鸡肠道Cajal间质细胞与肠神经系统的特性研究
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摘要
新近的观点认为正常的肠道蠕动依赖于肠壁中肠神经系统(Enteric Nervous System, ENS)、Cajal间质细胞(interstitial cells of cajal, ICC)和平滑肌细胞之间的相互作用。ICC是位于自主神经末梢和消化道平滑肌细胞之间的一类特殊的间质细胞,它在三个方面起作用：胃肠道平滑肌活动的起搏,推进电活动的传播和调节神经递质。以往关于ENS和ICC的研究主要集中在哺乳动物胃肠道。鸟类与哺乳动物的胃肠道在结构上存在一定差异性,同时,鸟类的肠神经系统不仅包括肠壁内的神经丛,还包括位于肠管外肠系膜缘的肠神经(又称Remak's nerve).这种差异是否导致ENS和ICC结构及其作用机制的不同?本研究以鸡作为试验动物,详细研究了鸡肠道内ICC的鉴定、分布以及与神经、平滑肌之间的联系,并对鸡NES中与ICC关系最为密切的NOS和AChE阳性神经元进行了胚后发育和分布比较研究。研究结果将从细胞和分子水平阐明鸡ENS和ICC的结构组成和相互联系,对揭示鸟类胃肠道功能有着重要意义,为鸟类胃肠动力学疾病治疗提供科学依据。
试验I鸡肠道Cajal间质细胞的超微结构研究本试验应用透射电镜系统观察了鸡肠道Cajal间质细胞(interstitial cells of cajal, ICC)的超微结构特点,结果显示在鸡肠道的不同部位均有ICC的存在,分布密度不等。根据分布位置的不同可将ICC分为肌间ICC、肌内ICC、粘膜下层ICC、深肌层ICC和固有层ICC。电镜下,这些ICC具有共同的超微特征：存在大量的线粒体、丰富的中间丝；细胞核呈梭形或椭圆形,内含大量异染色质。ICC胞体呈梭形或星形,有数量不等的胞质突起,这些突起与周围平滑肌细胞和肠道神经形成密切联系。部分ICC也显示了平滑肌细胞的一些结构特点,如不连续的基底膜、胞膜小凹和致密体。ICC之间,以及ICC与邻近的平滑肌细胞之间存在着明显的缝隙连接样结构。与哺乳动物不同,鸡回肠固有层也分布有ICC。在鸡肠道组织中,成纤维细胞的分布也较为丰富,它们与ICC的主要区别在于其具有发达的分泌性细胞器,包括囊泡和不连续的基底膜,但缺乏中间丝和胞膜小凹。鸡肠道ICC的分布、超微结构特点及其与其它细胞的联系表明,ICC在鸡消化道动力学机制中发挥着重要的调控作用。ICC在粘膜固有层的分布,提示鸡肠道ICC可能比哺乳类具有更多的亚型。
试验Ⅱ鸡肠道的C-kit阳性细胞表达与分布由于缺乏合适的标记物,Cajal间质细胞(interstitial cells of cajal, ICC)在禽类肠道的分布未见资料报道,本试验在第三章对鸡肠道ICC超微结构鉴定的基础上,应用合成的三相探针通过原位杂交方法,检测ICC的标记物c-kit mRNA在鸡肠道不同区域的分布规律；同时,应用实时荧光定量PCR(RT-PCR)分别对不同肠段粘膜组织和去粘膜肠壁组织的c-kit mRNA表达进行测定。原位杂交结果显示出两种类型的c-kit mRNA阳性细胞。第一种为纺锤形或星极细胞组成,主要位于内环形肌和外纵形肌之间的肌间层,粘膜下层、环形肌和纵形肌内也有发现。这种类型的细胞被界定为ICC。进一步根据此类细胞分布位置不同可分为肌间ICC、肌内ICC和粘膜下ICC三种,而且不同肠段内阳性细胞数目也不相同。第二种为圆形或颗粒型细胞组成,主要位于粘膜固有层内,粘膜下层也有少量发现,这类细胞被界定为肥大细胞,主要分布于回肠和空肠。RT-PCR结果显示在不同肠段均有c-kit mRNA的表达：在去粘膜肠壁组织中,回肠和结肠表达最高,空肠和十二指肠次之,盲肠最低；而在粘膜组织中,空肠表达最高,十二指肠和回肠其次,结肠和盲肠最低。实验结果揭示了鸡肠道ICC的分布模式,显示了鸡肠道不同部位中c-kitmRNA表达差异,这种差异可能与不同肠段的功能及其调节方式有关。
试验Ⅲ不同日龄鸡肠道c-kit与其配体SCF mRNA的表达研究本试验以β-actin为内参,采用SYBR染料法的实时荧光定量PCR (RT-PCR)技术。选取第1、10、20和40日龄三黄鸡去除粘膜后的十二指肠、空肠、回肠、盲肠和结肠肠壁中c-kit及其配体SCF mRNA的表达进行相对定量,初步阐明正常鸡肠道中c-kit及其配体SCF的mRNA表达量随着日龄变化的规律。结果显示所有肠段中c-kit mRNA的表达量从1日龄至40日龄逐渐降低,至20日龄时c-kit mRNA表达极显著低于1日龄。SCFmRNA的表达量从1日龄至10日龄逐渐升高,10日龄时mRNA表达最高,之后慢慢下降,至20日龄时的SCF mRNA表达显著低于10日龄。各肠段在不同日龄的c-kit和SCF mRNA表达量的变化表现如下：1日龄时,结肠中c-kit mRNA表达量最高,回肠最低；十二指肠中SCF mRNA表达量最高,盲肠最低。10日龄时,结肠中c-kit mRNA表达量最高,空肠最低；结肠中SCF mRNA表达量最高,盲肠最低。20至40日龄时,变化趋于稳定,都是回肠中c-kit和SCF mRNA表达量最高,盲肠最低。结果表明鸡肠道的胚后发育和功能形成是一个逐渐成熟的过程,而且c-kit及其配体SCF之间存在着密切的联系。
