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Molecular Characterization of Putative Calcitonin Gene-Related Peptide

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Molecular Characterization of Putative Calcitonin Gene-Related Peptide International Journal of Zoological Investigations Vol. 4, No. 2, 107-116 (2018) _______________________________________________________________________________________ International Journal of Zoological Investigations Contents available at Journals Home Page: www.ijzi.net ISSN: 2454-3055 Molecular Characterization of Putative Calcitonin Gene-Related Peptide Receptors and Expression of Calcitonin Gene-Related Peptide and its Receptor in the Early Development of Flounder, Paralichthys olivaceus Sekiguchi Toshio1, Suzuki Tohru2, Kurokawa Tadahide3, Amornsakun Thumronk4, Hai Tran Ngoc5, Srivastav Ajai K.6 and Suzuki Nobuo1* 1Noto Marine Laboratory, Division of Marine Environmental Studies, Institute of Nature and Environmental Technology, Kanazawa University, Ogi, Noto-cho, Ishikawa 927-0553, Japan 2Laboratory of Marine Life Science and Genetics, Graduate School of Agricultural Science, Tohoku University, Sendai 980-0845, Japan 3Hokkaido National Fisheries Research Institute, National Research and Development Agency, Kushiro Laboratory, Kushiro 085- 0802 Japan 4Fisheries Technology Program, Faculty of Science and Technology, Prince of Songkla University, Pattani Province 94000, Thailand 5College of Aquaculture and Fisheries, Can Tho University, Can Tho City, Vietnam 6Department of Zoology, D.D.U. Gorakhpur University, Gorakhpur 273009, India *Corresponding Author Received: 6th June, 2018 Accepted: 28th June, 2018 ______________________________________________________________________________________________________________ Abstract: Calcitonin gene-related peptide (CGRP) is a 37-amino acid neuropeptide that has been identified in teleosts and mammals. Calcitonin receptor-like receptor (CLR) forms the CGRP receptor. In teleost fish, CGRP plays a role in vasodilation and is associated with osmotic regulation. However, the embryonic function of CGRP remains unclear. Therefore, we have investigated CGRP function related to early development in the flounder, Paralichthys olivaceus. We first identified two CLRs from the flounder genome database. According to molecular phylogenetic analysis, two CLRs are classified in the group formed by teleost CLR1 and 2. They represented high amino acid sequence identity among mefugu CLR1-3 and human CLR. We also identified another CLR/CTR candidate from the flounder genome. Molecular phylogenetic analysis revealed that this gene was located at the clade formed by vertebrate CTRs that are CLR paralogs. The deduced protein of the gene also had four hormone-binding domains, which are the typical feature of teleost CTR, and was completely different from the CLR of teleosts, including flounder CLRs. These data strongly suggest that flounder possess two putative CGRP receptors. Finally, we performed the expression analysis of flounder CGRP and two CLRs in the flounder embryo. Both expressions of CGRP mRNA and two CLR mRNAs were detected from 96 hours post fertilization (hpf). These results demonstrate that CGRP has some functions at 96 hpf. At that stage, embryos have already hatched and the circulatory system functions, whereas digestive organs still continue to develop. Taken together, these findings suggest that the CGRP peptide may play a role in the circulatory system, as in adult fish, or it may be associated with the development of the digestive tract. Keywords: Calcitonin gene-related peptide, Calcitonin receptor-like receptor, Calcitonin receptor, Flounder ______________________________________________________________________________________________________________ 107 Introduction Calcitonin gene-related peptide (CGRP) is a CGRP level was induced by the transfer from 37-amino acid peptide. The CGRP gene freshwater to seawater in the eel, Anguilla encodes both CGRP and the calcitonin (CT) anguilla (Lafont et al., 2006). Although the peptide in the different exons (Amara et al., physiological function of adult teleosts has 1982; Rosenfeld et al., 1983). In mammals, been evaluated, but the function of CGRP in CGRP and CT mRNA are expressed in the early or larval development remains unclear central and peripheral nervous systems and in both mammals and teleosts. thyroid calcitonin cells (C cells), respectively, The canonical CGRP receptor comprises by alternative splicing (Amara et al., 1985; the CT receptor-like receptor (CLR) (Hay and Rosenfeld et al., 1983). CGRP genes and this Walker, 2017; Hay et al., 2018). On the other type of post-transcriptional process are also hand, the CT receptor (CTR) is paralogous to observed in teleosts (Takei et al., 2010). CLR. CLR and CTR belong to the secretin Mammalian CGRP plays a role in family G protein-coupled receptor (GPCR). vasodilation (Brain