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Galanin Immunoreactivity in the Brain of the Desert Lizard Uromastyx Acanthinura During Activity Season

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Galanin Immunoreactivity in the Brain of the Desert Lizard Uromastyx Acanthinura During Activity Season FOLIA HISTOCHEMICA ORIGINAL STUDY ET CYTOBIOLOGICA Vol. 51, No. 1, 2013 pp. 45–54 Galanin immunoreactivity in the brain of the desert lizard Uromastyx acanthinura during activity season Sadjia Benmansour Hammouche1, Mohammed Bennis2 1Aride Area Research Laboratory, Biological Sciences Faculty, University of Sciences and Technology of Houari Boumediene, Bab Ezzouar, Algeria 2Pharmacology, Neurobiology and Comportement Laboratory, Departement of Biology, Sciences Faculty, University of Cadi Ayyad, Marrakech, Morocco Abstract: The distribution of galanin immunoreactive perikarya and nerve fibers in the brain of the desert lizard U. acanthinura was studied by means of immunofluorescence using an antiserum against rabbit galanin. The animals were captured during the activity season in March (wet season) just before reproduction period and in June (arid season) after ovulation period. Immunoreactive neurons were mostly detected in the mediobasal and the infundibular recess nuclei, the nucleus of the paraventricular organ, the paraventricular organ, the periven- tricular nucleus and in the anterior hypothalamus at the level of the periventricular nucleus, the paraventricular nucleus and the supraoptic nucleus. The differences in brain galanin expression between animals collected un- der both sets of environmental conditions indicated changes which occur during the annual and reproductive cycles. The wide hypothalamic and extrahypothalamic distribution of galanin immunoreactive fibers suggests that this peptide may have hypophysiotropic, neuromodulator and neurotransmitter roles in the lizard U. acan- thinura. (Folia Histochemica et Cytobiologica 2013, Vol. 51, No. 1, 45–54) Key words: Galanin, lizard, brain, reproduction, immunohistochemistry, environment, desert, U. acanthinura Introduction gonadotropins [4], prolactin [5], growth hormone [6] and somatostatin [7]. These regulatory actions of gala- Galanin is a 29–30 amino acid neuropeptide, which is nin are further supported by many reports that show present in the central nervous system (CNS) and pe- galaninergic innervation of hypophyseal secretory ripheral tissues. Its primary structure is highly con- cells in several vertebrate groups including mammals served among vertebrates (almost 90%), indicating [8-11]. Moreover, the expression of galanin is elevat- the importance of the neuropeptide. Galanin has ed following estrogen administration, neuronal acti- multiple biological effects and many studies have dem- vation, denervation and/or nerve injury, as well as onstrated its involvement in several hypothalamic and during development [1]. The use of immunohis- hypophyseal functions including feeding, seizure, tochemical methods revealed a wide distribution of pain, learning, memory and depression [1]. This pep- galanin in the brain of several vertebrate groups: tide seems also to be involved in the osmoregulatory mammals [12–15], birds [16, 17], reptiles [9, 18], am- response to water-restriction [2, 3]. Galanin exerts phibians [19–22] and fish [8, 23–25]. neuroendocrine effects by modulating the release of In the lizard P. s. sicula, galanin seemed to inter- act in the oviduct with estrogen, vasoactive intestinal polypeptide and oviposition [26] and its systemic ad- ministration was able to stimulate the thyroid gland Correspondence address: S.B. Hammouche both at morphological and physiological level [27]. Aride Area Research Laboratory, Biological Sciences Faculty, University of Sciences and Technology of Houari Boumediene, However, there is no knowledge of the physiological P.B. 39, El-Alia, Bab Ezzouar, Algiers. Algeria; role played by galanin in the CNS of reptiles. The e-mail: [email protected] galanin stimulates food intake in the goldfish C. au- ©Polish Society for Histochemistry and Cytochemistry Folia Histochem Cytobiol. 2013 www.fhc.viamedica.pl 10.5603/FHC.2013.007 46 S. B. Hammouche, M. Bennis Table 1. Weight, mouth cloaca length and sexual state of U.acanthinura used in the study March June Mouth-cloaca length [cm] Weight [g] Mouth-cloaca length [cm] Weight [g] 20/Adult (beginning of spermatogenesis) 314.5 19/Adult (spermatogenesis) 348 20.5/Adult (spermatogenesis) 295 18/Adult (previtellogenesis) 201.9 22/Adult (at the egg retention) 371 18.2/Adult (previtellogenesis) 221 17/Sub-adult (immature) 175 17/Sub-adult (immature) 154.3 17.2/Sub-adult (immature) 163 17.5/Sub-adult (immature) 182.4 ratus [28], while it does not affect feeding in the neo- for 1 h at 4°C. They were then incubated overnight at 4°C in natal chicks [29]. Despite their crucial position among the primary antibody (polyclonal antiserum raised in rabbit vertebrate phylogeny, there have been only two stud- against porcine galanin generously gift by Prof. M. Tramu, ies published on the distribution of galanin in a rep- Bordeaux University, France) diluted 1:400 in PBS-gelatine- tilian species, the turtle M. caspica [9] and the snake Triton. After four washes (10 min each) in