试验Ⅳ鸡肠道Vimentin与NSE免疫荧光双标研究本试验采用ICC的标记物之一波形蛋白(vimentin)和神经元的特异标记物神经元特异性烯醇化酶(Neuron-specific Enolase, NSE),对肠道切片进行免疫细胞化学双标,观察鸡不同肠段内vimentin、NSE阳性细胞的分布特征,探究ICC与神经之间的联系。结果发现：vimentin阳性细胞及其突起主要分布在肌层、粘膜下层和固有膜内,特别是在内环形肌和外纵形肌之间的肌间层大量出现。阳性细胞在大肠与小肠的肌内分布稍有区别,前者主要见于肌间隔内,而后者主要见于肌束内。大部分NSE阳性细胞分布于肌间层和粘膜下层,肌层内有零散分布；固有膜NSE阳性细胞量较少,但小肠固有膜vimentin阳性细胞明显多于大肠。粘膜上皮附近也能观察到。通过双标观察,肠道肌间和粘膜下层vimentin阳性细胞紧密围绕在NSE阳性细胞周围。固有膜内vimentin阳性细胞也多位于NSE阳性细胞周围。表明鸡vimentin细胞与NSE阳性细胞在肠道各段呈交错分布,两者关系紧密,提示它们在肠道活动中存在一定调节关系。
试验V鸡肠道肌间层NOS阳性神经元胚后发育的结构研究肠神经系统(Enteric Nervous System, ENS)的正常发育对了解肠道功能是非常重要的。鸡胚是研究肌间神经丛发育和ENS功能的理想模型,但关于鸡胚后肌间神经丛的发育未见报道。本研究的目的是定性和定量分析胚后发育期间鸡肌间神经丛NOS阳性神经元(即氮能神经元)的形态和分布。分别对7日龄、15日龄和成年鸡不同肠段(十二指肠、空肠、回肠、盲肠和结肠)进行肌间铺片制作,还原型尼克酰胺腺嘌呤二核苷酸黄递酶(NADPH-d)组织化学反应后,利用IPP6.0软件对阳性神经元和神经节进行定量分析。结果显示NOS阳性神经元和神经纤维形成典型的三级网状结构,NOS阳性神经元形态各异,聚集在一起构成大小不等的神经节。NOS阳性神经元反应强度存在一定差异,数个阳性神经元经常可形成串珠形和U形结构。NOS阳性神经元和神经节的密度随着年龄的增长而降低,但神经节中的阳性神经元的数量却有所增加。在所有年龄组中,NOS阳性神经元数量在结肠最高,随后是回肠,空肠,十二指肠和盲肠。结果表明,鸡肠道肌间神经丛存在胚后发育现象,提示鸡ENS功能形成是一个逐渐成熟和完善的过程；另外在每个肠段之间的发育并无明显差异,提示不同肠段的功能执行在出生后就已确定。
试验Ⅵ鸡回肠粘膜下层胆碱能神经元的胚后发育研究乙酰胆碱能神经元释放的乙酰胆碱(ACh)是胃肠道最主要的兴奋性神经递质,能刺激平滑肌收缩,增强胃肠道分泌,促进肠道激素的释放。本研究分别制作0、5、10、20、40日龄三黄鸡回肠粘膜下层铺片样品,通过乙酰胆碱酯酶(AChE)组织化学反应,探究0日龄到40日龄鸡肠道粘膜下神经丛胆碱能神经元的构筑及其变化。结果显示AChE阳性神经元和神经纤维形成立体的三级网状结构,AChE阳性神经元形态各异,聚集在一起构成大小不等的神经节。随着年龄的增长神经节密度明显降低,直到40日龄组。相反,神经节中神经元大小逐渐增加,40日龄组中几乎是0日龄组的3倍。在胚后发育初期,每个神经节中的神经元数量从10日龄起,基本呈现稳定上升的趋势。鸡回肠粘膜下神经丛胚后发育变化的研究结果,将为鸡的肠神经系统失调引起的胃肠道疾病诊断提供重要的病理组织学参考。
试验Ⅶ鸡回肠段胆碱能和氮能神经元的组织化学定位分别制作鸡回肠冰冻切片、肌间和粘膜下层铺片,应用乙酰胆碱酯酶(AChE)和还原型尼克酰胺腺嘌呤二核苷酸黄递酶(NADPH-d)组织化学反应,分别对胆碱能和氮能神经元进行定位分析。冰冻切片结果显示AChE和NADPH-d阳性反应广泛分布于神经元胞体和神经纤维中。阳性神经元呈不规则或多边形,主要在肌间和粘膜下层零星或成簇出现,肌内也有一定的分布。肌间阳性神经纤维可明显插入环形肌内,阳性神经纤维经常围绕在血管周围,也有一些阳性神经纤维从肌层发出,穿过粘膜下层进入粘膜固有层,并且渗透到肠绒毛上皮下；肌内和粘膜中的AChE神经纤维的数量远超过NADPH-d阳性神经元和神经纤维。铺片结果显示AChE和NADPH-d阳性神经元位于神经节中,神经节之间通过阳性纤维相连形成致密的网络样结构,粘膜下网络明显比肌间密集,粘膜下神经节密度远大于肌间,但粘膜下每个神经节中的神经元数目要远小于肌间,粘膜下层神经元的大小也较肌间的小。AChE阳性神经元数目要远多于NADPH-d阳性神经元数目。结果表明鸡回肠广泛分布有兴奋性的胆碱能和抑制性的氮能神经元,可能在胃肠道功能调节中发挥重要作用。
At present, it is generally accepted that the normal intestinal peristalsis is dependent on the interaction among the enteric nervous system (ENS), the interstitial cells of Cajal (ICC) and smooth muscle cells. ICC is a special class of interstitial cells located between the autonomous nerve endings and gastrointestinal smooth muscle cells. Works mainly in three aspects:The ICC generate pacemaker activity throughout the gastrointestinal tract by initiating the slow wave activity that is integral to the coordination of gastrointestinal motility; In addition, ICC are thought to be involved in mediating neurotransmission and facilitating active propagation of electrical events.
