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Histochemical Differences Along the Intestine of Corydoras Paleatus (Siluriformes: Callichthyidae)

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Histochemical Differences Along the Intestine of Corydoras Paleatus (Siluriformes: Callichthyidae) Histochemical differences along the intestine of Corydoras paleatus (Siluriformes: Callichthyidae) Silvia E. Plaul1*, Claudio G. Barbeito1 & Alcira O. Díaz2 1. Laboratorio de Histología y Embriología Descriptiva, Experimental y Comparada, Facultad de Ciencias Veterinarias, UNLP. La Plata, Buenos Aires, Argentina; [email protected], [email protected] 2. Laboratorio de Histología e Histoquímica. Departamento de Biología. Instituto de Investigaciones Marinas y Costeras (IIMyC), Facultad de Ciencias Exactas y Naturales, CONICET. UNdMP. Mar del Plata, Buenos Aires, Argentina; [email protected] * Correspondence Received 27-II-2015. Corrected 20-VII-2015. Accepted 25-VIII-2015. Abstract: The Neotropical catfish Corydoras paleatus is a facultative air-breather and the caudal half of the intestine is involved in gas exchange. In South America, air-breathing fishes are found in tropical or sub-tropical freshwaters where the probability of hypoxia is high. The aim of this study was to characterize by traditional histochemical and lectinhistochemical methods the pattern of carbohydrate in the intestinal mucosa. Intestine samples were taken from 25 healthy adult specimens collected in Buenos Aires (Argentina). Samples were fixed by immersion in 10 % buffered formalin and routinely processed and embedded in paraffin wax. Subsequently, these sections were incubated in the biotinylated lectins battery. Labeled Streptavidin-Biotin (LSAB) system was used for detection, diaminobenzidine as chromogen and haematoxylin as a contrast. To locate and distinguish glycoconjugates (GCs) of the globet cells, we used the following histochemical methods: PAS; PAS*S; KOH/ PA*S; PA/Bh/KOH/PAS; KOH/PA*/Bh/PAS; Alcian Blue and Toluidine Blue at different pHs. Microscopically, the general structure of vertebrate intestine was observed and showed all the cell types characteristic of the intestinal epithelium. The cranial sector of catfish intestine is a site of digestion and absorption and its structure is similar to other fish groups. In contrast, enterocytes of the caudal portion are low cuboidal cells; and between these, globet cells and capillaries are observed, these latter may reach the mucosal lumen. Underlying the epi- thelium, observed a well-developed lamina propria-submucosa made of connective tissue; this layer was highly vascularized and did not exhibit glands. According to histochemistry, the diverse GCs elaborated and secreted in the intestine are associated with specific functions in relation to their physiological significance, with special reference to their role in lubrication, buffering effect and prevention of proteolytic damage to the epithelium together with other biological processes, such as osmoregulation and ion exchange. The lectinhistochemical analysis of the intestinal mucosa reveals the presence of terminal residues of glucose, mannose and galactose. In conclusion, this study has shown that GCs synthesized in the intestine of C. paleatus exhibit a high level of histochemical complexity and that the lectin binding pattern of the intestinal mucosa is characteristic of each species and the variations are related with the multiple functions performed by the mucus in the digestive tract. The information generated here may be a relevant biological tool for comparing and analyzing the possible glycosidic changes in the intestinal mucus under different conditions, such as changes in diet or different patho- logical stages. Rev. Biol. Trop. 64 (1): 327-340. Epub 2016 March 01. Key words: Corydoras paleatus, glycoconjugates, histochemistry, intestine, lectinhistochemistry, goblet cells. Teleosts have developed a great variety ecological variations, the general morphologi- of mechanisms to inhabit all existing aquatic cal pattern of teleost’s intestine corresponds environments and feed both on animals and with that of other vertebrates, which is divided plants (Wilson & Castro, 2011). Despite the into regions that cannot always be distin- differences among species resulting from these guished macroscopically (Cao & Wang, 2009; Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (1): 327-340, March 2016 327 Leknes, 2009; Cao, Wang, & Song, 2011). The aim of this study was to character- Microscopically, in the great majority of fish ize the carbohydrate pattern composition of species, intestinal wall is described by three mucins secreted by cells of the tunica mucosa (Çinar & Şenol, 2006) or four tunics (Olaya et in different regions of the intestine of C. palea- al., 2007; Wilson & Castro, 2011; Xiong et al., tus. Furthermore, lectinhistochemical (LHC) 2011), depending on whether the tunica