, China
ABSTRACT A low laying performance in goose is
one of the key factors preventing the industrial develop-
ment, and the laying performance is related to broody
behavior. However, the characteristics of broody behav-
ior in geese remain unclear. In this study, the total 144
geese (300 day old), including Zhedong geese (Anser
cygnoides), Sichuan geese (Anser cygnoides), and Car-
los geese (Anser anser) were selected and assigned to
1 of 3 groups/breed (including 4♂+12♀). Laying and
broody behaviors were recorded using the infrared video
cameras from 2016 November 11 to 2017 June 15. The
broody behavior was detected in 19.4% of Carlos geese,
33.3% of Sichuan geese, and 100% of Zhedong geese.
Different goose breeds showed similar behavior charac-
teristics. The low frequency of feeding, drinking, and
low body weight were observed in the middle of broodi-
ness. As the brooding progressed, the body temperature
showed a downward trend and then recovered, whereas
no difference was observed in Carlos goose. In addi-
tion, the plasma hormone concentration from different
breeds and stages of broodiness were compared. The
Key words: goose, broody behavior, hormones, egg production
INTRODUCTION
Broodiness, a common habit of most domestic fowls,
is associated with persistent nesting, turning, and re-
trieval of eggs, characteristic clucking, and defense of
the nest (Romanov et al., 2002). Broodiness is favor-
able in wild birds, which need to produce new offspring
through the natural incubating process, but it is unfa-
vorable in breeder flocks kept for maximum egg produc-
tion (Mohamed, 2013). Although the domestic birds are
used in the commercial production of hatching eggs, the
broody behavior still exist in some indigenous chicken
breeds and goose breeds. The rate of broodiness of silky
is over 60% (Geng et al., 2013). Almost over 90% in
some Chinese goose breeds, such as Zhedong geese and

2019 Poultry Science Association Inc.
C
Received February 27, 2019.
Accepted June 10, 2019.
1
Corresponding authors: ghchen@yzu.edu.cn (GHC); xuqi@yzu.
edu.cn (QX)
5226
contents of FSH (follicle-stimulating hormone) and LH
(luteinizing hormone) in geese were greater in the lay-
ing stage than that in the broody stage. Fewer FSH
and LH were detected in Zhedong geese and Carlos
geese, more in Sichuan geese. In broody goose, the PRL
(prolactin) concentrations of the 3 goose breeds peaked
in the middle of broodiness, and greater PRL was de-
tected in Sichuan geese than those in Carlos geese and
Zhedong geese. Finally, we compared egg production
between the broody and non-broody geese in the ob-
servation period. The egg production of broody Carlos
geese was 27, which was significantly higher than non-
broody geese (14 eggs), while in Sichuan geese there
was no significant difference between broody (24 eggs)
and non-broody geese (26 eggs). Finally, the higher egg
production was found with the more broody times in
Zhedong geese. Taken together, although the different
goose breeds showed similar broody behavior character-
istics, the broody rate and hormone secretion were dis-
similar, and the Zhedong geese exhibited strong broody
feature.
2019 Poultry Science 98:5226–5233
http://dx.doi.org/10.3382/ps/pez366
Magang geese, with annual 30 to 40 eggs (Yu et al.,
2016; Qin et al., 2013), whereas some goose breeds show
relative weak broodiness such as Huoyan geese with an-
nual 70 to 100 eggs (China National Commission of Ani-
mal Genetic Resouces, 2011). The strong broodiness of
goose has become key factor preventing the develop-
ment of the goose industry in China. Hence, it plays
an important role in understanding the broody rules
and characteristics to improve the egg performance of
different goose breeds.
Broody behavior not only reduces egg production,
but is also modulated by endocrine hormone. The
hypothalamic–pituitary–gonadal axis is a coordinated
neuroendocrine system that regulates initiation, devel-
opment, and maintenance of the broodiness process
in poultry. In detail, the hypothalamus is the coordi-
nating center of the endocrine system, which consol-
idates signals derived from upper cortical inputs, au-
tonomic functions, environmental cues such as light
and temperature, and peripheral endocrine feedback
 BROODY BEHAVIORS OF GEESE
(De and Schwartz, 2006). In turn, the hypothalamus
delivers precise signals to the pituitary gland, which
then releases hormones that influence most endocrine
systems in the body. For example, increased prolactin-
releasing hormones in hypothalamus acted on the an-
terior pituitary through the portal system, which re-
sulted in an increase in PRL secretion, and a decrease
in LH, FSH, and progesterone (PROG) secretion. High
levels of PRL affected the ovary and hypothalamic
body temperature regulation center through blood cir-
culation, and then inhibited the ovarian development
and adjusted body temperature (Burke and Dennison,
1980).