In the past, the studies on ENS and ICC are mainly concentrated in the mammals' gastrointestinal tract. The structure between birds and mammals is different, and the ENS of avian contains neurons that are in the wall of the gastrointestinal tract, as well as the intestinal nerve of Remak (INR) located in the mesenteric edge outside of the intestinal wall. Would this variation lead to the difference of the structure and mechanism of interactions between the ENS and the ICC? Chicken was selected as experimental animals on behalf of birds for detail researches of gastrointestinal tract, such as the identification and distribution of ICC, the relations between ICC and nerves and smooth muscle. In addition, the comparative study on the postnatal development and distribution of NOS and AChE positive neurons were revealed, because the two types of neurons were closely related with ICC in intestine. Those researches explain the structure and relation of cells and molecule in the ENS and ICC from chicken intestine, and analyze the functional significance of the ENS and ICC. This study provide basic datum for the works of the function and the protection of diseases in chicken gastrointestinal tract.
Experiment I Identification and distribution of interstitial cells of cajal in the intestine of chicken The present study has observed the ultrastructural feature of ICC in the chicken intestine by transmission electron microscopy (TEM). The results showed ICC could be found in different regions and the numbers were different. They were classified into several subtypes:ICC at the layer of myenteric plexus、ICC at the intramuscular level、 ICC in the deep muscular plexus、ICC on the submucosal plexus and ICC in the lamina propria mucosae. This cells share common ultrastructural features, such as the presence of numerous mitochondria, abundant intermediate filaments, their nuclei were fusiform, oval or indented with a dense band of peripheral heterochromatin, they also form close contacts with neighbouring SMC (SMC) and with enteric nerve. They are spindle-shaped or stellate, having a different number of ramified cell processes. In addition, some ICC also shows some typical features of SMC including a basal lamina, caveolae, and dense bodies. Ture gap junctions have been detected between with other ICC or with neighbouring SMC. We also identified a possible new subtype of ICC located in the lamina propria mucosae, which was different with mammals. Fibroblast-like cells (FLC) were abundant showing well-developed secretory organelles, including coated vesicles, a patchy basal lamina, but lacked prominent intermediate filaments and caveolae. The widespread distribution of ICC or equivalent cells in chicken intestine, together with the conservation of their ultrastructural features, suggests that they play same key regulatory roles in gastrointestinal movement with mammals. The observation of ICC in the lamina propria mucosae indicates that there may be more subtypes of these cells in the intestine than previously considered.
Experiment II The identification and expression of c-kit positive cells in the intestine of chicken The distribution of the interstitial cells of Cajal (ICC) remains obscure because of lacking suitable marker. In the present study on the basis of the third chapter about the ultrastructure identification of ICC in chicken intestine, the identification and expression of c-kit positive cells in the chicken intestine were demonstrated by means of in situ hybridization (ISH) histochemistry, and the expression of c-kit gene in the intestinal wall lacking mucosa and mucosa of normal chickens were also revealed by real-time fluorescence quantitative polymerase chain reaction (RT-PCR). Two types of cells stained positive for c-kit mRNA. The first group consisted of spindle-shaped or bipolar cells identified as ICC. ICC were found at a variety of locations-at the level of the myenteric plexus between the circular and longitudinal muscle, and intermingled with smooth muscle cells within muscle bundles in the circular and longitudinal muscle layers. ICC was also identified along the submucosal layer; they were classified into several subtypes. The second group was comprised of round-shaped cells, which resembled mast cells. Mast cells were mainly found in the lamina propria region as well as in submucosal layer. RT-PCR revealed the expression of c-kit mRNA throughout the intestinal wall lacking mucosa and mucosa of the total intestine. In intestinal wall lacking mucosa, c-kit mRNA was detected at high levels in all samples of the ileum and colon, moderate levels in the duodenum and jejunem, low levels in the caeca; in mucosa, c-kit mRNA was detected at high levels in all samples of the jejunum, following by ileum and duodenum. While the levels of mRNA in the caeca and colon was low. Conclusively, this study reveals for the first time the distribution of ICC, quantifies the expression of c-kit mRNA in the intestine of chicken and also compares the c-kit positive cell types on morphologically. The results indicate the difference may be related to different bowel functions.
