TRPV6在蛋鸡不同组织的表达分布及其在蛋壳腺钙离子转运中的作用

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英文题名：Expression and Localization of Trpv6in Different Tissues of Laying Hens and Its Effect on the Calcium Ion Transport in Eggshell Glang 作者：杨俊花 论文级别：博士 学科专业名称：临床兽医学 中文关键词：蛋鸡 ; TRPV6 ; 组织分布 ; 蛋壳腺 ; 钙离子 ; 产蛋循环 英文关键词：laying hen ; TRPV6 ; tissues distribution ; egg shell gland ; calcium ion ; oviposition cycle 学位年度：2010 导师：侯加法 学科代码：090603 学位授予单位：南京农业大学 论文提交日期：2010-11-01

摘要

TRPV6是瞬时性受体电位通道TRP超家族中的成员。广泛分布于哺乳动物Ca2+大量转运组织：肠道、肾脏、胎盘、骨骼等。是专门的上皮样通道,具有高度的Ca2+选择性,承担Ca2+向细胞内转运的限速通道。但TRPV6在蛋鸡上皮细胞Ca2+转运的重要作用还未曾报道,本文旨在研究TRPV6在蛋鸡体内的表达分布及其在蛋壳腺Ca2+转移中的重要作用,进而阐明产蛋期TRPV6在维持体内钙离子平衡的作用机制。试验Ⅰ钙离子跨细胞转运相关蛋白基因在蛋鸡不同肠段和肾脏中的表达
     10羽220日龄高产蛋鸡,断颈处死,采集不同肠段和肾脏组织。应用RT-PCR方法分别从蛋鸡不同肠段和肾脏组织扩增得到鸡TRPV6、CaBP-D28K和PMCA1b编码基因,然后将特异性片段连接到pMD18-T载体,测序分析扩增片段准确性；同时应用Real-time PCR方法检测不同肠段和肾脏中三种蛋白编码基因的表达量。结果表明：分别扩增得到426bp的TRPV6序列,789bp的CaBP-D28K序列以及644bp的PMCAlb序列,测序结果与GeneBank报道的预测序列一致。另外,TRPV6mRNA在十二指肠表达量最高,其次是空肠(P<0.05)、回肠(P<0.01)、盲肠(P<0.01)和直肠(P<0.01),’肾脏表达量显著低于十二指肠(P<0.05)；CaBP-D28K和PMCA lb mRNA在十二指肠、空肠、回肠段表达量较高,直肠和肾脏中表达量显著降低(P<0.05)。结果显示：在蛋鸡肠道和肾脏都存在钙离子跨细胞转运相关蛋白基因的表达,并提示蛋鸡肠道钙离子转移主要在小肠段完成；同时,TRPV6基因的成功扩增为深入研究蛋鸡肠道钙离子转运机制开辟了广阔的领域。
     试验ⅡTRPV6在蛋鸡不同肠段和肾脏中的表达分布
     试验动物同前。为探明TRPV6蛋白在蛋鸡肠道和肾脏中的组织学分布特点,采用石蜡切片、特异性的TRPV6抗体对高产蛋鸡不同肠段和肾脏组织进行免疫组织化学方法研究,同时采用western-blot方法研究TRPV6蛋白在这些部位的浓度变化。结果表明：在十二指肠、空肠、回肠、盲肠和直肠的刷状缘绒毛上皮均有TRPV6阳性表达分布,直肠阳性表达较少,肠隐窝或杯状细胞则偶有表达。在肾脏,TRPV6在近曲小管、远曲小管和连接小管均有阳性表达。western-blot结果指出十二指肠TRPV6含量最高,直肠含量最低,其他肠段位于二者之间。肾脏TRPV6含量也低于十二指肠。结果显示：TRPV6蛋白在蛋鸡整个肠道和肾脏上皮具有阳性表达,表明钙离子跨细胞转运在蛋鸡体内肠道钙离子吸收或肾脏重吸收都具有重要作用。
     试验ⅢTRPV6在蛋鸡不同生殖器官组织中的表达
     试验动物同前。观察TRPV6在蛋鸡卵巢、输卵管膨大部、峡部以及子宫的表达分布,本试验采用RT-PCR和免疫组化技术进行定性研究,同时采用Real-time PCR和western-blot方法观察其定量表达变化。结果表明：卵巢TRPV6蛋白主要分布在卵泡细胞、颗粒细胞以及卵泡膜细胞；在膨大部、峡部以及子宫主要分布在黏膜上皮的纤毛上皮顶端(面向管腔),在子宫黏膜上皮的基底层细胞也有少量分布；另外,与卵巢组织中TRPV6表达量相比,膨大部和峡部mRNA表达量显著降低(P<0.05),膨大部TRPV6蛋白含量显著降低(P<0.05),子宫组织中TRPV6mRNA水平低于卵巢组织,但蛋白表达量高于后者,且统计学分析没有显著差异。结果显示,TRPV6在卵巢、膨大部、峡部以及子宫位置均有表达,提示TRPV6不仅参与卵泡的发育、成熟,对蛋白以及蛋壳的形成也具有重要的意义。
     试验Ⅳ产蛋循环过程中TRPV6、CaBP-D28K(?)PMCA lb在蛋壳腺的动态表达
     研究钙离子跨细胞转运方式在蛋壳形成过程中的作用,本实验将40羽220日龄ISA蛋鸡分为5组,于产蛋后0h、2h、4.5h、8h、16h断头处死,采集蛋鸡蛋壳腺组织,运用Real-time PCR和western-blot方法检测产蛋循环过程中TRPV6、 CaBP-D28K、PMCA1b mRNA和蛋白浓度的动态变化。结果表明：卵子未进入蛋壳腺前(产蛋后0-4.5h),蛋壳腺内TRPV6、CaBP-D28K和PMCA lb mRNA表达水平较低,随后表达量逐渐升高,在蛋壳钙化过程中达到最大值(产蛋后16h),与0h相比,TRPV6和CaBP-D2gK mRNA表达差异均极显著(P<0.01)；另外,产蛋循环过程中,TRPV6、CaBP-D28K和PMCA lb蛋白浓度变化与mRNA变化一致,产蛋后16h到达最大值,其中CaBP-D28k蛋白浓度与0h相比,差异显著(P<0.05)。结论：产蛋循环可调控蛋壳腺内TRPV6、CaBP-D28K和PMCA1b的表达,并提示钙离子跨细胞转运途径在蛋壳钙化过程钙离子进入过程发挥重要作用。
     试验Ⅴ血浆钙磷浓度及其相关激素水平在产蛋循环中的动态变化
     40羽220日龄ISA蛋鸡分为5组,详细记录产蛋间隔时间,分别于产蛋后0h、2h、4.5h、8h、16h断头处死,采集血液,测定血浆总钙、总磷、甲状旁腺激素、降钙素、碱性磷酸酶和骨钙素水平。结果表明：在产蛋循环过程,血浆总钙、总磷浓度未发生显著变化；血浆甲状旁腺激素水平在产蛋后16h蛋壳钙化过程中达到最高水平,与产蛋后0h相比,差异显著(P<0.05)；血浆降钙素水平在产蛋后0h含量最高,后逐渐降低,在8h达到最低点(P<0.05)；产蛋后8h,血浆碱性磷酸酶活性达到最高,与Oh相比,差异显著(P<0.05)；另外,血浆骨钙素水平在产蛋后4.5h到达最高,但差异不显著。结论：在产蛋循环过程中,与钙磷代谢相关的钙调激素以及血液生化指标都呈现相应的动态变化。
The ion channel TRPV6is one member that belongs to the TRP super-family of cation channel, and distributed in calcium-transporting tissues such as intestine, kidney, placenta and bone of mammals, all of which are characterized by high Ca2+transport requirements. TRPV6is known as highly Ca+-selective channels, and serves as an important rate-limiting step in facilitating the entry of Ca2+into the epithelial cells. However, the presence of TRPV6in avian tissues is unclear. Therefore, the objective of the present study was to determine the distribution and expression patterns of TRP V6for different tissues in the laying hens, and investigate the role of TRPV6during the eggshell calcification. This information is important to fully understand the physiological role of this Ca2+channel in maintaining Ca2+homeostasis during eggshell mineralization in laying hens.
     Experiment I The mRNA expression of proteins involved in the Ca2+transcellular transport in the different intestinal segments and kidney in laying hens