Although broody behavior has been observed, and
endocrine hormones have been evaluated during broodi-
ness in most domestic fowl, such as chickens (Shimmura
et al., 2010; Geng et al., 2014) and Muscovy ducks
(ChIh and Ma, 1977; Mohamed, 2013), there have
been no reports on the details of broody behaviors of
geese. In our study, the 3 goose breeds (Zhedong geese,
Sichuan geese, and Carlos geese) with different broody
characteristics were selected, the broody behaviors were
systematically observed, and the concentrations of re-
productive hormones were investigated in the annual
reproductive cycles. The results obtained an invalu-
able information for understanding the differences in
broody patterns of some goose breeds, which provided
a scientific basis for improving the reproductive per-
formance and promoting the industrial development in
geese.
MATERIALS AND METHODS
Ethics Statement
All the experimental procedures were approved by
the animal care and use committee of Yangzhou Uni-
versity (approval number: 151–2014).
Experimental Design and Management
The experiment was conducted at the National Wa-
terfowl Germplasm Resources Pool (Taizhou, China).
A total of 144 geese (300 day old), including Zhe-
dong geese (Anser cygnoides), Sichuan geese (Anser
cygnoides), and Carlos geese (Anser anser), were se-
lected and assigned to 1 of 3 groups/breed (including
4♂+12♀) and raised in separate pens. All geese were
provided with the same diet, which was combined with
coarse and concentrated material and fed ad libitum
(Table 1). The geese were kept in a standard light reg-
imen of 14 h light (14L:10D) during the laying period
(Wang, 2009). The nests were used with rice husk as
the mattress, which was large enough (96 cm×85 cm)
to comfortably hold several standing geese, and allowed
geese to turn around easily. Colored paint on the neck
or/and back of the geese was used as a marker for ob-
serving behavior.
5227
Table 1. Composition and nutrient content of
experimental diet (air-dry basis).
Item Content(%)
Corn powder 40.25
Crushed wheat 25.00
Crushed barley 10.00
Bean cake powder 13.25
Green hay powder 4.00
Limestone 4.50
Calcium hydrogen phosphate 1.50
Salt 0.50
Vitamin and microelement additives 1.00
Total 100
Recording of Behaviors and Observation
An infrared video camera (Jindun, 720p,
resolution:1280×960, frame Rate: 25 frame/s, Nanjing,
China) was set up above laying nest in the pen of each
group. In the preliminary trail, when geese had laid
batch of eggs, some geese began to display broody
behaviors. This behavior persisted for more than 2
consecutive days, and we classified it as broodiness.
We found that the females sit on the nest for long
periods and ultimately incubated their eggs for almost
the whole day if eggs accumulated in the nest for 3
consecutive days. We also discovered the geese kept
incubating in the nest when we took all the eggs
away. Broody behaviors, including persistent nesting,
turning, and retrieval of eggs, clucking, and nest de-
fense, were observed. According to the characteristics
of behavior, the broody stage was divided artificially
into 3 stages. First, the early stage of broodiness, the
geese initiate broodiness, the geese have a nest-seeking
behavior, that is, they keep walking in the nest, looking
for suitable nests, and frequently using the husks to
build nests. Secondly, the middle stage of broodiness,
the geese enter this stages, they show continuous sleep
with half-closed eyes, do not accept mounting from
the gander. The broody time is up to 22 h every day,
and the drinking & feeding greatly reduce compared
with the early of broodiness. Finally, the later stage
of broodiness, the geese enter the later stage, the
feather color of the broody geese get dim. The time
of incubating gradually decreases with broodiness
prolongs. The geese occasionally walk out of the nest,
and the drinking and feeding greatly gradually increase
compared with the middle of broodiness.
In the formal trail, the broody behaviors were
recorded daily in the annual reproductive cycle (2016
November 11 to 2017 June 15), and the daily data were
collected and analyzed from 6 am to 6 pm. Eggs were
collected at 6 pm every day. The proportion of broodi-
ness, the number of broodiness, the peak of broodiness,
the duration of broodiness, and the daily broody times
of the 3 goose breeds were analyzed in the annual re-
productive cycle.