Experiment Ⅲ The study on c-kit and SCF mRNA expression in chicken intestine of different ages Tissue samples (the intestinal wall lacking mucosa of duodenum, jejunum, ileum, caeca, and colon) were made in1-days-old,10-days-old,20-days-old and adult (40-days-old) chickens (n=5) and identify the expression of c-kit and its ligand SCF mRNA in this tissues by real-time fluorescence quantitative polymerase chain reaction (RT-PCR) with SYBR green and were normalized against the housekeeping gene β-actin. The results showed that the expression patterns were different between c-kit and SCF genes. The expression of c-kit mRNA was decreased in different intestinal segments with ages. In1-days-old group, c-kit mRNA was detected at highest levels, the expression of c-kit mRNA detected in20-days-old and adult group were extremely significantly lower than that of1-days-old group. The expression of SCF mRNA was increased from1-days-old to10-days-old, and then the tendency was descend. The expression of SCF mRNA detected in20-days-old and adult group were significantly lower than that of10-days-old group. In different age groups, the expression of c-kit and SCF mRNA showed a dynamic change. In1-days-old group, c-kit mRNA was detected at highest level in colon and lowest level in caeca, SCF mRNA was detected at highest level in duodenum and lowest level in caeca. In10-days-old group, c-kit mRNA was detected at highest level in colon and lowest level in jejunum, SCF mRNA was detected at highest level in colon and lowest level in caeca. But in20-days-old and adult group, the change was similar, SCF and c-kit mRNA was detected at highest level in ileum and lowest level in caeca. Conclusively, this study reveals it is a close relationship between SCF and c-kit and indicates the functional maturation in development of postnatal intestine is a gradually process.
Experiment Ⅳ Double-labeled immunofluorescence study on vimentin and NSE protein in chicken intestine Different intestinal segments were used in this study in order to investigate the morphological relationship between the interstitial cells of Cajal and nerve using immunofluorescent study on vimentin and Neuron-specific Enolase (NSE) by double-labeled immunofluorescence combined laser-scanning confocal microscopy, and distribution of the two positive cells were analyzed. The results showed vimentin positive cells and their processes were mainly found within lamina muscularis, submucosal layer and lamina propria, specially in he myenteric plexus between the circular and longitudinal muscle. However, vimentin positive cells were distributed in the septum of lamina muscularis of the large intestine and within the muscle bundles of small intestine. NSE positive cells mainly located in the myenteric plexus between the circular and longitudinal muscle and submucosal layer, and scattered within muscle bundles in the circular and longitudinal muscle layers. NSE positive cells were lower in mucosa than that of lamina muscularis, and these cells were more in small intestinal mucosa than that of large intestine; these cells were also found near mucosal epithelium. We found the vimentin positive cells closely surround the NSE positive cells in lamina muscularis and submucosal layer by double-labelling, and similar relations seen in mucosa. Conclusively, the vimentin and NSE positive cells form close contacts with each other, suggesting they play an important biological role in the intestinal motility.
Experiment V The structural study on NOS positive neurons in myenteric plexus of the chicken intestine during postnatal development Information regarding the development of the enteric nervous system (ENS) is important for understanding the intestinal function. Because fertilized chicken eggs provide easy access to embryos chicken model have been widely used to study developing fetal myenteric plexus, however no study has been focused on the postnatal period. The aim of this study was to perform a qualitative and quantitative analysis of NOS positive cells (the nitrergic neurons) in the myenteric plexus of developing chickens in the postnatal period. Whole-mount preparations of the myenteric plexus were made in7-days-old,15-days-old and adult chickens (n=15). The myenteric plexus was studied after nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d) histochemistry using light microscopy, digital photography and IPP6.0software. The number of positively stained neurons and ganglia was counted in the duodenum, jejunum, ileum, cecum and colon in the different age groups. The results showed the nerve fiber bundles connecting the ganglia form a clear meshwork. The positively stained neurons showed various morphology, staining intensity and formed bead-shaped and U-shaped arrangements in the myenteric plexus. The number of positively stained neurons per area and the number of ganglia per area decreased with age; however the number of positive neurons per ganglia increased. The number of NADPH-d positive neurons was highest in the colon, following by the ileum, the jejunum, the duodenum and the cecum in all age groups. Conclusively, Developmental changes in the myenteric plexus of chickens continue in the postnatal period, this indicates that the formation of function needs a gradual process of maturation and improvement. In addition, no significant difference is seen among different intestinal segments during postnatal development, suggesting that the function of different intestinal segments had been determined after birth.
Experiment VI The study on morphology and distribution of AChE positive neurons in submucosal plexus of the chicken ileum Acetylcholine is recognized as a potent mediator of excitatory nerves responsible for the constriction of the smooth muscle of the gastrointestinal tract, enhance the secretion of the gastrointestinal tract, and stimulate the release of gut hormones. The aim of this study was to determine the normal cholinergic neuronal density and morphology in the submucosal plexus of the chicken ileum from0--days-old to40-days-old. In this experiment, whole-mount preparations of the submucosal plexus of chicken ileum were made in0-days-old,5-days-old,10-days-old,20-days-old and40-days-old chickens. The submucosal plexus was studied after acetylcholine esterase (AChE) histochemistry. The results showed that the positively stained neurons showed various morphology, staining intensity and formed a clear network by ganglia and nerve fibers. The number of ganglia per area decreased with age, however the size of positive neurons increased, and the size in40-days-old group was3times than that of0-days-old group. The change of number of positive neurons per ganglia was dynamic, but the tendency was stably increased from10-days-old to adult. Conclusively, developmental changes in the submucosal plexus of chickens was marked in the postnatal period by qualitative and quantitative analysis, this study will provide an important histopathological reference on the diagnosis of causes of enteric nervous system disorders.
