Domestic Animal Endocrinology 25 (2003) 93–99

Vasotocin and reproductive functions of the

domestic chicken
A. Jurkevich a , R. Grossmann b,∗
a Institute of Ecology, Vilnius University, Akademijos 2, Vilnius LT-2600, Lithuania
b Institute for Animal Science, Mariensee, Federal Agricultural Research Centre (FAL),
Höltystr. 10, 31535 Neustadt a. Rbge., Germany

Abstract
The neurohypophyseal hormone arginine vasotocin (AVT) combines both antidiuretic and re-
productive activities. In the domestic chicken AVT produces assimetric effects on the reproductive
functions of males and females. AVT synthesized in magnocellular diencephalic neurons is released
into circulation in a highly coordinated manner contributing to the peripheral control of oviposition
in hens. Conversely, parvocellular AVT cells located in the limbic system (bed nucleus of stria ter-
minalis (BST)) are quite different in their properties and, possible, functions. In domestic chickens
these cells express AVT in a sexually dimorphic manner and are found solely in males. This sexually
dimorphic part of the AVT system is sensitive to gonadal steroids. Experimental data demonstrated
that AVT modulates different aspects of reproductive behavior including courtship vocalization
and copulation. Sexual differentiation of these limbic vasotocinergic cells show striking correlation
with sexual differentiation of masculine behavior. Evidences coming from physiological, anatom-
ical and ethological studies suggest strong implication of the vasotocinergic system in the control
of reproductive functions.
© 2003 Elsevier Inc. All rights reserved.
Keywords: Vasotocin; Reproduction; Chicken; Behavior

1. Introduction

The neurohypophyseal hormone arginine vasotocin (AVT) in birds, likewise its ho-
mologue arginine vasopressin in mammals, has one of the most remarkable records of
documented autonomic and behavioral functions. Because of its important role played in
osmoregulation, AVT was defined as antidiuretic hormone in all terrestrial non-mammalian
species [1,2]. In birds, AVT shows vasomotor and thermoregulatory effects [3], stimulates

∗ Corresponding author. Tel.: +49-5034-871164; fax: +49-5034-871247.

E-mail address: roland.grossmann@fal.de (R. Grossmann).

0739-7240/$ – see front matter © 2003 Elsevier Inc. All rights reserved.
doi:10.1016/S0739-7240(03)00048-1
94 A. Jurkevich, R. Grossmann / Domestic Animal Endocrinology 25 (2003) 93–99

the release of adrenocorticotropic hormone from the pituitary [4,5] and reduces plasma al-
dosterone [3]. While the neurohypophyseal system of most mammalian species in addition to
the antidiuretic hormone produces also oxytocin that controls important reproductive func-
tions such as uterine contractility during partition and milk ejection, and induces some forms
of parental behavior, avian AVT combines both antidiuretic and reproductive functions.
Moreover, some recent neuroanatomical data point out on its role in regulation of sex-related
reproductive behavior [6,7]. The aim of this review is to summarize briefly the sex-related
functions of AVT emerging from recent experimental studies in the domestic chickens.

2. AVT and control of oviposition

Data obtained from a number of experimental studies have demonstrated that oviposition
in the hen is associated with an increase of AVT concentration in the circulation [8–10].
Plasma AVT increases sharply at the time of oviposition inducing the oviduct contractions
and decreases within 30 min after an egg has been laid [11]. Oviposition-related elevation of
plasma AVT is accompanied by a depletion of AVT concentration in the neurohypophysis
[12] suggesting that hormone is likely to be synthesized in hypothalamic magnocellular
neurons. However the neurohypophysis is not the only possible source of AVT release
during oviposition. AVT was also found to be expressed in the uterus and ovary of laying
hens [13]. It needs to be established whether this locally synthesized AVT acts in concert
with centrally produced hormone controlling egg lay or it may have a different role as a
paracrine regulator of the reproductive system.

3. Brain vasotocin system

In the avian neuroendocrine system, neurons producing AVT are obviously among the
most abundant. They are concentrated largely within the preoptic and supraoptic brain re-
gions and the paraventricular hypothalamic nucleus. Magnocellular vasotocinergic neurons
form the hypothalamo–neurohypophysial–neurosecretory system. AVT is synthesized in
these cells and transported along the axon terminals to the neurohypophysis where the pep-
tide is released into the blood stream. In the preoptic area and paraventricular nucleus of
the hypothalamus AVT is colocalized in the same cells with chicken gonadotropin releasing
hormone I [14] suggesting a possible coordinated response of both neuropeptides in control
of reproductive functions.
The AVT system includes also a sexually dimorphic population of parvocellular neurons
that are located around a conventional border dividing di- and telencephalon. These neurons
extend from the mediocaudal part of the preoptic region to the dorsolateral part of the bed
nucleus of stria terminalis (BST) (Fig. 1). The role of these brain regions in control of sexual
behavior in birds is well-documented [15–17]. Additionally, a dense sexually dimorphic
network of AVT-immunoreactive (AVT-ir) fibers is observed in the lateral septum (SL)
[18]. In a few bird species investigated so far, males have more AVT-ir cells in the BST and
more abundant AVT-ir innervation in the BST and SL than females. In chicken and quail
these sex differences are ultimate, with complete absence of the AVT-ir structures in the
BST and SL of adult females (Fig. 2) [18,19].
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Fig. 1. Low magnification photomicrograph of a 40-␮m-thick coronal brain section through the septo-preoptic
region of adult cockerel showing the distribution of arginine vasotocin immunoreactive cells and fibers. CO,
optic chiasm; BSTdl, bed nucleus of the stria terminalis, pars dorsolateralis; BSTvm, bed nucleus of the stria
terminalis, pars ventromedialis; LHy, lateral hypothalamus; OM, occipitomesencephalic tract; nCPa, nucleus of
pallial commissure; PVN, paraventricular nucleus; SL, lateral septal nucleus; SM, medial septal nucleus; VL,
lateral ventricle; 3V, third ventricle. Scale bar = 200 ␮m.