Another infrared video camera (Jindun, 720p, Nan-
jing, China) was set up above playground in the
5228
Table 2. Stastistical analysis of broody characteristics in the 3 goose breeds.
The percentage
Breeds of broodiness
Zhedong goose 100%a
Sichuan goose 33.3%b
Carlos goose 19.4%b
P (df) 0.000(2)
Note: Each breed contained 36 geese, and the numbers of broody geese of Zhedong geese, Sichuan
geese, and Carlos geese were 36, 12, and 7, respectively.
a,b
Different superscripts within columns indicate means are significantly different (P ≤ 0.05).
pen. The frequency of daily feeding and drinking was
recorded from 6 am to 6 pm daily in the observation
period.
Measurement of the Body Temperature and
Body Weight
The body temperature and body weight of geese were
investigated in the laying stage and the broody stage. A
special poultry infrared thermometer was used to mea-
sure the body temperature.
Determination of Reproductive Hormone
Blood samples were collected from the brachial veins
of geese during the laying stage and the broody stage.
The blood was centrifuged at 3,000 rpm for 10 min,
and samples were stored at –20◦ C. The concentrations
of PRL, AMH, FSH, PROG, and LH were detected by
infinite M200 Pro microtiter plate reader (Swiss Tecan
company) using the commercial enzyme immunoas-
say kits (Shanghai Taisheng Bioengineering Institute,
Shanghai, China). The minimal detectable doses of
PRL, AMH, FSH, PROG, and LH were 0.6 ng/L,
0.5 ng/mL, 0.3 U/L, 3.5 pmol/L, and 16pg/mL, respec-
tively.
Recording of Egg Production
In the observation period, the individual egg produc-
tion was recorded daily using the infrared video cam-
eras. Even if the geese laid eggs at night, the individual
could be identified by the depth and position of the
marked color.
Statistical Analysis
The comparisons among different breeds and differ-
ent measure stages pairwise were fullfilled using re-
peated measures and Mauchly’s Test of Sphericity in
SPSS statistical software (SPSS, Ver. 19.0), including
the broody behaviors, the body weight, body tempera-
ture, and reproductive hormone content. Mauchly’s test
of sphericity was judged whether there were relations
among the repeatedly measured data. If any (P ≤ 0.05),
the 2-way ANOVA was taken, if not (P > 0.05), the 1-
way ANOVA was used (Zhang, 2002). The data of the
YAO ET AL.
The number of broodiness The peak period of
(times/goose∗ cycle) broodiness
2.00 ± 0.47a Feb-Mar and Jun-Jul
1.25 ± 0.50b Jan- Feb and Apr-May
1.00 ± 0.00b Feb-Apr
0.000(2)
number of broodiness of different breeds were evaluated
with the 1-way ANOVA procedure, and the proportions
of broodiness were analyzed with chi-squared tests. The
mean ± SE are presented in the table and figures. A
level of P ≤ 0.05 was set as the criterion for statistical
significance.
RESULTS
Comparison of the Specific Broody
Behaviors of the Different Geese Breeds
In our study, we found the geese kept incubating in
the nest even if the eggs took away every day. All of the
Zhedong geese and 33.3% of the Sichuan geese displayed
broody behaviors after laying batch of eggs. How-
ever, 19.4% of Carlos geese displayed broody behavior
(Table 2). Zhedong geese displayed broody behav-
ior twice on average in the annual reproductive cy-
cle, whereas Sichuan geese and Carlos geese displayed
broody behavior only once (Table 2). Female geese sit
on the nest for more than 1 mo. The peak period of
broody behavior was different among the 3 breeds. Car-
los geese preferred to brood from February to April,
whereas Sichuan geese and Zhedong geese from Jan-
uary to February, April to May and from February to
March, June to July, respectively (Table 2).
Furthermore, specific broody behaviors of different
goose breeds were recorded. The significant main ef-
fect was observed for different stages on the duration
of broodiness and drinking frequency (Table 3). The
duration of broodiness in different goose breeds was
about 27 D during the middle of broodiness, which
was significantly longer than both early and later stages
(P = 0.000) (Table 3). The broody time lasted for more
than 20 h/d in the early and middle of broodiness, and
shortened to 2 to 3 hs/d in the later of broodiness.