Experiment Ⅶ Histochemical localization of the cholinergic and nitrergic neurons in the chicken ileum Histochemical localization and analysis of the cholinergic and nitrergic neurons in the chicken ileum were investigated by staining with acetylcholine esterase (AChE) and nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d), respectively. AChE and NADPH-d activity was demonstrated in neuronal cell bodies and nerve fibers in the chicken ileum. The positive neurons showed irregular or polygonal shape and were mainly present sporadic or clumped in the myenteric and submucosal plexus. The positive nerve fibers frequently surrounded the ileac blood vessels. They were abundantly present in myenteric and submucosal plexus of the ileum forming a network. Some positive nerve fibers traversed the submucosa into the lamina propria mucosae. Fine nerve fibers were found to penetrate into intestinal villi underneath the epithelium. Extensive networks of more intensely staining AChE positive nerve fibers were present in the mucosa as compared to that of NADPH-d positive fibers. Ganglia density of submucosal plexus was markedly higher than that of myenteric plexus, whereas neurons per ganglion and the number of neurons per mm2and the size of neurons of submucosal plexus were shorter than that of myenteric plexus. In addition, the number of AChE positive neurons was more than that of NADPH-d positive neurons. We concluded that the chicken ileum is characterized by abundance of nerve structures which may play a significant functional role in ileum of the chicken.
试验I鸡肠道Cajal间质细胞的超微结构研究本试验应用透射电镜系统观察了鸡肠道Cajal间质细胞(interstitial cells of cajal, ICC)的超微结构特点,结果显示在鸡肠道的不同部位均有ICC的存在,分布密度不等。根据分布位置的不同可将ICC分为肌间ICC、肌内ICC、粘膜下层ICC、深肌层ICC和固有层ICC。电镜下,这些ICC具有共同的超微特征：存在大量的线粒体、丰富的中间丝；细胞核呈梭形或椭圆形,内含大量异染色质。ICC胞体呈梭形或星形,有数量不等的胞质突起,这些突起与周围平滑肌细胞和肠道神经形成密切联系。部分ICC也显示了平滑肌细胞的一些结构特点,如不连续的基底膜、胞膜小凹和致密体。ICC之间,以及ICC与邻近的平滑肌细胞之间存在着明显的缝隙连接样结构。与哺乳动物不同,鸡回肠固有层也分布有ICC。在鸡肠道组织中,成纤维细胞的分布也较为丰富,它们与ICC的主要区别在于其具有发达的分泌性细胞器,包括囊泡和不连续的基底膜,但缺乏中间丝和胞膜小凹。鸡肠道ICC的分布、超微结构特点及其与其它细胞的联系表明,ICC在鸡消化道动力学机制中发挥着重要的调控作用。ICC在粘膜固有层的分布,提示鸡肠道ICC可能比哺乳类具有更多的亚型。
试验Ⅱ鸡肠道的C-kit阳性细胞表达与分布由于缺乏合适的标记物,Cajal间质细胞(interstitial cells of cajal, ICC)在禽类肠道的分布未见资料报道,本试验在第三章对鸡肠道ICC超微结构鉴定的基础上,应用合成的三相探针通过原位杂交方法,检测ICC的标记物c-kit mRNA在鸡肠道不同区域的分布规律；同时,应用实时荧光定量PCR(RT-PCR)分别对不同肠段粘膜组织和去粘膜肠壁组织的c-kit mRNA表达进行测定。原位杂交结果显示出两种类型的c-kit mRNA阳性细胞。第一种为纺锤形或星极细胞组成,主要位于内环形肌和外纵形肌之间的肌间层,粘膜下层、环形肌和纵形肌内也有发现。这种类型的细胞被界定为ICC。进一步根据此类细胞分布位置不同可分为肌间ICC、肌内ICC和粘膜下ICC三种,而且不同肠段内阳性细胞数目也不相同。第二种为圆形或颗粒型细胞组成,主要位于粘膜固有层内,粘膜下层也有少量发现,这类细胞被界定为肥大细胞,主要分布于回肠和空肠。RT-PCR结果显示在不同肠段均有c-kit mRNA的表达：在去粘膜肠壁组织中,回肠和结肠表达最高,空肠和十二指肠次之,盲肠最低；而在粘膜组织中,空肠表达最高,十二指肠和回肠其次,结肠和盲肠最低。实验结果揭示了鸡肠道ICC的分布模式,显示了鸡肠道不同部位中c-kitmRNA表达差异,这种差异可能与不同肠段的功能及其调节方式有关。