     Ten ISA laying hens at the peak stage (220days old) were decapitated for sampling. To obtain the encoding gene of gallus TRPV6, CaBP-D28K and PMCA lb from the different intestinal segments and kidney in laying hen, the RT-PCR method was carried out. Then these special DNA fragments were cloned into pMD18-T vector and the sequence was determined. In addition, TRPV6, CaBP-D28K and PMCA1b mRNA expression in the kidney and different intestinal segments were quantified by Real-time PCR. The results were as follows: standard RT-PCR with TRPV6, CaBP-D28K and PMCA1b primers yielded three single amplified fragments of the expected size of426bp,789bp and644bp from all intestinal segments and the kidney, respectively. Sequence analysis revealed a100%homology at the nucleotide level with the predicted mRNA sequence of Gallus Gallus TRPV6、CaBP-D28K and PMCA lb. Real-time PCR analysis demonstrated that, among these tissues examined, the highest level of TRPV6expression was in the duodenum, then jejunum (P<0.05), kidney (P<0.05) and ileum (P<0.01). The lowest level of expression was found in the cecum and rectum samples (P<0.01). Furthermore, the levels of CaBP-D28K and PMCA1b were high in duodenum, jejunum and ileum, but the lower levels of expression were showed in the rectal and renal samples compared with duodenum (P<0.05), respectively. In conclusion, three calcium-transporters involved in the Ca2+transcellular transport were expressed in the intestine and kidney in laying hens, suggestting that the upper intestine (duodenum and jejunum) is the main site of Ca2+absorption in the birds, and the expression of TRPV6supplys new basis to do research in mechanism of calcium transport in laying hen.
     Experiment Ⅱ Localization and expression of TRPV6in different intestinal segments and kidney in laying hens
     The experimental animals were the same with the above one. Immunohistochemical analysis was used to investegate the TRPV6localization and expression in different intestinal segments and kidney in laying hens during peak laying. In addition, changes of concentration of TRPV6protein among these tissues were obtained by the western-blot analysis. The results are as follows:TRPV6was localized to the brush-border membranes of the duodenum, jejunum, ileum, cecum and rectum. Expression was weaker in the rectum and little or no expression was found in crypt and goblet cells. In the kidney, distinct immunopositive staining for TRPV6was detected at the apical domain of the proximal convoluted tubules (PCT), distal convoluted tubules (DCT) and medullary connecting tubules (CNT). Furthermore, western-blot analysis indicated that the duodenum expressed the greatest amount of TRPV6and the rectum the least, the other segments expressing intermediate levels. Moreover, the kidney expressed lower TRPV6protein levels compared to duodenum. In conclusion, the epithelial Ca2+channel TRPV6is strongly expressed in the apical cells of the entire intestine and the renal tubules suggesting that active Ca2+transcellular transport plays a crucial role in dietary calcium (re)absorption in laying hens.
     Experiment Ⅲ The expression of TRPV6in the different reproductive organs in laying hens
     The experimental animals were the same with the above one. Immunohistochemical analysis and RT-PCR were used to investegate the TRPV6localization and expression in the ovarium, magnum, isthmus and uterus in laying hens. In addition, quantitation of mRNA level and protein concentration of TRPV6in these tissues were carried out by Real-time PCR and western-blot analysis, respectively. The results are as follows:positive expression of the TRPV6protein and the TRPV6mRNA was observed in the ovarium, magnum, isthmus and uterus in laying hens. TRPV6was localized to the granulose cells and theca cells and the oocytes in the ovarium. Furthermore, intense