It was only observed that Sichuan geese have longer
broody time in the early of broodiness than that of Zhe-
dong geese. For drinking frequency, broody geese only
drank around 2 times/day in the middle of broodiness,
whereas about 4 to 5 times in the early and late stages.
In addition, the low frequency of feeding was observed
in the middle of broodiness. Sichuan geese only feed
1.20 times/day, which was significantly lower than that
of Zhedong geese and Carlos geese (P < 0.05).
 BROODY BEHAVIORS OF GEESE 5229
Table 3. Main effect of breeds and stages of broodiness on specific broody behaviors.

Stages of the Duration of Daily broody Feeding frequency Drinking fre-

Breeds broodiness broodiness (day) time(h/day) (times/day) quency(times/day)

Zhedong goose Early 5.36 ± 0.26 20.06 ± 0.30c 3.66 ± 0.15a,b 3.87 ± 0.14
Middle 26.57 ± 1.41 23.29 ± 0.08a 2.31 ± 0.17c 2.13 ± 0.15
Later 7.82 ± 0.39 3.65 ± 0.30d 4.29 ± 0.19a 4.83 ± 0.27
Sichuan goose Early 7.00 ± 1.73 21.90 ± 0.81a,b 4.33 ± 0.67a 3.73 ± 0.18
Middle 34.67 ± 4.91 23.23 ± 0.03a 1.20 ± 0.06d 1.40 ± 0.06
Later 7.33 ± 0.88 3.83 ± 0.96d 3.47 ± 0.09a,b 3.90 ± 0.06
Carlos goose Early 6.86 ± 0.51 21.20 ± 0.60b,c 3.90 ± 0.30a 3.87 ± 0.19
Middle 27.57 ± 3.85 22.40 ± 0.41a,b 2.61 ± 0.20b,c 2.37 ± 0.36
Later 7.29 ± 0.61 2.37 ± 0.37d 4.33 ± 0.47a 5.60 ± 0.67
Breeds Zhedong goose 39.75 ± 1.53 15.67 ± 8.75 3.42 ± 1.21 3.61 ± 1.52
Sichuan goose 49.00 ± 4.04 16.32 ± 9.45 3.00 ± 1.52 3.01 ± 1.22
Carlos goose 41.71 ± 3.96 15.32 ± 9.47 3.61 ± 1.13 3.95 ± 1.77
Stages of broodiness Early 5.76 ± 1.64b 20.41 ± 1.67 3.76 ± 0.81 3.86 ± 0.67b
Middle 27.40 ± 8.15a 23.12 ± 0.67 2.28 ± 0.87 2.12 ± 0.80c
Later 7.68 ± 1.93b 3.43 ± 1.53 4.23 ± 1.01 4.90 ± 1.48a
P-value (Mauchly’s 0.000 < 0.05 0.294 > 0.05 0.220 > 0.05 0.001 < 0.05
test of sphericity)
P-value (2-way ANOVA)
Breeds 0.357 – – 0.123
Stages 0.000 – – 0.000
Breeds x stages 0.298 – – 0.592

Note: The daily data were collected and analyzed from 6 am to 6 pm in the annual reproductive cycle (2016 November 11 to 2017 June 15).
The data were recorded from 29 broody Zhedong geese (excluded 7 sick and disabled geese in later observation), 12 broody Sichuan geese, and
7 broody Carlos geese.
a–d
Different superscripts within columns indicate means are significantly different (P ≤ 0.05).

Changes in Weight and Body Temperature
During the Different Broody Stages
Although the 3 goose breeds have different body
types, they all showed significant weight loss in the
middle stage of broodiness compared with the laying
stage (P = 0.021), and recovered to a certain degree
in the later stage of broodiness (Table 4). The body
temperature of Zhedong geese and Sichuan geese also
changed significantly during the brooding process (P <
0.05). As the brooding progressed, the body temper-
ature showed a downward trend and then recovered,
whereas no difference was observed on the body tem-
perature in Carlos geese.
Plasma Hormone Concentrations During
the Different Broody Stages
The plasma hormone concentrations from different
breeds and stages of the broodiness were compared.