试验Ⅲ不同日龄鸡肠道c-kit与其配体SCF mRNA的表达研究本试验以β-actin为内参,采用SYBR染料法的实时荧光定量PCR (RT-PCR)技术。选取第1、10、20和40日龄三黄鸡去除粘膜后的十二指肠、空肠、回肠、盲肠和结肠肠壁中c-kit及其配体SCF mRNA的表达进行相对定量,初步阐明正常鸡肠道中c-kit及其配体SCF的mRNA表达量随着日龄变化的规律。结果显示所有肠段中c-kit mRNA的表达量从1日龄至40日龄逐渐降低,至20日龄时c-kit mRNA表达极显著低于1日龄。SCFmRNA的表达量从1日龄至10日龄逐渐升高,10日龄时mRNA表达最高,之后慢慢下降,至20日龄时的SCF mRNA表达显著低于10日龄。各肠段在不同日龄的c-kit和SCF mRNA表达量的变化表现如下：1日龄时,结肠中c-kit mRNA表达量最高,回肠最低；十二指肠中SCF mRNA表达量最高,盲肠最低。10日龄时,结肠中c-kit mRNA表达量最高,空肠最低；结肠中SCF mRNA表达量最高,盲肠最低。20至40日龄时,变化趋于稳定,都是回肠中c-kit和SCF mRNA表达量最高,盲肠最低。结果表明鸡肠道的胚后发育和功能形成是一个逐渐成熟的过程,而且c-kit及其配体SCF之间存在着密切的联系。
试验Ⅳ鸡肠道Vimentin与NSE免疫荧光双标研究本试验采用ICC的标记物之一波形蛋白(vimentin)和神经元的特异标记物神经元特异性烯醇化酶(Neuron-specific Enolase, NSE),对肠道切片进行免疫细胞化学双标,观察鸡不同肠段内vimentin、NSE阳性细胞的分布特征,探究ICC与神经之间的联系。结果发现：vimentin阳性细胞及其突起主要分布在肌层、粘膜下层和固有膜内,特别是在内环形肌和外纵形肌之间的肌间层大量出现。阳性细胞在大肠与小肠的肌内分布稍有区别,前者主要见于肌间隔内,而后者主要见于肌束内。大部分NSE阳性细胞分布于肌间层和粘膜下层,肌层内有零散分布；固有膜NSE阳性细胞量较少,但小肠固有膜vimentin阳性细胞明显多于大肠。粘膜上皮附近也能观察到。通过双标观察,肠道肌间和粘膜下层vimentin阳性细胞紧密围绕在NSE阳性细胞周围。固有膜内vimentin阳性细胞也多位于NSE阳性细胞周围。表明鸡vimentin细胞与NSE阳性细胞在肠道各段呈交错分布,两者关系紧密,提示它们在肠道活动中存在一定调节关系。
试验V鸡肠道肌间层NOS阳性神经元胚后发育的结构研究肠神经系统(Enteric Nervous System, ENS)的正常发育对了解肠道功能是非常重要的。鸡胚是研究肌间神经丛发育和ENS功能的理想模型,但关于鸡胚后肌间神经丛的发育未见报道。本研究的目的是定性和定量分析胚后发育期间鸡肌间神经丛NOS阳性神经元(即氮能神经元)的形态和分布。分别对7日龄、15日龄和成年鸡不同肠段(十二指肠、空肠、回肠、盲肠和结肠)进行肌间铺片制作,还原型尼克酰胺腺嘌呤二核苷酸黄递酶(NADPH-d)组织化学反应后,利用IPP6.0软件对阳性神经元和神经节进行定量分析。结果显示NOS阳性神经元和神经纤维形成典型的三级网状结构,NOS阳性神经元形态各异,聚集在一起构成大小不等的神经节。NOS阳性神经元反应强度存在一定差异,数个阳性神经元经常可形成串珠形和U形结构。NOS阳性神经元和神经节的密度随着年龄的增长而降低,但神经节中的阳性神经元的数量却有所增加。在所有年龄组中,NOS阳性神经元数量在结肠最高,随后是回肠,空肠,十二指肠和盲肠。结果表明,鸡肠道肌间神经丛存在胚后发育现象,提示鸡ENS功能形成是一个逐渐成熟和完善的过程；另外在每个肠段之间的发育并无明显差异,提示不同肠段的功能执行在出生后就已确定。
试验Ⅵ鸡回肠粘膜下层胆碱能神经元的胚后发育研究乙酰胆碱能神经元释放的乙酰胆碱(ACh)是胃肠道最主要的兴奋性神经递质,能刺激平滑肌收缩,增强胃肠道分泌,促进肠道激素的释放。本研究分别制作0、5、10、20、40日龄三黄鸡回肠粘膜下层铺片样品,通过乙酰胆碱酯酶(AChE)组织化学反应,探究0日龄到40日龄鸡肠道粘膜下神经丛胆碱能神经元的构筑及其变化。结果显示AChE阳性神经元和神经纤维形成立体的三级网状结构,AChE阳性神经元形态各异,聚集在一起构成大小不等的神经节。随着年龄的增长神经节密度明显降低,直到40日龄组。相反,神经节中神经元大小逐渐增加,40日龄组中几乎是0日龄组的3倍。在胚后发育初期,每个神经节中的神经元数量从10日龄起,基本呈现稳定上升的趋势。鸡回肠粘膜下神经丛胚后发育变化的研究结果,将为鸡的肠神经系统失调引起的胃肠道疾病诊断提供重要的病理组织学参考。
试验Ⅶ鸡回肠段胆碱能和氮能神经元的组织化学定位分别制作鸡回肠冰冻切片、肌间和粘膜下层铺片,应用乙酰胆碱酯酶(AChE)和还原型尼克酰胺腺嘌呤二核苷酸黄递酶(NADPH-d)组织化学反应,分别对胆碱能和氮能神经元进行定位分析。冰冻切片结果显示AChE和NADPH-d阳性反应广泛分布于神经元胞体和神经纤维中。阳性神经元呈不规则或多边形,主要在肌间和粘膜下层零星或成簇出现,肌内也有一定的分布。肌间阳性神经纤维可明显插入环形肌内,阳性神经纤维经常围绕在血管周围,也有一些阳性神经纤维从肌层发出,穿过粘膜下层进入粘膜固有层,并且渗透到肠绒毛上皮下；肌内和粘膜中的AChE神经纤维的数量远超过NADPH-d阳性神经元和神经纤维。铺片结果显示AChE和NADPH-d阳性神经元位于神经节中,神经节之间通过阳性纤维相连形成致密的网络样结构,粘膜下网络明显比肌间密集,粘膜下神经节密度远大于肌间,但粘膜下每个神经节中的神经元数目要远小于肌间,粘膜下层神经元的大小也较肌间的小。AChE阳性神经元数目要远多于NADPH-d阳性神经元数目。结果表明鸡回肠广泛分布有兴奋性的胆碱能和抑制性的氮能神经元,可能在胃肠道功能调节中发挥重要作用。
At present, it is generally accepted that the normal intestinal peristalsis is dependent on the interaction among the enteric nervous system (ENS), the interstitial cells of Cajal (ICC) and smooth muscle cells. ICC is a special class of interstitial cells located between the autonomous nerve endings and gastrointestinal smooth muscle cells. Works mainly in three aspects:The ICC generate pacemaker activity throughout the gastrointestinal tract by initiating the slow wave activity that is integral to the coordination of gastrointestinal motility; In addition, ICC are thought to be involved in mediating neurotransmission and facilitating active propagation of electrical events.
