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Redalyc.Influence of the Estrous Cycle on Some Non Reproductive Interdisciplinaria ISSN: 0325-8203 [email protected] Centro Interamericano de Investigaciones Psicológicas y Ciencias Afines Argentina González Jatuff, Adriana Silvia; Torrecilla, Mariana; Quercetti, Marcela; Zapata, María Patricia; Rodríguez Echandía, Eduardo Luciano Influence of the estrous cycle on some non reproductive behaviors and on brain mechanisms in the female rat Interdisciplinaria, vol. 29, núm. 1, 2012, pp. 63-77 Centro Interamericano de Investigaciones Psicológicas y Ciencias Afines Buenos Aires, Argentina Available in: http://www.redalyc.org/articulo.oa?id=18026124004 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative INFLUENCE OF THE ESTROUS CYCLE ON SOME NON REPRODUCTIVE BEHAVIORS AND ON BRAIN MECHANISMS IN THE FEMALE RAT* ADRIANA SILVIA GONZÁLEZ JATUFF**, MARIANA TORRECILLA***, MARCELA QUERCETTI****, MARÍA PATRICIA ZAPATA*****, AND EDUARDO LUCIANO RODRÍGUEZ ECHANDÍA****** *Supported by grants from Universidad Nacional de Cuyo (UNCuyo) and founds from the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina. **Doctor in Biochemistry. Member of the Scientific Researcher Career of Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). ***Doctor in Psychology. Scholar post Doctoral of the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). E-Mail: [email protected] ****Degree in Biology. Área de Farmacología of the Facultad de Ciencias Médicas of the Universidad Nacional de Cuyo (UNCuyo). *****Member of Technichal Career of the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). ******Professor Emeritus of the Universidad Nacional de Cuyo (UNCuyo). Instituto de Medicina y Biología Experimental de Cuyo (IMBECU - CONICET). Área de Farmacología de la Facultad de Ciencias Médicas Universidad Nacional de Cuyo (UNCuyo). Avenida Ruíz Leal s/n. Parque General San Martín. (5500) Mendoza. República Argentina The authors wish to thank Dr. G. Jahn for the revision of the present manuscript. RESUMEN ciclo estrual. La presente revisión se informa en tres secciones. (1) Se presenta una breve descrip- El rol de las hormonas sexuales sobre el com- ción de la variación de tres hormonas sexuales portamiento reproductivo ha sido extensamente principales con acción directa sobre el SNC: es- documentado. La fluctuación periódica de hor- trógenos (E2), progesterona (P4) y prolactina monas sexuales en hembras de numerosas espe- (PRL). Se han descripto propiedades ansiolíticas cies ha sido relacionada con cambios comporta- para P4 y anti-estrés para PRL; para E2, la bi- mentales no sexuales tales como humor, ansie dad, bliografía es controvertida, describiéndose accio- agresión y respuesta a estrés. El sustrato bioló- nes excitatorias y anti-estrés. (2) Se informan gico-neural de estos cambios se basa en los cam- algunos cambios cerebrales y comportamentales bios que estas hormonas inducen en el Sistema que tienen lugar en cada estadio del ciclo estrual. Nervioso Central. Este trabajo resume algunos Las fluctuaciones hormonales se consideran bá- cambios que afectan a receptores y neurotrasmi- sicas para la interpretación de tales cambios. (3) sores de los sistemas GABAérgico, serotoninér- Se mencionan algunos hallazgos acerca de la in- gico, dopaminérgico y péptido liberador de pro- fluencia hormonal cíclica sobre los sistemas de lactina del SNC de la rata. La rata hembra posee neurotransmisión del SNC y sobre un nuevo pép- un ciclo sexual de 4 días de duración, denominado tido propuesto como mediador de la respuesta a INTERDISCIPLINARIA, 2012, 29, 1, 63-77 63 González Jatuff, Torrecilla, Quercetti, Zapata, and Rodríguez Echandía estrés, el péptido liberador de prolactina (PrRP). stage. A novel peptide, the Prolactin Releasing Esperamos que este trabajo contribuya a un mejor Peptide (PrRP), which is sensitive to E2 fluctu - entendimiento de los mecanismos del comporta- ation, has also been linked to the lower activity of miento de las hembras y sus variaciones, y sus im- the HPA axis. Stress-induced activation of PrRP plicancias farmacológicas y diagnósticas. neurons is significantly decreased in E compared with P and D, suggesting that E2 suppresses the Palabras clave: Ciclo estrual; Comportamiento activation of PrRP neurons. The late luteal phase no sexual; Rata; Hormonas; Mecanismos cere- (D) correlates with the premenstrual phase in brales. women, commonly associated with psychological disturbances, including mood disorders and increas- ed aggression. Consistently, increased levels of ABSTRACT anxiety and aggression have been detected in rats during D. Sex hormone fluctuations in females are in - 3.