Some significant main effects were detected in differ-
ent breeds and stages on the contents of FSH and LH
(Table 5). The contents of FSH and LH were greater
in the laying stage than the broody stage (P = 0.019,
P = 0.007). Fewer FSH and LH were detected in
Zhedong geese and Carlos geese, and more in Sichuan
geese. In broody geese, greater AMH were detected
among the 3 breeds than that in laying geese. The con-
tent of AMH was the greatest from Sichuan geese in the
middle of broodiness, whereas that was the fewest from
the laying geese of Carlos geese. Additionally, there was
a significant interaction between factors (breeds and
stages) for the contents of PRL and PROG (P = 0.000).
As for the content of PRL, the concentration of 3 goose
breeds peaked in the middle stage of broodiness, and
greater PRL was detected in Sichuan geese than those
in Carlos geese and Zhedong geese, and there was no
difference among the 4 stages in Zhedong geese. Fewer
PROG tended to be present in the middle of the brood-
iness of Sichuan geese and Zhedong geese than that in
the other stages, while the fewest PROG in the later of
the broodiness of Carlos geese.
Effect of Broody Behaviors on Egg
Production
Egg production has a crucial effect on poultry re-
productive performance, and clutch size and broodiness
were taken into consideration in our study. Here, some
geese displayed broody behaviors after laying a clutch
of eggs, but some geese did not. Therefore, we com-
pared egg production between broody and non-broody
geese. The results showed that there was no significant
difference in egg production between broodiness and
non-broodiness of Sichuan geese (Figure 1, P = 0.394).
However, the egg production of non-broody geese (27
eggs) was significantly higher than that of broody in
Carlos geese (14 eggs) (Figure 1, P = 0.018).
In addition, all Zhedong geese displayed broody be-
haviors after laying a clutch of eggs. Meanwhile, the
broody time was different in the annual reproductive
5230 YAO ET AL.
Table 4. Main effect of breeds and stages of the broodiness on body weight and temperature.

Body
Breeds Stages of the reproduction Body weight (g) temperature(◦ C)

Zhedong goose Laying stage 3911.41 ± 581.80 41.37 ± 0.36a,b

Early stage of broodiness 3751.11 ± 501.13 40.99 ± 0.44a-c
Middle stage of broodiness 3451.00 ± 511.97 40.85 ± 0.52b-d
Later stage of broodiness 3953.78 ± 544.18 41.38 ± 0.44a,b
Sichuan goose Laying stage 3486.33 ± 263.08 41.43 ± 0.06a
Early stage of broodiness 3085.00 ± 177.10 41.03 ± 0.23a-c
Middle stage of broodiness 2812.00 ± 420.76 40.37 ± 0.25d
Later stage of broodiness 3412.67 ± 249.95 40.70 ± 0.56c,d
Carlos goose Laying stage 4406.86 ± 576.25 41.39 ± 0.43a,b
Early stage of broodiness 4633.71 ± 583.18 41.51 ± 0.21a
Middle stage of broodiness 4134.43 ± 629.62 41.20 ± 0.13a-c
Later stage of broodiness 4354.71 ± 589.28 41.54 ± 0.19a
Breeds Zhedong goose 3766.82 ± 564.14b 41.15 ± 0.50
Sichuan goose 3199.00 ± 375.76c 40.88 ± 0.50
Carlos goose 4382.43 ± 589.32a 41.41 ± 0.29
Stages of the reproduction Laying stage 3970.68 ± 602.36a 41.38 ± 0.35
Early stage of broodiness 3864.08 ± 644.31a 41.09 ± 0.44
Middle stage of broodiness 3528.47 ± 619.59b 40.88 ± 0.50
Later stage of broodiness 3985.76 ± 574.31a 41.35 ± 0.46
P-value (Mauchly’s test of 0.008 < 0.05 0.377 > 0.05
sphericity)
P-value (2-way ANOVA)
Breeds 0.000 –
Stages 0.021 –
Breeds x stages 0.775 –
Note: The data were recorded from 29 broody Zhedong geese, 12 broody Sichuan geese, and 7 broody Carlos geese.
a–d
Different superscripts within columns indicate means are significantly different (P ≤ 0.05).

Table 5. Main effect of breeds and stages of the broodiness on plasma hormone levels.