PBS, the sec- B. jararaca [18]. tions were incubated at room temperature with the second- Although physiological data have been obtained ary antibody (IgG anti-rabbit FITC Dako, France) diluted about the role of galanin in the lizard, a deserticole 1:200 in PBS-gelatine-triton for 2–3 h. They were washed in and seasonal breeder, there are no data describing dis- PBS, spread and mounted in Mowiol (Calbiochem, France) tribution of galaninergic system in the brain of lizard and observed in epifluorescence microscope (Olympus, Ja- species. Therefore, we undertook study on the distri- pan) equipped with an excitation filter HQ480/40, dichroic bution of galanin throughout the entire brain of the filter Q505/lp, and emission filter HQ535/50 allowing to U. acanthinura by means of immunohistochemistry. observe green fluorescence. The pictures were digitalized with D76 Olympus color camera coupled to microcomput- Material and methods er system with DP controller software. The specificity of the rabbit serum anti-galanin has Animals. Lizards U. acanthinura used in this study were previously been evaluated and used to locate the gala- collected during two periods of the active season, in nin in the sheep diencephalon [12]. To check the spec- March just before reproduction: adult male (n = 1), adult ificity of the labeling, adjacent sections were incubat- female (n = 2) and sub-adult female (n = 2) and in early ed without primary antiserum. June after ovulation: adult male (n = 2), sub-adult male The size of perkaryons, measured across the long (n = 2) and adult female with eggs retention (n = 1). Their axis of cell bodies, and diameter of nuclei were ob- weight and the mouth-cloaca length were summarized in tained in sections of 50 mm thickness. The average the Table 1. Animals were anesthetized by intraperitoneal (mean) and SD were obtained for 9 to 20 of cells per injection of 25% urethane solution (5,7 mL/kg body weight) section and according to the total number of neurons and subsequently transcardially injected with 0,9% NaCl observed for each area. solution. Specimens were then perfused with 4% paraform- aldehyde in phosphate buffer 0.1M, pH 7.4 for 30 to 40 min. Results The brains were removed and postfixed for 24 h in the same fixative, then they were cryoprotected in 30% sucrose in The galanin immunoreactivity in lizard’s brain showed phosphate buffer 0.1M, pH 7.4 at 4°C during 24 h. After- bilateral symmetry. Since animals were killed at dif- wards, the fixed brains were frozen at –40°C, cut at –20°C ferent times of the active period of the annual cycle, on a freezing microtome on serial frontal sections of 50 mm seasonal variations had to be evaluated. thickness. The free-floating sections were collected in phos- phate buffer 0.1 M, pH 7.4. Neuronal perikarya Immunohistochemistry. The serial frontal sections were Two areas with high densities of galanin immunore- immunostained according to indirect immunofluorescence active neurons were observed in the brain of U. acan- method. Every free-floating section was pre-incubated in thinura. The first one was detected in the mediobasal phosphate buffered saline 0.1M, pH 7.4 (PBS) containing hypothalamus in the infundibular recess nucleus 0.1% of gelatine and 0.3% of Triton (PBS-gelatine-Triton) (RIN), the nucleus of the paraventricular organ ©Polish Society for Histochemistry and Cytochemistry Folia Histochem Cytobiol. 2013 www.fhc.viamedica.pl 10.5603/FHC.2013.007 Galanin immunoreactivity in the brain of the desert lizard Uromastyx acanthinura during activity season 47 (NPO), the paraventricular organ (PO) and the poste- (Figure 3). Some perikarya displayed an apical pro- rior periventricular nucleus (PPvN). The galanin im- cess directed toward the 3rd ventricle (Figure 4). We munoreactive perikarya were generally bipolar, large noted absence of labeling in the ependymal cells. At and mostly pear-shaped. The form of their nucleus the level of PO, the galanin perikarya were localized varied between round and oval with the occurrence of parallel to each other, with a vertical orientation to- a nucleolus. Some galaninergic perikarya displayed an ward the wall of the 3rd ventricle with majority dis- apical process directed toward the ventricle. In the playing an apical process directed toward the ventri- outer layer of the median eminence (ME), some im- cle (Figure 5). At the level of ME, some immunore- munoreactive perikarya were detected. active perikarya were detected in the outer cellular The second area of galanin-immunoreactive neu- layer (Figure 6). These last two structures were rela- rons, with lower density as compared to the first one, tively less densely innervated by galaninergic fibers was present in the anterior hypothalamus, above the than in RIN and NPO. optic chiasma at the level of the anterior PvN (APvN), In sub-adult females, the galanin-labeled perikarya the paraventricular nucleus (PVN) and the supraop- appeared less numerous than in the adult animals tic nucleus (SON). The majority of the periventricu- (Figure 7), in fact, their number was about 29 per sec- lar galanin-immunoreactive perikarya displayed an tion in an area of 8683 mm2, which was 53.7% less apical process directed toward the ventricle.
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