Contrary to the magnocellular neurosecretory system, parvocellular vasotocin neurons

may release peptide directly into brain interstitial fluid where it modulates the neural circuits
controlling behavior. Such functions were demonstrated by vasopressinergic parvocellular
neurons located in the homologous limbic structures (amygdala, BST) of the rat brain [20].
The synthesis of AVT in the sexually dimorphic parvocellular neurons of the Japanese
quail is sensitive to testosterone but not to non-aromatisable androgen, 5␣-dihydrotesto-
sterone. Lately strong evidences were provided that aromatization of testosterone into
estradiol is essential to stimulate AVT synthesis [21].

4. Development of sex differences in the vasotocin system and sexual behavior

Developmental studies in chickens have demonstrated that AVT neurons arise within
the BST during embryogenesis in both sexes [22]. First conspicuous AVT-ir cells can be
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Fig. 2. Medium magnification photomicrographs of coronal brain sections illustrating the distribution of AVT-ir
cells and fibers in the brain of adult male (A) and female (B) domestic chicken. For abbreviations see Fig. 1. Scale
bar = 100 ␮m.

detected already around day 12 of embryonic (E) development. During further development
the number of ir cells increases progressively reaching the maximal values both in males
and females around hatching. The number of AVT-producing cells in the BST completes
differentiation by day 35 posthatch.
Recent studies indicated that sexual differentiation of the AVT system is likely to be in-
duced by estrogen-dependent mechanisms. Adult male quail or chickens treated as embryos
on E8–E9 with an in ovo injection of estradiol benzoate have virtually completely lost the
AVT-ir structures in the BST, medial preoptic nucleus and septum, i.e. they demonstrated a
female-like distribution of vasotocinergic fibers and parvocellular perikarya [23,24]. Con-
versely, the females during the same period of development received single in ovo injections
of an aromatase inhibitor that prevents synthesis of endogenous estradiol from testosterone
displayed a male-typical distribution of AVT-ir structures. Injections of either estradiol or
aromatase inhibitor are not effective when given on E18, i.e. much beyond the critical period
for sexual differentiation of behavior [24]. Moreover, experimental manipulations to alter
estrogen milieu do not significantly affect the magnocellular component of the AVT system
outside of the BST. These facts indicate that embryonic effects of estrogens on the AVT
system are time- and region-specific.
It is important to note that the embryonic differentiation of the AVT system in chickens
and quail demonstrates striking correlation with the differentiation of sexual behavior in
these species. In gallinaceous birds, estrogens play a key role in differentiation of sexual
 A. Jurkevich, R. Grossmann / Domestic Animal Endocrinology 25 (2003) 93–99 97

behavior. The high levels of estrogens typical to female embryos permanently suppress
female capacities to display masculine behavior at adulthood [7]. Male embryos exposed
to estradiol as earlier as E12–E13 loss permanently their abilities to copulate at adulthood
[25,26]. Detail ethological analysis in chickens revealed no significant changes in the ap-
petitive components of male sexual behavior (courtship) while consummatory components
(mounting attempts, treading, and copulation) were suppressed. Curiously, the frequency
of aggressive displays towards females has increased. Moreover, estradiol reduced the
frequency of crowing and changed some acoustic characteristics of this call.
These results along with extensive sexually dimorphic AVT innervation of the brain
areas known to be involved in the control of reproductive behavior strongly suggest a
role of this peptide in sexual behavior. Available data indicate indeed that AVT regulates
male sexual behavior in chickens, pigeons and Japanese quail [27,28], controls courtship
singing [29,30], facilitates aggressive behavior in colonial songbirds but inhibits aggression
in songbirds with a territorial social organization [30,31].

5. Conclusion

It is a scientific tradition to consider AVT as endocrine regulator of osmotic homeosta-

sis. The results reviewed here obviously demonstrate that AVT plays a significant role in
different aspects of reproduction of domestic chickens by modulation of both peripheral
and central mechanisms. A surge of AVT released from the hypothalamus evokes, probably
in interrelation with some other regulatory factors (see [32]), the oviposition in hen. On
the other hand, AVT expression in parvocellular extrahypothalamic neurons depends from
the levels of gonadal hormones and correlates with male-typical sexual behavior. These
central regulatory effects on the reproductive axis may be mediated by interactions with
other neuroendocrine systems such as GnRH-I and/or galanin.

Acknowledgements

Supported by H.W. Schaumann Stiftung.

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