Breeds Stages of the reproduction PRL(ng/L) AMH(ng/mL) FSH(U/L) PROG(pmol/L) LH(pg/mL)

Zhedong goose Laying stage 34.81 ± 2.84e 4.19 ± 0.51d,e 3.84 ± 0.37 46.19 ± 6.34b-d 116.06 ± 12.69
Early stage of broodiness 50.00 ± 6.11e 6.47 ± 0.84b-d 2.00 ± 0.23 29.65 ± 2.80d,e 71.98 ± 9.32
Middle stage of broodiness 61.97 ± 5.76e 6.73 ± 0.79b-d 2.13 ± 0.23 33.97 ± 4.13c-e 68.22 ± 8.63
Later stage of broodiness 41.76 ± 4.80e 7.17 ± 0.78b,c 2.54 ± 0.26 49.94 ± 4.35b,c 76.24 ± 6.74
Sichuan goose Laying stage 39.55 ± 7.65e 6.56 ± 1.82b-d 4.32 ± 1.05 72.75 ± 5.88a 89.86 ± 3.97
Early stage of broodiness 110.32 ± 16.94d 7.48 ± 2.44b,c 3.78 ± 1.73 27.91 ± 2.52d,e 75.46 ± 18.10
Middle stage of broodiness 180.78 ± 9.21a 11.18 ± 3.27a 4.10 ± 1.36 26.07 ± 7.36e 71.63 ± 18.92
Later stage of broodiness 164.22 ± 10.77a,b 8.96 ± 1.43a,b 3.96 ± 1.21 56.28 ± 5.05a,b 76.15 ± 6.08
Carlos goose Laying stage 36.58 ± 5.64e 3.83 ± 0.41e 4.75 ± 0.85 37.89 ± 3.22b-e 69.89 ± 4.86
Early stage of broodiness 107.98 ± 7.23d 4.43 ± 0.96d,e 3.84 ± 0.85 26.84 ± 4.29d,e 53.70 ± 7.30
Middle stage of broodiness 141.75 ± 12.96b,c 5.55 ± 1.17c-e 3.07 ± 0.57 37.41 ± 1.91b-e 42.78 ± 4.63
Later stage of broodiness 128.60 ± 10.24c,d 3.78 ± 0.95e 3.75 ± 0.79 7.00 ± 0.90f 53.03 ± 6.16
Breeds Zhedong goose 51.39 ± 31.19 6.59 ± 4.19 2.42 ± 1.47b 38.78 ± 27.66 77.02 ± 48.65a
Sichuan goose 127.38 ± 66.79 8.32 ± 4.24 4.08 ± 2.24a 46.67 ± 26.01 79.56 ± 22.48a
Carlos goose 103.73 ± 55.08 4.49 ± 2.65 3.76 ± 2.12a 25.16 ± 16.96 52.70 ± 17.66b
Stages of the Laying stage 36.91 ± 19.82 4.94 ± 2.79 4.31 ± 2.01a 53.66 ± 26.21 95.38 ± 48.09a
reproduction
Early stage of broodiness 78.48 ± 48.36 6.01 ± 4.08 2.87 ± 2.10b,c 28.73 ± 17.75 66.84 ± 41.47b
Middle stage of broodiness 104.19 ± 63.04 6.87 ± 4.35 2.58 ± 1.54c 33.95 ± 24.36 60.93 ± 39.02b
Later stage of broodiness 103.91 ± 61.55 6.42 ± 3.96 3.08 ± 1.80b,c 40.18 ± 31.14 69.27 ± 30.95b
P-value (Mauchly’s test of 0.000 < 0.05 0.547 > 0.05 0.000 < 0.05 0.000 < 0.05 0.000 < 0.05
sphericity)
P-value (2-way ANOVA)
Breeds 0.013 – 0.002 0.376 0.001
Stages 0.029 – 0.019 0.395 0.007
Breeds x stages 0.000 – 0.479 0.000 0.619

Note: Plasma collected from 29 broody Zhedong geese, 12 broody Sichuan geese, and 7 broody Carlos geese at 4 stages (laying stage, early stage
of the broodiness, middle stage of the broodiness, and later stage of the broodiness) of the reproduction. The 5 individuals from the 3 goose breeds
were selected and detected hormone.
a–e
Different superscripts within columns indicate means are significantly different (P ≤ 0.05).
 BROODY BEHAVIORS OF GEESE
Figure 1. Comparison of egg production of broody and non-
broody geese. Note: The eggs were recorded for every day (from 2016
November 11 to 2017 June 15). Non-broody: some geese showed non-
incubating eggs after they laid a clutch. Broody goose: some geese
performed to incubate eggs after they laid a clutch. The asterisk (∗ )
indicates a significant difference (P < 0.05), df = 1.