staining was observed in villus epithelial cell (face to lumen) in mucous epithelium of the magnum, isthmus and uterus, and the weaker staining was also found in the basilar layer of secretory cell of endometrium in the uterus. Moreover, the levels of TRPV6mRNA in the magnum and isthmus were significantly decreased compared with that in the ovarium (P<0.05). The expression of TRPV6in the magnum was also lower than that in the ovarium (P<0.05). However, the expression of TRPV6mRNA in the uterus was lower than that in the ovarium, but the change of protein concentration was in contrast, and the difference was not significent. In conclusion, the expression of TRPV6in these tissues implyed that the TRPV6is not only involved in the development and maturation, but also plays an important role in albumen secretion and eggshell calcification.
     Experiment IV Dynamic expression of TRPV6, CaBP-D28K and PMC A lb in the ESG during the oviposition cycle in laying hens
     Forty ISA laying hens at the peak stage (220days old) were assigned to five reproductive stages for sampling. The samples of ESG were obtained by decapitated at0,2,4.5,8and16h after post-oviposition, respectively. Thus, the aim of this study is to investigate the effect of active Ca2+transcellular transport on the eggshell calcification. In addition, quantitation of mRNA level and protein concentration of TRPV6, CaBP-D28K and PMCA lb at different stages in the ESG were carried out by Real-time PCR and western-blot analysis, respectively. The results are as follows:the expression levels of TRPV6, CaBP-D28K and PMCA lb mRNA in the ESG were retained very low until the egg movement into the shell gland (0-4.5h after ovulation), then significantly increased at16h during eggshell calcification. In addition, TRPV6, and CaBP-D28K indicated significant statistical difference (P<0.01), respectively. Furthermore, western blotting showed that the expression of TRPV6and PMCA1b reached the maximum at16h after ovulation, but the statistical difference was not significant. The change of CaBP-D28K expression was very similar to that of TRPV6, but the concentration was significantly increased at16h than that at0h after ovulation (P<0.05). In conclusion, the expression of TRPV6, CaBP-D28K and PMCA lb in the ESG was regulated by the oviposition cycle, suggestting that active Ca2+transcellular transport exerted significant effects in calcium delivering in the ESG.
     Experiment V Dynamic changes of levels of plasma total Ca, P and calcium-regulated hormones during the oviposition cycle
     Forty ISA laying hens at the peak stage (220days old) were assigned to five reproductive stages for sampling. Production records were kept before the study. Blood were obstained by decapitated at0,2,4.5,8and16h after post-oviposition, respectively. Thus, the aim of this study is to investigate the concentration changes of plasma total calcium, phosphorus, alkaline phosphatase (ALP), parathyroid hormone (PTH), calcitionin (CT) and osteocalcin (BGP) during the oviposition cycle. The results are as follows:there were not significant changes of total calcium and phosphorus after oviposition. The level of plasma CT was the highest at0h and the lowest at8h (P<0.05) after oviposition, the change of plasma PTH level was antithesis to that of plasma CT, which was constant slightly increase after oviposition, and reached the maximum at16h during the eggshell calcification (P<0.05). In addition, the level of plasma ALP activity was notablely increased at8h after oviposition (P<0.05). Moreover, the concentration of plasma BGP reached the maximum at4.5h after oviposition. In conclusion, the calcium-regulated hormones and biochemical indicators involved in the metabolism of calcium and phosphorus revealed the dynamic changes during the oviposition cycle.

引文

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