- Findings about cyclic hormone influences volv ed in some behavioral states such as mood, on the CNS neurotransmitters and on the stress anxiety, aggression and stress response, due to mediator, prolactine-releasing peptide (PrRP) functional changes in the central nervous system were described. GABA is the main inhibitory (CNS) activity induced by the cyclic sex hormone system in the brain. Estrous-cycle-dependent in- fluctuation. This review includes three sections. creases in δ-GABAA receptors were reported; 1.- A description of the three major hormone this subtype underlying a tonic inhibitory current, fluctuations in the estrous cycle: estrogens (E2), is the most sensitive target of P4 and ALLO. E2 progesterone (P4) and prolactine (PRL). E2 a - causes a reduction in GABAergic inhibition, le- chieves the maximum circulation levels during P. ading to an increase in the excitatory tone. It also E2 is mainly excitatory and has been considered to acts on hippocampus causing a transient lowering have an antidepressive action. The highest plasma of GABA synthesis in the interneuron. These fin- levels of P4 and its metabolite allopregnenolone dings suggest an excitatory role for E2 through in- (ALLO) occur in P. Ovulation takes place in the hibition of the GABA system. The serotonin night of P, and the resulting corpora lutea produces system, involved in behavioral responses such a secondary increase in P4 (and ALLO) on D. The as stress, anxiety and depression, also exhibits va- P4 peak level occurring in D1 and in the evening riations along the estrous cycle, in part dependant of P was shown to exert benzodiazepine-like of GABA-receptor changes. effects, including sedation. It has been proposed Gender differences were described for DA that ALLO, rather than P4 is the one acting on func tion in the brain, due to E2 and P4 modula- GABA systems. Circulating levels of ALLO tion. DA release and reuptake fluctuate with chan- parallel those of P4 across the estrous cycle and is ges in circulating steroid levels. E2 enhances DA known to have anxiolytic properties. release and DA-mediated behaviors, such as ge- PRL is produced mainly in the adeno hy poph - neral activity stimulation, food seeking behavior ysis, though synthesis in other sites of the brain also and sexual motivation. PrRP is produced in hypo- occurs. Its regulation is mostly inhibitory and is thalamic and extra hypothalamic structures. It has exerted by dopamine (DA) released in the hy - been propos ed as a mediator of stress responses. pothalamus. A surge in PRL secretion occurs during Though gender differences have been shown, dis- P. PRL would be a modulator of the HPA-axis, and tribution of PrRP does not change during the es- is considered as a stress hormone. trous cycle. 2.- Some behavioral and neural changes occur We hope this review may contribute to at each stage of the cycle. Lower anxiety level in understand the mechanism of female behav ioral P females was described. This has been correlated variations and their pharmacological and diagnostic with increased circulating levels of endogenous implications. ALLO in P. PRL is another hormone that may cause the lower scores of anxiety observed at P, Key words: Estrous cycle; Non sexual behav iour; since the peak of plasma PRL is observed at this Rats; Hormone; Brain mechanisms. 64 INTERDISCIPLINARIA, 2012, 29, 1, 63-77 Behavioral and brain changes in rats INTRODUCTION FLUCTUATIONS IN E2, P4 AND PRL LEVELS Behavioral responses in animals depend on Increasing number of reports are concerned external stimuli (experience) and endogenous with the cyclic fluctuations of the main ovarian factors. Age, gender and physiological state hormones, i.e. estrogen (17β-estradiol , E2) such as sexual receptivity, pregnancy or lac - and progesterone (P4), and the pituitary tation are among the main sources of normal peptide prolactin (PRL) in connection with the behavioral variations. In the majority of these functional changes occurring in some of the situations, a particular hormonal profile un - central neurotransmitter systems. Some of the derlie the changes in behavior; hormones major behavioral changes associated with modulate many brain receptors, their signaling these hormones are summarized in Table 1. mechanisms and neuro transmitter metabolism, E2 is produced by the ovarian follicles thus inducing changes in neural transmission under the follicle-stimulating hormone (FSH) and in synaptic and neural plasticity. In this stimulus and achieves the maximum circula- review we will focus on sexual hormones, tion levels during P. E2 is mainly excitatory, it particularly three hormones of the cycling
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