Figure 2. Effect of frequency of broody behavior on egg production
in the annual reproductive cycle of Zhedong geese. Note: Fbro = 1: The
only one brooding period in the annual reproductive cycle. Fbro = 2:
The 2 brooding periods in the annual reproductive cycle. Fbro = 3:
The 3 brooding periods in the annual reproductive cycle. The same
lowercase letters indicate no significant differences (P > 0.05), while
different lowercase letters indicate significant differences (P < 0.05),
df = 2.
cycle. The egg production was compared among the
geese with different broody times (Figure 2). The re-
sults indicated that the egg production of geese that
displayed brooding once (Fbro = 1), twice (Fbro = 2),
and 3 times (Fbro = 3) averaged 9 eggs, 18 eggs, and 25
eggs, respectively. The broody frequency had a signifi-
cant effect on egg production, with higher frequencies
resulting in a greater egg production in Zhedong geese.
DISCUSSION
At present, geese generally were reared on floor,
and the cage rearing has not yet appeared. Therefore,
5231
goose broodiness is still the main problem of commer-
cial production of goose in China. Some investigations
have confirmed this phenomenon (Qin et al., 2013; Yu
et al., 2016). However, few studies have evaluated the
frequency of specific brooding behaviors, and the egg
laying and incubation patterns of different waterfowl.
Jiang et al (2010) analyzed the correlation between
broody behaviors and egg production in indigenous
chickens, and Geng et al (2014) studied the effects of
photoperiod on broody behavior of Beijing You chick-
ens. Our study aimed to observe the behavior of dif-
ferent goose breeds during the broody stages, and ex-
plore the egg laying broody pattern of these geese. In
our study, different geese performed similar broody be-
haviors, but there were differences in the frequency of
broody behaviors in the various goose breeds tested.
In the long-term breeding process, Sichuan geese and
Carlos geese often eliminated individuals with rela-
tively strong broodiness, and the percentage of broody
geese was reduced. Conventional breeding has a slower
progress in controlling broody traits due to the low her-
itability (h2 = 0.116) (Romanov et al., 2002), and it is
difficult to completely eliminate broodiness, so some in-
dividuals are always retained broodiness. Zhedong geese
has strong incubation tendency and the broodiness rate
is almost 100%, which make it difficult to eliminate
broodiness by selecting non-broody individuals.
As broodiness progressed, feeding and drinking de-
creased gradually. However, chickens consume almost
the same amount of food during broodiness at other
times (Shimmura et al., 2015). The difference in behav-
ior between geese and chickens may revealed that the
goose has strong fat storage capacity. Basheer (2013)
found that if eggs are continuously removed from the
nests of turkeys, it shortened the time spent in the nest,
and also stopped the broody behavior of turkeys. How-
ever, in our study, geese continued to nest if eggs were
removed out of the nest, which indicates that geese have
stronger maternal behaviors than turkeys.
The body weight and temperature of geese dropped
to minima in the middle stage of broodiness, which is
in accordance with another study (Savory, 1979). Fe-
males spent almost of their time in the nest warming
the eggs, and had reduced activity and feeding behav-
iors, which account for the decreases in body weight and
temperature. All geese gradually regained the weight
and temperature during the later stage of broodiness.
Feed intake during the later stage of egg incubation
was greater, which may be related to the body weight
gained when the geese were near the end of broodi-
ness. These results hinted that we should pay atten-
tion to the feed intake of broody geese during the
feeding and management process, and properly feed
the geese with their preferred food to avoid excessive
weight loss, which may cause difficulties in recovery of
body weight later (Yang and Cheng, 1996). We also
found that the body temperature of the Carlos geese
did not decrease significantly during the brooding pro-
cess, which may be due to the strong subcutaneous
5232 YAO ET AL.
fat deposition ability compared with the other 2 goose
breeds.
Broody behaviors are regulated by a variety of hor-
mones. Studies have demonstrated that PRL played a
key role in starting and maintaining broody behaviors.