In the past, the studies on ENS and ICC are mainly concentrated in the mammals' gastrointestinal tract. The structure between birds and mammals is different, and the ENS of avian contains neurons that are in the wall of the gastrointestinal tract, as well as the intestinal nerve of Remak (INR) located in the mesenteric edge outside of the intestinal wall. Would this variation lead to the difference of the structure and mechanism of interactions between the ENS and the ICC? Chicken was selected as experimental animals on behalf of birds for detail researches of gastrointestinal tract, such as the identification and distribution of ICC, the relations between ICC and nerves and smooth muscle. In addition, the comparative study on the postnatal development and distribution of NOS and AChE positive neurons were revealed, because the two types of neurons were closely related with ICC in intestine. Those researches explain the structure and relation of cells and molecule in the ENS and ICC from chicken intestine, and analyze the functional significance of the ENS and ICC. This study provide basic datum for the works of the function and the protection of diseases in chicken gastrointestinal tract.
Experiment I Identification and distribution of interstitial cells of cajal in the intestine of chicken The present study has observed the ultrastructural feature of ICC in the chicken intestine by transmission electron microscopy (TEM). The results showed ICC could be found in different regions and the numbers were different. They were classified into several subtypes:ICC at the layer of myenteric plexus、ICC at the intramuscular level、 ICC in the deep muscular plexus、ICC on the submucosal plexus and ICC in the lamina propria mucosae. This cells share common ultrastructural features, such as the presence of numerous mitochondria, abundant intermediate filaments, their nuclei were fusiform, oval or indented with a dense band of peripheral heterochromatin, they also form close contacts with neighbouring SMC (SMC) and with enteric nerve. They are spindle-shaped or stellate, having a different number of ramified cell processes. In addition, some ICC also shows some typical features of SMC including a basal lamina, caveolae, and dense bodies. Ture gap junctions have been detected between with other ICC or with neighbouring SMC. We also identified a possible new subtype of ICC located in the lamina propria mucosae, which was different with mammals. Fibroblast-like cells (FLC) were abundant showing well-developed secretory organelles, including coated vesicles, a patchy basal lamina, but lacked prominent intermediate filaments and caveolae. The widespread distribution of ICC or equivalent cells in chicken intestine, together with the conservation of their ultrastructural features, suggests that they play same key regulatory roles in gastrointestinal movement with mammals. The observation of ICC in the lamina propria mucosae indicates that there may be more subtypes of these cells in the intestine than previously considered.
Experiment II The identification and expression of c-kit positive cells in the intestine of chicken The distribution of the interstitial cells of Cajal (ICC) remains obscure because of lacking suitable marker. In the present study on the basis of the third chapter about the ultrastructure identification of ICC in chicken intestine, the identification and expression of c-kit positive cells in the chicken intestine were demonstrated by means of in situ hybridization (ISH) histochemistry, and the expression of c-kit gene in the intestinal wall lacking mucosa and mucosa of normal chickens were also revealed by real-time fluorescence quantitative polymerase chain reaction (RT-PCR). Two types of cells stained positive for c-kit mRNA. The first group consisted of spindle-shaped or bipolar cells identified as ICC. ICC were found at a variety of locations-at the level of the myenteric plexus between the circular and longitudinal muscle, and intermingled with smooth muscle cells within muscle bundles in the circular and longitudinal muscle layers. ICC was also identified along the submucosal layer; they were classified into several subtypes. The second group was comprised of round-shaped cells, which resembled mast cells. Mast cells were mainly found in the lamina propria region as well as in submucosal layer. RT-PCR revealed the expression of c-kit mRNA throughout the intestinal wall lacking mucosa and mucosa of the total intestine. In intestinal wall lacking mucosa, c-kit mRNA was detected at high levels in all samples of the ileum and colon, moderate levels in the duodenum and jejunem, low levels in the caeca; in mucosa, c-kit mRNA was detected at high levels in all samples of the jejunum, following by ileum and duodenum. While the levels of mRNA in the caeca and colon was low. Conclusively, this study reveals for the first time the distribution of ICC, quantifies the expression of c-kit mRNA in the intestine of chicken and also compares the c-kit positive cell types on morphologically. The results indicate the difference may be related to different bowel functions.
Experiment Ⅲ The study on c-kit and SCF mRNA expression in chicken intestine of different ages Tissue samples (the intestinal wall lacking mucosa of duodenum, jejunum, ileum, caeca, and colon) were made in1-days-old,10-days-old,20-days-old and adult (40-days-old) chickens (n=5) and identify the expression of c-kit and its ligand SCF mRNA in this tissues by real-time fluorescence quantitative polymerase chain reaction (RT-PCR) with SYBR green and were normalized against the housekeeping gene β-actin. The results showed that the expression patterns were different between c-kit and SCF genes. The expression of c-kit mRNA was decreased in different intestinal segments with ages. In1-days-old group, c-kit mRNA was detected at highest levels, the expression of c-kit mRNA detected in20-days-old and adult group were extremely significantly lower than that of1-days-old group. The expression of SCF mRNA was increased from1-days-old to10-days-old, and then the tendency was descend. The expression of SCF mRNA detected in20-days-old and adult group were significantly lower than that of10-days-old group. In different age groups, the expression of c-kit and SCF mRNA showed a dynamic change. In1-days-old group, c-kit mRNA was detected at highest level in colon and lowest level in caeca, SCF mRNA was detected at highest level in duodenum and lowest level in caeca. In10-days-old group, c-kit mRNA was detected at highest level in colon and lowest level in jejunum, SCF mRNA was detected at highest level in colon and lowest level in caeca. But in20-days-old and adult group, the change was similar, SCF and c-kit mRNA was detected at highest level in ileum and lowest level in caeca. Conclusively, this study reveals it is a close relationship between SCF and c-kit and indicates the functional maturation in development of postnatal intestine is a gradually process.