sub- technique is used to locate, identify and dif- mucosa is considered together with the tunica ferentiate variations in the saccharide residues mucosa or not. present in the glycoconjugates (GCs) with the Mucus, the product of the hydratation aim to discriminate different cell populations of mucins, is a common feature in the tunica (Barbeito, Ortega, Matiller, Gimeno, & Sal- mucosa of vertebrates; it is secreted by the vetti, 2013). goblet cells or mucous cells (MC) (Yashpal, Kumari, Mittal, & Mittal, 2007; Díaz, García, MATERIALS AND METHODS Figueroa, & Goldemberg, 2008a; Díaz, García & Goldemberg, 2008b; Tano de la Hoz, Flami- Animals: Twenty-five healthy adult speci- ni, & Díaz, 2012). These cells secrete a variety mens (15 females and 10 males) of C. paleatus of mucins that contribute to the generation (LS: 62.22 ± 1.84 mm for females and 51.70 of a gelatinous layer, to protect and lubricate ± 1.54 mm for males) were used. The field the intestinal surface to facilitate the transport work was carried out in streams situated in La of food and reduce the epithelial mechanical Plata, Buenos Aires, Argentina. The handling, stress. Recent studies of molecular biology collection and slaughter process of all individu- indicate that MC also participates in signaling als followed the guidelines of the American pathways leading to a coordinated response for Fisheries Society (A.F.S., 2004). Fish were proliferation, differentiation, apoptosis and cell transported alive in plastic bags, and were kept adhesion (Pérez-Sánchez et al., 2013). Several in an aquarium at 22° C and pH 7.0 (Gómez, 1996) for a minimum of three weeks, they were authors have studied the mucins distribution fed at sunset with food for bottom fish (Tetra and their role in the process of digestion, in min tropical tablet, Germany). The animals the local immune responses and antimicrobial were slaughtered by an anesthesia overdose action for defense against a wide range of inju- using a Eugenol (30 mg/L) (García-Gómez, de ries (Carraway, Ramsauer, Haq, & Carraway, la Gándara, & Raja, 2002). 2003; Sasaki, Ikeda, & Nakanuma, 2007). The identification of the different types of GCs Sampling: The necropsies were carried present in tissue sections can be made using out immediately after sacrifice, the digestive conventional histochemical techniques (Díaz tract was rapidly removed and samples were et al., 2008b). fixed by immersing in 10 % buffered formalin Air-breathing fishes are commonly found before the dehydration in an ethanol series and in tropical or sub-tropical fresh waters that are xylene, and posteriorly embedded in paraffin. ambient ion-poor and prone to hypoxia and Samples of intestine were longitudinally divid- hypercarbia (Shartau & Brauner, 2014). The ed into three parts (cranial, middle and caudal Neotropical catfish Corydoras paleatus Jenyns portions) considering the shape, transparency, 1842 is a facultative air-breather and the caudal thickness of the wall and the width of the lumen half of the intestine is involved in gas exchange in transverse sections. The cranial portion or (Podkowa & Goiakowska-Witalinska, 2002). intestine 1 (I1) has a typical tubular form and To date, despite the commercial importance its wall is opaque; the middle area, which was of this species like ornamental fish, there subdivided into two portions, intestine 2 (I2) have been few studies on the digestive tract and intestine 3 (I3), has a broader lumen and its of C. paleatus. wall became thinner and more translucent than 328 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (1): 327-340, March 2016 I1; in the caudal portion or intestine 4 (I4), the decreasing ethanol series, and incubated in lumen diameter kept increasing and the wall 0.03 % H2O2 in methanol for 30 min at room was even thinner. Histological sections were temperature, to inhibit the endogenous per- cut using a sledge microtome, prepared accord- oxidase. Sections then were treated with 0.1 ing to standard protocol, and then stained using % bovine serum albumin in PBS for 30 min routine techniques haematoxylin and eosin and incubated with fifteen Biotinylated lectins (H-E) and Masson trichrome staining for mor- for 2 h at room temperature. After that, sec- phology (Martoja & Martoja, 1970). tions were washed separately and subsequently treated with an avidin-biotin-peroxidase com- Traditional histochemistry: Sections of plex (ABC kit, Vectastain Elite PK 6 200). tissue were also subjected to histochemical Diaminobenzidine (DAB) 0.02 % (Biogenex, procedures for GCs identification, as detailed San Ramón, CA) was used with chromo- in Table 1. The combined technique AB pH 2.5/ gen. Slides were counterstained with Mayer’s PAS allowed the identification of three groups haematoxylin. Staining intensity was graded of goblet cells. With regard to this, was calcu- according to a semiquantitative range (-)
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