In our study, the PRL concentrations of the 3 goose
breeds peaked in the broody stage. The onset of broody
behavior in chicken and turkey is correlated with an in-
crease of PRL in the plasma (Sharp et al., 1988; Be-
decarrats et al., 1997), and that the blood PRL level
remains high throughout the broody stage in chicken
and turkey (Zadworny et al., 1988; Kuwayama et al.,
1992). Further research confirmed that the sagittal area
that immunostained for PRL was significantly greater
in the adenohypophysis of broody turkey hens (Ramesh
et al., 1996). Besides, American king pigeons in the
broody stage were found PRL concentrations rose by
110.28% (Zhang, 2011). In addition, we also discovered
greater PRL was detected in the middle of broodiness of
Sichuan geese than those in Carlos geese and Zhedong
geese, which might be related to the depth of broodi-
ness. Our results also revealled the lower frequency of
drinking water and feeding and the longer broody dura-
tion in Sichuan geese comapred with the other 2 breeds.
We also discovered high concentrations of AMH dur-
ing the broody stage. AMH is secreted by antral folli-
cles and small sinusoidal follicle granulosa cells (Rooij
et al., 2002; Fanchin et al., 2003). Many studies have
shown that AMH is related to ovarian reserve function
(Carlsson et al., 2006). Low concentrations of AMH will
accelerate the conversion rate of primordial follicles to
growing follicles, and reduce the function of ovarian re-
serve (Sun et al., 2016). Besides, it was believed that
the high expression level of AMH in duck pre-follicles
maintained the undifferentiated granulosa cells, and in-
hibited the initial recruitment of primordial follicles
(Chen et al., 2018). Therefore, it was reasonable that
the concentrations of AMH in the 3 goose breeds were
significantly higher in the broody stage than that in
the laying stage, and AMH concentrations peaked in
the middle stage of the broodiness. At the same time,
we also found that Sichuan goose had the highest AMH
content in the middle of the broodiness, which might
mean that Sichuan geese better maintained the ovarian
reserve function during the broody process.
In addition, we also found that concentrations of FSH
and LH were lower in the broody stage, and were higher
in the laying stage. Another studies also revealed that
the concentrations of FSH and LH in chicken and turkey
were high in the laying stage and low in the broody
stage (Porter et al., 1991; Sharp et al., 1979; Khalil
et al., 2009). FSH and LH acted as 2 gonadotropins,
which promoted follicular maturation and estrogen se-
cretion. It was also reasonable that a large number
of follicle atresia and cell apoptosis resulted in low
secretion of FSH and LH during the broodiness. At
the same time, we found that these 2 hormones in-
creased to some extent in the later stage of broodiness,
which also seemed to mean that the follicles begun to
develop again in the stage. Fewer FSH and LH were
detected in Zhedong Geese and Carlos Geese, which
suggested more follicle atresia and cell apoptosis were
happened in the broody stage. As for the PROG, it
was reported that the highest level of PROG was found
in laying turkey, while the lowest value was in broody
turkey (Khalil et al., 2009). While we found the concen-
tration appeared different tendency with different goose
breeds, especially fewest in the later stage of the brood-
iness of Carlos geese and highest in the laying stage of
Sichuan geese. The results might be concerned with the
difference of broody behaviors in the 3 geese breeds.
Although broody behavior of poultry is significant
for their offspring, it also affects egg production (Geng
et al., 2013). Our study showed that egg production of
broody Carlos geese was significantly lower than that
of non-broody geese. Shimmura et al. (2010) found that
a high proportion of broody chicken resulted in low
egg production. Tsuyoshi et al. (2015) also revealed egg
production was lower in brooded than in non-brooded
chickens (P < 0.05). However, there was no significant
difference in egg production between broody and non-
broody Sichuan geese. In our observation, some geese
neither brooded their eggs nor continued to lay after
finishing a clutch of eggs. It was necessary to increase
egg production by removing broody geese or stopping
the broody behavior for some breeds like Carlos geese,
but this method was not suitable for all breeds.
Another important result was that all females of Zhe-
dong geese brooded after laying eggs, and most of the
geese had more than 2 laying-broodiness cycles. Sur-
prisingly, the broody time had a significant effect on
egg production, with a higher frequency of broody be-
havior resulting in more eggs. As such, our results in-
dicated that it was not feasible for Zhedong geese to
increase egg production by eliminating broody geese.
ACKNOWLEDGMENTS
This work was supported by the Agricultural ma-
jor new species creation project of Jiangsu Province
(PZCZ201735); and the Key Research and Develop-
ment Program of Jiangsu province (BE2017381); and
the Natural Science Foundation of China (31772583).
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