Experiment Ⅳ Double-labeled immunofluorescence study on vimentin and NSE protein in chicken intestine Different intestinal segments were used in this study in order to investigate the morphological relationship between the interstitial cells of Cajal and nerve using immunofluorescent study on vimentin and Neuron-specific Enolase (NSE) by double-labeled immunofluorescence combined laser-scanning confocal microscopy, and distribution of the two positive cells were analyzed. The results showed vimentin positive cells and their processes were mainly found within lamina muscularis, submucosal layer and lamina propria, specially in he myenteric plexus between the circular and longitudinal muscle. However, vimentin positive cells were distributed in the septum of lamina muscularis of the large intestine and within the muscle bundles of small intestine. NSE positive cells mainly located in the myenteric plexus between the circular and longitudinal muscle and submucosal layer, and scattered within muscle bundles in the circular and longitudinal muscle layers. NSE positive cells were lower in mucosa than that of lamina muscularis, and these cells were more in small intestinal mucosa than that of large intestine; these cells were also found near mucosal epithelium. We found the vimentin positive cells closely surround the NSE positive cells in lamina muscularis and submucosal layer by double-labelling, and similar relations seen in mucosa. Conclusively, the vimentin and NSE positive cells form close contacts with each other, suggesting they play an important biological role in the intestinal motility.
Experiment V The structural study on NOS positive neurons in myenteric plexus of the chicken intestine during postnatal development Information regarding the development of the enteric nervous system (ENS) is important for understanding the intestinal function. Because fertilized chicken eggs provide easy access to embryos chicken model have been widely used to study developing fetal myenteric plexus, however no study has been focused on the postnatal period. The aim of this study was to perform a qualitative and quantitative analysis of NOS positive cells (the nitrergic neurons) in the myenteric plexus of developing chickens in the postnatal period. Whole-mount preparations of the myenteric plexus were made in7-days-old,15-days-old and adult chickens (n=15). The myenteric plexus was studied after nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d) histochemistry using light microscopy, digital photography and IPP6.0software. The number of positively stained neurons and ganglia was counted in the duodenum, jejunum, ileum, cecum and colon in the different age groups. The results showed the nerve fiber bundles connecting the ganglia form a clear meshwork. The positively stained neurons showed various morphology, staining intensity and formed bead-shaped and U-shaped arrangements in the myenteric plexus. The number of positively stained neurons per area and the number of ganglia per area decreased with age; however the number of positive neurons per ganglia increased. The number of NADPH-d positive neurons was highest in the colon, following by the ileum, the jejunum, the duodenum and the cecum in all age groups. Conclusively, Developmental changes in the myenteric plexus of chickens continue in the postnatal period, this indicates that the formation of function needs a gradual process of maturation and improvement. In addition, no significant difference is seen among different intestinal segments during postnatal development, suggesting that the function of different intestinal segments had been determined after birth.
Experiment VI The study on morphology and distribution of AChE positive neurons in submucosal plexus of the chicken ileum Acetylcholine is recognized as a potent mediator of excitatory nerves responsible for the constriction of the smooth muscle of the gastrointestinal tract, enhance the secretion of the gastrointestinal tract, and stimulate the release of gut hormones. The aim of this study was to determine the normal cholinergic neuronal density and morphology in the submucosal plexus of the chicken ileum from0--days-old to40-days-old. In this experiment, whole-mount preparations of the submucosal plexus of chicken ileum were made in0-days-old,5-days-old,10-days-old,20-days-old and40-days-old chickens. The submucosal plexus was studied after acetylcholine esterase (AChE) histochemistry. The results showed that the positively stained neurons showed various morphology, staining intensity and formed a clear network by ganglia and nerve fibers. The number of ganglia per area decreased with age, however the size of positive neurons increased, and the size in40-days-old group was3times than that of0-days-old group. The change of number of positive neurons per ganglia was dynamic, but the tendency was stably increased from10-days-old to adult. Conclusively, developmental changes in the submucosal plexus of chickens was marked in the postnatal period by qualitative and quantitative analysis, this study will provide an important histopathological reference on the diagnosis of causes of enteric nervous system disorders.
Experiment Ⅶ Histochemical localization of the cholinergic and nitrergic neurons in the chicken ileum Histochemical localization and analysis of the cholinergic and nitrergic neurons in the chicken ileum were investigated by staining with acetylcholine esterase (AChE) and nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d), respectively. AChE and NADPH-d activity was demonstrated in neuronal cell bodies and nerve fibers in the chicken ileum. The positive neurons showed irregular or polygonal shape and were mainly present sporadic or clumped in the myenteric and submucosal plexus. The positive nerve fibers frequently surrounded the ileac blood vessels. They were abundantly present in myenteric and submucosal plexus of the ileum forming a network. Some positive nerve fibers traversed the submucosa into the lamina propria mucosae. Fine nerve fibers were found to penetrate into intestinal villi underneath the epithelium. Extensive networks of more intensely staining AChE positive nerve fibers were present in the mucosa as compared to that of NADPH-d positive fibers. Ganglia density of submucosal plexus was markedly higher than that of myenteric plexus, whereas neurons per ganglion and the number of neurons per mm2and the size of neurons of submucosal plexus were shorter than that of myenteric plexus. In addition, the number of AChE positive neurons was more than that of NADPH-d positive neurons. We concluded that the chicken ileum is characterized by abundance of nerve structures which may play a significant functional role in ileum of the chicken.
引文
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