et al., 1985; Brain and They contain seven transmembrane domains Grant, 2004). It is also associated with pain and hormone-binding domains (HBDs) that transmission because of CGRP synthesis in the are responsible for binding to the ligand sensory nerve (Winston et al., 2005). Notably, peptide (Parthier et al., 2009). Both receptors the CGRP peptide of the trigeminal nerve has show close phylogenetic relationships in garnered attention as a cause of migraines vertebrates (Martins et al., 2014). Therefore, (Hay and Walker 2017; Edvinsson et al., discrimination between CLR and CTR is 2018). The teleost CGRP peptide also required to clarify the CGRP receptor. represents vasodilatory function (Kagstrom We previously cloned and determined the and Holmgren, 1998; Shahbazi et al., 2009). In nucleotide sequence of a CGRP gene in the rainbow trout, adrenaline-inducing flounder, Paralichthys olivaceus (Suzuki et al., precontracted small diameter gut arteries 2001). The flounder CGRP gene encoded both were relaxed by treatment with chicken CGRP. CGRP and the CT peptide in different exons as This reaction was antagonized by the N- in another teleost. In addition, we also terminal truncated CGRP8–37 (Kagstrom and identified the nucleotide sequence of a Holmgren, 1998). Cod CGRP induced dose- putative CLR-type gene and CTR-type gene dependent relaxation of celiac arteries precontracted with adrenaline in Atlantic cod from a flounder gill cDNA library and clarified Gadus morhua (Shahbazi et al., 2009). their tissue distribution (Suzuki N et al., Interestingly, CGRP is implicated in fluid 2000). Although the deduced amino acid homeostasis (Suzuki et al., 2002; Lafont et al., sequences of both receptors were similar to 2006). In flounder, CGRP mRNA was that of vertebrate CLR, the CTR-type receptor expressed in the gill in seawater conditions. resembled vertebrate CTR because of its Transferring flounder from seawater to expression in the vertebral bone (Suzuki N et freshwater decreased the expression of CGRP al., 2000). However, there was no evidence mRNA to an undetectable level (Suzuki et al., that the CLR-type and CTR-type genes are 2002). Moreover, an increase in the plasma vertebrate CLR and CTR orthologs, 108 respectively. Therefore, the CGRP receptor has obtained by the data mining were used for yet to be determined. molecular phylogenetic analysis and sequence comparison. In the present study, we comprehensively annotated CLR and CTR from the flounder Molecular phylogenetic analysis of flounder genome. First, candidates of two CLRs were CLR/CTR extracted from the flounder genome database We performed molecular phylogenetic using two previously unannotated sequences analysis with modifications of the previous as queries. Moreover, a CTR candidate also study (Sekiguchi et al., 2016). Multiple was detected from the flounder genome. To alignments of amino acid sequences were identify the CGRP receptor candidate, generated by using the Muscle program of flounder CLR and CTR were classified by MEGA 7.0.26 software (Kumar et al., 2016). molecular phylogenetic analysis and Next, we manually removed the gap and molecular characteristic analysis. To gain constructed unrooted molecular phylogenetic clues as to the CGRP function of early flounder trees using the neighbor-joining method with development, we finally performed an MEGA 7.0.26 software. The bootstrap value expression analysis of flounder CGRP and CLRs in flounder embryos. was calculated by 500 pseudo-replications using MEGA 7.0.26 software. We performed Materials and Methods molecular phylogenetic analysis of CLR/CTR. Experimental Materials The accession numbers of the CLR and CTR Fertilized eggs of the flounder, Paralichthys are as follows: cattle CLR, NP_001095577.1; olivaceus, were maintained in a 50 L tank elephant shark CLR, XP_007888311; flounder supplied with running seawater (17 ± 1 C) CLR1, BAA92817; flounder CLR2, Q9IB86; circulating at a flow rate of 500 ml/min. human CLR, NP_005786; medaka CLR1, Embryos were collected at 0 (immediately XP_020560071; mefugu CLR1, BAE45312; after fertilization), 24, 48, 72, 96, 120, and 144 mefugu CLR2, BAE45313; mefugu CLR3, hours post fertilization (hpf). Embryos were BAE45314; mouse CLR, NP_061252;pig CLR, frozen at -80 C for reverse transcription- Q867C1; red stingray CLR-like, BAL63062; polymerase chain reaction (RT-PCR). tropical clawed frog CLR, Q28DX2; zebrafish Data mining of flounder CLR and CTR CLR1, NP_001004010; cattle CTR, NP_001069737.2; chicken CTR, XP_425985; Candidates of flounder CLR were searched by elephant shark CTR, XP_007903520; flounder TBLASTN algorithms from RefSeq RNA in the CTR, XP_019945977; green anole CTR, Japanese flounder genome database (https://blast.ncbi.nlm.nih.gov/Blast.cgi?PRO XP_008110735.1; human CTR, NP_001733; GRAM=tblastn&PAGE_TYPE=BlastSearch&BL medaka CTR, XP_011479129; mefugu
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